Fern Grower’s Manual

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Fern Grower’s
Manual
Revised and Expanded Edition

Barbara Joe Hoshizaki and

Robbin C. Moran

TIMBER PRESS
Portland, Oregon
The in-text Adiantum line drawings are by Lynn Vander Velde; unless otherwise
credited in the captions, all other line drawings are by Barbara Joe Hoshizaki.

Unless otherwise credited in the captions, the in-text black-and-white photographs are
by Arthur Takayama.

Unless otherwise credited in the captions, the color plates are by Barbara Joe Hoshizaki.

Copyright © 2001 by Barbara Joe Hoshizaki and Robbin C. Moran. All rights reserved.

Published in 2001 by
Timber Press, Inc.
The Haseltine Building
133 S.W. Second Avenue, Suite 450
Portland, Oregon 97204, U.S.A.

Printed in Hong Kong

Library of Congress Cataloging-in-Publication Data

Hoshizaki, Barbara Joe.

Fern grower’s manual / Barbara Joe Hoshizaki and Robbin C. Moran.—
Rev. and expanded ed.
p. cm.
Includes bibliographical references (p. ).
ISBN 0-88192-495-4
1. Ferns, Ornamental. I. Moran, Robbin Craig, 1956– II. Title.

SB429. H64 2001

635.9′373—dc21
00-059999
 Contents

Preface 7

1. About Ferns 11
2. The Structure of Ferns 13
3. Obtaining Ferns 22
4. Cultural Needs 25
5. Soils and Fertilizers 37
6. Through the Year with Ferns 48
7. Planting 51
8. Propagation 63
9. Landscaping 87
10. Growing Special Ferns 99
11. Troubles with Growing Ferns 118
12. How Ferns Get Their Names 140
13. Ferns and Fern Allies in Cultivation 151

Appendix I: Measuring Light 541

Appendix II: Fern Societies 542
Appendix III: Importing Ferns 543
Appendix IV: Names of Pest and Disease Control Substances 547
Appendix V: Family Classification of Fern Genera Treated in the Text 553
Glossary 558
Literature Cited 565
Subject Index 573
Plant Name Index 581

Color plates, including USDA Plant Hardiness Zone Map, follow page 240

5
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 Preface

Ferns continue to hold the interest of gardeners as we enter a new millennium. After the Victorian fern
craze, interest in ferns focused mainly on the native species and some of those common in the trade. After
World War II, interest in exotic ferns gradually took hold, and this fostered the formation of new fern
societies and the offering of a greater selection in the trade.
It has been more than 25 years since the first edition of Fern Grower’s Manual was published (Hoshizaki
1975). At the time, the author thought it would interest the horticultural public for only a few years before
ferns would fall out of vogue. She was wrong. A strong interest in ferns continues today as evidenced by the
great increase in fern societies all over the world for both amateurs and professional botanists. Also, the
number of fern species in cultivation has increased tremendously. The contributions of fern botanists in
writing floras and monographs, clarifying species, and supporting amateur interest are important factors in
this growth. The horticultural area has also made great contributions, particularly in pest control, tissue
culture, plug culture, new products, and determining the growing needs of certain ferns.
All this interest in ferns has created a need for a revised edition of Fern Grower’s Manual—one that
updates the information on pest control, propagation, new materials and products, cultural requirements,
and many other topics. But unlike other recent fern books, this revised edition emphasizes identifying cul-
tivated ferns by way of the text and illustrations for the 124 genera and more than 700 species treated (each
species is illustrated). Because diagnostic features are often unclear in photographs, this edition relies heav-
ily on silhouettes and line drawings. With additional help from the book’s glossary and a good hand lens,
the reader should be able to identify most of the cultivated species.
This revised edition treats all the commonly cultivated (and some rarely cultivated) fern species, vari-
eties, and many cultivars in the United States. Besides identification, this new edition will help readers learn
more about a particular fern by including its geographical distribution and author citations after the species
names. This information enables readers to have a better idea of which floras, texts, or journals to consult
for further information about a fern. An expanded bibliography also points to other literature sources.
This book is mainly written for people seriously interested in growing ferns, knowing their names and
what makes them similar or different, and appreciating their diversity. It is not a coffee-table book, nor a
chatty type of garden book meant for light reading. Beginning fern amateurs may find more information
than they need, but they will also find information useful at their level. Although this book primarily is a
reference, it is also for browsing and gleaning bits of information not readily found elsewhere.
The core information in this book will be particularly helpful to plant people who want to grow or iden-
tify different ferns and fern allies. Gardeners, nursery people, commercial growers, plant inspectors, botan-
ical garden staff, horticulturists, botanists, biology teachers, and others will find the information they need
in this one volume. The alternative would be to have a library of horticultural books and floras from the

7
8 P

world over and/or access to a fern botanist with a large herbarium. Botanists needing to grow ferns for their
research will find specific information on growing genera of ferns as well as general cultural information.
Students in horticulture, botany, and biology will find that this book gives an overview of fern infor-
mation that is basic and up-to-date, but they also should note that much remains to be discovered. In addi-
tion to examining the growth response of ferns to various environmental conditions, there is also much to
clarify on the relationships of genera and species. It is hoped that this book will stimulate students to study
horticultural and botanical problems of ferns or use them in their research. Although this type of problem
solving has been traditionally left to the scientist, observant, thoughtful, and innovative growers have made
and will continue to make valuable contributions to studying ferns (pteridology). Gardeners without sci-
entific training have introduced new species to horticulture, produced new cultivars and hybrids, and
found the right combination of conditions to grow difficult species. Much is unknown about temperature
tolerances of ferns, and this too is information that may be observed by growers and added to the store of
knowledge about ferns. Commercial growers have worked out detailed cultural information on growing
Boston ferns and leather ferns. Cultural details for other important trade groups have yet to be fully
reported.
Keeping an accurate list of the species in cultivation has become increasingly difficult, if not impossi-
ble. Many name changes since the first edition of this book have greatly complicated the task. Moreover,
new ferns are constantly being introduced to horticulture. There are about 10 fern societies in the world
whose members often bring new ferns into cultivation, and the number of species that can still be intro-
duced is sizable. An annual list recently offered by a fern society listed 775 packets of spores for exchange.
With so many possible species to list, it was thought better to err on the side of including too many instead
of too few. Several new and promising introductions came to our attention too late to be fully treated.
Several species listed as cultivated could not be verified, and others may have been overlooked. Since few
monographs have been written about problem ferns, identification was sometimes difficult and often com-
plicated by not knowing the country of origin.

Acknowledgments
Writing this book has been a prolonged and immense undertaking, and many people have helped and
encouraged us to reach the finish line. We especially wish to thank those people who read portions of the
manuscript for accuracy and gave helpful suggestions. They are Dr. Wade L. Berry, a plant physiologist and
soil scientist of the University of California, Los Angeles, who edited the cultural information; Dr. A. James
Downer, Farm Advisor, of the University of California Extension, Ventura, who edited the section on
insect pests and diseases; Dr. Ole Becker, of the University of California, Riverside, who edited the section
on nematodes; and botanist Sharon Ishikawa, of the United States Department of Agricultural, Animal and
Plant Health Inspection Service, who edited the section on importing plants.
We are indebted to the kindness of Dr. Alan R. Smith, of the University of California, Berkeley, and
Dr. John T. Mickel, of The New York Botanical Garden, for their assistance and time while we worked at
their herbaria. We also thank Dr. Art Gibson, of the University of California herbarium, for use of their
facilities. For identifying difficult ferns, we wish to thank the following people: Dr. Christopher R. Fraser-
Jenkins for Dryopteris; Dr. Richard Hauke for Equisetum; Dr. Blanca León for Campyloneurum; Dr. John
T. Mickel for Elaphoglossum and hardy ferns; Dr. Alan R. Smith for Thelypteris and other genera; Dr. Rolla
M. Tryon for cheilanthoids and suggestions on genera designations; and Dr. Iván Valdespino for Selaginella.
We were fortunate to have the help of other pteridologists from around the world. For Asian ferns we
were helped by many Japanese botanists. We are particularly grateful for identifications from Dr. Toshiyuki
Nakaike, Prof. Keisuke Yasuda, and Prof. Norio Sahashi. Also of assistance were Prof. Masahiro Kato,
 P 9

Prof. Reiko Yoroi, Prof. Ryoko Imaichi, Dr. Haruki Hirabayashi, Dr. Mitsuyasu Hasebe, and Dr. Noriaki
Murakami. The late Dr. Warren H. Wagner Jr., Dr. Dan Palmer, and Dr. Barbara Parris were valuable
sources of information on Pacific Island ferns. Dr. David B. Lellinger supplied information about Costa
Rican ferns, Dr. Lynn Raulerston about Guam and Palau ferns, Martin Rickard about English ferns,
Michael Price about Philippine and Malaysian ferns, and Dr. Peter Bostock and Chris Goudey about Aus-
tralian ferns. Yolanda Orta and Martin Grantham shared their expertise about fern propagation. For keep-
ing us informed of ferns in their area and for generous assistance in other ways, we are indebted to Mari-
lyn Johnson of Florida, Judith Jones of Washington, John and Carol Mickel of New York, and Nancy
Swell of Virginia.
We also thank the following for obtaining specimens and answering specific questions: the late Virginia
Ault, Ed and Phyllis Bates, Betsy Feuerstein, Robin Halley, the late Gerda Isenberg, Hildegard and Bruce
Jackson, Al and Marilyn Johnson, Judith Jones, Miriam Leefe, Robert Manthorne, Dr. Bruce McAlpin,
Duane Petersen, Milton Piedra, Ruth Radcliffe, Martin Rickard, the late Dorothy Skula, Nancy Swell, Roy
Vail, Helen Woodley, and Reggie Whitehead. In addition, we learned much from interacting with many
members of the South Florida Fern Society, the Tampa Fern Society, the New York Chapter of the Amer-
ican Fern Society, the Los Angeles International Fern Society, the San Diego Fern Society, the Hardy Fern
Society, and the Southwestern Fern Society.
Many commercial growers were generous with their help and information. We thank Ades and Gish
Nursery, Barford’s Hardy Ferns, California Ferns, Casa de Flora, Clark’s Nursery, Charles Alford Plants,
Daisy Plant Farm, Eisenbraun Ferns, Fancy Fronds, Fender’s Nursery, Fern Farm, Foliage Gardens, Frank
and Dorothy Skula, Freshwater Fauna and Flora, Glasshouse Works, H. Marugame, Half-moon Bay Nurs-
ery, Henry’s Plant Farm, Huntley Greens, Jerry Horne, Lyndon Horticultural Enterprises, Marilyn John-
son, Monrovia Nursery, MSK Rare Plant Nursery, Plants Unlimited, Santa Rosa Tropicals, Siskiyou Rare
Plant Nursery, Swell’s Nursery, Talnadge Fern Gardens, Tropical Gardens, Varga’s Nursery, Weather-
wood Nursery, Wedel’s Nursery, Weeks Nursery, Yerba Buena Nursery, and Yolanda’s Ferns.
For helping with computers, locating citations, assisting with the illustrations, and just making things
work smoother, we would like to acknowledge the help of Joan DeFato, Carol Brooks-Hoshizaki, Jon and
Madeleine Hoshizaki, Janet Keyes, Harry Le Vine Jr., Alvin Lee, Sylvia Louie, Vickie Milane, Mamie
Moy, Damon Woods, and Leo Yamanaka. We wish to acknowledge Arthur Takayama for taking and
preparing most of the black-and-white photographs and Lynn Vander Velde for the Adiantum line draw-
ings. When the senior author desperately needed editing assistance, Ed and Phyllis Bates, Ann Herrington,
Marilyn Johnson, and Susan MacQueen came to her rescue. Dr. Kenneth Wilson has been a source of
unflagging help, ranging from providing botanical expertise to trouble-shooting computer problems. The
senior author’s husband, Dr. Takashi Hoshizaki, a plant physiologist, suggested doing this revision more
than 15 years ago. He helped with the editing and artwork, but most importantly gave cheerful support and
encouragement over the years.
The authors apologize for any oversights in our acknowledgments and wish to assure all that their con-
tributions are nevertheless deeply appreciated.
The senior author also expresses her gratitude and deepest thanks to the junior author, Dr. Robbin
C. Moran, for consenting to join in producing this book. Not only is he one of today’s leading fern scien-
tists, he has also demonstrated his interest and ability in writing for general fern audiences. He has added
valuable information to the text, clarified complicated topics, and improved the book in many ways. With-
out his broad knowledge of ferns, his patience, and his energy to check on innumerable details, make the
required changes, do the editing, and work with Timber Press, this book would have languished much
longer in manuscript form.
We wish to especially acknowledge and thank our editor, Josh Leventhal. He made our work so much
easier and the book so much better with his careful editing and good judgment.
10 P

Although we relied heavily on the counsel of many people and are greatly indebted to their contribu-
tions, any discrepancies or errors in this publication are entirely our own. The information and advice con-
tained in this work have been brought together with care, but because of the large number of variables we
cannot guarantee complete accuracy or results. The reader acts on his or her own responsibility.

Barbara Joe Hoshizaki

Los Angeles, California

Robbin C. Moran
The New York Botanical Garden
 1 About Ferns

Ferns bring to mind pleasant, cool glens and shaded forests. We recall such things when we use ferns in our
gardens and homes. The ferny look is well known and loved, and some of us are content to sit back and relax
as we enjoy the soft green array of patterns and textures in ferns. But there is more to appreciate about ferns
than their mere appearance.
Ferns are usually recognized by their finely divided leaves, a type of leaf so characteristic that it is called
a “ferny leaf.” There are, however, many plants with ferny leaves that aren’t ferns, and many ferns that don’t
have typical ferny leaves. What, then, makes a fern a fern?
Ferns are spore-bearing plants; they lack flowers, fruits, and seeds. The so-called asparagus fern (Aspara-
gus setaceus) is not a fern despite its finely divided appearance because it bears flowers and seeds (it is actu-
ally a member of the lily family; its fruits are the orange berries often seen on the plant). Spores and seeds
greatly differ. Spores are simple one-celled microscopic structures, whereas seeds are complex, many-celled
structures usually visible to the unaided eye. Although plants such as algae, liverworts, and mosses produce
spores, they differ from ferns by lacking the large, thin, true leaves. Ferns further differ from these spore-
bearers by their development of specialized tissue (xylem and phloem) to conduct food and water. These
tissues also strengthen the stem and enable ferns to grow taller than other spore-bearing plants.
Unlike seed plants, ferns depend on water to complete their typical life cycle. They grow in places
where, when the time comes to reproduce, enough water is available for the sperm to swim to the egg.
Seed plants, such as pines and flowering plants, produce cones or flowers that use wind or insects to com-
plete their life cycle. They do not need water in the external environment for fertilization, and therefore they
can grow in drier conditions and dominate more of the landscape.
Nevertheless, some ferns have the surprising ability to thrive in extreme climates. Desert ferns, of which
there are few, often grow in the shade of rocks and boulders, using every bit of available water. Their roots
grow deep in the soil between the cool rocks, and their fronds are often covered with woolly hairs or scales
to protect them from water loss. When water is insufficient for new growth, many desert ferns curl their
leaves or shed their leaflets and suspend growth until the next rain.
Like desert ferns, alpine ferns are also adapted to climatic extremes. They tend to be small and have
hard-textured fronds that can endure the cold, dry winds. They grow only during the short summers as
weather permits.
Most ferns of temperate regions grow in the ground or on rocks, and only a few species grow on trees.
Approximately 200 species of ferns are native to the temperate areas of the United States. Temperate to sub-
tropical areas noted for their abundance of ferns include parts of the Himalayas, Australia, New Zealand,
and Japan. The tropics, however, harbor the largest number of fern species, particularly at middle elevations
in the mountains. Here the mountain fog encourages an abundance of ferns.

11
12 C 1

New ferns are still being discovered in well-known parts of the world, and in the lesser known parts
many more remain to be discovered. There are about 12,000 named species, and many of these have never
been introduced into horticulture. Temperate places like Japan and the cool, mountain slopes of the trop-
ics have many ferns suitable as new introductions for temperate gardens in the United States.
Compared to other plants, ferns have few economic uses. Nevertheless several are important. Ferns and
fern allies dominated the extensive coal-forming swamps of the Carboniferous period 280 to 345 million
years ago. Coal, therefore, is often composed of the compressed remains of ancient ferns and fern allies. If
you are reading this book by electric light and your local power plant is fueled by coal, chances are that the
energy used to generate the electricity for your bulb was captured millions of years ago by a fern or fern ally!
An important economic fern is Azolla, the mosquito fern. It is a rich source of nitrogen, a nutrient usu-
ally limiting to plant growth. Azolla is often used as an organic fertilizer in the rice paddies of southeastern
Asia. Considering how many people rice feeds in that part of the world, Azolla must rank among the world’s
most important economic ferns. It is also used to a lesser extent to supplement the diet of chickens, pigs,
and fish.
Perhaps the best-known use of ferns is the edible fiddleheads of species such as the ostrich fern of the
eastern United States and the bracken fern of Asia. (An early Oregon agricultural bulletin gave recipes on
how to cook bracken, including one on making “creamed bracken.” Subsequent research, however, has
shown that bracken contains carcinogens and should not be eaten.) The foliage of the water fern (Ceratop-
teris) and Diplazium esculentum are eaten in the tropics.
Various ferns with fibrous stems have been used as a substrate to cultivate epiphytic plants. The fibrous
stems of Osmunda and tree ferns have been shredded to make a planting medium for orchids. More recently,
tree-fern trunks have been cut and shaped to make containers for hanging-basket ferns. Because of conser-
vation concerns, however, the export of tree ferns and tree-fern products is restricted. In areas where they
abound, tree ferns have also been used to line roads and to construct small buildings because their trunks
are resistant to rot.
Other uses of ferns include thatching roofs and making tea and dyes. Some fern rhizomes and rachises,
such as those of the Gleicheniaceae and Lygodium, are flexible and tough enough to be used in basketry.
Folklore ascribes many dubious uses to ferns, such as restoring hair, making oneself invisible, and provid-
ing lucky amulets. The use of the male fern to treat worm infections, however, is an example of one such
tale being true. Some ferns, such as the bracken fern, gleichenias, and the hay-scented ferns, are considered
weeds. Ferns have also been used to investigate basic problems in plant biology, such as organ development,
hormones, genetics, evolution, and responses to light. But of all their uses, ferns are by far most important
for their ornamental value.
 2 The Structure of Ferns

Like sunflowers, oaks, pines, and other familiar plants in the landscape, the ferns have stems, roots, and
leaves. Although these parts are familiar to most of us, their character and function are sometimes unusual.
Much of this chapter might seem complicated, but you will find that a basic understanding of fern struc-
ture is extremely useful in making use of the rest of this book. Figure 2.1 illustrates the parts of a fern.

Stems
The stems of ferns can become tree-like and conspicuous, or they may be horizontal and go unnoticed.
Inconspicuous stems might be short, partly buried in the soil, or densely covered with leaf bases, hairs, or
scales. If the stem is erect and stout, it may be called a rootstock (Figure 2.2a). If it lies horizontally on the
ground and is anchored to the soil by roots, it is called a rhizome (Figure 2.2b–e). Some rhizomes are long,
freely branched, and climbing or scrambling over soil, rocks, or trees. They can bear fronds close together
or far apart. Some rhizomes may grow into the soil and send out new fronds in unexpected places. Others
branch frequently and form dense clumps. These clumps usually produce fronds in whorls, in irregular clus-
ters, or arranged in two ranks. Erect stems generally produce fronds in a whorl or vase-like cluster. Semi-
erect stems may have less perfect whorls. The slender, long stems on Boston ferns (Nephrolepis exaltata) and
other species are called stolons (Figure 2.2f). These spring from the main stem or rootstock and produce
new plantlets at their tips or from side buds. The stems of tree ferns are so large and wide that they are rightly
called trunks (Figure 2.2g). Some grow up to 18 m (60 ft.) tall. Where abundant, tree ferns have been used
to construct small buildings, and the starch in their stems has been used for food. Some ferns have thick,
hollow stems that house ants. The ants protect the fern and provide some nutrients to the fern in exchange
for shelter.
Stems serve the plant in several ways. First, the stem tip produces new stems, leaves, and roots. If this
tip is damaged, it might kill the fern, particularly if the stem does not branch laterally. Given its importance,
the stem tip is often protected by a covering of hairs or scales (Figure 2.2j, k). Second, the stem supports the
leaves. Leaves can be supported in large clusters at great heights, as in tree ferns, but most ferns have their
leaves supported on short- to long-creeping stems. Whatever the pattern of support, the leaves are always
held up to the light and air. Third, stems conduct water and nutrients between the roots and leaves. For this
purpose they are equipped with strands or bundles of special tissue known as xylem and phloem or, col-
lectively, as vascular or conducting tissue. The vascular tissue can have different patterns depending on the
species or taxonomic group to which the fern belongs (Figure 2.2h, i). Fourth, stems store food and water,

13
 costa
costule
segment
rachis costule

segment

Pinnule Detail

rachis

blade

pinna

frond

pinnule

segment

stipe

fiddlehead
scales

rhizome

roots

Figure 2.1. Parts of a fern. After Taylor (1984).

 c

a d

g
i

h
j
k

Figure 2.2. Kinds of stems: a. erect stem; b. rhizome, short-creeping (Asplenium bulbiferum); c. rhizome, wide-creeping
and branching (Gymnocarpium dryopteris); d. rhizome, wide-creeping and branching from stipe base (Hypolepis
tenuifolia); e. rhizome creeping, branching and fleshy (Phlebodium); f. stolons (Nephrolepis exaltata); g. trunk of a tree
fern; h. vascular or conducting tissue within rhizome (Hypolepis); i. vascular or conducting tissue within rhizome
(Polypodium); j. hairs covering the rhizome of h; k. scales covering the rhizome of i.
16 C 2

particularly if they are thick and fleshy (Figure 2.2e). Some stems, such as stolons and branching rhizomes,
reproduce plants or increase the size of the colony.

Roots
The roots of adult ferns grow from the stem. There is never a main root system consisting of a taproot and
lateral branch-roots. Fern roots are mostly fine, fibrous, densely branched, thick masses; rarely are they
thick and fleshy. They grow close to the soil surface and are easily injured when the soil is tilled or disturbed.
Young, actively growing roots have whitish or yellowish tips. Older portions of the roots are dark brown or
black. The presence of a large number of young roots is a good sign of active growth.
Creeping or reclining rhizomes produce new roots as they grow forward. These roots are usually located
on the underside of the rhizome just behind the tip. Ferns with erect stems produce new roots from the stem
base. Erect-stemmed ferns may have their bases covered with old leaf bases, making it hard for the emerg-
ing roots to establish themselves into the soil (Figure 2.2a). If, however, the old fronds decay and a fine layer
of forest litter accumulates around the stem, emerging roots will find a rooting medium. In cultivation it
may be necessary to remove the old leaf bases of erect stems and replant the fern deeper into the soil to give
the emerging roots a better chance to establish themselves and prolong the life of the fern.
Old tree ferns often have masses of roots on their stems. These roots are called aerial roots and form a
fibrous outer layer around the stem. They absorb water and add strength and rigidity to the stem, which is
necessary to support the heavy crown of fronds. The trunks of some tree ferns—for example, the Hawai-
ian tree fern Cibotium glaucum—readily send out new roots when planted.
The roots of some ferns house fungi in a symbiotic relationship. The fungi (mycorrhizae) help gather
mineral nutrients for the fern in exchange for food. The fungi reside in the cells of the root or the root hairs
that often become swollen and mucilaginous at their tips. The mycorrhizae that have been studied on ferns
are reported to be cup fungi (Ascomycetes) and Glomales (Moteetee et al. 1996). Mycorrhizae of agricultural
crops studied have been found to enhance growth when soil nutrients are low and not when sufficient or
high. Whether these findings apply to ferns is not known.
The primary function of roots is to absorb water and minerals as well as hold the plant in place. Some
ferns have roots with other functions, such as producing new plants or storing food and water.

Leaves
The leaves of ferns are called fronds and are produced at the stem tip. They develop by uncoiling and while
doing so are called fiddleheads or monkey tails (see Figure 2.1). Technically, they are called croziers, from
the term for a shepherd’s crook. Croziers are tender and vulnerable to drying and damage. They are often
protected by a covering of hairs or scales. A few ferns have emerging fronds that are hook-shaped or straight
rather than coiled.
When a crozier uncoils, it produces a frond that typically has two parts: the stipe (petiole) and blade (see
Figure 2.1). The stipe is the stalk, and the thin leafy part it supports is the blade. The stipe commonly
bears hairs or scales that are often important in identification of a fern. In some ferns the stipes may be dark
and highly polished, as in the maidenhair ferns (Adiantum). The stipes may also be long, short, or entirely
absent (sessile). If the stipe naturally breaks or separates from the stem along a predetermined line or joint,
it is said to be jointed or articulate. The tissue found in or near the middle of the stipe is called the vascu-
lar or conducting tissue (stipe bundles), which can be seen if you cut through the stipe with a razor blade.
Many fern genera can be identified by the patterns of this tissue.
 T S  F 17

Blades come in many shapes and may be undivided, little divided, or much divided into smaller parts.
Names are given to all these shapes, states of division, and their parts. The undivided state is called simple
(Figure 2.3). If the blade is divided to the central vein or midrib, the frond is said to be compound (Figure
2.4). If each resulting leaflet is again divided to its midrib, the frond is twice compound. Tree ferns are often
two- or four-times compound. If the blade is cut so that the leaflets form along the midrib, the frond is said
to be pinnately compound or pinnate. If the blade is cut so the leaflets join at the same point on the stipe,
the frond is said to be palmately compound or palmate. If the lateral leaflets on a palmate frond are cleft into
two or more segments the frond is called pedate. Most ferns are pinnately compound.

a b c d

Figure 2.3. Simple fronds: a. simple and entire; b. bifurcate; c. pinnatifid, with pedate base; d. pinnatifid.

Whether the blade is pinnate, palmate, or pedately compound, the leaflet of the first division is called
a pinna (pinnae is the plural). If the pinna is divided, its divisions are called pinnules or secondary pinnae,
and the frond is two-pinnate (bipinnate). If the secondary pinnae are divided, their divisions are called ter-
tiary pinnae, and the frond is three-pinnate (tripinnate). A general name for these parts, whether primary,
secondary, or tertiary in rank, is leaflet. If a frond is pinnately compound, its midrib appears like a contin-
uation of the stipe and is then called a rachis. The margins of the blade or its leaflets may be variously mod-
ified (Figure 2.5).
Venation patterns differ greatly among ferns (Figure 2.6). Veins may be unforked (simple) or forked.
If forked, they can be branched distinctively. Their ends may be free or joined with others to form meshes,
in which case they are said to be netted (or areolate or anastomosed). Sometimes free veins are enclosed in
a mesh, and the ferns are said to have included veinlets. Some ferns have veins that are deeply immersed in
the blade and hardly visible.
The great variety in frond shape, size, and venation does not affect the ability of the frond to fulfill its
main function—to make food for the plant with the help of sunlight.

Fruiting Bodies
The lower surface of the frond may bear rusty patches or black dots or lines known as sori (sorus is the sin-
gular). These are clusters of spore cases or sporangia. The sori may be round, oblong, linear, or of some other
 r

p p
p p

r r r
sp

d
a c

sp

sp p

r p

g
f

tp
sp
e

Figure 2.4. Compound fronds (r = rachis; p = pinna; sp = secondary pinnule; tp = tertiary pinnule): a. pinnatifid above
to pinnate below; b. pinnate; c. pinnate-pinnatifid; d. bipinnate; e. tripinnate; f. pedate, the pinnae pinnate; g. palmate
or radiate, the pinnae pinnate.
 T S  F 19

d e
a c
b g
f

Figure 2.5. Leaf margins: a. entire; b. undulate; c. crenate; d. serrate; e. lobed; f. pinnatifid; g. incised.

shape (Figure 2.7). Some ferns have loosely arranged sori, not definite clusters. The sporangia may be scat-
tered along the main veins, or they may completely cover the lower surface of the frond. The sori of some
ferns are protected by an indusium, a bit of tissue that, depending on the species, is formed in various ways
from various parts of the blade (Figure 2.8). In some ferns it consists only of the margin of the leaf rolled
over the sori (Figure 2.8c). Such an enrolled leaf margin is called a false indusium. The shape and location
of the sorus and the kind of indusium (if present) are important in identifying ferns.
The spore cases or sporangia are easily seen with a ten-power (10×) magnifying glass, but to see their
finer details requires a microscope. They will be seen as stalked capsules or cases containing spores. If the
sporangia have already shed their spores, they will appear frayed or cracked (Figure 2.9). If the sporangia still
contain spores, the case will be round and intact. In this condition you may be able to see a slightly raised
dark ring of cells around the case. This ring is called the annulus and acts like a catapult to open the case and
hurl out the spores. The annulus may vary in position on the spore case, and in some genera it is only a patch
of slightly thickened cells instead of a well-defined ring (Figure 2.10). In some species all the sporangia in
a sorus open at about the same time, but in most ferns the sporangia open at different times over an
extended period.

b c
a d

Figure 2.6. Veins: a. simple; b. forked with free vein endings; c. netted or areolate; d. netted with included veinlets.
 a b
c
d

Figure 2.7. Sori: a. along the veins (Coniogramme); b. round (Macrothelypteris); c. linear and marginal (Pteridium); d.
linear and medial (Athyrium).

b
a c
d

Figure 2.8. Indusia: a. peltate or umbrella-shaped (Polystichum); b. scale-like (Davallia); c. false indusium, formed from
an enrolled leaf margin (Adiantum); d. cup-shaped (Dennstaedtia).

Figure 2.9. Sporangium shedding spores.

 T S  F 21

a b c d

Figure 2.10. Annulus positions: a. lateral (Todea); b. apical (Lygodium); c. oblique (Gleichenia); d. vertical
(Polypodium).

A single frond may produce hundreds of thousands of spores. To the unaided eye, spores appear as a
fine dust. Under the microscope, they can be seen to have two basic shapes, depending on the species (Fig-
ure 2.11). Some have bean-shaped spores (monolete), whereas others have more or less pyramidal or slightly
three-angled globose spores (trilete). Both shapes usually have patterns or embellishments on their sur-
faces, and these markings often characterize certain families or genera. The spores of most ferns are brown
or black, but yellow spores are characteristic of the polypods, and green spores are found in some groups.
Green spores are short-lived, usually lasting only one or two weeks. In contrast, nongreen spores usually
remain viable for several years, but with age they germinate more slowly and less dependably.

a b c d e

Figure 2.11. Kinds of spores: a–c. monolete; d–e. trilete.

The life cycle of ferns is complex but of great importance to growers. It is discussed further in Chapter
8 in connection with propagation techniques.
 3 Obtaining Ferns

Fern Nurseries and Fern Organizations

Most plant nurseries carry at least a few ferns and usually can direct you to specialty nurseries or mail-order
businesses that carry a wider variety. Local newspapers announce garden club meetings and flower shows.
These activities will give you the opportunity to meet local botanists and gardeners who can help you to
locate ferns. Botanical gardens usually know what plant societies exist in your area and whom to contact for
information. They may also have a file of more distant and foreign nurseries that might ship ferns. Fern soci-
eties have been organized in various parts of the world. These organizations mail newsletters, print journals,
sponsor spore exchanges, conduct field trips, provide programs, and maintain a fern library or Web sites for
their members. For information on contacting these fern groups, see Appendix II, “Fern Societies.”

Purchasing Ferns
Before you buy a fern, check its climatic needs to determine whether you can provide it with the proper tem-
perature, shade, and humidity. Also determine whether it will produce the desired landscape effect, espe-
cially given its size when fully grown. Some ferns take up more space than you intended.
Once at the nursery, purchase the fern in a 10 cm (4 in.) to 3.8 liter (1 gal.) size container, unless you
are in a hurry to have a larger plant. Leave the smaller sizes for a time when you have gained more experi-
ence in growing them. Plants larger than the 3.8 liter (1 gal.) size are more expensive, and some may have
passed the stage of rapid growth.
Examine the growing tip. Select a fern that has a firm, large, growing tip. Avoid any that seem discol-
ored, shriveled, or undersized. If the fern is actively growing, see that the tip is forming healthy new fronds.
Avoid ferns with shriveled or deformed growth. If you do not want to repot the fern in the near future, pick
one that has not already overgrown its pot. The growing tip should not be pushing against the edge of the
pot. If you are obtaining a fern that forms clumps of rhizomes, select the one that has the largest number
of healthy rhizome tips.
Nurseries use different soil mixes. Be aware that some mixes require more watering than others. Also,
the smaller the pot and the larger the fern, the more water it will require. If the fern is accustomed to grow-
ing in high humidity at the nursery, it may wilt when you get it home. If you give it just enough humidity
to keep it from wilting, new fronds should adapt to the drier conditions without difficulty. Otherwise, see
the section in Chapter 4 entitled “Air Circulation and Humidity.”

22
 O F 23

Collecting Ferns
Certain parts of the world are particularly rich in ferns, and it is tempting for you or your friends to collect
some exotic ferns during travels to these areas. But given the concern about the environment and rare or
endangered species, collecting wild plants is discouraged. Collecting wild plants is forbidden in nearly all
countries unless special collecting permits are obtained. In addition, export permits are also required, and
customs or agricultural inspectors may want to see those permits when you exit a country or re-enter your
country. See Appendix III for more information on permits and restrictions.
If collecting ferns from private property, seek out the owner’s permission. Before collecting wild plants,
be sure to have the proper permits and check that the plants are not on the rare, threatened, or endangered
list. Plants so listed are protected by law (see Appendix III for details). Some plants, such as certain species
of Azolla and Salvinia, are deemed potential pests and are barred from entry into the United States and
within the country cannot be transported without a permit. Even with nonthreatened species, resist the
temptation to collect indiscriminately, wastefully, and without regard for the environment. Thoughtless
collectors have given all collectors a bad reputation. Take only the number of plants that you can grow.
If you live in an area where native ferns abound, you will have access to species suited for the climate
but not necessarily for garden culture. Temperate, deciduous ferns are usually collected in the late fall or
early spring, and other ferns just before the growing season, usually in early or mid-spring. Although ferns
may be collected at any time, their chances of survival are best when they are transplanted just before their
active growing period so that they have a full season to grow and establish themselves. Recently matured
ferns (as opposed to older or long-mature plants) have the best chance of re-establishing themselves. Also,
plants with thick rhizomes generally have a better chance of survival than those with thinner ones. If the fern
is branched or clumped, one piece of the branch or clump is enough to start a new plant. When collecting
a rhizome, take as many of the roots as possible and try not to disturb the soil attached to them. Firm the
soil around the exposed roots of the remaining plants. While transporting the plant, keep the soil and
foliage cool and moist, and replant the fern as soon as possible.
By visiting local nurseries, gardens, and members of plant societies, you might be able to obtain inter-
esting species and establish valuable contacts as well. Above all, keep in mind that spores usually do not
require permits for import and are easier to collect, pack, and mail. Also, spore-grown plants tend to adapt
more readily to cultivation.

Holding Ferns until Departure Time

If you have collected ferns and will not return home for a week or so, be sure to remove the plants with as
little disturbance to the roots as possible and put them into a plastic bag kept open for ventilation. Water
the fern sparingly, and provide drainage in the bag if needed. Find a cool, shady place to store the plants as
you travel. Prior to departure, prepare and package the plants as discussed in the following section.

Packing Ferns
Water the ferns a day before packing so that excess water has time to drain away. Ferns to be transported
with soil on their roots need only be wrapped in newspaper. If the ferns are in pots, they may be tapped out
of the pots to reduce the weight. When bringing plants into the United States, all soil must be removed from
the plant. Dip the roots into a bucket of water so the soil will fall away, but try to leave intact as many of
the roots as possible. A number of dips and a gentle agitation in the water should get the soil off. Remove
24 C 3

all dead and broken fronds. Double-check to see that all insects are removed. The tip of the stem or rhizome
is the most important part of the fern and must be well protected. If fronds must be removed for packing,
do so, but keep the roots and particularly the rhizome or crown intact. Protect the rhizome and roots from
being crushed by surrounding them with packing material. If packing material other than newspaper is used
and ferns are to be sent into the United States, make certain that the material is on the approved list issued
by the Plant Import Permit Station. If you are going to remote areas, bring your own packaging and wrap-
ping supplies. Stringed tags are useful labels, as are gummed labels that will stick on plastic. Write the
name and place of collection on the labels in pencil or permanent ink.
Some plants can be wrapped for protection in newspaper rolled into a cylinder or a cone and placed in
a plastic bag to retain moisture. Make certain that the plants are not too wet, especially if they are put into
plastic bags without any newspaper to cushion the plant and absorb extra water. Plants that are too wet will
rot readily in plastic bags, especially in warm climates. Air may be blown into plastic bags and sealed to serve
as a cushion. Plants sent to the United States should be in sealed bags. If pests are found in any one open
bag, then all the other open bags in the shipment might be considered infested and would be quarantined
by agricultural inspectors.
Empty plastic water bottles can be made into excellent containers for transporting plants. Cut the bot-
tle in half around the middle. One piece will fit snugly inside the other piece with a slight squeeze when you
wish to close the container.
Cardboard cartons lined with wax paper or plastic can be used to ship ferns. In some areas, woven bas-
kets are easier to obtain and can be used, unless they are made of willow or other wood likely to carry bor-
ing insects. Such baskets are barred from entry into the United States. Secure the individually wrapped ferns
so they will not slip or slide in the box.
Commercial growers have reported that potted, established plants may show stress after 7 to 10 days if
left in the dark, even when other conditions were favorable.

Interstate Transport of Ferns

Regulations about the interstate movement of plants vary between states and even between counties within
a state. Check with your local state and county agricultural offices before shipping plants. Many states
regulate interstate commercial activities involving legally acquired protected species. States that have large
agricultural businesses also have many restrictions. Contact the appropriate state fish and wildlife agency for
further information.
 4 Cultural Needs

Understanding Fern Needs

Ferns have the same basic growing requirements as other plants and will thrive when these are met. There
is nothing mysterious about the requirements—they are not something known only to people with green
thumbs—but the best gardeners are those who understand plant requirements and are careful about satis-
fying them. What, then, does a fern need?
All plants need water. Water in the soil prevents roots from drying, and all mineral nutrients taken up
by the roots must be dissolved in the soil water. Besides water in the soil, most plants need water in the air.
Adequate humidity keeps the plant from drying out. Leaves need water for photosynthesis and to keep from
wilting.
All green plants need light to manufacture food (sugars) by photosynthesis. Some plants need more light
than others, and some can flourish in sun or shade. Most ferns, however, prefer some amount of shade.
For photosynthesis, plants require carbon dioxide, a gas that is exhaled by animals as waste. Carbon
dioxide diffuses into plants through tiny pores, called stomata, that abound on the lower surface of the
leaves. In the leaf, carbon dioxide is combined with the hydrogen from water to form carbohydrates, the
plant’s food. This process takes place only in the presence of light and chlorophyll, a green pigment found
in plant cells. To enhance plant growth, some commercial growers increase the carbon dioxide level in
their greenhouses to 600 ppm (parts per million), or twice the amount typically found in the air.
Plants need oxygen. The green parts of a plant do not require much oxygen from the air because plants
produce more oxygen by photosynthesis than they use. The excess oxygen liberated from the plants is used
by all animals, including humans. What do plants do with oxygen? They use it just as we do, to release the
energy stored in food. We use energy to move about, to talk, to grow, to think—in fact, for all our life proc-
esses. Although plants don’t talk or move much, they do grow and metabolize and must carry on all their
life processes using oxygen to release the stored energy in their food.
Roots need air all the time. They get it from the air spaces between the soil particles. Overwatering dis-
places the air between soil particles with water, thereby removing the oxygen needed by the roots. This
reduces the root’s ability to absorb mineral nutrients and can foster root-rot.
Plants need minerals to grow properly. The minerals are mined from the soil by the plant’s root system.
If a certain mineral is missing, such as the calcium needed for developing cell walls, then the plant will be
stunted, discolored, or deformed.
Some plants tolerate a wide range of temperatures, whereas others are fussy. If the temperature is too
high or too low, the machinery of the plant will not operate satisfactorily or will cease entirely.

25
26 C 4

The basic needs of plants are not hard to supply, but growing success depends on attending to these
needs with care and exactitude. The remainder of this chapter is devoted to a discussion of these require-
ments, with the exception of mineral needs, which are discussed in Chapter 5.

Temperature
Ferns vary greatly in their temperature requirements. Most grow best at daytime temperatures of 18 to 27°C
(65–80°F). Tropical ferns generally prefer ranges from 21 to 27°C (70–80°F) during the day, and some may
even tolerate similar nighttime temperatures. Most ferns grow best when the nighttime temperature is
about 5°C (10°F) cooler than the daytime temperature. Some ferns need seasonal periods of cooler tem-
peratures to grow well. You cannot predict a fern’s temperature tolerance simply by the looks of the plant.
Knowledge of its native habitat and the temperature tolerances of its relatives will help, but there are some-
times surprising exceptions. When the hardiness of a species is unknown, it is best to use larger individu-
als because they generally withstand temperature extremes better than small plants. Since the minimum
temperature tolerance is not known for many ferns, gardeners can make useful contributions to horticul-
ture by reporting such data from their garden to appropriate fern societies and journals.
Floras, which are books for identifying the plants of a particular region, sometimes provide informa-
tion on determining the temperature tolerance of ferns in a given area. Some relatively recent fern floras
helpful to gardeners have been written for Australia (ABRS/CSIRO 1998; Andrews 1990; Jones and Cleme-
sha 1981), New Zealand (Brownsey and Smith-Dodsworth 1989), South Africa (Burrows 1990; Jacobsen
1983), and Japan (Iwatsuki et al. 1995; Kurata and Nakaike 1964–1997, fully illustrated with maps).
North America and Canada have a flora (Flora of North America Editorial Committee 1993) and there are
many smaller ones. In tropical America, fern floras have been written for the West Indies (Proctor 1977,
1985, 1989), Chihuahua, Mexico (Knobloch and Correll 1962), Oaxaca, Mexico (Mickel and Beitel
1988), Chiapas, Mexico (Smith 1981), Central America (Moran and Riba 1995), Costa Rica and Panama
(Lellinger 1989), the Venezuelan Guayana (Smith 1995), Peru (Tryon and Stolze 1989–1994), and Chile
(Marticorena and Rodríguez 1995). Except for Taiwan (Huang 1994) and Hong Kong (Edie 1978), China
and Korea have not been adequately covered (at least not in English). The ferns of the western Himalayas
have been treated (Khullar 1994) as well as those of Thailand (Tagawa and Iwatsuki 1979–1989), and
western Malaysia (Piggott 1988). Many of the Pacific Islands have floras, but noticeably lacking are floras
for eastern Malaysia, Indonesia, and New Guinea. The British Isles have many recent floras, but those for
Continental Europe in English are not updated.
Ferns may be classified as hardy, semi-hardy, semi-tender, or tender. This classification refers to the
coolest temperature range that a fern can tolerate. It is somewhat arbitrary, since many ferns fall on the bor-
derline between categories or can endure short periods of lower temperatures. The classification, however,
ensures that the temperature preferences of ferns can be more easily known. See the section “Outdoor Ferns”
in Chapter 9 and the individual entries in Chapter 13 for temperature requirements of particular species.

H F
Hardy ferns can be grown outdoors in temperate or colder areas of the United States. They tolerate win-
ter temperatures well below freezing. Also belonging to this group are alpine species that can endure very
cold temperatures for a long time.
Hardy garden species are mostly native to northern and central Europe, North America, and northern
Asia. Many ferns native to the eastern United States are suitable for cultivation. Some adapt to gardens in
warmer climates, but others do not. The cultivation of hardy ferns is treated in several references, includ-
 C N 27

ing those by Dyce (1991), Foster (1993), Grounds (1974), Kaye (1968), Mickel (1994), Rush (1984a), and
Rickard (2000).

S- F
Semi-hardy ferns usually tolerate nighttime temperatures above 4°C (40°F) during the cool season.
They survive periods of freezing temperatures that are short and not too severe. These ferns come from
many parts of the world, mostly where warm-temperate climates prevail, such as Japan, Korea, and China.
Semi-hardy ferns grow outdoors in the southeastern United States and warmer northern areas. They are
good choices for inland valleys along warmer coastal regions.

S- F
Semi-tender ferns grow well where nighttime temperatures are mostly above 10°C (50°F) during the
cool season and where freezing temperatures are rare. Frost may burn the foliage or kill the plant, although
some species can survive near-freezing temperatures for a few hours. The semi-tender ferns are native to sub-
tropical and warmer areas with typical daytime temperatures of about 18°C (65°F). Many come from
Latin America, Australia, and New Zealand. Coastal southern California is particularly well suited for
semi-tender ferns, whereas those that tolerate cooler temperatures can be grown inland or farther north
along the coast.

T F
Tender ferns (also known as warm or “stove” species) usually grow poorly when temperatures drop to
16°C (60°F) or below for successive nights. These ferns are mostly native to the lowland tropics where
year-round warmth and humidity prevail. Southern Florida and the Hawaiian lowlands are particularly
favorable places to grow tender ferns outdoors. Here, nighttime temperatures are closer to 21°C (70°F) or
above for most of the year, and daytime temperatures are near 24°C (75°F) or above. In other areas of the
United States, most lowland tropical species must be grown under glass, at least during the cool months,
especially the very tender species that do not like successive nights of temperatures below 21°C (70°F).
Some semi-tender and tender ferns prefer cooler temperatures at night and do not do well in areas
with warm nighttime temperatures. These ferns are usually native to the cooler upland areas of the tropics
and include most of the tree ferns. Conversely, some lowland tropical ferns may not grow in places where
the nighttime temperatures are consistently much lower than the daytime temperatures.

L T V

In coastal areas where the temperatures are ameliorated by the sea, a diversity of climates will be found
within a few miles, especially if the area is hilly or mountainous. In these places it is especially important to
know the temperature conditions of your specific locale. In southern California, for example, climates
ranging from subalpine to subtropical occur within a distance of 64 kilometers (40 miles). Many non-
coastal areas also may have microclimates due to local geography. For information on your local climate,
write the United States Department of Commerce, National Climatic Data Center, Federal Building,
Asheville, North Carolina 28801, or call 704-CLIMATE (254-6283). Also consult the U.S. Department
of Agriculture’s map of hardiness zones, which gives minimum winter temperatures throughout the United
States and Canada (last page of the color plates section).
28 C 4

L-T D
Low temperatures (but above freezing) can damage plants. In such instances the plant usually turns yel-
low, produces little growth, and wastes away. But before you dig up a fern to move it to a warmer place, be
sure it is not merely taking its normal rest period. Many ferns take a rest period, especially during the cooler
winter months, when they normally turn yellow and produce little growth. They vigorously renew growth
usually in spring, whereas those that have suffered from low temperatures slowly recover and often have
stunted, undersized rootstocks or rhizomes.
Semi-tender and tender ferns growing in marginally favorable temperatures need to be watered lightly
during the cooler months. At this time growth is slow, and overwatering can often result in root-rot. All
ferns, tender or hardy, are more vulnerable to low temperatures if they are young, weak, recently trans-
planted, or have much new growth.
Various steps can be taken to prevent or reduce damage when temperatures drop, but success will ulti-
mately depend on the fern’s sensitivity to cold and the severity and duration of the low temperatures.
Harden the plants for several weeks or more by giving them less water and exposing them to low but not
injurious temperatures prior to winter weather. Some plants are easier to harden than others. Avoid heavy
pruning or fertilizing late in the season to discourage vulnerable, tender new growth. Keep withered fronds
on the plant to provide some protection through the winter. Transplant ferns early in the growing season
so they will be well established by fall. Protect weak, young, or susceptible plants by moving them to
warmer places such as against buildings, under eaves or trees, or on slopes where the cold air drains away.
Temporary coverings of burlap, straw, leaves, or newspaper will provide some protection during cold spells.
Various foams are available that are nontoxic and work by insulating the plant. Commercial sprays that pro-
tect foliage from frost are also available, but their effectiveness on ferns is unknown.
Once frost or freezing is imminent, increase heat and reduce heat loss from the ground, plant, and sur-
roundings. On clear, still nights the coldest temperatures are likely to come an hour or so before sunrise, and
that is when protective measures may be needed. Shortening the time a plant is exposed to lethal low tem-
peratures from hours to minutes can prevent injury or death. When adding heat with various heaters, be
sure to guard against dehydration, especially from electric heaters in small enclosures. Air movement by
means of fans may bring in warmer air if a low inversion layer is present (that is, a low-lying layer of warm
air above the cold). Inversion layers can be expected when the night is calm and clear but the preceding day
was warm and sunny. This causes the warm air of the day to rise during the night and the heavier colder air
to move downward and settle around the plants.
Fog can be used to protect crops during cold spells with temperatures as low as −2°C (28°F). Farmers
place fog-producing nozzles about 1 m (3 ft.) apart and 2 to 3 m (6.5–10 ft.) above the ground over the aisle
to prevent any dripping water from damaging the plants. The dense fog is confined by a roof of shade
cloth and plastic curtains. The cold temperature is absorbed by the fog, leaving less to damage the plants.
True fog differs from mist in that fog droplets are so small that they do not settle out of the air, whereas mist
droplets are larger and settle out. These larger droplets are less efficient than true fog. Fogging or misting
should be started as soon as temperatures drop below 2°C (35°F), and then maintained continuously or at
regular intervals. It should be continued until the sun has warmed the air above 0°C (32°F). Stopping the
treatment too early may cause the lethal temperature of the plant to be reached.
The formation of frost or ice gives off heat. The layer of frost or ice covering or encasing the plant also
confers some insulation advantage, but the weight of the ice might damage or discolor the plant. Furrow irri-
gation is another means of increasing the temperature, when the water in the furrow freezes, but plants can
become overwatered by this method, and the rise in temperature may be insufficient to protect the plant.
Plants that have been super-cooled (that is, cooled so quickly that damaging ice crystals have not
formed on the plant tissue) should not be moved or disturbed because the movement will cause the sudden
 C N 29

formation of ice crystals that, as they grow, pierce and kill the cells of the plant. Super-cooling is apt to occur
when plant surfaces are dry. Allow super-cooled plants to warm slowly.
If plants are frozen, allow them to thaw gradually. This can be done by placing newspaper over the plant
to insulate and shade it from the sun, the warmth of which would otherwise melt the ice crystals within the
plant too rapidly.
Ferns damaged by frost or freezing generally wilt (unless hard tissue is present) and turn black. Less
severely damaged fronds have spots or areas that appear scorched or burned, especially on the more exposed
parts. Damage from desiccation can also occur. For tree ferns and species with woody tissue, particularly
subtropical species, a common practice is to keep the damaged parts on the plant until there are signs of new
growth. Herbaceous ferns that collapse and have completely black fronds may recover if the rhizome is still
firm. Damaged stems or rhizomes often produce distorted, weak growth and will recover slowly, if at all.
Any cold-damaged plants should be watered sparingly because the loss of foliage reduces their water needs.
Assessing the extent of damage may take as long as one growing season on tree ferns. For more details, see
Reasoner (1982).

H-T D
In arid areas, high-temperature damage is difficult to distinguish from damage caused by dry air. Even
with sufficient humidity, high temperatures can scorch or wilt fronds. If the wilted parts do not recover, they
turn brown and brittle. On thicker fronds the scorched areas are usually near the margin or exposed places.
During heat waves, shade the ferns and keep the air humid and the soil moist. In greenhouses, open the
vents and keep the mist nozzle operating. Water sprayed on the roof of the greenhouse will reduce the
temperature appreciably. Never keep plants in a closed car or its trunk on hot days.

C R
Certain temperate and alpine ferns need cold temperatures during the winter to grow well. Without a
cold period these ferns will not renew growth in the spring, or if new growth is produced, it is weak and ulti-
mately wastes away. Growers have established the specific cold requirements of several ferns (see the section
“Forcing Ferns” in Chapter 10). Not all temperate ferns, however, have chilling requirements, and species
that have populations in the southern parts of their ranges may not require cold. Growers in warm climates
should be aware of the difficulties of growing species that require a cold period. Even tropical ferns taken from
higher elevations can prove difficult to grow in hot, lowland tropical climates because of this requirement.
Experiments on Michigan ferns (Hill 1976) showed that different species require different cold peri-
ods, if at all, to renew growth in the spring. Osmunda claytoniana, for example, needs 8 to 12 weeks at 4°C
(40°F) in the dark to achieve 100% bud-break or renewed growth. If the plants are removed from the cold
and dark and placed in a warm, sunny greenhouse (at 20 to 30°C [68–86°F] with 24 hours of light), those
that received the longest cold treatment emerge the fastest (plants with 12 weeks of cold treatment emerged
within 10 days once in the greenhouse). In contrast, bud-break in Adiantum pedatum and Thelypteris palus-
tris was less influenced by the length of the cold treatment, and in Cystopteris fragilis, Onoclea sensibilis,
Deparia acrostichoides, and Diplazium pycnocarpon, no cold treatment was required for bud-break. The lat-
ter species began growth without cold treatment soon after they were brought into the greenhouse. These
experiments explain why it is difficult to grow Osmunda claytoniana in warm climates whereas it is easy to
grow Diplazium pycnocarpon.
30 C 4

Light
H M L
Most ferns grow best in shade or filtered light. The optimal amount of light for most ferns is that pro-
vided by the sun on a dim, overcast day. This measures between 200 and 600 foot-candles. Foot-candles
(f.c.) is a unit for measuring light intensity; another more recent system uses micromole (µmol) units. (See
Appendix I, which describes a method for measuring light intensity with a photographic light meter.)
Adult ferns prefer more light than younger ones, and most spores germinate and grow best in low light.
Contrary to what many people believe, dense shade is unsuitable for most ferns. In dense shade, fronds grow
spindly, are less frequently produced, and tend to yellow and die early.
Ferns growing in low but adequate light are usually large and luxuriant. Under high light, the fronds
become firmer, thicker, more soriferous, and more tolerant of environmental changes. Ferns suffering from
too much light are smaller, less luxuriant, yellowish green, and may have brown margins.
In coastal areas with typically overcast weather, ferns can be planted in full sun. Inland areas may have
brighter days, and the ferns will require more shade. When planting ferns, do not overlook the added light
reflected from walks and buildings. This light can become intense at certain times of the year as the sun’s
angle changes with the seasons.

S F
Trees, laths, painted glass, fiberglass, and plastic cloth all may be used to shade ferns. Plastic cloths made
of polypropylene (sold as Prop-a-Lite, which superseded Saran or Lumite) are much used by the nursery
trade. Different grades give from 30 to 90% shade. In sunny areas such as southern California, 73% shade
cloth (giving 73% shade) has been used with good results. If ferns must be grown in direct sun, they should
be shielded from the sun between noon and 3 .., the brightest and hottest hours of the day.

L Q
Sunlight is composed of a variety of colored lights. These colors are seen separately in a rainbow or
through a glass prism. The red and blue parts of this color spectrum are absorbed by the plant’s pigments
and are used in photosynthesis. Most of the remaining colors in the spectrum are not used by plants and are
passed through the leaf or are reflected. Green light in particular is reflected, which explains why leaves
appear green. Light intensity (discussed earlier under “How Much Light”) should not be confused with light
quality. Light intensity refers to how strong the emitted light is, whereas light quality refers to the kinds of
colored light present. Gardeners who grow plants in sunlight need not be concerned about whether their
plants get enough red and blue light, as they can do little to modify the sunlight’s quality or spectrum. How-
ever, gardeners who grow plants under artificial light to increase the light intensity may also want to know
whether the lamps they are using gives the proper light quality or light spectrum.

A L
Ferns grown indoors, especially in more northerly areas, often suffer from too little light. Supplemen-
tal or full artificial light must be given. Although fluorescent (cool-white) lamps are the usual choice, they
are deficient in red light. Fluorescent lights can be used alone, though they are often used in combination
with incandescent lights (ordinary light bulbs) to supplement the red light. Incandescent lights may also be
used alone, but they produce more heat that can injure plants and are very low in blue light. Gro-Lux
 C N 31

lights give a slightly better light quality than cool-white fluorescent but are twice as expensive. High-inten-
sity lights have a good light spectrum and are more energy efficient. They are increasingly easier to install
and more economical. Because they have higher light intensity than other lamps, they can be hung farther
away from the plants. Of the two types of high-intensity lights, the metal halide lamps provide a better spec-
trum for plants and a more pleasing light to the human eye. High-pressure sodium lamps produce a yel-
lowish cast that some people find objectionable. Old lamps may need to be replaced as their intensity
diminishes. Light intensity can be increased by 20 to 50% by walls and benches with reflecting surfaces,
such as by lining them with aluminum foil.
Table 4.1 shows light intensities from two 4-foot fluorescent tubes of 40 watts placed at various heights
above the plants. Light intensity is given at 15 cm (6 in.), 30 cm (12 in.), 45 cm (18 in.), and 60 cm (24 in.)
below the light source. High-intensity lamps will give approximately twice the foot-candle readings shown
in Table 4.1. Use of a reflector will also increase the intensity of light. Most ferns grow best at 200 to 600
foot-candles. Light intensity is measured at the top of the plant. (Also see Appendix I.)

Table 4.1. Light intensity, in foot-candles (f.c.), below two 4-foot, 40-watt cool-white fluorescent tubes at various
distances above the plant material. The light intensity was measured on a horizontal surface with a light meter (Weston
illumination meter model 756).

P R
Little has been reported about the photoperiodic responses of ferns—that is, how they respond to
changes in day length. Such changes can affect growth and the ability to form spores (Patterson and Free-
man 1963). Some ferns seem to become deciduous, resume growth, or produce dimorphic fronds in
response to different day lengths.
32 C 4

Watering
H M W
Most cultivated ferns prefer moist—not soggy—soil. Continuously saturated soil should be avoided as
much as dry soil. A few cultivated ferns might prefer soil that is drier or moister than the average, but these
are exceptions. Ferns should receive less water during cool weather, especially when they are not actively
growing. Chapter 13 gives the water requirements for specific ferns.

H O
There is no precise answer to the question of how often to water ferns. Average conditions require
watering every two or three days, less during cool weather and more during warm. Plants in small pots dry
faster and need to be watered more frequently than those in larger pots. Plants in clay pots need to be
watered more frequently than those in plastic or glazed pots. Because water drains faster through sandy or
coarse soil mixes, plants growing in such soils will need to be watered more frequently than those in silt- or
clay-soil mixes. It is wisest to water a plant when it needs it rather than on a regular time schedule.

D W W I N

Do not wait for the plant to wilt. Feel the soil surface. If it is dry or starting to dry, it is time to water.
Another indication that the plant needs water is if the pot feels lighter than usual. A useful test for ferns in
clay pots is to tap the pot: if it is very dry, there will be a ringing sound. If you still are unable to decide
whether more water is needed, probe the soil to a depth of about 4 cm (1.5 in.) below the surface in a spot
where you will not injure the plant, and feel if the soil is dry. Peat-based soil mixes become lighter in color
when dry. In any case, water before the soil shrinks and pulls away from the sides of the pot.

H  W
Once you have determined that water is needed, thoroughly water the plant with a gentle sprinkle or
flow. Avoid forceful streams because large droplets will compact the soil or wash the soil away from the
roots. Stop when you see water coming through the pot’s drainage hole. If the plant is in the ground, be sure
that the water has reached the soil around the roots. Overhead watering, which wets the foliage, is not rec-
ommended for some ferns, particularly xerophytic species or those with finely divided leaves. These ferns
should be watered at soil level. In areas with high humidity and poor air circulation, you should water
plants in the early part of the day so that the foliage does not remain wet for long. This helps discourage cer-
tain diseases. Scant sprinklings of water will increase the humidity but are seldom sufficient for medium-
sized or larger ferns. Such sprinklings can also cause an accumulation of salts that are injurious to plants.

T  W P

A wilted plant needs water immediately. As you water, be sure that the water reaches the roots and is
not running down between the pot wall and the ball of dry soil. If this happens, submerge the pot in water
and soak it until it is thoroughly wet. Then remove it and drain. Thereafter water only to keep the soil moist.
Do not overwater. Wilted fronds that do not recover will turn brown in a few days and should be removed.
Keep as many green fronds on the plant as possible. Recovery of a wilted plant will depend on the extent
of damage to the roots and the plant’s innate vigor. Generally, the thicker the rhizome, the greater the
 C N 33

chances of recovery. Thicker rhizomes contain more stored food, and this fosters new growth to replace the
damaged tissue.

T  O P

Sometimes a plant wilts even though you have been watering it. The most common cause of this is over-
watering, which will damage the roots. Soil that feels wet even two or three days after watering may be too
wet for the plant. With extreme overwatering, the soil will feel soggy wet and may smell bad. When lifted,
the pot will feel heavy for its size because of the waterlogged soil. When tapped, it gives a dull thud.
Damage to the plant may be gradual or sudden. If gradual, the fronds will first yellow and then grow
poorly. The fronds might wilt, at which point examination of the roots usually reveals that they have rot-
ted away. If the plant hasn’t yet wilted, reduce the watering. If it has wilted, determine how badly the roots
and rhizomes are damaged. If most of the roots are rotted and the rhizome is soft and discolored, the plant
will probably not survive and should be discarded. Otherwise, trim off the rotted parts and replant the
healthy part in fresh, well-aerated soil. Keep the soil moist and hope for the best. Be careful not to overwa-
ter again. The cut ends of rhizome and roots may be dusted with a fungicide, such as Captan, to discour-
age further rotting.

H O C D

Roots need air. The oxygen contained in the air is used by the roots to metabolize their stored-food
energy, and this energy is essential to the life processes within the roots. Overwatering displaces the air
within the soil and replaces it with water. As a result, the roots suffocate. In addition, the excess water pro-
motes the growth of bacteria and fungi that can attack the plant. (Aquatic plants avoid suffocation by hav-
ing internal air canals that supply their underwater parts with oxygen.)

W Q  S I

Water your plants with a water temperature that feels comfortable to your hands, generally between 19
and 27°C (66–80°F). Be careful not to use hot water from garden hoses or pipes left in the sun; let the water
run and cool down first. Very cold water also should be avoided because it slows the uptake of nutrients.
Fluoride and chlorine are commonly added to tap water but not in amounts harmful to ferns. You
should not, however, water your plants with water from swimming pools. Its chlorine content and other
additives kills plants.
Salts in fertilizers provide the minerals plants need, but too much salt can slow growth or even kill
plants. Excess salts accumulate in the soil from repeated watering with water that is high in salts. If the salt
in the soil becomes too concentrated, the roots cannot absorb water, and in extreme cases water will move
out of the roots and back into the soil. When this happens, the result is known as root burn.
To reduce the risk of root burn, do not use water containing softeners because it contains sodium.
Excess sodium not only burns the roots but also harms soil structure and replaces desirable nutrients. Per-
haps the most common cause of root burn is overfertilizing. Do not add too much manure, ashes, or com-
post of sedge peat, which may have a high salt content. When salt becomes excessive, a crust of salt, usually
whitish gray, accumulates on the soil surface or flowerpot. Salt damage is common in arid areas, where the
soil is high in salts because there is insufficient rain to wash it away. Water supplies coming from arid areas
may also contain undesirable levels of salt.
Salt injury can be reduced by proper watering. Water ferns thoroughly until the amount of water equal
to about 20% of the pot’s volume passes through the pot. Provide good drainage to allow for adequate
34 C 4

leaching. Salt-encrusted pots should be cleaned or discarded (see the section in Chapter 7 on “Planting in
Pots”). Remember, light sprinklings of water will add salt to the soil, not leach it out.

A T  W

Sometimes watering a hanging-basket fern will damage the ferns planted below it because the water
draining from above batters the plants below. The plants underneath might also become overwatered. To
avoid this, the basket can be taken down and watered, or the plants underneath can be removed while the
baskets above are watered. Another alternative is to provide catch basins in the basket by building a sub-
stantial rim of moss around the upper edge of the baskets. This will hold more of the water and reduce the
spillage. Also, before planting, you can place a saucer at the bottom of the basket on top of the moss lining
to act as a hidden catch basin (see “Planting in Baskets” and “Special Care” in Chapter 7). Watering with
a fine gentle spray takes longer but will result in a better soaking of the basket and less runoff.
To prevent potted ferns from drying too quickly, plant the smaller pot into a bigger one. Fill the spaces
between the pots with sand, planting mix, or coarse sphagnum moss. This method is known as double pot-
ting (see Figure 4.1f ). Placing a watered plant in a clear plastic bag will keep it moist for a week or more
depending on how wide the bag is left open. If closed, the bag should have some ventilation holes, unless
ample air space surrounds the fern inside the bag. This is a handy way to care for ferns when you are away
for a week or so.

Air Circulation and Humidity

A M
Ferns need protection from drafts and hot, dry, battering winds. Although some ferns tolerate wind,
their fronds usually look ragged. A small amount of air circulation and ventilation is desirable in greenhouses
or similar enclosures. Air circulation discourages fungi and distributes humidity and temperature evenly.
Small fans, partially open windows, and vents circulate air adequately in the average home greenhouse.
Some outside air flowing into the greenhouse also can be beneficial, as it replenishes the carbon dioxide nec-
essary for plant growth. Incoming air is detrimental, however, when it is extremely dry, hot, cold, smoggy,
or apt to bring in insect pests.

H M H

Most ferns prefer conditions of 60 to 80% relative humidity during the day. Too much humidity will
cause condensation that promotes bacterial and fungal disease on the foliage. Older and more robust ferns
can tolerate less humidity, whereas more delicate or younger ones prefer higher levels. Most epiphytes and
all xerophytes can tolerate less humidity than other ferns.

L-H D
Low humidity is one of the most common causes of poor indoor growth. During the winter, the rela-
tive humidity in a typical, centrally heated living room is 40 to 50%—much lower than what most ferns
prefer. Increasing the air temperature worsens the problem because it lowers the relative humidity. Low
humidity can damage ferns in several ways. Emerging fronds tend to shrivel and older fronds yellow more
quickly. Typically, the fronds wilt or the margins become scorched. Barely adequate humidity can result in
 a
b

d f
e

Figure 4.1. Increasing the humidity: a. mister or humidifier; b. grouping plants; c. placing pots just above the water
level on pebbles in a tray or pot; d. and e. enclosing plants in plastic frames or glass; f. double potting.
36 C 4

thicker, smaller, harder-textured fronds that may be heavier spore-bearers. An indirect problem of insuffi-
cient humidity is increased infestations of red spider mite.

I  H

Humidity can be increased several ways (Figure 4.1). Mechanical devices for increasing humidity
indoors and in greenhouses are available on the market. Indoor units are controlled by electricity; green-
house and outdoor units are generally attached to the water line. All units produce a fine mist. In these
mechanical devices, the minute openings in the valve frequently become plugged and must be cleared reg-
ularly. Intermittent spraying of the air, foliage, and walkways with a fine mist from a garden hose is effec-
tive in increasing the humidity outdoors. A single misting is inadequate during hot, dry days because the
moisture evaporates quickly. Depending on the amount of dryness, misting might be needed several times
during the driest part of the day. Venting the household clothes drier into an adjacent greenhouse makes
good use of the warm humid air on days when it is needed. Grouping potted plants together or planting
them in the ground will keep them in more humid conditions. Increasing the humidity is desirable, but
always avoid overwatering.
A simple procedure to increase the humidity indoors is to place potted plants on pebbles spread out in
a shallow tray containing water. The bottom of the pot should not touch the water. Planting smaller plants
in glass or plastic containers or enclosing larger plants on the sides with plastic sheets or glass frames will also
help retain the moisture and humidity. The foliage can be misted during the hotter, drier part of the day.
Double potting (see “Additional Tips on Watering,” earlier in this chapter) will also increase the humid-
ity around plants. Kitchens and bathrooms are usually the most humid areas in the home, and if light is suf-
ficient, they are good places to grow plants.

H-H D
Excessive humidity, especially accompanied by condensation, is harmful for most ferns and should be
controlled, particularly during the night when the temperature may drop quickly. Control can be accom-
plished by watering or misting early in the day or by opening vents to increase air circulation. Excessive
humidity encourages fungal diseases and algae and results in the plants growing soft, weak, and succulent,
making them vulnerable to sudden climatic changes. Under high humidity and warm temperatures, some
ferns with firm, smooth fronds (such as polypodiums) develop brownish-purplish areas, particularly where
the fronds arch or bend. Finely divided fronds, as in some cultivars of Nephrolepis exaltata, might yellow and
shed their leaflets if the humidity is too high.

S  O H G

Heavy smog is known to damage young plants of Blechnum gibbum and Microlepia platyphylla (and it
probably damages others). The damage appears in about one day. The first signs are tan or purple spots near
the smaller veins on the fronds. These spots then dry and turn brown in about two days. Parts of the frond
or the whole frond may die. Installing anti-smog devices is impractical for home growers, and the only cor-
rective measure is to close outside air vents during periods of heavy smog or avoid planting smog-sensitive
species.
Other air pollutants such as ethylene, carbon monoxide, sulfur dioxide, and natural and manufactured
gas can injure ferns and other plants. Fern foliage suffering from toxic gases tends to curl under and/or turn
yellow. Typical symptoms in flowering plants are curling of parts, growth of dormant buds, and early petal
drop or closure. A sensitive test for the presence of ethylene is the early folding of carnation petals.
 5 Soils and Fertilizers

Soils
There are probably as many soil mixes as there are growers. Experienced growers prepare their own mixes
to best suit their particular style of gardening. If the grower waters frequently, a coarser mix will be used; if
less frequently, a finer mix. If the grower fertilizes regularly, a soil mix without fertilizer will be used. If fer-
tilizers are applied irregularly, the grower will add fertilizer to the mix or use a mix that contains rich gar-
den soil. Whatever mix is used, however, ferns need a growing medium that will hold moisture, provide aer-
ation, support the roots, and provide nutrients or retain added ones.

S  P

Beginners and even commercial growers may find it easier to purchase commercially prepared planter
mixes. Most of these contain 50 to 75% organic matter (peat moss, leaf mold, wood shavings, bark chips,
or various combinations of these), with the balance being inorganic matter (sand, perlite, vermiculite, soil,
or various combinations of these). Some potting mixes might also contain additional nutrients and wetting
agents. Soilless mixes are favored by most commercial growers because the mixes with soil are heavier and
can vary in quality depending on the type of soil used. Some standard mixes for ferns (with or without soil)
are given in Table 5.1.
A simple mix consists of one part peat moss and one part fine sand, to which is added 10 to 15 g per 3.8
liters (2–3 tsp. per gal.) of a 10-10-10 general fertilizer. The mix may need to be sieved with a screen to get
fine-textured soil for very young or small ferns. For basket ferns and terrestrial ferns requiring good drain-
age, add another part or two of coarse material, such as chipped or ground bark, coarse leaf mold, or per-
lite. (Also see the Cornell epiphytic mix in Table 5.1.)

S  O P B

Consider yourself fortunate if the soil in your planting bed is loam and at least half organic matter. From
time to time the organic matter might need to be replaced, but such a soil is ideal and easy to work for out-
door plantings. If the soil is on the sandy side, it can be improved by adding water-holding materials such
as peat moss. Sandy soils require more watering and fertilizing than other soils, but they are easy to work,
provide excellent aeration, and do not accumulate salts. Plants grown in sandy soils respond quickly to
fertilizer.

37
38 C 5

Table 5.1. Commonly Used Planting Mixes

Planting Mixes with Soil

G S M
Leaf mold 4 parts
Garden loam 1 part
Sand 1 part

G S M  L-L F

Leaf mold 3 parts
Garden loam 1½ parts
Sand 1½ parts
Calcium carbonate (ground limestone) 1 gram per liter of mix (or 1 ounce per cubic foot) or
more to bring pH to 7–8

Soilless Planting Mixes

U  C S M C (Matkin and Chandler 1957)
Peat moss (Canadian or German sphagnum or California 0.14 cubic meter (5 cubic feet)
hypnum)
Sand, fine 0.14 cubic meter (5 cubic feet)
Potassium nitrate 43 grams (1.5 ounces)
Potassium sulfate 43 grams (1.5 ounces)
20% superphosphate 397 grams (14 ounces)
Dolomite lime 78 grams (2.75 ounces)
Calcium carbonate (ground limestone) 397 grams (14 ounces)
If a moderate amount of reserve nitrogen is desired, add 397 grams (14 oz.) of hoof and horn or blood meal to the
above, but use the mixture within a week of preparation.
For limestone-loving plants, add more limestone, up to 397 grams (14 oz.).

Silty soils hold more water and nutrients than sandy soils, but they might need improved aeration for
good fern growth. Adding organic matter, sand, or perlite will improve the aeration.
Partial or light clay soils need additional coarse and resilient material to improve aeration and to keep
the clay from packing and becoming hard. Add coarse-textured organic matter until it makes up about half
or more of the soil. Perlite or gypsum (calcium sulfate) will also be helpful additives to reduce packing of
the clay. Be cautious about adding sand to certain clays, as binding reactions may occur.
If the soil is primarily dense clay, replace it with potting mix. The replacement is only needed in pock-
ets where the ferns are to be planted. Clay soils are difficult to dilute. Even if an equal amount of organic
matter is added to heavy clay soils, the soil will be suitable for ferns for only about three years. As the added
organic matter gradually decays, clays predominate more and more, and the result is poor aeration and
drainage. These conditions slow plant growth and promote rot, particularly during cool weather. Top
applications of mulch are usually too thin to improve a clay soil, and thicker mulch layers may bury the
ferns. In either case, little organic matter will reach the roots. If the soil has reverted to heavy clay, dig out
the ferns in spring, add organic matter, perlite, or gypsum, and replant. Tree ferns and large ferns are dif-
ficult to replant, so be certain that the soil is suitable before planting in the first place.
 S  F 39

Table 5.1. Continued

C F P M (Boodley 1972)

Peat moss (Canadian or German sphagnum) 0.14 cubic meter (5 cubic feet)
Vermiculte, #2 grade 0.07 cubic meter (2.5 cubic feet)
Perlite, medium-fine grade 0.07 cubic meter (2.5 cubic feet)
Dolomite lime 1.4 kilograms (3 pounds)
20% superphosphate 340 grams (12 ounces)
10-10-10 fertilizer, inorganic 0.45 kilogram (1 pound)
Iron sulfate 113 grams (4 ounces)
Potassium nitrate 170 grams (6 ounces)
Peter’s Soluble Trace Element Mix 5 grams (1 teaspoon)
For limestone-loving plants, add 1.4 kilograms (3 lbs.) of dolomite lime.

C E M (Boodley 1972)

Peat moss (Canadian or German sphagnum), shredded or 0.14 cubic meter (5 cubic feet)
screened through 6 mm (0.2 in.) mesh
Douglas fir bark, screened through 3–6 mm (⅛–¼ in.) 0.14 cubic meter (5 cubic feet)
mesh
Perlite, medium-fine grade 0.14 cubic meter (5 cubic feet)
Dolomite lime 1.4 kilograms (3 pounds)
20% superphosphate 1.2 kilograms (2.7 pounds)
10-10-10 fertilizer, inorganic 0.6 kilogram (1.3 pounds)
Iron sulfate 170 grams (6 ounces)
Potassium nitrate 227 grams (8 ounces)
Peter’s Soluble Trace Element Mix 15 grams (1 tablespoon)

Commercially Prepared Mixes

A number of commercially prepared mixes are available on the market. Some may need additions of perlite or sand for
better drainage.

Clay soils hold more nutrients and water than sandy soils. The main problems with clay soils are their
harsh texture, tendency to expand when wet and shrink when dry, and lack of aeration. Although tropical
ferns grow on clay, they are usually located on slopes with good drainage and in a climate where the clay sel-
dom dries and shrinks.
Coarse granitic soils or decomposed granite formed from the breakdown of granite are unsuitable for
growing ferns. They contain little or no organic matter and the main particles are coarse, resembling crum-
bly gravel. Although granitic soils contain some nutrients, water drains through them so quickly (due to the
coarse particle size) that the roots have little time to absorb the water. The fine roots of ferns will dry rap-
idly in such soil. For most ferns, growing on granitic soil requires incorporating large amounts of water-
holding, humus-forming material such as peat moss and compost.
40 C 5

R P B

Where appropriate, raised planting beds can be a good solution for poor soil or a lack of topsoil. Raised
beds may be erected over poor soil and filled with planter mix. If commercial planter boxes cannot be used,
redwood planks or rocks are suitable as retaining walls. Be certain that the soil is well drained, and if not,
provide drain holes to the sides and add a layer of pot shards, gravel, or coarse sand to the bottom of the
planting bed.

M
Mulches are loose materials placed over the soil, such as peat moss, leaf mold, and wood or bark prod-
ucts. Mulches serve several functions. They reduce evaporation from the soil, prevent mud from splattering
on the foliage, discourage weeds and some pests, protect exposed roots, and impart a neater appearance to
the garden. On foot paths, medium-sized bark or wood chips reduce soil compaction. Ferns benefit from
mulching, and spring is a good time to do so. The major precaution is to avoid burying the crown of the fern.

S A
Soil additives are incorporated into the soil to improve its quality or suitability for plants. New addi-
tives are constantly being developed. Some are made from recycled products such as rubber or Styrofoam,
and others are old materials that have had their character changed, such as baked clay. Some are products
that are given a new use, such as lichen (reindeer moss). The following are common soil additives used for
growing ferns.
Bark products. Bark can be shredded, ground, or chipped. Bark products decay more slowly than wood by-
products or peat and are used when a well-drained, long-lasting organic material is needed. Composted bark
(partly decayed) is often used in commercial planting mixes. Nursery growers favor using bark products
from fir trees.
Charcoal. Charcoal is added to mixes that might become waterlogged. It absorbs toxic materials produced
when the soil is poorly aerated. There is, however, a limit to what charcoal can do for a soil, and it is best
to avoid waterlogged soils from the start.
Fertilizers. Fertilizers contain minerals needed by the plant. Fish emulsion, blood meal, hooves, and horns
are examples of organic fertilizers often used in soil mixes for ferns. Planting mixes with added organic fer-
tilizers should be used within a week after being prepared. Inorganic fertilizers that can be added to plant-
ing mixes include potassium nitrate, potassium sulfate, superphosphate, dolomite lime, and calcium car-
bonate. Mixes containing inorganic fertilizers can usually be stored for a long time.
Humus. Humus is the dark, resilient, water-holding material that results from the partial decay of plants.
A “humusy” soil is one that contains much partially decayed plant material. Any plant material can be
made into humus. Compost, of course, is very high in humus.
Inorganic matter. Inorganic matter lacks substances derived directly from plants or animals. Inorganic
matter commonly used in planting mixes for ferns includes perlite, vermiculite, sand, and silt.
Leaf mold. Next to peat moss, leaf mold (or partly decayed leaves) is the most popular substance used in soil
mixes for ferns. Quality oak-leaf mold is increasingly difficult to obtain. In California it is illegal to remove
leaf mold from many areas, and if taken from lowland areas it may be infected with Armillaria or oak root-
rot fungus.
 S  F 41

Lime. See the discussion under “Acidic and Basic Soils” later in this chapter.
Loam. Loam is a soil that has a mixture of different grades of sand, silt, and clay in such proportions that
the characteristics of no one component predominate. An average loam soil contains about 1 part sand, ⅔
part silt, and 1 part clay, although the proportions can vary considerably depending on the grade (size) of
the sand, silt, and clay. Loam is an ideal general garden soil with sufficient drainage, aeration, water-hold-
ing capacity, workability, and nutrient-holding capacity.
Organic matter. Organic matter is anything derived from plants or animals or their remains. It adds nutri-
ents and humus to the soil. Organic matter commonly used in planting mixes for ferns includes peat moss,
leaf mold, sawdust, wood shavings, ground or shredded bark, compost, and manure. Manure and compost
are not favored for commercial planting mixes because of their variable quality and possible contamination
with plant diseases.
Oyster shell. See the discussion under “Acidic and Basic Soils ” later in this chapter.
Peat moss. Peat moss is partly or wholly decomposed moss that provides resilience and aeration to the soil
as well as water-holding capabilities. It may be composed of a mixture of moss species or of individual spe-
cies such as sphagnum moss or hypnum moss. Sedge or black peat does not contain moss and is not rec-
ommended for use with ferns because it might contain excessive salts. Milled peat moss is the common form
used in planting mixes.
Peat moss can be difficult to wet. Try moistening it a day or two before use. It will absorb warm water
faster than cold water, and adding a wetting agent to the water also speeds absorption. Milled peat moss may
need to be mixed and squeezed to ensure an even distribution of moisture. Certain brands of milled peat
moss come in a coarser grade suitable in soil mixes for growing larger ferns and epiphytes. Fine-textured,
milled peat moss breaks down more rapidly but is suitable in mixes used for growing spores or small plants.
Uncut moss consists of intact strands or clumps that have been harvested and dried. It is coarse and
stringy and is often used to line hanging baskets or to plant epiphytes. The best kind comes from various spe-
cies of sphagnum moss. Other uncut moss, usually taken from tree trunks, is a mixture of various moss spe-
cies and has a harsher texture and holds water less evenly. This kind is sometimes sold as green moss and
should not be confused with the live sphagnum moss used to grow insectivorous plants. Uncut moss holds
moisture yet provides excellent aeration if not kept too wet. When used for epiphytes mounted on boards
or tree trunks, pads of uncut moss are placed around the roots and rhizomes. When used in pots, the moss
should not be packed too tightly because it might hold too much water. The purity of commercially avail-
able uncut moss has deteriorated over the years, and nowadays it often contains twigs, grass, and leaves or
is very fragmented. Uncut sphagnum moss from New Zealand is expensive but of excellent quality. Span-
ish moss (Tillandsia usneoides) should not be confused with the uncut sphagnum moss of horticultural use.
Uncut moss that is soaked and drained before use is easier to work. Use gloves when handling wet
uncut moss. Cuts in the skin occasionally become infected by a fungus that lives in the moss. Coconut fiber
and tree-fern trunk fibers are sometimes substituted for uncut moss in lining baskets, but they are stiffer and
retain less water than peat moss.
Perlite. The drainage and aeration of soil can be improved by adding perlite, which is also known by the
brand name Sponge Rok. Perlite is a sterile inorganic material derived by heating and expanding siliceous
rock. It is light, porous, loose, and firm. The particles come in various sizes.
Rockwool (Gordan). Rockwool is a sterile substance made from coal, carbonate, and basalt heated and
blown into fibers. It is 97% air, chemically inert, and does not decompose. Rockwool is used in soilless
mixes. Compressed forms come in blocks, mats, or tubes for special horticultural or floral use. Do not
reuse rockwool that is designed for floral use because it might contain formaldehyde.
42 C 5

Sand. Sand refers to rock particles in a particular size range. The fine grade, commercially designated as
United States Screen Mesh size #30 (approximately 0.5 mm in diameter), is recommended for potting
mixes, although finer sizes to #270 (0.05 mm in diameter) may be used. Coarser grades of size #12, #16,
or #20 (1.6 mm, 1.2 mm, and 0.85 mm in diameter, respectively) are added to soil to increase aeration and
drainage. Size #8 (2.5 mm in diameter) and lower number sizes are designated as gravel. Quartz sand and
washed builder’s sand are suitable for gardening. Do not use sand from ocean beaches or other saline areas
unless it is washed to remove excess salts that might otherwise burn roots.
Sawdust and other wood by-products. Adding wood by-products such as sawdust, shavings, or chips
increases the humus content and improves the soil structure. Wood by-products, however, contain few min-
eral nutrients.
Fresh or partly decayed sawdust (or other wood waste) may be used. Most fresh sawdust and wood
waste contain no appreciable amounts of toxic substances, and those toxins that might be present will
decompose within a few weeks after incorporation into the soil. Reports of toxicity are associated with
finely ground sawdust that rots under conditions of low oxygen, producing toxins in the process. Sawdust
is converted to humus by microorganisms that need air and mineral nutrients to grow. Therefore it is best
to aerate the compost and provide additional minerals for the microorganisms. Add 5.4 kg (12 lbs.) of 10-
6-4 fertilizer for every 45.4 kg (100 lbs.) of sawdust. Other fertilizers rich in nitrogen and phosphorous may
be used. Nitrogen is the most important mineral, and calculations are based on supplying 0.5 kg (1.1 lbs.)
of nitrogen per 45.4 kg (100 lbs.) of sawdust. Apply the minerals at intervals throughout the composting
period. Nitrified sawdust has nitrogen added, sometimes in forms or amounts that can burn fern roots.
Soil. Typically, soil is decomposed, finely divided rock material with a small amount of organic matter.
Sand, silt, and clay (listed from largest to smallest size) are the most important rock particles in garden soil.
Sand provides excellent drainage, aeration, and workability, but retains little water or nutrients. Clay, the
finest textured particle, has poor drainage, poor aeration, and difficult workability, but retains water and
adsorbs and retains nutrients. Silt, being intermediate in size to sand and clay, has essentially intermediate
properties but lacks clay’s chemical ability to attract and retain nutrients.
Vermiculite. Vermiculite is a sterile inorganic material produced by heating bits of mica and causing them
to expand. It comes in various particle sizes and contains some potassium and calcium. Vermiculite adsorbs
minerals and makes them available to plant roots. It also retains water but tends to collapse in a relatively
short time. Wet vermiculite should not be pressed or compacted because its pore structure will be ruined.

A  B S

Growers place much importance on whether a soil is acidic or basic (alkaline). The importance is not
in the degree of acidity itself, but in the acidity’s effect on the availability of soil nutrients to plants. If your
plants are growing well, do not worry about soil acidity. If they are languishing and unresponsive to bal-
anced fertilizers, check first for unfavorable growing conditions, then check for acidity. In general, regions
with arid climates have basic soils and ones with rainy climates tend to have acidic soils.
Determining acidic or basic soil. The range from an acid to base condition is known as the pH range and
is measured on a logarithmic scale from 1 to 14. A pH of 7 is neutral, less than 7 is acidic, and more than
7 is basic. The greatest amount and variety of nutrients are available to plants at pH values between 5.5 and
7. Therefore, most plants grow best somewhere in this range. Most ferns flourish between pH 6 and 7.
Many plants show iron deficiency (yellowing of the leaves) at pH levels of 5.5 to 6, although this has not
been reported for ferns. An inexpensive way to determine soil pH is with pH paper, which turns a color cor-
responding to a particular pH value.
 S  F 43

Limestone-loving ferns or ferns of basic soil (pH 7–8). Ferns native to basic soils or limestone need more
calcium than other ferns. Chapter 9 includes a list of ferns for such soil. To grow these ferns it might be nec-
essary to add calcium, usually dolomite (CaCO3 · MgCO3), ground limestone (CaCO3), or oyster shells.
Burned or quick lime (CaO) and hydrated or slaked lime (Ca[OH]2) should be used sparingly because small
amounts can greatly increase the pH and harm the plants. Where limestone is available, ferns can be grown
in soil pockets between these rocks, or crushed limestone may be added to the soil. Some growers have had
success using pieces of concrete instead of limestone.
Testing for calcium in the soil is easy. Obtain a teaspoon of a strong acid, such as sulfuric acid or muri-
atic acid (hydrochloric acid or hydrogen chloride). Sulfuric acid may be obtained from a car battery. Muri-
atic acid may be obtained from some toilet bowl cleaners. Place 5 ml (1 tsp.) of the acid on a cup or less of
soil and watch for bubbling. Bubbling indicates the presence of calcium in the form of lime (calcium car-
bonate). If no bubbling occurs, you may not have enough calcium in the soil, or you may have one of the
few soils that has calcium but does not respond to this test, in which case you might have to have the soil
tested commercially.
Acid-loving ferns (pH 4–7). Some acid-loving ferns grow in wet, marshy areas where decaying organic mat-
ter produces carbonic acid. Other acid-loving ferns may grow in well-drained soil that is acidic because its
calcium and basic salts have been leached away. A peaty or highly organic soil will provide acidic conditions
for ferns. Also, soils can be acidified by adding aluminum sulfate, ammonium sulfate, ammonium nitrate,
or finely ground sulfur. The latter is the most economical and efficient additive. Chapter 9 includes a list
of ferns for acid soils.

S S
In the strict sense, soil sterilization means killing all microorganisms, but in general horticulture it
usually refers to destroying disease organisms, insects, and weeds. It will here be referred to as soil “treat-
ment” to distinguish it from true sterilization.
Except for commercial growers, planting mixes for ferns are usually not given soil treatment unless dis-
eases are present or the mix is used for growing fern spores or young ferns. The use of compost, manure,
soils, or leaf mold in soil mixes increases the need for treatment. Soil mixes can be treated using chemicals
or heat (complete soil sterilization is usually done in an autoclave or oven). For information regarding soil
treatments, the commercial grower will find the publication by Matkin and Chandler (1957) particularly
helpful.
Chemical treatments. Chemicals such as methyl bromide and formaldehyde can be used to treat soil but
not, of course, around living plants. However, such chemicals are restricted or no longer available or have
been discontinued for health and environmental reasons. Home growers might find it easier, although not
as effective, to use fungicide drenches such as pentachloronitrobenzene or PCNB (Terraclor), thiram
(Arasan), captan, or quaternary ammonium compounds such as Consan or Physan 20. (See Chapter 11 and
Appendix IV.) These products can be used around live plants.
Steam treatment. Most commercial growers use steam to treat the soil. The steam is applied through var-
ious types of equipment until all parts of the soil have been heated to 71°C (160°F) for 30 minutes. For a
better margin of safety, heat at 82°C (180°F) for 30 minutes. High heat, however, might produce toxic con-
ditions. If this happens, let the soil stand for a few weeks or leach it thoroughly with water.
For small amounts of soil, a homemade steam sterilizer can be devised for treatment. Get a large cook-
ing pot or roasting pan and fill with about 1 cm (0.5 in.) or more of water. Place a rack in the pan to raise
the soil (or to raise pots containing the soil) above the water level. Cover and heat this over the stove or in
44 C 5

the oven at about 120°C (250°F), or high enough to produce steam. Be sure that the steam penetrates all
parts of the soil for at least 30 minutes. This might require placing the oven controls at about 150°C
(300°F) for half an hour if there is much mass to be penetrated. Very moist soil wrapped in aluminum foil
can also be heated in this manner.
Since most plastic pots or boxes cannot be treated with high heat, they have to be disinfected separately
by soaking for 30 minutes in a 10% bleach solution, after which they are rinsed with previously boiled water
and then filled with the treated soil.
Microwave treatment. Treatment in a microwave oven is suitable for small amounts of soil. Add generously
moistened soil to a 2 liter (2 quart) casserole dish, cover, and set the oven on high for 5 to 15 minutes, or
until the steam thoroughly penetrates all the soil. Keep it covered while cooling.
Boiling water treatment. Many growers find that gently pouring boiling water over pots filled with small
amounts of soil and provided with good drainage is a satisfactory treatment, especially if the soil will be used
for growing spores. For a given amount of soil, pour at least twice as much boiling water gently through it.
By placing a clean paper towel over the moist soil before pouring, you can prevent much of the lighter mate-
rial in the mix from floating up to the surface.
Direct boiling treatment. Although it kills soil pathogens, boiling soil in a cooking pot is not recom-
mended. This treatment ruins the soil’s structure, reducing it to a soggy mush that takes a long time to drain
and, upon drying, lacks aeration.
Steam sterilization. Complete sterilization kills all organisms including their spores. Except for scientific
work, such as growing ferns in agar, full sterilization is unnecessary. In steam sterilization, the material to
be sterilized is placed in an autoclave at a pressure of 1 kilogram per square centimeter (15 lbs. per sq. in.)
for 15 minutes or longer (depending on the amount) at a temperature of 121 to 123°C (250–254°F).
Small amounts of soil, pots, dishes, or other items can be sterilized in a home pressure-cooker at 1 kilogram
per square centimeter (15 lbs. per sq. in.) for 20 to 30 minutes.
Dry heat sterilization. In dry heat sterilization, soil is placed in an oven at 160°C (320°F) for two hours or
longer, depending on the amount of soil. This method has the disadvantage of making the soil difficult to
wet; however, it is useful for sterilizing paper envelopes used to store sterilized spores. To sterilize envelopes,
put them in a heavier box before heating them to the oven temperature given above.

Fertilizers
Ferns growing in rich soils are well supplied with mineral nutrients and do not need fertilizers or other sup-
plements. Although most gardeners do not have such fertile soils, they want the best-looking plants possi-
ble. Ferns that receive ample fertilizers or mineral nutrients grow faster and bigger and more luxuriant.
Ferns, however, are extremely sensitive to overfertilizing, and the novice grower, being aware of this fact,
is often hesitant to fertilize. It is therefore important to learn how to select the proper fertilizers and apply
them correctly.

W F N F

You can tell when a fern needs fertilizer because it grows slowly, is smaller than normal, and produces
poor-quality foliage. Ferns producing healthy green fronds do not need fertilizers. If you do not want a fern
to grow large or quickly, do not fertilize unless it shows poor foliage color. In that case, light applications
of fertilizer will improve the color.
 S  F 45

C  F
Beginners are advised to select fertilizers that have a reputation for “low burn.” These contain a low per-
centage of highly soluble salts (that is, salts that dissolve quickly in water). Liquid fertilizers with a low-burn
reputation include fish emulsion and those containing urea. Dry fertilizers with low-burn properties include
fish meal, cottonseed meal, castor-bean oil, and activated sewage sludge.

L V D F

Liquid fertilizers are probably safer and easier to use for the beginning grower. They can be applied
while watering, and this saves labor. Also, there is less likelihood of accidental overdose as a fertilizer is
diluted for application. A disadvantage of liquid fertilizer is that some of it quickly passes beyond the root
zone and is wasted. That which remains will also leach out of the soil more rapidly than dry fertilizers, and
so liquid fertilizers need to be applied more often. With dry fertilizers, you must spread the powder or
granules thinly and evenly over the soil surface. If the dry fertilizer contains much soluble salt and is acci-
dentally concentrated in one spot, it will injure the plant.
Fertilizers have been developed that slowly release soluble salts for three to nine months. Some release
a bit of their salts each time the plant is watered. Because ferns require more frequent watering than other
plants, be careful not to overfertilize when using these fertilizers. New fertilizers (such as Nutricote) release
nutrients at certain temperatures, making frequent watering less of a problem. Dry and slow-release fertil-
izers are not recommended for staghorn ferns (Platycerium) because the granules can get caught behind the
base fronds and injure the tissue.

O V I F

Organic fertilizers are derived from organic substances, whereas inorganic fertilizers are mostly derived
from chemicals or mining processes. Fertilizers such as fish emulsion, bone meal, or castor-bean meal are
organic; ammonium sulfate and superphosphate, among others, are inorganic fertilizers.
Beginners are advised to use organic fertilizers with low-burn reputations and defer use of inorganic
types unless they also have a low-burn reputation. Generally, organic types contain fewer soluble salts than
inorganic types and result in less burn. They also release their nutrients more slowly than inorganic types and
tend to last longer in the soil. Inorganic fertilizers, however, produce quicker plant responses. Exceptions occur
in both categories of fertilizer in regard to soluble salt content. Blood meal, an organic fertilizer, contains a high
amount of soluble salt and may readily burn plants if applied carelessly. Some inorganic fertilizers have salts
that dissolve slowly in water and therefore burn plants less readily. Do not use manures in places such as
greenhouses where warmth and high humidity prevail. Their use encourages bacteria and molds.

C  I F

Beginners should use complete fertilizers, which contain the three most important elements needed by
a plant: nitrogen (N), phosphorus (P), and potassium (K). Incomplete fertilizers have only one or two of
these elements. Complete fertilizers are generally preferred over incomplete ones, although results vary
according to the plant, soil, and cultural conditions.
The approximate percentages of each of the three main elements contained in a fertilizer is cited on the
package as a series of three numbers. The first number refers to nitrogen, the second to phosphorus, and the
third to potassium. Most fish emulsions have percentages of 5-2-2, while hoof and horn is listed as 15-0-
0. Fish emulsion is a balanced fertilizer whereas hoof and horn is not.
46 C 5

Plants require several elements, but nitrogen is the most important. Trace elements (those needed by
plants in minute amounts) are usually abundant enough in the soil or in fertilizers as impurities. An excep-
tion is calcium. Non-limestone-derived soils in areas of high rainfall often must have calcium added, usu-
ally in the form of dolomite or ground limestone. The amount of calcium in a fertilizer is usually indicated
by percent on the label.

F A
Always follow the manufacturer’s directions for application of a fertilizer. Never give more than the rec-
ommended dosage. If you are uncertain about the fern’s sensitivity to the fertilizer, there is no harm in
reducing the concentration. Some growers regularly use only half the recommended dosage. It is more
beneficial to give smaller amounts over several applications than to give the full dosage all at once. Although
this involves more work, the plant makes better use of the fertilizer.
Fertilizers that must be diluted should be diluted accurately. Dissolve as much of the fertilizer as pos-
sible before application. For watering large numbers of plants, a range of mechanical devices are available.
Great time-savers are those that mix the fertilizer with the water as it flows through the hose. Watering
devices that operate only with a strong stream of water are not desirable, as the force of the water may dam-
age the fronds and wash the soil from the roots.
Powders or granular forms must be sprinkled evenly and thinly over the moist soil surface. Do not mix
these fertilizers into the soil by tilling. Fern roots are fine and fibrous and grow close to the surface. Tilling
will injure the roots. Water thoroughly immediately after applying the fertilizer powder or granules. Any
fertilizer accidentally spilled on the foliage should be washed off immediately. If too much fertilizer has been
accidentally applied to one spot, particularly if near the crown of the fern, remove it or spread it out thinly
to reduce the possibility of burning the fern and encouraging undesirable mold growth. Although organic
fertilizers require microorganisms such as molds to decompose them and release their nutrients, a high
concentration of mold may damage the fern.

F  A
The frequency of fertilizer application depends on the manufacturer’s recommendations. On the aver-
age, one application every three weeks is needed for liquid fertilizers and longer intervals for dry fertilizers.
Fertilize infrequently if you do not want the fern to grow too large. Ferns that do not enter any noticeable
rest period but continue to grow actively throughout the year benefit from fertilizers applied at any time.
If you want to discourage new growth prior to winter or for fear of frost damage, do not fertilize late in the
season. Also, do not fertilize ferns during their periods of rest or dormancy because there is little intake of
nutrients at that time.
Where there is heavy watering or frequent rainfall on porous soils, the water will leach the fertilizer.
Ferns growing in such areas should be fertilized more often. On the other hand, soils that do not drain read-
ily retain fertilizer and can be fertilized less frequently. For commercial growers, the recommended nitro-
gen level for maintaining the best fern foliage color is about 150 ppm, which amounts to about 2.5 ml (½
tsp.) of a fertilizer in 3.8 liters (1 gal.) of water if the fertilizer contains a 20% nitrogen content. This should
be applied as needed.

T O F

Overfertilization results in burned or wilted foliage. Damage to roots and rhizomes may also occur. If
you have accidentally overfertilized, immediately remove all fertilizer that might be on the surface and
 S  F 47

make sure that the soil is well drained. Then water heavily to leach out any remaining fertilizer. Later water-
ing should be sparing so that the soil becomes well aerated. Increase the humidity and remove badly burned
and wilted, dead fronds. Recovery of the plant usually depends on how badly the roots and rhizomes have
been damaged. Young ferns or those with thin, nonfleshy rhizomes are more difficult to save than older ferns
or those with fleshy rhizomes.
Successful fertilization works on the principle that a little soluble fertilizer salt is good but too much is
bad. Damage from overfertilization is caused by the release of too much soluble salt. This causes water to flow
out of the roots and into the soil, thus dehydrating, wilting, and eventually killing the plant. If only some
of the roots are injured, there may be a partial wilting and/or the characteristic leaf scorch or burned appear-
ance. The partial loss of roots or foliage sets the plant back, and it may be stunted for some time. Fertilizer
salts on the leaf can also dehydrate and burn the tissue. (See “Water Quality and Salt Injury,” in Chapter 4.)

Ferns and Plant Hormones

The effects of plant hormones, such as auxin and gibberellic acid, on ferns are complex and reported in sci-
entific journals. Gibberellic acid has been shown to increase the growth rate of young walking ferns (Camp-
tosorus rhizophyllus). Other hormones are known to interact in complex ways on the germination of spores
and the early development of prothalli (gametophytes) and sporelings. The quality of light and various
chemicals also interact with plant hormones to affect spore germination and prothallial development.
 6 Through the Year with Ferns

Year-round Needs

Where ferns grow throughout the year, it is necessary to water, fertilize, and groom year-round. Elsewhere,
for best growth, fertilize more frequently during the growing seasons and less so or not at all during the cool
months. Watch for slugs and snails as well as other pests and for sudden and damaging changes in the
weather.
Remove old or badly damaged fronds from vigorously growing plants. Cut them off at the base of the
stipe. Removing these fronds maintains a neat appearance and reduces hiding places for pests. If the plant
is growing poorly and has only a few fronds, avoid removing any fronds except those that are totally brown
or yellow. Sometimes only the tip or margin of an otherwise lovely frond is disfigured. Instead of remov-
ing the whole frond, the disfigurement may be trimmed away with scissors and the remaining part cut
into a shape that is close to the natural one.
There are advantages and disadvantages to the grooming practice of cutting off all the tattered, leggy,
misshapen fronds along with the undamaged fronds. The advantages are
1. In dense frond clusters it is easier to cut off all the fronds than to single out only the
damaged ones.
2. In a few weeks, fresh new compact growth will appear.
3. The frond shapes will be symmetrical due to greater space for development.
The disadvantages are
1. Certain species are harmed by having all their fronds removed at the same time.
2. Encouraging vulnerable new growth just before a frost or cool weather may bring about the
demise of non-deciduous ferns.
3. The bare look, even if temporary, is unattractive to some gardeners.
If the advantages outweigh the disadvantages, then be sure that the plant is a vigorous grower that ordi-
narily produces many fronds during the growing season (as are the common species of Adiantum, Pteris, and
Microlepia). Also, make sure that the fronds are removed when plenty of growing season lies ahead. In areas
with short growing seasons, cutting back all the old fronds is best done in early spring before the fiddleheads
have emerged. In areas with longer growing seasons, this may be done later. In areas with year-round grow-
ing conditions, the time to cut back fronds is not critical for most non-deciduous ferns.
Ferns benefit from an occasional fine spray to remove dust, dirt, and lurking insects. Fronds with hairy
surfaces tend to catch debris that does not wash off easily. Usually the debris can be loosened with a soft
brush.

48
 T  Y  F 49

Leaf spots left by salts in the water are harmless if not too numerous. If conspicuous and unattractive,
the spots can be removed by misting them gently with a liberal amount of water to wash off the salts. About
half a tablespoon of mild household detergent added to a quart of water (about 15 grams per liter) helps dis-
solve the salt faster. This treatment must be followed with a thorough water rinse. When applying the
detergent solution and rinse, tilt the pot to prevent the detergent from dripping on the soil. Spots can be
avoided if rain or de-ionized water is used instead of tap water. Remember that liquid fertilizers applied on
the leaves will also produce water spots. Commercial leaf polishes can be used on thick, leathery ferns to
remove spots and to shine the surface, but the shine does not necessarily help the plant.

Winter Care
If you live in a temperate area, your outdoor ferns will be dormant during the winter and require little care
other than checking that their protective coverings or shelters are not disrupted. Outdoor ferns in sub-
tropical climates grow slowly during the winter, and there is little to be done besides watering, removing
withered fronds, and fertilizing once or twice a month (or not at all). Winter is the low point in an outdoor
subtropical fern garden, but if the fronds are taken care of in late summer and fall, they will grace the
garden throughout the winter. Watch for early spring growth and remove old fronds before the emerging
fiddleheads become hopelessly entangled in them. Indoor ferns, greenhouse ferns, and those in tropical gar-
dens require their usual care, although species with a rest period should be watered and fertilized less.

Spring Care
In temperate and subtropical areas, ferns grow vigorously in the spring as new fiddleheads emerge and
uncoil. In temperate areas, the winter shelters can be removed in spring. In all areas, dead and old fronds
should be removed before the new fiddleheads emerge. This minimizes injury to the new fiddleheads and
gives them room to develop. Aphids commonly appear on ferns in early spring, so watch for them as well
as other pests. If the winter rains have washed some of the soil away from the roots, replace this soil or cover
with mulch, particularly the area around the tip of the rhizome where new roots emerge. Resume fertiliz-
ing every two or three weeks if maximum growth is desired. All ferns must be protected from early heat
waves, particularly while new fronds are uncoiling since they are soft and easily dehydrated. Spring is a good
time to divide, transplant, or repot ferns, and this should be done before the fiddleheads uncoil.
Some ferns will shed all or most of their fronds just before putting forth new ones. The squirrel’s-foot
fern (Davallia mariesii var. stenolepis) drops its fronds in late winter or early spring; the Canary Island daval-
lia (D. canariensis) drops its fronds in spring or early summer; and the knight’s polypody (Goniophlebium
subauriculatum ‘Knightiae’) drops its fronds in late spring. These ferns look forlorn with their withered
fronds, but new fronds will soon clothe the basket again. Remove any lingering old fronds to make way for
the new.

Summer Care
Most ferns produce new fronds well into the summer. Their older fronds that turn yellow or brown should
be removed. Fertilization should be continued for maximum growth. Watering and keeping the air humid
will be the major problem in some areas, especially for hanging baskets and pots. Watch for thrips and scale
insects that usually appear at this time. Snails, slugs, and other pests will also be active. In many parts of the
50 C 6

country that have a long growing season, it is still possible to divide, replant, and repot in summer. Trim
the more robust ferns when they threaten to shade the smaller and slower growing species. Be ready to mist
your ferns when the weather gets too hot. Avoid overwatering the soil when increasing the humidity; sprin-
kle the walkways instead of saturating the soil. Fern spores are most frequently formed at this time and in
the fall.

Fall Care
From late summer into fall, fern growth wanes and less water is required. Fertilizers should be reduced or
withheld. Deciduous ferns will yellow and brown, and new growth stops until spring. Leave the old fronds
on the plant to protect them through the winter in areas where frost and freezes occur. In very cold climates,
some ferns need mulches and even screens or frames built over them. It is important to mark or remember
where deciduous species are planted so that they will not be accidentally dug up or injured during winter
garden activities.
In subtropical gardens with a long growing period, the ferns will still need to be groomed and watered.
Where there is danger of frost, do not remove green fronds, even if they are tattered, because removing them
will encourage new growth that could be damaged by frost. Resist the urge to trim until the danger from
frost has passed. Also reduce fertilizers to discourage new growth before winter. As the weather cools, ferns
need less fertilizer and water. Most of the fronds that remain on the plant in late summer and fall will stay
on the plant throughout the winter; however, if temperatures are mild, some ferns will continue to produce
a few smaller fronds in the months ahead. If the temperature drops very much, the fronds will yellow.
 7 Planting

Planting Time
Spring is generally the best time to plant, replant, transplant, or propagate ferns. It allows for a long grow-
ing period so that the fern can establish itself before cool weather. Ferns that form only one set of fronds per
year—for instance, some osmundas—are best moved in early spring before the new fronds have appeared,
or in fall after the old ones have died. If they must be moved in leafy condition, preserve as much of the
foliage as possible to ensure good growth the following year. Ferns that produce new fronds throughout a
long growing season can be safely moved while in leaf. Avoid moving ferns with soft, newly formed fronds
because they wilt and damage easily. If a fern must be moved at this time, avoid disturbing the roots and
keep the root ball intact (as is done for all transplants). After replanting, water well and keep the plant
shaded and the air humid.

Beds, Pots, or Baskets?

Deciding whether to plant ferns in the ground, in pots, or in hanging containers is determined by the cli-
mate, setting, personal preference, and species being planted. Ferns planted in the ground usually require
less care than those in pots or baskets. They do not dry out as fast, and the soil acts as a buffer to absorb extra
moisture, excess fertilizer, temperature extremes, and other unfavorable conditions. Also, ferns planted in
the ground do not have to be replanted as frequently as those grown in pots. Hanging containers, however,
show cascading or drooping fronds to best advantage. Trough gardens (discussed in Chapter 10) may be
large enough to have conditions between those of ground plantings and container plantings.
If you want to move your plants around for decorative purposes or winter protection, plant them in
containers. If you want to keep a natural effect with potted plants, sink the pots in the ground to hide
them. Many ferns that grow naturally in the ground (terrestrial ferns) can be planted in baskets or in pots
subsequently placed in a basket. Terrestrial ferns in baskets or pots need to be kept moist. Ferns that natu-
rally grow on trees (epiphytic ferns) or on rocks (epilithic or saxicolous ferns) are generally planted in pots
or baskets with a loose soil mix, but they can also be planted in the ground if the drainage is good.
Terrestrial or epiphytic ferns with long-creeping rhizomes bearing fronds that grow far apart are gen-
erally unattractive in pots. They soon grow to the edge of the pot and can die from salt burn if salt has accu-
mulated around the edge, or if the plant grows beyond the edge of the pot it might die from drying. Some-
times the rhizome will grow down into the pot and become trapped at the bottom. In any case, the old
portions of the rhizomes are left exposed and in full view at the top of the pot. With age they become unat-
tractive. For these reasons, ferns with long-creeping rhizomes should be planted in beds or moss-lined bas-

51
52 C 7

kets where they can root over a broader surface. Nevertheless, they are usually planted to best effect in the
ground where they have ample space to grow and are free to climb up trees or moss-covered posts.
Ferns with closely spaced fronds, erect or semi-erect stems, or short-creeping rhizomes are best for
planting in pots where they can display their foliage patterns. Ferns native to dry or seasonal climates are best
planted in pots or isolated places in the garden where their particular watering needs may be more easily
met. Trough gardens are especially suitable for such purposes (see Chapter 10). Bottle gardens or terrari-
ums are used to grow ferns and selaginellas requiring high humidity.

Planting in Beds
S  S
Before planting in the ground, check the site for proper light, wind exposure, and soil conditions.
Anticipate the full size of the fern you are planting. Will it fit your landscape needs? It is easier to consider
these factors now rather than later. If possible, start with hardened plants—that is, ones acclimated to
exposed conditions. Greenhouse plants moved outdoors can wilt, especially if they have soft succulent
growth. These plants should be gradually exposed to drier air over a week or more.

P  B

Prepare the soil so that it is moist and loose (friable). Use soil that contains at least half organic matter
(see Chapter 5). Soil in the planting bed does not have to be deep because fern roots generally grow near the
surface. About 20 to 45 cm (8–18 in.) of soil is sufficient for small to medium ferns. Tree ferns grow best
with 60 cm (24 in.) of soil or more. These requirements do not mean the entire bed needs to be deepened.
If the soil is unsuitable for ferns and you do not want to change the whole bed, plant the ferns in pockets
of suitable soil and provide good drainage, especially in heavy clay soils, by adding coarse gravel or shards
to the bottom of the planting hole. An area of suitable soil for ferns should be large enough to accommo-
date the expected spread of roots. Usually this equals about one-third to one-half of the spread of the
fronds. Once the soil has been prepared, dig the hole to the depth and width of the container. Gently com-
press the soil at the bottom of the hole so it will not settle too much after planting. Then proceed as follows.

B P P

Getting the plant out of the container. If the soil in the container is dry, water it. If possible, do this a day
before removal so that the soil will not be muddy. Moist soil will not fall away from the roots as easily as dry
soil and is less likely to stick to the inside walls of the pot. If the fern is in a can, have the nurseryman cut
the can. Ferns in small pots are easily tapped out. You can do this by supporting the top of the soil with one
hand, turning the pot over, and giving the rim a sharp tap on the edge of a bench (Figure 7.1). The ball of
soil and the fern should slide out intact. Lay larger pots on their side, protect the edges with padding, and
then tap with a mallet until the mass of soil slides away from the pot. Applying water pressure from a hose
into the drain hole can float out stubborn soil masses.
Digging ferns from the ground for transplanting. If a fern must be taken out of the ground and replanted,
the best time to do this is in early spring. Water the plant a day or two before transplanting so that the soil
will be moist enough to cling to the roots (but not muddy) when the plant is moved. The diameter of the
root ball and soil should be about half the spread of the fronds. The depth will depend on the size of the fern.
 P 53

The soil and roots should slide

out of the pot intact.

Have the soil moist.

Support the soil with one hand.
Invent the pot and tap it sharply
on the edge of a table.
b

Have the soil moist.

Place the pot on its side.
Protect the edges of the pot with padding.
Tap edges with a mallet.
Soil and roots should slide out intact.

Figure 7.1. Removing plants from pots: a. from a small pot; b. from a large pot.

Positioning and planting the fern. Normally, note the soil level of the fern you are moving and replant it
at the same level (Figure 7.2). If an erect or semi-erect fern has been tipping over, remove the old stipe bases
and bury the rootstock or rhizome deeper in the soil, leaving its crown or growing tip uncovered, not
buried (Figure 7.3). Measure the height of the soil ball. Add or remove soil from the hole (or pot) to bring
the crown to the proper level when set in place. For pots, the level of the soil should be about 1 cm (0.5 in.)
or more below the top edge to provide a catch basin for water (Figure 7.4). All added soil should be gently
firmed in place.
If the soil attached to the roots is poor, gently shake most of it loose before replanting. Remove dead,
old, and inactive roots. Cut old broken fronds and stipe bases at the rhizome. If the fern is badly root-
bound, gently loosen some of the surface roots and spread them out.
For creeping ferns, position the growing tip of the rhizome to face the direction in which you wish it
to grow. If rhizomes creeping over the soil surface refuse to stay in place after planting, anchor them with
pieces of bent wire or small rocks. When planting in pots, place the growing tip as far from the edge as pos-
sible (Figure 7.5). This might place the foliage at the edge of the pot instead of the center, but it will pro-
vide maximum room for future growth.
Once the plant is positioned, fill in the spaces around the roots. Gently press the soil into place (but do
not compact it) as it is added until you reach the desired level. Do not cover the crown or growing point if
it was growing exposed before. Water the plant well with a gentle sprinkle. A strong stream of water will flat-
ten the plants, wash soil from the roots, and compact the soil, destroying its aeration. Some growers rec-
ommend commercial vitamin B1 preparations for stimulating root growth of transplanted ferns, but this
treatment generally has no effect on intact plants. The leaves make enough of this vitamin, and it can be
 crown
old and new
soil level remove old
stipe bases crown

soil ball new soil level

spread roots old soil level

into new soil

spread roots
Figure 7.2. Transplanting an established plant: into new soil
soil ball
top, original planting; bottom, new planting.

Figure 7.3. Replanting a toppled plant: top, toppled

plant; bottom, replanted fern, with old stipe bases
removed and rootstock planted deeper, but without
covering the crown.
 top view
new pot

Place coarse gravel or

broken pieces of clay pots soil ball
over the drain hole

main growing tip

side view
main growing tip

Add fresh soil. Depending

on the size of the pot, the
soil ball should rest about
new pot
1–8 cm (0.5–3 in.) below
the top edge of the pot.
Loosen and spread out soil ball
dense or compacted roots.

side view

Center the plant. Add

more soil around the soil
ball and firm the soil in
place. Bring the soil level
to the level of the soil ball,
unless the roots are bent wire
exposed, in which case
raise the soil level slightly.
Remove old or broken
fronds, and water gently Figure 7.5. Potting ferns with
but thoroughly. creeping rhizomes, with the growing
tip placed as far from the edge of the
Figure 7.4. Repotting. pot as possible.
56 C 7

stored in the stem or passed to the roots. A temporary shade placed over recently transplanted plants will
help them recover and may reduce frond loss. Slug and snail control may also be in order. Within the first
few days after planting, some of the softer fronds might wilt, and if they do not recover should be removed.
New fronds will soon grow to replace them.

S C
Remove dead and damaged fronds. Rake leaves away from the crowns, particularly if slugs and snails
are a problem. Keep alert about trimming fast-growing plants that shade the slower growing ferns. Promptly
replace soil that has been washed away by rain or watering. Apply mulches if desired (see Chapter 5). Some
beds may need to be reworked every few years. Slower growth and packed soil indicate that replanting is
needed.

Planting in Pots
S  P
A well-selected pot will provide the best growing conditions possible and display the fern to its best
advantage. Some growers routinely repot in early spring to provide fresh soil and larger pots. If ferns are
growing well and not root-bound, they do not need to be repotted. A fern collection containing pots of
many shapes and sizes and variable soil mixes will need more attention during watering.
Pot size. In general, the diameter of the pot should be about one-third the height of the fern as measured
from the soil level. Repotting will generally require the next size up. For best appearance and growth, avoid
planting ferns in pots that are too big. Many ferns, especially the maidenhairs (Adiantum), grow poorly
when over-potted because the soil holds too much moisture in proportion to the active roots. The roots can-
not use up enough of the surrounding moisture to keep the soil well aerated.
Clay versus plastic pots. If proper adjustments are made, ferns do equally well in plastic or clay pots.
Remember that plastic or glazed pots retain more moisture than porous clay pots or wooden tubs. Thus,
water the plastic or glazed pots less or use a coarser, quicker-draining soil mix than you would with clay pots.
If the soil in a plastic pot has dried and pulled away from the sides, wet the soil thoroughly so that it
expands and the water will not trickle between the soil and the pot without penetrating to the roots. Unlike
clay pots, plastic ones are not as likely to support algae or become heavily encrusted with salt. Plastic pots also
have the advantage of weighing less, and holes for extra drainage are easily made with a hot ice pick or awl.
Pots without drainage holes. Beginners should use pots with drainage holes and avoid those without. If pots
without holes must be used, however, there are two ways to provide drainage. The first is to place one to two
inches of coarse material at the bottom of the pot before planting. Water the plant with distilled water just
enough to moisten (not wet) the soil—do not overwater. The second method is to find a clay or plastic pot
that has drainage holes and will fit inside the undrained pot. Plant the fern in the pot with the drainage hole
and place it within the undrained one. The fern can be removed as needed for watering, permitted to drain,
and then slipped back into the pot lacking the hole. Some gravel or drainage material may be placed at
the bottom of the hole-less pot to prevent the fern from standing in water that may have accumulated at the
bottom.
Pot shape. Most ferns do well in standard pot shapes, particularly the stout pots known in the trade as fern
or bulb pots. If drainage, aeration, and room for soil are adequate, the shape of the pot should not affect the
 P 57

growth of the plant. Also consider the aesthetic aspects of the pot. It should highlight the plant, not detract
from it. Bright colors and bold designs draw attention to the pot, not the plant.
When purchasing pots, look inside them to see that the drainage hole is large enough and is either
flush with the bottom or depressed to allow the water to drain freely. Pots with flat, broad bottoms tend to
drain slowly. Add extra gravel or other drainage material if more or larger holes cannot be made. Some nov-
elty containers have such a small capacity that frequent watering may be necessary to keep the plant from
wilting, and roots may fill the space too soon.

P   P

Providing drainage. Small pots 6.5 cm (2.5 in.) wide do not need anything to cover the drainage hole.
Larger pots 10 to 20 cm (4–8 in.) wide, however, need at least a piece of broken clay pot, gravel, or coarse
moss over the drainage hole. This lets the water drain and keeps the soil in the pot. Deep pots and those over
25 cm (10 in.) wide need a layer of drainage material at the bottom. Pot shards, gravel, or perlite can be used.
The amount of drainage material will depend on the pot size, frequency of watering, and soil mix. Hang-
ing pots that drain poorly in spite of sufficient drainage material can be induced to drain by inserting a cot-
ton wick through the drainage hole. Where slugs are a problem, a fine screen or plastic mesh placed over the
drainage hole before planting prevents slugs from entering the pot.
Cleanliness. Pots should be clean. Those previously used should be scrubbed thoroughly before reusing.
Basic planting procedures. After providing proper drainage, follow the “Basic Planting Procedures” dis-
cussed under “Planting in Beds” earlier in this chapter.

S C
Potted plants need to be watched for drying or excessive water.
Poor drainage. Never let a pot stand in water unless it contains a fern that grows naturally in swamps or
aquatic habitats. If the saucer below a pot accumulates water long after watering, it should be emptied. Pots
can be raised on pebbles above the water level to avoid becoming waterlogged. If your plant wilts without
standing in water, it probably needs repotting.
Poor drainage in small- to medium-sized pots can be detected by tapping the soil ball out of the pot a
day or two after watering. Examine the soil to see whether it is waterlogged. Replace the soil ball, and
examine it again the next day. If the soil is still soggy and the plant is not growing well, water it less fre-
quently or improve the drainage. Poor drainage can also be detected by the presence of soft decaying roots
and a strong odor of decaying vegetation (see “Watering,” in Chapter 4). In flower beds or large contain-
ers it may be necessary to check the drainage by digging a narrow hole into the soil to determine whether
it is waterlogged.
Root-bound ferns. A fern that is too big for its pot and becomes root-bound (filling the pot with roots) will
grow slowly, produce small fronds, and have leaves that rapidly turn yellow. Such plants need frequent
watering. If the rhizome branches or clumps, the center may be inactive or dead while the younger parts may
be growing over each other at the periphery of the pot. Such a plant can be kept growing for a time with
careful watering and fertilizing, but its vigor will gradually decline.
Root-bound ferns should be replanted or divided and replanted. If you want to increase the size of the
plant, cut off its dead parts and replant it into a next larger size pot. If you want to keep the fern in the same
size pot, it can be divided if it has many growing tips, and thin it to size by removing the smaller and weaker
rhizome branches and other dead parts. Keep several large rhizomes with their growing tips intact, and
58 C 7

replant in the same size pot. Root-bound tree ferns and ferns with unbranched rhizomes cannot be divided
and usually must be transplanted into larger pots. If there are many old and inactive roots, they can be
trimmed so that the root ball can be replanted in the same pot.
Salt crust on pots. A whitish salt crust on pots is unsightly and injurious to ferns. Unfortunately, it is hard
to dissolve. If possible, change the pot or soak and wash it. For soaking, add Physan (Consan) to the water
because it facilitates scrubbing. For badly encrusted pots, chip or scrape off as much of the crust as possi-
ble, then soak the pot and flush it with water. Vinegar dissolves some of the salt crust, but avoid getting vine-
gar on the soil or plant. If the pot is so heavily encrusted that it cannot be cleaned, then it should be dis-
carded. Remember that salt build-up on pots can be reduced by watering thoroughly (rather than
sprinkling), or by using rainwater, or by using plastic pots or cans instead of clay pots (see Chapter 4).

Planting in Baskets
Ferns that grow on trees are called epiphytes. They are not parasites but merely use the tree for support.
Because soil is absent from tree trunks and branches, epiphytic ferns secure nutrients and moisture by
sending their roots into the bark or the accumulated debris of leaves, mosses, and lichens. Epiphytic ferns
are adapted to rapid drainage and drier air than terrestrial ferns. Most ferns used in hanging containers are
epiphytes, although some terrestrial ferns also grow well in baskets. Ferns with drooping or cascading
fronds look particularly attractive in hanging baskets.

S  B

To simulate the native habitat of epiphytic ferns, plant them in open baskets that provide good drain-
age and have openings where creeping rhizomes can take root. Terrestrial ferns may be planted in open-
structured baskets but do better in more tightly woven ones that drain more slowly. Baskets may be made
of wire, wood, tree-fern trunks, ceramic, or plastic. Although clay or plastic pots can be used, they have to
be planted with a looser soil mix or watered less frequently. Also, ferns with creeping rhizomes might grow
into the pot and become trapped, or the rhizomes may grow over the edge of the pot where they are in dan-
ger of drying because there is no moist surface in which they can root.
Wire baskets. Commercially made wire baskets are inexpensive and readily available. There are, however,
drawbacks to their use. Their open structure requires a lot of lining material, and the wire rusts away in a
relatively short time, though commercial anti-rust compounds can be used to protect galvanized wire. Epi-
phytes with long-creeping rhizomes, especially those that tend to grow down the sides of a container, do best
in wire baskets that provide numerous places for the rhizomes to root. Never use copper wire for baskets
because it is toxic to plants.
Wood baskets. Wood baskets come in many designs and are generally less open and therefore require less
lining material. Redwood baskets are favored because they resist rot. Epiphytes that prefer moister condi-
tions and have short-creeping or clumped rhizomes grow best in wood baskets.
Tree-fern fiber baskets. Baskets made from tree-fern fibers are good for epiphytes because they drain well
yet retain sufficient moisture. They do not need lining material. Baskets made from Central American tree
ferns are more durable than those from the Hawaiian Islands. Because of environmental concerns, the
importing of tree fern products is restricted or prohibited by law.
Ceramic and plastic baskets. Ceramic baskets come in a variety of styles. The open-structured ones provide
excellent drainage and are well suited for ferns with long-creeping rhizomes. Keep in mind that cascading
 P 59

fronds will obscure the design of a basket. Plastic baskets are inexpensive and usually have large enough
meshes to allow thick rhizomes to emerge. Plastic baskets need less moss than wire baskets to line their inner
surfaces, and their meshes are just small enough that the moss does not fall out easily.

P  B

wire basket

soil ball

potting mix

moss lining

Figure 7.6. Planting basket ferns.

1. Line the basket with about 5 cm (2 in.) of moist, coarse or uncut moss (green moss or uncut
sphagnum moss), coarsely shredded tree-fern fiber (hapu), or coconut fiber. Shredded
redwood bark also works but is not commonly sold.
2. Add the potting mix to the basket and place the soil ball about 5 to 8 cm (2–3 in.) below
the rim of the basket (Figure 7.6).
If the fern is an epiphyte, plant it in the cavity with uncut or shredded sphagnum moss or
a very friable potting mix high in organic matter, such as coarse peat, or composted bark.
For terrestrial species, use ordinary potting mix in the cavity (see “Basic Planting
Procedures,” earlier in the chapter).
3. Press the soil firmly in place around the soil ball.
4. Cover the potting mix and the exposed part of the soil ball with about 2.5 cm (1 in.) of
lining material. This will prevent subsequent watering from dislodging the loose soil mix.
Extra pieces of rhizomes may be poked through the sides of the basket into the moss.
Rhizomes that will not stay in place can be anchored with bent wires.
5. Water the newly planted basket with a gentle but thorough spray or soak it for a few
minutes in a tub. Check for large leaks, which should be plugged up with additional moss.
 Remove the old basket.
Trim off dead rhizomes and old fronds.

Separate rhizomes into smaller clumps.

Select the healthiest clumps with the
most growing tips.

Trim off all the fronds from

the sides of the basket.
Loosen and spread out the
rhizomes.
Remove and discard old and
dead rhizomes.

Replant the clumps into a new basket

(see Figure 7.6)
Extra rhizomes of suitable length may
be thrust into the sides of the basket.
Water gently but thoroughly.

Replant into a new basket (see Figure 7.6).

Spread the loosened rhizomes into the
fresh potting mix and moss lining.
Water gently but thoroughly.

Figure 7.7. Replanting overgrown basket ferns.

 a

b c

Figure 7.8. Keeping basket ferns moist: a. plant a potted fern in a basket lined with moss; b. place a shallow clay saucer
over the moss lining before planting; c. place a piece of plastic sheet, with drainage holes punched into the plastic, over
the moss lining before planting.
62 C 7

R B F

It is time to replant a basket fern when the basket rusts or rots away or when the rhizomes begin to grow
over one another and become weak and thin (Figure 7.7). If the fern is growing out of the top of the bas-
ket only (not out of the sides or bottom), then it can be merely freed from the basket, cleaned of dead tis-
sue and old soil, and planted into a new basket as described above.
If the fern is growing out of the top, sides, and bottom of the basket, replanting is a bit more involved,
but it can be accomplished in several ways. One method is to cut away the basket and remove the rhi-
zomes in big pieces. The biggest and healthiest pieces can be replanted in a new basket as previously
described. Alternatively, you can remove the fronds from the sides and bottom of the basket, lift out the
plant, and drop it into a new basket lined with fresh moss. Any intervening spaces should be filled with loose
planting mix or moss. In time, new rhizomes will grow through the moss layer. Another way to revitalize
an old basket fern covered with rhizomes is to cut out the center of the basket from the top, without dis-
turbing the growth at the sides and bottom. Fill the hole with fresh potting mix and then cover with uncut
sphagnum or green moss. If the outer part of the wire basket has rusted away, mend it with wire or fashion
a wire sling to support the basket. (Remember: do not use copper wire.)
Some growers prefer to build a new basket around the old one. This requires suitable wire, much
patience, and care to minimize the damage to fronds and rhizomes on the sides and bottom of the basket.
Cut galvanized wire of suitable strength to form the rings of the new basket. Join each of the rings around
the old basket. To hold the rings in place, tie them with cross-wires or solder on cross-wires. Moss is then
stuffed into the space between the old and new baskets.

S C
If properly planted, a basket fern will last a few years. If it is an epiphyte, the fern must have good
drainage. Terrestrial ferns planted in baskets have a tendency to dry too quickly, and their fronds are sus-
ceptible to dryness, particularly if hung indoors. These plants should be substituted with more adaptable
plants or rotated with fresh ones. If you keep them in pots, they can be lifted out of the moss-lined baskets
and easily replaced (Figure 7.8). In arid climates, hanging baskets tend to dry quickly and can be difficult
to wet again without soaking. To slow the drying, place a small sheet of plastic over the moss lining at the
time of planting. A shallow clay saucer can also be used, but holes for better drainage are punched more eas-
ily into the plastic. A flowerpot may be sunk at the top of the basket, to one side of the fern, to act as a catch
basin for water. The rim of the basket can be built up to act as a catch basin as well. Keep alert for weak or
rusting wires and replace them immediately. As moss or lining material decays or falls away, replace it by
stuffing fresh lining material into the cavities.
 8 Propagation

Ferns can be propagated two basic ways: from pieces of the parent plant or from spores. The first involves
vegetative reproduction and gives rise to offspring genetically identical to the parent. The second method
involves sexual reproduction and gives rise to offspring genetically different from the parent and among
themselves. Before learning to propagate ferns by spores, you need to understand the fern life cycle.

Typical Life Cycle

Fern spores grow into new ferns indirectly (Figure 8.1). If a spore lands on a moist place with sufficient light,
it germinates. Several days or weeks later, it sends out a slender cell (a rhizoid) that anchors the spore in
place. It then produces a green cell that divides repeatedly in one direction to form a thread of cells. Even-
tually the cells at the tip of the thread divide laterally to widen the structure, and in about three months the
thread grows into a flat, heart-shaped piece of tissue about 5 mm (0.2 in.) wide. This small plant is called
the gametophyte or prothallus (plural, prothalli). On the underside of this tiny plant are male sex organs
(antheridia) containing sperm and the female sex organs (archegonia) each containing an egg. Mixed among
these are numerous rhizoids that secure the gametophyte to its substrate. When enough water is present in
the environment, the antheridia burst and release the sperm. The sperm swim to an egg and fertilizes it. The
egg can be on the same or a different prothallus. A film of water must be present in the environment to allow
the sperm to swim to the egg, which explains why ferns are most abundant in wet habitats.
The union of a sperm and egg is known as fertilization. It marks the start of the sporophyte—the
familiar fern that eventually bears the spores. The fertilized egg (zygote) divides and forms an embryo or
young sporophyte that develops roots, stems, and leaves. As the roots grow and anchor the sporophyte in
the soil, the prothallus withers, leaving the sporophyte on its own. If fertilization does not occur, the pro-
thallus continues to live, sometimes for many years.
The first series of leaves produced by the young sporophyte do not resemble those of the adult plant.
The successive leaves that are produced gradually assume the typical adult leaf form (Figure 8.2). About five
months to a year after the spores have germinated, a small but well-formed fern is developed. Some ferns
grow faster and can produce spore-bearing fronds in six or seven months. The fastest growing is the water
fern (Ceratopteris), which can grow from spore to mature spore-bearing fronds in only one month. Most
ferns, however, take two years or more to complete their life cycle. Normally once the sporophyte is estab-
lished, the spore-bearing fronds are produced year after year.

63
64 C 8

e
b c d

a i

k j
p

l
n
o
m

Figure 8.1. Fern life cycle: a. familiar fern plant or sporophyte; b. pinna bearing sori; c. sorus; d. indusium; e.
sporangia; f. spore; g. spore germinating; h. young prothallus or gametophyte; i. underside of mature prothallus or
gametophyte; j. rhizoids; k. antheridium; l. sperm; m. archegonium; n. egg; o. embryo; p. young sporophyte.

Variations of the Life Cycle

Ferns reproduce by other means as well. They can multiply directly from parts of the sporophyte such
as the root, rhizome, or frond. Taking branches from the rhizome is a common way to propagate ferns. On
some ferns, clumps of tissue known as buds or bulblets form and grow into new plants. Depending on the
species, these buds develop on the roots, blades, veins, or rachises (Figure 8.3). Less commonly, pieces of
the stipe base may be induced to form buds, such as in Cystopteris, Dryopteris carthusiana, Dryopteris filix-
mas, Matteuccia struthiopteris, Phyllitis scolopendrium, Pteridium aquilinum. The stipules at the stipe bases
of marattiaceous ferns (Angiopteris and Marattia) also produce buds if properly planted. Fronds from young
plants of some ferns can also be induced to form buds. In rare cases, prothalli grow directly from fronds of
young sporophytes (Osmunda regalis var. regalis ‘Cristata’, Polystichum setiferum, and others). This is known
as apospory, which means “without spores” (Figure 8.4).
Some ferns (especially those of dry habitats) produce spores but reproduce asexually by proliferating a
new plant directly from the prothallus instead of through the union of egg and sperm (Figure 8.5). This
process, called apogamy (without sex cells), produces ferns faster than the sexual method. Apogamous
ferns, such as many cheilanthoids, can produce sporophytes in six weeks after sowing (under 18 hours of
Figure 8.2. Age variation in fronds, showing fronds produced from a progressively (left to right) older plant of
Doryopteris nobilis.

Figure 8.3. Proliferous buds: a. root bud and a young plant established from Asplenium auritum roots; b. leaf bud on
the underside of a frond tip of Woodwardia radicans.
66 C 8

Figure 8.4. Apospory: prothallus developing from the tip of a pinnule in

Polystichum setiferum.

artificial light). Pteris cretica produced sporophytes in two and a half months after sowing without artificial
light. In a sowing of spores, the sporophytes of apogamous prothalli tend to appear at the same time,
whereas the sporophytes of sexual prothalli appear at different times. Ferns that develop apogamously also
tend to produce leaves before the roots, whereas ferns going through the sexual life cycle tend to produce
roots before the leaves. The leaves appear just below the notch of the prothallus. Ferns that reproduce
entirely by apogamy (those in which no sexual cycle is known) include such familiar cultivated species as
Cyrtomium falcatum, C. fortunei, Dryopteris cycadina, and Pteris cretica. Some ferns reproduce by apogamous
and sexual means.
A single prothallus can produce more prothalli. New ones may form along the margins of the old ones
(Figure 8.6a). If conditions are favorable and if fertilization does not occur, growth continues for years. If

c
first leaf

prothallus b
first leaf

first leaf
stem tip second leaf
stem tip
root
prothallus root

prothallus

Figure 8.5. Various stages of a fern developing apogamously: a. the first leaf emerging from the prothallus; b. stem tip
and root formation; c. the first leaf and roots expanding, as the second leaf emerges. The new fern is formed directly
from the surface of the prothallus, not from within an archegonium as in sexually reproducing ferns.
 P 67

fertilization does occur, the prothallus dies as the sporophyte develops. In filmy ferns, shoestring ferns,
grammitids, and some polypodiums, new prothalli are produced from spindle-shaped or rod-shaped struc-
tures (gemmae) borne on short stalks along the margin of the prothalli (Figure 8.6b).

Figure 8.6. Prothallus proliferation: a. an old prothallus forming eight new prothalli and one on the tip of a tapered
process, as shown on Platycerium quadridichotomum; b. gemmae borne on stalks growing from the edge of the
prothallus in Trichomanes, each gemma grows into a new prothallus.

Vegetative Propagation
Plant parts that reproduce new plants are the rhizome, root, and frond. The branching rhizome is
requently used in vegetative propagation. Ferns without branching rhizomes or buds generally cannot be
vegetatively propagated and must be grown from spore (this is true for most tree ferns). Success in vegeta-
tive propagation is greatest when moist soil and sufficient humidity are maintained with as little watering
as possible.

D
To divide a rhizome, look for the growing tips where new fronds arise. Before making the division, you
can stimulate new side buds to form on a thick rhizome by partly cutting through it and waiting for the side
buds to develop. Before you remove a tip or a clump of tips for dividing, be certain that enough growing
tips are left on the parent plant to keep it alive. Look for a joint or naturally weak spot to make the cut from
the parent plant (Figure 8.7a). The larger the rhizome or clump taken, the better the chances of a success-
ful division. Use a clean, sharp knife to make the cut. With a trowel, dig up the division. Avoid injuring the
growing tips, and keep as much of the soil around the roots of the division as possible. Cut off old or bro-
ken fronds and remove parts of the larger fronds to reduce water loss, particularly if the air is dry. Cut the
bladeless and withered stipes as close to the rhizome as possible and remove any old or broken roots.
Before planting the division, dust its cut ends with a fungicide such as Captan. Select a rooting medium
that is well-drained, such as a mixture of one part peat moss and one part perlite. Epiphytes often root well
68 C 8

b c

Figure 8.7. Dividing and planting rhizome divisions: a. dividing the rhizome (roots not shown) (Polypodium); b.
planting the rhizome at half its thickness (Davallia); c. planting the rhizome diagonally (Davallia).

in moist uncut sphagnum moss. Avoid using soil, manure, compost, or other substances that harbor a lot
of bacteria and fungi. Provide the box, flat, or pot with good drainage.
Replant the division at its former soil level. If its roots were previously exposed, set the division slightly
deeper so that the roots are covered. Firm the medium in place and water well. Keep the plant lightly
shaded for a few days.
Pieces of creeping rhizomes without roots should be secured to the rooting medium to keep them from
drying out. Plant these at half their thickness into the medium (Figure 8.7b); bent wire can be used to
hold them in place. If the rhizome pieces are a few inches long, thrust their cut ends diagonally into the soil
to about one-third their length, leaving the growing tip exposed (Figure 8.7c).
After planting, the divisions still need to be cared for. Keep them in a humid, shaded, warm place until
rooted enough to be transplanted. Guard against fungus gnats and other pests that feed on exposed tissue
or new roots. Above all, avoid overwatering that might foster mold or decay. If this develops, increase ven-
 P 69

tilation, stop overhead watering, and apply a fungicide drench or Physan. Apply inorganic fertilizer two to
three weeks after dividing.
Plants can be easily propagated by air-layering if they have rhizomes that are long and tend to grow away
from basket or soil. Wrap damp uncut sphagnum moss around the rhizome of the mother plant. Tie the
moss into place with string or strips of plastic. The moss may also be wrapped with a small sheet of plastic
before tying. When roots have developed in the moss, the rhizome can be separated from the mother plant
and planted.

B
Some ferns produce new plantlets from buds on their fronds or, more rarely, roots. Bud-producing
fronds can be anchored to the soil while still attached to the mother plant, or they may be detached and
planted (Figure 8.8). The benefit of leaving the bud on the mother plant is that it will have an adequate sup-
ply of water and food while taking root. The bud can be anchored to the soil with a bit of soil or clean sand.
If detached from the mother plant, bud-bearing fronds may be planted whole or in pieces. These are
anchored to the soil surface with a planting mix or clean sand, watered, and then covered to provide adequate
humidity. Once the buds have rooted, cut the mother frond to separate the plantlets. Some species root more
readily if the bud is left attached to the mother plant (this is particularly true for Adiantum caudatum).
A detached bud should be planted at half its thickness in a rooting medium. Water gently but thor-
oughly and then cover with a piece of glass or plastic. Keep the air humid, but avoid excess humidity and
overwatering.
Detached buds may also be grown conveniently in clear plastic shoeboxes or refrigerator boxes, prefer-
ably with holes for drainage. Line the bottom of the box with an inch of perlite. Place the planting medium
(usually half sand and half peat) over the perlite, then plant the buds. The lid might need to be opened to
control excessively high humidity.
Plants that can be propagated from detached buds include Asplenium bulbiferum, A. daucifolium, Poly-
stichum setiferum Divisilobum Group, Tectaria gemmifera, Woodwardia orientalis, and W. radicans. For
these species, apply a weak solution of inorganic fertilizer every two to three weeks, and gradually remove
the covering over the planting as the buds grow. Transplant the larger individuals to avoid overcrowding.
If mold develops, increase the air circulation and use a fungicide drench.
Root buds develop on ferns such as Ophioglossum, some Platycerium, Diplazium esculentum, and Asple-
nium auritum. They are usually unnoticed until the new plantlets emerge from the soil. When these plants
are several inches tall they can be separated from the mother plant and transplanted.

P   S

The leaf stalk or stipe can be used to propagate the hart’s-tongue fern (Phyllitis scolopendrium), lemon-
scented fern (Thelypteris limbosperma), lady fern (Athyrium filix-femina), and soft shield fern (Polystichum
setiferum). Probably many other ferns can be propagated by this method, but they have yet to be tried.
The stipes of the hart’s-tongue can be induced to form buds by sowing small green, plump, healthy
pieces over a sterilized planting mix and covered to raise the humidity. Another method is to select older
leaves with stipes that are still plump and green and snap them off near the rhizome. Wash the frond and
cut the stipe into 2.5 cm (1 in.) lengths. (The base of the frond with stipe attached may also be used.) The
pieces are planted upside down with the stipe about 3–6 mm (0.1–0.2 in.) above the soil. To prevent dry-
ing, cover the container with plastic, and mist as necessary. The appearance of small green blisters on the
stipe indicates bud formation. When the buds develop into plants about 1 cm (0.5 in.) tall, they can be care-
fully removed and treated as young ferns. The development of a bud into a plantlet takes about three
 b

Figure 8.8. Planting buds: a. layering or rooting a bud still attached to the mother plant (Adiantum caudatum); b.
rooting a bud attached to a section of a frond, top and side view (Polystichum setiferum, a proliferous variant); c.
rooting detached buds in a pot (Woodwardia orientalis); d. rooting detached buds in a plastic container (Tectaria
gemmifera).
 P 71

months. For the lady fern, a portion of the rhizome tissue should be left on the stipe. To prevent mold or
rot, treat the stipe pieces with Physan 20 or a similar fungicide. For more details see Dyce (1985) and
Rickard (1986).

P   S

Stipules are the somewhat fleshy, flap-like outgrowths that clasp each side of the stipe base in Angiopteris
and Marattia. A stipule may be cut and planted, usually upright, one-half to two-thirds its length in per-
lite or other clean, well-drained medium. Young plants will develop from the edges in a few months. Once
its roots develop, the young plant may be separated from the stipule and planted. If the stipule is still firm,
it can be induced to form additional plantlets. Maintaining humidity and controlling mold is the same as
that explained under “Propagation from the Stipe.”

T C
Tissue culture is the process of growing entire plants from bits of tissue by planting them in a growth
medium of nutrients and hormones. Aseptic conditions must be maintained throughout the procedure,
which involves transferring the tissue from one formulation to another at certain stages. Most Boston ferns
and many other species are commercially propagated by tissue culture. The advantage of growing ferns by
tissue culture is that large numbers of genetically uniform, disease-free plants can be produced in a small
area. Ferns for which there is limited vegetative propagating material, or those that do not produce viable
spores, are especially suitable for tissue culture.
Some ferns propagated by tissue culture are more attractive than those grown from spores. They tend
to be fuller and more compact due to greater branching of the rhizome. Shorter production time is not
always achieved through tissue culture—it depends on the species and growing conditions. It may take as
long to grow maidenhair (Adiantum) from tissue culture as it does from spores; however, staghorn ferns
(Platycerium) usually take less time. Ferns that have been divided and replanted in tissue cultures for three
or more generations tend to produce many mutations, most of which are misshapen and unattractive. The
more troublesome problems in tissue culture include obtaining uncontaminated tissue (explants) and get-
ting young plants acclimated to greenhouse conditions.
Tissue culture involves three basic stages. In the first or establishment phase, the explants are disinfec-
ted and planted in a growth-inducing medium. Fern explants are usually taken from leaves or young stems
(or stolons in Boston ferns). In the second stage, called the multiplication phase, the explants are planted
on a medium that induces shoot proliferations. Growth at this stage can be subcultured to increase the num-
ber of plantlets. In the final stage, or pre-transplant stage, the clumps of shoots are divided and transferred
to a medium that stimulates further development, particularly of the roots. Growth at this stage can also be
subcultured to increase the number of plantlets. After the plantlets have developed roots, stems, and leaves,
they are transplanted into a soil mix and placed in a greenhouse. The tubes or bottles containing the vari-
ous stages of growth are usually kept at 27°C (81°F) and given 16 hours of artificial light at 300 foot-can-
dles, although some growers use lower temperatures (to 16°C or 60°F) and lower light (100 foot-candles).
The main ingredients in the various culture media are water, inorganic nutrients, sugar, vitamins, and
plant hormones called auxins (which stimulate root growth) and cytokinins (which stimulate shoot growth).
Ferns grow best on a solid rather than liquid medium, which is why agar or agar substitutes are also added.
The recipes for growth media vary from stage to stage and from one species to another. Most ferns grow well
on only two formulations, with the same formulation used for the first and second stages. Murashige’s for-
mulations for in vitro propagation are satisfactory for a wide variety of plants and can be bought premixed.
For the beginner at tissue culture, it is easiest to buy a tissue-culture kit from a scientific or biological sup-
72 C 8

ply house. Occasionally, plant societies, colleges, and universities offer workshops or classes on tissue cul-
ture, and much literature is available for all levels of interest.

Spore Propagation
Although not difficult, growing ferns from spores takes time and patience, but the fascination and
pleasure derived from this activity is a reward in itself. The space and equipment needed are minimal.
Most ferns grown commercially from spore take about 6 to 10 months until they are large enough to be
planted into 5-cm (2-in.) pots. Other ferns take longer, such as filmy ferns, which take a few years to
develop sporelings. For raising fern allies from spores, see “Fern Allies” in Chapter 10. To grow gameto-
phytes for experimental purposes, see Dyer (1979).

C S
To collect spores you need to find the sporangia (spore cases). These are usually grouped in clusters,
called sori, on the underside of the frond. The sori are mostly 2 to 5 mm (about 0.1–0.2 in.) wide and may
be round, oblong, or linear. Less frequently, the sporangia are scattered over the entire lower surface of the
frond or borne on modified portions of the leaf.
When collecting spores, it is important to recognize whether the sporangia are ripe or have already
shed their spores. Immature sporangia usually appear whitish green. As they mature, they turn light brown
and finally medium or dark shiny brown, or in some species blackish (Figure 8.9). (In some ferns, such as
Polypodium, the mature sporangia are yellowish due to the color of the spores within.) The sporangia appear
dull brown and frayed after the spores are shed. Another clue to the ripeness of spores is the appearance of
the indusium. Green or whitish indusia indicate unripe spores; light brown, yellow, or nearly black indu-
sia indicate mature spores. Shed or shriveled indusia usually indicate that the spores have been shed.
Ripeness of the sporangia may be more accurately determined with a 10× hand lens (or jeweler’s loop).
Plump sporangia without a cracked wall contain spores, whereas sporangia frayed or with cracks or slits have
shed their spores. Shriveled sporangia will not produce viable spores. Pick fronds when most of the spo-
rangia are still plump and shiny.
To gather spores for sowing, place the picked frond in an envelope or over a clean piece of smooth paper
with the sporangia side down. Cover the frond to prevent the shed spores from blowing away or other
spores from contaminating your collection. Usually the spores will be shed on the paper within minutes,
but wait a day or two if you want to collect the maximum number of spores. They will resemble fine dust.
Some fronds produce large quantities of spores whereas others do not. The spores are often brownish but
may also be green, yellow, or black, depending on the species. Also shed are whole or fragmented sporan-
gia, bits of hair, scales, or other tissue. Tree ferns of the Cyathea group tend to shed whole sporangia that
later open to shed the spores. Examine the paper with a hand lens to be certain that spores are present. Store
the spores in an envelope or folded paper packet. For reference, write the name of the species, the source,
and the date on the outside of the packet.
Contamination by other fern spores can be a problem during collection. To reduce the risk of con-
tamination, pick only clean fronds, or if these are not available, brush or rinse away dirt and debris. Col-
lecting the spores on a cool or dim day reduces spore loss and contamination from other spores.
Another way to collect spores is to place leaflets bearing ripe sporangia in a plastic bag to keep them from
drying in transit. Then follow the directions given later in this chapter under “Sterilization of spores within
sporangia.”
 P 73

Figure 8.9. Determining when to collect spores

(Thelypteris puberula): left, too young; middle,
ripe; right, too old.

Spore envelopes. The best way to collect and store spores is in folded paper packets (Figure 8.10). These
are easy to make and are leak-proof. The problem with using ordinary envelopes is that the spores tend to
leak out, and if the leaks are covered with cellophane tape, many spores can be lost because they stick to the
tape. Plastic bags seem like an obvious alternative, but they have the disadvantage of the spores sticking to
the bag and being hard to remove. Fertile leaf material dries more quickly in paper packets, averting mold
problems.

2 3

4 5 6

Figure 8.10. Making a spore envelope: 1. Fold thin paper in half lengthwise (along the dotted line). 2. Double fold the
upper open edges. 3. The folded and sealed upper edge. 4. Turn the envelope over (keep folded edge on top) and fold
diagonally along the dotted lines. 5. Tuck the folded right corner under the band as shown. 6. To seal the envelope,
tuck the left corner under the band. Label with name, date, and source. Leaflets or spores are best enclosed at step 1
or 5.

C S
Removal of nonspore material (sporangia, scales, hairs) reduces contamination by algae, bacteria, fungi,
and mosses. The larger nonspore material can be picked out with forceps or brushed away with a fine
brush. If pure or nearly pure spores are desired, separate the nonspore material by slightly tilting the paper
74 C 8

upon which spores were collected and gently tapping it. The spores are left behind as the fluffier nonspore
material tumbles ahead and can be brushed away. The spores can then be stored until use.
Small bottles with a very fine (about 0.05 mm) screened mesh in place of the top can be used to sepa-
rate spore from nonspore material and to evenly sow the spores. The finest tea screen available, lens paper,
and even cheesecloth have been used as satisfactory screening material. Check the material with a hand lens
to be sure that the separation is adequate.

S S
Most hobbyists and commercial growers do not sterilize (disinfect) spores before sowing. Scientific
work, however, requires it. Most methods use bleach diluted with water and a wetting agent such as Tween
20, Aerosol OT, Alconox, or a mild household detergent, which will prevent the spores from sticking
together. The following methods of disinfecting spores are representative of the many kinds used. For
more details, see Ford and Fay (1990).
Sterilization of spores freed from the sporangia. Clean the spores to remove the larger nonspore material.
Place the spores in a 5 to 10% bleach solution and add a fraction of a drop of wetting agent (1 part Tween
20 to 2000 parts bleach solution). Soak the spores for one minute. Collect the spores on filter paper. Rinse
two to four times with sterile water. Sow from the water solution or dry and store until needed. Some spe-
cies are sensitive to bleach and may require a weaker solution and less soaking time. For such species, reduce
the concentration of bleach solution to 2% and soak for about 5 to 10 seconds. Researchers report that soak-
ing green spores in 2% bleach solution for two to four minutes gives satisfactory results.
Sterilization of spores within sporangia. Select leaflets or fronds with ripe but unopened sporangia. Soak
in a 5 to 10% (rarely 20%) bleach solution to which a wetting agent has been added as above. Remove all
air bubbles with a small brush, and keep the leaflets immersed 5 to 10 seconds. Do not rinse. Place in a clean
envelope. (Envelopes can be disinfected by placing them in a box and baking in an oven at 160°C [320°F]
for about two hours. Never heat the spores.) The envelopes containing the disinfected leaflets can be placed
between layers of clean paper to absorb the moisture. With regular changing of the absorbing paper, the
spores should be shed and ready for use or storage in about three days.

V  S
The fresher the spores, the greater and faster the germination. Depending on the species, fern spores can
remain viable from 2 days to 130 years. Storing nongreen spores under cool conditions, including refrig-
eration, prolongs their viability. Variation exists in the percent germination from spores taken at different
times from the same plant and from different plants of the same species. Spores collected from vigorously
growing plants usually have the highest germination rate. The spores of some species require dormancy
before germination. The dormancy can range from two weeks to one year. For more information, see Rush
(1984a, b).
Green spores, such as those in Osmunda, Todea, Equisetum, and Grammitis, generally have the short-
est viability, averaging about two months (the shortest recorded is two days, and the longest about one year).
Because of their short viability, green spores should be sown soon after collecting. For Osmunda (and prob-
ably other genera with green spores), viability can be prolonged by refrigeration at 4°C (40°F). Osmunda
spores stored in a freezer at −20°C (−4°F) for 2 to 3 years retained their viability (Rush 1984b).
Nongreen spores remain viable longer than green ones, generally from 1 to 48 years, with the extremes
being a few weeks (Cyathea) to 130 years (Marsilea). See Lloyd and Klekowski (1970) for a discussion of via-
bility and germination times in green versus nongreen spores.
 P 75

S M
There are many ways to grow ferns from spores, but only the most common methods are discussed here
(Figure 8.11). Before sowing it is helpful to sterilize or treat the soil. This discourages algae, mosses, bacte-
ria, fungi, and other pests that can crowd or attack the spores and prothalli (see “Sterilizing Soils” in Chap-
ter 5). Because fern spores are easily airborne and may contaminate other sowings, avoid successive sowings
of different species on the same day. Several methods have been developed to control the amount of spores
sown, such as mixing the spores with inert, dry material, spraying them through an atomizer, or delivering
aliquot amounts through a syringe (Basile 1973). For sowing small amounts, a cotton swab dipped in the
spore and tapped over the container gives better control than merely tapping the spores out from an enve-
lope. For more details, see Basile (1973).
Growing a few plants. The following methods are suitable for a hobbyist who wants to raise only a few
plants. Milled sphagnum moss is good for beginners, but any of the soil mixes listed under the commercial

glass
soil level, sow here
clear or
a translucent
lid

nutrient solution
glass level, sow here
d
soil level, sow here

clay pot
lid
agar level,
saucer sow here
e
b

clear or translucent lid

inverted sow on surfaces of an
soil level, glass jar inverted clay pot
sow here
perlite sphagnum
moss water level
c
saucer

Figure 8.11. Common methods of planting spores: a. in a flat; b. in a pot; c. in a plastic container; d. on a nutrient
solution; e. on an agar plate; f. on an inverted pot.
76 C 8

method can be used as a planting medium. For small sowings, adequate space is any small, clear container
that is wide enough to allow fingers or forceps to easily remove the prothalli or sporelings and is deep
enough to hold 5 cm (2 in.) of soil and provide 2.5 to 5 cm (1–2 in.) of overhead space. The container will
need a cover, and transparent lids are best, but plastic wrap or glass can also be used. If the soil is disinfec-
ted by pouring boiling water through it, drain holes will be necessary in the container. Allow enough over-
head space for growth; avoid filling pots too close to the top. If overhead space for the sporelings becomes
inadequate, an inverted clear-plastic cup placed over the pot will provide more room and retain humidity.
Containers without drain holes may be layered on the bottom with 1.5 to 2.5 cm (0.5–1 in.) of perlite
(Sponge Rok) to drain excess water from the soil above. Because additional watering after sowing is infre-
quently needed, many growers use cups or containers without drain holes or omit the layer of perlite. The
container should have about a 5 cm (2 in.) layer of planting medium.
If microwaveable planting containers are used, they can be disinfected along with the soil mix in a
microwave oven. Before microwaving, generously moisten the soil, gently firm it in place, loosely cover with
a lid or wax paper, and cook for about five minutes on high. Upon removal, the soil mix should be steamed
through and moist. Remoisten with boiled water if necessary. Keep these disinfected containers covered
until sowing time. With plastic containers that distort in heat, the soil will need to be disinfected previously
in heat-resistant containers and then spooned into the plastic container when cool. The containers can be
previously disinfected with a 10% solution of bleach or Physan 20 and rinsed with boiled water.
A simpler procedure, but one with greater risk of contamination, is to lightly pasteurize a soil mix in a
clean, undrained plastic cup. To do this, add just enough boiling water to moisten the soil for planting (plas-
tic cups generally do not distort with only this amount of boiling water). A novel use of the pressed-peat pots
(Jiffy-pot pellets, with mesh removed if present) is to place a pellet in a small (180–240 ml or 6–8 oz.) plas-
tic tumbler and moisten with about 120 ml (4 oz.) of boiling water. When the peat has expanded, loosen
it, gently firm it down, cover the tumbler, and let cool. Sow the spores thinly to avoid spindly, tangled pro-
thalli, and minimize crowding. A cotton swab dipped in the spores and gently tapped over the medium
helps to control the amount of spores sown. Although dense sowings can be thinned, the thinning process
tends to disrupt and damage growth. Therefore thin sowings are best. After sowing, the containers are cov-
ered with clear lids or glass or plastic wrap held in place with a rubber band.

Commercial method. The spore-planting media vary from grower to grower. Most growers use a mixture
of one to two parts peat moss and one part fine sand. Some growers add loamy soil, fine perlite, or other
additives to this mix. Commercial planting mixes formulated for growing seeds can also be used for spores.
Finely milled sphagnum moss used alone provides good results but might not be suitable for all ferns.
Whatever soil medium is used, it should be able to pass through a 3 mm (0.1 in.) mesh. Fine texture is
important because it ensures good contact with the prothalli and facilitates transplanting. Some growers use
a fine-textured mix on top of a coarser mix to promote drainage.
Glass or plastic containers may be used as previously discussed. New wood flats and larger clay pots with
drainage holes are traditionally used because they can be sterilized or disinfected easily.
Commercial growers have devised several ways of sowing spores as evenly as possible over a broad sur-
face. Salt shakers and spray bottles (atomizers) with spores suspended in water have been used. Depending
on the species and the transplanting procedures, some growers sow spores densely so that the growth soon
after germination is a solid green mat, which will need early separation. By sowing a small amount of spores
into each plug (a depression in a special plastic growing tray; see “Growing in Plugs” later in this chapter),
growers save labor in transplanting. After sowing, the containers are covered with clean plate glass. Plate
glass is better than plastic wrap, which will sag when stretched over a wide container and cause condensa-
tion to collect and drip on one spot. The resulting wet spots promote the growth of fungi, bacteria, and
algae.
 P 77

Nutrient solution method. Many different types of nutrient solutions are used commercially and are of
interest to amateurs. Some growers use distilled or tap water or dilute solutions of balanced inorganic fer-
tilizers (at one-quarter strength), though the more common nutrient solutions are not diluted (Table 8.1).
Other solutions include Beyerincks (Brown 1920) and Knudson C without the sugar and agar (Knudson
1946). The measures given in Table 8.1 are given in the metric and English systems to avoid the need for
special measuring equipment. However, since fractions of ounces or tablespoons are difficult to measure,
the formulation makes 95 liters (25 gal.) of solution—the surplus can be used as a liquid fertilizer. If smaller
amounts are desired, the measurements given in grams make one liter or approximately one quart of solu-
tion. The ingredients are inexpensive and can be obtained from local chemical supply houses, firms selling
fertilizers, and drugstores. Prepare the solution by adding the salts to the water in the order given. Impuri-
ties in the chemicals and from the water (tap water) will probably provide the trace elements needed. If you
want to add trace elements, however, use the formulation in Table 8.1. If the solution is to be stored for a
long time, add the iron or trace elements just before use.
These solutions will have a pH of about 6 if distilled water is used. A pH of 6 is optimum for germi-
nating most fern spores. In general, the pH range for germinating spores is 5–7. A more acidic solution can
be obtained by omitting calcium compounds. Some ferns of limestone habitats have spores that can ger-
minate at pH 7–9.
The nutrient solution can be sterilized by placing it in a pressure cooker for 15 minutes at 1 kilogram
per square centimeter (15 lbs. per sq. in.) or by boiling it for a few minutes. The solution is poured into pre-
viously sterilized dishes or plastic containers (which may be sterilized by boiling or washing with a 10%
bleach solution and rinsed). The depth of the solution may vary, but 6 mm (0.2 in.) is enough for germi-
nation. Cover the solution and allow it to cool. Then sow the spores and replace the cover. To keep the
spores floating on the solution, avoid moving the dish after sowing. Tilt the lid slightly to prevent con-
densation from dripping on the spores.
Nutrient agar solution. This method provides a solid nutrient medium for growing spores. It has no advan-
tage over other methods, and ferns are reported to grow slower by this method. It is used mainly for scien-
tific research or when the spore material is extremely valuable and sterile techniques are necessary to ensure
survival. Agar, the solidifying substance, can be obtained from scientific supply companies. Less expensive
substitutes are now available.
The medium is made by adding 14 g (approximately ½ oz.) of agar to 1 liter (approximately 1 quart)
of any suitable nutrient solution. (Seven drops of a general-purpose, water-soluble fertilizer to one liter of
water makes a convenient nutrient solution.) Dissolve the agar by heating it in the nutrient solution, stir-
ring occasionally. Then pour the dissolved solution 6 to 9 mm (0.2–0.4 in.) deep into shallow dishes, cover
loosely, and sterilize immediately in a pressure cooker for 15 minutes at 1 kilogram per square centimeter
(15 lbs. per sq. in.) pressure. After the dishes have been removed from the pressure cooker and the agar has
cooled and solidified, they are ready to be planted, preferably with disinfected spores. Before and after
planting, keep the dishes covered to avoid airborne contamination and moisture loss. Ordinarily the agar
will remain moist enough to support growth into the mature prothalli stage. Vaseline petroleum jelly can
be used to seal the edges to prevent drying. If the agar should dry or crack, transfer the growth to a fresh
medium as soon as possible. A nutrient solution originally used to grow orchid seeds and containing sugar
(sucrose) has been reported as good for growing fern spores. It works best at pH 5 (Knudson 1946).
Inverted pot method. The inverted pot method is ideal for exhibiting the growth of prothalli and young
ferns. It is inefficient for growing ferns commercially because scraping prothalli off the pot is tedious and
awkward.
Fill a small, clean, porous clay flower pot with uncut sphagnum or peat moss, then invert it into a
saucer or shallow dish. Sterilize the unit with steam or by pouring boiling water over it, or place the whole
Table 8.1. Nutrient Solutions

Grams per Ounces per Tablespoons

1 liter of 25 gallons per 25 gallons
Salt Formula Grade of salt water of water of water

B’ S
Ammonium nitrate NH4NO3 Technical 0.5 2 4
Potassium phosphate KH2PO4 Technical 0.2 ¾ 1½
Magnesium sulfate MgSO4 · 7H2O Technical 0.2 ¾ 2
Calcium chloride CaCl2 Technical 0.1 ½ 1
Ferric chloride FeCl3 Technical trace trace trace

K’ C S (after Knudson 1946)

Calcium nitrate Ca(NO3)2 · 4H2O Fertilizer 1.0 4 8
Ammonium sulfate (NH4)2SO4 Fertilizer 0.5 2 4
Magnesium sulfate MgSO4 · 7H2O Technical 0.25 1 2
Ferric sulfate FeSO4 · 7H2O Technical 0.025 1⁄ 10 1⁄ 5

Manganese sulfate MnSO4 · 4H2O Technical 0.0075 1⁄ 30 1⁄ 15

Potassium phosphate KH2PO4 Technical 0.25 1 2

K’ S
Potassium phosphate KH2PO4 Technical 0.2 ¾ 1½
Potassium nitrate KNO3 Fertilizer 0.2 ¾ 1½
Calcium nitrate Ca(NO3)2 · 4H2O Fertilizer 0.8 3 7
Magnesium sulfate MgSO4 · 7H2O Technical 0.2 ¾ 2
Ferric phosphate FePO4 Technical trace trace trace

H’ S #2 (after Hoagland and Aron 1950, except iron chelate replaces ferrous tartrate)
Ammonium phosphate NH4H2PO4 Technical 0.12 ½ 2
Potassium nitrate KNO3 Fertilizer 0.62 2½ 5
Calcium nitrate Ca(NO3 )2 · 4H2O Fertilizer 0.62 2½ 6
Magnesium sulfate MgSO4 · 7H2O Technical 0.38 1½ 4
Chelates of iron as Commercial trace or 2 ppm to the nutrient solution
Versene or Sequestrene
Trace elements (after Hoagland and Aron 1950). A stock solution is made by adding the following to 1 liter (1 qt.) of
water. Use 1 ml (12 drops or 0.034 oz) of the stock solution to 1 liter (1 qt.) of any of the nutrient solutions given
above. If tap water is used, the addition of trace elements is generally not necessary.
Boric acid H3BO3 2.86 g
Manganese chloride MnCl2 · 4H2O 1.81 g
Zinc sulfate ZnSO4 · 7H2O 0.22 g
Copper sulfate CuSO4 · 5H2O 0.08 g
Molybdic acid H2MoO4 · H2O 0.02 g
 P 79

unit in boiling water. When drained and cooled, sow it with spores and cover with a larger glass or jar.
Replace water in the saucer as needed. There is less likelihood of overwatering by this method, and if the
spores are not too densely sown, nicely formed heart-shaped prothalli will develop, free from bits of plant-
ing medium. A dilute solution of fertilizer applied in the saucer will hasten growth. Young ferns may be
loosened and lifted off the clay pot with forceps or a thin, flat blade and planted in potting mix.
Miscellaneous methods. Other media and methods are used to sow fern spores, such as plaster of Paris
blocks, broken clay pots, moistened bricks, tree-fern-trunk fibers, soil, peat moss, and uncut sphagnum
moss. Milled sphagnum is particularly good. The soil and container should be sterilized or treated with
steam or boiling water as a precautionary measure.

A-S C
After sowing, the planting should be placed in the proper light and temperature. The first sign of
growth is the appearance of a very thin green mat about 14 days after sowing. In the early stages of germi-
nation, the emerging green cells are too small to be seen without magnification. Germination usually starts
in 4 to 14 days but may take as long as a year. Ferns grown for the trade usually form mature prothalli in
several months. Others take more than a year to mature.
Light. Place the sown spores in filtered sunlight of low to medium intensity (150 to 500 foot-candles). If
artificial (cool-white fluorescent) light is used, it may be left on continuously or for 8 to 16 hours per day
as desired. Research indicates different cellular responses with blue, red, and far-red light, but these
responses are unimportant to the general growing of spores. Several weeks in total darkness are required for
spore germination in Botrychium dissectum and Ophioglossum, and they may need mycorrhizal fungi to
develop further unless grown in a nutrient medium.
Temperature. The optimal temperature for spores to germinate and grow is 20 to 28°C (68–86°F). Most
growers keep the temperature near 25°C (77°F) and lower it as the plants mature. Uniform temperatures
reduce problems associated with water condensation.
Watering. Watering is unnecessary after sowing if the medium was thoroughly moistened and the plant-
ing covered. The edges of the containers may be sealed with Vaseline petroleum jelly to retain moisture. If
water must be added, set the pots in a saucer of water. Flats and plastic boxes may be watered with a fine mist
or spray. Distilled or cooled boiled water is best during these early stages. Tap water can be an unwanted
source of algae.
Fertilizers. Two to three weeks after the green mat of germinated spores appears, weak solutions of fertil-
izer (about half the recommended strength) can be applied. Commercial growers differ on the concentra-
tion and timing of these applications. Concentrations vary from 100 to 200 ppm, given at every watering
or every other watering. Although researchers have applied sugar (2.5% sucrose) under sterile conditions
to hasten growth, this is not recommended for ordinary purposes because it encourages mold.
Transplanting. Where only a few plants are desired and the prothalli are healthy, transplanting may be
delayed until the sporelings have developed about three leaves. They are then picked out and transplanted,
usually to 6.5 cm (2.5 in.) pots containing a fine-textured potting mix.
In commercial operations, transplanting starts when the green mat of prothalli have formed, about two
months after sowing. Some growers transplant at an early stage when the mat is thick enough to lift off.
Small pieces of the mat 6 to 12 mm (0.2–0.5 in.) across are lifted off with forceps (or the tip of a spoon if
a liquid medium is used). Waiting longer with a liquid medium can result in tangled growth and make it
difficult to get the prothalli to drop off the spoon during transplanting.
80 C 8

The clumps of prothalli are spaced about 12 mm (0.5 in.) apart and firmed onto a moist, finely screened
potting mix of half peat and sand, or whatever mix is desired. Be certain the clumps are in good contact with
the planting medium, otherwise they will dry out. Keep a spray bottle of distilled or boiled cool water
handy, and use it while transplanting. Although not always done, the planting mix should be sterilized or
disinfected. Some growers treat the soil at 88°C (190°F) for three hours before use, and all subsequent
watering is done with cooled boiled water (until the covers are removed from over the planting). Some
growers use fungicide drenches on the transplants at this stage. After they are in place, the transplants
should be watered with a fine spray, preferably of cooled boiled water. Cover the planting with glass or
plastic.
The second transplant is done when the prothalli have enlarged to about 4 mm (0.2 in.) in diameter for
most species. For commercial ferns this is usually about three to four months after sowing. Clumps of pro-
thalli about 10 mm (0.4 in.) or more wide are firmed onto the soil mix and watered as on the first transplant.
Some commercial growers start the second transplant when sporelings appear, at which point a clump of
prothalli along with one or more sporelings is then transplanted, sometimes directly into cell packs or plug
trays, which are plastic trays with small depressions in which the plants are grown and eventually mar-
keted. Keeping the prothalli in small clumps seems to result in better growth than separating them too
finely.
Growers who prefer to sell larger plants usually do a third transplant. This is done when the ferns are
about 1.5 to 2.5 cm (0.5–1 in.) tall or have three well-developed leaves, which in commercial ferns is about
five to seven months after sowing. Sporelings may be planted in fresh flats of soil or 6.5 cm (2.5 in.) pots,
with the usual attention to firm them into place and protect them from drying.
Potted transplants are initially protected in frames under glass or plastic coverings. Once established,
they must be hardened. Hardening is achieved by lifting the glass or plastic covering more and more each
day to allow the foliage to adapt to drier air and higher light. Eventually a lath covering is put in place. The
hardening process may take three to six weeks. Ferns hardened in 6.5 cm (2.5 in.) pots still need protection
and should not be planted directly in the ground unless sheltered conditions can be provided.
During the third transplant, some growers separate the sporelings into uniform size classes before
replanting them in flats or pots. If replanted into flats, the fourth transplant (about nine months after sow-
ing) moves the sporelings into 6.5 cm (2.5 in.) pots, and they are hardened off and marketed in about a
month or two.
Early transplants are almost always made as clumps of prothalli, with or without attached sporelings.
No attempt is made to separate the clumps into individual prothalli because clumps produce more vigor-
ous sporelings than isolated prothalli. This is probably due to greater opportunities for out-crossing with
nearby prothalli. With apogamous ferns, clumps rather than individual prothalli are transplanted for ease
of handling. Between and during transplants it is essential to maintain high humidity and soil moisture
because the transplants are small and will dry quickly. Always use a fine spray of water. If sporelings are slow
to appear, spray the prothalli with enough water to leave a thin film on them for a few hours. This can be
repeated in a few days. Discoloration may indicate a lack of fertilizer or overwatering. Disease may develop
in spite of all precautions (see “Special Care”).

G  P
Home and commercial growers sometimes cultivate ferns entirely in plugs. Plugs are plastic trays with
depressions provided with drain holes. The depressions, or “plugs,” come in different sizes and numbers per
tray. Growing ferns in plugs instead of pots saves labor, planting media, and bench space. It also allows for
holding plants in cooling chambers for a limited time until needed. Cooling chambers hold back growth
to fit marketing schedules. Plug trays designed for the home propagator come with a lower tray to catch
 P 81

draining water and a clear plastic cover to maintain high humidity. The plug trays with small-diameter
depressions (25 mm [1 in.] or less) are suited to growing young ferns and even spores.
The commercial planting mix for plugs differs from most soilless planting mixes by its finer texture and
by containing a wetting agent. For the home gardener, a finer textured soilless mix suitable for planting seeds
will do.
Because the soil capacity in the plug is small, problems can develop rapidly. Greater care must be taken
with the amount and timing of watering. The even distribution and penetration of water are important.
Avoid allowing water to remain on the foliage by watering at a time of day that allows the foliage to dry.
Good air circulation is also important. Crowded wet plants and poor air circulation promote disease (espe-
cially Botrytis in cooler temperatures). Drying of the soil stresses plants and makes them more vulnerable to
diseases as Rhizoctonia and Pythium.
Etiolation, or weak spindly growth resulting from low light, is another problem. It can be prevented by
placing the trays under a strong light (but not so strong that it causes the foliage to turn yellow). Transplants
that are already etiolated make poor material for plug (or small pot) plantings. They often remain leggy and
do not achieve the sturdy growth required of commercial plants. Firmer tissue can also be promoted by ade-
quate ventilation and judicious use of fertilizers. Some growers prefer to use fertilizers sparingly. They also
use nitrate fertilizers, instead of ammonium ones, which they claim produce stockier plants.
Like other plants grown under protected conditions, those grown in plugs will need to be gradually
hardened off. Care must be taken in deciding whether to increase or decrease exposure during the process.
The ideal result among commercial growers is to have “toned plugs,” or plants that are compact, sturdy, of
good color, and hardened off enough to take the rigors of transport to the market.

S C
Mold may develop on the soil, prothalli, and young sporophytes in spite of all precautions. If this hap-
pens, take the following steps immediately. Stop overhead watering. Water with cooled boiled water. Check
that water is not dripping from condensation overhead. Remove the mold and at least 12 mm (0.5 in.) of
the plant tissue beyond the infected area. Finally, drench the area with fungicide and repeat as directed on
the product label (see “Fungi and Bacteria” in Chapter 11).
Water molds (Pythium and Phytophthora) usually cause the prothalli to turn dark and watery and sub-
sequently collapse. Gray mold (Botrytis) appears as gray tufts on the prothalli and foliage. It flourishes
under cool-moist conditions and can therefore be combated by increasing the temperature and decreasing
the humidity. A cyanobacterium (Oscillatoria) also forms a blackish to grayish mold-like growth, and the
threads form dense mats and inhibit growth (see “Algae, Mosses, and Liverworts” in Chapter 11). A pink,
nonpathogenic fungus (Pyronema) that is very resistant to steam treatment may grow over the prothalli and
crowd them out. Rhizoctonia appears as well-spaced threads. If persistent mold or alga infection occurs
even after materials and equipment have been sterilized and the proper cultural conditions are employed,
the sporing room may need to be washed down or drenched with a disinfectant such as formaldehyde,
bleach, or quaternary ammonium compounds (Physan 20, Consan). In sporing rooms and growing rooms,
it is important to reduce air movement from the outside because it might carry in fungus spores.
Insects are another problem (see Chapter 11). Fungus gnat larvae can attack prothalli. They can be
killed by sprays, but it is best to keep the adult flies out of the cultures by tightly sealing the containers with
plastic wrap, Vaseline petroleum jelly, or cotton batting placed between the pot and glass covering. Cultures
grown in terrariums are said to be undamaged by fungus gnats if an insectivorous plant, such as sundew
(Drosera), is also placed in the terrarium. Foliar nematodes can also damage prothalli.
The failure of prothalli to produce young ferns (sporophytes) long after sowing (6 to 18 months) can
be due to several causes. The most common cause is insufficient water for the sperm to swim to the egg. If
82 C 8

sporelings develop only near the edges of the container (where water condensation is greatest), this is a
good indication that more water is needed. If misting does not produce sporophytes, fertilization can usu-
ally be achieved by flooding the prothalli with a thin film of water for a few hours and then letting it drain
away. Other reasons for failure are more difficult to determine. Sometimes the sowing is so dense that the
crowded prothalli produce only antheridia, not archegonia, in which case sporelings cannot be produced
because there are no eggs to be fertilized. In other cases, ferns need a basic medium to grow healthy prothalli
(as in some Pellaea species). Some ferns just take a long time to form sporelings (two years is not uncom-
mon). Slow-growing cultures often become contaminated by algae and mosses (see Chapter 11).

F E  G  S

Some ferns are easy to grow from spores whereas others (such as Elaphoglossum and some Blechnum) are
difficult. Polystichum and many species of Polypodium, for example, germinate satisfactorily but produce
sporelings slowly. On the other hand, the following ferns are easy and quick to grow from spores and read-
ily form sporelings. They are highly recommended for the beginner.
Adiantum capillus-veneris Microlepia platyphylla
Adiantum hispidulum Osmunda, most species—fresh spores can
Adiantum raddianum germinate in one day
Asplenium platyneuron Pellaea viridis
Athyrium filix-femina Phlebodium aureum
Cheilanthes bonariensis Phlebodium pseudoaureum
Cheilanthes wrightii Pityrogramma austroamericana
Cyrtomium falcatum Pteris cretica
Doodia aspera Pteris multifida
Doodia media Pteris tremula
Dryopteris affinis Pteris vittata
Hypolepis tenuifolia Thelypteris dentata
Macrothelypteris torresiana Thelypteris parasitica
Matteuccia orientalis Thelypteris puberula
Microlepia strigosa Todea barbara

C T  S

Most fern species or cultivars produce relatively uniform offspring, but occasionally they produce
sporelings that appear quite different. The differences range from subtle to conspicuous (see “Variation
Within Species” in Chapter 12). Some commonly cultivated species that produce variable offspring include
Adiantum raddianum, Asplenium bulbiferum (the narrow-segment form), Athyrium filix-femina, Phlebodium
aureum, Phyllitis scolopendrium, and Polystichum setiferum. With these species it is necessary to cull by hand
those sporelings that vary greatly from the parents, but there is always the possibility of being rewarded with
a sporeling that has highly unusual but attractive characteristics. Such sporelings are mostly the result of
mutations or new hereditary combinations and might not reproduce true from spore. Rarely are they due
to hybridization between different species.
 P 83

Hybridizing Ferns
Ferns are famous (or infamous!) for hybridizing. They hybridize in nature, and many fern hybrids have been
artificially produced in the laboratory. Some genera that hybridize frequently are Asplenium, Dryopteris, and
Thelypteris.
Fern hybrids can be recognized because they are intermediate between their parents. If the parents
greatly differ, the hybrids tend to be irregular and vary widely in size, shape, or number of discrete struc-
tures (pinnae, pinnules, sori, and so forth) between the extremes set by the two parents. Hybrid interme-
diacy can usually be seen in frond shape, vein patterns, and sorus shape. Contrary to expectation, hybrids
made at different times from the same parents are not always identical.
Fern hybrids are not always superior to their parents, but some do display hybrid vigor. For example,
Adiantum ×tracyi has physiological qualities superior to both its parents (A. jordanii and A. pedatum). In cul-
tivation, the hybrid grows more robustly than either parent and, in California, it remains green most of the
year. An unfortunate feature of this hybrid (and many others like it) is that it is sterile and can only be prop-
agated by dividing its rhizome. Only rarely are fern hybrids fertile—that is, capable of producing viable
spores.
The simplest method for hybridizing ferns is sowing spores of the two parents together and hoping that
cross-fertilization will occur (Figure 8.12a). The resulting sporelings have to be grown to a fairly large size
to see whether they have the expected intermediate characteristics of the hybrid. This takes a lot of time and
material.
Another method involves sowing the spores of the parents separately and then transplanting the pro-
thalli close to each other in the hope that a hybrid will develop. With this method you can be certain that
spores of both parents have germinated and produced prothalli. For details about this method, see Rasbach
et al. (1994).
Hybrids can also be produced by planting different parts of two mature prothalli close together (Fig-
ure 8.12b). The notched end of the prothallus bears most of the archegonia that contain the eggs; the
remaining part of the prothallus (the tapered end) bears most of the antheridia that contain the sperm. By
cutting one-third of a prothallus at the notched end of one parent and one-third at the tapered end of the
second parent, and by planting these ends together, it is possible to obtain a hybrid. Any sporophytes
resulting from this method are not necessarily hybrids, however. It is possible that the sporeling resulted
from self-fertilization that occurred before separation, or from a selfing after separation (sometimes
antheridia are still present near the notched end of the prothallus), or from new proliferations of the pro-
thallus. Nevertheless, the chances of producing a hybrid by this method are much better than planting
complete prothalli of two species next to each other (Lovis 1968).
Hybrids can also be produced by concentrating the sperm from one parent in water and then adding pro-
thalli of the other parent (Figure 8.12c). The spores from the species intended to be the male parent are
densely sown to encourage antheridia to form. Prothalli used as the female parent are sown thinly or spaced
apart by transplanting to produce more archegonia. The prothallus cultures are sparingly and carefully
watered by soaking the soil from below, so as to prevent a film of water from developing between the soil and
prothalli—this film of water may cause premature dispersal of the sperm and unwanted self-fertilization. Care
must be taken to prevent condensation on the planting cover from falling on the prothalli. Because antheridia
develop before archegonia, the spores of the intended female parent should be sown four to six weeks earlier
so that both parents will be at the proper stage of development simultaneously. Because it is difficult to
know when the parents are at the optimum stage of development for hybridization, it is useful to have sev-
eral cultures of both parents sown on different dates. Prothalli with antheridia suitable for hybridization
usually develop three to four months after sowing, whereas archegonia will develop one to two months after
 a

male parents female parents

Cut the tapered end off the Cut the notched end off the
intended male parent. intended female parent.

Plant the tapered and notched ends next

to each other in a new planting.

Figure 8.12. Various methods of hybridizing ferns: a. planting spores or young prothalli of intended parents together;
b. exchanging notched and tapered ends of intended parents; c. growing parents separately, mixing them, and then
growing potentially fertilized prothalli separately.
 c

Sow spores of the intended

male parent densely.
Place cluster of male parents in Sow spores of the intended
a small, shallow dish of water. female parent thinly, or
Tease them apart. transplant to provide ample
developing room.

Place 30 cm (12 in.)

under a light until
sperm are seen in Clean 8 to 12 female
samples examined parents of debris.
under a microscope.

Place right side up in

dish with male parents.

Cover with glass and keep under a

light for ½ hour to overnight.

Remove female parents to a clean

dish, rinse several times to
remove any male parents.

Plant potentially fertilized female

parents in fresh separate plantings.
86 C 8

the antheridia are ready. Because unwanted self-fertilization may occur, the archegonia must be examined for
signs of sporophyte development before use. This is best done under a dissecting microscope.
The hybridization technique consists of placing about 1.5 square centimeters (¼ sq. in.) of the small
young prothalli bearing the antheridia in a dish (watch glass) with a small amount of warm water, about 25
to 35°C (77–95°F), and teasing the antheridia apart. The dish is then placed 30 cm (12 in.) beneath a 40-
watt light bulb or in sunlight. At 20-minute intervals, the water is checked under a microscope for active
sperm. Once active sperm have been found, 8 to 12 prothalli of the female parent, which have been cleaned
of debris and examined for the absence of sporophytes, are immediately placed, archegonia-side down, on
the water containing the sperm. The dish is covered with glass to prevent drying and returned to the light.
It is left undisturbed for a half-hour to overnight. At the end of this time the female prothalli are removed,
washed several times in fresh water to remove any of the small male prothalli, and planted in fresh soil.
Hybrid sporophytes usually appear four to six weeks after fertilization. Any sporophytes that appear
before this time generally are the result of self-fertilization. A success rate of about 80% has been reported
with this technique in producing hybrids between closely related and compatible strains. Fern sperm is
known to be attracted to malic acid (0.01%) and citric acid, and this may be of use in hybridization work.
Generally, only closely related ferns hybridize. Other abnormal plants are sometimes confused with
hybrids. Abnormal fronds are caused by injury, mutation, or unusual gene recombinations. If a suspected
hybrid can be duplicated by artificial hybridization in the laboratory, then it probably is a true hybrid.
Indirect proof can be found by comparing a suspected hybrid with its parents and looking for intermedi-
ate features. In all cases, keep records of the parents and the methods used in hybridizing because this infor-
mation is scientifically important.
 9 Landscaping

When landscaping with ferns, as with any plants, be sure that they receive enough warmth, sunlight,
humidity, protection from wind, and proper soil conditions. Keep in mind their size when fully grown and
any special problems that may arise in their upkeep. Attention to these considerations early on avoids dis-
appointment later.
The planting design you have in mind should be visualized in relationship to its surroundings. Will the
fern harmonize with its surrounding environment? Will it eventually be too shaded or crowded by sur-
rounding trees and shrubs? Are there too many elements in the design that create a sense of “busyness” or
confusion? Sometimes plants that differ greatly provide a refreshing sense of contrast; however, they can be
jarring or disturbing if carelessly selected. The placement of the plants in relationship to the surroundings
should provide a restful sense of balance. If unbalanced, the design can be disturbing to the eye’s sense of
equilibrium. These considerations are basic landscape design principles which when correctly applied pro-
vide the most pleasing aesthetic arrangement to your garden.

Solving Particular Landscaping Problems

Besides using ferns in general landscaping and for blending or contrasting with other plants, ferns can be
used to solve particular problems or needs. Lists of appropriate ferns to use for some of these purposes are
provided later in the chapter under “Ferns for Particular Landscape Purposes.”

C  I
Plants used to create center points of interest are called accent plants. The most effective accent plants
have either strong, bold, well-defined patterns to their foliage or, at the other extreme, are soft, fine, and deli-
cate. Accent ferns should not be too small or be crowded among other plants. The general outline of the
plant should have an interesting pattern rather than be just an uninteresting blob of pretty foliage.

C  I  D

Use fine-textured, airy-looking plants with soft lines to give the feeling of distance. Coarse-textured
plants give the feeling of closeness. Generally, soft-colored foliage gives the sense of distance, whereas bright
colors give a sense of closeness.

87
88 C 9

Figure 9.1. Ferns can be useful in hiding distracting views.

Figure 9.2. Using ferns as a center of interest.

H D V

Ferns can be used to hide unwanted views. Hanging baskets are good at blocking the top of a distract-
ing view. The lower part can be hidden with medium-sized ferns planted in back or in front of the hang-
ing baskets.

Figure 9.3. Creating the illusion of distance with ferns.

M  C  F

By carefully hanging baskets at different heights it is possible to produce a curtain of foliage across the
side of a patio or outside a window. The climbing ferns (Lygodium) are well suited to producing a screen of
foliage if they are provided with wires for support.

Figure 9.4. A curtain of ferns.

A H  W

Basket ferns can be placed on top of walls to give added height or to hide distracting views (such as the
neighbor’s clothesline). Wire fencing (2-by-1-inch mesh) can be cut, bent, and formed into a basket that
will fit nicely across the top of a concrete block wall. The lower front and back edge of the basket should be
extended to form a lip to secure the basket on the wall. The basket in effect straddles the wall. Wooden boxes
constructed to securely straddle the wall are also suitable.
 L 89

Figure 9.5. Using ferns to add height to walls.

P F A E L

Sometimes the effect of a small palm or tree is needed inside a small room, but there is not enough space
to put a large container on the floor. Hanging a basket fern from the ceiling will achieve the same effect and
keep the floor space open.

Figure 9.6. A hanging basket fern offers foliage above eye level.

A I   B W

A bare wall can be made attractive by hanging ferns in baskets or containers or with staghorn ferns
mounted on boards. A small piece of wood or a wad of aluminum foil wedged between the basket and the
wall will help prevent water stains on the wall.

Figure 9.7. Ferns used to decorate a bare wall.

S R S

Certain ferns creep over rocks and thereby have the effect of softening hard lines. Provide pockets of soil
in the rock to get these ferns started. Many species of Davallia and Polypodium are admirably suited for this
use.
90 C 9

Figure 9.8. Softening rock surfaces with ferns.

B  F P

Border ferns are useful for softening the hard lines of a concrete walk. Use small- to medium-sized
ferns that will not quickly grow out of bounds. Many maidenhair ferns (Adiantum) are the right size for this
and grow well along the concrete. Blechnum appendiculatum is suitable in warmer climates. Many ferns can
also be planted along the base of a bare foundation or wall to break the visual monotony. Using species with
short-creeping rhizomes will reduce maintenance needs. The ostrich fern (Matteuccia struthiopteris) is fre-
quently used as a foundation planting around homes in the eastern United States.

Figure 9.9. Ferns can effectively soften the border

along a wall or walkway.

G C
Where conditions are shady or semi-shady, ferns do well as a ground cover. In such uses, ferns also help
prevent erosion.

Figure 9.10. Ferns used as ground cover.

 L 91

Ferns for Particular Landscape Uses

A variety of landscape situations can be served by a range of fern species, depending on the climate and the
intended landscape effect. The lists presented here suggest some of the more common or adaptable ferns for
specific purposes. Chapter 10 also includes information on growing ferns for certain situations and on the
landscape use of specific types of ferns, such as the tree ferns. The entries in Chapter 13 provide more
detailed information on the requirements and uses of these ferns.

O F
The 10 ferns listed here for each section of the United States were selected because they are easy to grow,
attractive, and readily obtainable. Native ferns are favored on these lists. Most parts of the United States
require hardy to very hardy ferns. For additional outdoor species and their temperature tolerances, see
Chapter 13.
Northeastern United States. Hardy to very hardy ferns are the only ones suitable for the northeastern
United States, where prolonged periods of cold are common. This area includes mostly Zones 3–6. Sum-
mers are hot and humid in some areas and often difficult for ferns that prefer cooler summers.
Adiantum pedatum Matteuccia struthiopteris
Deparia acrostichoides Onoclea sensibilis
Dryopteris erythrosora Osmunda claytoniana
Dryopteris goldiana Osmunda regalis
Dryopteris marginalis Polystichum acrostichoides
Southeastern United States. Depending on the locality, ferns for the southeastern United States may be
hardy to semi-tender. This area includes mostly Zones 6–8. For the cooler areas of the Southeast, see the
listing for “Northeastern United States,” and for milder areas see “Southern Florida and Hawaii” and
“Coastal Central and Southern California.” Ferns adapted to hot, humid summers thrive in this area.
Southern Florida and Hawaii. This area includes mostly Zones 10 and 11. Tropical species are favored
here, but a wide range of ferns are adapted to the climate of southern Florida and Hawaii, except those ferns
that like cool summers or need cold temperatures to break dormancy. The occasional cold spells may
require moving the more tender species to protected areas.
Adiantum trapeziforme Microsorum punctatum
Aglaomorpha coronans Nephrolepis falcata ‘Furcans’
Asplenium nidus and relatives Platycerium bifurcatum and cultivars
Davallia, many species Selaginella, many tropical species
Microsorum grossum Sphaeropteris cooperi
Central United States. This area includes mostly Zones 3–6. In the northern part of this area, only very
hardy ferns are suitable; see “Northeastern United States.” For the southern area, see the listings for south-
eastern and southwestern states.
Northwestern United States. This area includes mostly Zones 5–8, with a few pockets of Zone 9. Some
semi-hardy ferns will grow in coastal areas, but the hardy species listed here will be more reliable on coastal
and inland areas. Coastal areas have cooler summers that may not favor species adapted to hot summers.
92 C 9

Adiantum aleuticum Dryopteris dilatata

Athyrium filix-femina Phyllitis scolopendrium
Athyrium niponicum ‘Pictum’ Polypodium vulgare
Blechnum spicant Polystichum andersonii
Dryopteris affinis Polystichum munitum
Southwestern United States. This area includes mostly Zones 6–9. In most areas, hardy or semi-hardy ferns
will do best. Some milder areas will support semi-tender species. High aridity and summer heat are usual
problems. The water or soil can be basic in certain locales, making it difficult to grow acid-loving ferns.
Adiantum capillus-veneris Osmunda cinnamomea
Asplenium platyneuron Pellaea atropurpurea
Athyrium filix-femina Polystichum setiferum
Dryopteris filix-mas Woodsia obtusa
Onoclea sensibilis Woodwardia virginica
Coastal central and southern California. This area includes mostly Zones 9 and 10. In southern Califor-
nia and the central coastal areas, semi-hardy or semi-tender species do best. Many temperate ferns adapt to
this area, but not those that need a cold winter to break dormancy. Aridity is a problem during heat waves,
and the water or soil in some areas are basic, making it difficult to grow acid-loving ferns.
Adiantum raddianum and cultivars Microlepia strigosa
Asplenium bulbiferum Polystichum polyblepharum
Cyrtomium falcatum and cultivars Pteris cretica
Davallia mariesii var. stenolepis Rumohra adiantiformis
Dicksonia antarctica Sphaeropteris cooperi

S-T F
The following ferns will grow in areas that are shadier than usual, but they still must have a certain
amount of light. Provide more light in climates where the weather is frequently overcast. Most of these ferns
will grow in brighter light, but they may be less luxuriant.
Adiantum aleuticum Dryopteris goldiana
Adiantum capillus-veneris Dryopteris marginalis
Adiantum pedatum Gymnocarpium dryopteris
Adiantum raddianum Nephrolepis cordifolia
Arachniodes standishii Osmunda cinnamomea
Asplenium bulbiferum Phegopteris connectilis
Athyrium filix-femina Phyllitis scolopendrium
Blechnum spicant Polystichum acrostichoides
Dennstaedtia punctilobula Polystichum munitum
Dryopteris dilatata Polystichum setiferum
Dryopteris filix-mas

S-T F
Ferns seldom look their best when growing in direct sun (if they grow at all in such intense light). The
following ferns appear reasonably attractive if grown in places with direct morning or late-afternoon sun.
Some tolerate full sun if the skies are often overcast. Elsewhere they need filtered light during the hottest part
 L 93

of the day. Those species marked with an asterisk (*) will tolerate full sun only if soil moisture and humid-
ity are adequate.
Aglaomorpha coronans Osmunda, all species*
Athyrium filix-femina Pellaea, most species
Blechnum appendiculatum Phlebodium pseudoaureum
Cheilanthes, most species Pityrogramma, most species
Cibotium glaucum* Platycerium veitchii
Cyrtomium falcatum Polystichum polyblepharum
Dennstaedtia punctilobula* Pteridium aquilinum*
Doodia media Pteris cretica
Dryopteris erythrosora Pteris tremula
Dryopteris ludoviciana Rumohra adiantiformis
Lygodium japonicum Sphaeropteris cooperi
Microlepia platyphylla Thelypteris noveboracensis
Microlepia strigosa Thelypteris puberula
Nephrolepis cordifolia Todea barbara
Onoclea sensibilis*

A F
Aquatic ferns are those that have their foliage submerged (or partly so) or floating on the surface. Such
ferns vary considerably in form compared to bog and wet-soil ferns. Some aquatic ferns can be purchased
from aquarium supply stores. For some species, water that is high in salts may stunt growth or may burn
or blotch the leaves. Changing the old water completely rather than adding more helps reduce salts and
increase aeration. Using a high proportion of sand and adding charcoal to the rooting medium will help
keep the water free of noxious chemicals and odors.
Acrostichum aureum Marsilea, all species
Azolla, all species Microsorum pteropus
Bolbitis heudelotii Salvinia, all species
Ceratopteris, all species

B  W-S F

Bog ferns grow in soggy or poorly drained areas and only incidentally have their foliage submerged in
water. Their roots tolerate submergence—though some species are more tolerant than others. In very wet
soils, avoid high amounts of organic material. Sharp sand and charcoal can be added to keep the soil sweet
(that is, to avoid anaerobic fermentation and the accumulation of its often toxic or unpleasant byproducts).
For ferns that float or grow partly or totally submerged, see the preceding discussion of “Aquatic Ferns.”
Some ferns may be both aquatic and bog-adapted, such as Ceratopteris and Marsilea.
Acrostichum aureum Marsilea, all species
Acrostichum danaeifolium Matteuccia struthiopteris
Blechnum serrulatum Microsorum pteropus
Ceratopteris, all species Onoclea sensibilis
Dryopteris ludoviciana Osmunda regalis
Equisetum, all species Thelypteris palustris
94 C 9

F  B  L S

These ferns need more calcium than others, preferring a soil pH of 7–8. Where limestone is available,
the plants can be grown in soil pockets in the rock. (See Chapter 5 for more information on limestone-
loving ferns.)
Adiantum tenerum Ceterach officinarum
Asplenium adiantum-nigrum Gymnocarpium robertianum
Asplenium ruta-muraria Phyllitis scolopendrium
Asplenium platyneuron Polypodium cambricum
Camptosorus rhizophyllus Polystichum aculeatum

F  A S

These ferns grow in acidic soils with pH 4–7. (See Chapter 5 for more information on acid-loving
ferns.)
Blechnum penna-marina Dryopteris tokyoensis
Blechnum spicant Osmunda claytoniana
Dennstaedtia punctilobula Phegopteris connectilis
Dryopteris clintoniana Polystichum, various species
Dryopteris expansa

D S F

The following ferns tolerate drier, more exposed areas than most ferns. They need regular watering and
protection until established. These are not xerophytic ferns, which are discussed in Chapter 10.
Aglaomorpha coronans Microsorum viellardii
Blechnum appendiculatum Nephrolepis cordifolia
Cyrtomium falcatum Phlebodium pseudoaureum
Davallia mariesii var. stenolepis Pteridium aquilinum
Davallia tyermannii Pteris cretica
Dennstaedtia punctilobula Pteris vittata
Dryopteris filix-mas Rumohra adiantiformis
Microlepia strigosa Sphaeropteris cooperi

R G F

In optimum conditions of temperature, light, and nutrients, the following species grow relatively
rapidly.
Adiantum raddianum Matteuccia struthiopteris
Anogramma chaerophylla Microlepia strigosa
Athyrium filix-femina Nephrolepis cordifolia
Cibotium schiedei Onoclea sensibilis
Dennstaedtia punctilobula Phegopteris connectilis
Goniophlebium subauriculatum ‘Knightiae’ Pteridium aquilinum
Hypolepis, most species Pteris cretica
Macrothelypteris torresiana Pteris tremula
 L 95

Pteris vittata Thelypteris, most species

Selaginella kraussiana Woodwardia radicans
Sphaeropteris cooperi Woodwardia virginica
Sphaeropteris excelsa

S G F

The following greenhouse or subtropical ferns are slow growing.
Alsophila tricolor Drynaria, all cultivated species
Angiopteris, all species Elaphoglossum, most species
Cibotium glaucum Marattia, all cultivated species
Dicksonia antarctica Platycerium, all species
Dicksonia fibrosa Pyrrosia lingua

C  F F

The young fronds of many species of Adiantum and Blechnum are red when they emerge and turn
green with age. Other ferns are white or yellow on the underside and remain so at maturity. Variegated ferns
—those with stripes or blotches of white, yellow, red, or other colors on their fronds—are mostly cultivars
or forms of green species and are generally more difficult to grow.
Adiantum hispidulum—young fronds red
Adiantum macrophyllum—young fronds red
Adiantum raddianum ‘Variegatum’—variegated white
Alsophila tricolor—underside white
Arachniodes simplicior—variegated white and yellow
Athyrium niponicum ‘Pictum’—variegated grayish and purple
Athyrium otophorum—stipe, rachis, and costa red
Blechnum appendiculatum—young fronds red
Blechnum brasiliense—young fronds red
Cheilanthes argentea—underside yellow or white
Dryopteris erythrosora—young fronds red, indusia typically red
Microlepia platyphylla—bluish green foliage
Phlebodium aureum ‘Mandaianum’—bluish green foliage
Pityrogramma austroamericana—underside yellow
Pityrogramma calomelanos—underside white or yellow
Pteris argyraea—variegated white
Pteris aspericaulis—stipe, rachis, and costa deep red
Pteris ensiformis ‘Evergemiensis’—variegated white
Pteris ensiformis ‘Victoriae’—variegated white
Pteris nipponica—variegated white
Selaginella, several species—red, gold, blue, iridescent, or variegated

E  D F

The question of whether ferns are evergreen or deciduous is of varying importance to different gar-
deners. In climates where the garden is used year-round, evergreen ferns are preferred. Where the garden
96 C 9

is used seasonally, deciduous species are more frequently cultivated, although some hardy evergreens might
be welcomed. Where seasonal changes are eagerly anticipated, the deciduous species are symbols of spring
as their crowns of beautiful fresh green fiddleheads emerge after a long winter.
One reason it is important to know whether a fern is deciduous is so that you do not think it is dying
when it goes into its rest period. Growing hardy, deciduous ferns in warm climates requires selecting adapt-
able species (sometimes a matter of trial and error), not giving them too much water in the winter (they will
rot), and remembering not to accidentally dig them up when working in the garden. Young plants of some
deciduous species can retain their greenness during their first one or two years of life.
A fern that is deciduous will have all its leaves decline at about the same time. In contrast, an evergreen
fern has leaves that do not decline all at the same time. Contrary to popular belief, evergreen plants do not
have leaves that stay green forever—the older leaves are eventually shed. Whether plants are deciduous or
evergreen, their declining leaves may be self-shedding (articulate) or they may wither in place (marces-
cent). These characteristics affect their landscape use and maintenance.
Deciduous ferns with self-shedding fronds have a special layer (the abscission layer) near the base of the
stipe that allows a clean break at the joint with the stem. Self-shedding fronds require mostly seasonal rak-
ing for maintenance.
Deciduous ferns with fronds that wither in place are more troublesome to groom because their fronds
will need to be cut from the plant to maintain neatness. Because withered, persistent fronds protect the
crown from winter cold, they are usually removed in spring before the new fronds have emerged. For neat-
ness, year-round grooming may be needed to remove any off-season, withered leaves.
Fronds of evergreen ferns generally live a few months to a few years, depending on the species and cul-
tural conditions. Spent fronds of evergreen ferns may be self-shedding (as in Polypodium) or marcescent (as
in Polystichum). Expect a light litter or light grooming problems year-round with most evergreen ferns. For
more details on grooming, see Chapter 6.
The distinction between being evergreen and deciduous is not always clear. Some ferns lose all their
leaves in cold climates but do so only partly in warmer climates. Reclined, marcescent fronds can still con-
tain functional conducting tissue and may stay green and photosynthetic for weeks. Other ferns gradually
reduce growth, and in the absence of new growth, the existing fronds become increasingly tattered but
stay erect. In certain climates the period of dormancy in some ferns is so short that the shedding of the old
fronds is simultaneous with the flush of new fronds, which makes it seem as if the plant is evergreen. Some
species have a rhythm of leaf loss that does not coincide with Northern Hemisphere winters. Many tropi-
cal species grow slowly during the cool months and resume active growth as soon as warm weather returns.
Given these intermediate conditions, discrepancies abound in the literature on the subject. To be clear,
deciduousness or evergreenness should be stated relative to a given climate.
Deciduous ferns. The following ferns are commonly available and obligately deciduous. Their fronds are
deciduous even in warm climates, although their dormancy period may be shorter. Those listed here are
marcescent.
Athyrium filix-femina Gymnocarpium dryopteris
Athyrium niponicum Matteuccia struthiopteris
Athyrium otophorum Onoclea sensibilis
Deparia acrostichoides Osmunda regalis
Diplazium pycnocarpon Phegopteris connectilis
Evergreen ferns. The following common ferns are evergreen even in cold-temperate climates, although their
new growth may be delayed until spring. Those listed here are marcescent.
Blechnum penna-marina Cyrtomium falcatum
Blechnum spicant Dryopteris erythrosora
 L 97

Dryopteris intermedia Polystichum setiferum

Phyllitis scolopendrium Polystichum tsus-simense
Polystichum acrostichoides

A F
Accent ferns have distinctive lines, shapes, textures, or patterns. They can be used as center points of
interest in landscaping.
Aglaomorpha coronans Matteuccia struthiopteris
Asplenium bulbiferum Microsorum punctatum ‘Grandiceps’
Asplenium nidus Nephrolepis exaltata and cultivars
Blechnum brasiliense Osmunda, all species
Blechnum gibbum Phlebodium aureum and cultivars
Cyrtomium falcatum Platycerium, all species
Didymochlaena truncatula Tree ferns, all species

B  F F

The ferns listed here spread sparingly or keep their general shape when planted along a flower border
or walk or foundation of a house. Ferns that are short-creeping or clumping or have erect rootstocks are
good for this use. Ferns with long-creeping rhizomes are unsuitable because they outgrow their boundaries
too quickly, and they are best used as ground covers.
Adiantum hispidulum Microlepia strigosa
Adiantum pedatum Pellaea rotundifolia
Adiantum raddianum Phyllitis scolopendrium
Athyrium filix-femina Polystichum acrostichoides
Athyrium niponicum ‘Pictum’ Pteris cretica and cultivars
Diplazium subsinuatum Pteris multifida and cultivars
Dryopteris erythrosora Thelypteris, many species

F  G C

Ferns useful as ground covers eventually carpet the ground by means of their widely creeping rhizomes
or stolons. In some cases, ferns with short-creeping rhizomes and clusters of arching fronds can produce the
same effect. The following ferns spread by rhizomes or stolons.
Blechnum appendiculatum Nephrolepis cordifolia and other species
Blechnum penna-marina Onoclea sensibilis
Davallia, all species Phegopteris connectilis
Dennstaedtia, all species Phegopteris hexagonoptera
Gymnocarpium dryopteris Selaginella, several species
Hypolepis, all species Thelypteris acuminata
Matteuccia struthiopteris Thelypteris palustris

B F
Ferns in hanging baskets or containers are commonly used in today’s house and garden decor. Both epi-
phytes and terrestrial ferns can be used in hanging displays. Species with long fronds that droop over the
98 C 9

edges of the container are particularly attractive. General directions on the selection, planting, and care of
basket ferns are given in Chapter 7, and details on growing specific groups of ferns are provided in Chap-
ter 10. Some of the more popular basket ferns are listed here.
Adiantum capillus-veneris Phlebodium aureum
Adiantum raddianum Phlebodium pseudoaureum
Aglaomorpha, all species Platycerium, all species
Campyloneurum, most species Polypodium, most species
Davallia, all species Pyrrosia lingua
Goniophlebium, all species Rumohra adiantiformis
Nephrolepis, all species

R G  W F

Many ferns can be grown among rocks, but rock walls or rock gardens are traditionally planted with
small to medium-small ferns. The rock background allows these ferns to be more easily seen yet safely
tucked away to avoid being stepped on or overly shaded by rapidly growing plants.
Ideally, some of the rocks should be limestone so that lime-loving ferns can be grown. If limestone can-
not be found, lime can be added to the soil mixes (see Chapter 5).
Using the proper soil before setting the rocks in place avoids future growth problems. The ferns should
be firmly planted with a loam or potting mix in an area between the rocks that is a few inches wide and deep
and has good drainage. Some ferns will need additions of coarse sand. Small rocks added during planting
will help secure the fern and soil in place. Protect the surface soil against erosion by firming it and placing
gravel or a few small stones on top of it. Generally, temperate species of rock ferns adapt poorly to warm-
climate gardens. Where rainfall is moderate to sparse, xerophytic ferns do well in rock plantings (see “Xero-
phytic Ferns” in Chapter 10). Also see “Trough Gardens” in Chapter 10. Kaye (1968) provides further
details on planting hardy ferns in rock gardens. Some suitable ferns for rock gardens are
Adiantum aleuticum subsp. subpumilum Davallia tyermannii
Adiantum capillus-veneris Dryopteris, small species or cultivars
Adiantum hispidulum Gymnocarpium dryopteris
Adiantum raddianum ‘Pacottii’ Lemmaphyllum microphyllum
Adiantum venustum Nephrolepis cordifolia ‘Lemon Buttons’
Asplenium ×ebenoides Nephrolepis exaltata, dwarf forms
Asplenium platyneuron Pellaea, all species
Asplenium ruta-muraria Phyllitis scolopendrium
Asplenium trichomanes Pityrogramma, all species
Astrolepis, all species Polypodium, all species of the P. vulgare
Athyrium filix-femina, small cultivars complex
Camptosorus rhizophyllus Polystichum lemmonii
Camptosorus sibericus Polystichum tsus-simense
Ceterach aureum Pteris cretica and cultivars
Cheilanthes, all species Pteris multifida and cultivars
Cryptogramma crispa Pyrrosia, all smaller species
Cystopteris bulbifera Selaginella, many species
Cystopteris fragilis Woodsia obtusa
Davallia mariesii Woodsia oregana
10 Growing Special Ferns

House Ferns
Ferns add grace to a room, and if the room is not too dry and has adequate light, many species can serve as
permanent decorations. Light and humidity often can be improved, but if not, it is best to use several ferns
in rotation, replacing a plant with a fresh one when it shows poor growth. Plants removed from display will
recover if given sufficient light, humidity, and care. Since the kitchen and bathroom are often the most
humid rooms in the house, they are good places to grow ferns if the light is adequate.
Generally, robust ferns with leathery fronds, such as the house holly fern (Cyrtomium falcatum) and
leather fern (Rumohra adiantiformis), grow well indoors. Most davallias are also good as house ferns, and
some are attractive for their finely cut foliage (Davallia fejeensis and D. mariesii var. stenolepis). The most fre-
quently used indoor ferns are the Boston ferns (Nephrolepis exaltata and its cultivars), which come in a
variety of textures.
If possible, start with nearly mature ferns that are acclimated to drier conditions. These “hardened”
plants will grow better in the drier air of a house than those recently removed from a humid greenhouse.
Avoid selecting plants that appear too soft and green.
Frequency of watering depends on the relative humidity of the room, the size of the fern, the type of
soil, and the kind of pot. Most indoor ferns require a thorough watering two to three times a week, more
if the air is dry and less if cool. Water the pot until the water runs out of the drain hole in order to wash out
the salts that accumulate in the soil. Do not permit pots to sit continually in saucers of water. If you are
watering pots by setting them in saucers, remove the pots as soon as the soil has become moistened through-
out. If the pots lack drain holes, use distilled water or rainwater to avoid salt accumulation, but be careful
not to overwater. Be sure that a layer of coarse drainage material such as gravel, pot shards, or perlite is at
the bottom of pots lacking drain holes. Charcoal added to the soil absorbs noxious chemicals produced by
microorganisms that thrive in the wetness. It is better, however, to avoid overwatering than to rely on the
charcoal’s absorptive power. Always remember that the soil should feel moist, not soggy.
Every two weeks or so, apply a fine spray of water to the foliage to wash off dust and insects. Watch
for scale insects, mealybugs, and aphids. These pests can spread quickly because they have few natural ene-
mies indoors. Furthermore, they are not washed off the plant by rain or overhead watering, as often hap-
pens outdoors.
The main challenge in growing ferns indoors is to provide enough humidity and light. Otherwise the
culture of indoor ferns is similar to that of ferns used elsewhere. The following are some favorite house ferns.

99
100 C 10

Adiantum hispidulum Microsorum grossum

Asplenium antiquum Microsorum punctatum and cultivars
Asplenium australasicum Nephrolepis exaltata and cultivars
Asplenium bulbiferum Pellaea rotundifolia
Asplenium daucifolium Phlebodium aureum
Asplenium nidus Phlebodium pseudoaureum
Cibotium schiedei Platycerium bifurcatum
Cyrtomium falcatum and cultivars Pteris cretica and cultivars
Davallia fejeensis Pteris tremula
Davallia mariesii var. stenolepis Rumohra adiantiformis

Trough Gardens
Trough gardens can be thought of as large, deep-dish gardens. Stone troughs used by farmers to water ani-
mals became much prized as planters for miniature plants. Trough and similar broad containers are ideal
for displaying small ferns, confining species with long-creeping rhizomes, and serving as garden ornaments.
Handsome troughs can be made from easily obtainable materials. Hypertufa troughs are made from one
part cement, one part coarse sand, and two parts shredded peat moss. These ingredients are mixed with
water to the consistency of creamed cottage cheese and poured into a mold to form 5-cm (2 in.) thick
walls. The mold can be fashioned from two cardboard cartons and reinforced with wire mesh. After two
days, the mold is peeled away and the cast allowed to cure for seven days. Any plant toxins in the cement
are neutralized by filling the trough with water and adding about 2.5 g (½ tsp.) of dissolved potassium per-
manganate. After two to three hours, the solution is discarded and the trough is rinsed out. Drainage holes
are drilled into the trough. Troughs are allowed to cure for another two or three days. See MacPhail (1990)
for details on reinforcing thin (less than 5 cm [2 in.] thick) troughs with hardware cloth, creating textural
refinements, using other materials, and readying the trough for final planting. Rock garden magazines
often list other ingredients and directions for building planting troughs. When preparing the soil for a
trough, remember that containers with broad bottoms tend to drain slowly, so use a well-drained mix or
provide ample-sized drain holes.

Terrariums or Bottle Gardens

Many fern species are suited for terrariums or glass bottles because they flourish under the high humidity
within the container and the low light found indoors.

S  C

Suitable containers come in many shapes and sizes. Avoid thick, dark-colored glass containers that
reduce the light. Shallow containers without covers are unsuitable for most ferns because they dry too
quickly. If such containers are used, place them in a protected, humid place and frequently check for mois-
ture. It is best to select containers that are deep enough to provide for a few inches of soil and sufficient over-
head growing space for the plants. Fish aquariums, large brandy snifters, or containers with similar shapes
are satisfactory. Some ferns require less humidity than others and grow well in containers with large open-
ings. A container with a wide opening makes planting and maintaining the terrarium easier. Wide open-
ings can always be effectively reduced by covering with plastic wrap.
 G S F 101

Plastic containers designed as terrariums are utilitarian. They are lightweight, less likely to break com-
pared to glass, easily opened, have a ventilation hole, and are shaped to accommodate plants. Some prob-
lems, however, are that plastic containers scratch easily and might be unsuitable in certain decors.

S  P

Small plants in 5 to 10 cm (2–4 in.) pots are best for terrariums. Young plants of large species are easy
to obtain but will soon outgrow the container. Some slower growing large species may be suitable if others
are unavailable. Always start with healthy plants. Do not mix moisture-loving plants with those that like
drier conditions. The following recommended plants are small to medium-small species or have small cul-
tivars adapted for terrariums. Because some are slow growers and tender, they are not common nursery
items and must be purchased in specialty shops. Many ferns of dry habitats are small and suitable for use
in terrariums; see the listing under “Xerophytic Ferns” later in the chapter.
Adiantum capillus-veneris Hemionitis, all species
Adiantum hispidulum Lemmaphyllum microphyllum
Adiantum raddianum ‘Gracillimum’, Microgramma vacciniifolia
‘Micropinnulum’, ‘Pacific Maid’, or Nephrolepis exaltata ‘Elsevier’ and
‘Pacottii’ ‘Mini-ruffles’
Anogramma chaerophylla Polystichum tsus-simense
Asplenium cuspidatum Pteris cretica ‘Wilsonii’
Asplenium daucifolium Pteris ensiformis ‘Victoriae’
Asplenium oligophlebium Pteris multifida
Asplenium trichomanes Pyrrosia, small species
Bolbitis heteroclita Difformis Group Selaginella, small species
Camptosorus rhizophyllus Tectaria zeylanica
Davallia parvula Vittaria lineata
Diplazium tomitaroanum

P  T

For a 4 to 8 liter (1–2 gal.) container, place about 2.5 to 5 cm (1–2 in.) of perlite at the bottom for
drainage. If you have charcoal granules, mix some into the perlite or place a layer over the perlite. Place
about 5 cm (2 in.) or more of moist potting soil over the perlite and charcoal. There is wide variety in soil
preferences, ranging from sand to peat-soil mixes to uncut sphagnum. The potting soil is best disinfected
to kill any algae, bacteria, or fungi and should be done a few days before use. The microwave, boiling water,
a soak in Physan (Consan), or some other method is usually adequate to disinfect the soil (see “Sterilizing
Soils” in Chapter 5). Let the soil drain. At planting time, use evenly moistened soil, not one that is too wet
or too dry.
Plant the fern in the container at the same soil level as it was previously planted. If necessary, remove
some of the soil around the roots or from the terrarium to accommodate the roots of the new fern. Tall ferns
should be planted in the center or back of the terrarium with the smaller ferns, selaginellas, or mosses in
front. Firm the soil around the roots with a blunt-tipped tool. Long sticks and tongs may be needed to plant
ferns in narrow-mouth jars. Wire tools made from bent coat hangers are also serviceable. Long forceps are
useful for picking off dead fronds.
After planting, moisten the soil mix thoroughly with distilled water, but avoid overwatering. The ter-
rarium might need to be covered or partly covered, depending on the humidity and water needs of the
102 C 10

plants, the size of the opening, the amount of soil moisture, and the extent of damage to the roots during
planting. If too much condensation appears on the sides of a covered container, uncover it temporarily
or enlarge the opening in the plastic wrap to allow evaporation. For the first few days after planting, keep
the terrarium in a well-shaded place. In covered containers, a decrease in condensation is usually a signal
that watering may be needed. The effects of using commercial products, usually containing silicon or glyc-
erine, for reducing condensation is mixed. In any case, avoid getting these products on the plant or in
the soil.

C  P T

Caring for terrariums is relatively easy, but several factors need attention. Light needs to be ample, but
terrariums should never be placed in direct sunlight. Too little light can sometimes be a problem, which is
revealed by the plants’ developing long, thin, weak, light-green growth (etiolation). Another problem
related to lighting is when the plants all lean to one side in the direction of the light. To avoid this, turn the
terrarium occasionally or use overhead fluorescent lights.
The temperature must also be carefully considered. Growth will be faster in warm rooms and slower in
cold ones. The best growth usually occurs between 21 and 27°C (70–80°F). Never place terrariums in hot
places such as on the top of a television or beside a heater.
Water and humidity in terrariums also need to be monitored. As long as the soil stays moist (not soggy)
the terrarium does not need extra water. Much of the water in a closed or nearly closed terrarium will evap-
orate from the soil and foliage, condense on the glass, and fall back on the soil. Expect some condensation
on the glass. This recycling of water often provides sufficient moisture for the plants, and extra water might
not be needed for six weeks or more. To be safe, however, check the water weekly. Open terrariums need
to be watered more frequently. Apply distilled water when the soil starts to feel slightly less than moist—
never let the soil dry beyond this point. Apply water as a gentle sprinkle. Very fine-textured ferns and most
xerophytes should not be watered on their foliage. If you accidentally overwater, carefully tilt the terrarium
and blot up the excess with paper towels or other absorbent material, or push a long eyedropper or turkey
baster to the bottom and suck up the excess water. Leave the terrarium uncovered to allow the excess water
to evaporate.
If growing well, terrarium ferns do not need fertilizer. If they become off-color, they can be fertilized
sparingly with a dilute solution (half strength or less) of liquid fertilizer. Do not fertilize too much because
the plants will grow too large for the terrarium, in which case they will need to be trimmed back or replaced.
If a white crust of fertilizer salts develops on the soil, remove it and replace with fresh soil.

Maidenhair Ferns
Maidenhair ferns (Adiantum) have a fine billowy appearance and shiny black stalks that, because they
resemble hair, give the group its name. They are found in many parts of the world but are most abundant
in the American tropics. Most of the cultivated species are finely divided, and many have ruffles, fringes,
crests, and other types of fancy foliage.
The common species are easy to cultivate. The three most important factors are moist soil, good drain-
age, and humidity. Let them go to the dry side of moist before rewatering, but not to the point of wilting.
Adiantum is best grown outdoors or in greenhouses instead of indoors, unless a humid place is available such
as the kitchen or bathroom. Shriveling of new fronds is usually a sign of low humidity, especially if the soil
is moist. The other common cause of shriveling is poor root development, possibly the result of over-
watering, lack of nutrients, or poor drainage. Maidenhairs are particularly sensitive to being planted in
 G S F 103

pots that are too big for them, as the roots are poorly aerated in oversized pots. Most maidenhairs slow their
growth by the fall and take a rest period until spring. Water them lightly during this inactive period because
too much watering causes the rhizomes to rot.
Dividing, transplanting, or repotting is best done in spring before growth is renewed. A suitable soil is
one consisting of one part sand and one to two parts peat moss or leaf mold. A handful or two of coarser
material (ground bark or perlite) may be added to a 15 to 20 cm (6–8 in.) pot for better drainage. The delta
maidenhair (Adiantum raddianum), Venus’s hair (A. capillus-veneris), and fan maidenhair (A. tenerum)
thrive in calcium-rich soils. The use of balanced fertilizers usually satisfies this requirement, but lime, lime-
stone, or oyster shell may also be added. Fish emulsion or other suitable fertilizers are applied every three
weeks to ferns planted in sand and peat moss alone.
To maintain attractive plants, remove old or discolored fronds. Sometimes, especially with species that
form dense clumps (such as Adiantum capillus-veneris and A. raddianum), the fronds in the center will die.
To avoid the need for complete replanting, remove the dead part and refill the space with fresh soil. In frost-
free climates, all the spent and unsightly fronds may be removed between fall and spring, but in areas with
frosts, wait until spring to remove the dead fronds. In either case, damage to new growth can be avoided by
trimming off all the old fronds in spring just before the new growth uncoils. Although the plant looks bare
after this procedure, the alternative—having to remove the old fronds carefully without injuring the new
ones—is far more tedious.
Maidenhairs are plagued by several pests that must be controlled. Slugs and snails are a constant prob-
lem in some areas, and sowbugs and pillbugs can also cause damage. Aphids usually appear in spring and
can be difficult to eliminate. These pests can be controlled by applying the proper insecticide, but be
extremely careful in doing so because maidenhairs are easily burned by most insecticides. If it is unknown
whether an insecticide will burn the plant, apply it only on a test plant. (Also see Chapter 11 for more
information on controlling pests.)
Some maidenhairs are best grown under particular horticultural conditions. The American or five-
finger maidenhair (Adiantum pedatum) is easy to grow in cold-temperate climates but not in warm ones.
The Venus’s hair (A. capillus-veneris) and the rough maidenhair (A. hispidulum) are good for warm, tem-
perate, and subtropical climates. The delta maidenhair (A. raddianum) is more tender than the Venus’s hair
but possesses the advantage of having many attractive cultivars. The delta maidenhair and Venus’s hair are
sometimes slow to establish, but once in place they grow rapidly, particularly among cement sidewalks
and foundations where presumably more calcium is present. In tropical areas and greenhouses, you can eas-
ily grow A. tenerum and the coarser-leaflet forms such as the silver-dollar fern (A. peruvianum), diamond
maidenhair (A. trapeziforme), two-edged maidenhair (A. anceps), and large-leaved maidenhair (A. macro-
phyllum). The rough maidenhair tolerates lower humidity than most and is suitable as a houseplant.
Several species of maidenhair are especially attractive but difficult to grow. The kidney-shaped fronds
of Adiantum reniforme are handsome and eye-catching, but the plant is tender and hard to cultivate. The
trailing maidenhair (A. caudatum) is interesting for its long fronds that produce a new plantlet at their
whip-like tips. It is easy to grow in well-drained soil but must be cultivated in a warm greenhouse. Adiantum
×tracyi, a semi-hardy, evergreen species, is rarely cultivated because it reproduces only from divisions, not
spores. Large elegant fronds are produced by A. formosum, but because of its wide-creeping habit this spe-
cies is not grown in pots, and few people have greenhouses with beds large enough to accommodate it. It
will grow outdoors in warmer subtropical gardens.
An old favorite in cultivation is the Farley maidenhair (Adiantum tenerum ‘Farleyense’). It has grace-
fully arching fronds and many ruffled segments. It develops most luxuriantly when grown in warm green-
houses with daytime temperatures near 21°C (70°F). Another maidenhair that is smaller but just as lovely
is green petticoats (A. capillus-veneris ‘Imbricatum’). It is a difficult fern for beginners but needs less heat
than ‘Farleyense’. Adiantum raddianum ‘Pacific Maid’ has broadly overlapping segments and the light
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fluffy look of the preceding two selections, but it is a more erect, stiffer plant. It is good for beginners and
readily available in the trade.
With a greenhouse or similar means, many of the subtropical and tropical species can be grown with-
out difficulty. Keep in mind their particular needs for humidity and consistently moist but well-aerated soil.
For more details on cultivating Adiantum, see Chapter 13 and also Hoshizaki (1970a).

Staghorn Ferns
Staghorn ferns (Platycerium) are prized for their striking appearance and usefulness as decoration on walls
or tree trunks. In subtropical climates the common ones are grown outdoors year-round, and once estab-
lished, they require minimum care. Because they require only moderate humidity, staghorn ferns are suit-
able indoors, if provided with plenty of filtered light. Plants established and properly mounted need only
be hung in place and watered and fertilized occasionally.
If plants are watered with a hose, water them thoroughly. Small plants may be soaked in water for 10
to 15 minutes. Do not be tempted to water or sprinkle staghorns every time you water other ferns.
Staghorns need less water. More water will be required in a hot spell, less if the weather is cool and cloudy.
Staghorns are susceptible to overwatering by beginner growers because the outer surfaces of the base
fronds may feel dry even if the spongy inner layers are completely saturated with water. To test the mois-
ture, press your fingers firmly against the brown (not green) base fronds. If they are too wet, water will ooze
out. If they feel moist but no water oozes out, the moisture level is just right and the plant should not be
watered. Do not press against the freshly developed green base fronds because this can cause damage. Some
growers wait until the moss at the bottom of the planting is dry and crumbly before watering. Others wait
until the fertile fronds start to become limp. The weight of the plant also indicates relative wetness and dry-
ness. You should learn to determine whether a plant needs water by lifting it and judging its weight.
Inadequate watering usually results in slow or no growth. A sudden and severe lack of water causes
young fronds to wilt and the older ones to become limp and take on a grayish cast. The inner layers of base
fronds will be dry. Sometimes drought-like symptoms are caused by sowbugs, pillbugs, slugs, and snails eat-
ing the emerging roots. These pests are best seen at night.
When deciding where to hang a staghorn fern, be sure that it will not receive a constant drip of water
from overhanging baskets or from the eaves or roof. The symptoms of overwatering include no or poor
growth, absence of new base fronds, soggy base fronds, algae developing on the moss, and (at worst) pur-
plish to blackish decay spots on the base fronds.
Beginners should start with the easily grown species, such as Platycerium bifurcatum, P. hillii, and P.
veitchii. Slightly more difficult to grow and needing warmer conditions are P. willinckii and P. alcicorne.
Staghorn ferns can be propagated from the buds or young plantlets that form on their roots and some-
times their rhizomes. These plantlets are called “pups” by some gardeners. When the pups are big enough,
they can be separated from the parent plant. There is a greater chance of success if the pups are not removed
until they have several brown base fronds. Spring is a good time to remove them, or just before fresh shield
fronds grow out. Use a garden fork or knife to pry or cut beneath and around the oldest shield frond of the
pup (Figure 10.1). If possible, take a little of the base frond from the mother plant. Slip your fingers into
the cut and lift the pup from the mother plant. Plant the pup with the growing tip on the upper side.
The pups are usually mounted on either a tree or a board (Figures 10.2, 10.3). How you mount the pup
on a board depends on the total thickness of the base fronds. If more than 2.5 cm (1 in.) thick, the pup can
be mounted directly on a board and secured with wire (not copper) or stout strips of plastic. If the base
fronds are less than 2.5 cm (1 in.) thick, then place a 2.5 to 5 cm (1–2 in.) pad of sphagnum moss between
the base fronds and the board. If you are mounting the pup on a tree, do not wrap wire around the trunk,
 G S F 105

Figure 10.1. Removing a plantlet, or pup, from a

staghorn fern.

Figure 10.2. Planting a staghorn fern on a tree.

Figure 10.3. Planting a staghorn fern on a board,

front and side view.

which might girdle the tree and kill it. Instead, use plastic strips, nylon fishing line, discarded hosiery, or
string. If wire must be used, hammer small nails into the trunk and tie the wire to the nails.
It is also possible to grow a large staghorn in such a way that will not require remounting soon. Fill a
broken clay pot, wire basket, or shallow box with a loose, coarse humusy soil, covering or lining it with
sphagnum moss to keep the mixture from falling out (Figures 10.4, 10.5, 10.6). The moss can be kept in
place by covering it with chicken wire in which a hole has been cut for the plantlet. The surface of the plant,
particularly the buds, should be level with or slightly above the surface of the sphagnum moss. Plants so
mounted will have the advantage of more rooting medium, but the disadvantage of being susceptible to
106 C 10

overwatering. Recently mounted pups should be kept moist but well drained. When foliage no longer
appears wilted, the plants can be placed in more exposed places. The wires or strips of plastic will soon be
covered with new base fronds. Remounting established plants is the same as for mounting pups, except that
old base fronds might need to be trimmed away. To grow large symmetrical staghorns, keep one plant and
remove all the pups that appear.

Figure 10.4. Planting a staghorn fern in a wire

basket, front and side view.

Figure 10.5. Planting a staghorn fern in a pot

to be hung, front and side view.

Figure 10.6. Planting a staghorn fern in a pot,

front and side view.

Fertilize the plants about every three weeks, although less so during cooler months. Use liquid fertiliz-
ers because solid fertilizers, such as granules, powders, or pearl types, do not dissolve readily and, if caught
between the base fronds, can burn the tissue or promote infection by mold.
Once fungi or bacteria infect a leaf they are difficult to control. Several control methods are available,
but they are not always effective. Some growers report limited success by removing the decaying spot before
it spreads. They cut at least 1.5 cm (0.5 in.) beyond the decayed area. Others soak the plant in fungicides
(see Chapter 11). Some growers reduce the watering drastically in hope of killing the infection before
killing the plant. Prevention is better than an uncertain cure. Do not overwater, and use only thoroughly
dissolved fertilizer.
Aside from requiring a bright airy place and sufficient water, the common species of staghorn ferns pose
no special problems and will reward you with their striking and unusual foliage. If you have experience
growing the common species, the less common ones are also worth trying, although they can be more dif-
ficult. Most of these other species are large plants that require more space. See the references listed under
Platycerium in Chapter 13 for more details.
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Davallia
The squirrel’s-foot fern, Davallia mariesii var. stenolepis (D. trichomanoides of trade), is the most commonly
cultivated Davallia. Its finely divided triangular fronds attract attention, as do its long, chaffy rhizomes that
can creep over an entire basket. Davallias are adaptable to a wide variety of situations. Although usually
grown in hanging baskets or pots, they are also suitable as ground covers and even over rocks if given
enough soil to secure a footing. They can also flourish indoors in sufficient light. Because they are epi-
phytes with firm-textured leaves, davallias can usually withstand the slightly drier air indoors better than
most other ferns. They should not be kept too moist. If fertilized a few times during the growing season,
Davallia plants will multiply readily from their creeping rhizomes.
Most davallias grow seasonally, shedding their fronds once a year between fall and spring and then
entering a period of slow growth. Sometime later a flush of new growth fully refoliates the plant. The exact
time of foliage shedding, and the length of time before new ones appear, varies with the species and climate.
Fronds persist longer on luxuriantly growing plants. In warmer climates such as southern California and
Florida, the deciduous habit of most species is hardly noticeable because new fronds appear before the old
ones have withered. The fronds sometimes turn a beautiful yellow before they wither and drop. If old
fronds have not fallen by the time new ones appear, they should be removed to provide room for the new
ones. The self-shedding of spent fronds is a decided advantage in dealing with high-hanging baskets.
Baskets that become covered with layers of rhizomes should be replanted. In mild climates, the best time
to do this is the fall or spring. Tear the clumps apart, saving those clumps with the most growing tips, and
then discard the dead or old parts of the reusable clumps. Davallia baskets should be lined with uncut
sphagnum moss and filled with a humusy, loose planting mix. Keep the basket moist but well drained, and
set it in a humid place until the plants are established. Further details on planting in baskets are provided
in Chapter 7.
The tender species Davallia fejeensis and D. solida are frequently seen in greenhouses and conservato-
ries. In warm-temperate climates the following species will grow and withstand light frost and short peri-
ods of freezing temperatures: D. canariensis, D. mariesii var. mariesii and var. stenolepis, D. pyxidata, and D.
tyermannii.

The Boston Fern and Its Relatives

The Boston fern and its relatives belong to the genus Nephrolepis and are among the most popular ferns for
home and garden. The original Boston fern was the Nephrolepis exaltata cultivar ‘Bostoniensis’, from which
numerous cultivars have been selected. Loosely speaking, the common name “Boston fern” has come to be
used to describe the many cultivars of N. exaltata. In warm climates these ferns are particularly favored as
indoor or outdoor pot or basket plants. They are mostly epiphytes and prefer bright airy places and good
soil aeration. Nephrolepis species as a group do not like soggy soil and so should be kept on the drier side of
moist, especially when they are not actively growing in cool weather. The fronds are fuller if provided with
ample humidity and the plants regularly fertilized.
The most common causes of poor growth in these ferns are under-watering or overwatering, low
humidity, and (for some species) cool winter temperatures. The well-known Boston fern cultivar ‘Fluffy
Ruffles’ seems to tolerate slightly cooler temperatures than the other cultivars of Nephrolepis exaltata. Some
of the fancy cultivars quickly succumb to poor cultural conditions and can soon yellow and shed their
leaflets. Even with good growing conditions, however, a few lower leaflets on older fronds will yellow and
should be trimmed off. Nephrolepis produces long, slender, scaly, string-like stems (stolons) from its cen-
108 C 10

tral stem. These can be removed or wound around the plant and tucked inside the pot to keep a tidy look.
Repotting Nephrolepis is best done before active growth, usually in the spring. The taller, older plants
may need to be set deeper into the soil than they were before, but be sure not to bury their stem tips. Cut
away most of the old leaf bases from the stem. Avoid planting them in oversized pots; they tolerate being
slightly root bound. Use an epiphytic soil mix or a mixture of half sand and half peat moss (see Chapter 5).
Most species of Nephrolepis can be propagated by sowing spores. All can be propagated by divisions or
by layering the stolons, which produce new plants when they touch the soil. Planting in benches or shal-
low, broad containers gives the stolons room to spread in all directions and take root. The resulting plants
can be separated and potted when they have two or more fronds. The Boston ferns usually do not produce
spores and must be propagated by rooting the stolons or by meristem culture. Provide bright airy places for
their culture. Commercial greenhouses use as much as 3000 to 3500 foot-candles of light or 73% shade
cloth (see Chapter 8 for details on propagation).
The numerous cultivars of Nephrolepis exaltata offer a wide variety of sizes, shapes, and textures. Some
cultivars produce fronds as long as 1.5 m (5 ft.), whereas others produce fronds that are extremely broad or
narrow. These and other fancy-foliage forms, especially the finely divided ones, often grow slowly.
Boston fern cultivars with finely divided or congested fronds must be watered carefully. Their foliage
cannot support water applied directly from above, which tends to mat and break the fronds. If allowed to
remain on the leaves, the water causes the leaflets to have a dark, water-soaked appearance and can result in
loss, disfigurement, or yellowing of the foliage. If water mats the foliage, gently shake it off or blot it up.
Cultivars with densely clustered leaves need to be groomed regularly. Remove the dead or declining
fronds and unwanted stolons. This will allow more air and light to reach the younger growth below and give
the plant a more attractive appearance.
Besides Boston fern, the tuber sword fern (Nephrolepis cordifolia) is easy to grow. It is probably the most
cold tolerant of all sword ferns, being able to withstand short periods near 0°C (32°F). In subtropical and
tropical climates it is commonly planted as an outdoor border plant—a purpose to which it is well suited
because of its usually stiff, erect fronds. It can even be clipped like a hedge. The tuber sword fern has sev-
eral cultivars, the most popular being ‘Tesselata’ (‘Plumosa’), which has divided leaflets. ‘Tesselata’ grows
more slowly than the species form. Even slower is ‘Duffii’, which has rounded leaflets and is best grown
under glass (except in warm, humid climates where it can be grown outdoors).
The scurfy sword fern (Nephrolepis hirsutula) is rank and generally unattractive except in warm, humid
places. It grows poorly where nighttime temperatures dip below 16°C (60°F) and has no advantage over the
other species except that it grows easily from spores.
Nephrolepis species or cultivars with long, drooping fronds are difficult to find commercially because
growers dislike the difficulties involved with growing and shipping them. They must be displayed in bas-
kets or elevated containers to be attractive. Some long-frond types sold by specialty nurseries include
Nephrolepis falcata, N. pendula, and N. exaltata and its cultivars ‘Gretnae’ and ‘Rooseveltii’.

Polypodium and Relatives

The genus Polypodium and its relatives (the family Polypodiaceae) form a large and diverse group that
varies greatly in cultural requirements. Most are epiphytes and grow best in baskets or hanging containers
with good drainage and aeration. Others can be grown in the ground if provided with good drainage and
not overwatered. Most cultivated species are tender or semi-tender and therefore must be grown in green-
houses in most parts of the United States. They need ample light but should never be placed in direct sun.
They prefer a loose, humusy soil mix. Creeping types are best planted in a moss-lined basket so that they
can root in the moss as they grow. (For a monographic treatment of the group, see Roos 1985.)
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The rabbit’s-foot ferns or golden polypodies (Phlebodium) are among the most popular polypodiums
grown in the United States. Phlebodium aureum and P. pseudoaureum include several cultivars, some of
which have attractive bluish gray foliage. Once established, the golden polypodies grow rapidly if given
plenty of light. They should be watered when the soil is nearly dry. Some selections tolerate cold better than
others. Variants with broader, thinner lobes and sori in two or more rows on each side of the segment need
warmer temperatures than those with the narrower, firm lobes and sori in one to nearly two rows on each
side of the segment. The bluish green P. aureum ‘Mandaianum’ has thin, ruffled lobes. If unblemished
fronds are desired, grow it indoors or in sheltered places.
Knight’s polypody (Goniophlebium subauriculatum ‘Knightiae’) is another favorite basket fern. The
long, pinnate fronds are fringed and arch down to form a beautiful hanging plant. Established plants tol-
erate irregular watering. Outdoors in warm areas, the old fronds are shed in spring but are soon replaced
with new ones. The cut fronds last a long time in flower arrangements.
Another polypody fern, Microsorum pustulatum, is not widely cultivated but should be. In England it
is a hardy fern, and it should be tried in cooler areas of the United States. Microsorum pustulatum is medium-
sized with deeply divided blades resembling those of the East Indian polypody (M. scolopendria).
Microsorum punctatum ‘Grandiceps’ makes a handsome houseplant. Its thick, glossy leaves, ruffled
and irregularly forked or notched, withstand the drier air of the indoors admirably well. Because it grows
slowly, it seldom needs repotting.
A coarse, large, durable member of the polypody group is Aglaomorpha coronans. Although it grows
slowly, the species tolerates periods of dryness and aridity once established. Remove the old fronds as they
start to decline, because otherwise the individual leaflets will drop and scatter about.
Most of the small polypodium relatives that have simple, entire fronds are tender species and must be
grown in humid greenhouses or terrariums. Because they are small, they are not particularly showy and
therefore seldom seen in collections. One of these small species, Microgramma squamulosa, is attractive
due to its dark veins.
The temperate species of Polypodium vary in their cultural needs. The more common ones are planted
in baskets or among rocks. The licorice fern (Polypodium glycyrrhiza) often volunteers in baskets lined with
moss collected from western Canada and the western United States. A choice species is the leathery polypody
(P. scouleri), which grows along the coast in the Northwest. Its fronds are small- to medium-sized, deeply
lobed, thick, and glossy. It grows slowly and is suitable for pots and baskets in cool humid climates. The res-
urrection fern (P. polypodioides), a common epiphyte in the southeastern United States, is a small fern that
curls its leaves during drought and unrolls them when wet weather returns; it is difficult to establish.

Tree Ferns
Whether planted individually or in groups, tree ferns are sure to be noticed. They can provide a back-
ground for other plants, and their trunks can be used to display small epiphytic orchids, tillandsias, and
ferns.
Most ferns that bear fronds at the top of a tall trunk are called tree ferns by gardeners, although such
growth forms are found in unrelated ferns such as Sadleria, Blechnum, and Ctenitis. Most of the ferns in the
Dicksonia and Cyathea families (Dicksoniaceae and Cyatheaceae) are tree-like and are the ones typically
referred to as tree ferns by botanists. They range from 1 to 15 m (3–50 ft.) tall. In many species, the girth
of the trunk is increased by producing a thick layer of hard, dark, tangled roots—the root mantle. This layer
forms a substrate on which epiphytic plants grow.
Ferns of the Cyathea and Dicksonia families, as well as some tree-fern-like members of Blechnum, are
mostly native to cool, moist mountainous regions of the tropics (a few species occur in warm-temperate
110 C 10

areas). There are about 700 species in these two families, but probably fewer than 50 have been tried in cul-
tivation in Europe and the United States.
Temperature tolerances of the tree ferns are the first consideration in selecting them for cultivation.
Some species tolerate occasional frost and mild freezes, whereas others are intolerant of prolonged periods
at temperatures of 15°C (59°F). Nighttime temperatures are also an important consideration because many
species prefer cooler nighttime than daytime temperatures. Those tree ferns native to lowland tropics where
nighttime and daytime temperatures are relatively constant do well outdoors in tropical gardens. The many
species native to Australia and New Zealand grow well outdoors in subtropical areas, and some (such as
Dicksonia antarctica) grow in mild-temperate areas. In cold-temperate areas, tree ferns must be moved
indoors during the winter.
Tree ferns should be planted in well-drained soil. Add peat and sand (or perlite) to heavy soils, or
replace parts of it. Windy sites must be avoided. Although some species can grow in direct sun, they are
more luxuriant in shade. Water from overhead leaves or hanging baskets should not be allowed to drip into
the crowns of tree ferns. Provide plenty of overhead space for development. Tree-fern diseases continue to
be reported on wild and cultivated species. Rhizoctonia and tip blight (see Chapter 11) infect tree ferns, and
there may be other diseases as well.
Although some tree ferns can be shortened by severing their trunks and replanting them, the risk
involved in doing this is high. Some species root from trunks more readily than others. The Australian tree
fern of the trade, Sphaeropteris cooperi, does not root from severed trunks, whereas Cibotium glaucum, C.
menziesii, and Dicksonia antarctica root fairly well.
Cutting the trunk is best done before the active growing period and before new fronds emerge (usually
in the spring). After cutting, dust the cut ends with a fungicide to protect against rot, then plant in a pot
with a well-drained soil mix, and water sparingly to discourage rotting. Avoid watering the crowns, which
are fleshy and prone to rot. The number of remaining fronds can be reduced by half to prevent too much
water being lost from the trunk. If the trunks are too fleshy and not in danger of drying, it might be best to
wait before planting until a protective “skin” develops over the cut tissue. To encourage this, put the trunk
in a cool, humid place and keep it from drying. After the skin develops, plant the trunks about one-fourth
to one-third their length into moist soil, making sure to keep the trunks shaded and humid. When new
fronds emerge, the plant can be watered more. A few months before cutting the trunk, wrap a layer of
moist uncut sphagnum moss above the cutting point. This encourages roots to form, which then grow into
the moss. The moss layer should be thick enough that it will not dry quickly. Ample humidity and warmth
hasten root formation.
The Australian-native Cooper’s tree fern (Sphaeropteris cooperi) is the most frequently grown tree fern
in the United States. It grows in sun or shade in coastal, central, and southern California, in southern
Florida, and in conservatories in cooler areas. It tolerates frost and short periods of freezing temperatures,
but its foliage might die back. On older plants the fronds tend to spread more horizontally than on younger
plants and may be 6 m (20 ft.) across. The trunks are relatively slender, and cultivated plants seldom form
extensive aerial roots enveloping the stem. Because these roots are usually absent, the trunks are weak and
can snap off in strong winds. Therefore Cooper’s tree fern is best grown under tall trees where their crowns
have room to spread but are protected from wind. The lower fronds will droop or drop several times a
year. This foliage drooping is usually accompanied or preceded by a heavy discharge of spores. For the
sake of neatness, drooping fronds close to the trunk can be removed, but if left on the plant they eventu-
ally dry and fall off on their own. Under optimum conditions this fern may grow 30 cm (1 ft.) per year when
young, and the cultivar S. cooperi ‘Brentwood’ grows even more rapidly. The fronds and trunk of Cooper’s
tree fern are covered with many small scales that can irritate the skin and especially the eyes. Washing the
skin removes the scales and itching. Goggles should be worn to protect the eyes when working with this
fern.
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The Tasmanian dicksonia (Dicksonia antarctica) is the second most frequently cultivated tree fern. It
is semi-hardy to semi-tender and has been reported to endure −7°C (20°F), or a bit lower for short periods,
as well as snow on its fronds. It prefers an average winter temperature of about 9°C (48°F). Its fronds are
shorter, narrower, stiffer, harsher, and more numerous than those of Sphaeropteris cooperi. The crown of
foliage reaches about 4 m (13 ft.) across and has a tufted appearance on the stout trunk. It grows more slowly
than Cooper’s tree fern—about one-third as fast. The Tasmanian dicksonia prefers climates influenced by
the ocean or cool and humid conditions. Dicksonia fibrosa and D. squarrosa are seldom cultivated because
they are more difficult to grow.
The Hawaiian tree fern (Cibotium glaucum) is seen now and then in outdoor plantings in southern Cali-
fornia. Bare-root trunks are imported from the Hawaiian Islands, and these can be rooted by planting
them about one-third their length in a well-drained soil mix. Spore-grown plants form more symmetrical
crowns but are infrequently found in the trade. The fronds of the Hawaiian tree fern tend to ascend and arch
more gracefully than those of Cooper’s tree fern. Their leaf bases and trunks are covered with silky, yellowish
tan hairs, unlike the chaffy scales of Cooper’s tree fern.
Occasionally, trunks of Cibotium menziesii and Sadleria cyatheoides are found in shipments of C. glau-
cum trunks due to their similar appearance. Cibotium menziesii is called the “man fern” by the Hawaiians,
possibly because its trunk is covered with stiff black hairs. It requires the same cultural conditions as the
Hawaiian tree fern. Sadleria cyatheoides tends to be more difficult to grow than the other Hawaiian species
and is sensitive to having its roots disturbed. Bare-root trunks of this species may be distinguished from oth-
ers by the presence of scales instead of hairs. Sadleria and Hawaiian cibotiums tend to grow more slowly
than Sphaeropteris cooperi.
The Mexican tree fern (Cibotium schiedei) was once widely sold as a houseplant, but it grows too big
for the average house. It hardly forms a trunk but produces offshoots freely around its base. Its many light
green fronds droop gracefully to give the plant a soft effect.
The silver tree fern (Alsophila tricolor) and the black tree fern (Sphaeropteris medullaris) need more pro-
tection from wind and sun than other tree-fern species to look their best.
Cultivated species that resemble tree ferns include Blechnum gibbum and B. brasiliense. These ferns are
medium-sized and form erect trunk-like stems. They are semi-tender to tender. The plant sometimes called
the American tree fern (Ctenitis sloanei) is native to southern Florida and might form a short, erect stem,
giving it a tree-fern–like appearance. Diplazium esculentum could be considered a small tree fern because it
produces an upright stem, but it is only 2 to 4 cm (0.75–1.5 in.) in diameter. This Asian species has become
naturalized in Louisiana and Florida. It is cultivated in Florida and California.

Xerophytic Ferns
Xerophytes are plants adapted to dry climates. Many xerophytic ferns grow in the arid southwestern United
States. In more humid regions of the country, they are found in exposed places such as on tree branches,
rocks, or cliffs.
Many fern genera have xerophytic species. Genera that are mostly xerophytes include Actiniopteris,
Astrolepis, Cheilanthes, Doryopteris, Notholaena, Pellaea, and Pityrogramma. Hairs, scales, or powder cover
the foliage of many xerophytic ferns to help retain moisture. These coverings can be white, silver, or yellow
and make the fern especially attractive. Ferns that are adapted to dry climates are often small and therefore
suitable for rock or trough gardens. Some xerophytic species are challenging to grow and may require sev-
eral attempts before the proper conditions are found.
In cultivation, xerophytic ferns grow best in bright but indirect sunlight, except along cloudy coastal
areas where they may be planted in full sun. They prefer a well-drained soil kept on the drier side of moist.
112 C 10

Most grow well in an evenly moist soil or one that does not rapidly fluctuate between wet and dry. In gen-
eral, the fronds should be kept free of water droplets resulting from condensation and overhead watering,
although fog does not seem to bother some xerophytic ferns.
Soil mixes range from simple to complex, but all must be well drained. A simple mix consists of one part
peat moss or leaf mold and one to two parts gravelly sand. A more complex mix consists of one to two parts
decomposed granite about 6 mm (0.2 in.) in diameter, two parts compost (leaf mold or fir bark), one part
perlite (#2 size), and one part sand (#20 size). Adding horticultural charcoal to the mix is optional, as is plac-
ing a top dressing of half-inch fine gravel on the soil surface after planting.
Xerophytes should be planted with their crowns slightly above the soil. If using pots, make sure that the
soil level is close to the rim in order to reduce the amount of water that could be caught in the pot during
watering. A process known as double potting maintains uniform soil moisture over a longer time. The
fern is planted in a porous clay pot, which in turn is planted in a larger clay pot, usually 5 to 7.5 cm (2–3
in.) wider than the first. The same soil mix is used in both pots. Xerophytes can be successfully grown in ter-
rariums if the soil moisture is carefully monitored and the humidity not excessive. Soil water evaporates
slowly in a terrarium, thus maintaining a more constant moisture level. In such a protected environment
fronds may develop more fully than in nature.
Outdoors, xerophytes are often planted in trough gardens (see “Trough Gardens” earlier in the chap-
ter), among rocks, or on well-drained sites (also see “Rock Garden and Wall Ferns” in Chapter 9). Xero-
phytic ferns are extremely sensitive to overwatering and can die if overwatered only a few times, and so they
should be planted away from plants that require more water. Whether in the ground or in pots, the plants
should be watered early in the morning so that any water settling on the fronds will evaporate during the day.
Most xerophytic ferns go dormant during the summer in their native habitats. In cultivation, however,
dormancy might not occur. In addition, many xerophytic ferns grow more slowly or go dormant as cool
weather approaches. Dormant or slow-growing plants need less water.

Filmy Ferns
Filmy ferns have been grown successfully in botanical gardens and private collections and were popularly
grown in Wardian cases in England during the Victorian era. These mostly small ferns are distinctive by
their membranous leaves usually one cell thick between the veins. Because of their thinness, the plants
need high humidity and shade. They also require good drainage. In nature most grow as epiphytes or ter-
restrially, and the group reaches its greatest development in tropical montane forests that are covered by
clouds most of the day and have an abundance of mosses on tree trunks and branches. The filmy-fern fam-
ily (Hymenophyllaceae) also includes the kidney fern (Trichomanes reniforme), an eye-catching species cul-
tivated in New Zealand and Australia but rarely in the United States. The elegant Prince-of-Wales plume
(Leptopteris superba) is not a filmy fern even though it has membranous leaves; however, it does require the
same cultural conditions as filmy ferns. It is rarely grown in the United States but is grown in Australia, New
Zealand, and England.
Filmy ferns from warm-temperate areas grow well in temperatures from 4 to 21°C (40–70°F), although
they also tolerate short periods of slightly lower or higher temperatures. Climates where temperatures are
consistently higher than 27°C (80°F) for days are unsuitable. Most tropical species come from cooler mon-
tane forests and grow best in temperatures ranging from 16 to 24°C (60–75°F). In favorable climates filmy
ferns can be grown in terrariums or bottles provided with uncut sphagnum moss or a well-drained potting
soil. In warm climates, air conditioners are needed to keep the temperature low.
Special chambers or houses, often located in a shady corner of a greenhouse, can be built to hold filmy
ferns. An intermittent misting system should be installed and, depending on the climate, possibly some
 G S F 113

means of cooling. The chamber’s top and one side should be made out of glass or translucent plastic. If the
chamber is situated in an area with abundant light, only the top needs to be glass. Line the walls with
uncut sphagnum moss held in place by nylon fishing line or chicken wire (do not use copper wire). Plant
the ferns in pots or on pieces of tree-fern trunk and hang them on the moss, or plant them directly into the
wall of moss. Humidity should be kept near 100% by misting several times a day or by automatic humid-
ifiers. Chamber walls can also be made out of hollow concrete blocks filled with sphagnum moss or suit-
able excelsiors. The filled blocks are placed with the open end up, and a pipe set on the top of the block
drips water into the moss to keep the walls moist. Ferns are hung in pots against the wall. Some means of
draining the water away from the enclosure should be provided. The quality of water can also pose prob-
lems. If the salt content is high, salt-free water will have to be used. Farrar (1968) achieved good growth
of filmy ferns in chambers at 100% relative humidity with temperatures between 18 and 24°C (65–75°F)
and indirect natural light at or below 300 foot-candles. The ferns were given a fine mist for 15 minutes
every hour.

Fern Allies
Fern allies are the whisk ferns (Psilotum), ground pines (Lycopodium), spike mosses (Selaginella), and horse-
tails (Equisetum). Like ferns, these plants reproduce by dispersing spores, but none has leaves like ferns.
Instead, the leaves of the fern allies are small, inconspicuous, and often scale-like with only a single vein.
Water clovers (Marsilea), mosquito ferns (Azolla), and water spangles (Salvinia) are ferns but might be
mistaken for fern allies because of their unfernlike appearance.

P  W F

A handful of stems from a Psilotum plant resembles a small whisk broom, hence the name whisk fern.
The green stems seem leafless but actually bear scattered, small, scale-like leaves. The upper branches may
have rounded, three-lobed sporangia (spore cases). The plants grow on trees, especially in the leaf axils of
old palm leaves, or in rock pockets in tropical or subtropical areas. They grow easily in greenhouses with
strong light and respond well to bone meal and general fertilizers. Propagation is easy by division of the rhi-
zome clump, although transplants are sometimes slow to establish. In greenhouses the plants volunteer read-
ily from spores, but growing by the intentional sowing of the spores is difficult. Laboratory studies indicate
that it can take as long as 3 to 18 months for spore germination. The gametophytes (comparable to the pro-
thalli of ferns) are about 4 mm (0.2 in.) long, cylindrical, and grow beneath the soil.

L  G P

Lycopodiums are terrestrial or epiphytic species with erect, creeping, trailing, or drooping stems. They
range from a few inches tall to several feet long. The stems are amply covered with scale-like leaves, some of
which have small roundish to bean-shaped spore cases (sporangia) at their base.
The temperate species of Lycopodium are particularly difficult to grow from transplants because they do
not readily re-establish themselves in a new location. Transplants have been reported to establish success-
fully by planting in oak-leaf mold and applying aluminum sulfate to maintain acidity (amounts not speci-
fied). The plants were kept constantly moist. Transplants of Lycopodium selago have been successfully grown
in uncut sphagnum moss watered with distilled water or water treated by reverse osmosis.
A few tropical epiphytic lycopodiums, such as Lycopodium phlegmaria and L. phlegmarioides, are grown
in moss-lined hanging baskets or anchored on boards with a pad of uncut moss. Boards of tree-fern fiber are
114 C 10

used in Florida. Warmth, humidity, and constantly moist soil with good drainage are required for most epi-
phytic species.
Tropical epiphytic lycopodiums are usually propagated by divisions or layering because they are diffi-
cult to start from cuttings. Species such as Lycopodium phlegmaria can be propagated by layering or by
anchoring the stem tips in vermiculite or sand and waiting for rootlets to develop. Rooting should take place
in six to eight months, after which the plantlets can be separated from the mother plant.
Besides layering, cuttings are another method of propagating, although it tends to be less successful. The
usual procedure is to take 8 cm (3 in.) long pieces from the tips of stems bearing sterile leaves (that is, those not
bearing sporangia). The leaves are removed from the lower third, and the cutting is rooted in clean potting soil
mixed with four to five parts sand. Be careful not to overwater the potting soil, otherwise rot can easily result.
Provide the cuttings with only enough moisture and humidity to prevent rotting. The temperate Lycopodium
selago has been successfully propagated from cuttings rooted in uncut sphagnum moss and watered with dis-
tilled water. Whether propagating by layering or cutting, some bottom heat will hasten the process.
Growing lycopodiums from spores is rarely done, but the process would be essentially the same as for
selaginellas, discussed in the following section.

S  S M

Selaginellas are more widely cultivated than lycopodiums. The name spike moss refers to the spike-like
cluster of fertile leaves at the branch tips. About 700 species of Selaginella exist worldwide, some of which
are adapted to deserts (where they are often nestled between rocks) and others are found on rain-forest
floors. Almost all the species are terrestrial. Some resemble lycopodiums, but the cultivated species are gen-
erally softer, mossier, or fernier compared to the stiffer, harder-textured lycopodiums. The two genera also
differ by their spores: Selaginella produces two kinds of spores (separate male and female), whereas
Lycopodium produces only one (bisexual).
The resurrection plant (Selaginella lepidophylla) is one of the few desert species that are grown. It
requires excellent drainage and less humidity than other selaginellas. Most of the temperate species are
small plants, moss-like in growth and useful as ground covers. Selaginella apoda often invades moist, shaded
lawns in the eastern United States. In humid warm-temperate areas, S. kraussiana can also be invasive.
Most tropical species thrive in warm, humid greenhouses, growing luxuriantly at temperatures of 21°C
(70°F) or more. They tend not to be too fussy about their soil mix as long as it retains adequate moisture
and is well drained. They respond well to regular applications of fertilizer. The erect forms (such as S.
umbrosa) are used as taller ground covers or in pots.
Most selaginellas can be propagated by cuttings or layering (pegging the foliage to the soil and waiting
for it to root). All can be propagated by dividing the clump. Species that root all along the underside of the
stem are easily propagated by about 4 cm (1.5 in.) long cuttings placed in clean builder’s sand and finely
misted three or four times a day until rooted. Species that root only near the base of the stem are more dif-
ficult to root by cuttings. All species may be propagated sexually.
To propagate by spores, select recently matured fertile spikes or leaf clusters that have unopened spo-
rangia—the sporangia appear as plump, rounded, yellowish structures at the base of each fertile leaf. Cut
the spikes into short lengths and scatter them over a clean, firmed, well-drained mix of one part garden soil
and four to five parts sand. (Other mixes such as uncut sphagnum moss have also been used.) Some of this
finely screened soil should be sprinkled over the spike pieces to anchor them. The container or flat is then
watered gently and covered with a piece of glass or plastic wrap and kept at 21°C (70°F). About nine
months later, small plants should appear.
For this method it is important to use mature fertile branches containing male and female spores. If the
maturity of the fertile branches is difficult to determine, pick and sow them at intervals to increase the
 G S F 115

probability that both kinds of spores will be present. The methods for hybridizing selaginellas are somewhat
involved due to the presence of male and female spores. For details, see Webster (1979).

E, H,  S R

Horsetails (Equisetum) are reed-like plants with jointed, hollow stems that range from a few centime-
ters to about 6 m (20 ft.) tall. Because they grow near or in wet areas and their stems contain silica, pioneers
used them for scrubbing pots, hence the name scouring rush. They are useful in wet parts of the garden such
as pools, or as novelties in pots. Some species are deciduous. They are well suited to full sun or bright light
and wet soil or standing water. Some species spread aggressively from underground stems, sending up iso-
lated erect stems. The winter scouring-rush (Equisetum hyemale) is a particularly aggressive spreader. It is
difficult to eradicate once established and should be planted in sunken containers to limit its spread.
Horsetails can be propagated by rhizome divisions, cuttings, or spores. Division works best with large
rhizome pieces that have new shoots or buds attached. The divisions can be planted in various kinds of soil
but do best in soil that is low in organic matter, about 25% (wet organic soils spoil or sour readily). To prop-
agate from cuttings, cut the stem into pieces containing at least three joints. Plant the pieces upright with
about one-half to two-thirds of their length in sand. Keep the cuttings moist until lateral shoots appear; if
drying is a problem, pots may be placed in a shallow saucer of water. Guarding against poor aeration and
sour soil is particularly important during the rooting process. Some species can be propagated by bending
the stems of the mother plant and anchoring them in place in the soil or in a pot of water until well rooted.
The spores of equisetums are contained in cones at the stem tips. In some species the cones appear on
green stems, and in other species they are produced only in the spring on ephemeral, nongreen, erect
shoots. The hexagonal plates on the cone separate when the spores are ready to be shed, and this is the time
to collect the spores for propagation. The spores are green and remain viable for only a few days. They
should be sown soon after collecting. The procedures for sowing and growing are the same as for ferns (see
Chapter 8). The germinated spores will develop into disk- or cushion-shaped gametophytes typically about
3 mm (0.1 in.) long. When transplanting the gametophytes, keep them in small clumps to ensure a mix of
male and female gametophytes. (Gametophytes are at first either male or female, but females can later
become bisexual.) Young sporelings are transplanted by the same procedures as for ferns.

Ferns for Shows and Exhibits

G S F
The beautiful ferns seen in shows and exhibits can be grown by any gardener willing to give a little time
on a regular schedule. Consistency and staying alert for sudden changes in weather or growing conditions
are especially important. Large, showy plants need plenty of space to produce symmetrical growth, but
small- to medium-sized plants can be grown in limited space. Although large ferns make spectacular show
subjects, the smaller species are often more interesting and too frequently overlooked by amateurs. Partic-
ularly for show plants, you need to provide the following conditions:
1. Ample growing space protected from temperature extremes and wind. Crowded plants
compete for light and usually produce lopsided foliage. Space lacking adequate protection
from the weather is of no avail because the fronds will become disfigured or damaged.
2. Light on all sides. Evenly distributed light will help produce symmetrical growth. If the
light is stronger on one side of the fern, turn the fern each week to encourage symmetrical
growth. For large baskets, attaching a swivel to the wire-hang will assist in turning the plant.
116 C 10

3. Frequent, dilute applications of fertilizers. Make frequent applications of fertilizer at one-

quarter strength rather than monthly applications at full strength. Apply weak solutions
every one to two weeks during periods of active growth. If water spots (salt deposits) appear
on the leaves, apply the fertilizer solution to the soil, not the foliage.
4. Groom plants as necessary. Grooming encourages the development of perfect fronds.
Remove yellowed, damaged, and misshapen fronds, as well as those that crowd perfectly
formed and positioned fronds.
5. Produce one large plant. Most show entries are based on one plant per container or entry,
unless you enter under a multiple-plant category. Offshoots produced in close proximity to
the crown of the main plant should be removed if one large plant is desired. Use a sharp
knife to cut the rhizome, but pull the roots of the two plants apart with a fork or your
fingers. Tearing the roots apart will leave more roots on the offshoot and the parent than
cutting them will. Ferns with creeping, branched, mat-forming rhizomes are considered a
single plant and need not be reduced to one creeping rhizome.
6. Be alert to unfavorable growing conditions such as pest damage, sudden dry weather, heavy
rain, strong winds, or other unexpected changes. Cope with such situations immediately.
Spots and blemishes are often due to plant stresses that are avoidable. Do not procrastinate!
7. Before the show, groom the plants carefully. Remove water spots and dust from the foliage.
Commercial polishes may be used on coarse, leathery ferns, but they give an artificial shine
that is objectionable to some judges. Before show time, clean the pots of algae and salt. If it
is customary to cover the soil in the pot for entry, do so with materials that will not detract
from the effect of the plant. To prevent damage during transport, wrap the plant in a cone
of newspaper. Ferns with spreading fronds might need to have their fronds tied back to
avoid damage. Soft strips of fabric or women’s hosiery can be used to tie back the fronds.
Do not transport the plants in an open vehicle unless they are well protected. Do not leave
them in a closed vehicle on a hot day. Anchor them securely in place for transport.
8. Register plants carefully. Follow the registration procedures for the show and enter the
plants in the proper category. Be certain the exhibitor’s ticket is secured to the plants.
Before you leave the plants, check that they have enough water, and make arrangements to
have them watered during the show if this is not done by the show authorities.

F F
Some hardy deciduous ferns that are to be exhibited out of season may be forced into early growth for
the show. Others, such as Woodsia glabella and Polystichum braunii, apparently do not take kindly to forc-
ing. Some will develop deformed fronds, and others will collapse soon after producing fronds.
Thurston (1939) successfully forced many ferns from New England and mid-Atlantic states for a flower
show in March. The ferns were planted in pots or flats in fall or earlier. They were placed in outdoor frames
and covered with lath. As the weather cooled, the ferns were gradually covered with leaves. The ferns were
permitted to freeze in November but under the protection of layers of leaves, branches, lath, glass sash, and
straw so as to be easy to dig out at the end of December. Plants were then removed from the frames and
placed in a dark shed to thaw for four days at temperatures between 4 and 10°C (40–50°F). Afterward they
were moved to a greenhouse and given light and exposed to temperatures of 13°C (55°F) during the day and
4 to 10°C (40–50°F) during the night. After five days, the temperature was raised to 21°C (70°F) during
the day and 10°C (50°F) at night. The water, heat, light, humidity, and ventilation for each species was judi-
ciously adjusted in the greenhouse. The water given was at 21°C (70°F) and was not permitted to touch the
foliage of most of the ferns; the exceptions were Camptosorus and others that like humidity, which were
 G S F 117

misted to create relative humidity of 80%. Hardening off of the ferns began in the second week of Febru-
ary, and temperatures dropped to 13 to 16°C (55–60°F) during the day and to 7 to 10°C (45–50°F) at
night. The ferns were ready for the show by the first week of March. Thurston observed that forced ferns
produced good growth for the show, but the continued new growth was not as vigorous as in nonforced
ferns.
Most semi-hardy and more tender ferns can be kept in show condition by keeping them in warm
greenhouses and protecting the fronds from damage. Little is known about forcing tender species that are
deciduous.
11 Troubles with Growing Ferns

Recognizing Cultural Troubles

The earlier you detect and correct a problem, the better. As soon as your fern starts to decline, consider its
basic conditions. Have you been giving it too much or too little water? Has the weather turned too cool for
the fern? Have you forgotten to fertilize, or have you fertilized incorrectly? Is the light sufficient? Is the plant
entering its rest period? Most troubles are caused by poor cultural conditions, unless, of course, there are
pests. Common symptoms and causes of illnesses are outlined here. (See also Chapter 4 for more details on
providing proper cultural conditions.)

S: S ;   ;    

  .
Common Cause Possible Solution
Plant is entering its normal rest period or Check species for deciduous habit.
dormancy.
Plant is overwatered, especially if the lower Decrease water and/or increase drainage.
leaves yellow and the soil is wet most of Use coarser soil mixes. Use clay instead of
the time. plastic pots. If the plant is overpotted,
plant it in a smaller pot.
Temperatures are too low. Growth improves when the plant is moved
to a warmer location, or as the cool
weather turns warmer.
Too much light, especially if the plant is in Reduce light.
direct sun or very bright indirect light.
Not enough fertilizer, especially if the new Fertilize.
leaves are small and slow to grow.
Plant is root-bound, especially if it wilts Repot.
between waterings and roots can be seen
to have filled the pot.

118
 T  G F 119

S: F  , ,  ;  

  .
Common Cause Possible Solution
Air is too dry. Plant may be in draft or wind, Increase humidity or move the plant to a
especially if damage is along the edges of more protected place.
the frond and appears after a windy or
dry period.
Temperatures are too high. If this is the case, Provide more shade and ventilation. Lower
the damaged spots will be found where the temperature in greenhouses.
the leaf is most exposed to the heat source.
The frond’s margins may also be burned.
Damage appears after or during a heat
wave.
Soil was or is too dry. Parts of a wilted frond may recover, but
other parts may be permanently damaged.
Trim off dead parts, water well, then keep
the plant moist, not wet.
Too much light. Reduce light.
Insecticide, fungicide, or fertilizer burn, Trim away badly damaged parts.
especially if treatment has been applied
around the plant.
Water-soaked tissue, resulting in brown Trim away badly damaged parts. Do not
spots or areas. permit water to sit on fronds.
Salt damage, especially if a thick white crust Use fertilizers sparingly. Leach the soil
accumulates on the pot or soil and the thoroughly. Water less frequently but
water is known to contain high amounts more thoroughly.
of salt.
Frost or freeze damage, especially if the Trim back dead parts.
weather has been cold.

S: E     ;  .

Common Cause Possible Solution
Dryness when fronds were developing. Provide more soil moisture and humidity.
Damaged fronds cannot be restored.
Earlier gas, insect, or insecticide damage. Remove toxic gas or insects; change
insecticides. Damaged fronds cannot be
restored.

S: F        .

Common Cause Possible Solution
Root damage caused by drying or Water well, then keep plant moist but not
overfertilization. wet.
120 C 11

Root damage caused by overwatering or If roots have deteriorated, trim dead parts
noxious materials. The soil may smell bad. away, replant the rhizome into fresh soil,
keep moist but not wet, and hope for the
best.

S: F   ,  -.

Common Cause Possible Solution
Not enough light. Give more light.

S: F   ,   ,    
.
Common Cause Possible Solution
Too much humidity. Reduce humidity.
Too much nitrogen fertilizer. Reduce nitrogen fertilizer.

Recognizing Pests and Diseases

If you cannot find any problems with the basic cultural conditions, inspect the plant for signs of insects,
fungi, bacteria, or other pests. A hand lens is helpful. Nursery growers generally find that cultural problems
are uniformly distributed in nursery plots and tend to produce more regular patterns on plants, whereas
pests and diseases are spotty and localized and spread in a gradient from the original infection, making more
irregular patterns on the plant crop. Home growers, not having the uniformity of a large crop to check, must
look at individual plants with the eyes of a detective.
For signs of pest or disease damage, check the roots for abnormalities, particularly decay. Look carefully
on the undersides of the fronds for specks, dots, or unusual-looking structures—do not confuse these with
the clusters of spore cases or sori. Most sori are regularly placed on the underside of the frond, whereas
insects are unevenly scattered and often favor nesting in the angle of the veins. Sucking insects can cause
poor growth, puckered foliage, distortion, bleached spots, and discoloration (discoloration is also caused by
nematodes). The most common sucking insects on ferns are aphids, scale insects, mealybugs, and thrips.
Biting insects and pests chew parts of the frond away. Some of the common examples are grasshoppers,
caterpillars, cutworms, pillbugs, sowbugs, slugs, and snails. Their presence may often be indicated by frass
on the foliage.
Fungal or bacterial infections usually accompany overwatering or excessive humidity. They usually
produce symptoms such as tissue that is rotted, slimy, or water-soaked. Sometimes colonies of the organ-
isms can be seen as tufts or mats of mold, sooty spots, and circular spots of dead tissue, particularly those
with small dots or concentric patterns. The organisms can be found anywhere on the plant. Do not over-
look the base of the stipe near the soil; infections sometimes start there. Fungi can cause sudden wilting of
the plant, marginal browning of fronds, and distortion of emerging fronds. These symptoms, however, are
difficult to distinguish from culturally caused troubles.
 T  G F 121

Alternatives to Chemical Controls

Some pesticides and fungicides are toxic to humans, pets, and many beneficial organisms. Alternatives to
the use of these chemicals are being actively researched by scientists. Some of the alternatives are cultural
and physical methods, biological control, and use of nontoxic or low-toxic controls. These methods can
control a specific pest with one or more applications. Repeating the application as needed will be satisfac-
tory for most cases. Attaining long-term control with few repeat applications and still avoiding the use of
toxic chemicals requires integrating and analyzing a range of information. This approach is called Integrated
Pest Management (IPM). The identity of the pest, its life cycle, the cultural conditions that can thwart them
or favor their enemies, use of predators or parasites, and when, how much, and under what conditions to
release these predators are types of information that must be known and assessed for each situation. In its
fullest sense, IPM is a holistic approach. More information on IPM can be obtained from local libraries,
county agricultural offices, and organizations or journals devoted to the subject.

B C
Biological control uses beneficial organisms to kill pests. Familiar control organisms include the pray-
ing mantis, lady bugs, lacewings, ant lions, spiders, lizards, toads, and birds. Less well known are bacteria,
fungi, nematodes, parasitic mites, and wasps. All these controls are harmless to humans and pets. New
control organisms for specific pests are constantly being offered, and inquires should be made to county
agricultural extension services for restrictions and suppliers.
To work effectively, control organisms must be released before infestation becomes severe and well after
application of any long-lasting insecticides. Results take time, and close monitoring might be required to
determine effectiveness. Several releases may be necessary. Control rather than annihilation is the objective.
If control organisms are microscopic (such as bacteria), they can be formulated as a dust or liquid solu-
tion. Insects or animals are released as eggs, juveniles, or adults, depending on the species. Severe infesta-
tions are best treated first with a quickly biodegradable pesticide before the release of the controls. Because
healthy plants resist pests better than unhealthy ones, it is essential that you maintain proper fertilization
and optimum cultural conditions. You can also encourage naturally occurring controls such as spiders,
lizards, and toads.
To maintain beneficial organisms after they have controlled the pest might require advanced plan-
ning, unless you intend to make new releases. Ask the supplier how to provide for the control organisms
until the next pest attack or season. It might require alternative food sources and suitable habitats and pro-
tection from certain pesticides.

S, O,  F A S  A

Sprays of soaps, neem extracts, oils, and fatty acids are safe for humans and less harmful to the envi-
ronment. Insecticidal soaps are effective on soft-bodied pests. Their main drawback is that they can dam-
age the plant or leave behind an unsightly white residue on the leaves. It might be necessary to test spray to
determine the fern’s sensitivity. Plants should not be sprayed when temperatures are above 32°C (90°F). If
plants are sensitive, they could wilt, spot, or burn (particularly along the margin), symptoms which may be
seen within hours or days. Soft young foliage is most susceptible. If possible, delay spraying until young tis-
sue matures. If wilting is noted within a few hours, rinse immediately with water.
Insect sprays made from liquid dishwashing detergent and water are effective on soft-bodied insects but
usually must be reapplied periodically. The recommended proportion of dishwasher detergent is between
122 C 11

0.4 and 2.0%, or about 5 to 10 ml per 480 ml (1–2 tsp. per 2 cups) of water. The more detergent, the more
effective the spray and the more risk of burning the fronds. One test using a 3% solution found no dam-
age to maidenhair ferns (Adiantum). Studies indicate that soap used with acephate (Orthene), pyrethroids,
and chlorohydrocarbon pesticides increases the effectiveness of the spray. Commercial insecticidal soaps are
also available, but commonly used brands are not recommended for most ferns.
Alcohol also kills insects. It must be applied laboriously with a cotton swab by touching it to the insect.
Generally concentrations of 35 to 40% are used.

P-D I
Although plant-derived insecticides are rapidly biodegradable and generally safer to use than synthetic
chemicals, they are not selective and could harm beneficial insects and fish. Sprays and dusts are available,
and protective gear should be used to prevent inhalation. Several new plant-derived insecticides, such as
neem (Margosan), seem promising. Old standbys include rotenone, best if ingested by the pest, and
pyrethrin (pyrethrum), which is best for adult insects. These pesticides kill ants, aphids, caterpillars, cut-
worms, leaf hoppers, mites, thrips, whiteflies, earwigs, mealybugs, millipedes, and sowbugs. They are inef-
fective against adult scale insects.
Nicotine compounds are also plant-derived insecticides. Although they degrade quickly, they are toxic
to mammals and therefore unavailable to home growers. Commercial growers use them against pests such
as aphids, young scale, mealybugs, thrips, and fungus-gnat larvae.
Several synthetic plant-derived insecticides (as pyrethroids modified from pyrethrin) are on the mar-
ket. They have the advantages of being more effective on specific pests or longer lasting. Some are men-
tioned under the pest that they best control.

I G R

Insect growth regulators (IGR) interrupt the normal development of pests and thus prevent them from
maturing or laying eggs. Most target certain insects and do not harm the beneficial ones. Kinoprene (Enstar
5E) affects aphids, mealybugs (ground and foliar), scale insects, whiteflies, and fungus gnats. It is, however,
known to damage maidenhair fern. See the further discussion under aphids. Some growers reportedly have
obtained satisfactory results by combining IGR and synthetic pyrethrin.

P
Pheromones are substances given off by plants or animals that cause specific behavioral responses in
other individuals of the same or different species. Some pheromones are sex attractants used to lure insects
into traps. Most pheromones available in the trade are used to control moths, flies, and beetles whose
young feed on plants and may only incidentally feed on ferns. Pheromones are nontoxic to humans, pets,
and the environment.

P C: T  B

Sticky traps are frequently used in pest control. Flypaper, sticky paste, and sticky colored paper can be
used to trap insects and detect early infestations. For sticky colored paper, blue attracts thrips, whereas yel-
low attracts aphids, whiteflies, leaf hoppers, moths, and fungus gnats better than white paper. Yellow plas-
tic lids or containers coated with SAE 90 grade oil are also effective traps. Nowadays the effectiveness of traps
is often enhanced by adding pheromones and baits with pest-attracting odors. Insects are also attracted by
 T  G F 123

ultraviolet light, and such lights are used in “bug zappers” with electrically charged grids that kill adult fly-
ing insects on contact. Ultraviolet light can be useful to control moths that in their caterpillar or grub stage
eat plants.
Mulches may help reduce surface-crawling pests, and a layer of ashes often discourages slugs and snails,
but use sparingly because too many ashes could harm the soil. A commercially available diatomaceous
earth dust is reportedly effective against soft-bodied insects. The microscopic needles of silica in the dust
puncture the insect’s guts after ingestion. It should not be applied when beneficial insects are active. Report-
edly, this talc-like powder can be handled safely without gloves.
Spraying with water dislodges some pests and gives a certain measure of control. Adding a few drops of
liquid detergent to the water helps loosen pests attached to stems or leaves. Picking off insects by hand or
dabbing them with an alcohol- or oil-soaked cotton swab is labor intensive. The best time to look for slugs,
snails, and cutworms is at night, especially when the ground is moist. Providing good sanitation will reduce
hiding and breeding places of pests.

Chemical Pesticides and Fungicides

The Federal and State Environmental Protection Agencies regulate and proscribe the label contents and
directives for all pesticides. In effect, the label is the law. The availability of and methods of applying pesti-
cides are regulated by governmental agencies. In some cases the regulations restrict the sale of certain chem-
icals to licensed agricultural growers, in others they restrict the use of pesticides to certain regions of the
United States. The following pages mention some of these restricted pesticides and fungicides along with
their more widely available alternatives. Government regulations frequently change, and producers may
modify their products as well. Therefore, readers should obtain updated information from their local
county agricultural service.
The chemicals of choice are given under each pest discussed in this chapter. Further details on most of
the insecticides may be found under “Aphids.” Also see Appendix IV for cross-references to chemical and
trade names and notes. No endorsement is intended by the inclusion of trade names, nor is criticism
implied of products not mentioned. Although the advice contained has been carefully compiled, the authors
cannot guarantee results. The reader acts on his or her own responsibility.

P P  F D

If you must use pesticides or fungicides, observe the following cautions. These will help you better
control the pest or disease, avoid or reduce damage to plants, and minimize the dangers to humans, pets,
and the environment.
1. Read the manufacturer’s label and directions carefully. Follow recommended follow-up
treatments. The label is the law!
2. New products and brands should be used with care. Note that brands vary according to
the grade of the pesticides used, the percentage of active ingredient, the formulation, and
whether they are used as a liquid, powder, granule, or aerosol.
3. Pesticides have a shelf life, and some lose their effectiveness in two to three years. Always
buy and use fresh pesticides. Store as directed on the label.
4. Use dust or sprays made from wettable powders instead of emulsifiable concentrates, if a
choice is available. Granules cause less damage than emulsions, which contain oils that
damage ferns.
124 C 11

5. Test the products on a few plants before applying them to other plants. Damage usually
appears after one day to two weeks and is evident as wilting, yellowing, spotting, marginal
or surface burning, or abnormal growth.
6. Proper dosage is given on the label, but only a few fern species have been tested at the
dosages listed by the manufacturer. The recommended dosage will often damage
susceptible ferns. Reducing the dosage, however, violates the label law, even if a lower
dosage is effective and does not damage the fern.
7. Sprays and dusts are most effective when thinly and thoroughly applied to all surfaces of
the foliage. Fine mist sprays are the best. Water at the soil level after spraying or dusting so
that the residue is not washed off the foliage.
8. Aerosols or smoke generators should be used only in greenhouses or tightly sealed plastic
houses.
9. Spray, dust, or apply granules when the leaves are dry to avoid or minimize damage.
10. Spray early in the day so that plants can dry rapidly.
11. Spray when the air temperature will stay below 29°C (85°F) for at least two hours after
spraying.
12. Move houseplants outdoors for treatment if possible.
13. The healthier the plant, the less likely the possibility of spray damage. Plants most easily
damaged are those that are root bound, in need of water, or very young. Any damage that
does occur might only affect the existing foliage, and any new growth may be unharmed if
the roots and rhizomes have not been damaged.
14. Assume that all pesticides and fungicides are toxic. To reduce the possibility of damage to
your (and others’) health, proper procedures, protective clothes, and equipment must be
used, and re-entry periods observed.
15. For the commercial grower, the suitability of a pesticide or fungicide for large-scale use
should be checked with local county agents, state experiment stations, extension specialists,
or the manufacturer’s technical representatives. These specialists can determine the
suitability of a specific pesticide for local soils, temperatures, moisture conditions, cultural
practices, and dosage rates.
Systemic pesticides are those taken into the plant tissue and are effective in controlling the pest or dis-
ease from within the plant. A drench is a solution applied by sprinkling it over the soil.

Insect and Other Pests

A
Ants feed on honeydew, a sweet secretion deposited by pests such as scale insects, aphids, and mealy-
bugs. The ants not only protect these pests, but also disperse them!
P  C. If nests are found, pour enough boiling water to reach the main
chambers.
Chemical controls of choice are chlorpyrifos (Dursban), diazinon, or malathion. Carbaryl (Sevin),
bendiocarb (Dycarb, Turcam), and pyrethrin are other choices. Several other chemicals are used for ant con-
trol, but their availability is restricted in some states.
 T  G F 125

A
Aphids are small, soft-bodied insects that are red, green, yellow, orange, or black. The fern aphid is black
with whitish legs. Aphids weaken plants by sucking juices and causing distortion and stunting of foliage.
They also produce a secretion called honeydew that is sipped by ants. Because the ants encourage and
protect the aphids, they must also be controlled (see above). Some aphids are easy to kill but difficult to
control because reinfestation often occurs from nearby areas. The fern aphid is common on new growth,
particularly in the spring (Figure 11.1).

Figure 11.1. Aphids (8×).

P  C. For nonchemical control, see the various measures discussed under
“Alternatives to Chemical Controls,” earlier in the chapter.
Acephate (Orthene) is a systemic, broad-spectrum insecticide. Fern growers prefer the 75% soluble
powder used at the manufacturer’s rate of 5 ml (1 tsp.) to 3.8 liters (1 gal.) of water. Because damage has
been reported on Blechnum gibbum, Phlebodium aureum, Polystichum, and Pteris ensiformis, you should do
a sample test before using acephate powder on your ferns. Emulsions are more easily available and should
also be tested before wider use. The manufacturer recommends an acephate emulsion of 9.4% used at 30
ml per 3.8 liters of water (2 tbsp. per gal.) for aphids and 45 ml per 3.8 liters of water (3 tbsp. per gal.) for
mealybugs and scale insects on ornamentals in general. This dosage, however, could burn some ferns, so test
first. Orthene aerosol might also burn some ferns.
Carbaryl (Sevin) is a broad-spectrum insecticide that can be used around edible plants. Carbaryl is
registered for aphid use, but only for the apple aphid and rose aphid. For the fern aphid, other insecticides
126 C 11

are more effective, though carbaryl is also useful for other pests that attack ferns. It is available as a dust,
emulsion, or wettable powder. Emulsions burn sensitive ferns.
Diazinon is another broad-spectrum insecticide. Diazinon emulsions of 25% are recommended by
the manufacturer to be used at 10 ml to 3.8 liters of water (2 tsp. per gal.) for foliage plants, but not for cer-
tain ferns. Dosages reduced one-quarter to one-half the recommended concentration gave adequate con-
trol with little damage, but such dosages violate label laws. Wettable diazinon powder of 50% at 15 ml per
3.8 liters of water (1 tbsp. per gal.) is recommended for foliage plants; however, the manufacturer also
reports that maidenhairs and other ferns are burned at this concentration. In all cases, sample test on a few
plants before wider use.
Dimethoate (Cygon 2E), a systemic, broad-spectrum insecticide, is effective for three weeks after appli-
cation. When used as a drench to minimize foliage burn, Cygon 2E (23.4%) is used at 5 ml to 3.8 liters of
water (1 tsp. per gal.) for foliage plants in general. As a spray for foliage plants, the recommended dosage is
7.5 ml per 3.8 liters of water (1½ tsp. per gal.). Treatments of 2.5 to 5 ml per 3.8 liters of water (½–1 tsp.
per gal.) have been successful in tests on ferns. This product has restrictions and is unavailable in certain
states.
Disulfoton (Di-syston) granules applied to the soil provide systemic protection against aphids and
other insects. It is often included in rose-care products and is effective against scale insects, red spider-
mite, and thrips.
Fluvalinate (Mavrik Aquaflow) is a synthetic pyrethrin-like chemical. The water-based formulation
makes it safe for plants. Recommended dosage ranges from 1.3 to 3.3 ml per 3.8 liters of water (¼–⅔ tsp.
per gal.) for foliage plants.
Imidacloprid (Marathon) is a broad-spectrum systemic insecticide. It comes in wettable powder or
granular form. The active ingredient does not readily move in the soil, therefore drenches or granules must
be applied to the root zone to be effective. It reportedly remains active in the soil for 8 to 12 weeks after one
application. Some ferns reportedly do not respond to granular forms even if generously applied. It is not
available to home growers.
Kinoprene (Enstar 5E) is safe for Boston fern (Nephrolepis exaltata ‘Bostoniensis’) and its cultivar
‘Fluffy Ruffles’ and for Pteris cretica ‘Parkeri’ when used at the manufacturer’s recommended rate. It severely
damages maidenhair fern (Adiantum), however, and its effect on other ferns is unknown.
Malathion is a broad-spectrum insecticide available as a wettable powder but more commonly as a
dust or emulsion. The latter is most commonly used, and the manufacturer’s recommended dosage for 50%
malathion concentration on foliage plants in general is 10 ml per 3.8 liters of water (2 tsp. per gal.). This
dosage severely burns or injures a variety of ferns, although commercial fern growers encountered little or
no damage when the dosage was reduced to 2.5 ml per 3.8 liters of water (½ tsp. per 1 gal.), though this
reduction violates the label laws.
Nicotine sulfate is no longer available in the United States, although a nicotine alkaloid aerosol is avail-
able to commercial growers. It is highly toxic.
Oxydemetron-methyl (Metasystox-R) is a systemic insecticide. The 25% emulsion is used at 7.5 to 10
ml per 3.8 liters of water (1½–2 tsp. per gal.) for foliage plants. It is reportedly safe for maidenhair and other
ferns when applied to the soil.
Oxamyl (Vydate) controls aphids and a variety of insects, nematodes, and mites; however, it is available
only to commercial growers.
Pyrethrin kills aphids and comes in many formulations. Synthetic pyrethrins (pyrethroids) are also
available.
Resmethrin kills aphids and other pests and comes as a spray or aerosol. The aerosol form is favored by
some commercial growers.
Soap sprays are discussed under “Alternatives to Chemical Controls.”
 T  G F 127

E
Earwigs are reddish brown insects about 13 mm (0.5 in.) in length, with prominent pinchers at the end
of their tails. They ordinarily eat decaying vegetation, but they can damage ferns by nibbling at tender
new growth. During the day earwigs hide under flower pots and plant debris. They forage at night.
P  C. Keep a clean culture. Trap earwigs during the night in a roll of damp
newspaper, and destroy it in the morning.
Propoxur (Baygon) bait is the insecticide of choice. Baits or dusts of carbaryl (Sevin), chlorpyrifos
(Dursban), and diazinon also control earwigs. Scatter the bait where earwigs congregate but not directly on
the fern foliage.

F G
Fungus gnats resemble small mosquitoes less than 3 mm (0.1 in.) long. With a hand lens the feelers
(antennae) are seen to be longer than the head, and the veins of the wing form a Y-shaped pattern. These
features are absent on shore flies, a similar gnat. Fungus gnats frequent damp places and soils with high
organic matter. When disturbed, the adults often run rapidly on the plant or soil before taking flight.
Both shore flies and fungus gnat adults can spread disease, but the fungus gnat causes the most dam-
age when in its larval stage. In greenhouses and enclosed places they eat roots, root hairs, stems, and pro-
thalli. Shore-fly larvae are maggot-like and eat detritus, whereas fungus gnats eat living plant tissue. Fun-
gus-gnat larvae are small, whitish, translucent, legless worms with black heads. They can be found beneath
the soil level or in the plant tissue being eaten. Affected plants grow slowly, lack vigor, may wilt, show yel-
low leaves, and have rotted roots. Prothalli appear wilted or have small but noticeable holes where the lar-
vae have fed. The greatest damage is seen during plant growth stages in which there are few roots and the
larvae have fed on them.
P  C. Keep a clean culture. Remove weeds and plant debris, and take steps
to avoid excessive moisture in or on the soil and surrounding area. To control algae, which often flourish
in unclean cultures and attract fungus gnats, use clean or new flats and pots (see “Algae, Mosses, and Liv-
erworts”). On spore pots and flats covered with glass, seal any space between the glass cover and pot or flat,
preferably with plastic foam strips instead of cotton. If feasible, reduce the organic content in the soil mix.
Ammonium nitrate used as a drench at three-quarters the normal fertilizer concentration works for larval
control, and acts as a fertilizer as well.
If drenches or sprays are used, remove surrounding debris before application. Avoid irrigation for one
day after application. Drench, dust, or spray the plant, the soil, under benches, and any places where the
gnat or its larvae might be. Yellow sticky cards are useful for trapping adult gnats and for monitoring the
effectiveness of other control measures.
Biological controls include Bacillus thuringiensis and other organisms (see “Alternatives to Chemical
Controls”). Bacillus thuringiensis H-14 (Gnatrol) kills larvae. The drench works within 24 hours after
ingestion. Ferns are unharmed by the recommended dosages of 5 to 10 ml per 3.8 liters of water (1–2 tsp.
per gal.) for light infestations, or 20 to 40 ml (4–8 tsp.) for heavy infestations. A nematode that kills the lar-
val stages of the fungus gnat is sold as Exhibit.
Chemical treatments of choice include diazinon (also used in a capsule form called Knox-out), kino-
prene (Enstar 5E), oxamyl (Vydate 10% granules for the larvae and Vydate 2L spray for the larvae and
adults), and resmethrin (a synthetic pyrethrin). One percent resmethrin aerosol (PT1200 Resmethrin) did
not damage Boston ferns, and growers reported no damage to the commonly grown species of Pteris and
Pellaea. Resmethrin spray (SBP-1382-2EC) is recommended for foliage plants at 5 ml per 3.8 liters of
water (1 tsp. per gal.); its toxicity to ferns is unknown. Less effective, but also registered as a control for fun-
128 C 11

gus gnats, are dimethoate (Cygon 2E), carbaryl (Sevin), and malathion. See “Aphids” for details and
precautions.
Although unregistered against fungus gnat larvae, fluvalinate (Mavrik Aquaflow) seems a promising
control. Others products that are reportedly effective on fungus gnats include Fenoxycarb (an insect growth
regulator), Citation, Orthene, and Dycarb.

M
Several species of mealybugs attack fern fronds and (less frequently) roots. These small insects are char-
acterized by a white mealy look (Figure 11.2). They weaken a plant by feeding on its sap. Ants are attracted
to the mealybugs’ honeydew secretion.

Figure 11.2. Mealybug (18×).

P  C. Adult mealybugs are more difficult to control than the immature
stages (crawlers). Therefore you should inspect plants frequently for infestation and employ control meas-
ures promptly. Control the ants (as discussed earlier); also see “Alternatives to Chemical Controls.” For root
mealybugs, use the spray as a drench.
Acephate (Orthene), diazinon, and malathion are the insecticides of choice. Also used are dimethoate
(Cygon 2E), imidacloprid (Marathon), kinoprene (Enstar 5E), oxydemetron-methyl (Metasystox-R), and
oxamyl (Vydate). If you have a choice, use systemics. See “Aphids” for more details.
 T  G F 129

M
Millipedes generally feed on decaying plant debris but also eat healthy fern foliage. They appear worm-
like but have many legs. The garden species is reddish brown, about an inch long, and emits a sweet, pun-
gent odor when crushed (reminiscent of toasted almonds). Millipedes may be confused with predatory
centipedes, which differ by having only one instead of two pairs of legs per body segment, and by generally
moving faster. Millipedes are most active at night. During the day look for them under flowerpots and
debris.
P  C. Keep a clean culture. Remove debris.
Propoxur (Baygon) bait is the pesticide of choice. Carbaryl (Sevin) dust or a spray from 50% wettable
powder, at 30 ml per 3.8 liters (2 tbsp. per gal.), will also control millipedes. Spray into cracks, under con-
tainers, and in other hiding places, thoroughly wetting these surfaces. Avoid spraying on fern foliage.
Pesticides such as acephate (Orthene), chlorpyrifos (Dursban), diazinon, pyrethrin, and rotenone also
kill millipedes. Be sure to follow the directions on the label, and avoid getting the insecticide on foliage.

M, R S-M, S M, T-S M

Although many species of mites attack plants, red spider-mites are the most pernicious. They are about
the size of a grain of salt and vary in color from red to green to cream. The adults have eight legs. When
shaken onto white paper, the mites can be seen as tiny crawling specks. They leave a delicate webbing on
the leaves. Plants damaged by mites decline in vigor and show a yellow, stippled pattern or larger yellow-
ish areas that can eventually turn brown. By the time damage is noticed, the mites could have moved else-
where. Since mites are attracted by dusty dry conditions, they seldom attack ferns.
P  C. Do not let the plants suffer from moisture stress and low humidity.
Spray with water to wash off mites and accumulated dust and increase the humidity. Spraying is necessary
only if the mites persist in large numbers. Spray thoroughly on the lower surface of the foliage. Soap sprays
and biological controls are effective (see “Alternatives to Chemical Controls”).
Chemical controls of choice include dicofol (Kelthane), which is relatively harmless to ferns.
Dienochlor (Pentac, using the 50% wettable powder spray) is safe on Boston fern (Nephrolepis exaltata
‘Bostoniensis’) and its cultivar ‘Fluffy Ruffles’. Other pesticides that may be used on mites include acephate
(Orthene), diazinon, disulfoton (Di-syston), fluvalinate (Mavrik Aquaflow), oxydemetron-methyl (Meta-
systox-R), and oxamyl (Vydate). See “Aphids” for details and precautions. Carabyl (Sevin) and malathion
seem to cause subsequent outbreaks of mite infestations; therefore, carabyl is often combined with dicofol
in sprays. Sulfur dust is ineffective and might burn the plant.

N  N
Leaf nematode blight is caused by minute worms (Aphelenchoides fragariae). The nematodes form red-
dish brown or black spots that are typically sharply marked on the leaf. They typically feed on the leaf tis-
sue between the veins, seldom across the veins. Therefore the spots have an angular shape. On bird’s-nest
fern (Asplenium nidus) the discoloration is limited to tissue between the parallel veins. In other species and
on young ferns, the affected area may be more irregular due to the pattern of the veins. Nematodes can be
seen by placing a bit of infected tissue in a drop of water, then lightly mincing and examining it under about
20× magnification. With good light, you can see the characteristic thrashing movement of these nearly
transparent, slender worms (about 0.5 mm long). Leaf nematodes thrive under cool moist conditions.
They travel in a film of water or in drops from splashed water. Besides weakening and discoloring the
plant, nematodes spread disease. Root-knot nematodes have not been reported to attack ferns.
130 C 11

It is unlikely that nematodes spread through fern spores, unless fragments of infected leaf tissue are sown
with the spores. Rapid drying of the leaf tissue prevents nematodes from entering their rest stage. Other-
wise, they can remain alive for a year or more.
Certain ferns are more susceptible to leaf nematode damage than others. These include Asplenium
(A. bulbiferum and A. nidus in particular), Blechnum, Pteris, Tectaria, and Woodwardia (W. fimbriata and
W. radicans in particular). Also susceptible are Adiantum, Cyrtomium falcatum, Dicksonia antarctica, Diplaz-
ium dilatatum, Diplazium proliferum, Dryopteris filix-mas, Dryopteris hondoensis, Nephrolepis cordifolia,
Polypodium, Polystichum munitum, Rumohra adiantiformis, Sphaeropteris cooperi, and Thelypteris.
P  C. Prevention is easier than control. Start with disease-free stock and use
only clean flats or pots. Beware of soil, materials, or tools that might harbor nematodes. Disinfect con-
taminated materials or tools, pasteurize suspected soil with steam at 82°C (180°F) for about 30 minutes,
or treat with a soil fumigant. Because nematodes thrive in debris, discolored leaves should be discarded to
where they will not reinfect other plants. Remove infected leaves and burn them. Avoid overhead watering
and other conditions that wet the leaves. Remove weeds that might harbor nematodes. For the fern hob-
byist with only a few infected plants, the best course is to dispose of them properly and enact preventative
measures to protect against further spread. Bird’s-nest fern (Asplenium nidus) has been successfully freed of
nematodes by immersing the plant in hot water at 46°C (115°F) for 10 to 15 minutes.
If the infection is from the ground, the plants should be grown in pots on raised benches. Do not allow
the infected soil to be splashed on benches, pots, or foliage. If planting the ferns in the ground, allow the
soil to remain fallow and weed-free for a season or so. This will decrease but not eliminate the nematode
population. Drying the soil also reduces the nematode population. Live mature marigolds (Tagetes patula)
will release from their roots a chemical (ozone) that kills nematodes. This control practice is only effective
if many marigolds are planted.
The chemicals used to control nematodes are generally unavailable to home growers because of their
toxicity to humans and pets and their ability to remain in the soil or plants for a long time if improperly
applied. Nematicides used on ferns include ethoprop (Mocap), oxamyl (Vydate), and phenamiphos
(Nemacur). All of these have restrictions on their use. With the constant changes in labeling and laws, it is
best to contact local agricultural agencies for current information. For more details, see Krusberg (1992) and
anonymous (1986).

P  S

Psocids or book lice are small, pale insects that scurry around on damp soil or decaying vegetation.
Springtails (Collembola) are similar in appearance but jump when disturbed. Both thrive in warm moist
environments. Neither harms ferns directly, but they can spread fungi and bacteria.
P  C. If you encounter problems with a spreading disease, spray the soil
with a soap spray or with the chemicals listed for controlling aphids.

S I
Adult scale insects live in a stationary, wax-like shell under which they suck plant juices. Extra juices are
secreted as honeydew, which attracts ants and serves as the medium for the growth of a blackish mold.
Several species of scale insects feed on ferns. The most common is the brown soft-scale. It has black or
brown spots on its back. Young scale insects (crawlers) continually emerge from the mother scale for about
two months and grow to full size in two more months (Figure 11.3).
P  C. Adult scale insects are difficult to control, and therefore frequent
inspection and enacting controls at an early stage are important. Although biological controls such as kino-
 T  G F 131

Figure 11.3. Scale insects (10×). Note the very

young scale insects emerging from the older scale
at the bottom.

prene (Enstar 5E) are available, natural predators can be encouraged by spraying the plants with water to
keep off dust. Soaps may be effective against crawlers. Adults can be controlled by dabbing with alcohol-
or oil-soaked swabs. Chemicals of choice to control the adults are diazinon, dimethoate (Cygon 2E), imi-
dacloprid (Marathon), malathion, acephate (Orthene, more effective against the crawlers). Other pesticides
that kill scale insects include oxydemetron-methyl (Metasystox-R), oxamyl (Vydate), and disulfoton (Di-
syston) granules. See “Aphids” for more details.

S  S

The brown garden snail and various slugs can quickly devastate ferns. Other species that usually eat
decaying vegetation can also damage certain ferns. Small snails, such as the rotund disc snail, do not leave
a conspicuous slime trail and easily go unnoticed. Their damage causes small holes in the foliage and col-
lapsed fiddleheads. Slugs do not have a shell like that of a snail, but they do leave a mucous trail. New
growth and tender foliage are most vulnerable, as are certain fern genera (especially Asplenium). Slugs and
snails come out of hiding and feed at night. During the day they hide in dark, moist, cool places.
P  C. Examine the plants and surrounding areas (especially if moist) in the
late evening to handpick and destroy the snails and slugs. During the day, search under flower pots or in
other hiding places. With snails, be certain that they are completely killed as they can easily repair cracked
shells. Keep a clean culture by removing debris and other potential slug hideouts. Fill in cracks and crev-
ices in walls, fences, and building foundations with mortar to reduce winter hiding places. Beer in cups or
various traps attract snails, but not as well as methaldehyde baits. Coarse bark chips and various mulches
discourage slug and snail movement. In particular, ashes inhibit the movement of slugs and snails but
should not be applied in large amounts to the soil. Sharp sand, perlite, gritty materials, and ground peat
132 C 11

moss are other mulches that snails dislike to traverse. Slugs do not seem to object to sand. Snails are stopped
by vertical copper bands at least 4 cm (1.5 in.) wide, though slugs are less inhibited by this. Insecticidal soaps
reportedly kill slugs and snails.
Decollate snails are predators of brown garden snails but will eat tender foliage, including that of some
ferns. Slugs are rarely attacked by decollate snails. Toads, opossums, and other wildlife feed on snails and
slugs.
For chemical controls, various baits, granules, and sprays can be applied on less exposed areas of flats,
fences, and walls where snails and slugs travel and feed. The pests tend to return to the same hiding and feed-
ing areas.
Methaldehyde attracts and stuns slugs and snails, and death is dependent on subsequent dehydration.
Slugs and snails can recover and may live to develop resistant strains. Keep pets, particularly dogs, away from
methaldehyde baits, which are poisonous. Baits in pellet form can become unsightly due to mold; other
forms do not mold or the mold is less conspicuous. Methaldehyde is less effective at temperatures below
20°C (68°F) and loses potency in sunlight and high humidity. A special formulation (which can be obtained
as Deadline, Corry’s, and other brand names) resists sunlight and humidity damage and lasts longer. Sprays
have the advantage of penetrating the soil where slugs may be hiding, but earthworms will also be harmed.
Avoid spraying fern foliage.
Methiocarb (Mesurol) kills snails or slugs instead of stunning them. It also kills earthworms. The gran-
ular form should be applied when the foliage is dry. Increased humidity increases its toxicity; where humid-
ity fluctuates widely, using methiocarb in combination with methaldehyde will give better control. For use
in commercial greenhouses, methiocarb comes in aerosol form. It also kills aphids, scale insects, spider
mites, moths, and whiteflies.
Mexacarbate (Zectran) is infrequently sold. Because direct sprays will burn maidenhair and possibly
other ferns, it should be applied only to the soil and walkways. The dehydrating effect of aluminum sulfate
applied as a drench of 56.7 grams per 3.8 liters of water (2 oz. per gal.) is reported to kill snails and slugs,
but its effect on ferns is unknown. It might acidify the soil. For further details, see Parrella et al. (1985).

S  P

Sowbugs and pillbugs are segmented, oval-shaped pests that eat decaying plants and the tender new
growth of fronds, rhizomes, and roots. Pillbugs roll into a ball when disturbed; sowbugs do not. Both are
also known as woodlice.
P  C. Keep a clean culture by removing debris and hiding places. Pour boil-
ing water into hiding places, or attract the bugs to one spot with half a potato, apple, or turnip pulped on
one side and placed cut side down. A small inverted flowerpot filled with damp moss can also be used.
The chemical controls are the same as for millipedes (which see). Diazinon is particularly effective, as
are Sevin dust (carabyl) and pyrethroids.

T
Thrips are small, slender insects that cause noticeable white or bleached areas on ferns by sucking the
plant juices. Bleached areas may become especially conspicuous during the summer months. Tiny, scattered
black or brown dots (mold-infected honeydew) left by the thrips are visible with a hand lens on the under-
side of the damaged area of the frond (Figure 11.4). By the time the bleached areas appear, none or only a
few lingering thrips will be seen. Greenhouse thrips, the species that attacks ferns, have dark brown to
black adults, about 1 to 1.5 mm long. This species is common and found worldwide. It may attack ferns
indoors or outdoors. The flower thrip does not form black dots on the foliage and has not been reported to
attack ferns.
 T  G F 133

Figure 11.4. Thrips (6×). Note the dots of secretion and

bleached leaf tissue.

P  C. Keep the surrounding grass mowed and remove weeds and badly dam-
aged fronds. Ferns susceptible to thrips (such as Polystichum and some Dryopteris) should be placed in
cooler, more humid sites of the garden. If in a greenhouse, use 0.178 mm (0.007 in.) brass mesh screen
across openings. The chemical controls of choice are acephate (Orthene), disulfoton (Di-syston), malathion,
and oxydemetron-methyl (Metasystox-R). Others include chlorpyrifos (Dursban), dimethoate (Cygon
2E), fluvalinate (Mavrik Aquaflow), oxamyl (Vydate), and imidacloprid (Marathon). See “Aphids” for
more details; also see “Alternatives to Chemical Controls.”

W
Whiteflies are sucking insects about 3 mm (0.1 in.) long that fly upward when the foliage is disturbed.
Examination of the leaf’s underside will reveal many flattish, oval larvae and pupae adhering to the surface.
Whiteflies are uncommon on ferns, although in some areas newly introduced species are attacking some
ferns and are difficult to control.
P  C. See “Alternatives to Chemical Controls” for biological controls, soaps,
and insect growth-regulators.
Chemical controls for greenhouse whiteflies are acephate (Orthene), diazinon, dimethoate (Cygon
2E), fluvalinate (Mavrik Aquaflow), malathion, oxydemetron-methyl (Metasystox-R), and resmethrin,
among other registered controls. See “Aphids” for more details. Insecticides used for greenhouse whiteflies
may not be effective against other whitefly species.

O A P

Many insect pests feed on fern foliage or prothalli. They include cutworms, caterpillars, beetles (includ-
ing weevils), crickets, grasshoppers, and cockroaches. Their presence is often indicated by damaged foliage,
disturbed soil, or droppings. Earthworms and other soil dwellers do not directly damage ferns but may indi-
rectly damage prothalli or young sporophytes by churning the soil around them. Mice may eat ferns.
134 C 11

P  C. The chemical control of choice is carbaryl (Sevin), but also used are
acephate (Orthene), diazinon, and chlorpyrifos (Dursban). See “Aphids” for more details. Also see “Alter-
natives to Chemical Controls.”

Fungi and Bacteria Attacking Ferns

Generally, cultivated ferns are not susceptible to fungal or bacterial diseases. These usually appear when the
plants have been kept too wet at the roots or on the foliage. Prevention is better than attempting a cure.
Ferns properly planted and cared for are less likely to succumb to disease. A number of symptoms common
to virus infections on ferns have been noted but not well studied.
Prevention starts with healthy plants that show resistance to disease. Provide them with good drainage,
and keep the crown of the plant above the soil. Foliage should not be allowed to stay wet for long periods.
Avoid overwatering and splashing, and be sure to water in the early part of the day to allow time for the foliage
to dry. Space the plants far enough apart so that there is adequate air circulation around and between them.
Fronds should be gently shaken to remove the excess water, especially with finely divided foliage that becomes
matted with water; better yet, avoid frequent overhead watering with such ferns. Keep a clean culture,
because old leaves, plant debris, and weeds might harbor disease. Potted plants should be kept raised above
the soil level to prevent possible contamination. Little is known about biological controls for fungal or bac-
terial diseases. Fungi (Trichoderma and Gliocladium virens) that control water molds are being studied.
Preventative measures involve sterilizing the soil, pots, and tools used in planting (see “Sterilizing Soils”
in Chapter 5). Pots, tools, and benches can be disinfected with alcohol, 10% bleach solutions, quaternary
ammonium compounds (Physan 20, Green Shield), or carbolic acid (Lysol). Formaldehyde is unsafe for
people, pets, and the environment and has been withdrawn from the market for such use.
When disease symptoms appear, cut the diseased leaves back to healthy tissue and destroy the diseased
parts (see “Leaf Spots” for more details). Fungal diseases tend to produce spots with a halo pattern that is
lighter colored at the edge. Bacterial diseases generally cause discoloration that is shiny and water-soaked.
Diseased parts should be discarded to where they will not infect other plants.
A last resort for home growers is to use chemicals (see Appendix IV). Most fungicides will only suppress
the disease, not eradicate it. Some injure the plant, others are expensive, and most have not been tested on
ferns. If they have been tested, it is usually only on the Boston fern and leather fern.
Fungicides should be sprayed to provide good coverage, especially on the lower leaf surfaces favored by
fungi. Apply drenches to moist soil. Fungi that attack ferns fall into two categories: water molds and non-
water molds. No spray exists to control both, but combinations of compatible fungicides are available to
home growers, such as Banrot (contains etridiazole and thiophanate-methyl). If the specific fungus causing
the disease is unknown, a broad-spectrum fungicide (usually compatible mixes) must be used. Compatible
mixes sometimes work better than if the fungicides are used separately. Most broad-spectrum fungicides are
effective against Botrytis, Rhizoctonia, and water molds that cause most of the blights and damping-off
symptoms. Commercial growers routinely spray fungicides and apply drenches as preventative measures,
but some fungicides accumulate in the soil to levels that are toxic to plants, and sometimes fungi will
develop resistance. Rotating fungicides may be needed to avoid these problems.
Identifying pathogens to species requires time-consuming collaboration with a plant pathology labo-
ratory, but commercial growers need quick identifications to begin treatment as soon as possible. To fill this
need, kits (such as Alert) are available to identify certain diseases within minutes. Presently, these kits detect
only three soil-borne diseases: Rhizoctonia, Pythium, and Phytophtora (the latter two are water molds).
Routine preventative fungicide treatments used by commercial growers include Chipco 26019 (or
Cleary’s 2226) and Subdue. They are registered for use in certain situations (such as outdoors, greenhouses,
 T  G F 135

interiorscapes), for certain types of plants, and for certain users. As with pesticides, use of these chemicals
other than as described on the label violates the law.
In the following section the common names of fungicides and some of their trade names are given.
Trade names are used for convenience and do not imply endorsement. See Appendix IV for cross-references
between trade names and the active ingredient.

Armillaria R-R
Armillaria root-rot has been reported to kill Dicksonia fibrosa in southern California. It is widespread
and attacks woody tissue, but some woody plants are more susceptible than others. The infected plants
decline and eventually die. Prying into a dead stem will reveal the whitish growth of the mold. Eventually
the characteristic honey-colored mushrooms may emerge, usually around the base of the stem.
P  C. Avoid planting Dicksonia fibrosa or woody plants in areas susceptible
to Armillaria unless the species are resistant. The soil can be fumigated before planting, but recontamina-
tion is likely to occur.

B
Blights are caused by fungi or bacteria that attack young growing tissue. Affected young plants will
wither and die, and the fronds of older plants will dry at the tips and margins (especially in Boston ferns).
Blights can appear as water-soaked spots on bird’s-nest ferns (Asplenium nidus). Also see the section “Tip
Blight” later in the chapter.
P  C. Remove and destroy blighted parts and disinfect tools. Avoid getting
the foliage wet, reduce humidity, and space the plants farther apart to facilitate drying and improve air cir-
culation. If the specific disease organism is unknown, use a broad-spectrum spray (such as Banrot). Chem-
icals with the active ingredients propiconazole (Banner), chlorothalonil (Daconil 2787), iprodione (Chipco
26019), or mancozeb (Dithane M-45) might also be effective. Also see “Rhizoctonia” and “Water Molds.”

D-o
Damping-off diseases attack young plants near the soil surface and cause them to fall over, wither, and
die. Prothalli often reveal the symptoms of damping-off. Several different pathogens cause the disease.
P  C. Sterilize the soil if possible. Provide good ventilation, keep tops of
young plants dry, and reduce the moisture level to a minimum. Broad-spectrum fungicides must be used
unless the pathogen is known. Commercial growers favor Banrot or a combination drench of iprodione
(Chipco 26019) and metalaxyl (Subdue). See also “Blights,” “Rhizoctonia,” and “Water Molds.”

G M (Botrytis)

The fungus Botrytis forms delicate tufts of grayish white mold on dead, injured, or dying fronds. Infec-
tion typically starts from the upper part of the plant and proceeds toward the soil. Young ferns may survive
for a short time but eventually produce deformed fronds or stop growing altogether. Botrytis spores are air-
borne and common. Cool moist conditions favor the fungus.
P  C. Remove dead and injured leaves. Maintain a warmer, drier environ-
ment. Promote free air circulation and use fans if necessary. The following fungicides are effective against
gray mold: chlorothalonil (Daconil 2787), copper sulfate pentahydrate (Phyton 27), dimethyl 4,4′-O-
phenylenebis (3-thioallophanate) (Cleary’s 2226), iprodione (Chipco 26019), mancozeb (Dithane M-45),
136 C 11

maneb (Dithane M-22), thiophanate-methyl (Zyban), and zineb (Dithane Z-78). All are registered for
use on foliage plants, but often the only ferns that have been properly tested are the Boston fern and leather
fern. Spray test before widely applying any fungicide to your ferns.

L S
Spots and blotches on leaves are caused by factors such as poor air quality (see Chapter 4), physical dam-
age, excessive moisture or shade, fungi, bacteria, and nematodes. Healthy ferns in well-ventilated places sel-
dom become infected. Blemishes usually appear on older fronds, but new fronds can also become infected.
P  C. With minor infections, the disfigured fronds should be removed to
prevent spreading. Cut the foliage back to healthy tissue, destroy the infected parts, and disinfect all tools.
Reducing humidity, spacing the plants farther apart, increasing air circulation, and avoiding overhead
watering or splashing will help prevent further infection. If the spots are few, smearing them with a cotton
swab dipped in a strong solution of baking soda, a 10% bleach solution (avoid healthy tissue), or a quater-
nary ammonium compound (Physan 20 or Green Shield) may help stop the spots from enlarging. This
treatment is particularly effective for the larger spots appearing on staghorn ferns. Experiments indicate that
insecticidal soaps can increase or decrease leaf spots, depending on the disease and host. A 0.5% baking-
powder solution has successfully been used to treat leaf spots but is not registered for such use. It is report-
edly more effective when combined with a horticultural oil, such as Volck Oil Spray, a petroleum oil.
Chemical sprays are a last resort. A broad-spectrum spray (see under “Blights”) must be used unless the par-
ticular pathogen is known.
Some specific leaf-spot pathogens that have been identified on ferns include Cercospora, Cladosporium,
Colletotrichum, Cylindrocladium, Fusarium, Myrothecium, Peyronellaea, Phoma, Pseudomonas, and Xan-
thamomas campestris. The latter two are bacteria that produce rot with a wet appearance (especially on base
fronds of Platycerium). They can be controlled by Physan 20 applied with a cotton swab. Leaf blister (Taph-
rina), which forms a blister on both sides of the leaf, attacks mostly native ferns.

Rhizoctonia
Rhizoctonia is a fungus that usually attacks plants near the soil surface. The stems or stipes rot at the
point of infection, and the weakened stalk cannot support the frond. The infection can also spread below
the soil. Thicker roots turn yellow or brown and have cankers. Prothalli and young plants wither and damp
off. Fronds may wilt, grow slowly, be deformed, or stop emerging altogether. This condition is called “hard
crown” by some growers. Rhizoctonia often causes black-brown lesions on stems, leaf stalks, and roots.
The base fronds (shields) of staghorn ferns may show wrinkly brown lesions.
The decaying parts of infected plants have brownish threads that can be seen with a hand lens. These
threads are coarse enough to cling to soil particles (water molds have fine threads that do not cling to soil par-
ticles). Under the microscope, the characteristic T-shaped cells where the filaments branch are diagnostic.
Rhizoctonia also attacks tree ferns. The new fronds do not uncoil but seem to dry in place. Mature
fronds will wilt, and the crown hardens and turns brown. Eventually the plant dies.
Overwatered plants or ones growing in poorly drained soils are most susceptible to infection. Infected
plants continue to grow but usually languish and succumb to minor stresses. High humidity and temper-
atures of 21 to 27°C (70–80°F) favor the disease.
P  C. Discard infected plants. Make sure contaminated soil or equipment
does not touch clean soil or equipment. Sterilize the soil before replanting contaminated areas. The disease
is soil-borne and can easily spread by splashing or flowing water. Several chemicals can be applied as a
drench, mainly as a preventative. They will inhibit the disease but not destroy it. The chemical of choice is
 T  G F 137

pentachloronitrobenzene (PCNB, Terraclor), but it burns certain ferns. Also effective are chlorothalonil
(Daconil 2787), Cleary’s 2226, iprodione (Chipco 26019), captan, etridiazole (Truban), and the broad-
spectrum Banrot. In all cases it is a good idea to test spray first.

R
Many species of fungi and bacteria can cause decay or rots. Rots caused by bacteria appear wetter and
slimier than those caused by fungus.
P  C. Cut away rotting spots. Disinfect tools and treat as for blights (which
see). Bacterial rots may respond to such bactericides as streptomycin (Agri-strep), copper sulfate pentahy-
drate (Phyton 27), and others. Some bactericides are expensive, and ferns may be sensitive to them. Small
spots of rot can be dabbed with a cotton swab, as mentioned under “Leaf Spots.” As a preventative meas-
ure, dust cut surfaces of the plant with fine sulfur or dip in a solution of general disinfectants (Physan 20
or Green Shield).

R
Rust diseases have long been known to affect wild ferns, but they are less frequently encountered on cul-
tivated ones. The disease produces orange-brown pustules or white spots on the underside of the leaf. Dis-
coloration can appear on the upper surface of the leaf opposite the pustules. Rust spores may be airborne.
Moist, cool nights and warm days favor rust. The rust Milesia is common in England on hart’s-tongue fern
(Phyllitis scolopendrium).
P  C. Remove infected leaves to minimize airborne rust spores that can rein-
fect other leaves. Winter cleanup, particularly the removal of old fronds, will reduce infection in the spring.
Propiconazole (Banner), folpet (Phaltan), chlorothalonil (Daconil 2787), mancozeb (Dithane M-45),
maneb (Dithane M-22), and zineb (Diathane Z-78) are chemical sprays that kill rusts. Their effect on
ferns is not known.

S M
Sooty mold is a fungus (usually Fumago) that appears as black smutty patches on leaves. It also grows
on the honeydew drops secreted by aphids and mealybugs. Although unsightly, sooty molds do not harm
the plant. They do, however, intercept light to the fronds.
P  C. Control scale insects, aphids, mealybugs, and ants. Promote air
circulation.

T B
The development of irregular, deformed fronds and a subsequent slowing or cessation of growth may
appear among young and well-established species of tree ferns (such as Dicksonia antarctica and Sphaeropteris
cooperi). Laboratory tests on diseased Dicksonia plants have tentatively identified the pathogen as the fun-
gus Phyllosticta. This fungus is sometimes called tip blight because it causes the leaf tissue to appear sun-
burned along the margins, but the brown coloration can spread throughout the leaflet. Leaflets become dis-
torted, undersized, appear moth-eaten, or are absent in places. Emerging fiddleheads can be irregularly
twisted, and leaf-stalk cross sections sometimes reveal a central area of dead, darkened tissue. This disease
has appeared in southern California, and similar symptoms have been reported in the Hawaiian Islands.
P  C. Generous applications of fertilizer may retard the disease symptoms for
a time, but they eventually return. Broad-spectrum fungicides are reportedly effective against tip blight,
138 C 11

including chlorothalonil (Daconil 2787) for home growers, and iprodione (Chipco 26019), mancozeb
(Dithane M-45), and zineb (Dithane Z-78) for commercial growers. Overhead watering of tree ferns
should be avoided.

W M  R-R

Plants overwatered or allowed to stand in water are usually killed by water molds of the genera Pythium
and Phytophthora. These organisms thrive in cool temperatures. They first infect the roots, then the stem.
Symptoms of water molds include stunted growth, yellow or wilted fronds, and reduced root growth. In
addition, the stems, if not rotted away, become so soft that when washed the outermost tissue (cortex)
readily separates from the tougher central strand. Both genera of water molds also attack prothalli and
sporelings; these appear water-soaked when infected.
A few differences between the two types of water molds are evident, however. Generally, Phytophthora
tends to attack woody plants, whereas Pythium favors herbaceous ones such as ferns. Phytophthora discol-
ors the conducting tissue of the stem; Pythium discolors the softer tissue (the pith and cortex), often reduc-
ing it to a mush.
Water molds differ from Rhizoctonia by their tendency to infect the root tips and then progress upward,
whereas Rhizoctonia infects the stem at the soil surface and progresses up and down. Also, the filaments of
water molds, unlike those of Rhizoctonia, are so fine that they are invisible even with a hand lens, and they
do not cling to soil particles.
P  C. Prevention is the best measure. Provide good drainage and do not
overwater. Avoid splashing water, which will spread the disease from contaminated water and soil. As
always, do not permit contaminated soil or other equipment to be mixed with clean soil or equipment.
Infected plants should be discarded. Before replanting the contaminated areas, sterilize the soil. Peat moss
may be a source of Pythium.
A biological control fungus, Trichoderma, kills water molds but is not widely available. A weak (light
pink) solution of potassium permanganate used as a drench inhibits water molds. Metalaxyl (Subdue) is a
systemic fungicide used as a drench. Commercial growers sometimes use it as a preventative measure at rates
of 0.5 ml per 3.8 liters of water (about 5 drops per gallon). Granular forms are also available. Fenaminosulf
(Dexon, Lesan) kills only water molds but degrades in light and must be used at once. Etridiazole (Terra-
zole, Truban), fosetyl-Al (Aliette), and captan are also used. All are registered for use on foliage plants, but
sometimes only for Boston fern or leather fern, so test spraying should be done. Be careful with Aliette: it
can burn ferns.

Algae, Mosses, and Liverworts

Algae, mosses, and liverworts can shade or crowd fern prothalli and sporelings. Some species of cyanobac-
teria (blue-green algae) produce toxins, and this may explain the decline in cultures so contaminated.

A
Of all the algae found in spore pans, the green algae and cyanobacteria (blue-green algae) cause the most
trouble. Oscillatoria (a cyanobacteria) forms a blackish slime when growing under bright light, and a gray-
ish mold-like mat under high humidity and dimmer light. It also emits a distinct odor. Studies indicate that
some toxins produced by cyanobacteria reduce respiration in mammals and seed plants, which might
account for the inhibition of sporeling growth when these algae are present. Nevertheless, some of the
 T  G F 139

diminished growth probably results from the cyanobacteria robbing the plant of light, crowding it, and
attracting fungus gnats. Ferns that are inherently slow growing or those growing in poor conditions in
spore pans are most susceptible to algae. By providing optimum conditions for rapid fern growth, some of
the algae problems may be averted.
P  C. Practice sanitation in planting and growing areas. Sterilize the soil
before sowing spores. Use only distilled water because tap water might be contaminated with algae, and
avoid any other possible contamination sources, such as unclean watering cans or pots. If algae appear, pick
out the growth. In small plantings a light dab with a cotton swab will usually pick up the algae mat with-
out disturbing the ferns. Dry soil additives such as pumice and sawdust placed on top of soilless mixes can
reduce algae growth.
Algae can be controlled with quaternary ammonium compounds (Physan 20 or Green Shield). Sprays
of these algaecides were harmless to small plants of Cyrtomium falcatum, Dicksonia antarctica, Pteris cretica,
Sphaeropteris cooperi, and Sphaeropteris medullaris when used at a rate of 400 ppm (1.67 ml per 3.8 liters of
water, or ⅓ tsp. per gal.). With either chemical, reapplications will probably be necessary.
A pale blue solution (0.1 ppm) of copper sulfate (bluestone) used as a drench will also inhibit algae
growth. Do not allow any of the solution to get on the ferns. Various algaecides have been found to hurt
young ferns, whereas others (such as Algamycin) do not. The synthetic antibiotic chloramphenicol (Parke-
Davis or Rochelle) has been used to control cyanobacteria as a drench (at 20–40 ppm) but is not registered
for this use. Chlorothalonil (Daconil 2787) and mancozeb (Dithane M-45) are other fungicides that have
been reported as effective against some algae.
To prevent the spread of algae and to destroy the habitat of fungus gnats and other pests, the algae on
benches, walkways, and walls should be controlled. Many commercial preparations are available for this
purpose, but some are toxic to plants and must be used with care. Drenches or sprays of hydrated lime (at 681
g to 3.8 liters of water, or 1.5 lbs. per gal.) or copper sulfate (454 g to 3.8 liters, or 1 lb. per gal.) are effective.

M  L

Mosses and liverworts can become so abundant that they crowd and shade prothalli, especially in old
cultures under bright light. Do not allow liverworts (particularly Lunularia) to establish among your ferns.
They spread rapidly, carpeting the soil surface and preventing water, fertilizer, and air from getting into
the soil.
P  C. Sterilize the soil before sowing spores, and water with distilled water.
If mosses are a problem, do not put charcoal on the soil because many mosses flourish on it. Remove mosses
and liverworts by hand if they grow over the ferns. When removing liverworts, try not to fragment them
because the pieces will regenerate. Fern prothalli should be grown under lower light to weaken mosses. Most
species of greenhouse mosses need more light to grow robustly than do ferns. Provide optimum conditions
to hasten the growth of ferns, because slow-growing cultures tend to develop more mosses and liverworts.
Marchantia, a liverwort, can be killed with a spray of quaternary ammonium compound (Physan 20)
applied at 800 ppm (3.3 ml per 3.8 liters of water, or ⅔ tsp. per gal.). The effect of this spray on ferns is
unknown; however, it is known that strong concentrations damage some plants.
12 How Ferns Get Their Names

There are about 12,000 species of ferns worldwide, but only about 30 are commonly cultivated. Most peo-
ple know these cultivated species by easy-to-remember common names that are sometimes charmingly
appropriate or even whimsical. Common names are useful when dealing with only a few species and when
local usage is uniform, but they can cause confusion if many species and people from distant lands are
involved. One reason for the confusion is that several different common names might apply to the same
plant. Phlebodium aureum, for example, is known variably as the rabbit’s-foot fern, the bear’s-foot fern, the
serpent fern, and golden polypody. Conversely, several different species might be referred to by the same
common name; the name “lace fern” is used to describe plants in five different genera (Cheilanthes gracil-
lima, Microlepia strigosa, Nephrolepis exaltata ‘Smithii’, Odontosoria chinensis, and Paesia scaberula). No
rules are established for designating the correct common name for a species. Any number of names might
exist, and new ones can be coined by anyone. Another problem with common names is that most plants
don’t have them.
Scientific names have the disadvantage of often being long, difficult to pronounce, and hard to remem-
ber. Because they are in Latin, their meaning is usually obscure to most people. But scientific names offer
several advantages over common names. They are understood internationally. A trained botanist in Indo-
nesia, Russia, or Mexico will know exactly what you mean when you utter “Dryopteris.” Using scientific
names reduces confusion and increases accuracy in communication. The other great advantage is that they
indicate a degree of evolutionary relationship. You can tell by the name, for example, that Dryopteris inter-
media and Dryopteris marginalis are more closely related to each other than either is to Polystichum acrosti-
choides. If the common names of these three species were used—glandular wood fern, marginal shield fern,
and Christmas fern—relationships are not apparent. If you deal with many species and want to be precise
when communicating with other plant people, then it is worth the time to learn about scientific names.
Every species has only one correct scientific name. The situation is complicated, however, because
many species have synonyms, or alternative scientific names, that arise as a result of new discoveries or
hypotheses about relationships. For example, in the 1800s the Massachusetts fern was called Dryopteris
simulata, but this name is now considered a synonym of Thelypteris simulata because the plant belongs
with the thelypterioid ferns instead of the dryopteroids. The change resulted from greater knowledge based
on new evidence about the relationships of the plant.
In the encyclopedic portion of this book, commonly encountered synonyms are given after the accepted
scientific name, which is the one currently used in most floras, manuals, and monographs. The synonyms
are given because they continue to be used by some gardeners and trade people. There is no mandate that
new names be immediately adopted. Sometimes they are controversial, and even professional botanists
disagree as to which name is correct. To avoid confusion, plant labels can give both names, placing the syn-

140
 H F G T N 141

onym in parentheses. The word “formerly” is sometimes added before the synonym. Adding this extra
name to labels or catalog lists might be an inconvenience to the commercial growers, but gardeners are
becoming more knowledgeable about species names and appreciate greater accuracy in labeling.

Naming Species
The scientific name of a species consists of two words. The first is like the surname of a person (Doe), the
second is like the given name (Jane). The first word denotes the genus or larger group to which the plant
belongs and indicates some degree of relationship. The second word denotes a particular kind of plant in
the group. For instance, Asplenium bulbiferum is the scientific name for the mother fern. The first word,
Asplenium, is the genus to which the mother fern belongs. The second word, bulbiferum, indicates the par-
ticular kind of Asplenium. There is a similar fern called Asplenium daucifolium. It is apparent from this
name that the species is related to the mother fern because both are in Asplenium. It is also clear that the fern
is not the same species as the mother fern because it has a different second word, or specific epithet. These
two species differ not only in size and color but more significantly by the shape of the leaflets and the rhi-
zome scales (Figure 12.1). Having the same specific epithet means nothing. Botrychium virginianum and
Polypodium virginianum have the same specific epithet, but they belong to different genera and families. In
this case the specific epithet indicates that the original specimens from which these species were described
were collected in the State of Virginia.
How does a plant get a scientific name? A plant suspected as new to science is studied by a botanist who
compares it to its relatives. If no match is found, and the plant is not a freak or variant of a known species,
then the botanist names the plant according to the rules set forth in the International Code of Botanical
Nomenclature, or the Code for short.
If the plant differs greatly from known plants, the botanist might decide to give it a genus name all its
own. If, on the other hand, the botanist believes the plant belongs to an established genus, a specific epithet
is added to the genus name to complete the species name of the plant. The botanist can name the plant after
anything; it may be the name of the person who discovered it, the place where it was discovered, or a struc-
tural feature.
The botanist who names the plant is known as the author or authority. The author’s name appears after
the scientific name, although this is sometimes omitted in general use. The author’s name may be given in
full, or sometimes only the initials or a surname abbreviation are used. The author citations are important
to botanists in determining the priority and application of names. The scientific name for the bird’s-nest
fern, for example, is Asplenium nidus Linnaeus (sometimes abbreviated “L.” or “Linn.”) for the famous
Swedish botanist who began the consistent practice of giving species two names. This practice is called the
binomial system, and it permits a systematic arrangement of species into a filing system.

P S N

It matters little how one pronounces scientific names as long as they are understood by the listener. No
one knows exactly how Latin was pronounced, and even in the heyday of the Roman Empire several dialects
were probably spoken in different regions. So who’s to say who’s right? The best advice is to pronounce the
names as the people around you do. When someone presumes to correct your pronunciation, a knowing
smile is the appropriate response.
142 C 12

Figure 12.1. The fronds of the closely allied species Asplenium bulbiferum (left) and A. daucifolium (right).

Variation Within Species

Variation is a fundamental property of all living things. Just as species vary throughout their range, so do
individuals within a given population. This variation might be in plant structure or function, such as pro-
ducing fuller fronds or producing spores later in the season. The variation could be caused by growing
conditions, such as the amount of shade or soil moisture, or by genetic differences, such as mutations or
novel gene combinations. Variants with a hereditary basis can be named. Such a name appears after the spe-
cies name. The common categories for these names are subspecies (subsp.), botanical varieties (var.), forms
(forma or f.), and cultivated varieties (cultivar).

C V
A cultivated variety, or cultivar, is a distinctive plant arising through selection from cultivated or wild
stock or by hybridization in cultivation. Its distinctive attributes are uniform and stable and are retained
when propagated. Whether or not the variant is distinct and worthy of being assigned a name is left to the
individual naming the cultivar to decide. Not every fern variation is named, and many are unworthy of
receiving a name. Some common variations seen in cultivated varieties of ferns include the following:
 H F G T N 143

• Changes in the leaf margin, such as fringing, lobing, ruffling, cresting, or forking.
• Changes in size of the parts, such a depauperate, narrower, wider, smaller, or large pinnae or
pinnules.
• Changes in position of the parts, such as overlapping or more distant pinnae.
• Changes in overall size of the plant, such as dwarfism or gigantism.
• Changes in color, such as variegation.
• Miscellaneous changes, such as formation of buds, failure to form spores, maintenance of
juvenile foliage, and even physiological differences such as greater resistance to drought or
cold.
Figure 12.2 shows the variations possible in cultivars of the lady fern.
The name of a plant cultivar is enclosed in single quotation marks and follows the species name. Older
usage might have the abbreviation “cv.” preceding the cultivar name. Valid cultivar names given after 1959
are in non-Latin form and in roman type, not italic. The main words of the name have the first letter cap-
italized, as in Adiantum raddianum ‘Pacific Maid’. The species epithet can be omitted if there is no likeli-
hood of confusion with another cultivar bearing the same name in the genus. Thus Adiantum ‘Pacific
Maid’ is permissible.
Cultivar-groups are used to name an assemblage of two or more similar, named cultivars. (The word
“named” was added to the definition of cultivar-group in the 1995 International Code of Nomenclature for
Cultivated Plants. This precludes using the cultivar-group as a catchall category for variants belonging to the
group but unworthy of recognition. This wording is currently being met with considerable controversy.)
This category may also be used to recognize distinct groups within a species when the species itself is no
longer recognized. For instance, plants known as Davallia fejeensis have been reduced by some botanists to
D. solida. Plants known as D. fejeensis are prized by horticulturists and in this reduction to D. solida would
be left without a name. However, it may continue to be recognized by using the cultivar-group category D.
solida Fejeensis Group. If a particular cultivar in the Fejeensis Group needs to be identified, the group
name is enclosed in parentheses, followed by the cultivar name in single quotation marks, as in Davallia sol-
ida (Fejeensis Group) ‘Plumosa’.
Naming of cultivated plants is governed by rules set forth in the International Code of Nomenclature for
Cultivated Plants, commonly referred to as the Cultivated Plant Code or ICNCP (Trehane 1995). The
Cultivated Plant Code promotes uniformity, accuracy, and stability in naming horticultural plants. Grow-
ers, plant businesses, and producers of cultivated varieties should use these rules to minimize confusion. The
assignment and use of cultivar names is a voluntary matter. It should not be confused with trademark
names or patents, which are legal matters involving the government (discussed later in the chapter).
Unfortunately, despite these rules, the naming of fern cultivars in the trade and among hobbyist grow-
ers often makes it difficult to determine whether a given cultivar is new or a renamed older one. Most cul-
tivars in circulation have not been named according to the rules in the International Code of Nomenclature
for Cultivated Plants, and thus, technically speaking, have invalid names. Yet it is impossible to ignore these
names because they are an important part of the commercial trade and of many gardens. Until a Fern Reg-
istration Authority is established, the problems with the valid naming and identification of many cultivars
will be difficult to solve.

B V  F

Species that show significant variation in different parts of their geographic range are said to have bo-
tanical varieties. These variations may be given botanical variety status (var.) and named. For example, the
bracken fern (Pteridium aquilinum) has several botanical varieties (Figure 12.3). In the western United
 1
3
2
10

11

5 12
6

8
7 13

Figure 12.2. Variation in cultivars of the lady fern (Athyrium filix-femina): 1. ‘Caput-Medusae’; 2. ‘Foliosum
Grandiceps’; 3. ‘Grandiceps’ (dwarf form); 4. ‘Gemmatum Bolton’ (original clone); 6. ‘Acroladon’ (original clone); 7.
‘Frizelliae’ (original clone); 8. ‘Frizelliae Cristatum’ (from spore); 9 ‘Corymbiferum’; 10. ‘Pritchardii’ (original clone);
11. ‘Angustocruciatum Cristatum’; 12. ‘Fieldii’ (original clone); 13. ‘Victoriae’ (original clone);
14. ‘Victoriae Foliosum’; 15. ‘Flabellipinnulum’; 16. ‘Minutissimum Congestum’; 17. ‘Congestum Minus’; 18.
‘Crispum Coronans’; 19. ‘Veroniae’ (original clone); 20. ‘Plumosum Stansfield’ (original clone); 21. ‘Plumosum
Druery’ (original clone); 22. ‘Plumosum Furcillans’; 23. ‘Plumosum Penny’ (original clone); 24. ‘Plumosum
Multifidum’; 25. ‘Howardii’ (original clone); 26. ‘Clarissima Jones’ (original clone); 27. ‘Cristatum’; 28 ‘Clarissima
Cristatum Garnett’ (original clone). After Kaye (1965).
 14
21
20

22
15

23

24 25

17
16

26

27

19

28

18
146 C 12

States there is P. aquilinum var. pubescens, the western bracken; common in the northeastern United States
is P. aquilinum var. latiusculum, the eastern bracken. Besides geographical differences, the western bracken
has pinnules nearly at right angles to the rachis and hairs on the indusia, while the eastern bracken has
obliquely placed pinnules and hairless indusia. A third and more southeastern variety, P. aquilinum var.
pseudocaudatum, the tailed bracken, has long, narrow terminal segments. A fourth variety occurs in south-
ern Florida: P. aquilinum var. caudatum, the lacy bracken, with very stiff, wiry fronds. (Some botanists now
believe var. caudatum to be a separate species and call it P. caudatum.) The abbreviations “subsp.” or “ssp.”
stand for subspecies, a category used to imply an incipient species.

Figure 12.3. Comparison of the foliage of the botanical varieties of Pteridium aquilinum (left to right): var. latiusculum,
var. pubescens, and var. caudatum.

A variation that appears sporadically among plants in nature is placed in a category known as a form
(forma or f.). Additionally, the variation should be found sporadically throughout the range of the species.
An example is Blechnum spicant f. bipinnatum. The typical form of the species (B. spicant f. spicant) has
entire pinnae, but f. bipinnatum has deeply lobed pinnae. The lobed form, which is atypical, was found
growing among typical plants in nature (Figure 12.4).
Sometimes botanical forms can be environmentally induced. Examples are the lobed types of sensitive
fern (Onoclea sensibilis) and Christmas fern (Polystichum acrostichoides). In certain individuals of these spe-
cies, abnormally deeply lobed pinnae are produced on plants that also produce typical pinnae. The deep
lobes are caused by trauma from frost, mowing, fire, or transplanting (Beitel et al. 1981; Wagner 1942).
These variants are often referred to as Onoclea sensibilis f. obtusilobata and Polystichum acrostichoides f.
incisum, but they are unworthy of taxonomic recognition. If they were named, then two different form
names would apply to the same plant—an illogical situation.
 H F G T N 147

Figure 12.4. Comparison of the pinnae of the botanical forms of Blechnum spicant: forma spicant (left) and forma
bipinnatum (right).

Early use of the category “varietas” did not distinguish between cultivated varieties, botanical varieties,
and forms; hence, many plants listed as “var.” are nowadays considered cultivars or forms. Actually, the dis-
tinction between cultivated varieties, botanical varieties, and forms is poorly defined and overlapping. A few
plants might end up having both a botanical name and a cultivar name. The naming of subspecies, varieties,
and forms is governed by rules set forth in the International Code of Botanical Nomenclature.

H
A hybrid is the offspring from a cross between two different species. Fern hybrids are often sterile
because the chromosomes from the two parents cannot pair with each other during meiosis—the kind of
cell division that gives rise to spores. The result is spore abortion, and under the microscope the spores
appear irregular, misshapen, and often blackened. Ferns with aborted spores must be propagated by divi-
sion of the rhizome or by buds.
When sterile, hybrids are designated by the hybrid sign “×” placed between the genus name and spe-
cific epithet. Examples are Adiantum ×tracyi (= A. aleuticum × A. jordanii), Dryopteris ×triploidea (= D.
carthusiana × D. intermedia), and Polystichum ×potteri (= P. acrostichoides × P. braunii ). A name such as
“Adiantum ×tracyi” is the binomial; “A. aleuticum × A. jordanii” is the formula name.
Occasionally a sterile hybrid doubles its chromosome number, a process referred to as polyploidy.
Polyploidy automatically confers fertility to the hybrid. The plant can now produce normal (not mis-
shapen) spores that germinate upon sowing. Such fertile hybrids, also known as “nothospecies,” take on an
existence of their own, sometimes spreading far beyond the ranges of the parents. In this book, fertile spe-
cies of hybrid origin are given binomials without using the hybrid sign “×.” Examples of such plants are
Cystopteris tennesseensis, which arose from a cross involving C. bulbifera and C. protrusa, and Dryopteris
clintoniana, which arose from a cross involving D. celsa and D. ludoviciana.
Crosses between different genera may be placed in hybrid genera (Figure 12.6). To form the name of
a hybrid genus, the generic names of the two parents are combined, and the hybrid sign “×” is placed before
the name. For example, a hybrid between a species of Aglaomorpha and a species of Drynaria would be
placed in the hybrid genus ×Aglaonaria.
148 C 12

a × b
Figure 12.5. Hybrid of species in the same genus:
a. parent Adiantum jordanii; b. parent A.
aleuticum; c. hybrid offspring A. ×tracyi.

If naturally occurring hybrids are brought into cultivation, they must keep their hybrid name unless
they were given an additional cultivar name at the time of publication (this rarely, if ever, happens). For
example, Adiantum ×tracyi must be listed as such in horticultural use (Figure 12.5). But had this hybrid
originated in cultivation (it did not), it would be permissible to list it as Adiantum ‘Tracyi’.

Why Fern Names Change

Changing the accepted scientific name of a fern annoys growers and even professional botanists. But with-
out name changes, new knowledge about ferns would be difficult to assimilate and organize. Maintaining
existing names would be like keeping a static file. Newly discovered plants would have to be added to the
existing categories in the file. Some categories would contain such large numbers of plants that they would
be unwieldy to work with and difficult to understand. Unrelated plants would be inadvertently placed
together, while related ones could be scattered in separate categories. It would be like trying to classify
vehicles of today into categories made for the vehicles of 1900.
To accommodate our increasing knowledge of plants and to organize it into a meaningful relationship,
files occasionally have to be changed. New categories might need to be added, and older ones combined or
divided. As a result, there will be name changes. But of greater importance and broader interest is that the new
knowledge of plants and their relationships is incorporated into the file, and the way the plants are organized
in the file will reflect our updated knowledge. There are still many gaps in our knowledge of plant relation-
ships, and until these plants are thoroughly studied and understood, name changes are sure to continue.
 H F G T N 149

a × b
Figure 12.6. Hybrid of species of different genera: a. parent Asplenium
adiantum-nigrum; b. parent Phyllitis scolopendrium; c. hybrid offspring
c ×Asplenophyllitis jacksonii. After Lovis and Vida (1969).

Trademarks and Plant Patents

A grower or originator of a cultivar may obtain a trademark or a trademark name for his or her plant. If
approved by government authorities, the trademark does not prevent others from growing the particular
plant, only from using the mark or name. Such trade names are not technically considered cultivar names,
since the International Code of Nomenclature for Cultivated Plants stipulates that cultivar names must be
available to all users. According to the Cultivated Plant Code, the trade names should be printed in capi-
tal letters to distinguish them from the true cultivar name and to indicate their special status. Trademarked
names and plant patents are usually recognized by notations or symbols next to the names.
Patents are separate legal entities from trademarks and must be applied for separately. United States law
grants a patent to anyone who has invented or discovered and asexually reproduced any distinct and new
variety of plant. A plant patent excludes others from growing, using, or selling the plant for 17 years in the
United States and its territories. Foreign countries may have reciprocal agreements with the United States,
or they may have their own laws. Fees, illustrations, and descriptions of the plant and how it differs from
others are some of the requirements of applying for a plant patent. Patented plants are primarily identified
by a patent number and secondarily by a name. For added protection, patented plants may also have a trade-
mark name, although growers often forego this additional expense. The issuing of trademarks and patents
in the United States is governed by the Patent and Trademark Office of the U.S. Department of Commerce
in Washington, D.C.
The Plant Variety Protection Act (Public Law 91-577), administered by the U.S. Department of Agri-
culture in Beltsville, Maryland, provides for a system of protection for cultivated varieties produced from
seeds. Since ferns reproduce from spores, this law is not applicable to cultivated fern varieties.

Learning the Fern Genera

When looking at the diversity of cultivated ferns, we somehow know that a particular fern is a staghorn fern
or a maidenhair fern. Some ferns are so distinct that we have little difficulty recognizing them. With these
and other ferns, the overall appearance is enough to reveal to us its fern group. Other ferns require closer
observation. For instance, if you look only at their foliage characteristics, the maidenhair spleenwort (Asple-
nium trichomanes) and the bird’s-nest fern (A. nidus) appear very different—yet both are members of the
150 C 12

b
a

Figure 12.7. These two seemingly dissimilar

ferns belong to the same genus on the basis of
the sorus shape and the presence of indusia:
a. Asplenium trichomanes, the maidenhair
spleenwort; b. A. nidus, the bird’s-nest fern.

genus Asplenium (Figure 12.7). What distinguishes this genus are the sori and indusia, not the shape of the
frond. The sori and indusia of the two Asplenium species are long and narrow, and although longer in the
bird’s-nest fern, they are basically the same in both species. The indusium in both species is attached to the
upper fork of a vein branch, and the rhizome scales are latticed or resemble a stained-glass window
(clathrate) when observed through a hand lens. (They also all have two vascular bundles in the stipe.) Thus
the relationships between ferns is not always obvious to the naked eye.
The fern student interested in learning how to recognize ferns in general should start by studying the
genera. Start with a fern that you know, such as the maidenhair ferns of genus Adiantum. Select a fertile
frond, and look up the scientific name of the plant in the index of this book. The discussion in Chapter 13
will describe a few of the more conspicuous features that distinguish this genus from others. Locate on
your fern the features mentioned, then check your observations with the illustrations of the genus. A hand
lens of 8 to 15× magnification will assist you greatly. Once you are satisfied that you can recognize the fea-
tures mentioned, keep them in mind; the next time you see another kind of maidenhair fern, look for these
features again. Soon you will associate maidenhairs with a certain set of features. You will also recognize fea-
tures that do not belong with the maidenhairs. Select other fern genera that you know and repeat the proc-
ess: (1) look up the scientific name; (2) locate the distinguishing features on your plant; (3) check your
observations with the diagrams; and (4) examine other members of the genus for the same features to rein-
force your observations. At this point you may be able to recognize some of the more technical differences
between the species within the genus, but it is most important to concentrate on the features of the genus
as a whole. The features used to distinguish a genus are generally less diversified and less confusing than
those used to distinguish species. Besides, the species will be easier to recognize once you have become
familiar with the genera. Most of the generic features are associated with the position, shape, and type of
sorus and indusium. Also important is the rhizome, which may be erect or creeping, hairy or scaly. The
venation and the shape of the frond may also be important. Chapters 2 and 13 and the glossary discuss or
illustrate most of these distinguishing features.
13 Ferns and Fern Allies
in Cultivation

This chapter describes the ferns and fern allies commonly cultivated in the United States. It omits those spe-
cies found only in special collections and botanical gardens. Some species treated here may no longer be in
cultivation; conversely, it was impractical to include all recent introductions. A few genera not cultivated
in the United States are included because they were deemed worthy of culture, challenging to grow, or im-
portant in teaching botany. Most of the entries have been verified against botanical specimens and/or de-
scriptions; however, specimens or sufficient data were not always available, and many species are poorly
known. The correctness of the names must be considered in view of these problems.

Explanations of the Entries

G E
Common alternative names (synonyms), if any, are listed after each genus name. Synonyms are fol-
lowed by common names, which are given only if widely used. The text that follows provides a general in-
troduction to the genus, followed by a botanical description. Any special name problems encountered
within a genus are also discussed.
If botanists agree on a particular genus name and there is little possibility of a new name change, the
agreed-upon name is usually adopted. When a name has been changed, however, the traditional genus
name will be cross-referenced.
Large genera such as Athyrium, Cheilanthes, Davallia, Polypodium, Selaginella, Thelypteris, and others
have been subdivided formally or informally into smaller groups. The diagnostic characters and species are
listed for each subgroup. This will assist the reader in identifying a fern to a particular subgroup without
having to study each fern in the species entry.

S E
Species within each genus are listed alphabetically by scientific name. Synonyms or misapplied names
and common names, if any, follow. Cultural information is followed by a brief botanical description and
information on the geographical area to which the fern is native.
Where species important in horticulture were reduced by botanists to another species (as is the case in
the genus Davallia), the horticulturally important species name was retained with a note in the entry indi-

151
152 C 13

cating the discrepancy. Keeping such names is the least disruptive stance to take at this time. If necessary,
they may later be given cultivar status.
Only the most important synonyms are listed for a species; many synonyms not used in horticulture are
omitted. You can get an idea of the other genera under which a species might be synonymized by looking
at the generic synonyms after the genus name, or by referring to the Index of Plant Names. If the fern is
widely known under an incorrect name, this name will be noted as being “misapplied to” or as a “trade
name.”
Subspecies, varieties, and cultivars. Within the species entries, plants below the rank of species, including
subspecies, varieties, forms, and cultivars, are also discussed. Due to the tremendous number of cultivars—
there are at least a thousand—including all cultivars is beyond the scope of this book and many are omit-
ted. We have focused on those cultivars that are more common in the trade or among hobbyists and have
a fairly distinct appearance. Cultivars with names that are believed to be in agreement with the International
Code of Nomenclature for Cultivated Plants are also favored. As discussed in the previous chapter, the culti-
var names circulating in the trade are not always reliable because they are often invalid or inconsistently or
erroneously used. Where a name is known to be commonly misapplied to a particular plant, that is indi-
cated in the text.

C I
For each species, information is provided on the temperature, light, soil, and water requirements of the
plants. Ornamental attributes, including height and habit, and landscape uses are also discussed. The cul-
tural information is meant only as a guide. Often little is known about a fern’s requirements for growth, and
what is known can vary greatly with different gardens, microclimates, and genetic races of the plants.
Height. As used in this chapter, fern size refers to the height of the whole plant, not the frond length. Be-
cause some ferns vary greatly, the given heights may have exceptions. The following general terms are used
to describe fern height:

Very small: less than 15 cm (6 in.) tall

Small: less than 30 cm (1 ft.) tall
Medium: 30–100 cm (1–3 ft.) tall
Large: over 100 cm (3 ft.) tall

Temperature. It is difficult to give the minimum temperature tolerances of most introduced ferns. Little
data exist on the subject, and what is reported is often insufficiently documented. Knowing the tempera-
tures of the fern’s native habitat helps, but ferns often grow well in colder or hotter places than might be ex-
pected from their natural distribution. The categories of hardiness used in this book are fairly broad and may
overlap, but they offer workable categories for sorting out the hardiness of different ferns. The range of av-
erage annual minimum temperatures in the following section is based on the USDA Plant Hardiness Zone
Map (United States Department of Agriculture, Agricultural Research Service, Miscellaneous Publication
Number 1475; also see the map at the end of the color plates section). These ratings indicate the minimum
temperature range at which a plant may be expected to survive. They do not necessarily represent the op-
timal winter temperature for the plant. Also keep in mind that the timing and duration of the minimum
temperatures can profoundly affect a fern’s survival, as can many other factors. Microclimates and genetic
variations within a species often create exceptions to hardiness ratings. Two ferns might be listed as toler-
ating the same minimum temperatures, yet one might survive only with cool nights whereas the other only
with warm nights.
 F  F A  C 153

Very hardy: Severe and extended winters, such as in alpine, tundra, taiga, and northern
coniferous forest. The range of annual minimum temperature is −46°C and below to −
29°C (−50°F and below to −20°F).
Hardy: Cold-temperate areas; tolerates long periods below freezing. The range of average
annual minimum temperature is −29 to −15°C (−20 to 5°F).
Semi-hardy: Warmer temperate areas; tolerates short periods below freezing. The range of
average annual minimum temperature is −15 to −10°C (5 to 15°F).
Semi-tender: Subtropical areas, with frost or subfreezing temperatures very rare; winter night
temperatures often reaching 4 to 10°C (40 to 50°F). The range of average annual
minimum temperature is −10 to −4°C (15 to 25°F).
Tender: Tropical areas, indoors, or greenhouses not below 16°C (60°F) at night, 18°C (65°F)
or higher during the day. The range of average annual minimum temperature is −4 to 4°C
and above (25 to 40°F and above).
Very tender: Temperatures should be maintained at about 21°C (70°F) or higher most of the
time; some species will tolerate infrequent dips to 18°C (65°F). The range of average
annual minimum temperature is 4°C and above (40°F and above).
With the dearth of data on introduced ferns, we have used broader categories of hardiness than the U.S.
Department of Agriculture’s hardiness zone map. Where USDA zones are known from reliable sources,
they are cited. The rough equivalent between the USDA zones and those used in this book are given in
Table 13.1.

Table 13.1. Geographical Areas and Their Hardiness Ratings

Geographical area Range of average minimum temperature USDA zones Ratings in this text

Alpine, tundra below −46°C to −40°C (below −50°F 1, 2 very hardy

to −40°F); extremely cold, arctic climate
Coniferous forest, northern −40°C to −29°C (−40°F to −20°F); very 3, 4 very hardy
Midwest cold temperate
Northeast and central Midwest −29°C to −18°C (−20°F to 0°F); cold 5, 6 hardy
temperate
Far West inland areas, south to −29°C to −18°C (−20°F to 0°F); cold 5, 6 hardy
northern New Mexico temperate
Southeastern states, most of −18°C to −7°C (0°F to 20°F); temperate 7, 8 hardy to semi-hardy
Texas to southern New Mexico or semi-tender
Coastal Northwest −12°C to −1°C (10°F to 30°F); 8, 9 semi-hardy to semi-
warm temperate tender or tender
Pacific states slightly inland to −12°C to −1°C (10°F to 30°F); 8, 9 semi-hardy to semi-
southern Arizona warm temperate tender or tender
Coastal southern California −1°C to 4°C and above (30°F to 40°F 10, 11 tender to very tender
and above); subtropical and
Mediterranean climate
Southern Florida, Hawaii above 4°C (above 40°F); tropical climate 10, 11 tender to very tender
154 C 13

Light. Because light intensity interacts with humidity and temperature, these factors must be considered
along with the light ratings given in this book. Light intensity is usually measured in units called foot-can-
dles. Estimating foot-candles by eye is difficult. See Appendix I for information on calculating foot-candles
with a camera light meter.
Low light: Dense shade, about 200 to 400 foot-candles.
Medium light: Partial shade and sun or medium shade, about 400 to 600 foot-candles.
High light: Bright shade, about 600 to 1000 foot-candles.
Direct sunlight: On a clear day, direct sunlight measures from 5,000 to 10,000 foot-candles.
Plants tolerating direct sun usually can withstand higher temperatures as well.

Soil. The following terms are used in the plant entries to categorize soil types:
Garden soil: Garden soil, preferably a loam with a lot of humus.
Potting mix: Mostly soilless mixes; must be well-drained if containing soil.
Uncut moss: Coarse, uncut, or unground moss (green moss or sphagnum moss). Used in
hanging baskets or tied to boards to give good drainage.
Drained: Provide with good drainage, which may be increased by planting on boards, hanging
baskets, or mixing material such as bark pieces, coarse perlite, or coarse sand in the
medium.
Acidic: Prefers acidic conditions (pH 4–7).
Basic: Prefers basic or limestone conditions (pH 7–8).

Water. Most ferns should be watered deeply but not too often; avoid light sprinkles. Expect the soil to be
saturated after a thorough watering. Fortunately for most plants, this saturated condition does not last
long as the extra water will drain away, the soil will start to dry, and much of the water will be replaced by
air. The basic requirement is to keep the soil at the moisture level preferred by the plant for most of the time
and not let it dry out. Water less during cool weather and more frequently during hot, dry weather.
Wet: The soil is saturated with water and the excess allowed to drain away (a condition known
as field capacity). Plants that prefer such conditions have to be watered more frequently to
maintain this level of wetness.
Moist-wet: The soil is not saturated with water but feels wet and tends to be sticky when
handled. It is a condition between those described for wet and moist.
Moist: The soil feels moist but is not so wet as to be sticky when handled. Most ferns prefer
this range of moisture.
Moist-dry: The soil feels damp but is not sticky to the hands. It is a condition between those
described for moist and dry.
Dry: The soil feels dry and is light in weight due to the low water content.

E  D F

As discussed in Chapter 9, the distinction between evergreen and deciduous is not always sharp. If the
deciduous behavior of a species is known and is consistent in different climates, it is indicated as “decidu-
ous” in the text. Otherwise, gardeners should evaluate each species for their particular climate. Most tropi-
cal and subtropical ferns grow slowly during cool months, and their fronds may become tattered and dis-
colored, but once the temperature rises they resume active growth.
 F  F A  C 155

Evergreen. As used in this text, evergreen refers mainly to hardy ferns that remain green even in freezing
temperatures. These ferns grow primarily in temperate climates. Tender ferns can be assumed to be ever-
green (or nearly so) unless noted otherwise.
Deciduous. As used in this text, deciduousness indicates fronds that markedly wither and fall from the plant
or wither in place (marcescent) even if temperatures remain constant. Some subtropical or tropical species
(mainly native to seasonal climates) are deciduous at certain times of the year regardless of temperature
changes.

D F
Each species entry includes the main diagnostic characteristics, not a full botanical description. Because
these descriptions are short, the accompanying illustration of the species should be examined. Refer to the
preceding chapters and the glossary for explanations of unfamiliar terms.

Examining Ferns for Identification

When examining a fern, you should select recently matured fronds from adult plants. Fresh, clean plant ma-
terial is especially important when looking at the indusia and scales. A good hand lens with 10× magnifi-
cation is essential for observing smaller features. Hold the lens close to your eye to get the widest field of view
and close-up of the subject. Keep the subject in full light, not in your shadow when examining it. Charac-
teristics that require a microscope to be seen are omitted in this book.
Where vein patterns must be examined, use strong back lighting. If the blade is too thick, look for dis-
colored portions where the veins might be more visible. Failing this, a small piece of the blade can be
soaked in lye—a kitchen drain-opener such as Draino works well. Soak the tissue until it clears and the veins
become visible; this will usually take one or two days unless the blade is very thick. Drain and rinse the tis-
sue thoroughly because lye and drain-openers are extremely caustic. Veins may become more visible on
fronds soaked in water for a week or two as the softer tissue disintegrates.
Sometimes you need to examine cross sections of the stipes to see the number and arrangement of the
vascular bundles (conducting tissue). To do this, cut the stipe near the base and examine its cut surface. Use
a sharp blade because a dull one will tear and distort the tissue.
Ferns can be identified by using floras, or books that treat the plants growing in a given geographical
area. For fern genera from anywhere in the world, try Kramer and Green (1990); for the American tropics,
try Tryon and Tryon (1982); for ferns in Canada and the United States, see Mickel (1979), Lellinger
(1985), and Flora of North America Editorial Committee (1993). Lellinger (1994) includes a list of floras
in print.
Sometimes botanists at universities, botanical gardens, and museums will agree to identify ferns or
know of someone who will. Contact them first and ask for permission to send ferns for identification, and
ask how the ferns should be prepared for shipment. The task of identification is greatly simplified if a por-
tion of the rhizome with intact hairs or scales is included with the sterile and fertile fronds. Moreover, it is
important to know the fern’s country of origin and (especially in the tropics) its elevation. Small pieces of
fronds are unsuitable for identification.
 156 C 13

Acrostichum  13.1.1, 2 sake of “acrostichoid sori,” a term used to describe sori in
Leather fern which the sporangia are spread across the lower surface of
The genus Acrostichum includes bold-looking ferns with the leaf, making the entire sorus resemble a mat of brown
large, erect fronds up to 4 m (13 ft.) tall. Although the felt. Acrostichoid sori always lack indusia. This type of
plants grow tall in the wild, they tend to be shorter in cul- sorus characterizes Acrostichum and other, unrelated genera
tivation. In tropical climates or large indoor spaces with ad- such as Elaphoglossum and Platycerium.
equate light, they can be grown in ponds or in pots kept The genus occurs throughout the tropics, thriving pri-
wet or set in water. Acrostichum is subject to damage by marily in mangroves and brackish marshes. Its name comes
scale insects and slugs. from the Greek akros, summit, and stichos, row, referring to
All three species of Acrostichum have fronds that are one- the fertile pinnae borne toward the leaf apex in Acrostichum
pinnate and leathery, with veins closely and uniformly net- aureum.
ted. The fertile pinnae are slightly narrower than the sterile
ones. The sporangia cover the entire lower surface of each Acrostichum aureum Linnaeus  13.1.3
pinna, and the sori lack indusia. Acrostichum is the name- Leather fern, coastal leather fern
Tender, thrives in Zone (9)10
A very large fern with erect rhizomes and fronds in clus-
ters. Best grown under high light in garden soil or potting
mix kept constantly wet. Acrostichum aureum can grow
with its stems submerged but is typically found rooting in
mud with the foliage held above water. It grows natively in
brackish water but can be cultivated in fresh water.

b
Figure 13.1.1. Acrostichum danaeifolium: habit.

Figure 13.1.2.
Acrostichum
danaeifolium: fertile Figure 13.1.3. Acrostichum
pinna (left) and sterile aureum: a. habit, bar = 25 cm
pinna (right). (10 in.); b. vein pattern, bar =
5 mm (0.2 in.). After Proctor
(1985).
 F  F A  C 157

The blades of this species have up to 30 pairs of pinnae together than those of A. aureum. If the plants are fertile,
that are glabrous on their lower surfaces. The veins form another distinction can be seen: the fertile pinnae of A.
areoles along the costae, and the areoles are about three danaeifolium typically occur along the entire length of the
times longer than wide. See Acrostichum danaeifolium for a leaf (on the basal, middle, and apical portions) whereas
comparison with that species. Acrostichum aureum occurs those of A. aureum occur only toward the apex (usually the
throughout the tropical regions of the world. apical one to seven pairs). In nature A. danaeifolium grows
along coasts as well as farther inland—thus the common
Acrostichum danaeifolium Langsdorff & Fischer name inland leather fern—whereas A. aureum grows only
 13.1.1, 2, 4 near the coast.
Giant fern, inland leather fern
Tender, Zone 9 Actiniopteris
A very large fern with erect rhizomes and fronds in clus- Grown as novelty plants for the attractive fan-shaped
ters. Best grown under high light in garden soil or potting fronds, Actiniopteris is a genus of small, terrestrial ferns with
mix kept constantly wet. short-creeping rhizomes and clustered fronds. Although at-
The fronds of Acrostichum danaeifolium tend to be taller tractive, the plants are difficult to grow. They seem to pre-
and more erect than those of A. aureum. The blades have fer moderately humid conditions, but excessive moisture
40–60 pairs of pinnae. The areoles formed by the veins around their roots should be avoided.
along the costae are less than three times longer than broad. Actiniopteris is characterized by fan-shaped blades re-
This species occurs only in the tropics of the New World. peatedly forked into linear segments. Along the segment
Acrostichum danaeifolium resembles A. aureum but can margins are the sori, which are protected by the enrolled
be distinguished by hairs on the lower surface of the blade. edges of the segments. In the wild the species grow in dry
The hairs are less than 0.5 mm long and are best seen with habitats, and the blades, when dry, bend toward the leaf
a hand lens. The pinnae of this species are also usually closer stalk.
The genus consists of five species and is native to Africa
and the arid parts of tropical Asia. Its name comes from the
Greek aktis, ray, and pteris, fern, alluding to the radiating
segments of the blade. For additional information on this
genus, see Pichi-Sermolli (1962) and the section on “Xero-
phytic Ferns” in Chapter 10 of this book.

Actiniopteris semiflabellata Pichi-Sermolli

 13.2.1, 2
Tender to semi-tender
b
A small fern with short-creeping rhizomes and close-set
stipes. Grows under high light in well-drained, sandy gar-
den soil; avoid overwatering and excessive humidity. Actini-
opteris semiflabellata is difficult to grow in cultivation.

Figure 13.1.4. Acrostichum

danaeifolium: a. habit, bar = 25 cm
(10 in.), after Proctor (1984); b. vein
pattern, bar = 5 mm (0.2 in.), after
Proctor (1985).

Figure 13.2.1. Actiniopteris semiflabellata: habit.

 158 C 13

leaving a clean break. The segments have free veins ending

in enlarged tips (hydathodes) on the upper surface. The
b sori are marginal, short, and covered by a round to cres-
cent-shaped indusium that differs in texture and color from
the blade.
Because of the dark stipes and rachises, the shape of the
pinnules, and the marginal sori, Adiantopsis resembles cer-
tain species of Adiantum. The genus name, in fact, means
“Adiantum-like,” and comes from adding the Greek suffix
-opsis, meaning like, to Adiantum. Adiantopsis consists of
about 10 species and is indigenous to tropical America and
Africa.

Adiantopsis radiata (Linnaeus) Fée  13.3.1

Tender to semi-tender
A small fern with short-creeping rhizomes. Best grown
under medium to high light in garden soil with sand or pot-
ting mix kept moist but well drained. Avoid excessive wa-
tering and poor drainage. This species is moderately diffi-
a cult to cultivate.

Figure 13.2.2. Actiniopteris semiflabellata: a. frond, bar =

5 cm (2 in.); b. sori and indusia, bar = 2 mm.

The base of the blade forms an angle of about 55 to 70

degrees, and the fertile blades consist of 10–24 closely
placed, linear segments, with the apex of each segment end-
ing in a tooth. The species occurs in eastern Africa, Arabia,
Madagascar, Mauritius, and Nepal.
Actiniopteris semiflabellata is often misidentified in the
trade as A. radiata (Swartz) Link, a species not cultivated in
the United States. Actiniopteris radiata differs by its wide,
fan-shaped blades that form a 150- to 180-degree angle at
their base, and the apices of the broadest fertile segments
have two to six teeth.

Adiantopsis
This genus consists of small- to medium-sized, terrestrial or
rock-inhabiting ferns. Only one species is cultivated, grown
primarily for the attractive radiating arrangement of its pin-
nae; other species have pedate or pinnate fronds. All species
in the genus have erect or short-creeping and decumbent
rhizomes. Adiantopsis is closely related to Cheilanthes but
can be distinguished by the grooved upper surface of the
rachis and the sharp wings on either side of the groove. The Figure 13.3.1. Adiantopsis radiata: fronds, bar = 5 cm
ultimate segments are asymmetrical and eventually fall off, (2 in.).
 F  F A  C 159

Adiantopsis radiata has blades that consist of four to nine Hoshizaki (1970a), and the section on “Maidenhair Ferns”
one-pinnate pinnae radiating from the top of the stipe. The in Chapter 10.
species occurs throughout most of the American tropics.
Adiantum aethiopicum Linnaeus  13.4.2
Adiantum  13.4.1 Semi-tender, Zone 8
Maidenhair fern A small to medium, clump-forming fern with fronds in
Most maidenhairs are small- to medium-sized, terrestrial loose-spreading clusters on medium- to short-creeping rhi-
ferns prized for their fine, billowy foliage and shiny dark zomes. Best grown under medium light in moist garden
stalks. They generally require medium light, humidity, and soil or potting mix. This species is easy to grow.
moist soil but will not tolerate soggy soil. For this reason, Adiantum aethiopicum is characterized by finely divided
keep the plants well drained, and do not overpot. Adiantum blades up to three- or four-pinnate and by ultimate seg-
is mostly tender, but a few species are hardy or semi-hardy. ments that are fan-shaped to broadly obovate, with up to
three deep notches on the distal margin. Typically each seg-
ment has one to three sori. The plants are completely
glabrous, and in fresh material the base of the segment
stalks have an inconspicuous pale green swelling. The in-
dusium is kidney-shaped with a deep, C-shaped sinus. The
species is native to South Africa, Australia, New Zealand,
and New Caledonia. The specific epithet aethiopicum sug-

Figure 13.4.1. Adiantum capillus-veneris: habit.

Adiantum is characterized by shiny black or chestnut-

brown stipes and sori borne at the margins of the segments.
The indusium is formed by the enrolled margin of the seg- b
ment, and sporangia are borne on its inner surface. This
modified leaf margin is called a false indusium, in contrast
to a true indusium which is formed by an outgrowth of the
epidermis on the lower surface of the leaf. False indusia
occur in such related genera as Cheilanthes, Pellaea, and
Pteris, but only in Adiantum are the sporangia borne on the
false indusium, not beneath it on the opposing leaf tissue.
Another distinctive character of Adiantum is that the
leaflets often lack midribs, and the veins are repeatedly and
evenly forked. Adiantopsis, a similar genus, is distinguished
by the grooving on the upper surfaces of the rachises and
costae, with sharp wings on either side of the groove. In
contrast, Adiantum species (except for Adiantum flabellula-
tum) have rounded and ungrooved rachises and costae.
Adiantum occurs worldwide and consists of about 200
species, most of which grow in the American tropics. The a
fronds of these species have the unusual property that,
when wetted, water beads-up into silvery droplets that
quickly roll off. The genus name comes from the Greek
adiantos, meaning unwettable. For a further account of the Figure 13.4.2. Adiantum aethiopicum: a. frond, bar =
cultivated species and their cultivars, see Goudey (1985), 5 cm (2 in.); b. fertile leaflets, bar = 1 cm (0.4 in.).
 160 C 13

gests that the species occurs in northern Africa; however, in The bluish green fronds are pedate with one-pinnate
Africa it occurs only near the Cape. branches. The blade is fan-shaped or funnel-shaped in out-
Most of the plants sold as Adiantum aethiopicum in the line. The medial segments are more than three times longer
United States are actually A. raddianum ‘Triumph’, which than they are broad or somewhat long-triangular or reni-
can be distinguished by its lack of a swelling at the base of form, their tips are sharply denticulate-lobed, and the lobes
the segment stalk and by veins that end in minute sinuses are separated by deep sinuses, 0.6–4.0 mm deep. Adiantum
instead of teeth. Adiantum capillus-veneris, a similar spe- aleuticum is indigenous to the western United States and
cies, differs by oblong indusia and the absence of swellings western Canada.
on the segment stalk base. This species is very close to Adiantum pedatum, an east-
ern United States species that differs in having a rounder
Adiantum aleuticum (Ruprecht) C. A. Paris blade outline and shorter medial segments with rounded,
 13.4.3 crenulate or crenulate-toothed lobes. The lobes are sepa-
syn. Adiantum pedatum subsp. aleuticum Ruprecht rated by shallow sinuses (mostly 0.1–2.0 mm deep).
Western five-finger fern In the wild this species often grows on serpentine, a
Hardy, Zone 4(5) metamorphic rock that is often green and slippery to the
A small to medium fern with fronds in clusters on a clump- touch. Serpentine has several traits inimical to plant
forming rhizome. Grows in low to medium light in moist growth. It is low in essential nutrients, such as nitrogen,
potting mix. Adiantum aleuticum can be difficult to grow in calcium, potassium, and phosphorous, and high in toxic
arid climates. elements such as nickel and chromium. Its pH may be
either highly acidic or basic. Soils derived from serpentine
are often sterile and support unusual endemic plants.
subsp. subpumilum (W. H. Wagner) Lellinger (forma
imbricatum Hoshizaki; ‘Imbricatum’). Dwarf western five-
finger fern. Smaller than the typical subspecies, with
densely overlapping leaflets.

Adiantum anceps Maxon & C. V. Morton

 13.4.4
Two-edged maidenhair
Tender
A medium-sized fern with fronds in loose clusters from a
thick, short-creeping rhizome. Grows under low to me-
dium light in moist potting mix.
Adiantum anceps has two- to three-pinnate blades with
large, ovate to rhomboid segments that have a concavely
long-acuminate apex. It resembles A. peruvianum but has
larger, less rhomboid segments and concavely elongate tips.
Adiantum anceps is native to the Andes of Colombia, Ecua-
dor, and Peru.

Adiantum bellum T. Moore  13.4.5

b Bermuda maidenhair
Semi-tender
A small to medium fern with fronds in loose clusters from
a clump-forming rhizome. Grows in low to medium light
in moist potting mix.
Adiantum bellum resembles A. capillus-veneris (and
a might be the same) but apparently differs in that the oblong
indusia have irregularly jagged, rather than entire, margins.
It is native to Bermuda and Guyana.
Some forms superficially resemble Adiantum tenerum
‘Farleyense’. Some of the trade material circulating as A.
bellum is actually A. raddianum ‘Pacottii.’
Figure 13.4.3. Adiantum aleuticum: a. habit, bar = 5 cm
(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
 F  F A  C 161

Adiantum capillus-veneris Linnaeus

 13.4.1, 6, 7
Southern maidenhair, Venus’s hair
Semi-hardy, Zone 7
A small to medium fern with fronds in loose clusters from
a much-branched, medium- to short-creeping rhizome.
Grows best under low to medium light in moist, basic gar-
den soil or potting mix. Adiantum capillus-veneris is easy to
b grow and is commonly cultivated. The plants die back with
frost. They reportedly do not grow well in the Seattle area
(Zone 8), perhaps due to soil that is too acidic.
Adiantum capillus-veneris is characterized by three- to
four-pinnate blades and wedge-shaped segments with veins
ending in marginal teeth. The color of the segment stalk
gradually merges into the green of the veins, and the indu-
sia are oblong and entire. Adiantum capillus-veneris is one of
the most widely distributed ferns in the world, occurring
primarily in warm-temperate to subtropical areas. It often
a grows on calcareous soil or rocks, especially around springs
or caves. It is sometimes cultivated on limestone. This spe-
cies is often confused with the commonly grown A. raddi-
Figure 13.4.4. Adiantum anceps: a. frond, bar = 5 cm anum and A. tenerum.
(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.). ‘Fimbriatum’. Figure 13.4.7a. The distal margins of the
segments are fimbriate.

a Figure 13.4.5. Adiantum

bellum: a. habit, bar = 5 cm
(2 in.); b. fertile leaflets,
bar = 1 cm (0.4 in.).
Figure 13.4.6. Adiantum capillus-veneris: a. frond, bar =
5 cm (2 in.); b. fertile leaflets, bar = 1 cm (0.4 in.).
 162 C 13

c
b

Figure 13.4.7. Adiantum capillus-veneris cultivars:

a. ‘Fimbriatum’; b. ‘Imbricatum’; c. ‘Mairisii’. Bar =
5 cm (2 in.). a

‘Imbricatum’. Green petticoats. Figure 13.4.7b. Fronds

tend to be decumbent and have ruffled, densely overlapping Figure 13.4.8. Adiantum caudatum: a. fronds, bar =
segments. A beautiful plant, but more difficult to grow than 5 cm (2 in.); b. fertile leaflets, bar = 1 cm (0.4 in.).
the other cultivars. Avoid overhead watering. Semi-tender.
‘Mairisii’ (‘Banksianum’). Figure 13.4.7c. Sparingly
branched rhizomes up to 9 mm (0.4 in.) thick—thicker
than those of the other cultivars. Stipes are erect and the pinnate, and bear a bud at the end of a whip-like tip. The
blades broadly triangular to triangular, mostly three-pin- pinnae are often coarsely lobed. This species is native to
nate, with segments asymmetrical, long-subrhomboid to tropical Africa, India, and Asia.
triangular. Appears to be more cold hardy than the species.
This selection apparently arose in the 1800s at the nursery Adiantum concinnum Humboldt & Bonpland ex
of Mairis & Co., England, and was described as Adiantum Willdenow  13.4.9
×mairisii T. Moore. Rush (1983) suggests that it was of hy- Brittle maidenhair
brid origin, resulting from a cross between typical A. capil- Tender
lus-veneris and A. aethiopicum. A small to medium fern with clump-forming rhizomes,
glabrous stipes, and broadly triangular fronds. Grows in
Adiantum caudatum Linnaeus  13.4.8 low to medium light in moist potting mix.
Trailing maidenhair Adiantum concinnum can be distinguished by the in-
Tender nermost segments of the pinnae overlapping the rachis. It is
A small to medium fern with nearly erect rhizomes and native to tropical America, where it is common and some-
fronds that arch downward. Grows in medium light in times weedy. The plants often grow on stone walls and
moist potting mix. The plants are best displayed in a hang- steep banks.
ing container. ‘Edwinii’. Fronds and segments broader and larger than
Adiantum caudatum has extremely distinctive fronds: the typical species, with the inner upper pinnules hardly
they are pendent, up to 50 cm (20 in.) long, narrow, one- overlapping the rachis. The cultivar is easier to grow and is
 F  F A  C 163

Figure 13.4.10. Adiantum cultratum: a. frond, bar =

5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).

a Adiantum diaphanum Blume  13.4.11

Filmy maidenhair
Semi-tender
Figure 13.4.9. Adiantum concinnum: a. frond, bar = A small fern with fronds in tufts on a short, semi-erect rhi-
5 cm (2 in.); b. leaflet pair at base of pinna, bar = 1 cm zome. Grows under low to medium light in moist potting
(0.4 in.). mix with a lot of peat.

less tender than the species form. It might be a hybrid of

Adiantum concinnum and A. raddianum.

Adiantum cultratum J. Smith ex Hooker

 13.4.10
Tender b
A medium-sized fern with fronds in a loose cluster from
slender, creeping rhizomes. Grows under high-medium
light in potting mix kept moist.
Adiantum cultratum resembles A. trapeziforme but dif-
fers by the slender, creeping rhizomes, narrower segments
two to two-and-a-half times longer than broad, and shorter
stalks (2–3 mm long). It is native to Mexico.
This species needs more study. It was originally de- a
scribed from the Lesser Antilles (St. Vincent) but has not
been found there since. Adiantum cultratum is currently
known from Mexico and has been confused with the plant
traditionally called A. trapeziforme.

Figure 13.4.11. Adiantum diaphanum: a. frond, bar =

5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
 164 C 13

Adiantum diaphanum is characterized by smooth stipes, Adiantum flabellulatum Linnaeus  13.4.13
and blades that are pinnate to some having one pair of pin- Tender or slightly hardier
nae at the base. The membranous segments bear black or A small to medium fern with fronds in a loose cluster from
reddish brown hairs on the lower surface. The indusium a clump-forming rhizome. Grows under medium light in
varies from round to nearly kidney-shaped and usually has potting mix kept moist. This species is easy to start from
dark hairs on the surface. An extremely unusual character- spores.
istic is the small barrel-shaped tubers borne by the roots.
The species is native to Asia, Australia, New Zealand, and b
the Pacific Islands. It is a delicate fern that in nature usually
thrives near water.

Adiantum excisum Kunze  13.4.12

Chilean maidenhair
Tender
A small fern with clustered fronds from clump-forming rhi-
zomes. Grows in high to medium light in moist potting
mix. The plants need humidity and can be difficult to grow.
Adiantum excisum resembles a delicate A. raddianum but
differs by having dark green and airy foliage, often zigzag
rachises, and broad segments with deep, spreading lobes.
It is native to Chile.

Figure 13.4.13. Adiantum flabellulatum: a. fronds, bar =

5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).

Adiantum flabellulatum has pedate blade architecture,

which only a few species of maidenhair have (such as A.
pedatum), and its stipes are grooved on the upper surface,
unlike those of all other species of Adiantum. This species
can be further distinguished by the short, stiff, brownish
hairs on the upper (but not lower) side of the stipe and
rachis, and by the glabrous segments. The plants resemble
a stiff A. pedatum, but the blades are not bluish green as in
that species, and the segments are entire, not incised. The
similar A. hispidulum differs by the presence of whitish hairs
on the segment surfaces. Adiantum flabellulatum occurs
from India west to China, Japan, and the Philippines.

Adiantum formosum R. Brown  13.4.14

a Australian maidenhair
Semi-hardy to semi-tender
A medium to large fern with fronds spaced far apart on a
Figure 13.4.12. Adiantum excisum: a. frond, bar = 5 cm long-creeping, branched rhizome. Grows under medium
(2 in.); b. fertile leaflets, bar = 1 mm (0.4 in.). to low light in moist garden soil or potting mix. Because of
 F  F A  C 165

b b

Figure 13.4.15.
Adiantum fragile:
a. frond, bar = 5 cm
(2 in.); b. leaflets
a shedding, bar = 1 cm
a (0.4 in.).

Figure 13.4.14. Adiantum formosum: a. frond, bar =

5 cm (2 in.); b. fertile leaflets, bar = 5 mm (0.2 in.).
Adiantum fructuosum Poeppig ex Sprengel
 13.4.16
its long-creeping rhizome, Adiantum formosum is best kept Tender
confined in a container. Withhold fertilizer to keep the A medium-sized fern with fronds clustered on a short,
fronds smaller. Divisions must be made with care because knotted rhizome. Grows under low to medium light in
the deep-growing rhizomes and stipe bases break easily. moist garden soil or potting mix.
Adiantum formosum is characterized by large, broadly Adiantum fructuosum is characterized by bicolorous rhi-
triangular fronds that are up to five-pinnate. The pinnae zome scales; stipes adjacent to each other; blades two-
get smaller as they go from the base to the apex of the blade.
The basal pinnae are by far the largest and are usually nar-
rowly triangular. The species is native to Australia and New
Zealand.

Adiantum fragile Swartz  13.4.15

Tender
A small to medium fern with fronds in a loose cluster from
a clump-forming rhizome. Grows under medium light in
moist, basic, well-drained garden soil or potting mix.
Adiantum fragile is characterized by bright yellowish
brown rhizome scales; lance-ovate blades that are three- to
four-pinnate at the base; delicately herbaceous, light green,
segments that easily fall off; veins ending in marginal teeth;
and subreniform to oblong indusia. It resembles A. tenerum
but can be distinguished by the color of the rhizome scales.
Adiantum fragile is native to the West Indies, where it often a
grows on limestone ledges, cliffs, and crevices of old stone b
walls.

Figure 13.4.16. Adiantum fructuosum: a. frond, bar =

5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
 166 C 13

pinnate with a terminal pinna resembling the lateral ones; Some young plants of Adiantum hispidulum are circu-
fertile pinnules with tips mostly straight and obtuse, not lating as Adiantum ‘Birkenheadii’, a name originally used in
falcate; and round apices. The blade features often overlap early English horticultural literature for a plant thought to
with A. tetraphyllum, which is distinguished by a long- have originated from A. diaphanum.
creeping rhizome and more falcate segments. Adiantum
fructuosum is native to the American tropics. Adiantum jordanii K. Müller  13.4.18
California maidenhair
Adiantum hispidulum Swartz Semi-hardy to semi-tender
 13.4.17;  9 A small to medium fern with fronds in a cluster from a
Rosy maidenhair, rough maidenhair clump-forming rhizome. Best grown under high light in
Semi-hardy or hardier, Zone 7 moist-dry garden soil or potting mix. The leaves usually
A small fern with clustered fronds from a clump-forming wither in summer, and the soil should be kept drier at this
rhizome. Grows in low to high light in moist to moist-dry time. This species is difficult to grow.
garden soil or potting mix. Adiantum hispidulum tolerates
drier conditions than other maidenhairs. The plants are
easy to cultivate and frequently seen in the United States
trade. Young fronds are reddish.
Adiantum hispidulum is characterized by hairy stipes, pe-
date blades that are firm and have whitish hairs on the sur-
face, and rounded indusia with slender, reddish brown hairs
on the surface. The species is native to southern India, east-
ern Africa, and the Pacific Islands. It has escaped from cul-
tivation in the United States (Connecticut and Georgia). b
The name Adiantum pubescens Schkuhr applies to a vari-
ant with thinner and laxer hairs.

b a

Figure 13.4.18. Adiantum jordanii:

a. frond, bar = 5 cm (2 in.);
b. fertile leaflet, bar = 1 cm (0.4 in.).

Adiantum jordanii resembles A. capillus-veneris but has

broad, fan-shaped segments that are cleft or entire. Adi-
antum jordanii is native to the western United States (Cali-
fornia and Oregon) and Mexico (Baja California). It hy-
bridizes with A. aleuticum to form the sterile hybrid A.
×tracyi.
a
Adiantum latifolium Lamarck  13.4.19
Tender
A medium-sized fern with fronds widely spaced along a
Figure 13.4.17. Adiantum hispidulum: a. fronds, bar = slender, creeping rhizome. Grows under low to medium
5 cm (2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.). light in moist garden soil or potting mix.
 F  F A  C 167

b
b

Figure 13.4.19. Adiantum

latifolium: a. habit, bar = 5 cm
a (2 in.); b. fertile leaflets, bar =
1 cm (0.4 in.).
a

Figure 13.4.20. Adiantum lucidum: a. fronds, bar = 5 cm

(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
Adiantum latifolium bears two-pinnate blades with up to
four pairs of lateral pinnae and a terminal pinna that re-
sembles the lateral ones. The segments are often glaucous
beneath with a faint midrib basally, and the sterile segments Adiantum lunulatum N. L. Burman  13.4.21
are evenly serrate or denticulate. The oblong indusia are on syn. Adiantum philippense Linnaeus
the upper and sometimes lower margins of the segments. Tender
This species is native to tropical America and has escaped in A small to medium fern with erect or suberect rhizomes
Thailand and peninsular Malaya. and arching fronds. Grows under medium light in well-
drained garden soil or potting mix. The plants do best in
Adiantum lucidum (Cavanilles) Swartz humid conditions and are difficult to grow.
 13.4.20 A delicate fern, Adiantum lunulatum sometimes roots
Tender at the leaf tips. It is characterized by one-pinnate blades,
A small to medium fern with clustered fronds from a long-stalked pinnae with entire or bluntly toothed sterile
clump-forming rhizome. Grows under medium light in margins, and oblong-linear indusia. It grows throughout
moist potting mix. the tropics of the world.
Only a few species of Adiantum have a single long sorus
along both the upper and lower margins of the pinnae, and Adiantum macrophyllum Swartz  13.4.22
Adiantum lucidum is one such species. It can be distin- Large-leaved maidenhair
guished by the combination of this soral characteristic and Tender
one-pinnate blades, scaly rachises, and free veins. Other A medium-sized fern with clustered fronds from a short-
characteristics include short-creeping rhizomes, asymmet- creeping to semi-erect rhizome. Grows under low to me-
rical pinna bases, and serrate pinna tips (where sterile). This dium light in moist potting mix. New fronds are often red
species is native to lowland rain forests in tropical America. or pink.
 168 C 13

Figure 13.4.23. Adiantum

monochlamys: a. frond, bar =
Figure 13.4.21. Adiantum lunulatum: 5 cm (2 in.); b. fertile
frond, bar = 5 cm (2 in.). leaflets, bar = 1 cm (0.4 in.).

b
a
a b

Figure 13.4.22. Adiantum

macrophyllum: a. frond, bar = Adiantum patens Willdenow  13.4.24
5 cm (2 in.); b. fertile leaflet, Tender
bar = 1 cm (0.4 in.). A small to medium fern with clustered fronds from a
clump-forming rhizome. Grows under medium light in
potting mix kept moist. This species is difficult to cultivate.
Like Adiantum lucidum, A. macrophyllum has a single Adiantum patens resembles A. pedatum because of its
long sorus on both the upper and lower margins of the pin- pedate blades but differs by having crenate (not incised)
nae, but it can be distinguished from A. lucidum by its segment margins, rachises beset with minute whitish hairs
glabrous rachis. In addition, A. macrophyllum has one-pin- (not glabrous), and round-reniform (not oblong) indusia.
nate blades and wide, sessile (or nearly so) pinnae that are The species occurs in the American tropics and Africa. The
glaucous beneath. It is native to the American tropics. African plants are often distinguished as subspecies oatesii
(Baker) Schelpe, but they hardly differ and should be con-
Adiantum monochlamys D. C. Eaton sidered the same.
 13.4.23;  10
Hardy, Zone 6 Adiantum pedatum Linnaeus  13.4.25
A small fern with fronds clustered on short-creeping, American maidenhair, five-finger fern
clump-forming rhizomes. Grows under low to medium Hardy to −37°C (−35°F), Zone 3
light in moist potting mix. A small to medium fern, generally 30–60 cm (12–24 in.)
Adiantum monochlamys resembles A. capillus-veneris but tall, with fronds in clusters from the clump-forming rhi-
has narrower wedge-shaped segments and usually only one zome. Grows under low to medium light in moist garden
sorus per segment. It is native to China, South Korea, and soil or potting mix. This species is difficult to grow in arid
Japan. climates.
 F  F A  C 169

The most distinctive characteristics of Adiantum peda-

tum are its pedately divided blades, which are circular or
nearly so in outline, and its incised segments. The leaf tis-
sue is usually bluish green. The species resembles A. patens
but differs by its glabrous stipes and rachises. It occurs in
the central and eastern United States, Canada, and eastern
Asia. Related plants of the western United States are now
called A. aleuticum.

Adiantum pentadactylon Langsdorff & Fischer

 13.4.26
Tender
A medium-sized fern with fronds clustered on a clump-
forming rhizome. Grows under low to medium light in
moist potting mix.
Adiantum pentadactylon resembles A. trapeziforme but
a
b differs in that the dark color of the segment stalk merges
into the veins, rather than stopping abruptly, and its seg-
ments are more deeply lobed and have attenuate apices. It
is native to Brazil.

Figure 13.4.24. Adiantum patens: a. frond, bar = 5 cm

(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).

Figure 13.4.25. Adiantum pedatum: frond, bar = 5 cm Figure 13.4.26. Adiantum pentadactylon: a. frond, bar =
(2 in.). 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
 170 C 13

Adiantum peruvianum Klotzsch  13.4.27

Silver-dollar fern, Peruvian maidenhair
Tender
A medium-sized fern with fronds clustered from a short-
creeping to clump-forming rhizome. Grows under medium
light in moist potting mix.
Adiantum peruvianum has large, rhombic, shiny seg-
ments borne on black slender stalks. The color of the stalk
stops abruptly at the base of the segment; it does not enter
the segment and gradually merge with the green of the
veins as in some other species. Adiantum peruvianum is b
more common in collections than the similar A. anceps,
from which it differs by having acute or rounded segment
tips. It differs from A. trapeziforme, another similar species,
by its inconspicuous serration on the pinnules. The species
a
is native to Ecuador, Peru, and Bolivia.

Adiantum petiolatum Desvaux  13.4.28

Tender
A small to medium fern with fronds distributed along a
creeping rhizome. Grows under medium light in moist pot-
ting mix.

Figure 13.4.28. Adiantum

petiolatum: a. frond, bar =
5 cm (2 in.); b. fertile leaflet,
bar = 1 cm (0.4 in.).

The distinctive characteristics of Adiantum petiolatum

are the long-creeping rhizomes, usually well-spaced fronds,
b
one-pinnate blades, pinnae that are dull or glaucous be-
neath, and sterile segments with margins evenly serrate.
The pinna bases tend to be broadly humped on the for-
ward margin and often overlap the rachis. The species is
widespread in tropical America.

Adiantum poiretti Wikström  13.4.29

Tender
A small to medium fern with fronds in loose clusters from
a long-creeping rhizome. Grows under medium light in
a moist garden soil or potting mix.
Adiantum poiretti belongs to a group of species charac-
terized by finely divided, billowy blades with stalked and
small ultimate segments. This species can be distinguished
from others in the group (such as A. concinnum and A.
tenerum) by its long-creeping rhizomes and oblong-lunate
sori. A yellow powder is frequently mixed among the spo-
rangia, and if present the color is usually visible through
the indusium. Other traits include golden-brown rhizome
scales, blades up to three-pinnate, jointed segments that
are wider than they are long, and sterile segments with
Figure 13.4.27. Adiantum peruvianum: a. frond, bar = poorly defined teeth. Adiantum poiretti is native to tropical
5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.). America.
 F  F A  C 171

Adiantum pulverulentum Linnaeus  13.4.31

Tender
A medium to large fern with clustered fronds from a short-
creeping rhizome. Grows in medium light in moist garden
soil or potting mix.
Adiantum pulverulentum has two-pinnate blades with a
b terminal pinna resembling the lateral ones. This character-
istic, plus the single sorus along the upper margin of the
pinnules, makes A. pulverulentum one of the easiest maid-
enhairs to identify. (Other species with similar two-pinnate
blades have more than one sorus along a pinnule margin.)
The species is widespread in tropical America.

a Figure 13.4.29. Adiantum

poiretti: a. frond, bar = 5 cm
(2 in.); b. fertile leaflet, bar =
b
1 cm (0.4 in.).

Adiantum polyphyllum Willdenow  13.4.30

Giant maidenhair
Tender
A large fern with clustered fronds from a clump-forming
rhizome. Grows under medium light in moist potting soil.
Adiantum polyphyllum is a large, stiff fern with many
jointed segments. The blades are broadly triangular and up
to four-pinnate at the base. The lower pinnae have the pin-
nules on the basiscopic side with a pair of longer branches.
Up to six indusia are found on the upper margin of the seg-
ment. The species occurs from Venezuela to Peru.

b
Figure 13.4.31. Adiantum pulverulentum: a. frond, bar =
5 cm (2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.).

Adiantum pyramidale (Linnaeus) Willdenow

 13.4.32
syn. Adiantum cristatum Linnaeus
Tender
A medium-sized fern with fronds clustered from a short-
creeping, clump-forming rhizome. Grows under medium
a to high light in garden soil or moist potting mix.
Adiantum pyramidale can be identified by the small,
hard bumps on the stipes; blades to three-pinnate at the
base; the basal pinnae, which normally have a short to elon-
gate branch on the basiscopic side; and oval to oblong,
Figure 13.4.30. Adiantum polyphyllum: a. frond, bar = nearly straight indusia, mainly on the upper margin and
5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.). sometimes around the apex. It is native to the West Indies.
 172 C 13

b
b

Figure 13.4.32. Adiantum pyramidale: a. frond, bar = Figure 13.4.33. Adiantum raddianum: a. frond, bar =
5 cm (2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.). 5 cm (2 in.); b. fertile leaflets, bar = 1 cm (0.4 in.).

Adiantum raddianum C. Presl  13.4.33, 34

syn. Adiantum cuneatum Langsdorff & Fischer a
Delta maidenhair
Semi-tender
A small to medium fern with clustered fronds from a
clump-forming rhizome. Thrives in low to high light in
moist garden soil or potting mix. The foliage dies back with
light frosts.
Adiantum raddianum is characterized by mostly trian- b
gular, three- to four-pinnate blades, wedge-shaped seg-
ments without hairs, veins ending in a sinus between the
teeth, and rounded indusia. It is native to the American
tropics.
The delta maidenhair is the most commonly cultivated
maidenhair in the United States. It has numerous, often
poorly defined and confused cultivars. Some cultivars in
the current trade not listed here include ‘Fragrans’, ‘Fra-
grantissimum’, ‘Grandiceps’, ‘Lady Geneva’, ‘Micropin-
nulum’, and ‘Weigandii’. d
‘Croweanum’. Crowe maidenhair. Fronds more trian-
gular, more foliaceous; fronds and segments larger than
those of the species type. Easy to grow. c
‘Fritz Luth’. Figure 13.4.34a. Fronds triangular with
even edges, flattened or with pinnae spreading over the next
upper one, growth moderately compact, yet lacy and airy.
‘Gracillimum’. Figure 13.4.34b. Blades up to five-pin- Figure 13.4.34. Adiantum raddianum cultivars: a. ‘Fritz
nate, with very small segments, 2–4 mm (0.1–0.2 in.) long. Luth’; b. ‘Gracillimum’; c. ‘Pacific Maid’; d. ‘Pacottii’.
Growth is loose but fine-textured. Bar = 5 cm (2 in.).
 F  F A  C 173

‘Ocean Spray’. Resembles ‘Fritz Luth’, but the blades

are flatter and the pinnae are nearly all in the same plane.
Slower, more compact growth, to 46 cm (18 in.) tall.
‘Pacific Maid’. Pacific maid. Figure 13.4.34c. Fairly dis-
tinct by its crowded pinnae, pinnules, and segments. The
segments are broadly fan-shaped. It grows rapidly and is
looser and taller than ‘Pacottii’.
‘Pacottii’. (Adiantum bellum of trade, not of T. Moore).
Double maidenhair. Figure 13.4.34d. Small, triangular,
erect fronds; segments very densely placed; rarely more than
25 cm (10 in.) tall. Compared to other cultivars, it grows
slowly from spores and transplants.
‘Triumph’. Triumph maidenhair. Small-medium
fronds, usually about 20 cm (8 in.) long, often with tear-
drop-shaped segments. An extremely variable plant. This a
cultivar is sometimes misnamed as Adiantum aethiopicum. b
‘Variegatum’. Variegated maidenhair. Segments spar-
ingly variegated a cream-white. Grows slowly compared to
other cultivars.

Adiantum reniforme Linnaeus  13.4.35 Figure 13.4.36. Adiantum seemannii: a. frond, bar =
Tender to semi-tender 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
A small fern with fronds clustered from a short-creeping
rhizome. Grows under low to medium light in moist pot-
ting soil. This species is usually difficult to grow.
are usually several sori per pinna. This species resembles A.
Adiantum reniforme has small, kidney-shaped fronds
anceps, but its blade is whitish green (glaucous) beneath.
with entire margins. It is native to Africa (Kenya), Madeira,
Adiantum seemannii is native from Mexico to Panama and
the Canary Islands (Tenerife), and Comoros.
Colombia.

Adiantum tenerum Swartz  13.4.37, 38

Brittle maidenhair, fan maidenhair
Tender
b A small to medium fern with fronds clustered on a short-
creeping rhizome. Best grown under low to medium light
in moist, basic potting mix.
Adiantum tenerum resembles A. raddianum but is larger,
and it further differs by the dark color of its segment stalk
a
ending abruptly at the segment where it is enlarged into a
disk. The segments fall away freely (especially when dry),
Figure 13.4.35. Adiantum reniforme: a. frond, bar = and the veins end in a tooth. This species is native to trop-
5 cm (2 in.); b. indusia, bar = 5 mm (0.2 in.). ical America.
A number of cultivars exist, ‘Farleyense’ being the best
known.
‘Farleyense’. Farley maidenhair. Figure 13.4.38a. Fronds
Adiantum seemannii Hooker  13.4.36 tend to be more arching than the typical species; the seg-
Seemann’s maidenhair ment margins are ruffled and crowded. Avoid overhead wa-
Tender tering. This cultivar originated in about 1865 on a Barba-
A medium-sized fern with fronds clustered from a clump- dos sugar plantation named Farley Hill. The plant won
forming rhizome. Grows in low to medium light in moist instant acclaim in British horticultural circles and rapidly
potting mix. spread to gardens around the world. Rogers (1998) re-
Adiantum seemannii is characterized by broadly trian- viewed the history and classification of this cultivar.
gular, one- or two-pinnate blades with a terminal pinna re- ‘Fergusonii’. Figure 13.4.38b. Blades to three-pinnate
sembling the lateral ones. The pinnae are 5–8 cm (2–3 in.) but commonly two-pinnate, with pinnae long-stalked and
long, usually ovate, and have serrate sterile margins. There pinnules crowded toward the pinna tips.
174 C 13

Adiantum tetraphyllum Humbold & Bonpland ex

Willdenow  13.4.39
Four-leaved maidenhair
Tender
A medium-sized fern with fronds spread along a short- to
medium-creeping rhizome. Grows in moist garden soil or
b potting mix.
Adiantum tetraphyllum has two-pinnate blades with a
terminal segment resembling the lateral pinnae. The rhi-
zome scales are bicolorous, with dark centers and lighter
margins. Fronds have two to six pairs of lateral pinnae, and
these usually have slightly falcate apices. The pinnules are
1–4 cm (0.5–1.5 in.) long and have coarsely serrate margins
when sterile. The indusia are oblong to oblong-lunate with
short hairs on the upper surface, and they number three to
six along the upper margin of the segment. This species is
similar to A. fructuosum. It is widespread in the American
tropics.

Figure 13.4.37. Adiantum tenerum: a. frond, bar = 5 cm

(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.). b

b
a
a

Figure 13.4.38. Adiantum tenerum cultivars: Figure 13.4.39. Adiantum tetraphyllum: a. frond, bar =
a. ‘Farleyense’; b. ‘Fergusonii’. Bar = 5 cm (2 in.). 5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
 F  F A  C 175

Adiantum ×tracyi C. C. Hall ex W. H. Wagner

 13.4.40
Semi-hardy
A small fern with loosely clustered fronds from a clump-
forming rhizome. Grows under low to medium light in gar-
den soil or potting mix kept moist. b
Adiantum ×tracyi is characterized by broadly triangular
blades that taper abruptly from a four- to five-pinnate base
to a one-pinnate tip. It is native to the western United
States (California and Oregon).
Adiantum ×tracyi is a naturally occurring sterile hybrid
of A. aleuticum and A. jordanii. Its sterility is evinced by
aborted spores that appear misshapen and irregular when
seen under a microscope. These spores do not germinate,
and the plant can be propagated only by divisions of the
rhizome.

Figure 13.4.41. Adiantum trapeziforme: a. frond, bar =

5 cm (2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).
b

Adiantum venustum G. Don  13.4.42

Himalayan maidenhair
Hardy
A small fern with fronds slightly distant along a medium- to
short-creeping rhizome. Grows under low to medium light
in moist potting mix. Adiantum venustum is slow to estab-
lish and is deciduous in colder climates. It forms dense, uni-
form colonies that make a good border planting.
a Adiantum venustum is characterized by triangular blades
up to three-pinnate, obovate segments, and sterile segment
margins that are evenly and deeply dentate. The sori are

Figure 13.4.40. Adiantum ×tracyi: a. frond, bar = 5 cm

(2 in.); b. fertile leaflet, bar = 5 mm (0.2 in.).

b
Adiantum trapeziforme Linnaeus  13.4.41
Diamond maidenhair
Tender
A medium-sized fern with clustered fronds from a short-
creeping rhizome. Grows under low to medium light in
moist potting mix.
Adiantum trapeziforme is characterized by triangular Figure 13.4.42.
blades up to four-pinnate, with a terminal pinna similar to Adiantum venustum:
the lateral ones. The segments are trapeziform and stalked, a. frond, bar = 5 cm
with the stalks 4–7 mm (0.2–0.3 in.) long. The dark stalk (2 in.); b. fertile
a
color ends abruptly at the segment blade. This species is leaflets, bar = 1 cm
native to Central America and the West Indies. (0.4 in.).
 176 C 13

reniform and generally occur two per segment. This species

resembles some forms of A. raddianum, but the sterile seg-
ments are serrulate along the distal margin, not deeply
lobed. It grows along the Afghanistan-Indian border.

Adiantum villosum Linnaeus  13.4.43

Tender
A medium-sized fern with fronds clustered along a short-
creeping rhizome. Grows under medium light in moist gar-
den soil or potting mix.

Figure 13.5.1. Aglaomorpha coronans: habit.

b
organic debris that eventually decay to provide a soil for
the roots. The dry, papery texture of this humus-collecting
leaf-base resists decay.
Aglaomorpha coronans and the hybrid ×Aglaonaria
robertsii are the most frequently cultivated ferns in the
genus. They are usually planted in hanging baskets but can
also be grown in pots or even in the ground if given suffi-
cient drainage by using a coarse medium. Though large,
they are suitable as indoor plants because they tolerate low
humidity and irregular watering. When the fronds become
old, the pinnae fall off, but the rachises persist as long,
a brown spines. The shed pinnae litter the floor and can be-
come untidy.

Figure 13.4.43. Adiantum villosum: a. frond, bar = 5 cm

(2 in.); b. fertile leaflet, bar = 1 cm (0.4 in.).

Adiantum villosum has strongly nodose rhizomes and

dark green, shiny blades that are up to two-pinnate and
have terminal segments that are similar to but longer than
the lateral pinnae. The pinnules are oblong to narrowly
rhombic, with acuminate or acute tips and margins coarsely
serrate when sterile. The indusia are oblong or linear and
appear mostly on the upper margin. Adiantum villosum re-
sembles A. latifolium but differs by its short-creeping rhi-
zome. The species is native to the American tropics.

Aglaomorpha  13.5.1–3

syn. Merinthosorus, Photinopteris, Pseudodrynaria,
Thayeria
Aglaomorpha is a genus of handsome and large epiphytic
ferns that typically form a spreading cluster or nest of
fronds. The blades are coarsely cut and form a bold pat-
tern. The frond bases are often widened and turn brown Figure 13.5.2. Aglaomorpha coronans: frond
and papery with age. They collect falling leaves and other base.
 F  F A  C 177

Aglaomorpha cornucopia (Copeland) M. C. Roos

 13.5.4
syn. Thayeria cornucopia Copeland
Very tender to tender
A medium-sized fern with branched, long-creeping rhi-
zomes that tend to climb. Grows best under high light in
potting mix or uncut moss kept moist-dry and well
drained. It forms a nice display if allowed to climb.
The fronds of Aglaomorpha cornucopia are up to 60 cm
(24 in.) long, sessile, and borne singly on special rhizome
branches. The blade base wraps around the rhizome and is
shaped like a cornucopia or oblong basket, becoming dry
and papery. The blade above is green and pinnatifid. Fertile
pinnae occur in the apical portion of the frond and resem-
ble a string of beads, being constricted between the round
sori. The species is native to the Philippines.

Figure 13.5.3. Aglaomorpha coronans: sori.

The genus is recognizable by the following characteris-

tics: rhizomes short- to long-creeping, thick, fleshy, and
scaly; fronds variable, some species with fronds entirely fo-
liaceous, some with the basal part of the frond usually en-
larged, dry, and papery (humus-collecting) and the upper
part foliaceous; the foliaceous part one-pinnate or more
often pinnatifid; pinnae jointed (articulate), with a true ter-
minal pinna (segment) present; main veins raised and con- Figure 13.5.4. Aglaomorpha cornucopia: habit with fertile
nected nearly at right angles by cross-veins; sporangia in frond, bar = 10 cm (4 in.). Courtesy of T. Hoshizaki.
discrete sori or spreading; no indusia.
Aglaomorpha was formerly defined as including only
those plants bearing widened frond bases that turn brown
and papery, deeply pinnate lobing at the upper part of the Aglaomorpha coronans (Wallich ex Mettenius)
fronds, and discrete sori (not fused or acrostichoid). Nowa- Copeland  13.5.1–3, 5
days, however, it is defined in a broader sense to include Semi-tender or hardier
plants formerly placed in Merinthosorus, Photinopteris, and A medium to large fern with branched, short- to medium-
Thayeria. The characteristics of Merinthosorus are discussed creeping rhizomes. Best grown under high light in well-
under Aglaomorpha drynarioides; those of Photinopteris, drained, moist-dry potting mix or uncut moss. The species
under A. speciosa; and Thayeria, under A. cornucopia. Some is easy to grow and thrives in both humid and drier envi-
botanists still recognize Photinopteris because of its fully ronments. It can be cultivated in the ground and in hang-
one-pinnate fronds and the absence of humus-collecting ing baskets. Aglaomorpha coronans is the most commonly
frond bases. In this book we distinguish Drynaria, which is cultivated species in the genus, a testament to its durability
closely related to Aglaomorpha, by the separate humus-col- and robust growth.
lecting and foliaceous fronds and the absence of a true ter- The fronds of Aglaomorpha coronans are stiff, pinnati-
minal pinna or lobe on the foliaceous fronds. Drynaria can fid, and sessile. They become brown and papery at the base
hybridize with Aglaomorpha, and one such hybrid, ×Aglao- but are shiny dark green above. The pinnae are connected
naria robertsii, is commonly cultivated. to each other by wings, and the fertile pinnae are unmodi-
Aglaomorpha contains 14 species and occurs from the fied. The sori vary from oblong to linear, and several occur
Asiatic tropics to New Guinea and nearby islands. See in a row between the lateral veins. This species is native to
Chapter 10, the section “Polypodium and Relatives,” for Bangladesh, India, China, Taiwan, southeastern Asia, Ma-
more information on culture; for a monographic treatment laysia, and the Ryuku Islands of Japan.
of the group, see Roos (1985). The genus name comes from ‘Angeli’. Fronds small, to about 30 cm (12 in.) tall, sim-
the Greek aglaios, splendid, and morphe, shape. Aglaia was ple and entire or deeply and irregularly lobed.
one of the Graces of Greek mythology.
 178 C 13

b
Figure 13.5.6. Aglaomorpha
drynarioides: fertile frond, bar
= 10 cm (4 in.).

Figure 13.5.5.
Aglaomorpha
coronans: a. frond,
bar = 10 cm
(4 in.); b. part of
fertile pinna, bar
= 2.5 cm (1 in.).

Aglaomorpha drynarioides (Hooker) M. C. Roos

 13.5.6
syn. Merinthosorus drynarioides (Hooker) Copeland b
Tender
Large to very large, with branched, short-creeping rhi-
zomes. Grows under high light in well-drained, moist-dry
potting mix or uncut moss.
Aglaomorpha drynarioides has soft rhizome scales, and
its fronds grow to nearly 1.75 m (5.5 ft.) long. The fronds
are sessile, pinnatifid, and brown-papery at the base but fo-
liaceous above. The fertile pinnae toward the tip of the
blade are long and linear, with sporangia covering the un-
dersides or in linear patches on wider pinnae. The species is
native to Malaysia, Indonesia, and New Guinea.
Figure 13.5.7.
Aglaomorpha heraclea (Kunze) Copeland
Aglaomorpha
 13.5.7
heraclea: a. frond,
Tender
bar = 10 cm (4 in.);
Very large, with branched, short-creeping rhizomes. Grows b. part of fertile
best under high light in moist-dry potting mix or uncut pinna, bar = 2.5
moss kept well drained. cm (1 in.).
The fronds of Aglaomorpha heraclea are huge, up to 2 m
(6.5 ft.) long. They are also pinnatifid, sessile, and brown-
papery at the base but foliaceous distally. The fertile pinnae
are about the same width as the sterile ones—not con-
tracted, as is common in the genus. The sori are numerous,
small, and scattered. This species is native from Sumatra to a
Malaysia, Java, Borneo, and New Guinea.
In the trade Drynaria morbillosa (C. Presl) J. Smith is
often misapplied to Aglaomorpha heraclea.
 F  F A  C 179

Aglaomorpha meyeniana Schott  13.5.8

Tender
A medium-sized fern with branched, short-creeping rhi-
zomes. Best grown under high light in well-drained, moist- b
dry potting mix or uncut moss.
Aglaomorpha meyeniana is characterized by orange rhi-
zome scales and sessile fronds that are brown-papery at the
base but foliaceous distally. The fertile pinnae are narrow
and constricted between the sori, appearing like a string of
beads. The sori are large and round. This species is native to
Taiwan and the Philippines.

a a

b Figure 13.5.9. Aglaomorpha pilosa: a. fronds, fertile (left)

and sterile (right), bar = 10 cm (4 in.); b. part of fertile
pinna, bar = 2.5 cm (1 in.).

Figure 13.5.8. dle-like hairs. The soral patches resemble those of A. splen-
Aglaomorpha dens, but the latter species differs by having larger fronds,
meyeniana: a. fertile shorter-creeping rhizomes, humus-collecting frond bases,
frond, bar = 10 cm and hairless sporangial capsules. Aglaomorpha pilosa is na-
(4 in.); b. part of tive to the Philippines.
fertile pinna, bar = Some plants circulating in gardens are misidentified as
2.5 cm (1 in.). Aglaomorpha brooksii Copeland.

Aglaomorpha speciosa (Blume) M. C. Roos

 13.5.10
Aglaomorpha pilosa (J. Smith) Copeland syn. Photinopteris speciosa (Blume) C. Presl
 13.5.9 Tender
Tender A large fern with branched, short-creeping rhizomes.
A medium-sized fern with branched, long-creeping rhi- Grows best under medium to high light in well-drained,
zomes. Grows best under high light in a well-drained, moist-dry potting mix or uncut moss.
moist-dry potting mix or uncut moss. Provide ample room Aglaomorpha speciosa bears one-pinnate fronds that lack
to accommodate the creeping rhizomes. humus-collecting bases. The ovate-oblong pinnae often
The rhizome scales of Aglaomorpha pilosa are appressed, have white dots on their upper surfaces and apices that are
ovate-acuminate or triangular, pale-margined, and ciliate long-tapered and often drooping. A conspicuous nectary
along the distal margins. The deeply pinnatifid blades lack gland can be seen at the base of the pinna stalk. The fertile
any modifications for collecting humus. The pinnae are pinnae, which occur in the apical part of the frond, are ex-
acuminate and hairy and are separated by mostly narrow tremely narrow, often drooping, and completely covered
sinuses; the lower pinnae gradually taper to a wing along with sporangia on their undersides. This species is native to
the stipe. The fertile pinnae are narrow and occur only in southeastern Asia, Malaysia, Indonesia, and the Philippines.
the apical portion of the blade. The sori are oblong or occur Aglaomorpha speciosa resembles certain species of Poly-
in four-sided patches. Sporangial capsules bear four nee- podium because of its fully one-pinnate fronds and the ab-
 180 C 13

Figure 13.5.10. Aglaomorpha

speciosa: fertile frond, bar =
10 cm (4 in.).

sence of humus-collecting frond bases. Some authors still Figure 13.5.11.

classify this species in Photinopteris. Aglaomorpha
splendens: a. fertile
Aglaomorpha splendens (J. Smith) Copeland frond, bar = 10 cm
 13.5.11 (4 in.); b. part of
Tender fertile pinna, bar =
Large to very large, with branched, short-creeping rhi- 2.5 cm (1 in.).
a
zomes. Grows best under medium to high light in moist-
dry, well-drained potting mix or uncut moss.
The fronds of Aglaomorpha splendens grow to 2 m (6.5
ft.) long and are sessile. They have brown, papery bases but
are foliaceous above. They are deeply pinnatifid into trian-
gular-lanceolate pinnae. The fertile pinnae occur toward
the frond tips and are contracted about one-third to one-
fifth the width of the sterile pinnae. The sori are quadran-
gular patches borne in rows on each side of the midrib. This b
species is native to the Philippines.
The plants are often confused with Aglaomorpha brook-
sii Copeland, a species from Borneo that differs mainly in
having the quadrangular soral patches variously subdivided
into smaller, sometimes irregular patches. Aglaomorpha
brooksii has not been found in cultivation. Aglaomorpha
splendens also often circulates in gardens as A. pilosa.
Figure 13.5.12.
×Aglaonaria robertsii Hoshizaki  13.5.12 ×Aglaonaria robertsii:
syn. Aglaomorpha ‘Roberts’ a. frond, bar = 10 cm
Semi-tender or hardier (4 in.); b. part of fertile
A medium to large fern with branched, short- to medium- pinna, bar = 2.5 cm
creeping rhizomes. Grows best under high light in well- (1 in.). After Hoshizaki
drained, moist-dry potting mix or uncut moss. The plants (1991).
do well in humid or drier environments and are easy to cul-
tivate.
×Aglaonaria robertsii is a hybrid of Aglaomorpha coro-
nans and Drynaria rigidula. It resembles A. coronans but is a
softer textured and lighter green. The fronds become
 F  F A  C 181

brown and papery at the base but remain green and folia-
ceous above. The blades are pinnatisect with apices pin-
nately lobed to a small terminal segment (or the terminal
segment is absent). The pinnae are connected by a thin,
cartilaginous margin, and each pinna is slightly narrowed
above the broadly adnate base. The fertile pinnae resemble
the sterile ones and are not narrowed as in most Aglaomor-
pha species. The sori vary from round to sublinear, with
one or two sori united or separated and placed between the
lateral veins.
This plant is a hybrid of horticultural origin and does
not grow in the wild. Its parental species belong to the gen-
era Aglaomorpha and Drynaria; thus the generic name of
the hybrid is a compounding of those two names. In ac-
cordance with the International Code of Nomenclature for
Cultivated Plants, the hybrid is also assigned the cultivar
name ‘Santa Rosa’. Santa Rosa Tropicals, California, was
the nursery that first tissue cultured the plant.
‘Starburst’. The pinna margins are sharply and narrowly
incised; less robust than ‘Santa Rosa’. Figure 13.6.2. Alsophila tricolor: frond.

Alsophila  13.6.1–5

syn. Nephelea
The tree ferns of genus Alsophila are related to Cnemidaria
(not cultivated), Cyathea, and Sphaeropteris in the family
Cyatheaceae. Alsophila differs from these other genera by
the black spines or sharp conical bumps on its stipes and the

Figure 13.6.3. Alsophila tricolor: sori, with an indusium

visible on detached sorus, upper right.

Figure 13.6.4. Alsophila tricolor: tip of

stipe scale, ending in a dark stiff bristle,
bar = 0.1 mm.

Figure 13.6.1. Alsophila tricolor: habit.

 182 C 13

along the undersides of the pinnule and segment midrib

are small, whitish, and puffy. The sori are partly covered
with fringed scales and lack a true indusium. The species is
native to Australia and Tasmania.
Almost all the plants listed as Alsophila australis in the
United States trade (usually under the name Cyathea aus-
Figure 13.6.5. tralis) are actually Sphaeropteris cooperi. True A. australis is
Alsophila tricolor: rarely grown in the United States but is often cultivated in
margin of stipe mild areas of England. The glossy, dark brown scales are
scale, marginal cells distinctive of this species.
different from
central cells, bar = Alsophila tricolor (Colenso) R. M. Tryon
0.1 mm.  13.6.1–5, 7;  11
syn. Cyathea dealbata Colenso
Silver tree fern
Semi-hardy
A large tree fern, 3–9 m (10–30 ft.) tall, with blades white
beneath. Grows best under medium light in moist garden
soil or potting mix. The plants tend to grow slowly and
should be placed in areas sheltered from the wind.
minute black bristles on its stipe scales (at least at the apex).
Alsophila tricolor is characterized by glaucous, slightly
The genus name comes from the Greek alsos, grove, and
philein, to love, alluding to certain species that grow in
shady groves. Alsophila contains about 235 species and is
native to the Old and New World tropics. See Cyathea for
a further comparison of the cyatheaceous genera and for
general comments about tree ferns.
Figure 13.6.7. Alsophila tricolor:
Alsophila australis R. Brown  13.6.6 a. pinnule, bar = 1 cm (0.4 in.);
syn. Cyathea australis (R. Brown) Domin b. stipe scales, bar = 1 cm
Rough tree fern, Australian tree fern (0.4 in.).
Semi-hardy to semi-tender
A large tree fern. Best grown under medium light in moist, b
acidic(?) garden soil or potting mix. The plants grow slowly
and do best in cool, moist climates.
a
The stipe bases of Alsophila australis are beset with con-
ical spines to 3 mm (0.1 in.) long and are covered by dark
brown, glossy, slender, brittle scales. The tips of the scales
are slightly contorted and bear a dark bristle. The scales

prickly stipes and shiny, slightly twisted, slender stipe scales

to 7 cm (3 in.) long and tipped with a dark bristle. The glo-
c bose indusia have an opening at the top. This species is na-
b
tive to New Zealand and is, in fact, the national fern of that
country. It is depicted on New Zealand’s coins, bills, pass-
a ports, official seals, national airline planes, and most tour-
ist posters.

Anemia  13.7.1, 2

Flowering fern
Figure 13.6.6. Alsophila australis: a. pinnule, bar = 1 cm These small- to medium-sized terrestrial or epilithic ferns
(0.4 in.); b. stipe scales, bar = 1 cm (0.4 in.); c. bullate are particularly attractive for their erect, long-stalked, and
scale from the underside of the frond, bar = 0.1 mm. branch-like fertile leaflets. The rhizome is erect or creeping
 F  F A  C 183

and is covered by hairs, not scales. The blades can be up to

three-pinnate but are mostly one- to two-pinnate-pinna-
tifid. Fertile fronds typically have the two lowermost pinnae
fully contracted (skeletonized), erect, long stalked, and
bearing sporangia. The sporangia are pear-shaped, with the
annulus encircling the narrow end.
Anemia occurs in the tropics and subtropics of the
world, though most of its 100 species grow in the Americas.
The genus name comes from the Greek aneimon, naked,
alluding to the exposed sporangia that lack an indusium.

Anemia adiantifolia (Linnaeus) Swartz

 13.7.3
Pine fern
Semi-tender, Zone 9
A small to medium fern with an erect, seldom-branched
rhizome. Grows best under medium to high light in basic,
moist-dry garden soil or potting mix.
Anemia adiantifolia has fully three-pinnate fronds. It is
native to tropical America and grows as far north as Florida.

Anemia phyllitidis (Linnaeus) Swartz

 13.7.1, 2, 4
Semi-tender, Zone 9
A small fern with erect, seldom-branched rhizomes. Grows
Figure 13.7.1. Anemia phyllitidis: habit. best under medium to high light in moist-dry garden soil or
potting mix.

Figure 13.7.3. Anemia

Figure 13.7.2. Anemia phyllitidis: stalk-like fertile adiantifolia: frond, bar =
pinnae. 5 cm (2 in.).
 184 C 13

Angiopteris  13.8.1–4

Mule’s-foot fern
Members of Angiopteris are large, coarse, fleshy terrestrial
ferns with massive stems. Their large size limits them to
conservatories and outdoor gardens in the tropics. The
plants need good drainage and must be protected from
wind and frost. Do not let them dry, as wilted fronds re-
cover only with difficulty.

Figure 13.7.4. Anemia phyllitidis:

frond, bar = 5 cm (2 in.). Figure 13.8.1. Angiopteris evecta: habit.

Anemia phyllitidis has one-pinnate fronds with a termi- These ferns bear fronds up to 7 m (23 ft.) long, although
nal pinna that resembles the lateral ones. The veins are net- the fronds tend to be smaller in cultivation. The roots and
ted. This species is native to tropical America. stems are usually thick and fleshy, often massive; stems
sometimes slender and creeping. The middle of the stipe
Anemia rotundifolia Schrader  13.7.5 often has a swelling or bump. The blades vary from one- to
Semi-tender
A medium-sized fern with erect rhizomes and buds at the
tips of the fronds. Grows best under medium light in moist
garden soil or potting mix.
The fronds of Anemia rotundifolia are arching and one-
pinnate with roundish pinnae, and the rachis is greatly ex-
tended, whip-like, and roots at the tip. This species is native
to Brazil.

Figure 13.7.5. Anemia

rotundifolia: frond, bar =
5 cm (2 in.).

Figure 13.8.2. Angiopteris evecta: clusters of sporangia.

 F  F A  C 185

Angiopteris, which contains about 200 species, is native

to the Old World tropics, where it grows in wet habitats. It
has become naturalized in Jamaica and Hawaii. The genus
name comes from the Greek angeion, case or capsule, and
pteris, fern, referring to the sporangia that are larger than
those of many other ferns.
Plants of the genus Angiopteris and the allied Marattia
are so large that few adequate scientific specimens have
been made, making study of the species particularly diffi-
cult. Although new Angiopteris continue to be introduced,
their identities are uncertain, and many are named as vari-
eties of Angiopteris evecta.

Figure 13.8.3. Angiopteris evecta: swollen pinna bases. Angiopteris angustifolia C. Presl  13.8.5
Tender
Very large, with erect, stout, massive stems and whorled
two-pinnate and often have a swelling at the juncture of fronds. Best grown under medium light in well-drained
the rachis and pinnae. The sori consist of a double row of garden soil or potting mix. Angiopteris angustifolia prefers a
free (unfused) sporangia, and an indusium is absent. humid environment; do not permit the soil to dry.
Angiopteris resembles the closely related Marattia, and The pinnules are 12–20 mm (0.5–0.75 in.) wide and
sterile specimens are difficult to assign to either genus. have translucent false veins that run from the margin to
Plants from both genera have fleshy flaps of tissue called nearly the midrib of the pinnule. This species occurs from
stipules on either side of the stipe base. The stipules hug Malaysia to the Philippines.
the swollen base of the fleshy stipe. The genera have similar
cultural requirements.
New plants can be propagated from stipules cut from
the stipe. The stipules should be planted in a moist, loose
medium (gravel, perlite, sphagnum, or a loose, well-drained a
planting mix) at a slant or flat on the medium with either
side up. Some growers notch the thin edges slightly to en-
courage plantlet formation at these points; otherwise,
plants tend to form at the base of the stipule, usually one on
each side. Apply bottom heat at 27°C (80°F) to hasten the
appearance of buds, which may take several months or up
c
to 2 years to appear; large stipules are slower to produce
buds. The buds should be kept moist, but not so wet as to
rot. After the buds produce roots and two or three leaves,
they can be transplanted. Angiopteris can also be propagated
by offshoots (more commonly produced on young plants)
or spores. On all established plants, new roots that emerge
from the stem base should be kept moist with a covering of
mulch. For more information, see Hill (1984).

Figure 13.8.4.
Angiopteris evecta:
stipules around Figure 13.8.5. Angiopteris angustifolia: a. pinna, bar =
bases of leaf stalks, 5 cm (2 in.); b. pinnule, bar = 2 cm (0.8 in.); c. veins
bar = 5 cm (2 in.). with long false vein (left) and sporangia (right), bar =
1 cm (0.4 in.).
 186 C 13

Angiopteris evecta (G. Forster) Hoffmann powder on the lower surface of the blades. The life cycle of
 13.8.1–4, 6 Anogramma is highly unusual because the gametophytes
Mule’s-foot fern are perennial and the sporophytes are annual. In contrast,
Tender to marginally semi-tender the gametophytes of all other ferns are ephemeral, dying
Very large, with erect, stout, massive stems and whorled after the sporophyte is formed. Thus, the leaves of Ano-
fronds. Best grown under medium light in well-drained gramma plants live less than a year and reappear the fol-
garden soil or potting mix. The plants prefer a humid en- lowing growing season.
vironment; do not permit the soil to dry. Anogramma grows in tropical and warmer temperate
Angiopteris evecta is characterized by pinnules that are areas throughout the world and consists of five species. The
about 25 mm (1 in.) wide and translucent false veins that genus name comes from the Greek ano-, upward, and
extend less than halfway from the margin to the midrib. gramme, line, referring to the elongate sori on the distal seg-
The species occurs from Malaysia to Polynesia. One of the ments.
most spectacular plantings of A. evecta can be seen in the
Foster Botanical Garden in Honolulu, Hawaii. Anogramma chaerophylla (Desvaux) Link
 13.9.1–3
Semi-tender
a
A small fern with erect rhizomes. Grows best under me-
dium light in moist potting mix. Volunteer plants appear in
spring, densely and luxuriantly, and usually wither within
a few months. In terrariums it reproduces readily from
spores.
c

Figure 13.8.6. Angiopteris evecta: a. pinna, bar = 5 cm

(2 in.); b. pinnule, bar = 2 cm (0.8 in.); c. veins with
short false vein (left) and sporangia (right), bar = 1 cm
(0.4 in.).

Figure 13.9.1. Anogramma chaerophylla: habit.

Anogramma
Anogramma has special appeal in terrariums because of its
small size and delicate appearance. Anogramma chaerophylla Anogramma chaerophylla can be distinguished from
should not be planted where volunteer ferns are not de- closely related species by its glabrous stipes, broadly trian-
sired, such as in commercial greenhouses, because it readily gular and three- to four-pinnate blades, and white to tan
spreads by spores. spores. In nature the fronds can reach 40 cm (16 in.) long,
The genus consists of terrestrial or epilithic ferns with but only about 15 cm (6 in.) in cultivation. The species is
erect rhizomes and mostly small, clustered fronds. The native to tropical America.
blades vary from one- to four-pinnate and are elongate-tri- Anogramma leptophylla (Linnaeus) Link is cultivated in
angular, membranous, and glabrous (except for one species, England and can be distinguished by its dark brown spores
Anogramma leptophylla, which is hairy). The veins are free, and 1 mm long hairs on the stipes. It is native to the Old
and the sporangia run along the veins. Indusia are absent. World and Mexico.
Anogramma is closely related to Pityrogramma, differing
mainly by its smaller size, thinner texture, and the lack of
 F  F A  C 187

shaped indusium. Many species in the genus resemble

broad-fronded polystichums, but their kidney-shaped in-
dusia, anadromous frond plan, and (in most species) creep-
ing rhizomes distinguish them. Arachniodes might be con-
fused with certain species of Dryopteris, but that genus
usually differs by having a catadromic frond plan and de-
cumbent or erect rhizomes.
Arachniodes, which contains about 50 to 70 species and
hybrids, grows in the tropics and subtropics throughout
Figure 13.9.2. the world but is most diverse in Asia. The genus name
Anogramma comes from the Greek arachnion, spider’s web, and -odes,
chaerophylla: meaning “having the form or nature of.” No one knows
sporangia along why the fern received this name.
veins.
Arachniodes aristata (G. Forster) Tindale
 13.10.1, 2
syn. Polystichum aristatum (G. Forster) C. Presl
East Indian holly fern
Hardy, Zone 6
A medium-sized fern with long-creeping rhizomes. Best
grown under medium light in garden soil or potting mix.
This species does well in moist to moist-dry soil. The plants
are evergreen, easy to cultivate, and best displayed in wide
pots or beds. They grow slowly from spores.
Arachniodes aristata is recognizable by its dark green,
broad, five-sided sterile blades, which are up to four-pin-
nate. The fertile fronds are taller than the sterile ones and
have slightly more contracted segments. This species is
native to Africa, Australia, New Zealand, and the Pacific
Islands.

Figure 13.9.3. Anogramma chaerophylla: habit, bar =

5 cm (2 in.).

Arachniodes  13.10.1

The arachniodes are medium-sized terrestrial ferns useful in
pots or planting beds. One species in particular, the East
Indian holly fern (Arachniodes aristata), is valued for its
attractive shiny, dark green, divided foliage.
The rhizomes vary from short- to long-creeping; rarely
are they erect. The blades are two-pinnate or more divided,
broad, often triangular or pentagonal, anadromous, and
dimorphic or not. The round sori are covered by a kidney- Figure 13.10.1. Arachniodes aristata: habit.
 188 C 13

fertile fronds are taller and more contracted than the sterile.
This species is native to Japan and China and has become
naturalized in the southeastern United States.
b A variegated form sometimes circulates under the mis-
applied name of Arachniodes aristata ‘Variegatum’.

Arachniodes standishii (T. Moore) Ohwi

 13.10.4
c
syn. Polystichum standishii (T. Moore) C. Christensen
Hardy, Zone 6
A medium-sized fern with slightly erect to short-creeping
rhizomes. Grows best in humid environments under me-
dium light in moist garden soil or potting mix. The plants
Figure 13.10.2. are evergreen and slow-growing.
Arachniodes aristata: Arachniodes standishii is the largest and most finely dis-
a. habit, bar = 5 cm sected (up to four-pinnate) cultivated species of Arach-
(2 in.); b. basal pinna, niodes. The blades are elongate-triangular, monomorphic,
bar = 5 cm (2 in.); dull, medium green, and softer textured than those of other
a c. segment, bar = species in cultivation. The sori, which are covered by broad
5 mm (0.2 in.). reniform indusia, obscure most of the lower surface of the
segments. The species is native to Japan, Korea, southern
China, and Vietnam.
Arachniodes simplicior (Makino) Ohwi
 13.10.3 Argyrochosma
Hardy, Zone (6)7 syn. Notholaena in part, Cheilanthes in part
A medium-sized fern with long-creeping rhizomes. Best Argyrochosma is a genus of small, wiry ferns that grow in
grown under medium light in moist-dry garden soil or pot- dry rocky places. In the past the species have been classi-
ting mix. Some forms are especially attractive for their var- fied with either the cloak ferns (Notholaena), lip ferns
iegated foliage, bearing yellow-green lighter-colored tissue. (Cheilanthes), or rock brakes (Pellaea). See Cheilanthes for a
Arachniodes simplicior has evergreen fronds up to three- comparison of Argyrochosma to these and other genera.
pinnate. The fronds are usually too few and far apart on Windham (1987) showed that Argyrochosma is most closely
the creeping rhizome to make a full, leafy pot plant. The

c
a

b b

Figure 13.10.4. Arachniodes standishii: a. habit, bar =

Figure 13.10.3. Arachniodes simplicior: a. frond, bar = 5 cm (2 in.); b. basal pinna, bar = 5 cm (2 in.);
5 cm (2 in.); b. basal pinna, bar = 5 cm (2 in.). c. pinnule, bar = 1 cm (0.4 in.).
 F  F A  C 189

related to Pellaea but can usually be distinguished by the Argyrochosma dealbata is distinguished from A. jonesii
presence of a waxy powder (farina) on the lower surfaces of and A. microphylla by the white powder on the lower sur-
the blades (this is absent in Argyrochosma jonesii and A. mi- face of the blades. It is further characterized by dark reddish
crophylla) and other technical characteristics. brown, shiny stipes and triangular blades two- to three-pin-
The blade surfaces of Argyrochosma lack hairs or scales. nate and blue-green on the upper surface. The dark color of
The segments are untoothed and distinctly stalked. The the stalk enters the segment bases and merges with the
sporangia run along the veins, usually covering the outer green of the blades. The ultimate segments are round to
one-third to two-thirds of the segments. The fertile margins ovate-oblong, 3 mm (0.1 in.) or less long, and their margins
are flat or slightly enrolled and have the same color and tex- enroll, covering the dark sporangia that are borne on the
ture as the rest of the blade—that is, the fertile margins are distal third of the secondary veins. The species is native to
not texturally modified like those of some other cheilan- the south-central United States, where it grows on calcare-
thoid ferns. ous cliffs and ledges.
About 20 species occur in the genus, which is entirely
native to the New World. Its name comes from the Greek Argyrochosma jonesii (Maxon) Windham
argyros, silver, and chosma, powder, referring to the whitish  13.11.2
powder found on the lower blade surfaces of most species. syn. Cheilanthes jonesii (Maxon) Munz, Notholaena
jonesii Maxon
Argyrochosma dealbata (Pursh) Windham Jones’s cloak fern
 13.11.1 Semi-hardy, Zone 8(9)
syn. Cheilanthes dealbata Pursh, Notholaena dealbata A small fern with compact, erect to ascending, usually un-
(Pursh) Prantl branched rhizomes and clustered fronds. Best grown under
Powdery cloak fern high light in drained, basic(?) garden soil kept moist-dry to
Hardy, Zone 5 dry.
A small fern with compact, erect to ascending, usually un-
branched rhizomes and clustered fronds. Apparently prefers
medium light with basic(?), moist-dry to dry, drained gar-
den soil with sand.

b b

Figure 13.11.1.
Argyrochosma dealbata:
a. frond, bar = 2.5 cm
a (1 in.), after Tryon Figure 13.11.2. Argyrochosma jonesii:
(1956); b. segments a. frond, bar = 2.5 cm (1 in.);
with sori and indusia, b. fertile segment, bar = 1 mm.
bar = 1 mm. After Tryon (1956).
 190 C 13

Argyrochosma jonesii has dark brown stipes and shiny, four-pinnate, glaucous (but not white-powdery) blades;
ovate-lanceolate, leathery blades that are two- to three-pin- and rachises grooved on the upper surface. The segment
nate. The dark color of the segment stalks extends into the margins are enrolled, and the sporangia are borne on the
base of the segments and merges with the green of the outer third of the veins. The species is native to the south-
blade; the stalks are not jointed to the segments, nor does western United States and northern Mexico, where it in-
the stalk color stop abruptly as in A. microphylla. The lower habits rocky, limestone hillsides and cliffs.
surface of the blade lacks white powder and has flat or
slightly enrolled margins, with sporangia borne on outer Arthropteris  13.12.1, 2
third to half of the veins. The species is native to the south- Joint fern
western United States and Mexico. Some species of Arthropteris look like a small Thelypteris
with a long-creeping rhizome. Nevertheless, one species,
Argyrochosma microphylla (Mettenius ex Kuhn) Arthropteris tenella, is distinct for its small- to medium-
Windham  13.11.3 sized, one-pinnate fronds. This small fern is handsomely
syn. Cheilanthes parvifolia (R. M. Tryon) Mickel, displayed in baskets or by training it to grow up the trunks
Notholaena parvifolia R. M. Tryon of trees. The latter growth habit can be started by planting
Small-leaved cloak fern the fern in the soil around the tree and training the rhi-
Semi-hardy, Zone 7 zomes up the trunk, holding them in place with string or
A small fern with compact, erect to ascending, usually less visible plastic fishing line. A collar of uncut sphagnum
unbranched rhizomes and clustered fronds. Grows under moss placed around the base of the tree encourages rooting
high light in basic(?), moist-dry, well-drained garden soil and more vigorous growth.
with sand. These terrestrial or epiphytic ferns have long-creeping,
Unlike the previous two species, Argyrochosma micro- slender, scaly rhizomes, and the stipes are jointed. The joint
phylla has its ultimate segments jointed to their stalks. This is slightly raised and sometimes distant from the stipe base,
can be seen by the dark color of the stalk stopping abruptly often appearing as a faint dark line. Blades vary from one-
at the segment base, not passing into the segment and pinnate to one-pinnate-pinnatifid and have sessile pinnae
merging with the green of the blade. The species is further jointed to the rachis. The veins are either simple or forked,
characterized by brown stipes; triangular to ovate, three- to never netted. The round sori are covered by rounded, kid-
ney-shaped indusia (this is absent in some species). The
genus resembles certain species of Thelypteris but can be
distinguished by the tiny, jointed, usually reddish (when
dry) hairs on the upper surfaces of the rachises and costae;
Arthropteris does not bear needle-shaped hairs, as is often
the case in Thelypteris. Another distinction is that
b Arthropteris stipes have three or more vascular bundles,
whereas Thelypteris stipes have only two.

a Figure 13.11.3.
Argyrochosma microphylla:
a. frond, bar = 2.5 cm
(1 in.); b. segment with
sori and indusia, bar =
1 mm. After Tryon
(1956).

Figure 13.12.1. Arthropteris tenella: habit.

 F  F A  C 191

Figure 13.12.3. Arthropteris monocarpa: a. habit, bar =

Figure 13.12.2. Arthropteris tenella: sori. 5 cm (2 in.); b. joint near middle of stipe, bar = 1 cm
(0.4 in.).

This genus is native to the Old World tropics, Japan, A similar species, Arthropteris orientalis (J. F. Gmelin)
Polynesia, and temperate areas of Australia and New Zea- Posthumus, is not yet known in cultivation but has been
land. One species occurs in South America (Juan Fernán- confused with A. monocarpa. It differs by having stipes
dez Islands); the remaining 12 to 15 species are found in the jointed in the upper (distal) half, one to eight sori per lobe,
Old World. The genus name comes from the Greek and the presence of whitish (not dark) hydathodes on the
arthron, joint, and pteris, fern, alluding to the jointed stipes upper surface of the blade. The joint on the stipe appears as
of some species. a faint dark line and is easily overlooked. This species is also
native to tropical Africa. Introduced plants came from
Arthropteris monocarpa (Cordemoy) C. Christensen Zimbabwe.
 13.12.3
Semi-tender to semi-hardy Arthropteris tenella (G. Forster) J. Smith ex Hooker f.
A small to medium fern with medium- to long-creeping  13.12.1, 2, 4
rhizomes. Best grown under medium light in moist-dry syn. Polypodium tenellum G. Forster
garden soil or potting mix. The plants can be used as a Joint fern
ground cover or confined in a wide container. They can Semi-hardy
also be used over logs or tree trunks if the substrate is suffi- A small to medium fern with medium- to long-creeping
ciently moist. The fronds are deciduous. rhizomes. Best grown under low to medium light in moist
Arthropteris monocarpa is characterized by stipes jointed to moist-dry garden soil or potting mix or uncut moss. The
in the lower (basal) half, one-pinnate-pinnatifid blades, usu- plants are versatile: they can form handsome displays if
ally one sorus per lobe, and usually dark hydathodes on the trained to grow up tree trunks, or they can be grown in bas-
vein tips of the upper surface of the blade. The indusium is kets or as a ground cover. Any divisions should be made
kidney-shaped. This species is native to tropical Africa. during periods of active growth.
 192 C 13

teeth. They occur one to three per segment. The species is

native to the United States (California) and Mexico (north-
a ern Baja California).

Aspidotis carlotta-halliae (W. H. Wagner & E. F.

Gilbert) Lellinger  13.13.2
syn. Cheilanthes carlotta-halliae W. H. Wagner & E. F.
Gilbert
Semi-hardy, Zone 7
A small fern with compact, short-creeping to ascending rhi-
zomes and clustered fronds. Grows under high light in
well-drained, moist-dry to dry garden soil.
Aspidotis carlotta-halliae is a hybrid of A. californica and
A. densa. It is best distinguished from its parents by the false
indusia with 6–10 coarse irregular teeth or lobes. It is native
b to northern California, where it typically grows on serpen-
tine rock.

Figure 13.12.4. Arthropteris tenella: a. habit, bar = 5 cm

(2 in.); b. joint at stipe base, bar = 1 cm (0.4 in.).

Arthropteris tenella is characterized by one-pinnate

blades with a terminal pinna resembling the lateral ones.
The sori are non-indusiate, round, and close to the mar-
gin. It is native to New Zealand and Australia.

Aspidotis
Lace fern
These small ferns typically grow in dry, rocky areas. Aspi-
dotis species are sometimes included in Cheilanthes, though
they can be distinguished by their elongate, distally toothed
segments with shiny, striated upper surfaces and broad, pa-
pery indusia. See Cheilanthes for a comparison with related
genera. Aspidotis consists of four species, three of which
occur primarily in western North America, and the re-
maining one in Mexico. The generic name comes from the
Greek word aspidotes, shield-bearer, in allusion to the
shield-like false indusia.

Aspidotis californica (Hooker) Nuttall ex Copeland

 13.13.1
syn. Cheilanthes californica (Hooker) Mettenius
California lace fern
Semi-hardy, Zone 8 Figure 13.13.1. Aspidotis
A small fern with compact, short-creeping to ascending rhi- californica: frond, bar =
zomes and clustered fronds. Grows under high light in 2.5 cm (1 in.).
well-drained, moist-dry to dry garden soil.
The blades of Aspidotis californica are pentagonal to tri-
angular, four- to five-pinnate, with segments mostly lance-
olate, 1.5–4 mm long, and pointed apices and margins
coarse and sparsely toothed. The indusia are papery, light- Figure 13.13.2. Aspidotis carlotta-
colored, semi-circular, and entire or have two to six coarse halliae: frond, bar = 2.5 cm (1 in.).
 F  F A  C 193

Aspidotis densa (Brackenridge in Wilkes) Lellinger

 13.13.3
syn. Cheilanthes siliquosa Maxon, Pellaea densa
(Brackenridge) Hooker
Indian’s dream, cliff brake
Very hardy, Zone 4
A small fern with a more-or-less compact, short-creeping to
ascending, branching rhizome and clustered fronds. Grows
under high to medium light in well-drained, moist-dry to
dry garden soil with sand. Avoid getting water on the
fronds.

Figure 13.14.1. Asplenium daucifolium: habit.

bird’s-nest ferns, with fronds more than one meter long.

a The small species are favorites for rock gardens or terrari-
ums. The larger subtropical and tropical species are well
b adapted to growing indoors in pots. Several species of
bird’s-nest ferns are in cultivation, mainly Asplenium an-
tiquum, A. australasicum, and A. nidus. They are popular
Figure 13.13.3. Aspidotis densa: a. frond, bar = 2.5 cm indoor plants and are easy to grow if not overwatered. They
(1 in.); b. fertile segment, bar = 1 mm. tend to get overwatered when planted in large pots, but for-
tunately they can thrive in relatively small pots. All species
of Asplenium must be protected from slugs and snails, es-
pecially emerging fronds and young plants.
An evergreen fern, Aspidotis densa is distinguished by A fern can usually be assigned to this genus if it has elon-
small triangular fronds with long segments. The blades vary gate sori (which vary from oblong to linear, depending on
from pentagonal to triangular, three- to four-pinnate, and the species) and clathrate rhizome scales. Another helpful,
have linear (not bead-like), entire segments. A glabrous but less easily seen, characteristic is the two vascular bundles
midrib is conspicuous on the lower surface. The fertile in the stipe. These bundles unite in the distal part of the
fronds often have longer stipes and narrower segments than stipe to form an X-shape. The sori have a thin, flap-like in-
the sterile ones. The indusium is linear, with 10–15 shal- dusium, although this trait is sometimes difficult to see be-
low, regular or irregular teeth. The species is native to the cause the indusium gets pushed aside and covered by the
northwestern United States and southwestern and eastern sporangia as they mature. Unfortunately, frond form can-
Canada. not be used to distinguish this genus because almost every
frond shape and degree of cutting, from simple to several
Asplenium  13.14.1–3 times pinnate, occur in Asplenium. The veins are free in
Spleenwort nearly all the species.
Asplenium is a large, diverse genus of terrestrial or epiphytic Most botanists now classify Camptosorus, Ceterach, Phyl-
ferns. They range in size from the tiny European spleen- litis, and Schaffneria in Asplenium, since these genera hy-
worts, with fronds only a few centimeters long, to the giant bridize with Asplenium and are distinguished from it by
194 C 13

only a few minor characteristics. In this book, however,

these genera are kept separate, partly for convenience and
partly because current studies indicate newer realignments
of the genera.
Asplenium occurs worldwide and is one of the largest
fern genera, containing about 650 species. The genus name
comes from the Greek splen, spleen. The Greek physician
and botanist Dioscorides thought that plants of this genus
cured diseases of the spleen.

Asplenium adiantum-nigrum Linnaeus

 13.14.4
Black spleenwort
Semi-hardy, Zone 8
A small fern with suberect rhizomes. Grows best under me-
dium light in moist, basic garden soil with sand or potting
mix.
Asplenium adiantum-nigrum is characterized by dark or
black stipes and (typically) triangular, two-pinnate-pinna-
tifid blades. The basal pinnae are the largest and most di-
vided. The species is cosmopolitan, growing mostly in tem-
perate areas.

Asplenium aethiopicum (N. L. Burman) Becherer

 13.14.5
Semi-hardy, perhaps more tender (?)
A medium-sized fern with short-creeping to suberect rhi-
zomes. Grows under low to medium light in moist garden
Figure 13.14.2. Asplenium daucifolium: frond. soil or potting mix.

Figure 13.14.4. Asplenium

adiantum-nigrum: frond, bar =
5 cm (2 in.).

Figure 13.14.3. Asplenium daucifolium: sori and indusia.

 F  F A  C 195

Figure 13.14.5. Asplenium

aethiopicum: frond, bar =
5 cm (2 in.). b

a
c

Figure 13.14.6. Asplenium antiquum: a. fronds, bar =

5 cm (2 in.); b. sori on upper vein fork, bar = 1 cm
Asplenium aethiopicum can be distinguished by its dark (0.4 in.); c. midrib cross section, bar = 5 mm (0.2 in.).
stipes and rachises that bear numerous blackish, thread-like
scales. The pinnules are distinctive by being wedge-shaped
and deeply and irregularly incised, the incisions irregularly
toothed. The blade is narrowly lanceolate to oblong and upper fork of forked veins. The indusium is more than 0.5
up to two-pinnate-pinnatifid. The species is native to tropi- mm wide. This species is native to Taiwan and Japan.
cal Africa, Madagascar, and Australia. Asplenium antiquum is sold widely. It was first intro-
This species is closely related to Asplenium praemorsum duced into the United States in 1969 from the spores of
Swartz, an American tropical species, and further research plants collected near Hakone, Japan. This species is a ro-
may show that they are the same. As presently recognized, bust grower with a neater outline than Asplenium nidus. As-
A. aethiopicum is highly variable and includes five ploidy plenium nidus and A. australasicum differ from A. antiquum
levels. by having the veins closer, the sori on both forks of a vein,
and the indusia less than 0.5 mm wide. Asplenium aus-
Asplenium antiquum Makino  13.14.6 tralasicum also has the lower side of the midrib keeled, not
Japanese bird’s-nest fern rounded. Asplenium serratum, a similar species from the
Semi-tender, perhaps hardier (?) American tropics, differs by having free, not connected,
A medium-sized fern with erect, stout, unbranched rhi- veins at the tips.
zomes. Best grown under medium-high light in well- ‘Victoria’. Plate 12. Margins ruffled and fringed. A sim-
drained, moist to moist-dry potting mix or uncut moss. ilar cultivar with narrower fronds is also in the trade.
The plants do well in pots but are sensitive to overwatering.
Asplenium antiquum resembles the common bird’s-nest Asplenium auritum Swartz  13.14.7
fern (A. nidus) but has more compact growth and narrower Eared spleenwort
fronds with more obvious short stipes. The midrib on the Tender
underside of the leaf is rounded, not keeled, and the veins A small fern with erect rhizomes. Best grown under me-
in the middle of the blade are more than 1 mm apart. The dium light in moist potting mix.
vein tips are united into a connecting strand that runs par- Asplenium auritum has thick, lanceolate fronds, up to
allel to the blade margin. The sori usually occur on the one-pinnate-pinnatifid at the base, with scattered clathrate
 196 C 13

Figure 13.14.8. Asplenium

australasicum: a. frond, bar =
5 cm (2 in.); b. veins at base of
frond, bar = 1 cm (0.4 in.);
c. midrib cross section, bar =
5 mm (0.2 in.).

Figure 13.14.7.
Asplenium auritum:
frond, bar = 5 cm (2 in.).

c
scales on the lower surfaces. The indusium is inconspicu-
ous. The species is widespread in tropical America, where it a
typically grows as an epiphyte on tree trunks.

Asplenium australasicum (J. Smith) Hooker Asplenium bulbiferum G. Forster  13.14.9
 13.14.8 Mother fern
syn. Asplenium australis Horticulture Semi-hardy to semi-tender
Semi-tender A medium-sized fern with short-creeping to suberect rhi-
A medium to large fern with erect, stout, unbranched rhi- zomes and pale green, arching foliage. Grows under low to
zomes. Grows well under medium light in well-drained, medium light in moist potting mix. This species is recom-
moist to moist-dry potting mix or uncut moss. The plants mended for places with low light. It is easy to grow and can
do particularly well in pots, but be careful not to overpot. be easily propagated from the buds on the fronds; however,
Collectively, the fronds form a funnel or bowl. This species it has the disadvantage of being particularly susceptible to
tends to be more cold hardy and less vulnerable to slugs attack by slugs and snails. Narrowly segmented forms prop-
than Asplenium nidus, which it resembles. agated from spores do not breed true.
The fronds of Asplenium australasicum vary greatly in Asplenium bulbiferum is characterized by highly divided
outline but are generally oblanceolate. They differ from blades (mostly two-pinnate-pinnatifid but sometimes up
those of A. nidus by having keeled undersides of the mid- to nearly four-pinnate) that may bear numerous buds on
ribs, and the sori are usually spread along more than half the the upper surfaces. The blades are oblong-ovate and thick.
length of the vein. The species is native to Australia and the Each incurved lobe of the pinnules contain a single sorus.
Pacific Islands. The species is native to India, Malaysia, Australia, New
Whether fronds are arranged in a narrow funnel or wide Zealand, and the Pacific Islands.
bowl is not consistent in Australian plants or those in cul-
tivation. The most reliable feature for separating Asplenium Asplenium ×crucibuli Horticulture  13.14.10
australasicum from A. antiquum and A. nidus is the keeled syn. ×Asplenosorus crucibuli Horticulture
rather than rounded midribs. Although infrequently Hardy to semi-hardy, Zones 5
grown, A. serratum also has simple leaves and a keeled mid- A small fern with erect rhizomes. Grows best under low to
rib, but it can be distinguished because its veins are free at medium light in moist garden soil or potting mix. These
the tips, not connected. evergreen plants are excellent for rock gardens.
 F  F A  C 197

Asplenium ×crucibuli is a sterile hybrid characterized by

linear fronds that are mostly pinnatifid but with a few pin-
nae at the base, and by elongate, tail-like blade tips. It was
produced in horticulture from a cross between A. platyneu-
ron and Camptosorus sibericus.

b Asplenium cuspidatum Lamarck  13.14.11

Tender
A small fern with erect rhizomes and proliferous roots.
Grows best under low-medium light in moist uncut moss.
This species usually forms tufts of very small, delicate fronds
in terrariums or bottle gardens. Like the filmy ferns, it pre-
fers humidity and grows slowly during the winter.
Asplenium cuspidatum has triangular fronds up to three-
pinnate, with ultimate segments narrowly obovate, some-
times forked at the tip. It is widespread in tropical America.

Figure 13.14.9. Asplenium bulbiferum: a. frond, bar =

5 cm (2 in.); b. leaf bud, bar = 1 cm (0.4 in.).

Figure 13.14.11. Asplenium cuspidatum: habits,

bar = 5 cm (2 in.).

Asplenium daucifolium Lamarck

 13.14.1–3, 12
syn. Asplenium viviparum C. Presl
Figure 13.14.10.
Mauritius spleenwort, dwarf mother fern
Asplenium ×crucibuli:
Semi-tender
fronds, bar = 5 cm
(2 in.). A small to medium fern with short-creeping to suberect
rhizomes and bud-bearing fronds. Grows under low to me-
dium light in moist potting mix. Though somewhat slow
growing, this species is recommended for sites with low
light. It is easy to start new plants from buds on the frond.
Asplenium daucifolium resembles A. bulbiferum but has
smaller, stiffer, darker green fronds, and the underside of
the rachis is beset with linear or thread-like scales. Similar
scales also occur in A. bulbiferum, but they are ovate and
cupped. The species is endemic to Madagascar and the
Macarene Islands.
 198 C 13

Figure 13.14.13.
Asplenium
×ebenoides:
fronds, bar =
5 cm (2 in.).

Figure 13.14.12. Asplenium

daucifolium: frond, bar =
5 cm (2 in.).
Asplenium flabellifolium Cavanilles  13.14.14
Necklace fern
Semi-tender
Asplenium ×ebenoides R. R. Scott  13.14.13 A small fern with short-creeping to suberect rhizomes and
syn. ×Asplenosorus ebenoides (R. R. Scott) Wherry fronds bearing buds at their tips. Grows best in humid en-
Scott’s spleenwort, dragon-tail fern vironments in well-drained, moist potting mix. The plants
Hardy, Zone (4)5 prefer cool, moist, shaded areas.
A small fern with erect rhizomes. Best grown under me- The fronds of Asplenium flabellifolium are one-pinnate
dium light in moist potting mix. Growth may be improved with round to fan-shaped pinnae, the larger pinnae three-
with ample humidity, as in terrariums.
Asplenium ×ebenoides is of hybrid origin, resulting from
a cross between A. platyneuron and Camptosorus rhizophyl-
lus, and the plants are variably intermediate between those
species in frond shape and cutting. The intermediacy is best
seen in the cutting of the blade, which is one-pinnate at the
base, as in A. platyneuron, and entire or nearly so distally or
long tapered, as in C. rhizophyllus. The central portion of
the blade is lobed, often irregularly, expressing the inter-
mediate condition. The venation is also intermediate: A.
platyneuron has free veins, C. rhizophyllus has netted veins,
and the hybrid has both. Figure 13.14.14. Asplenium
This hybrid occurs naturally in the eastern United States flabellifolium: frond, bar =
where the two parents mingle. It produces aborted spores 5 cm (2 in.).
and is therefore sterile; however, at one site in Georgia,
plants have doubled their chromosome number to become
fertile, producing viable spores. Some commercial green-
houses have propagated these fertile plants from spores, and
the fern is becoming more widely available in the trade.
 F  F A  C 199

lobed or more. The rachis is long and trailing, often rooting

at the tip. The species is native to Australia and New
Zealand.

Asplenium fontanum (Linnaeus) Bernhardi b

 13.14.15
Smooth rock spleenwort
Hardy, Zone 5
Very small, to about 15 cm (6 in.) long, with clumped rhi-
zomes. Grows under medium light in well-drained, moist,
basic garden soil or sand and peat mix. The plants should
be kept moist at all times.
Asplenium fontanum is characterized by lanceolate, two-
pinnate-pinnatifid blades. It is native to Europe and the
Himalayas.

Figure 13.14.15. Asplenium fontanum:

a. frond, bar = 5 cm (2 in.); b. pinna,
bar = 5 mm (0.2 in.).
a
Figure 13.14.16. Asplenium a
longissimum: a. frond, bar = 5 cm
(2 in.); b. pinna variations, bar =
2 cm (0.8 in.).
Asplenium longissimum Blume  13.14.16
syn. Blechnum indicum N. L. Burman
Tender
Asplenium marinum Linnaeus  13.14.17
A medium to large fern with rhizomes erect or nearly so.
Sea spleenwort
Best grown under medium light in moist potting mix or
Semi-hardy
uncut moss. The plants grow robustly and are best dis-
played in hanging baskets, but they must be watered fre- A small fern with clumped rhizomes. Best grown under me-
quently. The fronds can reach 1 m (3 ft.) long or more and dium light in moist potting mix. The plants are slow-grow-
often bear buds at the apex. ing and need humidity. In nature they grow along coasts on
Asplenium longissimum is characterized by long, one- rocks subject to sea spray. This species is sensitive to frost.
pinnate fronds narrowed at the base and by numerous nar- Asplenium marinum is distinguished by lanceolate, one-
rowly triangular, sessile pinnae with coarse, doubly serrate pinnate fronds with fleshy, glossy blades. The pinnae vary
margins. The pinna bases are mostly truncate or wedge- from ovate-lanceolate to oblong and have crenate-serrate
shaped on the lower pinnae, whereas the middle pinnae are margins and blunt apices. This species is native to Europe
unequal-sided, with the upper side slightly enlarged and and northern Africa.
eared and the lower side more oblique. The species occurs
from Vietnam to Indonesia, Borneo, and the Philippines. Asplenium nidus Linnaeus  13.14.18
In the United States this species is often confused with Bird’s-nest fern
Asplenium pellucidum; however, it differs from that species, Tender
and also from A. polydon, by the buds at the frond tips. A medium to large fern with erect, stout, unbranched rhi-
zomes. Best grown under medium to low light in moist-
dry potting mix or uncut moss. The plants grow relatively
slowly. Do not overpot, and keep well drained.
200 C 13

The fronds of Asplenium nidus are up to 1.5 m (5 ft.)

long, and most form a vase-shaped cluster. The stipe is
short and inconspicuous, and the blades are simple, entire,
and variable in shape but mostly oblanceolate. The midrib
on the underside of the blade is rounded, not keeled as in
related species. The vein tips unite in short arcs before the
cartilaginous margin. The sori usually occupy less than half
Figure 13.14.17. Asplenium the vein length and are borne on both branches of the vein.
marinum: frond, bar = 5 cm The species is native to the tropics of the Old World and to
(2 in.). Hawaii.
This variable species has several named varieties, which
further research may prove to be valid species. Plants called
Asplenium nidus in cultivation that have midribs keeled on
the underside are mostly A. australasicum. Plants with mid-
ribs rounded on the underside and sori often only on the
upper vein fork are A. antiquum.
A number of garden variants of Asplenium nidus exist,
including plants with unusual characteristics such as lobed
or fimbriate margins and narrow fronds. The most strik-
ing and well known is the lasagna fern.
var. plicatum van Alderwerelt van Rosenburgh (‘Crispa-
folium’, ‘Crispum’, and possibly ‘Undulatum’ of the trade).
Lasagna fern. A smaller plant with pleated or wavy blades.
The plants have thick, rigid fronds that twist slightly, re-
sembling lasagna noodles. They grow slowly and need pro-
tection during winter in subtropical climates.

Asplenium oblongifolium Colenso  13.14.19

syn. Asplenium lucidum G. Forster
Glossy spleenwort
Semi-hardy
A medium-sized fern with short-creeping to suberect rhi-
b
zomes. Grows best under medium light in moist garden
soil or potting mix. The plants have attractive deep green,
glossy fronds and are easy to cultivate.
Asplenium oblongifolium has one-pinnate blades with an
apical pinna that resembles the lateral ones. The pinnae are
short-stalked, entire, and tapered to the tip. The sori form
Figure 13.14.18. Asplenium nidus:
a herringbone pattern and occupy most of the area between
a. frond, bar = 5 cm (2 in.); b. sori on
the midrib and margins. This species is endemic to New
both vein forks, bar = 1 cm (0.4 in.);
Zealand.
c. midrib cross section, bar = 5 mm
(0.2 in.). Asplenium oligophlebium Baker  13.14.20
Probably semi-tender
A small fern with erect rhizomes and fronds bearing buds at
the apex. Seems to grow best under low light in moist pot-
ting mix. The plants are delicate, prefer humidity, and grow
extremely well in terrariums. They root readily from the
frond tips.
Asplenium oligophlebium is characterized by narrowly
lanceolate, one-pinnate fronds with many linear-lanceolate
to semi-triangular pinnae. The margins are irregularly
lobed, especially the acroscopic margins, and the basal lobes
c of the pinnae are enlarged, often forked. The species is en-
a
demic to Japan.
 F  F A  C 201

Asplenium pellucidum Lamarck  13.14.21

Tender
A medium to large fern with suberect rhizomes and pen-
dent fronds. Grows under medium light in moist potting
mix or uncut moss.
The stipes of Asplenium pellucidum are short, and the
pinnae are shorter toward the base of the blades. The pinna
margins are serrate, and the middle pinnae are very unequal
at the base. The rachis is beset by numerous dark scales that
are rounded at the base and abruptly tapered to a bristle. As-
plenium pellucidum is native to the Old World tropics ex-
cept Africa. The plants cultivated in the United States are
often misidentified as A. longissimum.

Figure 13.14.19. Asplenium

oblongifolium: frond, bar =
5 cm (2 in.).

Figure 13.14.20. Asplenium

oligophlebium: frond, bar =
5 cm (2 in.).

Figure 13.14.21. Asplenium pellucidum: a. frond, bar =

5 cm (2 in.); b. pinnae, bar = 2 cm (0.8 in.).
 202 C 13

Asplenium pinnatifidum Nuttall  13.14.22

syn. ×Asplenosorus pinnatifidus (Nuttall) Mickel
Lobed spleenwort
Hardy
A small fern with erect rhizomes and clustered fronds. Best
grown under medium light in moist gardening soil or pot-
ting mix. An excellent rock garden or, in arid climates, a
good terrarium fern.
Asplenium pinnatifidum has narrowly lanceolate blades
that are pinnatifid for most of their length except for one or Figure 13.14.23. Asplenium
two pinnae at the base. The blade apex is often long-atten- platyneuron: frond, bar =
uate. The species is native to eastern North America, where 5 cm (2 in.).
it usually grows in shaded sandstone crevices.
Asplenium pinnatifidum is a hybrid of A. montanum and
Camptosorus rhizophyllus, and the lobing of the blade is vari-
ably intermediate between these two species. Although of
hybrid origin, the plants are polyploid, having doubled
their chromosome number, which means that they have
normal, viable spores.

Asplenium polyodon G. Forster  13.14.24

syn. Asplenium falcatum Lamarck
Sickle spleenwort
Semi-tender or hardier
Short-creeping, stout rhizomes and often pendent fronds to
130 cm (52 in.) long. Grows under low to medium light in
Figure 13.14.22. Asplenium moist potting mix or uncut moss. The species is extremely
pinnatifidum: habit, bar = 5 cm (2 in.). variable in frond length. The long-fronded forms are best
grown in baskets.
The stipe of Asplenium polyodon is dark brown, scaly,
Asplenium platyneuron (Linnaeus) Britton, Sterns & and about one-third to one-half the blade length. The
Poppenburg  13.14.23 larger stipe and rachis scales are long-triangular and smoky
Ebony spleenwort colored. The blades are narrowly oblong, one-pinnate, and
Hardy, Zones 4 and 5 widest at the base. The pinnae have unequal bases (exca-
A small to medium fern with suberect rhizomes. Grows vate on the lower side), coarsely, doubly serrate margins,
under low to high light in basic garden soil or potting mix and gradually tapered tips. This species might be confused
kept moist-dry. The plants are easy to grow once established. with A. longissimum or A. pellucidum. It is native from
Asplenium platyneuron resembles A. trichomanes but dif- Madagascar to the Pacific Islands.
fers by having larger and more erect fronds that are oblan-
ceolate, and oblong pinnae eared near the base on the upper Asplenium prolongatum Hooker  13.14.25
margin. The “ear” on the pinna often overlaps the rachis, Semi-hardy
which is ebony-colored. The fertile fronds are often tall and A small fern with erect rhizomes and fronds bearing buds at
erect, whereas the sterile ones are smaller and spreading, the tips. Grows under low to medium light in moist soil or
sometimes forming a basal rosette. This species is common potting mix with good drainage. The plants grow well in
throughout the eastern United States; it also occurs in terrariums or in rock crevices with adequate humidity.
South Africa. Asplenium prolongatum is characterized by flattened
Asplenium platyneuron has been hybridized in cultiva- stipes and oblong to narrowly triangular blades. The two-
tion with the Asian Camptosorus sibericus to form A. ×cru- pinnate fronds have rachises extended into a whip-like tail
cibuli. It is also one of the parents involved in the naturally bearing buds at the tip. It is native to India, China, and
occurring hybrids A. ×ebenoides and A. pinnatifidum. Japan.
 F  F A  C 203

Asplenium kenzoi S. Kurata is presumably a hybrid of

this species and A. antiquum. It resembles A. prolongatum
but is less divided and more leathery.
b
Asplenium resiliens Kunze  13.14.26
Black-stem spleenwort
Hardy, Zone 6
A small fern with erect rhizomes. Grows best under me-
dium light in moist, basic potting mix. The plants are mod-
erately easy to grow.
Asplenium resiliens resembles A. platyneuron but has
smaller, rounder pinnae, and the blades are often bluish
green in color. It is apogamous, producing 32 diploid
spores per sporangium. Native to the United States, Mex-
ico, the West Indies, and South America, it grows on lime-
stone rocks.

Asplenium ruta-muraria Linnaeus  13.14.27

Wall-rue
a
Very hardy, Zone 4
A small fern with clumped rhizomes. Grows under me-
dium light in basic garden soil with sand or potting mix.
This species tolerates moist to dry soil but needs good
drainage and air circulation. It is difficult to establish.
Asplenium ruta-muraria is characterized by irregularly
Figure 13.14.24. two-pinnate fronds, and the pinnules vary from rounded to
Asplenium polyodon: fan-shaped. This species is native to North America, Eu-
a. frond, bar = 5 cm rope, and Asia. It typically grows on limestone.
(2 in.); b, c. pinna
c variations, bar =
2 cm (0.8 in.). Figure 13.14.26. Asplenium resiliens:
fronds, bar = 5 cm (2 in.).

Figure 13.14.25.
Asplenium
prolongatum: habit,
bar = 5 cm (2 in.).

Figure 13.14.27. Asplenium ruta-muraria:

habits, bar = 5 cm (2 in.).
 204 C 13

Asplenium septentrionale (Linnaeus) Hoffmann

 13.14.28
Forked spleenwort
Very hardy, Zone 4
A small fern with clumped rhizomes. Grows under me-
dium light in moist-dry potting mix. Water sparingly and a
give good drainage. The species is sensitive to root disturb-
ances and is difficult to establish. It is infrequently offered
by specialty nurseries as a novelty. Figure 13.14.29.
Asplenium septentrionale is characterized by linear blades Asplenium serratum:
and narrow, ascending segments. In general aspect, the a. fronds, bar = 5 cm
plants resemble tufts of grass. They usually grow on lime- (2 in.); b. sori,
stone. This species is native to North America, Europe, and indusia, and free
Asia. vein tips, bar = 2 cm
(0.8 in.).

Figure 13.14.28. Asplenium

septentrionale: fronds, bar =
5 cm (2 in.). b

Asplenium serratum Linnaeus  13.14.29

Tender, Zone 10
A medium-sized fern with erect, stout, unbranched rhi-
zomes. Grows under medium light in moist potting mix
or uncut moss.
Asplenium serratum resembles the Old World bird’s-nest
ferns (such as A. antiquum, A. australasicum, and A. nidus)
but differs by having veins that are free at the tips, not con-
nected. It is not as robust as the Old World species and is
infrequently grown. As presently circumscribed, A. serra-
tum is extremely variable, apparently consisting of several
closely related species that need to be distinguished through
further research. It is native to tropical America.

Asplenium surrogatum P. S. Green  13.14.30

syn. Asplenium mayii Horticulture, A. pteridoides of
authors, not Baker
Semi-tender or hardier
A medium-sized fern with short-creeping to suberect rhi-
zomes. Grows under medium light in moist garden soil or
potting mix.
The fronds are oblong to ovate-oblong, one-pinnate-
pinnatisect, leathery, and shiny. Asplenium surrogatum has
linear-triangular and deeply pinnatisect pinnae, with the Figure 13.14.30.
tips of the lobes irregularly cleft into sharp teeth. The species Asplenium surrogatum:
is endemic to Lord Howe Island in the southwest Pacific. frond, bar = 5 cm (2 in.).
 F  F A  C 205

The plant called Asplenium mayii in horticulture is the The fronds of Asplenium thunbergii are narrowly oblong,
same as this species. In 1894 the original young plant of A. up to two-pinnate near the base, with numerous pinnae di-
mayii was presumed to be a horticultural hybrid of A. bap- vided into narrowly oblong, one-veined segments. There
tistii and A. pteroides, a species sometimes confused with A. is one sorus per segment. The rachis is broadened by nar-
surrogatum. row wings on both sides, and buds form at the rachis-costa
junctions. The species is native from southeastern Asia to
Asplenium thunbergii Kunze  13.14.31 Indonesia and Borneo.
syn. Asplenium belangeri (Bory) Kunze
Tender Asplenium trichomanes Linnaeus
A medium-sized fern with suberect, branched, clumped  13.14.32;  13
rhizomes and bud-bearing fronds. Grows under medium Maidenhair spleenwort
light in moist, well-drained garden soil or potting mix. The Very hardy, Zone 2?
plants are readily attacked by slugs and snails. Damage from A small fern with erect rhizomes. Grows under low light in
these pests can be lessened by growing the plants in hang- moist potting mix. Some plants adapt more easily to culti-
ing pots. The species is easily propagated by leaf buds. vation than others. Plants from China have adapted well
to gardens in southern California. Asplenium trichomanes
is an evergreen fern. The fronds often grow in a rosette pat-
tern.
Asplenium trichomanes is characterized by dark rachises
and one-pinnate blades with rounded to oblong pinnae. It
resembles A. resiliens, but the pinnae are less elongate and
the blade tissue is thinner and a brighter green (not dull
blue-green). A cosmopolitan species, A. trichomanes occurs
on every continent but mainly in temperate areas.
An infrequently seen but closely related species is Asple-
nium tripteropus Nakai, from eastern Asia, which differs by
rooting at the frond tip.
In North America Asplenium trichomanes consists of two
chromosomal races: one with two sets of chromosomes
(diploid) and another with four (tetraploid). The two races
differ in habitat, with the diploids tending to occur on
acidic rocks such as sandstones, granites, and basalts, and
the tetraploids occurring on basic rocks such as limestones
and dolomites (Moran 1982).
‘Cristatum’. A crested form.
‘Incisum’. Pinna margins incised.

Figure 13.14.32. Asplenium

trichomanes: fronds, bar =
5 cm (2 in.).

Figure 13.14.31.
Asplenium thunbergii:
frond, bar = 5 cm (2 in.).
 206 C 13

Astrolepis beitelii (Mickel) D. M. Benham & Windham

 13.15.1
syn. Cheilanthes beitelii Mickel
Beitel’s cloak fern
Figure 13.14.33. Asplenium Semi-hardy
trichomanes-ramosum: habit, A medium-sized fern with compact to short-creeping rhi-
bar = 5 cm (2 in.). zomes and clustered fronds. Grows under high light in
well-drained, moist to moist-dry garden soil with sand.
This species prefers heat.
Astrolepis beitelii resembles A. sinuata but usually differs
by the larger pinnae, 13–35 mm (0.5–1.5 in.) long, with
three to five pairs of lobes. The upper surface of the blade
has star-shaped scales to 1 mm long, which are usually de-
ciduous. The species grows from central Mexico to Guate-
Asplenium trichomanes-ramosum Linnaeus mala, often on rocky slopes.
 13.14.33
syn. Asplenium viride Hudson Astrolepis sinuata (Lagasca ex Swartz) D. M. Benham
Green spleenwort  13.15.2
Very hardy, Zone 2 syn. Cheilanthes sinuata (Lagasca ex Swartz) Domin,
Notholaena sinuata (Lagasca ex Swartz) Kaulfuss
A small fern with erect rhizomes. Grows under low light in
Semi-hardy, Zone 7(8)
basic, well-drained, moist potting soil. This species is in-
frequently cultivated and difficult to grow. A small to medium fern with compact to short-creeping
Asplenium trichomanes-ramosum resembles A. trichoma- rhizomes and clustered fronds. Grows under high light in
nes but differs by having a green rachis. It usually grows on
limestone and is native to North America, Europe, Asia,
and Africa.

Astrolepis
Star-scale cloak fern
The genus Astrolepis consists of small to medium ferns that c
typically grow in dry, rocky places. They tolerate dryness
and make excellent rock garden plants.
The blades of star-scale cloak ferns are linear to linear-
oblong, one-pinnate to pinnate-pinnatifid, and somewhat
leathery. The segments are shallowly lobed or entire and
covered with star-shaped scales on the upper surface. The
sporangia run along the veins and lack an indusium, the
segment margins being more or less flat, not strongly en-
rolled, and of the same color and texture as the rest of the
blade tissue (not differentiated as in other cheilanthoid
ferns). b
Until recently, the species of Astrolepis were classified
with either Notholaena or Cheilanthes; however, they can
be separated from these genera by the star-shaped scales on a
the upper surfaces of the blades and the two vascular bun-
dles in the stipe (the other genera have only one). See Chei-
lanthes for a comparison to that genus and other cheilan-
thoid ferns.
Astrolepis is native to the Americas and contains about
eight species. The common and generic names refer to the
star-shaped scales on the upper surfaces of the blades: astro Figure 13.15.1. Astrolepis beitelii: a. frond, bar = 2.5 cm
is Greek for “star” and lepis means “scale.” (1 in.); b. upper surface of lobe with scales, bar = 1 mm,
by H. Fukuda; c. star-shaped scale, bar = 1 mm, by H.
Fukuda. Courtesy of John T. Mickel.
 F  F A  C 207

Athyrium  13.16.1

The lady fern (Athyrium filix-femina) and Japanese painted
fern (A. niponicum ‘Pictum’) are well-known representa-
tives of the genus Athyrium. The lady fern comes in a be-
wildering array of garden variations that are eye-catching in
c their unusual patterns. For example, the foliage can be reg-
a ularly crisscrossed, finely divided, or bunched to form a
dense, ball-like tuft. Athyrium is a favorite in temperate cli-
mates and is easy to grow, but the plants tend to have brit-
tle fronds that break easily. In subtropical or warmer cli-
mates some species languish and die within a few seasons.
Fertilizing and periodically replanting in fresh humusy soil
can prolong the life of these plants. In arid climates it is
necessary to keep the soil moist for best growth. Many spe-
cies grow readily from spores.
The genus consists of terrestrial ferns, not epiphytes.
They have long- to short-creeping, ascending, or erect rhi-
b zomes with nonclathrate scales. The blades are linear to
broadly triangular, anadromic at least toward the base, one-
pinnate or more divided, and usually herbaceous. The
rachises have a V-shaped groove on the upper surface, and
this groove is continuous with that of the costae. The
grooves are often minutely hairy within. In many species a
small green spine is found on the edge of the groove at the
base of the costae or costules. The veins can be free or
united. The sori are single, not in pairs (or rarely so), and
they vary from linear to oblong to J-shaped or, less fre-
quently, round. An indusium is present, rarely absent.

Figure 13.15.2. Astrolepis sinuata: a. frond, bar = 2.5 cm

(1 in.); b. upper surface of lobe with scales, bar = 1 mm,
by H. Fukuda; c. star-shaped scale, bar = 1 mm, by H.
Fukuda. Courtesy of John T. Mickel.

drained, moist-dry to dry garden soil with sand. The plants

seem to prefer heat and are relatively easy to grow.
The lower surface of the pinnae bear minute glandular
hairs beneath the scales. Astrolepis sinuata resembles Chei-
lanthes bonariensis due to the linear, one-pinnate-pinnatifid
blades, but it differs by having ciliate scales instead of long
matted hairs on the lower surface of the pinnae. It also re-
sembles A. beitelii but has smaller pinnae, only 12–20 mm
(0.5–0.75 in.) long, with four to six pairs of lobes. It further
differs by the star-shaped scales on the upper surface of the
pinnae being shorter, to 0.5 mm long, and usually persist-
ent. The species is native to the southwestern United States,
Central America, the West Indies, and South America.

Figure 13.16.1. Athyrium filix-femina: habit.

208 C 13

A good characteristic to use for distinguishing Athyrium

from similar genera such as Arachniodes, Dryopteris, and
Polystichum is the presence of only two vascular bundles in
the stipe. These bundles are concave, with the inner sides
facing each other, and unite upward to form a U shape, not
an X shape as in Asplenium. Thelypteris also has only two
vascular bundles in the stipe, but it usually differs by having
one-pinnate-pinnatifid blades and (usually) needle-shaped
hairs.
Athyrium occurs worldwide, especially at higher eleva-
tions, and contains about 180 species. The genus name
comes from the Greek athyros, doorless. What this refers to
Diplazium: Some sori (usually the basal ones) paired back-
is uncertain.
to-back along the same vein, mostly linear; rachis grooves
Certain species in this genus are sometimes united with
U-shaped or flat-bottomed and continuing into the costae;
Deparia or Diplazium, which are recognized separately in
conspicuous multicellular hairs absent on the blade.
this work. To assist in identification, the salient features of
(Diplazium australe, D. dilatatum, D. esculentum, D. lon-
the genera are summarized here.
chophyllum, D. proliferum, D. pycnocarpon, D. subsinua-
tum, D. tomitaroanum)

Athyrium alpestre (Hoppe) Clairville  13.16.2

syn. Athyrium distentifolium var. americanum (Butters)
Cronquist
American alpine lady fern
Hardy, Zone (4)5
A medium-sized fern with short-creeping to suberect rhi-
zomes and deciduous leaves. Grows under medium light
in acidic, moist to wet garden soil or potting mix. This spe-
cies is difficult to grow.
Athyrium: Sori single, not paired back-to-back on the same
Athyrium alpestre resembles a small lady fern except that
vein, usually linear to oblong, sometimes J-shaped; rachis
the sori are round and the indusia are reduced or absent. It
grooves V-shaped and opening to admit the groove of the
is native to alpine and subalpine areas in North America,
costae; conspicuous multicellular hairs absent on the blade.
Europe, and Asia.
(Athyrium alpestre, A. delavayi, A. distentifolium, A. filix-
femina, A. niponicum, A. otophorum, A. vidalii)
Athyrium delavayi H. Christ  13.16.3
Semi-hardy (?)
A medium-sized fern with erect, sparingly clump-forming
rhizomes. Grows under medium light in moist garden soil
or potting mix. The plants have a delicate aspect and at-
tractive reddish costae.
Athyrium delavayi has ovate-oblong fronds with linear
pinnae and small, nearly rhomboid pinnules. The basal
pinnule is longer than the others and often pinnatifid and
overlapping the rachis. The species is native to China.

Athyrium distentifolium Tausch ex Opiz

 13.16.4
Alpine lady fern
Very hardy, Zones 1–4(?)
Deparia: Sori paired or not, variously shaped; rachis This species resembles Athyrium alpestre but is more
grooves variously shaped but not opening into the costae; coarsely cut, shorter, and obviously indusiate. It is easier to
conspicuous multicellular hairs present on the blade. (De- cultivate than A. alpestre. Athyrium distentifolium is native
paria acrostichoides, D. japonica, D. petersenii) to Europe and Asia.
 F  F A  C 209

Figure 13.16.4. Athyrium

Figure 13.16.2. Athyrium alpestre: distentifolium: frond, bar =
a. habit, bar = 5 cm (2 in.); b. pinnule 5 cm (2 in.).
with roundish sori and poorly developed
indusia, bar = 5 mm (0.2 in.).

Athyrium filix-femina (Linnaeus) Roth

 13.16.1, 5–9
Lady fern
Hardy, Zone 4
A medium-large fern with short-creeping to erect, clump-
forming rhizomes and deciduous fronds. Grows best in low
to high light in garden soil or potting mix kept amply
moist-wet during the growing season. Most varieties are
easily cultivated.
Athyrium filix-femina is extremely variable but generally
is characterized by lanceolate to ovate, two-pinnate-pin-
natifid blades and elongated or hooked indusia. It is wide-
spread, occurring in North America, South America, Eu-
rope, and Asia.
The many cultivars of lady fern are derived from the Eu-
ropean variety, var. filix-femina, and they differ in frond
shape: highly branched, twisted, widened, narrowed, re-
duced, crested, crisscrossed, congested, rounded, or jagged
(see Figure 12.2). Many cultivars must be propagated by
divisions because they do not breed true from spores. Of
the hundreds of cultivars, only a few are listed here. Some
pteridologists raise the varieties listed here to the rank of
species.
Figure 13.16.3. Athyrium var. angustum (Willdenow) G. Lawson (Athyrium an-
delavayi: frond, bar = gustum (Willdenow) C. Presl). Northern lady fern. Figure
5 cm (2 in.). 13.16.5. This variety is hardy to Zone 2. It is distinguished
by stipes nearly as long as the blades, elliptic to lanceolate
blades broadest near or just below the middle, and yellow
spores. It is native to central to northeastern Canada and
the United States.
 Figure 13.16.7. Athyrium
filix-femina var. cyclosorum:
Figure 13.16.5. frond, bar = 5 cm (2 in.).
Athyrium filix-
femina var.
angustum: frond,
bar = 5 cm
(2 in.).

Figure 13.16.8.
Figure 13.16.6. Athyrium filix-femina var. asplenioides: Athyrium filix-
a. frond, bar = 5 cm (2 in.); b. pinnule with elongate sori femina var. filix-
and indusia, bar = 3 mm (0.1 in.), courtesy of Knobloch femina: frond, bar
and Correll (1962); c. rhizome, bar = 2 cm (0.8 in.), = 5 cm (2 in.).
courtesy of Knobloch and Correll (1962).
 F  F A  C 211

‘Frizelliae’. Figure 12.2.7. Fronds small-medium, lin-

ear, with the pinnae reduced to a congested cluster of fan-
shaped, deeply toothed pinnules or segments.
‘Minutissimum’. Dwarf lady fern. Resembles var. filix-
femina but differs by having denser foliage with light green
fronds up to 15 cm (6 in.) tall.
‘Veroniae-cristatum’. Zone 4. Stipes crimson, blades
broadly triangular, pinnae strongly ruffled, overlapping,
and ending in a slender tassel.
‘Victoriae’. Figure 12.2.13. Blades small and narrow
with crested apices; the pinnae appearing crisscrossed,
forming an X shape with the rachis at the center.

Athyrium niponicum (Mettenius) Hance

 13.16.10;  14
Very hardy, Zone 3(4)
A medium-sized fern with short-creeping rhizomes and de-
ciduous fronds. Grows best under high light in moist-wet
garden soil or potting mix.
Athyrium niponicum is characterized by broad, two-pin-
Figure 13.16.9. Athyrium nate-pinnatifid, soft herbaceous fronds, with the lower pin-
filix-femina ‘Cristatum’: nae amply stalked. A herbaceous spine projects upward
frond, bar = 5 cm (2 in.). from the base of costae on the upper surface. The indusia
are elongate and often hooked or J-shaped. This species is
native to China and Japan.

var. asplenioides (Michaux) Farwell (Athyrium asple-

nioides (Michaux) Hultén). Southern lady fern. Figure
13.16.6. This variety is hardy to Zone 5. It is distinguished
by stipes shorter than the blades, ovate-lanceolate to lance-
olate blades that are broadest just above the base, and dark
brown spores. Native to the southeastern United States.
var. californicum Butters. California lady fern. This va-
riety resembles var. cyclosorum but differs in having brown
spores and narrower, more acute pinnules unequal at the
base. It is slightly more difficult to grow than var. cyclosorum
and is rarely cultivated.
var. cyclosorum Ruprecht. Northwestern lady fern. Fig-
ure 13.16.7. This variety is hardy to Zone 3(4). The stipe is
half the length of the blade or less, blades are elliptic to ob-
lanceolate, and spores yellow. It commonly volunteers in
peat moss. Native from Alaska to the northwestern United
States.
var. filix-femina. European lady fern. Figures 13.16.1, 8.
This variety is hardy to Zone 3. It resembles var. cycloso-
rum except that the fronds are more lanceolate. It is easy to
grow but the fronds break readily. Native to Europe. Nu-
merous cultivars have been derived from this variety, some
of which are shown in Figure 12.2. The more commonly
cultivated ones are the following: Figure 13.16.10.
‘Cristatum’. Crested lady fern. Figures 12.2.27, 13.16.9. Athyrium niponicum
Fronds medium-sized, the apices and pinnae heavily crested. ‘Pictum’: frond, bar =
‘Cruciato-cristatum’. Pinnae forked at their base, with 5 cm (2 in.).
the divisions crisscrossing over adjacent pinnae; tips crested.
 212 C 13

Plants of Athyrium niponicum are often misidentified in Athyrium otophorum is characterized by broad, triangu-
the trade as A. iseanum Rosenstock or its synonym A. goe- lar, two-pinnate-pinnatifid, slightly leathery fronds with
ringianum (Kunze) T. Moore. Athyrium iseanum is not long, narrowly triangular pinnae. The stipe, rachis, and
known in cultivation in the United States. costa are reddish tinged. The species is native to Japan,
‘Pictum’. Plate 14. Japanese painted fern. A variegated Korea, and China.
form with reddish and gray-silver colors. The colors de- Athyrium epirachis (H. Christ) Ching, a species from
velop best in high light. Crested, ruffled, and other forms China and Japan with similar red-tinged rachises and
have been reported. The ghost fern, a suspected hybrid be- costae, has been introduced into cultivation. It differs from
tween ‘Pictum’ and Athyrium filix-femina, is becoming A. otophorum by narrower, less foliaceous blades and erect
available commercially and shows great horticultural prom- rhizomes. It is a more vigorous grower and more evergreen
ise. Its medium-sized leaves are a dull, “ghostly” grayish than A. otophorum.
green.
Athyrium vidalii (Franchet & Savatier) Nakai
Athyrium otophorum (Miquel) Koidzumi  13.16.12
 13.16.11;  15 Semi-hardy to hardier, Zone 6
Hardy, Zone 4(5) A medium-large fern with ascending to suberect rhizomes
A medium-sized fern with short-creeping, clump-forming and clustered, deciduous fronds. Grows well under me-
rhizomes and deciduous fronds. Grows under medium dium light in moist garden soil or potting mix.
light in moist garden soil or potting mix. Athyrium vidalii is characterized by triangular, two-pin-
nate to two-pinnate-pinnatifid blades, sessile to short-
stalked pinnae, spines present on the costule or base of the
pinnule midribs (upper surface), and elliptic to oblong sori.
The indusium is persistent, conspicuous, lunate, hooked
or horseshoe-shaped, with margins irregularly toothed or
entire. The species is native to China, Japan, Korea, and
Taiwan.

b
b

a
a

Figure 13.16.11. Athyrium otophorum: a. frond, bar =

5 cm (2 in.); b. pinnule with elongate sori and indusia, Figure 13.16.12. Athyrium vidalii: a. frond, bar = 5 cm
bar = 5 mm (0.2 in.). (2 in.); b. spine at base of pinnule, bar = 5 mm (0.2 in.).
 F  F A  C 213

Azolla  13.17.1–3

Mosquito fern, fairy moss
Azolla consists of floating ferns that are grown as a novelty
in aquariums, bowls, and ponds. These small, moss-like
plants grow in warm, sluggish water or nearby muddy soils.
They tend to die in cold weather after forming buds, which
sink to the bottom of the pond and resume growth when
warm weather returns. Mosquito ferns prefer full sunlight,
warm temperatures, and silty water containing organic mat-
ter. High light encourages reddish growth. The common
name is derived from the fern’s supposed ability to discour-
age mosquito reproduction by densely carpeting the water’s
surface, thereby preventing the adults from laying eggs and
preventing the larvae from getting air at the surface. Figure 13.17.2. Azolla filiculoides: habit, enlarged.
The plants have minute, plump, scale-like leaves. Fila-
mentous roots extend down from the thread-like rhizome,
which branches repeatedly and bears fronds in two rows. This genus occurs worldwide and has about seven spe-
The fronds are bilobed, with the upper lobe fleshy and the cies. Its name comes from the Greek azo, to dry, and olluo,
lower lobe inconspicuous, thin, colorless, and submerged. to kill, presumably alluding to the fern’s tendency to be
The sori, which are rarely present, are round, whitish struc- killed by drying. Azolla is the world’s most important eco-
tures borne on the lower surface of the plant. There are two nomic fern because it is used as an organic fertilizer in rice
kinds of sori, one bearing female spores (megaspores) and paddies in China and Vietnam. The plants are a rich source
the other male spores (microspores). of nitrogen, a nutrient often limiting to plant growth. The
Identifying the species of Azolla is difficult because the
distinguishing characteristics are microscopic. The num-
ber of cells composing the leaf hairs is helpful in distin-
guishing the species, but a magnification of at least 40× is
needed to see this characteristic. A scanning electron micro-
scope is needed to see surface details on the female spores,
and even when these details can be seen, they can be diffi-
cult to interpret. Moreover, Azolla plants rarely bear spores,
so these structures are often unavailable for identification.

Figure 13.17.3. Azolla, bar = 2 mm: a. Azolla

Figure 13.17.1. Azolla filiculoides: habit. caroliniana; b. Azolla filiculoides; c. Azolla pinnata.
 214 C 13

nitrogen enters the plant by the action of a cyanobacterium Azolla pinnata is broadly triangular and regularly pin-
(blue-green algae), Anabaena azollae, that lives within the nately branched. The roots are branched and feathery, un-
leaves of Azolla. The cyanobacterium combines, or “fixes,” like the other species in the genus, all of which have un-
atmospheric nitrogen with hydrogen to make ammonia, a branched, nonfeathery roots. It occurs in tropical Asia,
molecule that can be taken up by organisms and used to Africa, Australia, New Zealand, and New Caledonia.
form nitrogen-containing compounds. Besides its use as
fertilizer, Azolla is also fed to livestock and certain fish. For Belvisia
a popular account of the economic importance of Azolla, Members of this genus are small- to medium-sized, epi-
see Moran (1997); more technical accounts can be found in phytic ferns with undivided, narrow fronds bearing a fertile
Moore (1969), Lumpkin and Plucknett (1982), van Hove spike at the tip. They are infrequently cultivated but are
(1989), and Wagner (1997). generally grown in greenhouses or, in tropical areas, out-
doors in pots or baskets. The gracefully arching fronds are
Azolla caroliniana Willdenow  13.17.3a closely spaced and best displayed in an elevated position.
Hardy, Zone (4)5; survives frost and ice The most distinctive characteristics of Belvisia are the
Small-sized. Prefers high light. This floating fern is the best simple leaves with narrowed, tail-like fertile tips. The rhi-
species for growing on mud. The plants turn crimson in zome is short- to long-creeping, unbranched, and covered
the fall, and several weeks later they die and sink to the bot- by black or red-brown, clathrate scales. The veins are netted
tom. with irregular areoles containing forked veinlets. The sori
Azolla caroliniana plants are roughly circular to trian- lack indusia and are spread on the lower surface of the tail-
gular and about the size of a dime. The largest hairs (papil- like tip. Minute, scale-like paraphyses are mixed among the
lae) on the upper surfaces of the leaves are two-celled. The sporangia.
megaspores lack angular bumps or pits and are densely cov- The genus, which consists of eight species, occurs in the
ered with tangled filaments. Compared to A. filiculoides, Old World tropics, northeastern Australia, and Polynesia in
the clumps of A. caroliniana tend to fragment more often tropical and warm-temperate climates. It is named for
and are more compact and rounder in outline. The species A. M. F. J. Palisot de Beauvois (1752–1820), a French trav-
occurs in the Americas, Europe, and Asia. eler and agrostologist (grass specialist) whose Latinized
Azolla mexicana C. Presl is less hardy but may be in cul- name is Belvisius. For a technical treatment of Belvisia, see
tivation. It differs by having megaspores sparingly covered Hovenkamp and Franken (1993).
with tangled filaments.
Belvisia mucronata (Fée) Copeland  13.18.1
Azolla filiculoides Lamarck  13.17.1, 2, 3b syn. Belvisia callifolia (H. Christ) Copeland
syn. Azolla filiculoides var. rubra (R. Brown) Strasburger, Tender
A. rubra R. Brown A medium-sized fern with short-creeping rhizomes. Best
Hardy, Zone (6)7 grown under medium light in well-drained, moist potting
Small-sized. Grows best in high light. This floating species mix or uncut moss.
forms dense mats on the water’s surface. Growth peaks in The blades of Belvisia mucronata vary tremendously, es-
late spring and noticeably declines in winter. The plants pecially in width, ranging mostly from 1 to 5 cm (0.5–2
can survive under thin ice. Most of the cultivated material in.) wide, and they are gradually narrowed below the fertile
in southern California is Azolla filiculoides. spike. The rhizome scales have dentate to ciliate margins,
Clumps of Azolla filiculoides are less likely to fragment and the cells of the main body of the scales are all thick.
than those of A. caroliniana, and they have a more open, The species is native to Sri Lanka, southeastern Asia, Ma-
branched aspect. The largest hairs on the upper leaf lobe laysia, Australia, and the Pacific Islands.
are one-celled. The megaspores are warty with raised an-
gular bumps. This species occurs in the Americas, Europe, Blechnum  13.19.1, 2
northeastern Asia, southern Africa, and the Pacific Islands. syn. Lomaria
Most Blechnum species are terrestrial plants with leaves that
Azolla pinnata R. Brown  13.17.3c;  16
are reddish when young but turn green with age. Nearly all
Ferny azolla
the species have pinnatifid or one-pinnate blades; never are
Tender
they finely cut. The trunk-forming species, such as Blech-
Small-sized. Best grown under high light. This floating fern num brasiliense and B. gibbum, are particularly valued in
has been used for centuries in China and Vietnam as a pots for accent use. The species with creeping rhizomes are
green manure for growing rice. The federal and some state useful as ground covers or bedding plants. Blechnum aus-
governments list Azolla pinnata as an aquatic weed and pro- trale, B. penna-marina, and B. spicant are favored in warm-
hibit its import, possession, and distribution. temperate regions, whereas B. appendiculatum is used in
 F  F A  C 215

Figure 13.19.2. Blechnum appendiculatum: sori and

indusia.

b subtropical climates. Many species apparently favor acidic

a
soils, such as those suitable for azaleas and rhododendrons.
Spores are often difficult to germinate.
The Australian and New Zealand species are widely cul-
Figure 13.18.1. Belvisia mucronata: tivated in their native countries, and some of these are being
a. habit, bar = 5 cm (2 in.); b. sporangia tried in the United States. Blechnum fraseri (Cunningham)
spreading on the frond tip, bar = 1 cm Luerssen is a small, attractive tree-like species from New
(0.4 in.). Zealand but is, unfortunately, difficult to grow in the
United States. A large, trunk-forming species from Costa
Rica has also been tried but without success. Blechnum
tabulare (Thunberg) Kuhn seems to be established in the
South African trade. It resembles a cycad and would make
an attractive addition to United States gardens.
The genus usually can be recognized by the pinnatifid or
one-pinnate fronds, with long sori running parallel to and
on both sides of the costae. Nearly all the species are terres-
trial, although some are hemiepiphytes, bearing long-creep-
ing rhizomes that climb tree trunks. The veins are free. If
the sterile and fertile leaves are dimorphic, the fertile ones
are often longer stalked and have narrower pinnae than the
sterile ones.
The species with upright trunks seldom form offshoots,
but others can be propagated by rhizome divisions or from
stolons (if produced). Blechnum stolons are long and slen-
der, and they are produced from a short, erect rhizome.
When their tips emerge from the soil, they produce new
plantlets that can, after they have rooted, be detached from
the main plant and transplanted.
Blechnum consists of about 150 to 200 species world-
wide, most of which grow in the Southern Hemisphere.
The genus name comes from the Greek blechnon, which is
an ancient name for ferns.

Blechnum amabile Makino  13.19.3

Hardy to semi-hardy, Zones 7 to 8
A medium-sized fern with creeping rhizomes and clustered
Figure 13.19.1. Blechnum appendiculatum: habit. fronds. Grows best under medium light in moist garden
 216 C 13

Figure 13.19.3.
Blechnum amabile:
habit, showing
dimorphic fertile
frond (center) and
sterile fronds, bar =
5 cm (2 in.).

a b
soil or potting mix. Fronds gradually wither in place upon
light frost. This species is a slow grower.
Blechnum amabile bears dimorphic, one-pinnate fronds.
The sterile blades are lanceolate and narrow, 2–4 cm (0.75–
1.5 in.) wide, and the fertile fronds are barely longer. Ster-
ile pinnae are numerous, linear, and decurrent on the
rachis, with the lower pinnae shorter and triangular. The Figure 13.19.4. Blechnum appendiculatum variants:
fertile pinnae are narrower and more contracted than the a. monomorphic sterile frond of the less red form with
sterile ones. The species is native to China and Japan. glandular hairs; b. monomorphic sterile frond of the
This plant is similar to Blechnum niponicum, which see. redder form with needle-like hairs. Bar = 5 cm (2 in.).

Blechnum appendiculatum Willdenow

 13.19.1, 2, 4;  1
syn. Blechnum glandulosum Link, B. occidentale var. Blechnum australe Linnaeus  13.19.5
minor Hooker Hardy to semi-hardy, Zone 7(8)
Semi-hardy, Zone 8 A small to medium fern with erect, stolon-forming rhi-
A small to medium fern with whorled fronds. Grows best zomes and fronds usually in an uneven whorl or cluster.
under medium light in moist-dry garden soil or potting Best grown under medium to high light in moist garden
mix. This species spreads readily by stolons. soil or potting mix. The plants spread easily from stolons
Blechnum appendiculatum greatly resembles B. occiden- and are easy to cultivate.
tale, and cultivated plants have long been misidentified as Blechnum australe has slightly dimorphic, one-pinnate
that species. Blechnum appendiculatum differs, however, by fronds. The sterile fronds are narrowly elliptic with lance-
being hairy on the rachis underside. It is native to the olate pinnae, and the pinnae bear a mucronate lobe or “ear”
southern United States and American tropics. on the upper side at the base. The fertile fronds are con-
Two kinds of Blechnum appendiculatum, apparently rep- tracted, though the lower pinnae may be similar to the ster-
resenting different species, are cultivated. The first (Figure ile pinnae. The typical variety, var. australe, is native to
13.19.4b) has pointed hairs on the lower surface of the southern Africa, Madagascar, and the South Atlantic is-
rachis, and the red of the young stipes and rachises is mostly lands.
retained as the frond matures. The second type (Figure Blechnum australe var. auriculatum (Cavanilles) de la
13.19.1, 2, 4a) has minute capitate-glandular hairs (the Sota occurs in Brazil, Uruguay, and Argentina.
glands pale yellow) on the lower surface of the rachis and
blade tissue. Although reddish when young, the stipe is less Blechnum brasiliense Desvaux  13.19.6
intensely red than that of the first type, and the color is lost Semi-tender to tender
as the fronds mature, so the stipes and rachises become A medium to large fern with a trunk-like stem (in older
greenish. In addition, the first type has darker green blades plants) and whorled fronds that collectively form a funnel
than the second. or basket. Best grown under medium to high light in moist
 F  F A  C 217

garden soil or potting mix. This species is sensitive to trans-

planting. Some variants have redder juvenile fronds than
others.
Blechnum brasiliense is characterized by numerous
sharp-pointed, ascending pinnae and oblanceolate fronds
c gradually tapered to the base. The stems form a short trunk
and are covered by dark brown or blackish scales. The ster-
ile and fertile fronds are monomorphic. The species is na-
tive to Guatemala, Ecuador, Peru, and Brazil.
‘Crispum’. Pinna margins slightly ruffled.

Blechnum capense (Linnaeus) Schlechtendal

 13.19.7
a Semi-hardy
A large fern with short-creeping, branched rhizomes and
clustered fronds. Grows under medium to high light in
(acidic?) garden soil or potting mix kept moist or moist-wet.
This species has proven difficult to grow in some gardens.
Blechnum capense has one-pinnate, dimorphic fronds.
Figure 13.19.5. Blechnum The sterile fronds are elliptic with 10–50 pairs of crowded,
australe: a. sterile frond, bar = oblong to linear pinnae. The basal pinnae are highly reduced
5 cm (2 in.); b. dimorphic and rounded. The pinnae on the fertile fronds are linear but
fertile frond, bar = 5 cm are rounded and sterile at the base of the blade. The species
(2 in.); c. pinnae with (as recognized here) is endemic to New Zealand.
mucronate tip, bar = 5 mm b The name Blechnum capense applies to several poorly
(0.2 in.). known species. As used here, the name applies to the com-
mon New Zealand plant known in that country as kiokio.

Figure 13.19.6. Blechnum brasiliense:

a. monomorphic fertile frond;
b. sterile frond from young plant.
Bar = 5 cm (2 in.).

a b

Figure 13.19.7. Blechnum capense: a. sterile frond;

a b b. dimorphic fertile frond. Bar = 5 cm (2 in.). After
Brownsey and Smith-Dodsworth (1989).
 218 C 13

Plants circulating as B. capense in the United States may gins strongly enrolled when old. The plant is native to
not be the New Zealand species. South America.
In the United States and England, plants circulating as
Blechnum cordatum (Desvaux) Hieronymus Blechnum chilense (Kaulfuss) Mettenius, B. tabulare (Thun-
 13.19.8 berg) Kuhn, and B. magellanicum (Desvaux) Mettenius all
Semi–hardy to hardy, Zone 7 seem to be B. cordatum. Blechnum wattsii Tindale of Aus-
A medium-large fern with erect fronds that may be spaced tralia is rarely cultivated in the United States. It can be dis-
along a creeping rhizome as well as clustered on an erect tinguished from B. cordatum by the absence of aerophores,
rhizome. Grows under medium light in moist-wet garden more oblong pinnae that are abruptly long acuminate at
soil or potting mix. In the northwestern United States it the tip, and fertile pinnae that are farther apart.
may take full sun if well watered. Rhizomes and stolons
below the ground usually survive freezing weather. The Blechnum falciforme (Liebmann) C. Christensen
plants spread by stolons.  13.19.9
Blechnum cordatum has dimorphic fronds. The sterile Semi-tender
fronds have oblong, dark green blades that are somewhat A large fern with ascending to erect rhizomes and whorled
hard and leathery, with the terminal pinna resembling the fronds. Grows under medium light in moist garden soil or
lateral pinnae; the pinnae are oblanceolate and finely potting mix.
toothed. On the lower surface of the rachis opposite each The fronds of Blechnum falciforme are one-pinnate, di-
pinna stalk is a small, elongated bump, a type of aerophore. morphic, and noticeably scaly. The stipe has a dense mat-
Aerophores may be inconspicuous or absent on dried fronds ting of twisted hair-like scales, and the sterile pinnae have
but are easy to see on fresh fronds. They are usually whitish cartilaginous, serrate margins. The basal pinnae are not
yellow and contrast strongly with the color of the sur- reduced, and the undersurfaces of the costae are scaly. The
rounding tissue. The fertile pinnae are linear, with the mar- species is native to Central and South America.

Blechnum gibbum (Labillardière) Mettenius

 13.19.10;  17
Semi-tender to tender
A medium-sized fern with erect, trunk-like stems (in older
plants) and fronds borne in a rosette-like whorl. Grows best
in medium light in moist, well-drained garden soil or pot-
ting mix. This species is somewhat difficult to grow because

a b

Figure 13.19.8. Blechnum cordatum: a. sterile frond from

young plant; b. old dimorphic fertile frond. Bar = 5 cm Figure 13.19.9. Blechnum falciforme: a. sterile frond;
(2 in.). b. dimorphic fertile frond. Bar = 5 cm (2 in.).
 F  F A  C 219

b
a

Figure 13.19.11. Blechnum gracile: a. sterile frond;

b. monomorphic fertile frond. Bar = 5 cm (2 in.).

segment that is equal to or two times as long as the lateral

pinnae. The sterile and fertile fronds have the same form
b (monomorphic), although the fertile ones tend to be more
a erect. The species is native to tropical America.

Blechnum niponicum (Kunze) Makino

 13.19.12
Hardy
Figure 13.19.10. Blechnum gibbum: a. dimorphic fertile
A medium-sized fern with erect rhizomes and fronds ar-
frond; b. sterile frond from young plant. Bar = 5 cm
ranged in a rosette. Grows under medium light in moist
(2 in.).
garden soil or potting mix.

of its sensitivity to transplanting, irregular watering, and

wind.
Blechnum gibbum resembles B. brasiliense but has black
rhizome and stipe scales, dimorphic fronds, and generally
narrower sterile pinnae. It is native to New Caledonia and
islands of the South Pacific.
An unnamed variant with wider and blunter pinnae is a b
often misidentified as Blechnum moorei C. Christensen. Figure 13.19.12.
True B. moorei, which is unknown in cultivation, differs Blechnum niponicum:
by having spinulose-ciliate margins and pinna apices that a. dimorphic fertile
are truncate, notched, or forked. frond; b. sterile frond.
Bar = 5 cm (2 in.).
Blechnum gracile Kaulfuss  13.19.11
syn. Blechnum fraxineum of authors, not Willdenow
Semi-tender
Small-sized. Grows best under medium light in moist gar-
den soil or potting mix. The plants spread readily by sto-
lons.
Blechnum gracile is distinctive by its blade form: one-
pinnate with only one to four pinna pairs and a terminal
 220 C 13

Blechnum niponicum has dimorphic fronds, the fertile latum. The species is widespread in tropical America.
fronds taller than the sterile ones. The sterile fronds are one- Most plants identified as Blechnum occidentale in gar-
pinnate and 5–10 cm (2–5 in.) wide. The scales on the base dens in the United States are actually B. appendiculatum.
of the stipe are linear. This species is native to Japan.
Blechnum spicant resembles B. niponicum but differs by Blechnum penna-marina (Poiret) Kuhn
narrower, less oblanceolate sterile fronds and much taller  13.19.14;  18
fertile fronds. The similar B. amabile has a creeping rhi- Hardy, Zone 5
zome, narrower sterile fronds, shorter fertile fronds, and A small fern with fronds spaced along a much-branched,
ovate stipe scales. creeping, stoloniferous rhizome. Grows under medium or
high light in moist but well-drained, acidic garden soil or
Blechnum occidentale Linnaeus  13.19.13 potting mix. The plants spread by stolons and make a use-
Hammock fern ful ground cover in areas with mild winters.
Probably tender Blechnum penna-marina bears one-pinnate, dimorphic
A small to medium fern with erect rhizomes and clustered fronds, the fertile ones much contracted and about twice as
fronds. Grows under medium light in moist garden soil or tall as the sterile. The species is native to South America,
potting mix. The plants spread by stolons. southeastern Australia, New Zealand, and the circum-
Blechnum occidentale is characterized by blades that are antarctic islands.
one-pinnate at the base but pinnatisect above and blade An unnamed dwarf cultivar circulates in the trade, as
apices that are gradually tapered to the tip. The blades are well as the following:
usually narrowly triangular, with the lowest pinnae point- ‘Cristata’. Tip of frond crested.
ing downward and heart-shaped at the base. The rhizomes
usually bear several stolons. The lower surface of the rachis Blechnum polypodioides Raddi  13.19.15
is glabrous, not hairy as in the closely related B. appendicu- syn. Blechnum unilaterale Swartz
Tender
A small to medium fern with erect, stoloniferous rhizomes
and clustered fronds. Grows under medium light in moist
garden soil or potting mix.
Blechnum polypodioides is characterized by pinnatisect
to one-pinnate, lanceolate fronds tapered at both ends. The
sterile and fertile fronds are monomorphic and minutely
hairy on the lower surfaces. The pinnae tend to be numer-
ous, oblong to linear, and essentially entire, and the lower
ones are distant and much reduced. The species is wide-
spread in tropical America.

Figure 13.19.14.
Blechnum penna-
marina: habit,
showing sterile
frond (shorter
outer fronds) and
dimorphic fertile
fronds (taller
a b inner fronds), bar
= 5 cm (2 in.).

Figure 13.19.15. Blechnum

Figure 13.19.13. Blechnum occidentale: a. monomorphic polypodioides: monomorphic sterile
fertile frond; b. sterile frond. Bar = 5 cm (2 in.). frond, bar = 5 cm (2 in.).
 F  F A  C 221

Blechnum punctulatum Swartz  13.19.16

Hardy to semi-hardy
A medium-sized fern with branched, ascending rhizomes
and clustered fronds. Grows under medium light in moist
garden soil or potting mix. This species is slow growing.
Blechnum punctulatum has dimorphic, one-pinnate
fronds. The sterile fronds vary from lanceolate to linear-el-
liptic and are reduced to short pinnae below. The fertile
fronds are more oblanceolate compared to the sterile ones.
Sori are absent from the lower, shorter pinnae of the fertile
fronds. The blades resemble those of B. australe but have
blunter pinna tips. Blechnum punctulatum is native to
southern Africa.

a b

Figure 13.19.16.
Blechnum
punctulatum: Figure 13.19.17. Blechnum serrulatum: a. monomorphic
a. dimorphic fertile fertile frond; b. young sterile frond. Bar = 5 cm (2 in.).
frond; b. sterile
a
frond. Bar = 5 cm
(2 in.).
leaves are monomorphic. Sori are borne only on the distal
pinnae and are covered by erose-lacerate indusia. This spe-
cies is native to the United States (Florida), the West Indies,
Blechnum serrulatum L. C. Richard  13.19.17 Central and South America, India, Malaysia, and Australia.
syn. Blechnum indicum of authors, not N. L. Burman In the Old World this species has long been called Blech-
Saw fern, swamp fern num indicum N. L. Burman; however, that name is a syn-
Tender, Zone 9 onym of Asplenium longissimum Blume (Morton 1967).
A large fern with creeping rhizomes and clustered, shiny,
thick fronds. Grows under medium to high light or direct Blechnum spicant (Linnaeus) Smith  13.19.18
sun. This species prefers moist garden soil or potting mix Deer fern
but tolerates wet soils. The fronds tend to grow upright in Hardy to semi-hardy, Zone 7
full sun. During the dry season, the pinnae are shed from A medium-sized fern with erect, sometimes-branched rhi-
the rachis by means of the abscission layer, leaving behind zomes and fronds in a rosette or cluster. Best grown under
erect, naked rachises. The species often forms dense colo- low-medium light in moist, well-drained, acidic potting
nies in the wild. mix. Be sure to provide the plants with humidity and acidic
Blechnum serrulatum is characterized by fronds one-pin- soil, and avoid using hard water. This evergreen species has
nate throughout, with 20–40 pinna pairs and a single api- proven difficult to grow in the southwestern United States,
cal segment. The pinna margins are finely serrate, and care- possibly because of high summer heat and humidity. It
ful inspection will reveal that the pinnae are jointed to the grows readily from spores.
rachis, with the joint visible as a faint dark line or swelling. Blechnum spicant bears fronds that are one-pinnate and
The stipes are often densely clustered. The sterile and fertile dimorphic. The sterile fronds vary from lanceolate to nar-
 222 C 13

rowly elliptic, and fertile fronds have taller and narrower

pinnae. The species is native to western North America and
Europe.
Several cultivars exist, including the following, which
are common in the trade:
‘Crispum’. Margins ruffled.
‘Redwood Giant’. Plants large.
‘Serratum’. Margins serrate.

Figure 13.20.1. Bolbitis portoricensis: habit. Photo

courtesy of G. Hampfler.

Figure 13.19.18. Blechnum spicant:

a. sterile frond; b. dimorphic fertile
frond. Bar = 5 cm (2 in.).

Bolbitis  13.20.1–3

syn. Edanyoa
This genus is infrequently cultivated. It consists mostly of
medium-sized, coarse plants that typically grow along rocky
streambanks in the tropics and less frequently in soil or on
tree trunks.
Bolbitis is characterized by creeping rhizomes, dimor-
phic sterile and fertile fronds, and acrostichoid sori. The
sterile fronds vary from simple to one-pinnate (or rarely Figure 13.20.2. Bolbitis portoricensis: sterile frond (left)
two-pinnate). The veins can be either free or netted and, if and contracted fertile frond (right).
netted, with or without veinlets. The fertile fronds have
long stipes that elevate the blades above the sterile fronds.
Bolbitis heteroclita includes a tiny terrestrial (formerly
Edanyoa difformis) that differs greatly from other Bolbitis
species in general appearance. Its small size makes it popu-
 F  F A  C 223

Figure 13.20.3. Bolbitis portoricensis: sporangia spreading Figure 13.20.4. Bolbitis heteroclita: sterile frond, bar =
on under surface. 5 cm (2 in.).

lar for terrariums. Bolbitis heudelotii in the United States plants. Many of the species reduced to B. heteroclita are rec-
and B. fluviatilis in Europe are sometimes sold in aquar- ognized in horticulture and are here given cultivar status.
ium supply stores as underwater plants. Some species are Difformis Group (Edanyoa difformis Copeland). Figure
easy to grow but are rarely seen in cultivation, such as B. 13.20.5. A diminutive creeping fern often used in terrari-
aliena, which needs warm temperatures year-round. ums and easy to grow. Fronds are usually less than two-pin-
Bolbitis contains 85 species and occurs throughout the nate-pinnatifid and lack the terminal elongated pinna of
tropics of the world. The genus name comes from the the species. Native to the Philippines and long known as
Greek bolbition, diminutive of bolbos, bulb, alluding to the the species Edanyoa difformis. Hennipman believes this
small bulblets borne on the leaves of some species. For a plant is a neotenous form (represents juvenile characteris-
technical monograph of the genus, see Hennipman (1977). tics) of Bolbitis heteroclita. The International Code of No-

Bolbitis heteroclita (C. Presl) Ching

 13.20.4–8
syn. Bolbitis simplicifolia (Holttum) Ching, Edanyoa b
difformis Copeland
Tender
a
A medium-sized fern with medium- to long-creeping rhi-
zomes and whip-like blade apices. Grows well under me-
dium light in moist to moist-wet garden soil or potting mix.
The blades of typical Bolbitis heteroclita (Figure 13.20.4)
are narrowly triangular, with entire to coarsely dentate-
lobed lateral pinnae. The terminal pinna is often tailed or
whip-like and bears a bud at the tip. The veins are netted in
irregular areoles with few included veinlets. The species is
native to Asia and Malaysia. Figure 13.20.5. Bolbitis heteroclita Difformis Group:
Bolbitis heteroclita is a variable and complex species, and a. habit, bar = 5 cm (2 in.); b. sterile frond (left) and
several name changes have been required for the cultivated fertile frond (right), bar = 5 cm (2 in.).
224 C 13

menclature for Cultivated Plants allows listing it as a cultivar

in a group category.
‘Malacca’. Figure 13.20.6. This variant is known among
growers in the United States only as a small Bolbitis hetero-
clita. The small, triangular or oblong-triangular fronds and
pinnatifid lower pinnae tend to be consistent features for
recognition. ‘Malacca’ is easy to grow. It seems to be the
same as the cultivated plants Hennipman (1977) lists as
‘malaccensis’, a smaller plant from the Philippines. Because
the latter name is in Latin, it cannot be used as a cultivar
name. Thus the plant is here listed as ‘Malacca’ to meet the
requirement of the International Code of Nomenclature for
Cultivated Plants.
Simplicifolia Group (Bolbitis simplicifolia (Holttum)
Ching). Figure 13.20.7. Plants small-medium; rhizome
creeping; fronds in compact clusters, erect, usually sym-
metrical; pinnae broad-elliptic; terminal pinnae resemble
the lateral pinnae, though usually longer; seldom more than
one pair of lateral pinnae; margins undulate to nearly en-
tire. The plants in cultivation were reportedly introduced
from Malaysia about 30 years ago. Long known as B. sim-
plicifolia, this plant has been reduced to synonymy under B.
Figure 13.20.6. Bolbitis heteroclita ‘Malacca’: habit, heteroclita. The International Code of Nomenclature for Cul-
showing fertile frond third from left, bar = 5 cm (2 in.). tivated Plants of 1995 allows listing it as a cultivar in a
group category.
Tenuissima Group. Figure 13.20.8. Produces highly
variable fronds even on the same plant. The fronds are
often asymmetrical with puckered surfaces. Hennipman

Figure 13.20.7.
Bolbitis heteroclita
Simplicifolia
Group: habit
with sterile Figure 13.20.8. Bolbitis heteroclita Tenuissima Group:
fronds, bar = habits, showing variations in sterile fronds, bar = 5 cm
5 cm (2 in.). (2 in.).
 F  F A  C 225

(1977) listed the plant as ‘tenuissima’, a Latin name mean- Bolbitis portoricensis (Sprengel) Hennipman
ing “more slender.”  13.20.1–3, 10
syn. Bolbitis cladorrhizans (Sprengel) Ching
Bolbitis heudelotii (Bory ex Fée) Alston Tender
 13.20.9 A medium to large fern with medium-long-creeping rhi-
Tender zomes and elongate blade apices bearing buds at their tips.
A small fern with medium- to long-creeping rhizomes. Can grow submerged in an aquarium or out-of-water in
Grows under medium light on rocks or in soil. This species pots with garden soil or a sand-peat mix in medium light.
can be used as an underwater plant for aquariums, or it can If grown in pots, the soil must be kept wet.
be potted in a peat-sand-soil mix and kept wet. It is rarely The fronds of Bolbitis portoricensis are triangular and
cultivated. one-pinnate, with the lower pinnae longer and lobed on
Bolbitis heudelotii has one-pinnate fronds, usually with the basiscopic side. The veins are netted. The apical por-
ascending pinnae. If the plants are submerged, the sterile tions of well-developed fronds often are elongate (some-
pinna margins are entire, but when held above water, they times whip-like or tailed) and bud-bearing at the tips. The
are irregularly and shallowly serrate. The fertile frond has species is native to tropical America.
narrower pinnae and emerges above the water during non-
flood periods. This species is native to Africa, where it often Bommeria
grows along streambanks prone to flooding. Copper fern
Bommeria is a genus of attractive small ferns that produce
triangular fronds covered with silky white hairs. They are
rarely cultivated and are native to dry, rocky places.
The most distinctive characteristics of this genus are its
small, triangular to pentagonal blades and sori that run

Figure 13.20.9.
Bolbitis heudelotii: Figure 13.20.10. Bolbitis
habit with sterile portoricensis: sterile frond
frond, bar = 5 cm and rhizome, bar = 5 cm
(2 in.). (2 in.).
 226 C 13

along the veins between the midribs and margins, often

mostly near the margins. The blades are lobed, only two- to
three-pinnatifid (never fully pinnate), whitish, hairy, and
slightly scaly. The rhizomes vary from short- to long-creep-
ing and are scaly, not hairy. The veins may be free or partly
netted, and indusia are absent.
Bommeria contains only four species, and these are na-
tive to the southwestern United States, Mexico, and Cen-
tral America. It is named for Joseph Edouard Bommer
(1829–1895), a Belgian pteridologist.

Bommeria hispida (Mettenius ex Kuhn) Underwood

 13.21.1
Semi-hardy, Zone 8
A small fern with medium-creeping rhizomes. Grows
under medium light in moist-dry, well-drained potting mix
with coarse sand. The plants are somewhat difficult to
grow.
The fronds of Bommeria hispida range from 4 to 15 cm
(1.5–6 in.) long and have pentagonal blades with surfaces
covered with long, white hairs. The veins are free, not net-
ted. It is native to the United States, Mexico, and Nicaragua.

Botrychium  13.22.1, 2

Grape fern
These very small- to medium-sized terrestrial ferns are Figure 13.22.1. Botrychium virginianum: frond.
named for the resemblance of their sporangia to clusters of
grapes. In addition to the allusion in the common name,
the genus name comes from the Greek botrys, meaning
grape ferns are used mostly in natural plantings, although
bunch or cluster, as in grapes. In the eastern United States
the medium-sized species are used as potted plants in other
parts of the world. Grape ferns are hard to establish in gar-
dens and are difficult if not impossible to grow from spores.
A well-drained loam mixed with compost is best for cul-
Figure 13.21.1. Bommeria tivating Botrychium species. Water about once a week, but
hispida: a. habit, bar = 5 cm avoid overwatering, which can promote root-rot. On the
(2 in.); b. underside of fertile other hand, it is important to not let the roots dry out when
pinnule, bar = 5 mm (0.2 in.). the plants are in their deciduous period. Guard against slugs
After Knobloch and Correll and snails, which readily attack the newly emerging fronds.
(1962). Grape ferns also should be protected from wind. Trans-
plant them with extra care, disturbing the roots as little as
possible. Because each plant produces only one frond per
year, the loss of a leaf is a potentially serious setback.
The most distinctive characteristic of the genus is its
fronds, which are divided into two parts: a sterile part con-
sisting of a green leafy blade, and a fertile part consisting of
a long stalk bearing a branched cluster of globose sporangia
at its apex. All members of Botrychium are terrestrial and
have fleshy roots and short, erect, fleshy rhizomes. The
genus, which consists of about 55 species, is cosmopolitan.
a

b
 F  F A  C 227

Botrychium dissectum Sprengel  13.22.4

syn. Botrychium obliquum Muhlenberg
Dissected grape fern
Very hardy, Zone 4
A small fern with erect rhizomes. Grows under medium
light in moist garden soil or potting mix. The blades typi-
cally turn bronze or coppery in late fall or winter, especially
in exposed habitats.
Botrychium dissectum is characterized by having sterile
blades with terminal segments divided all the way to the
tip. The blades differ greatly in the amount of dissection,
even in the same population. Two extreme forms are rec-
ognized: forma dissectum, with blades lacerate throughout
and a highly dissected appearance, and forma obliquum
(Willdenow) Fernald, with blades merely lobed or pinnati-
fid and a less cut appearance. Forma dissectum is not found
in Florida and the Gulf States. The species is native to the
eastern United States and southeastern Canada.

Botrychium virginianum (Linnaeus) Swartz

 13.22.1, 2, 5
Rattlesnake fern
Very hardy, Zone 3
A small-medium fern with erect rhizomes. Grows under
medium light in moist garden soil or potting mix. This spe-
cies is difficult to establish.

Figure 13.22.2. Botrychium virginianum: branches

bearing sporangia.
Figure 13.22.3. Botrychium
biternatum: frond, bar =
5 cm (2 in.).
Botrychium biternatum (Savigny) Underwood
 13.22.3
syn. Botrychium dissectum var. tenuifolium (Underwood)
Farwell
Sparse-lobed grape fern, southern grape fern, winter
grape fern
Hardy, Zone 6
A small fern with erect rhizomes. Grows under medium
light in moist garden soil or potting mix. The sterile blade
stays green throughout the winter and often into spring,
and a new frond develops in early summer. The fertile part
of the frond, which is not always present, withers soon after
the spores are shed.
The terminal segments of the sterile blade are elongate
and nearly parallel-sided. The sterile blades of Botrychium
biternatum remain green throughout the winter, in con- Figure 13.22.4.
trast to B. dissectum, which tends to turn bronze or coppery Botrychium
during the winter. The species is native to the southeast- dissectum:
ern United States. frond, bar =
5 cm (2 in.).
 228 C 13

Camptosorus consists of only two species, one of which

occurs in North America and the other in northeastern
Asia. The genus name comes from the Greek camptos,
curved, and soros, mound or heap, referring to the bent or
crooked sori that distinguish it from Asplenium.

Camptosorus rhizophyllus (Linnaeus) Link

 13.23.1
syn. Asplenium rhizophyllum Linnaeus
North American walking fern
Very hardy, Zone 3(4)
A small fern with erect rhizomes and leaves that bear buds
at the apices. Grows under low to medium light in moist,
basic potting mix. In arid climates, grow in a terrarium.
The fronds of Camptosorus rhizophyllus are about 10–20
cm (4–8 in.) long, long-tapered at the tip, and cordate or
hastate at the base. This evergreen fern is native to eastern
Figure 13.22.5. Botrychium North America, where it reaches its greatest frequency and
virginianum: habit, bar = 5 cm abundance on limestone. It grows less frequently on sand-
(2 in.). stone.

Camptosorus sibericus Ruprecht  13.23.2

syn. Asplenium ruprechtii Sa. Kurata
Asian walking fern
Very hardy, Zone 4
Botrychium virginianum is characterized by finely di- A small fern with erect rhizomes and long-tapered leaves
vided, green, triangular blades that are deciduous in late bearing buds at the tips. Grows under low to medium light
summer, not evergreen throughout the winter as in the pre- in moist, basic potting mix. In arid climates, grow in a
vious two species. The fertile spike attaches just below the terrarium.
green blades, but it is not always fully developed and some-
times is represented only by an aborted stub. The spike
withers in midsummer as soon as the spores are shed. The
species is common in the forests of North America. It also
occurs in Central America, South America, Europe, and
Asia.

Camptosorus
syn. Asplenium, in part
Walking fern
The walking ferns are small, lime-loving terrestrials used in
rock gardens or terrariums. The common name derives
from the plant’s ability to form plantlets at the tips of its
long-tapered fronds, making the plant appear to “walk”
over the rock surface. Although it can be grown in warmer
climates outdoors or in terrariums, Camptosorus must have
humidity and drainage. The plants grow best in temperate
climates.
The genus is characterized by netted veins and fronds
that taper to a long, whip-like apex, which roots at the tip. a
b
The rhizome is short and erect. The sori are elongate and
scattered (not regularly arranged as in the closely related
genus Asplenium) and covered by a flap-like indusium. This Figure 13.23.1. Camptosorus rhizophyllus: a. habit, bar =
genus is often classified in Asplenium because it hybridizes 5 cm (2 in.); b. sori and indusia, bar = 5 mm (0.2 in.),
with several species of that genus. courtesy of T. Hoshizaki..
 F  F A  C 229

that form between the primary lateral veins. Campyloneu-

rum contains about 50 species restricted to the American
tropics. Most of the species are epiphytes. Lellinger (1988)
gives a provisional key to the species. See Polypodium for a
comparison of this genus to other polypodiaceous genera.

Campyloneurum angustifolium (Swartz) Fée

 13.24.1
Narrow-leaved strap fern
Semi-tender, Zone 9
A medium-sized fern with short-creeping rhizomes and
arched or pendent fronds borne in a cluster. Thrives under
high-medium light in drained, moist to moist-dry potting
mix.
Campyloneurum angustifolium is characterized by linear
a blades only 4–12 mm (0.2–0.5 in.) wide, with stipes short
or absent. The veins are hard to see because of the thick-tex-
b tured blade, but there are less than four areoles between
midrib and margin, with the costal areoles more elongate
Figure 13.23.2. Camptosorus sibericus: a. habit, bar = than the marginal areoles. The sori are borne in one or two
5 cm (2 in.); b. sori and indusia, bar = 5 mm (0.2 in.), (rarely three) rows between midrib and margin. The species
courtesy of T. Hoshizaki. is widely distributed throughout tropical and subtropical
America, often growing as an epiphyte on tree trunks.
Campyloneurum angustifolium varies greatly, and several
unnamed variants are in cultivation. Some forms have con-
sistently narrow fronds only 4 mm (0.2 in.) wide. Wild
Camptosorus sibericus has stouter and smaller evergreen
plants with twisted fronds and undulate margins have also
fronds than C. rhizophyllus. They are usually up to about 5
been brought into cultivation from Costa Rica.
cm (2 in.) long and often constricted above the tapered
base. The species is native to Siberia, Japan, Korea, and
northern China.
Asplenium ×crucibuli is a hybrid produced in horticul-
ture from a cross between this species and A. platyneuron.

Campyloneurum
Strap fern
The more frequently cultivated strap ferns, such as Campy-
loneurum angustifolium and C. phyllitidis, are handsomely
displayed in hanging containers, where their dense cluster
of narrow fronds form a spreading or gracefully arching
pattern. The frond width can vary greatly within a species.
Nearly all species of Campyloneurum have simple, en-
tire, strap-shaped leaves. Their blades are glabrous except
along the rachis, where there may be scattered scales. The
veins are always netted, although this may be difficult to
see in species with thick blades. In many species the main a b
lateral veins are parallel and connected by curved cross-
veins to form areoles, and the areoles enclose veinlets that
point toward the margin. The sori are round, lack indusia,
and in most species are borne in one to four rows between Figure 13.24.1. Campyloneurum angustifolium: a. habits
the main veins. The rhizome varies from short- to long- of two plants, bar = 5 cm (2 in.); b. sori and vein
creeping and may be clothed by clathrate or nonclathrate pattern, bar = 1 cm (0.4 in.).
scales.
The genus name comes from the Greek kampylos,
arched, and neuron, nerve, referring to the arched veinlets
 230 C 13

Campyloneurum brevifolium (Link) Link

 13.24.2
syn. Campyloneurum latum T. Moore c
Tender, Zone 10
A medium-large fern with short-creeping rhizomes and
erect fronds borne in a cluster. Grows under medium light
in moist, well-drained potting mix.
Campyloneurum brevifolium resembles C. phyllitidis but
differs by often having three (versus two) veinlets within
each areole. The stipes are 5–18 cm (2–7 in.) long, and the
blades are mostly 5–13 cm (2–5 in.) wide, dark green, and
abruptly contracted to the stipe. Sori are borne in two or
more rows between the lateral veins, with up to 14 sori in a
row between midrib and margin. The species is widespread
in tropical America.

Campyloneurum costatum (Kunze) C. Presl

b
 13.24.3
Tender, Zone 10
A medium-sized fern with short-creeping rhizomes and
arched or pendent fronds borne in a cluster. Grows under
medium light in well-drained, moist potting mix. a
The stipes of Campyloneurum costatum are 4–14 cm
(1.5–5.5 in.) long, and the blades are elliptic or oblong,
Figure 13.24.3. Campyloneurum costatum: a. habit, bar =
5 cm (2 in.); b. frond, bar = 5 cm (2 in.); c. sori and vein
pattern, bar = 1 cm (0.4 in.).

mostly 5 cm (2 in.) wide with acute apices. The main lateral

veins are easily visible but the smaller, secondary ones are
obscure. There are four to eight areoles between midrib and
b
margin. The sori are in two rows between lateral veins, with
four to eight sori in a row between midrib and margin. The
species is widespread in tropical and subtropical America.

Campyloneurum phyllitidis (Linnaeus) C. Presl

 13.24.4
Semi-tender, Zone 9
A medium-sized fern with short-creeping rhizomes and
c spreading to slightly arching fronds borne in a cluster.
Grows under medium light in well-drained, moist to
moist-dry potting mix. This species is the most common
Campyloneurum in cultivation. It needs protection from
slugs and snails.
a The stipes of Campyloneurum phyllitidis are absent or
up to 5 cm (2 in.) long. The yellowish green blades are 4–8
cm (1.5–3 in.) wide, gradually tapering toward the base or
a short stipe, with acute to obtuse apices. The main lateral
veins are clearly visible, but the smaller secondary ones that
Figure 13.24.2. Campyloneurum brevifolium: a. frond, form areoles are hard to see. Two sori are usually borne be-
bar = 5 cm (2 in.); b. sori distribution, bar = 1 cm tween the main lateral veins, with 5 to 14 sori in series be-
(0.4 in.), after Hoshizaki (1982); c. sori and vein tween the midrib and margin. This species is widespread in
pattern, bar = 1 cm (0.4 in.), after Hoshizaki (1982). tropical and subtropical America.
 F  F A  C 231

Figure 13.24.5. Campyloneurum tenuipes: a. habit and

c frond, bar = 5 cm (2 in.); b. frond apex with sori and
vein pattern, bar = 1 cm (0.4 in.).

Figure 13.24.4. Campyloneurum

phyllitidis: a. frond, bar = 5 cm (2 in.); Campyloneurum xalapense Fée  13.24.6
b. sori distribution, bar = 1 cm (0.4 in.), Tender to semi-tender
after Hoshizaki (1982); c. sori and vein A medium-sized fern with short-creeping rhizomes and
pattern, bar = 1 cm (0.4 in.), after fronds borne in a cluster. Grows under medium light in
Hoshizaki (1982). moist potting mix.
Campyloneurum xalapense is characterized by short or
absent stipes, oblanceolate blades, acuminate to abruptly
The broader fronds of Campyloneurum phyllitidis are
often mistaken for C. brevifolium. The short or absent stipe
and the more regularly shaped areoles of C. phyllitidis
help distinguish it from C. brevifolium, which has a well-
defined stipe and irregularly shaped areoles with three in-
cluded veinlets. Some plants, however, are difficult to dis-
tinguish. Crested and fimbriate forms of C. phyllitidis occur
in cultivation.

Campyloneurum tenuipes Maxon  13.24.5

Tender to semi-tender
A medium-sized fern with short-creeping rhizomes and
erect to spreading fronds borne in a cluster. Grows under
medium light in moist potting mix with good drainage.
The fronds of Campyloneurum tenuipes reach 30–80 cm
a
(12–32 in.) in length, with stipes about one-third the length
of the frond. The blade is narrowly elliptic with a tapered b
base and abruptly narrowed to a tail-like apex. The main
lateral veins are visible, but the finer veins are usually hard
to see. The sori are borne in two rows between the main lat- Figure 13.24.6. Campyloneurum xalapense: a. habit, bar
eral veins, in series of 6–10 between the midrib and margin. = 5 cm (2 in.); b. white dots on upper blade surface, bar
The species occurs from southern Mexico to Honduras. = 1 cm (0.4 in.).
 232 C 13

caudate apices, and white dots on the upper blade surfaces. to go from spores to mature, spore-bearing plants, though
Veins are less obscure on the lower surfaces. The areoles are three or four months is more typical. Because of this quick
irregularly shaped, with generally five to eight between the cycle, the plants are favorites of laboratory geneticists who
costa and margin, and the sori are in two rows between the study differences in heritable traits from one generation to
main lateral veins. The species occurs from southern Mex- the next.
ico to Costa Rica. The plants live about one year and are usually perpetu-
ated by the numerous buds that form on the blade surface,
Ceratopteris  13.25.1 typically in the margins of the sinuses. The buds detach
Water fern, water sprite and float away from the parent plant. Mature plants can be
Ceratopteris is commonly used as an aquarium plant. These propagated by division of the rhizome, which may be al-
ferns either float on water, grow fully or partially sub- lowed to float or, if grown submerged, held in place by
merged, or root in mud. The fertile leaves are erect and are coarse gravel. If plants are to be rooted in mud, garden soil
produced above water. In Asia the foliage of Ceratopteris with little organic matter will do. Some growers recom-
thalictroides is used in salads and is said to have a peppery mend a mixture of about half peat and half sand mixed with
taste. 10% top soil. Plants that are rooted in mud need their
Ceratopteris is characterized by short, erect, inconspicu- fronds kept moist. Temperatures must be maintained close
ous rhizomes bearing a few thin scales. The sterile and fer- to 27°C (80°F) for good growth. The plants and buds de-
tile fronds are dimorphic—the fertile ones are more finely cline when temperatures are below 20°C (68°F), and if lost,
cut with narrower segments—but both have triangular, new plants must be started from spores.
glabrous blades. The veins are netted, not free. The large, The genus, which consists of four species, is native to
marginal sporangia are protected by a continuous enrolled the tropics and subtropics worldwide. The name comes
indusium. from the Greek keras, horn, and pteris, fern, alluding to the
This genus has the distinction of having the fastest life antler-like fertile leaves. The species are sometimes difficult
cycle of any fern. The plants can take as little as one month to identify because the fronds vary considerably in form.
Only Ceratopteris thalictroides and C. pteridoides are com-
monly cultivated. For more details on cultivation, see Curt-
wright (1995) and Lloyd (1974).

Ceratopteris pteridoides (Hooker) Hieronymus

 13.25.2, 3
American water fern, triangular water fern
Tender, Zone 9
A small to medium fern with short, inconspicuous rhi-
zomes. Grows under high light in wet garden soil or float-
ing on water. In nature the plants float; rarely do they root
in mud. This species reproduces readily from buds on the
blades.
The best distinguishing characteristics of Ceratopteris
pteridoides are the inflated stipe bases and the opposite pin-
nae or lobes of the sterile blade. The inflated stipe base is
composed of aerenchyma, a type of tissue related to paren-
chyma that contains large intercellular air spaces that im-
part buoyancy to the plant. The buds generally develop into
plantlets on the actively growing fronds, whereas in C. thal-
ictroides the buds generally develop on older, senescing
fronds. The species is native to the southern United States
(Florida and Louisiana), tropical America, and southeastern
Asia.
Ceratopteris cornuta Le Prieur, an Old World species,
might be sold in aquarium stores. It differs from C. pteri-
doides by lacking an inflated stipe and by having mostly al-
ternate basal pinnae and triangular, one- to two-pinnate
blades with many buds.

Figure 13.25.1. Ceratopteris habit.

 F  F A  C 233

Figure 13.25.4. Ceratopteris thalictroides: habit, with a

Figure 13.25.2. Ceratopteris pteridoides: sori and indusia. more divided pinna in the upper right, bar = 5 cm
(2 in.). After Beddome (1864).

late to lanceolate-ovate and two- to three-pinnate, with

buds that tend to grow on dying fronds. The fertile fronds
are usually two- to three-pinnate with narrow, linear seg-
ments. This species is found throughout the tropics of the
world, except Africa.
Ceratopteris richardii Brongniart (syn. C. deltoidea Bene-
dict), a species of the southern United States, tropical
America, and Africa, is possibly in cultivation. Some au-
thors consider it the same as C. thalictroides, from which it
is sometimes difficult to distinguish. Ceratopteris richardii is
said to have larger sterile fronds with more buds, and fertile
fronds to three-pinnate. The only sure way to distinguish
the species is by the number of spores per sporangium: C.
richardii has 16 spores, whereas C. thalictroides has 32.

Ceterach  13.26.1, 2

Rusty-back fern
Figure 13.25.3. Ceratopteris pteridoides: habit, bar = 5 cm
Ceterach is a genus of small, terrestrial ferns with numerous
(2 in.). After Tryon and Stolze (1989).
overlapping scales on the undersides of the fronds. It is used
mostly in rock gardens in temperate climates (see “Xero-
phytic Ferns” in Chapter 10). During dry weather, the
Ceratopteris thalictroides (Linnaeus) Brongniart fronds enroll, exposing their scaly lower surfaces. During
 13.25.4;  19 wet weather, they expand quickly. The scales on the lower
Oriental water fern surface of the blade absorb water to rehydrate the frond.
Tender, Zone 9 Ceterach is characterized by pinnatifid or one-pinnate
A small to medium fern with short, inconspicuous rhi- blades that are densely scaly beneath, obtuse segment tips,
zomes. Grows under high light in wet garden soil or float- and elongate sori with inconspicuous indusia. The scales
ing on water. Typically, the plants root in mud. are broad and clathrate. The veins join near the margin of
Unlike the preceding species, Ceratopteris thalictroides the frond, and the sori are often obscured by the scales and
has narrow (not inflated) stipe bases and mostly alternate therefore are difficult to see. The rhizomes of all species are
basal pinnae or lobes. The sterile fronds are mostly lanceo- short and erect. Many pteridologists classify Ceterach with
 234 C 13

Figure 13.26.3.
Ceterach aureum:
fronds, bar =
5 cm (2 in.).

Figure 13.26.1. Ceterach officinarum: habit.

The species is native to the Madeira and Canary Islands.

Some pteridologists considered this species a variety of C.
officinarum.

Ceterach officinarum de Candolle

 13.26.1, 2, 4;  20
syn. Asplenium ceterach Linnaeus
Scale fern
Hardy
A small-medium fern with erect rhizomes and clustered
fronds. Grows under medium light in moist-dry, basic pot-
Figure 13.26.2. Ceterach officinarum: fertile pinnae. ting mix. This stout fern is excellent for rock gardens. In na-
ture it grows on calcareous rocks and mortared walls.

Asplenium because it hybridizes with that genus. The two

genera are distinguished in this book for convenience.
Only three species occur in Ceterach, and all are native
to the Old World. The genus name comes from the Greek
sjetrak, an old name applied to the plant by Persian physi-
cians.

Ceterach aureum (Cavanilles) L. V. Busch

 13.26.3
Rusty-back fern
Hardy
A small-medium fern with erect rhizomes and clustered
fronds. Grows under medium light in moist-dry potting
mix. This species is excellent for rock gardens. Figure 13.26.4.
Ceterach aureum resembles C. officinarum but has larger Ceterach officinarum:
and broader fronds. The blades are generally two times fronds, bar = 5 cm
longer than broad, and the scales beneath are toothed. (2 in.).
 F  F A  C 235

The fronds of Ceterach officinarum are up to four times

longer than broad, less than 30 cm (12 in.) long, and the
blade scales are entire, not toothed. The species is native
from the United Kingdom to the Pyrenees.

Cheilanthes  13.27.1–3

Lip fern
Cheilanthes is a genus of small, evergreen, terrestrial or rock-
inhabiting ferns of dry climates. The plants often have
woolly or scaly frond undersides. A number of species exist
in special collections, and only a few are in general use in
rock gardens and trough gardens. Although attractive for
their small size and (often) fine texture, lip ferns can be dif-
ficult to grow. Care is needed to provide the proper amount
of water, humidity, and drainage through the seasons. The
plants prefer a medium with about 50% coarse sand or per-
lite. The soil should be kept moderately moist during peri-
ods of active growth but drier during inactivity. Avoid wet-
ting the foliage but maintain moderate humidity. Wide,
frequent fluctuations in moisture level seem harmful to lip Figure 13.27.2. Cheilanthes gracillima: leaflets with
ferns. Fluctuations can be reduced by locating plants next indusia.
to large rocks or by planting them in clay pots that are in
turn planted into larger pots (double potting) or directly
in the ground. Excessive shade can cause the fronds to be- Cheilanthes is large, variable, and hard to characterize.
come weak and abnormally elongated; sites located to re- All species have free veins, and the sori are typically found
ceive direct sun in the early morning and late afternoon can along the margins and protected by a false indusium. The
help prevent this. Some species grow well in full sun. For rhizomes vary from compact and erect to decumbent or
more details on the culture of Cheilanthes species, see the long-creeping. The fronds have one- to five-pinnate blades
section on “Xerophytic Ferns” in Chapter 10.

Figure 13.27.3. Cheilanthes lanosa: sori with indusia

Figure 13.27.1. Cheilanthes gracillima: habit. pushed back.
236 C 13

that may be glabrous, glandular, hairy, scaly, or covered Cheilanthes: Upper surface of the blade with-
with a colored powder on the lower surface. Some species out stellate scales, not striate; lower surface
have numerous bead-like segments that are circular or of the blade not powdery; margin of the fer-
nearly so and often cupped, with the open end of the cup tile segments slightly enrolled to cover the
facing downward. sporangia; ultimate segments entire, stalked
Cheilanthes contains about 180 species and is wide- or sessile. (Cheilanthes alabamensis, C. argen-
spread in the temperate and tropical regions of the world. tea, C. arizonica, C. bonariensis, C. buchtienii,
The genus name comes from the Greek cheilos, lip, and an- C. covillei, C. eatonii, C. farinosa, C. feei, C.
thos, flower, alluding to the position of the sporangia be- fendleri, C. gracillima, C. hirta, C. lanosa, C.
neath the lip-like false indusium. lendigera, C. lindheimeri, C. newberryi, C. tomentosa, C. vil-
Some species of Notholaena and Pellaea are difficult to losa, C. wrightii)
distinguish from Cheilanthes. In view of this problem,
Notholaena has been redefined to include only those species Mildella: Upper surface
with a dense white or yellow waxy powder on the lower sur- of the blade without stel-
faces of the blades and with sporangia at the tips of the veins late scales, not striate;
in a more or less continuous marginal band covered by an lower surface of the blade
enrolled but texturally unmodified frond margin. This def- not powdery; margin of
inition, accepted here, places all the cultivated species for- the fertile segments en-
merly listed as Notholaena into Cheilanthes. Many Cheilan- rolled to cover the spo-
thes species go in and out of cultivation rather frequently, rangia; ultimate segments
and therefore all could not be treated here. The reader is entire, sessile, adnate; in-
referred to floras, particularly of drier regions, for more dusium linear, attached
information. below the margin of the segment and thus leaving a nar-
Pteridologists now distinguish several genera formerly row band of blade tissue between the edge and the indu-
included in Cheilanthes. To help with identification, these sium. (Mildella intramarginalis)
genera are described here.
Notholaena: Upper surface
Argyrochosma: Upper surface of the blade of the blade without stel-
without stellate scales, not striate; lower sur- late scales, not striate;
face of the blade usually has a yellowish or lower surface of the blade
whitish powder; margin of the fertile seg- often with a yellowish or
ments enrolled to cover the sporangia or not; whitish powder; margin of
ultimate segments entire and stalked. (Argy- the fertile segments en-
rochosma dealbata, A. jonesii, A. microphylla) rolled to cover the sporan-
gia; ultimate segments en-
Aspidotis: Upper tire, sessile to subsessile.
surface of the (Notholaena standleyi)
blade without stel-
late scales, striate;
lower surface of the blade not Cheilanthes alabamensis (Buckley) Kunze
powdery; margin of the fertile  13.27.4
segments en- Alabama lip fern
rolled to cover Hardy, Zone 7
the sporangia, A small to medium fern with short-creeping to compact
white-papery; ul- rhizomes and clustered fronds. Grows best under medium-
timate segments entire and short-stalked. high light in moist-dry to dry, basic garden soil and sand.
(Aspidotis californica, A. carlotta-halliae, A. Cheilanthes alabamensis has black stipes, rachises, and
densa) costae, with the color entering the pinnules. The hairs on
the upper rachis surface are dense, short, and twisted; those
Astrolepis: Upper surface of the blade with beneath are long and sparse or absent. The blade is lanceo-
stellate scales, not striate; lower surface of late to linear-oblong, two-pinnate to two-pinnate-pinnati-
the blade not powdery; margin of the fertile fid, with segments elliptic to elongate-triangular, not bead-
segments flat, not enrolled to cover the spo- like. The blade surface varies from hairy to nearly glabrous.
rangia; pinnae stalked to subsessile. (As- The species is native to the United States and Mexico,
trolepis beitelii, A. sinuata) where it typically grows on limestone.
 F  F A  C 237

Figure 13.27.4. Cheilanthes

alabamensis: a. frond, bar = 2.5 cm
(1 in.); b. lower surface of fertile a
pinna, bar = 1 cm (0.4 in.).

Figure 13.27.5.
b Cheilanthes argentea:
a. frond, bar = 2.5 cm
(1 in.); b. lower surface of
fertile pinna, bar = 2 cm
a (0.8 in.).

Cheilanthes argentea (J. F. Gmelin) Kunze

 13.27.5;  21
syn. Aleuritopteris argentea (J. F. Gmelin) Fée
Hardy, Zone 5
A small fern with compact rhizomes and clustered fronds. b
Grows under medium to high light in moist to moist-dry,
well-drained garden soil with sand. The plants are easy to
grow from spores. One variant in cultivation has white
lower blade surfaces.
Cheilanthes argentea is distinctive by its pentagonal, one-
pinnate-pinnatifid to two-pinnatifid blades covered by a
yellow or white powder on the lower surfaces. The basal
pinnae are asymmetrical, being larger on the basiscopic c
side, and the stipes are dark purplish and shiny. The spo-
rangia are covered by a continuous, entire indusium. The
species is native from northern Asia to Malaya and India.

Cheilanthes arizonica (Maxon) Mickel

 13.27.6
syn. Cheilanthes pyramidalis subsp. arizonica Maxon
Arizona lip fern
Semi-hardy, Zone 8
A small fern with short-creeping rhizomes and clustered
a
fronds. Grows under high light in well-drained, moist-dry
to dry garden soil and sand.
Cheilanthes arizonica is distinguished by elongate-pen-
tagonal blades up to four-pinnate, basal pinnae conspicu- Figure 13.27.6. Cheilanthes arizonica: a. frond, bar =
ously larger than the above pair, and the ultimate segments 2.5 cm (1 in.); b. underside of sterile segment, note
elliptic to linear-oblong (not bead-like), the largest 4–10 glands, bar = 5 mm (0.2 in.); c. underside of fertile
mm (0.2–0.4 in.) long. The lower surface of the blade bears segment, bar = 5 mm (0.2 in.). After Knobloch and
scattered reddish glands, but the upper surface is glabrous. Correll (1962).
 238 C 13

The species is native to southeastern Arizona, Mexico, Gua-

temala, and Honduras. In nature it occurs on quartzite or
igneous rocks.

Cheilanthes bonariensis (Willdenow) Proctor b

 13.27.7
syn. Notholaena aurea (Poiret) Desvaux
Bonaire lip fern, slender cloak fern
Hardy to semi-hardy, Zone 7
A small-medium fern with short-creeping to compact rhi-
zomes and clustered fronds. Grows under high light in Figure 13.27.8.
moist-dry to dry, well-drained garden soil with sand. Cheilanthes
Cheilanthes bonariensis has narrow, one-pinnate-pin- buchtienii: a. frond,
natifid blades with densely hairy lower surfaces. The pinnae bar = 2.5 cm (1
are numerous (often about 40 pairs) and have four to seven in.); b. lower
pairs of lobes. They may be shed from the rachis (jointed). surface of fertile
The hairs on the lower surface are dense, long, and matted, pinna, bar = 1 cm
varying from white to creamy yellowish in color. The spe- (0.4 in.).
cies is widespread in the American tropics and subtropics.

Cheilanthes buchtienii (Rosenstock) R. M. Tryon

 13.27.8
syn. Notholaena buchtienii Rosenstock
Buchtien’s lip fern
Semi-hardy, to Zone 9 or colder
A small fern with short-creeping rhizomes and clustered a
fronds. Seems to thrive in gardens under high light in
moist-dry, well-drained garden soil with sand.

The stipe of Cheilanthes buchtienii is about half as long

as the blade or shorter. The blade is narrowly linear and
one-pinnate-pinnatifid or two-pinnate at the base. The pin-
nae vary from triangular to triangular-lanceolate and are
b densely hairy beneath with matted, pale brown hairs. The
species is native to Bolivia and Argentina.

Cheilanthes covillei Maxon  13.27.9

Figure 13.27.7.
Coville’s lip fern
Cheilanthes bonariensis:
Semi-hardy, Zone 7
a. frond, bar = 2.5 cm
(1 in.); b. upper surface A small fern with compact, short-creeping rhizomes and
of pinna, bar = 5 mm clustered fronds. Grows under high light in well-drained
(0.2 in.). garden soil kept moist-dry to dry.
Cheilanthes covillei has dark brown stipes, and the lan-
ceolate to ovate-triangular blades are three- to four-pinnate
with bead-like segments. Both surfaces of the blade are
glabrous, or a few small scales may be near the base on the
underside. The rachises, costae, and costules are densely
scaly, the scales broadly ovate and ciliate toward the base,
darker in the center, and nearly covering the segments on
a the undersides. This species resembles C. fendleri, but its
rhizome scales are strongly appressed, instead of loosely
overlapping, and the costal scales are entire to erose. It is na-
 F  F A  C 239

Cheilanthes farinosa (Forsskål) Kaulfuss

 13.27.11
c Figure 13.27.9. Cheilanthes syn. Aleuritopteris farinosa (Forsskål) Fée
covillei: a. frond, bar = 2.5 cm Semi-tender (?)
(1 in.); b. upper surface of
segments, bar = 1 mm, after A small to medium fern with compact rhizomes and clus-
Hickman (1993); c. scale on tered fronds. Grows under high light in drained, moist-dry
lower side of segments, bar = garden soil or potting mix.
1 mm, after Hickman (1993). Cheilanthes farinosa is characterized by triangular to nar-
rowly triangular blades that are two-pinnate and have a
b white or yellow powder on the lower surface. The stipe is
a dark and shiny. The basal pinnae are enlarged on their
lower side, and the segments are not bead-like. This species
tive to the southwestern United States and Mexico (Baja is native to the tropics of the world.
California), where it grows on rocky cliffs and ledges, usu-
ally on igneous rocks.

Cheilanthes eatonii Baker in Hooker & Baker

 13.27.10
Eaton’s lip fern a
Hardy, Zone 6
A small fern with compact rhizomes and clustered fronds.
Thrives under high light in moist-dry to dry, well-drained
garden soil with sand. The plants are relatively easy to grow.
Cheilanthes eatonii has dark brown stipes. The blades
are oblong-lanceolate, three- to four-pinnate, and hairy on
both surfaces (more so beneath). The segments are oval to
round and bead-like, and the rachis has scattered linear b
scales and is hairy. This species resembles C. tomentosa. It is
native to the southern United States, Mexico, and Costa
Rica, usually growing on rock slopes and ledges.

Figure 13.27.10. Figure 13.27.11.

Cheilanthes eatonii: Cheilanthes farinosa:
a. habit, bar = 2.5 cm a. frond, bar = 2.5
(1 in.); b. lower cm (1 in.); b. lower
surface of fertile surface of fertile
pinnule, bar = 5 mm pinna, bar = 5 mm
(0.2 in.). After (0.2 in.), after
Knobloch and Correll Knobloch and
(1962). Correll (1962).

Cheilanthes feei T. Moore  13.27.12

Slender lip fern
Very hardy, Zone 4
A small fern with compact to short-creeping rhizomes and
small, rosette-like clusters of blue-green fronds. Grows
under high light in moist-dry to dry, basic, well-drained
garden soil with sand.
The most distinctive characteristic of Cheilanthes feei is
its small, blue-green, finely divided blades with numerous
b a bead-like segments. The stipe bears multicellular (septate)
hairs. The lower surface of the segment is dense with long,
 240 C 13

b
Figure 13.27.12.
Cheilanthes feei: a. frond,
bar = 2.5 cm (1 in.);
b. upper surface of
Figure 13.27.13.
pinnule, bar = 5 mm
Cheilanthes fendleri: a.
(0.2 in.), after Knobloch
habit, bar = 2.5 cm (1 in.);
and Correll (1962);
b. lower surface of fertile
c. lower surface of pinnule,
pinnule, bar = 2 mm, after
bar = 5 mm (0.2 in.), after
Knobloch and Correll
a Knobloch and Correll
(1962).
(1962).
c

lax hairs; the upper surface is sparsely hairy to glabrous.

This species is native to North America (mostly the central
United States), where it grows on limestone cliffs.

Cheilanthes fendleri Hooker

 13.27.13;  22
Fendler’s lip fern
Hardy, Zone 5 b
A small fern with evergreen fronds scattered along a long-
creeping rhizome. Grows under high light in well-drained,
moist-dry garden soil. The plants are relatively easy to grow.
Cheilanthes fendleri is distinctive by the combination of
long-creeping rhizomes, dark brown stipes, and bead-like
segments. The blades are lanceolate to ovate-triangular, a
three- to four-pinnate, and glabrous on both sides or with
a few scales near the base on the lower surface. The primary
and secondary costae, however, are densely scaly, the scales
nearly covering the segments. This species resembles C. cov- ments, which are oblong, or rarely oval, and bead-like. The
illei, which differs by having rhizome scales strongly ap- lower surfaces of the blades are densely covered with
pressed rather than loosely so, and costal scales ciliate near branched hairs and small ciliate scales. The margins are
the base rather than entire to erose. Cheilanthes fendleri is hardly or not modified into a false indusium. This species
native to the southwestern United States and Mexico. is native to the northwestern and central United States.
This species is sometimes misnamed in the trade as
Cheilanthes lanosa or C. tomentosa.

Cheilanthes gracillima D. C. Eaton a

 13.27.1, 2, 14
Lace fern
Hardy, Zone 5 b
A small fern with short-creeping rhizomes and clustered,
dark green fronds. Grows under high light in drained, Figure 13.27.14. Cheilanthes
moist-dry garden soil with sand. This is a compact, ever- gracillima: a. habit, bar =
green fern. 2.5 cm (1 in.); b. lower
Cheilanthes gracillima has dark brown stipes, and the surface of fertile pinnule,
blades are linear-oblong, two- to three-pinnate, 1–2.5 cm bar = 5 mm (0.2 in.).
(0.5–1 in.) wide, and densely woolly beneath. The rachis
bears scattered scales but no hairs. The scales along the costa
of the pinnae are long-ciliate and do not conceal the seg-
 F  F A  C 241

Cheilanthes hirta Swartz  13.27.15

Parsley fern
Semi-hardy to hardier (?) b
A small to medium fern with short-creeping rhizomes bear-
ing fronds in a loose cluster. Best grown under medium-
high light in well-drained, moist to moist-dry garden soil or
potting mix. This species is relatively easy to grow.
The short, dark brown stipes of Cheilanthes hirta are
covered with reddish brown, long, stiff, hair-like scales. The a
blades vary from linear-elliptic to narrowly oblanceolate
and are two- to three-pinnate-pinnatifid. The segments
have rounded lobes (not bead-like) and scattered, long,
c
gland-tipped hairs on both surfaces. The costa is dark
brown with reddish brown hairs. The marginal lobes are
curved over discrete sori, and the false indusium is minute
or absent. The species is native to southern Africa.

Cheilanthes lanosa (Michaux) D. C. Eaton in Emory

 13.27.3, 16
syn. Notholaena vestita Desvaux
Hairy lip fern
Hardy, Zone 5
A small to medium fern with short-creeping rhizomes bear- Figure 13.27.16. Cheilanthes lanosa: a. frond, bar =
ing clustered, grayish fronds. Grows under medium-high 2.5 cm (1 in.); b. lower surface of fertile pinnule, bar =
light in well-drained, acidic garden soil or potting mix kept 5 mm (0.2 in.); c. upper surface of pinna, bar = 1 cm
(0.4 in.).

moist-dry to dry. Cheilanthes lanosa is easy to cultivate.

Some of the plants sold by this name in the trade are C.
a fendleri. It usually grows on the ground in or among sand-
stone; it does not grow on vertical cliff faces.
Cheilanthes lanosa has narrow-triangular to lanceolate,
b two-pinnate-pinnatifid blades that are sparsely hairy on
both surfaces. The stipe, rachis, and costae are purple-
brown and have long hairs, but no scales, and the ultimate
segments are oblong to lanceolate (not bead-like) and pin-
nately lobed. The species is native to the eastern United
States.

Cheilanthes lendigera (Cavanilles) Swartz

 13.27.17
Beaded lip fern
Semi-hardy or hardier, Zone 8
A small fern with fronds scattered or clustered along a long-
creeping rhizome. Grows well under high light in drained,
moist-dry to dry garden soil or potting mix. The plants are
relatively easy to grow.
Cheilanthes lendigera has dark brown stipes with long,
soft, whitish hairs. The blades are triangular-lanceolate, to
three-pinnate, with the costae nearly scaleless beneath. The
segments are small and bead-like, with the upper surfaces
Figure 13.27.15. Cheilanthes hirta: a. frond, bar = smooth and the lower surfaces covered with long, tawny
2.5 cm (1 in.); b. pinna, bar = 2 cm (0.8 in.). hairs. The sporangia are covered by a prominent, broad,
 242 C 13

b
Figure 13.27.18.
Cheilanthes
lindheimeri: a.
habit, bar =
2.5 cm (1 in.);
b. upper surface
of pinnule, bar =
2 mm, after
Knobloch and
Correll (1962);
c. lower surface
Figure 13.27.17. of pinnule and
Cheilanthes lendigera: a enlargement of
a. frond, bar = 2.5 cm scale, bar =
(1 in.); b. lower surface 2 mm, after
of fertile pinnule, Knobloch and
a Correll (1962).
showing pouch-like
indusia, bar = 2.5 mm c
(0.1 in.).

translucent false indusium that nearly covers the underside

of the segment. The species occurs from the southwestern
United States to South America and the West Indies.
The fronds of Cheilanthes newberryi are a distinctive cot-
Cheilanthes lindheimeri Hooker  13.27.18 tony, whitish gray. The stipes are wiry, and the blades are
Lindheimer’s lip fern linear to ovate-lanceolate, two- to four-pinnate, and 1–5
Semi-hardy, Zone 7 cm (0.5–2 in.) wide. The rachis is hairy on both surfaces,
A small fern with fronds scattered or clustered on a long- and the segments are round and flat, only somewhat bead-
creeping rhizome. Grows well under high light in drained, like, up to 3–5 mm wide, with the lower surface conspicu-
moist-dry to dry garden soil. This species is relatively easy to ously woolly with fine, cobwebby hairs (some of these also
grow. occur on the upper surface). The species is native to the
The blades of Cheilanthes lindheimeri are oblong-lance-
olate to ovate-triangular, to four-pinnate, and 2–5 cm
(0.75–2 in.) wide. The rachis has some scales and hairs, and
the costal scales are lanceolate with truncate or subcordate
bases. The margins have fine, curly hairs that form a tangled
b
mass partly concealing the segments. The minute, bead-
like segments are nearly smooth on both surfaces but ap-
pear hairy due to the hairy costal scales. The species is native
to the southwestern United States and Mexico.

Cheilanthes newberryi (D. C. Eaton) Domin

 13.27.19 Figure 13.27.19. Cheilanthes
syn. Notholaena newberryi D. C. Eaton a newberryi: a. frond, bar = 2.5 cm
Newberry’s lip fern, cotton fern (1 in.); b. pinna, bar = 1 cm
Semi-hardy to hardy, Zone 7 (0.4 in.).
A small fern with short-creeping to compact rhizomes and
clustered fronds. Grows under high light in drained, moist
to moist-dry garden soil with sand.
 F  F A  C 243

United States (southern California) and Mexico (Baja Cal-

ifornia) and usually grows on igneous rocks.

Cheilanthes tomentosa Link  13.27.20

Woolly lip fern
b
Hardy, Zone 6
A small-medium fern with compact rhizomes and clustered
fronds. Grows well under high light in well-drained, moist-
dry to dry garden soil with sand. The plants usually bear
attractive gray-green fronds. This species is relatively easy to
grow.
Cheilanthes tomentosa has dark stipes and oblong-lance-
olate, four-pinnate blades, 1.5–8 cm (0.6–3 in.) wide. The
rachis bears scattered inconspicuous linear scales and hairs,
and the scales on the costae are narrow, not concealing the
segments. The ultimate segments are oval, rarely oblong,
bead-like, up to 1–2 mm wide, with the lower surface
densely hairy with matted hairs and the upper surface with c
fine hairs. The species resembles C. eatonii, which differs by
having more prominent, wider scales on the rachis and
costae. Cheilanthes tomentosa is native to the southwestern
United States and Mexico, where it grows on a variety of a
rock types.

Cheilanthes villosa Davenport ex Maxon

Figure 13.27.21. Cheilanthes villosa:
 13.27.21
a. frond, bar = 2.5 cm (1 in.);
Villous lip fern
b. upper surface of fertile pinnule,
Semi-hardy
bar = 2 mm, after Knobloch and
A small fern with erect to ascending, compact rhizomes and Correll (1962); c. lower surface of
clustered fronds. Grows best under high light in drained, fertile pinnule, bar = 2 mm, after
moist to moist-dry, basic garden soil with sand. This species Knobloch and Correll (1962).
is relatively easy to grow.

Because of its bead-like segments, Cheilanthes villosa re-

sembles C. lendigera. It differs by its dense covering of
Figure 13.27.20. Cheilanthes tomentosa:
broadly ovate scales on the costae and crinkled hairs on the
a. frond, bar = 2.5 cm (1 in.); b. lower
upper surface of the segments. It is native to the south-
surface of fertile pinnule, bar = 5 mm
western United States and northern Mexico.
(0.2 in.), after Knobloch and Correll
(1962).
Cheilanthes wrightii Hooker  13.27.22
Wright’s lip fern
Semi-hardy, Zone (7)8
A small fern with long-creeping rhizomes and clustered to
scattered fronds. Grows under high light in drained, moist-
dry to dry garden soil with sand.
Cheilanthes wrightii is characterized by two-pinnate-pin-
natifid, lanceolate to ovate-triangular blades that are 1–4
cm (0.5–1.5 in.) wide. The segments vary from oblong to
linear and are glabrous on both surfaces. The species is na-
a b tive to the southwestern United States and Mexico. It usu-
ally grows on igneous rocks.
 244 C 13

b
Figure 13.27.22.
Cheilanthes wrightii:
a. habit, bar =
2.5 cm (1 in.);
b. lower surface of
fertile pinnule, bar =
2.5 mm (0.1 in.).
After Knobloch and
Correll (1962).

Cibotium  13.28.1–3

Cibotiums are large to huge tree ferns, usually with tall
trunks. The Mexican tree fern long known in cultivation,
Cibotium schiedei, does not form an erect trunk but has a
massive crown that produces lacy, gracefully arching fronds.
Figure 13.28.2. Cibotium glaucum: stipes and trunk with
Hairs taken from the trunks and stipes of the Hawaiian tree
hairs.
fern (C. glaucum), called hapu, were used to stuff pillows
and mattresses, but they had the disadvantage of deterio-
rating with age. During the mid-1950s, plants of C. glau-
cum from the island of Hawaii were sold in the continental of the trunks would occasionally contain trunks of the man
United States under the incorrect name “C. chamissoi.” In fern (C. menziesii ), which can be distinguished by the
southern California they are grown outdoors, and in colder blackish, stiff hairs on its stipes and trunk. Man fern has
climates they are used as indoor pot plants. They are still never been abundant in the trade and is harder to grow than
popular and nowadays are propagated from spores instead C. glaucum.
of from trunks collected from wild populations. Shipments The stems of Cibotium vary from prostrate to crown-
like to tall trunks, depending on the species. They bear
long, densely matted hairs, which also occur on the stipe
bases. The blades are two-pinnate-pinnatifid or more di-
vided, broadly triangular, and lightly to strongly whitish
on the undersides. Some species have eared segments; that
is, they have a slightly longer basal lobe that points forward.
The secondary costa and its midribs are ridged on the upper
surface. As in all tree ferns of the family Dicksoniaceae, the

Figure 13.28.3. Cibotium chamissoi: sori and indusia

Figure 13.28.1. Cibotium glaucum: habit. details, bar = 5 mm (0.2 in.). Courtesy of T. Hoshizaki.
 F  F A  C 245

sori are marginal and enclosed by a two-lobed indusium,

the lower lobe fitting inside the upper one. This arrange- c
ment gives the sorus a clam-like appearance.
A member of the family Dicksoniaceae, Cibotium is a
readily separated from the cyatheaceous tree ferns by its
marginal sori and the presence of hairs (not scales) on the
stipe and trunk. Dicksonia, which also has hairs, differs by
having fronds that are narrowed at the base and green (not b
whitish) on the undersides; the outer indusial lobe more
like the leaf segment in color and texture; and the inner in-
dusial lobe thin and brownish.
Cibotium contains 10 species and is native to Central
America, Asia, and Hawaii. The genus name comes from
the Greek kibotos, a small chest or casket, alluding to the
shape of the indusium. For more on culture, see the sec-
tion in Chapter 10 on “Tree Ferns.” For a technical treat- Figure 13.28.4. Cibotium barometz: a. pinnae from
ment of the species, see Maxon (1912) and Palmer (1994). young plant, bar = 5 cm (2 in.); b. primary pinnules, bar
= 1 cm (0.4 in.); c. prostrate rhizome from mature plant,
bar = 5 cm (2 in.).
Cibotium barometz (Linnaeus) J. Smith
 13.28.4
Scythian lamb, vegetable lamb
Tender grew and remained attached to the soil by a “trunk” from
its navel. It fed on the surrounding vegetation and died
A large tree fern with prostrate stems, not upright trunks.
when this was exhausted. In the late 1700s botanists con-
Grows under medium light in moist potting mix. This spe-
cluded that C. barometz was the source of this legendary
cies is rarely cultivated.
plant, but the legend is actually based on the cotton plant.
The rhizomes of Cibotium barometz are covered with
See Moran (1992a) and Lee (1887) for a fuller account of
golden-brown hairs, and similar hairs occur on the rachis
the legend.
and costae but are matted and woolly. The sori are limited
to the lower half of segments. This species is native to
Cibotium chamissoi Kaulfuss  13.28.3, 5
southeastern Asia.
Tender
Because it is a tree fern, Cibotium barometz is banned
from import in countries abiding by the CITES treaty, A large tree fern. Grows well under medium light in moist
which bans trade in rare or endangered plants and animals potting mix.
(see Appendix III). In the pre-treaty days of the 1930s, C. The stipes of Cibotium chamissoi are woolly only at the
barometz was listed in trade catalogs in the United States, base, and the blades are dull green to slightly glaucous on
but it is unknown whether the plants were actually this spe- the lower surfaces and noticeably covered with cobweb-
cies. like, tan hairs. The segments are not eared. Though en-
The stems are occasionally sold in oriental markets for demic to the Hawaiian Islands, this species is rarely culti-
medicinal use. Concoctions from the rhizome are used to vated in Hawaii, and plants sold by this name in the contin-
treat arthritis and joint pains, and the hairs are used to ental United States are mostly C. glaucum.
staunch bleeding. In some parts of China it is common for
households to have a piece of Cibotium trunk with attached Cibotium glaucum (Smith) Hooker & Arnott
stipe bases from which the hairs can be pulled off and put  13.28.1, 2, 6;  3
on cuts and scrapes. The pieces of stem are fashioned into Hapu, Hawaiian tree fern
an article that resembles a small dog or lamb, with the stipe Semi-tender
bases serving as the legs. Sometimes fiddleheads are at- A large tree fern, with trunks to 5 m (16 ft.). Grows well
tached to the rhizome to represent ears or the tail. In south- under medium to high light in moist garden soil or potting
ern China, where Cibotium barometz is native, it is called mix. This species is the most common Cibotium in cultiva-
the “golden-haired dog.” tion. It may root from bare-root trunks, but most trade
Cibotium barometz is associated with the legend of the plants are grown from spores. It grows slowly and is not as
vegetable lamb of Tartary, sometimes also called the cold hardy as Sphaeropteris cooperi, another very common
Scythian lamb. During the Middle Ages it was believed that tree fern.
there existed a half-plant, half-lamb creature somewhere in The stipes of Cibotium glaucum are pale and woolly at
the region north of the Black Sea (Tartary or Sythia) that the base or throughout. The segments are glaucous on the
246 C 13

lower surface, glabrous or with cobweb-like white hairs,

a b
and one or both segments are eared at the base of at least
some pinnules. The species is endemic to the Hawaiian
Islands.
In the trade Cibotium glaucum is often misidentified as
C. chamissoi Kaulfuss.

Cibotium menziesii Hooker  13.28.7

Man fern
Marginally semi-tender
A large tree fern, with trunks to 5 m (16 ft.). Grows well
under medium light in moist potting mix. The plants re-
quire more heat and humidity than Cibotium glaucum.
Cibotium menziesii has stipes clothed with thick, waxy,
stiff hairs. The segments are glossy and lack ears. Minute
tufts of hairs are present on the lower surface. The species is
endemic to the Hawaiian Islands.
c

a b

Figure 13.28.5. Cibotium chamissoi: a. pinna, bar = 5 cm

(2 in.); b. primary pinnule, bar = 1 cm (0.4 in.); c. hairs
on lower segment surface, bar = 5 mm (0.2 in.).

a b

Figure 13.28.7. Cibotium menziesii: a. pinna, bar = 5 cm

(2 in.); b. primary pinnules, bar = 1 cm (0.4 in.);
c. lower segment surface dotted with minute tufts of
hair, bar = 5 mm (0.2 in.).

Cibotium schiedei Schlechtendal & Chamisso

c  13.28.8
Mexican tree fern
Figure 13.28.6. Cibotium glaucum: a. pinna, bar = 5 cm Semi-tender
(2 in.); b. primary pinnule, bar = 1 cm (0.4 in.); c. eared A large tree fern with stout crowns and stems that form a
segments at base of primary pinnule, bar = 5 mm trunk when very old. Grows well under medium light in
(0.2 in.). moist, well-drained potting mix. This fast-growing species
 F  F A  C 247

a c

Figure 13.29.1. Coniogramme japonica: habit.

Figure 13.28.8. Cibotium schiedei: a. pinna, bar = 5 cm

(2 in.); b. primary pinnule, bar = 1 cm (0.4 in.); c. sori
protruding beyond segment margins, bar = 5 mm
(0.2 in.).

has gracefully arching fronds, but older plants are often too
big for most homes. Plantlets sometimes emerge as off-
shoots at the base of the crown. These may be separated
and rooted, but are usually hard to establish.
The stipe of Cibotium schiedei is clothed with yellow-
brown silky hairs, and the blades are large, 2–3 m (6.5–10
ft.) long, and light green on the upper surface but glaucous
on the lower. The segments lack ears, and the sori often pro-
trude beyond the margin. The species is native to Mexico.
A similar species, Lophosoria quadripinnata (J. Gmelin)
C. Christensen, is becoming more frequently cultivated. It
might be identified as a Cibotium because it also has woolly
hairs on the trunk and stipe base and large, finely divided
leaves that are glaucous beneath. It can be distinguished
from Cibotium schiedei by having sori borne on the lower
surface of the leaf, rather than at the margin, and by the
lack of indusia.

Coniogramme  13.29.1, 2

Coniogramme makes a good ground cover but needs pro-
tection from slugs and snails. These medium- to large-sized,
creeping, terrestrial ferns have strap-shaped pinnae. The
stipes contain one large vascular bundle, and the blades are
mostly one-pinnate, except for the basal pair (or two) of Figure 13.29.2. Coniogramme japonica: sporangia along
pinnae, which tend to be more divided. The blade always veins.
 248 C 13

terminates in an apical segment that resembles the lateral Coniogramme japonica (Thunberg) Diels
pinnae, and the veins tips are conspicuously enlarged on  13.29.1, 2, 4
the upper surface of the leaf. The sori run along the veins Bamboo fern
between the midrib and margin of the segments. Indusia Semi-hardy, Zone (7)8
are absent. The cultivated species of Coniogramme mimic a A medium-large fern with deciduous fronds and medium-
large Pteris cretica but differ by having longer-creeping rhi- to long-creeping rhizomes. Grows well under low light in
zomes, deciduous fronds, and sori that run between the moist garden soil or potting mix.
midrib and margin of the segments, not along the margin Coniogramme japonica resembles the preceding one but
as in Pteris. has netted veins. It is native to China, Taiwan, Korea, and
The genus consists of about 20 species native to the Old Japan.
World tropics, Japan, and in the Pacific to Hawaii. Its name ‘Variegata’. Segments with yellow along the midribs.
comes from the Greek konios, dusty, and gramme, line, re-
ferring to the lines of sori along the veins.

Coniogramme intermedia Hieronymus

 13.29.3
syn. Coniogramme fraxinea (G. Don) Diels
Hardy
A medium to large fern with deciduous fronds and me-
dium- to long-creeping rhizomes. Grows well under low
light in moist garden soil.
Coniogramme intermedia is characterized by free veins
and strap-shaped ultimate segments. It is native to India,
China, Korea, Japan, Malaysia, and Hawaii.

Figure 13.29.4. Coniogramme japonica: frond, bar =

5 cm (2 in.).

Cryptogramma  13.30.1–3

Rock brake, parsley fern
A coarse, parsley-like foliage characterizes Cryptogramma,
which consists of small ferns used in rock gardens in cold-
temperate climates. The plants are difficult to grow.
The fronds vary from two- to three-pinnate (rarely four-
pinnate) and have toothed sterile segments and free veins.
The segments of the fertile fronds appear pod-like due to
the strongly enrolled margins (the false indusium) that
cover the sporangia.
Cryptogramma contains about 10 species and occurs in
North America, Europe, Asia, southern Chile, and Argen-
tina. The genus name comes from the Greek kryptos, hid-
Figure 13.29.3. Coniogramme intermedia: frond, bar = den, and gramme, line, alluding to the line of sori hidden by
5 cm (2 in.). the enrolled leaf margin.
 F  F A  C 249

Cryptogramma acrostichoides R. Brown

 13.30.1–4
American parsley fern
Very hardy, Zone 2
A small fern with suberect to erect rhizomes and tufted
evergreen fronds. Grows well under medium light in well-
drained garden soil. The plants seem to grow best if the soil
is kept moist in the spring but moist-dry other times of the
year.
The rhizomes of Cryptogramma acrostichoides are hard,
short, scaly, and much branched. The sterile fronds remain
green during the winter and senesce the following spring.
The stipes are dark brown toward the base, and the blades
are up to three-pinnate and somewhat leathery. Minute,
cylindrical hairs occur on the upper blade surface and in
the grooves of the stipe, rachis, and costae. The sterile seg-
ments vary from oblong to ovate-lanceolate, with 6–12
teeth or shallow lobes. The species is native to North Amer-
ica and Asia.
The Cascade parsley fern, Cryptogramma cascadensis E.
R. Alverson, resembles C. acrostichoides and might be cul-
tivated in the northwestern United States. It differs from C.
acrostichoides by having sterile fronds deciduous in autumn
and by the lack of hairs on the upper surfaces of the blades
and in the grooves of the stipes and rachises. It is native to
the northwestern United States and adjacent Canada.

Figure 13.30.1. Figure 13.30.2.

Cryptogramma Cryptogramma acrostichoides:
acrostichoides: sterile fertile frond. b
frond.

Figure 13.30.4. Cryptogramma acrostichoides: a. fertile

Figure 13.30.3. Cryptogramma acrostichoides: fertile frond (left) and sterile frond (right), bar = 5 cm (2 in.);
segments. b. fertile pinnule, bar = 5 mm (0.2 in.).
 250 C 13

Some botanists consider Cryptogramma acrostichoides to Cryptogramma crispa resembles C. acrostichoides but has
be a variety of the European species, C. crispa, but they dif- thinner, deciduous leaves, and sterile blades up to four-pin-
fer consistently in blade cutting and chromosome number nate. It is native to Europe.
(2n = 60 in C. acrostichoides and 2n = 120 in C. crispa).
They also differ in habitat. Cryptogramma acrostichoides Cryptogramma sitchensis (Ruprecht) T. Moore
grows on noncalcareous rocks, often in slightly drier habi-  13.30.6
tats than is usual for other species in the genus. In contrast, Alaska parsley fern
C. crispa grows on calcareous rocks in more humid places. Very hardy, Zone 2
A small fern with suberect to erect rhizomes and tufted
Cryptogramma crispa (Linnaeus) R. Brown evergreen fronds. Grows well under medium light in gar-
 13.30.5;  23 den soil with good drainage. Probably does best if the soil is
European parsley fern moist in spring and drier at other times.
Very hardy, Zone 2 Cryptogramma sitchensis greatly resembles C. acrostichoi-
A small fern with erect to suberect rhizomes. Grows best des and is believed to have arisen by hybridization between
under medium light in well-drained garden soil. The plants that species and possibly C. raddeana Formin of eastern
seem to do best if the soil is kept moist in the spring but Asia. It differs from C. acrostichoides by its obovate seg-
moist-dry other times of the year. ments with two to six deep lobes and at least some three- to
four-pinnate sterile fronds. It is native to northwestern
Canada and Alaska, where it typically grows among rocks
in acid soils.

Figure 13.30.6. Cryptogramma sitchensis: a. habit, with

Figure 13.30.5. Cryptogramma crispa: habit, with sterile two sterile fronds (shorter) and two fertile fronds (taller),
fronds (left and right) and fertile frond (center), bar = bar = 5 cm (2 in.); b. sterile pinnule, bar = 3 mm
5 cm (2 in.). After Sowerby (1855). (0.1 in.). After Cody and Britton (1989).
 F  F A  C 251

Cryptogramma stelleri (S. G. Gmelin) Prantl Ctenitis contains 150 species and is native to tropical re-
 13.30.7 gions of the world. The genus name comes from the Greek
Slender rock brake, cliff brake, fragile rock brake kteis, comb, referring to the comb-like lobes of the pinnae
Very hardy, Zone 2 in Ctenitis submarginalis (Langsdorff & Fischer) Ching, the
A small fern with deciduous fronds distributed along a slen- type species of the genus.
der (1–1.5 mm wide), creeping rhizome with few branches.
Thrives under medium light in well-drained, moist, basic Ctenitis sloanei (Poeppig ex Sprengel) C. V. Morton
garden soil. The fronds die back by late summer and need  13.31.1–4
less water at that time. syn. Ctenitis ampla auct. non (Humboldt & Bonpland ex
Cryptogramma stelleri bears stipes that are dark at the Willdenow) Ching
base but greenish above. The blades are up to two-pinnate, American tree fern, Florida tree fern
herbaceous to membranous, and thin. The segments of Tender, Zone 10(11)
sterile leaves are ovate-lanceolate to fan-shaped and lobed A large tree fern with erect stems that can form trunks up to
halfway to the midrib. The species is native to northern 1.3 m (4 ft.) tall, although it infrequently does so in culti-
North America, Europe, and Asia, where it typically grows vation. Grows under medium light in moist, drained pot-
on calcareous rocks, often forming dense colonies in crev- ting mix. This species is sensitive to transplanting and is
ices or ledges. difficult to establish.
The stipe scales of Ctenitis sloanei are dense, linear or
thread-like, orange (rarely brown), and toothed. The veins
reach all the way to the margin, and the indusium falls away
early. The species is native to the United States (Florida)
and American tropics.

Figure 13.30.7. Cryptogramma

stelleri: habit, sterile frond
(left) and fertile frond (right),
bar = 5 cm (2 in.). After Cody
and Britton (1989).

Figure 13.31.1. Ctenitis sloanei: habit.

Ctenitis
These medium- to large-sized, terrestrial ferns are infre-
quently cultivated, except in Florida. The Florida tree fern
(Ctenitis sloanei ) can form a trunk up to 1.3 m (4 ft.) tall.
Ctenitis can usually be distinguished by the following
combination of characteristics: scaly rhizomes; three or
more vascular bundles in the stipe; minute, jointed, often
reddish hairs on the upper surfaces of the rachises and
costae; free veins (not netted); and round sori. Usually the
rachis is grooved on its upper surface and the costae are not.
In those rare instances where both are grooved, the grooves
are interrupted at the junctures, not continuous. This is in
contrast to dryopteroid ferns such as Cyrtomium, Dryop- Figure 13.31.2. Ctenitis: rachis, groove not opening into
teris, and Polystichum, in which the grooves are prominent the costae, costae not grooved (left), costae grooved
on both the rachises and costae and run into one another. (right).
252 C 13

Cyathea
syn. Hemitelia
Tree fern
Cyathea is a genus of elegant tree ferns that typically bear a
crown of large fronds on top of a tall, erect trunk. They can
reach 15–20 m (50–65 ft.) tall in the wild, and some
mountain slopes support dense stands of these stately trees.
In temperate areas this genus is seen mainly in conservato-
ries or large greenhouses. Tree ferns can be grown as house-
plants, but most species seem to do best outdoors in sub-
tropical or tropical gardens, where they can grow up to 5 m
(16 ft.) in height. Most thrive under filtered light, but they
will grow in full sun in coastal areas where heat and light are
not too intense. Growth is best in rich, organic, moist soil.
(See Chapter 10 for more information.)
Tree ferns have several uses. In some cultures they are
used for food, fodder, or building material. The trunks are
highly valued in backwoods construction because they are
strong and resist decay. The thick layer of tough aerial roots
(the root mantle) found around the trunk of some species
is used as a growing medium for epiphytes, especially or-
chids. The scales on the trunk and stipes can shed readily,
and care should be taken to avoid getting the scales into
your eyes and on your skin. Although not toxic, the scales
can cause severe irritation.
Figure 13.31.3. Ctenitis sloanei: sori and indusia. Most species of Cyathea have upright, scaly trunks and
a layer of tough aerial roots, at least toward the base. The
fronds are borne in a whorl at the top of the stem, forming
a palm-like growth habit, and they usually have finely di-
vided, two- to three-pinnate blades. The stipe scales lack
a dark bristles at the apex, in contrast to those of the similar
genus Alsophila, and the marginal cells differ in orientation
and often in color from the central cells (when viewed
under 20× magnification). The round sori are borne on the
undersides of the blades, and the indusia, when present,
vary from scale-like to cup-like or globose.
To distinguish cyatheaceous ferns from other tree-fern
b groups, look at the indument or covering on the stem apex
or stipe bases. Cyathea has a covering of scales, whereas
dicksoniaceous tree ferns such as Cibotium and Dicksonia
have hairs. Sadleria has scales on the trunk and stipe, but its
fronds are narrow and the sori are covered by a conspicuous
oblong indusium.
Cyathea is native to the Old and New World tropics,
occurring mainly in cloud forests. It consists of about 115
species. The genus name comes from the Greek kyathos, a
wine cup, referring to the cup-like shape of the indusium in
certain species.
c Cyathea is defined here in the narrow sense, excluding
Alsophila, Cnemidaria, and Sphaeropteris. Although pteri-
Figure 13.31.4. Ctenitis sloanei: a. frond from mature dologists agree that these genera represent natural groups,
plant, bar = 5 cm (2 in.); b. frond from immature plant, they disagree over whether the groups should be recognized
bar = 5 cm (2 in.); c. veins reaching the margin, bar = as separate genera. Some pteridologists classify the species
5 mm (0.2 in.). of all four groups into one large, inclusive genus, Cyathea.
 F  F A  C 253

For a reference to tree-fern taxonomy, see Kramer and Cyathea arborea (Linnaeus) Smith  13.32.1, 2
Green (1990). The following guide to the cultivated genera West Indian tree fern
of Cyatheaceae is provided to facilitate identification: Tender to very tender
A large tree fern. Grows best under medium light in moist
potting mix. This species grows in southern Florida and
seems to require warm nights.
The fronds of Cyathea arborea eventually fall away from
the trunk. The stipe scales are mostly whitish, and the base
of the segment midrib on the underside have one or two
white, puffy scales. The saucer-shaped indusium forms a
cup beneath the sporangia. The species is native to the West
Indies, Venezuela, and Colombia.

Cyclopeltis
These medium-sized terrestrial ferns are grown in south-
ern Florida and are sometimes used in public parks in the
West Indies. The rhizome is thick, densely scaly, and as-
cending or erect. The blades are one-pinnate with many
Alsophila: Stipe scales tipped with a dark bristle; marginal sessile pinnae and a terminal one resembling the laterals.
cells of scales are different from the central cells in orienta-
tion, size, and often color; stipes spiny, with spines black
and shiny. (Alsophila australis, A. tricolor)

Cyathea: Stipe scales not tipped with a dark bristle; mar-

ginal cells of scales are different from the central cells in ori-
entation, size, and often color; stipes spiny or not, but if
spiny, the spines are not black and shiny. (Cyathea arborea)

Sphaeropteris: Stipe scales tipped with a short, dark, spine-

like hair, not a bristle; marginal cells of scales are similar to Figure 13.32.1. Cyathea arborea: a. habit, bar = 1 m
those in the center, except for occasional black teeth; stipes (3 ft.), after Little and Wadsworth (1964); b. tip of stipe
spiny or not, but if spiny, the spines are not black and scale ending in several cells (not a bristle or hair), bar =
shiny. (Sphaeropteris aramaganensis, S. cooperi, S. excelsa, S. 0.1 mm; c. margin of scale, with outer border of cells
horrida, S. medullaris) oriented at right angles to the central cells, bar =
0.1 mm.
 254 C 13

b
a

b
c

Figure 13.33.1.
Figure 13.32.2. Cyathea arborea: a. pinnule, bar = 1 cm Cyclopeltis
(0.4 in.); b. stipe scale, bar = 1 cm (0.4 in.); c. detail of semicordata: a
segment with bullate scale at base, bar = 1 mm, a. habit, bar =
illustration by P. J. Edwards, used with permission; 10 cm (4 in.);
d. enlargement of bullate scale, bar = 0.1 mm, b. sori with peltate
illustration by P. J. Edwards, used with permission. indusia, bar = 5
mm (0.2 in.).

The veins are free, and the sori are round and covered by
peltate indusia. Cyrtomium  13.34.1, 2
The peltate indusium immediately separates Cyclopeltis syn. Phanerophlebia
from Dryopteris and Thelypteris, which it superficially re- This genus of terrestrial or rock-inhabiting ferns is well rep-
sembles. It differs from Polystichum by having entire, not resented by the house holly fern (Cyrtomium falcatum),
spiny, margins and pinnae that lack an inner, upward- widely grown indoors in pots for its leathery, glossy green,
pointing ear. durable foliage. The most common cultivar in nurseries is
The genus, which consists of five species, is native to the Rockford fern (C. falcatum ‘Rockfordianum’), which
tropical America and southeastern Asia. The genus name has irregularly fringed pinna margins. This plant is easy to
comes from the Greek kyklos, circle, and pelte, shield. grow, but its soil should be kept on the moist-dry side. It
has become naturalized in parts of the United States with
Cyclopeltis semicordata (Swartz) J. Smith mild winters. In the wild Cyrtomium grows in the soil or on
 13.33.1 rocks, not as epiphytes.
Tender Cyrtomium can usually be distinguished by erect, scaly
A medium-sized fern with erect rhizomes. Best grown rhizomes; one-pinnate, evergreen fronds; netted veins; and
under medium light in moist garden soil. The plants are round sori covered by an umbrella-shaped indusium. The
occasionally grown in Florida. lateral pinnae tend to be nearly equal in size, and the ter-
Cyclopeltis semicordata has one-pinnate fronds with ses- minal pinna usually resembles the lateral ones, except that
sile pinnae that have a semicircular basal lobe overlapping the base can be lobed.
the rachis. The upper surfaces of the rachises and costae Some pteridologists classify the New World species of
bear short hairs, as is typical for ferns related to Ctenitis and Cyrtomium (which are rarely cultivated) in Phanerophlebia
Tectaria. It is widespread in the American tropics where it because those species have casually to sparingly netted veins
typically grows on wet forest floors. like Phanerophlebia and in contrast to the Old World spe-
cies of Cyrtomium. At the other extreme, Cyrtomium and
 F  F A  C 255

Figure 13.34.1. Cyrtomium falcatum: habit.

Figure 13.34.3. Cyrtomium caryotideum: a. frond, bar =

5 cm (2 in.); b. serrate-spinulose margin, bar = 5 mm
(0.2 in.); c. indusium, bar = 1 mm.

Figure 13.34.2. Cyrtomium falcatum: veins, sori, and

indusia, bar = 1 cm (0.4 in.). and densely spiny-serrate along the margins. The sori are
often sparse or absent near the margins, and the indusium
margin is distinctly erose-dentate to fimbriate. The species
is native to Japan, China, Taiwan, Vietnam, India, the Phil-
Phanerophlebia are sometimes merged into Polystichum. For ippines, and Hawaii.
more information on the classification of these ferns, see A putative hybrid with Cyrtomium falcatum occurs in
Kurata (1963) and Yatskievych (1996). the trade and needs further study. Its frond and pinnae re-
Cyrtomium is native to the tropics and subtropics of the semble C. falcatum, but the pinnae are fewer and further
world and contains about 15 species. The genus name apart and have sparingly spiny-serrate margins and erose-
comes from the Greek kyrtoma, arch, referring to the veins dentate to fimbriate indusia, as in C. caryotideum.
that anastomose in arches in some species.
Cyrtomium falcatum (Linnaeus f.) C. Presl
Cyrtomium caryotideum (Wallich ex Hooker &  13.34.1, 2, 4, 5;  24
Greville) C. Presl  13.34.3 House holly fern
Dwarf holly fern Semi-hardy or hardier, reportedly to −12°C (10°F),
Hardy, Zone 6 Zone (6)7
A medium-sized fern with erect rhizomes. Grows well A medium-sized fern with erect rhizomes. Grows well
under medium light in moist-dry, basic(?) garden soil or under medium to high light in moist-dry garden soil or
potting mix. In some areas of the United States, such as the potting mix. The plants are easy to cultivate and have at-
Southeast and southern California, the crown seems to suc- tractive evergreen or semi-evergreen fronds. They tolerate
cumb to a disease. For some reason the crowns often pro- drier air than most ferns.
duce aborted fronds and eventually stop growing. The fronds of Cyrtomium falcatum are about 75 cm (30
Cyrtomium caryotideum is characterized by three to six in.) tall and bear 10 (rarely 8) or more pinna pairs. The pin-
pairs of large pinnae. The pinnae are hastate or have a tri- nae are thick, leathery, and dark glossy green. In wild plants
angular acuminate ear or are earless. They are dull green the margins are nearly entire, although a few obscure,
256 C 13

minute teeth may be present. The indusia margins are

erose. This species is native to India, China, and Japan.
The major cultivars are
b ‘Butterfieldii’. Butterfield holly fern. Figure 13.34.5a.
Margins coarsely serrate.
‘Mayi’. May fern. Figure 13.34.5b. Fronds often forked
or branched; pinna apices widened, somewhat ruffled, and
irregularly lacerate-toothed.
‘Rockfordianum’. Rockford holly fern. Figure 13.34.5c.
Pinna margins regularly fringed, somewhat like a holly leaf.
This cultivar is common in the trade.
Figure 13.34.4. Cyrtomium
An unnamed dwarf form (Figure 13.34.5d) is reported
falcatum: a. habit, bar = 5 cm
a to be sexual whereas the other cultivars are apogamous.
(2 in.); b. indusium, bar =
This dwarf form is a choice plant presently in limited cir-
1 mm.
culation.

Cyrtomium fortunei J. Smith  13.34.6

Hardy, Zone 5(6)
A medium-sized fern with erect rhizomes. Grows well
under medium light in moist-dry garden soil or potting
mix. The plants are easy to grow from spores and have at-
tractive evergreen fronds.
Cyrtomium fortunei resembles C. falcatum but has gray-
green, less glossy, less leathery fronds. There are 10–25 pairs
of pinnae and these have serrate margins, at least distally.
The terminal pinna is smaller than the lower lateral ones,
and the indusium margins are entire or subentire. The spe-
cies is native to Japan, South Korea, southeastern China,
and Indochina.
This species varies tremendously and its varieties some-
times intergrade.

b
b

c
d

Figure 13.34.6. Cyrtomium

fortunei: a. frond, bar = 5 cm
(2 in.); b. indusium, bar =
1 mm.

Figure 13.34.5. Cyrtomium falcatum, frond variants: a

a. ‘Butterfieldii’; b. ‘Mayi’; c. ‘Rockfordianum’;
d. unnamed variant. Bar = 5 cm (2 in.).
 F  F A  C 257

var. clivicola (Makino) Tagawa. Upper surfaces of the tured blades about 30 cm (12 in.) long and 9 cm (3.5 in.)
blades matte yellow-green; pinnae typically in about 10 wide. The pinnae are distant from one another, nearly op-
(15) pairs, usually to 4 cm (1.5 in.) long, mostly eared at the posite or alternate, and crenate-serrate. The apex is unusual
base; indusium margins somewhat erose. Native to China in being short and deeply pinnatifid. The indusium mar-
and Japan. gins are erose. The species is native to China.
var. fortunei. Upper surfaces of the blades deep green,
shiny; pinnae in 15–25 pairs, less than 2–3 cm (0.75–1.2 Cyrtomium macrophyllum (Makino) Tagawa
in.) wide, generally truncate and nearly earless; indusium  13.34.8, 9
margins entire. The indusia are grayish white throughout Big-leaf holly fern
(concolorous). Native to Japan, South Korea, southeastern Semi-hardy, Zone 6(7)
China, Indochina, and Thailand. A medium-sized fern with erect rhizomes. Grows under
var. intermedium Tagawa. Pinnae about 15 (8–19) pairs, medium light in moist-dry garden soil or potting mix. The
bases widely wedge-shaped, weakly eared or not, gradually evergreen fronds are a lighter green than those of Cyrto-
narrowed toward an attenuate apex. The indusia are black- mium falcatum.
ish in the center. Native to Japan. Some of the trade mate- Cyrtomium macrophyllum has 2–8 (12) pinna pairs, with
rial in the United States named as this variety probably rep- the lateral pinnae more than 4 cm (1.5 in.) wide and entire
resents var. clivicola. or slightly toothed toward the tip. The pinna bases are
rounded to broadly wedge-shaped, without ears, and the
Cyrtomium lonchitoides (H. Christ) H. Christ terminal pinna is larger than or nearly the same size as the
 13.34.7 lateral pinnae. The indusia are nearly entire. The species is
Semi-hardy (?) native to Japan, China, and the Himalayas.
A medium-sized fern with erect rhizomes. Best grown var. tukusicola (Tagawa) Tagawa (Cyrtomium tukusicola
under medium light in moist garden soil or potting mix. Tagawa). Figure 13.34.9. Separated from var. macrophyl-
Cyrtomium lonchitoides is characterized by stipes only lum by wedge-shaped pinnae and indusia that are black-
5–10 cm (2–4 in.) long and elongate, lanceolate, thin-tex- brown in the center but pale at the margins. Native to
Japan (Honshu and Kyushu). Evergreen and hardy in Salt
Lake City, Utah.

Figure 13.34.7. Cyrtomium

lonchitoides: habit, bar =
5 cm (2 in.).

Figure 13.34.8. Cyrtomium macrophyllum var.

macrophyllum: frond, bar = 5 cm (2 in.).
 258 C 13

Figure 13.34.9.
Cyrtomium
macrophyllum var.
tukusicola: frond,
bar = 5 cm (2 in.).
Figure 13.35.1. Cystopteris fragilis: habit.

ing rhizomes and medium-sized, triangular fronds that are

three-pinnate-pinnatifid. The plants are easy to grow from
spores and may readily volunteer. This native of Japan and
Taiwan prefers partial shade and moist garden soil. It is
listed as hardy to Zone 8. In warmer zones, care must be
taken not to overwater during the cool months and when
the plant is in its deciduous period.
Cystopteris, which contains about 20 species, is native to
Cystopteris  13.35.1, 2
the temperate zones and to montane areas in the tropics.
Bladder fern
The bladder-like indusia are the source of the common
name of the genus Cystopteris. These mostly small ferns
with delicate foliage are used in shaded rock gardens and on
damp banks. They are hardy in cold-temperate areas, and a
few species are adapted to boreal or alpine climates. Cystop-
teris is seldom cultivated in warmer climates because of its
deciduous nature and need for a cold rest period. Many
species thrive in basic soils.
The genus is characterized by two vascular bundles in
the stipe; thin, herbaceous blades one- to three-pinnate-
pinnatifid; free veins; and round sori covered by an arched-
inflated indusium. The latter trait is often difficult to see
because it gets pushed aside by the sporangia as they ma-
ture. On young leaves, the indusium appears scale-like or
hood-shaped and is attached beneath the sorus on the side
closest to the midrib.
The characteristics used to distinguish the species of Cys-
topteris overlap considerably, and therefore the species can
be difficult to identify. The widespread Cystopteris fragilis is
particularly variable and is considered by some botanists to
comprise several species. Cystopteris might be confused with
Woodsia, which differs by having the indusium surrounding
the underside of the sorus rather than attached to one side.
Cystopteris also resembles Acystopteris, a genus repre-
sented in the trade by Acystopteris japonica (Luerssen) Na-
kai. It can be separated from Cystopteris by the multicellu-
lar hairs on its fronds and by the small or absent indusia.
Acystopteris japonica is a deciduous species with long-creep- Figure 13.35.2. Cystopteris fragilis: sori and indusia.
 F  F A  C 259

The genus name comes from the Greek kystos, bladder, and
pteris, fern, and refers to the inflated indusium covering the
sorus.

Cystopteris alpina (Lamarck) Desvaux  13.35.3 b

syn. Cystopteris fragilis subsp. alpina (Lamarck) Hartman,
C. regia of authors, not (Linnaeus) Desvaux
Hardy
A small fern with short-creeping rhizomes and deciduous
leaves. Grows under low-medium light in moist, basic gar-
den soil or potting mix. c
Cystopteris alpina resembles C. fragilis in growth habit
and leaf shape but differs by the pinnules being narrower
and wedge-shaped, more finely divided, and the veins end-
ing in a marginal notch. In nature it usually grows on lime-
stone. It is native to the mountains of Europe, Asia Minor,
and northeastern Africa.

Cystopteris bulbifera (Linnaeus) Bernhardi

 13.35.4
Berry bladder fern, bulblet bladder fern
Very hardy, Zone 3 a
A small to medium fern with short-creeping rhizomes and
deciduous fronds bearing buds along the rachises and (less
frequently) the costae. Thrives under low-medium light in

Figure 13.35.4. Cystopteris bulbifera: a. habit, bar = 5 cm

(2 in.); b. leaf bud, bar = 1 cm (0.4 in.); c. pinnule with
sori and indusia, bar = 3 mm.

moist-wet, basic garden soil or potting mix. The plants are

easy to grow in temperate climates and are readily propa-
gated from bulblets.
Cystopteris bulbifera is the only species of this genus that
bears large, green, fleshy bulblets along the rachises and
costae, or has reddish stipes (although this color often
Figure 13.35.3. Cystopteris fades with age). The blades are thin and one- to two-pin-
alpina: habit, bar = 5 cm nate-pinnatifid, and the rachis and costa usually bear tiny
(2 in.). gland-tipped hairs, especially while young (magnification
needed). The veins end in teeth, not in sinuses. This species
is native to (primarily) the eastern United States and adja-
cent Canada.
‘Crispa’. Margins ruffled.

Cystopteris dickieana Sim  13.35.5

syn. Cystopteris fragilis subsp. dickieana (Sim) Hylander
Dickie’s bladder fern
Hardy
A small fern with deciduous leaves clumped along a short-
creeping rhizome. Grows well under low-medium light in
moist-wet, drained garden soil or potting mix.
 260 C 13

Figure 13.35.6.
Cystopteris fragilis:
a. habit, bar = 5 cm
(2 in.); b. sorus and
indusia, bar = 1 mm,
after Knobloch and
Correll (1962).

Figure 13.35.5.
Cystopteris dickieana:
habit, bar = 5 cm (2 in.).
a

Cystopteris montana (Lamarck) Bernhardi ex Desvaux

 13.35.7
Mountain bladder fern
Very hardy, Zone 1
A small fern with deciduous fronds and medium-creeping
rhizomes. Grows well under low-medium light in basic,
moist-wet, well-drained garden soil or potting mix.
Cystopteris dickieana greatly resembles C. fragilis, and Cystopteris montana is easily distinguished from other
some pteridologists argue that it is not distinct. It is said to bladder ferns by its nearly black, medium-creeping rhi-
differ primarily by its wrinkled, warty (not spiny) spore sur- zomes and wide, pentagonal blades. It is native to the
faces. The blades and pinnules also tend to be narrower, mountains of North America, Europe, and Siberia.
and the pinnae are typically overlapping. The veins end The fronds of this species are cyanogenic; when crushed,
mostly in marginal notches. The species is native to north- they emit the pungent odor of prussic acid, which resem-
ern North America, Greenland, Iceland, Europe, Asia, and bles the smell of burnt almonds. Presumably this deters her-
northern Africa. bivores.

Cystopteris fragilis (Linnaeus) Bernhardi Cystopteris protrusa (Weatherby) Blasdell

 13.35.1, 2, 6  13.35.8
Brittle bladder fern, fragile bladder fern syn. Cystopteris fragilis var. protrusa Weatherby
Very hardy, Zone 1 Southern fragile fern, woodland bladder fern, lowland
A small fern with short-creeping rhizomes and deciduous bladder fern
leaves. Grows well under low-medium light in garden soil Hardy, Zone 5
or potting mix kept moist-wet. This species is easy to grow. A small fern with creeping rhizomes and clumped, decidu-
The blades of Cystopteris fragilis are thin and herbaceous, ous fronds. Thrives under low-medium light in moist-wet
with nonoverlapping pinnae and veins ending mostly in garden soil or potting mix.
teeth, not shallow notches. The pinnules are roughly trian- Cystopteris protrusa resembles C. fragilis, but its creep-
gular and sessile or very short-stalked. Minute gland-tipped ing rhizomes protrude 1–4 cm (0.5–1.5 in.) beyond the
hairs, present in some other bladder ferns, are absent. The current year’s cluster of fronds. Another difference is that
species is extremely variable and one of the most widespread the pinnules on large, fertile leaves tend to be distinctly
ferns in the world, occurring on every continent. stalked, whereas in C. fragilis they are sessile or nearly so.
An unnamed, crested form is reported in cultivation. The rhizomes bear yellowish hairs not found in any other
 F  F A  C 261

species of Cystopteris. This species is native to the eastern

United States, where it grows on shaded forest floors.

Cystopteris reevesiana Lellinger  13.35.9

syn. Cystopteris fragilis subsp. tenuifolia Clute
Southwestern brittle fern
Hardy, Zone 5
A small fern with short-creeping rhizomes and clumped,
deciduous leaves. Thrives under medium light in moist gar-
den soil or potting mix.
Cystopteris reevesiana resembles C. fragilis but has longer
creeping rhizomes, stipes that are mostly dark purple at
base, and more finely divided blades, with the proximal
pinnae one-pinnate-pinnatifid to two-pinnate. It is native
to the southwestern United States.

Figure 13.35.7. Cystopteris

montana: habit, bar = 5 cm
(2 in.).

Figure 13.35.9.
Cystopteris reevesiana:
frond, bar = 5 cm
(2 in.).

Cystopteris tennesseensis Shaver  13.35.10

Tennessee bladder fern
Hardy, Zone 6
Figure 13.35.8. Cystopteris A small fern with short-creeping rhizomes and clumped,
protrusa: habit, bar = 5 cm deciduous leaves. Thrives under low-medium light in moist-
(2 in.). wet, basic garden soil or potting mix.
Cystopteris tennesseensis is a fertile hybrid resulting from
a cross involving C. bulbifera and C. protrusa. It can be dif-
ficult to distinguish from its parents. Like C. bulbifera, C.
tennesseensis bears bulblets on the lower surfaces of the
 262 C 13

Figure 13.35.11. Cystopteris tenuis:

habit, bar = 5 cm (2 in.).

Figure 13.35.10.
Cystopteris tennesseensis:
habit, bar = 5 cm (2 in.).

this takes practice to recognize. Cystopteris tenuis is a fertile

hybrid originating from a cross between C. fragilis and C.
protrusa. It is native to the United States and Canada.

rachis and (sometimes) costae, but the bulblets are smaller, Davallia  13.36.1–3
scalier, and less fleshy. Minute gland-tipped hairs occur on syn. Araiostegia, Humata, Pachypleuria, Scyphularia,
the rachises and costae (magnification needed), a trait im- Trogostolon
parted from C. bulbifera. The veins end in notches or teeth. Davallias are among the most beautiful and treasured cul-
A characteristic of the Tennessee bladder fern not shared by tivated ferns. Their broad, lacy, often shiny fronds are usu-
other bladder ferns is that nearly all the leaves, even the ally displayed in hanging baskets in conservatories or green-
smallest, bear sori. This species is native to the central houses, where the entire basket may be hidden by the
and eastern United States, where it grows on limestone. It foliage. The most sought-after davallias are those with
is the common bladder fern in the Ozarks of Missouri and finely divided fronds, such as certain selections of the Fiji
Arkansas. davallia (Davallia fejeensis) that bear numerous segments
about the width of a pencil lead. The warm-temperate spe-
Cystopteris tenuis (Michaux) Desvaux cies are grown outdoors in mild climates of the United
 13.35.11 States, although most of the nursery stock is used indoors.
syn. Cystopteris fragilis var. mackayi G. Lawson Besides hanging baskets, davallias are planted over rocks
Mackay’s brittle fern or as ground covers so that their creeping rhizomes can be
Very hardy, Zone 4 seen—the rhizomes are often described as “furry feet.” One
A small fern with short-creeping rhizomes. Grows best species with such rhizomes is the bear’s-foot fern (Davallia
under medium light in moist soil or potting mix. The leaves tyermannii). It is a choice fern of small to medium size with
are deciduous and absent during the winter months. durable, triangular, moderately fine-divided fronds. Al-
Cystopteris tenuis resembles C. fragilis and can be diffi- though slower growing than many other common species,
cult to distinguish. It typically has pinnules that are wedge- D. tyermannii less frequently needs repotting. Resembling
shaped at the base, whereas those of C. fragilis are broadest the bear’s-foot fern, the squirrel’s-foot fern, D. mariesii var.
at the base and triangular. The pinnae and pinnules also stenolepis (D. trichomanoides of trade) is the most common
tend to be more ascending than those of C. fragilis, although species in the trade. It has light brown, furry rhizomes and
 F  F A  C 263

Figure 13.36.3. Davallia mariesii var. stenolepis: sori and

indusia.
Figure 13.36.1. Davallia mariesii var. stenolepis: habit.

is faster growing and softer in appearance than the bear’s-

foot fern, but it is just as robust and adaptable.
The small Japanese ball fern (Davallia mariesii var.
mariesii ) is now rarely seen in the United States. During
the 1960s it was imported from Japan with the rhizomes
tied and shaped into balls, monkeys, pergolas, and other
figures. These were hung in gardens and patios, and when
the fronds emerged, the plants would take on the outline of
the ball or figure. In warm-temperate gardens the Japanese
ball fern can be planted as a ground cover. Its small, trian-
gular, deciduous fronds are attractive, and some forms ap-
pear densely leafy due to overlapping pinnae.
The diminutive Davallia parvula is another attractive
species. It has finely dissected fronds and is often used in
terrariums or bottle gardens where humidity is high and
the tiny plant is safe from being stepped on.
Temperature tolerances of davallias are incompletely
known, but Davallia mariesii var. mariesii can be grown
outdoors where the average minimum January tempera-
ture is about −1°C (30°F). Davallia tyermannii probably
tolerates a similar temperature minimum. Other warm-
temperate species that are likely to tolerate short, infrequent
freezes are D. canariensis, D. griffithiana, D. pyxidata, D.
mariesii var. stenolepis, D. tasmanii, and (marginally) D. tri-
Figure 13.36.2. Davallia mariesii var. stenolepis: frond. chomanoides. Almost all the others are tender outdoors, ex-
cept in areas of Florida and the Hawaiian Islands. For more
information about cultivation, see Hoshizaki (1981) and
the section in Chapter 10 titled “Davallia and Relatives.”
The genus Davallia has several distinctive characteris-
tics. Depending on the species, plants can be epiphytic,
epilithic, or terrestrial. The rhizomes are always long-creep-
264 C 13

ing and scaly, the scales being mostly ovate to lanceolate Araiostegia was delineated primarily by its thin blades,
and peltate; if basally attached, the scales are reniform or finely divided fronds, scaly rachises and costae, and deli-
cordate with overlapping bases. The stipes are jointed to cate, roundish or kidney-shaped indusia attached only at
the rhizomes and fall off cleanly, leaving a circular scar. The the base. The species in this group are rarely cultivated but
blades are anadromous and vary from simple to finely dis- include Davallia hymenophylloides and D. pulchra.
sected (up to four-pinnate) and from lanceolate to triangu- In his monograph on the genus Davallia, Nooteboom
lar in shape. The rachis is raised and grooved on the upper (1994) changed the names of several well-known cultivated
surface, with the grooves running into those of the costae. species. These changes leave many plants with distinct and
The veins are free, not netted. Some species have false veins consistent characteristics (at least in horticulture) essentially
that run parallel to the main veins but are not connected to without names. Because this will bring confusion to gar-
them. Unlike true veins, the false veins do not conduct deners, many of the species synonymized by Nooteboom
water or mineral nutrients; presumably they lend mechani- are still recognized in this book. Eventually, most of these
cal support to the blade tissue. The sori are close to the mar- reduced species should be given cultivar status under the
gin and covered by an indusium that may be round, scale- cultivar-group category discussed in Chapter 12.
like, shell-shaped, pouch-shaped, or more often cup- or This genus is named for Edmond Davall (1763–1798),
tubular-shaped. a Swiss plant collector. It contains about 35 species native
The two recent studies on the delineation of davalliod to the Old World tropics, mainly in Asia and the eastern
genera differ considerably. Kato (1985) recognized several Pacific.
genera whereas Nooteboom (1994) united most of them.
In this work we have followed Nooteboom, but for those Davallia canariensis (Linnaeus) Smith  13.36.4
wishing to use the genera recognized by Kato, we list their syn. Davallia portugal Horticulture
differences below. Canary Island davallia, hare’s-foot fern
Davallia, in the traditional sense, has indusia that are Semi-hardy to semi-tender
attached all the way up the sides, forming a pocket with a
A medium-sized fern with medium- to long-creeping rhi-
single opening toward the leaf margin. The cultivated spe-
zomes. Grows well under medium light or full sun in
cies that belong here are Davallia canariensis, D. denticu-
drained, moist-dry garden soil, potting mix, or uncut moss.
lata, D. divaricata, D. embolstegia, D. epiphylla, D. fejeensis,
The plants are easily cultivated but are moderately slow
D. griffithiana (though with the indusia attached only
growing. The new growth is seasonal.
slightly up the sides), D. lorrainii, D. mariesii, D. pyxidata,
Davallia canariensis is characterized by thick rhizomes
D. solida, and D. tasmanii.
up to 18 mm (0.75 in.) wide that are white-waxy beneath
Humata was separated from Davallia by its leathery
the scales. The blades are broadly triangular, about 40 cm
fronds and indusia attached only at the base, not along the
(16 in.) long and wide, and have a fleshy lip present at the
sides. The cultivated species that belong here include Da-
junction of the costa and rachis on the upper ridge. The
vallia falcinella, D. heterophylla, D. parvula, D. repens, and
indusia are cup-shaped. This species is native to Portugal,
D. tyermannii.
Spain, the Canary Islands, and Morocco.
Kato (1985) recognized several segregate genera from
Humata, such as Trogostolon and Pachypleuria. Under this
system, the genus Trogostolon includes Davallia falcinella. Davallia denticulata (N. L. Burman) Mettenius ex
The characters of Trogostolon are dark brown, peltate, Kuhn  13.36.5
toothed rhizome scales with needle-like apices; fronds syn. Davallia elegans Hedwig
finely dissected with one vein in the ultimate segments; sori Toothed davallia
at a vein fork; and round indusia attached by the base and Tender
halfway up the sides. If Kato’s findings are followed, Pachy- A medium-large fern with medium- to long-creeping rhi-
pleuria includes D. parvula and D. repens. The characters zomes. Grows well under medium light in moist-dry pot-
that define Pachypleuria are peltate, ciliate rhizome scales ting mix or uncut moss with good drainage. The plants are
with wart-like outgrowths on the lower surface; fronds easy to grow and tend to volunteer in suitable climates.
mostly small, leathery, linear to deltate, simple to bipin- The rhizome scales of Davallia denticulata are toothed
natifid, rarely finely dissected; sori at a vein fork or bend; and narrowed abruptly from a broad base. The blade seg-
and firm indusia attached at the base or slightly up the ments are oblong, often with a faint line (false vein) paral-
sides. lel to and between the true veins. The indusia are small and
Scyphularia was segregated from Davallia mainly by the cup-shaped. This species is native to the Old World tropics.
needle-like and bristly scale tips of its rhizome. Cultivated A crested form with somewhat narrower fronds is pres-
species include Davallia pentaphylla and D. triphylla, al- ently cultivated.
though the latter’s rhizome scales are not strongly needle-
like and bristly.
 F  F A  C 265

c c

d
d

a
b

b a

Figure 13.36.4. Davallia canariensis: a. frond, bar = 5 cm

(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile Figure 13.36.5. Davallia denticulata: a. frond, bar = 5
segments, bar = 5 mm (0.2 in.); d. swollen lips at rachis- cm (2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c.
costa junctions, bar = 5 mm (0.2 in.). fertile segments, bar = 5 mm (0.2 in.); d. sterile segment
with false veins, bar = 5 mm (0.2 in.).

Davallia divaricata Blume  13.36.6 scales of Davallia embolstegia are mostly narrowly triangular
Tender to lanceolate and long-acuminate. Indusia are cup-shaped
A medium-large fern with medium- to long-creeping rhi- with an elongated free lip. This species resembles D. epi-
zomes. Grows well under low to high light in moist-dry phylla but is larger and has ungrooved or only weakly
potting mix or uncut moss with good drainage. The young grooved rachises and nonpeltate rhizome scales. Davallia
fronds are red. embolstegia is native to Borneo and the Philippines.
Davallia divaricata has rhizome scales that are thin,
large, and uniformly reddish brown. The rachises and cos- Davallia epiphylla Sprengel  13.36.8
tae are reddish; the indusia, cup-shaped. This species is na- syn. Davallia denticulata var. elata (G. Forster)
tive to tropical Asia and Malaysia. Mettenius ex Kuhn
Tender
Davallia embolstegia Copeland  13.36.7 A medium-sized fern with medium- to long-creeping rhi-
Tender zomes. Grows well under medium light in drained, moist-
A medium-large fern with medium- to long-creeping rhi- dry potting mix or uncut moss.
zomes. Grows well under medium light in well-drained, Davallia epiphylla has peltate rhizome scales, and the
moist-dry potting mix or uncut moss. This species needs larger stipe scales are broadly heart-shaped at the base and
heat and humidity to grow well. It is rarely cultivated. abruptly narrowed above to a thread-like tip. The indusia
The rhizome scales attach at a notch in the cordate base are cup-shaped with an elongated free lip. This species is
(not peltate), and the segments lack false veins. The stipe native to Malaysia, New Guinea, and the Pacific Islands.
266 C 13

b c
c

b
Figure 13.36.6. Davallia divaricata: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile a
segments, bar = 5 mm (0.2 in.).
Figure 13.36.8. Davallia epiphylla: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile
e segments, bar = 5 mm (0.2 in.); d. indusia, bar = 1 mm;
c e. stipe scales, bar = 2 mm.

This species is readily separated from all the others, ex-

cept Davallia embolstegia, by the free lip on the indusium.
The rarely grown D. embolstegia is much larger, more finely
divided, and has nonpeltate rhizome scales.

d Davallia falcinella C. Presl  13.36.9

syn. Humata falcinella (C. Presl) Copeland, Trogostolon
Figure 13.36.7.
falcinellus (C. Presl) Copeland
Davallia
Very tender
embolstegia:
a. frond, bar = A small fern with medium-creeping rhizomes and finely di-
5 cm (2 in.); vided fronds. Grows best under medium light in well-
b. rhizome scales, drained, moist-dry uncut moss. The plants are sensitive to
bar = 5 mm overwatering.
(0.2 in.); c. fertile The rhizome scales of Davallia falcinella are peltate, but
segments, bar = they narrow abruptly above the base into a firm, dark
5 mm (0.2 in.); brown bristle. The blades are up to four-pinnate, and the
d. indusium, bar indusia are round and attached halfway up the sides. The
= 1 mm; e. stipe species is endemic to the Philippines.
scales, bar =
2 mm.
b a
 F  F A  C 267

c d

a
d

Figure 13.36.9. Davallia falcinella: a. habits, sterile frond

(left) and fertile frond (right), bar = 5 cm (2 in.);
b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile
b c
segments, bar = 5 mm (0.2 in.); d. rhizome with bristly
scales, bar = 5 mm (0.2 in.).

Figure 13.36.10. Davallia fejeensis: a. frond, bar = 5 cm

(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.);
c. rhizome scale hairs, bar = 1 mm; d. fertile segments,
Davallia fejeensis Hooker  13.36.10, 11
bar = 5 mm (0.2 in.).
syn. Davallia solida var. fejeensis (Hooker) Nooteboom
Fiji davallia
Tender
A small to medium fern with medium- to long-creeping ‘Plumosa’. Figure 13.36.11i. Fronds medium-sized,
rhizomes. Grows well under medium light in drained, pendulous, finely divided, the segments linear, to 0.5 mm
moist-dry potting mix or uncut moss. wide.
Davallia fejeensis produces fronds that are variable in size
and degree of dissection. The blades can be up to five-pin- Davallia griffithiana Hooker  13.36.12
nate, with linear segments supplied by a single vein. The syn. Humata griffithiana (Hooker) C. Christensen
rhizome scale margins bear long hairs. The indusia are tu- Semi-tender
bular and occupy all or nearly all of the segment width. The A small to medium fern with medium- to long-creeping
plants resemble D. solida in other details. This species is na- rhizomes. Grows well under medium light in drained,
tive to Fiji and the Austral Islands. moist-dry potting mix or uncut moss. This species is slow
This variable plant (see Figure 13.36.11) intergrades growing.
into Davallia solida (also highly variable) and is often clas- Davallia griffithiana resembles D. tyermannii but its rhi-
sified as a variety of that species (Nooteboom 1994). The zome scales are triangular rather than ovate-attenuate, and
most distinctive cultivars include: the blade is somewhat more foliaceous. In addition, the
‘Major’. Figure 13.36.11j. Fronds larger, to 60 cm (2 rounded indusia are attached a short ways up the sides, not
ft.) long, moderately finely divided; the segments oblance- at the base. Nooteboom (1994) considered D. tyermannii a
olate, 1 mm at the narrowest part. The thick rhizomes grow synonym of D. griffithiana. This species is native to north-
straight and away from the substrate. ern India, China, and Taiwan.
 268 C 13

c
b
a

Figure 13.36.11. Davallia fejeensis variations: all lower pinnae (except h, a middle pinna), all unnamed cultivars, except d.
‘Dwarf Ripple’, e. ‘False Plumosa’, i. ‘Plumosa’, and j. ‘Major’. Bar = 5 cm (2 in.).

Davallia heterophylla J. E. Smith  13.36.13 Davallia hymenophylloides (Blume) Kuhn

syn. Humata heterophylla (J. E. Smith) Desvaux  13.36.14
Tender to very tender syn. Araiostegia hymenophylloides (Blume) Copeland,
A small fern with medium-creeping rhizomes that often be- Leucostegia hymenophylloides (Blume) Beddome
come hanging and vine-like. Grows well under humid con- Tender to very tender
ditions and medium light in drained, moist potting mix or A medium-large fern with medium- to long-creeping rhi-
uncut moss. The plants are frequently used for terrariums zomes and deciduous, broad, thin, finely divided fronds.
or hanging baskets. Grows best in drained, moist potting mix or uncut moss.
Davallia heterophylla has dimorphic fronds. The sterile Davallia hymenophylloides has fleshy rhizomes covered
ones are simple, lanceolate-ovate, entire and often truncate by thin, nearly entire scales. The blades are up to four-pin-
at the base. In contrast, the fertile fronds are narrower and nate-pinnatifid, sometimes elongate and narrowing toward
lobed. Semicircular indusia are attached by a broad base. the base, with ultimate segments narrowly oblong. The in-
The species is native to India, southeastern Asia, Malaysia, dusium is kidney-shaped, to 1 mm wide, and attached at
Indonesia, the Philippines, New Guinea, and the Pacific the cordate base. This species is native to India, southeast-
Islands. ern Asia, Malaysia, Indonesia, and the Philippines.
 d

Figure 13.36.12. Davallia griffithiana: a. habit, with a variant frond on left, bar = 5 cm (2 in.); b. rhizome scales, bar =
5 mm (0.2 in.); c. fertile segments, bar = 5 mm (0.2 in.); d. indusium, bar = 1 mm.
270 C 13

Davallia lorrainii Hance  13.36.15

syn. Davallia trichomanoides var. lorrainii (Hance)
Holttum
Tender
A small to medium fern with medium- to long-creeping
Figure 13.36.13. rhizomes and deciduous fronds. Grows well under medium
Davallia light in drained, moist-dry potting mix or uncut moss.
heterophylla: Avoid overwatering during dormancy.
habit, fertile Davallia lorrainii has a strongly deciduous growth habit
frond (left) and and rhizome scales that are dark brown to black with many
two sterile even, pale hairs borne at nearly right angles to the margin.
fronds (right), The round-ovate stipe scales have pale margins bearing
bar = 5 cm hairs similar to those on the rhizome scales. Davallia lor-
(2 in.). rainii resembles D. trichomanoides Blume (not of trade),
and Nooteboom (1994) considered it a variety of that spe-
cies. It is native to Asia and the Malayan Archipelago.

Davallia mariesii T. Moore ex Baker

 13.36.1–3, 16, 17;  5
syn. Davallia bullata Hayata
Semi-hardy fern
A small to medium fern with medium- or longer creeping
rhizomes and deciduous to semi-evergreen fronds. Best
grown under medium light in drained, moist-dry garden
b soil, potting mix, or uncut moss.

Figure 13.36.15. Davallia lorrainii: a. fronds, bar = 5 cm

(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.);
Figure 13.36.14. Davallia hymenophylloides: a. frond, bar c. rhizome scale cilia, bar = 1 mm; d. stipe scales, bar =
= 5 cm (2 in.); b. fertile segments, bar = 2 mm. 2 mm.
 F  F A  C 271

e d

c
b

a
a

Figure 13.36.16. Davallia mariesii: a. habit and larger

frond (left), bar = 5 cm (2 in.); b. rhizome scales, bar =
5 mm (0.2 in.); c. rhizome scale cilia, bar = 1 mm; b
d. fertile segments, bar = 5 mm (0.2 in.); e. stipe scales,
bar = 2 mm.

Davallia mariesii has triangular fronds up to three-pin- Figure 13.36.17. Davallia mariesii var. stenolepis: a.
nate at the base, with stipe scales triangular to ovate or ob- habit, bar = 5 cm (2 in.); b. rhizome scales, bar = 5 mm
long, mostly narrowing abruptly to a long, twisted apex. (0.2 in.); c. rhizome scale cilia, bar = 2 mm; d. fertile
The whitish tan to brown rhizome scales are lightly pressed segments, bar = 5 mm (0.2 in.); e. stipe scales, bar =
to the rhizome, and the margins bear short cilia and teeth. 2 mm.
The indusia are short and tubular. This species is native to
Korea, Japan, Taiwan, and China.
Nooteboom (1994) considered Davallia mariesii to be
the same as D. trichomanoides var. trichomanoides. The foot fern. Figures 13.36.1–3, 17. Fronds to about 20 cm (8
plants called D. mariesii in horticulture, however, are dis- in.) long, nearly evergreen or with a short dormant period
tinct and important enough in the trade to require main- between shedding of the old fronds and emergence of the
taining the name. new ones. The pinnae and pinnules do not noticeably over-
var. mariesii. Japanese ball fern. Figure 13.36.16. Fronds lap. Variety stenolepis resembles var. mariesii but is only
to 15 cm (6 in.) long, triangular, deciduous, the pinnae slightly deciduous, larger, and has less crowded, more at-
closely placed, and the pinnules sometimes overlapping. tenuate pinnae. Native to the Ryuku Islands, South Korea,
This variety intergrades with var. stenolepis. Native to Korea China, and Taiwan. The variety is easily cultivated but less
and Japan. In Japan the live rhizomes are gathered when cold hardy than var. mariesii. It is the most widely grown
the plant is in its leafless period and tied into balls or other Davallia in the United States and is usually sold, incor-
figures and subsequently hung out to grow as a basket fern. rectly, as D. trichomanoides. Most Japanese botanists con-
var. stenolepis (Hayata) Hoshizaki (Davallia stenolepis sider var. stenolepis merely a semi-evergreen, southern vari-
Hayata, D. trichomanoides of trade, not Blume). Squirrel’s- ant of D. mariesii.
 272 C 13

Davallia parvula Wallich ex Hooker & Greville c

 13.36.18
syn. Humata parvula (Wallich) Mettenius, Pachypleuria
parvula (Wallich) M. Kato
Tender
Very small, with medium-creeping rhizomes and finely di-
vided, triangular blades. Grows well under low-medium
light in drained, moist potting mix or uncut moss. The
plants need humidity and are best used in terrariums. They
are easily harmed by too much fertilizer.
The most distinctive characteristic of Davallia parvula is
its small size: the fronds are only to 2.5 cm (1 in.) long and a
broad, with segments less than 1 mm wide. The indusia are
rounded and attached at the base. This species is native
from Malaysia and Indonesia to New Guinea.

b b

a Figure 13.36.19. Davallia pentaphylla: a. habits, with the

former D. pycnocarpa at lower left, bar = 5 cm (2 in.);
b. rhizome and scales, bar = 5 mm (0.2 in.); c. sori and
indusia, bar = 5 mm (0.2 in.).
Figure 13.36.18. Davallia parvula: a. habit and frond,
bar = 5 cm (2 in.); b. fertile segments, bar = 5 mm
(0.2 in.).
Davallia pulchra D. Don  13.36.20
syn. Araiostegia pulchra (D. Don) Copeland
Davallia pentaphylla Blume  13.36.19 Tender
syn. Davallia pycnocarpa Brackenridge, Scyphularia A small-medium fern with medium- to long-creeping rhi-
pentaphylla (Blume) Fée, S. pycnocarpa (Brackenridge) zomes. Grows best under medium light in well-drained,
Copeland, S. sinusora Copeland moist-dry potting mix or uncut moss.
Tender Davallia pulchra resembles D. mariesii var. stenolepis but
A small fern with medium- to long-creeping rhizomes. has more-delicate blades and bean-shaped or rounded in-
Grows best under medium light in well-drained, moist-dry dusia attached only at the base. It is native from India to
potting mix or uncut moss. This species is easy to grow. Tibet, southeastern Asia, and southern China.
Davallia pentaphylla is distinctive for its one-pinnate
blades, typically with five leaflets, and a terminal leaflet that Davallia pyxidata Cavanilles  13.36.21
resembles the lateral ones. The leaflets, particularly the basal syn. Davallia solida var. pyxidata (Cavanilles)
pair, are sometimes lobed or forked. The rhizome scales are Nooteboom
also distinctive by their widely spreading habit and black, Australian davallia, Australian hare’s-foot fern
needle-like tips, which imparts a bristly appearance. The Semi-hardy to semi-tender, Zones 8–9
indusia are tubular. This species is native to Malaysia, In- A small-medium fern with medium- to long-creeping rhi-
donesia, New Guinea, and the Pacific Islands. zomes. Best grown under medium-high light in well-
Nooteboom (1994) did not recognize Scyphularia pyc- drained, moist-dry potting mix or uncut moss. The plants
nocarpa, a variant that consistently produces lobed or are sometimes used as bedding plants. The stiff, upward-
forked lower leaflets (see Figure 13.36.19 lower left). growing rhizomes do not make for a good basket plant.
 F  F A  C 273

d
c
e

b
a
b
a

Figure 13.36.21. Davallia pyxidata: a. frond, bar = 5 cm

Figure 13.36.20. Davallia pulchra: a. habit, bar = 5 cm (2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c.
(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile ascending rhizome, bar = 10 cm (4 in.); d. fertile
segment, bar = 5 mm (0.2 in.); d. sori and indusia, bar = segments, bar = 5 mm (0.2 in.); e. apex of sterile
1 mm. pinnule, bar = 5 mm (0.2 in.).

The rhizomes of Davallia pyxidata tend to grow stiffly Davallia repens (Linnaeus f.) Kuhn  13.36.22
upward and are covered by easily detached, nearly black syn. Humata repens (Linnaeus f.) Diels, H. vestita
scales with tapered apices and long-hairy margins. The (Blume) T. Moore, Pachypleuria repens (Linnaeus f.)
stipes contain four (sometimes three or five) vascular bun- M. Kato
dles. The secondary pinnules are mostly pointed, with the Semi-tender to tender
apex ending in a dominant tooth. The indusia are short A small fern with medium-creeping rhizomes. Grows best
and tubular. This species is native to Australia. under warm, humid conditions, preferring medium light
Nooteboom (1994) classified Davallia pyxidata as a va- and moist-dry, drained potting mix or uncut moss.
riety of D. solida; however, D. solida var. solida differs most The rhizomes of Davallia repens are white-waxy beneath
conspicuously by having rhizomes that generally do not the scales. The blades are triangular and deeply pinnately
grow stiffly upward. Other differences are the lighter-col- lobed to two-pinnate at the base, with the lower lobes
ored rhizome scales, which are difficult to detach whole; broader and lobed on the lower margin. The indusium is
stipes with six (three to nine) vascular bundles; more leath- semi-circular and attached only at the base. This species is
ery and glossy blades; more obtuse secondary pinnules with native from northern India to Japan, and the Mascarene
apices not ending in a dominant tooth; more obscure veins; Islands to Australia.
and longer indusia. Furthermore, D. pyxidata is hardier Nooteboom (1994) pointed out the tremendous varia-
than the tender D. solida. Given these differences, D. pyxi- tion in the group and reduced Davallia vestita and other
data merits recognition as a distinct species. species to D. repens.
 274 C 13

Figure 13.36.22. Davallia repens: a. habit and frond

variations, with the former D. vestita at far right, bar =
5 cm (2 in.); b. fertile segment, bar = 5 mm (0.2 in.).

Davallia solida (G. Forster) Swartz

 13.36.23, 24 b
Polynesian davallia
Tender
Figure 13.36.23. Davallia solida: a. frond, bar = 5 cm
A small to medium fern with medium- or longer creeping (2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. fertile
rhizomes. Grows well under medium light in potting mix segments, bar = 5 mm (0.2 in.); d. stipe scales, bar =
or uncut moss kept moist-dry and given good drainage. 2 mm.
The blades of Davallia solida are coarsely cut and varia-
ble in size and shape. The rhizome scales are long-hairy on
the margin and difficult to detach whole. The stipe scales
vary greatly in shape, and their margins may have a sparse
tangle of hairs. Sometimes the scales are so reduced that they Figure 13.36.24.
appear as a fine scurf. The segment veins end short of the Davallia solida
margin. The indusia are cup-shaped, longer than wide. This ‘Ruffled Ornata’:
species is native to Asia, Malaysia, Australia, and Polynesia. frond, bar = 5 cm
Many cultivated variants circulate under a variety of (2 in.).
names, and often these are difficult to distinguish, except
for the following:
‘Ruffled Ornata’. Figure 13.36.24. Segments broad,
margins ruffled, tips drooping.

Davallia tasmanii Field  13.36.25

Tasman’s davallia
Semi-tender
A small fern with medium-creeping rhizomes. Grows well
under medium light in drained, moist-dry potting mix or
uncut moss. The plants are slow growing and are rarely cul-
tivated.
The rhizome scales of Davallia tasmanii bear shaggy
hairs, and the blades are often pentagonal and leathery. The
veins are obscure, and the indusia are cup-shaped. This spe-
cies is endemic to New Zealand.
 F  F A  C 275

c d
e

b
a

a c

Figure 13.36.25. Davallia tasmanii: a. fronds, bar = 5

cm (2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c.
fertile segments, bar = 5 mm (0.2 in.). Figure 13.36.26. Davallia trichomanoides: a. habit, bar =
5 cm (2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c.
rhizome scale cilia, bar = 1 mm; d. fertile segments, bar
= 5 mm (0.2 in.); e. stipe scales, bar = 2 mm.
Davallia trichomanoides Blume  13.36.26
Squirrel’s-foot fern
Semi-hardy to semi-tender
A small to medium fern with medium- or long-creeping Davallia triphylla Hooker  13.36.27
rhizomes. Grows well under low to high light in drained, syn. Scyphularia simplicifolia Copeland, S. triphylla
moist-dry potting mix or uncut moss. For trade material (Hooker) Fée
under the name Davallia trichomanoides, see D. mariesii Tender
var. stenolepis. A small fern with medium- to long-creeping rhizomes. Best
The reddish brown rhizome scales of Davallia tri- grown under low to high light in drained, moist-dry pot-
chomanoides spread widely or curl backward, imparting a ting mix or uncut moss. The plants grow slowly. They in-
ruffled appearance. The stipe scales are ovate-triangular, frequently produce fronds with three leaflets.
long-attenuate, and ciliate. The pinnae are long-attenuate, A distinctive characteristic of Davallia triphylla is the
and the indusia are cup-shaped. This species is native to flattened rhizome covered by tightly appressed scales, the
Malaysia, New Guinea, and Java. The plants of D. tricho- margins of which bear long, white hairs. The older scales
manoides in cultivation came from New Guinea. tend to turn black. The fronds are simple and lanceolate, or
Nooteboom (1994) broadly construed Davallia tri- trifoliate with entire leaflets. The indusia are longer than
chomanoides to include D. lorrainii, D. mariesii var. mari- they are wide and stoutly tubular but slightly tapered. The
esii, D. mariesii var. stenolepis, and others. Davallia trichom- species is native to Malaysia and Indonesia.
anoides resembles the common trade plant D. mariesii var.
stenolepis but has larger, softer, more finely cut fronds and
long-tapering pinna apices. The rhizome scales of D. tri-
chomanoides also differ by being red-brown (not whitish
brown) and widely spreading (not pointing forward).
 276 C 13

a
d

b
a

Figure 13.36.27. Davallia triphylla: a. habit, bar = 5 cm

(2 in.); b. rhizome scales, bar = 5 mm (0.2 in.); c. sori
and indusia, bar = 5 mm (0.2 in.).
Figure 13.36.28. Davallia tyermannii: a. habits, with
dwarf form at upper left, bar = 5 cm (2 in.); b. rhizome
scales, bar = 5 mm (0.2 in.); c. fertile segments, bar =
Davallia tyermannii (T. Moore) Hooker & Baker 5 mm (0.2 in.); d. indusium, bar = 1 mm.
 13.36.28
syn. Humata tyermannii T. Moore
Bear’s-foot fern, white rabbit’s-foot fern
Semi-hardy Davallodes
A small to medium fern with medium or longer creeping Davallodes plants are medium-sized epiphytes with elliptic
rhizomes. Best grown under medium to high light in well- blades. They are mainly seen outdoors in Florida as basket
drained, moist-dry garden soil or potting mix or uncut ferns or in the greenhouses of fern collectors. Rarely they
moss. The species is easy to grow (although somewhat slow are offered in specialty catalogs.
growing) and extremely variable. It is a popular houseplant. Davallodes differs from the similar Davallia by having
Davallia tyermannii is characterized by whitish rhizome catadromous blades (at least on the larger pinnae) some-
scales and triangular blades up to three-pinnate-pinnatifid. what to strongly narrowed toward the base, not anadro-
The smallest segments are dilated. Roundish-oval indusia mous and widest at the base as in Davallia. Furthermore,
are attached only at the base. The species is native to China. the rachises, costae, and often the blade tissue are hairy,
Nooteboom (1994) considered Davallia tyermannii a whereas Davallia is glabrous at maturity. Davallodes has
synonym of D. griffithiana. These two species are very close, creeping, fleshy rhizomes covered by needle-like, peltate
separated mainly by the indusia being slightly attached on scales. The roots are produced on the underside of the rhi-
the sides in D. griffithiana and only at the base in D. tyer- zome at the nodes, not along the internodes. The stipes are
mannii. The great variation, particularly in D. tyermannii, jointed to the rhizome. The blades vary from one-pinnate-
needs further study. Crested and dwarf forms of D. tyer- pinnatifid to two-pinnate-pinnatifid, but they are always
mannii are prized by gardeners. thin. Sori occur mostly at the vein fork and are covered by
 F  F A  C 277

small, inconspicuous indusia that are either scale-like, reni-

form, or pouch-shaped. The attachment of the indusium
varies from basal to half-way up the sides.
Davallodes consists of about 10 species (only 7 were rec-
ognized by Nooteboom 1994) that occur in wet forests
from Sumatra to the Philippines and New Guinea. The
genus name means “Davallia-like,” adding the Greek suffix
-odes, meaning like, to Davallia.

Davallodes hirsutum (J. Smith ex C. Presl) Copeland

 13.37.1
Tender
A medium-sized fern with medium- to long-creeping rhi-
zomes. Grows well under medium light in drained, moist
potting mix or uncut moss. The plants are easy to grow if
given sufficient warmth and humidity throughout the year.
A distinctive characteristic of Davallodes hirsutum is the
Figure 13.38.1. Dennstaedtia davallioides: habit.
spreading, needle-like tips of the blackened rhizome scales.
The blades are hairy and lanceolate to narrowly elliptic.
The indusia are small and inconspicuous, about 0.5 mm
The wide-creeping habit makes Dennstaedtia unsuitable as
wide. The species is native to Malaysia, the Philippines, Su-
a pot plant. The rhizomes branch readily, and their growth
lawesi, and New Guinea.
can be controlled by cutting away unwanted parts of the
rhizome and foliage.
Dennstaedtia  13.38.1–3
The genus is characterized by creeping, hairy (not scaly)
Cup fern
rhizomes, stipes containing a single U-shaped vascular bun-
The cup ferns are medium- to large-sized terrestrial ferns dle, and cup-shaped indusia borne at the margins of the
with creeping rhizomes and finely divided foliage. They blades. The fronds vary from medium-sized to huge, usually
may be used in foundation plantings or as bedding ferns.

Figure 13.37.1. Davallodes

hirsutum: a. habit, bar =
5 cm (2 in.); b. fertile
segment, bar = 2 mm.
After Hoshizaki (1981).

a
Figure 13.38.2. Dennstaedtia davallioides:
segments bearing sori.
 278 C 13

Figure 13.38.3. Dennstaedtia bipinnata: sori and cup-

shaped indusia.

two- to three-pinnate and broadest at the base. The veins

are free, not netted. The sori are usually turned slightly
downward. Many species of Dennstaedtia bear branch-buds
on the stipe base. These buds, called “epipetiolar buds,” de-
velop into creeping stems. They are not always present, but
if so, they can be up to three or four per stipe. Figure 13.38.4.
The genus is native to the tropics and subtropics of the Dennstaedtia bipinnata:
world and contains about 45 species. It is named for August frond, bar = 5 cm
Wilhelm Dennstaedt (1776–1826), a German botanist. (2 in.).

Dennstaedtia bipinnata (Cavanilles) Maxon

 13.38.3, 4
Couplet fern
Tender, Zone 9
A medium-sized fern with long-creeping rhizomes. Prefers
medium light in moist garden soil. This species does best in
a climate with warm nights.
Dennstaedtia bipinnata can be distinguished from other
cultivated species in the genus by the form of the upper
sides of the rachises, costae, and costules. These axes are
grooved, and the grooves of the costules run into those of
the costae, which in turn run into that of the rachis. Fur-
thermore, the grooves of the pinnules are bordered by an
erect wing of green tissue. The leaf axes are ungrooved in
other Dennstaedtia species or, if grooved, the grooves do
not run into each other or have the green wings. Other
characteristics of this species are triangular, glossy blades
that are glabrous except for dark hairs present on the rachis
opposite the larger pinnae. Epipetiolar buds are absent. It is
native to the American tropics and southern Florida.

Dennstaedtia cicutaria (Swartz) T. Moore

 13.38.5
Semi-tender Figure 13.38.5. Dennstaedtia
A medium-large fern with long-creeping rhizomes. Grows cicutaria: frond, bar = 5 cm
well under medium light in moist garden soil. (2 in.).
 F  F A  C 279

Dennstaedtia cicutaria can be distinguished from the Dennstaedtia punctilobula (Michaux) T. Moore
other cultivated Dennstaedtia by the numerous needle-  13.38.7
shaped hairs on the lower surfaces of the blades. The rhi- Hay-scented fern
zomes are 12 mm (0.5 in.) thick or more, and branch-buds Very hardy, Zone 3
arise near the stipe bases. The triangular blades have dull A medium-sized fern with long-creeping rhizomes and de-
green surfaces and are densely hairy within the rachis ciduous fronds. Prefers low to high light in moist, acidic
grooves. The species is native to the American tropics. garden soil. This species spreads well in gardens, sometimes
becoming a weed, but it is easy to control by pulling up the
Dennstaedtia davallioides (R. Brown) T. Moore shallow rhizomes. New fronds are produced throughout
 13.38.1, 2, 6 the growing season. When crushed, the fronds emit an odor
Lacy ground fern of freshly cut hay. The substance that produces the odor is
Semi-hardy to semi-tender contained within the enlarged, glandular cell of the hair
A medium-large fern with long-creeping rhizomes. Grows covering the blades.
well under medium light in moist or moist-dry garden soil. The blades of Dennstaedtia punctilobula are narrowly
The rhizomes of Dennstaedtia davallioides grow to about triangular to lanceolate and two-pinnate-pinnatifid, with
6 mm (0.2 in.) thick, and the blades are triangular, me- gland-tipped hairs on the lower surface. Epipetiolar buds
dium green, with hairs more or less evenly distributed over frequently occur at the stipe bases. The species is native to
the rachis. The species is native to Australia. eastern North America.

Figure 13.38.7.
Dennstaedtia punctilobula:
frond, bar = 5 cm (2 in.).

Deparia  13.39.1

syn. Athyrium in part, Diplazium in part, Dryoathyrium,
Lunathyrium
These terrestrial ferns are related to the lady ferns of genus
Athyrium. They are easy to cultivate and can be used in beds
or for background foliage in warm-temperate areas. The
cultivated species of Deparia grow readily from spores.
Deparia consists of mostly medium-sized ferns with
Figure 13.38.6. Dennstaedtia davallioides: habit, bar = erect or short- to long-creeping rhizomes. The scales are
5 cm (2 in.). not clathrate, in contrast to those of Asplenium, a genus
280 C 13

Deparia acrostichoides (Swartz) M. Kato

 13.39.2
syn. Athyrium thelypteroides (Michaux) Desvaux,
Diplazium acrostichoides (Swartz) Butters
Silver glade fern, silvery spleenwort
Very hardy, Zone (3)4
A medium-large fern with short-creeping rhizomes and de-
ciduous fronds. Grows well under medium light in moist
garden soil or potting mix.
Deparia acrostichoides has swollen stipe bases, blades
one-pinnate-pinnatifid, broadly lanceolate, noticeably nar-
rowed below, and silvery on the lower surface. The indusia
are linear or hooked. The species is native to the southeast-
ern United States.

Deparia japonica (Thunberg) M. Kato

 13.39.3
syn. Athyrium japonicum (Thunberg) Copeland
Hardy, Zone 6
a A medium-sized fern with deciduous fronds from a short-
creeping, branched rhizome. Grows well under medium
b light in moist-wet garden soil or potting mix. The plants
tend to volunteer.
The sterile blades of most cultivated plants are triangu-
lar, one-pinnate-pinnatifid, and sometimes blackish green.
Figure 13.39.1. Deparia: a. segment with typical indusia;
The fertile fronds are taller, narrower, and more erect than
b. rachis and costae grooves not continuous.
the sterile ones. The indusia are linear with laciniate mar-
gins and are arranged in a herringbone pattern. This species
is native from the Himalayas to China, Korea, and Japan,
with which Deparia could be confused. The stipes contain where it favors warm-temperate or temperate climates.
two vascular bundles at the base. The blades vary from one-
pinnate to two-pinnate-pinnatifid. They are at least partly
anadromous, and the costal grooves are interrupted at the
juncture with the rachis—that is, they do not run into the
groove of the rachis. The hairs of the rachises and costae
are multicellular and jointed. The veins are free (not net-
ted), and the sori vary from round to U- or J-shaped to lin-
ear. Sometimes the sori are paired, as in Diplazium.
Deparia species were formerly included in either Athyr-
ium or Diplazium. The easiest way to distinguish this genus
is by the multicellular hairs on the fronds (Athyrium and
Diplazium lack such hairs). Furthermore, the costal grooves Figure 13.39.2. Deparia
in Deparia do not run into the rachis groove, whereas in acrostichoides: frond, bar =
Athyrium and Diplazium the grooves are continuous. This 5 cm (2 in.).
characteristic is usually best developed in the middle of the
blade. For further comparison, see Athyrium; for a techni-
cal treatment of many Deparia species, see Kato (1984).
Deparia contains about 40 species native mostly in the
Old World east to Hawaii, North America, and are natu-
ralized in Europe and tropical America. The genus name
comes from the Greek depas, cup or beaker, alluding to the
shape of the indusium in certain species.
 F  F A  C 281

Figure 13.39.3. Deparia japonica:

frond, bar = 5 cm (2 in.).

Deparia japonica is related to D. petersenii and is con-

sidered the same by some authors. It is said to differ from
D. petersenii by its gradually acuminate, serrate blade and a b
pinnae, oblique serrate-toothed segment, and incurved in-
dusia. Some cultivated material is difficult to distinguish.

Deparia petersenii (Kunze) M. Kato  13.39.4

syn. Athyrium petersenii (Kunze) Copeland Figure 13.39.4. Deparia petersenii: a. frond, bar = 5 cm
Semi-hardy or hardier, Zone 8 (2 in.); b. pinna with elongate sori and indusia, bar =
1 cm (0.4 in.).
A medium-sized fern with a short-creeping, branched rhi-
zome. Grows well under medium light in moist garden soil
or potting mix. This species is easy to grow, robust, and
tends to volunteer. The fiddleheads are reportedly edible. southwestern England. The tree-fern trunk fibers used in
In cultivated material, the blades of Deparia petersenii horticulture come from D. sellowiana of Latin America and
vary from triangular to ovate-lanceolate and are up to two- should not be confused with the hapu fiber, which come
pinnate, with the basiscopic pinnule next to the rachis re- from Cibotium glaucum of Hawaii.
duced to a somewhat angular lobe. The indusia are linear All dicksonias have trunk-like stems that are clothed,
with laciniate margins and arranged in a herringbone pat- along with the stipe bases, by bristly hairs. The old, dead
tern. This species is extremely variable and is closely related fronds usually persist on the trunk and hang downward,
to D. japonica. It favors subtropical and tropical climates in forming a “skirt.” The blades vary from two-pinnate-pin-
Asia and Australia and has been introduced in the Pacific Is- natifid to four-pinnate and are oblong or lanceolate, nar-
lands and the southeastern United States. rowed at the base, and have a stiff, leathery texture. The
sori are borne at the margins of the blades and appear clam-
Dicksonia  13.40.1, 2 like because of the two-valved indusium. The outer lobe of
Tree fern the indusium is formed from the enrolled leaf margin and
The genus Dicksonia consists of large tree ferns with nar- is greenish, resembling the leaf tissue in color and texture.
row, leathery fronds. The Tasmanian dicksonia (Dicksonia The inner lobe, however, is brownish and thinner.
antarctica) and Cooper’s tree fern (Sphaeropteris cooperi ) Dicksonia is readily separated from the cyatheoid tree
from Australia are the most commonly cultivated tree ferns ferns by having hairs rather than scales on the trunk and
in the United States. Both species are grown outdoors in stipe and by sori borne on the blade margins, not the lower
Florida and in coastal California at least as far north as San surfaces. The related genus Cibotium differs from Dicksonia
Francisco. A handsome grove of Tasmanian tree ferns can by its broadly triangular blades with slightly glaucous lower
be seen at Golden Gate Park in San Francisco. Tasmanian surfaces, and by the inner and outer indusial lobes being of
dicksonias also thrive in estate gardens in mild parts of similar color and texture.
 282 C 13

Figure 13.40.3.
Dicksonia antarctica:
a. pinna, bar = 5 cm
(2 in.); b. primary
pinnule, bar = 1 cm
(0.4 in.).

Figure 13.40.1. Dicksonia antarctica: habit.

Figure 13.40.2. Dicksonia antarctica: sori and indusia.

or greenish rachises and costae. The blade is harsh and stiff,
with central pinnae three to four times longer than broad
and central pinnules long and acute to acuminate. It is na-
The name Dicksonia comes from James Dickson (1738– tive to Australia and Tasmania.
1822), a Scottish physician and cryptogamic botanist. The Cold weather easily damages or kills the fronds but not
genus contains about 20 species and occurs mostly in wet the trunks. Several nurseries in California reported that the
montane forests (especially cloud forests) of the tropics and frond tips were burnt when the temperature dropped to
warm-temperate regions of the world. For more informa- −7°C (19°F), and the fronds died completely when the
tion about these plants, see the section in Chapter 10 on temperature stayed this low for one week. The trunks, how-
“Tree Ferns.” ever, were not killed and soon recovered to produce new
fronds. Dormant trunks in shipments have been known to
Dicksonia antarctica Labillardière survive −5°C (23°F) for two months. English gardeners re-
 13.40.1–3;  3 port that the plants do not survive if the frost days number
Tasmanian dicksonia 20–40 per season.
Semi-hardy to semi-tender
A large tree fern, with a trunk up to about 6 m (20 ft.) tall. Dicksonia fibrosa Colenso  13.40.4
Grows well under medium light or (if not too intense) full Woolly tree fern, fibrous dicksonia
sun in moist garden soil or potting mix. This species is easy Semi-tender to semi-hardy; frost hardy
to grow and is by far the most common Dicksonia in the A large tree fern, to 6 m (20 ft.) or more tall. Grows under
trade. The plants generally grow best in areas with cool, medium light or direct sun (if not too intense) in moist gar-
humid climates year-round, without dry, cold winds. den soil or potting mix. The plants prefer cool, humid cli-
Dicksonia antarctica is characterized by dark brown mates, are rarely grown, and reportedly are susceptible to
trunks, stipes that soon loose their covering of hair, and tan armillaria rot.
 F  F A  C 283

Figure 13.40.4.
Dicksonia fibrosa:
a. pinna, bar = 5 cm
(2 in.); b. primary
pinnule, bar = 1 cm b
(0.4 in.).

Dicksonia fibrosa has reddish brown trunks. The stipes

retain some of their hairs, and the rachises and costae are
tan or greenish. The blades are smaller, glossier, and more
prickly than those of D. antarctica. The central pinnae are
five to six times longer than broad, and the central pinnules a
short-acute. This species is endemic to New Zealand. b

Dicksonia sellowiana Hooker  13.40.5 Figure 13.40.5. Dicksonia sellowiana: a. pinna, bar =
syn. Dicksonia gigantea H. Karsten 5 cm (2 in.); b. primary pinnule, bar = 1 cm (0.4 in.);
Semi-tender to slightly hardier c. sori and indusia, bar = 1 mm.
A large tree fern, with trunks to 6 m (20 ft.) tall in its native
habitat. Grows best under medium light in moist garden
soil or potting mix. This species is particularly attractive for
its dark, glossy fronds, which are generally broader and and sculpted into animals such as ducks, swans, squirrels,
glossier than those of Dicksonia antarctica. In the coastal and goats. These sculptures are used to adorn lawns and
areas of central and southern California this species can be patios, much like cement statuary is used in the United
grown year-round outdoors without difficulty. States. The Mexicans call this type of sculpture maquique.
The trunk of Dicksonia sellowiana tends to be thickest at Segments of the trunk are also carved into pots for growing
the base because of the wide, dense layer of black rootlets. plants or sawed into squares and used as a substrate for
The fronds are widest above the middle and taper toward growing epiphytic orchids. In nature, many ferns and other
the base to a short stipe. The trunk apex and stipes are cov- epiphytes grow on the root mantles, some species being al-
ered with long hairs. The blades are two- to three-pinnate- most entirely restricted to it.
pinnatifid, up to 3 m (10 ft.) long, and slightly leathery. The similar Dicksonia berteriana (Colla) C. Christensen,
The pinnae are sessile, and the ultimate fertile pinnules are an endemic to the Juan Fernández Islands, is cultivated in
narrowly oblong, each bearing six or more sori. This species a few special collections. It differs from D. sellowiana by
is widespread in the American tropics, where it generally the fertile segments lacking a leafy apex or having only a
grows at elevations above 2000 m (6500 ft.). small one.
In Latin America (particularly Mexico and Central
America) the trunks of Dicksonia sellowiana are harvested
 284 C 13

Dicksonia squarrosa (Forster f.) Swartz (many irregular areoles that mostly lack included free vein-
 13.40.6 lets). Lepisorus further differs by the presence of peltate
Slender tree fern, rough tree fern, New Zealand paraphyses, at least on young sori. See Polypodium for a fur-
dicksonia ther discussion of related genera.
Semi-tender; tolerates light frost Only two species are found in Dictymia, occurring from
A large tree fern, 5–6 m (16–20 ft.) tall. Grows best under eastern Australia to Fiji. The genus name comes from the
medium light in moist to moist-wet garden soil or potting Greek diktyon, net, alluding to the netted veins.
mix. This species prefers cool, humid climates. It tends to
form offshoots from stolons and can be propagated by re- Dictymia brownii (Wikström) Copeland
moving the offshoots.  13.41.1
The trunk of Dicksonia squarrosa is slender and black, Semi-tender (?)
and the stipes are rough with stiff hairs when young. The A small-medium fern with creeping rhizomes. Grows un-
rachises and costae vary from red-brown to blackish, and der medium light in moist potting mix.
the undersides of the blades are glaucous green. This species Dictymia brownii has stiff, leathery, and erect fronds.
is smaller than D. antarctica. It is endemic to New Zealand. The sori are large, round or oblong, and raised. The species
is native to eastern Australia.

a a
b

Figure 13.41.1. Dictymia brownii: a. habit, bar = 5 cm

(2 in.); b. sori and veins, bar = 1 cm (0.4 in.).
Figure 13.40.6. Dicksonia squarrosa: a. frond (from a
young plant), bar = 5 cm (2 in.); b. pinnae, bar = 5 cm
(2 in.); c. primary pinnule, bar = 1 cm (0.4 in.).
Didymochlaena
Mahogany fern
Dictymia The mahogany ferns are medium- to large-sized plants with
These small- to medium-sized ferns are suitable for use in erect rhizomes and fronds that form a basket-shaped whorl.
pots, baskets, or rock gardens. The rhizomes of Dictymia The new fronds are a striking reddish brown (hence the
plants are creeping and scaly, as is typical of polypodiaceous common name), and the rectangular segments and horse-
ferns. The fronds vary from linear to lanceolate and are al- shoe-shaped sori give the plants added interest. They are
ways dark green, glabrous, and leathery, with entire or wavy sold as pot plants for indoor use but can be grown outdoors
margins. Netted veins are immersed in the blade tissue, in warmer climates.
with many irregular areoles usually without included vein- The genus is characterized by two-pinnate fronds with a
lets. The sori are large, round to elliptic or narrowly ob- terminal pinna resembling the lateral ones. The segments
long, and lack indusia. are slightly leathery and rectangular. A minute spine occurs
Dictymia can be confused with Polypodium, Microsorum, at the bases of the segment stalks on the upper surfaces of
or Lepisorus but differs from those genera by its venation large, mature fronds. The sori are horseshoe-shaped.
 F  F A  C 285

Didymochlaena is generally thought to consist of one ex-

tremely variable species occurring throughout the tropics of
the world. The genus name comes from the Greek didy-
mos, double, and chlaina, cloak, apparently referring to the
indusium that wraps around to cover both sides of the vein.

Didymochlaena truncatula (Swartz) J. Smith

 13.42.1, 2
Semi-tender
A medium to large fern with erect rhizomes and clustered
fronds. Grows well under medium light in moist garden
soil or potting mix. Didymochlaena truncatula is easily
grown and sometimes forms side crowns that can be sepa-
rated for propagation. Older plants tend to topple unless re-
planted deeper or mulched at the base.

b
a

Figure 13.42.2. Didymochlaena truncatula: a. immature

frond, bar = 5 cm (2 in.); b. pinnules with sori and
indusia, bar = 1 cm. (0.4 in.).

Diplazium  13.43.1–3

syn. Allantodia
The small species of Diplazium are useful in borders,
whereas the larger ones are grown in beds as background fo-
liage or in pots. Generally the best growth is achieved under
constant moist, cool, humid conditions; however, a few
species are adapted to warm- or cold-temperate climates.
All species of Diplazium are terrestrial (never epiphytic),
with rhizomes varying from long-creeping to erect. The
rhizome scales are not clathrate as in Asplenium, a genus
that has a similar soral shape and might be confused with
Diplazium. The stipes contain two vascular bundles at the

Figure 13.42.1. Didymochlaena truncatula: mature

frond. Figure 13.43.1. Diplazium tomitaroanum: habit.
 286 C 13

Figure 13.43.2. Diplazium tomitaroanum: indusia.

base. The blades vary from simple and entire to four-pin-

nate-pinnatifid. The rachis groove is generally U-shaped,
continuous with the grooves of the costae, and typically
glabrous within. The veins can be either free or netted, de-
pending on the species. The sori are indusiate, linear, and
borne either singly or paired back-to-back along a vein; the
paired sori are most frequent on the basal veins.
Diplazium species are sometimes included in Athyrium
or Deparia. For a comparison of these three genera, see the
discussion under Athyrium. The cultivated species of Dipla-
zium tend to have more coarsely cut foliage than Athyrium.
The genus consists of about 400 species native to trop-
ical regions throughout the world. Its name comes from
the Greek diplazios, double, referring to the sori that some-
times occur on both sides of a vein.

Diplazium australe (R. Brown) N. A. Wake

Figure 13.43.4. Diplazium australe: frond, bar = 5 cm
 13.43.4
(2 in.).
Austral lady fern
Semi-hardy, Zone 8
A medium-large fern with short-creeping to suberect,
well under low to medium light in moist garden soil or pot-
clump-forming rhizomes and deciduous fronds. Grows
ting mix. This species is easy to grow.
Diplazium australe is characterized by broadly ovate or
triangular, three-pinnate-pinnatifid blades. The sori are
only 2–3 mm long, crescent-shaped, and only occasionally
a are they paired back-to-back along the veins. The species is
native to Australia and New Zealand.

b Diplazium dilatatum Blume  13.43.5

Semi-tender or somewhat hardier
A medium to large fern with short-creeping to suberect,
branched rhizomes. Grows well under medium light in
moist garden soil or potting soil. The plants are easily
grown from spores.
Diplazium dilatatum has large, two-pinnate fronds with
minute, stalked, transparent to reddish glands on the un-
dersurface. The scales of the rhizomes and stipe bases have
black margins and forked teeth (use hand lens). It is native
to southern Asia, Malaysia, Australia, and New Guinea.
Figure 13.43.3. Diplazium: a. sori and indusia; b. U- Diplazium maximum (D. Don) C. Christensen is closely
shaped rachis groove continuous with costa groove. related to this species, if not the same.
 F  F A  C 287

Figure 13.43.5.
Diplazium
dilatatum: frond,
bar = 5 cm (2 in.).

Figure 13.43.6.
Diplazium esculentum:
frond, bar = 5 cm (2 in.).
Diplazium esculentum (Retzius) Swartz
 13.43.6
syn. Callipteris esculenta (Retzius) J. Small
Vegetable fern
Semi-tender, Zone 9 pinnatifid above. The species is native to the southern
A medium-sized fern, to 60 cm (2 ft.) tall, with erect rhi- United States (Louisiana) and Mexico to northern South
zomes that can become trunk-like on older plants. Grows America.
well under medium to high light in moist-wet garden soil
or potting mix. Colonies of this species resemble a grove of Diplazium proliferum (Lamarck) Thouars
short-stemmed, coarse-leaved tree ferns. The fiddleheads  13.43.8
are edible. New plants frequently arise from root buds. syn. Callipteris prolifera (Lamarck) Bory
Diplazium esculentum is characterized by broad, two- Tender
pinnate fronds and veins that unite along the costae. The A large fern with erect rhizomes that form offshoots. Grows
scales of the rhizomes and stipe bases have black margins well under medium light in moist-wet garden soil or pot-
and forked teeth (use hand lens). It is native to Africa, ting mix; it does best with warm nights. The plants are large
southeastern Asia, and Polynesia; it has been introduced in and coarse but interesting for the large buds on the midrib.
Florida and Louisiana. Stems of older plants may become trunk-like.
Diplazium proliferum has one-pinnate fronds with
Diplazium lonchophyllum Kunze  13.43.7 coarsely serrate-lobed pinnae and netted veins. The scales of
Semi-tender, Zone 9 the rhizomes and stipe bases have black margins and forked
A medium-sized fern with suberect rhizomes that often teeth (use hand lens). This species is native to Africa, Mada-
produce offshoots. Grows well in moist garden soil or pot- gascar, and from Malaysia to Polynesia.
ting mix; not adapted to climates with cool nights.
The blades of Diplazium lonchophyllum are triangular-
lanceolate and two-pinnate at the base but one-pinnate-
288 C 13

Diplazium pycnocarpon (Sprengel) M. Broun

 13.43.9
Figure 13.43.7. Diplazium American glade fern, narrow glade fern
lonchophyllum: frond, bar = Very hardy, Zone 4
5 cm (2 in.). A medium-large fern with short-creeping, branched rhi-
zomes and deciduous fronds. Grows well under medium
light in moist, basic garden soil or potting mix. The leaves
are often light green.
Diplazium pycnocarpon is characterized by one-pinnate
fronds and nearly entire pinnae. It is native to eastern
North America.

Figure 13.43.9. Diplazium pycnocarpon: fronds, bar =

5 cm (2 in.).

Figure
13.43.8. Diplazium subsinuatum (Hooker & Greville) Tagawa
Diplazium  13.43.10
proliferum: syn. Athyrium dubium (D. Don) Ohwi, Diplazium
a. frond, bar lanceum (Thunberg) C. Presl
= 5 cm Semi-hardy, Zone 7(8)
(2 in.); A small fern with branched, medium- to long-creeping rhi-
b. leaf buds, zomes. Grows well under medium light in moist garden
bar = 5 cm soil or potting mix. The plants are useful for edging or as
(2 in.). ground cover.
 F  F A  C 289

Doodia  13.44.1

Hacksaw fern, rasp fern
This genus of small- to medium-sized terrestrial ferns in-
cludes the commonly cultivated hacksaw fern, Doodia me-
dia. This species tolerates drier soil and less humidity than
most ferns and is used in warm-temperate and subtropical
gardens as an edging plant or indoors as a pot or terrarium
plant. Doodia is easy to grow from spore, but the offspring
vary, with some individuals more erect or having redder
fronds. The species may spread by stolons.
Doodia is characterized by erect, narrow, pinnatifid or
Figure 13.43.10. one-pinnate, harsh-leathery fronds and oblong indusia that
Diplazium subsinuatum: run parallel and close to the costae. The rhizome varies
habit, bar = 5 cm (2 in.). from erect to ascending and bears dark scales. The numer-
ous close-set pinnae are ovate or linear, sessile or broadly
attached, and sharply toothed. The fertile fronds are often
taller than the sterile ones and have narrower pinnae. The
indusia open toward the costa, not the margin.
The fronds of Diplazium subsinuatum are simple, lin- Doodia consists of 12 species. It occurs from Sri Lanka
ear-lanceolate, thick, dark green, and glossy. The species is to Polynesia, Australia, and New Zealand. The genus is
native to India, eastern Asia, the Philippines, and Borneo. named for Samuel Doody (1656–1706), keeper of the
Chelsea Physic Garden, London, and the first cryptogamic
Diplazium tomitaroanum Masamune botanist in Britain.
 13.43.1, 2, 11
Semi-hardy, Zone 7(8) Doodia aspera R. Brown  13.44.2
A small fern with branched, medium- to long-creeping rhi- Prickly rasp fern
zomes. Grows well under medium light in moist to moist- Semi-hardy
wet garden soil or potting mix. The plants can be used for A small-medium fern with suberect rhizomes. Best grown
edging, as ground covers, or as an attractive plant for small under medium light in moist-dry garden soil or potting
pots. It does grow slowly. mix.
The blades are narrowly lanceolate, usually one-pinnate The pinnae of Doodia aspera are broadly attached (ad-
at the base and pinnatifid to lobed above, dark green, and nate) to the rachis, except the lowest pair is occasionally
glossy. Diplazium tomitaroanum is native to China, Japan, stalked. The blades are hairless. This species is native to
and Taiwan. Australia and New Zealand.

Figure 13.43.11. Diplazium

tomitaroanum: habit, bar =
5 cm (2 in.).

Figure 13.44.1. Doodia media: habit, young plant.

 290 C 13

a b

Figure 13.44.2. Doodia aspera: a. sterile frond (left) and

fertile frond (right), bar = 5 cm (2 in.); b. frond base, bar
c
= 1 cm (0.4 in.).

Doodia media R. Brown  13.44.1, 3

Hacksaw fern, rasp fern
Semi-hardy, Zone 8; tolerates light frost Figure 13.44.3. Doodia media: a. sterile frond (left) and
A small-medium fern with suberect rhizomes. Grows best fertile frond (right), bar = 5 cm (2 in.); b. frond base, bar
under high light in moist-dry garden soil. New growth = 1 cm (0.4 in.); c. sori and indusia, bar = 5 mm
tends to be redder if plants are placed in the sun. This spe- (0.2 in.).
cies is easy to cultivate. When sown, the spores produce
large prothalli.
Doodia media has fine, short hairs on the stipes and
blades. The pinnae in the lower third of the blade are
stalked or sessile but not adnate as in the preceding species.
Doodia media is native to Australia and New Zealand. It is
often confused with D. aspera in the United States trade.
Most of the trade material is subsp. australis Parris, Figure 13.44.4.
which has pinnae gradually changing from decurrent to Doodia mollis:
stalked over at least six pairs of pinnae, instead of abruptly a. sterile frond
changing as in subsp. media. (left) and fertile
frond (right), bar
Doodia mollis Parris  13.44.4 = 5 cm (2 in.);
Semi-hardy b. frond base,
A small fern with erect, clump-forming rhizomes. Grows bar = 1 cm
well under medium light in moist garden soil or potting (0.4 in.).
mix. This species is softer-textured than Doodia media and
is rarely cultivated.
Doodia mollis has dimorphic sterile and fertile fronds, a b
with the fertile ones taller and with narrower pinnae than
the sterile. In the lower half of the fronds, the pinnae are
 F  F A  C 291

stalked and have short “ears” at their base. Sometimes the

lowermost pinnae appear trilobed because of the ears. This
species is endemic to New Zealand.

Doryopteris  13.45.1–3

The eye-catching Doryopteris offers maple-like leaves borne
on dark, shiny stalks. Most species are small- to medium-
sized and usually grown in pots. In nature they grow in soil
or on rocks. Some species produce buds at the base of the
blade on either side of the stipe, and these buds can be used
to propagate the plants. The cultivated species are tender
and need warm temperatures year-round. Take care not to
overwater the plants during the cool months, when their
growth slows. These attractive ferns can be grown indoors
if they are gradually hardened before being placed in a drier
atmosphere.
The genus Doryopteris is characterized by dark, shiny
stipes, broadly triangular blades that are basally lobed and
then lobed again, and sori that run along the margin of the
Figure 13.45.2. Doryopteris concolor: basal part of fertile
blade. The rhizome is scaly and varies from decumbent to
frond.
suberect; rarely is it long-creeping. The sterile and fertile
blades are usually the same size and shape, but in some spe-
cies the fertile ones can be narrower and longer-petiolate. Doryopteris contains about 30 species, most of which
The sterile fronds can vary considerably, even on the same occur in tropical America. The genus also occurs in Africa,
plant. The veins are free or netted, and the indusium is Madagascar, Australia, southern India, Malaysia, and the
formed by the enrolled margin of the blade. islands of the South Pacific. The genus name comes from
the Greek, dory, lance, and pteris, fern, alluding to the shape
of the blade in certain species.

Figure 13.45.3. Doryopteris palmata: sori and

Figure 13.45.1. Doryopteris palmata: habit. indusia.
 292 C 13

Doryopteris concolor (Langsdorff & Fischer) Kuhn in

Decker  13.45.2, 4
syn. Cheilanthes concolor (Langsdorff & Fischer) R. M.
Tryon & A. F. Tryon
Tender
A small fern with erect rhizomes. Grows well under me-
dium light in moist potting soil.
Doryopteris concolor is characterized by free veins and
glabrous stipes grooved on the upper surface, with sharp,
raised ridges on either side of the groove. The sterile blades
lack buds and are more deeply divided than those of D.
palmata, a similar species. Doryopteris concolor is native to
Central and South America, the West Indies, tropical Af-
rica, Asia, Malaysia, Australia, and Polynesia.

Figure 13.45.5.
Doryopteris ludens:
habit, bar = 5 cm
(2 in.).

Doryopteris nobilis (T. Moore) C. Christensen

 13.45.6
Tender
A small-medium fern with short- to medium-creeping rhi-
zomes. Grows well under medium-high light in moist pot-
ting mix.
Doryopteris nobilis is the largest species in the genus, with
Figure 13.45.4. Doryopteris concolor:
sterile blades up to 65 cm (26 in.) long. The blades vary
frond, bar = 5 cm (2 in.).
tremendously, ranging from lanceolate–arrowhead-shaped
to palmately lobed to deeply pinnately lobed with lower
pinnae again lobed (often only basiscopically). The blade
bears buds at the base, and the veins are netted. The lobes
of the fertile fronds often have serrate tips beyond the sori.
This species is native to Colombia, Bolivia, Brazil, and
northern Argentina.
Plants circulating among collectors as Doryopteris sagit-
tifolia (Raddi) J. Smith are juvenile forms of D. nobilis.
True D. sagittifolia has arrowhead-shaped blades, or if
Doryopteris ludens (Wallich ex Hooker) J. Smith (rarely) five-lobed, the terminal lobe is the largest and the
 13.45.5 lateral lobes each bear a smaller basiscopic lobe.
Tender
A small-medium fern with slender, long-creeping rhizomes. Doryopteris palmata (Willdenow) J. Smith
Grows well under medium light in moist potting mix.  13.45.1, 3, 7
Doryopteris ludens resembles D. nobilis in blade shape syn. Doryopteris pedata var. palmata (Willdenow) Hicken
and netted veins but differs by the long-creeping rhizomes Spear-leaved fern, hand fern
and smaller sterile blades, only 4–45 cm (1.5–18 in.) long, Tender
that lack buds. It is native to northern India and southern A small fern with short-creeping to suberect rhizomes.
China to Malaysia. Grows well under high light in moist potting mix.
Doryopteris palmata, which has netted veins, has previ-
ously been treated as a variety of D. pedata; however, it dif-
 F  F A  C 293

Drynaria  13.46.1–3

Oak-leaf fern
Drynaria is a genus of spectacular medium- to large-sized
epiphytic ferns. The medium-sized species are suitable in-
doors with adequate light because the foliage usually toler-
ates the drier air indoors. During the dry season, or when
fronds become old, the pinnae (or lobes) are shed from the
midrib, but the midrib itself remains attached to the rhi-
zome, sticking outward like a long spine. The green fronds
may yellow when nighttime temperatures drop below 15°C
(59°F) for more than a few weeks. If the temperature has
not been too cold, growth resumes when warm weather re-
turns. For more details on cultivation, see the section in
Chapter 10 on “Polypodium and Relatives.” For a technical
monograph of the genus, see Roos (1985).
Drynaria has creeping, thick, fleshy, scaly rhizomes and
netted veins. The genus bears two kinds of fronds (it is di-
morphic). One type of frond is short and broad and turns
brown and papery. These fronds sit close to the stem and
collect falling debris that eventually decomposes to form
Figure 13.45.6. Doryopteris nobilis: fronds, bar = 5 cm
humus from which the roots can absorb nutrients. They
(2 in.).
are always sterile and resemble oak leaves. The second kind
of frond is longer, green, herbaceous, and fertile. Its func-
fers from that species by the presence of buds at the base of tion is to photosynthesize and produce spores, not collect
the blades, rounded and puberulent stipes (not grooved and humus. The fertile herbaceous fronds are pinnatifid or one-
glabrous), and wrinkled spores. It also has a different geo- pinnate and lack a true terminal pinna or lobe, although
graphical distribution, occurring from Mexico to Venezu- the terminal position may be occupied by a slightly skewed
ela and Bolivia, whereas D. pedata grows only in the West lateral pinna or lobe. (The true apex aborts, and occasion-
Indies. These differences warrant recognition at the species ally a partly aborted terminal pinna is produced.) The sori
level. are round or nearly so and lack indusia.
This genus resembles Aglaomorpha, which differs by
having a well-developed, true terminal pinna or lobe on the
foliaceous frond. It also lacks the separate small, brown, pa-
pery, humus-collecting fronds characteristic of Drynaria.
The close relationship of the two genera is evident in their
ability to hybridize. One such hybrid, ×Aglaonaria robertsii,
is important in horticulture.

Figure 13.45.7. Doryopteris

palmata: frond, bar = 5 cm (2 in.).

Figure 13.46.1. Drynaria sparsisora: habit.

 294 C 13

Figure 13.46.2. Drynaria rigidula: habit. a

Figure 13.46.4. Drynaria bonii: a. habit, with a humus-

collecting frond on the right, bar = 10 cm (4 in.);
b. pinna and sori, bar = 2.5 cm (1 in.).

only a few pinnae, which are separated by broad sinuses.

The pinnae are 2.5–4 cm (1–1.5 in.) wide, adnate to the
rachis, and form wings on the rachis above and below their
base. The sori are in two irregular rows between secondary
Figure 13.46.3. Drynaria rigidula: sori.
veins. The species is native to India, southeastern Asia, and
China.
Drynaria contains 16 species and is native to the Old Drynaria bonii is often confused with D. descensa Cope-
World tropics. The genus name comes from the Greek dry- land, a Philippine species differing by crenate, humus-col-
inos, of oaks, and refers to the plant’s oak-like sterile fronds. lecting fronds and smaller, less than 2.5 cm (1 in.) wide,
and undulate lobes on the foliaceous fronds.
Drynaria bonii H. Christ  13.46.4
Tender Drynaria propinqua (Wallich ex Mettenius) Beddome
A medium-sized fern with short-medium-creeping,  13.46.5
branched rhizomes. Grows well under high light in moist- Probably tender
dry, drained potting mix or uncut moss. A medium-sized fern with branched, creeping rhizomes.
Drynaria bonii is one of the smallest species in the genus. Grows well under high light in drained, moist-dry potting
Its humus-collecting fronds are rounded and nearly entire, mix or uncut moss. This species is reportedly difficult to
whereas the green, herbaceous fronds are pinnatisect with grow.
 F  F A  C 295

Figure 13.46.5. Drynaria propinqua: a. foliaceous frond

(left) and humus-collecting frond (right), bar = 10 cm
(4 in.); b. sori, bar = 2.5 cm (1 in.).
a

The humus-collecting fronds of Drynaria propinqua are

deeply pinnatifid with narrowly acute lobes, and the folia-
Figure 13.46.6. Drynaria quercifolia: a. foliaceous frond
ceous fertile fronds are deeply pinnatifid into lanceolate
(left) and humus-collecting frond (right), bar = 10 cm
pinnae. The sori are borne in one row on both sides of, and
(4 in.); b. rhizome scale, bar = 3 mm (0.1 in.); c. sori,
close to, the costae. The species is native to India, south-
bar = 2.5 cm (1 in.).
eastern Asia, and China.

Drynaria quercifolia (Linnaeus) J. Smith

 13.46.6 This species is native from India to southeastern Asia, Ma-
Oak-leaf fern laysia, Indonesia, the Philippines, New Guinea, and Aus-
Tender tralia.
A large fern with medium- to short-creeping, branched rhi-
zomes and spreading fronds. Grows well under high light in Drynaria rigidula (Swartz) Beddome
drained, moist-dry potting mix, or uncut moss. The plants  13.46.2, 3, 7
are easy to cultivate and are fast-growing. Semi-tender to slightly hardier
The rhizome scales of Drynaria quercifolia are brownish, A large fern with medium- to short-creeping, branched rhi-
narrowly triangular, and spreading at about a 45-degree zomes and usually gracefully arching fronds. Prefers high
angle from the rhizome surface. The humus-collecting light and drained, moist-dry potting mix or uncut moss.
fronds vary from wavy to lobed. The foliaceous ones are The humus-collecting fronds develop poorly if the soil is
deeply pinnatifid and up to about 90 cm (36 in.) long. The kept too moist. The plants grow slowly.
sori vary from round to oblong, but sometimes adjacent Drynaria rigidula is the only species of Drynaria with
ones fuse into an elongate sorus. They are arranged in two one-pinnate fronds and sessile, wedge-shaped pinnae nar-
fairly regular rows or scattered between the lateral veins. rowly winged to the rachis. It grows in southeastern Asia,
 296 C 13

b c

Figure 13.46.7.
Drynaria rigidula:
a. foliaceous frond
(center) and
humus-collecting
frond (right), b
foliaceous pinnae
of ‘Whitei’ (lower
left), bar = 10 cm
(4 in.); b. pinna
base and sori, bar =
2.5 cm (1 in.).

a
a

China, Malaysia, Indonesia, the Philippines, Australia, Figure 13.46.8. Drynaria sparsisora: a. foliaceous frond
New Guinea, and southeastern Pacific Islands. (left) and humus-collecting frond (right), bar = 10 cm
‘Whitei’. Plate 25. Pinnae often broader, lacerate to ir- (4 in.); b. rhizome scales, bar = 3 mm (0.1 in.); c. sori,
regularly lobed. bar = 2.5 cm (1 in.).

Drynaria sparsisora (Desvaux) T. Moore

 13.46.1, 8
is sought after for its copper-red new foliage and bright red
Tender
indusia. Generally, deciduous species from cold-temperate
A large fern with medium- to short-creeping, branched rhi- climates tend to have a shorter dormancy when grown in
zomes. Grows well under high light in drained, moist-dry warmer climates. Deciduous fronds wither in place (mar-
potting mix or uncut moss. cescent) but may or may not lose their green color.
Drynaria sparsisora resembles D. quercifolia but differs About 50 species of Dryopteris are cultivated in the
by blackish rhizomes scales that are bent above their base United States, and new ones are constantly being intro-
into a 90-degree angle to form a stiffly erect bristle. The fo- duced, replacing older ones that often disappear from the
liaceous fronds are smaller, harder-textured, and more trade. Natural hybrids are common, and Dryopteris com-
stiffly erect, and the sori are smaller and irregularly scat- plexa (D. affinis × D. filix-mas), in particular, is widely sold,
tered between lateral veins. It is native to southeastern Asia, often under the trade name D. filix-mas ‘Robusta’. Most of
Malaysia, Indonesia, New Guinea, and Australia. the cultivars originated from the European species, prima-
rily D. affinis and D. filix-mas. As a group, the shield ferns
Dryopteris  13.47.1, 2 pose no special problems in cultivation, although some of
Shield fern, buckler fern, wood fern the species native to colder climates do not adapt well to
These familiar ferns abound in the woods, fields, and wet gardens in warm climates, and some species thrive only in
areas of the eastern United States, Europe, Japan, and many acidic soils. Most species are easy to propagate from spores
other areas. Dryopteris is widely used in temperate gardens or divisions. Offshoots come from the rhizome bases.
as accents, in borders, or for background foliage. The many Dryopteris consists of terrestrial plants with thick, erect
species available in the trade range from small to large, or suberect rhizomes that are surrounded by close, spirally
coarsely to finely cut, and deciduous to evergreen. One gar- arranged old leaf bases. Rarely, the rhizomes are short-
den favorite is the ornamental autumn fern (Dryopteris ery- creeping. The rhizome scales are nonclathrate. The stipes
throsora) from Japan. It is widely available in the trade and are grooved, scaly, and have 3 to 7 (9–10) vascular bundles
 F  F A  C 297

lated groups of similar species. Furthermore, the fronds can

vary even on a single plant. Identification requires examin-
ing large, mature leaves. Name changes also complicate the
situation. Many such changes have occurred in certain Eu-
ropean species (Dryopteris affinis complex) and North
American ones (D. spinulosa complex). For a fuller treat-
ment of the cultivated species, see Hoshizaki and Wilson
(1999).
Due to the large number of cultivated species of Dry-
opteris, the following key is given to aid identification. After
you limit the unknown species to a smaller group using the
key, you can then compare it to the descriptions and illus-
trations of those species. The groups used in this key are
the sections of Dryopteris recognized by Fraser-Jenkins
(1986).

1. Scales on the lower surface of the blades strongly to

slightly inflated or puffy (at least at the base).
Figure 13.47.1. Dryopteris uniformis ‘Cristata’: habit. 2. Basal pinnae with the downward-pointing basal pin-
nule longer than the adjacent pinnules: Section Variae
(Dryopteris bissetiana, D. formosana, D. sacrosancta, and
D. varia).
2. Basal pinnae with the downward-pointing basal pin-
nule slightly shorter or about the same size as the others:
Section Erythrovariae (Dryopteris championii, D. cysto-
lepidota, D. decipiens, D. erythrosora, D. fuscipes, D. hon-
doensis, and D. purpurella).
1. Scales on the lower surface of the blades flat or nearly so.
3. Blades one-pinnate to one-pinnate-pinnatifid.
4. Pinnae entire: Section Pycnopteris (Dryopteris sieboldii).
4. Pinnae pinnatifid.
5. Pinnae cut less than halfway to the midrib: Sec-
tion Hirtipedes (Dryopteris cycadina and D. kuratae).
5. Pinnae cut nearly to the midrib: Section Fibril-
losae (Dryopteris affinis, D. complexa, D. crassirhi-
zoma, D. lepidopoda, D. polylepis, D. pseudo-filix-mas,
and D. wallichiana).
3. Blades two-pinnate to four-pinnate.
6. Blades mostly narrowed toward the base (except
slightly so in Dryopteris clintoniana and D. goldiana):
Figure 13.47.2. Dryopteris erythrosora: indusia. Sections Pandae and Dryopteris (Section Pandae: Dry-
opteris ×australis, D. ×bootii, D. celsa, D. clintoniana, D.
cristata, D. ludoviciana, and D. tokyoensis; Section Dry-
opteris: D. caucasica, D. filix-mas, D. fragrans, D.
arranged in a C-shaped pattern. The blades are one- to goldiana, D. oreades, and D. sichotensis).
four-pinnate, bearing scales but no hairs. The blades are 6. Blades mostly widest at the base (except narrowed in
usually catadromous but occasionally otherwise. The mid- Dryopteris lacera, variable in D. arguta).
rib is grooved, and the groove usually continues uninter- 7. Segment teeth not sharp or spine-tipped (varia-
rupted into the costae. Round sori are covered by kidney- ble in Dryopteris stewartii ): Sections Aemulae, Palli-
shaped indusia, which are attached at a sinus (rarely the dae, and Remotae. (Section Aemulae: Dryopteris ae-
indusia are absent). The genus, which contains about 225 mula; Section Pallidae: D. arguta, D. lacera, D. mar-
species, is cosmopolitan, but most of the species occur in ginalis, D. mindshelkensis, D. stewartii, D. sublacera,
temperate regions. and D. uniformis; Section Remotae: D. remota).
The species of Dryopteris are difficult to identify because 7. Segment teeth sharp-pointed or hair-tipped: Sec-
there are so many of them, and they tend to form closely re- tion Lophodium (Dryopteris amurensis, D. campylop-
 298 C 13

tera, D. carthusiana, D. dilatata, D. expansa, and D. Dryopteris affinis (Lowe) Fraser-Jenkins

intermedia).  13.47.4, 5
syn. Dryopteris abbreviata (de Candolle) Newman, D.
Dryopteris aemula (Aiton) O. Kuntze  13.47.3 pseudomas (Wollaston) Holub & Pouzar
Hay-scented wood fern Yellow-golden-scaled male fern, common golden-scaled
Hardy, Zone 6 male fern, scaly male fern, hard male fern
A medium-sized fern. up to 80 cm (32 in.) long, with erect Hardy, Zone 4(5)
to suberect rhizomes and semi-evergreen fronds. Grows A medium to large fern (some cultivars are dwarf), to 130
under medium light in moist, acidic garden soil or potting cm (52 in.) tall, with erect, stout rhizomes. Grows well
mix. This species grows slowly and prefers high humidity. under medium light in moist garden soil or potting mix.
Dryopteris aemula resembles D. dilatata but has smaller, The fronds are deciduous in cold-temperate climates but
paler green, moderately ruffled blades, and the stipe scales nearly evergreen in warmer ones. The plants are easy to
are of one color, not darker in the center. The stipes are grow.
dark purple-brown at the base and as long as the blade, The stipe scales are golden, dense, and conspicuous, es-
which is triangular-ovate, to three-pinnate-pinnatifid, with pecially on new fiddleheads. Dryopteris affinis has glossy,
pinnules curving upward and appearing ruffled. When lanceolate blades to two-pinnate. The undersides of the
crushed, the blades emit an odor of drying hay. The species costae next to the rachis have a black blotch. The indusia
is native to western Europe. are tucked under the sori when young. This species is native
from Europe to the Caspian Sea and northwestern Africa.
The subspecies of Dryopteris affinis are difficult to dis-
tinguish. The many cultivars are confused as to their names
and the subspecies from which they originated. In some

d
a

e
b

b
Figure 13.47.4. Dryopteris affinis: a. frond, bar = 5 cm
(2 in.), after Hoshizaki and Wilson (1999); b. stipe
scales, bar = 1 cm (0.4 in.), after Hoshizaki and Wilson
Figure 13.47.3. Dryopteris aemula: a. frond, bar = 5 cm (1999); c. pinnules from medial pinnae, bar = 1 cm
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After (0.4 in.); d. subsp. borreri, bar = 5 cm (2 in.); e. subsp.
Hoshizaki and Wilson (1999). cambriensis, bar = 5 cm (2 in.).
 F  F A  C 299

‘Crispa Gracilis’. Figure 13.47.5c. Dwarf, congested,

b d g
upright leathery fern with pinna apices curved and hooked.
c
‘Cristata’. (‘Cristata the King’, ‘The King’). Figure
13.47.5a. Fronds arching, blade apex and pinnae ending
in a tassel or large crest. Zones 4–9.
‘Cristata Angustata’. Figure 13.47.5e. Narrow frond, to
7.5 cm (3 in.) wide, apex and pinnae crested.
‘Polydactyla’. Figure 13.47.5f. A group of crested forms
with long, finger-like, flat tassels on the pinna tips. Trade
plants usually listed as ‘Polydactyla Dadd’s’.
‘Revolvens’. Figure 13.47.5g. Plants by this name vary.
The current trade plant has longer, wider fronds with the
margins curved downward; the original plant has shorter,
narrower fronds with the pinna tips curved downward.
‘Stableri’. Fronds narrow, erect to slightly arching.
‘Stableri Crisped’. Figure 13.47.5h. Like ‘Stableri’ but
with ruffled pinnae and pinnules.

Dryopteris amurensis H. Christ  13.47.6

Very hardy, Zone 4
A medium-sized fern, to 50 cm (20 in.), with short-creep-
h ing rhizomes and evergreen fronds. Grows under medium
light in moist garden soil or potting mix.
e Dryopteris amurensis has stipes that are longer than the
a f
blades. The blades are broadly triangular (wider than long),
three-pinnate pinnatifid at the base, membranous, and
Figure 13.47.5. Dryopteris affinis cultivars: a. ‘Cristata’; bearing small scales along the veins on the lower surface.
b. ‘Congesta’; c. ‘Crispa Gracilis’; d. ‘Congesta Cristata’; The species is native to northeastern Asia.
e. ‘Cristata Angustata’; f. ‘Polydactyla’; g. ‘Revolvens’,
after Druery (1912); h. ‘Stableri Crisped’. Bar = 5 cm
(2 in.).

Figure 13.47.6.
Dryopteris
case the names may correctly apply to D. filix-mas. In the
amurensis:
United States the hybrid between D. affinis and D. filix-
a. frond, bar =
mas, known as D. complexa (misnamed D. filix-mas ‘Un-
5 cm (2 in.);
dulata-Robusta’) was widely sold.
b. stipe scales,
subsp. borreri (Newman) Fraser-Jenkins. Figure
bar = 1 cm
13.47.4d. Fronds lanceolate to oblong, bases truncate, sur-
(0.4 in.). After
faces not glossy; deciduous in early winter in temperate
Hoshizaki and
climates.
Wilson (1999).
subsp. cambriensis Fraser-Jenkins. Figure 13.47.4e.
Fronds oblanceolate to linear-elliptic, bases tapering, upper
surfaces slightly glossy; deciduous in temperate climates.
‘Congesta’. Figure 13.47.5b. Fronds dwarf, the pinnae
and pinnules congested but neatly overlapping. a
‘Congesta Cristata’. Figure 13.47.5d. Fronds dwarf,
congested, crested.
‘Crispa’. Crisped male fern. In Victorian times this b
name applied to plants with dwarf and broad fronds, to 20
cm (8 in.) long and 14 cm (5.5 in.) wide, with ruffled and
congested foliage. Plants in the current trade are normal-
sized with twisted segments. They are perhaps correctly
called ‘Paleaceo Crispa’.
 300 C 13

Dryopteris arguta (Kaulfuss) Maxon  13.47.7

Coastal wood fern, California shield fern
Semi-tender, Zones 8–9
A medium-sized fern, to 65 cm (26 in.), with short-creep-
ing to suberect rhizomes. Grows under low to medium
light in moist-dry garden soil or potting mix with good
drainage. This species is difficult to grow. The plants are
usually semi-dormant in the dry season and sensitive to
overwatering at that time.
The blades of Dryopteris arguta vary from triangular to
ovate-lanceolate and are mostly pinnate-pinnatifid with ser-
rate segments bearing spreading, spine-tipped teeth. The
species is native to western North America.

Figure 13.47.8. Dryopteris

×australis: rhizome and frond,
bar = 5 cm (2 in.). After
Hoshizaki and Wilson (1999).

ents in appearance, but fertile parts tend to be contracted.

The species is native to the southeastern United States.

Dryopteris bissetiana (Baker) C. Christensen

 13.47.9
a b Beaded wood fern
Hardy, Zone 5
A medium-sized fern, to 45 cm (18 in.), with erect rhi-
zomes and dark green, glossy, evergreen fronds with a bead-
like aspect to the foliage. Grows well under medium light in
moist garden soil or potting mix. Easy to grow.
Dryopteris bissetiana bears triangular, two-pinnate-pin-
natifid, leathery blades, with the lowest basiscopic pinnule
Figure 13.47.7. Dryopteris arguta: a. habit, bar = 5 cm
next to rachis elongate. The segment margins turn slightly
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
downward and lack short spines or teeth. The scales on the
Hoshizaki and Wilson (1999).
costa undersides are weakly inflated. The species is native to
eastern Asia.
Most of the current trade material called Dryopteris bis-
Dryopteris ×australis (Wherry) Small  13.47.8 setiana is actually D. erythrosora.
Dixie wood fern
Hardy, Zone 5 Dryopteris ×boottii (Tuckerman) Underwood
A medium-large fern, to 115 cm (45 in.), with short-creep-  13.47.10
ing rhizomes and deciduous fronds. Grows well under me- Boott’s wood fern
dium light in moist garden soil or potting mix. The plants Very hardy, Zone 3
are easy to grow. They cannot be propagated from spores A medium-sized fern, to 70 cm (28 in.) tall, with short-
because the spores are aborted. All plants sold in the trade creeping rhizomes that form offshoots. Grows well under
are propagated by tissue culture. medium light in moist garden soil or potting mix. The
Dryopteris ×australis is a sterile hybrid of D. celsa and D. plants are deciduous. It cannot be propagated by spores be-
ludoviciana. The plants are intermediate between the par- cause these are aborted.
 F  F A  C 301

Dryopteris ×boottii is a sterile hybrid of D. cristata and D.

Figure 13.47.9. intermedia. It mostly resembles D. cristata, but it bears the
Dryopteris bissetiana: small, stalked glands typical of D. intermedia. The pinnae
a. frond, bar = 5 cm often twist horizontally to give a “venetian-blind” effect, as
(2 in.); b. stipe in D. cristata. Dryopteris ×boottii is native to the northeast-
scales, bar = 1 cm ern United States and is commonly found where the two
(0.4 in.). After parents mingle.
Hoshizaki and
Wilson (1999). Dryopteris campyloptera Clarkson  13.47.11
syn. Dryopteris spinulosa var. americana (Fischer) Fernald
Mountain wood fern, eastern spreading wood fern
Very hardy, Zone (3)4
A medium-sized fern, to 60 cm (2 ft.), with ascending to
erect rhizomes and deciduous fronds. Grows under low to
medium light in moist garden soil or potting mix. The
plants need ample humidity and are unsuited for subtropi-
cal climates. Although this species is of hybrid origin, the
b plants are fertile and therefore can be propagated by spores.
Dryopteris campyloptera is a fertile hybrid of D. expansa
and D. intermedia. It mostly resembles D. expansa but has
a more erect habit and firmer texture. The basiscopic basal
pinnule is two to four times longer than the opposite up-
ward-pointing pinnule (the acroscopic pinnule). The spe-
a
cies is native to northeastern North America.

b
Figure 13.47.10. Dryopteris
×boottii: frond, bar = 5 cm
(2 in.). After Hoshizaki
and Wilson (1999).

Figure 13.47.11. Dryopteris campyloptera: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
Hoshizaki and Wilson (1999).
 302 C 13

Dryopteris carthusiana (Villars) H. P. Fuchs Dryopteris caucasica (A. Braun) Fraser-Jenkins & Corely
 13.47.12  13.47.13
syn. Dryopteris spinulosa (O. F. Müller) Watt Hardy, Zone 5
Spinulose wood fern, toothed wood fern, narrow buckler A medium-sized fern, to 80 cm (32 in.) tall, with deciduous
fern fronds and erect rhizomes that form offshoots. Grows well
Very hardy, Zone 2 under medium light in moist garden soil or potting mix.
A medium-sized fern, to 40 cm (16 in.), with ascending to Dryopteris caucasica resembles D. filix-mas (one of its
erect rhizomes. Grows well under medium light in moist, parents) but differs by paler blades, pinnae widest at the
acidic garden soil or potting mix. The plants are deciduous middle (not the base), acute segments that are doubly
and unsuited to subtropical climates. toothed at the apex with acute teeth, and white, lacerate in-
The blades of Dryopteris carthusiana are two-pinnate- dusia. It is native to the Middle East.
pinnatifid to three-pinnate and narrowly triangular-lance-
olate. The lowermost, downward-pointing pinnule is less
than two times as long as the basal upward-pointing pin- a
nule above it. Dryopteris carthusiana is distinguished from
the similar D. dilatata by its medium to light green fronds,
flat margins, and stipe scales of one color. It also resembles
D. intermedia but lacks the minute glandular hairs charac-
teristic of that species. The species is native to North Amer- Figure 13.47.13.
ica, Europe, and Asia. Dryopteris caucasica:
A foliose form of Dryopteris carthusiana is often sold in a. frond, bar = 5 cm
the trade and incorrectly identified as D. stewartii. (2 in.); b. stipe
scales, bar = 1 cm
(0.4 in.). After
Hoshizaki and
Wilson (1999).

Dryopteris celsa (W. Palmer) Knowlton, W. Palmer &

Pollard  13.47.14
Log fern
Hardy, Zone (4)5
A medium-large fern, to 130 cm (52 in.), with medium- to
a
short-creeping rhizomes and deciduous fronds. Grows well
under medium light in moist-wet, acidic soil or potting
mix. Though deciduous, it grows satisfactorily in southern
California.
b Dryopteris celsa is a fertile hybrid of D. goldiana and D.
ludoviciana. It resembles D. clintoniana but differs by the
dark brown stipe scales with a darker central stripe; in ad-
Figure 13.47.12. Dryopteris carthusiana: a. frond, bar = dition, on the basal pinnae, the basiscopic pinnule next to
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After the rachis is equal to or shorter than the adjacent pinnules.
Hoshizaki and Wilson (1999). It is native to the eastern United States.
 F  F A  C 303

Figure 13.47.14.
Dryopteris celsa:
a. frond, bar = c
5 cm (2 in.);
b. stipe scales, bar
= 1 cm (0.4 in.).
After Hoshizaki
and Wilson
(1999).

b
a
b
Figure 13.47.15. Dryopteris championii: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. pinnule from medial pinna, bar = 5 mm (0.2 in.);
Dryopteris championii (Bentham) C. Christensen ex d. bullate scales from costa, bar = 1 mm. After Hoshizaki
Ching  13.47.15 and Wilson (1999).
Champion’s wood fern
Hardy, Zone 5
A medium-sized fern, to 90 cm (36 in.), with erect rhi-
zomes and evergreen fronds. Grows well under medium to Dryopteris clintoniana is a hybrid of D. cristata and D.
high light in moist-dry garden soil or potting mix. In the goldiana. It resembles D. cristata, but its basal pinnae are
northeastern United States the fertile leaves are not pro- longer than they are wide and not twisted into a horizontal
duced until the late fall. This species is particularly attrac- plane. The species is native to northeastern North America.
tive in winter because its evergreen fronds remain upright,
not reclining as in many species with evergreen fronds. Dryopteris complexa Fraser-Jenkins  13.47.17
Dryopteris championii has narrowly triangular fronds Robust male fern
and pinnules that are eared on both sides at the base. The Hardy, Zone 4(5)
stipe, rachis, and costae bear a dense covering of shiny red- A medium-large fern, to 130 cm (52 in.), with erect rhi-
brown scales, and those on the costae become strongly in- zomes that form offshoots and deciduous to semi-evergreen
flated. The species is native to eastern Asia. fronds (in warmer climates). Grows well under medium
light in moist to moist-dry garden soil or potting mix. The
Dryopteris clintoniana (D. C. Eaton) Dowell plants are easy to cultivate and become rapid, vigorous
 13.47.16 growers with full foliage. The fronds are deciduous, but less
Clinton’s wood fern so in warmer climates.
Very hardy, Zone 3 Dryopteris complexa is a fertile hybrid of D. affinis and D.
A medium-sized fern, to 95 cm (38 in.), with short-creep- filix-mas. The stipe scales are narrowly triangular and mem-
ing rhizomes and deciduous fronds. Grows well under me- branous. The middle and distal pinnae often overlap
dium light in acidic, moist-wet garden soil or potting mix. slightly, and the pinna midrib has a blackened spot on the
304 C 13

underside next to the rachis. Unlike D. affinis, which also

has the blackened spot, this species has segments that are
toothed on the sides and has rounded apices. It is native to
Europe.
The names Dryopteris filix-mas ‘Undulata-Robusta’ and
D. filix-mas ‘Robusta’ are often misapplied in the United
States trade to this species.
Figure 13.47.16.
Dryopteris clintoniana: Dryopteris crassirhizoma Nakai  13.47.18
a. frond, bar = 5 cm Thick-stemmed wood fern
(2 in.); b. stipe scales, Hardy, Zones 5–8
bar = 1 cm (0.4 in.).
After Hoshizaki and A medium-sized fern, to 1 m (3 ft.), with deciduous fronds
Wilson (1999). and erect rhizomes that form offshoots. Grows well under
medium light in moist-dry garden soil or potting mix. This
species is easy to grow.
Dryopteris crassirhizoma forms an attractive vase-shaped
cluster of fronds. Mickel (1994) reported that upon the on-
set of cold weather the fronds abruptly recline to the ground
overnight (not gradually) and remain green throughout the
winter. The erect, scaly crown of this fern is broad and mas-
sive, measuring 10–15 cm (4–6 in.) across. The fronds re-
a b semble those of D. affinis but differ by thick, glossy, brown
stipe scales up to 2 cm (0.75 in.) long, one-pinnate-pin-

c
c

a
b
a

Figure 13.47.18. Dryopteris crassirhizoma: a. frond, bar =

Figure 13.47.17. Dryopteris complexa: a. frond, bar = 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); c. pinnules, bar = 1 cm (0.4 in.); d. medial pinna (above)
c. pinnules from a medial pinna, bar = 5 cm (2 in.). and lower pinna (below), bar = 2 cm (0.8 in.). After
After Hoshizaki and Wilson (1999). Hoshizaki and Wilson (1999).
 F  F A  C 305

natifid blades, longer and narrower segments with margins Dryopteris cycadina (Franchet & Savatier)
entire or weakly crenate-serrate, and rounded and entire C. Christensen  13.47.20
apices. The costae lack a black blotch at the base. The spe- Shaggy wood fern, black wood fern
cies is native to northeastern Asia. Hardy, Zone 6
A medium-sized fern, to 1 m (3 ft.), with erect rhizomes
Dryopteris cristata (Linnaeus) A. Gray and semi-evergreen fronds. Grows well under medium
 13.47.19 light in moist-dry garden soil or potting mix.
Crested wood fern The common names of Dryopteris cycadina come from
Very hardy, Zone 3 the dense, dark scales that cover the stipes and rachises. The
A medium-sized fern, to 50 cm (20 in.) tall, with deciduous blades are one-pinnate and leathery with about 30 pairs of
fronds and short-creeping to erect rhizomes that form off- pinnae, these mostly coarsely serrate to lobed one-third of
shoots. Grows well under medium light in moist-wet gar- the way to the costa. The basal pinnae point downward.
den soil or potting mix. The plants are easy to cultivate. The species is native to eastern Asia.
The fronds of Dryopteris cristata are one- to two-pin- The cultivated plants of Dryopteris cycadina are often
nate, narrowly lanceolate, and often have pinnae oriented misidentified as D. atrata (Wallich) Ching or D. hirtipes
more or less horizontally—the “venetian blind” effect. The Kuntze.
basal pinnae are triangular, and fertile fronds are taller and
more erect than sterile ones. The fertile pinnae occur mostly Dryopteris cystolepidota (Miquel) C. Christensen
toward the apex of the blade and are slightly contracted.  13.47.21
The species is native to the northern and southeastern syn. Dryopteris nipponensis Koidzumi
United States. It commonly hybridizes in nature with D. Hardy, Zone (5)6
intermedia to produce D. ×boottii. A medium-sized fern, to 60 cm (2 ft.), with branched,
short-creeping rhizomes and evergreen, glossy fronds.
Grows well under medium light in moist garden soil or pot-
ting mix. The plants are easy to grow.
The blades of Dryopteris cystolepidota are broadly trian-
gular, up to nearly three-pinnate at base, and thin-leath-
ery. The segments have small spines turned up from the
surface. The species is native to Japan and Korea.

Figure 13.47.19. Dryopteris a Figure 13.47.20. Dryopteris

cristata: a. frond, bar = cycadina: a. frond, bar = 5 cm
5 cm (2 in.); b. stipe scales, (2 in.); b. stipe scales, bar =
bar = 1 cm (0.4 in.). After 1 cm (0.4 in.); c. medial
Hoshizaki and Wilson pinna, bar = 1 cm (0.4 in.).
(1999). After Hoshizaki and Wilson
(1999).

b
a

b
 306 C 13

a Figure 13.47.22. Dryopteris

decipiens var. diplazioides:
a. frond, bar = 5 cm (2 in.);
b. stipe scales, bar = 1 cm
(0.4 in.); c. lowest pinna, bar
= 2 cm (0.8 in.). After
Hoshizaki and Wilson (1999).

a
b

b
Dryopteris dilatata (Hoffmann) A. Gray
 13.47.23, 24
Broad wood fern, broad buckler fern
Hardy, Zone 4(5)
A medium to large fern, to 1 m (3 ft.) or more, with decid-
c uous fronds and ascending to erect rhizomes that form off-
shoots. Grows well under low to medium light in acidic,

Figure 13.47.21. Dryopteris cystolepidota: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); a
c. lowest pinna, bar = 2 cm (0.8 in.). After Hoshizaki
and Wilson (1999).

Dryopteris decipiens (Hooker) O. Kuntze

 13.47.22
Hardy, Zone 5(7)
A small fern, to 30 cm (1 ft.), with ascending to erect rhi-
zomes that form offshoots. Grows well under medium light
in moist garden soil or potting mix. The fronds are decid-
uous in cold climates. They are coppery brown when
young.
Dryopteris decipiens has one-pinnate-pinnatifid fronds
with, generally, 15–18 pairs of shallowly lobed pinnae. The b
apices of the pinnae are curved slightly upward, toward the
apex of the blade. The scales at the base of the stipes are
purplish brown. The lower surfaces of the rachises and
costae are beset with inflated scales. The species is native to
Japan and China.
Plants in which the lower pinnae have a nearly free, Figure 13.47.23. Dryopteris dilatata: a. frond, bar = 5 cm
round lobe next to the rachis are referred to as var. diplazi- (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
oides (H. Christ) Ching (Figure 13.47.22). Hoshizaki and Wilson (1999).
 F  F A  C 307

together, slightly ruffled, blue-green in color. Hardy to

a
Zone 6.
b ‘Lepidota Cristata’. Figure 13.47.24b. Fronds finely dis-
sected and crested, to 50 cm (20 in.), stipe and rachis with
many brown, puffy scales. Hardy to Zone 4 and evergreen.
Not as robust a grower as other cultivars.
‘Recurved Form’. Segment margins strongly curved
downward. Not as robust a grower as many of the other
cultivars.

Dryopteris erythrosora (D. C. Eaton) O. Kuntze

 13.47.2, 25;  26
Autumn fern
Hardy, Zone 5(6)
A medium-sized fern, to 1 m (3 ft.), with evergreen fronds
and ascending to erect or prostrate, branching rhizomes.
Grows well under low to medium-high light in moist to
moist-dry garden soil or potting mix. The plants are easy to

e
e
c
d

Figure 13.47.24. Dryopteris dilatata cultivars: a. ‘Crispa

Whiteside’; b. ‘Lepidota Cristata’; c. ‘Grandiceps’; d
d. ‘Cristata’; e. ‘Jimmy Dyce’. Bar = 5 cm (2 in.).

moist garden soil or potting mix. The plants are easy to

grow. They are semi-evergreen and less vigorous in sub-
tropical climates.
Dryopteris dilatata is a fertile hybrid of D. expansa and
D. intermedia. It is often confused with D. carthusiana,
which differs by darker green fronds with margins turned
downward, larger stipe scales with a dark central stripe, and
the downward-pointing pinnule next to the rachis two or b
more times as long as the opposite upward-pointing one. It a
is native to Europe.
Dryopteris dilatata is often mislabeled as D. austriaca.
‘Crispa Whiteside’. Whiteside’s crisped broad-buckler
fern. Figure 13.47.24a. Ruffled foliage, lighter color than c
the species.
‘Cristata’. Figure 13.47.24d. Frond and pinna tips
crested when fully developed. Figure 13.47.25. Dryopteris erythrosora: a. frond, bar =
‘Grandiceps’. Figure 13.47.24c. Frond apex crested into 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
a dense tassel, pinnae crested. c. lowest pinna, bar = 2 cm (0.8 in.); d. pinnule from
‘Jimmy Dyce’. Figure 13.47.24e. Fronds shorter, to medial pinna, bar = 5 mm (0.2 in.); e. forma prolifica,
60 cm (24 in.), usually less stiff and erect, the pinnae close bar = 5 cm (2 in.). After Hoshizaki and Wilson (1999).
 308 C 13

cultivate. New growth has an attractive reddish coppery Dryopteris expansa has tan stipe scales with a dark central
color, turning dark, shiny green at maturity. New fronds stripe. The blades lack glands and are broadly triangular
are produced throughout the growing season, and in win- and three-pinnate-pinnatifid at the base, with the down-
ter the fronds remain upright, not reclining as in many ward-pointing pinnule next to the rachis two to three times
evergreen ferns. longer than the opposite upward-pointing pinnule. The
Dryopteris erythrosora is highly variable but can be char- pinnule margins are flat, not enrolled. The species is native
acterized as follows: the stipe scales are blackish and the to North America, Europe, and eastern Asia. It is a parent
blades are two-pinnate, mostly oblong-ovate, broad, and of D. campyloptera and is difficult to distinguish from the
arching. The lowest basiscopic pinnules next to the rachis offspring.
are short and have marginal teeth spinulose-tipped and in-
curved. The costal scales are strongly inflated and dark to- Dryopteris filix-mas (Linnaeus) Schott
ward the tip. Set close to the midrib, the indusia are red or,  13.47.27, 28
rarely, greenish white (forma viridosora (Nakai ex H. Ito) Common male fern
H. Ito). The species is native to eastern Asia. Hardy, Zone 4(5)
The variability of this species has caused considerable A large fern, to 130 cm (52 in.), with deciduous fronds and
confusion in the trade. Most of the current plants circulat- short-creeping to erect rhizomes that form offshoots.
ing as Dryopteris bissetiana and D. purpurella are actually Grows well under low to medium light in moist garden soil
D. erythrosora. or potting mix. The usually deciduous fronds become semi-
forma prolifica (Maximowicz ex Franchet & Savatier) evergreen if the plants are grown in warmer climates. This
H. Ito. Blades shorter, 37 cm (15 in.), more triangular, usu- species is easy to grow and has many garden variations.
ally with buds; pinnules narrow-linear, pointed. Dryopteris filix-mas is of hybrid origin (but fertile) from
a naturally occurring cross between D. caucasica and D. ore-
Dryopteris expansa (C. Presl) Fraser-Jenkins & Jermy ades. The fronds are one-pinnate-pinnatisect (nearly two-
 13.47.26 pinnate) with segments slightly tapered and toothed along
syn. Dryopteris austriaca (Jacquin) Woynar
Arching wood fern, northern spreading wood fern
Very hardy, Zone 3(4)
A medium-sized fern, to 60 cm (2 ft.), with ascending or
erect rhizomes and deciduous fronds. Grows well under
low to medium light in acidic, moist garden soil or potting
mix. The plants do not do well in subtropical climates.

d
b

a b
a

Figure 13.47.27. Dryopteris filix-mas: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
Figure 13.47.26. Dryopteris expansa: a. frond, bar = 5 cm c. pinnules from medial pinna, bar = 1 cm (0.4 in.);
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After d. other frond variations, bar = 10 cm (4 in.). After
Hoshizaki and Wilson (1999). Hoshizaki and Wilson (1999).
 b

a c
d

h i
f g

Figure 13.47.28. Dryopteris filix-mas cultivars: a. ‘Linearis Polydactyla’, an extreme form; b. ‘Linearis Polydactyla’; c.
‘Cristata’; d. ‘Grandiceps’; e. ‘Linearis Congesta’; f. ‘Barnesii’; g. ‘Crispa Cristata’; h. a dwarf, crisped, crested cultivar; i.
‘Ramo Cristata’. Bar = 5 cm (2 in.).
 310 C 13

the sides and apex. The teeth are acute and typically point
toward the segment apex. The upper surface is puckered
opposite the sori, and the young indusia have a flat, thin
rim all around the edge. The species is native to North
America and Europe. c
‘Barnesii’. Figure 13.47.28f. Barne’s male fern. Fronds
upright, narrow; pinnae short, to 5 cm (2 in.) long, wide,
blunt; pinnules narrowed at base.
‘Crispa Cristata’. Figure 13.47.28g. Frond apices and
pinnae crested, pinnules ruffled.
‘Cristata’. Figure 13.47.28c. Frond apices and pinnae
crested; many different types.
‘Grandiceps’. Figure 13.47.28d. Frond apices heavily
tasseled, pinna tips less so.
‘Linearis Congesta’. Figure 13.47.28e. Fronds to 37 cm
(15 in.), pinnae crowded, pinnules narrow. b
‘Linearis Polydactyla’. Slender crested male fern. Figure
13.47.28a, b. Blades broad and elliptic, the blade and pinna
apices crested, the segments finger-like, the segments of the a
pinnae skeletonized, mostly linear to nearly filiform. May
grow better in acidic soil.
‘Ramo Cristata’. Figure 13.47.28i. Fronds to 60 cm (24
in.), branched and crested.
‘Undulata-Robusta’. See Dryopteris complexa.
Figure 13.47.29. Dryopteris formosana: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
Dryopteris formosana (H. Christ) C. Christensen c. pinnule from medial pinna, bar = 5 mm (0.2 in.).
 13.47.29 After Hoshizaki and Wilson (1999).
Hardy, Zone (6)7
A medium-sized fern, to 45 cm (18 in.), with short-creep-
ing to erect rhizomes and evergreen fronds. Grows well
under medium light in moist garden soil or potting mix.
Dryopteris formosana has broad, pentagonal fronds up
to three-pinnate. The lowest basiscopic pinnule next to the
rachis is quite long, and the segment teeth are short-aristate
and tend to turn upward from the plane of the blade. The
scales on the costa undersides are weakly inflated, appearing
puffy (not flat). The species is native to Japan, Taiwan, and
the Philippines.

Dryopteris fragrans (Linnaeus) Schott a

 13.47.30
Very hardy, Zone 2
A small fern, to 20 cm (8 in.), with erect rhizomes and mar-
cescent fronds. Grows well under medium light in moist
garden soil or potting mix. This species does best in cold-
temperate areas. The plants grow in rock crevices and talus b
slopes in the wild.
Dryopteris fragrans has narrowly elliptic, one-pinnate-
pinnatifid blades, the stipes only about one-third as long Figure 13.47.30. Dryopteris fragrans: a. habit, bar = 5 cm (2
as the blades. The brown, withered fronds persist on the in.); b. stipe scales, bar = 1 cm (0.4 in.). After Hoshizaki
rhizome, sometimes accumulating as a thick mat. Yellow- and Wilson (1999).
ish, round glands dot the blade, especially on the lower sur-
face, and they emit a distinctive fragrance when crushed.
The indusia are circular, large, 1–2 mm wide, and overlap-
 F  F A  C 311

ping when young. The species is circumboreal, occurring in

North America, Europe, and Asia.
a
Dryopteris fuscipes C. Christensen  13.47.31
Semi-hardy (?)
A medium-sized fern, to 50 cm (20 in.), with ascending to
erect rhizomes and semi-evergreen fronds. Grows well
b
under medium light in moist garden soil or potting mix.
Dryopteris fuscipes has triangular blades up to two-pin-
nate with narrow pinnae and stout, oblong segments. The
scales on the lower surface of the costae are strongly in-
flated, appearing puffy (not flat). The species is native to
eastern Asia.

Figure 13.47.31.
Dryopteris fuscipes:
a. frond, bar =
5 cm (2 in.); b. Figure 13.47.32. Dryopteris goldiana: a. frond, bar =
stipe scales, bar = 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
1 cm (0.4 in.). Hoshizaki and Wilson (1999).
After Hoshizaki
and Wilson
(1999).
lie close to the midrib. The species is native to eastern
North America.

Dryopteris hondoensis Koidzumi  13.47.33

Very hardy, Zone 4
A medium-sized fern, to about 60 cm (2 ft.), with ever-
a green fronds and short-creeping, ascending, or erect rhi-
zomes that form offshoots. Grows under low to medium
light in moist to moist-dry garden soil or potting mix.
b Dryopteris hondoensis is very similar to D. erythrosora ex-
cept for its thinner leaf texture and short-stalked pinnae.
In addition, the undersides of the costae have nonbullate or
very weakly bullate scales. Plants with pinkish indusia are
named forma rubisora Sa. Kurata. The species is native to
Dryopteris goldiana (Hooker ex Goldie) A. Gray Japan, Korea, and China.
 13.47.32
Giant wood fern, Goldie’s fern Dryopteris intermedia (Muhlenberg ex Willdenow)
Very hardy, Zone 3(4) A. Gray  13.47.34
A large fern, to 130 cm (52 in.), with ascending to erect syn. Dryopteris spinulosa var. intermedia (Muhlenberg)
rhizomes and deciduous fronds. Grows well under low to Underwood
medium light in moist to moist-wet garden soil or potting Evergreen wood fern, glandular wood fern, fancy fern
mix. The blades are slightly bluish green. Very hardy, Zone 3
The stipe scales of Dryopteris goldiana are narrowly tri- A medium-sized fern, to 70 cm (28 in.), with erect rhi-
angular with thin, fragile, membranous margins and dark zomes and evergreen fronds. Grows well under low to me-
centers. The blades are mostly large, widely ovate, and one- dium light in moist garden soil or potting mix. The plants
pinnate-pinnatifid to two-pinnate, with wide truncate are easy to grow in temperate areas but are not suitable in
bases. The pinna apices are abruptly pointed, and the sori subtropical ones.
 312 C 13

a c Figure 13.47.34.
Dryopteris
intermedia:
a. frond, bar =
5 cm (2 in.);
b. stipe scales, bar
= 1 cm (0.4 in.).
b After Hoshizaki
and Wilson
(1999).

a b

Figure 13.47.33. Dryopteris hondoensis: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. pinnule from medial pinna, bar = 5 mm (0.2 in.);
d. lowest pinna, bar = 2 cm (0.8 in.).
b

Two characters will distinguish Dryopteris intermedia

from similar ones (such as D. carthusiana) in the north-
eastern United States. First, the lower surfaces of the blades,
especially the rachises and costae, are glandular pubescent,
the glands appearing as minute hairs with swollen, head-
like tips (a hand lens is needed to see this). Second, the in-
nermost pinnule on the basal pinnae is shorter than the ad-
jacent, outer ones (they are longer in similar species). The a
species is native to eastern North America.
c
Dryopteris kuratae Nakaike  13.47.35
Semi-hardy, Zone 6
A medium-sized fern, to about 1 m (3 ft.), with deciduous
fronds. Grows well under medium light in moist garden
soil or potting mix. Easy to grow.
Dryopteris kuratae bears one-pinnate-pinnatilobed Figure 13.47.35. Dryopteris kuratae: a. frond, bar = 5 cm
blades, and the lobes have wide V- or U-shaped sinuses. (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); c. medial
The sori are scattered close to margin. The species is native pinna, bar = 1 cm (0.4 in.). After Hoshizaki and Wilson
to eastern Asia. (1999).
 F  F A  C 313

This plant is often confused with Dryopteris pycnopter- as wide at the base as in the middle, and the pinnae are pin-
oides (H. Christ) C. Christensen of Japan, a larger plant natisect with many oblong, shiny segments. Segments have
with sori scattered over the pinnae. rounded apices bearing sharp teeth. The species is native
from eastern India to China and Taiwan.
Dryopteris lacera (Thunberg) O. Kuntze
 13.47.36
Hardy, Zone 5
Figure 13.47.37.
A medium-sized fern, to about 60 cm (2 ft.), with ascend- Dryopteris
ing to erect rhizomes and deciduous fronds. Grows well lepidopoda: a.
under medium light in moist garden soil or potting mix. frond, bar = 5 cm
This species is easy to cultivate. The normally deciduous (2 in.); b. stipe
fronds tend to be slightly evergreen in warmer climates. scales, bar = 1 cm
The broad lanceolate blades of Dryopteris lacera are one- (0.4 in.). After
pinnate-pinnatifid to two-pinnate and fertile only toward Hoshizaki and
the tip. The segments or lobes are entire or nearly so, unlike Wilson (1999).
many Dryopteris species that are lobed. At maturity the
fronds are glabrous, without scales on the rachises or costae.
The fertile pinnae are more contracted than the sterile and
wither early. The species is native to eastern Asia.

Figure 13.47.36.
Dryopteris lacera: a.
frond, bar = 5 cm (2 in.); a
b. stipe scales, bar = 1 cm
(0.4 in.). After Hoshizaki
and Wilson (1999).

b Dryopteris ludoviciana (Kunze) Small

 13.47.38
a Southern wood fern
Hardy, Zone (5)6
A medium-large fern, to 130 cm (52 in.), with short-creep-
ing to erect rhizomes that readily form offshoots. Grows
well under medium light in moist-wet garden soil or pot-
ting mix. It is semi-evergreen. The plants are easy to grow
Dryopteris lepidopoda Hayata  13.47.37 in southern California.
Hardy, Zone (5)6 The shiny dark green blades are one-pinnate-pinnati-
A medium-sized fern, to 1 m (3 ft.), with erect rhizomes fid, with the fertile pinnae only in the apical part of the
and evergreen fronds. Grows well under medium light in blade and markedly contracted. Dryopteris ludoviciana is
moist garden soil or potting mix. According to Mickel native to the southeastern United States.
(1994), the young fronds are reddish gold. This species hybridizes with Dryopteris celsa to form the
Dryopteris lepidopoda has relatively long stipes with nar- sterile hybrid D. ×australis.
row, black, ciliate scales. The blades are oblong-triangular,
 314 C 13

a
a

b
c
Figure 13.47.38. Dryopteris ludoviciana: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
Hoshizaki and Wilson (1999). Figure 13.47.39. Dryopteris marginalis: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. medial pinna, bar = 1 cm (0.4 in.). After Hoshizaki
and Wilson (1999).
Dryopteris marginalis (Linnaeus) A. Gray
 13.47.39
Eastern wood fern, marginal shield fern
Very hardy, Zone 2(3)
A medium-sized fern, to 60 cm (2 ft.), with erect rhizomes
and evergreen fronds. Grows well under medium light in
moist garden soil or potting mix. The plants are easy to
grow and have bluish green blades. They do not do well in
subtropical climates.
Dryopteris marginalis can be distinguished from many Figure 13.47.40. Dryopteris
other Dryopteris by having sori that are close to the margin. mindschelkensis: a. frond, bar =
The blades are one-pinnate-pinnatifid to two-pinnate and 5 cm (2 in.); b. stipe scales,
slightly leathery. The species is native to northeastern bar = 1 cm (0.4 in.). After
North America, where it grows on or among rocks. Hoshizaki and Wilson (1999).

Dryopteris mindshelkensis N. V. Pavlov

 13.47.40
syn. Dryopteris submontana (Fraser-Jenkins) Fraser-
Jenkins, D. villarii subsp. submontana Fraser-Jenkins
& Jermy a
Rigid buckler fern, limestone wood fern
Hardy, Zone (5)6
b
A medium-sized fern, to 60 cm (2 ft.), with ascending to
erect rhizomes and deciduous fronds. Grows well in moist,
basic garden soil or potting mix.
 F  F A  C 315

Dryopteris mindshelkensis has narrowly triangular fronds

densely covered on both surfaces with stalked, yellow
glands that emit a fragrance when crushed. The species is
native to Europe and northern Africa.

Dryopteris oreades Fomin  13.47.41

Mountain male fern
Hardy, Zone 4 (5–6)
A medium-sized fern, to 50 cm (20 in.), with deciduous
fronds and erect rhizomes forming many offshoots. Grows c
well under medium to high light in acidic, moist garden
soil or potting mix. The fronds arise in a strongly ascending
fashion, forming shuttlecock-like clusters.
Dryopteris oreades resembles a small D. filix-mas but dif-
fers by gray-green, ruffled blade margins, segment apices
with blunt teeth pointing away from the pinnule apex in a
fan-shaped pattern, stipe scales dull, and young indusia
thick and green. The species is native to the British Isles
and western Europe.

Dryopteris polylepis (Franchet & Savatier) b

C. Christensen  13.47.42
Scaly wood fern
Hardy, Zone (5)6 a
A medium-sized fern, to about 60 cm (2 ft.), with decidu-
ous fronds. Grows well under medium light in moist gar-
den soil or potting mix.
Figure 13.47.42. Dryopteris polylepis: a. frond, bar =
5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. pinnules with unbranched veins, bar = 2 cm (0.8 in.).
After Hoshizaki and Wilson (1999).

The pinnae are pinnatisect, and the veins are unforked,

except for those in the basiscopic segments next to the
rachis. Dryopteris polylepis is native to northeastern Asia.
c
Dryopteris pseudo-filix-mas (Fée) Rothmaler
 13.47.43
Mexican male fern
Hardy, Zone 5(6)
A medium-sized fern, to 1 m (3 ft.), with semi-evergreen
fronds and erect rhizomes forming offshoots. Grows well
under medium light in moist garden soil or potting mix.
The plants are easy to cultivate.
Dryopteris pseudo-filix-mas resembles D. affinis but dif-
fers by the basiscopic pinnule of the lowest pinnae usually
a b elongate and pinnatifid on older plants (those of D. affinis
are not elongated and are shallowly lobed). As in D. affinis,
a black spot is present on the lower surface of the costa base.
Figure 13.47.41. Dryopteris oreades: a. frond, bar = 5 cm It is native to Mexico and Guatemala.
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.); c. pinna, bar
= 1 cm (0.4 in.). After Hoshizaki and Wilson (1999).
 316 C 13

a a d

b
c

Figure 13.47.44. Dryopteris purpurella: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. lowest pinnae, bar = 2 cm (0.8 in.); d. pinnule from
Figure 13.47.43. Dryopteris pseudo-filix-mas: a. frond, medial pinna, bar = 5 mm (0.2 in.). After Hoshizaki and
bar = 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); Wilson (1999).
c. lowest pinnae with variations in cut of the basiscopic
pinnule, bar = 2 cm (0.8 in.). After Hoshizaki and
Wilson (1999).
Dryopteris remota (A. Braun) Druce  13.47.45
Scaly buckler fern
Very hardy, Zone 4
Dryopteris purpurella Tagawa  13.47.44 A medium-sized fern, to 1 m (3 ft.), with ascending to erect
Hardy, Zone 5 rhizomes and deciduous fronds. Grows well under medium
A medium-sized fern, to about 60 cm (2 ft.), with erect rhi- light in moist garden soil or potting mix. In the northeast-
zomes and evergreen fronds. Grows well under low to me- ern United States this species volunteers when planted in
dium light in moist garden soil or potting mix. gardens.
Dryopteris purpurella resembles D. erythrosora, but it dif- Dryopteris remota is of hybrid origin between D. affinis
fers by having purplish stipes and rachises; two-to three- and D. expansa. Both normal and aborted spores are pro-
pinnate-pinnatifid, smaller, coarser, and flatter blades; ses- duced. The blades are narrowly triangular-lanceolate, two-
sile, fewer (about six), and more distant pinnae; lower pinnate, and nonglandular. The pinnules are acute. The
pinnae at nearly right angles to the rachis; and pinnules species is native to Europe.
overlaying the rachis. The red color of the rachis and costae
fades slowly. The underside of the costa bears strongly in- Dryopteris sacrosancta Koidzumi  13.47.46
flated scales, and the sori are submarginal. The species is Hardy, Zone (6)7
native to Japan, Korea, and China. A medium-sized fern, to about 60 cm (2 ft.), with ascend-
Trade plants of Dryopteris erythrosora are often misiden- ing to erect rhizomes. Grows well under low to medium
tified as D. purpurella. light in moist garden soil or potting mix.
 F  F A  C 317

a
Figure 13.47.46.
Dryopteris
sacrosancta: a.
frond, bar = 5 cm
Figure 13.47.45. Dryopteris remota: a. frond, bar = 5 cm
(2 in.); b. stipe
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
scales, bar = 1 cm
Hoshizaki and Wilson (1999).
(0.4 in.). After
a Hoshizaki and
Wilson (1999).
Dryopteris sacrosancta has large stipe scales with pale bor-
ders, and the scales on the undersides of the costae are
weakly inflated (puffy) and dark tipped. The blades are tri-
angular, three-pinnate, slightly glossy and leathery, and flat
margined. The lowermost downward-pointing pinnule
next to the rachis is elongate. The species is native to Japan,
China, and Korea.

Dryopteris sieboldii (T. Moore) O. Kuntze

 13.47.47
Japanese wood fern, Siebold’s wood fern
Hardy, Zone 6
A medium-sized fern, to 50 cm (20 in.), with erect rhi-
zomes and semi-evergreen fronds. Grows well under me-
dium light in moist garden soil or potting mix. The plants
are somewhat slow growing and tend to emerge rather late
in the spring (late June in New York).
Dryopteris sieboldii is distinct from nearly all other wood
ferns because it has a terminal segment that resembles the
lateral pinnae (most wood ferns have a tapered pinnatifid b
apex). The blades are one-pinnate with two to seven large, a
leathery, oblong-lanceolate, entire pinnae. The species is
native to Japan, Taiwan, and China.

Dryopteris sichotensis V. Komarov  13.47.48

syn. Dryopteris coreano-montana Nakai
Hardy, Zone 5(?) or hardier (?) Figure 13.47.47. Dryopteris sieboldii: a. frond, bar =
A medium-large fern, to 130 cm (52 in.), with erect rhi- 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After
zomes forming offshoots and upright, semi-evergreen Hoshizaki and Wilson (1999).
 318 C 13

c Figure 13.47.49.
Dryopteris stewartii:
a a. frond, bar = 5 cm
(2 in.); b. stipe
scales, bar = 1 cm
(0.4 in.); c. basal
pinna, bar = 4 cm
(1.5 in.). After
Hoshizaki and
Wilson (1999).

Figure 13.47.48. Dryopteris sichotensis: a. frond, bar = c

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); c.
pinnules, bar = 5 mm (0.2 in.). After Kurata and
Nakaike (1979).
Dryopteris sublacera H. Christ  13.47.50
Hardy to semi-hardy, Zone 7
fronds. Grows under medium or stronger light in moist A medium-sized fern, to 30 cm (1 ft.), with evergreen fronds
garden soil or potting mix. and erect rhizomes forming offshoots. Grows well under
Dryopteris sichotensis resembles D. oreades, but its pin- medium light in moist garden soil or potting mix. The
nule teeth are slightly acute, not splayed out, the pinnule plants are easy to cultivate.
apices are more or less acute or rounded, the stipe scales Dryopteris sublacera bears abundant reddish to dark
more or less glossy, and the fronds yellow-green. The spe- brown, minutely fringed scales on the stipes and rachises,
cies is native to northeastern Asia. and these scales leave a roughened surface when shed. The
pinnules have rounded tips bearing short teeth. The species
Dryopteris stewartii Fraser-Jenkins  13.47.49 is native to India and eastern Asia.
Hardy, Zone 6
A medium-large fern, to about 130 cm (52 in.), with de- Dryopteris tokyoensis (Matsumoto ex Makino)
ciduous fronds and ascending to erect rhizomes that form C. Christensen  13.47.51
offshoots. Grows well under medium light in moist garden Tokyo wood fern
soil or potting mix. The plants are easy to grow. The nor- Hardy, Zone 5
mally deciduous fronds become semi-evergreen in warmer A medium-sized fern, to 1 m (3 ft.), with deciduous fronds
climates. and ascending rhizomes that form offshoots. Grows
The blades of Dryopteris stewartii are triangular-lanceo- well under medium light in moist, acidic garden soil or pot-
late and two-pinnate-pinnatifid, and the segments are ting mix. This species is not suited for gardens in southern
rounded and often bear spine-tipped teeth. The species is California.
native to the Himalayas. Dryopteris tokyoensis is characterized by narrow, erect
Plants circulating in the United States trade as Dryopteris fronds that form a slender, vase-shaped cluster. The blades
stewartii are foliose forms of D. carthusiana. True D. stew- are widest above the middle and taper gradually toward the
artii is circulating in the trade incorrectly as D. goeringiana base, with 20–40 pairs of pinnae; the basal-most pinnae are
Koidzumi, a species not cultivated in the United States. nothing more than highly reduced, ear-like lobes. The cen-
 F  F A  C 319

Figure 13.47.51. Dryopteris

tokyoensis: a. frond, bar = 5 cm
(2 in.); b. stipe scales, bar =
1 cm (0.4 in.). After Hoshizaki
and Wilson (1999).

b
c

Figure 13.47.50. Dryopteris sublacera: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.); c. basal
pinnae, bar = 2 cm (0.8 in.). After Hoshizaki and
Wilson (1999). d

tral pinnae are sessile or nearly so, with an auricle or “ear”

on both sides of the base. The species is native to China,
Japan, and Korea.

Dryopteris uniformis (Makino) Makino

 13.47.1, 52
Hardy, Zone 5 (6)
A medium-sized fern, to 60 cm (2 ft.), with deciduous
fronds and erect rhizomes that sometimes form offshoots. c
Grows in moist garden soil or potting mix. Although
deciduous, this species is easy to grow, even in warmer
climates.
Dryopteris uniformis is characterized by one-pinnate-
pinnatifid to two-pinnate blades and densely scaly stipes
and rachises. The scales are dark brown to black and have
entire or fringed margins. The sori are borne on the upper b
a
pinnae. The species is native to Japan, Korea, and China.
‘Cristata’. (Dryopteris uniformis forma cristata Ogata).
Figures 13.47.1, 52d. Frond apices sometimes forked, pin-
nae crested. Said to volunteer readily from spores. Briefly Figure 13.47.52. Dryopteris uniformis: a. frond, bar =
deciduous in southern California. 5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. medial pinna, bar = 2 cm (0.8 in.); d. ‘Cristata’, upper
part of frond, bar = 5 cm (2 in.). After Hoshizaki and
Wilson (1999), except d.
 320 C 13

Dryopteris varia (Linnaeus) O. Kuntze

 13.47.53
Hardy to semi-hardy, Zone 6(7)
A medium-sized fern, to 60 cm (2 ft.), with short-creeping
to ascending rhizomes and semi-evergreen fronds. Grows
well under medium light in moist garden soil or potting
mix. The plants are easy to cultivate but grow slowly. The
new fronds are bronze.
The blades of Dryopteris varia are triangular or pentag-
onal, abruptly narrowed in the distal third, stiff, thick,
leathery, and dull gray-green. The lowest basiscopic pin-
nule next to the rachis is the longest. The costal scales are c
weakly inflated, flattened, or shallowly convex. The species
is native to India and eastern Asia.

Dryopteris wallichiana (Sprengel) N. Hylander

 13.47.54;  27
syn. Dryopteris paleacea (Swartz) Handel-Mazzetti, D.
parallelogramma (Kunze) Alston b
Wallich’s wood fern
Hardy, Zone 5(6)
A medium-large fern, to 130 cm (52 in.), with semi-ever-
green fronds and erect rhizomes that form occasional off-
shoots. Prefers medium light and moist garden soil or pot- a

Figure 13.47.54. Dryopteris wallichiana: a. frond, bar =

5 cm (2 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. pinnules from medial pinnae, bar = 1 cm (0.4 in.).
After Hoshizaki and Wilson (1999).

ting mix. This species grows best in mildly cool, humid cli-
mates. The fronds are arranged in a handsome shuttlecock
pattern.
Dryopteris wallichiana is characterized by one-pinnate-
pinnatifid blades with 20–35 pairs of sessile pinnae. The
segments of the pinnae are squared or oblong. The rachis is
conspicuously covered by black, linear or thread-like scales.
The species is native to subtropical and tropical mountains
of the world.

a Elaphoglossum  13.48.1, 2

syn. Hymenodium, Peltapteris
Paddle fern
The genus Elaphoglossum consists of mostly medium-sized
ferns with simple, entire, often paddle-shaped blades. Fer-
b tile fronds are usually taller and narrower than the sterile
ones and are produced infrequently. The sori are acrosti-
choid, appearing as if spread across the lower surface of the
Figure 13.47.53. Dryopteris varia: a. frond, bar = 5 cm blade, not in discrete dots. Many species bear clustered
(2 in.); b. stipe scales, bar = 1 cm (0.4 in.). After fronds that are attractive in pots or baskets. Others are at-
Hoshizaki and Wilson (1999). tractive for their metallic hues or unusual covering of hairs
 F  F A  C 321

material is the lack of information about the country of ori-

gin. The genus name comes from the Greek elaphos, stag,
and glossa, tongue, alluding to the resemblance of the blades
to a deer’s tongue.

Elaphoglossum apodum (Kaulfuss) Schott ex J. Smith

 13.48.3
Tender
A small-medium fern with erect, clustered fronds. Grows
well under low to medium light in drained, moist potting
soil or uncut moss.
The rhizome scales of Elaphoglossum apodum are dense,
linear-attenuate, twisted, glossy, and pale orange-brown.
The stipes are nearly absent, and the blades are widest above
Figure 13.48.1. Elaphoglossum crinitum: habit.
the middle, about 2.5–4 cm (1–1.5 in.) wide, and thin-her-
baceous. Brownish orange, hair-like scales are found on
both surfaces of the blade. The fertile fronds are short and
narrow. This species is native to tropical America.
or scales. Most of the species are epiphytes and need green- Elaphoglossum erinaceum (Fée) T. Moore, which is rarely
house or indoor protection during the cold months. The cultivated, can be distinguished from E. apodum by longer
spores are often difficult to germinate. stipes and more oval blades.
Elaphoglossum species can usually be recognized by sim-
ple and entire fronds, free veins, and acrostichoid sori. The
rhizomes are scaly and creeping or, rarely, erect. In many
species the stipe base is jointed to a darker, stump-like ex-
tension (phyllopodium) of the rhizome. The blades vary in
shape from broadly ovate to linear; rarely are they pinnati-
fid or deeply dichotomously cut, as is the case with Elapho-
glossum peltatum. The blade tissue varies from glabrous to
densely scaly and is mostly firm to leathery.
Peltapteris differs from Elaphoglossum by its deeply cut
blades but is closely related to typical Elaphoglossum species
and therefore is treated here in the latter genus. The name
Hymenodium has been used for Elaphoglossum crinatum,
which is distinct by having netted veins, but this character-
istic alone is insufficient to distinguish another genus.
Elaphoglossum occurs in the tropics and subtropics of
the world, though most of the species are American. It con- Figure 13.48.3.
tains about 500 species, making it one of the world’s most Elaphoglossum
species-rich fern genera. Due to the large number, the spe- apodum: habit, bar
cies are difficult to identify, and many species have not re- = 5 cm (2 in.).
ceived scientific names. Another obstacle with cultivated

Figure 13.48.2. Elaphoglossum: sporangial area.

 322 C 13

Elaphoglossum crinitum (Linnaeus) H. Christ and heart-shaped. The blades of Elaphoglossum decoratum
 13.48.1, 4 are linear-lanceolate to oblong, dark, thick, leathery,
syn. Hymenodium crinitum (Linnaeus) Fée abruptly short-acuminate at the apex, and roundish to
Elephant-ear fern acute at the base. The surfaces of the blade are covered by
Tender minute, star-shaped scales; these are much less conspicu-
A small-medium fern with short-creeping rhizomes. Grows ous than the marginal scales. The species is native to trop-
best with year-round heat under low to medium light in ical America.
well-drained, moist-wet potting mix or uncut moss.
Elaphoglossum crinitum is a striking species, character- Elaphoglossum herminieri (Bory ex Fée) T. Moore
ized by large, oblong blades with scattered, erect, black  13.48.6
scales on both surfaces. Unlike most Elaphoglossum species, Tender
the veins are netted (without included veinlets). The species A medium to large fern with short-creeping rhizomes and
is native to the American tropics. often long, pendulous fronds. Grows under low to medium
light in drained, moist potting mix or uncut moss. The
Elaphoglossum decoratum (Kunze) T. Moore fronds are attractive for their bluish green color and pen-
 13.48.5 dulous habit.
Tender Elaphoglossum herminieri is characterized by pendent,
A small-medium fern with short-creeping rhizomes. Grows nearly sessile fronds up to 1 m (3 ft.) or more long. Linear,
under low to medium light in well-drained, moist potting metallic blue-green, leathery blades are dotted with minute,
mix or uncut moss. This species is difficult to grow. Showy star-shaped, glandular scales on both surfaces. The fertile
scales along the stipes and margins of the blades make for fronds, which are rarely produced, are rounded or kidney-
attractive foliage.
The scales on the stipes and blade margins are about 3.5
mm (about 0.1 in.) long, overlapping, yellowish brown,
Figure 13.48.5.
Elaphoglossum
decoratum: a. habit,
bar = 5 cm (2 in.);
b. marginal scale,
bar = 1 mm.

b a

Figure 13.48.4. Elaphoglossum

a crinitum: a. habit, bar = 5 cm a
(2 in.); b. marginal scales and
blade scales, bar = 1 mm.

Figure 13.48.6. Elaphoglossum

herminieri: a. fertile frond;
b. sterile frond. Bar = 5 cm
(2 in.).
 F  F A  C 323

shaped and much smaller than the sterile fronds. The spe- The creeping rhizomes of Elaphoglossum maxonii are 8–
cies is native to tropical America. 15 mm (0.3–0.6 in.) wide, with phyllopodia hidden among
the rhizome scales, the scales being narrowly triangular,
Elaphoglossum luridum (Fée) H. Christ golden or light brown, acuminate, and irregularly ciliate.
 13.48.7 The fronds are erect, leathery, and have ovate to narrowly
syn. Elaphoglossum schomburgkii (Fée) T. Moore elliptic blades, decurrent at the base. Minute and irregular
Tender star-shaped scales often with blackened centers are scattered
A medium-sized fern with short-creeping to nearly erect on both blade surfaces. The fertile fronds are smaller than
rhizomes. Grows well under low to medium light in well- the sterile ones. The species is native to the Greater Antilles.
drained, moist potting mix or uncut moss.
Elaphoglossum luridum is easy to recognize by its ob-
lanceolate, medium-sized blades and dark, shaggy scales on
the stipes and midribs. The lower surfaces of the blades
have scattered small scales, and the rachises have dark star-
shaped scales with twisted rays. The species is native to low- Figure 13.48.8. Elaphoglossum
land forests of the American tropics. maxonii: habit, bar = 5 cm (2 in.).

Elaphoglossum maxonii Underwood ex Maxon

 13.48.8
Semi-tender
A small to medium fern with medium-creeping rhizomes.
Grows well under medium light in drained, moist garden
soil or potting mix. This species is easy to cultivate.

Elaphoglossum metallicum Mickel  13.48.9

a Semi-tender
A medium-sized fern with short-creeping, branched rhi-
zomes and fronds close together. Grows well under me-
b dium light in drained, moist potting mix or uncut moss.
The fronds are attractive because of their iridescent, metal-
lic blue blades.
Elaphoglossum metallicum bears phyllopodia on its
Figure 13.48.7. Elaphoglossum luridum: a. fronds, bar = creeping rhizome. The stipes are densely covered with
5 cm (2 in.); b. base of fronds, showing dark scales, bar = blackened and some lighter colored scales, and the blades
5 cm (2 in.). are leathery and vary from lanceolate to broadly elliptic.
 324 C 13

The scales on the undersurface are scattered, dark reddish

brown, skeletonized, and about 1 mm long. There are fewer
on the upper surface, and they tend to fall away. This spe-
cies is known only from Peru.

Figure 13.48.9. Elaphoglossum

metallicum: frond, bar = 5 cm (2 in.).
b

Figure 13.48.10. Elaphoglossum peltatum: a. habit, with

fertile frond on left, sterile frond on right; b. forma
standleyi habit, with fertile frond on left, sterile fronds
on right. Bar = 5 cm (2 in.).

Elaphoglossum peltatum (Swartz) Urban

 13.48.10
syn. Peltapteris peltata (Swartz) C. V. Morton, Elaphoglossum sartorii (Liebmann) Mickel
Rhipidopteris peltata (Swartz) Fée  13.48.11
Tender Semi-tender or hardier
A small fern with thin, long-creeping rhizomes and erect, A small-medium fern with short- to medium-creeping rhi-
peltately divided fronds. Grows well under low to medium zomes and erect, clustered to slightly distant fronds. Grows
light in drained, moist potting mix or uncut moss. The well under medium light in moist garden soil or potting
plants are suitable for use in bottle gardens. mix. This species grows slowly and is often used as a ground
Elaphoglossum peltatum is easily distinguished by the cover or potted plant.
sterile blades typically deeply divided into narrow segments, The rhizome of Elaphoglossum sartorii bears distinct
although some forms are less divided. The rhizome and phyllopodia and is covered by linear-lanceolate, entire
stipe scales are broadly ovate, thin, and tan or pale brown. scales. The fronds are about 30 cm (12 in.) long and 5 cm
Unlike the other cultivated species of Elaphoglossum, this (2 in.) wide. The stipes are about one-quarter to one-third
species lacks phyllopodia. The fertile fronds are rounded the frond length, glabrous or sparsely scaly with irregularly
and slightly taller than the sterile. This species is widespread toothed scales. The blades are mostly elliptic, leathery,
in the American tropics. acuminate, the bases wedge-shaped, the surfaces glabrous
or with minute, black, star-shaped scales on the lower sur-
face. The fertile fronds are slightly narrower and taller than
the sterile ones. The species is endemic to Mexico.
Elaphoglossum sartorii varies greatly and probably con-
sists of several different species. The plant illustrated was
 F  F A  C 325

Figure 13.48.11. Elaphoglossum

sartorii: a. habit; b. mature frond.
Bar = 5 cm (2 in.).

a
b
collected in Mexico. Plants from elsewhere should be re-
ferred to the E. latifolium complex, which also has not been
fully studied. The rarely grown E. maxonii, also of the E.
latifolium complex, differs from E. sartorii by its thicker
rhizome and phyllopodia hidden by scales.

Elaphoglossum vestitum (Schlechtendal & Chamisso) Figure 13.48.12. Elaphoglossum vestitum: a. habit, with
Schott ex T. Moore  13.48.12 fertile frond on left, sterile fronds on right, bar = 5 cm
Semi-tender to tender (2 in.); b. blade scales, bar = 1 mm.
A small-medium fern with short-creeping rhizomes. Grows
well under low to medium light in drained, moist potting
mix or uncut moss.
Elaphoglossum vestitum has distinct phyllopodia and rhi- Equisetum  13.49.1
zome scales that are red-brown, shiny, linear-lanceolate, Horsetail, scouring rush
toothed, and 4–6 mm (0.2 in.) long. The stipes are slender Equisetum is usually planted around ponds, bogs, and
and covered with narrowly triangular, red-brown, ciliate marshes, or it can be grown in pots, tubs, and small dish-
scales. The narrowly elliptic, gray-green blades are densely gardens. Garden soil or potting mix are suitable if the plants
covered on the upper surface with whitish scales bearing are grown in pots and the soil is kept moist and drained. If
filiform rays. Similarly shaped hairs on the undersurface are the pots are set in standing water, a sand and peat-moss mix
reddish brown. The species is native to southern Mexico. should be used to discourage the soil from “souring.” Vig-
This species has been misidentified in horticulture as orously growing species such as Equisetum hyemale become
Elaphoglossum muelleri (Fournier) C. Christensen, which weedy in gardens and are difficult to eradicate because of
lacks phyllopodia and is not cultivated. their deep rhizomes. Equisetum is often used in Japanese
flower arrangements.
The plants can be propagated by dividing the rhizomes
or by burying the stem segments shallowly in the soil.
When buried in soil, new roots are eventually produced at
326 C 13

leaves united by their bases. The teeth on the sheath repre-

sent the tips of the leaves.
The common name of scouring rush was given to the
reed-like (unbranched) species of Equisetum by American
pioneers who used the silica-roughened stems to scrub pots
and pans. The common name horsetail was given to those
species with whorled branches, in allusion to their bushy,
bristly appearance resembling a horse’s tail. The genus
name comes from the Greek equis, horse, and seta, bristle.
Equisetum occurs worldwide, except New Zealand and
Australia, and contains 23 species. The section on “Fern
Allies” in Chapter 10 contains additional information on
the genus.

Equisetum arvense Linnaeus  13.49.2

Common horsetail, field horsetail
Very hardy, Zone (1)2
A medium-sized plant with creeping rhizomes that send up
irregularly spaced or clumped stems. Grows well under me-
dium to high light in moist to wet garden soil. The above-
ground stems are deciduous and have the general aspect of
fine, feathery, bright-green foliage. This species is weedy in
some parts of the United States and Europe.

Figure 13.49.2. Equisetum

arvense: a. sterile stem, bar =
5 cm (2 in.); b. habit of
dimorphic fertile stems with
cones, bar = 5 cm (2 in.).

Figure 13.49.1. Equisetum hyemale: stems.

the joints of the stem segments, and new stems soon follow.
This method of producing new growth is how sterile hy-
brids of Equisetum, which cannot reproduce by spores be-
cause they abort, can become locally abundant or spread to
distant sites where their parents do not grow. b
The genus is easy to recognize by its green, usually hol-
low, jointed stems and spore-bearing cones at the stem tips;
the cones, however, are not always present. The sheath
around the stem joints represents a whorl of greatly reduced
 F  F A  C 327

The sterile stems of Equisetum arvense have dense whorls Equisetum ×ferrissii Clute  13.49.4
of branches from the joints, and sheaths of the main, erect Ferriss’s scouring rush
stems tend to flare slightly outward and bear 8–12 teeth, Very hardy, Zone 3
some adhering in pairs. The fertile stems appear in the A medium-sized plant with creeping rhizomes that send up
spring before the sterile ones. They are smaller, nongreen, irregularly spaced or clumped stems. Grows under medium
and short-lived, withering soon after shedding their spores. to high light in moist-wet to dry garden soil. When grown
This species is widely distributed in temperate North in pots, this hybrid produces fewer stems compared to its
America, Europe, and Asia. parents.
Equisetum ×ferrissii is of hybrid origin between E. hye-
Equisetum diffusum D. Don  13.49.3 male and E. laevigatum, and its characteristics are interme-
Semi-tender and perhaps hardier, Zone (8)9 diate between those two species. For example, E. hyemale
A medium-sized plant with creeping, branching rhizomes. has evergreen stems, and E. laevigatum has deciduous ones;
Grows in medium to high light in moist to wet garden soil. the hybrid’s stems are intermediate, being evergreen toward
The plant’s closely placed stems produce an attractive pot the base but deciduous in the upper part. The stem sheaths
plant. of the hybrid are also intermediate: those on the lower stem
The aerial stems of Equisetum diffusum have branches are ashy-white with black borders above and below (as in E.
in regular or irregular whorls. The stems have four ridges, hyemale), but those on the upper part of the stem are green
with each ridge grooved and each side of the groove typi- with a black border only at the top (as in E. laevigatum).
cally lined with a row of closely set, minute, blocky saw The spores of the hybrid are whitish and aborted, not green
teeth (magnification needed). Ovoid-cylindrical cones may and full as in the parental species. Equisetum ×ferrissii is na-
be produced at the tips of the stem late in the growing tive to and widespread in North America.
season (July–November). Tattered late-season stems are
quickly replaced by new growth. This species is endemic
to the Himalayas.
Figure 13.49.4. Equisetum
×ferrissii: a. habit of small
plant, bar = 5 cm (2 in.);
a b. stem with cone, bar =
1 cm (0.4 in.); c. lower stem
sheaths, bar = 1 cm (0.4 in.).

a
b
Figure 13.49.3. Equisetum diffusum: a. habit of stems
from a young plant, bar = 5 cm (2 in.); b. outline of
stem cross-section, enlarged to show stem with grooves
in each of the four ridges.
b c
 328 C 13

Equisetum hyemale Linnaeus  13.49.1, 5 Equisetum laevigatum A. Braun  13.49.6
Scouring rush, rough horsetail, winter scouring rush Smooth horsetail
Very hardy, Zone 2 Very hardy, Zone 3
A medium to large plant with short- to long-creeping rhi- A medium-sized plant with creeping rhizomes that send up
zomes. Grows well under medium to high light in moist irregularly spaced or clumped stems. Grows well under me-
to wet garden soil. It is easy to grow but becomes weedy in dium to high light in moist-wet garden soil.
gardens and is difficult to eradicate because of its deep rhi- Equisetum laevigatum is characterized by smooth, usu-
zomes. When grown in full sun the plants are more likely to ally unbranched (although sometimes sparsely and irregu-
produce cones. larly branched, especially if injured), deciduous stems and
Equisetum hyemale is characterized by rough, evergreen green (not ash-colored) sheaths with a black rim. The teeth
stems, and sheaths with a wide, ash-gray band bordered by at the rim typically fall off early and are therefore absent.
a lower black band and an upper black rim. The stems are The cones are blunt at the apex. The species is native to
unbranched, but injury to the apex will result in lateral North America, often occurring in open, sunny habitats.
branches developing at some of the lower joints. The teeth
are 14 or more at a joint, but most of them fall off at matu-
rity. The cones have a minute point at the tip. This species
is native to North America, Europe, and Asia. It typically
forms large colonies along shaded riverbanks, streams, and
in railroad ballast.
The thick-stemmed selection from the Washington area
reverted to typical stem diameters when pot cultivated in
southern California, whereas the slender-stemmed variants
from Alaska maintained their distinctness.

a
b
b d

Figure 13.49.6. Equisetum laevigatum: a. habit of

a unbranched plant, bar = 5 cm (2 in.); b. habit of
branched plant, bar = 5 cm (2 in.); c. stem with cone,
bar = 1 cm (0.4 in.); d. sheath, bar = 1 mm.
Figure 13.49.5. Equisetum hyemale: a. habit, bar = 5 cm
(2 in.); b. stem with cone, bar = 1 cm (0.4 in.); c.
sheaths, bar = 1 cm (0.4 in.).
 F  F A  C 329

Equisetum scirpoides Michaux  13.49.7

Dwarf scouring rush
Very hardy, Zone 2
A small plant that grows in loose clumps of evergreen, curly
stems. Thrives under medium to high light in moist to wet
garden soil. The plants are cultivated in dish gardens or as
small novelty plants. Much of the trade material with this
name is actually Equisetum variegatum.
Equisetum scirpoides is easy to recognize by its small size,
clump-forming habit, and loosely twisted stems. Each
stem-joint bears three teeth. Unlike other species in the
genus, the stems are solid, not hollow. It is native to boreal
North America, Europe, and Asia.

b c b
a
Figure 13.49.7. Equisetum scirpoides: a. habit, bar = 5 cm
(2 in.); b. stem with cone, bar = 1 mm; c. sheath, bar =
1 mm. Figure 13.49.8. Equiseum telmateia: a. habit of sterile
stem, bar = 5 cm (2 in.); b. dimorphic fertile stem with
cone, bar = 5 cm (2 in.).

Equisetum telmateia Ehrhart  13.49.8

Giant horsetail
Semi-hardy, Zone 8 Equisetum variegatum Schleicher ex F. Weber & Mohr
A large plant with deciduous, branched, feathery stems that  13.49.9
form extensive colonies. Thrives under medium to high Hardy, Zone 5
light in moist-wet garden soil. The plants are generally too A small-medium plant with evergreen stems from short-
large and invasive for gardens. creeping rhizomes. Thrives under medium or high light in
Equisetum telmateia has deciduous stems with joints moist-wet garden soil. Cultivated plants are smaller, up to
producing a whorl of slender branches. The teeth that oc- 12.5 cm (5 in.) long, and more compact than those in the
cur at the rim of the sheath are pale or green at the base and wild. This species is useful for small dish or water gardens.
dark at the tip. The fertile stems appear in early spring and The stems of Equisetum variegatum are unbranched and
are tan or brown. They wither soon after they shed their minutely grooved, and the sheaths have 3–12 teeth with
spores. The species is native to North America, Europe, conspicuous white margins. The species is native to the
northern Africa, and western Asia. northern parts of North America, Europe, and Asia.
 330 C 13

b c
Figure 13.50.1. Gleichenia
dicarpa: habit, bar = 5 cm
(2 in.).

Figure 13.49.9. Equisetum variegatum: a. habit, bar =

5 cm (2 in.); b. stem with cone, bar = 1 cm (0.4 in.);
c. sheath, bar = 1 mm.

Gleicheniaceae  13.50.1, 2

Forking fern family
The family Gleicheniaceae includes five genera: Dicranop-
teris, Diplopterygium, Gleichenella, Gleichenia, and Stich-
erus. These genera are so distinctive in appearance and so poor, wet, acid soil, does not re-establish from mature
widespread and abundant in the tropics that they are dis- plants. The umbrella fern, a more delicate and dainty plant,
cussed here even though they are not cultivated in the is more difficult to grow. It requires damp conditions and
United States. Gleicheniaceae consist of about 140 species can be propagated by divisions thusly: a tip division with
in tropical and subtropical regions worldwide. Most of the two fronds and attached roots is carefully lifted and re-
species colonize poor soils and are too big and invasive for planted in a peaty soil mix to about 2.5 cm (1 in.) deep.
most greenhouses. But the family is eye-catching, and the The transplant is kept damp until established (Best 1980).
smaller-sized species can be cultivated, as they are in New Other growers have had further luck with the family.
Zealand and Australia. California hobbyists raised young plants of Gleichenia mi-
Members of this family, however, are difficult to culti- crophylla from spore in 1995, but the plants have grown
vate. Victorian fern-growers apparently had some success, slowly. The Royal Botanic Gardens at Kew, England, suc-
and one grower in Manchester, England, had an entire cessfully grew a species in a loose, humusy woodland soil,
greenhouse devoted to 13 species. The grower said he and New Zealand growers have cultivated Sticherus flabel-
placed the rhizomes in fibrous peat and silver sand, provid- latus (R. Brown) H. St. John (Figure 13.50.2). One grower
ing good drainage for the shallow roots. started young transplants in pure sand until established,
More recently, growers in Australia successfully raised then transferred them to a sand-soil-peat mix containing
the following species from spores: pouched coral fern, Glei- bone and blood meal.
chenia dicarpa R. Brown (Figure 13.50.1); umbrella fern, Although plants may be grown from spores, growth is
G. microphylla R. Brown; and G. rupestris R. Brown. The slow. Perhaps, like bracken fern (Pteridium), transplants of
growers reported that the pouched coral fern, which prefers Gleicheniaceae do not establish readily but grow well once
 F  F A  C 331

on its fronds, and for this reason is widely used in Japan for
New Year’s Day decoration. In parts of southeastern Asia,
baskets are woven from the tough, flexible rachises of several
species. Sticherus flabellatus (R. Brown) H. St. John, S. lo-
batus N. A. Wakefield, and Dicranopteris linearis (N. L. Bur-
man) Underwood are harvested in Australia and exported to
Japan and elsewhere for cut-foliage use.

Goniophlebium
syn. Polypodium, in part; Schellolepis
Most gardeners know the genus Goniophlebium as Poly-
podium. It differs, however, by having jointed pinnae and
goniophleboid venation—that is, several rows of regularly
shaped, angular areoles with one free, included veinlet
pointing toward the pinna margin. Some authors disregard
the jointed pinnae and instead emphasize the presence of
scattered, black, fibrous strands in the rhizome, clathrate
rhizome scales, and (usually) hairy and scaly paraphyses in
the sori. In this book, the species with nonjointed pinnae
Figure 13.50.2. Sticherus are left in Polypodium (P. amoenum and P. formosanum),
flabellatus: habit, bar = except for the hybrid Goniophlebium ×ekstrandii. The
5 cm (2 in.). plants with long fronds, such as the knight’s polypody (Go-
niophlebium subauriculatum ‘Knightiae’), are handsomely
displayed in hanging baskets, where their fronds may reach
2.6 m (8.5 ft.) long.
The rhizome of Goniophlebium is creeping and often
glaucous, and when viewed in cross section, many scat-
tered, black fibers appear among the circle of vascular bun-
dles. The rhizome scales are clathrate, and the blades vary
from pinnatifid to one-pinnate. The pinnae are jointed,
rarely scaly, with entire or toothed margins. The veins are
netted, and the areoles have a regularly angular shape, those
rooted. Because the United States Department of Agricul-
along the midribs large and containing a veinlet that points
ture requires that soil be removed from imported plants,
outward (toward the pinna margin). The sori are round-
any imported Gleicheniaceae stand little chance of success.
ish, in one row on either side of the costa, and terminal on
It is generally thought that gleichenias need a mycorrhizal
the veinlet contained within the areole. Usually hairs or
fungus in their roots to grow. The successes with spore-
scales are mixed among the sporangia. Like all polypodia-
grown plants, however, raise questions about this need.
ceous ferns, this genus lacks indusia.
Members of this family can usually be recognized by
Goniophlebium, which contains about 23 species, is na-
their forked pinnae with a dormant bud in the fork’s angle.
tive to Asia, Australia, and some Pacific Islands. The genus
In nature the plants are terrestrial, not epiphytic, although
name comes from the Greek gonia, angled, and phleps, a
their leaves may rest on other plants for support. The rhi-
vein, referring to the angular mesh of veins.
zomes are always long-creeping, and the fronds are scram-
bling or trailing, one to many times forked, with the stipe
containing a single, C-shaped vascular bundle. The pinnae Goniophlebium ×ekstrandii Hoshizaki
are one to several times forked and have free veins. The sori  13.51.1
are always round, located on the undersurface of the leaf, Semi-tender
and lack an indusium. A medium-large fern with long-creeping rhizomes. Grows
In Alabama and Florida the native Dicranopteris flexuosa under medium light in moist, drained potting mix.
(Schrader) Underwood has not become well established, Goniophlebium ×ekstrandii has one-pinnate fronds with
and colonies are known to have died out. The Hawaiian pinnae broadly attached to the rachis. The foliage is inter-
Islands have several native species of gleichenoids, some of mediate in shape between that of its parents, Polypodium
which form extensive stands. formosanum and Goniophlebium subauriculatum. This ster-
Plants of this family have only a few uses. A Japanese spe- ile hybrid is known only in horticulture, and its cultivar
cies of Diplopterygium has yearly growth that is well marked name is ‘Nola’.
 332 C 13

a a

Figure 13.51.1. Goniophlebium ×ekstrandii ‘Nola’:

a. frond, bar = 5 cm (2 in.), after Hoshizaki (1991); Figure 13.51.2. Goniophlebium percussum: a. frond, bar
b. pinna, bar = 1 cm (0.4 in.). = 5 cm (2 in.); b. pinna, bar = 1 cm (0.4 in.), after
Hoshizaki (1982); c. sorus circled with dark scales, bar =
1 mm, after Hoshizaki (1982).
Goniophlebium percussum (Cavanilles) W. H. Wagner
& Grether  13.51.2
syn. Goniophlebium verrucosum (Hooker) J. Smith,
Polypodium verrucosum (Hooker) Mettenius
Tender
A large fern with short- to medium-creeping rhizomes.
Grows well under medium light in moist, drained potting
mix. With marginal tropical conditions, the fronds tend to
remain short.
Goniophlebium percussum bears fronds that are long,
pendulous, and one-pinnate. The pinnae are jointed to the
rachis, wedge-shaped at the base, and short-stalked. Fine
white needle-like hairs are found on the pinna surface. The a
terminal pinna resembles the lateral ones or is longer.
The sori are deeply sunken, circled with darkly toothed,
stalked scales. The species is native from southeastern Asia
b
to Malaysia, Indonesia, New Guinea, Australia, and the
Philippines.

Goniophlebium persicifolium (Desvaux) Beddome

 13.51.3
syn. Polypodium persicifolium Desvaux Figure 13.51.3. Goniophlebium persicifolium: a. frond,
Tender bar = 5 cm (2 in.); b. pinna, bar = 1 cm (0.4 in.).
A large fern with short- to medium-creeping rhizomes.
Grows well under medium light in moist, drained potting
mix.
 F  F A  C 333

Goniophlebium persicifolium resembles G. percussum but The fronds of Goniophlebium subauriculatum are up to
differs by lacking needle-shaped hairs on the blades. In ad- 2 m (6.5 ft.) long and one-pinnate with a pinnatifid apex.
dition, the sori of this species are only slightly sunken, There are many pinna pairs and these are jointed, mostly
wider than 1.5 mm, and lack the ring of dark scales. Gonio- cordate at the base, and sessile or nearly so. The species is
phlebium persicifolium is native from India to China, south- native to China, southeastern Asia, Indonesia, Australia,
eastern Asia to Malaysia, Indonesia, and the Pacific Islands. and the Philippines.
‘Knightiae’. (Polypodium knightiae Horticulture).
Knight’s polypody. Figure 13.51.4c; Plate 28. Pinnae ir-
Goniophlebium subauriculatum (Blume) C. Presl
regularly and deeply incised. The plants tolerate more cold
 13.51.4
(semi-tender) and more irregular watering than the species.
syn. Polypodium subauriculatum Blume
Jointed polypody
Tender Grammitidaceae  13.52.1
A large fern with short- to medium-creeping rhizomes. Grammitid family
Grows well under medium light in moist, drained potting Although not cultivated in the United States, the family
mix. The fronds are long and pendulous, and the old fronds Grammitidaceae is included here because of its large size
are shed in early to mid-spring as new ones emerge. and horticultural potential. Most plants in the family are
small, tropical epiphytic ferns. Their attractive frond shapes
and delicate aspect are reminiscent of tiny polypodiums;
other species have fronds that resemble narrow strips
Figure 13.51.4. Goniophlebium trimmed by pinking shears. Among the genera in this fam-
subauriculatum: a. frond, bar = ily are Acrosorus, Adenophorus, Calymmodon, Cochlidium,
5 cm (2 in.); b. pinna, bar = Ctenopteris, Grammitis, Lellingeria, Melpomene, Micropoly-
1 cm (0.4 in.), after Hoshizaki podium, Prosaptia, Scleroglossum, Terpsichore, Themelium,
(1982); c. ‘Knightiae’ frond, and Zygophlebia. They are important in the tropics, espe-
bar = 5 cm (2 in.). cially in montane and cloud forests. Some species occur in
subtropical or warm-temperate forests where humid con-
ditions prevail. The Grammitidaceae contain about 450
species.
Although a few species were grown in Victorian times
and some have been recommended for cultivation, no one
seems to have cultivated the plants long-term. Usually they
languish in terrariums and eventually die. Their cultural
requirements are unknown. The proper combination of air
movement, humidity, and light seem to be the main factors
affecting their growth, but nutrients and mycorrhizal fungi
might also be important.
The senior author has established Cochlidium serrulatum
(Swartz) L. E. Bishop in a 3.8 liter (1 gal.) glass bottle (may-
onnaise jar type) covered with a piece of glass. Medium in-
direct light is supplied through a large north-facing win-
dow. The fern grows in moist uncut sphagnum moss, which
was lightly firmed into a 60-degree angle against one side of
the bottle. Room temperature ranges from 16 to 24°C
(60–75°F). Ordinary tap water is used but seldom required
c in the covered container. The plant appeared as a volunteer
in a small clump of moss (collected in Costa Rica) that was
planted at the top of the sphagnum moss slope. The plant is
now over a year old, thriving and producing spores.
a The plants resemble small polypodiums because of their
pinnatifid or one-pinnate blades and round sori that lack
b indusia. They differ by having green, tetrahedral spores
(not yellow, bean-shaped ones) and, usually, the rhizomes
are erect or short-creeping, not long-creeping, as in many
polypodiums.
 334 C 13

b c

d
a

Figure 13.53.1. Gymnocarpium disjunctum: habit.

f
e g

Figure 13.52.1. Grammitidaceae examples: a. Grammitis

palauensis, from Palau; b. Grammitis from Costa Rica;
c. Ctenopteris blechnoides, from Palau; d. Enterosora
trifurcata, from Puerto Rico; e. Grammitis adspersa,
from Malaya; f. Cochlidium graminoides (?), from Brazil;
g. Adenophorus pinnatifidus, from Hawaii. Bar = 5 cm
(2 in.).

Gymnocarpium  13.53.1, 2

Oak fern
These small- to medium-sized ferns are eye-catching be-
cause of their broadly triangular to pentagonal fronds,
which are spaced far enough apart on the rhizomes to show
off their pattern. The common oak fern (Gymnocarpium
dryopteris) found on the floors of temperate forests is used as
a ground cover or bed fern in the eastern United States and
is considered invasive by some gardeners.
Gymnocarpium can be distinguished by the combina-
tion of slender, long-creeping rhizomes, stipes with only Figure 13.53.2. Gymnocarpium disjunctum: sori.
 F  F A  C 335

two vascular bundles, broadly triangular to pentagonal Gymnocarpium dryopteris (Linnaeus) Newman
blades, and round (rarely elliptic), non-indusiate sori. The  13.53.4;  29
blades vary from pinnatifid to three-pinnate-pinnatifid and syn. Polypodium dryopteris Linnaeus
may have minute glands, but hairs are lacking. There is usu- Common oak fern
ally a slight swelling where the pinna joins the rachis. The Very hardy, Zone (3)4
veins are free, not netted. A small fern, 15–25 cm (6–10 in.) tall, with slender, long-
The genus consists of five species and is native to North creeping rhizomes and deciduous fronds. Thrives under
America, Europe, and Asia. Its name comes from the Greek low to medium light in moist garden soil or potting mix. Be
gymnos, naked, and karpos, fruit, referring to the sori careful not to allow the soil to dry.
(“fruit”) that lack an indusium. Gymnocarpium dryopteris has blades that are triangular-
pentagonal, two-pinnate-pinnatifid, 3–14 cm (1.2–5.5 in.)
Gymnocarpium disjunctum (Ruprecht) Ching long, and glabrous. The second pinna pair from the base is
 13.53.1–3 sessile, and the basal pinnules on both sides are essentially
Western oak fern equal in length. The sori are round. The species is native to
Very hardy, Zone 4 northern North America, north-central Europe, and north-
A medium-sized fern with slender, long-creeping rhizomes ern Asia to China and Japan.
and distant, deciduous fronds. Prefers low to medium light ‘Plumosum’. Pinnules thin, delicate, dense, the margins
in moist garden soil or potting mix. This species is unsuit- turned under.
able for areas with warm winters.
The blades of Gymnocarpium disjunctum are broadly tri- Gymnocarpium oyamense (Baker) Ching
angular-pentagonal, three-pinnate-pinnatifid, 8–24 cm  13.53.5
(3–9 in.) long, and glabrous. The second pinna pair from syn. Currania oyamensis (Baker) Copeland
the base is sessile, and its basiscopic basal pinnule is longer Hardy
than the acroscopic one. The sori are round. The species is A small-medium fern with slender, long-creeping rhizomes
native to northeastern Asia and the northwestern coast of and distant, deciduous fronds. Grows well under medium
North America. light in moist garden soil or potting mix.

Figure 13.53.3. Gymnocarpium disjunctum: a. frond, bar = 5 cm (2 in.); b. frond base, showing uneven basal pinnules in
second pinna pair, bar = 2 cm (0.8 in.).
336 C 13

Gymnocarpium oyamense has triangular-ovate, deeply

pinnatifid or one-pinnate blades and round to elliptic sori.
It is native to Nepal, China, Taiwan, Japan, the Philip-
pines, Moluccas, and New Guinea.

Gymnocarpium robertianum (Hoffmann) Newman

 13.53.6
Limestone oak fern
a Very hardy, Zone 3
b A small-medium fern with slender, long-creeping rhizomes
and distant, deciduous fronds. Thrives under low light in
basic, moist or moist-wet garden soil or potting mix.
The blades of Gymnocarpium robertianum are broadly
triangular and two- to three-pinnate-pinnatifid. Both blade
surfaces are minutely glandular (hand lens needed). The
second pinna pair from the base is usually stalked. The sori
are round. This species is native to eastern Canada and the
Figure 13.53.4. Gymnocarpium dryopteris: a. habit, bar = north-central United States, Europe, and Asia (Caucus
5 cm (2 in.); b. frond base, showing even basal pinnules Mountains). It grows on calcareous rocks such as limestone
in second pinna pair, bar = 2 cm (0.8 in.). and dolomite.

Hemionitis  13.54.1, 2

syn. Gymnopteris, Parahemionitis
Hemionitis is a genus of small, terrestrial or rock-inhabiting
tropical ferns. The best-known species is the strawberry fern
(Hemionitis palmata), named for a fancied resemblance of
the leaves to those of a strawberry plant. It is occasionally

Figure 13.53.5. Gymnocarpium oyamense: a. frond, bar = Figure 13.53.6. Gymnocarpium robertianum: a. frond,
5 cm (2 in.); b. sori, bar = 2 cm (0.8 in.). bar = 5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.).
 F  F A  C 337

times with a few scales, and they vary from simple and en-
tire to deeply palmately or pinnately lobed to one-pinnate
(noncultivated species can be up to three-pinnate). The
veins are either free or netted, and if the latter, the areoles
lack included veinlets. The sporangia run along the veins;
an indusium is absent. Some species of Hemionitis might be
confused with Bommeria, but the sori in that genus run
along the margin, not between the margin and midrib as in
Hemionitis.
The genus consists of seven species native to the Amer-
ican tropics. Hemionitis arifolia, which questionably be-
longs here, occurs in Asia. The genus name comes from the
Greek hemionos, mule, and is of uncertain derivation.

Hemionitis arifolia (N. L. Burman) T. Moore

 13.54.3;  30
syn. Parahemionitis arifolia (N. L. Burman) Panigrahi
Tender
A small fern with short-creeping to suberect rhizomes and
fronds bearing buds at the base of the blade where it joins
the stipe. Grows well under medium light in moist potting
mix.
Hemionitis arifolia has dimorphic fronds: the sterile
Figure 13.54.1. Hemionitis palmata: habit. blades are heart-shaped; the fertile, triangular-hastate. Both
fronds have hairs on the stipes and thinly on the blades,
and the veins are netted with polygonal, elongate areoles.
The species is native to the Old World tropics.
seen in fern collections, as is the closely allied H. pinnati- Some pteridologists classify this species in its own genus,
fida. The two species are attractive for their broad, maple- Parahemionitis (Panigrahi 1993). Its evolutionary relation-
like fronds. Both are grown in pots or terrariums or, in ships with the New World species are questionable and re-
tropical climates, outdoors. They are easy to cultivate but quire more study.
are often attacked by scale insects, slugs, and mealybugs.
The buds produced on the blade bases root readily when
layered, or when large enough, they may be detached and b
transplanted.
The rhizomes are scaly and vary from erect to ascending.
The stipes are often shiny and chestnut-brown to straw-
colored. The fronds are herbaceous, often hairy and some-

Figure 13.54.3. Hemionitis arifolia: a. fronds, bar = 5 cm

Figure 13.54.2. Hemionitis palmata: sporangia along (2 in.); b. sporangia along veins, bar = 1 cm (0.4 in.).
veins. After Beddome (1864).
 338 C 13

Hemionitis palmata Linnaeus  13.54.1, 2, 4

Strawberry fern
Tender
A small fern with short-creeping to suberect rhizomes and
bud-bearing fronds. Grows well under medium light in
moist potting mix.
In Hemionitis palmata, the sterile and fertile blades are
palmately lobed and similar in size and shape. Usually one
bud develops in a large marginal notch on the blade. The
veins are netted. This species is native to Central and South
America and the West Indies.

Figure 13.54.5. Hemionitis pinnatifida: fronds, bar =

5 cm (2 in.).

Histiopteris
Bat fern, bat-winged fern
Histiopteris is a medium- to large-sized terrestrial fern in-
frequently cultivated. Because of its long-creeping rhi-
zomes, it is best confined to large pots or tubs. The com-
mon name alludes to the wing-like appearance of the sessile
pinnules. Histiopteris incisa is widely distributed in the trop-
Figure 13.54.4. Hemionitis palmata: fronds, bar = 5 cm ics and subtropics and seems to vary in size, degree of blue-
(2 in.). gray greenness, and adaptability to cold.
Histiopteris is easily recognized by long-creeping rhi-
zomes; opposite, sessile (or nearly so) pinnae and sessile
pinnules; and enrolled leaf margins that serve as indusia.
Hemionitis pinnatifida Baker  13.54.5 The fronds are distant on the rhizome, and their tips grow
Tender continuously, never completely unfurling. The stipe con-
A small fern with short-creeping to suberect rhizomes and tains one vascular bundle, and the blades are two-pinnate or
bud-bearing fronds. Grows well under medium light in more divided, often glaucous beneath. The basal pinnules
moist potting mix. of each pinna are opposite, often reduced and touching the
Hemionitis pinnatifida resembles H. palmata and is of rachis. The veins vary from free to netted. The sori run
hybrid origin between it and another unknown species in along the blade margin.
the genus. Hemionitis pinnatifida is distinguished by deeply The genus is south-temperate to pantropical (mostly
pinnatifid terminal lobes and several buds that usually de- Malaysia) and contains five species. The genus name comes
velop in smaller marginal notches. It is native to Central from the Greek histion, sail, and pteris, fern, alluding to
America. shape of the basal pinnae.
 F  F A  C 339

Histiopteris incisa (Thunberg) J. Smith

 13.55.1
Bat fern
Semi-tender to semi-hardy or hardier
A large fern with long-creeping rhizomes and fronds about
6 cm (2.5 in.) apart. Grows well under medium to high
light in moist garden soil. This species often volunteers in
spore cultures and is sometimes weedy in its native habitat.
Plants from different geographical areas appear variable in
blade cutting and tolerance to cold.
Histiopteris incisa has all the characteristics given above
for the genus, except that the veins are netted. It is widely
distributed in the tropics and southern subtropics of the
world.

Hymenophyllum  13.56.1, 2

syn. Mecodium, Sphaerocionium
Filmy fern
The filmy ferns, or “filmies,” are mostly small epiphytes na-
tive to humid habitats in the tropics. They are rarely culti-
vated but can be used to good effect in terrariums. The

Figure 13.56.1. Hymenophyllum demissum: habit.

most commonly grown species are those adapted to cool,

humid climates. Some species adapt to cultivation readily
but require vigilance in tending to their moisture and tem-
perature needs. Where hot summer temperatures prevail,
cultivation is difficult but may be improved by using a mist
b

Figure 13.56.2.
a Hymenophyllum: sorus
and indusium.
Figure 13.55.1. Histiopteris
incisa: a. frond, bar = 5 cm
(2 in.); b. pinnule with sori
and indusia, bar = 1 cm
(0.4 in.).
 340 C 13

system. The delicate fronds of Hymenophyllum are only one

cell-layer thick except around the veins.
The rhizomes are creeping, slender, and hairy (not
scaly). The thin fronds are mostly small, with blades simple
to four-pinnate. The sori are marginal and surrounded by
a two-lipped or two-valved indusium. The receptacle is
(usually) included within the indusium, not protruding.
The spores are green and short-lived.
Only two genera of filmy ferns—Hymenophyllum and
Trichomanes—are recognized in this book, following the
practice of most pteridologists. A few pteridologists, how-
ever, divide these genera into many smaller ones (see the a
discussion under Trichomanes). Hymenophyllum can be dis-
tinguished from Trichomanes by the form of the indusium:
two-valved or lipped in Hymenophyllum, and tubular or
funnel-shaped in Trichomanes.
Hymenophyllum is native to the tropics and south-tem-
perate areas of the world. It contains about 300 species.
The genus name comes from the Greek hymen, membrane,
and phyllon, leaf, referring to the membrane-like texture of
the blades. For more information, see the section “Filmy
Figure 13.56.3.
Ferns” in Chapter 10.
Hymenophyllum
b demissum: a. frond
Hymenophyllum demissum (G. Forster) Swartz (from native habitat),
 13.56.1, 3 bar = 5 cm (2 in.); b.
syn. Mecodium demissum (G. Forster) Copeland sori and indusia, bar =
Semi-tender or hardier 5 mm (0.2 in.).
A small fern with long-creeping rhizomes. Grows under
low light in well-drained, moist to moist-wet potting mix
or uncut moss. Keep the plants cool and humid during hot
weather.
Hymenophyllum demissum has glabrous, three- or four- b
pinnate, mostly elliptic or ovate, 7–25 cm (3–10 in.) long
blades and unwinged stipes. The sori are often in pairs. The
species is endemic to New Zealand.

Hymenophyllum polyanthos (Swartz) Swartz

 13.56.4
syn. Mecodium polyanthos (Swartz) Copeland
Tender
A small fern with medium-creeping rhizomes. Grows un-
der low light in well-drained, moist to moist-wet potting
mix or uncut moss under humid conditions.
Hymenophyllum polyanthos has unwinged stipes (or
winged only toward the summit) and glabrous, three-pin-
nate blades. The sori are not in pairs. The species is native
to the tropics and subtropics of the world.
a
Hypoderris
This genus has trilobed fronds reminiscent of Tectaria, to
which it is related. Hypoderris differs from Tectaria only by
the indusium, which encircles the base of the sorus and is
strongly ciliate, as opposed to the indusia of Tectaria, which Figure 13.56.4. Hymenophyllum polyanthos: a. habit, bar
are absent or attached on one side of the sorus and arching = 5 cm (2 in.); b. sori and indusia, bar = 5 mm (0.2 in.).
 F  F A  C 341

above it. The indusium can be hard to see because it is often

appressed to the leaf surface and covered by sporangia.
Hypoderris consists of a single species native to moist
habitats of the West Indies and northern Venezuela. In the
United States it is mainly cultivated outdoors in southern
Florida, either in soil or pots. The genus name comes from
the Greek hypo, below, and derris, covering, referring to the
indusium below the sporangia.

Hypoderris brownii J. Smith  13.57.1

Tender
A small-medium fern with short- to medium-creeping rhi-
zomes. Grows well under low to medium light in moist gar-
den soil or potting mix. The plants respond well to nutri-
ents but require greenhouse conditions in all but humid
tropical climates. Figure 13.58.1. Hypolepis tenuifolia: habit.
Hypoderris brownii has scaly rhizomes that creep over
soil or rocks. Its blades vary from simple to trilobed, with
in parks of central and southern coastal California. Their
the central lobe the largest. The veins are netted with free,
spreading habit can be controlled by planting in large tubs.
included veinlets in the areoles. Native to moist habitats of
New plants are easily propagated by divisions or spores.
the West Indies and northern Venezuela.
The best features for recognizing Hypolepis are the long-
creeping, branched, hairy (not scaly) rhizomes, the finely
Hypolepis  13.58.1, 2 divided, large, triangular fronds, and the roundish sori usu-
Ground fern ally protected by tooth-like indusia formed by the blade
Hypolepis has long-creeping rhizomes and finely divided, margins (rarely the sori are below the margin and unpro-
triangular fronds. These medium- to large-sized ferns are tected). In addition, the blades are two-pinnate-pinnatifid
used as ground cover or, in warmer climates, as foundation to four-pinnate-pinnatifid, thin, and often hairy.
plantings. They are sometimes seen growing in thick stands

c
a

Figure 13.57.1. Hypoderris brownii:

a. habit, bar = 5 cm (2 in.); b. sori,
bar = 5 cm (2 in.); c. sorus above Figure 13.58.2. Hypolepis tenuifolia: sori and
indusium, bar = 1 mm. indusia.
 342 C 13

The genus is native to the tropics and subtropics of the

world and contains about 50 species. Hypolepis millefolium
Hooker, a smaller species from New Zealand, is cultivated
in the mild areas of England. Hypolepis punctata (Thun-
berg) Mettenius, an easily grown species introduced from
Taiwan, is rarely seen in cultivation.
b
Hypolepis repens (Linnaeus) C. Presl  13.58.3
Bramble fern
Semi-tender
A medium-large fern with long-creeping rhizomes. Grows
well under medium light in moist garden soil or potting
mix. This species is easy to grow. a
The stipes and rachises of Hypolepis repens are spiny and
vary from glabrous to hairy. The indusia are thin and often
inconspicuous. This species is native to Florida, Mexico to
South America, and the West Indies.
Most of the horticultural material in Florida identified
as Hypolepis repens is misidentified and actually represents Figure 13.58.4. Hypolepis tenuifolia: a. habit, bar = 5 cm
H. tenuifolia, which differs by lacking prickles on the stipes (2 in.); b. pinna, bar = 5 cm (2 in.).
and rachises.

Hypolepis tenuifolia has smooth stipes and rachises, with-

out prickles. The rhizomes are more than 5 mm (0.2 in.)
thick. The stipes and blades bear gland-tipped hairs, and
the sori are protected by tooth-like false indusia. The spe-
cies is native to Asia and Polynesia.
This species name is believed to apply to several differ-
ent species yet to be fully sorted out.

Lastreopsis  13.59.1–3

b The mostly medium-sized terrestrial ferns of genus Lastre-
opsis have broad, highly divided fronds. Some Australian
a and New Zealand species are suitable as a bedding plant in
warm-temperate climates. They are easy to grow and have
durable foliage. Lastreopsis microsora has been used for years
c
as bedding plants in southern California, although some
gardeners do not care for the dull, gray-green color of this
species. The creeping rhizome is moderately slow-growing
and easy to control. Lastreopsis acuminata is currently cir-
Figure 13.58.3. Hypolepis repens: a. pinna, bar = 5 cm culating among a few hobbyists and promises to be an ex-
(2 in.); b. rachis with small spines, bar = 1 cm (0.4 in.); cellent ornamental because of its fast growth and clustered,
c. sori and indusia, bar = 5 mm (0.2 in.). shiny leaves.
The rhizomes of Lastreopsis are creeping to nearly erect
and scaly. The stipes contain three or more vascular bun-
dles. The blades are usually pentagonal or triangular, several
Hypolepis tenuifolia (G. Forster) Bernhardi times pinnate, and the lowest pinna pair is enlarged on
 13.58.1, 2, 4 the lower side. The upper surface of the rachis and costae
Semi-tender or hardier bear short hairs. The veins are free, not netted. The sori
A large fern with long-creeping rhizomes. Grows well are round and, in most species, covered by a reniform
under medium or high light in moist garden soil or pot- indusium.
ting mix. This fern is easy to cultivate and is perhaps the Lastreopsis can be distinguished from similar genera such
most common species of Hypolepis cultivated in the United as Ctenitis, Dryopteris, and Tectaria by the upper surface of
States. the rachis, which has two thickened ridges and is densely
 F  F A  C 343

Figure 13.59.3. Lastreopsis microsora subsp.

Figure 13.59.1. Lastreopsis microsora subsp. pentangularis: sori and indusia.
pentangularis: habit.

Lastreopsis acuminata (Houlston) C. V. Morton

puberulent between the ridges. The ridges are continuous
 13.59.4
with the decurrent margins of the pinnules (also thick-
syn. Lastreopsis shepherdii (Kunze ex Mettenius) Tindale
ened). Unlike in dryopteroid ferns, the rachis and costae
Shiny shield fern
are rounded on their upper sides, not deeply grooved.
Semi-hardy
Lastreopsis is pantropical and contains 35 species. The
genus name means “resembling Lastrea,” formed by adding A medium-sized fern with clump-forming and ascending
the Greek suffix -opsis, like. For a taxonomic monograph of rhizomes. Grows well under low to medium light in moist
the genus, see Tindale (1965). or moist-wet garden soil or potting mix. The plants are easy
to grow from spores and establish readily.
Lastreopsis acuminata has tufted fronds often with pink-
tinged stipes. The blades are narrowly triangular and two-
to three-pinnate, coarser than those of most Lastreopsis. The
lowest pinna pair are larger on the basiscopic side. This spe-
cies is native to Australia and Tasmania.

Lastreopsis effusa (Swartz) Tindale  13.59.5

Semi-tender
A medium-large fern with clump-forming rhizomes.
Grows well under low to medium light in moist garden soil
or potting mix.
Lastreopsis effusa is characterized by highly dissected,
pentagonal or broadly lanceolate blades that bear a scaly
bud toward the tip of the rachis. Indusia are absent. It is
native to tropical America.

Figure 13.59.2. Lastreopsis microsora subsp.

pentangularis: grooves of rachis and costae.
344 C 13

Lastreopsis microsora (Endlicher) Tindale

 13.59.1–3, 6, 7
Semi-hardy to semi-tender
A small to medium fern with moderately long-creeping rhi-
zomes. Grows well under medium light in moist garden
soil or potting mix. Lastreopsis microsora is easy to grow.
Two subspecies are cultivated.
subsp. microsora. Figure 13.59.6. The rhizomes are
short- to medium-creeping and branched. The blades are
broadly pentagonal and bear a dense tuft of short pale hairs
at the base of the costules and costa. The tips of ultimate
segments are blunt, not acute, and the indusia are large and
reniform. This subspecies is native to Australia. In the
United States, plants of subsp. microsora have been con-
fused with Lastreopsis decomposita (R. Brown) Tindale,
which differs by having short-creeping rhizomes and in-
flated (bullate) scales at the base of the rachis. It is not
known to be cultivated in the United States.
subsp. pentangularis (Colenso) Tindale (Ctenitis pen-
tangularis (Colenso) Alston). Figures 13.59.1–3, 7. This
subspecies resembles the previous one but is smaller,
slightly less foliaceous, and more gray-green. It is native to
New Zealand.
Figure 13.59.4.
Lastreopsis acuminata:
fronds, bar = 5 cm
(2 in.).

Figure
13.59.5.
Lastreopsis
effusa: frond
and bud, bar =
5 cm (2 in.).

Figure 13.59.6. Lastreopsis microsora: frond, bar = 5 cm

(2 in.).
 F  F A  C 345

Solanopteris, the only other genus of ferns to have ant-

inhabited rhizomes, differs primarily by bearing globose
tubers sporadically along a long-creeping rhizome; the en-
tire rhizome is not enlarged and hollow, as in Lecanopteris.
Solanopteris is entirely restricted to the American tropics,
whereas Lecanopteris is native to southeastern Asia, mostly
Malaysia.
Lecanopteris contains 13 species. The genus name comes
from the Greek lekane, dish, and lepis, scale, referring to
the dish-like flaps of the blade associated with the sori.

Lecanopteris carnosa (Reinwardt) Blume

 13.60.1
Tender
A small-medium fern with medium-creeping rhizomes.
Grows under high light in well-drained, moist-dry uncut
moss. Culture is easier in constantly warm, humid, well-
lighted greenhouses with good ventilation.
Lecanopteris carnosa is characterized by rhizomes lack-
ing scales and pinnatifid, erect, leathery fronds. The rhi-
zomes are sometimes described as a “mat of lumps.” The
sori are borne in cup-like depressions with each cup borne
Figure 13.59.7.
on a small projection from the frond margin. The species is
Lastreopsis microsora
native from Sumatra to the Philippines.
subsp. pentangularis:
frond, bar = 5 cm
(2 in.). c

Lecanopteris b
syn. Myrmecophila, Myrmecopteris
Ant fern
The genus Lecanopteris has enlarged and hollow rhizomes
that provide homes for ants. The ants, in turn, protect the
plant by attacking anything that disturbs it. These small- to
medium-sized epiphytes are grown as novelties where tropi-
cal conditions can be provided.
The rhizomes of Lecanopteris are bluish green when
young and become blackish with age. The rhizome scales a
are absent or dense, round, peltate, and marginally mem-
branous. The stipes are jointed to a small raised projection
(phyllopodium), and the sterile blades vary from entire to
d
lobed to deeply pinnatifid. The veins are netted, not free.
The sori are round or oblong, usually sunken, and like all
polypodiaceous ferns, lack indusia. Figure 13.60.1. Lecanopteris carnosa: a. habit, bar = 5 cm
The species with scaly rhizomes, such as Lecanopteris (2 in.); b. fertile portion of frond, upper surface, bar =
sinuosa, have been placed in Phymatosorus (Phymatodes), 1 cm (0.4 in.); c. sori on projection bent to upper frond
but some pteridologists put them in Myrmecopteris (Myrme- surface, bar = 2 mm; d. swollen rhizome, bar = 1 cm
cophila). More study is needed to resolve these differences. (0.4 in.).
For details, see Johns (1995). See Polypodium for a com-
parison of Lecanopteris to the other polypodiaceous genera.
 346 C 13

Lecanopteris crustacea Copeland  13.60.2

syn. Myrmecopteris crustacea (Copeland) Pichi-Sermolli,
Phymatodes crustaca (Copeland) Holttum
Tender
A small-medium fern with 3–5 cm (1.2–2 in.) wide, me-
dium-creeping rhizomes. Grows well under high light in
well-drained, moist-dry uncut moss. Culture is easier in a b
constantly warm, humid, well-lighted greenhouses with
good ventilation.
Lecanopteris crustacea is characterized by round, peltate
rhizome scales, deeply pinnatifid fronds, and sunken sori. It
is native to Sumatra, Malaysia, and Borneo.

a b

Figure 13.60.3. Lecanopteris sinuosa: a. habit, bar = 5 cm

(2 in.); b. fertile portion of frond, bar = 1 cm (0.4 in.);
c. rhizome, bar = 1 cm (0.4 in.).

to linear-lanceolate, entire to coarsely crenate fronds. The

c
sori are round or oblong, large, and submarginal. It is native
from Thailand to the Solomon Islands, and to Taiwan.

Lemmaphyllum  13.61.1, 2

d The cultivated species of Lemmaphyllum are used as novel-
ties in terrariums, pots, and small hanging containers.
Figure 13.60.2. Lecanopteris crustacea: a. habit, bar = These small, creeping epiphytes have thick, oval, shiny, suc-
5 cm (2 in.); b. narrowed fertile segment, bar = 1 cm culent leaves. The production of fertile fronds can be in-
(0.4 in.); c. swollen rhizome, bar = 5 cm (2 in.); creased by growing the plants under higher light.
d. rhizome scale, bar = 1 mm, after Hoshizaki (1982). This genus has slender, creeping rhizomes covered by
peltate, ovate-lanceolate, clathrate scales. The stipes are
jointed to the rhizome, and the simple, entire blades vary
from papery to succulent and rounded to elliptic, obovate,
Lecanopteris sinuosa (Wallich ex Hooker) Copeland or lanceolate. Although often hard to see, the veins are net-
 13.60.3 ted, not free. The fertile fronds are narrower than the ster-
syn. Myrmecopteris sinuosa (Wallich ex Hooker) Pichi- ile ones. The sori are rounded or linear or running together,
Sermolli, Phymatodes sinuosa (Wallich ex Hooker) J. forming a row on each side of the midrib. As in all polypo-
Smith diaceous ferns, the sori lack indusia, but in this genus they
Tender are protected by umbrella-shaped hairs mixed among the
A small-medium fern with medium-creeping rhizomes 1– sporangia. Lemmaphyllum resembles Microgramma, though
1.5 cm (0.5–0.6 in.) wide. Grows well under medium to the commonly cultivated species differ by their thick, fleshy
high light in well-drained, moist-dry uncut moss. The cul- fronds and linear sori.
tural requirements are similar to those of Lecanopteris car- Lemmaphyllum is native to eastern Asia and contains six
nosa, but this species is easier to grow. species. The genus name comes from the Greek lemma,
Lecanopteris sinuosa is characterized by round, peltate scale, and phyllon, leaf, referring to the small, scale-like
rhizome scales with whitish margins, and simple, oblong structures mixed among the sporangia.
 F  F A  C 347

Lemmaphyllum microphyllum C. Presl

 13.61.1–3 a b
Semi-hardy
A small fern with creeping rhizomes. Grows well under
medium light in moist, drained potting mix. The plants
are easy to grow once established but tend to grow slowly.
Lemmaphyllum microphyllum has dimorphic sterile and
fertile fronds. The sterile fronds vary from circular to ob-
lanceolate and are 1–4 cm (0.5–1.5 in.) long. In contrast,
the fertile fronds are narrowly linear and up to 6 cm (2.5
in.) long. The sori are linear and in one row on either side
of midrib. The species is native to Japan, South Korea,
China, and Taiwan.

Figure 13.61.3. Lemmaphyllum microphyllum: a. habit,

bar = 5 cm (2 in.); b. fertile frond, bar = 5 cm (2 in.).

Lemmaphyllum subrostratum (C. Christensen) Ching

 13.61.4
Very tender
A small fern with medium-creeping rhizomes. Grows well
under medium light in moist, drained potting mix. The
plants grow slowly, possibly because of their heat require-
ment, and are rare in cultivation.
Lemmaphyllum subrostratum has only slightly dimorphic
sterile and fertile fronds. The sterile fronds have green and

a
Figure 13.61.1. Lemmaphyllum microphyllum: habit.

Figure 13.61.2. Lemmaphyllum microphyllum: sori on Figure 13.61.4. Lemmaphyllum subrostatum: a. habit,
narrowed fertile fronds. bar = 5 cm (2 in.); b. fertile frond, bar = 5 cm (2 in.);
c. rhizome scale, bar = 1 mm.
 348 C 13

well-defined stipes 1–3 cm (0.5–1.2 in.) long and blades

that vary from elliptic to lanceolate, 5–10 cm (2–4 in.) long
by 1–2 cm (0.5–0.8 in.) wide. In contrast, the fertile fronds c
have longer stipes and narrower blades. The sori are round.
The rhizome scales are broadly rhomboid at base. The spe-
cies is native to India and southeastern Asia.
Lemmaphyllum subrostratum was introduced commer-
cially from India, and the plants were originally misidenti-
fied as L. carnosum (J. Smith ex Hooker) C. Presl, a species
that differs by having ovate-lanceolate rhizome scales, ses-
sile (or nearly so) sterile fronds, narrowly linear fertile
fronds, and acrostichoid sori.

Lepisorus
syn. Polypodium in part, Pleopeltis in part
The species of Lepisorus grow in soil, on rocks, or on tree
trunks and branches. These are small- to medium-sized, a b
creeping polypodiaceous ferns. The widespread Lepisorus
thunbergianus is often seen on trunks and branches in the
Hawaiian Islands, and many cultivars of this species are
commonly grown in pots or baskets in Japan. The culti-
Figure 13.62.1. Lepisorus longifolius: a. habit, bar = 5 cm
vated species can be difficult to establish, but with patience
(2 in.); b. fertile portion of frond, bar = 1 cm (0.4 in.);
the second or third attempt may succeed. Other species of
c. peltate scale of sori, bar = 0.1 mm.
Lepisorus are extremely difficult to grow.
The rhizomes vary from short- to long-creeping and
contain scattered, blackish fiber strands (seen in cross sec-
tion). Peltate, clathrate, entire or toothed scales cover the peltate scales. This species is native from India to Malaysia
rhizomes, sometimes with one or two glandular projections and the Philippines.
on the margins. The stipes are short, and the blades are sim-
ple, entire, and lanceolate to linear, sometimes bearing Lepisorus thunbergianus (Kaulfuss) Ching
clathrate scales. The fertile fronds are nearly the same size  13.62.2
and shape as the sterile ones. The veins are netted with syn. Polypodium lineare Thunberg
many irregular areoles, within which are free veins. The sori Hardy to very hardy
vary from round to elongate and are protected when young
A small fern with medium-creeping rhizomes. Grows
by partly or fully peltate, clathrate, entire scales. Like all
under medium to high light in well-drained, moist-dry
polypodiaceous ferns, Lepisorus lacks indusia. The species of
potting mix or uncut moss. This species is difficult to estab-
Lepisorus were first classified in Polypodium and later in
lish but is a popular pot plant in Japan, with many named
Pleopeltis; see Polypodium for a comparison of this genus to
cultivars.
other polypodiaceous genera.
The rhizome scales of Lepisorus thunbergianus are dark
Lepisorus is native mainly to the Old World tropics and
brown, ovate-cordate at base, and long-attenuate. The
subtropics. The number of species is unknown, but is prob-
fronds are simple, leathery, tapered at both ends, linear-
ably less than 40. The genus name comes from the Greek
elliptic, 10–30 cm (4–12 in.) long, and 5–15 mm (0.2–0.6
lepis, scale, and soros, mound, referring to the scaly sorus.
in.) wide. Because of the leathery blades, the veins are hard
to see. The sori are round or oval and disposed in one row
Lepisorus longifolius (Blume) Holttum on either side of midrib. The species is native to Japan,
 13.62.1 South Korea, China, Taiwan, Indochina, the Philippines,
syn. Paragramma longifolia (Blume) T. Moore and the Hawaiian Islands.
Probably tender The species is extremely variable, not only in blade size
A medium-sized fern with short-creeping rhizomes. Grows and shape, but also in habitat. The plants grow on rocks,
well under medium light in moist, drained uncut moss or steep banks, tree trunks, stone walls, and the roofs of
potting mix. This species is reportedly the easiest Lepisorus houses.
to grow in Florida.
Lepisorus longifolius has linear-lanceolate fronds. Oval
to oblong, marginal sori are covered and surrounded by
 F  F A  C 349

Leptochilus amplus (F. V. Müller ex Bentham)

Nooteboom  13.63.1
syn. Colysis ampla (F. V. Müller ex Bentham) Copeland,
C. sayeri (F. V. Müller ex Bentham) Copeland
Tender
a A medium-sized fern with long-creeping rhizomes. Grows
well under low to medium light in moist, drained potting
b mix or uncut moss.
The stipe of Leptochilus amplus is winged to the base,
and on mature plants the blade is deeply pinnatifid with
about three pairs of oblong lobes, the terminal lobe resem-
bling the lateral ones. The apices of the lobes are acumi-
nate or long-attenuate, and the veins are conspicuous. The
sori are slender, linear, and run obliquely between the mid-
rib and margin. The species is native to Australia.
Leptochilus amplus circulates in horticulture as Colysis
sayeri, a synonym. Australian botanists have traditionally
recognized two species: (1) L. sayeri, with the stipe wing
not reaching the base and with sori shorter and located
midway between the midrib and margin of the lobes and,
(2) L. amplus, with the stipe wing reaching the base and
c with longer sori, usually extending from near the lobe mid-
rib to near the margin. According to Nooteboom (1997),
however, these characteristics intergrade and the two spe-
Figure 13.62.2. Lepisorus thunbergianus: a. habit, bar = cies should be considered the same.
5 cm (2 in.); b. peltate scale of sori, bar = 0.1 mm;
c. rhizome scale, bar = 1 mm.

b
Leptochilus Kaulfman
syn. Colysis C. Presl
These small- to medium-sized ferns have variously shaped
leaves and are attractive in pots, baskets, or (in warmer cli-
mates) the ground. Leptochilus, better known to gardeners
as Colysis, resembles Microsorum but differs by having elon-
gate sori located in an even row between two veins. They
are epiphytic or epilithic.
The genus has creeping rhizomes covered by peltate or
false-peltate, clathrate or subclathrate scales. The fronds are
simple or pinnatifid, with the fertile fronds contracted or
not. The veins are visible, netted, and form areoles con- a
taining branched included veinlets pointing in various di-
rections. The sori are mostly elongate or linear (rarely acros-
tichoid) and usually appear in one line or a fairly even row
that is parallel to the secondary veins. The indusium is
absent.
Leptochilus consists of about 10 species and is native to
Asia, Malaysia, Australia, and the Solomon Islands. The
genus name comes from the Greek lepto, meaning slender,
and -chilus, meaning lipped, alluding to the blade tips of
some species, which end in a slender, curled tip. For a tech-
nical comparison of Leptochilus to related genera, see
Nooteboom (1997); also see Polypodium for a comparison Figure 13.63.1. Leptochilus amplus: a. habit, bar = 5 cm.
of this genus to other polypodiaceous genera. (2 in.); b. sori and vein pattern, bar = 1 cm (0.4 in.).
 350 C 13

Leptochilus ellipticus (Thunberg) Nooteboom Leptochilus macrophyllus var. wrightii (Hooker)

 13.63.2 Nooteboom  13.63.3
syn. Colysis ellipitica (Thunberg) Ching, C. pothifolia syn. Colysis wrightii (Hooker) Ching
(Buchanan-Hamilton) C. Presl Semi-tender to slightly hardier
Semi-tender A medium-sized fern with medium- to long-creeping rhi-
A medium-sized fern with medium long-creeping rhi- zomes and evergreen fronds. Grows well under medium
zomes. Grows under medium to high-medium light in gar- light in moist to moist-dry garden soil or potting mix.
den soil or potting mix kept moist to moist-dry. This spe- Widely known among gardeners as Colysis wrightii, Lep-
cies is subject to slug damage. tochilus macrophyllus var. wrightii is characterized by simple,
The fronds of mature Leptochilus ellipticus plants are narrow, oblanceolate blades that are long-tapered toward
broad and deeply pinnately lobed to a very narrow wing the base. It is native to Japan, southern China, Indochina,
along the rachis or fully pinnate at the base. The lobes of and Taiwan. In the wild it often grows on rocks.
sterile fronds are lanceolate and the veins are only slightly ‘Monstrifera’. Blade tips widely fluted, tip and margins
visible. Compared to the sterile fronds, the fertile ones are undulate.
taller and have narrower lobes. The linear sori extend
obliquely between the midrib and margin. The species
is native to Japan, South Korea, China, Indochina, and
Taiwan. b
Leptochilus ellipticus was broadly construed by Noote-
boom (1997) and consists of diverse-appearing plants that
are considered species by other botanists.

Figure 13.63.3. Leptochilus macrophyllus var. wrightii: a.

habit, bar = 5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.).

Leptopteris
The genus Leptopteris resembles large filmy ferns with strik-
ing delicate, plume-like fronds. The filmy texture and
translucent appearance are due to the blades being only a
few cell layers thick. These ferns are native to moist, humid
habitats, where they can form woody trunks to 1 m (3 ft.)
a tall. Leptopteris should be grown in protected sites with
b above-freezing temperatures and ample humidity. If these
conditions are unavailable, the plants should be grown in-
doors in large, shaded terrariums kept cool and humid year-
round. They may be planted in high-quality uncut sphag-
num moss.
Figure 13.63.2. Leptochilus ellipticus: a. habit, bar = 5 cm Although rarely cultivated in the United States, Lep-
(2 in.); b. sori and vein pattern, bar = 1 cm (0.4 in.). topteris is so striking that it merits further attempts. The
 F  F A  C 351

genus is now on the endangered list in New Zealand, which is easier to grow than Leptopteris superba and tolerates
prohibits the export of mature plants. They can be grown slightly drier conditions.
from spores, but the spores are green and presumably have Leptopteris hymenophylloides has ovate to triangular blades
short viability. with flat segments (that is, in the same plane of the blade).
The rhizomes of Leptopteris are erect and trunk-forming. The species is endemic to New Zealand.
Medium- to large-sized fronds are arranged in a whorl and
have finely hairy rachises and costae. The blades may be up Leptopteris superba (Colenso) C. Presl  13.64.2
to three-pinnate, are very thin, and have ultimate divisions Crape fern, Prince-of-Wales plume
that are often bilobed. The sporangia are scattered on the Semi-tender to semi-hardy
lower surfaces of the blades. Indusia are lacking. A medium-sized fern with erect rhizomes and fronds that
Leptopteris, which contains six species, is native to Aus- collectively form a basket. Grows under low light and in
tralia, New Zealand, and New Guinea to Samoa and New moist-wet, humusy potting mix or uncut sphagnum moss.
Caledonia. The genus name comes from the Greek leptos, This species grows slowly and is difficult to cultivate, re-
thin, and pteris, fern, referring to the extremely thin fronds. quiring cool temperatures and constant high humidity. In
nature it grows near waterfalls.
Leptopteris hymenophylloides (A. Richard) C. Presl In contrast to the preceding species, Leptopteris superba
 13.64.1 has blades that taper gradually toward the base, ending in
Semi-tender to semi-hardy small pinnae. Another difference is that the segments turn
A medium-sized fern with erect rhizomes and fronds that up, out of the plane of the blade. The species is endemic to
collectively form a basket. Prefers low light and moist-wet, New Zealand.
humusy potting mix or uncut sphagnum moss. The plants
require high humidity and cool temperatures. This species Leucostegia
Although commonly used as a pot plant in Malaysia, Leu-
costegia is rarely cultivated in the United States. The genus
is related to Davallia, which it resembles by its much-di-
vided, broad fronds.
The plants are usually terrestrial and medium in size.
Their rhizomes are thick, fleshy, more or less long-creeping,

Figure 13.64.1. Leptopteris

hymenophylloides: frond, bar = Figure 13.64.2.
5 cm (2 in.). Leptopteris superba:
frond, bar = 5 cm
(2 in.).
 352 C 13

and root-bearing on all sides. They are covered by non- This fern resembles Leucostegia immersa C. Presl, which
peltate scales, and hairs are often present as well. Grooves is not cultivated in the United States but is reported to be
appear on the upper surfaces of the stipes. The blades are cultivated in Australia. That species differs by indusia at-
nearly three-pinnate to four-pinnate-pinnatifid, glabrous tached only by a narrow base.
when mature, anadromous, and herbaceous or papery
(chartaceous). The ultimate segments are rhombic-lanceo- Lindsaea  13.66.1
late. The sori are often impressed and borne at the vein tips. These attractive ferns are often seen in the tropics and
They are covered by delicate, kidney-shaped or rounded sometimes in temperate areas, but they are difficult to grow
indusia—if the indusium is reniform, then it is attached at and rarely cultivated in the United States. They are prima-
a point at the base; if rounded, then attached halfway up rily seen in botanical gardens and special fern collections. In
the sides. England, four species of Lindsaea are currently cultivated.
Leucostegia, which consists of two species, occurs in Some species establish themselves if provided with a well-
southern China, the Himalayas, and southern India to drained planting mix and anchored in place on uncut moss
Polynesia. The genus name comes from the Greek leucos, or a turf-like growing medium. The plants should be kept
white, and stege, cover, referring to the whitish indusia. evenly moist in terrariums. The genus is mentioned here
because of its horticultural potential and importance in the
Leucostegia pallida (Mettenius) Copeland tropics.
 13.65.1 Some of the species resemble maidenhair ferns (Adian-
Tender tum), but there are several differences. Lindsaea almost al-
A medium-sized fern with creeping rhizomes and stipes 2– ways has light-colored (tan, green, or yellowish) stipes,
3 cm (0.75–1.2 in.) apart. Grows well under medium to rachises, and costae, and these are grooved on their upper
high light in moist garden soil or potting mix. The plants surfaces, often with a sharp wing on either side of the
can be grown in baskets or pots. The fronds tend to spread groove. Furthermore, the indusium is formed on the lower
and should be given plenty of room.
Leucostegia pallida has triangular blades, up to four-pin-
nate, with obovate or rhombic ultimate segments. The sori
appear as raised bumps on the upper surface of the blade. b
Roundish indusia are attached at the base and halfway up
the sides. The species is native to the New Hebrides, Bor-
neo, and Malaya.

d
Figure 13.65.1.
a Leucostegia pallida:
a. habit, bar = 5 cm Figure 13.66.1. Lindsaea: a. frond (Lindsaea ensifolia);
(2 in.); b. segment b. habit (L. repanda); c. frond (L. stricta), bar for a,
with sori and indusia, b, and c = 5 cm (2 in.); d. sorus and indusium, bar =
bar = 2 mm. 2 mm.
 F  F A  C 353

surface of the blade and opens toward the margin. In con-

trast, Adiantum has brown or black rachises and costae that
are ungrooved, and the indusium is formed by the enrolled
margins of the leaf and therefore opens toward the center of
the segments.
Lindsaea contains about 150 species and occurs world-
wide. The genus is named for John Lindsay, a surgeon in
Jamaica at the turn of the 19th century who studied the
germination of fern spores.

Llavea
Llavea consists of one species, endemic to Mexico and Gua-
temala, that is rarely cultivated. Llavea cordifolia has an at-
tractive airy appearance and bears many gray-green, oval
segments. The plants are easy to grow from spores. They are Figure 13.67.1. Llavea cordifolia: habit.
sensitive to overwatering and susceptible to leaf-spot dis-
eases when grown in extremely humid environments.
The genus has several distinctive characteristics, the the other, Lonchitis occidentalis Baker, to Africa and Mada-
most obvious being the strong differentiation in the shape gascar. The genus name comes from the Greek lonche,
and position of its sterile and fertile segments. The sterile lance, and -itis, a suffix indicating a close connection. It
segments are oval or nearly so and borne throughout the comes from an ancient name that was used by the Greek
basal two-thirds of the blade, whereas the fertile ones are physician and botanist Dioscorides.
linear and borne only in the apical third. The sori are pro-
tected by the enrolled margin of the segments. The rhizome
scales are mostly black but grade into bright yellow ones at
the bases of the leaf stalks. The genus is named for Pablo de
la Llave (1773–1833), a traveler in Mexico. b

Llavea cordifolia Lagasca  13.67.1, 2

Tender to semi-tender
A medium-sized fern with short-creeping to suberect rhi-
zomes. Grows under medium to high light in moist, drained
potting mix. Llavea cordifolia is best used as a pot plant. Be
careful not to overwater, and do not allow water to remain
on the leaves. Provide adequate air circulation. See the
genus entry for a botanical description.

Lonchitis
syn. Anisosorus
These terrestrial ferns are rarely cultivated. Lonchitis has c
rather coarsely cut fronds with a soft, hairy appearance. The
rhizome is creeping, thick, and fleshy and bears hairs and
scales. The stipe contains two vascular bundles. The fronds
are slightly succulent, two- to three-pinnate-pinnatifid, a
broad, and hairy. The veins are free or occasionally netted.
The sori are marginal, being borne at the sides of the seg-
ments or lobes. The indusium is formed by the enrolled
leaf margin.
This genus is sometimes confused with Blotiella (not
cultivated), which has hard, erect rhizomes and fully netted
veins. Lonchitis is distinguished from the similar Pteris by
hairy (not scaly) rhizomes. Figure 13.67.2. Llavea cordifolia: a. frond, bar = 5 cm
The genus consists of two species that hardly differ. (2 in.); b. distal portion of a fertile frond, bar = 5 cm
One, Lonchitis hirsuta, is native to tropical America, and (2 in.); c. stipe base, bar = 5 cm (2 in.).
 354 C 13

Lonchitis hirsuta Linnaeus  13.68.1–3

Tender
A medium to large fern with short-creeping, succulent rhi-
zomes and softly hairy fronds. Grows well under medium
light in moist-wet potting mix.
All parts of Lonchitis hirsuta are hairy. The blade is soft
and broadly cut, reminiscent of tomato foliage in shape.
The veins are free. This species is native to the American
tropics.

a b

Figure 13.68.3. Lonchitis hirsuta: a. frond, bar = 5 cm

(2 in.); b. hairs, sori, and indusia, bar = 5 mm (0.2 in.).

Lycopodium  13.69.1, 2

Club moss, ground pine, princess pine, tassel fern
In the north-temperate zone, Lycopodium is often seen on
forest floors as small- to medium-sized, erect or trailing
plants with scale-like leaves. Many tropical species are epi-
phytes; less often are they scandent or trailing on the
Figure 13.68.1. Lonchitis hirsuta: habit. ground. The epiphytic species with pendent stems are
prized in horticulture. They can be handsomely displayed
in hanging containers or on boards. The temperate species
are difficult to grow and are seldom cultivated. They resent
disturbance and often require the presence of special fungi
(mycorrhizae) in the roots.
Lycopodiums have had several uses over the years. The
evergreen species were used in wreaths and table decora-
tions during the winter and Christmas season. The plants
were so intensively gathered for this use that laws were en-
acted to protect them in forest preserves and parks. Another
use involved the spores, which are full of volatile oils and
highly flammable. These spores were a primary ingredient
in fireworks and in flash powders used in photography. In
the days when pills were hand-made by pharmacists, Lyco-
podium spores were used to coat the pills and prevent them
from sticking. More recently, they have been used for the
same purpose on rubber surgical gloves.
To cultivate the North American species that grow on
forest floors, try transplanting them to woodland soils or
rock gardens with loose, moist soil. Those species that grow
in wetter areas need moist soil, but not so moist that the
plants rot. A sandy soil mixed with peat moss and loam,
carefully watered, might encourage rooting and new
Figure 13.68.2. Lonchitis hirsuta: pinnae. growth in transplants. Transplanting has been successful
 F  F A  C 355

Figure 13.69.2. Lycopodium squarrosum: fertile

leaves.

Figure 13.69.1. Lycopodium phlegmaria: habit.

Lycopodium is easy to recognize by its small, usually
scale-like leaves supplied by a single vein. Most plants are
in oak-leaf mold, using aluminum sulfate to maintain acid- evergreen, but a few species are deciduous. Many species
ity and preventing the plants from drying. A well-drained, have cones, but some bear sporangia in the axils of leaves
slightly acidic soil or coarsely cut sphagnum moss (not too that resemble vegetative leaves, thus cones are absent. The
wet) has worked with Lycopodium lucidulum. sporangium can be seen as a bean-shaped or rounder struc-
The epiphytic species are easy to grow, but most require ture, usually yellowish, on the upper surface of the scale (or
warm temperatures, humidity, and adequate air circula- leaf) near the base. The fertile leaves usually occur at the
tion. The planting medium must be coarse and well stem apex. The fertile leaves may be highly reduced and
drained. Care should be taken to guard against extremes: compact, thus forming a cone. The cones can also be sessile
never let the plants dry out nor get too wet. Poor air circu- or borne on specialized stalks. Only one kind of spore is
lation promotes leaf diseases and rot. Plants may need to produced.
have their roots covered by uncut sphagnum moss, which Some botanists divide Lycopodium into 3 to 12 genera,
can be tied to a board or a tree-fern-fiber planter. Hanging and it is only a matter of opinion how many of these, if any,
pots or baskets can also be used, in which case a coarser should be recognized. Selaginella is sometimes confused
planting mix (not soil) should be used. If lycopodiums are with Lycopodium, but the cultivated species of Selaginella
planted in pots with uncut sphagnum, the moss should be tend to have smaller leaves, softer foliage, and two kinds of
kept loose because packed moss holds too much water. sporangia: larger female ones at the base of the fertile zone
Lycopodium species can be propagated by divisions, tip and smaller male ones above.
cuttings, and layering. Propagators report that the new, lat- Lycopodium occurs worldwide and contains about 450
eral shoots tend to form between the sterile and fertile re- species. The genus name comes from the Greek lykos, wolf,
gions of the stem. Therefore when propagating by layer- and pous, foot, alluding to a fancied resemblance between
ing, this area should contact the growth medium. For the branch tips and a wolf’s paw.
further details on cultivation, see the section on “Fern Al-
lies” in Chapter 10.
 356 C 13

Lycopodium carinatum Desvaux  13.69.3

syn. Huperzia carinata (Desvaux) Trevisan,
Phlegmariurus carinatus (Desvaux) Ching
Keeled tassel fern
Tender
A medium-sized plant with slender, cord-like, pendulous,
evergreen stems. Grows well under medium light in
drained, moist potting mix or uncut moss attached to a
board or in a hanging container.
Lycopodium carinatum is a pendent epiphyte, several Figure 13.69.4. Lycopodium
times branched, with a gradual transition from sterile to cernuum: habit, bar = 5 cm
fertile leaves. The branches grow to about 80 cm (32 in.) (2 in.).
long and 8–10 mm (0.3–0.4 in.) wide including the leaves.
The sterile leaves are narrowly ovate, 10–13 mm (about 0.5
in.) long, 2 mm wide, thick, leathery, stiff, appressed, at
about a 30-degree angle from the stem, closely overlapping,
strongly keeled and folded along the midrib, entire, acute,
and decurrent on the stem. The species occurs from India
to Polynesia and Australia. Christmas tree with nodding branch tips. In the southeast-
ern United States the tips of the horizontal stems overwinter.
The sterile leaves of Lycopodium cernuum are narrowly
triangular, pale green, and entire. The cones are short, less
than 1 cm (0.5 in.) long, and borne at the tips of the nod-
ding branchlets. The fertile leaves are trowel-shaped and
fringed along the margins. This is one of the most wide-
b spread club mosses in the tropical and subtropical regions
of the world. It is often weedy in the wild, especially in dis-
Figure 13.69.3.
turbed habitats.
Lycopodium
carinatum:
Lycopodium clavatum Linnaeus  13.69.5
a. branches with
Staghorn club moss, running pine
sterile and fertile
c Very hardy, Zone (2)3
leaves in spikes,
bar = 5 cm A medium-sized plant with long-creeping, horizontal stems
(2 in.); b. sterile that periodically send forth erect stems. Grows under me-
leaf, bar = 3 mm dium light in moist-wet soil or potting mix. This species is
(0.1 in.); c; fertile difficult to grow.
leaf, bar = 3 mm The hair-like tip of the sterile leaves will distinguish Ly-
(0.1 in.). copodium clavatum from all others in the genus. The plants
are evergreen with antler-like branches and stalked cones.

Lycopodium cernuum Linnaeus  13.69.4

syn. Lycopodiella cernuua (Linnaeus) Pichi-Sermolli, Figure 13.69.5. Lycopodium
Palhinhaea cernua (Linnaeus) Vasconcellus & Franco clavatum: habit, bar = 5 cm
Nodding club moss (2 in.).
Tender to semi-tender, Zones 8–9
A medium to large plant with long, horizontal stems that
branch and scramble. Grows under medium to high light
in moist potting mix. The species resembles a miniature
 F  F A  C 357

The species is one of the most widely distributed pterido-

phytes in the world, occurring on every continent. Japa-
nese scientists report that spores can cause allergies.
a
Lycopodium complanatum Linnaeus  13.69.6
syn. Diphasiastrum complanatum (Linnaeus) Holub
Flat-branch club moss, ground cedar, running pine
Very hardy, Zone (1)2
A medium-sized plant with long-creeping horizontal stems
that send up much-branched erect stems with flattened
branchlets. Grows under medium light in moist soil or pot-
ting mix. The species is difficult to grow.
Lycopodium complanatum is an evergreen plant with flat-
tened lateral branchlets that have constrictions marking an- b
nual growth. The sterile leaves of the branchlets are four-
ranked (two laterals, one on top, and one on the bottom).
The cones are elevated on slender, candelabra-like stalks.
The species is native to North America, Europe, and Asia.
This species resembles Lycopodium digitatum A. Braun,
which also grows in eastern North America. Lycopodium Figure 13.69.7. Lycopodium dendroideum: a. habit, bar =
digitatum differs by the lack of growth constrictions on the 5 cm (2 in.); b. dorsal view of branchlet (above) and
branchlets and the fan-shaped arrangement of the hori- cross section (below), bar = 3 mm (0.1 in.), after Hickey
zontal branchlets. (1977).

triangular with an abruptly narrowing apex. The species is

native to northern North America and Asia.
This species resembles Lycopodium obscurum, which see
for comparison.
Figure 13.69.6. Lycopodium
complanatum: habit, bar = Lycopodium filiforme Swartz  13.69.8
5 cm (2 in.), after Beitel syn. Huperzia filiformis (Swartz) Holub, H. polytrichoides
(1979). (Kaulfuss) Trevisan, Lycopodium filiforme Kaulfuss
Tender
A medium-sized plant with pendulous, evergreen stems.
Grows well under medium light in moist, drained potting
mix or uncut moss attached to a board or in a hanging
container.
Lycopodium dendroideum Michaux  13.69.7
syn. Lycopodium obscurum var. dendroideum (Michaux)
D. C. Eaton
Prickly-tree club moss, princess pine
Very hardy, Zone 2 a b
Figure 13.69.8.
A medium-sized plant with long-creeping stems. Grows Lycopodium filiforme:
under medium light in moist soil or potting mix. The spe- a. branches with
cies is difficult to cultivate. sterile and fertile
Lycopodium dendroideum is an evergreen species. The leaves in spikes, bar
main stems are upright and branched into a miniature tree- = 5 cm (2 in.);
like form. The lateral branches are round in cross section, b. sterile leaf above,
and the sterile leaves are lanceolate and all of nearly equal fertile leaf below, bar
size. Leaves of branchlets are arranged in two dorsal, two = 3 mm (0.1 in.).
ventral, and two lateral rows, with those on the main stems
beneath the first branch spreading at a 30- to 90-degree
angle. The cones are sessile, and the fertile leaves are ovate-
 358 C 13

Like Lycopodium carinatum, L. filiforme has cord-like

stems, but they are narrower, only 2–3 mm wide (including
the leaves), to 60 cm (24 in.) long, and several times forked b
near the base. The sterile leaves are narrowly triangular to
lanceolate, to about 3.5 mm long, about 1.5 mm wide at
the base, keeled, with translucent and minutely irregularly a
toothed margins. The transition from the sterile to fertile
zone is gradual. The species is endemic to Australia.

Lycopodium lucidulum Michaux  13.69.9

syn. Huperzia lucidula (Michaux) Trevisan, Urostachys
lucidulus (Michaux) Nessel
Shining club moss
Very hardy, Zone 3
A small plant with erect stems that recline with age and take
root. Grows under medium light in moist, acidic potting
soil or coarsely cut moss. This species is difficult to grow. Figure 13.69.10. Lycopodium
The plants are evergreen with shiny leaves and usually grow nummulariifolium: a. branches,
in open tufts. The upper tips of the branchlets form buds bar = 5 cm (2 in.); b leaves, bar =
that are shed in the fall. 3 mm (0.1 in.).
Lycopodium lucidulum has erect, dichotomously branched
stems with constrictions marking annual growth. The ster-
ile leaves are dark green, shiny, and oblanceolate, with the
margins entire or papillate or with up to eight irregular Relatively large, circular leaves flattened against the stem
teeth. Most of the leaves point downward and impart a readily distinguish Lycopodium nummulariifolium from
shaggy appearance to the plant. The fertile leaves are simi- other species in the genus. An epiphyte, it is native to Thai-
lar to the sterile ones. The species resembles L. selago. It is land, Malaysia, Polynesia, and New Guinea.
native to eastern North America, where it grows on forest
floors and rarely on rocks. Lycopodium obscurum Linnaeus  13.69.11
Tree club moss, flat-branched tree club moss, ground
pine, princess pine
Very hardy, Zone 3
A medium-sized plant with long-creeping stems. Grows
under medium light in moist soil or potting mix. This spe-
cies is difficult to grow.
Lycopodium obscurum resembles L. dendroideum and is
Figure 13.69.9. Lycopodium sometimes difficult to distinguish. The easiest way to tell
lucidulum: habit, bar = 5 cm them apart is by looking at the leaves on the main, erect
(2 in.). stem below the first branch: these leaves are appressed or
slightly spreading-ascending in L. obscurum but widely
spreading in L. dendroideum. Another difference (although
you need to see the two species side by side to appreciate it)
is that lateral branches are slightly flattened in L. obscurum
Lycopodium nummulariifolium Blume but cylindrical in L. dendroideum. Lycopodium obscurum is
 13.69.10 native to the eastern United States and adjacent Canada.
syn. Huperzia nummulariifolium (Blume) Chambers,
Jermy & Crabbe, Phlegmariurus nummulariifolius Lycopodium phlegmaria Linnaeus
(Blume) Ching  13.69.1, 12
Tender syn. Huperzia phlegmaria (Linnaeus) Rothmaler,
A medium-sized plant with long, pendent, stiff branches Phlegmariurus phlegmaria (Linnaeus) Sen & Sen
arising from a clump of rhizomes. Best grown in a hanging Common tassel fern
container with good drainage. The planting media should Tender
be kept moist, but the surroundings should have good air A medium-sized plant with short-creeping rhizomes and
circulation and medium light. pendulous, evergreen stems. Grows well under medium
 F  F A  C 359

light in moist, drained potting mix or uncut moss attached

to a board or in a hanging container.
The fertile parts of Lycopodium phlegmaria are slender
and tassel-like, with fertile tips often continuing to grow
and forming sterile leaves. (In other species, the fertile tips
usually stop growing.) The sterile stems fork several times
and are 1.5–3 cm (0.6–1.2 in.) wide (including the leaves).
The leaves are spirally arranged, crowded, triangular to
ovate-lanceolate, stiff, leathery, dark green, entire, prickly,
short-stalked, and acute at the apex. The fertile branches
are slender and much branched, and the fertile leaves are
a much shorter than sterile leaves, with the sporangia about
as wide or wider than the fertile leaves. The species is native
to Africa, Asia, and the Pacific Islands.
b
Lycopodium phlegmarioides Gaudichaud
 13.69.13
syn. Huperzia phlegmarioides (Gaudichaud) Rothmaler
Layered tassel fern
Very tender, Zone 11
Figure 13.69.11. Lycopodium obscurum: a. habit, bar = A medium-sized plant with pendulous stems, to 80 cm (32
5 cm (2 in.); b. dorsal view of branchlet (above) and in.) long, that are several-times branching. Grows best in
cross section (below), bar = 3 mm (0.1 in.), after Hickey medium light in coarse potting mix or uncut moss in hang-
(1977). ing pots or on boards; keep moist but very well drained.
Lycopodium phlegmarioides is an epiphyte that resembles
L. phlegmaria but has more yellowish green, stiffer, and
fewer four-ranked leaves. The sterile leaves are broadly
ovate, twisted at the base and adnate to the stem. The spo-
b

c
c

Figure 13.69.12. Lycopodium phlegmaria: a. branches Figure 13.69.13. Lycopodium phlegmarioides: a. branches
with sterile and fertile leaves in spikes, bar = 5 cm with sterile and fertile leaves in spikes, bar = 5 cm
(2 in.); b. sterile leaf bar = 3 mm (0.1 in.); c. fertile leaf, (2 in.); b. sterile leaf, bar = 3 mm (0.1 in.); c. fertile leaf,
bar = 3 mm (0.1 in.). bar = 3 mm (0.1 in.).
 360 C 13

rangia are shorter than the fertile leaves. This species is na-
tive from Australia and Malaysia to New Caledonia and
the Pacific Islands.
Figure 13.69.15.
Lycopodium selago Linnaeus  13.69.14
b Lycopodium
syn. Huperzia selago (Linnaeus) C. Martius & Shrank
squarrosum:
Mountain club moss, fir club moss, alpine club moss
a. branches with
Very hardy, Zone 2
fertile and sterile
A small plant with evergreen, clump-forming stems. Grows leaves in spikes,
under medium light in moist-wet sandy soil or drained pot- bar = 5 cm
ting mix. This species is difficult to grow. (2 in.); b. sterile
a
The stems tend to be yellow-green (at least in cultiva- leaf, bar = 3 mm
tion), forked only once or twice, and produce buds near the (0.1 in.); c. fertile
tips. The sterile leaves are mostly narrowly triangular, leaf, bar = 3 mm
pressed close to the stem, and have papillate or entire mar- c
(0.1 in.).
gins. The fertile leaves resemble the sterile. Lycopodium se-
lago is similar to L. lucidulum but differs by having stomata
on both surfaces of the leaves, whereas in L. lucidulum these
are only on the lower surface. A hand lens is needed to see
the stomata, which appear as tiny white dots. Lycopodium se-
lago is native to northern North America (chiefly Canada).

Lygodium  13.70.1, 2

Climbing fern
Figure 13.69.14. Lycopodium Lygodium plants have twining, vine-like fronds and are
selago: habit, bar = 5 cm grown in pots or in the ground. Emerging fronds can be
(2 in.). trained on vertically strung wires or strings to produce a
lacy green curtain. The fronds can grow up to 7 m (23 ft.)
or more. The Japanese climbing fern (Lygodium japonicum)
and the small-leaved climbing fern (L. microphyllum) have
become naturalized in the southeastern United States. In
temperate areas the foliage of these species dies to the
Lycopodium squarrosum G. Forster ground in winter, and new fronds emerge in the spring. In
 13.69.2, 15 milder climates the fronds linger more than a year and may
syn. Huperzia squarrosa (G. Forster) Trevisan, produce new leaflets from dormant buds on the pinnae.
Phlegmariurus squarrosus (G. Forster) Löve & Löve Old fronds become untidy as their leaflets gradually turn
Water tassel fern, rock tassel fern brown and wither in place. Eventually the entire frond dies
Very tender, Zone 11 to the base and can only be removed, with difficulty, espe-
A medium-sized plant with clump-forming, pendulous or cially if it has become entangled with the new fronds. Prop-
erect, evergreen stems that are several-times branching. agation by rhizome divisions is unpredictable. Adult plants
Grows best in medium light in coarse potting mix or uncut generally do not transplant well.
moss kept moist but very well drained. Plant in hanging The rhizomes of lygodiums are short- to medium-creep-
containers or attach to boards. ing and hairy (not scaly). The fronds are vine-like, with the
The stems of Lycopodium squarrosum are up to 75 cm rachis twining and stem-like. The pinnae are one-pinnate
(30 in.) long, about 1.5–2.5 cm (0.5–1 in.) wide (including or palmate. The fertile leaflets (as used here, the smallest
the leaves). The sterile leaves are narrowly ovate to linear- stalked or free part of the blade) are narrower and have
lanceolate, 1–2 cm (0.5–0.75 in.) long, entire, thin but margins with spike-like extensions bearing two rows of
somewhat stiff, crowded, and widely spreading except at scale-like indusia, each indusium covering one sporangium.
the stem apex. The transition from the sterile to fertile re- Depending on the species, the veins can be either free or
gions is abrupt or gradual, but any cone-like structure is netted. The sporangia differ from those of other ferns be-
absent. The fertile leaves resemble the sterile ones, and their cause the annulus is located at the apex of the capsule, not
sporangia are inconspicuous, occupying less than one-fifth encircling it.
the length of the fertile leaf. The species is native to Africa, Lygodium consists of 45 species and, except for one tem-
Australia, Asia, and Polynesia. perate species (Lygodium palmatum), is native to the tropics
 F  F A  C 361

Figure 13.70.2. Lygodium japonicum: indusia.

Figure 13.70.1. Lygodium japonicum: habit.

and subtropics worldwide. In southern Thailand, Malaya,

Sumatra, the Philippines, and New Guinea, the twining
leaf rachises are split and flattened for weaving into baskets,
purses, and bracelets.

Lygodium circinnatum (N. L. Burman) Swartz

 13.70.3
Very tender
A large fern with short-creeping rhizomes. Grows well
under medium light in moist potting soil.
Lygodium circinnatum has sterile leaflets up to 20 cm (8 Figure 13.70.3. Lygodium circinnatum: leaflets, sterile
in.) long and about as wide. They are deeply palmately (above) and fertile (below), bar = 5 cm (2 in.).
lobed into four or five linear-lanceolate lobes to 25 cm (10
 362 C 13

in.) long. The species is native from India to southern

China, southeastern Asia, Malaysia, and Melanesia.

Lygodium flexuosum (Linnaeus) Swartz

 13.70.4
Tender
A large fern with short-creeping rhizomes. Grows well
under medium light in moist potting soil. The new growth
is often reddish.
Lygodium flexuosum resembles L. japonicum, but it has
less-divided sterile leaflets, lateral leaflets mostly eared on
one or both sides, and two- or three-lobed terminal leaflets
3–10 cm (1.2–4 in.) long. It is native to India, southern
China, Japan, and Malaysia to Australia to Melanesia. Figure 13.70.5. Lygodium
japonicum: leaflets, sterile
(above) and fertile (below),
bar = 5 cm (2 in.).

gate triangular leaflets are five- to seven-lobed, with the

terminal lobe the longest. The species is native to India,
eastern Asia, Australia, and the Philippines. It has become
naturalized in the southeastern United States. Its fronds are
deciduous in the middle Atlantic states but evergreen far-
ther south.
This species is often misidentified in the trade as Lygo-
dium scandens (Linnaeus) Swartz, a synonym of L. micro-
phyllum.
Figure 13.70.4. Lygodium flexuosum: leaflets, sterile
(above) and fertile (below), bar = 5 cm (2 in.). Lygodium microphyllum (Cavanilles) R. Brown
 13.70.6
syn. Lygodium scandens (Linnaeus) Swartz
Small-leaved climbing fern
Lygodium japonicum (Thunberg) Swartz Semi-tender, Zone 9
 13.70.1, 2, 5;  31 A large fern with short-creeping rhizomes. Grows well
Japanese climbing fern under medium-high light in moist-wet garden soil or pot-
Semi-hardy to hardier, Zone (6)7 ting mix. This species is easy to grow from spores.
A large fern with short-creeping rhizomes. Grows well un- Lygodium microphyllum is characterized by slender,
der high-medium light in moist garden soil or potting mix. creeping rhizomes and ovate to oblong-ovate sterile leaflets
This species is the most commonly cultivated of the climb- to 2.5 cm (1 in.) long. The base is heart-shaped, and the
ing ferns. It is easy to grow from spores. segments ovate and heart-shaped. It is native to Africa,
Lygodium japonicum is distinguished by its pinnately southeastern Asia, Australia, and Polynesia. It has become
lobed sterile leaflets up to 3 cm (1.2 in.) long. The elon- naturalized and weedy in Florida, especially in bald cypress
 F  F A  C 363

Figure 13.70.7.
Lygodium
palmatum: leaflets,
sterile (above) and
fertile (below), bar
= 5 cm (2 in.).

Figure 13.70.6. Lygodium

microphyllum: leaflets, sterile
(above) and fertile (below),
bar = 5 cm (2 in.).

that end before the leaf margin in a slightly swollen tip,

among other technical characteristics.
swamps, where it smothers the native vegetation and can be This genus resembles dryopteroid or ctenitoid ferns
a fire hazard by wicking fires up into the tops of trees. such as Dryopteris, Lastreopsis, and Megalastrum, which con-
tain species that have broadly triangular, two- to three-pin-
Lygodium palmatum (Bernhardi) Swartz nate-pinnatifid blades like Macrothelypteris. It differs from
 13.70.7 these others, however, by having stipes with only two vas-
Hartford fern cular bundles (not three or more), needle-like hairs on the
Very hardy, Zone 3 blades, and ungrooved upper surfaces of the rachises, cos-
A large fern with short-creeping rhizomes. Grows under tae, and costules. The pinnae of Macrothelypteris are free
low light in sandy, acidic soil. This species is difficult to from each other, not connected by a wing, thus differing
grow. Water the plants with rain water, especially in high- from the related genus Phegopteris. The needle-like hairs on
limestone areas. The plants can be propagated by spores, the lower surface of the costae are often more than 1 mm
which mature in November or December, but they loose long and colorless. The sori are round and covered by in-
their viability quickly. Mickel (1994) reports that sporel- conspicuous indusia.
ings are easy to obtain but hard to get to grow to maturity. Macrothelypteris consists of 10 species and is native to
The sterile leaflets of Lygodium palmatum are to 5 cm the tropics and subtropics of the Old World. One species,
(2 in.) long, about as wide, and deeply palmately lobed with Macrothelypteris torresiana, has become naturalized
three to seven elongate lobes. The fronds remain green dur- throughout the New World tropics and subtropics. The
ing the winter. This species is endemic to the eastern genus name comes from adding the Greek macros, large, to
United States. Thelypteris.
The evergreen fronds used to be collected during the
winter and used for Christmas decorations. This destroyed Macrothelypteris torresiana (Gaudichaud) Ching
so many populations that in 1869 the Connecticut legisla-  13.71.1
ture enacted a law to protect the plant—the first plant con- syn. Thelypteris torresiana (Gaudichaud) Alston, T.
servation law passed in the United States. uliginosa (Kunze) Ching
Torre’s fern, Mariana maiden fern
Macrothelypteris Semi-hardy, Zone (7)8
syn. Thelypteris subgenus Macrothelypteris A large fern with short-creeping to suberect rhizomes.
Only a single species of Macrothelypteris is cultivated: Grows well under medium light in moist to moist-dry gar-
Macrothelypteris torresiana. It was formerly classified in The- den soil or potting mix. The plants are easy to cultivate and
lypteris, but pteridologists now consider it sufficiently dis- volunteer readily from spore and often appear as contami-
tinct because of its large, highly divided leaves and veins nants in spore pans.
 364 C 13

Figure 13.72.1. Marattia: habit.

should be protected from frost, wind, and physical dam-

age, as the fleshy fronds break easily. Do not permit the
plants to wilt because the foliage recovers slowly.
The stems of Marattia are erect and bear fronds to 4 m
(13 ft.) long. In appearance, growth habit, and method of
propagation, Marattia resembles Angiopteris and is difficult
to distinguish from it when sterile. When fertile fronds are
present, the two genera are easy to distinguish: Marattia
has fused sporangia, whereas those of Angiopteris are free. In
Marattia, the middle of the stipe lacks the bump or swelling
that is present in Angiopteris, and the distal portion of the
a rachis is usually winged. The sorus consists of a double row
of fused sporangia (a synangium).
Figure 13.71.1. Macrothelypteris torresiana: a. frond, bar
= 5 cm (2 in.); b. winged costa and primary pinnule
bases, bar = 1 cm (0.4 in.).

Besides the characteristics listed under the genus, Ma-

crothelypteris torresiana has stipe bases with conspicuous
white lines (aerophores) along their sides, and the blades
are broadly triangular, light green, and two- to three-pin-
nate. The species is native to Asia and Africa, but is now
widely naturalized in the American tropics, subtropics, and
southeastern United States.
In the 1950s this species was commonly sold as Dry-
opteris setigera Kuntze and Thelypteris setigera (Blume)
Ching (a species not cultivated).

Marattia  13.72.1–3

Marattia is a genus of huge ferns with mostly fleshy, glo-
bose stems and large, coarse, somewhat fleshy fronds. The
succulent stems are full of starch that is sometimes eaten
by people and wild pigs, the latter of which have decimated
many populations. In the wild Marattia favors moist gullies
and grows at somewhat higher elevations and in denser Figure 13.72.2. Marattia fraxinea: fused
shade than Angiopteris, a related genus. Cultivated plants sporangia.
 F  F A  C 365

linear, with margins sharply serrate, rarely entire. The spo-

rangial capsules are oval. This species is native to Africa and
Madagascar.

Marattia salicina J. E. Smith  13.72.5

King fern, horseshoe fern
Tender to semi-tender; sensitive to frost
A large fern with erect stems. Grows well under medium
light in drained, moist garden soil or potting mix. The
plants need ample water. Do not permit the soil to dry be-
cause the fronds wilt readily and take a long time to recover.
Set the plant in a place protected from winds.
Figure 13.72.3. Marattia fraxinea: pinna. The fronds of Marattia salicina are up to 2–3.5 m (6.5–
11 ft.) long, and the segments are oblong to narrowly ovate
with shallowly toothed to almost entire margins. The spor-
angial capsules are oblong to subglobose. This species is na-
Marattia contains 60 species native to the tropics
tive to Australia, New Zealand, and Polynesia.
throughout the world. The genus is named for Giovanni
Francesco Maratti (1723–1777), a Benedictine abbot and
later professor and head of the botanical garden in Rome.

Marattia fraxinea J. E. Smith ex J. F. Gmelin

 13.72.2–4 a
Tender to semi-tender
A large fern with erect stems that tend to form trunks with
age. Grows well under medium light in moist, drained gar-
den soil or potting mix. Do not let the soil dry out, and be
sure to protect plants from wind.
Marattia fraxinea has two-pinnate fronds with ultimate
segments narrowly oblong-lanceolate to narrowly oblong b

Figure 13.72.5. Marattia salicina: a. fused sporangia, bar

= 1 mm; b. pinnules, fertile (above) and sterile (below),
b bar = 1 cm (0.4 in.); c. dentate margin, bar = 1 mm.

Marsilea  13.73.1

Water clover, pepperwort, nardoo
c
The four leaflets of Marsilea fronds resemble clover—thus
the common name water clover. These ferns are grown as
novelty plants in aquariums, ponds, and wide pots. If the
pots are submerged a few inches in water, the plants may
Figure 13.72.4. Marattia fraxinea: a. fused sporangia, produce floating leaves. Marsilea usually grows vigorously
bar = 1 mm; b. pinnules, fertile (above) and sterile and spreads by rhizomes. The plants do best in well-
(below), bar = 1 cm (0.4 in.); c. serrate margin, bar = watered, sunny locations. They are typically planted in gar-
1 mm. den soil or a mixture of half sand and peat. Mixes with high
366 C 13

rangement on the stipe base, their shape, and whether they

bear teeth. The position of the roots along the stem is also
taxonomically important. In some species the roots are pro-
duced only at the nodes, whereas in others they are pro-
duced on the nodes and internodes. For a technical treat-
ment of the American species, see Johnson (1986); for the
African species, see Launert (1968).
Marsilea occurs worldwide and contains about 60 spe-
cies. It is named for Count Luigi Ferdinando Marsigli
(1656–1730), an Italian botanist from Bologna.

Marsilea angustifolia R. Brown  13.73.2

Dwarf water clover, narrow-leaved nardoo
Hardy, Zone 6 and possibly colder areas
One of the smallest Marsilea species in cultivation. Grows
in full or filtered sun and responds well to ordinary soil or
a mix of soil, sand, and peat. The plants can grow sub-
merged and rooted in the mud with their leaves floating, or
Figure 13.73.1. Marsilea minuta: habit. they can grow on damp ground and hold their leaves erect.
With less soil water, the plants tend to produce tufts of fo-
liage at the nodes and form dense mats. When provided
with ample water and room, the rhizomes grow longer and
amounts of organic matter are apt to sour in standing the resulting mat of foliage is more open.
water. Marsilea angustifolia has 5–30 cm (2–12 in.) long fronds
Although the life cycle of Marsilea is the most complex in clusters, with narrowly cuneate leaflets (broader in culti-
of any fern, starting the plants from spores is easy and fun. vated plants) that are glabrous or scaly. The sporocarps are
The spores are of separate sexes (male and female) and con- in clusters, more or less globose, and hairy when young.
tained in brown, hard, bean-like structures called sporo- Each is borne on a short, unbranched stalk. The sporocarps
carps. These are extremely durable and remain viable for have a conspicuous blunt tooth on the upper edge and an
up to 100 years if kept dry. To germinate the spores, take a inconspicuous tooth-like bump formed from the stalk
piece of sand paper or a rough nail file and abrade the (raphe) tip. Marsilea angustifolia is reported from all states
sporocarp until the white inside is visible. Then put the of Australia except Tasmania.
sporocarp in shallow water under a bright light. Within The main distinguishing features of this species are the
minutes it will germinate and extrude a transparent, tail- short sporocarp stalk, the blunt sporocarp tooth, and the
like, gelatinous ring called the sorophore, to which the sori inconspicuous tooth-like tip of the stalk. Cultivated plants
are attached. (If the sporocarp does not germinate, try are smaller than wild plants, have more obovate leaflets, and
abrading it some more, or it might be too young, in which in live plants show a faint pale spot at the base of each leaflet.
case an older one should be used.) The sori resemble grains Also, the upper two leaflets are often smaller than the two
of white rice arranged in a row. Within a few more minutes lower leaflets. It is uncertain if this small cultivated plant,
the sori will release the small male and much larger female which appeared in a plant nursery, is a separate species.
spores that will quickly germinate and fertilize in the water.
After a week or so, the fertilized female spores, which will
appear as conspicuous white dots, can be picked up with an
eye dropper and released over wet sand or mud. Keep the
planting wet and in bright light. Young plants should
emerge and grow rapidly, maturing in 12 to 18 months. c
The water clovers are subaquatic ferns that root in mud.
New fronds are borne on long-creeping, branched rhi-
zomes. The shorter side branches of the rhizome often as-
cend and bear many crowded fronds, each of which bears Figure 13.73.2.
four leaflets and resembles a four-leaved clover. The sporo- Marsilea angustifolia:
carps are usually borne at the base of the stipe. a. habit, bar = 2.5 cm
The main characteristics used to distinguish the species (1 in.); b. sporocarp,
b
of Marsilea are the sporocarps and roots. The sporocarps a bar = 5 mm (0.2 in.);
provide useful diagnostic features in their number and ar- c. sporocarp, enlarged.
 F  F A  C 367

Marsilea drummondii A. Braun  13.73.3

Nardoo
Probably semi-hardy (?)
A small fern with medium-creeping to erect rhizomes.
Grows best under high light in aquatic conditions or in a b
moist to wet garden soil or sand-peat mix.
Marsilea drummondii is characterized by deciduous
fronds with greenish gray leaflets and (in nonaquatic forms)
conspicuous white, silky hairs. Roots are present on the
nodes and internodes. The sporocarps have two teeth, and
the sporocarp stalk usually bears one sporocarp. This spe-
cies is native to Australia, where it is called “nardoo.” It was a
eaten regularly by the Australian aborigines, and the early
Europeans explorers consumed it as a last resort to avoid
starvation. The sporocarps contain thiamase, an enzyme
that destroys Vitamin B (thiamine), and eating sporocarps
that were improperly prepared caused the death of several
early Australian explorers (Moran 1995b).

Marsilea macrocarpa C. Presl  13.73.4

Figure 13.73.4. Marsilea
Probably semi-hardy (?)
macrocarpa: a. habit, bar =
A small fern with medium-creeping to erect rhizomes. 2.5 cm (1 in.); b. sporocarp,
Grows best under high light in aquatic conditions or in a cross section on right, bar =
moist to wet garden soil or sand-peat mix. 5 mm (0.2 in.).

The distal margin of the leaflet is rounded and usually

shallowly lobed. The sporocarp is borne singly at the base of
the stipe and is oblong-rectangular, concave on both sides,
with one short tooth on the upper edge. Some roots are
present on internodes of the rhizome. Marsilea macrocarpa
is native to Africa. The cultivated plants were introduced
from Tanzania.

b Marsilea macropoda Engelmann ex A. Braun

 13.73.5
Semi-hardy
A small fern with medium-creeping to erect rhizomes. Pre-
fers high light or full sun. The plants grow well in a wet
sand-peat mix, but they can also be grown in drier soil if
used as a ground cover. In central Texas this species has
been used as a ground cover in full sun.
The stipes of Marsilea macropoda bear long, shaggy
hairs. The leaflets are hairy on both sides with long, lax,
whitish hairs (these sometimes lost on older leaves). The
Figure 13.73.3. sporocarp stalk is branched near the base and bears several
Marsilea sporocarps, which are densely covered with long, soft, fine
drummondii: reddish hairs. The lower tooth on the sporocarp is obtuse
a
a. habit, bar = and the upper one is inconspicuous or absent. The species
2.5 cm (1 in.); is native to Texas and Mexico (introduced in Alabama and
b. sporocarp, bar Louisiana), where it usually occurs on muddy clay soils or
= 5 mm (0.2 in.). in shallow water.
 368 C 13

Marsilea mutica Mettenius  13.73.7;  32

Semi-hardy
A small to medium fern with medium-creeping to erect rhi-
zomes. Grows well under high light in aquatic conditions
or in moist to wet garden soil or sand-peat mix.
Marsilea mutica can be distinguished by its variegated
leaflets that are pale green or yellowish green in the lower
(proximal) part and a dark green in the upper (distal) part,
often separated by a pale or brownish band. The sporocarp
stalk arises from the stipe base or slightly higher, and the
sporocarps are nearly globular and lack teeth. Roots are
a
present on the nodes and internodes. The species is native
to New Caledonia and Australia.
b

Figure 13.73.7.
Marsilea mutica:
a. habit, bar =
2.5 cm (1 in.);
Figure 13.73.5. Marsilea macropoda: a. habit, bar = b. sporocarp, bar =
2.5 cm (1 in.); b. sporocarps, bar = 5 mm (0.2 in.). 5 mm (0.2 in.).

b
Marsilea minuta Linnaeus  13.73.1, 6 a
Tender to semi-tender
A very small fern with medium-creeping to erect rhizomes.
Grows well under high light in aquatic conditions or moist
to wet sand-peat mix or garden soil. Marsilea quadrifolia Linnaeus  13.73.8
Marsilea minuta is one of the smallest cultivated species, Hardy, Zone 5
growing to 7 cm (3 in.) tall. Some roots are present on the A small fern with medium-creeping to erect rhizomes.
internodes. The distal margin of the leaflet tends to be cre- Grows well under high light in aquatic conditions or moist-
nate. The sporocarp stalk branches near the base, bearing wet garden soil or sand-peat mix. The plants are used in
two or three (rarely four) sporocarps that are small, 2.6–4.1 China for treating infections.
mm long, with one conspicuous tooth. Marsilea minuta is Unlike many species of Marsilea, Marsilea quadrifolia
a common, widespread, and often weedy species in Africa has leaves glabrous or nearly so. Also, the stipes of the non-
and India. It occurs in a few scattered locations in the New immersed fronds tend to recline on the soil instead of being
World tropics, but these populations probably represent stiffly erect as in many other species. The branched sporo-
escapes from cultivation. carp stalks of M. quadrifolia are found elsewhere only in
M. macropoda, a species that differs by its hairy leaves.
Roots are present on nodes and internodes. The species is
native to the northeastern United States, southeastern Eu-
a rope, and Asia.
Figure 13.73.6.
b Marsilea minuta:
Marsilea schelpeana Launert  13.73.9
a. habit, bar =
Notched water fern
2.5 cm (1 in.);
Semi-tender to (perhaps) hardier
b. sporocarp, bar =
5 mm (0.2 in.). A small fern with medium-creeping to erect rhizomes.
Grows well under high light in aquatic conditions or moist-
wet garden soil or sand-peat mix.
 F  F A  C 369

Marsilea vestita Hooker & Greville  13.73.10

Very hardy, Zone 3
A small fern with medium-creeping to erect rhizomes.
Grows well under high light in aquatic conditions or moist-
wet garden soil or sand-peat mix.
Marsilea vestita has an erect, unbranched sporocarp stalk
attached at the stipe base. The sporocarps are perpendicu-
lar or slightly nodding, thick, elliptic to nearly round, with
b a distal, acute tooth 0.4–1.2 mm long. Unlike most culti-
vated species, this one has roots that occur only at the nodes,
not along the internodes. The species is native to the
a
United States, Mexico, and Peru.
Figure 13.73.8. Marsilea
quadrifolia: a. habit, bar =
2.5 cm (1 in.); b. sporocarps,
bar = 5 mm (0.2 in.).

The leaflets of Marsilea schelpeana usually have two or

three deep notches but are reported to be sometimes only
slightly notched or, more rarely, not notched at all (entire).
The sporocarp stalks are undivided, bearing only one
sporocarp pointing upward at an oblique angle to the stalk.
This sporocarp bears one tooth and a second, obsolete one.
Roots are present on nodes and internodes. The species is b
endemic to South Africa.
This species is sometimes misnamed in the trade as Mar-
silea crenata C. Presl, a plant from eastern Asia with sporo-
carps in groups of two to seven and veins forming long are-
oles. It is unknown in cultivation in the United States.

Figure 13.73.10.
Marsilea vestita: a.
habit, bar = 2.5 cm
(1 in.); b. sporocarps,
bar = 5 mm (0.2 in.).

b
Matteuccia  13.74.1, 2
syn. Pteretis, Struthiopteris
Ostrich fern
The ostrich fern native to the United States and Canada,
a Matteuccia struthiopteris, is a medium- to large-sized plant
Figure 13.73.9. Marsilea that produces an attractive vase-shaped cluster of fresh
schelpeana: a. habit, bar = green fronds in spring. It forms dense stands in damp places
2.5 cm (1 in.); b. sporocarp, as it sends out new plants from stolons. The common name
bar = 5 mm (0.2 in.). refers to the feathery, plume-like fronds that resemble os-
trich feathers.
370 C 13

The genus Matteuccia is characterized by erect, stolon-

producing rhizomes and dimorphic fronds. The sterile
fronds are one-pinnate-pinnatifid. The veins are free, not
netted. The fertile fronds are much shorter than the sterile
ones, erect, brown, and woody. They persist over the win-
ter. The pinnae of the fertile fronds are greatly contracted
and strongly ascending. The sori are borne within the en-
rolled, leathery pinnae. Spores appear green under a micro-
scope but black when viewed against a white background.
Matteuccia contains three species and is native to North
America, Europe, and Asia. The genus name honors Carlo
Matteucci (1800–1863), a physicist at the University of
Florence in Italy.

Matteuccia orientalis (Hooker) Trevisan

 13.74.3
Figure 13.74.1. Matteuccia struthiopteris var. Oriental ostrich fern
pensylvanica: habit. Photo courtesy of R. Lloyd. Hardy
A medium-large fern with erect, stoloniferous rhizomes and
deciduous fronds. Thrives under medium to high light in
The fiddleheads of the ostrich fern are edible, and gath- moist-wet garden soil or potting mix.
ering them is a local industry in New England and Canada. The sterile fronds of Matteuccia orientalis have a stipe
The fiddleheads are boiled or steamed for 10 minutes and about half the length of the blade. Both the stipe and rachis
can be served many ways, but they are usually eaten with are scaly, and the blades are broadly ovate with the basal
just salt and pepper or garlic and butter. They can some-
times be found in specialty food stores, either pickled or
canned. The ostrich fern is the state vegetable of Vermont.

Figure 13.74.2. Matteuccia struthiopteris var. Figure 13.74.3. Matteuccia orientalis: fertile frond (left)
pensylvanica: fertile pinnae. and sterile frond (right), bar = 10 cm (4 in.).
 F  F A  C 371

pinnae slightly shorter or nearly the same as those above.

The species is native to China and Japan.
Figure 13.74.5. Matteuccia
Matteuccia struthiopteris (Linnaeus) Todaro struthiopteris var. struthiopteris:
 13.74.1, 2, 4, 5;  33 sterile frond (left) and fertile
syn. Matteuccia pensylvanica (Willdenow) Raymond, frond (right), bar = 10 cm
Pteretis nodulosa (Willdenow) Fernald, Struthiopteris (4 in.).
pensylvanica Willdenow
Ostrich fern
Very hardy, Zone 2
A large fern with fronds to 1.75 m (5.5 ft.) long and erect,
stoloniferous rhizomes. Grows well under medium light in
moist-wet garden soil or potting mix. This species is easy to
grow and is one of the most frequently sold ferns in the
eastern United States, where it is most often used as a foun-
dation planting around houses. The sterile leaves are de-
ciduous, but the fertile ones persist throughout the winter
and shed their spores in early spring, sometimes over snow.

Usually one or two new plants are produced each year from
the stolons.
The sterile leaves of Matteuccia struthiopteris are green
and oblanceolate, shaped like an ostrich feather, and grad-
ually taper toward the base, ending in highly reduced pin-
nae. The fertile leaves are brown, smaller than the sterile
ones, and have strongly ascending pinnae with enrolled
margins that protect the sporangia within. The species is
native to Europe, Asia, and North America.
var. pensylvanica (Willdenow) C. V. Morton. American
ostrich fern. Figures 13.74.1, 2, 4. Stipe scales about the
same color throughout (not darker in the center); pinna
lobes rounded. This is the common and widely planted va-
riety in North America.
var. struthiopteris (Figure 13.74.5). European ostrich
fern. Stipe scales bicolorous, darker in the center; pinna
lobes more truncate. This variety has been recently intro-
duced into cultivation in the United States.

Megalastrum
syn. Ctenitis section Subincisae (C. Christensen) Tindale
These medium- to large-sized ferns have decumbent to
erect rhizomes. The genus Megalastrum was previously clas-
sified in Ctenitis, but its basal basiscopic veins on the distal
pinnules spring from the costa, not the costule, and the lobe
supplied by this vein is widely adnate to the costa. Another
Figure 13.74.4. Matteuccia struthiopteris var. difference, best seen on the upper surface of the blade, is
pensylvanica: sterile frond (left) and fertile frond (right), that the veins end behind the margin in slightly enlarged,
bar = 10 cm (4 in.). club-shaped tips—that is, they do not run all the way to
 372 C 13

margin, as is typical in Ctenitis. Finally, the hairs on the Microgramma

upper surfaces of the rachises, costae, and costules are These appealing, small epiphytes are occasionally seen in
coarse, whitish, and strigose, not fine, reddish, and erect as fern collections. Their diminutive size makes them popular
in Ctenitis. Megalastrum contains about 40 species and is terrarium ferns, and they can also be grown on moss poles,
native to the American tropics, Africa, and Madagascar. bark, or in hanging baskets. Some species of Microgramma
The origin of the genus name is unknown. require more humidity than others, and these species do
best in terrariums or greenhouses.
Megalastrum subincisum (Willdenow) A. R. Smith & Microgramma can be identified by the following com-
R. C. Moran  13.75.1 bination of characteristics: rhizomes long-creeping and
syn. Ctenitis subincisa (Willdenow) Ching densely scaly, the scales not clathrate; fronds simple, entire;
Tender veins netted; sori in one row between the midrib and mar-
A large fern with an erect rhizome that sometimes forms a gin; indusia absent. See Polypodium for a comparison of
short trunk. Grows well under medium light in moist pot- this genus to other polypodiaceous genera.
ting mix. This species is rarely grown and is not widely The genus contains about 20 species and is entirely na-
available. tive to the New World tropics, with the exception of two
Megalastrum subincisum has straw-colored to brown species that are native to Africa and Madagascar. The genus
scales with curved marginal teeth. The blades are two- to name comes from the Greek mikros, small, and gramme,
three-pinnate-pinnatifid with sessile pinnules. The indu- line, referring to the slightly elongate sori in the species
sium tends to fall away early. The species is native to the from which the genus was first described—although most
American tropics. species in the genus have round sori.

Microgramma heterophylla (Linnaeus) Wherry

 13.76.1
Vine fern, climbing vine fern
b Tender, Zone 10
A small fern with long-creeping rhizomes. Grows well
under medium light in moist, drained potting mix or uncut
moss. The plants are sometimes grown in terrariums.
a The rhizomes of Microgramma heterophylla are slender,
0.5–1 mm wide, and the sterile blades reach 6 cm (2.5 in.)
long and are usually about 1.5 cm (0.6 in.) wide. In cultiva-
tion the upper blade surface is wrinkled and glabrous. The
species is native to southern Florida and the West Indies.

a
c

Figure 13.76.1. Microgramma heterophylla: a. habit, wild

Figure 13.75.1. Megalastrum subincisum: a. frond from plant, bar = 5 cm (2 in.); b. habit, cultivated plant, bar =
mature plant, bar = 5 cm (2 in.); b. frond from 5 cm (2 in.); c. fertile frond, bar = 5 cm (2 in.).
immature plant, bar = 5 cm (2 in.); c. pinnule, showing
thickened vein tips, bar = 5 mm (0.2 in.).
 F  F A  C 373

Microgramma lycopodioides (Linnaeus) Copeland

 13.76.2
Tender
A small fern with long-creeping rhizomes. Grows well
under medium light in moist, drained potting mix or uncut
moss.
Microgramma lycopodioides is characterized by rust-col-
ored rhizome scales and glabrous sterile blades that are lin-
ear to narrow-lanceolate, to 10 cm (4 in.) long, and acute at
the apex. The species is native to tropical America.
Figure 13.76.3.
Microgramma
megalophylla: habit,
Figure 13.76.2. bar = 5 cm (2 in.).
Microgramma
lycopodioides: a. habit,
bar = 5 cm (2 in.);
b. fertile frond, bar =
5 cm (2 in.).

a b

Microgramma megalophylla (Desvaux) de la Sota

 13.76.3
syn. Polypodium megalophyllum Desvaux
Tender Exceptionally large plants of Microgramma nitida might
A small-medium fern with long-creeping rhizomes. Grows be confused with M. megalophylla. The rhizome scales of
well under medium light in moist, drained potting mix or the latter, however, are triangular-ovate, centrally dark
uncut moss. from the base to the tip, and have a broad, thin, whitish
The most distinctive characteristic of Microgramma border. The rhizome scales of M. nitida are linear-lanceo-
megalophylla is its strongly flattened rhizomes about 10–30 late, centrally dark at the base and infrequently so to the
mm (0.5–1.2 in.) wide. This species also has the largest upper part of the scale, which is usually entirely whitish.
fronds in the genus: up to 24–50 cm (9–20 in.) long, 3–9
cm (1.2–3.5 in.) wide. The sori are round or slightly oblong
and large, to 6 mm (0.2 in.) in diameter, and arranged in a
medial, somewhat irregular row on both sides of the mid-
rib. The species is native to South America, especially Ama- b
zonia. It is reportedly used as a remedy for coughs.

Microgramma nitida (J. Smith) A. R. Smith

 13.76.4
syn. Polypodium palmeri Maxon Figure 13.76.4.
Tender Microgramma
nitida: a. habit,
A small fern with long-creeping rhizomes. Grows well bar = 5 cm
under medium light in moist-dry, drained potting mix or (2 in.); b. fertile
uncut moss. This species is easy to grow. frond, bar =
The rhizomes of Microgramma nitida are conspicuously 5 cm (2 in.).
flattened, as in the preceding species, but narrower, only
3–5 mm (0.1–0.2 in.) wide. The sterile blades are about 12
cm (5 in.) long, glabrous, slightly thick, and shiny. The sori a
are round and borne in a medial row on both sides of the
midrib. This species is native to Mexico, Central America,
Barbados, and Jamaica.
 374 C 13

Microgramma percussa (Cavanilles) de la Sota Microgramma piloselloides greatly resembles M. reptans

 13.76.5 but differs mainly by its slightly dimorphic sterile and fer-
syn. Pleopeltis percussa (Cavanilles) Hooker & Greville, tile fronds (as opposed to strongly dimorphic ones in M.
Polypodium percussum Cavanilles reptans). The fronds are typically oblong and densely whit-
Tender or slightly hardier ish scaly on the lower surface. The sori do not project be-
A small fern with long-creeping rhizomes. Grows well yond the margin. It is native to the West Indies.
under high light in moist-dry, drained potting mix or uncut
moss. This species is easy to grow, unlike Pleopeltis, from
which it has been separated.
The thick-textured fronds of Microgramma percussa are a b
narrowly linear-elliptic and have attenuate bases and apices.
The young sori lack peltate scales but have a dense mass of
persistent, branched, reddish filaments. The sporangia often
appear as yellow dots among the reddish filaments. The spe-
Figure 13.76.6. Microgramma piloselloides: a. habit, bar =
cies is widespread and common in tropical America.
5 cm (2 in.); b. sterile frond (left) and fertile frond
Ferns circulating as Pleopeltis macrocarpa and Poly-
(right), bar = 5 cm (2 in.).
podium lanceolatum in the U.S. trade were found to be Mi-
crogramma percussa.

Microgramma piloselloides (Linnaeus) Copeland Microgramma reptans (Cavanilles) A. R. Smith

 13.76.6  13.76.7
Tender syn. Microgramma ciliata (Willdenow) Alston,
A small fern with long-creeping rhizomes. Grows well Polypodium ciliatum Willdenow
under medium light in moist, drained potting mix or uncut Tender
moss. A small fern with long-creeping rhizomes and dimorphic
sterile and fertile fronds. Grows well under medium light in
moist, drained potting mix or uncut moss.
The sterile fronds of Microgramma reptans are up to 5
cm (2 in.) long, vary from ovate to oblong, and are beset
with many small, red-brown scales on the lower surface.
c The fertile fronds are linear and much narrower than the
sterile ones, 2–4 mm wide, with sori projecting beyond the
margin. The species is widespread throughout tropical
America.

a b

b
Figure 13.76.7. Microgramma reptans: a. habit of a small
plant, bar = 5 cm (2 in.); b. sterile fronds (three on the
Figure 13.76.5. Microgramma percussa: a. habit, bar = left) and fertile frond (right), bar = 5 cm (2 in.), after
5 cm (2 in.); b. fertile portion of frond, bar = 1 cm Hoshizaki (1982); c. scales on frond, bar = 1 cm
(0.4 in.); c. blade scales, bar = 1 mm. (0.4 in.), after Hoshizaki (1982).
 F  F A  C 375

Microgramma squamulosa (Kaulfuss) de la Sota

 13.76.8 b
Tender or semi-tender a
A small fern with long-creeping rhizomes. Grows well
under medium light in moist-dry, drained potting mix or
uncut moss. The plants have attractive dark green veins that
contrast strongly with the lighter green of the blade tissue.
Microgramma squamulosa is characterized by whitish Figure 13.76.9. Microgramma vacciniifolia: a. habit, bar
rhizome scales and scaly lower blade surfaces (especially the = 5 cm (2 in.); b. fertile frond, bar = 5 cm (2 in.). After
midrib) with highly dissected, scattered, whitish scales. The Hoshizaki (1982).
fronds grow to 16 cm (6.5 in.) long, are often reddish
tinged, and have the veins conspicuously darker green. The
species is native to tropical South America. lined veins. Microgramma lycopodioides differs by its rust-
colored rhizome scale and larger size.

Microlepia  13.77.1, 2

Microlepia consists of medium- to large-sized terrestrial
ferns found mostly in tropical or subtropical climates, with
a few species extending into warm-temperate climates. The
cultivated species have medium-creeping, clump-forming
rhizomes that bear fronds close enough to give a full-fo-
liage look to the plants. This appearance and their robust
growth habit and tolerance to irregular watering makes
a them popular garden plants in warmer climates. They also
make suitable houseplants.
c Microlepia strigosa is widely used in beds and foundation
b plantings in southern California. This vigorous grower re-
quires grooming to remove old fronds. It is the most com-
d
mon species in the trade but is often sold mixed with an-
other species tentatively identified as M. substrigosa. A tall
grower, M. platyphylla has large, bluish green segments and
Figure 13.76.8. Microgramma squamulosa: a. habit, bar = is useful in pots or as a screen in the landscape. Most species
5 cm (2 in.); b. fertile frond (left) and sterile frond
(right), bar = 5 cm (2 in.), after Hoshizaki (1982);
c. vein pattern, bar = 1 cm (0.4 in.), after Hoshizaki
(1982); d. rachis scale, bar = 1 mm, after Hoshizaki
(1982).

Microgramma vacciniifolia (Langsdorff & Fischer)

Copeland  13.76.9
Tender to semi-tender
A small fern with long-creeping rhizomes. Grows well
under medium light in moist, drained potting mix or uncut
moss.
Microgramma vacciniifolia has white rhizome scales and
sessile or nearly sessile fronds. The sterile fronds are round-
ish-ovate or longer, only 4–6 cm (1.5–2.5 in.) long, with
bases often obtuse-truncate. The fertile fronds tend to be
narrower and longer than the sterile ones. The species is
native to tropical America.
Microgramma vacciniifolia resembles M. squamulosa, a
species that differs by having larger fronds and darkly out- Figure 13.77.1. Microlepia strigosa: habit.
 376 C 13

Figure 13.77.2. Microlepia platyphylla: sori and

indusia.

of Microlepia are grown easily from spore, and mature

plants can be readily propagated by division of the rhizome.
The rhizomes of Microlepia are hairy, creeping,
branched, and clump-forming. The fronds are two-pinnate
to four-pinnate (rarely one-pinnate), anadromous, and usu-
ally have minute reddish hairs and a dense covering of short,
transparent hairs on the rachis or costae. Longer hairs may
Figure 13.77.3. Microlepia platyphylla: frond, bar = 5 cm
also be present on various parts of the blade. The veins are
(2 in.).
free, not netted. The sori vary from marginal to submar-
ginal and are covered by a shallow pouch or hood-shaped
indusium attached by its base and sides.
Microlepia, which contains 45 species, is native to Asia Microlepia speluncae (Linnaeus) T. Moore
and the Pacific basin, with one species, Microlepia strigosa,  13.77.4
extending into Africa and the American tropics. The genus Semi-tender to tender
name comes from the Greek mikros, small, and lepis, scale, A mostly medium-large fern with medium-creeping,
referring to the small, scale-like indusia. clump-forming rhizomes. Grows well under medium light
in moist garden soil or potting mix. In the United States the
Microlepia platyphylla (D. Don) J. Smith crested form is more common in cultivation than the typ-
 13.77.2, 3 ical form. Because it is not as cold tolerant as Microlepia
Semi-hardy to semi-tender strigosa, which it resembles, M. speluncae is infrequently
A large fern with medium-creeping, clump-forming rhi- seen in the trade.
zomes and blue-gray leaves. Grows well under medium to Microlepia speluncae resembles M. strigosa and M. sub-
high light in moist-dry garden soil or potting mix. This spe- strigosa but differs by having more herbaceous blades that
cies is easy to cultivate. It has erect, coarse foliage and can wilt quickly after picking, veins not raised on the lower
form large, dense clumps if not regularly divided. blade surface, and smaller indusia attached mainly at the
Microlepia platyphylla is characterized by triangular base and more submarginal (mostly about two lengths dis-
blades up to two-pinnate-pinnatifid with narrowly trian- tant from the margin). This species is widely distributed in
gular, pinnatifid to pinnately lobed, serrate pinnules. The the tropics throughout the world.
midribs of the ultimate segments bear fine hairs, but other- ‘Corymbifera’. (Microlepia pyramidata Horticulture).
wise the plants are glabrous. The species is native from Figure 13.77.4. Tips of blades and pinnae tasseled. Fronds
India and southeastern Asia to the Philippines. light green, arching gracefully.
 F  F A  C 377

submarginal or reaching the leaf margin. The species is na-

tive to the Himalayas, Sri Lanka, and from southeastern
Asia to Japan and Polynesia.
The native Hawaiian plant (also cultivated in Hawaii)
and some trade plants imported from Holland are some-
times called Microlepia strigosa but are here referred to M.
substrigosa.
Microlepia strigosa is a highly variable plant, but few of
the cultivated variants are distinct except for forma mac-
faddeniae C. V. Morton (Figure 13.77.5d, e), which has
narrowly linear fronds and pinnae greatly reduced and con-
densed, consisting of a cluster of fan-shaped pinnules.

Microlepia substrigosa Tagawa  13.77.6, 7

Semi-hardy to semi-tender
A large fern with medium-creeping, clump-forming rhi-
zomes and fronds more than 1.5 m (5 ft.) tall. Grows well
a under medium to high light in moist-dry garden soil or
potting mix. This species is easy to cultivate and is a rapid
grower. Microlepia substrigosa is often misidentified as M.
strigosa and M. firma.
Microlepia substrigosa resembles M. strigosa, but its
blades are nearly triangular and up to three-pinnate-pin-
natifid, with the lower pinnae triangular. Many indusia ap-
b pear to be on a short lobe of the leaf margin, and the edges
of most indusia do not reach the leaf margin. The species is
native to Japan, Taiwan, China, and possibly the Hawaiian
Islands (where it circulates as M. strigosa).
Two slightly different plants are encountered in culti-
vation, and they are tentatively identified here as Microlepia
Figure 13.77.4. Microlepia speluncae ‘Corymbifera’: substrigosa. Both are mistakenly sold as M. strigosa, which
a. frond, bar = 5 cm (2 in.); b. lower pinna, bar = 2 cm has ovate, predominately two-pinnate to two-pinnate-pin-
(0.8 in.). natifid blades. One of these plants is widely distributed by
a European grower and the other circulates in Hawaiian
and southern California gardens (under the misapplied
Microlepia strigosa (Thunberg) C. Presl name of M. firma). The European plant has an erect habit
 13.77.1, 5 and broadly triangular lower pinnae (Figure 13.77.6). The
Lace fern Hawaiian and southern California plant has a more spread-
Semi-hardy to semi-tender; reportedly can grow in Zone ing habit and narrower lower pinnae abruptly widened at
6 if winter-protected the base (Figure 13.77.7). Although the identity of these
plants is uncertain, they are not M. strigosa as generally
A medium-large fern with medium-creeping, clump-form-
known.
ing rhizomes. Grows well under medium or high light in
moist-dry garden soil or potting mix. The plants are easy to
cultivate and grow rapidly, although the species is known to Microsorum Link
tip-burn when exposed to temperatures of −7°C (19°F) or syn. Diblemma, Phymatodes, Phymatosorus, Polypodium in
lower for several days. The fronds spread or arch gracefully. part
Microlepia strigosa has yellowish brown or whitish rhi- The polypodiaceous genus Microsorum is best known for
zome hairs. The blades are two-pinnate-pinnatifid to three- the so-called cobra plant, a form of Microsorum punctatum
pinnate (at the base of the larger pinnae), ovate, and her- that is heavily short-crested at the tips of its erect, strap-
baceous. The lower one or two pairs of pinnae are slightly shaped fronds. The common name probably alludes to a
reduced and narrowly linear-triangular. The veins are dis- fanciful resemblance to a cobra ready to strike. The cobra
tinctly raised on the lower blade surface. The indusia are plant is widely used in Florida and the tropics as a ground
attached at the base and up the sides, and many appear at cover or edging. In other parts of the United States it is
the base of a leaf-margin notch. Most have their upper edge used as a potted plant and does well indoors. Several forms
 d

Figure 13.77.5. Microlepia strigosa: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar = 2 cm (0.8 in.); c. lower pinnae,
showing variations, bar = 2 cm (0.8 in.); d. forma macfaddeniae frond, bar = 5 cm (2 in.); e. forma macfaddeniae frond
detail, bar = 1 cm (0.4 in.).
 F  F A  C 379

a a

b b

Figure 13.77.6. Microlepia substrigosa from European Figure 13.77.7. Microlepia substrigosa from Hawaiian
growers: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar gardens: a. frond, bar = 5 cm (2 in.); b. lower pinna, bar
= 2 cm (0.8 in.). = 2 cm (0.8 in.).

have long, arching fronds, and these are attractive as hang- Most species of Microsorum have included veinlets
ing plants. pointing in all directions, not just toward the blade margin
Besides the cobra plant, the East Indian polypody (Mi- as in many other polypodiaceous ferns. The large sunken
crosorum scolopendria) is commonly cultivated. In tropical sori of Phymatosorus (Phymatodes) intergrades into Micro-
climates this species and its close relatives are used as sorum, and because other reliable distinguishing characters
ground covers, basket plants, or over rock walls. Although have not been found, Nooteboom (1997) reduced Phyma-
the branching rhizome is capable of growing to 6 m (20 ft.) tosorus to Microsorum. For more details, see Bosman (1991)
in length, cultivated plants are seldom allowed to grow this and Nooteboom (1997). Microsorum species were formerly
long. As with many ferns, the fronds may differ in shape listed under Polypodium and may be more familiar to gar-
on the same plant, ranging from entire to pinnatifid. deners under that name. See Hoshizaki (1982) for details
The genus consists of medium- to large-sized epiphytes on cultivated species. The entry for Polypodium in this
with long- or short-creeping rhizomes bearing peltate or chapter includes a comparison of Microsorum with other
false-peltate, clathrate or subclathrate scales. The blades are polypodiaceous genera.
either entire, lobed, or pinnatifid, or (in Microsorum lu- Microsorum consists of about 50 species native to tropi-
cidum) pinnate. The veins are netted (this is hard to see in cal and subtropical regions of Africa, southeastern Asia,
species with thick blades) and have within the areoles northern Australia, and the Pacific Islands. It has become
branched veinlets that may point in any direction. The sori naturalized in several areas in the New World. The genus
are separate, small to larger, round or less often elongate, name comes from the Greek mikros, small, and soros,
and irregularly scattered or in partly irregular rows, and mound, referring to the small, scattered sori, although some
they lack an indusium. species have larger sori.
 380 C 13

Microsorum commutatum (Blume) Copeland

 13.78.1
syn. Phymatosorus commutatus (Blume) Pichi-Sermolli, b
Polypodium vitiense Baker
Tender
A medium-sized fern with moderately-creeping rhizomes.
Grows well under medium light in moist, well-drained pot-
ting mix or uncut moss.
The Microsorum commutatum stipe is narrowly winged
toward the blade base; the blade is herbaceous, broad, and
deeply lobed. Both sides of the frond midrib have a few
small, dark, persistent, ovate-rounded scales. The sori are
small, in two irregular rows, and not sunken. The species is
native to Sumatra, Java, Borneo, Philippines, Sulawesi,
New Guinea, and certain Pacific Islands.
‘Southern Cross’. Lobe margins irregularly and shal-
lowly to deeply toothed, lobed, or fringed. Sori absent. This
is the cultivar grown in the United States. Sometimes cir-
culates incorrectly as Polypodium australe ‘Cambricum’.

Figure 13.78.2. Microsorum fortunei: a. habit, bar = 5 cm

(2 in.); b. sori, bar = 1 cm (0.4 in.).
b

Microsorum fortunei has strap-shaped fronds to 60 cm (2

ft.) long and 3–5 cm (1.2–2 in.) wide. They are attenuate
on both ends. The sori are not sunken, round, up to 4 mm
(0.2 in.) wide, in one (rarely two) slightly irregular row on
either side of the midrib. The species is native to Indochina,
China, Taiwan, and Japan.
a
Figure 13.78.1. Microsorum
commutatum ‘Southern
Microsorum grossum (Langsdorff & Fischer)
S. B. Andrews  13.78.3
Cross’: a. frond, bar = 5 cm
syn. Microsorum scolopendria auct. non (N. L. Burman)
(2 in.); b. midrib and costa
Copeland, Phymatosorus grossus (Langsdorff &
scales, bar = 1 cm (0.4 in.).
Fischer) Brownlie, Polypodium grossum Langsdorff &
Fischer
Tender
Microsorum fortunei (T. Moore) Ching A medium-sized fern with long-creeping rhizomes. Grows
 13.78.2 well under high light in drained, moist to moist-dry gar-
syn. Polypodium normale D. Don den soil or potting mix. The plants are widely cultivated in
Semi-hardy the tropics, where they are often used as a ground cover.
A medium-sized fern with short-creeping rhizomes. Grows The foliage can be trimmed like a hedge. The rhizomes
well under medium light in moist-dry potting mix. The tend to outgrow pots.
plants tend to become disfigured when grown under con- Microsorum grossum resembles M. scolopendria but dif-
ditions of low humidity. fers by having up to 10 pairs of lobes (instead of 4) per
 F  F A  C 381

grower and less tender than M. grossum and M. scolopen-

dria, it is favored in subtropical gardens. Otherwise all three
plants have similar cultural requirements. Nooteboom
(1997) considered M. grossum and P. banjeriana synonyms
a b of M. scolopendria.

Microsorum insigne (Blume) Copeland

 13.78.4
syn. Colysis insignis (Blume) J. Smith, Microsorum
dilatatum (Beddome) Sledge, M. hancockii (Baker)
Ching
Tender to semi-tender
A medium-sized fern with medium-creeping rhizomes.
Grows best under medium to high light in moist-dry pot-
c ting mix or uncut moss kept well drained. This species is a
robust grower, thriving in humid or dry environments.
Once established, it tolerates some irregular watering.
The fronds of mature Microsorum insigne are oblong to
lanceolate and deeply pinnately lobed. The lobes are few
to many and oblong-lanceolate with acuminate apices. The
terminal lobe resembles the lateral ones. The sori are round-
ish, small, and scattered. The species is native to India,
southeastern Asia, the Malayan Archipelago, China, and
Japan.
This species varies in the size and number of lobes.
Plants with many lobes are said to intergrade with the
e

Figure 13.78.3. Microsorum grossum: a. frond, bar = a Figure 13.78.4.

about 5 cm (2 in.), after Langsdorff and Fischer (1810); Microsorum insigne:
b. sori, upper frond surface, bar = 1 cm (0.4 in.), after a. frond, bar = 5 cm
Hoshizaki (1982); c. rhizome scales, bar = 1 mm, after (2 in.); b. sori, bar =
Hoshizaki (1982); d. Phymatodes [= Microsorum] 1 cm (0.4 in.).
banjeriana frond, bar = 5 cm (2 in.); e. Phymatodes
[= Microsorum] banjeriana rhizome scale, bar = 1 mm.

blade, and the plants tend to be larger and epiphytic rather

than terrestrial. The species is native to Australia and cen-
tral and eastern Polynesia.
Most plants cultivated in Hawaii and Florida are Micro-
sorum grossum. A similar species that sometimes circulates
under the incorrect name of Polypodium alternifolium is
tentatively identified as Phymatodes [= Microsorum] banjeri-
ana N. Pal & S. Pal (Figure 13.78.3d, e). This species is
difficult to separate from M. grossum; the most reliable char- b
acter is in the rhizome scale. Those of Phymatodes banjeri-
ana are rounded at the base and abruptly narrowed into a
long, slender, tapered apex, whereas the rhizome scales of
M. grossum are triangular. Because P. banjeriana is a robust
 382 C 13

smaller, more triangular form previously known as Micro-

sorum hancockii (Baker) Ching. b

Microsorum linguiforme (Mettenius) Copeland

 13.78.5
Tender or slightly hardier
A small to medium fern with medium- to long-creeping
rhizomes. Grows well under medium light in moist pot-
ting mix or uncut moss. This species is a rapid and robust
grower.
The long-creeping rhizome of Microsorum linguiforme is
somewhat flattened and often has branches that are oppo-
site or nearly so. The fronds are broadly ovate, entire, and
often contracted above the base. The blade apex is round-
ish to acuminate, and the base varies from tapered to cor- a
date. The veins are darkened. The sori are round and few.
This species is native from India and the Malayan Archi-
pelago to Fiji.
Figure 13.78.6. Microsorum longissimum: a. frond, bar =
5 cm (2 in.); b. vein pattern, bar = 1 cm (0.4 in.).

wide. Microsorum samarense also has strap-shaped fronds,

but its veins are hard to see and the upper surfaces of the
fronds are dull, not shiny. Microsorum longissimum is native
to Borneo, Sarawak, and the Philippines.

Microsorum lucidum (Roxburgh) Copeland

 13.78.7
syn. Phymatosorus cuspidatus J. Smith, P. lucidus
(Roxburgh) Pichi-Sermolli
Semi-tender
Figure 13.78.5. Microsorum
A medium-large fern with medium- to long-creeping rhi-
linguiforme: habit, bar =
zomes. Grows well under medium light in drained, moist-
5 cm (2 in.).
dry garden soil or potting mix. This species is easy to culti-
vate once established, and it may overgrow surrounding
plants.
The rhizome of Microsorum lucidum is green, fleshy, and
2.5 cm (1 in.) or more in diameter. The fronds are one-
pinnate and dark green with obscure veins. The sori are ar-
Microsorum longissimum J. Smith ex Fée ranged in one row, closer to the costa than the margin; they
 13.78.6 are not sunken. The species is native to India, southwestern
syn. Polypodium myriocarpum C. Presl ex Mettenius; not China, and southeastern Asia.
Polypodium longissimum Blume
Tender Microsorum membranifolium (R. Brown) Ching
A medium-sized fern with short-creeping rhizomes and  13.78.8
handsome shiny fronds bearing an attractive vein pattern. syn. Microsorum alternifolium of authors, Phymatosorus
Grows well under medium light in drained, moist to moist- nigrescens (Blume) Pichi-Sermolli, Polypodium
dry potting mix or uncut moss. nigrescens Blume
Microsorum longissimum has blackish rhizome scales. Pimple fern
The shiny fronds are strap-shaped, to 35–95 cm (14–38 Tender
in.) by 1–9 cm (0.5–3.5 in.), with the primary veins con- A large fern with medium-creeping rhizomes. Grows well
spicuous and forming one series of large areoles next to the under medium light in drained, moist potting mix or uncut
midrib. The sori are round or elongate and 0.5–1.5 mm moss. The plants often volunteer from spores. This species
 F  F A  C 383

a
b

Figure 13.78.8. Microsorum membranifolium: a. frond,

bar = 5 cm (2 in.); b. sori and vein pattern, upper
surface, bar = 1 cm (0.4 in.); c. raised sorus, upper frond
surface, bar = 2 mm.
b

Figure 13.78.7. Microsorum lucidum: a. frond, bar = b

5 cm (2 in.); b. pinna, bar = 5 cm (2 in.), after
Hoshizaki (1982).

has long been known as Polypodium nigrescens and is some-

times mislabeled in the trade as P. diversifolium Willdenow
and as P. alternifolium Willdenow.
Microsorum membranifolium has large, deeply lobed
fronds with distinct dark veins. The sori are deeply em-
bossed on the upper surfaces of the blades, forming a stout
cylindrical protuberance to 2 mm high. They are borne in
one row between the costae and margins. The species is na-
tive from India to southeastern Asia and Australia.

Microsorum musifolium (Blume) Copeland

 13.78.9
Tender
A large fern with short-creeping rhizomes and spreading
fronds clustered into an irregular basket shape. Grows well Figure 13.78.9. Microsorum musifolium: a. frond, bar =
under medium to high light in drained, moist potting mix 5 cm (2 in.); b. sori and vein pattern, bar = 1 cm
or uncut moss. Keep the plants out of direct sun and avoid (0.4 in.).
 384 C 13

overhead watering, which can cause leaf spots. The veins Microsorum pteropus (Blume) Copeland
form a handsome pattern.  13.78.11
The fronds of Microsorum musifolium are oblanceolate, syn. Colysis pteropus (Blume) Bosman
to 1 m (3 ft.) long, 10 cm (4 in.) wide, and sessile; the fronds Tender
are truncate to obtuse at the base and often narrowed A small fern with short-creeping rhizomes. Grows well
above. The venation is regular and conspicuous, with 6–10 under medium light in wet, coarse garden soil or potting
primary connective veins between the pairs of secondary mix with sand; can also be planted in drained pots or pots
veins. The species is native to southeastern Asia and the set in water. The plants can be grown submerged and are
Malayan Archipelago. often used in aquariums. They can remain submerged for
We have not followed Nooteboom (1997), who con- years without ill effects; however, sori are formed only on
cluded that this species was the same as Microsorum punc- emergent fronds, not those under water. The fronds are
tatum. Microsorum musifolium differs by its conspicuous vulnerable to leaf spots, which usually can be controlled by
veins and firm but thin translucent blades. In contrast, live fungicide sprays.
plants of M. punctatum have inconspicuous veins and leath- The fronds of Microsorum pteropus are simple to
ery blades. trilobed, with the center lobe the largest. Submerged fronds
are small and entire. The buds often form on the lower sur-
Microsorum papuanum (Baker) Parris face where sori would normally form. The species is native
 13.78.10 from India to southern China, the Malay Archipelago, New
Tender, Zone 9 Guinea, and Japan. In nature it typically grows along stream
A medium-sized fern with creeping rhizomes bearing fronds banks prone to flooding, or in the spray of waterfalls.
close or more distant from each other. Grows under high to
medium light in well-drained, moist to moist-dry potting
mix.
Microsorum papuanum differs from the closely related
Microsorum scolopendria by its leathery and often crisped
blade that is simple and entire or has one to three lobes,
with the central lobe the longest. The sori are round to oval,
often irregularly so, and deeply sunken into the frond (em-
bossed on the upper surface). In cultivated plants, the sori
reach 13 mm (0.5 in.) in length and to 8 mm (0.3 in.) in Figure 13.78.11. Microsorum
width. It is native to Malaysia, New Guinea, and the Pacific pteropus: habit, bar = 5 cm
Islands. (2 in.), after Hoshizaki (1982).

Figure 13.78.10. Microsorum

papuanum: a. frond, bar = 5 cm
(2 in.); b. sori, bar = 1 cm (0.4 in.). Microsorum punctatum (Linnaeus) Copeland
 13.78.12, 13
syn. Microsorum irioides (Poiret) Fée, M. polycarpon
(Cavanilles) Tardieu, Polypodium integrifolium Lowe,
P. iriodes Poiret, P. punctatum (Linnaeus) Swartz
Tender to semi-tender
A medium-sized fern with very short-creeping rhizomes.
a
Grows well under medium light in drained, moist-dry pot-
ting mix or uncut moss. It is easy to grow.
Microsorum punctatum has fronds that are strap-shaped
b or wider, to 1 m (3 ft.) long or more by 1.5–10 cm (0.6–4
in.) or more wide. They are mostly subcoriaceous, their
apex round to acuminate, their base cordate to tapered. The
leaves are often ruffled, fringed, or forked near or at the tips.
The veins are hard to see because of the thick blades. The
species is native to the Old World tropics and subtropics.
Wild plants are variable, and some may represent dif-
ferent species. Microsorum whiteheadii A. R. Smith & Hos-
 F  F A  C 385

kizaki is such a species and has recently been introduced

into cultivation. It differs from M. punctatum by its thick,
b succulent, relatively broader oblanceolate blade.
Microsorum punctatum is the most common Microso-
rum in the trade and is available in many forms, such as
wide or narrow, short or erect or nearly decumbent, crested,
and variously emarginate. New selections are constantly
being introduced, and the trade plants are loosely named.
Some common cultivars are:
‘Climbing-bird’s-nest fern’. Fronds like the bird’s-nest
fern (Asplenium nidus) but softer and arching.
‘Grandiceps’. Figure 13.78.13a. Similar to ‘Climbing-
bird’s-nest fern’ but loosely and thinly crested.
‘Ramosum’. (‘Cristatum’) Figure 13.78.13b; Plate 34.
Cobra plant. Fronds erect and densely crested.

Microsorum pustulatum (G. Forster) Copeland

 13.78.14
syn. Microsorum diversifolium (Willdenow) Copeland,
Phymatosorus diversifolius (Willdenow) Pichi-Sermolli,
Phymatosorus pustulatus (G. Forster) M. F. Large,
Polypodium diversifolium Willdenow, Polypodium
Figure 13.78.12. Microsorum pustulatum of authors, not G. Forster, Polypodium
punctatum: a. frond variations, scandens of authors, not G. Forster
bar = 5 cm (2 in.); b. sori, bar Kangaroo fern
= 1 cm (0.4 in.). After Semi-hardy
a Hoshizaki (1982).
A medium-sized fern with long-creeping rhizomes. Grows
well under medium light in drained, moist potting mix or
uncut moss. This species, long known as Phymatosorus di-
b versifolius, grows slowly but tolerates cold better than oth-
ers in the genus. It is grown outdoors in the warmer parts of
England.
Microsorum pustulatum has ovate to narrowly ovate rhi-
zome scales. The fronds are entire to broadly and deeply
lobed, dark green, and leathery. The veins are clearly visible.
The sori are in one row between the costa and margin. The
b
species is native to Australia and New Zealand.
Microsorum scolopendria and its allies differ by ciliate or
spiny rhizome scales, obscure veins, and the sori in one or
b two rows between the costa and margin.

Microsorum samarense (J. Smith ex Mettenius) Bosman

 13.78.15
syn. Diblemma tenuilore (J. Smith ex Mettenius) Ching,
Polypodium tenuilore J. Smith ex Mettenius
Tender
A small to medium fern with short-creeping rhizomes.
a Grows well under medium light in moist, drained potting
mix or uncut moss.
Microsorum samarense is characterized by linear-lanceo-
Figure 13.78.13. Microsorum punctatum, variations late fronds, obscure veins, and (often) elongate sori. The
in cultivars: a. ‘Grandiceps’, fronds pendent; leathery fronds are generally 25–45 cm (10–18 in.) long
b. ‘Ramosum’, fronds erect, all from the same plant. Bar and 5–10 mm (0.2–0.4 in.) wide. The margins tend to roll
= 5 cm (2 in.). After Hoshizaki (1982). inward when dry. The sori vary from round to (often) elon-
386 C 13

gate and are often concentrated on the marginal veins. The

species is native to the Philippines, where it grows as an epi-
phyte and on limestone.
a The plants vary tremendously in width. The narrowest
form circulates in horticulture as Diblemma tenuilore and
seems be shorter than usual, with many fronds having
forked tips. The wider form is rarely grown.

Microsorum scandens (G. Forster) Tindale

 13.78.16;  5
syn. Phymatosorus scandens (G. Forster) Pichi-Sermolli,
Polypodium pustulatum of authors, not G. Forster,
Polypodium scandens G. Forster
Fragrant fern
Semi-hardy
d
A small-medium fern with slender, long-creeping rhizomes.
b
Grows well under medium light in moist garden soil, pot-
ting mix, or uncut moss. The plants are well suited to bas-
ket culture and will grow up tree trunks if initially provided

Figure 13.78.14. Microsorum pustulatum: a. frond, bar = a

5 cm (2 in.); b. habit and frond variation, bar = 5 cm
(2 in.); c. visible veins, bar = 1 cm (0.4 in.); d. rhizome
scale, bar = 1 mm.

a b

Figure 13.78.16. Microsorum scandens:

Figure 13.78.15. Microsorum samarense: a. habits, bar = a. habit, showing sterile frond (left) and
5 cm (2 in.); b. sori and vein patterns, bar = 1 cm fertile frond (right), bar = 5 cm (2 in.);
(0.4 in.). b. sori, bar = 1 cm (0.4 in.).
 F  F A  C 387

with some moisture-retaining pockets in which to root. green, leathery, and flat. The veins are hard to see. The sori
The fronds are evergreen and fragrant when crushed. vary from round or oblong and are usually arranged in two
The fronds of Microsorum scandens are entire or pinnat- irregular rows on each side of the midrib. The species is na-
ifid, herbaceous, narrowly elliptic, to about 60 cm (24 in.) tive to the tropics of the Old World.
long, 16 cm (6.5 in.) wide. The frond base is long-decur- As construed by Nooteboom (1997), Microsorum scolo-
rent on the stipe. The lobes of the blade are narrow, 4–10 pendria consists of a single variable species; however, other
mm (0.2–0.4 in.) wide, and separated by broad, U-shaped botanists believe it consists of several. Plants of M. scolopen-
sinuses. The sori are round or elongate, borne in one row dria with more than four pairs of lobes are recognized by
between costa and margin, and slightly raised on the upper some authors as M. grossum, whereas those with four or
surface of the frond. The species is native to Australia and fewer are called M. scolopendria (and these plants are by no
New Zealand. means uniform). Most plants cultivated in the United States
are M. grossum.
Microsorum scolopendria (N. L. Burman) Copeland
 13.78.17 Microsorum steerei (Harrington) Ching
syn. Phymatosorus scolopendria (N. L. Burman) Pichi-  13.78.18
Sermolli, Polypodium scolopendria N. L. Burman Tender
East Indian polypody A small-medium fern with short-creeping rhizomes and
Tender thick and leathery fronds up to 40 cm (16 in.) long, though
A medium-large fern with long-creeping rhizomes. Grows usually shorter in cultivation. Prefers low to medium light
well under high light in drained, moist to moist-dry gar- and a well-drained, moist potting mix. Plants grow some-
den soil, potting mix, or uncut moss. what slowly and seem to do best at temperatures of 21°C
The long-creeping rhizome bears triangular or ovate- (70°F) or more year-round. This species is striking because
triangular scales that are spiny or ciliate on the margins. of its blue-green iridescent fronds.
The blade of Microsorum scolopendria is deeply lobed into The fronds of Microsorum steerei are like those of a small,
four or fewer pairs of segments that are somewhat yellowish fleshy M. punctatum but are more pointed and distinctly

a c

Figure 13.78.17.
Microsorum
scolopendria: a. habit,
bar = 5 cm (2 in.); b
b. sori, bar = 1 cm
(0.4 in.); c. rhizome
scale, bar = 1 mm.
 388 C 13

Once established, this species is a robust grower and may be

trained to grow up tree trunks or branches.
Microsorum superficiale is characterized by simple, lan-
ceolate, entire, herbaceous fronds that are borne apart on a
slender rhizome. The sori are scattered on the lower sur-
face. It is native to China, Japan, and Vietnam.

Microsorum viellardii (Mettenius) Copeland

 13.78.20
syn. Phymatodes viellardii (Mettenius) Fournier
Semi-tender
Figure 13.78.18. Microsorum
steerei: habit, bar = 5 cm A medium-large fern with long-creeping rhizomes. Grows
(2 in.). well under medium to high light in drained, moist to
moist-dry garden soil or potting mix. The plants are fairly
robust growers in baskets or in the ground. This species cir-
iridescent. The plants in cultivation were introduced from
Chumphon Province in southern Thailand by Chanin
Thorut, who reported the plants growing on rocks in a a
deep, shady, humid forest. The fern has been reported to
grow on limestone in China, Taiwan, and Vietnam.
b
Microsorum superficiale (Blume) Ching
 13.78.19
syn. Leptochilus buergerianum (Miquel) Bosman,
Microsorum buergerianum (Miquel) Ching,
Neocheiropteris superficialis (Blume) Bosman
Semi-tender
A small fern with medium- to long-creeping rhizomes.
Grows well under medium light in moist to moist-dry gar-
den soil, potting mix, or uncut moss kept well drained.

Figure 13.78.19.
Microsorum superficiale:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
1 cm (0.4 in.).

Figure 13.78.20. Microsorum viellardii: a. frond, bar =

a 5 cm (2 in.); b. sori and vein pattern, bar = 1 cm
(0.4 in.), after Hoshizaki (1982); c. rhizome scale, bar =
1 mm, after Hoshizaki (1982).
 F  F A  C 389

culated among growers as “Mt. Elgan fern,” a name of un- long-pentagonal blades are one-pinnate-pinnatifid to two-
certain origin. pinnate, with linear, entire or serrate, adnate segments. The
Microsorum viellardii has rounded rhizome scales up to surfaces of the blades are smooth except for some fine hairs
6 mm (0.2 in.) wide. The blade is deeply lobed, and the on the upper surface. The veins are prominent and free.
lobes are narrowed toward the base. The sori are submar- The indusium is linear, distinctly submarginal, and up to
ginal and in one row between the midrib and margin. This 0.8 mm wide. The species occurs from Mexico to Central
species is endemic to New Caledonia. America and the Galápagos Islands.

Mildella Nephrolepis  13.80.1, 2

Mildella is barely distinct from Cheilanthes but is recog- Sword fern, fish-bone fern
nized here because it has been used in by most major floras. The genus Nephrolepis is best known for the Boston fern
It differs from Cheilanthes by having a submarginal indu- (Nephrolepis exaltata ‘Bostoniensis’) and the numerous cul-
sium with the green segment margin extending beyond it. tivars derived from it. It is widely grown, and in temperate
The genus, which consists of about eight terrestrial species, parts of the United States it is used mainly indoors but may
occurs in Central America, the Galápagos Islands, Haiti, be placed outdoors during warm weather. Some Nephro-
and from northern India to China. It is named in honor of lepis plants can be grown outdoors year-round in subtrop-
Carl August Julius Milde (1824–1871), a German student ical areas, but they may show cold sensitivity. The tuber
of ferns and bryophytes. See Cheilanthes for a comparison sword fern (N. cordifolia) is commonly grown as a ground
of this genus to other cheilanthoid genera. cover in tropical and subtropical areas and tolerates cool
weather much better than the Boston fern. Other species of
Mildella intramarginalis (Kaulfuss ex Link) Trevisan Nephrolepis are rarely grown because they are tender and
 13.79.1 too large or coarse for ornamental purposes. All species
syn. Cheilanthes intramarginalis (Kaulfuss ex Link) grow better with strongly filtered light. The most common
Hooker problem in caring for established plants is overwatering
Semi-tender (?) combined with poor drainage. These ferns generally toler-
A small to medium fern with clump-forming rhizomes and ate short periods of dryness.
clustered fronds. Grows well under medium to high light in Nephrolepis reproduces vegetatively by runners or sto-
moist garden soil or potting mix. lons. These parts are produced just behind the rhizome
Mildella intramarginalis resembles a small Pteris. It has apex, and if their large numbers create an untidy appear-
short, triangular, sessile pinnae, the lowermost of which are ance, they can be removed without harming the plant.
prolonged on their basiscopic side. The narrowly ovate to Some stolons, however, should be encouraged to root so
that fresh new growth can eventually replace older growth,
which may last a few years or less.
The dividing and repotting of Nephrolepis is best done in
the spring when new growth appears. In temperate or sub-

Figure 13.79.1. Mildella intramarginalis: a. fertile fronds

(left) and sterile frond (right), bar = 2.5 cm (1 in.); b. tip
of fertile segment, bar = 1 mm. Figure 13.80.1. Nephrolepis cordifolia: habit.
390 C 13

Nephrolepis contains about 30 species and occurs

throughout the tropics and subtropics worldwide. The
genus name comes from the Greek nephros, kidney, and
lepis, scale, referring to the kidney-bean-shaped indusia.

Nephrolepis acutifolia (Desvaux) H. Christ

 13.80.3
Tender
A large fern with short, erect, stoloniferous rhizomes and
fronds over 1 m (3 ft.) long. Grows well under high light in
moist, well-drained potting mix.
Nephrolepis acutifolia can be distinguished by its leathery
fronds, reddish scaly rachises, and linear, marginal indusia.
It is native to tropical Africa and from southeastern Asia to
Figure 13.80.2. Polynesia.
Nephrolepis fronds:
N. exaltata (left) and
N. cordifolia (right).

tropical climates most new divisions made in cooler b

weather will not receive enough heat to allow them take
hold; however, robust growers may take hold readily in
cooler weather.
Nephrolepis can be identified by the combination of one-
pinnate blades, numerous pinnae (generally 20–50 pairs)
jointed to the rachis, and sori that are round, bean-shaped,
or crescent-shaped (rarely linear) and covered by a similar- Figure 13.80.3. Nephrolepis acutifolia:
shaped indusium attached at its base. Another distinguish- a. frond, bar = 10 cm (4 in.); b.
ing trait is the many long, slender, scaly runners (stolons) pinna, bar = 1 cm (0.4 in.).
formed at the base of the plant. An unusual characteristic is
that the fronds are usually coiled at the apex, never com-
pletely unrolling. Most species are epiphytes, but some
grow on rocks or, rarely, in soil. The main rhizomes are Nephrolepis biserrata (Swartz) Schott  13.80.4
more or less short and erect, never long-creeping. Giant sword fern
Some species of Nephrolepis are difficult to distinguish. Tender, Zone 9
The characteristics described in the species entries apply A medium to large fern with short, erect, stoloniferous rhi-
only to mature fronds. The scales and hairs on young zomes. Grows well under medium to high light in moist
fronds often differ from those of older plants and are unre- garden soil or potting mix. The pendent or arching fronds
liable for identification. Also, weathering can cause loss of may reach 5 m (16 ft.) long, but the pinnae are too far apart
hairs and scales, which are important identifying features. for this plant to be an attractive ornamental. The fronds
 F  F A  C 391

is widely used as a ground cover in tropical and subtropical

climates, second only to the Boston fern (Nephrolepis exal-
tata) in United States use. Tuber sword fern is less sensitive
to cool temperatures than Boston ferns—although the tips
burn if temperatures last one week or more at −7°C (19°F)
—and it maintains better color and active growth. This
species is often mislabeled in the trade as N. exaltata.
Nephrolepis cordifolia has two particularly distinctive
characteristics. First, whereas most species in the genus have
pendulous or arching fronds, those of N. cordifolia are stiff
and erect (except for a few cultivars), which is why it can be
used as a low border hedge in the tropics. Second, on the
runners are produced large, globose, scaly tubers, which are
often underground. Other characteristics of the species in-
clude stipe bases with spreading whitish or tan scales; pin-
a nae less than 4 cm (1.5 in.) long, usually overlapping, and
mostly glabrous; and the upper surface of rachis scaly, the
scales with a dark center. The indusia are bean-shaped to
semi-circular, with a broad sinus or none, mostly opening
b toward the pinna apex. The species is native to tropical

a
c

Figure 13.80.4. Nephrolepis biserrata: a. frond, bar =

10 cm (4 in.); b. pinna, bar = 1 cm (0.4 in.); c. pinna
margin, bar = 3 mm (0.1 in.).

b
are lax and hairy in general aspect. Plants volunteer readily
from spore, accounting for most of the plants in cultiva-
tion. Requires winter protection in southern California.
Nephrolepis biserrata can be separated from many others
in Nephrolepis by the hairy lower surfaces of the pinnae,
rachises, and costae. The stipe bases bear loose, spreading,
narrow, light brown, concolorous scales, and the pinnae
are typically 7 cm (3 in.) long or more with doubly crenate
margins (large crenations alternating with small ones). The
c
indusia are submarginal and round-reniform. The species is
native to tropical America, Africa, and southeastern Asia.
Trade plants by this name are actually Nephrolepis
falcata.

Nephrolepis cordifolia (Linnaeus) C. Presl

 13.80.1, 2, 5, 6;  35 d
syn. Nephrolepis auriculata (Linnaeus) Trimen, N.
tuberosa (Bory ex Willdenow) C. Presl
Tuber sword fern, erect sword fern
Semi-tender or slightly hardier, Zone 9 Figure 13.80.5. Nephrolepis cordifolia: a. frond, bar =
A medium-sized fern with erect, stoloniferous rhizomes. 10 cm (4 in.); b. middle part of frond, bar = 2.5 cm
Grows well under medium to high light in moist to dry (1 in.); c. tubers on stolons, bar = 2.5 cm (1 in.);
garden soil or potting mix. This species is easy to grow and d. pinna, bar = 1 cm (0.4 in.).
392 C 13

e
d

Figure 13.80.6. Nephrolepis cordifolia, fronds of cultivars: a. ‘Duffii’, pinnae bar = 1 cm (0.4 in.); b. ‘Kimberly Queen’;
c. ‘Lemon Buttons’, pinnae bar = 1 cm (0.4 in.); d. ‘Plumosa’; e. ‘Pom-pom’. All fronds, bar = 10 cm (4 in.).

America, Africa, Asia, Australia, Japan, and New Zealand ‘Lemon Buttons’. Figure 13.80.6c. Fronds to about 25
(where it reportedly lacks tubers). cm (10 in.) long, to 2.5 cm (1 in.) wide; pinnae reduced,
‘Duffii’. Figure 13.80.6a. Fronds less than 30 cm (12 oblong to roundish, not symmetrical at the base, not dou-
in.), erect, forked; pinnae small, to about 1 cm (0.5 in.) bled as in ‘Duffii’.
long, rounded or fan-shaped, often in pairs. This plant does ‘Petticoats’. Fronds to about 70 cm (28 in.) long, ends
not produced sori, and although considered a cultivar of forked several times in different planes; pinnae mostly ob-
Nephrolepis cordifolia, its parentage is doubtful, in part be- long-rectangular, sometimes shallowly and bluntly forked
cause it does not produce tubers and is more tender than N. at their tips. Forms with long fronds have been called ‘Den-
cordifolia. nis Petticoats’; those with short fronds, ‘Pom-pom’ (Fig-
‘Ecuadorian Fern’. Pinnae ruffled, undulate to irregu- ure 13.80.6e); and a shorter, more erect form, ‘Can-can’.
larly lobed, finely toothed. A recent introduction from cul- The weight of the terminal tassel, especially on long fronds,
tivation in Ecuador. causes the blades to bend and become pendent.
‘Kimberly Queen’. Figure 13.80.6b. Fronds to 1 m (3 ‘Tesselata’. Fronds to about 75 cm (30 in.) long; pin-
ft.) or more long and over 12.5 cm (5 in.) wide, sometimes nae one-pinnate in the distal half, except for lower pinnae;
slightly narrowly obovate; more arching and softer in ap- pinnules oblong-ovate. Variations in this cultivar include
pearance than the species. This cultivar grows vigorously. It pinna and pinnule length, the degree to which the pinnae
is often misidentified in the trade as Nephrolepis obliterata are bent backward, and the degree of emargination.
(R. Brown) Carruthers, an Australian fern with narrow, Whether these variations are due to cultural conditions or
distantly spaced pinnae, not known in cultivation. are stable is not certain. Related (if not the same) plants are
 F  F A  C 393

circulating as ‘Plumosa’ (Figure 13.80.6d) and ‘Mildred round to round–kidney-shaped with a U-shaped sinus,
Murray’. mostly opening toward the margin. This species is native to
Florida, the West Indies, and the Pacific Islands.
Nephrolepis exaltata (Linnaeus) Schott Young plants of Nephrolepis exaltata are difficult to dis-
 13.80.2, 7, 8;  5 tinguish from those of N. cordifolia (which are often mis-
Common sword fern, Boston fern takenly sold as N. exaltata). In N. exaltata the scales on the
Tender to semi-tender upper surfaces of the rachises are uniform in color or, if
A medium-large fern with short, erect, stoloniferous rhi- darker in the center, then also with irregular darker spots on
zomes. Grows best under medium to high light in moist- the rays of the scales. On young plants of N. cordifolia, these
dry potting mix. The wild form is stiff, upright, and has a scales are usually whitish with a dark center sharply marked.
rank appearance. Although the wild form is rarely culti- The fronds are also narrower than those of N. exaltata. Ster-
vated, the cultivars derived from it are among the most ile, adult plants of N. cordifolia are distinguished by their
loved and widely grown ferns. Natural hybrids between erect, stiff, narrower fronds, crowded pinnae, and tuber-
Nephrolepis exaltata and other species may be cultivated, producing runners. For commercial purposes N. cordifolia
especially a hybrid with N. multiflora in Hawaii. More in- is grown mainly from spores, whereas the sterile Boston
formation on cultivating Boston ferns is provided in Chap- ferns are cloned mainly by tissue culture.
ter 10. Nephrolepis exaltata has given rise to numerous culti-
The stipe bases of Nephrolepis exaltata have spreading, vars, and these generally have the same cultural require-
completely brown (concolorous) scales. The pinnae are ments as the wild form. So commonly grown are these ferns
usually less than 5 cm (2 in.) long, with the surfaces essen- that commercial growers have established the precise cul-
tially glabrous or with a few hair-like scales on the upper tural conditions for optimum growth: 2000 foot-candles
surface. The rachises are sparsely to densely covered with of light, temperatures between 18 and 35°C (65–95°F),
hair-like scales on the upper surfaces, and the indusia are fertilizers with a 3-1-2 composition, micronutrients, and
dolomite to adjust the pH to 5.5. Use a mix with good
drainage, and water when the soil surface is slightly dry. To
prevent breakage and decay of fronds, water should be
applied at the soil level, not on the foliage. Poor foliage re-
sults when temperatures are below 18°C (65°F) for more
than a month. Repotting just before the active growth in
spring is recommended.
Those ferns with more finely cut or congested fronds
might grow more slowly and need more care. You can ex-
pect cultivars with dense, compact foliage to show aborted
pinnae where light does not penetrate into the center of the
cluster. This does not harm the plant as long as the outer fo-
liage is healthy. All declining fronds or pinnae, however,
should be removed to promote air circulation and light
a penetration to underlying foliage, discouraging decay. The
cultivars with long fronds are seldom found in the trade be-
cause these large ferns are not well accommodated to
smaller homes, and nursery growers find it difficult to trans-
port long fronds without damage.
b English growers report that buds form on the foliage of
their cultivars, but in the United States this has been doc-
umented only for ‘Trevillian’. The buds are reported to
produce plants that revert to the ordinary Boston fern,
‘Bostoniensis’.
c The names of Boston fern cultivars are greatly confused.
Both old and new names have been loosely used and are
poorly documented, except for the work of Sessions (1978).
The delineation of cultivars is further complicated by their
tendency to intergrade. Sometimes a given cultivar will pro-
Figure 13.80.7. Nephrolepis exaltata: a. frond, bar = 10 duce fronds with different characteristics on the same plant.
cm (4 in.); b. middle part of frond, bar = 2.5 cm (1 in.); Cultivar variations include size and shape of divisions,
c. pinna, bar = 1 cm (0.4 in.). number of times pinnate, margin patterns, and degree of
 g

b c
a

h i

d
e f

m n

Figure 13.80.8. Nephrolepis exaltata cultivars: a. ‘Bostoniensis’, middle part of frond; b. ‘Dallas’; c. ‘Elsevier’ (frond
enlarged at right); d. ‘Fluffy Ruffles’; e. ‘Gretnae’, middle part of frond; f. ‘Massii’; g. ‘Mini-ruffles’ (frond enlarged at
right); h. ‘Norwoodii’, showing pinna; i. ‘Rooseveltii’, upper half of frond; j. ‘Sassy’; k. ‘Smithii’; l. ‘Splendida’; m.
‘Verona’, showing pinna; n. ‘Wanamaka’. Bar = 10 cm (4 in.).
 F  F A  C 395

ruffling, twisting, forking, or overlapping. Some of these times forked and heavily crested. The pinna apex is acute or
variants tolerate cold better, tend to revert less, and retain to three times forked. A large spectacular fern when well
their pinnae longer. grown.
Only a small number of the hundreds of cultivars of ‘Verona’. Figure 13.80.8m. Fronds three-pinnate, to
Nephrolepis exaltata are mentioned here. They are selected about 40 cm (16 in.) long and half as wide, spreading to
to show the great diversity within the group. pendent, the segments ovate. All divisions are distant from
‘Bostoniensis’ (var. bostoniensis Desvaux). Boston fern. each other, hence the blade appears open and imparts an
Figure 13.80.8a. Fronds one-pinnate, up to about 120 cm airy appearance to the plant.
(4 ft.) long, narrow, arching; pinnae slightly wavy; sori (if ‘Wanamaka’. Figure 13.80.8n. Fronds one-pinnate,
present) with aborted sporangia. ‘Bostoniensis’ is the parent dwarf, less than 30 cm (12 in.) long, narrow, stiff, upright,
of the many Nephrolepis exaltata cultivars. It is more com- leathery, dark green; pinnae closely overlapping, curled and
pact and graceful than the wild plant, with the pinnae twisted, crenulate.
broader and not as flat as those of the wild form. It was first
noticed in a Boston nursery in 1821, hence the name Bos- Nephrolepis falcata (Cavanilles) C. Christensen
ton fern.  13.80.9
‘Dallas’. Dallas fern. Figure 13.80.8b. A compact plant Broad sword fern, macho fern
with short fronds bearing oblong pinnae that are notice- Tender
ably serrate with round apices.
A large fern with erect, stoloniferous rhizomes and erect or
‘Elsevier’. Figure 13.80.8c. Fronds two-pinnate, minia-
pendulous, long fronds. Grows well under medium to high
ture, less than 30 cm (12 in.) long. Resembles ‘Mini-ruffles’
light in moist-dry potting mix or uncut moss.
but with narrowly elliptic, slightly twisted fronds.
The older stipe bases on Nephrolepis falcata have scales
‘Fluffy Ruffles’. Figure 13.80.8d. Fronds two-pinnate,
that are spreading to appressed, lanceolate or triangular,
dwarf, less than 30 cm (12 in.) long, elliptic, stiff, upright;
and nearly black with a pale margin. The pinnae are up to
stipes thick; pinnae and pinnules overlapping, with entire
about 10 cm (4 in.) long and often curved toward the apex
to crenate margins. Tends to stay greener in winter.
(falcate). The fertile pinnae are slightly narrowed compared
‘Gretnae’. Figure 13.80.8e. Like ‘Bostoniensis’ but with
to the sterile ones. A few scattered scales are found on both
stouter pinnae forked at the apex into blunt lobes, the frond
surfaces of the blades, but these tend to fall off with age.
apex often several times forked.
The upper surface of the pinna midrib is glabrous or with a
‘Golden Boston’ (‘Aurea’). Like ‘Bostoniensis’ but with
few scales. The sori are near the margin, and the indusia
yellow-green fronds.
are round with a narrow sinus. The species is native to Ma-
‘Gracillima’ (‘Irish Lace’). Resembles ‘Smithii’, but the
laysia and extends northward into Burma and Indochina.
fronds are three-pinnate and more ovate, and the pinnae
Nephrolepis falcata is often misidentified as N. biserrata
arise uniformly, one behind the other.
and N. ensifolia C. Presl.
‘Massii’. Figure 13.80.8f. Short frond form with pinnae
‘Furcans’. Fishtail sword fern. Figure 13.80.9b. Fronds
slightly twisted at apex.
shorter and less pendent than typical for the species; pinna
‘Mini-ruffles’. Figure 13.80.8g. Fronds miniature, to
tips once- or twice-forked. In the tropics this plant is com-
about 10 cm (4 in.) long, three-pinnate, broadly triangular,
monly grown in pots in full sun, although it produces bet-
and erect, with overlapping pinnae.
ter color in the shade.
‘Norwoodii’. Figure 13.80.8h. Fronds three-pinnate,
‘Ram’s Horn’. Figure 13.80.9c. Pinnae twisted and
less than 40 cm (16 in.) long, ovate, spreading; pinnae and
curled backward.
pinnules more or less in regular layers, overlapping and
congested, pinnules divided into long narrow segments.
The dense foliage on the blade has a fluffy but layered ap- Nephrolepis hirsutula (G. Forster) C. Presl
pearance with delicate details. The two-pinnate form also  13.80.10
circulating as ‘Norwoodii’ is closer to ‘Fluffy Ruffles’. Rough sword fern, scurfy sword fern
‘Rooseveltii’. Figure 13.80.8i. Fronds to 1 m (3 ft.) long, Tender
pinnae wavy, usually eared on both sides of the base. A medium-large fern with short, erect, stoloniferous rhi-
‘Sassy’. Figure 13.80.8j. Fronds short; pinnae moder- zomes. Grows well under medium to high light in garden
ately twisted, ruffled, and irregularly finely dentate-serrate. soil or potting mix kept moist-dry.
‘Smithii’. Lace fern. Figure 13.80.8k. Fronds to four- The older stipe bases of Nephrolepis hirsutula bear ap-
pinnate, dwarf, about 30 cm (12 in.) long, widely ovate, pressed, convex scales with nearly black centers and pale
arching to pendent, lacy; segments ovate, small, 2 mm or margins. The pinnae are about 9 cm (3.5 in.) or more long,
less long, spaced apart. closely placed, and usually sharply eared. They bear fila-
‘Splendida’. Figure 13.80.8l. Fronds one- to two-pin- mentous scales on the lower surface. The upper surface of
nate, pendent, to 1 m (3 ft.) long, with the distal half many the pinna midrib has long hairy scales or is glabrous. The
 396 C 13

Figure 13.80.9. Nephrolepis

falcata: a. frond, bar = 10 cm
(4 in.); b. ‘Furcans’ frond, bar
= 10 cm (4 in.); c. ‘Ram’s
Horn’ frond, bar = 10 cm
(4 in.); d. pinna, bar = 2 cm
(0.8 in.).

a
b c
c

d Figure 13.80.10. Nephrolepis hirsutula: a. frond, bar =

10 cm (4 in.); b. ‘Superba’ frond, bar = 10 cm (4 in.);
c. pinna, bar = 2 cm (0.8 in.).
rachis appears scurfy-scaly. Round indusia have a narrow
sinus and are arranged close to the margins. This species is
native to Australia and tropical Asia to the islands of the Nephrolepis lauterbachii somewhat resembles N. cordi-
Pacific; it has become naturalized in the American tropics. folia but has smaller, softer, thinner-textured fronds. The
The scurfy-scaly rachis is the source of the common rhizomes do not produce tubers, and the stipes are about 1
name scurfy sword fern. The species greatly resembles mm thick and bear tan-brown scales. The pinnae are sepa-
Nephrolepis multiflora, a species that bears a stubble of rate (not overlapping), less than 1.5 cm (0.6 in.) long,
short, erect hairs on the upper surface of the pinna midrib, somewhat oblong-rectangular, and glabrous. The pinna
whereas N. hirsutula has the hairs sparse or absent. bases are asymmetrical, being truncate or cordate on the
‘Superba’. Figure 13.80.10b. Pinnae irregularly lacini- acroscopic side and narrowly tapered on the basiscopic side.
ate-lobed, ruffled. The plants are sterile. This cultivar is The rachis is glabrous or has a few concolorous scales. The
often confused with Nephrolepis multiflora ‘Florist Fantasy’. indusia are round-reniform to lunate with an extremely
broad sinus (or none), and they face the pinna apices. Up to
Nephrolepis lauterbachii H. Christ  13.80.11 five sori occur on the upper margin opposite or nearly op-
Tender posite each major crenation. Nephrolepis lauterbachii is en-
A small-medium fern with short, erect, stoloniferous rhi- demic to New Guinea.
zomes. Grows well under medium to high light in moist-
dry garden soil or potting mix.
 F  F A  C 397

Figure 13.80.11. Nephrolepis b

lauterbachii: a. frond, bar =
10 cm (4 in.); b. pinna, bar
= 2 cm (0.8 in.).

Figure 13.80.12. Nephrolepis multiflora: a. frond, bar =

Nephrolepis multiflora (Roxburgh) F. M. Jarrett ex 10 cm (4 in.); b. ‘Florist Fantasy’ frond, bar = 10 cm
C. V. Morton  13.80.12 (4 in.); c. hairs on upper surface of pinna base, bar =
Asian sword fern 1 cm (0.4 in.); d. pinna, bar = 1 cm (0.4 in.).
Tender
A medium to large fern with short, erect, stoloniferous rhi-
zomes. Grows well under medium to high light in moist-
dry garden soil or potting mix. The plants have a rank ap- what twisted, ruffled, serrate, and attenuate. This cultivar
pearance and tend to be weedy. This species is widely has long been confused with Nephrolepis hirsutula ‘Su-
cultivated outdoors in Florida and Hawaii. perba’, which has laciniate-lobed, not ruffled, margins. An
Nephrolepis multiflora has pinnae about 10 cm (4 in.) unnamed dull dark green cultivar resembling ‘Florist Fan-
long. The scales on the stipe bases are appressed, black in tasy’ but with stiffly erect fronds has recently been circulat-
the center with pale tan borders, and minutely fibrillose ing in the trade.
along the margins (only N. hirsutula shares this character-
istic). The pinnae may be eared or not, and their surfaces Nephrolepis pendula (Raddi) J. Smith
are covered by hair-like scales or ciliate-margined scales.  13.80.13
The upper surface of the pinna midrib has a dense stubble Tender to semi-tender (?)
of short, erect hairs 0.2–0.3 mm long. The sori are close to A large fern with short, erect, stoloniferous rhizomes.
the margin, and the indusium is round with a narrow sinus. Grows well under medium light in moist potting mix. This
This species is native to the tropical regions of Africa and species grows slowly and is less robust than Nephrolepis
Asia and has become naturalized (and often weedy) in the cordifolia, which it resembles. It is a choice fern for hanging
American tropics. baskets because of its long, narrow, pendent fronds.
‘Florist Fantasy’. Figure 13.80.12b. Fronds shorter and Nephrolepis pendula differs from N. cordifolia by its flat,
more erect than the wild form; pinnae close together, some- longer, pendent fronds and shorter pinnae. The pinnae are
 398 C 13

Niphidium contains 10 species and is native to the New

World tropics. The genus name comes from the Greek
nipha, snow, and eidos, like, referring to the whitish indu-
ment on the leaves of the type species. For a monographic
a treatment of the genus, see Lellinger (1972). Also see Poly-
podium for a comparison of Niphidium to other polypodi-
aceous genera.

Niphidium crassifolium (Linnaeus) Lellinger

 13.81.1
Semi-hardy to hardier
A medium-large fern with short- to long-creeping rhizomes
and clustered fronds. Does well under medium light in
drained, moist-dry garden soil or potting mix. This species
grows easily in the ground, pots, and hanging baskets. It
b can tolerate several consecutive days below freezing to
−7°C (19°F).
The rhizome scales of Niphidium crassifolium are
clathrate centrally but nonclathrate toward the margins.
The fronds vary from narrowly oblanceolate to lanceolate,
and from acute to rounded or sometimes mucronate at the
tips. The upper surfaces of the blades, especially on older
fronds, usually bear white dots that are accumulations of
Figure 13.80.13. Nephrolepis pendula: a. frond, bar = calcium carbonate secreted by the hydathodes beneath.
10 cm (4 in.); b. pinna, bar = 1 cm (0.4 in.). This species is widely distributed in tropical America.

a
about twice as long as broad and have rounded apices and
mostly entire margins. The upper part of stipe and rachis
(except at the point of pinna attachment) essentially lacks
hairs or scales. The species is native to tropical America.

Niphidium
syn. Pessopteris, Polypodium in part
These epiphytic, epilithic, or terrestrial plants have me-
dium- to large-sized, stiff, undivided, upright fronds. They
are reminiscent of the bird’s-nest fern (Asplenium nidus),
but instead of the latter’s neat rosette of light green fronds,
Niphidium fronds are dark green and irregularly clustered. b
The two genera can be distinguished by the sori, which are
linear and indusiate in Asplenium but round and non-
indusiate in Niphidium. Niphidium also resembles the trop-
ical American genus Campyloneurum, but that genus has
two or more rows of sori between the main lateral veins, c
whereas Niphidium only has one. d
The rhizomes of Niphidium vary from short- to long-
creeping. The fronds are simple, entire, elliptic-lanceolate
to oblong, and leathery, often with white dots on the upper
surfaces. The main lateral veins are prominent and oblique Figure 13.81.1. Niphidium: a. N. crassifolium frond, bar
to the rachis. In the cultivated species the sori are large, = 5 cm (2 in.); b. N. crassifolium sori, bar = 2.5 cm
about 3 mm (0.1 in.) wide, usually round, and produced in (1 in.), after Hoshizaki (1982); c. N. crassifolium
a single row between the lateral veins. Each sorus is served rhizome scale, bar = 1 mm, after Lellinger (1972); d. N.
by a small vein ring. As in all polypodiaceous ferns, indusia albopunctatissimum rhizome scale, bar = 1 mm, after
are lacking. Lellinger (1972).
 F  F A  C 399

A few cultivated plants examined under the name

Niphidium crassifolium turned out to be N. albopunctatissi-
mum (Figure 13.81.1d), and these two species are difficult
to distinguish. They have distinct rhizome scale character-
istics—those of N. albopunctatissimum are irregular (erose)
in the nonclathrate part of the margin toward the tip—but
this trait is not always easy to use for identification, and
when the scales are absent the species cannot be distin-
guished. In cultivated and wild plants, both species can
have white dots on the upper surface of the frond (the name
albopunctatissimum means “with many white dots”), so this
is not a reliable characteristic to separate the two, either.
The fronds of N. albopunctatissimum are slightly smaller
and narrower than those of N. crassifolium, and in nature
N. albopunctatissimum is mostly found on the ground or
on rocks from Colombia to Bolivia, whereas N. crassifolium
is occurs mostly throughout tropical America.

Notholaena
Cloak fern
A mostly rock-inhabiting fern of dry climates, Notholaena
is difficult to grow and is found only in special collections.
These small ferns have short-creeping, compact rhizomes.
The genus resembles certain species of Cheilanthes and Pel-
laea but differs by the dense covering of white, cream, or Figure 13.82.1. Notholaena standleyi: fronds, the
yellow waxy powder (farina) on the undersides of the underside (left) with dense powder, and the upperside
fronds and by the marginal, hardly enrolled, narrow indu- (right), bar = 2.5 cm (1 in.).
sia. The stipes are frequently longer than the blades, and
the fronds may be linear-lanceolate to pentagonal and pin-
nate-pinnatifid to four-pinnate. Many species formerly
classified in Notholaena have been transferred to Cheilanthes Odontosoria
(which see for a comparison). syn. Sphenomeris
Notholaena consists of about 25 species native to North
The cultivated species of Odontosoria are finely cut terres-
America, Central and South America, and the West Indies.
trial plants of rocky areas. They are used in tropical gardens
The genus name comes from the Greek nothos, false, and
and are rarely seen elsewhere. Like many ferns that grow
chlaena, cloak, referring to the blade margins, which are
between rocks, these plants are difficult to remove without
not reflexed as in the similar genus Cheilanthes. For addi-
damaging the roots. Transplants with damaged roots often
tional information on growing cloak ferns, see the section
do not reroot readily, hence the genus is best propagated by
on “Xerophytic Ferns” in Chapter 10.
spores.
The short-creeping to suberect rhizomes are covered
Notholaena standleyi Maxon  13.82.1 with narrow or hair-like dark scales. The fronds are set close
Hardy together and have two- to four-pinnate blades. The ulti-
A small fern with compact rhizomes and fronds in a cluster. mate segments are linear or wedge-shaped and dichoto-
Requires high light in moist-dry, well-drained garden soil mously lobed, with a single vein or the vein once or twice
preferably mixed with coarse sand or gravel. forked. Marginal sori are borne at the vein tips or, more
Notholaena standleyi has pentagonal blades densely cov- rarely, are supplied by two or three veins. The rounded to
ered with a whitish powder on the lower surface, and the oval indusium is attached by the base and more or less by
hardly enrolled indusium is narrow. It might be confused the sides. It opens toward the margin of the segments.
with the more frequently grown Cheilanthes argentea, The genus, which consists of 22 species, is native to the
which is less divided, less densely powdery, and has a more tropics and subtropics of America, Africa, and Japan. Its
enrolled, broader indusium. Notholaena standleyi is native name comes from the Greek odous, tooth, and soros, mound,
to the southwestern United States and Mexico. and refers to the sori borne at the extreme tips of small,
marginal teeth.
 400 C 13

Odontosoria chinensis (Linnaeus) J. Smith Oleandra

 13.83.1 These primarily terrestrial ferns often require ample space
syn. Sphenomeris chinensis (Linnaeus) Maxon, S. chusana for their wide-creeping or scrambling habit. Oleandra pis-
(Linnaeus) Copeland tillaris grows erect and shrub-like but is unusual in appear-
Lace fern ance because its leaves are borne in whorls at intervals along
Tender the stems.
A medium-sized fern with short-creeping to suberect rhi- The genus is characterized by long-creeping rhizomes,
zomes. Grows well under medium to high light in well- jointed stipes, simple and entire blades, free veins, round
drained, moist garden soil or potting mix. The species is sori scattered close to the midrib, and kidney-shaped to
commonly grown in Hawaii but only rarely in the conti- horseshoe-shaped indusia. The joint in the stipe can usually
nental United States. be seen as a faint dark line or a slight swelling. The fronds
The blades of Odontosoria chinensis vary from lanceo- break off cleanly from these joints when they are shed.
late to ovate and are up to four-pinnate. The pinna midrib Oleandra occurs throughout the tropical regions of the
is straight, not zigzag (as in many other species in the world and contains about 40 species. The genus name
genus), and the ultimate segments are wedge-shaped or comes from oleander (Nerium), a flowering plant in the
lobed. The sori are often two or more per segment. The dogbane family (Apocynaceae), the leaves of which resem-
species is native from Madagascar to Polynesia and Japan. ble those of several species in Oleandra.

Oleandra articulata (Swartz) C. Presl  13.84.1

syn. Oleandra nodosa (Willdenow) C. Presl
Figure 13.83.1. Tender
Odontosoria chinensis: a.
habit, bar = 5 cm (2 in.); A medium to large fern with long-creeping, scrambling rhi-
b. sori and indusia, bar = zomes. Grows well under high to medium light in moist
5 mm (0.2 in.). garden soil or potting mix. This species requires ample
room in the garden to spread; ideally, plant it in baskets to
contain it.
Oleandra articulata is characterized by widely spreading
rhizome scales, blades glabrous or nearly so on both sur-

c
a

Figure 13.84.1. Oleandra articulata: a. habit, bar = 5 cm

(2 in.); b. rhizome scale, bar = 2 mm; c. joint on stipe,
bar = 1 cm (0.4 in.).
 F  F A  C 401

faces, and glabrous indusia. It is native to the American

tropics.

Oleandra pilosa Hooker  13.84.2

Tender
A medium-sized fern with climbing, ascending to erect rhi-
zomes. Grows well under high to medium light in moist
garden soil or potting mix. This plant’s growth habit is too
upright for planting in baskets; it is probably best suited
for growing against walls and other vertical surfaces.
Oleandra pilosa is characterized by tightly appressed,
hairy-margined rhizome scales, blades hairy on both sur-
faces, and ciliate indusia. It is native to the American trop-
ics.
Oleandra pilosa circulates among some gardeners as O.
neriiformis Cavanilles, a species unknown in recent culti-
vation and native to Australia, tropical Asia, the Philip-
pines, and Polynesia. a
Oleandra pistillaris (Swartz) C. Christensen
 13.84.3
Tender
A medium to large fern with long-creeping rhizomes and a
shrubby or straggly appearance. Grows well under medium b
light in moist garden soil or potting mix. The species as-
sumes a shrub-like appearance in tropical gardens. It can
be suitably grown in large pots or tubs. Figure 13.84.3. Oleandra pistillaris: a. habit, bar = 5 cm
(2 in.); b. sori and indusia, bar = 5 mm (0.2 in.).

Oleandra pistillaris bears its fronds in distinct whorls

Figure 13.84.2. Oleandra pilosa:
along the stem, with the stem tip extending far beyond the
a. habit, bar = 5 cm (2 in.);
last whorl. The sori are disposed in a row on both sides of
a b. rhizome scale, bar = 2 mm.
the midrib. This species is native from Malaysia and Java to
Polynesia(?).
This species might be confused with Oleandra neri-
iformis, which has creeping rhizomes that climb and pro-
duce aerial roots. The rhizomes of O. pistillaris, however,
are erect, woody, and shrub-like.

Olfersia
Rarely cultivated, Olfersia grows in soil in the wild or
scrambles over fallen logs or rocks. It occasionally climbs
tree trunks, in which instances the rhizomes become long-
creeping. Younger plants tend to maintain a short rhizome
for some time and therefore can be confined to a wide pot.
Olfersia is medium-sized and can be distinguished from
all other ferns by its creeping, densely scaly rhizomes and
b strongly dimorphic sterile and fertile fronds. The sterile
fronds are one-pinnate with a terminal segment resembling
the lateral pinnae. The veins are long-parallel and free for
most of their length, joining just before the margin to form
a submarginal connecting strand. The fertile fronds are bi-
 402 C 13

pinnate with linear pinnae and sori that merge. The indu- Onoclea can be identified by its netted veins, triangular
sium is absent. and deeply pinnatifid or (more often) one-pinnate-pinnat-
This genus consists of one species widespread in the ifid sterile blades, and strongly dimorphic sterile and fer-
American tropics. Its name honors Ignaz Franz Werner von tile fronds. Other characteristics are medium-wide-creep-
Olfers (1793–1871), a professor who collected in Brazil. ing rhizomes and fertile pinnae that are strongly ascending
See Moran (1986) for a technical treatment of the species. and contracted into rounded, bead-like lobes enclosing the
sori.
Olfersia cervina (Linnaeus) Kunze  13.85.1 The genus, which consists of only one species, is native
syn. Polybotrya cervina (Linnaeus) Kaulfuss to North America and Asia. The genus name comes from
Tender the Greek onos, vessel, and kleiein, to close, referring to the
A medium-sized fern with medium- to long-creeping rhi- pinnules of the fertile leaf, which roll up into bead-like seg-
zomes. Grows well under medium light in moist garden ments to enclose the sori.
soil. Olfersia cervina is easy to grow from spores. See the de-
scription of the genus for distribution and additional char- Onoclea sensibilis Linnaeus  13.86.1–3
acteristics. Very hardy, Zone 2
A medium-sized fern with medium- to long-creeping rhi-
zomes and deciduous fronds. Grows well under medium
to high light in moist-wet garden soil. This species is easy to
grow.
Onoclea sensibilis is characterized by pinnatisect fronds
with lobed pinnae narrowed toward the base, and veins
netted throughout. The margins of the blade are finely
serrulate. The fertile fronds are brown, erect, and rigid
with strongly ascending pinnae. The species is native to
eastern North America and eastern Asia, often growing in
wet habitats.
Onoclea sensibilis has been around for a long time. Fos-
sils identical to modern plants have been found in Paleo-
cene rocks in Canada—rocks that date back more than 54
million years (Rothwell and Stockey 1991).

Onychium
Claw fern
Onychium is a genus of terrestrial, clump-forming ferns that
can be planted in the ground or in pots. Their finely di-
vided fronds have a delicate, lacy appearance.
The rhizomes are short- to long-creeping and scaly. The
Figure 13.85.1. Olfersia cervina: sterile frond (left) and fronds are dimorphic or monomorphic. The sterile blades
fertile frond (right), bar = 5 cm (2 in.).

Onoclea
Sensitive fern, bead fern
The bead-like appearance of the fertile fronds accounts for
this genus’s common name of bead fern. Some say that the
name sensitive fern originates from the fronds’ sensitivity to
frost (they wither after the first subfreezing temperatures).
These medium- to large-sized natives of temperate areas
have coarsely cut but attractive foliage. The plants spread by
creeping, branching rhizomes. Onoclea is useful in wet,
temperate areas but will grow well in warmer climates even
though it is deciduous. The fertile fronds are often used in
dried flower arrangements. Figure 13.86.1. Onoclea sensibilis: habit.
 F  F A  C 403

vary from three- to five-pinnate and are anadromous and

glabrous. The veins are free, not netted. Linear sori run
along the margin. The indusium is formed by the enrolled
leaf margin, and because the segments are extremely nar-
row, the indusium on one side of the segment meets the
edge of the indusium on the opposite side.
Onychium contains about eight species native to the
tropics and subtropics of Africa, the Near East, India, Asia,
and New Guinea. The genus name comes from the Greek
onychion, a small claw or nail, and refers to the shape of the
ultimate segments of the blade.

Onychium japonicum (Thunberg) Kunze

 13.87.1–3
Japanese claw fern, carrot fern
Semi-hardy, Zone 8
A small-medium fern with short-creeping, branched rhi-
zomes. Grows well under high light in moist garden soil or
potting mix. The plants are easy to cultivate and have at-
tractive carrot-like foliage.

Figure 13.86.2. Onoclea sensibilis: fertile pinnae.

Figure 13.87.1. Onychium japonicum: habit.

Figure 13.86.3. Onoclea sensibilis: sterile frond (left) and

fertile frond (right), bar = 5 cm (2 in.). Figure 13.87.2. Onychium japonicum: sori and indusia.
 404 C 13

Figure 13.87.3. Onychium japonicum: frond, bar = 5 cm

(2 in.).
Figure 13.88.1. Ophioglossum petiolatum: habit.

The stipes of Onychium japonicum are straw-colored to-

ward the base, and the blades are three- to four-pinnate lected from the wild. This species is difficult to grow and
with segments narrowly lanceolate and acute. The sori are rarely survives transplanting. The hand fern can be recog-
about 5 mm (0.2 in.) long. This species is native to India, nized by its fan-shaped sterile blade, which grows to 45 cm
the Himalayas, eastern Asia, the Philippines, and Java. (18 in.) long and is deeply palmately lobed. The species is
epiphytic and often grows in the old humus-filled leaf axils
Ophioglossum  13.88.1, 2 of palms.
syn. Cheiroglossa Nearly all the species of Ophioglossum are terrestrial;
Adder’s-tongue fern, serpent’s-tongue fern only a few are epiphytic. Most have short, erect, fleshy rhi-
Ophioglossum is a genus of mostly small ferns that are grown zomes and whitish or tan, fleshy roots. The fronds are di-
in natural gardens or in pots as novelties. They are hard to vided in two parts: a green, photosynthetic sterile blade and
find in the wild, and some species are rare. The temperate an erect, elongate stalk (the fertile spike). The fleshy blades
ones are often overlooked because they grow in open grassy lack a midrib, and they vary from simple and entire to
areas and resemble other plants, especially plantains (Plan- strap-shaped and branched to fan-shaped and palmately
tago). One sure way to distinguish Ophioglossum from other lobed. The veins are netted, and the areoles may or may
plants is by the lack of a midrib on its blades—all other not contain veinlets. At the base of the sterile blade arises a
plants have a midrib or at least prominent parallel veins. long-stalked spike or (in the segregate-genus Cheiroglossa)
One of the most eye-catching tropical species is Ophio- several short spikes or a stalk branched into spikes at the
glossum palmatum (sometimes called Cheiroglossa palmata top. Sporangia are borne at the apices of the fertile spikes.
(Linnaeus) C. Presl), commonly known as the hand fern. It The sporangia are sunken on both sides of the spike and
is an endangered species in Florida and should not be col- open by a horizontal slit.
 F  F A  C 405

a c

Figure 13.88.3. Ophioglossum pendulum: a. frond with

two spikes, bar = 5 cm (2 in.); b. habit of mature plant,
bar = 15 cm (6 in.), courtesy of T. Hoshizaki; c. frond
with flared apex, bar = 5 cm (2 in.).

Figure 13.88.2. Ophioglossum petiolatum: fertile

spike. Photo courtesy of G. Hampfler.
Ophioglossum petiolatum Hooker
 13.88.1, 2, 4
Semi-hardy, Zone 7
Ophioglossum occurs worldwide and consists of about 30 A small fern with erect rhizomes. Grows well under high
species. The common names are derived from the fancied light in moist garden soil. Fronds emerge during wet peri-
resemblance of the erect fertile stalk to a snake’s tongue. ods. The plants reproduce vegetatively from buds on the
The genus name also alludes to this resemblance, being de- roots and may become weedy or volunteer in pots of other
rived from the Greek ophis, snake, and glossa, tongue. plants. The succulent leaves are readily attacked by milli-
pedes, slugs, and snails.
Ophioglossum pendulum Linnaeus
 13.88.3;  36
Tender, Zone 10
A medium to large fern with short-creeping or suberect rhi- Figure 13.88.4.
zomes and pendulous fronds. Grows under low to medium Ophioglossum
light in moist uncut sphagnum moss. The plants are typi- petiolatum: habit,
cally grown on a suspended board or basket so that the bar = 5 cm (2 in.).
fronds can hang downward as they do in nature (the plants
are epiphytic). When grown in pots, however, the fronds
tend to be more upright. A variant with two fertile spikes
per blade is in cultivation.
Ophioglossum pendulum has long, pendent, strap-shaped
sterile blades, up to 2 m (6.5 ft.) long, and occasionally sev-
eral times branched. The veins form large areoles that lack
included veinlets (the other species have included veinlets).
The species is native to the Old World tropics.
 406 C 13

Ophioglossum petiolatum produces two or three fronds

per stem, instead of one as in other species. The fronds ap-
pear in one or more flushes of growth per year. The sterile
blades have acute apices and the veins are netted, forming
large areoles that contain free veins. The species is native
to the southeastern United States, tropical America, Asia,
and the Pacific Islands. In the Gulf States (Florida to Lou-
isiana), the species is often abundant in sandy roadsides.

Ophioglossum vulgatum Linnaeus  13.88.5

Hardy to semi-hardy, Zone 6
A small fern with erect rhizomes. Grows under high light in
moist garden soil. The plants reproduce by root buds and
go dormant in hot weather.
The veins of Ophioglossum vulgatum resemble those of
O. petiolatum, but the sterile blades have a rounded apex
and the fronds are usually one per stem and appear once a
year, not in several flushes. The species is native to the
southeastern United States, Mexico, and temperate Eurasia.

Figure 13.89.1. Osmunda regalis var. brasiliensis: habit.

Figure 13.88.5. Ophioglossum

vulgatum: habit, bar = 5 cm
(2 in.).

Osmunda  13.89.1–3

Osmundas are familiar United States natives that produce
fresh green growth in the spring and early summer. The
fronds of these stately ferns form medium- to large-sized
clusters or whorls. In gardens they are used as background
foliage or accent plants. Although most osmundas grow in
wet, acidic soil, the more common species can be cultivated
in moist, neutral or very slightly acidic soil, although they
do not become as massive as the wild plants. The royal fern
(Osmunda regalis) attracts attention when contracted fer-
tile areas develop at the frond tips and contrast with the fo-
liaceous part of the frond. Because the thick, compact root
mantle of the cinnamon fern (O. cinnamomea) does not
decay rapidly, that species has been used as a substrate to Figure 13.89.2. Osmunda regalis var. brasiliensis: fertile
grow orchids and other epiphytes. pinnae.
 F  F A  C 407

Osmunda cinnamomea Linnaeus

 13.89.4;  8
Cinnamon fern
Very hardy, Zone (2)3
A medium to large fern with erect rhizomes that form oc-
casional offshoots. Grows well under low to high light in
moist-wet to wet, acidic garden soil. The plants are robust
growers with deciduous leaves. The fiddleheads are covered
with a dense mat of tawny or cinnamon-colored hair.
Osmunda cinnamomea is characterized by completely
dimorphic sterile and fertile fronds. The sterile blades are
one-pinnate-pinnatifid, oblong-lanceolate, and bear a
dense tuft of hair at the base of each pinna. The fertile fronds
are two-pinnate with strongly ascending pinnae bearing
globose sporangia. They wilt soon after the spores are shed.
The species is cosmopolitan.

a
b

Figure 13.89.3. Osmunda regalis var. brasiliensis:

sporangia.

The tropical species Osmunda vachellii Hooker, which

resembles a cycad, has recently been introduced to cultiva-
tion in southern California from the area around Hong
Kong. Although not treated here, it seems promising as an
ornamental. It has evergreen, one-pinnate fronds borne on
an erect stem and entire, narrowly lanceolate pinnae. The
fertile pinnae appear at or near the middle of the blade.
All species of Osmunda have erect rhizomes that can be-
come massive and occasionally branch. They bear fronds Figure 13.89.4. Osmunda cinnamomea: a. sterile frond
in a spiral tuft. The lower part of the stipe bears wing-like (left) and fertile frond (right), bar = 5 cm (2 in.);
outgrowths called stipules, although these are not equiva- b. pinnae base with tuft of hair, bar = 1 cm (0.4 in.).
lent to the stipules in marattiaceous ferns (Angiopteris, Ma-
rattia) or in angiosperms. The sterile blades vary from one-
to two-pinnate, and mature blades are usually smooth ex-
cept for a tuft of wool on the rachises and costae, usually at Osmunda claytoniana Linnaeus
the pinna base. The blades are always catadromous. The  13.89.5;  37
pinnae are often jointed to the rachis, but the joints are Interrupted fern
non-functional. The veins are free, not netted. All or some Very hardy, Zone (2)3
of the fertile fronds are strongly contracted, lack green leafy A medium to large fern with erect rhizomes that form oc-
tissue, and usually wither after the spores are shed. The spo- casional offshoots. Grows well under low to medium light
rangia are large and globose and not protected by an indu- in moist-wet to wet, acidic garden soil. The plants have de-
sium. The spores are green and short-lived. ciduous fronds and do poorly in the Gulf States and sub-
The genus consists of about 10 species and occurs tropical climates.
worldwide. Osmunda is named for Osmunder, the Saxon Osmunda claytoniana has one-pinnate-pinnatifid fronds
equivalent of the god Thor. with brown fertile pinnae borne in the middle of the blade
—they appear to interrupt the green, sterile pinnae. The
 408 C 13

Figure 13.89.5.
Osmunda claytoniana:
frond, middle pinnae
fertile, bar = 5 cm
(2 in.).

Figure 13.89.6. Osmunda japonica: sterile frond (left)

and fertile frond (right), bar = 5 cm (2 in.).

species is native to northeastern North America and Asia.

Osmunda ×ruggii R. M. Tryon is a naturally occurring
hybrid between this species and O. regalis. Although the Osmunda regalis Linnaeus
two parental species frequently grow together, the hybrid is  13.89.1–3, 7, 8;  6
extremely rare, known from only two localities, one in Royal fern, flowering fern
Connecticut and the other in West Virginia. The hybrid is Very hardy, Zone 2(3)
unavailable in the trade but would make an interesting ad- A medium to large fern with erect rhizomes that form oc-
dition to horticulture because it demonstrates hybrid in- casional offshoots. Grows well under medium light in
termediacy. It must be propagated vegetatively by division acidic, moist-wet to wet garden soil. The fronds are decid-
of the rhizome because the spores are aborted. uous or semi-deciduous, and the new growth is sometimes
reddish or purplish. When young, the sporangia appear
Osmunda japonica Thunberg  13.89.6 green from the color of the spores within. After the spores
Japanese osmunda are shed, the sporangia are rusty brown. Young leaves ap-
Hardy pear either purplish or green in spring—all the leaves on a
A medium-sized fern with suberect rhizomes and decidu- given plant will be of the same color. Leaves that are pur-
ous fronds. Grows well under medium light in moist gar- plish when young usually turn green with age.
den soil. The fiddleheads of this species are edible and are Osmunda regalis is characterized by two-pinnate fronds
available in markets in Japan. The cut rhizomes have been that, when fertile, bear sporangia only in the apical third
used as a substrate to grow epiphytes, much like tree-fern or so of the blade. The pinnules are oblong-lanceolate. This
trunks, in horticulture. species occurs worldwide and varies geographically. The
Osmunda japonica resembles O. regalis but has com- following geographic varieties are cultivated:
pletely dimorphic sterile and fertile fronds with finely ser- var. brasiliensis (Hooker & Greville) Pichi-Sermolli. Fig-
rate margins on the sterile pinnae. The fertile fronds are ures 13.89.1–3, 8. Fronds are slowly deciduous in temper-
erect and emanate from the center of the plant, whereas the ate areas and nearly evergreen in subtropical ones. In green-
sterile ones tend to spread, forming a wide, loose basket. houses, this variety produces new fronds throughout the
The species is native to eastern Asia. year. The rachis is glabrous. The fertile parts are conspicu-
ously stalked and distant from the sterile pinnules below.
Unlike var. regalis, the fertile pinnae are strongly ascend-
 F  F A  C 409

ing prior to withering and appear bunched together. The var. regalis. Figure 13.89.7. Fronds are completely de-
plants seem adapted to neutral or only slightly acidic soil. ciduous. The fronds are more leathery than those of the
The cultivated material of this fern was collected in Brazil. other varieties and have thicker stipes and rachises. The
Some botanists do not consider this variety distinct from pinnae and pinnules are sessile (or nearly so) and closer to-
var. spectabilis. gether than in var. spectabilis. The hairs on the rachis are
ample and darker than those of var. spectabilis. The fertile
pinnae spread at nearly right angles to the rachis and are
borne in the same plane as the sterile pinnae. This robust
grower is native to Europe and Asia. The following cultivars
are derived from it: ‘Cristata’, pinnules or segments crested
at the tips; ‘Purpurescens’, with purplish stipes, rachises,
and costae.
var. spectabilis (Willdenow) A. Gray. Fronds are com-
pletely deciduous. The pinnae and pinnules are stalked and
farther apart than in var. regalis. Also, the hairs on the
rachises are fewer and lighter colored than in var. regalis.
The fertile pinnae are slightly falcate and ascending, spread-
ing in the same plane as the sterile pinnae. This variety is
native to North America and might not be distinct from
var. brasiliensis.

Paesia
Plants of the genus Paesia typically have finely divided
fronds and zigzag rachises that allow them to scramble over
other plants. Paesia scaberula, however, the only cultivated
Figure 13.89.7. Osmunda
species, has only a slightly a zigzag rachis and less elongate
regalis: fertile frond, bar =
fronds.
5 cm (2 in.).
Paesia has slender, 3–5 mm (0.1–0.2 in.) long, creep-
ing, hairy rhizomes. Other characteristics of the genus are
two- to four-pinnate blades, free veins, and rachises and
costae that are strongly ridged and grooved on the upper
surfaces. The sori of Paesia have two indusia, an outer one
Figure 13.89.8. Osmunda formed from the modified, enrolled margin of the blade,
regalis var. brasiliensis: fertile and an inner one that is thin, colorless, and hidden by the
frond, bar = 5 cm (2 in.). outer one and, typically, by the sporangia. The sori are
borne on an outer, marginal vein supplied by two or more
lateral veins. In contrast, Hypolepis and Odontosoria (also
in the family Dennstaedtiaceae) have only one indusium,
and the sori are supplied by a single vein. Paesia is closely re-
lated to Pteridium but differs by having only one vascular
bundle in the stipe, whereas Pteridium has several arranged
in a horseshoe pattern.
Paesia consists of 12 species distributed in the Ameri-
can tropics, eastern Asia, and the western Pacific. It is
named for Duke Fernando Diaz Paes Leme, who visited
Brazil in 1660 on government service from Portugal.

Paesia scaberula (A. Richard) Kuhn  13.90.1

Hard fern, scented fern, lace fern
Semi-hardy, frost tolerant
A small to medium fern with hairy, long-creeping rhizomes
and stiff fronds. Grows best in high to medium light in
moist to moist-dry soil. Although difficult to transplant,
this species grows vigorously once established. In its native
 410 C 13

doors in Zone 9 in southern California, where they need

the heat and humidity of a greenhouse to thrive. The fo-
liage is particularly susceptible to attack from slugs, snails,
and scale insects.
The numerous (25 or more) narrow segments borne
perpendicular, or nearly so, to the rachis are a distinctive as-
pect of Pecluma. Most species are medium-sized epiphytes.
The plants of this genus are among the few ferns capable of
spreading from root proliferations. The rhizomes are short-
creeping and unbranched, never glaucous (as is sometimes
the case in the related genus Polypodium), and the scales are
nonclathrate and attached across the width of the base. The
black to reddish brown stipes are round in cross section
(not grooved) and jointed to the rhizome. Unlike in many
species of Polypodium, the narrowly elliptic blades of Pec-
luma are never scaly, although a few scales may be present
on the rachis. The veins are free or netted, simple or forked,
but in some species are difficult to see because of thick
blades. The sori are round and appear in one row between
the costae and margins, and like all polypodiaceous ferns,
they lack indusia.
The genus was formerly classified under Polypodium and
Figure 13.90.1. Paesia informally called the “Polypodium pectinatum-plumula
scaberula: habit, bar = complex.” Pecluma can be distinguished from Polypodium
5 cm (2 in.). by its short, unbranched rhizomes, nonclathrate and basally
attached rhizome scales, and the distinctive comb-like cut-
ting of the blades.
Pecluma consists of about 25 species native to the Amer-
New Zealand, it is weedy and volunteers readily in dis- ican tropics and subtropics. The genus name was formed by
turbed open areas, even in heavy soil. It will also volunteer compounding the specific epithets of two common, wide-
in greenhouses and spore pans. When rubbed or crushed, spread species: Pecluma pectinata and P. plumula. For a
the fronds emit a sweet odor, which is contained within the monographic treatment of this genus, see Evans (1969);
many glandular hairs on the surface of the blades. also see Polypodium for a comparison of Pecluma with re-
The stipes and rachises of Paesia scaberula are straight lated genera.
or only slightly zigzag and bear red-brown, short, bristly
hairs. The ovate or elliptic blades are three- to four-pin-
Pecluma dispersa (A. M. Evans) M. G. Price
nate, finely cut into segments about 1 mm wide. They are
 13.91.1
yellow green, and the lower surfaces bear many short, yel-
Tender, Zone 9; does poorly outdoors in Zone 9 of
lowish, sticky, glandular hairs. The species is endemic to
southern California
New Zealand.
A medium-sized fern with short-creeping rhizomes. Pre-
Pecluma fers medium light and moist but well-drained potting mix.
Comb fern This species can survive in Zone 9 but will not do well out-
doors in Zone 9 of southern California.
The numerous narrow, comb-like segments give Pecluma
Pecluma dispersa is characterized by the combination of
its distinctive appearance. The genus, which is only occa-
black rachises bearing conspicuous scales, blades essentially
sionally grown and mainly in the collections of fern hob-
glabrous on the lower surfaces, and twice-forked veins. It is
byists, consists of handsome ferns of warm, humid climates.
widespread in the American tropics.
In nature the plants typically grow on tree trunks or
branches, out of contact with the soil and its moisture. Dur-
ing the dry season the plants curl up and go dormant; how- Pecluma pectinata (Linnaeus) M. G. Price
ever, they revive within minutes after the first rains of the  13.91.2
next season. Most comb ferns are slow to establish in culti- Tender
vation, requiring more humidity than most ferns. They re- A medium-sized fern with short- to medium-creeping rhi-
spond well to frequent applications of nutrients. Although zomes. Grows well under medium light in moist, drained
they grow in Zone 9 in Florida, the plants do not grow out- potting mix. This species responds well to fertilization.
 F  F A  C 411

Figure 13.91.1. Pecluma dispersa:

a. habit and larger frond on right,
bar = 5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.), after Evans (1969);
c. rachis scales, bar = 1 mm, after
Evans (1969).

a
c

Pecluma pectinata is characterized by reddish brown,

minutely hairy (but nonscaly) rachises, blades hairy on the
lower surfaces with short-erect hairs, and twice-forked c
veins. The pinnae are greatly reduced toward the base of
the blade but are not eared or pointing downward as in
some species. Pecluma pectinata is widespread in the Amer-
ican tropics.

Pecluma plumula (Humboldt & Bonpland ex

Willdenow) M. G. Price  13.91.3
Tender, Zone 9; does poorly outdoors in Zone 9 of
southern California
A medium-sized fern with short- to medium-creeping rhi-
zomes. Grows under medium light in moist, drained pot- a
ting mix.
Pecluma plumula has black rachises beset with minute
hairs and scales. The blades are glabrous on the lower sur-
b
d

Figure 13.91.3. Pecluma plumula: a. fronds, bar = 5 cm

(2 in.); b. rhizome and frond base, bar = 1 cm (0.4 in.);
a Figure 13.91.2. Pecluma pectinata: a. habit, bar c. pinna, bar = 1 cm (0.4 in.), after Evans (1969);
= 5 cm (2 in.); b. pinnae, sterile (above) and d. rachis scales, bar = 1 mm, after Evans (1969).
fertile (below), bar = 1 cm (0.4 in.), after Evans
(1969).
 412 C 13

faces, and the pinnae at the base of the blades are reduced
abruptly to small, downward-pointing lobes. The veins are
once-forked. The species is widespread in the American
tropics.

Pellaea  13.92.1, 2

syn. Platyloma
Cliff brake
The cliff brakes are small- to medium-sized ferns with
mostly dark, polished stalks and gray-green, typically small
but broad leaflets. Pellaea usually grows on rock crevices,
ledges, and dry places, often in full sun. Semi-hardy spe-
cies, such as the button fern (Pellaea rotundifolia) and the
Australian cliff brake (P. falcata), are often sold as house-
plants because they are adapted to the low humidity usually
found indoors. Both species have narrow fronds and dark
green foliage, and they are easy to grow outdoors in sub-
tropical or warm-temperate climates. The taller, more di-
vided green cliff brake (P. viridis) is also easy to grow but
uncommon in the trade. All Pellaea species tolerate dryness
and, to some extent, irregular watering. They are best
grown in well-drained pockets of soil among rocks. Most of
the native American species are propagated by spores be-
cause they do not transplant readily.
The genus Pellaea is characterized by short- to long-
creeping rhizomes that bear bicolorous scales. The stipes Figure 13.92.2. Pellaea viridis: fertile pinnule.
are hard, often dark or blackish (some tan), and usually
shiny. The blades vary from one- to three-pinnate, their
apices bear a terminal leaflet (imparipinnate). The pinnules
is difficult to define because many of its characteristics are
or segments are broad, stalked or sessile, naked or with only
found in related genera. Pteridologists believe some species
a few scattered hairs. Several to many sori are borne on the
of Pellaea are not closely related and have accordingly trans-
margins. They may be oblong or linearly joined. The spo-
ferred them to other genera. It is likely that the familiar
rangia are covered by false indusia (rarely absent) formed
trade species Pellaea falcata and P. rotundifolia will eventu-
from the enrolled margins of the segments.
ally be transferred to Platyloma, a genus characterized by
Pellaea can usually be identified by the combination of
slender, long-creeping rhizomes, jointed segments, false in-
dark shiny stipes, pinnately divided blades ending in a ter-
dusia differentiated from the margin, and spiny spores.
minal leaflet, and the false indusium. The genus, however,
Pellaea is nearly cosmopolitan, but most of its 40 species
occur in the southwestern United States and Mexico, with
a secondary center of species richness in southern Africa.
The genus name comes from the Greek pellos, dusky, and
refers to the dull, bluish gray leaves of some species. See the
section on “Xerophytic Ferns” in Chapter 10 for more in-
formation on cultivating cliff brakes.

Pellaea andromedifolia (Kaulfuss) Fée

 13.92.3
Coffee fern
Semi-hardy, Zone (7)8; reportedly can endure
temperatures as low as −13°C (9°F)
A small to medium fern with slender, creeping rhizomes
and loosely clustered fronds. Grows under high light in
well-drained, moist-dry garden soil with coarse sand. The
common name alludes to the mature leaflets, which bear a
Figure 13.92.1. Pellaea falcata: habit. resemblance to coffee beans.
 F  F A  C 413

Pellaea andromedifolia is characterized by bicolorous rhi- narrower than the sterile ones. This species occurs from
zome scales, tan stipes, up to three-pinnate blades, and oval, Canada to Guatemala.
noncordate segments. The fiddleheads are densely scaly. In the wild this species grows on calcareous rocks such as
This species is native to the United States (California and dolomite and limestone. It resembles Pellaea glabella, with
Oregon) and Mexico (Baja California). which it often grows, but can be easily distinguished by the
hairy stipe and stalked pinnae and pinnules.
Pellaea atropurpurea (Linnaeus) Link  13.92.4
Purple cliff brake Pellaea brachyptera (T. Moore) Baker  13.92.5
Very hardy, Zone 4 Sierra cliff brake
A small to medium fern with stout, ascending rhizomes and Hardy, Zone 7
evergreen, clustered fronds. Prefers medium to high light A small fern with stout, ascending rhizomes and clustered
and moist-dry, basic garden soil with coarse sand. This spe- evergreen fronds. Grows under medium light in drained,
cies can tolerate annual minimum temperatures of −28 to moist-dry garden soil with coarse sand. The plants are dif-
−35°C (−23 to −31°F). ficult to cultivate, but they have been recorded to endure
Pellaea atropurpurea has rusty-tan and matted rhizome temperatures as low as −14°C (7°F).
scales. The stipe and rachis vary from purplish to black and Pellaea brachyptera has bicolorous rhizome scales, and
have a scruf consisting of minute, twisted hairs. The blade the stipes are dark brown and shiny. The blades are linear-
is bluish gray and two-pinnate, with the sterile segments oblong, two- to three-pinnate, and bluish green. The basal
oval to ovate-lanceolate. The fertile segments are longer and pinnae are one-pinnate, with the largest ones divided into
3–11 segments. The linear, tufted segments are crowded
on a short costa. The segment margins are greenish and
strongly enrolled, concealing nearly the entire lower sur-
Figure 13.92.3. Pellaea face of the pinnule. This species is native to the western
andromedifolia: frond, United States (California, Washington, and Oregon).
bar = 5 cm (2 in.).
Pellaea bridgesii Hooker  13.92.6
Bridges’ cliff brake
Hardy, Zone (5)6
A small fern with stout, ascending rhizomes and clustered,
deciduous fronds. Grows under high light in moist-dry,
drained garden soil with coarse sand. This species is difficult
to grow.

Figure 13.92.5. Pellaea brachyptera:

habit, bar = 5 cm (2 in.). After A. F.
Tryon (1957).

Figure 13.92.4. Pellaea

atropurpurea: habit, bar = Figure 13.92.6. Pellaea
5 cm (2 in.). After A. F. bridgesii: frond, bar =
Tryon (1957). 5 cm (2 in.).
 414 C 13

Some of the rhizome scales of Pellaea bridgesii are weakly Pellaea cordifolia is one of the few species in the genus
bicolorous. The stipes and rachises are dark brown and that has tan stipes and rachises. The blades are ovate-trian-
shiny. The linear blades are one-pinnate throughout, with gular and up to three-pinnate, with rachises and costae
the pinnae round to ovate, entire, and leathery. The fertile straight, not zigzag. The segments vary from round-cor-
pinnae are often folded lengthwise along the midvein. Spo- date to triangular-cordate, and the segment stalks are glab-
rangia are borne in a marginal band, but the margins re- rous. Hairs occasionally occur on the enrolled margin of
main flat, not enrolled as in many other species; P. bridge- indusium. This species is native to the southern United
sii therefore lacks a false indusium. The species is native to States (Texas) and Mexico.
the western United States.
Pellaea falcata (R. Brown) Fée  13.92.1, 8
Pellaea cordifolia (Sessé & Mociño) A. R. Smith syn. Platyloma falcata (R. Brown) J. Smith
 13.92.7 Australian cliff brake
syn. Pellaea cordata (Cavanilles) J. Smith, P. sagittata var. Semi-hardy, Zone (7)8
cordata (Cavanilles) A. F. Tryon A small-medium fern with dark green fronds in loose clus-
Semi-hardy, Zone 8 ters from short- to medium-creeping rhizomes. Grows well
A medium-sized fern with short-creeping rhizomes and lax, under medium to high light in drained, moist-dry garden
scrambling, bluish gray-green fronds. Grows well under or potting mix. This species can withstand temperatures
high light in drained, moist-dry garden soil with coarse down to −7°C (19°F) for one week.
sand. The longer fronds tend to become tangled and will The stipes and rachises of Pellaea falcata bear spreading
need support if grown upright. scales, and the blades are one-pinnate. The pinnae, which

b a

Figure 13.92.7. Pellaea cordifolia: a. frond, older, bar = Figure 13.92.8. Pellaea falcata fronds: a. typical form;
5 cm (2 in.); b. frond, younger; bar = 5 cm (2 in.). After b. var. nana; c. ‘Star Glow’, older (left) and younger
A. F. Tryon (1957). (right). Bar = 5 cm (2 in.).
 F  F A  C 415

are jointed to the rachis, are oblong and leathery, tapering

to an obtuse or acute apex that is sometimes mucronate. Figure 13.92.9. Pellaea
The indusia are thin and narrow. This species is native to glabella: habit, bar = 5 cm
India, Australia, New Zealand, Norfolk Island, and New (2 in.). After A. F. Tryon
Caledonia. (1957).
Some plants in cultivation are intermediate with Pellaea
rotundifolia; these have also been reported from New
Zealand.
var. nana Hooker. Figure 13.92.8b. This variety resem-
bles the type but is smaller and more compact, with pinnae
close or overlapping. The pinnae are also smaller, about
5–20 mm long and 2–6 mm wide. It is endemic to Aus-
tralia. Some botanists treat this variety as a species, Pellaea
nana (Hooker) Bostock.
‘Star Glow’. Figure 13.92.8c. Fronds dark green, shiny;
pinnae margins subdentate, slightly enrolled, the larger pin-
nae sharply eared at the base on the upper margin. It differs
from var. nana, from which it was apparently derived, by
having emarginate pinnae. This cultivar is often misidenti-
fied as Pellaea paradoxa (R. Brown) Hooker, a species not in
cultivation.

Pellaea glabella Mettenius ex Kuhn  13.92.9

Smooth cliff brake
Very hardy, Zone 3
A small fern with stout, ascending rhizomes and clustered
evergreen fronds. Grows under medium light in well-
drained, basic, moist-dry garden soil with coarse sand.
Pellaea glabella has rusty-brown rhizome scales that are
spreading, not matted. The stipes are brown, shiny,
smooth, and brittle. The bluish green blades vary from lin- Figure 13.92.10. Pellaea
ear to ovate-lanceolate and one- to two-pinnate. The basal mucronata: habit, bar =
pinnae are deeply divided into three to seven oblong to lin- 5 cm (2 in.). After A. F.
ear-ovate lobes or pinnules. The species is native to eastern Tryon. (1957).
North America, where it typically grows on limestone.

Pellaea mucronata (D. C. Eaton) D. C. Eaton

 13.92.10 Pellaea ovata (Desvaux) Weatherby  13.92.11
Bird’s-foot fern Flexuous cliff brake
Hardy, Zone 7 Semi-hardy, Zone 8
A small fern with stout, ascending rhizomes and evergreen, A medium-sized fern with short-creeping rhizomes and lax
clustered fronds. Grows well under high light or full sun fronds. Grows well under medium-high light in moist-dry,
in coastal areas in well-drained garden soil with coarse sand. drained garden soil with coarse sand. The longer fronds
This species is difficult to grow. The common name bird’s- tend to become tangled and need support.
foot fern alludes to the segment clusters, which resemble a Pellaea ovata can be distinguished from all others in the
bird’s foot. genus by its tan, zigzag stipes and rachises. Other charac-
Pellaea mucronata has bicolorous rhizome scales, and teristics are bicolorous rhizome scales and elongate-trian-
the stipes are dark brown and shiny. The ovate-triangular to gular blades to 84 cm (34 in.) long and 30 cm (12 in.)
lanceolate blades are up to three-pinnate toward the base, broad, mostly three-pinnate and gray-green. The segments
with costae much longer than the ultimate segments. The vary from elliptic to ovate and are sagittate to cordate at the
larger pinnae have more than 10 segments. The indusia base. The segment stalks bear short, reddish hairs. The en-
margins are uneven or erose. This species is native to the rolled indusium margin is glabrous. This species is native to
western United States and Mexico. the United States (Texas), Central and South America, and
the West Indies.
 416 C 13

Pellaea ternifolia (Cavanilles) Link  13.92.13

Semi-hardy, Zone 8
A small to medium fern with ascending, stout rhizomes and
clustered fronds. Grows under high light in well-drained,
b moist-dry garden soil or potting mix.
Pellaea ternifolia has particularly distinctive pinnae,
which are three-parted and sessile or nearly so. These char-
acteristics easily separate this species from others in the
genus. Additional traits include bicolorous rhizome scales;
black or dark purple, shiny stipes and rachises; and linear,
one-pinnate-pinnatifid blades. The pinnae are entire or
deeply three-lobed with mucronate apices. The species is
native to the United States (Texas), Central and South
America, and the Hawaiian Islands.

Pellaea viridis (Forsskål) Prantl  13.92.2, 14

syn. Cheilanthes viridis (Forsskål) Swartz, Pellaea hastata
(Thunberg) Prantl
Green cliff brake
Semi-hardy
A medium-sized fern with short-creeping rhizomes and
clustered fronds. Grows well under high light in well-
drained, moist-dry garden soil or potting mix.
Pellaea viridis has bicolorous rhizome scales and shiny

a Figure 13.92.12. Pellaea

rotundifolia: frond, bar =
5 cm (2 in.).

Figure 13.92.11. Pellaea ovata: a. frond, bar = 5 cm

(2 in.); b. fertile pinnule, bar = 1 cm (0.4 in.), after
Vareschi (1968).

Pellaea rotundifolia (G. Forster) Hooker

 13.92.12
syn. Platyloma rotundifolia (G. Forster) J. Smith
Button fern, New Zealand cliff brake
Semi-hardy, Zone 8; can tolerate temperatures as low as
−7°C (19°F) for one week
A small-medium fern with short-medium-creeping rhi-
zomes and fronds in a loose cluster. Grows well under me-
dium to high light in moist-dry, well-drained garden soil
with coarse sand. The plants are easy to grow and thrive in-
doors. Do not water until the soil is nearly dry.
Pellaea rotundifolia resembles P. falcata (which see) but
has smaller and narrower fronds. The round to oval pin- Figure 13.92.13. Pellaea
nae are up to 2 cm (0.75 in.) long and 1.3 cm (0.5 in.) ternifolia: habit, bar =
wide, with margins minutely but shallowly toothed and 5 cm (2 in.). After A. F.
apices rounded but abruptly narrowed to a sharp point. Tryon (1957).
The species is endemic to New Zealand.
 F  F A  C 417

Figure 13.92.14. Pellaea

viridis: frond, bar = 5 cm
(2 in.).

Figure 13.92.15. Pellaea

wrightiana: habit, bar =
5 cm (2 in.).

dark brown to black stipes and rachises grooved on upper

surface. The blades are lanceolate to triangular, commonly
two- to three-pinnate, and have basal pinnae that are more wrightiana is native to the southwestern United States and
developed on the lower side. The segments vary from cor- Mexico.
date to hastate to wedge-shaped but are most commonly
lanceolate to ovate. The veins are free (not netted) and eas- Pentagramma
ily visible. The false indusium is thin and subentire. This Goldback fern, silverback fern
species is native from Africa to India. The genus Pentagramma consists of small ferns of dry habi-
Pellaea viridis is extremely variable in blade cutting. tats. They are distinguished by powdery yellow or white
Most cultivated plants are typical P. viridis, but a few are in- undersides and pentagonal fronds. Mature plants grow best
termediate with var. macrophylla (Kunze) Sim (P. adian- in pots or rock gardens but are sensitive to overwatering
toides J. Smith), which is one-pinnate, or sometimes two- and must be transplanted with care. If landscape effect is of
pinnate at the base, with large pinnae. Some botanists prime importance, a similar growth form can be obtained
consider this species best placed in Cheilanthes. by planting Cheilanthes argentea, which is much easier to
grow. All species of Pentagramma germinate easily from
Pellaea wrightiana Hooker  13.92.15 spore.
syn. Pellaea ternifolia var. wrightiana (Hooker) A. F. The ascending rhizomes of Pentagramma are covered
Tryon with sharply bicolorous scales. The stipes are dark brown to
Hardy, Zone 6 black and round or nearly so in cross section; they contain
A small to medium fern with ascending, stout rhizomes and only one vascular bundle. The blades are triangular-pen-
clustered grayish fronds. Grows well under high light in tagonal and one- to two-pinnate-pinnatifid, and the pinnae
well-drained, moist-dry garden soil or potting mix. are sessile or adnate, usually with white or yellow powdery
Pellaea wrightiana resembles P. ternifolia, but it has undersides. Sporangia are borne along the free, dichoto-
more narrowly triangular blades, longer and more divided mously branched veins. An indusium is absent. The two
pinnae lobed or divided into 3–11 pinnules, and fully two- species of Pentagramma were formerly included in the
pinnate basal pinnae. It differs from P. brachyptera by hav- genus Pityrogramma because of their powdery lower blade
ing longer costae and whitish bordered segments. Pellaea surfaces and sori that run along the veins. Pteridologists
 418 C 13

currently consider these genera distantly related within the Pentagramma pallida (Weatherby) Yatskievych, Wind-
Pteridaceae and best placed in different subgroups of the ham & E. Wollenweber differs by having dull and pow-
family. dery stipes and blades that are white-powdery on both sur-
Pentagramma contains two species and is native to the faces, appearing grayish when fresh. It is endemic to the
southwestern United States and northwestern Mexico. The foothills of the Sierra Nevada in California and may be cul-
genus name comes from the Greek penta, five, and gramme, tivated by local hobbyists. Its cultural requirements are the
line, alluding to the pentagonal outline of the blade. same as for P. triangularis.

Pentagramma triangularis (Kaulfuss) Yatskievych, Phegopteris

Windham & E. Wollenweber  13.93.1 syn. Thelypteris subgenus Phegopteris
Semi-hardy, Zone 7 Beech fern
A small fern with short-creeping rhizomes. Grows under These medium-sized, terrestrial ferns have creeping rhi-
medium-high light in dry, well-drained garden soil. This zomes. Phegopteris was previously considered a subgenus of
species is difficult to cultivate, and the plants are sensitive to Thelypteris but can be distinguished from that genus by pin-
overwatering. nae that are broadly adnate in the distal half of the blade
The stipes of Pentagramma triangularis are shiny and and connected by wings along the rachis. Like all thelypte-
generally not powdery. The blades are white- or yellow- riod ferns, Phegopteris has only two vascular bundles in the
powdery below, appearing bright green or sometimes stipe (most other similar fern groups have four or more)
yellowish green when fresh. The species is extremely varia- and needle-shaped hairs. Indusia are absent. The species
ble and four subspecies have been described to accommo- with broadly triangular blades might be confused with
date the variation. It is native to western North America
Gymnocarpium, but that genus lacks the needle-like hairs.
and northwestern Mexico. In nature the plants occurs in
Phegopteris contains three species, all of which are culti-
rock crevices, at the base of overhanging boulders, and on
vated, native to the temperate regions of North America,
roadbanks.
Europe, and Asia. The genus name comes from the Greek
phegos, beech, and pteris, fern, since Phegopteris commonly
grows under beech trees.

Phegopteris connectilis (Michaux) Watt

 13.94.1
syn. Thelypteris phegopteris (Linnaeus) Slosson
Narrow beech fern, long beech fern
Very hardy, Zone 2
A medium-sized fern with medium-creeping rhizomes and
deciduous fronds. Grows well under low light or full shade
in moist, acidic garden soil or potting mix.
Phegopteris connectilis has slender, creeping rhizomes and
triangular blades. The basal pair of pinnae are sessile or ad-
nate to the rachis but not connected to those above by a
wing. The rachises and costae bear conspicuous brown
b scales, and the veins in the segments of middle pinnae are
mostly simple. This species is native to the northern regions
of North America, Europe, and Asia.

Phegopteris decursive-pinnata (H. C. van Hall) Fée

 13.94.2
syn. Thelypteris decursive-pinnata (H. C. van Hall) Ching
a Winged beech fern
Very hardy, Zone 4
A medium-sized fern with suberect rhizomes and erect, de-
ciduous fronds. Grows well under medium light in moist-
Figure 13.93.1. Pentagramma triangularis: a. frond, bar wet garden soil or potting mix. This species is easy to grow
= 2.5 cm (1 in.); b. sporangia along veins, bar = 3 mm from spores and generally does well in gardens in the east-
(0.1 in.). ern United States. Several new plantlets are produced each
 F  F A  C 419

Phegopteris hexagonoptera (Michaux) Fée

 13.94.3
b a
syn. Thelypteris hexagonoptera (Michaux) Nieuwland
Southern beech fern, broad beech fern
Very hardy, Zone 3
A medium-sized fern with medium-creeping rhizomes and
deciduous fronds. Grows well under low light or full shade
in moist, slightly acidic garden soil or potting mix. It grows
readily from spores.
Phegopteris hexagonoptera has slender, creeping rhizomes
and broadly triangular blades. The rachises and costae are
hairy and bear a few inconspicuous, narrow scales. A wing
along the rachis connects the basal pinnae to those above.
The veins are always forked or pinnate. This species is na-
tive to eastern North America.

Figure 13.94.1. Phegopteris connectilis: a. habit, bar =

5 cm (2 in.); b. incompletely winged rachis, bar = a
2.5 cm (1 in.).

year from short runners. The green of the fronds lasts until
the first hard frost.
The rhizomes of Phegopteris decursive-pinnata are erect
or suberect, and the blades are narrowly elliptic and tapered
toward both ends, rather than broadly triangular as in the
other two species. The pinnae are hairy on both surfaces,
and their bases are connected to the rachis by a broadly
lobed wing, but the lower pinnae are often free. The rachis
is also hairy and bears ciliate scales. The species is native to b
eastern Asia.

Figure 13.94.3. Phegopteris hexagonoptera: a. habit, bar =

5 cm (2 in.); b. winged rachis, bar = 2.5 cm (1 in.).

Phlebodium
syn. Polypodium in part
Rabbit’s-foot fern, golden polypody
Phlebodium aureum is well known in the trade as a pot or
basket plant, though it can also be grown in soil with good
drainage. Its fronds and those of P. pseudoaureum vary con-
siderably, from gray-green to blue-green shades, from stiff
and erect to arching or pendant, from medium to large in
b size, from entire to fringed lobes, from sparsely to heavily
crested tips, and so on. These characteristics are affected by
the amount of nutrients and exposure to light. Phlebodium
a aureum is less cold hardy and needs more protection than P.
pseudoaureum, which is also a robust grower. For more de-
tails on Phlebodium cultivars, see Hoshizaki (1982).
Phlebodium is closely related to Polypodium, and some
Figure 13.94.2. Phegopteris decursive-pinnata: a. habit, pteridologists prefer to classify it in that genus. Like Poly-
bar = 5 cm (2 in.); b. winged rachis, bar = 1 cm (0.4 in.). podium, it has creeping rhizomes with two ranks of fronds
 420 C 13

on the upper surface, round, non-indusiate sori, and yellow

bean-shaped spores. But Phlebodium differs by its venation:
the areoles contain two veinlets that supply the sorus, and
the areoles next to the costae are long, narrow, and lack in-
cluded veinlets. The frond shape also differs from that of
many polypodiums, being large and broadly cut with wide,
adnate segments and a terminal segment that resembles the b
lateral ones. See Polypodium for a comparison with other
polypodiaceous genera.
Phlebodium contains four species native to tropical
America. The genus name comes from the Greek phlebodes,
full of veins, in reference to the highly branched veins.
a
Phlebodium aureum (Linnaeus) J. Smith
 13.95.1 c
syn. Polypodium aureum var. aureum
Hare’s-foot fern, rabbit’s-foot fern, bear’s-paw fern,
serpent fern, golden polypody
Tender to semi-tender, Zone 9(10)
A medium to large fern with moderately long-creeping rhi-
zomes. Grows well under medium light in moist to moist-
dry garden soil or potting mix kept well drained. This spe-
cies is easy to cultivate and has more gracefully arching
fronds than Phlebodium pseudoaureum, which it resembles,
although it is less cold hardy. Figure 13.95.1. Phlebodium
Phlebodium aureum resembles P. pseudoaureum but dif- aureum: a. habit, bar =
fers by having the sori arranged in two rows between the 10 cm (4 in.); b. sori and
costa and margin. The fronds are deeply lobed into narrow vein pattern, bar = 1 cm
segments. Phlebodium aureum is believed to be a fertile hy- (0.4 in.), after Hoshizaki
brid of a cross involving P. decumanum and P. pseudoau- d
(1982); c. sori distribution,
rum. It is native to tropical America. bar = 1 cm (0.4 in.);
Some cultivated plants may be hybrids between Phle- d. ‘Mandaianum’, bar =
bodium aureum and P. pseudoaureum, as evidenced by their 10 cm (4 in.).
irregular venation and irregular rows of sori (inconsistently
ranging from 1 to 2 rows). See Figure 13.95.1c.
‘Cristatum’. Frond tips several times branched, tips of
pinnae less so. Phlebodium decumanum can be distinguished from the
‘Ekstrand’. Pinnae irregularly lobed, the lobes crowded, other cultivated species by its sori, which are arranged in
twisted, and irregularly and coarsely cut into teeth; blades three to seven rows (not one or two) between the segment
strongly ruffled. midrib and margins. The fronds are generally larger, often
‘Mandaianum’. Figure 13.95.1. Pinnae coarsely and over 1 m (3 ft.) long. The species is widespread in tropical
irregularly incised to laciniate, weakly ruffled. America.
‘Mayi’. Blades ruffled and fringed.
‘Undulatum’. Blades wavy. Phlebodium pseudoaureum (Cavanilles) Lellinger
 13.95.3;  5, 38
Phlebodium decumanum (Willdenow) J. Smith syn. Phlebodium areolatum (Humboldt & Bonpland ex
 13.95.2 Willdenow) J. Smith, Polypodium areolatum
syn. Polypodium decumanum Willdenow Humboldt & Bonpland ex Willdenow
Tender, Zone 10 Tender to semi-tender, Zone (8)9
A large fern with moderately long-creeping rhizomes. A medium to large fern with moderately long-creeping rhi-
Grows well under medium light in moist, well-drained pot- zomes. Grows well under low to high light in well-drained,
ting mix. This species is easy to cultivate and volunteers moist to moist-dry garden soil or potting mix. The plants
readily from spores. The orange rhizome scales are highly are easy to cultivate and robust growers. The fronds are typ-
attractive. ically firmer and stiffer than those of Phlebodium aureum.
 F  F A  C 421

Figure 13.95.2. Phlebodium decumanum: a. frond, bar = 10 cm (4 in.), after

Hoshizaki (1982); b. sori distribution, bar = 1 cm (0.4 in.); c. sori and vein
pattern, bar = 1 cm (0.4 in.), after Hoshizaki (1982).
a

The sori of Phlebodium pseudoaureum are arranged in

one row between the costa and margins, but a few may be
scattered in a second row. The species is widespread in trop-
ical America.
Within Phlebodium pseudoaureum, the blue-green (glau-
cous) plants vary considerably and circulate in the trade c
under a variety of loosely applied cultivar names under P. a
glaucum and P. glaucopruniatum. Some of the blue-green
cultivars include the following:
‘Mexican Tasseled’. Segment apices densely tasseled.
‘Leatherman’. Pinnae falcate, attenuate; fronds lax.

Phlebodium ×schneideri Horticulture

 13.95.4 b
Schneider’s rabbit-foot fern
Semi-tender (?)
A medium to large fern with moderately long-creeping rhi-
zomes. Grows under medium light in well-drained, moist
to moist-dry potting soil or uncut moss. The plants are no-
table for their cascading fronds. Avoid watering the ruffled
foliage because the extra weight might snap the stipe.
Phlebodium ×schneideri is reported to be a sterile hybrid
of horticultural origin resulting from a cross between Phle-
bodium aureum (?) and Polypodium vulgare ‘Elegantissi-
mum’, but it resembles other cultivars of Phlebodium au- Figure 13.95.3. Phlebodium pseudoaureum: a. frond
reum and P. pseudoaureum, only with more divided blades. variations, bar = 10 cm (4 in.); b. sori distribution, bar =
The pinnae are deeply lobed (almost fully divided), and the 1 cm (0.4 in.); c. sori and vein pattern, bar = 1 cm
lobes are in turn shallowly lobed and strongly ruffled. (0.4 in.), after Hoshizaki (1982).
 422 C 13

Figure 13.95.4.
Phlebodium
×schneideri: frond,
bar = 10 cm (4 in.).

Figure 13.96.1. Phyllitis scolopendrium: habit.

Phyllitis
Hart’s-tongue fern
The hart’s-tongue ferns are small, terrestrial, evergreen
plants with tongue-shaped fronds. The fiddleheads of Phyl-
litis were used as the model for the scrolls at the ends of vio-
lins. There is wide variation in spore-grown plants, and
many garden forms are seen in cultivation. Phyllitis may
also be propagated from basal pieces of the stipe, a process
described in Chapter 8. The genus is susceptible to root-
rot, so avoid overwatering and provide good drainage. For
details about cultivars, see Druery (1912), Kaye (1968),
and Dyce (1972).
The distinguishing characteristics of the genus are its
simple, entire blades and paired sori in which each sorus in
the pair opens toward its partner. Additional traits include
the heart-shaped blade bases, leathery texture, once- or
twice-forked veins, and linear indusiate sori. Phyllitis is
closely related to Asplenium and hybridizes with it. Some
pteridologists prefer to classify Phyllitis under Asplenium;
it is only a matter of opinion whether to do so. Phyllitis is
used here because of its long use by horticulturists.
The genus contains five species and is native to North
America, Europe, and Asia. The name comes from the
Greek phyllos, meaning leaf, which was an ancient Greek
name for the fern.

Phyllitis scolopendrium (Linnaeus) Newman

 13.96.1–4;  39
syn. Asplenium scolopendrium Linnaeus
Hart’s-tongue fern
Hardy, Zone 4(5)
A small-medium fern with suberect rhizomes and evergreen Figure 13.96.2. Phyllitis scolopendrium: fronds.
fronds. Grows well under low light in well-drained, basic,
moist-dry potting mix. The plants are sensitive to poor
drainage and root-rot.
 F  F A  C 423

1890). For representative examples of the numerous culti-

vars, see Figure 13.96.4 and those listed here:
‘Crispum’. Blade margins frilly.
‘Cristatum’. Frond apices crested.
‘Laceratum’. Blade margins deeply and irregularly cut.
‘Muricatum’. Blade surfaces roughened by ridges.
‘Ramosum’. Fronds branched.
‘Undulatum’. Blade margins wavy.

Pilularia
Figure 13.96.3. Phyllitis scolopendrium: sori Pillwort
and indusia. The pillworts are small, inconspicuous, sedge-like or grass-
like plants. They can be distinguished from grasses and
sedges by the coiled tips of their young leaves. Pilularia is
Phyllitis scolopendrium has tongue-shaped fronds with related to the water clover (Marsilea) but lacks the distinc-
cordate bases. The stipes are short and scaly. The species is tive clover-like leaves. Of little ornamental value, this genus
native to North America and Europe. is best used as part of a small, aquatic dish-garden or in bog
Only the European variety (var. scolopendrium) is culti- or marsh plantings.
vated. The smaller, American variety (var. americana Fer- The rhizome is slender and creeping, bearing a filamen-
nald) is rare and difficult to grow. It tends to bear sori in the tous stipe at each node. The fertile part, which is not al-
distal half of the frond but is otherwise similar to the Euro- ways present, is a stalked, bean-like structure (sporocarp)
pean variety. borne at the base of the stipe. The sporocarp encloses the
The hart’s-tongue fern is extremely variable. In the late sporangia, which contain either male or female spores.
1800s, one British fern grower listed 445 varieties (Lowe Pilularia is cosmopolitan and consists of five species.
The genus name comes from the Greek pilula, little ball,
and refers to the globose sporocarps.

Pilularia americana A. Braun  13.97.1

Hardy, Zone 5
A small fern with medium-creeping rhizomes. Grows well
b under high light in sandy or silty garden soil kept wet. The
plants can also grow submerged or partly submerged. Do
not let them dry completely.
a c Pilularia americana resembles P. globulifera by its small,
grass-like leaves but differs by having sporocarps borne

e Figure 13.97.1.
Pilularia americana:
habit, bar = 1 cm
(0.4 in.).

Figure 13.96.4. Phyllitis scolopendrium fronds: a. wild

form; b. ‘Ramo-cristatum’; c. ‘Kaye’s Lacerate’;
d. ‘Marginato-multifidum’; e. ‘Capitatum’. Bar = 5 cm
(2 in.).
 424 C 13

below, rather than above, the soil. It is native to the United

States and Mexico.

Pilularia globulifera Linnaeus  13.97.2

Hardy, Zone 5
A small fern with medium-creeping rhizomes. Grows well
under high light in sandy or silty garden soil kept wet. The
plants can also grow submerged or partly submerged. Do
not let them dry completely. This species is a rapid and ro-
bust grower in moist to wet soil.
Pilularia globulifera resembles the preceding species, but
its sporocarps are borne above the soil. Submerged plants
do not form sporocarps. Native to Europe.

Figure 13.98.1. Pityrogramma argentea: habit.

Pityrogramma has short-creeping to erect rhizomes cov-

ered with scales of one color or with a poorly defined darker
central portion. The stipes are dark and shiny, often stiff,
Figure 13.97.2. grooved on the upper surface, and have two or more vas-
Pilularia globulifera: cular bundles. The blades vary from linear-lanceolate to tri-
habit, bar = 1 cm angular, and their lower surfaces usually have a white, yel-
(0.4 in.). low, or orangish powder (farina). The lower pinnae are
stalked, and the veins are free, not netted. The sori run
along the veins, forming dark lines that sometimes become
confluent as the sporangia mature. An indusium is absent.
The genus presents several taxonomic and nomenclat-
ural problems that might confuse gardeners. The species

Pityrogramma  13.98.1, 2

syn. Trismeria
Goldback fern, silverback fern
The genus Pityrogramma is noted for its powdery yellow or
white lower blade surfaces. These small- to medium-sized
terrestrial ferns grow quickly; some species are short-lived.
On older plants, the roots often fail to support the ascend-
ing rhizome and fronds. Support can be provided by add-
ing mulch and rocks around the base of the plant and re-
moving some of the older fronds. The roots are sensitive to
disturbance, and so these ferns should be transplanted with
care. Many species of Pityrogramma are easy to grow from
spores and may become weedy in the greenhouse or spore Figure 13.98.2. Pityrogramma argentea: underside
pans. Avoid overwatering and check that the soil is well of pinnule, showing powder and sporangia (dark
drained. spots) along vein.
 F  F A  C 425

vary morphologically and hybridize readily, which compli-

cates identification. Many previously accepted species have
been reduced to varietal rank, and most pteridologists now Figure 13.98.3.
merge Trismeria with Pityrogramma. Pityrogramma triangu- Pityrogramma
laris, often called the goldback fern (as are other species of argentea: frond, bar =
Pityrogramma), has long been placed in this genus but is 5 cm (2 in.).
now classified in Pentagramma (which see). For taxonomic
details on the species of Pityrogramma, see Tryon (1962).
Pityrogramma, which contains 16 species, is native to
tropical and subtropical America, Africa, and Madagascar.
One species, Pityrogramma calomelanos, has become natu-
ralized in many parts of the Old World tropics. The genus
name comes from the Greek pityron, scurf, and gramme,
line, referring to the powdery lower blade surface, which
has sporangia in lines along the veins.

Pityrogramma argentea (Willdenow) Domin

 13.98.1–3
Semi-tender
A small to medium fern with suberect rhizomes. Grows
under high light in moist-dry, drained garden soil or pot-
ting mix.
Pityrogramma argentea is characterized by silver or gold
lower blade surfaces and somewhat wedge-shaped ultimate
segments that lack a well-defined midrib. The basal pinnae
are the largest. It is native to Africa and Madagascar.

Pityrogramma austroamericana Domin

 13.98.4;  40, 41
syn. Pityrogramma calomelanos var. aureoflava of authors,
not (Hooker) Weatherby ex L. H. Bailey, P.
calomelanos var. austroamericana (Domin) Farwell
Semi-tender, Zone 9
A medium-sized fern with suberect rhizomes. Grows well
under high-medium to high light in moist-dry garden soil
or potting mix kept well drained. The plants volunteer Figure 13.98.4. Pityrogramma
readily from spores. austroamericana: frond, bar =
Pityrogramma austroamericana differs from P. calome- 5 cm (2 in.).
lanos by its bright yellow blade undersides and firmer tex-
ture in live plants. It is native to the New World tropics
and naturalized in the Hawaiian Islands.
For many years this fern was misidentified as Pityro-
gramma hybrida Domin, a hybrid that is not cultivated. pinnate, white on the underside, and has ascending distal
pinnae and segments. This species resembles P. ebenea. It is
Pityrogramma calomelanos (Linnaeus) Link common and widespread in the American tropics and has
 13.98.5 become naturalized in many parts of the Old World tropics.
Silver fern
Tender, Zone 10 Pityrogramma ebenea (Linnaeus) Proctor
A medium-sized fern with suberect rhizomes. Grows well  13.98.6
under high light in drained, moist-dry garden soil or pot- syn. Pityrogramma tartarea (Cavanilles) Maxon
ting mix. It prefers some air circulation. Tender
Pityrogramma calomelanos has scales on the rhizome and A medium-sized fern with suberect rhizomes. Grows well
stipe tipped with a long conical cell. The stipe is often as under high light in moist-dry, well-drained garden soil or
long as the blade, which is two-pinnate-pinnatifid to three- potting mix.
 426 C 13

Figure 13.98.5.
Pityrogramma
calomelanos: Figure 13.98.6. Pityrogramma
a
underside of frond, ebenea: a. frond, bar = 5 cm
bar = 5 cm (2 in.). (2 in.); b. underside of
pinnule, bar = 2.5 cm (1 in.).

The scales of the rhizomes and stipes are tipped with

several single cells ending in a short, round cell. The pin-
nules of Pityrogramma ebenea are white to cream or color-
less on the lower surface and broadest at the base. They are
inserted at right angles to the pinna rachis. This species is
common and widespread in the American tropics.
Pityrogramma ebenea resembles P. calomelanos but dif-
fers by having wider basal pinnae on the lower side and the
distal pinnae perpendicular to the rachis. In contrast, the
basal pinnae of P. calomelanos are equally wide on both
sides, and the distal pinnae are ascending. An elevational
difference between the two species occurs in the wild: P.
ebenea grows above 1000 m (3300 ft.), whereas P. calome-
lanos generally grows below that elevation.

Pityrogramma trifoliata (Linnaeus) R. M. Tryon

 13.98.7
syn. Trismeria trifoliata (Linnaeus) Diels
Tender; survives Zone (9)10 with warm nights
Figure 13.98.7. Pityrogramma
A large fern with suberect rhizomes. Grows well under high
trifoliata: frond, bar = 5 cm
light in moist-wet garden soil or potting mix.
(2 in.).
Pityrogramma trifoliata has bicolorous rhizome scales,
and the fronds are large, erect, coarse, linear or narrowly
 F  F A  C 427

lanceolate, and usually white on the lower surfaces. The

pinnae are usually two- to three-parted (rarely up to seven-
parted) and spread in three dimensions around the rachis,
thus the blade is not flat. This species is native to southern
Florida and tropical and subtropical America.
This species differs from all others in the genus by hav-
ing pinnae deeply parted into two or three (or more) seg-
ments. (The pinnae are not always three-parted as the
specific epithet suggests.) Because of this difference, Pityro-
gramma trifoliata is sometimes classified in the genus Tris-
meria. Nowadays, however, pteridologists classify it with
Pityrogramma, partly because it readily hybridizes with sev-
eral species in the genus. The hybrids are usually common
where the species grow together, such as along roadsides.

Platycerium  13.99.1–3

Staghorn fern, elkhorn fern
Staghorn ferns are bizarre-looking, medium- to large-sized
epiphytes grown primarily on walls, tree trunks, baskets,
and pots. The base of the plant is composed of highly mod-
ified, thick, spongy fronds that provide anchorage and root-
ing media for the fertile foliage fronds. Australian species,
such as Platycerium bifurcatum and its relatives, are easy to
grow outdoors in subtropical climates. In many parts of the
United States they can be grown outdoors until the first
frost. Indoors, they should be kept in a well-lighted, airy
place but out of direct sun. The thick foliage fronds of
Platycerium tolerate the lower humidity indoors. Some spe-
cies form new plants (pups) from root buds, which can be
separated from the parent plant when they produce base
fronds about 13 cm (5 in.) wide. For more information,
see “Staghorn Ferns” in Chapter 10. Figure 13.99.1. Platycerium willinckii: habit.
Platycerium can be identified by the star-shaped hairs
on its blades and the striking frond dimorphism, with the
layers of sessile brown fronds appressed to the substrate and
larger, green leafy fronds that are upright, arching, or pen-
dulous. The latter type of frond bears sori; the former does
not. Other characteristics of the genus are scaly, very short-
creeping, stout rhizomes that are concealed by the base
fronds, except at the tip. The thick and spongy base fronds
are green at first but soon turn brown and persist on the
plant. The edges are thinner, and in some species the upper
margin is extended and foliaceous, lobed or not. The fo-
liage fronds vary from simple and entire to dichotomously
lobed or forked into strap-shaped segments. The sporangia
are formed in large patches on the lower surface and are
covered with star-shaped hairs. Indusia are absent.
The genus contains 15 to 18 species and occurs entirely
in the Old World tropics, except for a single South Amer-
ican species, Platycerium andinum. The genus name comes
from the Greek platys, flat, and keras, horn, referring to the
flattened, antler-like leaves.
A large number of loosely named platyceriums have ap- Figure 13.99.2. Platycerium willinckii: sporangial
peared in the trade in recent years. If you come across a area.
 Malayan-Asiatic species African-American species

P. quadridichotomum
stipe
Base frond
cross-
P. wallichii of Malayan-
section
Asiatic species
P. andinum
except
P. ridleyi

P. grande

P. superbum
P. stemaria
P. elephantotis

P. wandae
P. madagas-
cariense

P. ellisii
P. holttumii
P. alcicorne

P. coronarium
Javan-Australian species

P. veitchii
P. ridleyi

stipe cross section

stipe cross section P. willinckii

P. bifurcatum P. hillii
Figure 13.99.3. Frond and stipe (in cross section) characters of Platycerium species: Malayan-Asiatic species, with a dark
ring of tissue in the stipe, vascular bundles present or absent in the center; African-American species, with a pale ring of
tissue in the stipe, vascular bundles scattered in the center; Javan-Australian species, with a pale ring of tissue, the vascular
bundles in a circle, not scattered. After Hoshizaki (1972).
 F  F A  C 429

plant not named here, it probably represents a variant of

a
the Platycerium bifurcatum complex (P. bifurcatum, P.
hillii, P. veitchii, and P. willinckii), the members of which
are widely grown and noted for their variability. For a dis- c b
cussion of this complex, see P. bifurcatum. For more horti-
cultural information on the species and cultivars, see Joe
(1964), Franks (1969), and Vail (1984); for scientific treat-
ments, see Hoshizaki (1972), Hennipman and Roos
(1982), and Hoshizaki and Price (1990).
To identify the species, use the outline of the three
groups given here and in Figure 13.99.3. Once an uniden-
tified species is limited to one of the groups, it can be com- d
pared to the illustrations and descriptions of the individ-
ual species in that group. Figure 13.99.4. Platycerium alcicorne: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome
Malayan-Asiatic species. Some species may extend to Indo-
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
nesia, New Guinea, and Australia. Stipe cross section has a
1 cm (0.4 in.).
dark ring of tissue, and vascular bundles are arranged in a
circle with some bundles scattered within the circle or not;
base fronds spreading, upper margin deeply lobed (except
for Platycerium ridleyi, which has reniform base fronds); Platycerium alcicorne resembles P. bifurcatum, which dif-
roots do not produce buds; mostly large plants. (Platy- fers by the circular (not scattered) arrangement of the vascu-
cerium coronarium, P. grande, P. holttumii, P. ridleyi, P. lar bundles in the stipe and often lobed (not entire) upper
superbum, P. wallichii, P. wandae) margins of the base fronds. Some pteridologists consider P.
vassei the same as P. alcicorne, but in horticulture the former
African-American species. Stipe cross section does not have
name has been applied to a plant that has lighter green blades
a dark ring of tissue, and vascular bundles are arranged in a
and very convex, smooth base fronds. The main radiating
circle with some scattered within the circle; base fronds var-
veins on the base fronds of older plants of P. alcicorne from
iable, and if spreading, seldom as wide as the Malayan-Asi-
Madagascar are often raised, but the main cross-veins are
atic species; roots produce buds; mostly medium-sized or
not. In the related species P. madagascariense the main cross-
smaller plants compared to the Malayan-Asiatic species.
veins are also raised, forming a waffle-like pattern.
(Platycerium alcicorne, P. andinum, P. elephantotis, P. ellisii,
P. madagascariense, P. quadridichotomum, P. stemaria)
Platycerium andinum Baker  13.99.5
Javan-Australian species. Stipe cross section does not show American staghorn
a dark ring of tissue, and vascular bundles are arranged in a Semi-tender to tender
circle with no bundles within the center of the circle; base A large fern with a short rhizome and bud-forming roots.
fronds variable, and if spreading, seldom as wide as the Grows well under high light in drained, moist-dry, coarse
Malayan-Asiatic species; roots produce buds; mostly me- potting mix or uncut moss.
dium-sized or smaller plants compared to the Malayan-Asi- In Platycerium andinum the distal part of the base fronds
atic species. (Platycerium bifurcatum, P. hillii, P. veitchii, P. are extended, arching forward, forked, and lobed. The de-
willinckii) cumbent foliage fronds are more than 1 m (3 ft.) long,
hairy, and gray-green. The sporangial patch is borne on the
Platycerium alcicorne Desvaux  13.99.4 strap-shaped segments, typically near the middle of the
syn. Platycerium vassei Poisson frond but may be more distal. The species is native to Bo-
Tender to slightly hardier livia and Peru.
A medium-sized fern with a short rhizome and bud-form-
ing roots. Grows well under high light in drained, moist- Platycerium bifurcatum (Cavanilles) C. Christensen
dry, coarse potting mix or uncut moss. The plants are easy  13.99.6;  1, 5, 42
to cultivate. Common staghorn
Platycerium alcicorne has round–kidney-shaped base Semi-tender
fronds with entire margins. The center of the stipe in cross A medium to medium-large fern with a short rhizome and
section shows scattered vascular bundles. The foliage fronds bud-forming roots. Grows well under medium to high light
are erect and evenly forked several times into narrow seg- in well-drained, moist-dry to dry, coarse potting mix or
ments, with the soral patch on the ultimate fork. This spe- uncut moss. It is easy to cultivate and is a common but vari-
cies is native to eastern Africa and Madagascar. able species.
430 C 13

a d

Figure 13.99.5. Platycerium andinum: a. habit, bar = Figure 13.99.6. Platycerium bifurcatum: a. habits, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome 10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.). 1 cm (0.4 in.); e. stipe bases, bar = 2 cm (0.8 in.).

The base fronds of Platycerium bifurcatum typically have ations of the members of this complex. Some of these trade
the distal part slightly to moderately extended and shal- plants may be recognized as belonging to one of the four
lowly to moderately lobed. The foliage fronds are erect to species in the complex, whereas others are more difficult to
arching or pendent, typically with a short stipe that has the identify because of intergrading or inconstant characters.
vascular bundles arranged in a circle. The blades are several The particular frond form of a plant may not be the same in
times forked into narrow segments, and the sporangial later growth or under different cultural conditions. Fronds
patches are borne on the ultimate forks. Emerging fronds can vary by season and age.
soon turn to a horizontal position, instead of remaining Platycerium willinckii var. venosum Hughes is closer to P.
vertical for some distance as in the similar P. willinckii. The bifurcatum than to P. willinckii because of its strongly
species is native to eastern Australia. stalked (not sessile) foliage fronds. It differs from typical P.
The preceding description excludes Platycerium hillii, bifurcatum by having widely spreading base fronds, usually
P. veitchii, and P. willinckii, which are sometimes consid- longer foliage fronds, and strongly raised major veins on the
ered varieties or subspecies of P. bifurcatum but are here upper surface (but depressed on the lower) especially near
treated as separate species. Collectively, these species and the blade base. It was collected in northeastern Australia.
P. bifurcatum are referred to as the “P. bifurcatum com- Native var. venosum reportedly intergrades into typical P.
plex.” They all have a circular arrangement of vascular bun- bifurcatum plants (Roy Vail, personal communication).
dles in the stipes and lack dark tissue encircling the bundles. ‘Netherlands’. Fronds arching, grayish, the base fronds
(This trait can be seen by making a clean cut across the stipe swept upward, lobed on the upper margins. Common in
with a razor blade or sharp knife.) Most of the numerous, the trade.
loosely given Platycerium names in the trade represent vari-
 F  F A  C 431

Platycerium coronarium (König ex Müller) Desvaux Platycerium ×elemaria Hoshizaki & M. G. Price
 13.99.7  13.99.8
syn. Platycerium biforme (Swartz) Blume Tender
Disk staghorn A medium-large fern with a short rhizome and bud-form-
Tender ing roots. Grows well under medium to high-medium light
A large fern with a short rhizome that may branch and be- in drained, moist to moist-dry, coarse potting mix or uncut
come longer; new plants form when the tip of the branch moss.
reaches the surface. Grows well under high light in drained, Platycerium ×elemaria is a sterile hybrid of P. elephanto-
moist-dry, coarse potting mix or uncut moss. tis and P. stemaria. Its base fronds resemble those of the
The base fronds of Platycerium coronarium have the dis- parents, and the foliage fronds resemble those of P. elephan-
tal part extended, deeply lobed, and forward-arching. The totis but have the apical margins shallowly and irregularly
foliage fronds are long, pendent, and several times forked. forked or lobed. Hydathodes are absent. This plant origi-
The sporangial patch is typically a kidney-shaped lobe. This nated as a hybrid in horticulture and does not occur in the
species is native to southeastern Asia, Malaysia, Indonesia, wild. The cultivar name of this hybrid is ‘Sanchez’.
and the Philippines.

c
Figure 13.99.8. Platycerium ×elemaria: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.).

Platycerium elephantotis Schweinfurth

 13.99.9
syn. Platycerium angolense Wellwitsch ex Baker
d Angola staghorn
Tender
A medium-large fern with short rhizomes and bud-forming
Figure 13.99.7. Platycerium coronarium: a. habit, bar = roots. Grows well under medium to high-medium light in
10 cm (4 in.); b. sporangial area placement; c. rhizome drained, moist to moist-dry, coarse potting mix or uncut
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = moss. The plants quickly show water stress and are prone to
1 cm (0.4 in.). attack by scale insects.
The base fronds of Platycerium elephantotis are much ex-
tended, unlobed, and spreading slightly forward distally.
 432 C 13

Platycerium grande (Fée) Kunze  13.99.11

b c Tender
A large fern with short rhizomes. Grows under high light in
drained, moist-dry, coarse potting mix or uncut moss.
The base fronds of Platycerium grande have much-ex-
tended, deeply lobed, and usually forward-arching distal
parts. The foliage fronds have two sporangial patches per
frond, each typically borne on a semicircular extension jut-
ting out into the sinus of the second fork. The species is
d endemic to the Philippines.
Platycerium superbum is often misidentified in the trade
as P. grande, but it differs in having only one sporangial
patch per frond. The similar but smaller P. wallichii has
a outer branches that are shorter than the inner ones and
green instead of brown spores.
b
Figure 13.99.9. Platycerium elephantotis: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.).
a
c

The foliage fronds are spreading to pendent, asymmetri-

cally obovate, and entire; they have hydathodes. The spo-
rangial patch is apical. The species is native to Africa.

Platycerium ellisii Baker  13.99.10

syn. Platycerium diversifolium Bonaparte
Tender
A medium-sized fern with short rhizomes and bud-forming d
roots. Grows under high light in drained, moist to moist-
dry, coarse potting mix or uncut moss. The plants are Figure 13.99.11. Platycerium grande: a. habit, bar =
somewhat difficult to grow. 10 cm (4 in.); b. sporangial area placement; c. rhizome
Platycerium ellisii resembles P. alcicorne but has typical scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
foliage fronds usually shallowly once-forked at the tip. It is 1 cm (0.4 in.).
endemic to Madagascar.

b Platycerium hillii T. Moore  13.99.12

syn. Platycerium bifurcatum var. hillii (T. Moore) Domin
Green staghorn
Semi-tender
A medium-sized fern with short rhizomes and bud-forming
roots. Grows well under medium to high light in drained,
moist-dry, coarse potting mix or uncut sphagnum moss.
a This species is easy to grow.
Platycerium hillii has entire, round to kidney-shaped
c d base fronds. The wedge-shaped or fan-shaped foliage fronds
are erect with short stipes. They are several times forked,
Figure 13.99.10. Platycerium ellisii: a. habit, bar = 10 cm usually in the distal third of the frond. The forks are short
(4 in.); b. sporangial area placement; c. rhizome scales, and stout. The species is native to eastern Australia.
bar = 5 mm (0.2 in.); d. stipe cross section, bar = 1 cm This species is distinguished by the entire base fronds
(0.4 in.). and the short-stalked, erect foliaceous fronds that are two or
more times shallowly forked distally. All other members of
 F  F A  C 433

a
c

a c

d
Figure 13.99.12. Platycerium hillii: a. habit, bar = 10 cm
(4 in.); b. sporangial area placement; c. rhizome scales,
bar = 5 mm (0.2 in.); d. stipe cross section, bar = 1 cm
(0.4 in.).

the Platycerium bifurcatum complex have extended base

fronds and are shallowly to deeply lobed. See also the dis-
cussion under P. bifurcatum. Some of the choice cultivars of
P. hillii include ‘Mona Loa’, ‘Panama’, and ‘Talnadge’. d

Platycerium holttumii de Joncheere & Hennipman

 13.99.13
Tender
A large fern with short rhizomes. Grows under medium
light in drained, moist-dry, coarse potting mix or uncut Figure 13.99.13. Platycerium holttumii: a. habit, bar =
moss. 10 cm (4 in.); b. sporangial area placement; c. rhizome
Platycerium holttumii resembles P. grande, but its foliage scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
frond has the outer primary branch and its divisions short- 1 cm (0.4 in.).
ened. It is also resembles P. wandae, which differs by hav-
ing the outer primary branch wedge-shaped without any
further branching and by the presence of fringes on the base have denticulate margins and short stipes. The foliage
frond above or around the bud. Platycerium holttumii is na- fronds are wedge-shaped with the apical margins mostly
tive to southeastern Asia and the Malay Peninsula. shallowly forked one to three times, and the sporangial
patches occur in the apical part of frond. This species is en-
Platycerium madagascariense Baker demic to Madagascar.
 13.99.14 The trade plant circulating as ‘Horne’s Surprise’ is a
Tender presumed hybrid of Platycerium madagascariense and P.
A medium-sized fern with short rhizomes and bud-forming alcicorne.
roots. Grows under medium light in drained, moist-dry to
moist, coarse potting mix or uncut moss. This species is Platycerium ×mentelosii Hoshizaki  13.99.15
difficult to grow and has a dormant period. The base fronds Tender
are not thick and spongy and are loosely layered. There- A medium-large fern with short rhizomes. Grows well
fore, water drains readily from the plant and extra care must under medium light in drained, moist-dry, coarse potting
be taken to provide sufficient water. Control the many pests mix or uncut moss. Although a hybrid, this fern reportedly
that like to live between the loose layers of base fronds. produces some viable spores from which the plant can be
Platycerium madagascariense is distinctive for its waffle- propagated.
like depressions on the base fronds. By contrast, the main Platycerium ×mentelosii has characteristics intermediate
veins of P. alcicorne are raised, but not the cross-veins. The between those of its parents, P. stemaria and P. superbum.
base fronds of P. madagascariense are thick and papery The stipe cross section has the dark band of the P. superbum
when dry (not spongy as in P. alcicorne) and sometimes parent and the scattered vascular bundles of P. stemaria.
434 C 13

This sterile hybrid originated in horticulture; it does not

b
occur in the wild. The cultivar name for this hybrid is ‘Fan-
a tastic Gardens’.

Platycerium quadridichotomum (Bonaparte) Tardieu

 13.99.16
Tender
A medium-sized fern with a short rhizome and bud-form-
ing roots. Grows under high-medium light in drained,
c moist-dry to dry, coarse potting mix or uncut moss. This
species is difficult to grow and prone to fungal attacks. The
plants have a distinct dormant period.
Platycerium quadridichotomum is extended, irregularly
lobed, and foliaceous in the distal part of the base fronds.
The foliage fronds are spreading to pendent, several times
e
forked, and up to 20–50 cm (8–20 in.) long. The sporan-
d gia are borne on strap-shaped branches and typically oc-
cupy a medial position on the frond. Platycerium quadridi-
chotomum is endemic to Madagascar.
Figure 13.99.14. Platycerium madagascariense: a. habit,
Platycerium andinum has the same general structure but
bar = 10 cm (4 in.); b. sporangial area placement;
is a much larger plant with long, thick, densely hairy fo-
c. rhizome scales, bar = 5 mm (0.2 in.); d. stipe cross
liage fronds.
section, bar = 1 cm (0.4 in.); e. depressions between
veins of base frond and minutely dentate margin, bar =
b
5 cm (2 in.).

a c

d
b

a Figure 13.99.16. Platycerium quadridichotomum:

a. habit, bar = 10 cm (4 in.); b. sporangial area
placement; c. rhizome scales, bar = 5 mm (0.2 in.);
d. stipe cross section, bar = 1 cm (0.4 in.).

c
Platycerium ridleyi H. Christ  13.99.17
Tender
A medium-sized fern with short rhizomes. Grows under
high light in drained, moist-dry to dry, coarse potting mix
or uncut moss. This species is difficult to grow and is at-
tacked by a variety of insect pests.
Platycerium ridleyi has round to kidney-shaped base
fronds with unlobed or coarsely dentate margins. The fo-
liage fronds are erect, and the sporangial patch is borne on
an obovate lobe. This species is native to western Malaya
Figure 13.99.15. Platycerium ×mentelosii: a. habit, bar = and Indonesia.
10 cm (4 in.); b. sporangial area placement; c. stipe cross
section, bar = 1 cm (0.4 in.).
 F  F A  C 435

sinus of the second fork. The typical variety, var. stemaria,

b
is native to central and western Africa.
a var. laurentii de Wilde. Differs from the typical variety
by having irregularly dentate or laciniate apical margins on
the broad primary fork. It is native to Africa.

Platycerium superbum de Joncheere & Hennipman

 13.99.19;  43
syn. Platycerium grande var. tambourinense Domin
Giant staghorn
c Semi-tender
A large fern with short rhizomes. Grows well under me-
dium or high light in drained, moist-dry to dry, coarse pot-
d ting mix or uncut moss. This species is a choice but
extremely large fern that may be grown outdoors in sub-
tropical climates.
Figure 13.99.17. Platycerium ridleyi: a. habit, bar = In Platycerium superbum the distal part of the base fronds
10 cm (4 in.); b. sporangial area placement; c. rhizome are much extended, deeply lobed, and usually forward-
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = arching. The foliage fronds are several times forked. One
1 cm (0.4 in.). sporangial patch is produced per frond, typically on a semi-
circular extension jutting out into the sinus of the first fork.
The species is native to Australia.
Platycerium stemaria (Beauvaux) Desvaux
 13.99.18 Platycerium veitchii (Underwood) C. Christensen
Triangular staghorn  13.99.20
Tender syn. Platycerium bifurcatum subsp. veitchii (Underwood)
Hennipman & Roos
A medium-large fern with a short rhizome and bud-form-
Semi-tender
ing roots. Grows well under medium to high-medium light
in drained, moist to moist-dry, coarse potting mix or uncut A medium-sized fern with short, ascending rhizomes and
moss. The plants quickly show water stress. bud-forming roots. Grows well under medium to high light
The base fronds of Platycerium stemaria have the upper
margin much extended, unlobed, and spreading slightly b
forward. The spreading to pendent foliage fronds are twice-
forked, with the sporangial patches typically below the

b a

a d
d

Figure 13.99.18. Platycerium stemaria: a. habit, bar = Figure 13.99.19. Platycerium superbum: a. habit, bar =
10 cm (4 in.); b. sporangial area placement and frond 10 cm (4 in.); b. sporangial area placement; c. rhizome
variations; c. rhizome scales, bar = 5 mm (0.2 in.); scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
d. stipe cross section, bar = 1 cm (0.4 in.). 1 cm (0.4)in.
 436 C 13

hibited at the 1997 International Fern Society show in Los

Angeles produced an offshoot—something that rarely hap-
b pens with this species in cultivation. It is uncertain whether
c the buds came from a root or from a rhizome branch.
Platycerium wallichii has base fronds that are extended
and deeply lobed distally. The foliage fronds are several
times forked, with the outer primary fork shorter, and typ-
a ically two, sometimes three, sporangial patches are usually
d below the sinuses of the second fork. This species is native
to eastern India, Burma, Thailand, and China.

Platycerium wandae Raciborski  13.99.22

Figure 13.99.20. Platycerium veitchii: a. habit, bar =
syn. Platycerium willhelminae-reginae van Alderwerelt
10 cm (4 in.); b. sporangial area placement; c. rhizome
van Rosenburgh
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
Queen staghorn
1 cm (0.4 in.).
Tender
A large fern with short rhizomes. Grows well under high-
in drained, moist-dry to dry, coarse potting mix or uncut medium light in moist-dry, coarse potting mix or uncut
moss. The plants tolerate considerable heat. The whitish moss. This is the largest Platycerium species, and plants may
gray foliage and lobes on the base fronds develop best in spread to 2 m (6.5 ft.) across.
high light.
Platycerium veitchii resembles P. bifurcatum but differs
by the base frond typically extended on the upper margin
into an erect, narrow, deep lobe that browns readily, and
the foliage fronds are more erect, densely hairy, and gray-
green. When the narrow lobes of the base frond do not de-
velop, the plant may be difficult to distinguish from P. bi-
furcatum. It is endemic to Australia.
b
Platycerium wallichii Hooker  13.99.21 a
Indian staghorn
Tender
A medium-large fern with short rhizomes. Grows under
high-medium to high light in moist to moist-dry, coarse
potting mix or uncut moss. This species is difficult to cul-
tivate, and new growth is seasonal. A mature specimen ex-

a
b c

d
c d

Figure 13.99.21. Platycerium wallichii: a. habit, bar = Figure 13.99.22. Platycerium wandae: a. habit, bar =
10 cm (4 in.); b. sporangial area placement; c. rhizome 10 cm (4 in.); b. sporangial area placement; c. rhizome
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar =
1 cm (0.4 in.). 1 cm (0.4 in.).
 F  F A  C 437

Platycerium wandae resembles P. holttumii, but the mar- Polybotrya

gins of the base fronds around the growing point or above The rhizomes of these rarely cultivated tropical ferns be-
it have small fringes. The outer branch of the foliage fronds come long and scandent and may climb tree trunks up to
is reduced to a wedge. There are two sporangial patches, over 8 m (26 ft.) high. Because they are tender plants with
one on the wedge and the other on the inner primary long-creeping rhizomes, their use in the landscape is limited
branch (around the sinus of the next fork). This species is to tropical climates or large greenhouses, where they are
endemic to New Guinea. usually trained on tree trunks. Polybotrya is difficult to
maintain in baskets because of the long-creeping, scandent
Platycerium willinckii T. Moore habit.
 13.99.1, 2, 23 Polybotrya is characterized by long-creeping, densely
syn. Platycerium bifurcatum subsp. willinckii (T. Moore) scaly rhizomes and strongly dimorphic sterile and fertile
Hennipman & Roos, P. sumbawense H. Christ fronds. The apex of the blade is gradually tapered, not with
Java staghorn a terminal pinna resembling the lateral ones. The fertile
Tender to semi-tender fronds resemble a highly dissected, skeletonized version of
A medium-large fern with short rhizomes and bud-forming the sterile ones, and the sori always lack indusia.
roots. Grows well under high-medium light in moist-dry, The genus is native to the New World tropics and con-
coarse potting mix or uncut moss. This species is easy to tains 35 species. Its name comes from the Greek poly,
grow if protected from cold. The leaves are thick and gray- many, and botrys, bunch or cluster, alluding to the sporan-
green. gia clustered on the fertile leaves. For a technical account of
The distal part of the base fronds of Platycerium wil- the species, see Moran (1987).
linckii are extended and swept upward, deeply lobed, and
arched forward. The foliage fronds are long and pendent, Polybotrya osmundacea Humboldt & Bonpland ex
typically sessile, and several times forked, usually with the Willdenow  13.100.1
most forking in the distal part of the frond. Platycerium wil- Tender
linckii is native to Java and the Lesser Sunda Islands.
A medium-large fern with long-creeping rhizomes that can
Platycerium bifurcatum differs in having shorter, less-
climb to about 8 m (26 ft.). Grows well under high-me-
pendent foliage fronds, short stipes, and emerging fronds
dium light in moist garden soil or potting mix.
that are more horizontal, rather than remaining in a verti-
Polybotrya osmundacea has roughly triangular, two- to
cal position for some time and distance.
three-pinnate-pinnatifid sterile fronds with free veins. The
fertile fronds are contracted and have hardly any green tis-
sue. The sporangia form linear or oblong sori. The fertile
a fronds are only produced on the climbing portion of
b the rhizome. The species is widespread in the New World
tropics.

Polypodium  13.101.1, 2

Polypody
The common polypodies of genus Polypodium are probably
the most widely known ferns. They occur throughout the
world and are frequently depicted in designs and men-
tioned in folklore of Western cultures. In gardens they can
be used among rocks, on walls, or in pots or baskets. The
c larger species are usually grown in hanging baskets or pots.
Most polypodies are tender. Some can be grown in the
ground in warm climates if given good drainage. Certain
species make suitable indoor plants. The spores may be
slow to germinate but give many sporelings.
e The genus Polypodium can be identified by its long-
creeping rhizomes with two ranks of fronds on the dorsal
d
surface. The fronds are jointed to the rhizome and fall away
Figure 13.99.23. Platycerium willinckii: a. habit, bar = cleanly to leave a low, elevated base (the phyllopodium).
10 cm (4 in.); b. sporangial area placement; c. rhizome The blades are typically pinnatisect to one-pinnate (rarely
scales, bar = 5 mm (0.2 in.); d. stipe cross section, bar = more finely divided) with pinnae continuous with, not
1 cm (0.4 in.); e. stipe bases, bar = 2 cm (0.8 in.). jointed to, the rachis. The large, round sori lack indusia.
438 C 13

Figure 13.101.1. Polypodium cambricum: habit.

e
d

Figure 13.100.1. Polybotrya osmundacea: a. sterile frond

from immature plant, bar = 5 cm (2 in.); b. apical
portion of fertile frond, bar = 5 cm (2 in.); c. two fertile
pinnules (tertiary), bar = 5 mm (0.2 in.); d. sterile pinna
from mature plant, bar = 5 cm (2 in.); e. rhizome and
stipe from mature plant, bar = 5 cm (2 in.).

The spores are yellow, at least when fresh. No other fern

genus has this combination of characteristics.
Polypodium consists of about 125 species worldwide,
with most of the species in the American tropics. The genus
name comes from the Greek poly, many, and podion, foot, Figure 13.101.2. Polypodium cambricum: frond apex and
alluding to the resemblance of the rhizome’s numerous sori.
branches to feet. For more information on this and related
genera, see “Polypodium and Relatives” in Chapter 10.
Early pteridologists placed any fern with round, non-
indusiate sori in Polypodium. Defined as such, Polypodium
included species in unrelated genera such as Ctenitis, Dryop-
teris, and Thelypteris. Nowadays Polypodium is subdivided
into many smaller genera, and these are compared here.
The comparisons are only for the cultivated species and do
not cover all the species within the genus.
 F  F A  C 439

Campyloneurum: Rhizomes short- veins pointing toward the margins. Native to Africa and
creeping; rhizome scales clathrate; Asia. (Lepisorus longifolius, L. thunbergianus)
fronds medium-sized, simple, en-
tire, strap-shaped or linear-lance- Leptochilus (Colysis): Rhizomes
olate, monomorphic; veins net- short- to long-creeping; rhizome
ted, lateral veins parallel and scales peltate, clathrate; fronds
connected by curved cross-veins medium-sized; stipes often
to form the areoles, the included winged; blades simple to one-
veinlet unbranched and pointing pinnate, monomorphic or di-
toward the margin; sori round, morphic; veins netted, included
borne below the free veinlet tip. veinlet pointing in various direc-
Native to tropical and subtropical tions; similar to Microsorum ex-
America. (Campyloneurum angustifolium, C. brevifolium, cept sori are mostly elongate to
C. costatum, C. phyllitidis, C. tenuipes, C. xalapense) linear and oblique to the axis.
Native to Asia. (Leptochilus amplus, L. ellipticus, L. macro-
Dictymia: Rhizomes short-creep- phyllus var. wrightii )
ing; rhizome scales peltate, broadly
ovate, clathrate, usually with glan- Microgramma: Rhizomes long-creeping, often
dular marginal projections; fronds slightly flattened; rhizome scales elongate, not
medium-sized, simple, entire, lin- clathrate, attached well above their base; fronds
ear to lanceolate, subleathery, less than 39 cm (15 in.) long; blades simple, en-
smooth; veins netted, included tire, lanceolate to oblong, slightly dimorphic;
veinlets absent or rare; sori round veins netted with unbranched included veinlets
to narrowly oblong, slightly to pointing outward toward the margin; sori round.
strongly embossed on the blade. Native to Australia and Native to the American tropics and subtropics.
the islands of the southwestern Pacific. (Dictymia brownii ) (Microgramma heterophylla, M. lycopodioides, M.
megalophylla, M. nitida, M. percussa, M. pilosel-
Goniophlebium: Rhizomes medium- loides, M. reptans, M. squamulosa, M. vacciniifolia)
to long-creeping; rhizome scales pel-
tate, clathrate, toothed, often with Microsorum (Phymatosorus):
one- or two-celled, glandular marginal Rhizomes short- to long-
appendages; fronds medium- to large- creeping; fronds medium- to
sized; blades pinnatifid to pinnate large-sized, simple, margins
with a terminal pinna resembling the entire, lobed, or pinnatifid;
lateral ones; pinnae jointed; veins net- veins netted, included vein-
ted, areoles angular and in several lets simple or branched, vari-
rows, costal areoles large, with one included veinlet point- ously directed, always ending
ing outward. Native to Asia. (Goniophlebium ×ekstrandii, in a hydathode; sori scattered
G. percussum, G. persicifolium, G. subauriculatum) dots or larger, round to oval, immersed in the blade tissue
or not. Native to Africa, Asia, Australia, and islands of the
Lecanopteris: Rhizomes short-creep- southwestern Pacific. (Microsorum commutatum, M. for-
ing and swollen, with internal tunei, M. grossum, M. insigne, M. linguiforme, M. longissi-
chambers inhabited by ants; rhi- mum, M. lucidum, M. membranifolium, M. musifolium,
zome scales peltate, clathrate cen- M. pteropus, M. punctatum,
trally but not at the margin; fronds M. pustulatum, M. samarense,
small-sized; blades entire to deeply M. scandens, M. scolopendria,
pinnatifid; veins netted, the areoles M. steerei, M. superficiale, M.
with included free veinlets. Native viellardii)
to Malaysia. (Lecanopteris car-
nosa, L. crustacea, L. sinuosa) Niphidium: Rhizomes creep-
ing to ascending; fronds clus-
Lepisorus: Resembles Pleopeltis, tered, medium- to large-
except the rhizome scales are en- sized, simple, entire, upper
tirely clathrate and the outer row surfaces often have many
of areoles do not produce free white dots; sori round, large
440 C 13

(3–5 mm in diameter), in a single row between the main paraphyses (if present) not peltate. Cosmopolitan. (Polypo-
lateral veins, not sunken; sporangial capsules with minute dium amoenum, P. appalachianum, P. attenuatum, P. bom-
hairs. Native to the American tropics and subtropics. bycinum, P. brasiliense, P. calirhiza, P. cambricum, P. dis-
(Niphidium crassifolium) simile, P. formosanum, P. glycyrrhiza, P. guttatum, P.
hesperium, P. hirsutissimum, P. interjectum, P. ×leucospo-
Pecluma: Rhizomes very short-creep- rum, P. levigatum, P. loriceum, P. maritimum, P. menisci-
ing; rhizome scales attached along ifolium, P. pellucidum, P. plebeium, P. polypodioides, P.
their base; fronds small- to medium- ptilorhizon, P. pyrrholepis, P. rhodopleuron, P. sanctae-rosae,
sized; blades deeply pinnatisect, comb- P. scouleri, P. thyssanolepis, P. triseriale, P. virginianum, P.
shaped, the many divisions narrow vulgare)
and close together, with small one-
celled hairs on lower surfaces; sori Selliguea (Crypsinus): Rhizomes
round. Native to the American tropics long-creeping, conspicuous
and subtropics. (Pecluma dispersa, P. pectinata, P. plumula) black fiber strands within; rhi-
zome scales peltate; fronds me-
Phlebodium: Rhi- dium-sized, simple, entire, lan-
zomes medium- to ceolate to oblong-lanceolate,
long-creeping; rhi- some species (formerly in Crypsi-
zome scales usually nus) hastate, pinnatifid to pin-
orangish when freshly nate with cartilaginous and often
mature, not clathrate; notched margins; fertile fronds
fronds medium- to taller, more contracted; veins
large-sized, pinnati- netted, lateral veins conspicuous; sori brown, round but
lobed or pinnatifid; usually elongate or linear, borne between adjacent lateral
areoles next to the midrib mostly narrow and without in- veins in one or two rows. Native to southern Africa, Asia,
cluded veinlets; other areoles with included veinlets or not, Australia, and the islands of the Pacific. (Selliguea feei)
if present the veinlets are united or, if free, usually pointing
outward toward the margin; sori supplied by two veinlets. Solanopteris: Rhizomes long-creep-
Native to the American tropics and subtropics. (Phlebodium ing, with ant-inhabited tubers,
aureum, P. decumanum, P. pseudoaureum, P. ×schneideri) without black fiber stands within;
rhizome scales peltate, round;
Pleopeltis: Rhizomes long-creeping; rhi- fronds small-sized, simple to pin-
zome scales clathrate centrally but not at natifid; fertile fronds usually nar-
the margins; fronds small- to medium- rower and longer than the sterile;
sized, simple, entire (rarely dichoto- veins netted with included veinlets;
mously forked), surfaces with peltate sori round or linear. Native to the
scales; veins netted with included American tropics. (Solanopteris
branched(?) veinlets pointing in various bifrons, S. brunei)
directions, the marginal row of areoles
with free veins pointing toward Even with the removal of these
the margins; sori usually round, segregate genera, Polypodium con-
covered with peltate paraphyses tains many species. To facilitate identification of these spe-
when young. Native to the Amer- cies, Polypodium is divided into three groups.
ican tropics and subtropics, and
Africa. (No cultivated species; The Vulgare Group (Polypodium vulgare complex). This
plants brought into cultivation group is characterized by free veins (rarely with a few netted
have not survived.) ones) and pinnatifid (or pinnatilobed) blades with scaleless
surfaces. The sori are always in one row between the mid-
Polypodium: Rhizomes short- to rib and margin of the blade. The vulgare group contains
long-creeping; fronds small to about 25 species, most of which are native to the boreal and
large; blade pinnatifid or pinnate; temperate regions of North America, Europe, and Asia. A
pinnae not jointed; veins free or few occur in Mexico, and one species (Polypodium pellu-
netted, with or without a free in- cidum) is native to Hawaii.
cluded veinlet; sori round or ob- The most commonly cultivated species in the group are
long, at tip or junction of vein, of European origin. Other species may be cultivated but
 F  F A  C 441

are usually difficult to establish. Pteridologists have named Polypodium amoenum Wallich ex Mettenius
and described many species and hybrids that may be diffi-  13.101.3
cult for nonspecialists to identify, especially if the plant’s syn. Goniophlebium amoenum (Wallich ex Mettenius) J.
original habitat is unknown. Even if this is known, fertile Smith ex Hooker & Greville
fronds and a microscope are usually needed for accurate Semi-tender to hardier
identification. A small to medium fern with long-creeping rhizomes.
The cultivated species of the vulgare group treated here Grows well under low to high light in moist-dry, well-
include Polypodium appalachianum, P. calirhiza, P. cambri- drained garden soil or potting mix. The plants are subject
cum, P. glycyrrhiza, P. hesperium, P. interjectum, P. pelluci- to slug and snail attack and require a lot of humidity.
dum, P. scouleri, P. virginianum, and P. vulgare. They are The rhizomes of Polypodium amoenum contain scattered
illustrated together in Figure 13.101.34. dark fibers, and the fronds are generally 14–50 cm (5.5–20
The Loriceum Group (Polygoniophlebium). This group is in.) long and 8–27 cm (3–11 in.) wide, with pinnatifid,
characterized by pinnatifid to pinnate, rarely simple blades dull green blades. The segments typically have pentagonal
that have scaleless surfaces (a few scales might be present areoles next to the rachis. The sori are in one row between
on the rachis or costae). The veins form at least one row of the pinna midrib and margin. The species is native to the
regular areoles along the pinna midrib. Each areole con- Himalayas, southeastern Asia, China, and Taiwan.
tains a veinlet that points toward the blade margin. The Some pteridologists classify this species in Goniophle-
pinnae are continuous with the rachis, not jointed to it by bium, but it lacks jointed pinnae (a defining characteristic
an abscission layer. The sori appear in one to several rows of that genus) and is therefore retained in Polypodium until
between the costae and margins. The group contains about further study.
40 species, most of which are native to the New World.
Two widely grown species in the loriceum group are Polypodium appalachianum Haufler & Windham
Polypodium triseriale and P. menisciifolium. The former is  13.101.4, 34d
grown as a basket plant in gardens in Florida. Polypodium Appalachian polypody
menisciifolium, which is cultivated in baskets in southern Very hardy, Zone 3
California, grows vigorously and tolerates irregular water- A small fern with short- to medium-creeping rhizomes.
ing. Another species in the group, P. formosanum, is often Grows under medium light in drained, moist potting mix.
found in the trade and is prized for its attractive pale gray- This species adapts poorly to gardens in southern California.
ish green fronds and rhizomes. A cultivar with repeatedly
forked and congested rhizomes, ‘Cristatum’, adds further
interest to this species.
The cultivated species treated here are Polypodium amoe- b
num, P. attenuatum, P. brasiliense, P. dissimile, P. formo-
sanum, P. levigatum, P. loriceum, P. maritimum, P. meniscii-
folium, P. ptilorhizon, P. rhodopleuron, and P. triseriale.
The Scaly-Polypody Group. This group is characterized
by pinnatifid (or pinnatilobed) blades that are scaly, espe-
cially on the lower surface. Often the scales are dense and
conspicuous. The veins vary from free to netted, but they d
are usually hard to see because of the thick blade tissue or
because they are hidden by the scales. The sori are in one c
row between the costa and margin. All the species in the
scaly-polypody group, of which there are about 40, are na-
tive to tropical America. a
This group is closely related to Pleopeltis, and probably
all the species should be transferred to that genus, as has al-
ready been done with some species. The cultivated species
treated here are Polypodium bombycinum, P. guttatum, P.
hirsutissimum, P. ×leucosporum, P. plebeium, P. polypodi-
oides, P. pyrrholepis, P. sanctae-rosae, and P. thyssanolepis. Figure 13.101.3. Polypodium amoenum: a. habit, bar =
5 cm (2 in.); b. vein details along the rachis, bar = 1 cm
(0.4 in.), after Hoshizaki (1982); c. rhizome cross
section, showing vascular bundles and scattered fibers,
bar = 1 cm (0.4 in.); d. rhizome scale, bar = 1 mm.
 442 C 13

Polypodium attenuatum resembles P. triseriale but dif-

fers by deeply pinnatisect blades, falcate pinnae, a single
b
row of sori between the pinna midrib and margin, and a
dense mass of hairy, brown roots around the rhizome. The
species is native to lowland forests in tropical America.

Polypodium bombycinum Maxon  13.101.6

Tender
A medium-sized fern with short-creeping rhizomes. Grows
well under medium light in moist-dry, well-drained pot-
Figure 13.101.4. ting mix or uncut moss.
Polypodium Polypodium bombycinum has short-creeping, branched
appalachianum: rhizomes that are 3–5 mm (0.1–0.2 in.) thick and covered
a. habit, bar = 5 cm by rigid, dark brown scales. The stipes are short and close
a (2 in.); b. paraphysis, together. The blades are stiff, erect, and linear to narrowly
top view, bar = elliptic or oblanceolate, tapering in the lower half with basal
0.1 mm. pinnae reduced to small lobes. Appressed scales are so
abundant that they hide the lower blade surface. A small,
slightly swollen gland is found on the inside upper edge of
Polypodium appalachianum was recently separated from the segments, although this trait is often hard to see. The
P. virginianum and is distinguished from it by the elongate- species grows as an epiphyte in the tropical forests of Cen-
triangular blades and golden-brown rhizome scales nearly tral and South America.
of all one color. The species is native to eastern North
America. Polypodium brasiliense Poiret  13.101.7
Tender
Polypodium attenuatum Humboldt & Bonpland ex A large fern with short- to medium-creeping rhizomes.
Willdenow  13.101.5 Grows well under medium light in moist, drained potting
syn. Polypodium kuhnii Fournier mix.
Tender Polypodium brasiliense resembles P. triseriale but differs
A medium-sized fern with a short-creeping rhizome sur- by its larger size, pinnae with five to six series of areoles be-
rounded by a dense root mass. Grows well under medium
light in moist, drained potting mix.

a
c
b

a b

Figure 13.101.6. Polypodium bombycinum: a. fronds, bar

Figure 13.101.5. Polypodium attenuatum: a. frond, bar = = 5 cm (2 in.); b. scale from lower surface of blade, bar =
5 cm (2 in.); b. segment with sori, bar = 5 cm (2 in.); 1 mm, after Sota (1966); c. rhizome scale, bar = 1 mm,
c. rhizome scales, bar = 1 mm. after Sota (1966).
 F  F A  C 443

a
Figure 13.101.8. Polypodium
calirhiza: a. habit, bar = 5 cm
(2 in.); b. base of segment with
one areole, bar = 5 mm (0.2 in.).

Figure 13.101.7. Polypodium brasiliense: a. frond, bar = Polypodium cambricum Linnaeus

5 cm (2 in.); b. venation and sori, bar = 1 cm (0.4 in.);  13.101.1, 2, 9, 34a
c. rhizome scale, bar = 1 mm. After Hoshizaki (1982). syn. Polypodium australe Fée
Southern polypody
Semi-hardy to hardy
A small fern with short- to medium-creeping rhizomes.
tween the pinna midribs and margins, slightly scaly pinna Grows well under medium light in moist, drained, basic
midribs, and long-attenuate, sparsely and irregularly fim- garden soil or potting mix. The plants produce new fronds
briate rhizome scales. It is native to tropical America. in late summer.
‘Cristata’. See Polypodium triseriale ‘Cristatum’. Polypodium cambricum is characterized by broadly
ovate, pinnatifid blades, with the longest pair of pinnae
Polypodium calirhiza S. A. Whitmore & A. R. Smith often second from the base. Oval sori have paraphyses con-
 13.101.8, 34i sisting of branched hairs. It is native to Europe.
Semi-hardy, Zone 8 Polypodium cambricum, a member of the P. vulgare
A small fern with short- to medium-creeping rhizomes. complex, has long been considered a fringed (two-pinnati-
Grows well under medium light in drained, moist to moist- fid) variant of P. australe. Because P. cambricum is an earlier
dry garden soil or potting mix. The rhizome has an acrid or name, however, it must be used instead of the name P. aus-
slightly sweet flavor. trale. The unfringed form is typically found in nature (Fig-
Polypodium calirhiza has pinnatifid blades that are wid- ure 13.101.9a).
est above the base, with one or no areoles on each segment. Most of the English Polypodium cultivars belong to P.
The upper surface of the rachis has minute hairs, the lower cambricum, but they are uncommon in the United States
surface with lanceolate to ovate scales. This plant is a fertile trade and tend to grow slowly. The following are some cul-
hybrid of Polypodium glycyrrhiza and P. californicum Kaul- tivars that may be encountered in the United States:
fuss (Figure 13.101.34g). It is native to California and ‘Cambricum’ (var. cambricum). Figure 13.101.9b.
Mexico and is grown in gardens on the West Coast of the Fronds two-pinnatifid, sterile, never producing sori. Some
United States. plants circulating in the trade as ‘Cambricum’ are large,
tender plants belonging either to Microsorum commutatum
‘Southern Cross’ or Polypodium triseriale ‘Cambrioides’.
 444 C 13

Figure 13.101.10. Polypodium

dissimile: habit, bar = 5 cm
b (2 in.).

Figure 13.101.9. Polypodium cambricum: a. habit of

species, bar = 5 cm (2 in.); b. frond of ‘Cambricum’;
c. paraphysis (left) and sporangium (right), bar =
0.5 mm, after Shivas (1962).

The rhizomes of Polypodium formosanum are 7–8 mm

‘Cornubiense’. Fronds nearly three-pinnatifid, segments (0.3 in.) thick, conspicuously glaucous, and contain scat-
linear. tered dark fiber strands (seen in cross section). The pinnati-
‘Pulcherrimum’. Resembles ‘Cambricum’, but with fid, light gray-green blades are hairy on the margins and
blades to three-pinnatifid and fully fertile. both surfaces of the rachis. This species is native to south-
ern China, Japan, and Taiwan.
Polypodium dissimile Linnaeus  13.101.10 Some pteridologists classify this species in Goniophle-
syn. Polypodium chnoodes Sprengel bium, but it lacks the jointed pinnae characteristic of that
Tender genus. For this reason it is left in Polypodium pending fur-
A medium-sized fern with medium-creeping rhizomes. ther study.
Grows well under medium light in moist, drained potting ‘Cristatum’. Rhizomes forked many times and con-
mix. The fronds are arching, herbaceous, and gray-green. gested.
Polypodium dissimile is easily recognized by its distinctly
clathrate rhizome scales, hairy blades (both surfaces), and Polypodium glycyrrhiza D. C. Eaton
two to four rows of areoles between the pinna midrib and  13.101.12, 34h
blade margin. The lower pinnae are bent downward and syn. Polypodium vulgare var. occidentale Hooker
clasp the rachis. This species is native to tropical America. Licorice fern
Hardy, Zone 5(6)
Polypodium formosanum Baker A small fern with short- to medium-creeping rhizomes.
 13.101.11;  44 Grows well under medium light in moist, drained potting
syn. Goniophlebium formosanum (Baker) Rödl-Linder mix or uncut moss. This species is easy to cultivate and
Semi-tender to hardier often volunteers in the bales of green moss used to line
A small to medium fern with medium- to long-creeping hanging baskets.
rhizomes. Grows well under medium light in drained, The rhizomes of Polypodium glycyrrhiza have a distinct,
moist-dry garden soil or potting mix. The new fronds sweet, licorice taste. The upper surface of the rachis has
emerge at the end of the summer, and old fronds are shed minute hairs and the lower surface has linear or hair-like
in late spring. scales. The pinnae shape varies (a rock form has rounder
 F  F A  C 445

tips), and the sori vary from round to oval and lack para-
physes. The species is native to Asia and the western coast
of North America.

Polypodium guttatum Maxon  13.101.13

Semi-tender, Zone 9
b A small fern with short- to medium-creeping rhizomes.
Grows well under medium light in moist-dry, drained pot-
ting mix.
Polypodium guttatum lacks scales on the upper surface
of its blade, but scales are sparsely scattered on the lower
surface, imparting a speckled appearance. This species is
native to Mexico.

a a

Figure 13.101.11. Polypodium formosanum: a. habit, bar

c
= 5 cm (2 in.); b. vein details and hairs along the rachis,
bar = 1 cm (0.4 in.); c. rhizome with rhizome scales, bar
= 1 cm (0.4 in.).

Figure 13.101.12. Polypodium

glycyrrhiza: frond variations,
bar = 5 cm (2 in.), after
Hoshizaki (1982). Figure 13.101.13. Polypodium guttatum: a. habit, bar =
5 cm (2 in.); b. segment underside with scales, bar =
1 cm (0.4 in.); c. scale from lower blade surface, bar =
1 mm. Courtesy of Knobloch and Correll (1962).

Polypodium hesperium Maxon

 13.101.14, 34k
Western polypody
Very hardy, Zone (3)4
A small fern with short- to medium-creeping rhizomes.
Grows under medium light in moist-dry, drained potting
mix. This species bears evergreen fronds. It is difficult to
grow and doubtfully in cultivation.
Polypodium hesperium has pinnatilobed blades, and the
rachises are smooth above but bear linear-lanceolate scales
below. The oval sori lack paraphyses. It is a fertile hybrid of
Polypodium amorphum Suksdorf (Figure 13.101.34j) and
P. glycyrrhiza and is native to western North America from
Canada to Mexico.
 446 C 13

Polypodium interjectum Shivas

 13.101.16, 34c
Intermediate polypody
Hardy, Zone 5
A small fern with short- to medium-creeping rhizomes.
Figure 13.101.14. Polypodium hesperium: Grows well under medium light in drained, moist to moist-
habit, bar = 5 cm (2 in.). dry garden soil, potting mix, or uncut moss. This species is
easy to grow and produces new fronds in the late summer
and autumn.
Polypodium interjectum has firm, ovate, pinnatifid
fronds, often with longest pinnae in the fourth to sixth pair
from the base. The young sori are oval and lack paraphyses.
Polypodium hirsutissimum Raddi  13.101.15 The species is native to Europe.
Semi-hardy (?) This member of the Polypodium vulgare complex is a
fertile hybrid of P. cambricum and P. vulgare.
A medium-sized fern with short- to medium-creeping rhi-
zomes. Grows well under medium light in moist-dry,
drained potting mix or uncut moss.
b
Polypodium hirsutissimum has densely scaly, narrowly
oblong, pinnatisect blades. On the inside upper edge of the
segments is a small, slightly swollen gland. This species is
native to South America, where it grows epiphytically in
wet forests.

Figure 13.101.16.
Polypodium interjectum:
a. frond variations, bar
= 5 cm (2 in.);
b. sporangium, bar =
0.5 mm, after Shivas
(1962).

a b
a

Polypodium ×leucosporum Klotzsch

 13.101.17
syn. Pleopeltis ×leucospora (Klotzsch) E. G. Andrews,
×Pleopodium leucosporum (Klotzsch) Mickel & Beitel
d Semi-tender or hardier
A small-medium fern with medium-creeping rhizomes.
Grows well under medium light in drained, moist-dry pot-
ting mix.
c The ovate-elliptic fronds of Polypodium ×leucosporum
are deeply and coarsely lobed nearly to the midrib, with the
lower lobes sometimes much reduced. The sori are pro-
Figure 13.101.15. Polypodium hirsutissimum: a. frond, duced in a medial row. This fern is native to the American
bar = 5 cm (2 in.); b. scale from lower surface of blade, tropics.
bar = 1 mm, after Sota (1966); c. rhizome scale, bar = Polypodium ×leucosporum is a sterile hybrid of Pleopeltis
1 mm, after Sota (1966); d. scale from around the sorus, macrocarpa (Bory ex Willdenow) Kaulfuss and Polypodium
bar = 0.25 mm, after Sota (1966). thyssanolepis A. Braun ex Klotzsch. It was studied in detail
 F  F A  C 447

by Wagner and Wagner (1975). The spores are misshapen, Polypodium loriceum Linnaeus  13.101.19
irregular, and aborted. Sterile hybrids such as this one are Tender
one reason pteridologists believe that the scaly-polypody A small-medium fern with medium-creeping rhizomes.
group is more closely related to Pleopeltis than to other Grows well under medium light in drained, moist potting
groups of Polypodium. mix.
Polypodium loriceum resembles P. maritimum, but its
Polypodium levigatum Cavanilles  13.101.18 rhizome scales are less distinctly clathrate, its blades are
syn. Polypodium glaucophyllum Kunze ex Klotzsch glabrous, and only one row of sori is found between the
Tender pinna midrib and margin. This species is widespread in the
A small fern with creeping, gray-green rhizomes bearing a American tropics.
few minute, scattered scales. Grows in medium light in As presently defined, Polypodium loriceum is extremely
moist to moist-dry, well-drained potting soil or uncut moss. variable and includes several geographic entities that prob-
The simple and entire fronds of Polypodium levigatum ably represent different species. This matter, however,
are unusual. They are gray-green, elliptic-lanceolate, and needs further research.
smooth except for a few ovate-acuminate scales on the
lower surface of the rachis. The veins are netted. It is native
from Central America to South America.

Figure 13.101.17. Polypodium

×leucosporum: habit, bar =
5 cm (2 in.).

Figure 13.101.19. Polypodium loriceum: a. habit, bar =

5 cm (2 in.); b. rhizome scales, bar = 1 mm.

Polypodium maritimum Hieronymus

 13.101.20
Tender
A small-medium fern with medium-creeping rhizomes.
Grows well under medium light in drained, moist potting
mix.
Polypodium maritimum is distinguished by pinnatisect,
long-tapered blades and by two or three rows of areoles be-
tween the pinna midrib and margin. The rhizomes are
densely covered with roundish scales. The species is native
to Central America and northern South America.

Figure 13.101.18. Polypodium Polypodium menisciifolium Langsdorff & Fischer

levigatum: habit, bar = 5 cm  13.101.21
(2 in.). Semi-tender to hardier
A medium-sized fern with a medium- to long-creeping rhi-
zome. Grows well under medium light in drained, moist to
448 C 13

moist-dry potting mix or uncut moss. The plants are ro-

bust growers and easy to cultivate.
a Polypodium menisciifolium has one-pinnate fronds, and
the pinna bases are rounded or obtuse. The rachises are
glabrous on their upper surfaces. This species is native to
Figure 13.101.20. southern Brazil.
Polypodium maritimum: This species is sometimes misidentified as Polypodium
a. habit, bar = 5 cm fraxinifolium Jacquin, a species not in recent cultivation.
(2 in.); b. rhizome
scales, bar = 1 mm. Polypodium pellucidum Kaulfuss
 13.101.22, 34m
Pellucid polypody
Tender to semi-tender
A small-medium fern with short- to medium-creeping rhi-
zomes. Grows well under medium light in drained, moist
to moist-dry potting mix or uncut moss. When grown in
b high light, the fronds become short and compact.
The fronds of Polypodium pellucidum are shiny, bright
green, and evergreen. The veins are free (not netted), and a
false vein ends at each marginal notch. The sori vary from
round to oval and lack paraphyses. This species is endemic
to the Hawaiian Islands.

Polypodium plebeium Schlechtendal & Chamisso

 13.101.23
Semi-tender or hardier
A small fern with short- to medium-creeping rhizomes.
Grows well under medium light in drained, moist-dry pot-
ting mix. This species can be grown in soil or as an epiphyte.
a The rhizome scales of Polypodium plebeium have a dark
central strip and lighter margins. The dark green fronds

a b

Figure 13.101.21. Polypodium menisciifolium: a. habit,

bar = 5 cm (2 in.); b. pinna with sori, bar = 1 cm Figure 13.101.22. Polypodium pellucidum: a. frond, bar
(0.4 in.), after Hoshizaki (1982); c. rhizome scale, bar = = 5 cm (2 in.); b. segment with veins, bar = 3 mm
1 mm, after Hoshizaki (1982). (0.1 in.); c. rhizome scales, bar = 3 mm (0.1 in.).
 F  F A  C 449

a
c

c Figure 13.101.24. Polypodium polypodioides: a. habit, bar

= 5 cm (2 in.); b. blade scale, bar = 1 mm; c. rhizome
scale, bar = 1 mm. After Hoshizaki (1982).
b

var. aciculare Weatherby, of Central America; var. burchel-

lii (Baker) Weatherby, of South America; var. ecklonii
(Kunze) Weatherby, of southwestern Africa; var. michauxi-
d anum Weatherby, of the southeastern United States, Cen-
tral America, the West Indies, and northern South Amer-
ica; var. minus (Fée) Weatherby, of southern Brazil, Uru-
guay, Paraguay, and Argentina; and var. polypodioides, of
Central America, the West Indies, and northern South
America.
Figure 13.101.23. Polypodium plebeium: a. frond grown In the southeastern United States the resurrection fern
under garden conditions, bar = 5 cm (2 in.); b. frond variety michauxianum (sometimes called Pleopeltis polypo-
from the wild, bar = 5 cm (2 in.); c. base of segment dioides var. michauxianum (Weatherby) E. G. Andrews &
with sori and scales, bar = 1 cm (0.4 in.); d. hydathodes Windham) commonly grows on tree trunks or (more
along segment margin, upper surface, bar = 1 cm rarely) rocks and is occasionally cultivated. In the Gulf
(0.4 in.). States it is a nearly constant associate of the live oak (Quer-
cus virginiana). During dry weather the fronds curl into
rough C- and J-shapes and can lose up to 97% of their nor-
have many pinnae (up to about 18 pairs) that are narrow mal moisture content. They can survive for months in this
and close together. Scales are on the lower surface of the condition without any ill effects. The fronds rehydrate with
blade only, 0.5–1.0 mm long, few, and inconspicuous. The rain and expand within hours. Physiological studies have
veins are hard to see because the blades are thick. This spe- shown that the scales on the lower surface of the blade fun-
cies is native to Central America. nel water into the middle layer of the leaf, thus helping to
rehydrate the frond. For a nontechnical account of this, see
Polypodium polypodioides (Linnaeus) Watt Moran (1998).
 13.101.24
syn. Pleopeltis polypodioides (Linnaeus) E. G. Andrews Polypodium ptilorhizon H. Christ  13.101.25
Resurrection fern Tender
Hardy, Zone 6(7) A small fern with medium-creeping rhizomes. Grows well
A small fern with short- to (mostly) long-creeping rhi- under medium light in moist, drained potting mix.
zomes. Grows under medium light in drained, acidic, Polypodium ptilorhizon resembles a small P. loriceum but
moist-dry potting mix or uncut moss. The plants are slow- differs by having minute, scattered rhizome scales and fewer
growing and difficult to establish. Avoid disturbing the (generally 10–22) pairs of pinnae. The blades are glabrous,
roots during transplanting. and there is only one row of areoles and sori between the
Polypodium polypodioides has small fronds, to 20 cm (8 pinna midrib and margin. The species is native to tropical
in.) long and 4 cm (1.5 in.) wide. They are sparsely scaly America.
above and more densely scaly beneath. The sori are slightly
sunken and partially hidden by the covering of scales; they Polypodium pyrrholepis (Fée) Maxon
may appear slightly embossed on the upper surface of the  13.101.26;  45
blade. This species is native to the New World. Weatherby Semi-hardy
(1939) subdivided it into six varieties on the basis of frond A medium-sized fern with short- to medium-creeping rhi-
size and scale characteristics of the blades and rhizomes: zomes. Grows well under medium light in moist-dry gar-
450 C 13

den soil or potting mix kept well drained. The plants are
easy to cultivate in hanging baskets or in the ground.
Polypodium pyrrholepis has tufted, bright rusty-red rhi-
zome scales. The fronds are set close together, with blades
narrowly linear-oblong, pinnatisect, and densely scaly on
both surfaces. A small, slightly swollen gland is on the in-
side, upper edge of the segments. This species is native to
southern Mexico.

Polypodium rhodopleuron Kunze  13.101.27

Semi-tender
Figure 13.101.25. Polypodium A small fern with creeping rhizomes. Grows under medium
ptilorhizon: frond, bar = 5 cm light in moist, drained potting mix. This species can be dif-
(2 in.), after Hoshizaki ficult to cultivate but is attractive because of the reddish
(1982). rachises and (sometimes) other leaf parts. The color is best
developed on young fronds.
Polypodium rhodopleuron has whitish rhizome scales,
and the fronds are glabrous, pinnatisect, to about 30 cm
d (12 in.) long, and have red-tinged rachises. The veinlets
a next to costae have enlarged whitish tips. The species is na-
tive to Central America and Mexico.
e
a

b
b

Figure 13.101.27. Polypodium rhodopleuron: a. frond,

bar = 5 cm (2 in.); b. segment with veins, bar = 1 cm
(0.4 in.). After Hoshizaki (1982).

c Polypodium sanctae-rosae (Maxon) C. Christensen

 13.101.28
Semi-tender
A small-medium fern with short- to medium-creeping rhi-
zomes. Grows well under medium light in moist, drained
potting mix. The plants tend to wilt easily if the light be-
comes too intense. Older fronds are silvery white, gray-
Figure 13.101.26. Polypodium pyrrholepis: a. frond green.
grown in shade, bar = 5 cm (2 in.); b. habit, bar = 5 cm The rhizome scales of Polypodium sanctae-rosae are ovate
(2 in.); c. segment with scales, upper surface, bar = 1 cm to ovate-deltoid, small, appressed, and overlapping. The
(0.4 in.); d. scale from lower surface of blade, bar = stipes and rachises of young fronds are speckled with dark
1 mm, after Sota (1966); e. rhizome scale, bar = 1 mm, scales. On the upper surface of the blades the scales are pale
after Sota (1966). tan to white, whereas those on the lower surface are shiny,
pale rust colored. The sori are tightly enclosed by a rosette of
scales. The species is native to Mexico and Central America.
 F  F A  C 451

a
Figure 13.101.29. Polypodium
scouleri: a. frond, bar = 5 cm
(2 in.); b. rhizome scales, bar
= 3 mm (0.1 in.).
d c

b
a
b

The stipes of Polypodium thyssanolepis are longer than

the blades, which are deeply pinnatifid, about 10 cm (4 in.)
long, 7 cm (3 in.) wide, and broadest at the base. The pin-
Figure 13.101.28. Polypodium sanctae-rosae: a. habit, bar nae are few (three to seven pairs) and glabrous or sparsely
= 5 cm (2 in.); b. segment, upper surface, bar = 1 cm scaly on the upper surface but densely scaly on the lower.
(0.4 in.); c. scale from lower surface of blade, bar = The sori are not sunken. The species occurs from the south-
1 mm, after Sota (1966); d. rhizome scale, bar = 1 mm, western United States to South America and Jamaica. It
after Sota (1966). grows terrestrially or on or among rocks.

Polypodium scouleri Hooker & Greville

 13.101.29, 34l
Leathery polypody, coast polypody a
Semi-hardy, Zones 8–9
A small to medium fern with medium-creeping rhizomes.
Grows well under medium light in drained, moist-dry gar-
den soil or potting mix or in uncut moss. The plants are
evergreen, slow growing, and adapted to salt spray. c
Polypodium scouleri has glaucous, stout rhizomes that
produce stiff, leathery fronds with pinnae more than 12
mm (0.5 in.) wide. The veins are netted and form one row
of areoles. The sori are more than 3 mm (0.1 in.) wide. This
species is native to the West Coast of North America, from
Canada to Mexico.

Polypodium thyssanolepis A. Braun ex Klotzsch b

 13.101.30
syn. Pleopeltis thyssanolepis (A. Braun ex Klotzsch) E. G.
Andrews
Scaly polypody Figure 13.101.30. Polypodium thyssanolepis: a. habit, bar
Semi-hardy, Zone 8 = 5 cm (2 in.); b. segment with sori and scales, bar =
A small fern with medium-creeping rhizomes. Grows well 5 mm (0.2 in.), after Knobloch and Correll (1962);
under medium light in drained, moist-dry garden soil or c. blade scale, bar = 1 mm, after Knobloch and Correll
potting mix. (1962).
 452 C 13

Polypodium triseriale Swartz  13.101.31 Polypodium virginianum Linnaeus

Tender to slightly hardier  13.101.32, 34f
A medium-sized fern with medium-creeping rhizomes. Virginia polypody, rock polypody, American wall fern
Grows well under medium light in drained, moist garden Very hardy, Zone 2
soil or potting mix. The species is extremely variable A small fern with short- to medium-creeping rhizomes.
throughout its range. Grows under low-medium light in drained, moist potting
Polypodium triseriale has ovate rhizome scales with pale mix. The plants establish slowly and are susceptible to at-
margins. The fronds are one-pinnate at least at the base, tack from snails and slugs. The fronds are evergreen.
with linear-lanceolate pinnae; the lower pinnae are nar- The rhizome scales of Polypodium virginianum are dark
rowed toward the base and nearly free on the basiscopic brown and often weakly marked with a darker central
side, narrowly adnate on the acroscopic side. The rachises stripe. The pinnatisect, thin-leathery fronds bear round
are glabrous on the upper surface, and the veins form up to sori. The paraphyses are modified sporangia with glands.
five series of areoles. The sori are in one to three rows. This This species is native to the eastern half of North America.
species is common and widespread throughout tropical Polypodium virginianum is a fertile plant of hybrid origin
America. from an ancient cross involving P. appalachianum Haufler
‘Cambrioides’. Pinnae sterile, irregularly lacerate, and & Windham and P. sibericum Siplivinsky. It differs from P.
ruffled. This cultivar grows slowly. Its fronds tend to be too appalachianum by its more-linear fronds and light brown or
heavy for the stipes, which often break. Since the plant is bicolorous rhizome scales. The plants from boreal Canada
sterile, its placement with Polypodium triseriale is tentative. and Alaska previously called P. virginianum have been
For a similar plant, see Microsorum commutatum ‘South- transferred to P. sibericum, the Siberian polypody, which
ern Cross’. lacks glands on the paraphyses and has smaller spores.
‘Cristatum’. Pinnae several times forked to form a crest.
Reportedly found in Brazil by Barbara Varga of Varga Polypodium vulgare Linnaeus
Nursery, Pennsylvania. Circulating in the United States  13.101.33, 34b
trade as Polypodium brasiliense ‘Cristata’. Common polypody
Hardy, Zone (4)5
A small fern with short- to medium-creeping rhizomes.
Grows well under medium light in drained, moist potting
mix or uncut moss. The plants grow slowly, with new
growth appearing in early summer.

Figure 13.101.32. a
Polypodium virginianum:
a. habit, bar = 5 cm
(2 in.); b. paraphysis,
top view, bar = 0.1 mm.

b
b
c
b
Figure 13.101.33. Polypodium
Figure 13.101.31. Polypodium triseriale: a. habit, bar = vulgare: a. habit, bar = 5 cm
5 cm (2 in.); b. pinna with sori, bar = 1 cm (0.4 in.); (2 in.); b. sporangium, bar =
c. rhizome scale, bar = 1 mm. After Hoshizaki (1982). 0.5 mm, after Shivas (1962).
 a

b d e

c f

l
m
k
i

Figure 13.101.34. Frond silhouettes of species in the Polypodium vulgare complex. European species: a. P. cambricum;
b. P. vulgare; c. P. interjectum. Eastern North American species: d. P. appalachianum; e. P. sibericum; f. P. virginianum.
Western North American species: g. P. californicum; h. P. glycyrrhiza; i. P. calirhiza; j. P. amorphum; k. P. hesperium;
l. P. scouleri. Hawaiian species: m. P. pellucidum. Bars = 5 cm (2 in.). Plants c, f, i, and k of hybrid origin; see text entries
for details.
 454 C 13

The blades of Polypodium vulgare are flat, pinnatisect,

and lanceolate, and almost all the pinnae are equal in
length. The sori are round, without paraphyses. This spe-
cies is native to Europe.
Polypodium cambricum and P. interjectum differ from
this species by their broader fronds, their forward-project-
ing lower pinnae, and in sori details. For more informa-
tion, see Rickard (1981).
‘Ramosum Hillman’. Fronds branched near the base of
the blade; tips of the branches forked to form a crest.

Polystichum  13.102.1–3

Shield fern
The shield ferns of the genus Polystichum are small- to me-
dium-sized terrestrial ferns commonly grown in temperate
gardens. Many of the species are particularly attractive for
their dark green, glossy, evergreen foliage. The plants are
used in rock gardens, borders, or pots, and larger species
can be used as foundation plants or for background foliage. Figure 13.102.2. Polystichum pinnae: P. proliferum (left),
They are often slow to grow from spores. P. munitum (center), P. polyblepharum (right).
Eastern and western North America harbor distinctive
species of Polystichum. In the eastern region the Christmas
fern (Polystichum acrostichoides) is one of the most common glossy green. The soft shield fern (P. setiferum), a native of
native species and is widely cultivated. It is easily recog- Europe, and its bewildering array of more than 300 garden
nized by its one-pinnate fronds with narrowed fertile pin- varieties are grown throughout the United States. The wild
nae toward the tip. The western sword fern (P. munitum) is form of the species resembles the bristle fern but has dull,
abundant in the coastal forests of western North America, lighter green blades.
and it is commonly grown in gardens. Its narrow, one-pin- Western North American Polystichum species do not
nate fronds may grow to 1.6 m (5 ft.) tall but are usually grow as well on the East Coast, and vice versa, but the Eu-
30–100 cm (1–3 ft.) in cultivation. ropean and Asiatic species are adaptable in both areas, ex-
Besides the North American species, several European
and Asian ones are commonly planted in certain regions of
the United States. The bristle fern, Polystichum polybleph-
arum (widely sold as P. setosum), is cultivated in the South-
west and elsewhere. Its fronds are two-pinnate, dark, and

Figure 13.102.3. Polystichum proliferum: sori

Figure 13.102.1. Polystichum polyblepharum: habit. and indusia.
 F  F A  C 455

cept in very hot, humid places. The northwestern species Fronds two-pinnate-pinnatifid or up to three-pinnate, at
can be grown in the southwest if provided with high hu- least at the base of the lower pinnae: Polystichum proliferum,
midity and acid soils. Because only a few species produce P. richardii, P. silvaticum, P. tagawanum, P. tsus-simense,
multiple crowns that can be divided, most polystichums P. vestitum
are propagated from spores. Many species also form buds
along their rachises or at the tips of the fronds. These buds Polystichum acrostichoides (Michaux) Schott
are easily rooted by pegging the leaf to the soil. Thrip dam-  13.102.4
age is often conspicuous on fronds of Polystichum, appear- Christmas fern
ing as a grayish silvering on the blade. Very hardy, Zone 3
Polystichum has erect, ascending (rarely short-creeping) A medium-sized fern with ascending, clump-forming rhi-
rhizomes. The fronds are mostly one- to two-pinnate, less zomes and evergreen leaves. Grows well under low to me-
often three-pinnate, pinnatifid. They are elongate, often dium light in moist garden soil or potting mix. Although
narrowed at the base, with the apex gradually reduced. The easily cultivated in the eastern United States, the plants are
pinnae often have an upward-pointing “ear” at their base— difficult to grow on the West Coast. In winter, the fronds
a characteristic helpful in distinguishing the genus from recline to the ground, forming a rosette around the rhizome
Dryopteris, which often lacks the ear. Some one-pinnate apex.
species have terminal pinnae that resemble the lateral ones. Polystichum acrostichoides is characterized by one-pin-
The marginal teeth generally bear a short bristle or spine. nate blades with the distal, fertile pinnules narrower than
The veins are anadromic and free, not netted. Round sori the sterile ones. The pinnae have an upward-pointing “ear”
are formed in one row (or infrequently two) between the at the base, and the margins are spinulose. When the sori
midrib and margin. Indusia may be present or absent; when mature they tend to become confluent, resembling acrosti-
present, they are peltate. choid sori. The species is native to eastern North America.
Some authors believe that Cyrtomium (including Pha- A sterile hybrid, Polystichum ×potteri Barrington (= P.
nerophlebia) is so closely related to Polystichum that it acrostichoides × P. braunii ), is sometimes cultivated. The
should be combined with it. Cyrtomium, however, differs hybrid is native to Pennsylvania, New England, Nova Sco-
by having netted veins and sori in two or more series. tia, New Brunswick, and Quebec.
The genus Polystichum contains 180 species and is cos- Various cultivar names are listed in the trade, such as
mopolitan. For details on the many varieties, see Dyce ‘Crispum’, ‘Cristatum’, and ‘Incisum’.
(1963), Kaye (1968), and Mickel (1994). The genus name
comes from the Greek poly, many, and stichos, row, refer-
ring to the sori of the type species (Polystichum aculeatum),
which are arranged in many regular rows on the pinnae.
Because Polystichum contains so many species, the fol-
lowing subgroups are given to aid identification. Once an
unknown species is limited to one of these subgroups, it
can be compared to the descriptions and illustrations of Figure 13.102.4. Polystichum
other species in the group. The groups are based on typical acrostichoides: frond, bar =
fronds (not cultivars) from mature plants. 5 cm (2 in.).
Fronds one-pinnate, the pinnae entire, serrate, or crenate,
not pinnatifid: Polystichum acrostichoides, P. craspedosorum,
P. imbricans, P. lepidocaulon, P. lonchitis, P. munitum, P.
nepalense, P. stenophyllum, P. stimulans

Fronds one-pinnate-pinnatifid, lower pinnae may have a

free pinnule at their base: Polystichum andersonii, P. cali-
fornicum, P. kruckebergii, P. lentum, P. scopulinum, P. seti-
gerum, P. xiphophyllum

Fronds two-pinnate or nearly so in the proximal part: Poly-

stichum aculeatum, P. xiphophyllum

Fronds fully two-pinnate in most parts of the frond, not

more divided: Polystichum braunii, P. dudleyi, P. lemmonii,
P. makinoi, P. neolobatum, P. polyblepharum, P. proliferum,
P. rigens, P. setiferum, P. tripteron, P. tsus-simense
 456 C 13

Polystichum aculeatum (Linnaeus) Roth Polystichum andersonii M. Hopkins

 13.102.5  13.102.6
syn. Polystichum lobatum (Hudson) C. Presl Anderson’s sword fern, Anderson’s holly fern
Hard shield fern Hardy, Zone 5(6)
Very hardy, Zone 4 A medium-sized fern with ascending to erect rhizomes and
A medium-sized fern with ascending to erect rhizomes and evergreen, bud-bearing leaves. Grows well under medium
stiff, glossy, evergreen leaves. Grows well under medium light in moist garden soil or potting mix. The plants prefer
light in moist, basic garden soil or potting mix. This species moist, cool climates. This species adapts poorly on the East
is easy to grow. Coast and in southern California.
The stipes of Polystichum aculeatum are often short, and Polystichum andersonii has short stipes and elliptic-lan-
the fronds are lanceolate and nearly two-pinnate, with the ceolate, one-pinnate-pinnatifid blades that are narrowed
basal pinnae half the length of the middle ones. The pin- toward the base. The pinnules are adnate to the costae, the
nules are sessile or nearly so, and the spores are dark brown. adnation 2 mm wide or more. The bud(s) form on the dis-
This species is native to Europe. tal third of the rachis. The indusium is dentate. This species
Polystichum aculeatum is often difficult to distinguish is native to western North America.
from P. setiferum, which is softer, fully two-pinnate, often The buds of this species do not drop from the leaf as in
not or only slightly narrowed at the blade base, has pin- some ferns. Instead, they take root after the leaf has reclined
nules clearly stalked, and yellow spores. A sterile hybrid, in late fall and is prostrate on the soil. In the wild the plants
Polystichum ×illyricum (Burbas) Hahne (= P. aculeatum × P. form colonies by this method of reproduction.
lonchitis), is reported to be cultivated and appears interme-
diate between its parents.
a

c
Figure 13.102.5. Polystichum
aculeatum: a. frond, bar =
5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.). Figure 13.102.6. Polystichum
andersonii: a. frond, bar =
5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.); c. bud on
rachis, bar = 2 mm.
a

b
Polystichum braunii (Spenner) Fée
 13.102.7;  6
Braun’s holly fern, prickly shield fern
Very hardy, Zone 3(4)
A small to medium fern with clump-forming rhizomes and
dark green, shiny, evergreen fronds. Grows well under me-
dium light in moist potting mix. The plants do best if
 F  F A  C 457

Figure 13.102.7. Polystichum

braunii: frond, bar = 5 cm (2 in.).

b
placed in a cool site. The fiddleheads are particularly at-
tractive because they are densely covered by silvery scales,
which turn light brown with age.
Polystichum braunii has short stipes, usually about one-
quarter the length of the frond. The stipes are densely scaly,
and the blades are two-pinnate and elliptic-lanceolate, grad-
ually narrowed toward the base. The pinnules are short-
stalked and bristly. This species is native to North America,
Europe, and Asia.

Polystichum californicum (D. C. Eaton) Diels

 13.102.8
California sword fern, California holly fern
Semi-hardy, Zone 8
A medium-sized fern with ascending to erect rhizomes and a
evergreen fronds. Grows under low to medium light in
moist, drained potting mix. The plants are slow to estab-
lish. The new growth has silvery scales.
The blades of Polystichum californicum vary from one- Figure 13.102.8. Polystichum californicum: a. frond
pinnate-pinnatifid to almost two-pinnate and have adnate variations, bar = 5 cm (2 in.); b. pinna variations, bar =
pinnules, the adnation 2 mm or more wide. The species is 2 cm (0.8 in.), after D. Wagner (1979).
native to western North America.
This species is of hybrid origin, resulting from an an-
cient cross involving Polystichum dudleyi and either P. mu-
nitum or P. imbricans. According to Mickel (1994), some
forms are sterile, others fertile.

Polystichum craspedosorum (Maximowicz) Diels

 13.102.9
Hardy, Zone 5
A small fern with ascending to erect rhizomes and ever-
green fronds. Grows well under medium light in moist pot- Figure 13.102.9. Polystichum
ting soil. craspedosorum: frond, bar =
Polystichum craspedosorum has linear-lanceolate, one- 5 cm (2 in.), after Kurata and
pinnate blades, with the rachises prolonged and rooting at Nakaike (1983).
tip. It is native to eastern Siberia, China, Japan, and Korea.
 458 C 13

Polystichum dudleyi Maxon  13.102.10

Woodrustic fern, Dudley’s sword fern, Dudley’s holly
fern
Marginally semi-hardy, Zone 8
A medium-sized fern with ascending to erect rhizomes and
evergreen fronds. Grows well under medium light in moist,
drained potting mix. The plants do best in a cool, moist b
climate, but they establish slowly. The fronds are slightly
brittle.
Polystichum dudleyi has scaly stipes and rachises, and the
blades are lanceolate, two-pinnate, and hardly tapered to-
ward base. The pinnules are stalked and bear a tangle of
long, narrow, tan scales on their lower surfaces. The species
is native to California.

Figure 13.102.11.
Polystichum imbricans:
a. frond, bar = 5 cm
(2 in.); b. pinnae, bar
= 1 cm (0.4 in.), after
D. Wagner (1979).
b

Figure 13.102.10. Polystichum

dudleyi: a. frond, bar = 5 cm a
(2 in.); b. pinnae, bar = 2 cm
(0.8 in.), after D. Wagner
(1979).
subsp. curtum (Ewan) D. H. Wagner. Pinnae slender,
lanceolate, falcate, plane, larger than those of subsp. imbri-
cans; fronds up to 18 cm (7 in.) long. Native to California.
a
subsp. imbricans. Dwarf western sword fern. Pinnae
short, oblong, folded inward, oriented horizontally, over-
lapping; fronds dark green, stiff, erect, up to 50 cm (20 in.)
long. Native to southwestern Canada and the northwestern
Polystichum imbricans (D. C. Eaton) D. H. Wagner United States.
 13.102.11
syn. Polystichum munitum var. imbricans (D. C. Eaton) Polystichum kruckebergii W. H. Wagner
Maxon  13.102.12
Imbricate sword fern Kruckeberg’s holly fern
Hardy, Zone 6 Hardy, Zone 6
A small to medium fern with ascending to erect rhizomes. A small to medium fern with ascending to erect rhizomes
Grows well under medium to high light in moist garden and dark green, glossy, evergreen fronds. Grows under me-
soil or potting mix with good drainage. dium to high light in moist garden soil or potting mix with
The scales on the stipes and rachises of Polystichum im- good drainage. This species is difficult to grow.
bricans are shed early; the largest of these scales are less than Polystichum kruckebergii has narrow and one-pinnate-
1 mm wide. The blades are linear-lanceolate and one-pin- pinnatifid blades with ovate-triangular pinnae that tend to
nate with oblong to lanceolate pinnae that are tipped by a overlap. The species is native to western North America.
sharp, rigid, short point. The indusia are toothed or entire, This species is the fertile hybrid of Polystichum lemmonii
not ciliate. This species is native to the West Coast of Can- and P. lonchitis. Because it is fertile, it can be propagated by
ada and the United States. See also P. munitum. spores.
 F  F A  C 459

Figure 13.102.13. Polystichum

lemmonii: a. frond, bar = 5 cm
(2 in.); b. pinna, bar = 1 cm
(0.4 in.). After Cody and
Britton (1989).
b

Figure 13.102.12. Polystichum

kruckebergii: a. frond, bar = 5 cm
(2 in.), after Cody and Britton
a (1989); b. pinnae, bar = 1 cm
(0.4 in.).

Polystichum lemmonii Underwood  13.102.13

syn. Polystichum mohrioides var. lemmonii (Underwood)
Fernald
Shasta holly fern
Hardy, Zone 6
A small fern with ascending to erect rhizomes and semi-
evergreen(?) fronds. Grows under medium to high light in
moist potting mix with good drainage. The plants are dif-
ficult to grow.
Polystichum lemmonii has narrowly lanceolate, two-pin- Figure 13.102.14. Polystichum
nate-pinnatifid blades with stalked pinnules that lack a lentum: frond, bar = 5 cm (2 in.).
spine at their tips. The species is native to western North
America.

Polystichum lentum (D. Don) T. Moore

 13.102.14
Himalayan holly fern
Semi-hardy, Zone 7
A small-medium fern with ascending to erect rhizomes and
semi-evergreen(?) fronds. Grows well under medium light
in moist garden soil or potting mix. Polystichum lepidocaulon (Hooker) J. Smith
The blades of Polystichum lentum are narrowly elliptic  13.102.15
and one-pinnate-pinnatifid to two-pinnate at the base. The Semi-hardy, Zone 7
pinnae are only deeply toothed, although some of the lower A small-medium fern with ascending to erect rhizomes and
pinnae have a distinct basal pinnule. Buds are borne near dark green, leathery, evergreen fronds. Grows well under
the end of rachis. The species is native to Tibet, China, and medium light in moist garden soil or potting mix.
Burma. Polystichum lepidocaulon has scaly stipes and one-pin-
nate blades with long falcate, dark green, leathery pinnae.
 460 C 13

Figure 13.102.15. Polystichum Figure 13.102.16. Polystichum lonchitis: a. frond, bar =

lepidocaulon: fronds, bar = 5 5 cm (2 in.); b. lowermost pinna, bar = 1 cm (0.4 in.).
cm (2 in.).

Polystichum makinoi (Tagawa) Tagawa

 13.102.17
Makino’s holly fern
The densely scaly rachis extends well beyond the blade and Hardy, Zone 5
roots at the tip upon touching the ground. The species is A medium-sized fern with ascending to erect rhizomes and
native to Japan, Korea, eastern China, and Taiwan. dark, glossy, evergreen fronds. Grows well under medium
Mickel (1994) noted that this species was tried outdoors light in moist garden soil or potting mix.
at the New York Botanical Garden without success but that The stipes and rachises of Polystichum makinoi bear nu-
it should be hardy in milder climates to the south. merous brown scales with paler margins. The blades are
two-pinnate, the lower pinnae not or slightly reduced in
Polystichum lonchitis (Linnaeus) Roth size. The segments are rhombic and have soft, narrow, aris-
 13.102.16 tate teeth. The species is native to eastern India, southern
Northern holly fern China, Japan, and the Philippines.
Very hardy, Zone 3
A medium-sized fern with ascending to erect rhizomes and Polystichum munitum (Kaulfuss) C. Presl
dark green, evergreen fronds. Grows under medium light in s 13.102.2, 18
moist, basic garden soil or potting mix. The plants are dif- Western sword fern, common sword fern
ficult to grow even in cool climates. Hardy, Zone 7
The stipes are short, and the blades are linear-lanceo- A medium-large fern with ascending to erect rhizomes and
late, one-pinnate, and narrowed toward the base. The tri- evergreen fronds. Grows well under low to medium light in
angular basal pinnae are spinulose-margined, with the moist garden soil or potting mix with a lot of humus. This
spines close and spreading. Polystichum lonchitis is native species does best in moist, cool climates and does not grow
to northern North America and Greenland. well in the eastern of southeastern United States. The
Two hybrids involving this species are also cultivated: fronds are frequently used in floral decorations.
Polystichum ×illyricum (Burbas) Hahne (= P. aculeatum × P. The stipes and rachises of Polystichum munitum bear
lonchitis) and P. ×lonchitiforme (Halacsy) Becherer (= P. persistent ovate-lanceolate scales, to 1 mm wide or more in
lonchitis × P. setiferum). the distal part of the rachis. The blades are linear-lanceolate,
one-pinnate, and plane, and the pinna apex is acuminate
and short-spinulose (not with a sharp, short, rigid point).
 F  F A  C 461

The indusia are ciliate. Also see P. imbricans. The species is

native to the western United States, Canada, Alaska (Yu-
kon), and Mexico (Guadalupe Island); it is naturalized in
Europe.

Polystichum neolobatum Nakai  13.102.19

b Long-eared holly fern
Hardy, Zone 5
A medium-sized fern with ascending to erect rhizomes and
glossy, hard-textured, evergreen fronds that have a neatly
trimmed appearance. Grows well under low to medium to
high light in moist garden soil or potting mix. The plants
c
are easy to cultivate.
Polystichum neolobatum bears shaggy scales on its stipes
and rachises, and the blades are lanceolate, two-pinnate,
Figure and sparsely scaly on the undersides. The pinnule teeth are
13.102.17. stiff and spiny. The acroscopic pinnule next to the rachis is
Polystichum large and ear-like. This species is native to Nepal, the Him-
makinoi: a. alayas, Tibet, Burma, China, Taiwan, and Japan.
frond, bar =
5 cm (2 in.);
b. pinna, bar =
1 cm (0.4 in.);
c. pinnule, bar =
1 cm (0.4 in.).
a
Figure 13.102.19. Polystichum
neolobatum: a. frond, bar =
5 cm (2 in.); b. pinna, bar =
1 cm (0.4 in.).

b
Figure 13.102.18.
Polystichum munitum: a
a. frond, bar = 5 cm
(2 in.); b. pinna, bar =
1 cm (0.4 in.).

Polystichum nepalense (Sprengel) C. Christensen

 13.102.20
Nepal holly fern
a Hardy, Zone 5
A medium-sized fern with ascending to erect rhizomes and
evergreen fronds. Grows well under medium light in moist
potting soil with good drainage.
b Polystichum nepalense has narrowly lanceolate, one-pin-
nate blades with pinnae that are lanceolate, curved, eared,
and finely toothed. The species is native to the Himalayas,
China, Taiwan, and the Philippines.
 462 C 13

b
b

a c

Figure 13.102.21.
Polystichum
polyblepharum:
a. frond, bar =
5 cm (2 in.);
b. pinna, bar =
1 cm (0.4 in.);
c. pinnule, bar =
5 mm (0.2 in.).
a

Figure 13.102.20. Polystichum nepalense: a. frond, bar =

5 cm (2 in.); b. pinna, bar = 1 cm (0.4 in.); c. stipe scale,
bar = 1 mm. After Hu and Ching (1930).
States but might be a handsome addition to gardens. It is
hardy to Zone 5. This species is native to Japan and Korea.

Polystichum polyblepharum (J. Roemer ex Kunze) Polystichum proliferum (R. Brown) C. Presl
C. Presl  13.102.1, 2, 21  13.102.2, 3, 22
Bristle fern, Japanese sword fern Mother shield fern
Hardy, Zone 5(6) Hardy, Zone 5
A medium-sized fern with erect rhizomes and dark green, A medium-sized fern with erect rhizomes and dark green
glossy, evergreen fronds. Prefers high-medium light and fronds that are evergreen in warmer climates. Grows well
moist or moist-dry garden soil or potting mix. This species under medium light in moist garden soil or potting mix.
is easy to grow. This species is easy to grow and can be propagated from
Polystichum polyblepharum has densely scaly stipes and the bulbils on the fronds.
rachises. The blades are elongate-elliptic to oblong and two- The blades of Polystichum proliferum are lanceolate to
pinnate. The pinnules are eared, with the acroscopic ear broader at the base and two- to three-pinnate, with a bud
prominent and spine-tipped and the basiscopic ear incon- forming on the apical part. The pinnules are toothed. The
spicuous with its spine often bent over the upper pinnule species is native to Australia and New Zealand.
surface. This species is native to Japan, southern Korea, and This species resembles Polystichum vestitum but is iden-
eastern China. tifiable by the bulbils produced near the frond apices and by
The plants in the trade are widely misidentified as Poly- the glossy stipe scales with dark centers and pale borders.
stichum setosum Horticulture. The name Polystichum proliferum might be confused with
Polystichum retrosopaleaceum (Kodama) Tagawa some- P. setiferum cultivars that bear buds on their rachis and have
what resembles P. polyblepharum but differs by lower stipe circulated incorrectly under the same name.
scales broadly ovate with a long point (versus lanceolate-
triangular), the pinnae apices more acuminate-pinnatifid
(versus acute and subconform), and the foliage clear green
(versus dark green). It is rarely cultivated in the United
 F  F A  C 463

Figure 13.102.23. Polystichum

Figure 13.102.22. Polystichum richardii: frond, bar = 5 cm
proliferum: a. frond, bar = (2 in.), after Brownsey and
5 cm (2 in.); b. pinna, bar = Smith-Dodsworth (1989).
1 cm (0.4 in.).

Polystichum richardii (Hooker) J. Smith

 13.102.23 a
Richard’s holly fern
Semi-hardy, Zone 7 b
A small to medium fern with erect rhizomes and evergreen
fronds that vary from dark bluish green to olive green.
Grows under medium light in moist garden soil or potting
mix.
The stipes and rachises of Polystichum richardii are dark
and densely covered with blackish brown scales with pale
fringes at their base. The blades are narrowly oblong-ellip-
tic to narrowly triangular and two- to three-pinnate. The
indusia have a dark center and pale margins. The species is
native to New Zealand.

Polystichum rigens Tagawa  13.102.24

Hardy, Zone 5
c
A medium-sized fern with erect rhizomes and evergreen,
trim, rigid, glossy green fronds. Grows well under medium
light in moist garden soil or potting mix.
The stipe scales have an apical cluster of cilia, and the
blades are triangular and two-pinnate. Polystichum rigens
has lanceolate to narrow-triangular pinnae, the apices of
which have a spine-like tip. This species somewhat resem- Figure 13.102.24. Polystichum rigens: a. frond, bar =
bles P. tsus-simense, which has lanceolate stipe scales and is 5 cm (2 in.); b. pinnule, bar = 1 cm (0.4 in.); c. lower
a smaller plant. Polystichum rigens is native to Japan and stipe scales, bar = 1 cm (0.4 in.). After Kurata and
China. Nakaike (1979).
 464 C 13

Polystichum scopulinum (D. C. Eaton) Maxon

 13.102.25
Rock sword fern, western holly fern a
Very hardy, Zone 4
A small to medium fern with ascending to erect rhizomes
and leathery, semi-evergreen fronds. Grows under medium
to high light in moist garden soil or potting mix with good
drainage. This species is difficult to grow. Figure 13.102.26. Polystichum
Polystichum scopulinum has narrow, one-pinnate-pin- setiferum: a. frond, bar = 5 cm
natifid blades. The pinnae are deeply toothed distally and (2 in.); b. pinna, bar = 1 cm
have on the acroscopic side a large, free “ear” that is adnate, (0.4 in.).
the adnation 2 mm or more wide. The species is native to
western North America.
This species is a hybrid of Polystichum imbricans and P.
lemmonii.

Figure 13.102.25. Polystichum

scopulinum: a. frond, bar = 5 cm
(2 in.); b. pinna, bar = 1 cm (0.4 in.).
After Cody and Britton (1989).
Because it varies tremendously (mostly in garden plants),
this species is difficult to characterize. It is also confused
with the infrequently cultivated Polystichum aculeatum.
Many cultivars can be found in the trade and special col-
lections, and only a few are listed here. Sporelings fre-
quently vary, and many of these intergrade and are not par-
ticularly distinct or deserving of cultivar names. The
cultivar names are greatly confused in the trade and among
collectors, creating controversy among growers. The culti-
vars have been reorganized according to a different set of
a b characteristics than those previously used. For more infor-
mation, see Dyce (1963, 1987) and Kaye (1968).
‘Congestum’. Figure 13.102.27a. Dwarf soft shield fern.
Fronds 15–30 cm (6–12 in.) long, darkish gray-green; pin-
nae and pinnules close and overlapping. This cultivar dif-
Polystichum setiferum (Forsskål) T. Moore ex Woynar fers from typical plants by the smaller, upright, compact
 13.102.26, 27 blades that are thicker textured due to the congestion of
syn. Polystichum angulare (Willdenow) C. Presl pinnae and pinnules. It is suited for use in borders because
Soft shield fern of its small size.
Hardy, Zone 6 Divisilobum Group. Figure 13.102.27b. Blades narrow;
A medium to large fern with erect rhizomes and fronds that pinnae less foliaceous than those of the species; pinnules
are evergreen in warmer climates. Grows well under me- narrowly ovate, apices acute, base oblique and eared. This
dium light in moist to moist-dry garden soil or potting mix. selection includes the common trade cultivar that has nar-
Many variants of this species form buds along the rachis. row fronds and buds along the rachis. It is widely known in
The plants do not like very high humidity. the United States trade under the incorrect names Polysti-
The stipes of Polystichum setiferum are more than one- chum angulare ‘Proliferum’ and P. angulare ‘Alaska’ and
sixth the length of frond, and the blades are lanceolate and should not to be confused with the Australian and New
two-pinnate, with the lowest pinnae equal in length to the Zealand species P. proliferum.
middle pinnae or sometimes shorter. The pinnules have a Imbricatum Group. Figure 13.102.27c. Parts less con-
distinct stalk. This species is native to Europe. gested than ‘Congestum’, and fronds larger.
 F  F A  C 465

a b c

d
e f

Figure 13.102.27. Polystichum setiferum cultivars: a. ‘Congestum’; b. Divisilobum Group; c. Imbricatum Group; d.
Multilobum Group; e. Plumoso-Divisilobum Group; f. ‘Trilobum’. Bar = 5 cm (2 in.).

Multilobum Group. Figure 13.102.27d. Resembles Di- and congested; fronds bud-bearing. A very distinct, com-
visilobum Group but with pinnules stouter, nearly rectan- pact plant. Sometimes circulates as ‘Divisilobum Grandi-
gular, and not overlapping; some variants are bud-bearing. ceps’ or ‘Divisilobum Cristatum’ in the United States trade.
Plumoso-Divisilobum Group (‘Plumoso-Multilobum’).
Divided soft shield fern. Figure 13.102.27e; Plate 46.
Fronds 60–100 cm (2–3 ft.) long, horizontally spreading, Polystichum setigerum (C. Presl) C. Presl
to three-pinnate; pinnae overlapping; basiscopic pinnules  13.102.28
linear-lanceolate, longer than the acroscopic pinnules. This syn. Polystichum braunii subsp. alaskense (Maxon) Calder
beautiful fern has a feathery, neatly three-dimensional as- & R. L. Taylor
pect due to the spreading angle of the overlapping pinnae. Alaskan holly fern
‘Trilobum’ (‘Divisilobum Grandiceps’). Figure Hardy, Zone
13.102.27f. Pinnules have the same shape as those of Di- A medium-sized fern with ascending to erect rhizomes and
visilobum Group, but the distal half of the blade is a mass evergreen fronds. Grows under medium light in moist pot-
of branches. Blades to about 30 cm (12 in.) long, rachises ting mix. The plants do best in cool, moist climates.
branched, branches often irregular and branched one or The stipes of Polystichum setigerum are about half the
more times making the far (distal) part of the blade broad length of the fronds, and the blades are lanceolate and one-
 466 C 13

Figure 13.102.28. Polystichum

setigerum: a. frond, bar = 5 cm
(2 in.), after Cody and Britton
(1989); b. pinna, bar = 5 mm
(0.2 in.), after D. Wagner
(1979). Figure 13.102.29.
Polystichum silvaticum:
frond, bar = 5 cm
a (2 in.), after Brownsey
and Smith-Dodsworth
(1989).

b Polystichum stenophyllum H. Christ

 13.102.30
Hardy, Zone 6
A small fern with ascending to erect rhizomes and ever-
pinnate-pinnatifid to nearly two-pinnate. The pinnules are green leaves. Grows under medium light in moist potting
adnate, the adnation 2 mm or more, with bristle-toothed mix.
margins. This species is native to Alaska and Canada. Polystichum stenophyllum has short stipes clothed in
This species is of hybrid origin, resulting from a cross be- broadly ovate, ciliate scales with tailed apices. The linear-
tween Polystichum munitum and P. braunii. The trade ma- lanceolate, one-pinnate blades are up to 2.5 cm (1 in.) wide,
terial sold as Polystichum ‘Alaska’ is a cultivar of P. setiferum. and they bear buds below the apex. The many pinnae are
ovate, serrate, and eared, with the lower ones smaller and
Polystichum silvaticum (Colenso) Diels pointing downward. The species is native to India, Tibet,
 13.102.29 China, and Taiwan.
Semi-hardy, Zone 8(9)
Polystichum stimulans C. Presl  13.102.31
A small to medium fern with erect rhizomes and shiny, soft,
syn. Polystichum illicifolium T. Moore
evergreen fronds. Grows well under low to medium light in
Hardy, Zone 6
moist garden soil or potting mix. The plants grow best in
damp, shady sites. A small fern with ascending to erect rhizomes and ever-
The scales on the stipes and rachises are broad, glossy, green fronds. Grows under medium light in moist potting
and dark in the center. Polystichum silvaticum has lanceo- mix.
late, two-pinnate-pinnatifid blades. The blade and pinna Polystichum stimulans has linear-lanceolate, one-pinnate
tips are long and slender. Indusia are absent. The species is fronds up to 2.5 cm (1 in.) wide. The pinnae are very short
native to New Zealand. and deeply bristle-toothed, reminiscent of holly leaves. This
This species is closely related to Polystichum vestitum but species is native to the Himalayas, China, and Taiwan.
has more-finely dissected blades, narrowly winged costae,
and no indusium. In New Zealand the two species hy-
bridize freely whenever they grow together.
 F  F A  C 467

Polystichum tagawanum Sa. Kurata

b  13.102.32
a Tagawa’s holly fern
Very hardy, Zone 4
A medium-sized fern with ascending to erect rhizomes and
shiny, evergreen fronds. Grows well under medium light
in moist potting mix. The plants are easy to grow.
The stipe and rachis scales of Polystichum tagawanum
are reddish brown, ovate to lanceolate, and toothed. The
blades are ovate-lanceolate and two-pinnate to two-pin-
nate-pinnatifid. The pinnae are shorter at the base of the
blade and point downward. The pinnules are eared and ir-
regularly spinulose-lobed. This species is native to Japan.

b
Figure 13.102.30. Polystichum stenophyllum: a. frond, bar
= 5 cm (2 in.); b. pinnule, bar = 1 cm (0.4 in.); c. stipe
base scale, bar = 3 mm (0.1 in.). After Hu and Ching
(1930).

Figure 13.102.31. Polystichum Figure 13.102.32. Polystichum tagawanum: a. frond, bar

stimulans: frond, bar = 5 cm = 5 cm (2 in.); b. pinnule, bar = 5 mm (0.2 in.); c. stipe
(2 in.). scales, bar = 2 mm. After Kurata and Nakaike (1983).

Polystichum tripteron (Kunze) C. Presl

 13.102.33
Trifid holly fern
Hardy, Zone 5
A medium-sized fern with short-creeping to clump-form-
ing rhizomes and (in temperate climates) deciduous fronds.
Grows under medium light in moist potting mix. The
plants are moderately difficult to grow.
 468 C 13

Figure 13.102.34.
Polystichum tsus-simense
var. tsus-simense:
a. frond, bar = 5 cm a
(2 in.); b. pinna, bar =
1 cm (0.4 in.).

Figure 13.102.33. Polystichum tripteron: frond, bar =

5 cm (2 in.).

Figure 13.102.35.
Polystichum tsus-
Polystichum tripteron is distinctive by its greatly elon-
simense var.
gated basal pinnae, which impart an inverted T-shape to
mayebarae: a.
the blades. The blades are two-pinnate. The long apex is
frond, bar = 5 cm
one-pinnate above the lateral pinnae. The pinnules are dull
(2 in.); b. pinna,
green (not shiny) and coarsely toothed, without bristly
bar = 1 cm
teeth. This species is native to eastern Siberia, eastern
(0.4 in.). a
China, Korea, and Japan.

Polystichum tsus-simense (Hooker) J. Smith

 13.102.34, 35
Tsus-sima holly fern
Hardy, Zones (5)6
A small-medium fern with clump-forming rhizomes and
dark green, shiny fronds that are evergreen in warm cli-
mates. Grows well under medium light in moist-dry pot-
ting mix or garden soil. In the 1950s and 1960s this species
 F  F A  C 469

was widely grown in dish gardens. The plants, however,

were not as robust as the current trade material, some of
which has been identified as var. mayebarae (Tagawa) Sa. b
Kurata and sold as the “Korean rock fern.”
var. mayebarae (Tagawa) Sa. Kurata. Korean rock fern.
Figure 13.102.35. The trade plant is more foliaceous and
larger than var. tsus-simense. The larger fronds are two-pin-
nate-pinnatifid to three-pinnate on the acroscopic pinnule
of the lower pinnae. The larger stipe scales are dark brown
and lanceolate to ovate. Native to China and Japan.
var. tsus-simense. Figure 13.102.34. This variety has ob-
long-triangular, two-pinnate blades. The basal acroscopic
pinnule is conspicuously separate and larger than the oth-
ers. It is also eared; occasionally the ear is free. The larger
stipe scales are mostly blackish brown and narrow-lanceo-
late. This variety is native to the Himalayas, Thailand,
Indochina, China, Taiwan, Korea, and Japan.
c
Polystichum vestitum (G. Forster) C. Presl
 13.102.36
Prickly shield fern
Hardy, Zone 6
Figure 13.102.36. Polystichum
A medium-sized fern with erect rhizomes and harsh, prickly,
vestitum: a. frond, bar = 5 cm
semi-deciduous fronds that are dark green above, lighter
(2 in.), after Brownsey and
below. Grows under medium light in moist potting mix or
Smith-Dodsworth (1989); a
garden soil. This species grows in cool, moist climates.
b. pinna, upper surface, bar =
The stipes and rachises of Polystichum vestitum are con-
2 cm (0.8 in.); c. pinna, lower
spicuously and densely scaly with stiff, broad, shiny scales
surface, bar = 2 cm (0.8 in.).
that are black-brown with pale margins. The blades are two-
pinnate and narrowly oblong or elliptic. The pinnules have
a basal acroscopic lobe ending in a sharp point. This species
is related to P. proliferum. It is native to New Zealand.
a
Polystichum xiphophyllum (Baker) Diels
 13.102.37
Hardy, Zone 6
A medium-sized fern with erect rhizomes and evergreen
fronds. Grows well under medium light in moist garden
soil or potting mix. This species is easy to grow.
Polystichum xiphophyllum is characterized by stipes and
rachises with dark, narrowly lanceolate to linear scales. The
blades are narrowly triangular, and the pinnae are pinnate
in the basal half. The pinnae are lanceolate, triangular, short
stalked, serrate, and lobed-crenate to one-pinnate. The
acroscopic pinnule at the pinna base is free or adnate, large,
and ovate. The species is native to India and China.

Polystichum yaemonense Horticulture

Semi-hardy, Zone 8 b
A medium-sized fern with erect rhizomes. Grows best
under medium light in moist garden soil or potting mix.
Polystichum yaemonense has medium-sized fronds, the Figure 13.102.37. Polystichum xiphophyllum: a. frond,
blades of which are narrowly triangular and two-pinnate. bar = 5 cm (2 in.); b. pinna, variations, bar = 1 cm
The pinnules are ovate-rectangular, and the sori submar- (0.4 in.).
 470 C 13

ginal. The name is of uncertain origin, but it applies to a

fern cultivated in the Pacific Northwest. It is possibly a mis-
spelling of P. yokohamense T. Oka & Ohtani, a hybrid
whose parents are presumed to be P. fibrilloso-paleaceum
(Kodama) Sa. Kurata and P. tagawanum Sa. Kurata. It is
not P. yaeyamense (Makino) Makino, a one-pinnate species
not known in cultivation in the United States.

Psilotum  13.103.1, 2

Whisk fern
The whisk ferns are small- to medium-sized plants with re-
peatedly branched stems bearing inconspicuous scale-like
leaves. Psilotum is a primitive plant with no roots or vascu-
larized leaves, making for interesting study in botany
classes. It is cultivated mostly as a novelty plant. When
grown in greenhouses, the whisk fern tends to volunteer in
pots of other plants, either by spores or by underground Figure 13.103.2. Psilotum nudum: sporangia.
gemmae that spread as the potting soil is reused. The plants
respond well to applications of bonemeal. In the tropics
Psilotum is often found growing in the old leaf bases of naked or smooth, referring to the smooth, nearly naked
palms, where humus has accumulated. branches.
Psilotum is easily recognized by its repeatedly forked
green stems that lack broad, flat green leaves. It grows as an Psilotum nudum (Linnaeus) Palisot de Beauvois
epiphyte or terrestrial plant. Roots are lacking, but the  13.103.1–3
stems bear fine hairs (rhizoids) that absorb water and min- Semi-hardy, Zone 8
eral nutrients. The leaves are small, distant, and scale-like,
A small-medium plant with aerial stems in a cluster arising
without veins. Large, three-lobed sporangia are borne on
from a clump of root-like stems. Grows well under medium
upper branches, and they turn yellow when mature.
to high light in moist-dry garden soil, potting mix, or uncut
The genus, which contains two species, occurs in the
moss with good drainage. Hundreds of cultivars of Psilotum
tropics and subtropics of the world. For more information
nudum have been grown in Japan for several centuries.
on growing these plants, see the section on “Fern Allies” in
Plants are propagated mainly by divisions, as spores may
Chapter 10. The genus name comes from the Greek psilos,
take as long as 3 months to germinate, if at all.
Psilotum nudum has evergreen, erect to semi-pendent
stems, to 50 cm (20 in.) long and triangular in cross section.
The species is widespread in the tropics and subtropics as an
epiphytic or terrestrial plant.
The second species in the genus, Psilotum complanatum
Swartz, can be distinguished by its wider, flattened
branches. It is reportedly difficult to grow and is not found
in the trade, although a hybrid between the two species
from Hawaii, P. ×intermedium W. H. Wagner, shows hor-
ticultural promise.

Pteridium
Bracken
This well-known native of open fields and slopes is a me-
dium- to large-sized terrestrial fern that spreads from a slen-
der, wide-creeping, underground rhizome. Its triangular
fronds are spaced far apart and are several times divided.
Pteridium is rarely used as an ornamental because it spreads
invasively in the garden. It is a terrible weed in many parts
of the world.
The fiddleheads of bracken are harvested in Japan for
Figure 13.103.1. Psilotum nudum: habit. food, and in the United States the packages are sold in some
 F  F A  C 471

Pteridium aquilinum (Linnaeus) Kuhn

 13.104.1–3
Hardy to semi-tender, depending on the variety
A large fern with long-creeping rhizomes and large, trian-
gular, deciduous fronds. Prefers medium to high light or
full sun and moist-dry garden soil. Pteridium aquilinum,
defined in the broad sense, occurs worldwide.
var. caudatum (Linnaeus) Sadebeck. Lacy bracken. Fig-
ure 13.104.3b. Fertile ultimate segments narrow, mostly
only 1–2 mm wide. Semi-tender, Zone 9. Native to Florida
and tropical America.
var. latiusculum (Desvaux) Underwood ex A. Heller.
Eastern bracken. Figure 13.104.3c. Pinnule margins with
shaggy hairs. Very hardy, Zone 3. Native to the United
States, Mexico, Europe, and Asia.
var. pseudocaudatum (Clute) A. Heller. Tailed bracken.
Figure 13.104.3e. Terminal segment of pinnule long and
narrow, often 6–15 times longer than wide, mostly 2–5
mm wide. Grows in acid, sandy soil. Semi-hardy, Zone 7.
Figure 13.103.3. Psilotum nudum: habit, more arching Native to the eastern United States.
form, bar = 5 cm (2 in.). var. pubescens Underwood. Western bracken. Figures
13.104.1, 2, 3a, 3d. Blades densely hairy on the lower sur-
face. Very hardy, Zone 3. Native to North America.
Asian food markets. Nevertheless, eating the fiddleheads is
dangerous. They contain a carcinogen that causes stomach
cancer, and the disease has a higher incidence in areas where
bracken is consumed. Also, the fiddleheads contain thi-
aminase, an enzyme that destroys vitamin B (thiamine).
Bracken has several chemical defenses against insects. It
produces ecdysones, a hormone that promotes molting
(ecdysis) in insects, thus causing the insects to molt prema-
turely and die. It is also cyanogenic, producing hydrogen
cyanide locally where an insect is eating the plant. For a
popular account of the chemical defenses of bracken, see
Moran (1993).
Pteridium can usually be identified by its large (0.5–4.5
m [1.5–15 ft.] long), broadly triangular, two- to four-pin-
nate blades and its long-creeping, hairy rhizomes. It is also
characterized by sori that run along the segment margins
and are covered by a linear enrolled indusium. An unusual
trait is that the sterile segment margins are enrolled to form
a false indusium even when there are no sporangia—Pterid-
ium is the only fern genus that does this. Other character-
istics include fronds set far apart on the rhizome and sori
continuous around the tips and bases of the lobes and
smaller leaflets.
Pteridium traditionally has been construed as containing
a single species with 2 subspecies and 12 varieties world-
wide. Recently, however, pteridologists have recognized
some varieties as species because they do not intergrade Figure 13.104.1. Pteridium aquilinum var. pubescens:
with the other varieties and maintain their distinctness habit.
when growing in mixed populations. The genus name
comes from the Greek pteridion, meaning “a small fern.”
472 C 13

Pteris  13.105.1, 2

Brake fern
Pteris is a genus of small- to large-sized ferns that are popu-
lar and readily available in the trade. The plants grow in
soil or on rocks and are commonly used in terrariums,
planter beds, and indoor pots. They can also be grown out-
doors in mild climates. The fronds are variously divided,
but generally not finely so, and they grow in clusters to give
a full-foliage effect.
The most common species in the trade are the Cretan
brake (Pteris cretica), spider brake (P. multifida), Victorian
brake (P. ensiformis ‘Victoriae’), Australian brake (P. trem-
ula), and rusty brake (P. vittata). Most species are fairly
rapid, vigorous growers adapted to a range of soils. Brake
ferns generally do not like to be kept moist or wet, and
many prefer moist-dry conditions. The Cretan brake and
its cultivars tolerate low humidity indoors. Some of the me-
Figure 13.104.2. Pteridium aquilinum var.
dium-sized or larger species with sparingly branched rhi-
pubescens: sori and indusia.
zomes (mainly those in the P. quadriaurita complex) tend
to lose vigor with age. They also tend to topple, as the rhi-
zome and roots cannot support the foliage. Such plants can
be staked for support, and soil built up around the rhizome
b encourages root development. Alternatively, the entire
plant can be replanted deeper in the soil. Young plants re-
spond better to this replanting. Species with much-
branched rhizomes are easily propagated by divisions.
The rhizomes of Pteris are scaly (not hairy) and vary
from short-creeping to ascending and sparingly to amply
branched. The stipes are clustered, and the blades are usu-
c ally one to several times pinnately divided with narrow,
often sessile or adnate segments. The veins are usually
forked and free; less frequently they have a few areoles or are
completely netted. The sori run along the margins of the

d
e

Figure 13.104.3. Pteridium aquilinum varieties: a. var.

pubescens habit; b. var. caudatum frond tip; c. var.
latiusculum frond tip; d. var. pubescens frond tip; e. var.
pseudocaudatum frond. Bar = 5 cm (2 in.).
Figure 13.105.1. Pteris cretica: habit.
 F  F A  C 473

Figure 13.105.2.
Pteris vittata:
sorus and
indusium.

segments and are covered by an enrolled leaf margin (false

indusium). Several fern genera have marginal indusia that
resemble those of Pteris, namely Histiopteris, Mildella, Pel- Figure 13.105.3. Pteris altissima: frond, bar = 5 cm
laea, and Pteridium. It is easy to misidentify these genera as (2 in.).
Pteris.
Pteris contains about 250 species widespread in the trop-
ics and subtropics of the world. See Joe (1958) and Walker
(1970) for further details on the cultivated species. An an-
cient Greek term for ferns in general, the genus name
comes from the Greek pteron, wing or feather, derived from Figure 13.105.4. Pteris
a fancied resemblance of fern leaves to wings or feathers. argyraea: frond, bar =
5 cm (2 in.).
Pteris altissima Poiret  13.105.3
Forest brake
Tender
A large fern with clustered fronds. Grows well under me-
dium light in moist garden soil or potting mix.
The fronds of Pteris altissima are up to 2.5 m (8 ft.) long
and two- to three-pinnate-pinnatifid, with the basal pin-
nae larger on their lower side. The veins are netted. This
species is native to the American tropics.

Pteris argyraea T. Moore  13.105.4;  47

syn. Pteris quadriaurita var. argentea Beddome, P.
quadriaurita var. argyraea Horticulture
Silver brake, striped brake
Semi-tender
A medium-sized fern with clustered fronds. Grows well
under medium light in moist-dry garden soil or potting
mix.
 474 C 13

The blades of Pteris argyraea resemble those of P. quadri-

aurita but differ by a whitish band down the center of the
pinnae. It also resembles P. biaurita, but that species’s low-
est veins are fused to form an areole on either side of the a
midrib. Pteris argyraea is native to Sri Lanka and Java. The
cultivated plants are said to be robust like the plants from
Java and unlike those from Sri Lanka.

Pteris aspericaulis Wallich ex J. Agardh

 13.105.5
Semi-tender (?)
Medium-large fern with clustered fronds. Grows well un- b
der medium light in moist-dry garden soil or potting mix.
The blades of Pteris aspericaulis resemble those of P.
quadriaurita but differ by having deep red stipes, rachises,
and costae and by the small, nearly erect red spines along
the costae and costules on the upper surface. This species is Figure 13.105.6. Pteris
native to India and the Himalayan region. biaurita: a frond, bar = 5 cm
var. tricolor (Linden) T. Moore ex Lowe (Pteris tricolor (2 in.); b. netted veins along
Linden). The stipes, rachises, and costae are red, but the pinna midrib, bar = 1 cm
pinnules are white in their basal half and green distally. Na- (0.4 in.).
tive to India. The plants under this name in the recent trade
are typical var. aspericaulis. The true var. tricolor has not
been seen in United States cultivation for at least 25 years.

Pteris catoptera Kunze  13.105.7

Semi-hardy to semi-tender
A large fern. Grows well under medium light in moist-wet
to moist garden soil or potting mix.
The blades of Pteris catoptera resemble those of P. qua-
driaurita but differ by pinna apices acute to long-caudate
and the ultimate segments narrow and parallel sided. The
veins are free, not netted. This species is native to Africa.
b
Pteris cretica Linnaeus  13.105.1, 8, 9
Cretan brake
Semi-hardy, Zone 7; may die back in cooler temperatures
Figure 13.105.5. Pteris A small to medium fern with clump-forming rhizomes and
aspericaulis: a. frond, bar = densely clustered fronds. Grows best under high-medium
a 5 cm (2 in.); b. spines at light in moist-dry garden soil or potting mix. This species
pinnule bases, bar = 1 cm is easy to grow.
(0.4 in.). Pteris cretica has dimorphic sterile and fertile fronds. The
sterile blades are one-pinnate, except for the basal pinnae
that are forked once or twice. The rachis is winged between
the upper first or second pinna pairs. The pinnae are serrate,
Pteris biaurita Linnaeus  13.105.6 and the veins are free. The fertile blades have narrower pin-
Tender nae than the sterile ones and tend to be more erect and
Medium-large fern with clustered fronds. Grows well longer petiolate. This species is native to tropics worldwide.
under medium light in moist garden soil or potting mix. This highly variable species has numerous cultivars that
The blades of Pteris biaurita differ from those of P. qua- essentially breed true from spore. Some cultivars are con-
driaurita by having netted (not free) veins on either side of fused with Pteris multifida and other species. Only a few of
the pinna midrib. This species is native to the tropics the many cultivars are listed here.
throughout the world. ‘Albo-lineata’ (var. albolineata). See Pteris nipponica.
 F  F A  C 475

a b

Figure 13.105.8. Pteris cretica: a. fertile frond (left) and

sterile frond (right), bar = 5 cm (2 in.); b. rachis wing
Figure 13.105.7. Pteris catoptera: a. frond, bar = 5 cm limited to upper pinnae pair, bar = 1 cm (0.4 in.).
(2 in.); b. larger pinna with caudate tip, bar = 5 cm
(2 in.); c. veins and spines along midrib of pinna and
pinnules, bar = 1 cm (0.4 in.).
Pteris dentata Forsskål  13.105.10
syn. Pteris flabellata Thunberg
Toothed brake
‘Childsii’. Figure 13.105.9a. Pinnae irregularly fringed,
Semi-hardy to semi-tender
their tips often crested. The plants are sterile, but a similar,
fertile form has recently appeared in the trade. A medium-large fern with clustered fronds and sparingly
‘Distinction’. Blades more open, margins more fringed branched rhizomes. Grows well under medium light in
but with fewer crests than ‘Wimsettii’. moist-dry garden soil or potting mix.
‘Mayii’ (‘Cristata Mayii’). Figure 13.105.9b. Blades Pteris dentata has ovate, two-pinnate-pinnatifid to
weakly variegated, pinnae crested. nearly three-pinnate blades. The ultimate segments and
‘Ouvrardii’. Figure 13.105.9c. Resembles typical Pteris pinnules tend to fold inward along the midrib and have
cretica except larger and with more pinnae. It is a robust margins somewhat coarsely and irregularly serrate-toothed.
grower. This species is native to Africa.
‘Parkeri’. Figure 13.105.9d. Fronds larger than typical
Pteris cretica, the blades dark green, firm, leathery. Pteris ensiformis N. L. Burman  13.105.11
‘Rivertoniana’. Figure 13.105.9e. Margins with irregu- Semi-tender
lar broad, acute to acuminate fringes or segments. A small fern with short- or longer-creeping rhizomes that
‘Roweri’. Figure 13.105.9f. Plants compact, fronds may sparingly branch and form separate clusters of fronds.
crested. This cultivar is questionably distinct from early Grows well under medium light in moist-dry garden soil or
crested forms. potting mix. The typical form of the species is infrequently
‘Wilsonii’. Figure 13.105.9g. Plants compact, pinnae grown, although the cultivar ‘Victoriae’ is common.
crested. Pteris ensiformis has dimorphic sterile and fertile leaves.
‘Wimsettii’. Figure 13.105.9h. Pinnae irregularly The sterile blades are triangular to ovate and two-pinnate-
fringed, the fringes long to very long and narrow; the tips of pinnatifid, with the ultimate divisions oval-elliptic to more
the pinnae and some of the larger fringes are split or crested. elongate. The fertile blades are long-stalked with the ulti-
 476 C 13

c
a
d
b

h
e
f g

Figure 13.105.9. Pteris cretica cultivars: a. ‘Childsii’; b. ‘Mayii’; c. ‘Ouvrardii’; d. ‘Parkeri’; e. ‘Rivertoniana’; f. ‘Roweri’; g.
‘Wilsonii’; h. ‘Wimsettii’. Bar = 5 cm (2 in.).

mate divisions narrowly linear. This species is native to near the midrib but netted toward the margin. This spe-
India, China, Malaysia to Australia, and Polynesia. cies is native to the American tropics.
‘Evergemiensis’. Variegation more cream-white than in
‘Victoriae’, and the white extends between the lateral veins. Pteris longifolia Linnaeus  13.105.13
‘Victoriae’. Victorian brake. Figure 13.105.11b. Cream- Ladder brake
white band down the center of pinnae and pinnules, with Tender
only some white extending along the lateral veins. A medium fern with short-creeping rhizomes and clustered
fronds. Grows well under medium to high light in moist
Pteris grandifolia Linnaeus  13.105.12 garden soil or potting mix.
Tender Pteris longifolia has oblanceolate, one-pinnate blades.
A large fern with short-creeping rhizomes and clustered The stipes and rachises of unweathered plants are amply
fronds. Grows well under high-medium light in moist gar- covered with scales and hairs. The pinnae are numerous,
den soil or potting mix. Too large for most gardens, this simple, and jointed to the rachis. The veins are free. The in-
huge plant is grown only in south Florida. dusium is erose-fimbriate. This species is native to the
Pteris grandifolia has one-pinnate blades throughout, United States (Florida), Mexico, Guatemala, and the West
and the pinnae are simple and entire. The veins are free Indies.
 b

a b

a
Figure 13.105.10. Pteris Figure 13.105.12. Pteris grandifolia: a. frond, bar = 5 cm
dentata: a. frond, bar = (2 in.); b. pinnae, bar = 5 cm (2 in.).
5 cm (2 in.); b. pinna tip,
bar = 1 cm (0.4 in.).

b
a

Figure 13.105.13. Pteris

longifolia: frond, bar = 5 cm
(2 in.).

Figure 13.105.11. Pteris ensiformis: a. fertile frond (left)

and sterile frond (right), bar = 5 cm (2 in.); b. ‘Victoriae’
fertile frond (center) and sterile fronds (left and right);
bar = 5 cm (2 in.).
 478 C 13

Most of the plants sold in the United States under this winged only between two upper pinna pairs or less.) This
name are actually Pteris vittata. The two species are often species is native to China and Japan.
difficult to distinguish. The best characteristic to separate A few crested and compact-growing cultivars are some-
them is the jointed pinna bases of P. longifolia. These bases times seen.
have a dark abscission layer where the pinna attaches to the
rachis, and the juncture is usually thickened. (Pteris vittata Pteris nipponica W. C. Shieh  13.105.15
never has thickened rachises at the juncture with the pin- syn. Pteris cretica var. albolineata Hooker
nae.) These characteristics are often hidden by scales, which Semi-hardy, Zone 8
must be scraped away to identify the plant. A small to medium fern with short- to medium-creeping
rhizomes and fronds forming somewhat open clumps.
Pteris multifida Poiret  13.105.14 Grows well under medium light in moist to moist-dry gar-
syn. Pteris serrulata Linnaeus f. den soil or potting mix. This species is more commonly
Spider brake known by its synonym: Pteris cretica var. albolineata (or
Hardy, Zone 6 ‘Albo-lineata’).
A small fern with clump-forming rhizomes and fronds in Pteris nipponica is distinguished by a pale band of tissue
dense, irregular clusters. Grows well under medium light in on both sides of the pinna midrib. It differs from P. cretica
moist to moist-dry garden soil or potting mix. The plants by having a more open-spreading habit, by the absence of a
tend to be weedy. distinct wing on the distal part of the rachis, and by other
Pteris multifida resembles P. cretica but differs by nar- microscopic spore details. This species is native to Japan,
rower pinnae and the rachis fully winged between each of South Korea, and Taiwan.
the three upper pinna pairs. (Pteris cretica is unwinged or
Pteris quadriaurita Retzius  13.105.16
Semi-tender
A medium-large fern with clustered fronds and infrequently
branched, short-creeping rhizomes. Grows best under me-
dium light in moist-dry garden soil or potting mix.
The blades of Pteris quadriaurita are herbaceous and
one-pinnate-pinnatifid except for the basal pinnae that are
forked at the base. The costa (but not the costules) bears

a
b
b

Figure 13.105.15. Pteris nipponica: a. habit, with sterile

Figure 13.105.14. Pteris multifida: a. fertile frond (upper frond (left) and fertile frond (right), bar = 5 cm (2 in.);
left), others sterile, bar = 5 cm (2 in.); b. rachis winged b. rachis not winged below terminal pinna, bar = 1 cm
to or just above the base, bar = 1 cm (0.4 in.). (0.4 in.).
 F  F A  C 479

Figure 13.105.17. Pteris

semipinnata: frond, bar =
5 cm (2 in.).

a Figure 13.105.16. Pteris

quadriaurita: a. frond, bar =
5 cm (2 in.); b. free veins
along pinna midrib, bar =
1 cm (0.4 in.).

small appressed scales. The veins are free, not netted, along
the midrib. This species is native to tropics worldwide.
Pteris quadriaurita comprises a complex of variable
plants not fully studied. Plants examined in the trade la-
beled as P. fauriei Hieronymus are actually P. quadriaurita.
True P. fauriei, native to eastern Asia, has leathery fronds
and pinnae that are widest in the middle with caudate
apices.

Pteris semipinnata Linnaeus  13.105.17

Semi-hardy to semi-tender
A medium-sized fern with clustered fronds. Grows well
under medium light in moist garden soil or potting mix.
Pteris semipinnata has distinctive pinnae: the larger ones
are straight and entire on the upper side, but the lower side
is lobed. The blades are oblong-triangular and one-pinnate-
pinnatifid. This species is native to Japan, Korea, Taiwan,
southwestern China, the Himalayas, and Asian tropics.
var. dispar (Kunze) Hooker & Baker (Pteris dispar
Kunze). A form with lobed upper pinna margins.

Pteris tremula R. Brown  13.105.18

Australian brake
Semi-hardy
A large fern with densely clustered fronds and clump-form-
ing rhizomes. Grows well under low to high-medium light
or full sun in some areas in moist-dry garden soil or potting
mix. The plants grow rapidly and are easy to cultivate. They
often volunteer from spores. This is one of the more finely Figure 13.105.18. Pteris tremula: frond, bar = 5 cm
divided species of Pteris. (2 in.).
 480 C 13

The blades of Pteris tremula are triangular, membran- Pteris vittata Linnaeus  13.105.2, 20
ous, and two- to four-pinnate-pinnatifid. The ultimate seg- Chinese brake, rusty brake
ments are narrowly oblong to more linear and serrate with Semi-hardy, Zone (7)8
blunt teeth. The fertile segments are narrower than the ster- A medium-large fern with short-creeping rhizomes and
ile ones. The veins are free, not netted. This species is native clustered fronds. Grows well under high-medium light to
to Australia, Tasmania, New Zealand, and Fiji. full sun in moist-dry garden soil or potting mix. The plants
are easy to start from spore, but they tend to become weedy.
Pteris tripartita Swartz  13.105.19 They grow on old masonry and bricks, as can be seen in
Trisect brake the French Quarter in New Orleans.
Tender, Zone 9 Pteris vittata is characterized by one-pinnate blades wid-
A large fern, to 2 m (6.5 ft.) tall, with clustered fronds and est above the middle, with narrow pinnae and a terminal
clump-forming rhizomes. Grows well under medium light pinna that resembles the lateral ones. The stipes and
in continually moist-wet garden soil or potting mix. The rachises are sparsely scaly-hairy, and the pinnae are not
plants grow rapidly from spores but become too big for jointed to the rachis. The veins are free, and the indusia
most gardens. have entire margins. This species greatly resembles Pteris
Pteris tripartita has broadly triangular blades divided longifolia. It is native to eastern Asia but has become widely
into three parts: the central part is ovate-triangular and one- naturalized throughout the tropics and subtropics of the
pinnate-pinnatifid, and the two lateral parts (the basal pin- world.
nae) are as long as the central part, and each forks twice on
the lower side. The veins are netted. This species is native to
the Old World but has become naturalized in tropical
America.

Figure 13.105.20. Pteris vittata:

frond, bar = 5 cm (2 in.).

b
Pyrrosia  13.106.1–3
syn. Cyclophorus, Drymoglossum, Niphobolus
Felt fern
a
Most Pyrrosia species are small-medium-sized and have
simple, entire blades (a few are deeply lobed) with star-
shaped hairs. Be careful not to overwater pyrrosias. The
Figure 13.105.19. Pteris tripartita: a. small frond, bar = epiphytic species in particular need a coarser soil mix and
5 cm (2 in.); b. large frond, bar = 15 cm (0.5 ft). good drainage. The plants can be easily propagated by di-
vision of the rhizomes.
 F  F A  C 481

Figure 13.106.1. Pyrrosia lingua: habit.

Figure 13.106.2. Pyrrosia lingua: fronds.

Pyrrosia is best known in cultivation by the Japanese felt

fern or tongue fern (Pyrrosia lingua). Its tongue-shaped
fronds are densely covered beneath with a felt-like mat of
star-shaped hairs. This species is hardy outdoors in warm-
temperate areas, where it is usually grown in the ground. If
grown in pots, baskets, or on boards, it may need more pro-
tection during the winter. The fronds are dark green, leath- Figure 13.106.3.
ery, and durable, tolerating low humidity. The plants grow Pyrrosia lingua:
moderately slowly, which is an advantage because they do sori.
not require frequent repotting. Some of the cultivars have
an attractive crested habit.
Although less hardy, Pyrrosia hastata is also prized in
cultivation. It has mostly hastate fronds borne in loose clus-
ters. It can be grown outdoors in warm-temperate climates
but also makes a good indoor plant.
Pyrrosia is distinguished by simple or lobed blades with
star-shaped hairs on the surfaces, especially the lower sur-
face. The rhizomes are scaly and vary from short- to long-
creeping, bearing phyllopodia to which the fronds are at- Pyrrosia is native to the Old World tropics and temper-
tached and jointed. The fronds may be monomorphic or ate areas and contains about 50 mostly epiphytic or epi-
dimorphic, and they have a parchment-like to leathery or lithic species. The genus name comes from the Greek pyr,
succulent texture. The veins are netted, and the areoles con- fire, alluding to the dull reddish, scaly covering on the blade
tain included veinlets, although the veins are often hard to of the type species (Pyrrosia chinensis Mirbel).
see because of the thick blades. The sori vary from round to
elongate, or sometimes merging. An indusium, as in all Pyrrosia angustata (Swartz) Ching  13.106.4
polypodiaceous ferns, is absent. Tender
The fronds can vary tremendously within a species, due A medium-sized fern with medium-creeping rhizomes.
to cultural or genetic factors. Also, in those species with di- Grows well under medium light in moist-dry potting mix
morphic fronds, the sterile and fertile fronds can intergrade or uncut moss. This species is easy to grow.
with one another. For a technical treatment of the species of Pyrrosia angustata has dimorphic sterile and fertile
Pyrrosia, see Hovenkamp (1986). fronds, with the sterile blades oblong-lanceolate, 12–25 cm
 482 C 13

a b

Figure 13.106.4.
Figure 13.106.5. Pyrrosia confluens: a. var. confluens
Pyrrosia angustata:
habit, bar = 5 cm (2 in.); b. var. confluens sori, bar =
a. habit, bar = 5 cm
1 cm (0.4 in.); c. var. dielsii habit, bar = 5 cm (2 in.).
(2 in.); b. sori, bar =
1 cm (0.4 in.).

var. dielsii (C. Christensen) Hovenkamp (Pyrrosia diel-

sii (C. Christensen) Tindale). Silvery felt fern. Fronds dis-
tinctly dimorphic, leathery and often brittle when dry, apex
wide, often truncate or emarginate, hydathodes marginal;
sori separate, round to oval, rarely confluent. Native to Aus-
a
tralia. Though listed here, this variety is doubtfully culti-
vated in the United States. The fronds examined by the au-
thor were sterile, and the distinctness of the sori could not
be confirmed.

Pyrrosia costata (Wallich ex C. Presl) Tagawa &

(5–10 in.) long, and the fertile ones narrower and longer. K. Iwatsuki  13.106.6
Both kinds of fronds are covered with a loose, woolly de- syn. Pyrrosia beddomeana (Giesenhagen) Ching
ciduous down. The sori are 5–10 mm (0.2–0.4 in.) wide Tender
and vary from round to elongate, being deeply sunken in A medium-sized fern with short-creeping rhizomes. Grows
the lower surface of the blade and raised (embossed) on the under medium light in moist-dry potting mix or uncut
upper surface. This species is native to Thailand, Sumatra, moss with good drainage. The plants are slightly difficult to
Malaya, Borneo, and the Moluccas. grow. This species requires higher temperatures than most
species of Pyrrosia.
Pyrrosia confluens (R. Brown) Ching  13.106.5 Pyrrosia costata has stout rhizomes over 5 mm (0.2 in.)
Tender to slightly hardier thick. The stipes are absent or poorly developed, and the
A small fern with medium-creeping rhizomes. Grows well blades are crowded, widest above the middle, more than
under medium light in moist-dry potting mix or uncut 30 cm (12 in.) long and 3 cm (1.2 in.) wide, dark green
moss with good drainage. above, and rusty tan below. The sori are closely packed and
The rhizome scales of Pyrrosia confluens are ciliate to- usually apical on the blade. This species is native to south-
ward the apices. The sterile and fertile fronds vary from eastern Asia.
moderately to distinctly dimorphic. Both have narrowed Pyrrosia costata might be confused with P. stigmosa. The
apices (var. confluens), leathery texture, and hydathodes on latter has elongate rhizomes, long stipes, and costae and
or within the margin. The fertile fronds are longer. The sori secondary veins narrowly grooved on the upper surface in-
are apical. If separate, the sori are oblong and in a single stead of flat or shallowly and broadly grooved. Broad frond
row; if confluent, they form a continuous band around the forms of P. costata might be confused with P. princeps and
apex of the frond. This species is native to Australia, Nor- P. splendens. Pyrrosia princeps has dentate or ciliate rhizome
folk Island, New Caledonia, Loyalty Islands, and Tahiti. scales and a central bundle of dark support tissue in its stipe;
var. confluens. Fronds moderately dimorphic, leathery, in contrast, P. costata has entire rhizome scales and no cen-
apex narrow, hydathodes submarginal; sori separate or tral support tissue. Pyrrosia splendens has blade hairs with
confluent. long erect spines; these are absent in P. costata.
 F  F A  C 483

Figure 13.106.7.
Pyrrosia eleagnifolia:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
b 1 cm (0.4 in.).

b
Pyrrosia fallax (Alderwerelt) M. G. Price
 13.106.8
Tender (?)
A small fern with medium-creeping rhizomes. Grows well
a under medium light in moist-dry potting mix or uncut
moss with good drainage.
Figure 13.106.6. Pyrrosia costata: a. habit and larger Pyrrosia fallax has slender rhizomes covered by round-
frond, bar = 5 cm (2 in.); b. stipe cross section, bar = ish, dark scales with pale erose-ciliate margins. The phyl-
1 mm. lopodia are about 1–1.5 cm (about 0.5 in.) apart, and lat-
eral buds occurs on the rhizome about midway between
phyllopodia. The fronds are dimorphic. The sterile ones
are obovate or rounder, 1–4.5 cm (0.5–2 in.) long, and
Pyrrosia eleagnifolia (Bory) Hovenkamp nearly glabrous above but with pale hairs below. The fertile
 13.106.7 fronds are narrowly linear, about 4–10 cm (1.5–4 in.) long.
syn. Pyrrosia rupestris of authors, not (R. Brown) Ching, The sori are arranged in two or three irregular rows or are
P. serpens of authors, not (N. L. Forster) Ching
Semi-tender to hardier
A small fern with medium-creeping rhizomes. Grows
under medium light in moist-dry potting mix or uncut
moss with good drainage. The plants are difficult to estab- Figure 13.106.8. Pyrrosia fallax: a. habit,
lish but otherwise are tough and adaptable to drier condi- bar = 5 cm (2 in.); b. fertile frond with
tions. linear sori, bar = 1 cm (0.4 in.).
Pyrrosia eleagnifolia has entire to dentate rhizome scales,
and the fronds are dimorphic, thick-leathery, and densely
covered with tan hairs on the lower surfaces. Hydathodes
are absent. The sterile fronds are almost rounded, rarely to
3 cm (1.2 in.) long, and the fertile ones are longer and nar-
rower. The distinctly sunken sori are round, slightly spaced,
and appear in two or three rows. Native to New Zealand. a
This species was recently introduced from New Zealand
and has shown promise in warm-temperate areas. Botanists
have confused it with Pyrrosia rupestris (R. Brown) Ching of
Australia and P. serpens (N. L. Forster) Ching of the South b
Pacific. Pyrrosia rupestris differs in having hydathodes. Pyr-
rosia serpens differs in having two rows of sori with the sec-
ond row weakly developed. Pyrrosia serpens is doubtfully
cultivated in the United States.
 484 C 13

confluent along the distal margin of the frond. This spe-

cies is native to Ambon eastward to Solomon Islands.
The cultivated plants named as this species are only ten-
tatively identified as such. They are grown in scattered fern
collections in Florida and reportedly were collected in New
Guinea.

Pyrrosia hastata (Thunberg ex Houttuyn) Ching

 13.106.9
syn. Pyrrosia tricuspis (Swartz) Tagawa
Semi-hardy to hardier, Zone 8
A small-medium fern with short-creeping rhizomes. Grows
well under medium light in moist-dry garden soil or pot-
ting mix with good drainage. This species is easy to grow in
baskets or in the ground.
The stipes of Pyrrosia hastata are less than 1 cm (0.5 in.)
apart, and the blades are typically hastately lobed—they
may have up to five lobes, with the outer lobes much
smaller—and have star-shaped hairs to only 0.5 mm in di-
ameter. This species is native to South Korea and Japan.
Pyrrosia hastata plants with five-lobed fronds may be Figure 13.106.9. Pyrrosia hastata: frond variations, bar =
confused with P. polydactyla. In cultivated plants the stipe 5 cm (2 in.).
and main veins near the base of the blade are pale green in
P. hastata and black in P. polydactyla. The star-shaped hairs
of P. polydactyla are also slightly larger than those of P. has- submarginal, usually in a row on each side of the frond on
tata, up to 1 mm in diameter. A variant of P. hastata with the upper half. This species is native to southeastern China,
small hastate blades is in cultivation. Taiwan, North and South Korea, and Japan.

Pyrrosia lanceolata (Linnaeus) Farwell Pyrrosia lingua (Thunberg) Farwell

 13.106.10  13.106.1–3, 12
syn. Pyrrosia adnascens (Swartz) Ching, P. varia Japanese felt fern, tongue fern
(Kaulfuss) Farwell Hardy, Zone 6
Tender A small fern with medium- to long-creeping rhizomes.
A small fern with medium-creeping rhizomes. Grows well Grows well under medium to high (indirect) light in moist-
under medium-high light in moist-dry potting mix or dry potting mix or uncut moss with good drainage. The
uncut moss with good drainage. The plants are easy to
grow.
Pyrrosia lanceolata has slender rhizomes clothed by lan- b
a
ceolate scales with pale marginal hairs. The fronds are di-
morphic: the sterile ones are oval to ovate-lanceolate, to 24
cm (10 in.) long, 4 cm (1.5 in.) wide, whereas the fertile
ones are narrower and longer. The sori are about 1 mm
wide in weakly defined, oblique rows. This species is na-
tive to tropics of the Old World east to Polynesia.

Pyrrosia linearifolia (Hooker) Ching

 13.106.11
Hardy to semi-hardy, Zone 8
A small fern with medium-creeping rhizomes. Grows well
under low-medium light in moist-dry potting mix or uncut
moss with good drainage. The species can survive mini-
mum winter temperatures of −7°C (20°F)
The blades of Pyrrosia linearifolia are widest above the Figure 13.106.10. Pyrrosia lanceolata: a. habit,
middle and up to 9 cm (3.5 in.) long. The sori are round, bar = 5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.).
 F  F A  C 485

Figure 13.106.11. Pyrrosia

linearifolia: habit, bar =
5 cm (2 in.).

plants are easy to grow in pots, baskets, on boards, or in the

ground. The fronds are an attractive dark green.
Pyrrosia lingua has moderately slender rhizomes 1.2–3.7 a b
mm thick with phyllopodia spaced 2–8 cm (0.75–3 in.)
apart. The blades vary from lanceolate to oblong to elliptic-
lanceolate. They can grow up to 50 cm (20 in.) long and 5
cm (2 in.) wide but are usually only half this length. The
sori are many, round, 1 mm in diameter, and closely placed
between visible lateral veins. This species is native to China,
southeastern Asia, and India.
There are two botanical varieties of Pyrrosia lingua:
var. heteractis (Mettenius ex Kuhn) Hovenkamp (Pyr-
rosia heteractis (Mettenius ex Kuhn) Ching). Figure
13.106.12b. Blades broader than the type variety, apices
cuspidate, with star-shaped hairs of two types, those in the
upper layer with boat-shaped rays, those in the lower layer
with mainly woolly rays.
var. lingua. Figure 13.106.12a. Frond apices acute or c d
subobtuse, lower layer of star-shaped hairs with woolly rays
absent.
Many cultivars of Pyrrosia lingua var. lingua are grown,
differing in such characteristics as amount of cresting, type
of emargination, and blade width. Successive fronds can
differ in shape. The fancy-foliage forms often grow more
slowly than other forms. A few samples of the many culti-
vars include the following:
‘Cristata’. Crested tongue fern, Obake pyrrosia. Figure
13.106.12c. Blade tips several times irregularly forked, the
forks not greatly twisted. e f
‘Monstrifera’. Lacerate pyrrosia. Figure 13.106.12d.
Blade margins lacerate.
‘Nankin-shishi’. Peacock pyrrosia. Figure 13.106.12e.
Blade tips many times irregularly forked, the forks con-
gested and twisted.
‘Nokogiri-ba’. Figure 13.106.12f. Blades serrate-crenate. Figure 13.106.12. Pyrrosia lingua: a. var. lingua habit,
with fertile frond on left; b. var. heteractis habit;
c. ‘Cristata’; d. ‘Monstrifera’; e. ‘Nankin-shishi’;
f. ‘Nokogiri-ba’. Bar = 5 cm (2 in.). After Hoshizaki
(1981).
 486 C 13

Pyrrosia longifolia (N. L. Burman) C. V. Morton

 13.106.13
Tender to hardier (?)
A medium-sized fern with medium-creeping to sometimes
longer creeping rhizomes. Grows well under high light in
moist-dry to dry potting mix or uncut moss with good
drainage. The plants are easy to grow and tolerate irregular
watering. The long, strap-shaped, pale, leathery fronds are
distinct features of this fern in cultivation.
Pyrrosia longifolia has entire rhizome scales. The fronds
are long, leathery, and narrowly strap-shaped, to 60 cm (24
in.) long and 3.5 cm (1.5 in.) wide. This species is native
from southern China to southeastern Asia, to Indonesia, a
Micronesia, the Philippines, and Australia.
A smaller, slightly thinner textured form is also found in
cultivation. It reportedly was imported from New Guinea.
b

Figure 13.106.14. Pyrrosia nummulariifolia: a. habits of

different plants, bar = 5 cm (2 in.); b. rhizome scale, bar
a = 0.5 mm, after Hoshizaki (1981).

long, and the fertile ones are longer, narrower, and more
erect. The sori are mostly 1 mm wide, three or four on each
c
side of midrib, scattered, and sometimes fusing. This spe-
cies is native to India, southeastern Asia to Indonesia, and
the Philippines. The forms with longer fronds were for-
d merly called P. obovata.

Figure 13.106.13. Pyrrosia longifolia: a. fronds, bar = Pyrrosia piloselloides (Linnaeus) M. G. Price
 13.106.15
5 cm (2 in.); b. sori, bar = 1 cm (0.4 in.), after Hoshizaki
syn. Drymoglossum piloselloides (Linnaeus) C. Presl
(1981); c. rhizome and stipe bases, bar = 1 cm (0.4 in.),
Tender
after Hoshizaki (1981); d. rhizome scales, bar = 1 mm.
A small fern with medium-creeping rhizomes. Grows well
under medium light in moist-dry potting mix or uncut
moss with good drainage. The plants are easy to grow and
Pyrrosia nummulariifolia (Swartz) Ching suitable for terrariums.
 13.106.14 The rhizome scales of Pyrrosia piloselloides are dark with
syn. Pyrrosia obovata (Blume) Ching pale margins and bear long hairs. The fronds are dimor-
Tender phic, thick and fleshy, glossy green, and have deciduous
A small fern with medium-creeping rhizomes. Grows well hairs. The sterile fronds are roundish-oblong, sessile or
under medium light in moist-dry potting mix or uncut nearly so, and to 6 cm (2.5 in.) long by 2 cm (0.75 in.)
moss with good drainage. The plants are easy to grow and wide. The fertile fronds are short stalked, linear-oblong,
make an ideal terrarium plant. and to 12 cm (4.75 in.) long by 1 cm (0.5 in.) wide. The
Pyrrosia nummulariifolia has fleshy, dimorphic fronds. sori occur in a marginal band around the apex. This species
The sterile fronds are roundish, mostly to 2 cm (0.75 in.) is native from India to Malaysia and China (Hainan).
 F  F A  C 487

Figure
13.106.16.
a Pyrrosia
polydactyla:
frond, bar =
5 cm (2 in.).
c

Figure 13.106.15. Pyrrosia piloselloides: a. habit, bar =

5 cm (2 in.); b. habit, with sterile frond (left) and fertile
frond (right), bar = 1 cm (0.4 in.); c. rhizome scale, bar
= 0.5 mm. After Hoshizaki (1981).

This species might be confused with Pyrrosia nummula- b

riifolia, which has persistent instead of deciduous hairs on
the fronds and different rhizome scales.

Pyrrosia polydactyla (Hance) Ching Figure 13.106.17.

 13.106.16;  48 Pyrrosia porosa: a. habit;
Semi-tender b. fertile frond, with a
A small-medium fern with short-creeping rhizomes. Grows sori covering underside.
well under medium light in moist-dry garden soil, potting Bar = 5 cm (2 in.).
mix, or uncut moss with good drainage. The plants are easy
to grow and are suitable in baskets or the ground.
Pyrrosia polydactyla resembles P. hastata, but its blades
have six to eight deep, pedately divided lobes and the stipe The differences between Pyrrosia porosa and P. davidii
and main veins near the base of the blade are blackish. This are insufficient to recognize the latter (Hovenkamp 1986).
species is native to Taiwan. In their extreme forms, P. porosa is larger and inconsistently
has two kinds of star-shaped hairs, whereas P. davidii is
Pyrrosia porosa (C. Presl) Hovenkamp smaller and consistently has two kinds of star-shaped hairs.
 13.106.17
syn. Pyrrosia davidii (Giesenhagen ex Diels) Ching, P. Pyrrosia princeps (Mettenius) C. V. Morton
pekinensis (C. Christensen) Ching  13.106.18
Semi-tender or hardier Tender
A small fern with medium-creeping rhizomes. Grows well A medium-large fern, to 1 m (3 ft.) by 10 cm (4 in.), with
under medium light in moist-dry potting mix or uncut short-creeping rhizomes and tightly clustered fronds.
moss with good drainage. Grows well under medium light in moist-dry potting mix
The rhizomes of Pyrrosia porosa are 1.6–3.1 mm wide, or uncut moss with good drainage.
with the stipes spaced 3–7 mm (0.1–0.3 in.) apart. The lan- The rhizomes of Pyrrosia princeps are stout, compact,
ceolate, entire blades are 9–31 cm (3.5–12.5 in.) long and and covered by trough-like, dentate or ciliate scales. The
0.3–3.5 cm (0.1–1.5 in.) wide with scattered hydathodes. fronds are close together and sessile. Narrowly oblanceo-
The sori are 1–2 mm wide and clustered toward the apex late blades are 24–125 cm (10–50 in.) long by 3.5–12 cm
or spread all over the blade. This species is native from (1.5–4.75 in.) wide, gradually narrowed to the base, with
the Himalayan area to India, southeastern Asia, and the mostly acuminate apices. The upper layer of stellate hairs
Philippines. has spreading needle-like rays. The hydathodes are distinct
 488 C 13

b
b
a
Figure 13.106.19. Pyrrosia
rupestris: a. habit, bar = 5 cm
(2 in.); b. fertile frond with
sori, bar = 1 cm (0.4 in.).
After Hoshizaki (1981).

c
with conspicuous hydathodes in a row near the apex. The
fertile fronds are longer and narrower than the sterile ones,
mostly to 8–20 cm (3–8 in.) long. The sori are relatively
Figure 13.106.18. Pyrrosia princeps: a. habit, showing
large (1–2.5 mm wide) and slightly sunken, appearing in
frond with fertile area, bar = 5 cm (2 in.); b. trough-like
one to four rows on each side of the midrib. This species is
rhizome scale with marginal teeth, bar = 0.5 mm; c. stipe
native to Australia.
cross section, with the stippled area showing the dark,
thick-walled cells (sclerenchyma) that strengthen the
stipe, bar = 1 mm. Pyrrosia samarensis (C. Presl) Ching
 13.106.20
Tender
A small-medium fern with medium- to longer-creeping
and scattered. The round sori are 0.5 mm wide and densely rhizomes. Grows well under medium-high light in moist-
packed between the main veins in the apical part of blade. dry potting mix or uncut moss with good drainage. The
This species is native to Sulawesi, the Moluccas, and New plants are easy to grow.
Guinea. Pyrrosia samarensis has dimorphic sterile and fertile
Pyrrosia princeps can be confused with P. splendens, a fronds covered with loose white hairs that eventually fall
species that differs by having entire rhizome scales and off. The sterile fronds are narrowly lanceolate and tapered
blade hairs with a distinct, long, spine-like ray. at both ends; the fertile ones are narrowed apically and cov-
ered beneath with roundish sori running together. This
Pyrrosia rupestris (R. Brown) Ching species is native to the Philippines.
 13.106.19 Pyrrosia angustata also has loose, deciduous hairs but dif-
Rock felt fern fers by a more erect habit, thicker and grooved stipes, acute
Semi-tender to hardier (?) blade apices, and discrete, sunken sori.
A small fern with medium- to long-creeping rhizomes.
Grows well under medium light in moist-dry potting mix Pyrrosia sheareri (Baker) Ching  13.106.21
or uncut moss with good drainage. Semi-tender or hardier (?)
Pyrrosia rupestris has entire rhizome scales without mar- A medium-sized fern with short-creeping rhizomes. Grows
ginal hairs. The fronds are dimorphic, with the sterile ones well under medium-high light in moist-dry potting mix or
to about 7 cm (3 in.) long, typically angular-rhomboid, uncut moss with good drainage. The plants grow slowly.
 F  F A  C 489

The rhizomes of Pyrrosia sheareri are up to 7 mm (0.3

in.) thick. The stipes are about half to as long as the blades,
b which are narrowly ovate, to 50 cm (20 in.) long, leathery,
acuminate at the apex, and cordate-truncate to wedge-
shaped at the base, often with lobes or teeth. Hydathodes
are evident and scattered on the upper surface. The sori are
round, small, and densely scattered to confluent. This spe-
cies is native to China, Taiwan, and Vietnam.
a
Pyrrosia splendens (C. Presl) Ching  13.106.22
Tender
A medium-sized fern with short-creeping rhizomes. Grows
well under medium light in moist-dry potting mix or uncut
moss with good drainage.
Pyrrosia splendens has rhizomes up to 8 mm (0.3 in.)
thick. The rhizome scales are somewhat shiny red-brown
and entire. The fronds are close together, widest above the
middle, and 33–100 cm (13–40 in.) long by 3.2–11.5 cm
(1.2–4.5 in.) wide. The hairs on the upper surfaces of the
c
blades have a distinct, long, more or less erect, spine-like ray
about two to four times longer than the other appressed
rays. This species is native to the Philippines. It resembles P.
princeps.
Figure 13.106.20. Pyrrosia samarensis:
a. habit, bar = 5 cm (2 in.); b. sori,
a
bar = 1 cm (0.4 in.); c. rhizome and
stipe bases, bar = 1 cm (0.4 in.).

Figure 13.106.21. Pyrrosia c

sheareri: frond, bar = 5 cm
(2 in.).

Figure 13.106.22. Pyrrosia

splendens: a. frond with sori,
bar = 5 cm (2 in.); b. rhizome
scale, bar = 0.5 mm; c. stellate
hair of blade, with erect spine,
bar = 0.5 mm.
b
 490 C 13

Pyrrosia stigmosa (Swartz) Ching  13.106.23 creeping and bear fronds 1–3 cm (0.5–1.2 in.) apart. At
Tender the top of the stipe are two leaflets that have veins free or
A medium-sized fern with short-medium-creeping rhi- netted at their ends.
zomes. Grows well under medium-high light in moist-dry The genus has only one species, native to southern Bra-
potting mix or uncut moss with good drainage. zil and adjacent Argentina. The genus name honors André
The rhizomes of Pyrrosia stigmosa are about 4.5 mm (0.2 Frederick Regnell (1807–1884), a Swedish botanist who
in.) thick, with the stipes spaced 1–2 cm (0.5–0.75 in.) collected and studied plants in Brazil.
apart. The fronds are 11–48 cm (4.5–19 in.) long by 1.5–
5.2 cm (0.6–2 in.) wide. They are wedge-shaped toward Regnellidium diphyllum Lindman  13.107.1
the base and acuminate to rounded at the apex. The hyda- Very tender
thodes are distinct. The sori are about 0.5 mm wide and A small fern with medium-creeping rhizomes. Grows well
scattered. This species is native to southeastern Asia and under high light in moist-wet soil (a mixture of sand and
Indonesia. peat) or fully submerged. The plants are typically grown in
Pyrrosia stigmosa is similar to P. costata, but the latter pots set in water or in an aquarium with plants partly sub-
species has a stouter rhizome and little or no stipe. merged. Regnellidium diphyllum is easy to grow but apt to
die if the temperature drops below 21°C (70°F). Further
Regnellidium characteristics are described under the genus.
Horticulturists interested in aquatic plants might consider
growing Regnellidium. The genus is related to Marsilea, the Rumohra
clover fern, but differs by having two leaflets instead of four. Leather fern
The leaves are produced too far apart on the rhizome to The genus Rumohra consists of medium-sized, terrestrial or
make an attractive pot plant; the plants are mainly used as epiphytic ferns with broad, dark green, coarsely divided,
a novelty in aquariums. leathery fronds. The durable fronds are used extensively by
Regnellidium roots in mud, although sometimes it is florists in cut flower arrangements. The plants grow in beds
submerged and the fronds are floating. The rhizomes are or pots or baskets and require only moderate watering and
humidity. Over the years, commercial growers, mainly lo-
cated in Florida, have made many selections from their
stock, and today’s leather fern looks quite different from
wild plants, though the wild plants vary considerably as well.
The rhizomes of Rumohra are medium-creeping,
branched, and covered by peltately attached scales. The

Figure 13.106.23. Pyrrosia

stigmosa: habit, showing
fertile area distal on frond,
bar = 5 cm (2 in.). Figure 13.107.1.
Regnellidium
diphyllum: habit,
bar = 5 cm
(2 in.).
 F  F A  C 491

glabrous fronds are broadly triangular and three-pinnate,

with the basal pinnae the largest. The sori are covered by a
round, peltate indusium.
This genus is distinguished from the similar Polystichum
by its peltately attached rhizome scales, large basal pinnae,
and medium-creeping rhizomes. In contrast, Polystichum
has basally attached rhizome scales, fronds narrowed at the
base, and erect rhizomes. Rumohra is currently believed to
be related to the dryopteroid ferns, not to the davallioids as
previously thought.
Rumohra consists of seven species and is native to the
tropics of the Southern Hemisphere. The genus name hon-
ors Karl F. von Rumohr (1785–1843), an art student from
Dresden, Germany.
Figure 13.108.1. Rumohra adiantiformis: habit.
Rumohra adiantiformis (G. Forster) Ching
 13.108.1, 2
syn. Polystichum adiantiforme (G. Forster) J. Smith, P.
capense J. Smith, P. coriaceum Schott
Leather fern
Semi-hardy, Zone 8 a
A medium-sized fern with short- to medium-creeping rhi- b
zomes. Grows well under high-medium light in moist-dry
garden soil or potting mix. The plants are easy to grow.
When subjected to temperatures of −7°C (19°F) for seven
days, the fronds spot-burned or died, but the plants re-
sumed growth afterward. Unconfirmed reports indicate
that a skin rash may develop in response to contact with
this fern.
The wild and cultivated plants of Rumohra adiantiformis
are highly variable in the size and cutting of the fronds. The
young indusium arches over the sorus, forming an upside-
down cup, and is black when mature. The species is native
to the Southern Hemisphere.
‘Davis’. Pinnules slightly contracted.
c
Sadleria
Although Sadleria contains species with an arborescent
growth form, it is related to Blechnum instead of such tra- Figure 13.108.2. Rumohra adiantiformis: a. frond, bar =
ditional tree-fern genera as Cibotium, Cyathea, or Dickso- 5 cm (2 in.); b. sori and indusia, bar = 1 cm (0.4 in.);
nia. Sadleria differs from these genera by having relatively c. pinnae from different mature plants, bar = 5 cm
little-divided blades and linear sori. Some Sadleria species (2 in.).
are nonarborescent, although these are not cultivated.
Commercial nurseries used to import living trunks of Sad-
leria from the Hawaiian Islands; later, plants were com- are netted along the midvein but free beyond. The sori are
mercially grown from spores. Sadleria is grown outdoors as linear and continuous along both sides of the segment mid-
a tub or bedding plant in southern California, but the ribs. The indusium is shaped like the sorus and opens to-
plants need protection from the occasional freezing tem- ward the midrib.
peratures. They may also be used as houseplants. Young The genus contains six species, all endemic to the
fronds are often red. Since the genus is sensitive to root dis- Hawaiian Islands. It is named for Joseph Sadler (1791–
turbance, extra care must be taken when transplanting. 1849), a botany professor and student of the flora of Hun-
Sadleria consists of tree ferns and smaller plants with gary. See the section on “Tree Ferns” in Chapter 10 for
creeping rhizomes. The trunk or rhizome is scaly and bears more information on Sadleria; for a technical treatment of
one-pinnate-pinnatifid or two-pinnate fronds. The veins the species, see Palmer (1997).
 492 C 13

Sadleria cyatheoides Kaulfuss

Amaumau  13.109.1–3
Semi-tender to tender
A large tree fern, often with reddish new foliage. Grows
well under high light in moist garden soil or potting mix
with good drainage. This species is sensitive to root dis-
turbance; transplant with care. It will die at temperatures of
about −2°C (30°F).
The stipe of Sadleria cyatheoides is scaly mostly at the
base, not distally. The blades are one-pinnate-pinnatisect,
with the veins stopping short of the margins and immersed
(not visible) on the lower surface. The sori occupy more
than half the length of the segment. This species is native to
the Hawaiian Islands.
Sometimes small trunks are individually packaged and Figure 13.109.2. Sadleria cyatheoides: sori and indusia.
sold as novelties. Shipments of Sadleria cyatheoides trunks
occasionally contain a few trunks of S. pallida. This latter
species can be distinguished by scaly stipes throughout and
by obvious veins on the lower surfaces of the blades running
to the margin. Its culture is the same as that of S. cyatheoides.
a

Figure 13.109.3. Sadleria cyatheoides: a. frond, bar =

5 cm (2 in.); b. fertile segments, bar = 1 cm (0.4 in.).
Figure 13.109.1. Sadleria cyatheoides: habit.
 F  F A  C 493

Salvinia  13.110.1 Salvinia auriculata Aublet  13.110.2

Water spangle syn. Salvinia rotundifolia Willdenow
The water spangles are small floating ferns, although they Semi-tender to tender
also grow in mud around the shores of ponds, lakes, or A small aquatic fern with short-creeping rhizomes. Grows
swamps. Their small, rounded leaves add interest to ponds, well under direct sunlight. Possession of the plants is pro-
dish gardens, and aquariums. Salvinia does not tolerate hibited by governmental agencies; see S. molesta for details.
much cool weather and dies during the winter in most parts The roundish fronds of Salvinia auriculata bear hairs on
of the United States. Culture is the same as for another the upper surfaces that have four branches uniting at their
aquatic fern, Azolla. tips to form an egg-beater- or cage-like structure. The sori
Salvinia minima is the only indigenous species in the (sporocarps) are long-stalked, whitish, and spherical. Up
United States. A small clump of plants can grow rapidly to 30 are borne in a branched cluster. They all hang at
and carpet the water’s surface in short time. Salvinia molesta about the same level. The spores are normal, not aborted.
also grows rapidly and is a terrible weed in waterways in This species is native to tropical America.
the Old World tropics, and now in the United States,
where it hampers navigation and fishing. For this reason,
any member of the S. auriculata complex (species with egg-
beater-like hairs) may not be imported into the United c
States; see S. molesta for more details.
The fronds of this genus are 1–2 cm (0.5–0.75 in.) long b
and vary from round to oblong. Three fronds occur at each
node on the stem. Two of the fronds are green, floating,
and leaf-like, and the third is whitish, submerged, finely di- a
vided, and root-like (true roots are lacking). The upper sur-
faces of the green fronds bear water-repellent hairs. The Figure 13.110.2. Salvinia auriculata: a. habit, bar = 1 cm
submerged, root-like fronds rarely produce sori, but when (0.4 in.); b. hair from upper surface, bar = 1 mm;
they do, the sori consist of sporangia enclosed by a globose c. sporocarp, bar = 1 cm (0.4 in.), after Sota (1963).
indusium (the entire unit is called a sporocarp).
Salvinia is native to tropical and subtropical regions of
the world. It consists of 10 species. The genus name honors
Antonio Maria Salvini (1633–1729), an Italian professor of
Salvinia minima Baker  13.110.3
Greek.
Semi-tender, Zone (8)9
A small aquatic fern with short-creeping rhizomes. Grows
well under direct sunlight.
Salvinia minima is a generally smaller plant than S. au-
riculata and S. molesta. It can be distinguished from these
two species by hairs that are free at the tip, not united into
an egg-beater-like structure. The hairs are often sessile, not
borne on a cone-like base as in the other two species, and
they vary from one to four per group. The sori (sporocarps)
are sessile and four to eight in a chain-like cluster. This spe-

Figure 13.110.3. Salvinia minima: a. habit, bar = 1 cm

(0.4 in.); b. hairs from upper surface, typical (left) and
atypical (two on right), bar = 1 mm; c. sporocarps, bar =
Figure 13.110.1. Salvinia molesta: habit. 1 cm (0.4 in.), after Sota (1963).
 494 C 13

cies is native to the southeastern United States, West In- Web site at http://nas.er.usgs.gov/ferns for more informa-
dies, and Central and South America. tion. Moran (1992c) discusses the plant’s weediness and
At different times of the year, cultivated material may the biological control methods that have been used to com-
have many atypical hairs. These hairs might be lacking, rep- bat it.
resented by a bump, or stoutly cone-shaped. The free
branch-hairs borne at the tip of the cone-like hair may be Salvinia natans (Linnaeus) Allioni  13.110.5
poorly developed and number less than four. When sporo- Semi-tender to hardier
carps are not present, Salvinia minima might be confused A small aquatic fern with short-creeping rhizomes. Grows
with S. natans except for the roundish leaves of the former. well under direct sunlight.
Salvinia natans has oblong-truncate to ovate fronds. The
Salvinia molesta D. S. Mitchell  13.110.1, 4 hairs on the upper surfaces of the blades have four free
Semi-tender to tender branches, not united. The sori (sporocarps) are spherical,
A small aquatic fern with short-creeping rhizomes. Grows about six in a tight cluster. The species is native to Europe
well under direct sunlight. and Asia.
Salvinia molesta has rounded fronds, and the mature
ones are often folded lengthwise along the midrib, espe-
cially in crowded conditions. As in S. auriculata, stiff hairs a
b
are present on the upper surfaces of the blades. These hairs
consist of a stalk tipped by four hairs that unite at their c
apices to form an egg-beater- or cage-like structure. The
fertile part of the frond is chain-like (unbranched) and to-
ward the base has short-stalked sori (sporocarps) that are
ovate with a slight beak at the tip. The sori gradually be-
come sessile toward the apex of the fertile part. The spores
are mostly aborted. Salvinia auriculata can be distinguished
from this species by its branched, corymb-like fertile part. Figure 13.110.5. Salvinia natans: a. habit, bar = 1 cm
Salvinia molesta is native to southern South America (0.4 in.); b. hair from upper surface, bar = 1 mm;
(southern Brazil, Uruguay, and Paraguay) but has become c. sporocarps, bar = 1 cm (0.4 in.).
naturalized in Africa, India, Sri Lanka, Indonesia, Australia,
and New Zealand.
This species is a terrible weed in the Old World tropics
and is becoming a serious pest in the southern United Schaffneria
States from Texas to Florida. Although killed by hard frosts
This small, charming fern bears clusters of rounded, dia-
and prolonged subfreezing temperatures, it should not be
mond-shaped fronds. It grows on mossy rocks and ledges
cultivated anywhere in the United States because of its po-
(usually limestone) in tropical areas and is well suited for
tential to escape. In the United States it has been declared
terrarium culture.
a Federal Noxious Weed, which makes it illegal to import,
The rhizomes are erect and bear small, clathrate scales.
own, cultivate, transport, or sell the species. In some states,
The clustered fronds are 2.5–15 cm (1–6 in.) tall, and the
laws also prohibit its possession. Check the World Wide
blades are entire, rhombic-obovate to fan-shaped, glabrous,
and lacking a midrib. The veins spread and divide in a fan-
shaped pattern, forming areoles without included veins.
b c The sori are short to linear, with indusia shaped like the
sori and attached on one side to a vein. Some pteridologists
a classify this genus in Asplenium.
Schaffneria consists of only one species native to Central
America and Cuba. The genus is named for Wilhelm H. J.
Schaffner (1830–1882), a German pharmacist who col-
lected plants in Mexico.

Schaffneria nigripes Fée  13.111.1

Figure 13.110.4. Salvinia molesta: a. habit, bar = 1 cm syn. Asplenium nigripes (Fée) Hooker
(0.4 in.), after Moran (1992c); b. hair from upper Tender
surface, bar = 1 mm, after Moran (1992c); c. sporocarps, A small fern with erect rhizomes. Grows well under low-
bar = 1 cm (0.4 in.). medium to medium light in potting mix with good drain-
 F  F A  C 495

Figure 13.111.1. Schaffneria nigripes: a. habit, bar =

5 cm (2 in.); b. frond, bar = 1 cm (0.4 in.).
Figure 13.112.1. Selaginella kraussiana: mat-like growth
habit.

age but kept constantly moist. The plants are easy to grow
from spores.
See the characteristics of the genus for a description.

Selaginella  13.112.1–5

Spike moss
Selaginellas are mostly small- to medium-sized terrestrial
plants. They are moss-like or fern-like with many branches
bearing small, scaly leaves. They are used in rock gardens, as
ground covers, or as accent plants. The tropical species are
particularly eye-catching for their unusually colored leaves,
such as red or bronze shades or iridescent blue-greens. A
few tropical species are epiphytic, but they are rare and not
cultivated.
Different regions of the United States are associated
with certain species of native and cultivated Selaginella. In
the northeastern United States the mat-forming meadow
spike moss (Selaginella apoda) is often found in shady
lawns, especially in moist depressions. In the southern
United States S. kraussiana, a warm-temperate and sub- Figure 13.112.2. Selaginella kraussiana: branches with
tropical species with mossy, mat-like growth, is widely “cones.”
available in the trade and grows easily outdoors. It can in-
vade gardens and has become weedy in some areas. Selagi-
nella plana, a taller plant with frond-like branches, is easily
grown in more tropical areas and has also become widely
496 C 13

Figure 13.112.3. Selaginella umbrosa: frond-like growth

habit. Photo courtesy of B. Hoshizaki.

Figure 13.112.4. An example of a Selaginella with rosette

growth habit. Shown are several plants of S. tamariscina,
naturalized. The main requirements for promoting luxuri- which are frequently grown in Japan, but are very rarely
ant growth of the ornamental tropical and subtropical spe- cultivated in the United States and not treated in this
cies are ample humidity, soil moisture, and warm temper- book. They are native to Asia, where they usually grow
atures. In arid climates selaginellas are used mainly in rock on exposed rocks or hillsides.
gardens. The species adapted to such climates are generally
more difficult to grow and do not have wide ornamental
appeal. The resurrection plant (S. lepidophylla) is an excep-
tion: it uncurls from a ball of foliage to form an attractive most important distinction is the two kinds of sporangia
spreading rosette. For more cultural information, see “Fern and spores (male and female) in Selaginella versus only one
Allies” in Chapter 10. kind in Lycopodium. This difference, however, is not easily
In general, the plants of Selaginella are moss-like and observed. In aspect the cultivated species of Selaginella are
produce small leaves less than 5 mm (0.2 in.) long. The more herbaceous and delicate than Lycopodium and have
leaves contain only a single vein or midrib. The main stems inconspicuous “cones.”
may be erect, prostrate, creeping, trailing, or scrambling, Selaginella is a large genus, with about 750 species, and
but they sooner or later fork into successively smaller is extremely diverse in external form. It is cosmopolitan but
branches. In some species, such as Selaginella oregana, the primarily tropical. The genus name is derived from adding
leaves are spirally arranged around the stem, but in most the Latin suffix -ella, indicating a diminutive, to Selago, the
species they are arranged in four rows, with two lateral rows former name of a club moss in the genus Huperzia; thus,
pointing outward (the lateral leaves) and two dorsal rows Selaginella means “like a small Selago.”
pointing more or less forward (the median leaves). Also, a Identifying Selaginella species can be difficult due to
single leaf, known as the axillary leaf, occurs on the lower their large number, the small size of the distinguishing char-
surface of the stem at each fork (Figure 13.112.5). The acters, and (with cultivated plants) the lack of data about
leaves on the main part of the stem (the main-stem leaves) country of origin. For these reasons, selaginellas in horti-
may differ from those on the branchlets. The leaves vary in culture are often misidentified. There are many more spe-
such details as shape and type of margin or attachment. The cies in cultivation than those listed here, but they are un-
fertile leaves usually form a spike or cone-like cluster at the common and hard to obtain commercially. Native plants
tip of a branch, often appearing slightly or strongly angular. —such as Selaginella densa, native to midwestern states, S.
The lower leaves of the cone usually bear large female spo- rupestris, native to central and eastern states, and S. walla-
rangia, and upper leaves bear smaller male sporangia, but cei, native to northwestern states—are sometimes used in
this characteristic varies greatly among the species. rock gardens and are well described in United States floras.
Selaginella might be confused with Lycopodium, though To facilitate identification, we have divided the culti-
several technical differences separate the two genera. The vated species into three groups based on growth form. If a
 F  F A  C 497

lateral leaf
median
leaf

axillary stem
leaf
upper surface lower surface

Figure 13.112.5. Selaginella: leaf arrangement a

(Selaginella articulata), bar = 5 mm (0.2 in.).
Figure 13.112.6. Selaginella apoda: a. habit, bar = 5 cm
(2 in.); b. leaves, lateral (below) and median (above), bar
species has two growth forms (for example, Selaginella het- = 1 mm.
erodonta), it is listed under both groups.
Mat-like growth. These species resemble mosses. Their
main stems and branches are leafy and creeping, with a from gutters and in shallow depressions. It makes a good
mat-like or low-trailing habit. The growth is indefinite terrarium plant.
(that is, of unlimited growth), not determinate as in many The lateral leaves of Selaginella apoda are distant, ovate
upright species, and the roots are produced more or less all to ovate-lanceolate, slightly cordate at the base, green
along the stem at the branch forkings. Selaginella apoda, S. throughout but with one or two (rarely three) rows of trans-
diffusa, S. douglassii, S. heterodonta, S. kraussiana, S. oreg- parent cells along the margins, and minutely serrate with
ona, S. serpens, S. uncinata. acute apices. The medial leaves are ovate to ovate-lanceo-
late, either with an acute apex or attenuate without a mid-
Frond-like growth. These species have foliage that resem- rib extension; the base is slightly oblique on the inner side
bles fern fronds. The main stems are erect, resembling a but rounded and prominent on the outer side. This spe-
stipe, and produce roots at the base. They branch repeatedly cies is native to the eastern United States and Mexico.
above to form a blade-like structure resembling a fern frond
with numerous tiny, scale-like leaflets. This frond-like fo- Selaginella braunii Baker  13.112.7
liage arises from a long-creeping stem, or rarely scrambling Chinese lace fern
as in Selaginella willdenovii. The species in this group are S. Hardy, Zone 7
braunii, S. erythropus, S. heterodonta, S. involvens, S. marten-
A small-medium plant with erect, open, frond-like growth
sii, S. moellendorffii, S. oaxacana, S. plana, S. pulcherrima, S.
arising from medium-creeping horizontal stems. Grows
umbrosa, S. viticulosa, S. vogelii, and S. willdenovii.
well under medium light in moist garden soil or potting
Rosette growth. These species have frond-like branch sys- mix.
tems (as in the preceding group), but the fronds are ar- The lower, stalk-like part of the stem is somewhat stiff
ranged in a rosette. They curl into a ball when dry. The and firm with few leaves. The blade-like part of Selaginella
species in this group are Selaginella lepidophylla, S. harrisii, braunii is triangular to ovate and horizontal with short,
and S. pallescens (in dense shade the latter two may be more hairy branches. The lateral leaves of the ultimate branches
irregularly clustered than rosette-like). are subdistant, spreading, and decurrent on the basiscopic
side. This species is native to China but has become natu-
Selaginella apoda (Linnaeus) Spring  13.112.6 ralized in many parts of the world.
Meadow spike moss
Hardy, Zone 5 Selaginella diffusa (C. Presl) Spring  13.112.8
A small plant with mat-like growth from delicate, low, Tender
creeping, branched stems. Grows well under low to me- A small plant with delicate, mat-like growth and long-
dium light in moist to moist-wet garden soil. This species is creeping, branched stems. Grows well under low to me-
easy to cultivate. In the northeastern United States it can dium light in moist-wet to moist garden soil or potting mix.
become weedy in lawns, especially around downspouts The mat-like growth may also be trailing.
498 C 13

Selaginella diffusa varies tremendously in shape, size, and

amount of cilia on the leaves. The stems have a dark band
or swelling below each fork. The lateral leaves are distant to
close (but not overlapping), elliptical-lanceolate, acute,
sometimes white-bordered, and sparsely ciliate to entire.
The leaf has a prolonged ear with small cilia at the base on
the acroscopic side. The median leaves are elongate-ovate
b and broadest near the middle, with long-acuminate apices;
the base on the basiscopic side has a prolonged, tongue-
shaped ear with or without a whitish border. The margins
are finely denticulate and usually have multicellular cilia.
This species is native to the American tropics.

Selaginella douglassii (Hooker & Greville) Spring

 13.112.9
Douglas spike moss
Hardy, Zone 5
A small plant with delicate, mat-like growth and long-
creeping, branched stems. Grows well under medium light
in moist garden soil.
Selaginella douglassii has delicate leaves that resemble
those of S. apoda in size but are firmer in texture. The lat-
eral leaves are more or less spreading and distant. They are
Figure 13.112.7. Selaginella shiny green or turn brown with an orange to red spot at the
a braunii: a. branches, bar = base or are completely reddish. The apices are rounded, and
5 cm (2 in.); b. leaves, lateral the bases are eared and ciliate (elsewhere the leaf is entire).
(below) and median (above), The median leaves are entire, pointed, and unevenly eared
bar = 1 mm.

a a b

Figure 13.112.9. Selaginella douglassii: a. habit, bar =

5 cm (2 in.); b. leaf arrangement, bar = 2 mm.

b at the base, the ears ciliate. This species is native to the

northwestern United States.

Selaginella erythropus (C. Martius) Spring

 13.112.10
Tender
c A small plant with erect, frond-like growth arising from
short-creeping, horizontal stems. Grows well under me-
dium light in moist potting mix. This species is often used
in terrariums and bottle gardens because it likes humidity.
The lower surfaces of the leaves are burgundy and
Figure 13.112.8. Selaginella diffusa: a. habit, bar = 5 cm remain so with age, whereas the upper surfaces are dark
(2 in.); b. leaf arrangement, bar = 2 mm; c. leaves, lateral green. The margins have numerous minute teeth. Selagi-
(below) and median (above), bar = 1 mm. nella erythropus is native to northern South America.
 F  F A  C 499

b b

c
c
a

Figure 13.112.10. Selaginella erythropus: a. branches, bar Figure 13.112.11. Selaginella harrisii: a. branches, bar =
= 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
leaves, lateral (below) and median (above), bar = 1 mm. lateral (below) and median (above), bar = 1 mm.

Selaginella harrisii Underwood & Hieronymus Selaginella heterodonta (Desvaux ex Poiret)

 13.112.11 Hieronymus  13.112.12
Tender Tender
A small plant with erect, frond-like growth borne in a A small-medium plant with erect, frond-like or mat-form-
rosette or cluster; if the plants have clustered growth, they ing growth from long-creeping horizontal stems. Grows
arise from a short-creeping horizontal stem. Grows well well under medium light in moist garden soil or potting
under medium light in moist garden soil or potting mix. mix. This species is suitable as a ground cover in terrariums.
In Selaginella harrisii, the lower surfaces of the leaves are If mat-like in growth, Selaginella heterodonta has main
lighter colored than those of most selaginellas, often a sil- stems that are usually long and whip-like at the tips; if
very green. The lateral leaves are ovate and tightly imbricate frond-like in growth, the branches are erect-ascending or
near the stem tips, with the apex sharply acute and the base arched. The narrow branchlets are 3–4 mm (0.1–0.2 in.)
subauriculate with long cilia; the acroscopic margin is nar- wide. The lateral leaves vary from oblong-ovate to more el-
row, white, and short-ciliate to denticulate at the apex, and liptic and are acutish with subcordate bases; margins are
the basiscopic margin entire to denticulate. The older lat- strongly ciliate especially on the acroscopic side. The me-
eral leaves are green. The median leaves are ovate and aris- dian leaves are ovate-falcate, aristate or not, usually eared on
tate, with the base bearing a single short ear; the margin is the basiscopic side, and have long-ciliate margins. This spe-
short-ciliate and has a narrow white border. This species is cies is native to the Greater Antilles.
native to Mexico, Belize, Honduras, and Jamaica.
Selaginella harrisii was formerly included with S. palles- Selaginella involvens (Swartz) Spring
cens, which differs by the following characteristics: leaves  13.112.13
on the unbranched part of the main stem are long-ciliate, syn. Selaginella caulescens (Wallich ex Hooker &
not short-ciliate; the base of these leaves have long, almost Greville) Spring
overlapping ears, rather than a rounded base; the lateral Tree spike moss
leaves are short-ciliate, not long-ciliate; the basiscopic mar- Semi-tender to hardier
gin has a white border that is narrow to lacking, rather than A medium-sized plant with erect, frond-like growth aris-
a wider, white border; and the older lateral leaves are tan, ing from medium-creeping horizontal stems. Grows well
not green. under medium light in moist garden soil or potting mix. It
likes humidity.
500 C 13

The frond-like part of Selaginella involvens is triangular

with overlapping leaves. The leaves on the main stem be-
tween the middle and lower lateral branches have very un-
b evenly lobed bases and are extended on the basiscopic side
and often lobed again, with erose-ciliate margins. The lat-
eral leaves are ovate-triangular and ciliate-denticulate (less
so toward the apex), and the midrib and two longitudinal
grooves parallel to the midrib are indistinct. The median
leaves are ovate and have a more or less keeled midrib and
attenuate apices with two longitudinal grooves. This species
is native to Japan, Korea, China, and Vietnam.

Selaginella kraussiana (Kunze) A. Braun

 13.112.1, 2, 14
Spreading selaginella
Hardy, Zone (6)7
A small plant with mat-like growth and long-creeping (ex-
cept for ‘Brownii’), irregularly branching stems. Grows well
a under medium to low light in moist garden soil. The plants
are used for ground cover but may become weedy. It is the
most common species in the United States trade.
Figure 13.112.12. Selaginella heterodonta: a. habit, bar = The lateral leaves of Selaginella kraussiana are ovate,
5 cm (2 in.); b. leaves, lateral (below) and median finely dentate-serrate, and acute. The median leaves are
(above), bar = 1 mm. ovate and acuminate, with bases slightly cordate on the ba-
siscopic side. This species is native to Africa but has become
widely naturalized in the tropics and subtropics, including
parts of the southeastern United States.
Selaginella kraussiana is often misidentified in the trade
b as S. denticulata (Linnaeus) Spring, a species of central Eu-
rope and the Near East not cultivated in the United States.

b
c

Figure 13.112.13. Selaginella

a
involvens: a. branches, bar =
5 cm (2 in.); b. leaf
arrangement, bar = 2 mm; a
c. leaves, lateral (below) and
median (above), bar = 1 mm.

Figure 13.112.14. Selaginella kraussiana: a. habit, bar = 5

cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
lateral (below) and median (above), bar = 1 mm.
 F  F A  C 501

‘Aurea’. Plants yellow green or chartreuse; slightly less

robust growth form.
‘Brownii’. Pin-cushion moss. Growth very compact, a
cushion-like.
b
‘Gold-tips’ (‘Gold’). Branchlets lighter green at the tip.

Selaginella lepidophylla (Hooker & Greville) Spring

 13.112.15
Resurrection plant
Semi-hardy, Zone 8
A small plant with a rosette growth form. Grows well under
high light in basic, moist to moist-dry garden soil with sand c
and good drainage. The plants are popular because of their
ability to curl into a ball when dry and uncurl into a flat
rosette when watered. This uncurling is a mechanical ac-
tion brought about by the absorption of water. It does not
indicate that the plant is alive. If not too desiccated, the
plants can be re-established by placing in a well-drained
mix in a moderately humid environment.
All the leaves on Selaginella lepidophylla are imbricate,
broadly ovate, and pubescent, with transparent margins.
The lateral leaves are yellowish to reddish on the lower sur-
face and green on the upper surface, cordate and ciliate at
the base. The median leaves are cordate to truncate at the
base and ciliate toward base to dentate-ciliate toward the
apex. This species is native to the southwestern United
States (New Mexico and Texas) and Mexico.

Selaginella martensii Spring  13.112.16 Figure 13.112.16. Selaginella martensii: a. branches, bar
Semi-tender to hardier = 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c.
leaves, lateral (below) and median (above), bar = 1 mm.
A medium-sized plant with erect to suberect, frond-like
growth arising from short-creeping horizontal stems.
Grows well under medium light in moist to moist-wet gar-
den soil or potting mix. This species likes humidity. eral leaves are oblong, to 5 mm (0.2 in.) long. The apex is
Selaginella martensii has leaves that are dark green, rounded, the base unequal, acroscopic side rounded, basis-
smooth, and shiny above but lighter green below. The lat- copic side with a small ear, margins ciliate to denticulate at
the base to more entire toward the apex. The median leaves
are ovate, keeled, and usually long-aristate, the aristae to
b about half the blade length; the base has two uneven ears,
a and the leaf margins are ciliate, becoming more denticu-
late toward the apex, often with a white border. This species
is native to Central America.
‘Albolineata’. Snowy club moss. Leaves variegated with
scattered creamy-white to white streaks.

Selaginella moellendorffii Hieronymus

 13.112.17
Semi-tender
A medium-sized plant with erect to suberect, frond-like
growth arising from creeping horizontal stems. Grows well
under medium light in moist garden soil. It likes humidity.
The blade-like part of Selaginella moellendorffii is trian-
Figure 13.112.15. Selaginella lepidophylla: a. habit, bar = gular to ovate. The leaves on the main stem between main
5 cm (2 in.); b. leaf arrangement, bar = 2 mm. branches have larger leaves that are triangular, acuminate,
 502 C 13

c
a

a c

Figure 13.112.17. Selaginella

moellendorffii: a. habit, bar = 5 cm
(2 in.); b. leaf arrangement, bar = Figure 13.112.18. Selaginella oaxacana: a. habit, bar = 5
2 mm; c. leaves, lateral (below) and cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
median (above), bar = 1 mm. lateral (below) and median (above), bar = 1 mm.

dial leaves have fine streaks (idioblasts) on the upper sur-

faces, and their margins often curl upon drying. The lat-
and outward-pointing. The lateral leaves are ovate-falcate, eral leaves are obliquely oblong-triangular, obtuse at the
often with a faint false vein on either side of the midrib; the apex, truncate to rounded at the base, and serrulate. The
apex is acute, the margins minutely ciliate. The median medial leaves are smaller, ovate, acuminate, and also serru-
leaves are appressed, ovate, unequal sided, short-toothed late. This species is native to Central America.
apically, and ciliate. This species is native to China, Tai- Selaginella oaxacana is sometimes misidentified in the
wan, the Philippines, and Vietnam. trade as S. pulcherrima.
Selaginella moellendorffii reproduces vegetatively by
gemmae produced at the branch tips. The gemmae resem- Selaginella oregana D. C. Eaton  13.112.19
ble cones and tend to be slightly angular and redder than Oregon spike moss
the rest of the branchlet. They fall off easily, and as a result Semi-hardy, Zone 8
this species can become weedy in greenhouses. A small-medium-sized plant with hanging or trailing stems.
Grows well under medium light. The plants are cultivated
Selaginella oaxacana Spring  13.112.18 as hanging epiphytes, either without soil or over moist soil
Oaxaca spike moss and rocks in rock gardens. In the wild this species forms
Tender mats of hanging, stringy growth on trees, rarely on soil.
A medium-sized plant with erect, frond-like growth aris- Selaginella oregana is distinctive by its spirally arranged
ing from creeping horizontal stems. Grows well under me- leaves, all of one type, that are narrowly triangular to linear-
dium light in moist garden soil or potting mix in humid lanceolate. The leaves are not four-ranked, and therefore
environments. the branchlets do not appear flattened. This species is native
The frond-like part of Selaginella oaxacana is flabellate, to western North America.
and the leaves on the main stem below the first branch are
pressed to the stems and broadly ovate. The lateral and me-
 F  F A  C 503

b
b

c
a

c
a

Figure 13.112.20. Selaginella pallescens: a. habit, bar = 5

cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
lateral (below) and median (above), bar = 1 mm.

Figure 13.112.19. Selaginella

Selaginella plana (Desvaux ex Poiret) Hieronymus
oregana: a. branches, bar =
 13.112.21
5 cm (2 in.); b. spiral leaf
Tender to semi-tender
arrangement, bar = 2 mm;
c. similar lateral and median A small to medium plant with a frond-like growth that is
leaves, bar = 1 mm. often sprawling, branched, and rooting on the underside.
Grows well under medium light in moist garden soil. Ma-
ture plants may appear untidy because of their scrambling
habit. They sometimes become invasive and weedy. The
species is common in southern Florida fern gardens.
Selaginella pallescens (C. Presl) Spring The erect stem of Selaginella plana is either greenish,
 13.112.20 straw-colored, or (in Florida specimens) reddish brown. All
syn. Selaginella emmeliana Van Geert the leaves are entire, not ciliate. The lateral ones have an
Sweat plant acute to rounded apex, and the base is unequal; the base of
Semi-hardy, Zone 8 the acroscopic side has a narrow, often twisted ear, and the
A small plant with a rosette growth form, or if only loosely basiscopic side is truncate or rounded. The median leaves
clustered, then arising from a short, horizontal stem. Grows are narrowly ovate or oblong, eared or not. The axillary leaf
well under medium light in moist garden soil or potting is strongly eared on both sides, and the ears are often
mix. The plants form flat rosettes when young but tend to twisted. This species is native to southeastern Asia and In-
become loosely clustered with more erect branches with age. donesia and has become widely naturalized elsewhere in
Most plants of Selaginella pallescens can be distinguished the tropics.
by their rosette growth form and (mostly) white-bordered Two unidentified species resembling Selaginella plana
leaves. The lateral leaves of the smaller branches closely are also in cultivation.
overlap and are ovate-acuminate, mostly subfalcate at the
apex and short-ciliate. Their basiscopic margins may lack Selaginella pulcherrima Liebmann ex E. Fournier
the white narrow border. The median leaves are ovate and  13.112.22
serrate, with a small tooth at the apex. Older leaves of both Tender
forms are tan. This species is native to Mexico, Central A small-medium plant with frond-like growth arising from
America, Colombia, and Venezuela. long-creeping horizontal stems. Grows well under medium
Selaginella pallescens is often confused with S. harrisii. light in moist to moist-wet garden soil or potting soil. The
Another, unidentified species circulating in cultivation is plants like humidity.
also related to S. pallescens. It has sparsely ciliate leaves, and All leaves of Selaginella pulcherrima are dark green
the median leaves are ovate and long-attenuate. above, lighter below, and more or less smooth. The oldest
 504 C 13

b b
c

Figure 13.112.22. Selaginella pulcherrima: a. branches,

bar = 5 cm (2 in.); b. leaf arrangement, bar = 2 mm;
c. leaves, lateral (below) and median (above), bar =
Figure 13.112.21. Selaginella plana: a. habit, bar = 5 cm
1 mm.
(2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
median leaf (above), upper surface of lateral leaf (center),
under surface of lateral leaf (below), bar = 1 mm.

a
ones turn buff. The lateral leaves are ovate, acute, and cili-
ate to denticulate. The medial leaves are ovate, acuminate at
the apex, ciliate to denticulate, with or without an indis-
tinct white border. The axillary leaves lack ears. This species
is native to Mexico.
Selaginella oaxacana is sometimes misidentified in the
trade as S. pulcherrima.

Selaginella serpens (Desvaux ex Poiret) Spring c

 13.112.23
Tender
A small plant with low, mat-like growth from branched,
creeping stems. Grows well under low-medium light in b
moist garden soil or potting mix. The foliage undergoes a
diurnal color change from bright green in the morning to
pale silvery in the late afternoon. This change is caused by
a contraction of the contents of the epidermal cells, allow-
ing a layer of air to accumulate between the cuticle and the Figure 13.112.23. Selaginella serpens: a. habit, bar = 5 cm
outer cell wall. This layer diffracts light, making the leaf (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves, lateral
appear pale silvery. (below) and median (above), bar = 1 mm.
 F  F A  C 505

The lateral leaves of Selaginella serpens are ovate, obtuse, Selaginella uncinata (Desvaux ex Poiret) Spring
cordate at the base, and ciliate (less so toward the apex).  13.112.25;  49
The median leaves are ovate-acuminate, ciliate, and some- Blue spike moss, peacock plant, rainbow moss
times larger or broadly eared on the outer basiscopic side. Hardy, Zone 6
This species is native to the West Indies. A small to medium plant with mat-like growth from long-
creeping or trailing, branched stems. Grows well under me-
Selaginella umbrosa Lemaire ex Hieronymus dium light in moist garden soil or potting mix. The plants
 13.112.3, 24 reportedly overwinter well in coastal New York (Zone 6).
Red spike moss This species is attractive for its blue-green, iridescent fo-
Tender liage. This color is best developed when the plants are
A medium-sized plant with frond-like growth arising from grown in low light.
long-creeping, branched horizontal stems. Grows well Selaginella uncinata has ovate or ovate-oblong lateral
under medium light in moist-wet garden soil or potting leaves with transparent and entire margins. Some are
mix. It likes humidity. The species is attractive because of weakly stalked, and the leaf base is truncate to weakly cor-
its red stems. date, with the acroscopic side often overlapping the stem.
The main, erect stems of Selaginella umbrosa are red, The median leaves are overlapping, ovate-lanceolate, the
and the blade-like part is irregularly and densely flabellate, margins transparent and entire, the apex abruptly attenu-
flat, and bright green. The lateral leaves of the main ate, and the base often oblique and weakly cordate. This
branches are ovate-lanceolate, unequally sided, and strongly species is native to southern China and has become natu-
ciliate at the base. The median leaves are ovate-acuminate ralized in the southeastern United States.
and slightly keeled. This species is native to the American
tropics and has become naturalized in many parts of the Selaginella viticulosa Klotzsch  13.112.26
tropics throughout the world. Tender
A small-medium plant with frond-like growth arising from
creeping horizontal stems. Grows well under medium light
a in moist potting soil and in humidity. This species has
softer and laxer foliage than other selaginellas.

Figure 13.112.24. Selaginella umbrosa: a. habit, bar = 5

cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
lateral (below) and median (above), bar = 1 mm. Figure 13.112.25. Selaginella uncinata: a. habit, bar = 5
cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
lateral (below) and median (above), bar = 1 mm.
 506 C 13

c
b b

Figure 13.112.27. Selaginella vogelii: a. branches, bar =

Figure 13.112.26. Selaginella viticulosa: a. habit, bar = 5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves,
5 cm (2 in.); b. leaf arrangement, bar = 2 mm; c. leaves, lateral broadly attached to stem (below) and median
lateral (below) and median (above), bar = 1 mm. (above), bar = 1 mm.

The lateral leaves on the distal half of the main stem are Selaginella willdenovii (Desvaux ex Poiret) Baker
oblong, two times longer than wide, spreading (not imbri-  13.112.28
cate), and faintly streaked. The acroscopic margin is ciliate Peacock fern, vine spike moss
toward the base to denticulate toward the apex, and the Tender, Zone 9
basiscopic base is truncate. Selaginella viticulosa is native to A medium to large plant with sprawling to erect, frond-like
Costa Rica, Panama, Colombia, and Venezuela. growth. Grows well under medium-high light in moist gar-
den soil. Give the plants a lot of space to grow. The scram-
Selaginella vogelii Spring  13.112.27 bling stems may reach 6 m (20 ft.) long and typically bear
Tender small leaves with a strikingly beautiful sky-blue iridescence
A small plant with frond-like growth arising from short- on their upper surface. The iridescence, however, develops
creeping horizontal stems. Grows well under medium light only on plants grown in shade; those plants grown in full or
in moist garden soil or potting mix. This species likes partial sun are not iridescent. Iridescence in pteridophytes
humidity. tends to occur in species that grow on deep-shaded forest
The main, erect stems of Selaginella vogelii are about as floors, which suggests that this trait is an adaptation to low
long as the blade-like part. The latter is broadly triangular light. See Moran (1995a) for a popular account of irides-
and bears branches with many short, stiff hairs on the lower cence in ferns.
surface. The lateral leaves are narrowly triangular, con- All the leaves are entire on Selaginella willdenovii. The
stricted near the base then decurrent, and entire (not cili- lateral leaves are whitish eared on the acroscopic side, and
ate). The median leaves are long-acuminate, unequal at the the axillary ones have two elongate-rounded ears. This spe-
base, and entire. This species is native to tropical Africa. cies is native to tropical Asia and Indonesia and has occa-
sionally become naturalized elsewhere in the tropics.
 F  F A  C 507

name, which has priority). Species formerly placed in

Crypsinus are rarely grown or offered in the trade and there-
a fore are not treated here. Although the plants are attractive,
resembling small polypodiums, reports differ on their ease
b of culture.
Selliguea contains about 50 species or more and is native
to Africa, India, Malaysia to Australia, New Guinea, the
Pacific Islands, the Philippines, and Japan. The genus is
named for M. Selligue, a French optician and instrument
maker.

Selliguea feei Bory  13.113.1

Semi-tender
A small fern with medium- to long-creeping rhizomes.
Grows well under high-medium light in moist to moist-
dry potting mix or garden soil. The plants are easy to culti-
vate in pots or in the ground, but the fronds tend to be-
come brittle with age.
c Selliguea feei is characterized by broadly ovate rhizome
scales, broadly lanceolate sterile fronds, often narrower and
taller fertile fronds, and linear sori oblique to the costa. The
fronds have a slightly brittle, leathery texture, and the lateral
veins are conspicuous. The species is native to Java.

Figure 13.112.28. Selaginella willdenovii: a. branches,

bar = 5 cm (2 in.); b. leaf arrangement, bar = 2 mm;
c. leaves, lateral (below), median (middle), and axillary
Figure 13.113.1. Selliguea feei:
(above), bar = 1 mm.
a. habit, bar = 5 cm (2 in.);
b. fertile portion of frond, bar =
1 cm (0.4 in.).
Selliguea
syn. Crypsinus
The best-known cultivated species in this genus, Selliguea
feei, is grown as a novelty in pots or baskets or, in subtrop-
ical climates, in the ground. Selliguea consists of small- to
medium-sized epiphytic, epilithic, or terrestrial species.
The rhizomes are relatively long-creeping and contain con-
spicuous black fibers (seen in cross section). The rhizome
scales are peltately attached, lanceolate, attenuate or with
hair-like tips, and opaque or with a membranous margin; if
the margins are toothed or ciliate, the projections are glan-
dular. The fronds are distant along the rhizome and jointed
to it. The sterile and fertile fronds are dimorphic or not; if
dimorphic, the fertile fronds are narrowed or skeletonized
and taller than the sterile ones. The glabrous sterile fronds b
are simple, pinnatifid, hastate, or subpinnate, with carti-
laginous margins usually sinuate or minutely notched. The
veins are netted and form irregular areoles with free, in- a
cluded veinlets pointing in all directions. The main veins
are conspicuous and nearly reach the margin. The sori are
elongate or linear, and indusia are absent, as in all polypo-
diaceous ferns.
Because the distinguishing characters are inconsistent,
the genus Crypsinus is now included in Selliguea (the older
 508 C 13

Solanopteris
Potato fern Figure 13.114.1. Solanopteris
Comprising small epiphytes, the genus Solanopteris is bifrons: habit, bar = 5 cm (2 in.).
grown as a novelty for its globose tubers (modified stems),
which in nature are inhabited by ants. Growers report that
the plants often do well for a while then suddenly die, al-
though plants at the Marie Selby Botanical Garden in
Florida have been flourishing for a long time. Solanopteris
grows high in trees in its native, wet-forest habitat.
This genus and Lecanopteris are the only ferns that have
stems inhabited by ants. The ants fiercely attack anything
that disturbs the plant. In Solanopteris, they eventually fill
the tuber with organic debris and frass, at which point the
ants seek another tuber in which to live. The plant’s roots
then grow inside the tuber and extract water and mineral
nutrients from the organic matter left behind by the ants.
Thus, the plant receives two benefits from the ants’ activi-
ties: protection and nutrients. The plants seem to grow bet-
ter when inhabited by ants; however, the tubers will form
even if ants are absent. For more about the natural history
of this fern, see Moran (1992b).
In addition to the tubers, Solanopteris is characterized
by long-creeping rhizomes covered by minute, circular
scales. The sterile and fertile fronds are dimorphic, with the
sterile fronds entire to pinnately lobed, and the fertile ones
longer and narrower. The sori vary from round to elongate
and, like in all polypodiaceous ferns, they lack indusia.
Some pteridologists consider Solanopteris close enough
to Microgramma to be placed in that genus. It is kept dis-
tinct here because the name has been used in many floras
and is well established in horticulture.
The genus is native to the American tropics and con-
sists of four species. The name is derived from the potato
genus, Solanum, and the Greek pteris, fern, in reference to
the potato-like tubers.

Solanopteris bifrons (Hooker) Copeland

 13.114.1
Tender
A small fern with medium-creeping rhizomes. Grows
under high-medium light in moist to moist-wet potting
mix or uncut moss with good drainage. This species is usu-
ally planted in a pad of uncut moss and fastened to a hang-
ing board. It is difficult to grow.
The sterile fronds of Solanopteris bifrons are pinnately
lobed, and the tubers are covered by a glaucous bloom. This
species is native to the Amazonian lowlands of Colombia,
Ecuador, and Peru.

Solanopteris brunei (Wercklé ex H. Christ)

W. H. Wagner  13.114.2 Figure 13.114.2. Solanopteris brunei: habit, bar = 5 cm
Tender (2 in.).
A small fern with medium-creeping rhizomes. Grows
under high-medium light in moist to moist-dry(?) potting
 F  F A  C 509

mix or uncut moss with good drainage. Usually planted in

a pad of uncut moss and fastened to a hanging board. This
species is difficult to grow.
The sterile fronds of Solanopteris brunei are entire or
have slightly wavy margins. The tubers bear long, narrow
scales. This species is native from Costa Rica to Ecuador.

Sphaeropteris  13.115.1–6

Tree fern
This genus belongs to the Cyatheaceae, which is the major
family of tree ferns. The family also includes Alsophila, Cne-
midaria, and Cyathea (which see for a comparison of these
genera). Sphaeropteris can be distinguished from these oth-
ers by its stipe scales, which are typically white or brown,
not purplish or black, and are composed of cells that have
the same shape (narrowly rectangular) throughout the scale,
as compared to the differentiated border of diverging, ir-
regularly shaped cells in the other tree-fern genera. Such
scales are said to be “conform” or “nonmarginate.” In many
species of Sphaeropteris the scale margins bear short, dark
teeth (a hand lens is needed to see these).
The Australian Sphaeropteris cooperi is probably the most
common tree fern in the trade and is readily available from
nurseries in California and Florida. It grows rapidly in
many gardens, reaching 5 m (16 ft.) tall. The blond tree
fern (S. horrida), which has stipes neatly and densely cov- Figure 13.115.2. Sphaeropteris cooperi: scales on
ered with blond scales, has been reintroduced into United stipes and trunk.
States cultivation. New introductions and reintroductions
of other species continue.
The trunks and stipe bases of Sphaeropteris plants are
scaly (not hairy as in the dicksoniaceous tree ferns), and the
blades are two- to three-pinnate-pinnatifid. Sori are located
on the lower surfaces of the blades (not the margins) and are
either naked or protected by a globose indusium.
Sphaeropteris, which contains about 110 species, is na-
tive to the American tropics, India, southeastern Asia, New
Zealand, and the islands of the Pacific. The genus name

Figure 13.115.3. Sphaeropteris cooperi: sori.

Figure 13.115.1. Sphaeropteris cooperi: habit.

 510 C 13

Figure 13.115.5.
Sphaeropteris
cooperi: tip of
stipe scale,
not bristle tipped,
bar = 1 mm.

Figure 13.115.4. Sphaeropteris cooperi: frond.

comes from the Greek sphaera, sphere, and pteris, fern, al-
luding to the globose sori.

Sphaeropteris aramaganensis (Kanehira) R. M. Tryon

 13.115.7
syn. Cyathea aramaganensis Kanehira
Mariana Island tree fern
Semi-tender or hardier
A large tree fern. Grows well under medium or higher light
in moist-dry garden soil or potting mix. The plants are ro-
bust growers and occasionally form offsets on the sides of
the trunks. The offsets should be removed to maintain the
symmetry of the plant and to avoid crowding the main
crown. The removed offsets sometimes take root.
The stipes of Sphaeropteris aramaganensis are prickly on Figure 13.115.6. Sphaeropteris cooperi: central part of
the lower surfaces and covered by pale brown, slightly stipe scale, with the marginal cells same as the central
shiny, narrowly triangular scales that are long-tapered and cells, bar = 1 mm.
 F  F A  C 511

c b

a c b
a

Figure 13.115.7. Sphaeropteris aramaganensis: a. pinnule,

bar = 1 cm (0.4 in.); b. stipe scales, bar = 1 cm (0.4 in.);
c. scale from underside of segment midrib (side view),
Figure 13.115.8. Sphaeropteris cooperi: a. pinnule, bar =
bar = 0.1 mm.
1 cm (0.4 in.); b. large stipe scales (left) and small stipe
scales (right), bar = 1 cm (0.4 in.); c. scale from
underside of minor leaf axis, bar = 0.1 mm.
twisted at the apex. The segment midribs on the lower sur-
face near the base have small, whitish, puffy scales. The sori
have many hair-like paraphyses mixed among the sporangia
(magnification needed) and lack an indusia. The species is trunks thicker; the growth is more rapid and robust. It is
native to the Mariana Islands. said to withstand Rhizoctonia disease better than the usual
trade form.
Sphaeropteris cooperi (F. V. Müller) R. M. Tryon ‘Robusta’. Tall and rapid growing, otherwise not dis-
 13.115.1–6, 8;  2, 3 tinct. It was difficult to propagate and no longer in the
syn. Cyathea cooperi (F. V. Müller) Domin trade.
Australian tree fern, Cooper’s tree fern, coin-spot tree
fern Sphaeropteris excelsa (Endlicher) R. M. Tryon
Semi-tender  13.115.9
syn. Cyathea brownii Domin
A large tree fern. Grows well under medium to high light
Norfolk Island tree fern
and may tolerate nearly full sun. The plants prefer moist-
Semi-hardy to semi-tender
dry garden soil or potting mix, but they are relatively toler-
ant of aridity. This species survives occasional cold weather A large tree fern, up to 12 m (40 ft.) tall. Thrives under me-
to near −5°C (20°F) for 4–6 hours; plants die completely dium light in moist garden soil or potting mix. This species
at temperatures of −7°C (19°F) lasting one week. Cold seems to grow more rapidly and robustly than Sphaeropteris
weather can kill fronds, but the plants usually produce new cooperi.
ones. These plants grow rapidly. Mature plants shed their Sphaeropteris excelsa has a dark brown trunk, and the
old fronds, leaving numerous rounded to oval leaf scars on stipe bases are typically dark brownish and prickly on the
the trunks. The scars resemble coins, hence the name coin- lower surfaces. The stipes are covered by nonmarginate
spot tree fern. This species has become naturalized in the (conform) scales of two types: one is longer and wider,
wet tropics. It is the most common tree fern cultivated in
the United States. It is often confused with the seldom cul-
tivated Alsophila australis.
The stipes of Sphaeropteris cooperi vary from greenish to c b
tan and are more or less smooth beneath (not spiny). The
stipe scales are mostly of two types: one is long, linear-tri-
angular, to 4 cm (1.5 in.) long, mostly smoky white, with a
margins irregularly dark bordered or not and bearing
minute, dark teeth; the other is short, to 2 cm (0.75 in.),
linear, dark, and tending to be straight and bearing minute
teeth. The costules and midribs have very small, narrow-
triangular, spiny scales. The sori are surrounded beneath
by small, fringed scales, and a few hair-like paraphyses
occur among the sporangia. An indusium is absent. The Figure 13.115.9. Sphaeropteris excelsa: a. pinnule, bar =
species is native to Australia. 1 cm (0.4 in.); b. large stipe scales (left) and small stipe
‘Brentwood’. Plants more massive than the typical form scales (right), bar = 1 cm (0.4 in.); c. spiny, ovate scale
cultivated in the United States. The fronds are fuller and from underside of minor leaf axis, bar = 0.1 mm.
 512 C 13

whitish, long-linear triangular, and has a narrow, trim, dark

margin bearing small, dark teeth; the other is short, dark,
linear, often twisted, and bears minute spines (magnifica- c b
tion needed). The lower surfaces of the pinnules and seg-
ment midribs bear a mat of tangled, whitish, fringed, vari-
ously shaped small scales that fall early. The sori have a
paraphyses but lack indusia. This species is endemic to
Norfolk Island.
Sphaeropteris excelsa closely resembles S. cooperi but dif-
fers by more foliaceous fronds, prickly stipes, whiter stipe
scales, and the mat of whitish scales on the lower surfaces of
the pinnules and segment midribs. Figure 13.115.11. Sphaeropteris medullaris: a. pinnule,
bar = 1 cm (0.4 in.); b. stipe scales, bar = 1 cm (0.4 in.);
Sphaeropteris horrida (Liebmann) R. M. Tryon c. scale from underside of minor leaf axis, bar = 0.1 mm.
 13.115.10
syn. Cyathea princeps E. Mayer
Blond tree fern, monkey-tail fern
ing degrees. The species is found only occasionally in the
Semi-tender
trade.
A large tree fern. Best grown under medium or higher light A distinctive feature of Sphaeropteris medullaris is its
in moist garden soil or potting mix. This handsome fern is black trunk with patterned, hexagonal leaf scars left by the
conspicuous for its trim, blond scales covering the stipes fallen fronds. The stipes are black and covered by nonmar-
and rachises. ginate (conform), linear, black scales. The pinna midribs
The stipes of Sphaeropteris horrida bear a dense layer of on the lower frond surfaces are typically blackened, and the
pale, nonmarginate (conform) scales. The pinna midribs scales of costae and costules are ovate with spiny margins,
are roundish in cross section and bear tubercles at the bases the spines often black or black-tipped. The indusia are glo-
on the lower surfaces. The midribs of the pinnules and seg- bose and completely surround the sporangia. The species
ments have fine, pale, jointed hairs on the lower surfaces. occurs from New Zealand to the Pitcairns.
The sori are completely enclosed by globose indusia. This
species is native to Mexico and Central America. Stenochlaena
This fern has scandent rhizomes that climb trees. The fronds
Sphaeropteris medullaris (G. Forster) Bernhardi
are large, coarse, and somewhat far apart. The plants are best
 13.115.11;  1
displayed by allowing them to climb trees or by planting
syn. Cyathea medullaris (G. Forster) Swartz
them in big tubs provided with a moss-covered pole. When
Black tree fern, sago tree fern
planted in baskets or pots, Stenochlaena usually becomes too
Semi-hardy
scandent in growth to maintain a neat appearance.
A large tree fern. Best grown under medium or higher light. The long-creeping, climbing rhizomes are green and
This species grows well in moist garden soil or potting mix covered by peltate scales that fall away early. The fronds are
and should be planted in areas sheltered from the wind. distant and dimorphic. The sterile fronds are one-pinnate
The plants do especially well in cool, humid coastal cli- with veins forming a single row of narrow areoles next to
mates. They are conspicuous by their (usually) black the costa; the veins are otherwise free and forked. The fer-
trunks. The rachises and blade scales are also black to vary- tile fronds are one- or two-pinnate with contracted pinnae
and pinnules that are completely covered beneath with spo-
rangia. Indusia are absent.
b
Stenochlaena contains six species and occurs mostly in
the tropical regions of Africa and Malaysia. The genus
name comes from the Greek stenos, narrow, and chlaena,
a cloak, referring to the enrolled margins of the narrow fertile
pinnae.

Stenochlaena tenuifolia (Desvaux) T. Moore

 13.116.1, 2
Semi-hardy to semi-tender
Figure 13.115.10. Sphaeropteris horrida: a. pinnule, bar = A medium-large fern with long-creeping, climbing rhi-
1 cm (0.4 in.); b. stipe scales, bar = 1 cm (0.4 in.). zomes. Grows well under high light in moist garden
 F  F A  C 513

a b

Figure 13.116.1. Stenochlaena tenuifolia: habit.

Figure 13.116.2. Stenochlaena tenuifolia: a. habit with sterile

frond, bar = 5 cm (2 in.); b. fertile frond, bar = 5 cm (2 in.).
soil. Use care in transplanting. Plants may be propagated
from apical sections of the rhizome but are slow to estab-
lish. This species is able to survive three days of −8°C
(18°F) temperatures. tractive as accent plants. Most species must be protected
The sterile and fertile fronds of Stenochlaena tenuifolia during the winter (except in southern Florida) and seem to
are strongly dimorphic: the sterile fronds are one-pinnate need frequent watering and high humidity to grow well.
with numerous pairs of close-set, serrulate pinnae, and the They are generally easy to grow from spores. The often soft
fertile fronds are two-pinnate with greatly contracted pin- foliage of Tectaria is frequently attacked by scale insects and
nae. This species is native to tropical Africa. leaf-spot diseases–a situation that is exacerbated in some spe-
Stenochlaena tenuifolia is often misidentified in cultiva- cies by additional damage from insecticides and fungicides.
tion as S. palustris (N. L. Burman) Beddome, a closely re- Tectaria is a large and diverse genus, with more than
lated species that might be cultivated in the Hawaiian 150 species. It is difficult to characterize, but nearly all the
Islands. Stenochlaena palustris has sterile pinnae set far apart species have netted veins, basal pinnae that are broader on
on the rachis, inconspicuous marginal teeth, and one- the lower half than the upper, and minutely hairy upper
pinnate fertile fronds. It is native to Asia, Australia, and surfaces of the rachises and costae. The juvenile fronds typ-
Polynesia. ically differ greatly from the mature ones. Other character-
istics of the genus include scaly rhizomes that vary from
Tectaria  13.117.1, 2 creeping to erect, more than three vascular bundles in the
syn. Cionidium, Hemigramma, Quercifilix stipes, blades sinuate to lobed to three-pinnate-pinnatifid,
Most cultivated species of Tectaria are terrestrial, small- to and apices reduced to shallow lobes or hastate. The sori
medium-sized ferns grown in pots as house or greenhouse vary from small dots to elongate or (rarely) linear. The in-
plants. Some of the coarsely cut or trilobed species are at- dusia can be either present or absent. If present, the indu-
514 C 13

roof, and -aria, a substantive suffix, alluding to the roof-

like indusium of certain species.

Tectaria beccariana (Cesati) C. Christensen

 13.117.3
Tender or perhaps slightly hardier
A medium-large fern with erect rhizomes. Grows well
under low-medium to medium light in moist potting mix.
This species is best used as an accent plant.
The upper part of stipe has a narrow, ruffled wing, and
the blades are coarsely trilobed or more deeply lobed, with
the lower pinna pair free. The veins are netted, not free,
and the sori are elongate, numerous, scattered, and non-
indusiate. Tectaria beccariana is native to the Philippines
and New Guinea.

Tectaria decurrens (C. Presl) Copeland

 13.117.4
Tender
A medium-large fern with erect rhizomes. Grows well
under medium light in moist potting mix. This species is
best used as an accent plant.
The stipes of Tectaria decurrens are winged nearly to the
base, and the blades are deeply pinnatilobed. The veins are
Figure 13.117.1. Tectaria gemmifera: habit.

sia may be kidney-shaped, peltate, horseshoe-shaped, or

rarely linear.
Tectaria has not been thoroughly studied, and its species
are sometimes difficult to identify. It can be distinguished
from related ferns, such as Arachniodes, Dryopteris, and
Polystichum, by the pinna midribs, which have ungrooved
upper surfaces and are minutely pubescent; the other gen-
era have grooved pinna midribs, with the groove decurrent
into the main rachis, and are rarely pubescent.
Tectaria is widespread in the tropics and subtropics of
the world. The genus name comes from the Latin tectum,

Figure 13.117.3. Tectaria

beccariana: juvenile frond,
bar = 8 cm (3 in.).

Figure 13.117.2. Tectaria gemmifera: veins and sori, the

indusia shed.
 F  F A  C 515

b
b

Figure 13.117.5.
Figure 13.117.4. Tectaria decurrens: a. frond, bar = 8 cm a Tectaria gemmifera:
(3 in.); b. sori and indusia, bar = 5 mm (0.2 in.). a. frond, bar = 8 cm
(3 in.); b. bud, bar =
2 cm (0.8 in.).

netted, not free. The indusia are horseshoe-shaped. This

species is native to the Asian tropics, the Himalayas, Tai-
wan, and Polynesia. coarsely cut foliage is visually striking because of its broad
pentagonal outline.
Tectaria gemmifera (Fée) Alston Tectaria heracleifolia has one-pinnate fronds with three
 13.117.1, 2, 5 to five pairs of pinnae on mature plants. The lateral pinnae
Button fern are slightly cordate, the basal pinnae each with one large
Semi-tender lobe and other shallower lobes. The indusia are peltate.
A medium-sized fern with erect rhizomes. Grows well
under medium light in moist to moist-dry garden soil or
potting mix. Tectaria gemmifera is the most commonly cul- b
tivated species in the genus. The plants volunteer readily
from buds and tolerate subtropical temperatures with less
humidity. a
The fronds of this species are two-pinnate-pinnatilobed
and bud-bearing. The indusia are round-kidney-shaped
and minutely ciliate. Tectaria gemmifera is native to African
tropics.
In the trade Tectaria gemmifera is often misidentified as
T. cicutaria (Linnaeus) Copeland.

Tectaria heracleifolia (Willdenow) Underwood

 13.117.6
Halberd fern
Tender
A medium-large fern with erect rhizomes. Grows well Figure 13.117.6. Tectaria heracleifolia: a. juvenile frond,
under medium light in moist garden soil or potting mix. In bar = 8 cm (3 in.); b. indusia and veins, bar = 5 mm
greenhouses the plants readily volunteer from spores. The (0.2 in.).
 516 C 13

This species is native to the United States (Florida and

Texas), Mexico, Central and South America, and the West
Indies.
Young plants may be distinguished from the similar
Tectaria incisa by the absence of hairs in the rachis and costa
grooves. b

Tectaria hilocarpa (Fée) M. G. Price  13.117.7

syn. Hemigramma latifolia (Mejen) Copeland, Tectaria
latifolia (G. Forster) Copeland
Tender
A small fern with erect rhizomes. Grows well under low-
medium to medium light in moist potting mix. The plants
are grown as novelty items for their small size, mottled
fronds, and rosette arrangement. They respond well to fre-
quent applications of fertilizer.
The sterile fronds of Tectaria hilocarpa have light green
blotches, although plain green variants are also known; the
fertile fronds are contracted and linear and have irregular,
elongate sori. Indusia are absent. This species is native to
Java and the Philippines.

Tectaria incisa Cavanilles  13.117.8

Tender
A medium to large fern with erect rhizomes. Grows well
under medium light in moist soil or potting mix. a
Tectaria incisa is a coarse fern with one-pinnate blades
and three to six pairs of pinnae with oblique bases and
mostly subentire or undulate margins. Minute hairs are
present in the grooves of the rachises and costae. The main
Figure 13.117.8. Tectaria incisa: a. frond, bar = 8 cm
(3 in.); b. indusia, bar = 5 mm (0.2 in.).

Figure 13.117.7. Tectaria

a lateral veins are conspicuous and parallel. The indusium is
hilocarpa: a. frond, bar =
2 cm (0.8 in.); b. fronds, false-peltate. This species is native to the United States
fertile (left) and sterile (Florida), Mexico to South America, and the West Indies.
(right), bar = 8 cm (3 in.). Tectaria incisa is highly variable in blade cutting and pu-
bescence. As presently defined, it probably represents a
complex of several undescribed species.

Tectaria vivipara Jermy & T. G. Walker

 13.117.9
syn. Tectaria incisa forma vivipara (Jenman) C. V.
b Morton
Tender
Medium-sized fern with erect rhizomes. Grows well under
low to medium light in moist soil.
Tectaria vivipara resembles T. incisa, but it can be dis-
tinguished by the buds produced in the axils of the medial
and distal pinnae. This species also tends to have slightly
more pinna pairs than T. incisa (8–12 vs. 3–6), and both
surfaces of the blade are glabrous. The species is widespread
in the American tropics.
 F  F A  C 517

the sterile. The stipes are elongate and the blades greatly
contracted, with the sporangia acrostichoid, spreading
b across the lower surface and lacking indusia. This species is
native to Sri Lanka, India, southern China, Vietnam, and
c Taiwan.

Thelypteris  13.118.1, 2

syn. Amauropelta, Christella, Cyclosorus, Goniopteris,
Lastrea, Meniscium, Oreopteris, Parathelypteris,
Stegnogramma
Thelypteris is a large genus of mostly medium-sized, terres-
trial ferns. They are useful in borders, beds, backgrounds,
and natural plantings. Some species are adapted to wet
areas. Species with erect or semi-erect rhizomes are hand-
some in pots. Those with long-creeping rhizomes can be-
come weedy if planted in the ground. A wide choice of spe-
cies is available for use in warm-temperate and subtropical

Figure 13.117.9.
Tectaria vivipara:
a. frond, bar = 8 cm
(3 in.); b. bud at base
a
of pinna, bar = 5 cm
(2 in.); c. indusium,
bar = 5 mm (0.2 in.).

Tectaria zeylanica (Houttuyn) Sledge

 13.117.10
syn. Quercifilix zeylanica (Houttuyn) Copeland
Tender
A small fern with branching, short-creeping to ascending
rhizomes. Grows well under medium light in moist pot-
ting mix. Because of its small but attractive fronds, this spe-
cies makes an excellent terrarium or pot plant.
The sterile and fertile leaves of Tectaria zeylanica are
strongly dimorphic. The sterile fronds are shaped like small
oak leaves. The fertile fronds stand erect and are taller than

Figure 13.117.10. Tectaria

zeylanica: habit, with
contracted fertile frond,
bar = 8 cm (3 in.).

Figure 13.118.1. Thelypteris puberula: habit.

518 C 13

2. Blades narrowed toward the base (The-

lypteris limbosperma, T. nevadensis, T. noveboracen-
sis, T. resinifera, T. rudis, and T. quelpaertensis).
1. Veins meeting the margin at the bottom of the sinus;
blades simple to one-pinnate-pinnatifid.
3. Blades with a terminal pinna resembling
the lateral ones (Thelypteris poiteana, T. reticulata,
and T. serrata).
3. Blades tapered to the tip, pinnatifid toward
the apex, the apical part unlike the lateral pinnae
(Thelypteris acuminata, T. augescens, T. dentata, T.
guadalupensis, T. kunthii, T. ovata, T. parasitica, T.
Figure 13.118.2. Thelypteris puberula: sori and patens, T. puberula, and T. reptans).
indusia.
Thelypteris acuminata (Panz) C. V. Morton
 13.118.3
syn. Cyclosorus acuminatus (Houttuyn) Nakai, Thelypteris
climates. Generally, the species of Thelypteris are easy to ‘K. O. Sessions’
grow and establish rapidly from spore. Some species (such Semi-hardy
as Thelypteris dentata and T. parasitica) frequently volunteer A medium-sized, deciduous or semi-deciduous fern with
in greenhouses or appear as contaminants in spore pans. long-creeping rhizomes. Grows well under high light in
For a long time Thelypteris was included in Dryopteris, moist to moist-dry garden soil. The plants can tolerate
but it differs by typically having one-pinnate-pinnatifid short periods of drought.
blades, sharp-pointed hairs, and two vascular bundles in Thelypteris acuminata is characterized by long-creeping
the stipes. Other characteristics of the genus are scaly rhi- rhizomes, abruptly narrowed and long-tapered blade
zomes that vary from erect to creeping and rachises and
costae that are usually rounded, not grooved, on the upper
surface, but if grooved, the grooves are interrupted at the
rachis-costa junctures, not continuous. Whitish or yellow-
ish bumps or peg-like structures known as aerophores are
usually present at the base of the rachis-costa junctures on
the lower surface. Depending on the subgroup, the veins
are free or netted. Sori are borne on the lower surfaces of the
blades (not the margins) and are almost always round. In-
dusia may be present or absent. If present, they are mostly
kidney-shaped. b
Thelypteris is cosmopolitan and contains about 1000
species. Its name comes from the Greek thelys, female, and
pteris, fern.
Various authors have divided Thelypteris into as many as
30 genera, most of which are difficult to identify and are
based on obscure or microscopic characteristics such as
chromosome number, spores, and hair type. The nine gen-
era listed in the synonymy above are only those that pertain
to the commonly cultivated species of Thelypteris. Two seg-
regate genera are treated as distinct in this book: Macrothe-
lypteris and Phegopteris.
Because there are so many cultivated species of The-
lypteris, the following key to species-groups is provided to a
facilitate identification.

1. Veins free, meeting the margin above the sinus;

blades one-pinnate-pinnatifid.
2. Blades widest or nearly so at the base (The- Figure 13.118.3. Thelypteris acuminata: a. habit, bar =
lypteris palustris and T. simulata). 5 cm (2 in.); b. segments, bar = 1 mm.
 F  F A  C 519

apices, elongate pinnae, and veins of each segment united narrowed frond apices and irregularly crimped hairs on the
with the vein of an adjacent segment and forming an ex- lower pinna surfaces.
current vein running to the sinus. Native to eastern Asia.
Thelypteris dentata (Forsskål) E. St. John
Thelypteris augescens (Link) Munz & I. M. Johnston  13.118.5
 13.118.4 syn. Christella dentata (Forsskål) Brownsey & Jermy,
Abrupt-tipped maiden fern Cyclosorus dentatus (Forsskål) Ching
Tender, Zone 10 Downy thelypteris
A large fern, up to 120 cm (4 ft.) tall, with medium-creep- Semi-tender, Zone (7)8
ing rhizomes. Grows well under high light in moist, basic A medium-sized fern with short-creeping to suberect rhi-
garden soil or potting mix. zomes. Grows well under medium light in moist garden
The abruptly narrowed frond tip and long, narrow pin- soil or potting mix. The plants are easy to grow but do
nae are distinct features of Thelypteris augescens. The apical poorly in areas with cool nights. They often become weedy
portions of the leathery blades are about five times longer in greenhouses.
than they are broad, and the lateral pinnae are less than 1.5 Thelypteris dentata can be identified by the combination
cm (0.6 in.) wide. On the undersurfaces of the pinnae the of one-pinnate-pinnatifid blades, basal pinnae that are usu-
costae, costules, and veins are sparsely covered with hairs ally reduced and bent forward and downward, and basal
and scales. This species is native to Florida, Cuba, the Ba- veins that unite and form an excurrent vein that runs to the
hamas, and Central America. base of the sinus. Other helpful characteristics are the often
Much of the cultivated material by this name is actually
Thelypteris puberula, which differs by having less abruptly

b
Figure 13.118.5.
Thelypteris dentata:
a. habit, bar = 5 cm
(2 in.); b. segments,
bar = 1 mm.

a
b
Figure 13.118.4. Thelypteris augescens: a. frond, bar =
5 cm (2 in.); b. segments, bar = 1 mm.
 520 C 13

purplish stipes and rachises, and the lower surfaces of the

blades and costae are minutely but evenly hairy, without
glands. The species is native to the tropics of Africa and
Asia but has become widely naturalized in the American
tropics.

Thelypteris guadalupensis (Wikström) Proctor

 13.118.6
syn. Cyclosorus guadalupensis (Wikström) Ching b
Tender
A small fern with short-creeping to nearly erect rhizomes.
Grows well under medium light in moist garden soil or pot-
ting mix.
The stipes and rachises of Thelypteris guadalupensis are
minutely hairy with star-shaped hairs. The lanceolate
blades are mostly lobed or pinnatifid, except pinnate to- Figure 13.118.7.
ward the narrowed base, with the margins entire to crenate a Thelypteris kunthii:
or lobed. The fertile fronds are erect and long-stalked. This a. frond and
species is native to the West Indies. rhizome, bar = 5 cm
(2 in.); b. segments,
bar = 1 mm.

with even-length, long hairs. Hairs are also present on the

upper surfaces. The species is native to tropical and sub-
tropical America.

Thelypteris limbosperma (Allioni) H. P. Fuchs

 13.118.8
syn. Oreopteris limbosperma (Allioni) Holub
Hardy, Zone 5(6)
A medium-sized fern with short-creeping to suberect rhi-
zomes and deciduous fronds. Grows well under medium
light in moist, acidic garden soil or potting mix.
Figure 13.118.6.
The rhizomes of Thelypteris limbosperma are hidden by
Thelypteris guadalupensis:
old stipe bases that have persistent scales. The blades are ta-
habit, bar = 5 cm (2 in.).
pered toward the base and on the lower surfaces usually
have abundant yellow glands. Needle-like hairs (not scales)
occur on the lower surfaces of the costae. The veins meet
above the sinus base. This species is native to Europe.
A Thelypteris along the northwestern coast of North
America has been called this species but is actually The-
Thelypteris kunthii (Desvaux) C. V. Morton lypteris quelpaertensis (H. Christ) Ching, a species of eastern
 13.118.7 Asia. It differs by having few, small, colorless glands on the
syn. Thelypteris normalis (C. Christensen) Moxley lower surfaces of the blades and narrow, pale scales on the
Widespread maiden fern, southern shield fern, southern lower surfaces of the costae (rarely with needle-like hairs).
wood fern
Semi-tender to hardier, Zone (7)8 Thelypteris nevadensis (Baker) Clute ex C. V. Morton
A medium-sized fern with short- to medium-creeping rhi-  13.118.9
zomes. Grows well under medium light in moist garden syn. Parathelypteris nevadensis (Baker) Holttum
soil or potting mix. The plants are easy to grow. Sierra water fern
The basal veins of Thelypteris kunthii are free (not unit- Semi-hardy, Zone 8
ing with that of the adjacent segment) and run to the bases A medium-sized fern with short-creeping to suberect rhi-
of the sinuses. The lower surfaces of the blades are hairy zomes and fronds that wither and persist. Grows well under
 F  F A  C 521

Figure 13.118.8.
Thelypteris limbosperma:
a. habit, bar = 5 cm
(2 in.); b. segments, bar
= 1 mm.
Figure 13.118.9. Thelypteris
nevadensis: a. habit, bar =
5 cm (2 in.); b. segments
dotted with glands, bar =
1 mm.
a

a
moist garden soil or potting mix. The plants are easy to
b grow and may become invasive, tending to spread irregu-
larly throughout the garden if not contained. They are not
adapted to subtropical climates.
Thelypteris noveboracensis is characterized by one-pin-
nate-pinnatifid blades gradually tapered toward the base,
with the lowest pinnae appearing as mere stubs or nubbins.

medium light in moist-wet garden soil or potting mix. This

species does poorly in warm climates.
Thelypteris nevadensis has slender rhizomes 1–3 mm
wide. The blades are tapered toward the base and bear
stalked, yellow-orange glands on their lower surfaces. The
costae are glabrous or sparsely hairy beneath. The veins are
simple, free, and end above the sinus base. This species is
native to northwestern North America.
The commercial plants labeled as Thelypteris nevadensis
are often T. noveboracensis, which differs by lacking glands
on the blades, or if present, then sessile and sparse, and the
hairs on the costae are longer.

Thelypteris noveboracensis (Linnaeus) Nieuwland

 13.118.10 a
syn. Dryopteris noveboracensis (Linnaeus) Nieuwland,
Parathelypteris noveboracensis (Linnaeus) Ching b
New York fern
Very hardy, Zone (3)4
A medium-sized fern with long-creeping rhizomes and de- Figure 13.118.10. Thelypteris noveboracensis: a. habit, bar
ciduous, herbaceous leaves. Grows well under high light in = 5 cm (2 in.); b. segments, bar = 1 mm.
 522 C 13

The rhizomes are long-creeping, and the blades either lack apex and widest (or nearly so) at the base, and fertile pinnae
or are sparsely beset with sessile glands. The costae are con- narrower and more contracted than the sterile ones. The
spicuously hairy on the lower surface. The veins are un- veins are free, once-forked, and meet the margin above the
branched and meet the margin above the sinus. This species sinus. The lower surfaces of the costae bear scattered scales.
is native to eastern North America. The species is native to North America, Mexico, the West
Indies, and Europe.
Thelypteris ovata R. St. John  13.118.11 A crested variant (forma pufferae A. A. Eaton) is re-
Semi-hardy, Zone 8 ported in cultivation.
A medium-sized fern with medium-creeping rhizomes.
Grows well under medium light in moist to moist-dry gar- b
den soil or potting mix. The plants are easy to grow.
Thelypteris ovata is characterized by blade apices less
than five times long as wide, more or less attenuate and dif-
ferent from the lateral pinnae, and sessile pinnae with the
basal segment slightly longer than the other segments and
often toothed. Unlike in similar species, the costules and
veins on the upper surface are glabrous and the sori are sub-
marginal. The species is native to the southeastern United
States, Central America, and the West Indies.

b c

Figure 13.118.12. Thelypteris palustris: a. habit, bar =

5 cm (2 in.); b. sterile segments, bar = 1 mm; c. fertile
segments, bar = 1 mm.

Figure 13.118.11.
Thelypteris ovata: a.
habit, bar = 5 cm Thelypteris parasitica (Linnaeus) Fosberg
a  13.118.13
(2 in.); b. segments,
with elongate basal syn. Christella parasitica (Linnaeus) H. Léveillé,
segments, bar = Cyclosorus parasiticus (Linnaeus) Farwell
1 cm (0.4 in.). Semi-tender to slightly hardier
A medium-sized fern with medium-creeping rhizomes.
Grows well under medium light in moist garden soil or pot-
ting mix. The plants volunteer readily from spores.
In Thelypteris parasitica, the basal veins on a segment
unite with those of adjacent segments to form an excurrent
Thelypteris palustris Schott  13.118.12 vein that runs to the base of the sinus. Both surfaces of the
syn. Thelypteris thelypteroides (Michaux) Holub blade–particularly the veins on the lower one–have hairs
Marsh fern and stalked, orange glands. The indusia are long-ciliate.
Very hardy, Zone (2)3 The species is native to tropical Asia and the islands of the
A medium-sized fern with long-creeping rhizomes and de- Pacific.
ciduous fronds. Grows well under medium-high light in
moist-wet garden soil or potting mix.
Thelypteris palustris can be identified by long-creeping
rhizomes, one-pinnate-pinnatifid blades tapered toward the
 F  F A  C 523

b
Figure 13.118.13.
Thelypteris
parasitica: a.
habit, bar =
5 cm (2 in.); b.
segments with
minute glands on
segment midrib,
bar = 1 mm.

a
c

Figure 13.118.14.
Thelypteris patens: a. habit,
bar = 5 cm (2 in.); b. pinna
base, bar = 1 cm (0.4 in.);
b c. stipe scales, bar = 1 cm
(0.4 in.).

Thelypteris patens (Swartz) Small  13.118.14

Grid-scale maiden fern
Tender, Zone 10
A medium-large fern with stout, suberect to erect rhizomes.
Grows well under medium light in moist garden soil or pot-
ting mix.
The scales at the base of the stipe are distinctive in The-
lypteris patens: they are tan, broadly ovate, and persistent.
Also distinctive are the pinna bases, which are sessile and
have the basal segments slightly more elongate and toothed
than the adjacent ones. The species is native to Florida,
Central and South America, and the West Indies.
‘Lepida’. Blades skeletonized.

Thelypteris poiteana (Bory) Proctor  13.118.15 Figure 13.118.15. Thelypteris

syn. Goniopteris poiteana (Bory) Ching poiteana: habit, bar = 5 cm (2 in.).
Tender
A medium-sized fern with hard, medium-creeping rhi-
zomes and few fronds. Grows well under medium light in
moist garden soil or potting mix.
Thelypteris poiteana is characterized by netted veins and
one-pinnate blades with a terminal pinna that resembles the
 524 C 13

two to five lateral ones. The pinnae are elliptic with suben-
tire to coarsely serrate-crenate margins. Minute, forked, or
star-shaped hairs are present only on the rachis near the
junctions with the costae. The sori lack indusia and form a
double row between the main lateral veins. This species is
native to, and widespread in, the American tropics.
Figure 13.118.17.
Thelypteris puberula (Baker) C. V. Morton Thelypteris reptans:
 13.118.1, 2, 16 fronds, bar = 5 cm
Semi-tender, Zone (8)9 (2 in.).
A medium-sized fern with medium- to long-creeping rhi-
zomes. Grows well under high-medium light in moist-dry
garden soil or potting mix.
The blades of Thelypteris puberula are broadest at or near
the base, gradually or somewhat abruptly tapered toward
the apex, and glabrous on the upper surface except for
crimped hairs on the costae. The basal veins of each seg-
ment end at or above the sinus. The indusia are densely
hairy. This species is native to the southwestern United
States, Mexico, and Central America.

Thelypteris reptans (J. F. Gmelin) C. V. Morton

 13.118.17
syn. Goniopteris reptans (J. F. Gmelin) C. Presl
Creeping woodfern, creeping star-hair fern
moist-dry garden soil or potting mix. The plants are easy
Tender, Zone 9, with warm nights
to grow.
A small fern with short-creeping to suberect rhizomes and Thelypteris reptans is characterized by its small size and
evergreen fronds. Grows well under medium light in basic, elongate blade apices that bear buds. If you look along the
rachis with a hand lens, you will see minute star-shaped,
forked, or needle-like hairs. The indusium is absent or, if
present, minute and conspicuously ciliate. The fertile
fronds are erect, whereas the sterile ones are spreading. The
species is native to tropical America.

Thelypteris resinifera (Desvaux) Proctor

 13.118.18
b syn. Amauropelta resinifera (Desvaux) Pichi-Sermolli
Wax-dot maiden fern, glandular wood fern
Tender, Zone 9
A medium-sized fern with short-creeping to erect rhizomes
and evergreen, erect fronds. Grows well under medium
light in moist-wet garden soil or potting mix.
Thelypteris resinifera is characterized by sessile pinnae,
Figure 13.118.16. blades gradually narrowed toward the base, and resinous,
Thelypteris puberula: red, sessile glands on the lower surfaces of the blades. Other
a. frond and characteristics include rhizomes hidden by old stipe bases;
rhizome, bar = 5 cm erect, rigid, blades; and veins free and reaching the margin
(2 in.); b. segments, above the base of the sinus. The species is native to tropical
bar = 1 mm. America.
This species is sometimes mislabeled in gardens as The-
lypteris sancta (Linnaeus) Ching, a species unknown in
cultivation.
a
 F  F A  C 525

a
Figure 13.118.18. Thelypteris resinifera: a. habit, bar =
5 cm (2 in.); b. segments dotted with glands, bar =
1 mm.

Thelypteris reticulata (Linnaeus) Proctor

 13.118.19
syn. Meniscium reticulatum (Linnaeus) Swartz
Lattice-vein fern b
Tender, Zone 10
A medium-large fern with short-creeping to nearly erect
rhizomes. Grows well under low light in moist, slightly Figure 13.118.19. Thelypteris reticulata: a. habit, bar =
acidic garden soil or potting mix with good drainage. 5 cm (2 in.); b. margin and veins of pinna, bar = 1 mm.
Thelypteris reticulata has red-brown rhizome scales, and
the stipes of the fertile fronds are slightly longer than those
of the sterile ones. The sterile blades vary from triangular- the scales slightly clathrate. The veins are simple and meet
oblong to lance-oblong and are one-pinnate. The pinnae the margin above the base of the sinus. The sori are round
are oblong-attenuate to lanceolate-triangular, 2–6 cm (0.6– and lack an indusium. This species is native from Mexico to
2.5 in.) wide, and tailed at the apex, with margins wavy to Bolivia.
slightly crenate or entire and without hairs. The veins form
several parallel rows of areoles between costae and margins. Thelypteris serrata (Cavanilles) Alston
Curved-elliptic sori are borne at the middle of transverse  13.118.21
veins and lack indusia. This species is native to tropical and syn. Meniscium serratum Cavanilles
subtropical America. Serrate lattice-vein fern
Tender, Zone (9)10
Thelypteris rudis (Kunze) Proctor  13.118.20 A medium to large fern with short-creeping rhizomes.
syn. Amauropelta rudis (Kunze) Pichi-Sermolli Grows well under medium-high light in moist to moist-
Tender, Zone (9)10 wet garden soil or potting mix.
A medium-sized fern with short-creeping to suberect rhi- The blades of Thelypteris serrata are gradually reduced
zomes. Grows well under medium light in moist-wet gar- toward the apex, the terminal pinna usually smaller than
den soil or potting mix. the lateral ones. The pinnae are hooked-serrate, at least to-
Thelypteris rudis has one-pinnate-pinnatifid blades ward the tips. In mature plants, the veins form 10–18 are-
abruptly reduced toward the base to tiny, wart-like pinnae. oles between the costae and margins. Indusia are lacking.
The lower surfaces of the costae and rachises are hairy and This species is native to Florida, Central and South Amer-
sparsely scaly, the hairs stiff, short, forward-pointing, and ica, and the West Indies.
526 C 13

A similar species, Thelypteris reticulata, differs by hav-

ing nearly entire pinna margins.
Figure 13.118.20.
Thelypteris rudis: a. Thelypteris simulata (Davenport) Nieuwland
habit, bar = 5 cm  13.118.22
(2 in.); b. reduced syn. Parathelypteris simulata (Davenport) Holttum
lower pinnae, bar = Massachusetts fern, bog fern
5 cm (2 in.). Very hardy, Zone 4
A medium-sized fern with medium- to long-creeping, slen-
der rhizomes and deciduous fronds. Grows well under me-
dium light in wet, acidic garden soil or potting mix. This
species is excellent for bog gardens but is more difficult to
grow than Thelypteris palustris, a similar species that grows
in wet soils.
Thelypteris simulata has lanceolate blades with yellow-
ish, short-stalked glands on the lower surfaces and numer-
ous shiny, sessile, reddish to orangish, hemispherical glands
(use a hand lens). The veins are simple, ending above the
sinus. The fertile fronds are slightly longer than the sterile
ones due to longer stipes. The indusia are often glandular,
the glands reddish or orangish. The species is native to east-
ern North America.

Todea
a
A handsome accent or pot plant, Todea forms a large, erect
funnel of bright green foliage. In their native habitat these
terrestrial ferns can become large plants with stout trunks to
1 m (3 ft.) tall and fronds to 1.2 m (4 ft.) long. The trunks
b

Figure 13.118.22. Thelypteris

simulata: a. habit, bar = 5 cm
(2 in.); b. segments dotted
with glands, bar = 1 mm.
a

Figure 13.118.21.
Thelypteris serrata:
a. frond, bar = 5 cm b
(2 in.); b. margin and
veins of pinna (young
plant), bar = 1 mm.
 F  F A  C 527

can be massive and bear multiple crowns. The cultivated

plants in the United States, however, maintain moderate
growth and size.
The rhizomes are erect and, when old, become trunk-
like. The fronds are oblong, two-pinnate, and leathery. The
fertile pinnae are slightly contracted or not, and on their
lower surfaces they bear scattered, large sporangia. An in-
dusium is lacking. The spores are green and short-lived.
This genus, which contains only one species, is native to
southern Africa, Australia, and New Zealand. The name
Todea honors the German botanist Henrrich Julius Tode
(1733–1797).

Todea barbara (Linnaeus) T. Moore

 13.119.1–3 Figure 13.119.2. Todea barbara: sporangia
Semi-hardy to semi-tender; tolerates light frost scattered on pinnules.
A medium to large fern with erect rhizomes capable of
forming offshoots and fronds borne in a spiral. Grows well
under medium to high light in moist garden soil or potting
mix. Todea barbara is easy to grow from spores and usually
does well once established. The spores ripen nearly simul-
taneously and are shed when the sporangia are green. See
the characteristics listed for the genus.

Trachypteris
This genus of small, terrestrial or rock-inhabiting ferns has
short-creeping rhizomes, densely scaly leaves (at least on
the undersurface), and netted veins. The sori lack an indu-
sium. The genus name comes from the Greek word trachys,
rough, and pteris, fern, referring to the rough texture im-
parted by the scales to the fronds.

Figure 13.119.3. Todea barbara: frond, bar = 5 cm

Figure 13.119.1. Todea barbara: habit. (2 in.).
 528 C 13

Trachypteris pinnata (Hooker f.) C. Christensen

 13.120.1 a
Tender
A small fern, about 30 cm (12 in.) tall, that bears a rosette
of fronds from an erect or nearly erect rhizome. Seems to do
best in airy environments with well-drained soil and me-
dium light. Avoid overwatering and overpotting. This spe-
cies is an attractive fern recently introduced from Peru.
The sterile fronds of Trachypteris pinnata are widest
above the middle, blunt-tipped, and arranged in a rosette.
Their upper surfaces bear narrow, whitish scales, whereas
the lower surfaces are densely covered with rusty-red lan- b
ceolate scales. When dry, the fronds curl and expose their
scaly lower surfaces. The fertile fronds are erect and pin-
nate or pinnatifid. They bear sporangia on their undersides Figure 13.121.1. Trichomanes radicans: a. habit;
along the veins. The species is native to Ecuador, Peru, Bo- b. indusium and sorus with extended receptacle.
livia, Argentina, Paraguay, and Brazil.

Most Trichomanes species grow from slender, long-

creeping rhizomes, although the rhizome is erect in Tri-
chomanes holopterum and a few others. The fronds vary tre-
mendously in shape and size but are mostly small and
membranous. The sori are borne at the margin of the blade,
not on the lower surface as in many other ferns. Each sorus
has an elongated tissue (the receptacle) that protrudes as a
bristle above the trumpet- or tubular-shaped indusium.
Figure 13.120.1. Trachypteris
The mouth of the indusium is dilated or sometimes two-
pinnata habit: sterile fronds in
lipped. The spores are green and short-lived.
a rosette, fertile fronds long
Trichomanes and Hymenophyllum are the traditionally
stalked, bar = 5 cm (2 in.).
recognized genera of filmy ferns. They have been split into
as many as 34 segregate genera, and although these largely
represent natural groups, pteridologists generally recognize
only two genera. Hymenophyllum differs from Trichomanes
by the indusium, which in the former consists of two broad
lips and includes the receptacle within it, not protruding
out from the indusium.
Trichomanes occurs mostly in the tropics worldwide,
with a few species in the subtropics and temperate regions.
If defined in the broad sense, as is done here, the genus con-
tains about 300 species. The genus name comes from the
Trichomanes  13.121.1 Greek thrix, hair, and manes, cup, alluding to the recepta-
syn. Cardiomanes, Crepidomanes, Vandenboschia cles protruding from the cup-like indusium. For more in-
Filmy fern, bristle fern formation, see “Filmy Ferns” in Chapter 10.
Filmy ferns are mostly small, terrestrial or epiphytic ferns
with fronds one-cell-layer thick between the veins. This Trichomanes auriculatum Blume  13.121.2
thinness imparts a filmy texture to the blades–hence the syn. Crepidomanes auriculatum (Blume) K. Iwatsuki,
common name. The other common name, bristle fern, Vandenboschia auriculata (Blume) Copeland
refers to the bristle-like receptacle that protrudes beyond Semi-tender (?)
the mouth of the indusium. Because of the thin blades, the A small to small-medium fern with hairy-tipped, medium-
plants need high humidity and should be grown in terrari- creeping rhizomes and distant fronds. Grows well under
ums. Species seem to come and go in the United States low light in moist to moist-wet potting mix or uncut moss
trade, with old introductions often lost and replaced by with good drainage. This species does well in terrariums
new ones. but may go semi-dormant during the winter.
 F  F A  C 529

cies is native to Europe, Madeira, and the Canary Islands.

Trichomanes speciosum is sometimes confused in the lit-
erature with T. radicans Swartz, a pantropical species with
Figure 13.121.2. ovate to lanceolate blades.
Trichomanes
auriculatum: habit,
bar = 5 cm (2 in.).
a

The fronds of Trichomanes auriculatum are sessile or Figure 13.121.3. Trichomanes holopterum: a. frond, bar =
have stipes to 4 mm (0.2 in.) long. The blades are linear-ob- 5 cm (2 in.); b. indusium and sorus with extended
long, 10–40 cm (4–16 in.) long, and one-pinnate, with the receptacle, bar = 1 mm.
sterile pinnae lobed and crenate; the fertile pinnae are
deeply and sharply incised. This species is native from India
to southeastern Asia, Japan, Malaysia to Indonesia and
Micronesia.

Trichomanes holopterum Kunze  13.121.3

Tender
A small fern with short, erect rhizomes. Grows well under
low light in moist to moist-wet potting mix or uncut moss
with good drainage. The plants reliably produce fertile
fronds and are easy to grow; however, they need fresh
media every now and then.
Trichomanes holopterum has tightly clustered, erect
fronds 4–13 cm (1.5–5 in.) long with broadly winged stipes.
The blades are oblong or lanceolate-oblong, stiff, translu-
cent, and deeply pinnatifid into 5–10 pairs of oblong lobes
with coarsely sinuate margins. This species is native to
southern Florida, southern Mexico, and the West Indies.

Trichomanes speciosum Willdenow  13.121.4

syn. Vandenboschia speciosa (Willdenow) Copeland
Killarney bristle fern
Semi-hardy
A small fern with creeping rhizomes. Grows under low light
Figure 13.121.4.
in moist to moist-wet potting mix with good drainage.
Trichomanes
The rhizomes of Trichomanes speciosum are black and
speciosum: frond,
wiry, with bristly hair-like scales. The fronds are generally
bar = 5 cm (2 in.).
5–18 cm (2–7 in.) long with the stipes winged nearly to the
base, the blades triangular and to four-pinnatifid. This spe-
 530 C 13

Vittaria  13.122.1, 2

Shoestring fern
The genus Vittaria derives its common name from the
long, narrow, shoestring-like fronds of most species. These
infrequently cultivated epiphytes are usually grown on
boards or in baskets or pots to display their pendent foliage
and maintain good drainage. The fronds are close together
and make a handsome plant. Young fronds are sometimes
reddish. The genus is characterized by the usually grass- Figure
like, linear fronds and sori that appear in one long line on 13.122.2.
either side of the midrib, with the indusium absent. The Vittaria
rhizome has clathrate scales. lineata:
Crane (1997) showed that the Old World species of Vit- immersed
taria form a group distinct from the New World ones. linear sori.
They differ in technical characteristics, such as rhizome
symmetry, type of gemmae on the gametophytes, and para-
physes. Accordingly, the Old World species have now been
transferred to Haplopteris.
Vittaria is native to the tropics and subtropics world-
wide and contains about 50 species. The genus name comes
from the Latin vitta, ribbon or stripe, and -aris, resembling,
referring to the long, linear leaves of the type species.

Vittaria elongata Swartz  13.122.3

syn. Haplopteris elongata (Swartz) E. H. Crane
Tender
A medium-sized fern with short-creeping rhizomes and
pendent, grass-like leaves. Grows well under medium light
in moist potting mix or uncut moss with good drainage.
Vittaria elongata is distinguished by sori in a marginal
groove, not submarginal as in most other species. Other
characteristics include reddish brown to nearly black rhi-
zome scales, to 1 cm (0.5 in.) long; stipes present; blades to
70 cm (28 in.) by 3–10 mm (0.1–0.4 in.) wide, widest in
the middle; midribs distinct on the upper blade surfaces;
and paraphyses with tips enlarged to an inverted cone-
shape. This species is native to the tropics and subtropics
from Africa to Polynesia.

Vittaria lineata (Linnaeus) J. E. Smith

 13.122.1, 2, 4
Grass fern, shoestring fern
Tender
A medium-sized fern with short-creeping rhizomes and
pendent, grass-like leaves. Grows under medium light in
moist potting mix or uncut moss with good drainage; seems
to do best with high humidity. This species grows readily
from spores, and the prothalli (gametophytes) proliferate
copiously from minute, several-celled gemmae borne along
the margins. The prothalli can form thick mats in terrari-
Figure 13.122.1. Vittaria lineata: habit. ums and remain green for many years. Mature plants are
difficult to transplant.
 F  F A  C 531

obconic tips (a strong hand lens or microscope is needed to

see this). The species is native to tropical and subtropical
America.

a Woodsia  13.123.1–3

Cliff fern
The plants of the genus Woodsia are known as cliff ferns
because of their tendency to grow on or among rocks. They
are mostly small, delicate ferns used in rock gardens in cold-
Figure 13.122.3. Vittaria to cool-temperate climates. Most cliff ferns adapt poorly to
elongata: a. habit, bar = 5 cm warmer climates.
(2 in.); b. paraphysis, bar = The rhizomes are mostly short-creeping and scaly, pro-
0.2 mm. b ducing dense tufts of persistent stipes that may or may not be
jointed (articulate) in the middle. The stipes contain two
vascular bundles. The blades are one-pinnate-pinnatifid to
two-pinnate, usually ovate to linear-lanceolate, herbaceous
to firm, and deciduous. The lower pinnae are shorter and
increasingly placed further apart toward the base of the
frond. The sori are round, not elongate or linear. The indu-
sium is attached beneath the sporangia and comes up around
them, cup-like, to form a protective cover. They vary con-
siderably in shape at maturity, being either entire and glo-
bose or splitting into several lobes or fragile filaments.
Woodsia resembles Cystopteris and can be distinguished
from it by the persistent stipe bases and the indusia attached
completely beneath the sori (instead of only on one side).
a The persistent stipe bases may be all fairly even in length or
uneven, and this trait is helpful in identifying the species.

Figure 13.122.4.
Vittaria lineata:
a. habit, bar = 5 cm
(2 in.); b. sori, bar =
3 mm (0.1 in.).

Vittaria lineata has sori in grooves on the lower surfaces

of the blades, not on the margins. The rhizome scales are
particularly distinctive by their elongate, thread-like tips.
The fronds are up to 60 cm (24 in.) long and 3 mm (0.1
in.) wide. The paraphyses are slender, without expanded Figure 13.123.1. Woodsia scopulina: habit.
532 C 13

Woodsia alpina (Bolton) Gray  13.123.4

Northern woodsia
Very hardy, Zone (1)2
A small fern with erect to ascending rhizomes and decidu-
ous fronds. Grows well under medium light in moist gar-
den soil or potting mix.
A fertile hybrid, Woodsia alpina originated from a cross
of W. glabella and W. ilvensis. It resembles W. ilvensis, but
the hairs and scales on the blades are few or absent, and the
largest pinnae have one to three pairs of pinnules. The spe-
cies is native to northern North America, Greenland, and
northern Europe and Asia.

Woodsia ilvensis (Linnaeus) R. Brown

 13.123.2, 5
Rusty woodsia
Very hardy, Zone (1)2
A small fern with ascending to erect rhizomes and decidu-
ous fronds. Grows under medium light in moist-wet, acidic
garden soil or potting mix. This species is attractive because
of the rusty color of the scales covering the lower surfaces of
the blades, but it may be difficult to grow.
Figure 13.123.2. Woodsia ilvensis: fertile pinnae. The stipes of Woodsia ilvensis break at a swollen joint
near the base and, along with the blades, are copiously cov-
ered with many hairs and scales, especially on the lower sur-
The stipe bases of even length break at a joint that may be face. The largest pinnae have four to nine pairs of pinnules,
seen as a slight ridge or bump on the stipe. Stipe bases that and the indusium consists of slender hairs. This species is
are uneven lack joints. native to northern regions of North America, Europe, and
The genus Woodsia contains about 25 species and is na- Asia.
tive to alpine, boreal, and temperate regions, mostly in the
Northern Hemisphere. The genus is named for Joseph
Woods (1776–1864), a British architect and author of A
Tourist’s Flora.

Figure 13.123.3. Woodsia: sori from different species.

 F  F A  C 533

Woodsia manchuriensis Hooker  13.123.6

Manchurian woodsia
Very hardy, Zone 4
A small fern with short, erect rhizomes and deciduous
Figure 13.123.4. fronds. Grows well under medium light in moist garden
Woodsia alpina: soil or potting mix.
a a. frond, bar = Woodsia manchuriensis has shiny red-brown, minutely
2.5 cm (1 in.); hairy to glabrous stipes that are not jointed and hence break
b. sorus and unevenly. The blades are narrowly elliptic, one-pinnate-
indusium, bar = pinnatifid, and whitish beneath. The indusia are globose,
1 mm. thin, and shallowly and irregularly lobed, with the lobes
not ciliate. The species is native to northeastern Asia.

a
b

Figure 13.123.6. Woodsia

manchuriensis: a. frond, bar
= 2.5 cm (1 in.); b. sorus
Figure 13.123.5. Woodsia and indusium, bar = 1 mm.
ilvensis: a. frond, bar = 2.5 cm
(1 in.); b. sorus and indusium,
bar = 1 mm.

Woodsia obtusa (Sprengel) Torrey  13.123.7

Blunt-lobed woodsia
Very hardy, Zone 3
A small fern with short-creeping or ascending rhizomes and
b deciduous fronds. Grows well under medium light in basic
to neutral, moist-wet garden soil or potting mix. The plants
are easy to cultivate in rock gardens or open woods.
The stipes of Woodsia obtusa are not jointed and break
unevenly. They bear scattered scales or are nearly naked.
 534 C 13

Figure 13.123.8.
Figure 13.123.7. Woodsia Woodsia oregana:
obtusa: a. frond, bar = 2.5 cm a. frond, bar =
(1 in.); b. sorus and indusium, 2.5 cm (1 in.);
bar = 1 mm. b. sorus and
indusium, bar =
1 mm.

a
a

b
b

The blades are glandular, with the glands appearing as

minute hairs with swollen heads (magnification is needed
to see this), and the indusia consist of about four strap-
shaped lobes. This species is native to eastern North Amer-
ica. a

Woodsia oregana D. C. Eaton  13.123.8

Oregon woodsia, western cliff fern
Very hardy, Zone 4
A small fern with ascending to erect rhizomes and decidu-
ous fronds. Grows well under medium light in moist gar-
Figure 13.123.9.
den soil or potting mix with good drainage.
Woodsia polystichoides:
The stipes are not jointed and therefore break unevenly,
a. frond, bar = 2.5 cm
their surfaces smooth or bearing a few scattered scales. The
(1 in.); b. sorus and
blades of Woodsia oregana are naked or have minute glands,
indusium, bar =
and the indusia consist of short hairs about the length of the
1 mm.
sporangia. The species is native to western North America.

Woodsia polystichoides D. C. Eaton  13.123.9

Very hardy, Zone (3)4 b
A small fern with short, erect rhizomes and deciduous
fronds. Grows well under medium light in moist garden
soil or potting mix.
 F  F A  C 535

Woodsia polystichoides has stipes jointed near the top that

break to form an even stubble. The blades are one-pinnate,
and the pinnae are eared (thus making the plant resemble
Polystichum), entire to dentate, and sessile but not decur-
rent on the rachis. The rachis scales are lanceolate, and the Figure 13.123.10.
indusia are subglobose, shallowly lobed, and ciliate. The Woodsia scopulina:
species is native to China, Korea, and Japan. a. frond, bar =
2.5 cm (1 in.);
Woodsia scopulina D. C. Eaton  13.123.1, 10 b. sorus and
Mountain woodsia indusium, bar =
Hardy, Zone 5 1 mm.
A small fern with erect to ascending rhizomes and decidu-
ous fronds. Grows under medium light in moist-wet gar-
den soil or potting mix with good drainage. It is reportedly
difficult to grow.
The stipes of Woodsia scopulina lack joints and therefore
break unevenly. The stipes and blades have white hairs and
minute, short-stalked glands (use magnification). The in- a
dusium consists of narrow, strap-like lobes that fray at the
tips. The species is native to North America.

Woodsia subcordata Turczaninow  13.123.11

Hardy, Zone (5)6
A small fern with short-creeping to ascending rhizomes and
deciduous fronds. Grows well under medium light in moist b
garden soil or potting mix.
Although often inconspicuous, the joints of Woodsia
subcordata occur near the middle of the stipe. The stipe
scales are ovate-lanceolate and sparsely ciliate. The one-
pinnate blades are scaly and hairy on both surfaces, and the a
pinnae are lobed and eared at the base. The rachis scales are
linear, and the indusia are lobed, with the margins of the
lobes densely long-ciliate. The species is native to Korea,
Figure 13.123.11.
Japan, eastern China, and eastern Russia.
Woodsia subcordata:
The identification of this species is tentative. It is one
a. frond, bar = 2.5 cm
of the few Woodsia species that grows well in subtropical
(1 in.); b. sorus and
climates.
indusium, bar = 1 mm.
Woodwardia  13.124.1–3
syn. Anchistea, Lorinseria
Chain fern b
Woodwardia is a genus of medium- to large-sized, coarse,
mostly terrestrial ferns of cool-temperate and warmer areas.
Those species with cascading fronds can be used to cover
banks. Others do well in beds or pots, and some are favored pinnatifid and from firm to subleathery. The margins are
as accent plants. The commonly grown species are easy to minutely to coarsely serrate-dentate. The veins are netted,
culture and are adapted to a wide variety of soil types. The but the areoles lack included veinlets. The sori are mostly
young fronds are reddish. oblong and dorsal, appearing in a row on each side of the
The rhizomes are scaly and in most species short-creep- midribs. The indusia open toward the midribs, not the
ing or ascending, but a few species have slender, long-creep- margins.
ing ones. The sterile and fertile fronds are monomorphic, Woodwardia consists of 14 species and is native to
except for those of Woodwardia areolata, which are strongly North America, Central America, Mediterranean Europe,
dimorphic. The stipes contain more than three vascular and eastern Asia. The genus is named for Thomas Jenkin-
bundles. The blades vary from pinnatifid to one-pinnate- son Woodward (1745–1820), a British phycologist.
Figure 13.124.1. Woodwardia radicans: habit.

Figure 13.124.2. Woodwardia radicans: veins, sori, and

indusia.

Figure 13.124.3. Woodwardia orientalis: leaf buds.

 F  F A  C 537

Woodwardia areolata (Linnaeus) T. Moore

 13.124.4
syn. Lorinseria areolata (Linnaeus) C. Presl
Swamp fern, narrow-leaved chain fern, net-veined chain
fern
Very hardy, Zone 3
A medium-sized fern with slender, long-creeping rhizomes a
and erect, deciduous fronds. Grows well under low to me-
dium light in wet, acidic garden soil or potting mix.
Woodwardia areolata differs from the other cultivated
chain ferns by its strongly dimorphic sterile and fertile
fronds. The sterile blades are lanceolate and deeply lobed to
a narrow wing along the rachis, and they lack buds. The
fertile blades are one-pinnate with highly contracted pinnae
and soon wither after the spores are shed. The species is na-
tive to southeastern Canada and the eastern United States.

Figure 13.124.5.
a Woodwardia fimbriata:
a. frond, bar = 10 cm
(4 in.); b. lowest pinna,
Figure 13.124.4. bar = 5 cm (2 in.).
Woodwardia areolata:
a. habit with fertile
frond (right), sterile
frond (left), bar =
10 cm (4 in.); b. b
fertile pinnae, bar =
1 cm (0.4 in.).

b Once established, the plants need a lot of room because the

fronds tend to spread horizontally.
The emerging fiddleheads of Woodwardia orientalis are
densely scaly. The blades are triangular to ovate, flat, glossy,
Woodwardia fimbriata J. E. Smith  13.124.5 and can have many buds developed on the upper surface.
syn. Woodwardia chamissoi Brackenridge These buds bear small, obovate fronds while still attached
Giant chain fern to the main fronds. This species is native to the Himalayas,
Semi-hardy to slightly hardier, Zone (7)8 Japan, and China through Taiwan to the Philippines.
A large fern with short-creeping rhizomes and evergreen
fronds. Grows well under medium light in moist garden Woodwardia radicans (Linnaeus) Smith
soil or potting mix.  13.124.1, 2, 7;  50
Woodwardia fimbriata has erect-spreading fronds with European chain fern
broadly oblanceolate blades, without buds, glabrous on the Semi-hardy, Zone 8
lower surfaces. This species is native to southern Arizona A large fern with short-creeping rhizomes and evergreen
and the Pacific Coast from Oregon to Baja California. fronds. Grows well under high or medium light in moist
garden soil. Established plants need a lot of space for their
Woodwardia orientalis Swartz  13.124.3, 6 arching or cascading fronds. They are easy to grow. The
Oriental chain fern bud on the underside of the frond tip roots easily by itself
Semi-hardy to hardier, Zone 8 when left on the plant, or it may be removed and rooted.
A large fern with short-creeping rhizomes and evergreen Woodwardia radicans has oblong-ovate blades that bear
fronds. Grows well under high or medium light in moist a scaly, large, well-developed bud below the tip. On the
garden soil or potting mix. The plants are easy to grow and basal pinnae, the basiscopic segment next to the rachis is
can be propagated by detaching the buds and planting their reduced in size. This species is native to southern Europe.
basal ends in a fine soil mix kept in humid conditions.
 538 C 13

b
Figure 13.124.6. Woodwardia orientalis: a. frond, bar =
10 cm (4 in.); b. lowest pinna, bar = 5 cm (2 in.); c. leaf
buds, bar = 2 cm (0.8 in.). Figure 13.124.7. Woodwardia radicans: a. frond (left)
and leaf bud (right), bar = 10 cm (4 in.); b. lowest
pinna, bar = 5 cm (2 in.).
Woodwardia ×semicordata Mickel & Beitel
 13.124.8
Semi-tender, Zone 9 more pairs of decurrent pinnae. Woodwardia martinezii
The general culture of this fern is similar to that for Wood- Maxon ex Weatherby, native to Mexico, is unknown in
wardia spinulosa except that it might be more cold toler- cultivation. It differs from W. ×semicordata by long-creep-
ant. According to Mickel and Beitel (1988), W. ×semicor- ing rhizomes and sori along the costules and entire length of
data resulted from a cross between W. martinezii and W. the costae and rachis. Woodwardia ×semicordata is native
spinulosa, but further research is needed to determine to Mexico.
whether it is actually of hybrid origin. At least some spores
appear to be normal and well formed, and the plants might Woodwardia spinulosa M. Martens & Galeotti
be apogamous.  13.124.9
In cultivation, Woodwardia ×semicordata has been con- Mexican chain fern
fused with both of its putative parents. It resembles W. spin- Semi-tender, Zone (9)10
ulosa by having sori distributed along the costules (the mid- A large fern with short-creeping rhizomes and evergreen
ribs of the pinnules) but differs by having sori also fronds. Grows well under high light in moist garden soil.
distributed in the distal half or less of the costae (the mid- The plants are easy to grow.
ribs of the pinnae). Beneath the pinnatifid apex, only one Woodwardia spinulosa can be distinguished from W.
pair of free pinnae with decurrent bases is present (all other fimbriata by the triangular blades with fimbriate scales on
pinnae are nondecurrent), whereas W. spinulosa has two or the lower surfaces. The lower pinnae have basal segments
 F  F A  C 539

that tend to overlap the rachis. It can be distinguished from

W. radicans by the absence of buds on the fronds. This spe-
cies is native to Mexico and Central America.

Woodwardia unigemmata (Makino) Nakai

 13.124.10
Semi hardy or hardier, Zone 8
c A large, semi-deciduous to deciduous fern with short-creep-
ing rhizomes. Grows in medium light in moist garden soil
or potting mix. The plants may be propagated from the
bud near the frond tip.
Woodwardia unigemmata is similar to W. radicans ex-
cept it lacks the lowest downward-pointing (basiscopic)
a
segment immediately next to the rachis. It is native to east-
ern Asia.

a
b

Figure 13.124.8. Woodwardia ×semicordata: a. frond, bar

= 10 cm (4 in.); b. lowest pinna, bar = 5 cm (2 in.);
c. sori along costules and costae, bar = 2.5 cm (1 in.).
Figure 13.124.10.
Woodwardia
unigemmata: a. frond
(left) and leaf bud
(right), bar = 10 cm
(4 in.); b. lowest
pinna, bar = 5 cm
(2 in.).
c
a

b
Woodwardia virginica (Linnaeus) Smith
 13.124.11
syn. Anchistea virginica (Linnaeus) C. Presl
Figure 13.124.9. Woodwardia spinulosa: a. small frond, Virginia chain fern
bar = 10 cm (4 in.); b. lowest pinnae with basal pinnules Very hardy, Zone 3
overlaying the rachis, bar = 5 cm (2 in.); c. sori along A large, deciduous fern with long-creeping rhizomes up to
costules only, bar = 2.5 cm (1 in.). 1 cm (0.5 in.) thick. Grows in medium to high light in wet,
acidic garden soil. This species is considered invasive in
some areas.
540 C 13

Figure 13.124.11.
Woodwardia virginica:
a. habit, bar = 10 cm (4 in.);
b. pinnule with sori and
a indusia, bar = 4 mm.

The stipes of Woodwardia virginica are blackish at the

base. The fronds are one-pinnate-pinnatifid with narrow,
linear pinnae deeply pinnatifid and jointed (articulate) to
the rachis. The sori are elongate, borne along the costae and
costules, and covered by membranous indusia. It is native
to southeastern Canada and the eastern United States.
  
Measuring Light

The human eye often has difficulty judging light intensity. For this reason, special light meters can be pur-
chased to measure light intensity in foot-candle units. Photographic light meters can also be used, and the
readings can be converted to foot-candle units. The formula is as follows:
foot-candles = 20f 2 / TS
where f is the aperture in f-stop, T the shutter speed in seconds, and S the film speed in ASA units.

To measure light intensity using a photographic light meter, place a large sheet of white paper on the
surface to be measured. Set an appropriate ASA film speed on the meter and read the shutter speed required
for proper exposure at a given f-stop. Read the meter at about 30 cm (12 in.) from the paper, and use the
preceding formula to calculate the equivalent foot-candle measurement. For example, if the ASA film speed
setting is set at 100 and the proper exposure is ½ second at an f-stop of 16, the foot-candle value is 102:
20 × (16)2 / ½ × 100 = 102 foot-candles
This formula provides a workable approximation. Results will vary according to the accuracy of the light
meter and the cone of light measured by the meter.
For scientific work, precise light intensities need to be measured. A quantum meter accurately measures
the blue and red part of the spectrum (the photosynthetically active radiation, or PAR). Plants use the blue
and red light but not yellow and green light. With quantum meters, the intensity is measured in units
known as micromoles, instead of foot-candles. A reading of 200 micromoles (µmol m−2 sec−1) is equivalent
to about 1000 foot-candles. Quantum meters are specifically designed for plants, but the average fern
grower will find photometers sufficiently accurate for monitoring the light needs of ferns.
Other light-measuring units that may be encountered, and their approximate equivalency to foot-can-
dles, are as follows:
1 lumen/cm2 = 1000 foot-candles
1 lux = 1⁄ 10 foot-candle
1 micromole = 5 foot-candles

541
  
Fern Societies

Updated information and addresses for various fern societies in the United States and abroad can be
obtained from the American Fern Society. Contact the secretary of the American Fern Society:
Dr. W. Carl Taylor
Botany Department
Milwaukee Public Museum
800 W. Wells Street
Milwaukee, Wisconsin 53233-1478
The information is also available from the American Fern Society’s World Wide Web page:
http://www.amerfernsoc.org
Membership information for the American Fern Society can be obtained from the membership secretary:
Dr. David B. Lellinger
Botany Department, MRC-166
Smithsonian Institution
Washington, D.C. 20560
A highly informative Web site about ferns is maintained by the San Diego Fern Society. It has links to nearly
all other fern Web sites and is an excellent way to search for current information about the fern society near-
est you.
http://www.inetworld.net/sdfern/society.htm
Another helpful Web site for growers is that maintained by the British Pteridological Society.
http://www.nhm.ac.uk/hosted_sites/bps/
An e-mail exchange for fern information is available and called Fernet. Inquire about the service at
koning@escu.ctstateu.edu

542
  
Importing Ferns

The following information will be helpful for importing plants to the United States and knowing what to
expect as your plants pass through customs. When plants are carried through as personal luggage and they
comply with all regulations, a United States Department of Agriculture (USDA) Plant Import Permit is not
required for one-time or infrequent importers (though see the regulations and restrictions applicable to
ferns). Frequent importers need a plant-import permit. This permit also allows you to send plants to the
United States by mail or separate carrier to an inspection station. If the plants pass the inspection, they are
forwarded to you.
You must pay attention to the regulations and directions that come with the import permit. These rules
are not given here, except in a general way applicable to ferns. Because regulations and procedures are sub-
ject to change, always check with the agencies involved for the latest information.

Obtaining Permits for Collecting and Importing

The United States Department of Agriculture’s Plant Import Permit is free. To obtain one, you should con-
tact the USDA well before you travel. The address is
USDA, APHIS, PPQ Unit 136
4700 River Road
Riverdale, Maryland 20737-1235
telephone: (301) 734-7885; (301) 734-5948
fax: (301) 734-3560
e-mail: cbare@aphis.usda.gov
Web site: www.aphis.usda.gov
Your permit will come with the latest information on how plants must be prepared for entry into the
United States and which plants are forbidden or have special restrictions. Ferns currently forbidden entry into
the United States because of their potential to be serious weeds include the mosquito fern (Azolla pinnata)
and water spangles (Salvinia auriculata, S. biloba, S. herzogii, and S. molesta). Some genera of tree ferns
require special documents because of their endangered status, and this also applies to herbarium specimens
(see the section “Convention on International Trade in Endangered Species of Wild Fauna and Flora”).
Import permits are not required for fern spores or dried herbarium specimens of non-CITES species.
Each country has its own laws about collecting and exporting plants. Before you leave home, check with
the embassy, consulate, or tourist information office of countries you plan to visit for any plant collecting

543
544 A III

or export requirements. Also check with the CITES office for updated information and the address of the
CITES office abroad. Once at your destination, the local customs or plant protection or regulating agency
should be able to inform you of their regulations.

Preparing Plants for Import into the United States

Plants may be brought into the United States by various kinds of mail, freight, or in your luggage. In all cases
they must be prepared and packaged as directed in the papers that arrive with your import permit (also see
the section “Packing Ferns,” in Chapter 3). At minimum, the import permit requires that plants be free of
soil, pests, disease, and seeds. The plants must be packed in approved packing material, meet the size-age
factor, be labeled, and have certification papers from the proper plant quarantine officials in the country of
origin. They must not be on the forbidden list.
The size-age factor ruling applies only to tree ferns, not herbaceous plants. Tree ferns need to be less
than 45 cm (18 in.) long, as measured from the soil line to the terminal growing point. Plants should be
labeled, preferably with their scientific name and where they were collected. Phytosanitary certificates are
issued by local agricultural or quarantine officials. Although importing fern spores into the United States
is not restricted, they must be free of soil, pests, disease, and seed contaminants. Fragments of leaves in the
spore packet do not ordinarily present a problem if they are clean. When spores or herbarium specimens are
carried in your luggage, the plant inspectors may ask to see them. Green spores should be airmailed to
their destination and sown as soon as possible because they are short-lived.

E  M  F

You can mail or freight plants to the United States only if you have an import permit and have met all
the requirements. When plants are mailed using the yellow and green address labels issued with the permit,
the package will go directly to United States customs and a plant inspection station. If all is in order, the
package is sent by mail under the original postage to the address on your permit. Air-freight arrivals valued
above a certain price require the service and fee of a customs broker. The broker’s function is to meet the
shipment, complete the formalities at customs (pay the custom fee on your behalf), and convey the ship-
ment to the plant inspector. If the shipment is valued at less than a certain amount, you may process the
shipment yourself. Ask your local customs bureau for information on procedures.
Using the mail or freight has both advantages and disadvantages as compared to carrying the plants in
your luggage. A plant sent by mail can be lost or subjected to slow service, rough handling, and/or extreme
temperatures. Sending the plants by mail or freight, however, avoids the possible delays that can result
from inspection of the plants upon arrival at the airport and, perhaps, also avoids unexpected broker’s fees.
If you do not live near a plant inspection station, or if your point-of-entry into the United States does not
have a plant inspector on duty, it may be more convenient and economical to mail the plants. If stopovers
in other countries are necessary, it is best to send the plants directly to the United States before you leave
the country where the plants originated. If you carry plants to subsequent countries, they may be barred
from entry. In New Zealand, facilities are available to hold a small number of plants at the plant inspection
station until you can retrieve them when you leave the country. There is a small charge for holding fern
plants (including spores) for more than than 48 hours. You also must enter and leave the country through
the same airport. It is difficult, however, to determine in advance if such services are available.
 I F 545

E  Y L

If you enter the United States with plants in your luggage, you will be asked to declare them on your
customs form. After passing customs, you will be directed to an inspector who will see that the plants meet
the legal requirements. It is helpful but not necessary to have your import permit in hand. Generally, if there
are 12 or fewer plants, if there are no special restrictions, and if all plants are properly identified and free of
pests, then the inspector can clear them on the customs floor. Plants protected by CITES, more than 12
plants, or identification problems will require that the plants be sent to a plant inspection station. Expect
problems even if you have followed all procedures correctly. Inspectors frequently cannot identify the
plant, or regulations may have changed. The inspector may assist you (but is not required to) in getting the
plants to the plant inspection station. You may have to hire a bonded carrier to bring the plants to an
inspection station. If the cargo is large, it may be delivered to the airline’s cargo station, and the airline may
assist in getting the ferns to the station. Before your plants leave for the plant inspection station, be certain
that the invoice papers issued by customs indicating customs’ clearance (Post-entry papers or Manifest) are
securely attached to the plants or their wrappings. The entire inspection process can take anywhere from a
few hours to a few days. Unless arrangements are made to forward them to you, the plants will have to be
retrieved at the inspection station, which is not necessarily located near the airport.
Delays and inconveniences are also possible if the inspection station is closed (the hours are generally
8:00 .. to 4:30 ..), or if the plants arrive just before the weekend. Not all entry points into the United
States have plant inspection stations, and this can complicate carrying the plants in your luggage. In such
cases, the plants are sent to the closest plant inspection station. If possible, plan your entry at a customs facil-
ity that has a plant inspection station. A list of the plant inspection stations (called Ports of Quarantine
Clearance) is usually sent with your import permit.

Convention on International Trade in Endangered Species of

Wild Fauna and Flora
The permits required by the Convention on International Trade in Endangered Species (CITES) are a sep-
arate entity from the United States Department of Agriculture’s Plant Import Permit. If the plants you col-
lect are listed by CITES, you must get CITES permits, which often requires a fee. Presently, the only ferns
on the CITES list are tree ferns in the Cyatheaceae and Dicksoniaceae. To import these restricted ferns you
must obtain export permits from the country of origin, even if the plants are from cultivation or are dried
specimens for herbarium sheets. Tree fern spores are exempt from CITES permits. Approval for the export
permit is dependant on the species, where the plant was collected, and whether it is cultivated. The United
States Fish and Wildlife Service handles all CITES inquiries and will send you the addresses of foreign
CITES offices that issue export permits. Their address is
United States Fish and Wildlife Service
Office of Management Authority, CITES Permits
4401 N. Fairfax Drive
Arlington, Virginia 22203
telephone: (703) 358-2104
You can also contact any office of the USDA Animal and Plant Health Inspection Service, Plant Pro-
tection and Quarantine (USDA, APHIS, PPQ). They are located nationwide.
CITES regulations are subject to change, and therefore it is a good idea to contact CITES officials
early. When entering the United States with a CITES plant, be sure to pack the plant and its export per-
546 A III

mit separately from non-CITES plants. The plant inspector will routinely send any CITES plants to the
inspection station before release. Before relinquishing the plant, be certain that the CITES permit and the
plant are securely together, preferably in their own bag. Also make sure that the customs invoice (Post-entry
papers or Manifest) indicates that the plant passed customs. In addition to the usual USDA plant import
requirements, tree-fern trunks may not be more than 45 cm (18 in.) long measured from the soil level to
the tip of the stem (apical bud). The latest CITES regulations indicate that all tree ferns on the CITES list,
with or without trunks, are subject to the CITES regulations (formerly, those without trunks were exempt).
Avoid traveling to a second country with a CITES plant even if you have an export permit issued from
the first country you visited. You will need to get a second export permit (re-export) from the second coun-
try, and this can take several days. It may be better to change your itinerary or mail the ferns along with their
CITES export permit from the original country directly to the United States. CITES regulations specify
that CITES export permits must come from the country of origin, and that may be interpreted to mean the
last country visited. That last country may refuse to issue an export permit.
The preceding procedures and restrictions are meant to protect the United States from introduced
pests and diseases, and to prevent the extinction of threatened or endangered species. The restrictions may
be inconvenient and complicated, but they are imposed for the well being of agriculture, the environment,
and ultimately all of us.
  
Names of Pest and Disease
Control Substances

The active ingredients of common pest and disease control products are given in the following charts. The
trade names appear in upper case. Some products are available to commercial applicators only. Formula-
tions vary, and no attempt has been made to account for these variations. The Federal and State Environ-
mental Protection Agencies (EPA) regulate the label content and use of pesticides, fungicides, bactericides,
disinfectants, and other substances. Their directions are the law. EPA regulations may change, and new
products continue to be produced, so check with county agricultural services or other reputable sources for
the latest information. Always test a product on one plant before wider application. The reader acts on his
or her own responsibility. No endorsement is intended in the use of trade names, nor is criticism implied of simi-
lar products not mentioned.

Pesticides
Alphabetical listing Corresponding names Applications

acephate  broad spectrum, systemic, some formulations

for indoor use; aphids, mealybugs, millipedes,
psocids, mites, scales, sowbugs, pillbugs, thrips,
whiteflies, others
Bacillus thuringiensis  ,  bacterial formulation for larvae of moths,
butterflies
Bacillus thuringiensis  bacterial formulation for fungus gnats only
H-14
 see propoxur
bendiocarb ,  ants and others
- 40 see nicotine alkaloid
  see Bacillus thuringiensis
carbaryl  broad spectrum, easily available to home growers,
some formulations for indoor use; ants, some
aphids, earwigs, fungus gnats, millipedes, mites,
sowbugs, pillbugs, psocids, others

547
548 A IV

Alphabetical listing Corresponding names Applications

chlorpyrifos  some formulations for indoor use, easily available

to home growers; earwigs, fungus gnats,
millipedes, mites, sowbugs, pillbugs, psocids,
thrips, root mealybugs, ground-crawling pests
 2 see dimethoate
diazinon , - broad spectrum, easily available to home growers;
aphids, earwigs, fungus gnats, mealybugs,
psocids, mites, scale crawlers, thrips, whiteflies,
others
dicofol  mites only
dienochlor  mites only
dimethoate  2 broad spectrum, systemic; aphids, fungus gnats,
scales, thrips, whiteflies, others; law limits use to
injection equipment only in California (1993)
 see Bacillus thuringiensis
disulfoton - systemic, easily available to home growers, some
formulations for indoor use; aphids, mites, scales,
thrips, others
- see disulfoton
 see chlorpyrifos
 see bendiocarb
 5 see kinoprene
ethoprop  nematodes only; use restricted to commercial
applicators
fluvalinate   synthetic type of pyrethrin; aphids, fungus gnats,
mites, thrips, whiteflies, others
 see Bacillus thuringiensis
H-14
imidacloprid  broad spectrum, systemic for greenhouse and
nursery use; aphids, mealybugs, scales, thrips,
whiteflies, others; not available to home growers
 see dicofol
kinoprene  5 insect growth regulator; aphids, fungus gnats,
mealybugs, scales, whiteflies
- see diazinon an encapsulated diazinon
 see imidacloprid
malathion (maldison)  broad spectrum, easily available to home growers;
aphids, fungus gnats, mealybugs, scales,
whiteflies, others
 see neem
  see fluvalinate
 see methiocarb
-  see oxydemetron-methyl
methyl methaldehyde  slugs, snails
methiocarb  slugs, snails, others
 N  P  D C S 549

Alphabetical listing Corresponding names Applications

mexacarbate  slugs, snails

 see ethoprop
neem (azadirachtin)  general insecticide with some fungicidal qualities;
insect growth regulator
 see phenamiphos
nicotine alkaloid   very toxic, restricted use; aphids, thrips, adult
 whiteflies, others; nicotine sulfate is no longer
available in the United States
  see nicotine alkaloid

 see acephate
oxamyl  broad spectrum, more or less systemic; fungus
gnat larvae, nematodes, others; no longer labeled
for ornamental plants
oxydemetron-methyl -  broad spectrum, systemic; aphids, mealybugs,
others
 see dienochlor
 synthetic type of pyrethrin (pyrethroid);
sowbugs, pillbugs, others
phenamiphos  nematodes; restricted to commercial use
propoxur  ground-crawling pests
pyrethrin  short residual effect, easily available to home
(pyrethreum) growers; ants, aphids, earwigs, thrips, whiteflies,
others
pyrethroid see , synthetic types of pyrethrin
, or
fluvalinate
resmethrin  synthetic type of pyrethrin (pyrethroid), some
formulations for indoor use; mainly fungus
gnats, whiteflies, also aphids
rotenone  broad spectrum, short residual effect; beetles,
true bugs (Hemiptera), mites, scales, thrips,
others
 see soap
 see carbaryl
 see methyl methaldehyde
soap  , soft-bodied insects, mealybugs, not thrips

 see bendiocarb
 see oxamyl
 see mexacarbate
550 A IV

Fungicides, Bactericides, and Disinfectants

Alphabetical listing Corresponding names Applications

- see streptomycin

 see fosetyl-Al
 see thiram
 see propiconazole
 see etridiazole and very broad spectrum, mix of two fungicides
thiophanate-methyl
 see benomyl
benomyl  no longer labeled for ornamental plant use; see
thiophanate-methyl
 see copper sulfate
captan  broad spectrum; some blights, damping-off, leaf
spots, rots, water molds
carbolic acid (phenol)  general disinfectant
 26019 see iprodione
chloramphenical algaecide, but also bactericide
chlorothalonil  2787 broad spectrum, easily available to home growers;
some blights and damping-off, gray molds, some
leaf spots, Rhizoctonia, rusts, some algae
’ 2226 see dimethyl
4,4’-O-phenylenebis
(3-thioallophanate)
 see sodium hypochlorite
 see quaternary
ammonium
compounds
copper sulfate  algaecide
copper sulfate  27 bactericide and fungicide, systemic, leaves no
pentahydrate residue; some leaf spots, Botrytis
 2787 see chlorothalonil
 see fenaminosulf
dimethyl ’ 2226 some blights, damping-off, leaf spots and rots,
4,4′-O-phenylenebis gray molds, Rhizoctonia
(3-thioallophanate)
 -22 see maneb
 -45 see mancozeb
 -78 see zineb
etridiazole (ethazol) , , blights, damping-off, and rots caused by water
 in part molds; best mixed in soil before planting
fenaminosulf ,  blights, damping-off, and rots caused by water
(diazoben) molds; use drench at once, light sensitive
folpet  similar to captan; rusts, some leaf spots
fosetyl-Al  some blights, damping-off, and rots, water
molds; burns ferns
 N  P  D C S 551

Alphabetical listing Corresponding names Applications

  see quaternary

ammonium
compounds
iprodione  26019 somewhat systemic; some blights, damping-off,
and leaf spots, gray mold, Rhizoctonia
 see fenaminosulf
 see carbolic acid
mancozeb  -45 some blights, damping-off, and leaf spots, gray
mold, rusts, some algae; leaves a residue
maneb  -22 some blights, damping-off, and leaf spots, gray
mold, rust
metalaxyl  systemic; blights, damping-off, and rots caused
by water molds; use as spray, drench, or granules
 see pentachloronitro-
benzene
pentachloronitro- ,  some blights, damping-off, and rots, gray molds,
benzene Rhizoctonia; may burn ferns when used as a soil
fungicide, can persist in soil and cause plant
toxicity
 see folpet
 20 see quaternary
ammonium
compounds
 27 see copper sulfate
pentahydrate
potassium general disinfectant
permanganate
propiconazole  general fungicide, some systemic action; leaf
spots, rusts
quaternary ,  20, general disinfectant
ammonium  
compounds
quintozene see pentachloronitro-
benzene
sodium hypochlorite  general disinfectant
(bleach)
streptomycin - systemic; leaf spots, rots, and diseases caused by
bacteria
 see metalaxyl
 see pentachloronitro-
benzene
 see etridiazole
thiophanate-methyl ,  in part systemic, acts like benomyl; some blights,
damping off, and leaf spots, Rhizoctonia; use as a
drench
552 A IV

Alphabetical listing Corresponding names Applications

thiram  Rhizoctonia

 see etridiazole
zineb  -78 some blights, damping-off, and leaf spots, gray
molds, rusts
 see mancozeb and very broad spectrum, mix of two fungicides
thiophanate-methyl
  
Family Classification of Fern
Genera Treated in the Text

Aspleniaceae (spleenwort family)

Asplenium Phyllitis
Camptosorus Schaffneria
Ceterach

Blechnaceae (blechnum family; chain fern, hacksaw fern, and relatives)

Blechnum Stenochlaena
Doodia Woodwardia
Sadleria

Cyatheaceae (cyathea family)

Alsophila Sphaeropteris
Cyathea

Davalliaceae (davallia family; bear’s-foot fern and relatives)

Davallia Leucostegia
Davallodes

Dennstaedtiaceae (cup fern family)

Dennstaedtia Microlepia
Histiopteris Odontosoria
Hypolepis Paesia
Lindsaea Pteridium
Lonchitis

Dicksoniaceae (dicksonia family)

Cibotium Dicksonia

553
554 A V

Dryopteridaceae (wood fern family)

Arachniodes Hypoderris
Athyrium Lastreopsis
Ctenitis Matteuccia
Cyclopeltis Megalastrum
Cyrtomium Olfersia
Cystopteris Onoclea
Deparia Polybotrya
Didymochlaena Polystichum
Diplazium Rumohra
Dryopteris Tectaria
Gymnocarpium Woodsia

Equisetaceae (horsetail or scouring-rush family)

Equisetum

Gleicheniaceae (gleichenia family)

Grammitidaceae (grammitid family)

Hymenophyllaceae (filmy fern family)

Hymenophyllum Trichomanes

Lomariopsidaceae (elephant-ear family)

Bolbitis Elaphoglossum

Lycopodiaceae (ground pine family)

Lycopodium

Marattiaceae (mule’s-foot fern family)

Angiopteris Marattia

Marsileaceae (water-clover family)

Marsilea Regnellidium
Pilularia

Nephrolepidiaceae
Nephrolepis

Oleandraceae (oleandra family)

Arthropteris Oleandra
 F C  F G T   T 555

Ophioglossaceae (adder’s-tongue and grape-fern family)

Botrychium Ophioglossum

Osmundaceae (cinnamon fern family)

Leptopteris Todea
Osmunda

Polypodiaceae (polypody family)

Aglaomorpha Microgramma
Belvisia Microsorum
Campyloneurum Niphidium
Dictymia Pecluma
Drynaria Phlebodium
Goniophlebium Platycerium
Lecanopteris Polypodium
Lemmaphyllum Pyrrosia
Lepisorus Selliguea
Leptochilus Solanopteris

Psilotaceae (whisk fern family)

Psilotum

Pteridaceae (pteris family)

Acrostichum Cryptogramma
Actiniopteris Doryopteris
Adiantopsis Hemionitis
Adiantum Llavea
Anogramma Mildella
Argyrochosma Notholaena
Aspidotis Onychium
Astrolepis Pellaea
Bommeria Pentagramma
Ceratopteris Pityrogramma
Cheilanthes Pteris
Coniogramme Trachypteris

Salviniaceae (floating-fern family)

Azolla Salvinia

Schizaeaceae (climbing fern family)

Anemia Lygodium
556 A V

Selaginellaceae (spike moss family)

Selaginella

Thelypteridaceae (marsh fern, maiden fern family)

Macrothelypteris Thelypteris
Phegopteris

Vittariaceae (shoestring fern family)

Vittaria

Evolutionary Tree for Vascular Plants

Seed Plants

Ferns (incl. Psilotum)

Horsetails

Isoetaceae

Selaginellaceae Lycopods

Lycopodiaceae

An evolutionary tree (cladogram) showing the current ideas about the relationships of major groups of vascular plants.
The lycopod line of evolution has been distinct from other plants since the early Devonian, about 395 million years
ago. It is based on evidence from morphology, anatomy, and DNA sequences of living and fossil plants.
 F C  F G T   T 557

Grammitidaceae
Polypodiaceae
Polypodiaceae (Loxogramme)
Davalliaceae
Nephrolepidaceae
Lomariopsidaceae
Dryopteridaceae
Blechnaceae
Paleocene or more recent Thelypteridaceae
Aspleniaceae
Eocene
Pteridaceae
Pteridaceae (Adiantum)
Paleocene Vittariaceae
Dennstaedtiaceae
Dennstaedtiaceae (Lonchitis)
Lindsaeaceae
Triassic
Dicksoniaceae
Early Cretaceous Metaxyaceae
Cyatheaceae
Plagiogyriaceae
Late Cretaceous
Salviniaceae (Azolla)
Late Cretaceous Salviniaceae (Salvinia)
Middle Jurassic Marsileaceae
Permian Schizaeaceae
Gleicheniaceae
Permian Hymenophyllaceae
Osmundaceae
Ophioglossaceae
Marattiaceae
Late Carboniferous
Cycadaceae

An evolutionary tree (cladogram) showing the current ideas about the relationships of major families of ferns. The tree
is based on a combined analysis of data from morphology and DNA and is modified from Pryer et al. (1995). The
geologic periods given for some of the branches of the tree are times of earliest origin based on fossil evidence
(Collinson 1996).
 Glossary

acroscopic: the side facing the apex. The acroscopic margin of a pinna is that which faces the apex of the leaf.
The acroscopic margin of a pinnule is that which faces the apex of the pinna. Compare to basiscopic.
acrostichoid: with the sporangia spread throughout the surface, not in discrete lines or dots.
acuminate: tapering to a long point with the margins pinched or concave just before the tip.
acute: short-tapering to a point, the margins not pinched or concave just before the tip.
adnate: grown to or united with an unlike part.
aerophore: the aerating tissue of the leaf, usually in the form of a light-colored line running lengthwise along
the leaf stalk and sometimes down into the stem. In genera such as Blechnum and Thelypteris, aerophores
may occur as short white or yellowish pegs at the bases of the pinnae where they join the rachis. Stomata
are abundant in aerophores, and thus these structures allow air to diffuse into the leaf. Aerophores are
characteristic of fern leaves, being absent from leaves of other plants.
anadromic (anadromous): with the first basal branch or vein (as on a pinna) arising from the side toward
the frond tip. Compare to catadromic.
anastomosed: said of veins that unite to form areoles; net-veined.
annulus: a complete or partial ring or cluster of thick-walled cells on the spore case functioning to open the
spore case.
antheridium: the male sex organ containing the sperm. Borne on the underside of the prothallus.
apogamy: a form of asexual reproduction in which new sporophytes are produced directly from the pro-
thallus tissue instead of from the fertilized egg cell (zygote).
apospory: a form of asexual reproduction in which prothalli are produced directly from young sporophyte
tissue instead of from spores.
appressed: pressed close to the surface, not spreading.
archegonium: the female sex organ containing the egg.
areolate: having netted veins.
areole: the area enclosed by netted veins.
aristate: having stiff, bristle-like appendages.
articulate: having nodes or joints that naturally separate at maturity; jointed.
attenuate: gradually tapering to a very narrow, slender point.
axes: a collective term referring to the petiole, rachis, costae, costules, and midrib of a leaf.
basiscopic: the side facing the base. The basiscopic margin of a pinna is that which faces the base of the leaf.
The basiscopic margin of a pinnule is that which faces the base of the pinna. Compare to acroscopic.
bicolorous: having two colors, usually referring to scales in which the central part is darker than the mar-
gins. Compare to concolorous.

558
 anadromous veins
VEINS

FROND

catadromous veins
distal portion
of blade
PINNAE

medial
portion of
blade

anadromous pinna
acroscopic side
of pinna

basiscopic side
proximal of pinna catadromous
portion of pinna
blade

PINNULE

proximal portion of pinnule

acroscopic side
of pinnule

basiscopic side of pinnule distal portion

of pinnule

Location of plant parts. Courtesy of Carl Taylor, Paul Nelson, and the Milwaukee Public Museum.
560 G

Leaf shapes (left to right): linear, lanceolate, ovate, obovate, cuneate, triangular, pentagonal.

binomial: the species name, which is the two words consisting of the genus name and the specific epithet.
bipinnate: twice pinnate (two-pinnate). Both primary and secondary divisions of the frond are pinnate.
bipinnatifid: twice pinnatifid. The divisions of a pinnatifid frond are again pinnatifid.
blade: the thin, broad part of a leaf or frond. Also called the lamina.
bud: in ferns the term usually applies to a proliferous bud or lump of tissue that grows into a new fern plant.
bulbil or bulblet: a small bulb-like body borne upon a stem or leaf and serving to vegetatively reproduce the
plant.
bullate: puffy or inflated, humped. Usually refers to the base of a scale, especially in tree ferns.
catadromic (catadromous): with the first basal branch or vein (as on a pinna) arising from the side toward
the frond base. Compare to anadromic.
caudate: having a slender, tail-like appendage.
ciliate: having short hairs (cilia) along the margin.
clathrate: refers to scales composed of cells that have dark side cell walls and broad, clear central portions
(the “body” of the cell); thus the scale resembles leaded glass. A hand lens or microscope is needed to
see this characteristic.
compound: said of a leaf having two or more leaflets.
concolorous: having one uniform color throughout, usually referring to scales. Compare to bicolorous.
cone: a tight cluster of highly modified, spore-bearing leaves borne at the branch tips.
confluent: running together.
cordate: heart-shaped.
costa (plural, costae): the midrib of a simple frond, or of the pinna (secondary rachis) or pinnules of a
compound frond.
costule: the midrib of a pinnule.
crenate: having low, rounded teeth.
crested: having forked tips, usually many. Usually refers to the frond, pinnae, or segments.
crown: the tip of the stem where the leaves arise. Usually this term is applied to thick, upright stems.
crozier: a young coiled fern frond; the fiddlehead.
cultivar: a cultivated variety of plant, originating in cultivation or selected from among wild plants, gener-
ally chosen for a desirable trait(s).
decurrent: having the base of a blade or blade division extending down the petiole, rachis, or costa as a
winged expansion or ridge. A decurrent base is winged downward, at least for a short distance.
deltate: triangular.
dentate: toothed, usually with broad teeth directed outward.
denticulate: minutely dentate.
dichotomous: forked regularly into pairs.
 G 561

dimorphic: having two forms. In ferns the term usually refers to differences in size or shape of the sterile and
fertile leaves (or segments). Compare to monomorphic.
distal: the part farthest away from the main body or leaf axis. Compare to proximal.
dorsal: relating to the back or lower side of a leaf.
emarginate: having a shallow notch in the margin.
entire: having an unmodified margin (not divided, lobed, or toothed).
epilithic: growing on rocks; also epipetric, saxicolous, or rupestral.
epiphyte (epiphytic): a plant that grows upon another plant. An epiphyte uses its host plant only for sup-
port; it is not parasitic.
erose: appearing as if gnawed. Usually refers to an irregular margin.
excurrent vein: a vein running toward the margin, not the midrib.
falcate: scythe-shaped, curved and flat, tapering gradually.
false indusium: an indusium formed by the enrolled margin of the leaf. Characteristic of many genera in
the Pteridaceae, such as Adiantum, Cheilanthes, and Pellaea.
false peltate: appearing peltate but with a narrow, inconspicuous sinus running to the centrally attached
stalk; also referred to as pseudopeltate. See peltate.
family: a group of related genera. Plant family names end with the suffix -aceae.
farina: a meal-like powder, usually white or yellow and found on the lower surfaces of fronds. Character-
istic of such fern genera as Argyrochosma, Pentagramma, and Pityrogramma.
farinose: covered with a meal-like powder (farina).
fern allies: vascular plants that reproduce by spores and have a life cycle similar to that of true ferns, but dif-
fer by how they bear their sporangia and by having small leaves and usually simple, unbranched veins.
Most of the fern allies are not closely related to the true ferns. The living families of fern allies are the
Equisetaceae, Isoëtaceae, Lycopodiaceae, Psilotaceae, and Selaginellaceae.
fertile: in ferns usually referring to leaves that bear sori.
fiddlehead: a fern leaf, young or in bud, that is coiled in a spiral pattern. Also see crozier.
fimbriate: having a fringed margin, the fringe often consisting of numerous hairs or narrow segments.
flora: a list of all the species growing in a region, or a collective term for all the species growing in a region.
It also refers to books that identify the plants within a certain geographical area.
foot-candle: a unit of measuring light intensity. One foot-candle is equal to the amount of light cast by a
standard candle one foot away from the flame.
friable: describing soil that is moist and loose.
frond: the leaf of a fern, typically consisting of the petiole and blade.
fused: grown to or united with a similar part.
gametophyte: a small, usually flat plant bearing the sex organs (archegonia and antheridia) that in turn pro-
duce the gametes. Each cell in the body of the gametophyte has one set of chromosomes (1n). Game-
tophytes grow from spores.
gene: a unit on a chromosome that determines the inheritance of a particular trait.
genus: a group of related species.
glabrous: lacking hairs or scales, smooth.
glaucous: having a whitish, waxy bloom.
hardened plants: plants that have adjusted physiologically to harsher growing conditions.
hastate: arrowhead-shaped, but with the basal lobes pointing outward at wide angles.
heterosporous: having two kinds of spores, male and female, the males being smaller than the females and
produced in separate sporangia.
homosporous: having one kind of spore.
hybrid: an offspring of two different species.
562 G

hydathode: an enlarged vein tip on the upper surface of a blade. It often secretes water and minerals. In some
ferns, such as Nephrolepis, the minerals may accumulate as a white deposit over the hydathode.
imbricate: overlapping.
immersed: sunken into the surrounding tissue.
incised: cut sharply and irregularly, more or less deeply.
included veinlet: a vein within an areole or mesh formed by other veins.
indusium: the structure covering the sorus. Also see false indusium.
internode: the portion of a stem between two successive nodes.
jointed: able to separate naturally at a certain point, leaving a scar; articulate.
laciniate: slashed or cut into narrow-pointed lobes.
lanceolate: lance-shaped.
leaflet: one of the divisions in a compound leaf.
linear: long and narrow, usually 10 times longer than wide, the sides parallel or nearly so.
lunate: shaped like a crescent or half-moon.
marcescent: withering (senescing) without falling off; in contrast to jointed or articulate, which indicate
withering and falling off at a joint.
marginal: at or relating to the margin or edge of a leaf.
meiosis: the type of cell division that gives rise to spores. During meiosis, the cell replicates its chromosomes
once and divides twice; the result is four cells with only half the chromosome number of the original
cell.
mesh: refers to netted veins.
monomorphic: having only one form. In ferns the term usually refers to sterile and fertile leaves that are not
differentiated. Compare to dimorphic.
mucronate: tipped with a short, abrupt point or mucro.
mutation: a sudden heritable change appearing in animals or plants due to changes in the genes or chro-
mosomes.
node: the point on a stem where a leaf emerges.
nonclathrate: refers to scales in which the cells lack dark, easily visible cell walls outlining the body of the
cell. Compare to clathrate.
oblanceolate: lanceolate with the broadest part near the tip.
obovate: ovate or egg-shaped with the broader end apical.
obtuse: blunt or rounded at the tip.
ovate: egg-shaped in outline.
palmate: having main veins, segments, or lobes radiating from a common point.
paraphysis (plural, paraphyses): a sterile hair that is mixed among the sporangia.
pedate: palmate with the side lobes cleft into two or more segments.
peltate: having the stalk attached centrally rather than at the edge (like the handle on an open umbrella).
See false peltate.
petiole: a leaf stalk; also known as a stipe.
phyllopodium (plural, phyllopodia): a stump-like extension from the rhizome to which the fronds are
attached, usually by a distinct abscission layer.
pinna (plural, pinnae): the primary division of a pinnately divided frond; a leaflet.
pinnate: having a feather-like arrangement, with a single mid-vein from which leaflets arise.
pinnate-pinnatifid: referring to a blade that is once-pinnate and with the pinnae deeply lobed or cut, but
not to their midrib.
pinnatifid: cut half to three-fourths to the rachis.
pinnatisect: cut almost all the way to the rachis.
 G 563

pinnule: a secondary pinna.

prothallus (plural, prothalli): the gametophyte (which see).
proximal: the part nearest or toward the base or point of attachment. Compare to distal.
pteridologist: a student of pteridophytes.
pteridophytes: the ferns and fern allies. These plants have a life cycle in which the sporophyte and game-
tophyte generations grow independently of one another (that is, they are not attached). The sporophyte
is the large and conspicuous generation, the gametophyte is inconspicuous and ephemeral. (See Chap-
ter 8.)
puberulent: minutely hairy.
pubescent: hairy.
pups: a colloquial term referring to the young plants arising from older plants of Platycerium usually by root
proliferation.
rachis: the midrib of a compound frond.
receptacle: the tissue upon which the sporangia are borne. The receptacle is bristle-like in Hymenophyllum
and Trichomanes; in most other ferns it is flush with the leaf surface or slightly elevated.
reflexed: abruptly bent downward or backward.
reniform: kidney-shaped.
revolute: rolled backward from the margins or apex. In ferns the term usually refers to a leaf margin rolled
back to protect the sori.
rhizome: a stem that grows horizontally and is anchored to the soil by its roots.
rootstock: a short, erect stem.
rosette: a cluster of leaves arranged circularly.
rupestral: growing on rocks; epilithic.
saxicolous: growing on rocks; epilithic.
scale: an outgrowth of the epidermal layer that appears as a small, flat, usually dry structure two or more cells
wide. Hairs (trichomes) are similar but only one cell wide.
segment: the ultimate division into which a blade is divided.
serrate: having sharp, forward-pointing teeth, like the blade of a saw.
sessile: without a stalk.
seta (plural, setae): a sharp, stiff, bristle-like hair. In ferns these tend to be brown or black.
simple: not divided; not compound. Usually refers to blades that are not divided to the midrib, but can also
refer to unbranched veins.
sinus: a space or recess between two lobes of a frond or other expanded structure.
sorus (plural, sori): a cluster of spore cases (sporangia).
species: a category of individuals that usually interbreed freely and have many characteristics in common.
The word is used for both the singular and plural.
spinulose: having small spines or spine-like processes along the margin.
sporangium (plural, sporangia): the specialized structure within which the spores are produced; the spore
case.
spore: a reproductive cell produced in a sporangium, germinating and developing into a prothallus.
sporeling: a young fern plant (sporophyte) that arises from the prothallus.
sporocarp: a round structure that contains sporangia within. Characteristic of two fern families: the Salvini-
aceae and Marsileaceae. Sporocarps are globose and delicate in the Salviniaceae but bean-like and hard
in the Marsileaceae.
sporophyte: the familiar plant that bears roots, stems, and leaves (as opposed to a gametophyte or prothal-
lus). So-called because it is the phase of the life cycle that produces spores. Each cell in the body of a
sporophyte has two sets of chromosomes (that is, is 2n).
564 G

stellate: star-like; having arms or hairs that radiate from a central point. Usually refers to hairs or scales.
sterile: refers to leaves that do not produce sori and to hybrids in which spores are aborted.
stipe: the leaf stalk or petiole.
stipule: a basal appendage of a stipe or petiole, usually two. In ferns the term is applied to the Marattiaceae
and sometimes to the flared leaf bases in Osmunda.
stolon: a long, slender stem capable of producing a new plant at its tip or along its length.
strigose: having curved, sharp, forward-pointing hairs.
synonym: an alternative scientific name, but not the one currently accepted by taxonomists.
terrestrial: growing in the ground, not on trees.
tomentum: a dense covering of woolly hair.
ultimate segment: the final division of a frond.
undulate: having a wavy surface or margin.
vascular: pertaining to specialized tissue (xylem and phloem) that conducts water, mineral nutrients, and
sugars.
vascular bundle: a bundle or strand of vascular tissue.
veins: the strands of vascular tissue in a leaf. “Free veins” are those unconnected to other veins; “netted
veins” are those so connected.
wing: a thin expansion or flat extension of an organ or structure.
xerophyte: a plant adapted to dry habitats.
zygote: a fertilized egg cell. The first cell in the development of a sporophyte.
 Literature Cited

ABRS/CSIRO Australia. 1998. Flora of Australia. Vol. 48, Ferns, Gymnosperms and Allied Groups.
Melbourne.
Andrews, S. B. 1990. Ferns of Queensland: A handbook to the ferns and fern allies. Queensland Depart-
ment of Primary Industries Information series Z189008. Queensland Herbarium, Brisbane.
Anonymous. 1986. Guidelines for the control of plant diseases and nematodes. Agricultural Handbook No.
656. United States Department of Agriculture, Washington, D.C.
Basile, D. A. 1973. A simple method of initiating axenic cultures of pteridophytes from spores. American
Fern Journal 63: 147–151.
Beddome, R. H. 1864. Ferns of Southern India, Being Descriptions and Plates of the Ferns of the Madras
Presidency. Madras.
Beitel, J. M. 1979. The clubmosses Lycopodium sitchense and L. sabinaefolium in the upper Great Lakes area.
Michigan Botanist 18: 3–13.
Beitel, J. M., W. H. Wagner Jr., and K. S. Walter. 1981. Unusual frond development in sensitive fern Ono-
clea sensibilis L. American Midland Naturalist 105: 396–400.
Best, R. 1980? Growing Ferns. Bay Books Australian Gardening Library, Bay Books, Sydney.
Boodley, J. W. 1972. Soilless mixes. Horticulture 50: 38–39.
Bosman, M. T. M. 1991. A monograph of the fern genus Microsorum (Polypodiaceae). Leiden Botanical
Series 14: i–ix, 1–161.
Brown, E. D. W. 1920. The value of nutrient solutions as culture media for fern prothallia. Torreya 20:
76–83.
Brownsey, P. J., and J. C. Smith-Dodsworth. 1989. New Zealand Ferns and Allied Plants. David Bateman,
Aukland.
Burrows, J. E. 1990. Southern African Ferns and Fern Allies. Frandsen Publishers, Sandton, Republic of
South Africa.
Ching, R. C., and Hsen Hsu Hu. 1930. Icones Filicum Sinicarum. Bishen Singh Mahendra Pal, Singh,
India.
Cody, W. J., and D. M. Britton. 1989. Ferns and Fern Allies of Canada. Canadian Government Publish-
ing Center, Ottawa.
Collinson, M. E. 1996. “What use are fossil ferns?”—20 years on: With a review of the fossil history of
extant pteridophyte families and genera. Pages 349–394. In: Pteridology in Perspective, edited by J. M.
Camus, M. Gibby, and R. J. Johns. Royal Botanic Gardens, Kew.
Crane, E. H. 1997. A revised circumscription of the genera of the fern family Vittariaceae. Systematic
Botany 32: 509–517.

565
566 L C

Curtwright, D. 1995. Ceratopteris thalactroides and its relatives. Journal of the Los Angeles International
Fern Society, Inc. 22: 28–33.
Dittmer, H. J., E. F. Castetter, and O. M. Clark. 1954. The ferns and fern allies of New Mexico. Univer-
sity of New Mexico Publications in Biology 6: 1–139.
Druery, C. T. 1912. British Ferns and Their Varieties. Routledge, London.
Dyce, J. W. 1963. Variation in Polystichum in the British Isles. British Fern Gazette 9: 97–109.
———. 1972. British fern varieties: The Scolopendriums. Newsletter No. 10 (November). British Pteri-
dological Society, London.
———. 1985. New fern varieties—wild and cultivated. Pteridologist 1: 78–80.
———. 1987. Classification of fern variations in Britain. Pteridologist 1: 154–155.
———. 1991. The variation and propagation of British ferns. Special Publication No. 3. British Pterido-
logical Society, London.
Dyer, A. F. 1979. The culture of fern gametophytes for experimental investigation. In: The Experimental
Biology of Ferns, edited by A. F. Dyer. Academic Press, London.
Edie, H. H. 1978. Ferns of Hong Kong. Hong Kong University Press, Hong Kong.
Evans, A. M. 1969. Interspecific relationships in the Polypodium pectinatum–plumula complex. Annals of
the Missouri Botanical Garden 55: 193–293.
Farrar, D. R. 1968. A culture chamber for tropical rain forest plants. American Fern Journal 58: 97–102.
Flora of North America Editorial Committee. 1993. Flora of North America North of Mexico. Vol. 2,
Pteridophytes and Gymnosperms. Oxford University Press, Oxford.
Ford, M. V., and M. F. Fay. 1990. Growth of ferns from spores in axenic culture. Methods in Molecular
Biology 6: 171–180.
Foster, F. G. 1984. Ferns to Know and Grow. 3rd ed. Timber Press, Portland, Oregon.
Franks, W. 1969. Platycerium-Fern Facts. Published privately, Los Angeles, California.
Fraser-Jenkins, C. R. 1986. A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bul-
letin of the British Museum (Natural History), Botany Series 14: 183–218.
Goudey, C. J. 1985. Maidenhair Ferns in Cultivation. Lothian Publishing Co., Melbourne, Australia.
Grounds, R. 1974. Ferns. Pelham Books, London.
Hennipman, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Botanical Series
2: i–xiii, 1–331.
Hennipman, E., and M. C. Roos. 1982. A monograph of the fern genus Platycerium (Polypodiaceae). Ver-
handelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde,
Tweede Reeks 80: 1–126.
Hickey, R. J. 1977. The Lycopodium obscurum complex in North America. American Fern Journal 67:
45–49.
Hickman, J. C., ed. 1993. The Jepson Manual: Higher Plants of California. University of California Press,
Berkeley.
Hill, R. H. 1976. Cold requirements of several ferns in southeastern Michigan. American Fern Journal 66:
83–88.
Hill, R. S. 1984. Propagation of Marattiaceae. Australian Fern Journal 74: 11–15.
Hoagland, D. R., and D. I. Arnon. 1950. The water-culture method for growing plants without soil. Cali-
fornia Agricultural Experiment Station, Circular 347. College of Agriculture, University of California,
Berkeley.
Hoshizaki, B. J. 1970a. The genus Adiantum in cultivation (Polypodiaceae). Baileya 17: 97–196.
———. 1970b. Rhizome scales of Platycerium. American Fern Journal 60: 144–160.
———. 1972. Morphology phylogeny of Platycerium species. Biotropica 4: 93–117.
———. 1975. The Fern Growers Manual. Alfred A. Knopf, New York.
 L C 567

———. 1975. A staghorn fern (Platycerium) hybrid. American Fern Journal 65: 99–101.
———. 1981. The fern genus Davallia in cultivation (Davalliaceae). Baileya 21: 1–50.
———. 1982. The genus Polypodium in cultivation (Polypodiaceae). Baileya 22: 1–98.
———. 1991. An “intergeneric” hybrid: Aglaomorpha × Drynaria. American Fern Journal 81: 37–43.
Hoshizaki, B. J., and M. G. Price. 1990. Platycerium update. American Fern Journal 80: 58–69.
Hoshizaki, B. J., and K. A. Wilson. 1999. The cultivated species of the fern genus Dryopteris in the United
States. American Fern Journal 89: 1–100.
Hovenkamp, P. H. 1986. A monograph of the fern genus Pyrrosia. Leiden Botanical Series 9: i–xiii, 1–
310.
Hovenkamp, P. H., and N. A. P. Franken. 1993. An account of the fern genus Belvisia Mirbel (Polypodi-
aceae). Blumea 37: 511–527.
Hu, I. H., and R. C. Ching. 1930. Icones Filicum Sinicarum. Fascicle 1.
Huang, Tsen-Chieng, ed. 1994. Flora of Taiwan. 2nd ed. Vol. 1, Pteridophyta and Gymnospermae. Edi-
torial Committee of the Flora of Taiwan, Ta’an Taipei, Taiwan.
Iwatsuki, K., T. Yamazaki, D. Boufford, and H. Ohba. 1995. Pteridophyta and Gymnospermae. Vol. 1 of
Flora of Japan. Kodansha, Tokyo, Japan.
Jacobsen, W. B. G. 1983. The Ferns and Fern Allies of Southern Africa. Butterworth Publishers, Durban,
South Africa.
Joe, B. 1958. Pteris species cultivated in California. Lasca Leaves 8: 26–29.
———. 1964. A review of the species of Platycerium (Polypodiaceae). Baileya 12: 69–126.
Johns, R. J. 1995. Lecanopteris lomarioides, Polypodiaceae. Curtis’s Botanical Magazine 12: 89–95.
Johnson, D. M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Systematic Botany
Monographs 11: 1–87.
Jones, D. L. 1987. Encyclopaedia of Ferns. Timber Press, Portland, Oregon.
Jones, D. L., and S. C. Clemesha. 1981. Australian Ferns and Fern Allies. A. H. and A. W. Reed, Sydney,
Australia.
Kato, M. 1984. A taxonomic study of the athyrioid fern genus Deparia with main reference to the Pacific
species. Journal of the Faculty of Science, University of Tokyo, section 3, 13: 375–429.
———. 1985. A systematic study of the genera of the fern family Davalliaceae. Journal of the Faculty of
Science, University of Tokyo, section 3, 13: 553–573.
Kaye, R. 1965. Variations in Athyrium in the British Isles. British Fern Gazette 9: 197–204.
———. 1968. Hardy Ferns. Faber and Faber, London.
Khullar, S. P. 1994. An Illustrated Fern Flora of West Himalaya. International Book Distributor, Dehra
Dun, India.
Knobloch, I. W., and D. S. Correll. 1962. Ferns and Fern Allies of Chihuahua, Mexico. Texas Research
Foundation, Renner, Texas.
Knudson, L. 1946. A new nutrient solution for the germination of orchid seed. American Orchid Society
Bulletin 15: 214–217.
Kramer, K. U., and P. S. Green, eds. 1990. Pteridophytes and Gymnosperms. Vol. 1 of The Families and
Genera of Vascular Plants, edited by K. Kubitzki. Springer-Verlag, Berlin.
Krusberg, L. R. 1992. Best nematode control. Flower and Garden 36 (2): 8.
Kurata, S. 1963. On Japanese ferns belonging to the genus Cyrtomium. Science Report of the Yokosuka
City Museum 8: 23–47.
Kurata, S., and T. Nakaike. 1964–1997. Illustrations of Pteridophytes of Japan. Vols. 1–8. University of
Tokyo Press, Tokyo.
Langsdorff, G. H., and F. Fischer. 1810. Plantes recueillies pendant le voyage des Russes autour du monde.
Published privately by the authors, Tübingen.
568 L C

Launert, E. 1968. A monographic survey of the genus Marsilea Linnaeus. I: The species of Africa and
Madagascar. Senckenbergiana Biologica 49: 273–315.
Lee, H. 1887. The vegetable lamb of Tartary, a curious fable of the cotton plant. Sampson, Low, Marston,
Searle, and Rivington, London.
Lellinger, D. B. 1972. A revision of the fern genus Niphidium. American Fern Journal 62: 101–120.
———. 1985. A Field Manual of the Ferns and Fern Allies of the United States and Canada. Smithson-
ian Institution Press, Washington, D.C.
———. 1988. Some new species of Campyloneurum and a provisional key to the genus. American Fern
Journal 78: 14–35.
———. 1989. The ferns and fern allies of Costa Rica, Panama, and the Chocó. Part 1: Psilotaceae through
Dicksoniaceae. Pteridologia 2A: 5–364.
———. 1994. Useful fern books in print: 1993–1994. Fiddlehead Forum 21: 19–21.
Little, E. L., Jr., and F. H. Wadsworth. 1964. Common trees of Puerto Rico and the Virgin Islands. Agri-
cultural Handbook No. 249. United States Department of Agriculture, Forest Service, Washington,
D.C.
Lloyd, R. M. 1974. Systematics of the genus Ceratopteris Brongn. (Parkeriaceae). II: Taxonomy. Brittonia
26: 139–160.
Lloyd, R. M., and E. J. Klekowski Jr. 1970. Spore germination and viability in Pteridophyta: Evolution-
ary significance of chlorophyllous spores. Biotropica 2: 129–137.
Lovis, J. D. 1968. Fern hybridists and fern hybridising. II: Fern hybridising at the University of Leeds. Brit-
ish Fern Gazette 10: 13–20.
Lovis, J. D., and G. Vida. 1969. The resynthesis and cytogenetic investigation of ×Asplenophyllitis microdon
and ×A. jacksonii. British Fern Gazette 10: 53–67.
Lowe, E. J. 1890. British Ferns and Where Found. Swan Sonnenschein, London.
Lumpkin, T. A., and D. L. Plucknett. 1980. Azolla: Botany, physiology, and use as a green manure. Eco-
nomic Botany 34: 111–153.
MacPhail, J. 1990. Trough gardening. Pacific Horticulture (spring): 31–39.
Marticorena, C., and R. Rodríguez. 1995. Pteridophyta–Gymnospermae. Vol. 1 of Flora de Chile. Uni-
versidad de Concepción, Concepción, Chile.
Matkin, O. A., and P. A. Chandler. 1957. The U.C.-type soil mixes. In: U.C. System for Producing
Healthy Container-Grown Plants, edited by K. F. Baker. Extension Service Manual No. 23, Califor-
nia Agricultural Experiment Station.
Maxon, W. R. 1912. Studies of tropical American ferns. Contributions to the United States National
Herbarium 16: 54–58.
Mickel, J. T. 1979. How to Know the Ferns and Fern Allies. W. C. Brown, Dubuque, Iowa.
———. 1992. Pteridophytes. In: Gymnosperms and Pteridophytes, Vol. 17 of Flora Novo-Galiciana,
edited by R. McVaugh. University of Michigan, Ann Arbor.
———. 1994. Ferns for American Gardens. MacMillan Publishing Company, New York.
Mickel, J. T., and J. M. Beitel. 1988. Pteridophyte flora of Oaxaca, Mexico. Memoirs of the New York
Botanical Garden 46: 1–568.
Moore, A. W. 1969. Azolla: Biology and agronomic significance. The Botanical Review 35: 17–35.
Moran, R. C. 1982. The Asplenium trichomanes complex in the United States and adjacent Canada. Amer-
ican Fern Journal 72: 5–11.
———. 1986. The neotropical fern genus Olfersia. American Fern Journal 76: 161–178.
———. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Natural
History Survey Bulletin 34 (1): i–v, 1–138.
———. 1992a. The vegetable lamb of Tartary—A pteridological tale. Fiddlehead Forum 19: 2, 7, 8.
 L C 569

———. 1992b. The potato fern. Fiddlehead Forum 19: 18–20.

———. 1992c. The story of the molesting Salvinia. Fiddlehead Forum 19: 26–28.
———. 1993. Bracken, the poisoner. Fiddlehead Forum 20: 18–19, 22.
———. 1995a. Iridescent ferns and their shady behavior. Fiddlehead Forum 22: 2–4, 8.
———. 1995b. Nardoo. Fiddlehead Forum 22: 20–22.
———. 1997. The little nitrogen factories. Fiddlehead Forum 24: 9, 12–14.
———. 1998. Palea-botany. Fiddlehead Forum 25: 19–23.
Moran, R. C., and R. Riba, eds. 1995. Psilotaceae a Salviniaceae. In:Flora Mesoamericana, Vol. 1, edited
by G. Davidse, M. Sousa S., and S. Knapp. Universidad Nacional Autónoma de México, Ciudad de
México.
Morton, C. V. 1967. The fern herbarium of André Michaux. American Fern Journal 57: 166–182.
Moteetee, A., J. G. Duckett, and A. J. Russell. 1996. Mycorrhizas in ferns of Lesotho. In:Pteridology in Per-
spective: Proceedings of the Holttum Memorial Pteridophyte Symposium, Kew, edited by J. M.
Camus, M. Gibby, and R. E. Johns.
Nooteboom, H. P. 1994. Notes on Davalliaceae. II: A revision of the genus Davallia. Blumea 39: 151–214.
———. 1997. The microsoroid ferns. Blumea 42: 261–395.
Palmer, D. D. 1994. The Hawaiian species of Cibotium. American Fern Journal 84: 73–85.
———. 1997. A revision of the genus Sadleria (Blechnaceae). Pacific Science 51: 288–305.
Panigrahi, G. 1993. Parahemionitis, a new genus of Pteridaceae. American Fern Journal 83: 90–92.
Parrella, M. P., K. L. Robb, and P. Morishita. 1985. Snails and slugs in ornamentals. California Agricul-
ture (January–February): 6–88.
Patterson, P. M., and A. S. Freeman. 1963. The effect of photoperiodism on certain ferns. American Fern
Journal 53: 126–128.
Pichi-Sermolli, R. E. G. 1962. On the fern genus Actiniopteris Link. Webbia 17: 1–32.
Piggott, A. G. 1988. Ferns of Malaysia in Colour. Tropical Press, Kuala Lumpur SDN.BHD, Malaysia.
Proctor, G. R. 1977. Flora of The Lesser Antilles: Pteridophyta. Arnold Arboretum, Harvard University,
Jamaica Plain, Massachusetts.
———. 1985. Ferns of Jamaica. British Museum (Natural History), London.
———. 1989. Ferns of Puerto Rico and the Virgin Islands. New York Botanical Garden, Bronx, New
York.
Pryer, K. M., A. R. Smith, and J. E. Skog. 1995. Phylogenetic relationships of extant ferns based on evi-
dence from morphology and rbcL sequences. American Fern Journal 85: 205–282.
Rasbach, H., T. Reichstein, and R. L. Viane. 1994. Asplenium chiuahuenge (Aspleniaceae, Pteridophyta):
An allohexaploid species and the description of a simplified hybridization technique. American Fern
Journal 84: 11–40.
Reasoner, S. 1982. Play “Misty” for me. Greenhouse Manager (September): 39–42.
Rickard, M. 1981. Survey of variations in British polypodiums. British Pteridological Society Bulletin 1:
138–140.
———. 1986. Vegetative reproduction in ferns. Pteridologist 1: 112–115.
———. 2000. The Plantfinder’s Guide to Garden Ferns. Timber Press, Portland, Oregon.
Rogers, G. 1998. The history and classification of the Farley fern, Adiantum tenerum ‘Farleyense’. Ameri-
can Fern Journal 88: 32–46.
Roos, M. C. 1985. Phylogenetic systematics of the Drynarioideae (Polypodiaceae). Verhandelingen der
Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde, Tweede Reeks 85:
1–138.
Rothwell, G. W., and R. A. Stockey. 1991. Onoclea sensibilis in the Paleocene of North America: A dramatic
example of structural and ecological stasis. Review of Palaeobotany and Palynology 709: 113–24.
570 L C

Rush, R. 1983. A hardy hybrid Adiantum. British Pteridological Society Bulletin 2: 261–262.
———. 1984a. A guide to hardy ferns. Special Publication No. 1. British Pteridological Society, London.
———. 1984b. Raising pteridophytes from spores: The special cases. Pteridologist 1: 3–5.
Scamman, E. 1960. The maidenhair ferns (Adiantum) of Costa Rica. Contributions of the Gray Herbar-
iumof Harvard University 187: 3–22.
Sessions, A. 1978. Key to the cultivars of Nephrolepis exaltata ‘Bostoniensis’. Privately published.
Shivas, M. G. 1962. The Polypodium vulgare complex. British Fern Gazette 9: 65–70.
Smith, A. R. 1981. Pteridophytes. In:Flora of Chiapas, Part 2, edited by D. E. Breedlove. California Acad-
emy of Sciences, San Francisco.
———. 1995. Pteridophytes. In:Flora of the Venezuelan Guayana, Vol. 2, edited by J. Steyermark, P. E.
Berry, and B. K. Holst. Timber Press, Portland, Oregon.
Snyder, L. H., Jr., and J. G. Bruce. 1986. Field Guide to the Ferns and Other Pteridophytes of Georgia.
University of Georgia Press, Athens.
de la Sota, E. R. 1963. Contribución al conocimiento de las “Salviniaceae” neotropicales. IV. Darwiniana
12: 612–623.
———. 1966. Revisión de las especies Americanas del grupo “Polypodium squamatum” L. “Polypodi-
aceae” (s. str.). Revista del Museo de La Plata, Sección Botânica 10: 69–186, t. I–VII.
Sowerby, J. E., and C. Johnson. 1855. The Ferns of Great Britain. Self-published, London.
Tagawa, M., and K. Iwatsuki. 1979–1989. Flora of Thailand. Vol. 3, Pteridophytes. Chutima Press,
Bangkok.
Taylor, W. C. 1984. Arkansas Ferns and Fern Allies. Milwaukee Public Museum, Milwaukee, Wisconsin.
Thurston, S. H. 1939. Forcing a collection of native ferns of New England and the middle Atlantic states
for exhibition. American Fern Journal 29: 85–94.
Tindale, M. 1965. A monograph of the genus Lastreopsis Ching. Contributions to the New South Wales
National Herbarium 3: 249–339, t. 1–23.
Trehane, P., ed. 1995. International Code of Nomenclature for Cultivated Plants. Regnum Vegetabile
133: 1–175.
Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Annals of the Missouri Botanical
Garden 44: 125–193.
Tryon, R. M. 1956. A revision of the American species of Notholaena. Contributions of the Gray Herbar-
iumof Harvard University 179: 1–106.
———. 1962. Taxonomic fern notes. II: Pityrogramma and Anogramma. Contributions of the Gray
Herbariumof Harvard University189: 5–76.
Tryon, R. M., and R. G. Stolze. 1989. Pteridophyta of Peru. Part I: 1. Ophioglossaceae–12. Cyatheaceae.
Fieldiana, Botany, new series 20: 1–145.
———. 1989. Pteridophyta of Peru. Part II: 13. Pteridaceae–15. Dennstaedtiaceae. Fieldiana, Botany, new
series 22: 1–128.
———. 1991. Pteridophyta of Peru. Part IV: 17. Dryopteridaceae. Fieldiana, Botany, new series 27: 1–
176.
———. 1993. Pteridophyta of Peru. Part V: 18. Aspleniaceae–21. Polypodiaceae. Fieldiana, Botany, new
series 27: 1–176.
———. 1994. Pteridophyta of Peru. Part VI: 22. Marsileaceae–28. Isoëtaceae. Fieldiana, Botany, new
series 32: 1–190.
Tryon, R. M., and A. F. Tryon. 1982. Ferns and Allied Plants, with Special Reference to Tropical Amer-
ica. Springer-Verlag, Berlin.
Tutin, T. G., V. H. Heywood, N. A. Burges, D. H. Valentine, S. M. Walters, D. A. Webb. 1964. Flora
Europaea. Vol. 1, Lycopodiaceae to Platanaceae. Cambridge University Press, Cambridge.
 L C 571

Vail, R. 1984. Platycerium Hobbyist’s Handbook. Desert Biological Publications, Mena, Arkansas.
van Hove, C. 1989. Azolla and its multiple uses, with emphasis on Africa. Food and Agriculture Organi-
zation of the United Nations, Rome.
Vareschi, V. 1968. Helechos. In:Flora de Venezuela, Vol. 1, parts 1 and 2. Edición Especial del Instituto
Botânico, Caracas.
Wagner, D. H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia
1: 1–64.
Wagner, G. M. 1997. Azolla: A review of its biology and utilization. Botanical Review 63: 1–26.
Wagner, W. H., Jr. 1942. Bipinnate Christmas fern. American Fern Journal 32: 27–29.
Wagner, W. H., Jr., and F. S. Wagner. 1975. A hybrid polypody from the New World tropics. Fern
Gazette 11: 125–135.
Walker, T. G. 1970. Species of Pteris commonly in cultivation. British Fern Gazette 10: 143–151.
Weatherby, C. A. 1939. The group of Polypodium polypodioides. Contributions of the Gray Herbariumof
Harvard University 124: 22–35.
Webster, T. R. 1979. An artificial crossing technique for Selaginella. American Fern Journal 69: 9–13.
Wherry, E. T. 1972. The Southern Fern Guide. Doubleday, Garden City, New York.
Windham, M. D. 1987. Argyrochosma, a new genus of cheilanthoid ferns. American Fern Journal 77:
37–41.
Yatskievych, G. 1996. A revision of the fern genus Phanerophlebia (Dryopteridaceae). Annals of the Mis-
souri Botanical Garden 83: 168–199.
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 Subject Index

Please see page 581 for the Plant Name Index.

acid-loving ferns, 43 watering ferns in, 34, 62

acid soil, 42–43 wire, 58
aeration of soil, 37–38 wood, 58
agar, in nutrient solutions, 77 basket ferns, see also baskets
agricultural inspector, 24 keeping them moist, 62
air planting, 37, 58–62
circulation of, 34 planting-mix for, 37
dry-air injury, 34 popular species of, 97; see also Davallia and
fresh, 34 Polypodium in Plant Name Index
humidity, 34–36 replanting, 62
air layering, 69 special care for, 37, 62
algae, 138–139 watering, 34, 62
and high humidity, 36 beds, planting
Oscillatoria, 81, 138 planting procedures, 52
alpine ferns, 11 preparation of, 52
American Fern Society, Appendix II raised, 40
annulus, 19, 21 soil for, 40
antheridium (antheridia), 63–64 Beyerinck’s solution, 78
in hybridization, 83–86 binomial system of naming plants, 141
apogamy, 64 biological controls, 121–123
apospory, 64 blades, see leaves
aquatic ferns, 93 blight diseases, 135
archegonium (archegonia), 63 blood meal, 45
in hybridization, 83–86 bog ferns, 93
Boston ferns, 107–108; see also Nephrolepis exaltata in
bacteria, see fungi and bacteria Plant Name Index
bactericides, Appendix IV care of, 107–108
bark products, 40 grown from meristem cultures, 108
basic soil, 42–43 and high humidity, 36, 107
baskets, see also basket ferns for indoor planting, 99
ceramic, 58 light requirements, 107
drainage in, 34, 37 repotting, 108
linings for, 59–61 stems, 13, 108
planting in, 58–62 watering, 107–108
selecting basket, 58 Botanical Code of Nomenclature, 141
tree-fern fiber, 12, 16, 58 bottle gardens, see terrariums

573
574 S I

British Pteridological Society, 542 drainage

bud propagation in baskets, 37, 62
plants suitable for, 64, 65, 69 in beds, 52
staghorn ferns, 104–105 poor, special care for, 33
in pots, 55, 56–57
calcium, see also basic soil in terrariums, 101
addition to soil, 43, 46 for xerophytes, 111–112
forms of, 43
testing for, 43 egg(s), water needed for fertilization of, 63
carbon dioxide, 25 epiphytic ferns
charcoal, 40 in baskets, 58–62
chemical control and humidity, 34
alternatives to, 121–123 etiolation, 81
substances, Appendix IV evergreen ferns, 95–97, 154–155
CITES, 543
cladograms to family classification of ferns in text, Appendix V
fern families, Appendix V fatty acid sprays, 121
vascular plants, Appendix V fern(s)
classification of fern families, Appendix V accent ferns, 97
clay soil, 38 acid-loving, 43
collecting native plants, 23 acquisition of, 22–23; see also importing ferns;
conducting tissue, see vascular tissue transporting ferns
copper wire in hanging baskets, 62 alpine, 11
Cornell planting mixes, 39 aquatic, 93
costa (costae), 14 basket fern, see basket ferns
croziers, or fiddleheads, 12, 14, 16, 49 bog or wet soil, 93
cultivated varieties, see cultivars border ferns, 90, 97
cultivars (cv.), 142–143 buying, 22–23
Cultivated Code (International Code of characteristics of, 11
Nomenclature for Cultivated Plants), 143, classification of, see naming ferns; Appendix V
149 collecting native plants, 23
cultivation, see ferns; tilling with color in fronds, 95
cultural problems, 118–139 cultural needs of, 25–36; see also air; fertilizer;
grooming ferns; humidity; light; soil; soil mixes;
damage, see disease and damage temperature tolerances; watering
damping off, 135 cultural problems, recognition of, 118–120
davallia, 107 deciduous, 50, 95–96, 154
deciduous ferns, 50, 95–96, 154 desert, 11, 111–112; see also xerophytic ferns
desert ferns, 11, 111–112; see also xerophytic ferns fall care of, 50
disease and damage, see also, algae; fungi and bacteria; for drier soil, but needing regular watering, 94
insects and other pests; insecticides and other easy to grow from spores, 82
pesticides; liverworts, mosses economic uses of, 12
due to copper, 62 edible, 12
due to high humidity, 36 epiphytic, see epiphytic ferns
due to light, 30 evergreen, 95–97, 154–155
due to overfertilization, 46–47 filmy, 112
due to salts, 32 floras, 155
due to smog, and other gases, 36 folklore, 12
due to extreme temperatures, 28–29 forcing, 116–117
due to watering, 33 foundation ferns, 90
recognition of, 36 grooming of, 48–50
disinfectants, Appendix IV ground-cover ferns, 90, 97
dolomite, 38, 43 hybridization methods, 83–86
dormancy, see rest period identification of, 149–150, 155
double potting, 36 importing from abroad, 23, Appendix III
 S I 575

indoor, see house ferns fertilizer

interstate regulations on tranport, 23, Appendix III application, 46
life cycles of, 63–67 frequency of, 44, 46, 50
limestone-loving, 43, 94 complete, 45
naming ferns, see naming ferns incomplete, 45
outdoor, see landscaping with ferns; tree ferns liquid vs. dry, 45
pests and diseases, recognition of, 120; see also algae; low-burn, 45
fungi and bacteria; insects and other pests manure, 45
purchasing, 22 minerals (elements) in, 25
mosses and liverworts, damage by, 138–139 organic vs. inorganic, 40, 45
propagation, 63–86; see also propagation sandy soil and, 37
prothallium of, see prothallium in sawdust and woodwaste, 42
rapid-growing, 94 selection of, 45
rest periods of, 46, 49, 96 slow-release, 45
rock garden ferns, 98 to spores after germination, 79
root-bound, 57 in terrariums, 102
by section of country, 91–92 trace elements, 46, 78
shade-tolerant, 92 fiddleheads, or croziers, 12, 14
for shows, 115–117 filmy ferns, 112–113
slow-growing, 95 fish emulsions, 45
spring care of, 49 floras, 155
structure of, 13–21; see also fruiting bodies; leaves; foot-candles, Appendix I
roots; stems forcing plants, 116–117
summer care of, 49–50 forma (f.), 143–147
sun-tolerant, 92 forms, see forma
temperature damage, 28–29 fronds, see leaves
temperature tolerances of, 28–29, 152–153; see also frost damage, 28
hardy ferns; semi-hardy ferns; tender ferns fruiting bodies, 17, 19, 21; see also annulus; indusium;
for terrariums, 101 propagation; sorus; sporangia; spores
terrestrial, see terrestrial ferns fungi and bacteria, 134–138; see also algae; insects and
tree ferns, see tree ferns other pests; mosses and liverworts
tropical, 11 Armillaria root rot, 40, 135
uses of, 12 blights, 135
variations in, 142–147 Botrytis, 134, 135
wall ferns, 98 control of, 134–138, Appendix IV
for wet or boggy areas, 93 damping off, 135
winter care of, 49 Fumago, 137
xerophytic, see xerophytic ferns gray mold, 135
year-round care of, 49–50 and high humidity, 36
fern allies leaf spots, 136
ground pines, 113–114 manure and, 45, 68
horsetails, 115 Phytophthora, 81, 134, 138
Selaginella, 114–115 Pythium, 81, 134, 138
whisk ferns, 113 recognizing damage from, 134–138
fern societies, 22, Appendix II Rhizoctonia, 81, 134, 136
fertilization root rot, 138
during summer, 49 rots, 137
overfertilization, and its treatment, 46–47 rusts, 137
recognizing need for, 46 sooty molds, 137
reduced, or withheld in the fall, 50 symbiosis, 16
of sowed spore, 79 tip blight, 137–138
spingtime resumption of, 49 water molds, 134, 138
of staghorn ferns, 45, 106 fungicides and allied materials
fertilization, in life cycle; see also, spore propagation preventing damage from, 123–124
water needed for, 63, 64 use, 134–138, Appendix IV
576 S I

gametophyte, see prothallium baits, 122

gas injury, 36 chemicals and other controls, Appendix IV
gemmae, 67 detergent sprays, 121
genus (genera) fatty acid sprays, 121
identification of, 149, 155 insect growth regulators, 122
learning the, 149–150 oil sprays, 121
listing of genera in cultivation, Appendix V pheromones, 122
geographical zones and hardiness ratings, 153, plate 20 plant-derived, 122
Gordan, 41 soap sprays, 121
gray mold, 135 traps, 122
grooming ferns, 50 water spraying, 123
identification of, 149, 155 insects and other pests; see also algae; fungi and bacteria;
for shows and exhibits, 115–117 mosses and liverworts
growing tip ants, 124
condition of, when buying plant, 22 aphids, 49, 129
damage to, 22 beetles, 133
caterpillars, 133
hardened plants, 99 cockroaches, 133
hardiness zones, 27, 153, plate 20 crickets, 133
hardy ferns, 26, 153 cutworms, 133
height, 152 earwigs, 127
Hoagland’s solution #2, 78 fungus gnats, 127
hormones, plant, 47 grasshoppers, 133
house ferns; see also, terrariums larvae, 133
favorite species of, 99–100 mealybugs, 128
and humidity, 34–36 millipedes, 129
selecting plants, 99 mites, 129
watering, 32–33, 99 nematodes, or nemas, 129–130
humidity pillbugs, 132
amounts needed, 34–36 psocids, 130
high-humidity damage, 36 red spiders mites, 129
for filmy ferns, 113 scales, 130–131
for house plants, 34–36, 99 slugs, 131–132
how to increase, 36 snails, 131–132
low-humidity damage, 34 sowbugs, 132
need for, 34 spider mites, 129
in terrariums, 102 springtails, 130
humus, 40 thrips, 132–133
hybrid(s) white flies, 133
breeding true or variable, 82 International Code of Botanical Nomenclature, 141
naming, 147–148 International Code of Nomenclature for Cultivated
recognition of, 83 Plants, 143
sterility in, 83 interstate transport of ferns, 23, Appendix III
hybridization methods, 83–86
Knop’s solution, 78
importing ferns, 24–24 Knudson’s C solution, 78
holding until departure, 23
how to pack, 23 landscaping with ferns
regulations, 23–34, Appendix III accent ferns, 97
indoor ferns, see house ferns aquatic ferns, 93
indusium basket ferns, 97
false, 19, 20 bog or wet soil ferns, 93
in identification, 19, 149–150 border ferns, 90, 97
and ripeness of spores, 19 with color in fronds, 95
insect controls deciduous ferns, 50, 95–96, 154
 S I 577

drier soil ferns that need regular watering, 94 shading, 30

evergreen ferns, 95–97, 154–155 for sowed spore, 79
foundation ferns, 90, 97 in terrariums, 102
ground-cover ferns, 90, 97 limestone, 38, 43, 46, 94; see also calcium
placement, 87 limestone-loving ferns, 43, 94
rapid-growing, 94 liverworts, 138–139
rock garden ferns, 98 loam, 41
by section of country for outdoor use
California, coastal central and southern, 92 maidenhair ferns, 102–104
central states, 91 manure, 45
northeastern United States, 91 margins, leaf, 19
northwestern United States, 91 meristem culture, 71
southern Florida and Hawaii, 91 microwave sterilization of soil, 44
southeastern United States, 91 minerals in fertilizers, 45; see also fertilizer
southwestern United States, 92 molds
shade-tolerant ferns, 92 control of, 134–138, Appendix IV
slow-growing ferns, 95 gray mold, 135–136
to solve particular problems, 87–91, 118 overhead watering and formation of, 134
sun-tolerant ferns, 92–93 sooty, 137
tree ferns, see tree ferns water molds, 138
wall ferns, 89, 98 monkey tails, see croziers, or fiddleheads
wet-soil ferns, 93 monolete spores, 21
leaf mold, 38 mosses, 138–139
leaf spots, 136 mulches, 40
leaves, or fronds, 16–17; see also stipes mutations, 82, 86
abnormal, 118–120 mycorrhizae, 16
age variation in, 65
blades of, 17 naming ferns, 141–150
buds on, 65, 69, 70 author citation, 141
with color, 95 binomial system, 141
compound, 17 Botanical Code of Nomenclature, 141
croziers, or fiddleheads, 12, 16 classification, Appendix V
divisions of, 17 common names, 140
function of, 17 cultivars, 142–143
induced to form buds, 65, 69 cultivated Code, 143, 149
margins of, 17 forms ( formae), 146
planting buds from, 65, 69, 70 genus, 141
polishes for, 49 hybrids, 147–148
removing damaged ones, 48 name changes, 148
shapes of, 16 obtaining patents, 149
shedding of, 49 pronouncing names, 141
simple, 17 species, 141
stipes of, 16, 69 specific epithet, 141
veins of, 17 synonyms, 140
washing off dust and insects, 48 variations within species, 142
life cycle of ferns, 63–67 abbreviations used in names, 146–147
light botanical varieties (varietas), 143
artificial, 30–31 cultivated varieties (cultivars), 142–143
foot candles, 31, Appendix I forms ( formae), 143–146
intensities and duration of, 30–31, 154 subspecies, 146
materials for shading, 30 nitrogen
measurement of, 31, Appendix I in fertilizers, 45, 46
photoperiodic responses, 31 with sawdust and wood waste, 42
quality, 30 nomenclature, see naming ferns
requirements for specific ferns, 99, 153 nutrient-solution method of sowing, 77–78
578 S I

oil sprays, 121 potting, double, 36

outdoor ferns, 91–92 propagation
overpotting, 56, 102–103 air layering, 69
oxygen apogamous, 64
in photosynthesis, 25 apospory, 64
in roots, 33 from buds, 64, 65, 69, 70
of fern allies, 113–115
patents, plant, 149 life cycles of ferns, see life cycles of ferns
peat, black, 41 prothallium proliferation, 64
peat, sedge, 41 from rhizome divisions, 53, 67–69
peat moss, 40, 41 spore propagation, see spore propagation
perlite, 38, 41 tissue culture, 71
pesticides, 123–124, Appendix IV vegetative, 67–72
pests, see insects and other pests prothallium, or gametophyte
petiole, see stipe damage to by molds, algae, insects, 81, 138–139
pH failure of, to produce plants, 81
optimum of nutrient solutions, 77; of soil 42–43 growing from fronds, 64
measurement of, 42 in hybridizing, 83–86
pheromones, 122 produced from gemmae, 67
phosphorus proliferation, 64
fertilizers, 45 sporophytes produced from, 63
with sawdust and wood waste, 42 transplanting, 79–80
photosynthesis, 25
pinna, 17 rachis, 14
pinnule, 14 rest periods, 49, 96
planting and fertilization, 46
in baskets, 37, 58–62; see also baskets rhizoids, 63
in beds, 51, 52–53; see also beds, planting rhizomes
digging ferns out of ground for transplanting, kinds of, 13, 15
52–54 dividing and planting, 60, 67–69
positioning and planting fern, 53 rock garden ferns, 98
in pots, 51, 53, 56; see also pots, planting in rockwool, 41
removing plants from pots, 53 root(s)
time for, 51, 52 aerial, 16
planting beds, see beds, planting buds on, 69
planting mixes, see soil mixes characteristics, 16, 25
plugs, for growing ferns from spores, 80 emergence of, 16
potassium, in fertilizers, 45 functions of, 16
pots, see also terrariums and oxygen, 25, 33
clay vs. plastic, 32, 56 root-bound ferns, 57
salt-encrusted, 32, 33, 58 root mantle of tree ferns, 109
selecting the pot, 56 root rot, 33, 135; see also fungi and bacteria
shape of, 56–57 rots, 137
size, 56 rust diseases, 137
sterilization of, 44
without drain holes, 56, 57 salt injury
pots, planting in excess of salt in water, 33
basic procedures for, 55–58 overfertilization, 46
overpotting, 56, 102–102 scant watering, 32
sowing spores on, 77 prevention of, 58
potted plants San Diego Fern Society web site, Appendix II
poor drainage in, 33, 56, 57 sand, for potting mixes, 37, 42
providing drainage in, 55–57; see also drainage sandy soil, 37, 42
root-bound, 57 sawdust and wood waste, 42
special care for, 57–58 scientific names, 140–148
 S I 579

semi-hardy ferns, 26, 27, 153 species in cultivation, 151–540

semi-tender ferns, 26, 27, 153 specific epithet, 141
shade-tolerant ferns, 92 sperm(s), 63
shading materials, see light Shagnum moss, 41
silty soil, 38 sowing spores, see spore propagation
smog damage, 36 sporangia (spore cases)
soap sprays, 121 annulus positions, 21
soil(s), see also soil additives; soil mixes collecting spores from, 23
acidic, 42–43 shedding spores, 19
additives, see soil additives spore(s), 21; see also spore propagation
basic, 42–43 envelopes, 73
clay, 38, 42 green, 12, 74
granitic, 39 importation of, Appendix III
loam, 38, 40, 41 kinds of, 21
mulches, 40 length of life, 74
for outdoor planting beds, 37, 52 monolete, 21
pH of, 42–43 packets, 73
potting, 37 in life cycle, 63
and raised planting beds, 52 non-green, 21, 74
sandy, 37, 42, 42 trilete, 21
silty, 38, 42 viability of, 21, 74
sterilization of, 43–44 spore propagation; see also spores
tilling of, 46 after-sowing care
soil additives fertilizers, 79
for acid-loving ferns, 43, 94 light, 79
for basic-loving ferns, 43, 94 temperature, 79
bark products, 40 transplanting, 79–80
charcoal, 40, 101 watering, 79, 80, 81
compost, 40 breeding true or variable, 82
fertilizers, 40, 44; see also fertilizers cleaning spores, 73–74
Gordan, 41 collecting and storage of spores, 72–73
gypsum, 38 disinfecting spores, 74
humus, 40 ferns easy to grow by, 82
inorganic matter, 40; see also perlite; vermiculite germination time, 79
leaf mold, 40 hybridization, methods, 83–86
limestone, 38, 40, 43, 46; see also calcium plugs, 80–82
manure, 45 sowing methods and media, 74–82
mulches, 40 commercial method, 76
organic matter, 37, 38, 41; see also bark products; leaf inverted pot method, 77
mold; peat moss; sawdust and wood waste miscellaneous methods, 79
oyster shell, 43 nutrient agar solution, 77
peat moss, 41, 61, 62 nutrient solution method, 77
perlite, 38, 41, 101 pot method, 77
rockwool, 41 sowing methods and media, 74–82
sand, 37, 42 special care, 81–82
sawdust, 42 sterilizing, 74
sulfur, 43 staghorn ferns, 104–105
vermiculite, 42 steam sterilization of soil, 43
wood waste, 42 stems, 13
soil mixes, see also soil additives functions of, 13
Cornell, 39 rhizomes, 13
with soil, 38 rootstock, 13
soilless, 37, 38, 41 stolons, 13
sooty molds, 137 trunks, 13
sorus (sori), 17, 19–21, 150, 155 vascular, or conducting, tissue in, 11, 13
580 S I

sterilization propagation of, 110

of equipment, 43 roots of, 16, 109
of soil, 43–44 temperature tolerances of, 110
of spores, 74 trunks of, 13, 16, 109, 110
stipe(s), 16; see also leaves uses of, 13
buds formed on, 69 trilete spores, 21
characteristics of, 16 trimming, see grooming ferns
in identification, 16, 155 trough gardens, 100
stipules, induced to form buds, 71 trunks, tree-fern, 16, 109–111
stolons, 13
subspecies, 146 varietas (var.), 142–147
sun-tolerant ferns, 92–93 varieties, see varietas
synonyms, 140 vascular, or conducting, tissue, 15, 155
veins, 17
temperature vitamin B, 53
cold requirements, 29
damage to ferns, 28–29 water, see also watering
for forcing plants, 116–117 distilled, or rain, 99
and frond longevity, 96 need for, 25
local temperature differences, 26, 27, 153 overwatering, 33
preparation for low winter temperatures, 28–29, 49 quality of, and salt injury, 33; see also salt injury
after sowing spores, 79 salt in, 33
treatment of plants during heat wave, 29 softeners in, 33
in USDA zones, 152–153, color plate 20 temperature of, 33
tender ferns, 26, 27, 153 watering, see also water
terrariums, or bottle gardens, 100–102 damage due to, 33
care of, 102 during vacation, 34
condensation in, 102 ferns in baskets, 34, 97–98
containers for, 100 frequency of, 32
ferns for, 100 house ferns, 32–34, 99–99
and increased humidity, 102 methods, 32, 53, 134
planting of, 101–102 overhead, and mold formation, 81
watering, 102 of potted plants, 32, 53
terrestrial ferns and root rot, 33, 135
planting in baskets, 37, 62 of sowed spores, 79, 81
planting in beds, 51–53 staghorn ferns, 104
planting mix for, 37 terrariums, 102
special care for, in baskets, 62 treating wilted plants, 32
tip blight, 137–138 treating overwatered plants, 32, 33
tissue culture, 71 xerophytes, 111–112
transplanting water molds, 138
basket ferns, 62 water spots on foliage, 49
in beds, 52–54 removal of, 49
in pots, 53, 55 web sites about ferns, Appendix II
prothallia, 63, 79 wet-soil ferns, 93
time for, 51, 52 wetting agents, 74
transporting ferns, 23–24 wilted plant, treatment for, 32
foreign imports, 23, Appendix III winter, preparing plants for, 28–29, 49
holding until departure, 23
how to pack, 23 xerophytic ferns, 111–112; see also desert ferns; rock
interstate, 24 garden ferns
tree ferns, see in Plant Name Index: Alsophila; Cibotium; care of, 111–112
Ctenitis; Cyathea; Dicksonia; Sadleria; Sphaeropteris characteristics, 111–112
care of, 109–111 genera composed of, 111
fronds of, 109–111 and humidity, 112
genera containing, 111 watering, 112
 Plant Name Index

Plants listed in boldface have entries in Chapter 13, “Ferns and Fern Allies in Cultivation.” Cultivars and
common names are cross-referenced to species or genus. Cultivars are listed at the end of the genus.

abrupt-tipped maiden fern, see Thelypteris augescens, 519 Adiantum jordanii, 148, 166
Acrosorus, 333 Adiantum latifolium, 166
Acrostichum, 156 Adiantum lucidum, 167
Acrostichum aureum, 93, 156 Adiantum lunulatum, 167
Acrostichum danaeifolium, 93, 157 Adiantum macrophyllum, 95, 103, 167
Actiniopteris, 111, 157 Adiantum ×mairisii, see Adiantum capillus-veneris
Actiniopteris radiata, 158 ‘Mairisii’, 162
Actiniopteris semiflabellata, 157 Adiantum monochlamys, 168, color plate 10
Acystopteris japonica, 258 Adiantum patens, 168
adder’s-tongue fern, see Ophioglossum, 404 Adiantum pedatum, 29, 91, 92, 97, 103, 168
Adenophorus, 333 Adiantum pedatum subsp. aleuticum, see Adiantum
Adiantopsis, 158 aleuticum, 160
Adiantopsis radiata, 158 Adiantum pentadactylon, 169
Adiantum, 102–104, 159 Adiantum peruvianum, 103, 170
Adiantum aethiopicum, 159 Adiantum petiolatum, 170
Adiantum aleuticum, 92, 148, 160 Adiantum philippense, see Adiantum lunulatum, 167
Adiantum aleuticum subsp. subpumilum, 98, 160 Adiantum poiretti, 170
Adiantum anceps, 103, 160 Adiantum polyphyllum, 171
Adiantum banksianum, see Adiantum capillus-veneris Adiantum pubescens, 166
‘Mairisii’, 161 Adiantum pulverulentum, 171
Adiantum bellum, 160 Adiantum pyramidale, 171
Adiantum capillus-veneris, 82, 92, 98, 101, 103, 161 Adiantum raddianum, 82, 92, 94, 95, 97, 98, 103, 172
Adiantum caudatum, 69, 70, 103, 162 Adiantum reniforme, 173
Adiantum concinnum, 162 Adiantum seemannii, 173
Adiantum cristatum, see Adiantum pyramidale, 171 Adiantum tenerum, 94, 103, 173
Adiantum cultratum, 163 Adiantum tetraphyllum, 174
Adiantum cuneatum, see Adiantum raddianum, 172 Adiantum ×tracyi, 83, 103, 148, 175
Adiantum diaphanum, 163 Adiantum trapeziforme, 91, 103, 175
Adiantum excisum, 164 Adiantum venustum, 98, 175
Adiantum flabellulatum, 164 Adiantum villosum, 176
Adiantum formosum, 103, 164 Adiantum cultivars
Adiantum fragile, 165 ‘Banksianum’, see Adiantum capillus-veneris ‘Mairisii’,
Adiantum fructuosum, 165 162
Adiantum hispidulum, 82, 95, 97, 98, 100, 101, 103, ‘Birkenheadii’, see Adiantum diaphanum, 162
166, color plate 9 ‘Croweanum’, see Adiantum raddianum, 172

581
582 P N I

[Adiantum cultivars] Amauropelta rudis, see Thelypteris rudis, 525

‘Edwinii’, see Adiantum concinnum, 162 American alpine lady fern, see Athyrium alpestre, 208
‘Farleyense’, see Adiantum tenerum, 103, 173 American glade fern, see Diplazium pycnocarpon, 288
‘Fergusonii’, see Adiantum tenerum, 173 American maidenhair, see Adiantum pedatum, 168
‘Fimbriatum’, see Adiantum capillus-veneris, 161 American ostrich fern, see Matteuccia struthiopteris var.
‘Fragrans’, see Adiantum raddianum, 172 pensylvanica, 371
‘Fragrantissimum’, see Adiantum raddianum, 172 American parsley fern, see Cryptogramma acrostichoides,
‘Fritz Luth’, see Adiantum raddianum, 172 249
‘Gracillimum’, see Adiantum raddianum, 101, 172 American staghorn, see Platycerium andinum, 429
‘Grandiceps’, see Adiantum raddianum, 172 American tree fern, see Ctenitis sloanei, 251
‘Imbricatum’, see Adiantum aleuticum subsp. American wall fern, see Polypodium virginianum, 452
subpumilum, 160; Adiantum capillus-veneris, 103, American water fern, see Ceratopteris pteridoides, 232
162 Anchistea, see Woodwardia, 535
‘Lady Geneva’, see Adiantum raddianum, 172 Anchistea virginica, see Woodwardia virginica, 539
‘Mairisii’, see Adiantum capillus-veneris, 162 Anderson’s holly fern, see Polystichum andersonii, 456
‘Micropinnulum’, see Adiantum raddianum, 101, 172 Anderson’s sword fern, see Polystichum andersonii, 456
‘Ocean Spray’, see Adiantum raddianum, 173 Anemia, 182
‘Pacific Maid’, see Adiantum raddianum, 101, 103, Anemia adiantifolia, 183
173 Anemia phyllitidis, 183
‘Pacottii’, see Adiantum raddianum, 98, 101, 173 Anemia rotundifolia, 183
‘Triumph’, see Adiantum raddianum, 173 Angiopteris, 71, 95, 184
‘Variegatum’, see Adiantum raddianum, 173 Angiopteris angustifolia, 185
‘Weigandii’, see Adiantum raddianum, 172 Angiopteris evecta, 186
Aglaomorpha, 98, 176 Angola staghorn, see Platycerium elephantotis, 431
Aglaomorpha brooksii, 179 Anisosorus, see Lonchitis, 353
Aglaomorpha cornucopia, 177 Anogramma, 186
Aglaomorpha coronans, 91, 93, 94, 97, 177 Anogramma chaerophylla, 94, 101, 186
Aglaomorpha drynarioides, 178 Anogramma leptophylla, 186
Aglaomorpha heraclea, 178 ant fern, see Lecanopteris, 345
Aglaomorpha meyeniana, 179 Appalachian polypody, see Polypodium appalachianum,
Aglaomorpha pilosa, 179 441
Aglaomorpha robertsii, see ×Aglaonaria robertsii, 180 Arachniodes, 187
Aglaomorpha speciosa, 179 Arachniodes aristata, 187
Aglaomorpha splendens, 180 Arachniodes aristata ‘Variegatum’, see Arachniodes
Aglaomorpha cultivars simplicior, 188
‘Angeli’, see Aglaomorpha coronans, 177 Arachniodes simplicior, 95, 188
‘Roberts’, see ×Aglaonaria robertsii, 180 Arachniodes standishii, 92, 188
‘Santa Rosa’, see ×Aglaonaria robertsii, 181 Arachniodes ‘Variegatum’, see Arachniodes simplicior, 188
‘Starburst’, see ×Aglaonaria robertsii, 181 Araiostegia, see Davallia, 262
×Aglaonaria robertsii, 180 Araiostegia hymenophylloides, see Davallia
Alabama lip fern, see Cheilanthes alabamensis, 236 hymenophylloides, 268
Alaskan holly fern, see Polystichum setigerum, 465 Araiostegia pulchra, see Davallia pulchra, 272
Alaska parsley fern, see Cryptogramma sitchensis, 250 arching wood fern, see Dryopteris expansa, 308
Aleuritopteris argentea, see Cheilanthes argentea, 237, Argyrochosma, 188, 236
color plate 21 Argyrochosma dealbata, 189
Aleuritopteris farinosa, see Cheilanthes farinosa, 239 Argyrochosma jonesii, 189
Allantodia, see Diplazium, 285 Argyrochosma microphylla, 190
alpine club moss, see Lycopodium selago, 360 Arizona lip fern, see Cheilanthes arizonica, 237
alpine lady fern, see Athyrium distentifolium, 208 Arthropteris, 190
Alsophila, 181, 253 Arthropteris monocarpa, 191
Alsophila australis, 182 Arthropteris orientalis, 191
Alsophila tricolor, 95, 111, 182, color plate 11 Arthropteris tenella, 191
amaumau, see Sadleria cyatheoides, 492 Asian sword fern, see Nephrolepis multiflora, 397
Amauropelta, see Thelypteris, 517 Asian walking fern, see Camptosorus sibericus, 228
Amauropelta resinifera, see Thelypteris resinifera, 524 Aspidotis, 192, 236
 P N I 583

Aspidotis californica, 192 Asplenium trichomanes-ramosum, 206

Aspidotis carlotta-halliae, 192 Asplenium tripteropus, 205
Aspidotis densa, 193 Asplenium viride, see Asplenium trichomanes-ramosum,
Asplenium, 193 206
Asplenium adiantum-nigrum, 94, 149, 194 Asplenium viviparum, see Asplenium daucifolium, 197
Asplenium aethiopicum, 194 Asplenium cultivars
Asplenium antiquum, 100, 195, color plate 12 ‘Antiquum’, see Asplenium antiquum, 195
Asplenium aureum, see Ceterach aureum, 234 ‘Australis’, see Asplenium australasicum, 196
Asplenium auritum, 69, 195 ‘Crispafolium’, see Asplenium nidus var. plicatum,
Asplenium australasicum, 100, 196 199
Asplenium australis, see Asplenium australasicum, 196 ‘Crispum’, see Asplenium nidus var. plicatum, 199
Asplenium belangeri, see Asplenium thunbergii, 205 ‘Cristatum’, see Asplenium trichomanes, 205
Asplenium bulbiferum, 69, 92, 97, 142, 196 ‘Incisum’, see Asplenium trichomanes, 205
Asplenium ceterach, see Ceterach officinarum, 234, color ‘Undulatum’, see Asplenium nidus var. plicatum, 199
plate 20 ‘Victoria’, see Asplenium antiquum, 195, color plate
Asplenium ×crucibuli, 196 12
Asplenium cuspidatum, 101, 197 ×Asplenosorus crucibuli, see Asplenium ×crucibuli, 196
Asplenium daucifolium, 69, 100, 101, 142, 197 ×Asplenosorus ebenoides, see Asplenium ×ebenoides, 198
Asplenium ×ebenoides, 98, 198 ×Asplenosorus pinnatifidus, see Asplenium pinnatifidum,
Asplenium falcatum, see Asplenium polyodon, 202 202
Asplenium flabellifolium, 198 Astrolepis, 98, 111, 206
Asplenium fontanum, 199 Astrolepis beitelii, 206
Asplenium kenzoi, 203 Astrolepis sinuata, 206
Asplenium longissimum, 199 Athyrium, 207
Asplenium lucidum, see Asplenium oblongifolium Athyrium alpestre, 208
Asplenium marinum, 199 Athyrium angustum, see Athyrium filix-femina var.
Asplenium mayii, see Asplenium surrogatum, 204 angustum, 209
Asplenium montanum, 202 Athyrium asplenioides, see Athyrium filix-femina var.
Asplenium nidus, 91, 97, 100, 199 asplenioides, 211
Asplenium nidus var. plicatum, 200 Athyrium australe, see Diplazium australe, 286
Asplenium nigripes, see Schaffneria nigripes, 494 Athyrium delavayi, 208
Asplenium oblongifolium, 200 Athyrium distentifolium, 208
Asplenium officinarum, see Ceterach officinarum, 234, Athyrium distentifolium var. americanum, see Athyrium
color plate 20 alpestre, 208
Asplenium oligophlebium, 101, 200 Athyrium dubium, see Diplazium subsinuatum, 288
Asplenium pellucidum, 201 Athyrium epirachis, 212
Asplenium pinnatifidum, 202 Athyrium esculentum, see Diplazium esculentum, 12, 69,
Asplenium platyneuron, 82, 92, 94, 98, 202 287
Asplenium polyodon, 202 Athyrium filix-femina, 69, 82, 92, 93, 94, 96, 97, 98,
Asplenium praemorsum, 194 209
Asplenium prolongatum, 202 Athyrium filix-femina var. angustum, 209
Asplenium pteridoides, see Asplenium surrogatum, 204 Athyrium filix-femina var. asplenioides, 211
Asplenium resiliens, 203 Athyrium filix-femina var. californicum, 211
Asplenium rhizophyllum, see Camptosorus rhizophyllus, Athyrium filix-femina var. cyclosorum, 211
228 Athyrium filix-femina var. filix-femina, 211
Asplenium ruprechtii, see Camptosorus sibericus, 228 Athyrium goeringianum, 212
Asplenium ruta-muraria, 94, 98, 203 Athyrium iseanum, 212
Asplenium scolopendrium, see Phyllitis scolopendrium, 422, Athyrium japonicum, see Deparia japonica, 280
color plate 39 Athyrium niponicum, 92, 95, 96, 211, color plate 14
Asplenium septentrionale, 204 Athyrium otophorum, 95, 96, 212, color plate 15
Asplenium serratum, 204 Athyrium petersenii, see Deparia petersenii, 281
Asplenium sibericum, see Camptosorus sibericus, 228 Athyrium proliferum, see Diplazium proliferum, 287
Asplenium surrogatum, 204 Athyrium pycnocarpon, see Diplazium pycnocarpon, 288
Asplenium thunbergii, 205 Athyrium thelypteroides, see Deparia acrostichoides, 280
Asplenium trichomanes, 98, 101, 205, color plate 13 Athyrium vidalii, 212
584 P N I

Athyrium cultivars Blechnum australe var. auriculatum, 216

‘Cristatum’, see Athyrium filix-femina, 211 Blechnum brasiliense, 95, 97, 111, 216
‘Cruciato-cristatum’, see Athyrium filix-femina, 211 Blechnum capense, 217
‘Fieldii, see Athyrium filix-femina, 211 Blechnum chilense, 218
‘Frizelliae’, see Athyrium filix-femina, 144, 211 Blechnum cordatum, 218
‘Minutissimum’, see Athyrium filix-femina, 211 Blechnum falciforme, 218
‘Pictum’, see Athyrium niponicum, 97, 212, color Blechnum fraxineum, see Blechnum gracile, 219
plate 14 Blechnum gibbum, 97, 111, 218, color plate 17
‘Veroniae-cristatum’, see Athyrium filix-femina, 211 Blechnum glandulosum, see Blechnum appendiculatum,
‘Victoriae’, see Athyrium filix-femina, 211 216, color plate 1
Australian brake, see Pteris tremula, 479 Blechnum gracile, 219
Australian cliff brake, see Pellaea falcata, 414 Blechnum indicum, see Asplenium longissimum, 199;
Australian davallia, see Davallia pyxidata, 272 Blechnum serrulatum, 221
Australian hare’s-foot fern, see Davallia pyxidata, 272 Blechnum magellanicum, 218
Australian maidenhair, see Adiantum formosum, 164 Blechnum moorei, 219
Australian tree fern, see Alsophila australis, 182; Blechnum niponicum, 219
Sphaeropteris cooperi, 91, 92, 511, color plates 2, 3 Blechnum occidentale, 220
austral lady fern, see Diplazium australe, 286 Blechnum occidentale var. minor, see Blechnum
autumn fern, see Dryopteris erythrosora, 307, color plate glandulosum, 216
26 Blechnum penna-marina, 94, 96, 97, 220, color plate 18
Azolla, 12, 93, 213 Blechnum polypodioides, 220
Azolla caroliniana, 214 Blechnum punctulatum, 221
Azolla filiculoides, 214 Blechnum serrulatum, 93, 221
Azolla pinnata, 214, color plate 16 Blechnum spicant, 92, 94, 96, 221
Azolla rubra, see Azolla filiculoides, 214 Blechnum tabulare, 218
Blechnum unilaterale, see Blechnum polypodioides, 220
bamboo fern, see Coniogramme japonica, 248 Blechnum wattsii, 218
Barne’s male fern, see Dryopteris filix-mas ‘Barnesii’, 310 Blechnum cultivars
bat fern, see Histiopteris, 338 ‘Crispum’, see Blechnum brasiliense, 216; Blechnum
bat-winged fern, see Histiopteris, 338 spicant, 222
beaded lip fern, see Cheilanthes lendigera, 241 ‘Cristata’, see Blechnum penna-marina, 220
beaded wood fern, see Dryopteris bissetiana, 300 ‘Redwood Giant’, see Blechnum spicant, 222
bead fern, see Onoclea, 402 ‘Serratum’, see Blechnum spicant, 222
bear’s-foot fern, see Davallia tyermannii, 276 blond tree fern, see Sphaeropteris horrida, 512
bear’s-paw fern, see Phlebodium aureum, 420 blue spike moss, see Selaginella uncinata, 505, color plate
beech fern, see Phegopteris, 418 49
Beitel’s cloak fern, see Astrolepis beitelii, 206 blunt-lobed woodsia, see Woodsia obtusa, 533
Belvisia, 214 bog fern, see Thelypteris simulata, 526
Belvisia callifolia, see Belvisia mucronata, 214 Bolbitis, 222
Belvisia mucronata, 214 Bolbitis aliena, 223
Bermuda maidenhair, see Adiantum bellum, 160 Bolbitis cladorrhizans, see Bolbitis portoricensis, 225
berry bladder fern, see Cystopteris bulbifera, 259 Bolbitis fluviatilis, 223
big-leaf holly fern, see Cyrtomium macrophyllum, 257 Bolbitis heteroclita, 224
bird’s-foot fern, see Pellaea mucronata, 415 Bolbitis heudelotii, 93, 225
bird’s-nest fern, see Asplenium nidus, 199 Bolbitis portoricensis, 225
black spleenwort, see Asplenium adiantum-nigrum, 194 Bolbitis simplicifolia, see Bolbitis heteroclita, 224
black-stem spleenwort, see Asplenium resiliens, 203 Bolbitis cultivars
black tree fern, see Sphaeropteris medullaris, 512 ‘Malacca’, see Bolbitis heteroclita, 224
black wood fern, see Dryopteris cycadina, 305 ‘Slender Bolbitis’, see Bolbitis heteroclita, 224
bladder fern, see Cystopteris, 258 Bommeria, 225
Blechnum, 214 Bommeria hispida, 226
Blechnum amabile, 215 bonaire lip fern, see Cheilanthes bonariensis, 238
Blechnum appendiculatum, 90, 93, 94, 95, 97, 216, Boott’s wood fern, see Dryopteris ×boottii, 300
color plate 1 Boston fern, see Nephrolepis exaltata, 393
Blechnum australe, 216 Botrychium, 227
 P N I 585

Botrychium biternatum, 227 Cardiomanes, see Trichomanes, 528

Botrychium dissectum, 227 carrot fern, see Onychium japonicum, 403
Botrychium dissectum f. dissectum, 227 Cascade parsley fern, see Cryptogramma cascadensis, 249
Botrychium dissectum f. obliquum, 227 Ceratopteris, 12, 93, 232
Botrychium dissectum var. tenuifolium, see Botrychium Ceratopteris cornuta, 232
biternatum, 227 Ceratopteris deltoidea, see Ceratopteris richardii, 233
Botrychium obliquum, see Botrychium dissectum, 227 Ceratopteris pteridoides, 232
Botrychium virginianum, 227 Ceratopteris richardii, 233
bracken, see Pteridium, 470 Ceratopteris thalictroides, 233, color plate 19
brake fern, see Pteris, 472 Ceterach, 233
bramble fern, see Hypolepis repens, 342 Ceterach aureum, 98, 234
Braun’s holly fern, see Polystichum braunii, 456, color Ceterach officinarum, 94, 234, color plate 20
plate 6 chain fern, see Woodwardia, 535
Bridges’ cliff brake, see Pellaea bridgesii, 413 Champion’s wood fern, see Dryopteris championii, 303
bristle fern, see Polystichum polyblepharum, 462; Cheilanthes, 93, 98, 111, 235
Trichomanes, 528 Cheilanthes alabamensis, 236
brittle bladder fern, see Cystopteris fragilis, 260 Cheilanthes argentea, 95, 237, color plate 21
brittle maidenhair, see Adiantum concinnum, 162; Cheilanthes arizonica, 237
Adiantum tenerum 173 Cheilanthes beitelii, see Astrolepis beitelii, 206
broad beech fern, see Phegopteris hexagonoptera, 419 Cheilanthes bonariensis, 82, 238
broad buckler fern, see Dryopteris dilatata, 306 Cheilanthes buchtienii, 238
broad sword fern, see Nephrolepis falcata, 395 Cheilanthes californica, see Aspidotis californica, 192
broad wood fern, see Dryopteris dilatata, 306 Cheilanthes carlotta-halliae, see Aspidotis carlotta-halliae,
Buchtien’s lip fern, see Cheilanthes buchtienii, 238 192
buckler fern, see Dryopteris, 296 Cheilanthes concolor, see Doryopteris concolor, 292
bulblet bladder fern, see Cystopteris bulbifera, 259 Cheilanthes covillei, 238
Butterfield holly fern, see Cyrtomium falcatum Cheilanthes dealbata, see Argyrochosma dealbata, 189
‘Butterfieldii’, 256 Cheilanthes eatonii, 239
button fern, see Pellaea rotundifolia, 416; Tectaria Cheilanthes farinosa, 239
gemmifera, 515 Cheilanthes feei, 239
Cheilanthes fendleri, 240, color plate 22
California holly fern, see Polystichum californicum, 457 Cheilanthes gracillima, 240
California lace fern, see Aspidotis californica, 192 Cheilanthes hirta, 241
California lady fern, see Athyrium filix-femina var. Cheilanthes intramarginalis, see Mildella intramarginalis,
californicum, 212 389
California maidenhair, see Adiantum jordanii, 166 Cheilanthes jonesii, see Argyrochosma jonesii, 189
California shield fern, see Dryopteris arguta, 300 Cheilanthes lanosa, 241
California sword fern, see Polystichum californicum, 457 Cheilanthes lendigera, 241
Callipteris esculenta, see Diplazium esculentum, 12, 69, Cheilanthes lindheimeri, 242
287 Cheilanthes newberryi, 242
Callipteris prolifera, see Diplazium proliferum, 287 Cheilanthes parvifolia, see Argyrochosma microphylla, 190
Calymmodon, 333 Cheilanthes pyramidalis subsp. arizonica, see Cheilanthes
Camptosorus, 47, 228 arizonica, 237
Camptosorus rhizophyllus, 94, 98, 101, 228 Cheilanthes siliquosa, see Aspidotis densa, 193
Camptosorus sibericus, 98, 228 Cheilanthes sinuata, see Astrolepis sinuata, 206
Campyloneurum, 98, 229, 439 Cheilanthes tomentosa, 243
Campyloneurum angustifolium, 229 Cheilanthes villosa, 243
Campyloneurum brevifolium, 230 Cheilanthes viridis, see Pellaea viridis, 416
Campyloneurum costatum, 230 Cheilanthes wrightii, 82, 243
Campyloneurum latum, see Campyloneurum brevifolium, Cheiroglossa, see Ophioglossum, 404
230 Chilean maidenhair, see Adiantum excisum, 164
Campyloneurum phyllitidis, 230 Chinese brake, see Pteris vittata, 480
Campyloneurum tenuipes, 231 Chinese lace fern, see Selaginella braunii, 497
Campyloneurum xalapense, 231 Christella, see Thelypteris, 517
Canary Island davallia, see Davallia canariensis, 264 Christella dentata, see Thelypteris dentata, 519
586 P N I

Christella parasitica, see Thelypteris parasitica, 522 Coniogramme ‘Variegata’, see Coniogramme japonica, 248
Christmas fern, see Polystichum acrostichoides, 455 Cooper’s tree fern, see Sphaeropteris cooperi, 91, 92, 511,
Cibotium, 110, 111, 244 color plates 2, 3
Cibotium barometz, 245 copper fern, see Bommeria, 225
Cibotium chamissoi, 245 cotton fern, see Cheilanthes newberryi, 242
Cibotium glaucum, 16, 93, 95, 111, 245, color plate 3 couplet fern, see Dennstaedtia bipinnata, 278
Cibotium menziesii, 111, 246 Coville’s lip fern, see Cheilanthes covillei, 238
Cibotium schiedei, 94, 100, 111, 246 crape fern, see Leptopteris superba, 351
cinnamon fern, see Osmunda cinnamomea, 407, color creeping star-hair fern, see Thelypteris reptans, 524
plate 8 creeping wood fern, see Thelypteris reptans, 524
Cionidium, see Tectaria, 515 Crepidomanes, see Trichomanes, 529
claw fern, see Onychium, 402 Crepidomanes auriculatum, see Trichomanes auriculatum,
cliff brake, see Aspidotis densa, 193; Cryptogramma 528
stelleri, 251; Pellaea, 412 crested lady fern, see Athyrium filix-femina ‘Cristatum’,
cliff fern, see Woodsia, 531 212
climbing fern, see Lygodium, 360 crested tongue fern, see Pyrrosia lingua ‘Cristata’, 485
climbing vine fern, see Microgramma heterophylla, 372 crested wood fern, see Dryopteris cristata, 305
Clinton’s wood fern, see Dryopteris clintoniana, 303 Cretan brake, see Pteris cretica, 474
cloak fern, see Notholaena, 399; see also Argyrochosma, crisped male fern, see Dryopteris affinis ‘Crispa’, 299
188; Astrolepis, 206 crowe maidenhair, see Adiantum raddianum
club moss, see Lycopodium, 354 ‘Croweanum’, 172
coastal leather fern, see Acrostichum aureum, 156 Crypsinus, see Selliguea, 507
coastal wood fern, see Dryopteris arguta, 300 Cryptogramma, 248
coast polypody, see Polypodium scouleri, 451 Cryptogramma acrostichoides, 249
Cochlidium, 333 Cryptogramma cascadensis, 249
Cochlidium serrulatum, 333 Cryptogramma crispa, 98, 250, color plate 23
coffee fern, see Pellaea andromedifolia, 412 Cryptogramma raddeana, 250
coin-spot tree fern, see Sphaeropteris cooperi, 91, 92, 511, Cryptogramma sitchensis, 250
color plates 2, 3 Cryptogramma stelleri, 251
Colysis, see Leptochilus, 349 Ctenitis, 251
Colysis ampla, see Leptochilus amplus, 349 Ctenitis ampla, see Ctenitis sloanei, 251
Colysis ellipitica, see Leptochilus ellipticus, 349 Ctenitis decomposita, see Lastreopsis microsora, 344
Colysis insignis, see Microsorum insigne, 381 Ctenitis pentangularis, see Lastreopsis microsora subsp.
Colysis pothifolia, see Leptochilus ellipticus, 350 pentangularis, 344
Colysis pteropus, see Microsorum pteropus, 384 Ctenitis sloanei, 251
Colysis sayeri, see Leptochilus amplus, 349 Ctenitis subincisa, see Megalastrum subincisum, 372
Colysis wrightii, see Leptochilus macrophyllus var. wrightii, Ctenitis sect. Subincisae, see Megalastrum, 371
350 Ctenitis submarginalis, 251
comb fern, see Pecluma, 410 Ctenopteris, 333
common golden-scaled male fern, see Dryopteris affinis, cup fern, see Dennstaedtia, 277
298 Currania, see Gymnocarpium, 334
common horsetail, see Equisetum arvense, 326 Currania oyamensis, see Gymnocarpium oyamense, 335
common male fern, see Dryopteris filix-mas, 308 Cyathea, 71, 252
common oak fern, see Gymnocarpium dryopteris, 335, Cyathea aramaganensis, see Sphaeropteris armaganensis,
color plate 29 510
common polypody, see Polypodium vulgare, 452 Cyathea arborea, 253
common staghorn, see Platycerium bifurcatum, 429, Cyathea australis, see Alsophila australis, 182
color plates 1, 5, 42 Cyathea brownii, see Sphaeropteris excelsa, 511
common sword fern, see Nephrolepis exaltata, 393; Cyathea cooperi, see Sphaeropteris cooperi, 91, 92, 511,
Polystichum munitum, 460 color plates 2, 3
common tassel fern, see Lycopodium phlegmaria, 358 Cyathea dealbata, see Alsophila tricolor, 182, color plate
Coniogramme, 247 11
Coniogramme fraxinea, see Coniogramme intermedia, 248 Cyathea medullaris, see Sphaeropteris medullaris, 512
Coniogramme intermedia, 248 Cyathea princeps, see Sphaeropteris horrida, 512
Coniogramme japonica, 248 Cyclopeltis, 253
 P N I 587

Cyclopeltis semicordata, 254 Davallia embolstegia, 265

Cyclophorus, see Pyrrosia, 480 Davallia epiphylla, 265
Cyclosorus, see Thelypteris, 517 Davallia falcinella, 266
Cyclosorus acuminatus, see Thelypteris acuminata, 518 Davallia fejeensis, 100, 267
Cyclosorus dentatus, see Thelypteris dentata, 519 Davallia griffithiana, 267
Cyclosorus guadalupensis, see Thelypteris guadalupensis, Davallia heterophylla, 268
520 Davallia hymenophylloides, 268
Cyclosorus parasiticus, see Thelypteris parasitica, 522 Davallia lorrainii, 270
Cyrtomium, 254 Davallia mariesii, 98, 270, color plate 5
Cyrtomium caryotideum, 255 Davallia mariesii var. mariesii, 271, color plate 5
Cyrtomium falcatum, 66, 82, 92, 93, 94, 96, 97, 100, Davallia mariesii var. stenolepis, 92, 94, 100, 271, color
255, color plate 24 plate 5
Cyrtomium fortunei, 256 Davallia parvula, 101, 272
Cyrtomium fortunei var. clivicola, 257 Davallia pentaphylla, 272
Cyrtomium fortunei var. fortunei, 257 Davallia portugal, see Davallia canariensis, 264
Cyrtomium fortunei var. intermedium, 257 Davallia pulchra, 272
Cyrtomium lonchitoides, 257 Davallia pycnocarpa, see Davallia pentaphylla, 272
Cyrtomium macrophyllum, 257 Davallia pyxidata, 272
Cyrtomium macrophyllum var. tukusicola, 257 Davallia repens, 273
Cyrtomium tukusicola, see Cyrtomium macrophyllum var. Davallia simplicifolia, see Davallia triphylla, 275
tukusicola, 257 Davallia solida, 274
Cyrtomium cultivars Davallia solida var. fejeensis, see Davallia fejeensis, 267
‘Butterfieldii’, see Cyrtomium falcatum, 256 Davallia solida var. pyxidata, see Davallia pyxidata, 272
‘Mayi’, see Cyrtomium falcatum, 256 Davallia stenolepis, see Davallia mariesii var. stenolepsis,
‘Rockfordianum’, see Cyrtomium falcatum, 256 271
Cystopteris, 258 Davallia tasmanii, 274
Cystopteris alpina, 259 Davallia trichomanoides, 275
Cystopteris bulbifera, 98, 259 Davallia trichomanoides var. lorrainii, see Davallia
Cystopteris dickieana, 259 lorrainii, 270
Cystopteris fragilis, 29, 98, 260 Davallia triphylla, 275
Cystopteris fragilis subsp. alpina, see Cystopteris alpina, Davallia tyermannii, 94, 98, 276
259 Davallia vestita, see Davallia repens, 273
Cystopteris fragilis subsp. dickieana, see Cystopteris Davallia cultivars
dickieana, 259 ‘Major’, see Davallia fejeensis, 267
Cystopteris fragilis var. mackayi, see Cystopteris tenuis, 262 ‘Plumosa’, see Davallia fejeensis, 267
Cystopteris fragilis var. protrusa, see Cystopteris protrusa, ‘Ruffled Ornata’, see Davallia solida, 274
260 Davallodes, 276
Cystopteris fragilis subsp. tenuifolia, see Cystopteris Davallodes hirsutum, 277
reevesiana, 261 deer fern, see Blechnum spicant, 221
Cystopteris montana, 260 delta maidenhair, see Adiantum raddianum, 172
Cystopteris protrusa, 260 Dennstaedtia, 97, 277
Cystopteris reevesiana, 261 Dennstaedtia bipinnata, 278
Cystopteris regia, see Cystopteris alpina, 259 Dennstaedtia cicutaria, 278
Cystopteris tennesseensis, 261 Dennstaedtia davallioides, 279
Cystopteris tenuis, 262 Dennstaedtia punctilobula, 92, 93, 94, 279
Cystopteris ‘Crispa’, see Cystopteris bulbifera, 259 Deparia, 208, 279
Deparia acrostichoides, 29, 91, 96, 280
Dallas fern, see Nephrolepis exaltata ‘Dallas’, 395 Deparia japonica, 280
Davallia, 68, 97, 98, 103, 262 Deparia petersenii, 281
Davallia bullata, see Davallia mariesii, 270, color plate 5 diamond maidenhair, see Adiantum trapeziforme, 175
Davallia canariensis, 264 Diblemma, see Microsorum, 377
Davallia denticulata, 264 Diblemma tenuilore, see Microsorum samarense, 385
Davallia denticulata var. elata, see Davallia epiphylla, 265 Dickie’s bladder fern, see Cystopteris dickieana, 259
Davallia divaricata, 265 Dicksonia, 281
Davallia elegans, see Davallia denticulata, 264 Dicksonia antarctica, 92, 95, 111, 282, color plate 3
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Dicksonia berteriana, 283 Drynaria descensa, 294

Dicksonia fibrosa, 95, 282 Drynaria morbillosa, 178
Dicksonia gigantea, see Dicksonia sellowiana, 283 Drynaria propinqua, 294
Dicksonia sellowiana, 283 Drynaria quercifolia, 295
Dicksonia squarrosa, 284 Drynaria rigidula, 295, color plate 25
Dicranopteris, 330 Drynaria sparsisora, 296
Dicranopteris flexuosa, 331 Drynaria ‘Whitei’, see Drynaria rigidula, 296, color plate
Dicranopteris linearis, 331 25
Dictymia, 284, 439 Dryoathyrium, see Deparia, 279
Dictymia brownii, 284 Dryopteris, 296
Didymochlaena, 284 Dryopteris abbreviata, see Dryopteris affinis, 298
Didymochlaena truncatula, 97, 285 Dryopteris aemula, 298
Diphasiastrum complanatum, see Lycopodium Dryopteris affinis, 82, 92, 298
complanatum, 357 Dryopteris affinis subsp. borreri, 299
Diplazium, 208, 285 Dryopteris affinis subsp. cambriensis, 299
Diplazium acrostichoides, see Deparia acrostichoides, 280 Dryopteris amurensis, 299
Diplazium australe, 286 Dryopteris arguta, 300
Diplazium dilatatum, 286 Dryopteris atrata, 305
Diplazium esculentum, 12, 69, 287 Dryopteris ×australis, 300
Diplazium lanceum, see Diplazium subsinuatum, 288 Dryopteris austriaca, see Dryopteris expansa, 308
Diplazium lonchophyllum, 287 Dryopteris bissetiana, 300
Diplazium maximum, 286 Dryopteris ×boottii, 300
Diplazium proliferum, 287 Dryopteris campyloptera, 301
Diplazium pycnocarpon, 29, 96, 288 Dryopteris carthusiana, 302
Diplazium subsinuatum, 97, 288 Dryopteris caucasica, 302
Diplazium tomitaroanum, 101, 289 Dryopteris celsa, 302
Diplopterygium, 330 Dryopteris championii, 303
disk staghorn, see Platycerium coronarium, 431 Dryopteris clintoniana, 94, 303
dissected grape fern, see Botrychium dissectum, 227 Dryopteris complexa, 303
divided soft shield fern, see Polystichum setiferum Dryopteris coreano-montana, see Dryopteris sichotensis, 317
‘Plumoso-Divisilobum’, 465, color plate 46 Dryopteris crassirhizoma, 304
dixie wood fern, see Dryopteris ×australis, 300 Dryopteris cristata, 305
Doodia, 289 Dryopteris cycadina, 66, 305
Doodia aspera, 82, 289 Dryopteris cystolepidota, 305
Doodia media, 82, 93, 290 Dryopteris decipiens, 306
Doodia media subsp. australis, 290 Dryopteris decipiens var. diplazioides, 306
Doodia media subsp. media, 290 Dryopteris dilatata, 92, 306
Doodia mollis, 290 Dryopteris disjuncta, see Gymnocarpium disjunctum, 335
Doryopteris, 111, 291 Dryopteris erythrosora, 91, 93, 95, 96, 97, 307, color
Doryopteris concolor, 292 plate 26
Doryopteris ludens, 292 Dryopteris erythrosora f. prolifica, 308
Doryopteris nobilis, 65, 292 Dryopteris erythrosora f. viridosora, 308
Doryopteris palmata, 292 Dryopteris expansa, 94, 308
Doryopteris pedata var. palmata, see Doryopteris palmata, Dryopteris filix-mas, 92, 94, 308
292 Dryopteris formosana, 310
Doryopteris sagittifolia, 292 Dryopteris fragrans, 310
double maindenhair, see Adiantum raddianum ‘Pacottii’, Dryopteris fuscipes, 311
173 Dryopteris goeringiana, 318
Douglas spike moss, see Selaginella douglassii, 498 Dryopteris goldiana, 91, 92, 311
downy thelypteris, see Thelypteris dentata, 519 Dryopteris hirtipes, 305
dragon-tail fern, see Asplenium ×ebenoides, 198 Dryopteris hondoensis, 311
Drymoglossum, see Pyrrosia, 480 Dryopteris hondoensis f. rubisora, 311
Drymoglossum piloselloides, see Pyrrosia piloselloides, 486 Dryopteris intermedia, 97, 311
Drynaria, 95, 292 Dryopteris kuratae, 312
Drynaria bonii, 294 Dryopteris lacera, 313
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Dryopteris lepidopoda, 313 ‘Linearis Polydactyla’, see Dryopteris filix-mas, 310

Dryopteris ludoviciana, 93, 313 ‘Paleaceo Crispa’, see Dryopteris affinis ‘Crispa’, 299
Dryopteris marginalis, 91, 92, 314 ‘Polydactyla’, see Dryopteris affinis, 299
Dryopteris mindshelkensis, 314 ‘Polydactyla Dadd’s’, see Dryopteris affinis
Dryopteris nipponensis, see Dryopteris cystolepidota, 305 ‘Polydactyla’, 299
Dryopteris noveboracensis, see Thelypteris noveboracensis, ‘Ramo Cristata’, see Dryopteris filix-mas, 310
521 ‘Recurved Form’, see Dryopteris dilatata, 307
Dryopteris oreades, 315 ‘Revolvens’, see Dryopteris affinis, 299
Dryopteris paleacea, see Dryopteris wallichiana, 320, color ‘Robusta’, see Dryopteris complexa, 304
plate 27 ‘Stableri’, see Dryopteris affinis, 299
Dryopteris parallelogramma, see Dryopteris wallichiana, ‘Stableri Crisped’, see Dryopteris affinis, 299
320, color plate 27 ‘The King’, see Dryopteris affinis ‘Cristata’, 299
Dryopteris polylepis, 315 ‘Undulata-Robusta’, see Dryopteris complexa, 304
Dryopteris pseudo-filix-mas, 315 Dudley’s holly fern, see Polystichum dudleyi, 458
Dryopteris pseudomas, see Dryopteris affinis, 298 Dudley’s sword fern, see Polystichum dudleyi, 458
Dryopteris purpurella, 316 dwarf holly fern, see Cyrtomium caryotideum, 255
Dryopteris pycnopteroides, 313 dwarf lady fern, see Athyrium filix-femina
Dryopteris remota, 316 ‘Minutissimum’, 212
Dryopteris sacrosancta, 316 dwarf mother fern, see Asplenium daucifolium, 197
Dryopteris setigera, 364 dwarf scouring rush, see Equisetum scirpoides, 329
Dryopteris sichotensis, 317 dwarf soft shield fern, see Polystichum setiferum
Dryopteris sieboldii, 317 ‘Congestum’, 464
Dryopteris spinulosa, see Dryopteris carthusiana, 302 dwarf water clover, see Marsilea angustifolia, 366
Dryopteris spinulosa var. americana, see Dryopteris dwarf western sword fern, see Polystichum imbricans
campyloptera, 301 subsp. imbricans, 458
Dryopteris spinulosa var. intermedia, see Dryopteris
intermedia, 311 eared spleenwort, see Asplenium auritum, 69, 195
Dryopteris stewartii, 318 eastern bracken, see Pteridium aquilinum var.
Dryopteris sublacera, 318 latiusculum, 471
Dryopteris submontana, see Dryopteris mindshelkensis, 314 eastern spreading wood fern, see Dryopteris campyloptera,
Dryopteris tokyoensis, 94, 318 301
Dryopteris uniformis, 319 eastern wood fern, see Dryopteris marginalis, 314
Dryopteris varia, 320 East Indian holly fern, see Arachniodes aristata, 187
Dryopteris villarii subsp. submontana, see Dryopteris East Indian polypody, see Microsorum scolopendria, 387
mindshelkensis, 314 Eaton’s lip fern, see Cheilanthes eatonii, 239
Dryopteris wallichiana, 320, color plate 27 ebony spleenwort, see Asplenium platyneuron, 202
Dryopteris cultivars Ecuadorian fern, see Nephrolepis cordifolia ‘Ecuadorian
‘Barnesii’, see Dryopteris filix-mas, 310 Fern’, 392
‘Congesta’, see Dryopteris affinis, 299 Edanyoa, see Bolbitis, 222
‘Congesta Cristata’, see Dryopteris affinis, 299 Edanyoa difformis, see Bolbitis heteroclita, 223
‘Crispa’, see Dryopteris affinis, 299 Elaphoglossum, 95, 320
‘Crispa Cristata’, see Dryopteris filix-mas, 310 Elaphoglossum apodum, 321
‘Crispa Gracilis’, see Dryopteris affinis, 299 Elaphoglossum crinitum, 322
‘Crispa Whiteside’, see Dryopteris dilatata, 307 Elaphoglossum decoratum, 322
‘Cristata’, see Dryopteris affinis, 299; Dryopteris Elaphoglossum erinaceum, 321
dilatata, 307 Dryopteris filix-mas, 310; Dryopteris Elaphoglossum herminieri, 322
uniformis, 319 Elaphoglossum latifolium, 325
‘Cristata Angustata’, see Dryopteris affinis, 299 Elaphoglossum luridum, 323
‘Cristata the King’, see Dryopteris affinis ‘Cristata’, Elaphoglossum maxonii, 323
299 Elaphoglossum metallicum, 323
‘Grandiceps’, see Dryopteris dilatata, 307; Dryopteris Elaphoglossum muelleri, 325
filix-mas, 310 Elaphoglossum peltatum, 324
‘Jimmy Dyce’, see Dryopteris dilatata, 307 Elaphoglossum sartorii, 324
‘Lepidota Cristata’, see Dryopteris dilatata, 307 Elaphoglossum schomburgkii, see Elaphoglossum luridum,
‘Linearis Congesta’, see Dryopteris filix-mas, 310 323
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Elaphoglossum vestitum, 325 fragile bladder fern, see Cystopteris fragilis, 260
elephant-ear fern, see Elaphoglossum crinitum, 322 fragile rock brake, see Cryptogramma stelleri, 251
elkhorn fern, see Platycerium, 427 fragrant fern, see Microsorum scandens, 386
Equisetum, 74, 93, 115, 326 Fritz Luth maidenhair, see Adiantum raddianum ‘Fritz
Equisetum arvense, 326 Luth’, 172
Equisetum diffusum, 327
Equisetum ×ferrissii, 327 ghost fern, see Athyrium niponicum ‘Pictum’, 212
Equisetum hyemale, 328 giant chain fern, see Woodwardia fimbriata, 537
Equisetum laevigatum, 328 giant fern, see Acrostichum danaeifolium, 157
Equisetum scirpoides, 329 giant horsetail, see Equisetum telmateia, 329
Equisetum telmateia, 329 giant maidenhair, see Adiantum polyphyllum, 171
Equisetum variegatum, 329 giant staghorn, see Platycerium superbum, 435, color
erect sword fern, see Nephrolepis cordifolia, 391, color plate 43
plate 35 giant sword fern, see Nephrolepis biserrata, 390
European chain fern, see Woodwardia radicans, 537, giant wood fern, see Dryopteris goldiana, 311
color plate 50 glade fern, see Deparia acrostichoides, 280; Diplazium
European lady fern, see Athyrium filix-femina var. filix- pycnocarpon, 288
femina, 212 glandular wood fern, see Dryopteris intermedia, 311;
European ostrich fern, see Matteuccia struthiopteris var. Thelypteris resinifera, 524
struthiopteris, 371 Gleichenella, 330
European parsley fern, see Cryptogramma crispa, 250, Gleichenia, 330
color plate 23 Gleichenia dicarpa, 330
evergreen wood fern, see Dryopteris intermedia, 311 Gleichenia microphylla, 330
Gleichenia rupestris, 330
fairy moss, see Azolla, 213 Gleicheniaceae, 330
fancy fern, see Dryopteris intermedia, 311 glossy spleenwort, see Asplenium oblongifolium, 200
fan maidenhair, see Adiantum tenerum, 173 goldback fern, see Pentagramma, 417; Pityrogramma,
Farley maidenhair, see Adiantum tenerum ‘Farleyense’, 424
173 golden polypody, see Phlebodium, 419; Phlebodium
felt fern, see Pyrrosia, 480 aureum, 420
Fendler’s lip fern, see Cheilanthes fendleri, 240, color Goldie’s fern, see Dryopteris goldiana, 311
plate 22 Goniophlebium, 98, 331, 439
ferny azolla, see Azolla pinnata, 214, color plate 16 Goniophlebium amoenum, see Polypodium amoenum, 441
Ferriss’s scouring rush, see Equisetum ×ferrissii, 327 Goniophlebium ×ekstrandii, 331
fibrous dicksonia, see Dicksonia fibrosa, 282 Goniophlebium formosanum, see Polypodium formosanum,
field horsetail, see Equisetum arvense, 326 444, color plate 44
Fiji davallia, see Davallia fejeensis, 267 Goniophlebium percussum, 332
filmy fern, see Hymenophyllum, 339; Trichomanes, 528 Goniophlebium persicifolium, 332
filmy maidenhair, see Adiantum diaphanum, 163 Goniophlebium subauriculatum, 94, 333, color plate 28
fir club moss, see Lycopodium selago, 360 Goniophlebium verrucosum, see Goniophlebium percussum,
fish-bone fern, see Nephrolepis, 389 332
fishtail sword fern, see Nephrolepis falcata ‘Furcans’, 395 Goniophlebium cultivars
five-finger fern, see Adiantum pedatum, 168 ‘Knightiae’, see Goniophlebium subauriculatum, 333,
flat-branch club moss, see Lycopodium complanatum, 357 color plate 28
flat-branched tree club moss, see Lycopodium obscurum, ‘Nola’, see Goniophlebium ×ekstrandii, 331
358 Goniopteris, see Thelypteris, 517
flexuous cliff brake, see Pellaea ovata, 415 Goniopteris poiteana, see Thelypteris poiteana, 523
Florida tree fern, see Ctenitis sloanei, 251 Goniopteris reptans, see Thelypteris reptans, 524
flowering fern, see Anemia, 182; Osmunda regalis, 408, Grammitidaceae, 333
color plate 6 Grammitis, 74, 333
fluffy ruffles, see Nephrolepis exaltata ‘Fluffy Ruffles’, 395 grape fern, see Botrychium, 226
forest brake, see Pteris altissima, 473 grass fern, see Vittaria lineata, 530
forked spleenwort, see Asplenium septentrionale, 204 green cliff brake, see Pellaea viridis, 416
forking fern family, see Gleicheniaceae, 330 green petticoats, see Adiantum capillus-veneris
four-leaved maidenhair, see Adiantum tetraphyllum, 174 ‘Imbricatum’, 162
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green spleenwort, see Asplenium trichomanes-ramosum, Humata falcinella, see Davallia falcinella, 266
206 Humata griffithiana, see Davallia griffithiana, 267
green staghorn, see Platycerium hillii, 432 Humata heterophylla, see Davallia heterophylla, 268
grid-scale maiden fern, see Thelypteris patens, 523 Humata parvula, see Davallia parvula, 272
ground cedar, see Lycopodium complanatum, 357 Humata repens, see Davallia repens, 273
ground fern, see Hypolepis, 341 Humata tyermannii, see Davallia tyermannii, 276
ground pine, see Lycopodium, 113, 354; Lycopodium Humata vestita, see Davallia repens, 273
obscurum, 358 Huperzia carinata, see Lycopodium carinatum, 356
Gymnocarpium, 334 Huperzia filiformis, see Lycopodium filiforme, 357
Gymnocarpium disjunctum, 335 Huperzia lucidula, see Lycopodium lucidulum, 358
Gymnocarpium dryopteris, 92, 96, 97, 98, 335, color Huperzia nummulariifolium, see Lycopodium
plate 29 nummulariifolium, 358
Gymnocarpium oyamense, 335 Huperzia phlegmaria, see Lycopodium phlegmaria, 358
Gymnocarpium robertianum, 94, 335 Huperzia phlegmarioides, see Lycopodium phlegmarioides,
Gymnocarpium ‘Plumosum’, see Gymnocarpium 359
dryopteris, 335, color plate 29 Huperzia polytrichoides, see Lycopodium filiforme, 357
Gymnopteris, see Hemionitis, 336 Huperzia selago, see Lycopodium selago, 360
Huperzia squarrosa, see Lycopodium squarrosum, 360
hacksaw fern, see Doodia, 289 Hymenodium, see Elaphoglossum, 320
hairy lip fern, see Cheilanthes lanosa, 241 Hymenodium crinitum, see Elaphoglossum crinitum, 322
halberd fern, see Tectaria heracleifolia, 515 Hymenophyllum, 339
hammock fern, see Blechnum occidentale, 220 Hymenophyllum demissum, 340
hand fern, see Doryopteris palmata, 292; Ophioglossum, Hymenophyllum polyanthos, 340
404 Hypoderris, 340
Haplopteris, 530 Hypoderris brownii, 341
Haplopteris elongata, see Vittaria elongata, 530 Hypolepis, 94, 97, 341
hapu, see Cibotium glaucum, 245 Hypolepis millefolium, 342
hard fern, see Paesia scaberula, 409 Hypolepis punctata, 342
hard male fern, see Dryopteris affinis, 298 Hypolepis repens, 342
hard shield fern, see Polystichum aculeatum, 456 Hypolepis tenuifolia, 82, 342
hare’s-foot fern, see Davallia canariensis, 264;
Phlebodium aureum, 420 imbricate sword fern, see Polystichum imbricans, 458
Hartford fern, see Lygodium palmatum, 363 Indian’s dream, see Aspidotis densa, 193
hart’s-tongue fern, see Phyllitis scolopendrium, 422, color Indian staghorn, see Platycerium wallichii, 436
plate 39 inland leather fern, see Acrostichum danaeifolium, 157
Hawaiian tree fern, see Cibotium glaucum, 245 intermediate polypody, see Polypodium interjectum, 446
hay-scented fern, see Dennstaedtia punctilobula, 279 interrupted fern, see Osmunda claytoniana, 407, color
hay-scented wood fern, see Dryopteris aemula, 298 plate 37
Hemigramma, see Tectaria, 513
Hemigramma latifolia, see Tectaria hilocarpa, 516 Japanese ball fern, see Davallia mariesii var. mariesii, 271
Hemionitis, 101, 336 Japanese bird’s-nest fern, see Asplenium antiquum, 195
Hemionitis arifolia, 337, color plate 30 Japanese claw fern, see Onychium japonicum, 402
Hemionitis palmata, 338 Japanese climbing fern, see Lygodium japonicum, 362,
Hemionitis pinnatifida, 338 color plate 31
Hemitelia, see Cyathea, 252 Japanese felt fern, see Pyrrosia lingua, 484
Himalayan holly fern, see Polystichum lentum, 459 Japanese osmunda, see Osmunda japonica, 408
Himalayan maidenhair, see Adiantum venustum, 175 Japanese painted fern, see Athyrium niponicum ‘Pictum’,
Histiopteris, 338 212, color plate 14
Histiopteris incisa, 339 Japanese sword fern, see Polystichum polyblepharum, 462
holly fern, see Cyrtomium, 254; Polystichum, 454 Japanese wood fern, see Dryopteris sieboldii, 317
horseshoe fern, see Marattia salicina, 365 Java staghorn, see Platycerium willinckii, 437
horsetail, see Equisetum, 325 jointed polypody, see Goniophlebium subauriculatum,
house holly fern, see Cyrtomium falcatum, 255, color 333, color plate 28
plate 24 joint fern, see Arthropteris, 190
Humata, see Davallia, 262 Jones’s cloak fern, see Argyrochosma jonesii, 189
592 P N I

kangaroo fern, see Microsorum pustulatum, 385 Leptochilus amplus, 349

keeled tassel fern, see Lycopodium carinatum, 356 Leptochilus buergerianum, see Microsorum superficiale,
Killarney bristle fern, see Trichomanes speciosum, 529 388
Kimberly Queen, see Nephrolepis cordifolia ‘Kimberly Leptochilus ellipticus, 350
Queen’, 392 Leptochilus macrophyllus var. wrightii, 350
king fern, see Marattia salicina, 365 Leptochilus sayeri, see Leptochilus amplus, 349
Knight’s polypody, see Goniophlebium subauriculatum Leptochilus ‘Monstrifera’, see Leptochilus macrophyllus
‘Knightiae’, 333, color plate 28 var. wrightii, 350
Korean rock fern, see Polystichum tsus-simense var. Leptopteris, 350
mayebarae, 469 Leptopteris hymenophylloides, 351
Kruckeberg’s holly fern, see Polystichum kruckebergii, 458 Leptopteris superba, 112, 351
Leucostegia, 351
lace fern, see Aspidotis, 192; Cheilanthes gracillima, 240; Leucostegia hymenophylloides, see Davallia
Microlepia strigosa, 377; Nephrolepis exaltata hymenophylloides, 268
‘Smithii’ 395; Odontosoria chinensis, 400; Paesia Leucostegia immersa, 352
scaberula, 409 Leucostegia pallida, 352
lacerate pyrrosia, see Pyrrosia lingua ‘Monstrifera’, 485 licorice fern, see Polypodium glycyrrhiza, 44
lacy bracken, see Pteridium aquilinum var. caudatum, limestone oak fern, see Gymnocarpium robertianum, 336
471 limestone wood fern, see Dryopteris mindshelkensis, 314
lacy ground fern, see Dennstaedtia davallioides, 279 Lindheimer’s lip fern, see Cheilanthes lindheimeri, 241
ladder brake, see Pteris longifolia, 476 Lindsaea, 352
lady fern, see Athyrium filix-femina, 211 lip fern, see Cheilanthes, 235
large-leaved maidenhair, see Adiantum macrophyllum, Llavea, 353
167 Llavea cordifolia, 353
lasagna fern, see Asplenium nidus var. plicatum, 200 lobed spleenwort, see Asplenium pinnatifidum, 202
Lastrea, see Thelypteris, 517 log fern, see Dryopteris celsa, 302
Lastreopsis, 342 Lomaria, see Blechnum, 214
Lastreopsis acuminata, 343 Lonchitis, 353
Lastreopsis decomposita, 344 Lonchitis hirsuta, 354
Lastreopsis effusa, 343 long beech fern, see Phegopteris connectilis, 418
Lastreopsis microsora, 344 long-eared holly fern, see Polystichum neolobatum, 461
Lastreopsis microsora subsp. microsora, 344 Lorinseria, see Woodwardia, 535
Lastreopsis microsora subsp. pentangularis, 344 Lorinseria areolata, see Woodwardia areolata, 537
Lastreopsis shepherdii, see Lastreopsis acuminata, 343 lowland bladder fern, see Cystopteris protrusa, 260
lattice-vein fern, see Thelypteris reticulata, 525 Lunathyrium, see Deparia, 279
layered tassel fern, see Lycopodium phlegmarioides, 359 Lycopodiella cernuua, see Lycopodium cernuum, 356
leather fern, see Acrostichum, 156; Rumohra Lycopodium, 113-114, 354
adiantiformis, 491 Lycopodium carinatum, 356
leathery polypody, see Polypodium scouleri, 451 Lycopodium cernuum, 356
Lecanopteris, 345, 439 Lycopodium clavatum, 356
Lecanopteris carnosa, 345 Lycopodium complanatum, 357
Lecanopteris crustacea, 346 Lycopodium dendroideum, 357
Lecanopteris sinuosa, 346 Lycopodium digitatum, 357
Lellingeria, 333 Lycopodium filiforme, 357
Lemmaphyllum, 346 Lycopodium lucidulum, 358
Lemmaphyllum carnosum, 348 Lycopodium nummulariifolium, 358
Lemmaphyllum microphyllum, 98, 101, 347 Lycopodium obscurum, 358
Lemmaphyllum subrostratum, 347 Lycopodium obscurum var. dendroideum, see Lycopodium
lemon buttons, see Nephrolepis cordifolia ‘Lemon dendroideum, 357
Buttons’, 98, 392 Lycopodium phlegmaria, 113, 114, 358
lemon-scented fern, see Thelypteris limbosperma, 69, 520 Lycopodium phlegmarioides, 113, 359
Lepisorus, 348, 439 Lycopodium polytrichoides, see Lycopodium filiforme, 357
Lepisorus longifolius, 348 Lycopodium selago, 113, 360
Lepisorus thunbergianus, 348 Lycopodium squarrosum, 360
Leptochilus, 349, 439 Lygodium, 360
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Lygodium circinnatum, 361 Megalastrum subincisum, 372

Lygodium flexuosum, 362 Melpomene, 333
Lygodium japonicum, 93, 362, color plate 31 Meniscium, see Thelypteris, 517
Lygodium microphyllum, 362 Meniscium reticulatum, see Thelypteris reticulata, 525
Lygodium palmatum, 363 Meniscium serratum, see Thelypteris serrata, 525
Lygodium scandens, see Lygodium microphyllum, 362 Merinthosorus, see Aglaomorpha, 176
Merinthosorus drynarioides, see Aglaomorpha drynarioides,
macho fern, see Nephrolepis falcata, 395 178
Mackay’s brittle fern, see Cystopteris tenuis, 262 Mexican chain fern, see Woodwardia spinulosa, 538
Macrothelypteris, 363 Mexican male fern, see Dryopteris pseudo-filix-mas, 315
Macrothelypteris torresiana, 82, 94, 363 Mexican tree fern, see Cibotium schiedei, 246
mahogany fern, see Didymochlaena, 284 Microgramma, 372, 439
maiden fern, see Thelypteris, 517 Microgramma ciliata, see Microgramma reptans, 374
maidenhair fern, see Adiantum, 159 Microgramma heterophylla, 372
maidenhair spleenwort, see Asplenium trichomanes, 205, Microgramma lycopodioides, 373
color plate 13 Microgramma megalophylla, 373
Makino’s holly fern, see Polystichum makinoi, 460 Microgramma nitida, 373
male fern, see Dryopteris, 296 Microgramma percussa, 373
man fern, see Cibotium menziesii, 246 Microgramma piloselloides, 373
Manchurian woodsia, see Woodsia manchuriensis, 533 Microgramma reptans, 373
Marattia, 71, 95, 364 Microgramma squamulosa, 375
Marattia fraxinea, 365 Microgramma vacciniifolia, 101, 375
Marattia salicina, 365 Microlepia, 375
marginal shield fern, see Dryopteris marginalis, 314 Microlepia firma, 377
Mariana Island tree fern, see Sphaeropteris armaganensis, Microlepia platyphylla, 82, 93, 95, 376
510 Microlepia pyramidata, see Microlepia speluncae
Mariana maiden fern, see Macrothelypteris torresiana, 363 ‘Corymbifera’, 376
marsh fern, see Thelypteris palustris, 522 Microlepia speluncae, 376
Marsilea, 74, 93, 365 Microlepia strigosa, 82, 92, 93, 94, 97, 377
Marsilea angustifolia, 366 Microlepia strigosa f. macfaddeniae, 377
Marsilea crenata, 369 Microlepia substrigosa, 377
Marsilea drummondii, 367 Microlepia ‘Corymbifera’, see Microlepia speluncae, 376
Marsilea macrocarpa, 367 Micropolypodium, 333
Marsilea macropoda, 367 Microsorum, 93, 377, 439
Marsilea minuta, 368 Microsorum alternifolium, see Microsorum
Marsilea mutica, 368, color plate 32 membranifolium, 382
Marsilea quadrifolia, 368 Microsorum buergerianum, see Microsorum superficiale,
Marsilea schelpeana, 368 388
Marsilea vestita, 369 Microsorum commutatum, 380
Massachusetts fern, see Thelypteris simulata, 526 Microsorum dilatatum, see Microsorum insigne, 381
Matteuccia, 369 Microsorum diversifolium, see Microsorum pustulatum,
Matteuccia orientalis, 82, 370 385
Matteuccia pensylvanica, see Matteuccia struthiopteris, Microsorum fortunei, 380
371, color plate 33 Microsorum grossum, 91, 100, 380
Matteuccia struthiopteris, 90, 91, 93, 94, 96, 97, 371, Microsorum hancockii, see Microsorum insigne, 381
color plate 33 Microsorum insigne, 381
Matteuccia struthiopteris var. pensylvanica, 371 Microsorum irioides, see Microsorum punctatum, 385,
Matteuccia struthiopteris var. struthiopteris, 371 color plate 34
Mauritius spleenwort, see Asplenium daucifolium, 197 Microsorum linguiforme, 382
may fern, see Cyrtomium falcatum ‘Mayi’, 256 Microsorum longissimum, 382
meadow spike moss, see Selaginella apoda, 497 Microsorum lucidum, 382
Mecodium, see Hymenophyllum, 339 Microsorum membranifolium, 382
Mecodium demissum, see Hymenophyllum demissum, 340 Microsorum musifolium, 383
Mecodium polyanthos, see Hymenophyllum polyanthos, 340 Microsorum nigrescens, see Microsorum membranifolium,
Megalastrum, 371 382
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Microsorum papuanum, 384 Nephrolepis acutifolia, 390

Microsorum polycarpon, see Microsorum punctatum, 384, Nephrolepis auriculata, see Nephrolepis cordifolia, 391,
color plate 34 color plate 35
Microsorum pteropus, 93, 384 Nephrolepis biserrata, 390
Microsorum punctatum, 91, 97, 100, 384, color plate 34 Nephrolepis cordifolia, 92, 93, 94, 97, 391, color plate
Microsorum pustulatum, 385 35
Microsorum samarense, 385 Nephrolepis ensifolia, 395
Microsorum scandens, 386, color plate 5 Nephrolepis exaltata, 97, 100, 393, color plate 5
Microsorum scolopendria, 387 Nephrolepis exaltata var. bostoniensis, see Nephrolepis
Microsorum steerei, 387 exaltata ‘Bostoniensis’, 395
Microsorum superficiale, 388 Nephrolepis falcata, 91, 395
Microsorum viellardii, 94, 388 Nephrolepis hirsutula, 395
Microsorum vitiense, see Microsorum commutatum, 380 Nephrolepis lauderbachii, 396
Microsorum whiteheadii, 384 Nephrolepis multiflora, 397
Microsorum cultivars Nephrolepis obliterata, 392
‘Climbing-bird’s-nest fern’, see Microsorum Nephrolepis pendula, 397
punctatum, 385 Nephrolepis tuberosa, see Nephrolepis cordifolia, 391, color
‘Cristatum’, see Microsorum punctatum ‘Ramosum’, plate 35
385 Nephrolepis cultivars
‘Grandiceps’, see Microsorum punctatum, 97, 385 ‘Aurea’, see Nephrolepis exaltata ‘Golden Boston’, 395
‘Ramosum’, see Microsorum punctatum, 385, color ‘Bostoniensis’, see Nephrolepis exaltata, 395
plate 34 ‘Can-can’, see Nephrolepis cordifolia ‘Petticoats’, 392
‘Southern Cross’, see Microsorum commutatum, 380 ‘Dallas’, see Nephrolepis exaltata, 395
Mildella, 236, 389 ‘Dennis Petticoats’, see Nephrolepis cordifolia
Mildella intramarginalis, 389 ‘Petticoats’, 392
mini-ruffles, see Nephrolepis exaltata ‘Mini-ruffles’, 395 ‘Duffii’, see Nephrolepis cordifolia, 392
monkey-tail fern, see Sphaeropteris horrida, 512 ‘Ecuadorian Fern’, see Nephrolepis cordifolia, 392
mosquito fern, see Azolla, 12, 213 ‘Elsevier’, see Nephrolepis exaltata, 101, 395
mother fern, see Asplenium bulbiferum, 196 ‘Florist Fantasy’, see Nephrolepis multiflora, 397
mother shield fern, see Polystichum proliferum, 462 ‘Fluffy Ruffles’, see Nephrolepis exaltata, 395
mountain bladder fern, see Cystopteris montana, 260 ‘Furcans’, see Nephrolepis falcata. 395
mountain club moss, see Lycopodium selago, 360 ‘Golden Boston’, see Nephrolepis exaltata, 395
mountain male fern, see Dryopteris oreades, 315 ‘Gracillima’, see Nephrolepis exaltata, 395
mountain wood fern, see Dryopteris campyloptera, 301 ‘Gretnae’, see Nephrolepis exaltata, 395
mountain woodsia, see Woodsia scopulina, 535 ‘Irish Lace’, see Nephrolepis exaltata ‘Gracillima’, 395
mule’s-foot fern, see Angiopteris, 184 ‘Kimberly Queen’, see Nephrolepis cordifolia, 392
Myrmecophila, see Lecanopteris, 345 ‘Lemon Buttons’, see Nephrolepis cordifolia, 98, 392
Myrmecopteris, see Lecanopteris, 345 ‘Massii’, see Nephrolepis exaltata, 395
Myrmecopteris crustacea, see Lecanopteris crustacea, 346 ‘Mildred Murray’, see Nephrolepis cordifolia
Myrmecopteris sinuosa, see Lecanopteris sinuosa, 346 ‘Tesselata’, 393
‘Mini-ruffles’, see Nephrolepis exaltata, 101, 395
nardoo, see Marsilea, 365 ‘Norwoodii’, see Nephrolepis exaltata, 395
narrow beech fern, see Phegopteris connectilis, 418 ‘Petticoats’, see Nephrolepis cordifolia, 392
narrow buckler fern, see Dryopteris carthusiana, 302 ‘Plumosa’, see Nephrolepis cordifolia ‘Tesselata’, 393
narrow glade fern, see Diplazium pycnocarpon, 288 ‘Pom-pom’, see Nephrolepis cordifolia ‘Petticoats’, 392
narrow-leaved chain fern, see Woodwardia areolata, 537 ‘Ram’s Horn’, see Nephrolepis falcata, 395
narrow-leaved nardoo, see Marsilea angustifolia, 366 ‘Rooseveltii’, see Nephrolepis exaltata, 395
narrow-leaved strap fern, see Campyloneurum ‘Sassy’, see Nephrolepis exaltata, 395
angustifolium, 229 ‘Smithii’, see Nephrolepis exaltata, 395
necklace fern, see Asplenium flabellifolium, 198 ‘Splendida’, see Nephrolepis exaltata, 395
Neocheiropteris superficialis, see Microsorum superficiale, ‘Superba’, see Nephrolepis hirsutula, 395
388 ‘Tesselata’, see Nephrolepis cordifolia, 392
Nepal holly fern, see Polystichum nepalense, 461 ‘Trevillian’, see Nephrolepis exaltata, 395
Nephelea, see Alsophila, 181 ‘Verona’, see Nephrolepis exaltata, 395
Nephrolepis, 98, 107–108, 389 ‘Wanamaka’, see Nephrolepis exaltata, 395
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net-veined chain fern, see Woodwardia areolata, 537 Ophioglossum palmatum, 404
Newberry’s lip fern, see Cheilanthes newberryi, 242 Ophioglossum pendulum, 405, color plate 36
New York fern, see Thelypteris noveboracensis, 521 Ophioglossum petiolatum, 405
New Zealand cliff brake, see Pellaea rotundifolia, 416 Ophioglossum vulgatum, 406
New Zealand dicksonia, see Dicksonia squarrosa, 284 Oregon spike moss, see Selaginella oregana, 502
Niphidium, 398, 439 Oregon woodsia, see Woodsia oregana, 534
Niphidium albopunctatissimum, 399 Oreopteris, see Thelypteris, 517
Niphidium crassifolium, 398 Oreopteris limbosperma, see Thelypteris limbosperma, 69,
Niphobolus, see Pyrrosia, 480 520
nodding club moss, see Lycopodium cernuum, 356 Oriental chain fern, see Woodwardia orientalis, 537
Norfolk Island tree fern, see Sphaeropteris excelsa, 511 Oriental ostrich fern, see Matteuccia orientalis, 370
North American walking fern, see Camptosorus Oriental water fern, see Ceratopteris thalictroides, 233,
rhizophyllus, 228 color plate 19
northern holly fern, see Polystichum lonchitis, 460 Osmunda, 12, 74, 82, 93, 97, 406
northern lady fern, see Athyrium filix-femina var. Osmunda cinnamomea, 92, 407, color plate 8
angustum, 211 Osmunda claytoniana, 29, 91, 94, 407, color plate 37
northern spreading wood fern, see Dryopteris expansa, Osmunda japonica, 408
308 Osmunda regalis, 91, 93, 96, 408, color plate 6
northern woodsia, see Woodsia alpina, 532 Osmunda regalis var. brasiliensis, 408
northwestern lady fern, see Athyrium filix-femina var. Osmunda regalis var. regalis, 409
cyclosorum, 212 Osmunda regalis var. spectabilis, 409
notched water fern, see Marsilea schelpeana, 368 Osmunda ×ruggii, 408
Notholaena, 111, 236, 399 Osmunda vachellii, 407
Notholaena aurea, see Cheilanthes bonariensis, 238 Osmunda cultivars
Notholaena buchtienii, see Cheilanthes buchtienii, 238 ‘Cristata’, see Osmunda regalis var. regalis, 409
Notholaena dealbata, see Argyrochosma dealbata, 189 ‘Purpurescens’, see Osmunda regalis var. regalis, 409
Notholaena jonesii, see Argyrochosma jonesii, 189 ostrich fern, see Matteuccia, 369
Notholaena newberryi, see Cheilanthes newberryi, 242
Notholaena parvifolia, see Argyrochosma microphylla, 190 Pachypleuria, see Davallia, 262
Notholaena sinuata, see Astrolepis sinuata, 206 Pachypleuria parvula, see Davallia parvula, 272
Notholaena standleyi, 399 Pachypleuria repens, see Davallia repens, 273
Notholaena vestita, see Cheilanthes lanosa, 241 pacific maid, see Adiantum raddianum ‘Pacific Maid’,
173
oak fern, see Gymnocarpium, 334 paddle fern, see Elaphoglossum, 320
oak-leaf fern, see Drynaria, 293 Paesia, 409
Oaxaca spike moss, see Selaginella oaxacana, 502 Paesia scaberula, 409
Obake pyrrosia, see Pyrrosia lingua ‘Cristata’, 485 Palhinhaea cernua, see Lycopodium cernuum, 356
Odontosoria, 399 Paragramma longifolia, see Lepisorus longifolius, 348
Odontosoria chinensis, 400 Parahemionitis, see Hemionitis, 336
Oleandra, 400 Parahemionitis arifolia, see Hemionitis arifolia, 337, color
Oleandra articulata, 400 plate 30
Oleandra neriiformis, 401 Parathelypteris, see Thelypteris, 517
Oleandra nodosa, see Oleandra articulata, 400 Parathelypteris nevadensis, see Thelypteris nevadensis, 520
Oleandra pilosa, 401 Parathelypteris noveboracensis, see Thelypteris
Oleandra pistillaris, 401 noveboracensis, 521
Olfersia, 401 Parathelypteris simulata, see Thelypteris simulata, 526
Olfersia cervina, 402 parsley fern, see Cheilanthes hirta, 241; Cryptogramma,
Onoclea, 402 248
Onoclea sensibilis, 29, 91, 92, 93, 94, 96, 97, 146, 402 peacock fern, see Selaginella willdenovii, 506
Onoclea struthiopteris, see Matteuccia struthiopteris, 371, peacock plant, see Selaginella uncinata, 505, color plate
color plate 33 49
Onychium, 402 peacock pyrrosia, see Pyrrosia lingua ‘Nankin-shishi’, 485
Onychium densum, see Aspidotis densa, 193 Pecluma, 410, 439
Onychium japonicum, 403 Pecluma dispersa, 410
Ophioglossum, 69, 404 Pecluma pectinata, 410
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Pecluma plumula, 411 ‘Cristatum’, see Phlebodium aureum, 420

Pellaea, 93, 98, 111, 412 ‘Ekstrand’, see Phlebodium aureum, 420
Pellaea adiantoides, see Pellaea viridis, 416 ‘Leatherman’, see Phlebodium pseudoaureum, 421
Pellaea andromedifolia, 412 ‘Mandaianum’, see Phlebodium aureum, 420
Pellaea atropurpurea, 92, 413 ‘Mayi’, see Phlebodium aureum, 420
Pellaea brachyptera, 413 ‘Mexican Tasseled’, see Phlebodium pseudoaureum, 421
Pellaea bridgesii, 413 ‘Undulatum’, see Phlebodium aureum, 420
Pellaea cordata, see Pellaea cordifolia, 414 Phlegmariurus carinatus, see Lycopodium carinatum, 360
Pellaea cordifolia, 414 Phlegmariurus nummulariifolius, see Lycopodium
Pellaea densa, see Aspidotis densa, 193 nummulariifolium, 358
Pellaea falcata, 414 Phlegmariurus phlegmaria, see Lycopodium phlegmaria,
Pellaea falcata var. nana, 415 358
Pellaea glabella, 415 Phlegmariurus squarrosus, see Lycopodium squarrosum,
Pellaea hastata, see Pellaea viridis, 416 360
Pellaea mucronata, 415 Photinopteris, see Aglaomorpha, 176
Pellaea nana, see Pellaea falcata, 414 Photinopteris speciosa, see Aglaomorpha speciosa, 179
Pellaea ovata, 415 Phyllitis, 422
Pellaea paradoxa, 415 Phyllitis scolopendrium, 64, 92, 94, 97, 98, 149, 422,
Pellaea rotundifolia, 97, 100, 416 color plate 39
Pellaea sagittata var. cordata, see Pellaea cordifolia, 414 Phyllitis scolopendrium var. americana, 423
Pellaea ternifolia, 416 Phyllitis cultivars
Pellaea ternifolia var. wrightiana, see Pellaea wrightiana, ‘Crispum’, see Phyllitis scolopendrium, 423
417 ‘Cristatum’, see Phyllitis scolopendrium, 423
Pellaea viridis, 82, 416 ‘Laceratum’, see Phyllitis scolopendrium, 423
Pellaea viridis var. macrophylla, 417 ‘Muricatum’, see Phyllitis scolopendrium, 423
Pellaea wrightiana, 417 ‘Ramosum’, see Phyllitis scolopendrium, 423
Pellaea ‘Star Glow’, see Pellaea falcata, 415 ‘Undulatum’, see Phyllitis scolopendrium, 423
pellucid polypody, see Polypodium pellucidum, 448 Phymatodes, see Lecanopteris, 345; Microsorum, 377
Peltapteris, see Elaphoglossum, 320 Phymatodes banjeriana, 381
Peltapteris peltata, see Elaphoglossum peltatum, 324 Phymatodes crustaca, see Lecanopteris crustacea, 346
Pentagramma, 417 Phymatodes sinuosa, see Lecanopteris sinuosa, 346
Pentagramma pallida, 418 Phymatodes viellardii, see Microsorum viellardii, 388
Pentagramma triangularis, 418 Phymatosorus, see Lecanopteris, 345; Microsorum, 377
pepperwort, see Marsilea, 365 Phymatosorus commutatus, see Microsorum commutatum,
Peruvian maidenhair, see Adiantum peruvianum, 170 380
Pessopteris, see Niphidium, 398 Phymatosorus cuspidatus, see Microsorum lucidum, 382
petticoats, see Nephrolepis cordifolia ‘Petticoats’, 392 Phymatosorus diversifolius, see Microsorum pustulatum,
Phanerophlebia, see Cyrtomium, 254 385
Phegopteris, 418 Phymatosorus grossus, see Microsorum grossum, 380
Phegopteris connectilis, 92, 94, 96, 97, 418 Phymatosorus lucidus, see Microsorum lucidum, 382
Phegopteris decursive-pinnata, 418 Phymatosorus nigrescens, see Microsorum membranifolium,
Phegopteris hexagonoptera, 97, 419 382
Phlebodium, 93, 419, 439 Phymatosorus pustulatus, see Microsorum pustulatum, 385
Phlebodium areolatum, see Phlebodium pseudoaureum, Phymatosorus scandens, see Microsorum scandens, 386
420, color plates 5, 38 Phymatosorus scolopendria, see Microsorum scolopendria,
Phlebodium aureum, 82, 95, 97, 98, 100, 420 387
Phlebodium decumanum, 420 pillwort, see Pilularia, 423
Phlebodium glaucopruniatum, see Phlebodium Pilularia, 423
pseudoaureum, 420, color plates 5, 38 Pilularia americana, 423
Phlebodium glaucum, see Phlebodium pseudoaureum, 420, Pilularia globulifera, 424
color plates 5, 38 pimple fern, see Microsorum membranifolium, 382
Phlebodium pseudoaureum, 82, 94, 98, 100, 420, color pin-cushion moss, see Selaginella kraussiana ‘Brownii’,
plates 5, 38 501
Phlebodium ×schneideri, 421 pine fern, see Anemia adiantifolia, 183
Phlebodium cultivars Pityrogramma, 93, 98, 111, 424
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Pityrogramma argentea, 425 ‘Horne’s Surprise’, see Platycerium madagascariense,

Pityrogramma austroamericana, 82, 95, 424, color 433
plates 40, 41 ‘Mona Loa’, see Platycerium hillii, 433
Pityrogramma calomelanos, 95, 425 ‘Netherlands’, see Platycerium bifurcatum, 430
Pityrogramma calomelanos var. aureoflava, see ‘Panama’, see Platycerium hillii, 433
Pityrogramma austroamericana, 425 ‘Sanchez’, see Platycerium ×elemaria, 431
Pityrogramma calomelanos var. austroamericana, see ‘Talnadge’, see Platycerium hillii, 433
Pityrogramma austroamericana, 425 Platyloma, see Pellaea, 412
Pityrogramma ebenea, 425 Platyloma falcata, see Pellaea falcata, 414
Pityrogramma hybrida, 425 Platyloma rotundifolia, see Pellaea rotundifolia, 416
Pityrogramma tartarea, see Pityrogramma ebenea, 425 Pleopeltis, 439
Pityrogramma triangularis, see Pentagramma triangularis, Pleopeltis ×leucospora, see Polypodium ×leucosporum, 446
418 Pleopeltis macrocarpa, 446
Pityrogramma trifoliata, 425 Pleopeltis percussa, see Microgramma percussa, 374
Platycerium, 45, 69, 93, 97, 98, 104–106, 427 Pleopeltis polypodioides, see Polypodium polypodioides, 449
Platycerium alcicorne, 429 Pleopeltis thunbergiana, see Lepisorus thunbergianus, 348
Platycerium andinum, 429 Pleopeltis thyssanolepis, see Polypodium thyssanolepis, 451
Platycerium angolense, see Platycerium elephantotis, 431 ×Pleopodium leucosporum, see Polypodium ×leucosporum,
Platycerium biforme, see Platycerium coronarium, 431 446
Platycerium bifurcatum, 91, 100, 429, color plates 1, 5, Polybotrya, 437
42 Polybotrya cervina, see Olfersia cervina, 402
Platycerium bifurcatum var. hillii, see Platycerium hillii, Polybotrya osmundacea, 437
432 Polygoniophlebium, see Polypodium, 441
Platycerium bifurcatum subsp. veitchii, see Platycerium Polynesian davallia, see Davallia solida, 274
veitchii, 435 Polypodium, 68, 98, 108–109, 437
Platycerium bifurcatum subsp. willinckii, see Platycerium Polypodium alternifolium, see Microsorum grossum, 380
willinckii, 437 Polypodium amoenum, 441
Platycerium coronarium, 431 Polypodium amorphum, 445
Platycerium diversifolium, see Platycerium ellisii, 432 Polypodium angustifolium, see Campyloneurum
Platycerium ×elemaria, 431 angustifolium, 229
Platycerium elephantotis, 431 Polypodium appalachianum, 441
Platycerium ellisii, 432 Polypodium areolatum, see Phlebodium pseudoaureum,
Platycerium grande, 432 420, color plates 5, 38
Platycerium grande var. tambourinense, see Platycerium Polypodium attenuatum, 442
superbum, 435, color plate 43 Polypodium aureum var. aureum, see Phlebodium aureum,
Platycerium hillii, 432 420
Platycerium holttumii, 433 Polypodium australe, see Polypodium cambricum, 443
Platycerium madagascariense, 433 Polypodium bombycinum, 442
Platycerium ×mentelosii, 433 Polypodium brasiliense, 442
Platycerium quadridichotomum, 67, 434 Polypodium californicum, 443
Platycerium ridleyi, 434 Polypodium calirhiza, 443
Platycerium stemaria, 435 Polypodium cambricum, 94, 443
Platycerium stemaria var. laurentii, 435 Polypodium cambricum var. cambricum, 443
Platycerium sumbawense, see Platycerium willinckii, 437 Polypodium chnoodes, see Polypodium dissimile, 444
Platycerium superbum, 435, color plate 43 Polypodium ciliatum, see Microgramma reptans, 374
Platycerium vassei, see Platycerium alcicorne, 429 Polypodium costatum, see Campyloneurum costatum, 230
Platycerium veitchii, 435 Polypodium crassifolium, see Niphidium crassifolium, 398
Platycerium wallichii, 436, color plate 27 Polypodium crustaceum, see Lecanopteris crustacea, 346
Platycerium wandae, 436 Polypodium decumanum, see Phlebodium decumanum,
Platycerium willhelminae-reginae, see Platycerium wandae, 420
436 Polypodium dilatatum, see Microsorum insigne, 381
Platycerium willinckii, 437 Polypodium dispersum, see Pecluma dispersa, 410
Platycerium willinckii var. venosum, 430 Polypodium dissimile, 444
Platycerium cultivars Polypodium diversifolium, see Microsorum pustulatum,
‘Fantastic Gardens’, see Platycerium ×mentelosii, 433 385
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Polypodium dryopteris, see Gymnocarpium dryopteris, 335, Polypodium persicifolium, see Goniophlebium
color plate 29 persicifolium, 332
Polypodium ×ekstrandii, see Goniophlebium ×ekstrandii, Polypodium phyllitidis, see Campyloneurum phyllitidis,
331 230
Polypodium formosanum, 444, color plate 44 Polypodium phymatodes, see Microsorum scolopendria, 380
Polypodium fortunei, see Microsorum fortunei, 380 Polypodium piloselloides, see Microgramma piloselloides,
Polypodium fraxinifolium, 448 374
Polypodium glaucophyllum, see Polypodium levigatum, Polypodium plebeium, 448
447 Polypodium polycarpon, see Microsorum punctatum, 384,
Polypodium glaucopruniatum, see Phlebodium color plate 34
pseudoaureum, 420, color plates 5, 38 Polypodium polypodioides, 449
Polypodium glaucum, see Phlebodium pseudoaureum, 420, Polypodium polypodioides var. aciculare, 449
color plates 5, 38 Polypodium polypodioides var. burchellii, 449
Polypodium glycyrrhiza, 444 Polypodium polypodioides var. ecklonii, 449
Polypodium grossum, see Microsorum grossum, 380 Polypodium polypodioides var. michauxianum, 449
Polypodium guttatum, 445 Polypodium polypodioides var. minus, 449
Polypodium hancockii, see Microsorum insigne, 381 Polypodium pseudoaureum, see Phlebodium pseudoaureum,
Polypodium hesperium, 445 420, color plates 5, 38
Polypodium heterophyllum, see Microgramma heterophylla, Polypodium pteropus, see Microsorum pteropus, 384
372 Polypodium ptilorhizon, 449
Polypodium hirsutissimum, 446 Polypodium punctatum, see Microsorum punctatum, 384,
Polypodium integrifolium, see Microsorum punctatum, color plate 34
384, color plate 34 Polypodium pustulatum, see Microsorum pustulatum, 385;
Polypodium interjectum, 446 Microsorum scandens, 386
Polypodium irioides, see Microsoum punctatum, 384 Polypodium pyrrholepis, 98, 449, color plate 45
Polypodium knightiae, see Goniophlebium subauriculatum Polypodium rhodopleuron, 450
‘Knightiae’, 333, color plate 28 Polypodium sanctae-rosae, 450
Polypodium kuhnii, see Polypodium attenuatum, 442 Polypodium scandens G. Forster, see Microsorum scandens,
Polypodium latum, see Campyloneurum brevifolium, 230 386
Polypodium ×leucosporum, 446 Polypodium scandens Labill, see Microsorum pustulatum,
Polypodium levigatum, 447 385
Polypodium lineare, see Lepisorus thunbergianus, 348 Polypodium scolopendria, see Microsorum grossum, 380;
Polypodium linguiforme, see Microsorum linguiforme, 382 Microsorum scolopendria, 387
Polypodium longissimum, see Microsorum longissimum, Polypodium scouleri, 451
382 Polypodium sibericum, 452
Polypodium loriceum, 447 Polypodium sinuosum, see Lecanopteris sinuosa, 346
Polypodium lucidum, see Microsorum lucidum, 382 Polypodium squamulosum, see Microgramma squamulosa,
Polypodium lycopodioides, see Microgramma lycopodioides, 375
373 Polypodium subauriculatum, see Goniophlebium
Polypodium mandaianum, see Phlebodium aureum subauriculatum, 333, color plate 28
‘Mandaianum’, 420 Polypodium tenellum, see Arthropteris tenella, 191
Polypodium maritimum, 447 Polypodium tenuilore, see Microsorum samarense, 385
Polypodium megalophyllum, see Microgramma Polypodium thunbergianum, see Lepisorus thunbergianus,
megalophylla, 373 348
Polypodium menisciifolium, 447 Polypodium thyssanolepis, 451
Polypodium musifolium, see Microsorum musifolium, 383 Polypodium triseriale, 452
Polypodium myriocarpum, see Microsorum longissimum, Polypodium vacciniifolium, see Microgramma
382 vacciniifolia, 375
Polypodium nigrescens, see Microsorum membranifolium, Polypodium verrucosum, see Goniophlebium percussum,
382 332
Polypodium normale, see Microsorum fortunei, 380 Polypodium virginianum, 452
Polypodium palmeri, see Microgramma nitida, 373 Polypodium vitiense, see Microsorum commutatum, 380
Polypodium pectinatum, see Pecluma pectinata, 410 Polypodium vulgare, 92, 452
Polypodium pellucidum, 448 Polypodium vulgare var. occidentale, see Polypodium
Polypodium percussum, see Microgramma percussa, 374 glycyrrhiza, 444
 P N I 599

Polypodium xalapense, see Campyloneurum xalapense, 231 Polystichum ×potteri, 455

Polypodium cultivars Polystichum proliferum, 462
‘Cambricum’, see Polypodium cambricum, 443; Polystichum retrosopaleaceum, 462
Microsorum commutatum, 380 Polystichum richardii, 463
‘Cambrioides’, see Polypodium triseriale, 452 Polystichum rigens, 463
‘Cornubiense’, see Polypodium cambricum, 443 Polystichum scopulinum, 464
‘Cristata’, see Polypodium triseriale, 452 Polystichum setiferum, 64, 66, 69, 70, 92, 97, 464, color
‘Cristatum’, see Polypodium formosanum, 444; plate 46
Polypodium triseriale, 452 Polystichum setigerum, 465
‘Knightiae’, see Goniophlebium subauriculatum, 333, Polystichum setosum, 462
color plate 28 Polystichum silvaticum, 466
‘Nola’, see Goniophlebium ×ekstrandii, 331 Polystichum standishii, see Arachniodes standishii, 188
‘Pulcherrimum’, see Polypodium cambricum, 443 Polystichum stenophyllum, 466
‘Ramosum Hillman’, see Polypodium vulgare, 453 Polystichum stimulans, 466
polypody, see Polypodium, 437; see also Goniophlebium, Polystichum sylvaticum, see Polystichum silvaticum, 466
331; Microsorum scolopendria, 387 Polystichum tagawanum, 467
Polystichum, 93, 94, 454 Polystichum tripteron, 467
Polystichum acrostichoides, 91, 92, 97, 455 Polystichum tsus-simense, 96, 98, 101, 468
Polystichum aculeatum, 94, 456 Polystichum tsus-simense var. mayebarae, 469
Polystichum adiantiforme, see Rumohra adiantiformis, 491 Polystichum tsus-simense var. tsus-simense, 469
Polystichum alaskense, see Polystichum setigerum, 465, Polystichum vestitum, 469
color plate 46 Polystichum xiphophyllum, 469
Polystichum andersonii, 92, 456 Polystichum yaemonense, 469
Polystichum angulare, see Polystichum setiferum, 464, Polystichum yaeyamense, 469
color plate 46 Polystichum yokohamense, 470
Polystichum aristatum, see Arachniodes aristata, 187 Polystichum cultivars
Polystichum braunii, 456, color plate 6 ‘Alaska’, see Polystichum setiferum ‘Divisilobum’, 464
Polystichum braunii subsp. alaskense, 465 ‘Congestum’, see Polystichum setiferum, 464
Polystichum californicum, 457 ‘Crispum’, see Polystichum acrostichoides, 455
Polystichum capense, see Rumohra adiantiformis, 491 ‘Cristatum’, see Polystichum acrostichoides, 455
Polystichum coriaceum, see Rumohra adiantiformis, 491 ‘Divisilobum’, see Polystichum setiferum Divisilobum
Polystichum craspedosorum, 457 Group, 464
Polystichum dudleyi, 458 ‘Divisilobum Cristatum’, see Polystichum setiferum
Polystichum fibrilloso-paleaceum, 470 ‘Trilobum’, 465
Polystichum illicifolium, see Polystichum stimulans, 466 ‘Divisilobum Grandiceps’, see Polystichum setiferum
Polystichum ×illyricum, 456 ‘Trilobum’, 465
Polystichum imbricans, 458 ‘Imbricatum’, see Polystichum setiferum Imbricatum
Polystichum imbricans subsp. curtum, 458 Group, 464
Polystichum imbricans subsp. imbricans, 458 ‘Incisum’, see Polystichum acrostichoides, 455
Polystichum kruckebergii, 458 ‘Multilobum’, see Polystichum setiferum Multilobum
Polystichum lemmonii, 98, 459 Group, 465
Polystichum lentum, 459 ‘Plumoso-Divisilobum’, see Polystichum setiferum
Polystichum lepidocaulon, 459 Plumoso-Divisilobum Group, 465, color plate 46
Polystichum lobatum, see Polystichum aculeatum, 456 ‘Proliferum’, see Polystichum setiferum ‘Divisilobum’,
Polystichum ×lonchitiforme, 460 464
Polystichum lonchitis, 460 ‘Trilobum’, see Polystichum setiferum, 465
Polystichum makinoi, 460 potato fern, see Solanopteris, 508
Polystichum mohrioides var. lemmonii, see Polystichum pouched coral fern, see Gleichenia dicarpa, 330
lemmonii, 459 powdery cloak fern, see Argyrochosma dealbata, 189
Polystichum munitum, 92, 460 prickly rasp fern, see Doodia aspera, 289
Polystichum munitum var. imbricans, see Polystichum prickly shield fern, see Polystichum braunii, 456, color
imbricans, 458 plate 6; Polystichum vestitum, 469
Polystichum neolobatum, 461 prickly-tree club moss, see Lycopodium dendroideum,
Polystichum nepalense, 461 357
Polystichum polyblepharum, 92, 462 Prince-of-Wales plume, see Leptopteris superba, 351
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princess pine, see Lycopodium, 113, 354; Lycopodium ‘Ouvrardii’, see Pteris cretica, 475
dendroideum, 357; Lycopodium obscurum, 358 ‘Parkeri’, see Pteris cretica, 475
Prosaptia, 333 ‘Rivertoniana’, see Pteris cretica, 475
Pseudodrynaria, see Aglaomorpha, 176 ‘Roweri’, see Pteris cretica, 475
Psilotum, 113, 470 ‘Victoriae’, see Pteris ensiformis, 95, 101, 476
Psilotum complanatum, 470 ‘Wilsonii’, see Pteris cretica, 101, 475
Psilotum ×intermedium, 470 ‘Wimsettii’, see Pteris cretica, 475
Psilotum nudum, 470 purple cliff brake, see Pellaea atropurpurea, 412
Pteretis, see Matteuccia, 369 Pyrrosia, 480
Pteretis nodulosa, see Matteuccia struthiopteris, 371, color Pyrrosia adnascens, see Pyrrosia lanceolata, 484
plate 33 Pyrrosia angustata, 481
Pteridium, 93, 470 Pyrrosia beddomeana, see Pyrrosia costata, 482
Pteridium aquilinum, 94, 471 Pyrrosia confluens, 482
Pteridium aquilinum var. caudatum, 146, 471 Pyrrosia confluens var. confluens, 482
Pteridium aquilinum var. latiusculum, 146, 471 Pyrrosia confluens var. dielsii, 482
Pteridium aquilinum var. pseudocaudatum, 471 Pyrrosia costata, 482
Pteridium aquilinum var. pubescens, 471 Pyrrosia davidii, see Pyrrosia porosa, 487
Pteris, 472 Pyrrosia dielsii, see Pyrrosia confluens var. dielsii, 482
Pteris altissima, 473 Pyrrosia eleagnifolia, 483
Pteris argyraea, 95, 473, color plate 47 Pyrrosia fallax, 483
Pteris aspericaulis, 95, 474 Pyrrosia hastata, 484
Pteris aspericaulis var. tricolor, 474 Pyrrosia heteractis, see Pyrrosia lingua var. heteractis,
Pteris biaurita, 474 485
Pteris catoptera, 474 Pyrrosia lanceolata, 484
Pteris cretica, 66, 82, 92, 94, 98, 100, 474 Pyrrosia linearifolia, 484
Pteris cretica var. albolineata, see Pteris nipponica, 478 Pyrrosia lingua, 95, 484
Pteris dentata, 475 Pyrrosia lingua var. heteractis, 485
Pteris dispar, see Pteris semipinnata var. dispar, 479 Pyrrosia lingua var. lingua, 485
Pteris ensiformis, 95, 475 Pyrrosia longifolia, 486
Pteris fauriei, 479 Pyrrosia nummulariifolia, 486
Pteris flabellata, see Pteris dentata, 475 Pyrrosia obovata, see Pyrrosia nummulariifolia, 486
Pteris grandifolia, 476 Pyrrosia pekinensis, see Pyrrosia porosa, 487
Pteris longifolia, 476 Pyrrosia piloselloides, 486
Pteris multifida, 82, 97, 98, 101, 478 Pyrrosia polydactyla, 487, color plate 48
Pteris nipponica, 95, 478 Pyrrosia porosa, 487
Pteris quadriaurita, 478 Pyrrosia princeps, 487
Pteris quadriaurita var. argentea, see Pteris argyraea, 473, Pyrrosia rupestris, 488
color plate 47 Pyrrosia samarensis, 488
Pteris quadriaurita var. argyraea, see Pteris argyraea, 473, Pyrrosia serpens, 483
color plate 47 Pyrrosia sheareri, 488
Pteris semipinnata, 479 Pyrrosia splendens, 489
Pteris semipinnata var. dispar, 479 Pyrrosia stigmosa, 490
Pteris serrulata, see Pteris multifida, 478 Pyrrosia tricuspis, see Pyrrosia hastata, 484
Pteris tremula, 82, 93, 94, 100, 479 Pyrrosia varia, see Pyrrosia lanceolata, 484
Pteris tricolor, see Pteris aspericaulis var. tricolor, 474 Pyrrosia cultivars
Pteris tripartita, 480 ‘Cristata’, see Pyrrosia lingua, 485
Pteris vittata, 82, 94, 95, 480 ‘Monstrifera’, see Pyrrosia lingua, 485
Pteris cultivars ‘Nankin-shishi’, see Pyrrosia lingua, 485
‘Albo-lineata’, see Pteris cretica, 474; Pteris nipponica, ‘Nokogiri-ba’, see Pyrrosia lingua, 485
478
‘Childsii’, see Pteris cretica, 475 queen staghorn, see Platycerium wandae, 436
‘Cristata Mayii’, see Pteris cretica, 475 Quercifilix, see Tectaria, 513
‘Distinction’, see Pteris cretica, 475 Quercifilix zeylanica, see Tectaria zeylanica, 517
‘Evergemiensis’, see Pteris ensiformis, 95, 476
‘Mayii’, see Pteris cretica, 475 rabbit’s-foot fern, see Davallia 262; Phlebodium, 419
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rainbow moss, see Selaginella uncinata, 505, color plate scented fern, see Paesia scaberula, 409
49 Schaffneria, 494
rasp fern, see Doodia, 289 Schaffneria nigripes, 494
rattlesnake fern, see Botrychium virginianum, 227 Schellolepis, see Goniophlebium, 331
recurved broad-buckler fern, see Dryopteris dilatata Schneider’s rabbit-foot fern, see Phlebodium ×schneideri,
‘Recurved Form’, 307 421
red spike moss, see Selaginella umbrosa, 505 Scleroglossum, 333
Regnellidium, 490 Scott’s spleenwort, see Asplenium ×ebenoides, 198
Regnellidium diphyllum, 490 scouring rush, see Equisetum, Equisetum hyemale, 328
resurrection fern, see Polypodium polypodioides, 449 scurfy sword fern, see Nephrolepis hirsutula, 395
resurrection plant, see Selaginella lepidophylla, 501 Scyphularia, see Davallia, 262
Rhipidopteris peltata, see Elaphoglossum peltatum, 324 Scyphularia pentaphylla, see Davallia pentaphylla, 272
Richard’s holly fern, see Polystichum richardii, 463 Scyphularia pycnocarpa, see Davallia pentaphylla, 272
rigid buckler fern, see Dryopteris mindshelkensis, 314 Scyphularia simplicifolia, see Davallia triphylla, 275
robust male fern, see Dryopteris complexa, 303 Scyphularia sinusora, see Davallia pentaphylla, 272
rock brake, see Cryptogramma, 248 Scyphularia triphylla, see Davallia triphylla, 275
rock felt fern, see Pyrrosia rupestris, 488 scythian lamb, see Cibotium barometz, 245
Rockford holly fern, see Cyrtomium falcatum sea spleenwort, see Asplenium marinum, 199
‘Rockfordianum’, 256 Seemann’s maidenhair, see Adiantum seemannii, 173
rock polypody, see Polypodium virginianum, 452 Selaginella, 91, 95, 97, 114, 495
rock sword fern, see Polystichum scopulinum, 464 Selaginella apoda, 497
rock tassel fern, see Lycopodium squarrosum, 360 Selaginella braunii, 497
rosy maidenhair, see Adiantum hispidulum, 166, color Selaginella caulescens, see Selaginella involvens, 499
plate 9 Selaginella denticulata, 500
rough horsetail, see Equisetum hyemale, 328 Selaginella diffusa, 497
rough maidenhair, see Adiantum hispidulum, 166 Selaginella douglassii, 498
rough sword fern, see Nephrolepis hirsutula, 395 Selaginella emmeliana, see Selaginella pallescens, 503
rough tree fern, see Alsophila australis, 182; Dicksonia Selaginella erythropus, 498
squarrosa, 284 Selaginella harrisii, 499
royal fern, see Osmunda regalis, 408, color plate 6 Selaginella heterodonta, 499
Rumohra, 490 Selaginella involvens, 499
Rumohra adiantiformis, 92, 93, 94, 98, 100, 491 Selaginella kraussiana, 95, 500
Rumohra ‘Davis’, see Rumohra adiantiformis, 491 Selaginella lepidophylla, 501
running pine, see Lycopodium clavatum, 356; Lycopodium Selaginella martensii, 501
complanatum, 357 Selaginella moellendorffii, 501
rusty brake, see Pteris vittata, 480 Selaginella oaxacana, 502
rusty woodsia, see Woodsia ilvensis, 532 Selaginella oregana, 502
rusty-back fern, see Ceterach, 233 Selaginella pallescens, 503
Selaginella plana, 503
Sadleria, 491 Selaginella pulcherrima, 503
Sadleria cyatheoides, 111, 492 Selaginella serpens, 504
Sadleria pallida, 492 Selaginella umbrosa, 505
sago tree fern, see Sphaeropteris medullaris, 512 Selaginella uncinata, 505, color plate 49
Salvinia, 93, 493 Selaginella viticulosa, 505
Salvinia auriculata, 493 Selaginella vogelii, 506
Salvinia minima, 493 Selaginella willdenovii, 506
Salvinia molesta, 494 Selaginella cultivars
Salvinia natans, 494 ‘Albolineata’, see Selaginella martensii, 501
Salvinia rotundifolia, see Salvinia auriculata, 493 ‘Aurea’, see Selaginella kraussiana, 501
saw fern, see Blechnum serrulatum, 221 ‘Brownii’, see Selaginella kraussiana, 501
scale fern, see Ceterach officinarum, 233, color plate 20 ‘Gold’, see Selaginella kraussiana, 501
scaly buckler fern, see Dryopteris remota, 316 ‘Gold-tips’, see Selaginella kraussiana, 501
scaly male fern, see Dryopteris affinis, 298 Selliguea, 439, 507
scaly polypody, see Polypodium thyssanolepis, 451 Selliguea feei, 507
scaly wood fern, see Dryopteris polylepis, 315 sensitive fern, see Onoclea, 402
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serpent fern, see Phlebodium aureum, 420 spear-leaved fern, see Doryopteris palmata, 292
serpent’s-tongue fern, see Ophioglossum, 404 Sphaerocionium, see Hymenophyllum, 339
serrate lattice-vein fern, see Thelypteris serrata, 525 Sphaeropteris, 253, 509
shaggy wood fern, see Dryopteris cycadina, 305 Sphaeropteris armaganensis, 510
Shasta holly fern, see Polystichum lemmonii, 459 Sphaeropteris cooperi, 91, 92, 93, 94, 95, 110, 111, 511,
shield fern, see Dryopteris, 296; Polystichum, 454; color plates 2, 3
Thelypteris kunthii, 520 Sphaeropteris excelsa, 95, 511
shining club moss, see Lycopodium lucidulum, 358 Sphaeropteris horrida, 512
shiny shield fern, see Lastreopsis acuminata, 343 Sphaeropteris medullaris, 512
shoestring fern, see Vittaria lineata, 530 Sphaeropteris cultivars
sickle spleenwort, see Asplenium polyodon, 202 ‘Brentwood’, see Sphaeropteris cooperi, 511
Siebold’s wood fern, see Dryopteris sieboldii, 317 ‘Robusta’, see Sphaeropteris cooperi, 511
sierra cliff brake, see Pellaea brachyptera, 413 Sphenomeris, see Odontosoria, 400
sierra water fern, see Thelypteris nevadensis, 520 Sphenomeris chinensis, see Odontosoria chinensis, 400
silverback fern, see Pentagramma, 417; Pityrogramma, Sphenomeris chusana, see Odontosoria chinensis, 400
424 spider brake, see Pteris multifida, 478
silver brake, see Pteris argyraea, 473, color plate 47 spike moss, see Selaginella, 495
silver-dollar fern, see Adiantum peruvianum, 170 spinulose wood fern, see Dryopteris carthusiana, 302
silver fern, see Pityrogramma calomelanos, 425 spleenwort, see Asplenium, 193
silver glade fern, see Deparia acrostichoides, 280 spreading selaginella, see Selaginella kraussiana, 500
silver tree fern, see Alsophila tricolor, 182, color plate 11 squirrel’s-foot fern, see Davallia mariesii var. stenolepis,
silvery felt fern, see Pyrrosia confluens var. dielsii, 482 271, color plate 5; Davallia trichomanoides, 275
silvery spleenwort, see Deparia acrostichoides, 280 staghorn club moss, see Lycopodium clavatum, 356
slender cloak fern, see Cheilanthes bonariensis, 238 staghorn fern, see Platycerium, 427
slender crested male fern, see Dryopteris filix-mas star-scale cloak fern, see Astrolepis, 206
‘Linearis Polydactyla’, 310 Stegnogramma, see Thelypteris, 517
slender lip fern, see Cheilanthes feei, 239 Stenochlaena, 512
slender rock brake, see Cryptogramma stelleri, 251 Stenochlaena palustris, 512
slender tree fern, see Dicksonia squarrosa, 284 Stenochlaena tenuifolia, 512
small-leaved climbing fern, see Lygodium microphyllum, Sticherus, 330
362 Sticherus flabellatus, 330
small-leaved cloak fern, see Argyrochosma microphylla, Sticherus lobatus, 331
190 strap fern, see Campyloneurum, 229
smooth cliff brake, see Pellaea glabella, 415 strawberry fern, see Hemionitis palmata, 338
smooth horsetail, see Equisetum laevigatum, 328 striped brake, see Pteris argyraea, 473, color plate 47
smooth rock spleenwort, see Asplenium fontanum, 199 Struthiopteris, see Matteuccia, 369
snowy club moss, see Selaginella martensii ‘Albolineata’, Struthiopteris pensylvanica, see Matteuccia struthiopteris,
501 371, color plate 33
soft shield fern, see Polystichum setiferum, 464, color swamp fern, see Blechnum serrulatum, 221; Woodwardia
plate 46 areolata, 537
Solanopteris, 440, 508 sweat plant, see Selaginella pallescens, 503
Solanopteris bifrons, 508 sword fern, see Nephrolepis, 389; Polystichum, 454
Solanopteris brunei, 508
southern beech fern, see Phegopteris hexagonoptera, 419 Tagawa’s holly fern, see Polystichum tagawanum, 467
southern fragile fern, see Cystopteris protrusa, 260 tailed bracken, see Pteridium aquilinum var.
southern grape fern, see Botrychium biternatum, 227 pseudocaudatum, 471
southern lady fern, see Athyrium filix-femina var. Tasmanian dicksonia, see Dicksonia antarctica, 282,
asplenioides, 212 color plate 3
southern maidenhair, see Adiantum capillus-veneris, 161 Tasman’s davallia, see Davallia tasmanii, 274
southern polypody, see Polypodium cambricum, 443 tassel fern, see Lycopodium, 354
southern shield fern, see Thelypteris kunthii, 520 Tectaria, 513
southern wood fern, see Dryopteris ludoviciana, 313; Tectaria beccariana, 514
Thelypteris kunthii, 520 Tectaria cicutaria, 515
southwestern brittle fern, see Cystopteris reevesiana, 261 Tectaria decurrens, 514
sparse-lobed grape fern, see Botrychium biternatum, 227 Tectaria gemmifera, 69, 70, 515
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Tectaria heracleifolia, 515 Todea barbara, 82, 93, 526

Tectaria hilocarpa, 516 Tokyo wood fern, see Dryopteris tokyoensis, 318
Tectaria incisa, 516 tongue fern, see Pyrrosia lingua, 484
Tectaria incisa f. vivipara, see Tectaria vivipara, 516 toothed brake, see Pteris dentata, 475
Tectaria latifolia, see Tectaria hilocarpa, 516 toothed davallia, see Davallia denticulata, 264
Tectaria vivipara, 516 toothed wood fern, see Dryopteris carthusiana, 302
Tectaria zeylanica, 101, 517 Torre’s fern, see Macrothelypteris torresiana, 363
Tennessee bladder fern, see Cystopteris tennesseensis, 261 Trachypteris, 527
Terpsichore, 333 Trachypteris pinnata, 528
Thayeria, see Aglaomorpha, 176 trailing maidenhair, see Adiantum caudatum, 162
Thayeria cornucopia, see Aglaomorpha cornucopia, 177 tree club moss, see Lycopodium obscurum, 358
Thelypteris, 95, 97, 517 tree fern, see Alsophila, 181; Cibotium, 244; Ctenitis 251;
Thelypteris acuminata, 97, 518 Cyathea, 252; Dicksonia, 281; Sphaeropteris, 511
Thelypteris augescens, 519 tree spike moss, see Selaginella involvens, 499
Thelypteris decursive-pinnata, see Phegopteris decursive- triangular staghorn, see Platycerium stemaria, 435
pinnata, 418 triangular water fern, see Ceratopteris pteridoides, 232
Thelypteris dentata, 82, 519 Trichomanes, 67, 528
Thelypteris guadalupensis, 520 Trichomanes auriculatum, 528
Thelypteris hexagonoptera, see Phegopteris hexagonoptera, Trichomanes holopterum, 529
418 Trichomanes radicans, 529
Thelypteris kunthii, 520 Trichomanes speciosum, 529
Thelypteris limbosperma, 69, 520 trifid holly fern, see Polystichum tripteron, 467
Thelypteris subgenus Macrothelypteris, see trisect brake, see Pteris tripartita, 480
Macrothelypteris, 363 Trismeria, see Pityrogramma, 424
Thelypteris nevadensis, 520 Trismeria trifoliata, see Pityrogramma trifoliata, 426
Thelypteris normalis, see Thelypteris kunthii, 520 triumph maidenhair, see Adiantum raddianum
Thelypteris noveboracensis, 93, 521 ‘Triumph’, 173
Thelypteris ovata, 522 Trogostolon, see Davallia, 262
Thelypteris palustris, 29, 93, 97, 522 Trogostolon falcinellus, see Davallia falcinella, 266
Thelypteris palustris f. pufferae, 522 tsus-sima holly fern, see Polystichum tsus-simense, 468
Thelypteris parasitica, 82, 522 tuber sword fern, see Nephrolepis cordifolia, 391, color
Thelypteris patens, 523 plate 35
Thelypteris subgenus Phegopteris, see Phegopteris, 419 two-edged maidenhair, see Adiantum anceps, 160
Thelypteris phegopteris, see Phegopteris connectilis, 418
Thelypteris poiteana, 523 umbrella fern, see Gleichenia microphylla, 330
Thelypteris puberula, 73, 82, 93, 524 Urostachys lucidulus, see Lycopodium lucidulum, 358
Thelypteris quelpaertensis, 520
Thelypteris reptans, 524 Vandenboschia, see Trichomanes, 529
Thelypteris resinifera, 524 Vandenboschia auriculata, see Trichomanes auriculatum,
Thelypteris reticulata, 525 528
Thelypteris rudis, 525 Vandenboschia speciosa, see Trichomanes speciosum, 528
Thelypteris sancta, 524 variegated maidenhair, see Adiantum raddianum
Thelypteris serrata, 525 ‘Variegatum’, 173
Thelypteris setigera, 364 vegetable fern, see Diplazium esculentum, 12, 69, 287
Thelypteris simulata, 526 vegetable lamb, see Cibotium barometz, 245
Thelypteris thelypteroides, see Thelypteris palustris, 522 Venus’s hair, see Adiantum capillus-veneris, 161
Thelypteris torresiana, see Macrothelypteris torresiana, 363 Victorian brake, see Pteris ensiformis ‘Victoriae’, 476
Thelypteris uliginosa, see Macrothelypteris torresiana, 363 villous lip fern, see Cheilanthes villosa, 243
Thelypteris cultivars vine fern, see Microgramma heterophylla, 372
‘K. O. Sessions’, see Thelypteris acuminata, 518 vine spike moss, see Selaginella willdenovii, 506
‘Lepida’, see Thelypteris patens, 523 Virginia chain fern, see Woodwardia virginica, 539
Themelium, 333 Virginia polypody, see Polypodium virginianum, 452
thick-stemmed wood fern, see Dryopteris crassirhizoma, Vittaria, 530
304 Vittaria elongata, 530
Todea, 74, 526 Vittaria lineata, 101, 530
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walking fern, see Camptosorus, 228 woodrustic fern, see Polystichum dudleyi, 458
Wallich’s wood fern, see Dryopteris wallichiana, 320, Woodsia, 531
color plate 27 Woodsia alpina, 532
wall-rue, see Asplenium ruta-muraria, 203 Woodsia glabella, 532
water clover, see Marsilea, 365 Woodsia ilvensis, 532
water fern, see Ceratopteris 232; see also Marsilea Woodsia manchuriensis, 533
schelpeana, 368; Thelypteris nevadensis, 520 Woodsia obtusa, 92, 98, 533
water spangle, see Salvinia, 493 Woodsia oregana, 98, 534
water sprite, see Ceratopteris, 232 Woodsia polystichoides, 534
water tassel fern, see Lycopodium squarrosum, 360 Woodsia scopulina, 535
wax-dot maiden fern, see Thelypteris resinifera, 524 Woodsia subcordata, 535
western bracken, see Pteridium aquilinum var. pubescens, Woodwardia, 535
471 Woodwardia areolata, 539
western cliff fern, see Woodsia oregana, 534 Woodwardia chamissoi, see Woodwardia fimbriata, 537
western five-finger fern, see Adiantum aleuticum, 160 Woodwardia fimbriata, 537
western holly fern, see Polystichum scopulinum, 464 Woodwardia martinezii, 538
western oak fern, see Gymnocarpium disjunctum, 335 Woodwardia orientalis, 69, 70, 537
western polypody, see Polypodium hesperium, 445 Woodwardia radicans, 65, 69, 95, 537, color plate 50
western sword fern, see Polystichum munitum, 460 Woodwardia ×semicordata, 538
West Indian tree fern, see Cyathea arborea, 253 Woodwardia spinulosa, 538
whisk fern, see Psilotum, 470 Woodwardia unigemmata, 539
white-rabbit’s-foot fern, see Davallia tyermannii, 276 Woodwardia virginica, 92, 95, 539
Whiteside’s crisped broad-buckler fern, see Dryopteris woolly lip fern, see Cheilanthes tomentosa, 243
dilatata ‘Crispa Whiteside’, 307 woolly tree fern, see Dicksonia fibrosa, 282
widespread maiden fern, see Thelypteris kunthii, 520 Wright’s lip fern, see Cheilanthes wrightii, 243
winged beech fern, see Phegopteris decursive-pinnata, 418
winter grape fern, see Botrychium biternatum, 227 yellow-golden-scaled male fern, see Dryopteris affinis, 298
winter scouring rush, see Equisetum hyemale, 328
wood fern, see Dryopteris, 296; see also Thelypteris, 517 Zygophlebia, 333
woodland bladder fern, see Cystopteris protrusa, 260