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VOLUME 31 Nos.

1&2 2014
Volume 31 (2014)

EDITOR-IN-CHIEF : S. C. VERMA
EDITOR : S. P. KHULLAR
CO-EDITOR : H. K. CHEEMA
Editor-in-Chief : S. C. Verma
Co-editor : H. K. Cheema
Editor : S. P. Khullar
2014

THE INDIAN FERN SOCIETY


Correspondence Address :
Prof. S. P. Khullar, 1633 Sector 7-C, Chandigarh. E-mail : sp.khullar@gmail.com
Title of the Journal registered with Registrar, Newspapers for India vide his letter No. 35/7/8/6 RN-2
dated 11 February 1986 (Registration No. 42404/85)
THE INDIAN FERN JOURNAL
CONTENTS
Volume 31 Nos. 1-2 2014

New Frontiers of Fern Ecology


by Klaus Mehltreter 1-9
Re por
Repor ting in situ A po
porting pogg amous Abor ti
Aborti
tivv e Embr
Embryy os within Me
Megg aspor es of some Species of
aspores
Isoetes from Rajasthan.
by B. D. Sharma, D. R. Bohra, O. P. Suthar and R. Harsh 10-16
T he Gen us Ophio
Genus glossum fr
Ophioglossum om Wester
from esternn Gha ts of India
Ghats
by Sachin Patil and Meena Dongare 17-26
Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes of some Ferns of
Gang etic West Beng
Gangetic al
Bengal
by Ripan Paul, Tuhinsri Sen, U. Sen and S. Rahaman 27-39
Adiantum hispidulum Sw ar
Swar tz – A Ne
artz w Re
New por
portt fr
Repor om Sena
from pa
Senapa ti Distr
pati ict, Manipur
District,
by Th. Sunita Devi and Ajit Kumar Das 40-45
Ethno-Medico-Botanical Studies on some Pteridophytes of Rajasthan
by R. P. Kanther and Dilip Gena 46-52
Phenetics of South Indian Trichomanoid Ferns
by P. K. Vidya Varma, K. V. Mohanan and P. V. Madhusoodanan 53-68
Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) of South India
by P. K. Vidya Varma and P. V. Madusoodanan 69-89
Microstructural Observations on the Development of Gametophytes and Oogenesis in
Equisetum arvense
by Ge Wang, XI-Ling Dai, Quan-Xi Wang and Jian-Guo Cao 90-101
Some Ad ditions to the Pter
Additions idoph
Pteridoph yte Flor
idophyte Floraa of Himac hal Pr
Himachal adesh (W
Pradesh (W.. Himala
Himalayy a)
by S. P. Khullar and Sunil Verma 102-111
GC-MS Pr of
Prof ile of Lyg
ofile odium ffle
ygodium le xuosum L. (L
lexuosum yg
(Lyg odiaceae) – A Medicinal F
ygodiaceae) er
Fer
ernn fr om
from
Nor th Wester
North esternn Gha ts
Ghats
by Kanchan Dubal and Manisha Kale 112-117
Photosynthetic Pigments and their De
Degg r ada tion under Hyper
adation ther
Hyperther mia in Tw o Species of
thermia
Sela
Selagg inella Beauv
Beauv.. fr om Rajasthan
from
by Kanta Meena, Shahdab Hussain and B. L. Yadav 118-123
Discovery of Diploid Cytotype of Helminthostachys zeylanica (L.) Hook
(Helminthostachyaceae - Pteridophyta)
by Shashank Kumar Singh, S. Dominic Rajkumar, Shobhit Kumar Srivastava and
Ravi Pratap Gautam 124-131
Helminthostac
Helminthostachh ys zzee ylanica P opula
Popula tions Need Ser
opulations ious Attention : Disc
Serious losur
Disclosur es fr
losures om
from
Unpublished Observations
by Hit Kishore Goswami 132-135
Cytolo
Cytologg y of T hr ee Asplenium species fr
hree om Uttar
from akhand
Uttarakhand
by Anurita Sharma 136-138
Tectar ia puber
ectaria ula (Desv
puberula .) C. Chr
(Desv.) Chr.. (Dr
(Dryy opter idaceae : PTERIDOPHYT
opteridaceae A), A Ne
PTERIDOPHYTA), Neww Recor
Recordd ffor
or Asia
by Raju Antony, C. R. Fraser-Jenkins, N. Mohanan and Cheriyan P. Koshy 139-142
F er ns of the Aa
erns bshar F
Aabshar or
For est ar
orest ea (Kanda
area gha
(Kandagha t, Dist. Solan), Himac
ghat, hal Pr
Himachal adesh, West Himala
Pradesh, Himalayya
by Anurita Sharma 143-151
Some Comments on the P
Paa per "Na tur
"Natur al A pospor
tural posporyy in Pter is ar
Pteris g yr
arg aea T. Moor
yraea Mooree fr om
from
South INDIA" bby
y Raju Anton
Antonyy, S M Shar eef and N Mohanan in Indian F
Shareef er
Fer
ernn
Journal 29 : 148-152
Comments by No. 1 : Harvinder K. Cheema; No 2. Reply by Raju Antony 152-153
Is Huperzia hamiltonii (Spr eng
(Spreng .) Tr e vis. A Himala
eng.) Himalayy an Endemic?
An Empirical Evaluation using Species Distribution Modeling
by Nawal Shrestha and Xian-Chun Zhangi 154-161
Invasion of Salvinia molesta D. S. Mitchell. (African Payal) in Kerala and its Management
Madhusoodanan P. V., Sreerenjini V. K., Smitha R. B. and Prakash Kumar R. 162-172
Ecology of South Indian Pteridophytes
Madhusoodanan P. V., Prakash Kumar R., Sreerenjini V. K. and Smitha R. B. 173-186
Life Members of Indian Fern Society 2009-2014 187-189
Bibliography of Indian Pteridology (2013-2014)
Compiled by Dr. R. Mukhopadhyay and Prof. S. P. Khullar 190-200
Auditor's Report 201-202
Statement about IFJ Ownership 205

FR ONT CO
FRONT VER PICTURE
COVER
d
d

BA CK CO
BACK VER PICTURE
COVER
d
d
Indian Fern J. 3 1 : 1-9 (2014) ISSN 0970-2741

NEW FRONTIERS OF FERN ECOLOGY


FRONTIERS
KLAUS MEHLTRETER*
Instituto de Ecología, A. C., Xalapa, Ver., Mexico

(Received June 22, 2014; Revised Accepted July 14, 2014)

ABSTRACT
Fern ecology is a research discipline that received increasing attention during the last
20 years, contributing a large number of novel insights into ecological aspects of ferns and
their functional role in different ecosystems. In this contribution, new frontiers of fern ecology
are elucidated for three research areas: interactions of ferns with other organisms, nutrient
ecology, and applied fern ecology.

INTRODUCTION

Several prejudices refrained researchers from studying the ecology of ferns. Because
ferns and lycophytes have been considered as the oldest and least developed group of
vascular plants when compared to gymnosperms and angiosperms, it was erroneously
concluded that they cannot compete with coexisting seed plants, that they do not play any
major role in ecosystem functions, and that they also would have developed fewer
interactions with animals, especially insects (Mehltreter et al. 2010).
However, since the publication of Pryer et al. (2004), it became apparent that
lycophytes first diverged from euphyllophytes and those diverged into the two sister clades
of ferns (monilophytes) and seed plants during the Devonian about 380 mya. Although
angiosperms took over the dominant role in the vegetation since the late Cretaceous, slowly
replacing ferns, cycads and conifers, ferns may be still an important component of the flora
at some locations (e.g. remote tropical islands) and habitats (e.g., montane rainforests). Even
in some temperate or subtropical regions such as the Northwestern US or Eastern Australia
one might be overwhelmed by the dominant role ferns play in the forest understory. Some
fern species are also a lot more robust (e.g., tolerant against disturbance) than generally
assumed. In tropical disturbed sites, Gleicheniaceae such as Sticherus bifidus ,
Diplopterygium bancroftii in Mexico, and Dicranopteris linearis in Hawaii take over along
roadsides, landslides and disturbed ridges. Tree ferns in Australia survive fires of the
Eucalypt forests and are typically the first species to sprout (Fig. 1). With 9500 species of
ferns and 1380 species of lycophytes worldwide, both groups together comprise ca. 4% of
the species diversity of vascular plants, can reach 5-13% of the diversity in humid tropical
mountains, and contribute 40-70% of the ecosystems of remote tropical oceanic islands such
as Easter Island or Tristan da Cunha (Kessler 2010).
Fern research was strongly focused on systematics, phylogenetics, morphology,
anatomy, and developmental plant biology, and especially in Asia some applied research was
performed on the ethnobotany and medicinal uses of ferns. Future directions of ecological

* E-mail : kmehltreter@gmail.com
2 Indian Fern Journal Volume XXXI (2014)

studies of ferns were discussed in Walker et al. (2010) and here I will review some of the
current advances of three main subjects: interactions of ferns with other organisms, nutrient
ecology, and applied fern ecology.
INTERACTIONS OF FERNS WITH OTHER ORGANISMS

Interactions with plants


Which organisms foster or restrain the development of ferns? In the few cases
where ferns interact with algae these interactions seem to be symbiotic, favoring both
organisms (e.g. blue green algae of Anabaena in the floating fern Azolla, Fig. 2), but it
would be surprising that no negative interactions may occur between algae and ferns,
especially during the early developmental stages of the fern gametophytes.
Mosses and ferns share several features of their developmental biology (e.g.,
flagellate sperm cells that require water to swim) and their diversity peaks in humid tropical
and temperate forests. Consequently, ferns are commonly found growing together with
mosses, and we have to assume that they compete for the same resources. Mosses may indeed
colonize bare soil or tree trunks first and prepare the habitat by accumulation of organic
matter and storage of water, facilitating the subsequent establishment of ferns. On the other
hand, many boulders of mosses are free of any ferns. These fern-free moss boulders could
be the consequence of the exclusion of ferns by allelopathic inhibition of fern spore
germination or by competition of mosses with fern gametophytes. Clearly, the interactions
of mosses and ferns require further attention. Interactions with seed plants are mainly
competitive, affecting ferns negatively (Fig. 2), because ferns are growing mainly in the
understory of the larger gymnosperm and angiosperm trees. However, there is also a clear
positive interaction between trees and ferns, because trees provide a welcome substrate for
many epiphytic ferns, especially in tropical humid environments.
Interactions with fungi
Interactions of ferns and fungi have already received a lot more attention than
interactions of ferns with plants and animals. However, most studies have sampled the
mycorrhizal fungi of fern roots and currently we know that arbuscular mycorrhizae are very
common in ferns (e.g. 74% are colonized in Hawaii, Gemma et al. 1992), and more
recently, the less frequent ectendomycorrhizae with fern-ericoid mycorrhizae and dark
septate fungi, both Ascomycota, have been described (Siddiqui & Pichtel 2008). Although
most mycorrhizae are facultative, ferns benefit from these fungi by extending their water
absorption surface and mineral nutrient uptake (Mehltreter 2010a). Fungal parasites have
been studied mostly to control specific fungal diseases of ornamental ferns, such as
anthracnose on Rumohra adiantiformis (Leahy et al . 1995). The newest fern research,
however, is focusing on the supposedly neutral endophytes, symptomless fungi that develop
within the ferns without any signs of causing damage to the plant. However, once the fern
dies, the endophytes become apparent and develop into their reproductive stage (Petrini
Klaus Mehltreter : New Frontiers of Fern Ecology 3

Figures 1. Tree ferns ( Cyathea sp.) are the first plants to sprout after a fire in an Eucalyptus forest
in Australia.

1991). The number of those endophytic fungi has been completely underestimated, because
many of them are host and tissue-specific. Some endophytic fungi are even able to protect
ferns from infections of more harmful, parasitic fungi. In summary, fungi affect ferns in
all ways, but the positive effect of mycorrhizae and the negative effect of parasitic fungi
might be in a balance (Fig. 2).
Interactions with animals and humans
humans.
Most interactions of ferns with animals are affecting fern growth and reproduction
negatively (Fig. 2). However, since Brues (1920), it was hypothesized that ferns are less
attacked by animals than angiosperms, and Ehrlich and Raven (1964) assumed consequently
that ferns would have developed fewer defensive strategies against herbivorous animals.
Balick et al. (1978) counted fewer fern-insect interactions than angiosperm-insect interactions
and concluded that this bias is consequence of the lack of several morphological structures
4 Indian Fern Journal Volume XXXI (2014)

F igur es 3. Dicksonia antarctica is protected fern species, cultivated in Australia as ornamental plant
igures
and sold worldwide.
F igur es 4. Asplenium nidus in Taiwan is cultivated because of its edible young leaves.
igures
Klaus Mehltreter : New Frontiers of Fern Ecology 5

F i g u r e s 2 . Assumed quantitative interactions of ferns with other organisms: positive interactions


(empty white arrows), negative interactions (filled black arrows)

in ferns such as flowers, fruits and seeds. Coe et al. (1987) assumed that ferns are less
attacked because of their lower nutritional value, because shade plants have commonly lower
nutrient concentrations than sun plants.
Most of the former hypotheses, however, cannot withstand closer examination,
perhaps with exception of Balick´s idea of a lack of morphological complexity when ferns
and lycophytes (with their ca. 11.000 species) are compared with over 300.000 angiosperm
species. It seems that fern-insect interactions have been studied so infrequently, that our
current knowledge on plant-insect interactions is extremely biased towards angiosperms. In
fact, thousands of insects feed on ferns, and many of the fern feeding insect species are
still undiscovered.Two recent studies provide examples about the huge number of insects
that can be found on ferns. Fountain Jones et al. (2012) found 108 beetle species on 80
individual trunks of Dicksonia antarctica in Tasmania, and Fayle et al. (2012) identified 71
symbiontic ant species on 83 individual plants of the epiphytic Asplenium nidus in Borneo.
However, not all insects are damaging ferns; some are protecting them by attacking their
enemies. Because a lot of fern feeding insect species are specialists and feed on only a few
6 Indian Fern Journal Volume XXXI (2014)

fern species, some of the parasitoids of the fern feeders are even more specialized, and those
parasitoids are finally protecting the ferns from an overpopulation of the fern herbivorous
species. A good example has been reported by Kula et al. (2012), who found specific
parasitoids of the geometrid Psaliodes strigosa Warren, which feeds on Diplazium costale.
However, the trophic interactions do not stop at this level. In some cases, there are
hyperparasitoids, which can harm the parasitoid as well as the herbivore, so that their
positive or negative effect on the ferns cannot be easily predicted (Morse 2009). There is
no doubt that there are many more insect species to discover on ferns: herbivores,
parasitoids and hyperparasitoids.
Few mutualistic interactions of ferns with insects have been reported so far and most
of those point to ants. The nectary bearing ferns provide the ants with some reward,
whereas the ants protect the ferns from herbivores. However, our current knowledge of the
efficiency of the ant defense mechanism of nectary bearing fern species is restricted to only
two fern species, Pleopeltis plebeia (Koptur et al. 1998) and Pleopeltis crassinervata (Koptur
et al. 2013).Other new insights came from herbivory studies of three slug species on three
ferns, Athyrium filix-femina, Dryopteris filix-mas and Gymnocarpium robertianum. Slugs
may not digest all consumed fern spores and contribute to their endozoochoric dispersal in
a short range (Boch et al. 2013).
Few vertebrates have been reported to feed on ferns. Some exceptions are the
Azores bullfinch and the Japanese bat. The Azores bullfinch feeds on sporangia of Culcita
macrocarpa and Woodwardia radicans and leaves of Osmunda regalis and Pteridium
aquilinum during winter when its favorite food sources are scarce (Ramos 1994, 1995). The
Japanese bat Pteropus pselaphon feeds on the bird-nest fern Asplenium setoi and because
some fern spores are still viable in the bat feces, the endozoochoric dispersal of the fern
spores seems to be possible (Sugita et al. 2013). These new results on fern-animal
interactions will make it necessary to reconsider our current perception which hypothesized
that ferns are only wind dispersed.
Humans are clearly the worst threat to fern diversity by destroying and modifying
natural fern habitats. However, some fern conservation efforts finally protect and help some
of the endangered fern species to thrive.
NUTRIENT ECOLOGY

The most basic reasons to study the nutrient composition of ferns are to evaluate
species for their possible nutritional value as food source for humans and possible
applications for phytoremediation. However, ferns may be also a relevant food source to
herbivores and play an important role in nutrient allocation and nutrient cycling within
ecosystems (Richardson & Walker 2010). Historically, it was assumed that ferns have mainly
lower N, P and Ca concentrations than angiosperms, with exception of tree ferns
(Richardson & Walker 2010). In contrast to this assumption, ferns have been a valuable
Klaus Mehltreter : New Frontiers of Fern Ecology 7

food source to dinosaurs, the largest land animals on Earth (Hummel et al. 2008). A
screening of the N, P, K and Ca content in dry leaf biomass of 23 fern species in relative
nutrient rich riparian soils (Mehltreter, unpubl. data), demonstrated that ferns can reach high
nutrient levels (e.g. means of >3% N, 0.2% P, 1% K and 0.6% of Ca), but more
surprisingly their nutrient composition is highly variable even when soils are nutrient rich.
Consequently, we have to assume that some species discriminate some elements (e.g. cations)
during nutrient uptake. For instance, Asplenium riparium had 250% more Ca than the
average fern species at the riparian sites and Sticherus palmatus less than 10% of the average
Ca content. This enormous variation indicates that further studies are required to understand
the physiological reasons and ecological consequences of such nutrient discrimination. The
specific capacity of some fern species such as Pteris vittata to hyper accumulate arsenic (Ma
et al. 2001) is another example of nutrient discrimination, which is used for
phytoremediation, but we have to assume that such physiological changes should also result
in ecological consequences such as reduced herbivore damage.
APPLIED FERN ECOLOGY

Although the use of Pteris vittata for phytoremediation might be currently the most
innovative approach of applied fern ecology (Ma et al. 2001), many other practical uses of
ferns remain underestimated or underdeveloped. The market for ornamental ferns is huge,
but is restricted to varieties of a small percentage of fern species (e.g. Dicksonia antarctica;
Fig. 3). Relatively few ferns are used as food in Europe and the Americas (e.g., Matteuccia
struthiopteris in the United States), but there is a huge demand for new alternative
delicacies, and new methods of cultivation are required to cultivate those species (Thakur
et al. 1998). In Asia, especially China, India, Taiwan, Malaysia and the Philippines several
more species are collected from the wild and consumed (e.g. Asplenium nidus; de Winter
& Amoroso 2003; Fig. 4) but more species should be taken into cultivation and analyzed
for their chemical composition as well as nutritional, medicinal and possible industrial
properties.
About 60 fern species have become problematic as weeds (Robinson et al. 2010)
and there is an urgent need for ecological studies of those fern species to understand the
specific characteristics and environmental circumstances that make them a weed and to
develop methods that may improve current management strategies. On the other hand,
deforestation and changes in land use may cause a threat to a considerable number of ferns,
but a risk assessment has been completed for less than 2% of the approximately 11.000
fern and lycophyte species (Mehltreter 2010b). Fern conservation studies are imperative to
increase our knowledge about the reasons for the vulnerability of some species, to study
their habitat requirements, to cultivate them in laboratories and greenhouses, and to protect
them with efficient management strategies in situ.
8 Indian Fern Journal Volume XXXI (2014)

ACKNOWLEDGEMENTS

Research was supported by CONACYT (project no. CB2008-101542-F) and the


Instituto de Ecología, A. C. (project no. 20030-10796).
REFERENCES

B O C H S , B E R L I N G E R M , F I S C H E R M , K N O P E , N E N T W I G W, T Ü R K E M & P R AT I D 2 0 1 3 F e r n
and bryophyte endozoochory by slugs Oecologia 1 7 2 : 817-822
BA L I C K M J , F U RT H D G & C O O P E R - D R I V E R G 1 9 7 8 B i o c h e m i c a l a n d ev o l u t i o n a r y a s p e c t s o f
arthropod predation on ferns Oecologia 3 5 : 55-89
BRUES C T 1920 The selection of food plants by insects, with special reference tolepidopterous
larvae Amer Nat 5 4 : 313-32
C O E M L , D I L C H E R D L , FA R L OW J O , JA R Z E N D M & RU S S E L L D A 1 9 8 7 D i n o s a u r s a n d l a n d
plants. In : FRIIS E M, CHALONER W G & CRANE P R (eds) The Or igins of Angiosper ms
and their Biological Consequences Cambridge University Press, Cambridge, pp 225-58
DE WINTER W P & AMOROSO V B (eds) 2003 Crypto gams : Ferns and Fer n Allies, Plant Resour ces
of South-East Asia, 15 Bac khuys, Leiden
EHRLICH P R & RAVEN P H 1964 Butterflies and plants: a study in coevolution Evolution 1 8 : 586-
608
FAYLE T M, EDWARDS D P, TURNER E C, DUMBRELL A J, EGGLETON P & FOSTER W A 2012
Public goods, public services and by-product mutualism in an ant-fern symbiosis Oikos
1 2 1 : 1279-1286
FOUNTAIN-JONES N M, MCQUILLAN P B & GROVE S 2012 Beetle communities associated with the
tree fer n Dicksonia Antarctica La bill in Tasmania A ustral J Entomol 5 1 : 154-165
GEMMA J N, KOSKE R E& FLYNN T 1992 Mycorrhizae in Hawaiian pteridophytes : occurrence and
evolutionary significance Amer J Bot 7 9 : 843-52
H U M M E L J , G E E C T, S Ü D E K U M K H , S A N D E R P M , N O G G E G & C L AU S S M 2 0 0 8 I n v i t r o
digestibility of fern and gymnosperm foliage : implications for sauropod feeding ecology and
diet selection Proc Roy Soc, Ser B 2 7 5 : 1015-1021
L E A H Y R , S C H U B E RT T, S T R A N D B E R G J , S TA M P S B & N O R M A N D 1 9 9 5 . A n t h r a c n o s e o f l e a t h e r
leaf f ern Flor ida D epar tment of Agr icultur e and Customer Services, Plant Pathology Cir cular
3 7 2 4 pp
K E S S L E R M 2 0 1 0 B i o g e o g r a p h y o f f e r n s I n : M E H LT R E T E R K , WA L K E R L R & S H A R P E , J M
(eds) Fern ecology Cambridge University Press Cambridge pp 22-60
KO P T U R S , R I C O - G R AY V & PA L AC I O S - R I O S M 1 9 9 8 A n t p r o t e c t i o n o f t h e n e c t a r i e d f e r n
Polypodium plebeium in central Mexico Amer J Bot 8 5 : 736-739
KO P T U R S , PA L AC I O S - R I O S M , D Í A Z - C A S T E L A Z O C , M AC K AY W P & R I C O - G R AY V 2 0 1 3
Nectar secretion on fern fronds associated with lower levels of herbivore damage : field
e x p e r i m e n t s w i t h a w i d e s p r e a d e p i p hy t e o f M e x i c a n c l o u d f o r e s t r e m n a n t s A n n B o t 1 1 1 :
1277-1283
KULA R R, DIX-LUNA O & SHAW S R 2012 Review of Ila tha Fischer (Hymenoptera : Braconidae :
Alysiinae), including descriptions of six new species and a key to species Proc Entomol Soc
Washington 1 1 4 : 293-328
M A L Q , KO M A R K M , T U C , Z H A N G W H , C A I Y & K E N N E L L E Y E D 2 0 0 1 A f er n t h a t hy p e r
accumulates arsenic Nature 4 0 9 : 579
MEHLTRETER K 2010a Interactions of ferns with fungi and animals In : MEHLTRETER K, WALKER
Klaus Mehltreter : New Frontiers of Fern Ecology 9

L R &SHARPE, J M (eds) Fern ecology Cambridge University Press, Cambridge, pp 220-254


M E H LT R E T E R K 2 0 1 0 b F e r n c o n s e r va t i o n I n : M E H LT R E T E R K , WA L K E R L R & S H A R P E , J M
(eds) Fern ecology Cambridge University Press, Cambridge, pp 323-359
M E H LT R E T E R K , WA L K E R L R & S H A R P E , J M ( e d s ) 2 0 1 0 F e r n e c o l o g y C a m b r i d g e U n iv e r s i t y
Press Cambridge 444 pp
MORSE D H 2009 Four-level interactions: herbivore use of ferns and subsequent parasitoid –
h y p e r p a r a s i t o i d p e r f o r m a n c e Ecol Entomol 3 4 : 2 4 6 - 2 5 3
P E T R I N I O 1 9 9 1 F u n g a l e n d o p h y t e s o f t r e e l e ave s I n : A N D R E W S J A & H I R A N O S S ( e d s )
Microbial ecolo gy of leaves Spring er New Yor k pp 179–97
P RY E R K M , S C H U E T T P E L Z E , W O L F P G , S C H N E I D E R H , S M I T H A R & C R A N F I L L R 2 0 0 4
Phylogeny and evolution of ferns(Monilophytes) with a focus on the early leptosporangiate
diverg ences Amer J Bot 9 1 : 1582-1598
RAMOS J A 1994 Fern frond f eeding by the Azor es bullfinch J Avian Biol 2 5 : 344-347
RAMOS J A 1995 The diet of the A zor es bullfinch Pyr rhula m ur ina and flor istic var i ation within its
range Biol Conserv 7 1 : 237-249
RICHARDSON S & WALKER L R 2010 Nutrient ecolog y of ferns In : MEHLTRETER K, WALKER L
R &SHARPE, J M (eds) Fern ecology Cambridge University Press Cambridge pp 111-139
ROBINSON R C, SHEFFIELD E & SHARPE J M 2010 Problem ferns: their impact and management
I n : M E H LT R E T E R K , WA L K E R L R & S H A R P E , J M ( e d s ) F e r n e c o l o g y C a m b r i d g e
University Press Cambridge pp 256-322
SIDDIQUI Z A & PICHTEL J 2008 Mycor rhizae: an over view. In: SIDDIQUI Z A, AKHTAR M S &
F U TA I K ( e d s ) M y c o r r h i z a e : S u s t a i n a b l e A g r i c u l t u r e a n d F o r e s t r y S p r i n g e r S c i e n c e &
Business Media B V, pp 1–35
SUGITA N, OOTSUKI R, NURAKAMI T F N & UEDA K 2013 Possible spore dispersal of a bird-nest
f ern Asplenium setoi by Bonin f lying f o xes Pter opus pselaphon Mammal Study 3 8 ( 3 ) : 225-
229
T H A K U R R C , H O S O I Y & I S H I I K 1 9 9 8 R a p i d i n v i t r o p r o p a g at i o n o f M a t t e u c c i a s t r u t h i o p t e r is
(L.) Todaro – an edible fer n Pl Cell Rep 1 8 : 203-208
WALKER L R, MEHLTRETER K & SHARPE J M 2010 Current and future directions of fern ecology
I n : M E H LT R E T E R K , WA L K E R L R & S H A R P E , J M ( e d s ) F e r n e c o l o g y C a m b r i d g e
University Press Cambridge pp 360-378
Indian Fern J. 3 1 : 10-16 (2014) ISSN 0970-2741

REPORTING IN SITU APOGAMOUS ABOR


REPORTING TIVE EMBR
ABORTIVE EMBRYY OS WITHIN
MEGASPORES OF SOME SPECIES OF ISOETES FR OM RAJ
FROM ASTHAN
RAJASTHAN
B. D. SHARMA1*, D. R. BOHRA2**, O. P. SUTHAR3*** AND R. HARSH4****
1
Kath Mandi, Narnaul-123001
2
4B/7 Partap Nagar, Jodhpur-342001
3
Botany Dept., Govt. Girls College, Balotra-344022
4
Botany Dept., M.S. Govt. Girls College, Bikaner-334001

(Received November 4, 2013; Revised Accepted 16 May 2014)

ABSTRACT
Thin serial sections through the megasporangial portions of the four species of Isoetes
from Rajasthan namely - I. coromandelina, I. tuberculata , I. reticulata and I. rajasthanensis ,
show distinct three layers of the wall of trilete megaspores. Both functional (fertile) and sterile
spores occur intermingled in a megasporangium. A fertile megaspore has 'spore content' while
the sterile one is empty. The spore content either fills completely the spore cavity or it reduces
and changes to U or C or a triarch structure. Free floating nucleated and enucleated cells are
visible in the spore content. Some of the nucleated cell/cells make apogamous abortive embryos
which are of different shapes and sizes. Circular cavity or cavities and rhizoids are also visible
in many of the embryos. Archegonia are not yet seen in the sections of the megaspores. This
paper includes the detailed report of apogamous, abortive embryos in situ inside the megaspores
of Isoetes.
d s : Lycopod, Quillwort, Megaspores, in situ Abnormal Embryogeny
K e y Wo rrd

INTRODUCTION

A good number of pteridophytes occur during rainy season in the Aravalli Hills,
Chambal ravines and Harauti plateau in Rajasthan (Bir & Verma 1963, Sharma et al. 1988,
Sharma 2003, Yadav & Meena 2009, Gupta & Bhardwaj 2010). Isoetes grows at several
places in the Aravali Hills and South-Eastern portion of the state e.g. Mt. Abu, Daosa, Atru,
Jhalawar, Chittorgarh etc. It is represented by four species i.e. I. coromandelina L.f.,
I. tuberculata Gena and Bhardwaja, I. reticulata Gena and Bhardwaja and I. rajasthanensis
Gena and Bhardwaja (Sharma & Bohra 1978 Sharma et al. 1985, 2008, Gena & Bhardwaja
1984). Each leaf has in the basal adaxial portion a large sporangium (mega or
microsporangium) (Gifford & Foster 1989). In some of the species like I. coromandelina
and I. pantii mixed, bisexual sporangia are also reported but rare (Singh et al. 1984,
Goswami & Bhu 1998). A megasporangium has 200-600 or more trilete megaspores with
various kinds of ornamentations on the exine like tuberculate, reticulate, spinose etc. (Pant
& Srivastava 1962, Srivastava et al. 1993, 1995, Goswami 2004, Sharma et al. 2008).
Abnormal sterile spores also occur frequently like smaller spores, larger spores, jointed
spores, fused spores, enucleate spores (Bohra et al. 1980, Pant & Srivastava 1962, Panigrahi
1981, Sharma & Purohit 2013). The triploid I. coromandelina is reported to show a peculiar
* E-mail : bdsharma14@yahoo.com, ** drbohra14@gmail.com, *** opsuthar59@yahoo.in,
**** rharshpalaeobotanist@yahoo.com
B D Sharma et al. : Reporting in situ Apogamous Abortive Embryos within Megaspores of some Species 11

PLATE I
PLATE

Fig. 1-12 : Megaspores of Isoetes sp. with abnormal embryos. 1. Megaspore with a cavity in the spore
content. 2 , 3 . Megaspores with dark coloured embryos in C-shaped spore contents. 4 - 7 . Abnormal embryos
of different shapes and sizes. Rhizoids (?) in all. A cavity in 5 and a curved leaf like outgrowth in 6.
8-10. Irregular embryos and scattered cells. 11-12. Embryos in granular spore content. The three wall layers
of the megaspores also visible. (Bar 1–3 = μ 100, 4–12 = 40 μm).
12 Indian Fern Journal Volume XXXI (2014)

PLATE II
PLATE

Fig. 1-11. Sections of megaspores of Isoetes sp. with apogamous, abortive embryos. 1-6. Functional
megaspores with spore contents in various forms O, U, C, triarch. Small free floating nucleated cells and large
black bodies are abnormal embryos. 8-11. Enlargements of embryoids. Embryos of various shapes and sizes.
8. A cavity is visible. 9-11. Rhizoidal outgrowth the clearly seen (Bar 1–7 = 100 μm, 8–11 = 40 μm).
B D Sharma et al. : Reporting in situ Apogamous Abortive Embryos within Megaspores of some Species 13

mode of cytokinesis of the two unreduced nuclei, following abnormal meiosis, resulting in
two nucleate and two enucleate spores, both types being tetrahedral, and the latter being
small-sized (Verma 1960, 1961).
The life history of Isoetes has been studied by a number of earlier workers (like
Mettinius 1850, Hofmeister 1862, Kienitz-Gerloff 1881, Belajeff 1885, Farmer 1890,
Campbell 1891, Goebel 1897, La-Matte 1933, 1937 etc.). Majority of them studied spore
germination either in nature or by growing sporelings in petriplates (Taylor & Luebke
1986). La-Matte (1933) Tried to grow the megaspores of I. lithophila in the soil brought
from the collection site and used rainwater even then the success was not more than ten
percent. Even today despite enough advancement in biotechnology it is considered as a
difficult exercise to grow megaspores of Isoetes in the laboratory. Goebel (1897) had
reported for the first time the presence of apogamy in Isoetes. Ninan (1958) while working
on the cytology of pteridophytes specially Isoetaceae noticed apogamy in South Indian
material of Isoetes. Pant and Srivastava (1965) also reported apomictic germination of
spores in I. indica and I. panchananii. Srivastava et al. (1996) described in situ germination
of megaspores in I. panchganiensis and I. mahadevensis. Recently Sharma (2013) has
described in situ abnormal embryo formation inside the megaspores of the species of Isoetes
found in the state of Rajasthan. The present observations are based on the study of
microtome cut thin cross and longisections of megasporangial portions which have added
much to our knowledge about an abnormal embryogeny in Isoetes.
MATERIAL AND METHODS
ATERIAL

The material of the four species of Isoetes was collected from different places in
Rajasthan. I. tuberculata is acquatic while the other species are amphibious to terrestrial. In
I. tubersculata and I. reticulata microsporangia are yet to be reported (Gena & Bhardwaja
1984, Sharma et al. 1985). Microspores of the other two species are small and monolete.
However, in I. coromandelina a few trilete, microspores are also visible. Megasporangial
portions of the young and mature plants were collected and fixed in F.A.A. and then it was
preserved in 70% alcohol. The material was then processed for microtomy as suggested by
Johanson (1940). The slides were stained by the combination of safranin and haemotoxylin
and mounted in ditute canada balsom.
OBSERVAT I O N S
RV

A megasporangium of Isoetes encloses 200-600 or more trilete megaspores with


various kinds of ornamentations like tuberculate reticulate, spinose etc. on the exine. Both
functional (Plate fig. 1-6, Text fig. 2) and non functional spores (Platefig. 7) have identical
three layers of the wall i.e. the outermost is wide, light coloured and irregular in thickness
due to ornamentations (Text fig. 1, 2, 3, 12). The middle layer is thick, dark brown and
strong. The inner one is brown but comparatively thinner and encloses the ‘spore content’
in a functional spore. The sterile spores are empty (Plate fig. 7). The ratio between the
14 Indian Fern Journal Volume XXXI (2014)

fertile and sterile spores in a sporangium is variable in different species of Isoetes (Pant &
Srivastava 1962, Sharma et al. 1985, 2008, Panigrahi 1981). In I. coromandelina
Ekambaram and Venkatanathan (1933) found the ratio 50%-50%. Verma (1961) also
proposed 50% sterile (enucleated) and 50% fertile (nucleated) spores in a tetrad of this
species. Pant and Srivastava (1965) favoured this statement. But Sharma et al. (1985a)
noticed variations in the ratio of fertile and sterile spores in a megasporangium of I.
coromandelina and in other three species of Isoetes from Rajasthan. The spore content of a
functional megaspore consists of granular particles and starch grains. In the sections of some
of the megaspores the spore contents cover the entire space of spore cavity (Plate fig. 1).
While in others a small cavity (Text fig. 1) develops which increases in size and reduces
the spore content to U or C or a triarch structure (Plate fig. 1-6, Text fig. 2, 3). Spore
content has scattered nucleated and enucleated cells (Plate fig. 8-11, Text fig. 9, 10). The
former are circular and each has a dark coloured central nucleus. Sometimes these cells
occur in groups of 4 or 5 or more (Plate fig. 10). In some of the functional spores in
addition to free floating nucleated cells one or two large sized, regular or irregular shaped
dark coloured bodies are seen (Plate fig. 1-5, 8-11, Text fig. 2, 3). These are called
apogamous, abortive embryos in this paper. Out of hundreds of slides prepared, only in 25%
slides such embryoid bodies are visible. Further, out of the 20 sections present in a slide
only in 1-3 sections have the embryos. Shape and size of the embryoid bodies (Plate fig.
8-10, Text fig. 4-10) depend on the plane and position of sectioning of the megaspores. In
some of the embryos circular cavity/cavities (Plate fig. 8, Text fig. 5) are visible identical
to those figured by Hofmeister (1862, Plate XLVI). Small rhizoids like outgrowths are also
seen in many embryos (Plate fig. 9-11, Text fig. 4-10). The position of embryo is variable
i.e. it may be deep in the spore content (Text fig. 2, 3) or at the outer or inner periphery
(Plate fig. 9, 11, Text fig. 10, 11) or sometimes at the terminal position (Plate fig. 10). In
one or two embryos a long or curved (Text fig. 6, 8-10) root like structures has also been
seen. All these embryoid structures originate from the free floating nucleated cell/cells.
Archegonia could not be seen in any section studied for the present purpose.
The embryos are apogamous for want of microspores and spermatozoids. These are
also abortive because of their origin directly from the free floating nucleated cells of the
megagametophytes and absence of an archegenial structure which protects or covers the
normal embryos. Similarly there is no definite differentiation of organs like foot, root,
leaves etc. in these embryos. There are many ferns in which archegonia are not produced
and any cell of the prothallus may form an apogamous sporophyte e.g. Adiantum lunulatum
(Mehra & Gupta 1986, Sharma & Sharma 1992). Similar is the position in Isoetes, but
here the cells are free floating and not in a prothalloid form.
ACKNOWLEDGEMENTS

The authors wish to express thanks to Dr. Ravinder Singh for preparing the slides
and Ms Surya Shekhawat for assistance in photography.
B D Sharma et al. : Reporting in situ Apogamous Abortive Embryos within Megaspores of some Species 15

REFERENCES

BELAJEFF W 1885 Antheridien und spermatozoiden der hetrosporen Lycopodien Bot zeit 43 : 794-800
BIR S S & VERMA S C 1963 Ferns of Mount Abu Res Bull (N S) Panjab Univ Chandigarh India 1 4 : 187-
202
Bohra D R, Sharma B D & Singh R 1980 Isoetes in Rajasthan A study of spores Adv Pollen Spores Res 5 :
1-8
CAMPBELL D H 1891 Contribution to the life history of Isoetes Ann Bot 5 : 231-258.
EKAMBARAM T & VENKATANATHAN T N 1933 Studies on Isoetes coromandelina L I Sporogenesis J
Indian Bot Soc 1 2 : 191-225
FARMER J B 1890 Isoetes lacustris Ann Bot 5 : 37-62
GENA C B & BHARDWAJA T N 1984 Three new species of Isoetes L from Rajasthan J Bombay Nat Hist Soc
8 1 : 165-168.
GIFFORD E M & FOSTER S R 1989 Morphology and Evolution of Vascular Plants Freeman & Co New York
GOEBEL, K VON 1897 Ueber sprossbildung auf Isoetes blatten Bot Zeit 56 : 1-6
GOSWAMI H K 2004 Isoetes IV. Reproduction Bionature 24 : 13-24
Goswami H K & BHU I 1998 Bisexual sporangia in Isoetes pantii Bionature 18 : 31-36
HOFMEISTER W 1862 On the germination, development and fructification of higher cryptogamia and
fructification of the Coniferae Ray Soc London
JOHANSEN DA 1940 Plant Microtechnique Mc Graw Hill New York
KIENITZ–GERLOFF F 1881 Uber Wachatum and Zelltheilung in der Entwickelung der embryos von Isoetes
lacustris Bot Zeit 39 : 761-770, 785
LA MATTE C 1933 Morphology of the megagametophyte and the embryo-sporophyte of Isoetes lithophila
Am J Bot 2 0 : 217-233.
LA MATTE C 1937 Morphology and orientation of the embryo of Isoetes L Ann Bot (n s) 1 : 695-715
MEHRA P N & GUPTA A 1986 Gametophytes of the Himalayan ferns Botany Dept Punjab Univ Chandigarh
India
METTINIUS G 1850 Zur forlepflanger der geffasskryptogamen Beit Z Bot (from Hofmeister 1862)
NINAN CA 1958 Studies on the cytology and phylogeny of pteridophytes V Observations on Isoetaceae
J Indian Bot Soc 37 : 93-103.
PANIGRAHI G 1981 Systematics of the genus Isoetes L. (Isoetaceae) in Indian Biological Memoir 6 ( 2 ) :
129-138
PANT D D & SRIVASTAVA G K 1962 The genus Isoetes L in India Proc Natl Inst Sci India 28B : 242-280
PANT D D & SRIVASTAVA G K 1965 Cytology and reproduction of some Indian species of Isoetes L
Caryologia 3 0 : 239-251
SHARMA A & SHARMA B D 1992 Gametophytes of five species of Rajasthan ferns under cultural conditions
Bull Bot Surv India 32 : 151-161
SHARMA B D 2003 Fainting scenario of Rajasthan pteridophytes needs an attention Geophytology 3 2 :
25-28
SHARMA B D 2013 Discovery of in situ apogamous abortive embryos inside the megaspores of Isoetes L
Phytomorphology 6 3 : 153-156
SHARMA B D & BOHRA D R 1978 Isoetes in Rajasthan – anatomy of the rhizomorph of Isoetes
coromandelina Acta Bot Indica 6 : 122-127
SHARMA B D & PUROHIT S N 2013 Spores, germination and apogamy in Isoetes of Rajasthan India
Geophytology 43(2) : 177-182
16 Indian Fern Journal Volume XXXI (2014)

SHARMA B D, SINGH R & VYAS M S 1985 Isoetes in Rajasthan – A systematic study Bull Bot Surv India
27 : 58-62
SHARMA B D, BOHRA D R & VYAS M S 1985a Sporogenesis in Isoetes L Bionature 5 : 9-13
SHARMA B D, BOHRA D R, SUTHAR O P & HARSH R 2008 A brief account of the history and various
aspects of researchers on Isoetes Linn Indian Fern J 25 : 167-201
SHARMA S N, GENA C B & VERMA S 1988 A survey of the pteridophytic flora of Harauti plateau
Rajasthan Indian Fern J 5 : 58-62
SINGH R, BOHRA D R & SHARMA B D 1984 Mixed sporangia in Isoetes coromandelina Pollen et spores
25 : 41-47.
SRIVASTAVA G K, PANT D D & SHUKLA P K 1993 The genus Isoetes in India Amer Fern J 83 : 105-119
SRIVASTAVA G K, SHUKLA P K & WAGAI S O 1995 Megaspores of some Indian species of Isoetes L and
compared to living and fossil forms Palaeobotanist 4 4 : 215-224
SRIVASTAVA G K, SHUKLA P K, WAGAI S O & SRIVASTAVA M 1996 In situ germination of megaspores
of some Indian species of Isoetes L Nat Acad Sci Letters 19 : 191-194
TAYLOR W C & LUEBKE N T 1986 Germination of spores and growing sporelings of aquatic Isoetes Amer
Fern J 76 : 21-24
VERMA S C 1960 Enucleate spores in I. coromandelina L. Caryologia 13 : 117-123
VERMA S C 1961 Cytology of Isoetes coromandelina L. Amer Fern J 51 : 99-104
YADAV B L & MEENA D K 2009 The ferns and fern allies of the Bundi and Kota districts of Rajasthan,
India. Indian Fern J 26 : 149-160
Indian Fern J. 3 1 : 17-26 (2014) ISSN 0970-2741

THE GENUS OPHIOGLOSSUM FROM WESTERN GHA


FROM TS OF INDIA
GHATS
SACHIN PATIL1* AND MEENA DONGARE1
1
Department of Botany, Shivaji University, Kolhapur

(Received August 11, 2013; Revised Accepted Nov. 30, 2014)

ABSTRACT
The present paper deals with study of the genus Ophioglossum from Western Ghats
of India. It reports eight species of Ophioglossum along with a key to the species of
Ophioglossum is also provided.
Key Wor ds : Botrychium, Mankyua, Eusporangiate, AOO.
ords

INTRODUCTION

The Order Ophioglossales is a natural group of premature vascular plants which


reveal the most simple and most intricately mixture of characters compared to bryophytes,
progymnosperms, gymnosperms and angiosperms (Goswami, 2007). It includes a single
family Ophioglossaceae, which is of eusporangiate ferns discovered by Bauhin (1620) and
is a cosmopolitan in distribution. The family consists of 3 genera viz. Botrychium Sw.,
Helminthostachys Kaulf., Ophioiglossum L. (Clausen 1938) and the recently described genus
Mankyua (Sun et al., 2001). The present work focus on the genus Ophioglossum, was
described by Bauhin (1620) and validated the generic status by Linnaeus (1753) and included
it in his ‘Species Plantarum’. More than 45 species and a few varieties the genus
Ophioglossum are known so far worldwide (Pichi-Sermolli 1958, Yadav & Goswami, 2010).
In India the genus is represented by 12 species (Goswami et al., 2008).
As far as South India concerned, the genus Ophioglossum has been reported 5
species by Manickum and Irudayraj in 1992 from Southern Western Ghats, whereas 6 species
were reported by Bhuskute in 1999 from Maharashtra. For them, there is no authenticated
studied was carried out on the genus Ophioglossum . During the frequent survey of
petridophytes the authors collected eight species from Western Ghats (Fig. 1). A note on
the genus Ophioglossum along with the modified key (Panigrahi and Dixit, 1964) to the
species is also given for easy identification.
TA X O N O M I C T R E ATMENT
AT

Key to the Genera of Ophioglossaceae


1a. Sterile lamina simple or ribbon-like ........Ophioglossum
1b. Sterile lamina branched Spike branched ........2
2a. Spike simple, sporangia dehiscing longitudinal slit ........Helmmintostaches
2b. Spike branched, sporangia dehiscing transverse slit ........Botrychium

* E-mail : sach2010d@yahoo.co.in
18 Indian Fern Journal Volume XXXI (2014)

Key to species of Ophioglossum L.


1a. Plants up to 10 cm tall ........2
1b. Plants 10-30 cm tall ........5
2a. Sterile lamina linear, grass-like ........O. gramineum
2b. Sterile lamina ovate – elliptic or elliptic – lanceolate ........3
3a. Rhizome subglobose, trophophyll ovate – elliptic ........4
3b. Rhizome tuberous, trophophylls elliptic – lanceolate, apex acute
........O. lusitanicum
4a. Trophophills ovate or elliptic, apex obtuse, base cuneate
........O. nudicaule
4b. Trophophills ovate, apex acute - apiculate, base cuneate
........O. parvifolium
4a. Rhizome globose or bulbose ........5
4b. Rhizome not globose ........6
5a. Sterile trophophyll small, lanceolate, obtuse – ovate
........O. costatum
5b. Sterile trophophyll large, elliptical – oblong, glabrous
........O. polyphyllum
6a. Lamina codrate ........O. reticulatum
6b. Lamina lanceolate ........O. petiolatum
Ophio glossum ggrr amineum Willd. Nov. Act. Acad. Erfurt. 2: 18. t. f. 1 (1802); Beddome,
Ophioglossum
Handb. Suppl. Ferns Brit. India 108. 1892; Balakrishnan et al., Bull. Bot. Surv.
India 2: 337 (1960); Panigrahi & Dixit, Proc. Nat. Inst. Sci. India 35: 250 (1969);
Nayar and Kaur, Comp. Beddome, Handb. 106 91974); Dixit, Cens. Indian Pterid.
23. 1984; Nair et al., J. Econ. Tax. Bot. 12: 207 (1988); Manickam and Irudayaraj,
Pterid. Fl. W. Ghats 48. t. 26 (1992.
Ophioglossum clietrichiae Prantl, Ber. Deut. Bot. Ges. 1: 352 (1883);
Ophioglossum dietrichiae Prantl, Ber. Deut. Bot. Ges. 1: 352 (1883).
Ophioglossum gregarium Christ, Nova Guinea Bot. 8: 164 (1909);
Ophioglossum inconspicuum (Rac.) v.A.v.R., Bull. Dep. Agr. Ind. Neerl. 21: 9 (1908);
Ophioglossum indicum Khandelwal in Fras.-Jenk., New Sp. Synd. Ind.Pterid. 185 (1997).
Ophioglossum vulgatum var. gramineum (Willd.) Hook. f., Fl. Nov. Zel., 2: 50. (1854).
Ophioglossum moluccanum Schlechtend. f. incospicum, Rac. Nat. Tijd. Ned. Ind., 59: 237,
t. 2, f. 5 (1900)
Terrestrial herbs, 2 - 4 cm long; rhizome tuberous, 2 - 4 mm wide, with fleshy
roots; tropophylls 1 – 2 cm long, 0.5 – 3 mm dia., linear, grass-like, apex acuminate,
margin entire, soft, green; veins anastomosing; common stalk 1 – 1.5 cm long; fertile
segment 1 – 2 .5 cm long, 0.5 - 2 mm wide; fertile segment arising from the base of the
Sachin Patil & Meena Dongare : The Genus Ophioglossum from Western Ghats of India 19

Figure 1
20 Indian Fern Journal Volume XXXI (2014)

sterile blade; strobilli 0.7 – 1.5 cm long, 0.3 – 0.7 mm diam., 5 – 8 pairs of sporangia,
arranged in two alternate rows, globose; spores 25 – 40 µ dia., trilete, exine reticulate.
DISTRIB UTION : Wor
DISTRIBUTION ld – India, Sri Lanka, East Borneo, East Java, New Guinea,
orld
Vietnam and Africa.
India – Western Himalaya, West Bengal, Rajastan, Madhya Pradesh, Maharashtra,
Andhra Pradesh, Tamil Nadu, Karnataka, Kerela.
Wester n Gha
estern ts: Durgawadi, Bhimashankar, Mahabaleshwar, Patan, Sawantawadi, Amba,
Ghats:
Kolhapur, Panhala, Karad, Kas, Appachiwadi, Ajara, Amboli, Khanapur, Belgaum, Goa,
Molem, Castle Rock, Kasaragod, Thirunelveli.
CONSER
CONSERV VATION ST
STAATUS : Ophioglossum gramineum Willd. Has been collected
from many plateaus situated in Western Ghats of India. A population of about more than
2000 individuals was found. The area of occupancy (AOO) is 1-2 km2/per locality. It is
assessed as least concerned (LC) species following the IUCN Categories and Criteria
(IUCN 2001).
ECOLOGY : Collected from grassy plateaus associate with O. nudicaule , O.
petiolatum and O. costatum.
SPECIMEN EXAMINED : MAHARASHTRA: Kolhapur Dt., Shivaji University,
campus, alt. 700 m., Patil S. S. (SUK); KARN
KARNA ATAKA: Belgaum Dt., Adi, alt. 800 m.,
Mahamuni R. J. (SUK); ANDHRA PRADESH: Karimnagar Dt., Aklaspur, Subbarao
KERALA: Kasaragod Dt., Kasaragod, Ansari (MH67990); Idukki Dt.
(MH20238); KERALA Dt.,
Thekkady, Vivekananthan (MH67990); TAMIL N ADU: Tiruchirappally Dt., Nerthamalai,
NADU:
Ramamurthy (MH25941). Thirunelveli Dt. Dt., Main falls, Corutallum, alt. 160 m.,
Subramanyan (MH3748).
Ophio glossum lusitanicum Linn. sp. Pl., 2: 1063, 1753; Clausen, Mem. Torry Bot. Club,
Ophioglossum
19 (2): 159, 1938; Mahable, Bull. Bot. Surv. India, 4: 71, 1962; Panigrahi & Dixit,
Proc. Nat. Inst. Sci. India 35: 251, 1969.
Plant terrestrial; rhizome subterranean, very short, erect, tuberous, bearing many
long fleshy roots; trophophyll 2 – 3 cm long, 0.5 – 0.7 cm broad, green, long, elliptic -
lanceolate, entire margin, blunt or acute apex, cuneate at base; venation reticulate; fertile
segment 0.5 – 2 cm long without strobilii, unbranched, green at younge, yellow brown at
maturity; strobilli 0.5 – 1 cm long, 8 – 12 spornagia in two rows; sporangia dehiscing via
median slits transverse to the axis; spores 20 – 25 µm dia., trilete.
DISTRIB UTION : Wor
DISTRIBUTION ld – India, Algeria, Morocco; Portugal (Azores); Spain
orld
(Canary Is.); Tanzania, United Republic of; Tunisia; Uganda
India – Andhra Pradesh, Assam, Bihar, Goa, Jammu-Kashmir, Karnataka, Kerala,
Madhya Pradesh, Maharashtra, Orissa, Punjab, Tamil Nadu, Uttar Pradesh, West Bengal.
Wester n Gha
estern Ghatsts : Patan, Mahabaleshwar, Radhanagri, Trivandram.
CONSER
CONSERV VATION ST STAATUS : Ophioglossum lusitanicum Linn. is collected from
Sachin Patil & Meena Dongare : The Genus Ophioglossum from Western Ghats of India 21

Patan, Radhanagri, Mahabaleswar, Panchgani plateaus situated in Western Ghats of India. A


population of about 500 individuals was found. The area of occupancy (AOO) is 1-2 km2/
per locality. It is assessed as critically endangered (CE) species following the IUCN
Categories and Criteria (IUCN 2001).
ECOLOGY : Very rare species, collected from grassy plateaus, associated with O.
nudicaule and O. costatum.
SPECIMEN EXAMINED : INDIA – KERALA KERALA: Trivandram, forest between thura
and Bonasoore, alt., 525 m, J. Joseph (MH90512).
Ophio glossum nnudicaule
Ophioglossum udicaule L. F. Suppl. Pl. Syst. 443 (1781); Beddome, Handb. Ferns Br.
India, 464, t. 228 (1883); Panigrahi & Dixit, Proc. Nat. Inst. Sci. India 35. 252
(1969); Beddome Handb. 106 (1974); R.D. Dixit, Cens. Ind. Pterid. 23 (1984);
Manickam & Irudayaraj, Pterid. Fl. West Ghats: 48-49. t. 27 (1992); Chandra,
Ferns India 9 (2000); Pullaiah et al., Pterid. Andhra Pradesh. 29 (2003). Type -
South Africa: Cape of Good Hope.
Ophioglossum capense Sw., Schard. Journ. 1801(2): 308 (1803).
Ophioglossum capense Schlech. var. nudicaule (L.) Schlech., Fil. Prom. Bonae Sp.: 9
(1825).
Ophioglossum ellipticum Hook. & Grev.Icon. Filic. t. 40 A (1828).
Ophioglossum lineare Schlechter et Brause, Bot. Jerb., 49: 59, fig. 3F (1912).
Ophioglossum luersseni Prantl, Ber. deut. Bot. Ger. 1: 352 (1883).
Plant terrestrial, 4 – 8 cm height; rhizome subglobose, bearing many soft fibrous
roots; tropophylls 0.5 – 2 mm long, touft ovate or elliptic, apex obtuse, base cuneate;
veins reticulate, with free veinlets at margins; fertile segment 2 – 6 cm long, 0.1 – 0.4 mm
broad, green at young, yellow at maturity; strobilli 1 – 2 cm long, 10 – 20 sporangia; spore
30 – 40 µm dia., trilate, with reticulate ornamentation.
DISTRIB
DISTRIBUTION UTION : Wor ld – India, China, Indonesia; Malaysia; Thailand
orld
India: Darjiling, Himachal Pradesh, Jammu-Kashmir, Jharkand, Karnataka, Kerala, Sikkim,
Tamil Nadu.
We s t e rrn
n G h aatt s : Patan, Panchgani, Mahabaleshwar, Karad, Sawantawadi,
Apachiwadi, Castle Rock, Khanapur, Tilari Nagar, Radhanagri.
CONSER
CONSERV VATION ST ATUS : Ophioglossum nudicaule L. is collected from platues
STA
situated in Western Ghats of India. A population of about 500 individuals was found. The
area of occupancy (AOO) is 1-2 km2/per locality. It is assessed as vulnerable (V) species
following the IUCN Categories and Criteria (IUCN 2001).
ECOLOGY : Very common species, grows in patches on fully exposed localities of
open grassland habitats in Western Ghats of India.
SPECIMEN EXAMINED : INDIA – MADYA PRADESH, Hoshangabad Dist. Kesla,
alt. 500m., 22/07/1961, J. Joseph (MH25305); TAMIL NADU, Rmnad Dist., Erani forest,
22 Indian Fern Journal Volume XXXI (2014)

alt. 100m., 19/08/1964, R. D. Dixit (MH39459); Tirunelvel Dist., way to falls, alt. 400m.,
25/07/1951, K. Subrahamanyaum (MH17579); Coimbatore Dist., Aliyar, submergible area,
alt. 350m., 27/07/1962, K. M. Selastive (MH29569).
Ophio glossum par
Ophioglossum vif
parvif olium Grev. & Hook. Bot. Misc. 3: 218, 1833.
vifolium
O. nudicaule var. macrorrhizum (Kunze) Clausen, mem. Torry Bot. Club, 19 (2): 150, 1938;
Panigrahi & Dixit, Proc. Nat. Inst. Sci. India 35: 260 – 262, 1969; Bhuskute, Ind.
Fern Jour., 16: 51 – 54, 1999.
Terrestrial herbs, 4 – 5 cm height; rhizome 3-4 mm dia., erect, soft, tuberous,
branched, bearing few fertile and sterile leaves at the apex; trophophyll 0.4 – 1 cm long,
0.4 – 0.6 mm broad, ovate - lanceolate, apex acute to apiculate, margin entire, base cordate;
veins obscure, four or five veins passing up through the stalk of the blade; fertile segment
2 - 4 cm long, linear oblong; strobilli 0.5 – 1 cm long, 0.3 mm broad, 3 – 6 sporangia,
alternate on either side; spores 25 – 45 µm dia., trilete, exine reticulate.
DISTRIB UTION : Wor
DISTRIBUTION ld – India, China, South America, Sumatra, Malaysia,
orld
Thailand.
India : Madya Pradesh, Gujrat, Maharashtra.
Wester n Gha
estern ts : Panchgani, Kas, Pahhala, Kolhapur, Gadhinglaj, Apachiwadi, Ajara,
Ghats
Amboli, Sawantawadi, Tilari Nagar, Castle Rock.
CONSER
CONSERV VATION ST ATUS : Ophioglossum parvifolium Grev. & Hook.. is collected
STA
from platues situated in Western Ghats of India. A population of about 2000 individuals
was found. The area of occupancy (AOO) is 1-2 km2/per locality. It is assessed as least
concerned (LC) species following the IUCN Categories and Criteria (IUCN 2001).
ECOLOGY : Very common species, grows in patches on fully exposed in Western
Ghats of India.
Ophio glossum costa
Ophioglossum tum R. Br. Prod. Fl. Nov. Holl. 163 (1810). Panigrahi & Dixit, Proc.
costatum
Nat. Inst. Sci. India 35: 249 (1969); Frazer Jenkins Taxo. Rev. Three Hund. Ind.
Pterido. 19 (2007). Type - Australia:Queensland, Arnhem North Bay.
Ophioglossum brevipe Bedd., Ferns. Southern India 23. t. 72 (1863).
Ophioglossum bulbosum Bedd. Ferns. Brit. India Supl. t. 28, (1876).
Ophoglossum fibrosum Schum., Bedd., Handb. 465. t. 289 (1883); and Suppl. 109 (1892).
Ophioglossum pedunculosum Desv., Mag. Nat. Fr. Berlin 5: 306 (1811).
Rhizome 1 – 2 cm dia., tuberous, unbranched, bearing numerous long unbranched,
slender, fleshy roots; tropophylls 1.5 – 2.5 cm long, 1.5 – 2 cm broad, lanceolate, ovate
with obtuse apex - mucronate, base cuneate, margin entire, thick fleshy, green when fresh,
glabrous on both sides, coriaceous, midrib prominent, simple, tufted, all bearing fertile
segment; fertile segment 6 – 10 cm long, 1 – 3 mm thick; strobilli 3 – 5 cm long, 2.5 –
4 mm broad, 20-35 sporangia in two alternate rows; spores 25 – 30 µm dia, trilete, amb
circular, exine feveolate.
Sachin Patil & Meena Dongare : The Genus Ophioglossum from Western Ghats of India 23

DISTRIB UTION : Wor


DISTRIBUTION ld – India, Sri Lanka, Sumatra, Philippines, Thailand, South
orld
Africa, Malawi, Zambia, Zimbabwe, Mozambique, Madagascar, Australia, Peninsular
Malaysia, Cameroon, Ghana, Kenya, Tanzania, Guinea, Angola, Sudan, Congo, Ethiopia,
Brazil.
India – Madhya Pradesh, West Bengal, Maharashtra, S – India, to, NE-India.
Wester n Gha
estern ts : Idukki, Kas, Pahhala, Kolhapur, Amba, Vaibhavwadi, Goa, Molem,
Ghats
Gadhinglaj, Apachiwadi, Ajara, Amboli, Sawantawadi, Tilari Nagar, Castle Rock.
CONSER
CONSERV VATION ST ATUS : Ophioglossum costatum R. Br. is collected from
STA
platues situated in Western Ghats of India. A population of about 500 individuals was found.
The area of occupancy (AOO) is 1-2 km2/per locality. It is assessed as vulnerable (V)
species following the IUCN Categories and Criteria (IUCN 2001).
ECOLOGY : Rare fern, collected from grassy plateau of Western Ghats of India.
The population is varies according to the water content of the soils.
SPECIMEN EXAMINED : SPECIMEN STUDIED : INDIA – TAMIL N ADU:
NADU:
Tounelveli Dt., way to Thulukkamparair, alt. 300m., 06/09/1969, B. V. Shetty (MH62410);
Tirichinappalhi Dt, Narthamalai, alt. 333 m., 12/09/1958, K. Sulenamanyaml (MH17577);
KERALA: Idukki Dt., Pindimedu Pooyamkutty, Bhargavan (MH87404).
Ophioglossum petiolatum Hook., Exot. Fl. 1: t. 56 (1823); Panigrahi & Dixit, Proc. Natl.
Inst.Sci. 35. 260-61 (1969); Beddome Handb. 106 (1974); R.D. Dixit, Cens. Ind
Pterid. 24 (1984); Manickam, Fern Fl. Palani Hills 14. 1986; Manickam &
Irudayaraj Pterid. Fl. W. G. 52. t. 30 (1992);
Ophioglossum cordifolium Roxb. Hort. Bengal. 75 (1814).
Ophioglossum pedunculosum sensu Hotft; Rev. Fl. Malaya 2: 39 (1954).
Ophioglossum pedunculatum Nakai Bot. Mag. Tokyo. 39: 193 (1925).
Ophiglossum reticulatum (non Linn.) sensu Hosokawa. J. Jap. Bot. 18: 200 (1942).
Ophioglossum vulgatum (non Linn.) sensu Hosokawa, occasional paper
Bishop Museum 13: 608. 1937
Rhizome 0.5 – 1.3 mm diam., short erect, dark brown, cylindrical bearing one or
more tropophylls; tropophylls 2 – 4 cm long, 1 – 2 cm broad, erect to spreading, gray-
green, dull, shortly stalked, lanceolate, apex acute, margin slightly thickened, base cuneate
or subcordate; veins anastomosing, 6-10; fertile segment 3 – 12 cm long, 1 – 2.5 mm
thick; strobilli 3 – 4 cm, 1.5 – 3 mm, simple linear, lanceolate, up to 30 pairs of sporangia;
spores 35 – 45 µ dia., finely reticulate.
DISTRIB UTION : Wor
DISTRIBUTION ld – India, Nepal, Japan, Bhutan, Borneo, China, Philippines,
orld
Sumatra, Sri Lanka, England, Fiji, Japonica, Java, Madagascar, Florida, Mexico, Australia,
New Zealand, Africa.
India – Arunachal Pradesh, Sikkim, Nagaland, Kumaon, Kashmir, Uttar Pradesh,
Punjab, Madhya Pradesh Maharashtra.
Wester
esternn Gha ts : Kas, Dajipur, Amboli.
Ghats
24 Indian Fern Journal Volume XXXI (2014)

CONSER
CONSERV VATION ST ATUS : Ophioglossum petiolatum Hook.. is collected from
STA
platues viz. Kas, Dajipur and Amboli situated in Western Ghats of India. A population of
about 100 individuals was found. The area of occupancy (AOO) is 1-2 km2/per locality. It
is assessed as critically endangered (CE) species following the IUCN Categories and Criteria
(IUCN 2001).
ECOLOGY : Grows in moist exposed grassy areas.
SPECIMEN EXAMINED : TAMIL NADU: Nilgiris Dt., Kundha, Deb 31732 (MH);
Nilgiris Dt., Kundha, Deb (MH31732); MADHYA PRADESH, Kerkal, alt. 567m., 17/11/
1958, K. Subramaniayam (MH17583).
Ophioglossum reticulatum L. Sp. Pl. 2: 1063 (1753); Beddome, Ferns. Southern India 23.
t. 70. 1863; Beddome, Handb. Ferns. Bri. India, 465. t. 290 (1883); Panigrahi &
Dixit, Proc. Nat. Inst. Sci. India 35. 257 (1969); Bedd., Handb. 107. 1974; R.D.
Dixit, Cens. Ind. Pterid. 24 (1984); Manickam & Irudayaraj, Pterid. Fl. West Ghats:
51. t. 29 (1992). Type – Mauritius.
Ophioglossum peruvianum Presl, Suppl. Tent. Pterid 52. 1845.
Ophioglossum petiolatum sensu Wieffering, Blumea, 12: 327. 1964.
Ophiglossum cordifolium Roxb., Hort. Bengal, 75. 1814;
Ophioglossum vulgatum L. var. reticulatum (L.) D. C. Eaton., Mem. Amer. Acad. Arts 8:
218 (1861).
Rhizome 1 – 6 mm long, cylindrical, tuberous, bearing few long, thick fleshy
unbranched roots, with 1–2 tropophylls; tropophylls 7 – 12 cm long, subterranean, cordate,
broadly ovate, rarely broadly elliptic-oblong or broadly ovate, apex rounded, base cordate
to broadly cuneate; veins reticulate, several lateral veins passing to the base of the blade,
the ultimate areoles with or without included free veinlets; fertile segment 12 - 17 cm.
long, linear, lanceolate; strobilli 20 - 25 pairs, 0.5 – 1.5 mm thick, globose sunken arranged
in two alternate compact rows; spores 30 – 45 µ dia., spherical, trilete, exine hemispherical.
DISTRIB UTION : World – India, Malay Peninsula, Tropical America, Africa,
DISTRIBUTION
Philippines and Sri Lanka.
India – Andra Pradesh, Madya Pradesh, Kerala, Tamil Nadu, Maharashtra.
Wester n Gha
estern ts : Tirunelveli, Idukki, Nilagiri, Kas, Pahhala, Kolhapur, Gadhinglaj,
Ghats
Apachiwadi, Ajara, Amboli, Sawantawadi, Tilari Nagar, Castle Rock.
CONSER
CONSERV VATION ST STAATUS : Ophioglossum petiolatum Hook.. is collected from
platues viz. Kas, Dajipur and Amboli situated in Western Ghats of India. A population of
about 1000 individuals was found. The area of occupancy (AOO) is 1-2 km2/per locality. It
is assessed as least concerned (LC) species following the IUCN Categories and Criteria
(IUCN 2001).
ECOLOGY : Very common, found in under beneath of tree in the forest or grassy
plateaus.
Sachin Patil & Meena Dongare : The Genus Ophioglossum from Western Ghats of India 25

SPECIMEN STUDIED : INDIA – TAMIL N ADU : Tirunelveli Dt., near Kanni


NADU
forest rest house, alt. 800m., 28/08/1963, A. N. Henry (MH39280); Salem Dt.,Craigmoor
swamps, yercund, 06/06/1953, E. Vairaveha (MH140424); Kondaiampatti, alt. 270 m.,
Subramanyan (MH7780); Nilagiri Dt., Birola, way to Jhorapalli, alt. 850, 26/10/1972, K.
Uivekanathan (MH83036); Coimbatore Dt., Akkamalai, R.F., Joseph 15549 (MH);
KARN
KARNA ATAKA : Chamarajanagar Dt., Way to Bandipur, Naithani, (MH21230). ANDHRA
PRADESH : Kurnool Dt., alt. 270 m., Gundlabrahmeswaram (MH16950).
Ophio glossum pol
Ophioglossum yph
polyph
yphyllum
yllum A. Braun ex Seubert., Fl. Azor. 17 (1844); Pichi Sermolli,
Webbia 9: 632 (1954); Panigrahi and Dixit, Proc. Nat. Inst. Sci. India 35 B(3):
255 (1969); Dixit, Indian Fern J. 6: 146 (1989).
Ophioglossum cuspidatum Milde, Bot. Zeit. 22: 107 (1864);
Ophioglossum aitchisonii (Clarke) d’ Almeida, J. Indian Bot.Soc. 3: 63 f. 12-13 (1922);
Mahabale, Bull. Bot. Surv. India 4: 71 (1962).
Plant up to 15 cm long, terrestrial or semiaquitic; rhizome bulbous or tufted,
unbranched, rhizoids numerous, bearing three – four trophophylls; trophophyll 4 – 12 cm
long, 1 – 2 cm broad, touft, elliptical – oblong, glabrous, narrowed at base and apex, wider
medially, apex acute – mucronate, margin entire; texture thin but firm; veins without free
endings, anastomosing with simple included veinlets; fertile segments 4 – 6 cm long without
strobilii, arise from the base of each sterile leaves under the soil; strobilli 1 – 2.5 cm long,
green at young, yellow at maturity; spore trilate with circular amb. ca. 30 – 40 µm dia.
with fine reticulated exine.
DISTRIB UTION : Wo rrld
DISTRIBUTION ld : India, Ethopia, Kenya, South-West Africa, North
America.
India : Utter Pradesh, North-East Himalaya, Maharashtra.
Wester n Gha
estern ts : Appachiwadi.
Ghats
CONSER
CONSERV VATION ST ATUS : Ophioglossum petiolatum Hook. is collected from
STA
single locality situated in Western Ghats of India. A population of about 1000 individuals
was found. The area of occupancy (AOO) is 1-2 km2/per locality. It is assessed as critically
endangered (CE) species following the IUCN Categories and Criteria (IUCN 2001).
ECOLOGY : it grows in moist soil in grassland habitat of Sindudurg associated with
O. graminium, O. nudicaule, O. costatum and other Gramminae members of angiosperm.
DISCUSSION

The consistency and additional records of pteridophytes, keeps the update of ferns
and ferns-allies wealth of India. In India, like the Eastern Himalayas, the Western Ghats
and the Eastern Ghats having more diversity of pteridophytes. The Western Ghats is one of
the hottest hotspot of biodiversity. The floristic studies on pteridophytes have been carried
out time to time by workers viz. Beddome (1884), Blatter and d’Almedia (1922),
26 Indian Fern Journal Volume XXXI (2014)

Naiknaware (1994), Manickum and Irudayaraj (1992), however, in connection to the generic
level studied was not yet been carried out from Western Ghats of India. Therefore, the
present investigation was undertaken to study the genus Ophiglossum from Western Ghats
of India. It reports eight species of Ophioglossum from Western Ghats of India. Amongst
these three species are least concerned viz. Ophioglossum gramineum, O. parvifolium and
O. reticulatum, three species are critically endangered (CE) viz. Ophioglossum lusitanicum,
O. petiolatum and O. polyphyllum, and two species are vulnerable (V) viz. Ophioglossum
costatum and O. nudicaule.
ACKNOWLEDGMENT

The authors are thankful to The Head, Department of Botany, Shivaji University
Kolhapur for providing the laboratory facilities.
LITERATURE CITED
AT

B AU H I N C 1 6 2 0 Ophioglossum and Lunaria In Prodromus Theatri Botanici, Frankfurt, Germany


pp 354-355
BHUSKUTE S M 1999 Ophioglossums of Bhandara district Maharashtra state India Indian Fern
Journal 1 6 : 51-4
CLAUSEN R T 1938 A monograph of the Ophioglossaceae Memoir s of the Torrey Botanical Club 1 9
: 1–177
GOSWAMI H K, VERMA S C & SHARMA B D (eds) 2008 Biology of Pteridophytes–I Ophioglossum,
Linnaeus Bionature Monograph pp 1–135 Catholic Press Ranchi
PANIGRAHI G & DIXIT R D 1969 Studies in Indian Pteridophytes-IV The family Ophiglossaceae
India Proc Nat Inst Sci India 3 5 B : 230-266
PICHI SERMOLLI R E G 1958 The higher taxa of the Pteridophyta and their classification in
Hedberg O (ed) Systematics of Today Uppsala Univ Årsskrift 1958 ( 6 ) : 70-90
S U N B Y, K I M R D , K I M C H & P A R K C W 2 0 0 1 M a n ky u a ( O p h i o g l o s s a c e a e ) a n e w f e r n g e n u s
from Cheju Island Korea Taxon 5 0 : 1 0 1 9 – 1 0 2 4
YADAV B L & GOSWAMI H K 2010 Bull Natl Mus Nat Sci Ser B 3 6 ( 4 ) : pp 155–159
L I N NA E U S C 1 7 5 3 O p h i o g l o s s u m a n d O s m u n d a I n S p e c i e s P l a n t a r u m A d l a r d & S o n , B a rt h o l o m ew
Press , Dorking , Great Britain for Ray Society (1959) Ed 1, 2 : 1062-1064
Indian Fern J. 3 1 : 27-39 (2014) ISSN 0970-2741

EFFECT OF HEAVY MET


HEAVY AL POLLUTION ON ST
METAL OMA
STOMA TA OF JUVENILE
OMAT
SPOR OPHYTES OF SOME FERNS OF GANGETIC WESTBENGAL
SPOROPHYTES
RIPAN PAUL1*, TUHINSRI SEN1, U. SEN1** AND S. RAHAMAN2
1
Department of Botany, Kalyani University, Kalyani - 741235, India
2
Bangabasi Evening College, Kolkata - 700009, West Bengal, India

(Received Feb. 20, 2014; Revised Accepted June 18, 2014)

ABSTRACT
The environment of both urban and rural areas especially the industrial areas are
polluted by automobile discharges, industrial wastes and especially the rural areas by
agricultural wastes and various insecticide-sprays. Many ferns are excellent indicators of
environmental pollution especially caused by heavy metals like Lead, Mercury and Cadmium.
It has already been established that several fern-gametophytes can minimize the heavy metal
pollution by bioaccumulation and can act as phytoremediator (Paul et al. 2010). Dutta and Sen
(1992) reported the injurious effects caused by the toxic heavy metals on fern sporophytes in
nature. That the fern gametophytes can act as excellent indicators of heavy metal pollution, has
already been reported (Paul et al. 2011). The present study deals with effects of heavy metal
pollution on the stomata of the juvenile sporophytes of Adiantum philippense, Ceratopteris
thalictroides, Christella dentata (=Thelypteris dentata), Cyclosorus interruptus, Diplazium
esculentum, Drynaria quercifolia, Microsorum punctatum, Nephrolepis cordifolia, Pteris vittata
and Tectaria polymorpha. The micromorphological parameters examined here are the various
abnormalities of stomata of juvenile sporophytes developed from gametophytes cultured in the
media polluted with various concentrations of Pb, Hg and Cd. The stomatal abnormalities
observed in this study are the gradual decline of stomatal frequency and also shortening of
length and breadth of individual stoma due to increase of concentrations of heavy metal
pollution. In high concentrations of heavy metals stomata become distorted or even in extreme
concentrations their development is completely suppressed.
K e y Wo rrd
d s : Heavy metal pollution, juvenile fern sporophytes, stomata

INTRODUCTION

Our environment, now-a-days is surcharged with pollution. The pollution in air, soil
and water of both urban and rural areas are increasing dramatically due to toxic heavy
metals especially Lead, Mercury and Cadmium. These metals are emitted due to industrial
activities, automobile discharges, agricultural wastes, various pesticide-sprays, indiscriminate
dumping of industrial wastes. Though laws regarding air and water pollution are present in
our country, yet neither the wise nor the ignorant people are paying any importance to these
laws and thereby bringing their own miseries. Interestingly, it may be recalled that in
Sundarbans, several religious places remain unpolluted since these areas are debarred from
any human activities and the plants there dwell unpolluted (Sen et al. 2010).
Dutta and Sen (1992) reported the injurious effects caused by these pollutants on
adult fern sporophytes in nature. Cadmium is absorbed by Azolla caroliniana (Bennicelli et
al., 2004; Gupta & Devi, 1995) and also by Marsilea minuta (Singh et al., 2008). Panchanan

* E-mail : dr.ripanpaul@yahoo.in ** Prof. U. Sen, Block-B, P-15/96, Lake Row, Kalyani - 741235 - 02
28 Indian Fern Journal Volume XXXI (2014)

and Mondal (1998) have shown that with increase of pollutants in the substrate areas,there
is decrease of percentage of fertile spores production in Ceratopteris thalictroides. Both the
gametophytes and the sporophytes of some ferns can minimize the heavy metal pollution
by bioaccumulation and can act as phytoremediator (Paul et al. 2010). That fern
gametophytes can even act as excellent indicators of heavy metal pollution, has already been
reported (Paul et al. 2011).
In the present study, the effect of heavy metal pollution on the stomatal features of
juvenile sporophytes developing on gametophytes grown on polluted media have been
recorded.
MATERIAL AND METHODS
ATERIAL

Mature spores of Adiantum philippense L. (Adiantaceae), Ceratopteris thalictroides


(Linn.) Brongn. (Pteridaceae), Christella (=Thelypteris) dentata (Forssk.) Brownsey and
Jermy (Thelypteridaceae), Cyclosorus interruptus (Willd.) H. Ito (Thelypteridaceae),
Diplazium esculentum (Retz.) SW. (Athyriaceae), Drynaria quercifolia (Linn.) J. Sm.
(Polypodiaceae), Microsorum punctatum (Linn.) Copel. (Polypodiaceae), Nephrolepis
cordifolia (Linn.) Presl (Nephrolepidaceae), Pteris vittata (Linn.) (Pteridaceae), Tectaria
polymorpha (Wall. ex Hook.) copel. (Dryopteridaceae) were collected from several districts
of Gangetic West Bengal.
Spores were sterilized by 5% Calcium hypochlorite for 3-5 minutes and were sown
on modified Moore’s Medium supplemented with Nitsch’s trace elements (Kato 1969).
Different concentrations of pollutants (Pbcl2, Hgcl 2 & Cdcl2) were added to the control
medium. The tolerance level of taxa under investigation varies differently to different
pollutants and also in different concentrations of pollution levels. The germination
percentages and stages of development and abnormalities of the gametophytes in various
pollution levels were stated previously (see Ripan Paul et al. , 2011). However the
germination percentages were reported only up to 3 ppm concentration of different levels,
as beyond this concentration, excepting Christella (=Thelypteris) dentata , healthy
sporophytes do not develop. The germination percentages gradually decrease in higher
pollution levels. The percentages of sporophyte formation in control and in polluted
environment are recorded in Table 1. In a few taxa, the juvenile sporophytes borne on
gametophytes did not reach beyond first leaf stage. For this reason the data of stomata
were taken only of the first juvenile leaf. In all the cases, the stomata were compared
with those of control. The frequency of stomata, their length, breadth and other
characters were noted. The stomatal terminology is used following Cotthem, W. Van
(1970) and Sen & De (1992). The tables show the stomatal record only up to the level
of pollutions in which the sporophytes can tolerate and develop stomata. Interestingly,
beyond these tolerance levels, sporophytes either failed to develop fronds or formed
distorted stomata.
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 29

TABLE 1 : Toler ance le


olerance v els to pollution of dif
lev difff e rrent
ent ffee rrn
n taxa and per centa
percenta
centagg e of spor oph
sporoph yte ffo
ophyte o rrm
m aation
tion
in control and polluted media
media..

Sl. Name of taxa Name Conc. of heavy metal compounds (in ppm)
ppm)→→
N o . & percentage of
of sporophyte Heavy 0.001 0 . 0 1 0.1 0.5 1 1.5 2 3 4 5 6 7 8 9
formation in metal com-
Percentage (%) of sporophyte formation
formation→→
control pounds
1. Ceratopteris Pbcl2 1
thalictroides Hgcl2 2 2 1
20% Cdcl2 1
2. Christella Pbcl2 12 11 11 8 4 3 2 1
dentata Hgcl2 8 6 4 3 2 1 1
25% Cdcl2 20 18 18 16 15 14 13 11 9 7 4 3 2
3. Cyclosorus Pbcl2 6 5 2 1
interruptus Hgcl2 7 6 3 2 1
20% Cdcl2 5 4 2 1
4. Diplazium Pbcl2 2
esculentum Hgcl2 3
10% Cdcl2 4 1
5. Drynaria Pbcl2 4 1
quercifolia Hgcl2 2 1
15% Cdcl2 5 4 2 1
6. Microsorum Pbcl2 6 2
punctatum Hgcl2 1
8% Cdcl2 1
7. Nephrolepis Pbcl2 2 2 1
cordifolia Hgcl2 1
4% Cdcl2 2
8. Pteris Pbcl2 12 12 11 11 8 4 2
vittata Hgcl2 13 13 12 10 7 5 2 1
25% Cdcl2 15 15 13 12 11 11 8 2
9. Adiantum
philippense No sporophyte develops in these taxa in polluted environment.
15%
10. Tectaria
polymorpha
4%
30 Indian Fern Journal Volume XXXI (2014)

OBSER
RVVAT I O N

In the polluted environment the ferns similar to other types of plants, suffer very
much in their growth. In the polluted condition, ferns often exhibit distorted growth, and
in highly polluted state they do not grow at all. There are chromosomal abnormalities and
distortion and decline of chloroplast pigments. Even, there is adverse effect of the pollutants
on their micro morphological characteristics.
It has been noted that some ferns are extremely sensitive to Lead, Mercury and
Cadmium pollution, while some others can endure these heavy metal pollution to a great
extent. There are, however, some taxa which have medium-endurance capability. Stomatal
features of juvenile sporophytes of ten different ferns grown in culture media polluted with
Pb, Hg & Cd are discussed separately in this study.
z Adiantum philippense
This fern is extremely sensitive to environmental pollution especially to Hg-
pollution. Sporophytes in composite culture develop always apogamously even in control
medium. The first juvenile leaf of the sporophyte develop stomata quite frequently having
stomatal index of 32.43.The stomata are commonly polocytic, but copolocytic and
seppolocytic stomata also occur (Fig.1). Stomata are variable in size, measuring from 22.2
µm to 44.4 µm in length and 22.6 µm to 37 µm in width. Only in 0.001 ppm of Pbcl2 and
Cdcl2, very occasionally extremely ill-developed sporophyte appears with a single crooked
leaf and it is not properly unfolded and stomatal development cannot be recorded. In Hgcl2
polluted medium, sporophytes never appear.

Ta b les 2-9. Inde


Indexx & measur ement of stoma
measurement ta of fr
stomata onds of juv
fronds enile spor
juvenile oph
sporoph ytes ggrr own on polluted and contr
ophytes ol
control
medium
TABLE 2 : Name of taxon : Cer
Ceraa topter is thalictr
topteris oides
thalictroides

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 22.2-44.4 22.22-29.6 31.57


Hgcl 2 0.001 44.4-66.6 29.6-43.2 32.43
0.01 43-59.8 28-42 30.76
0.1 42-51.2 27.6-41.2 27.77
Cdcl 2 0.001 37-49 22.2-29.6 28.20
Contr ol (W
Control ithout hea
(Without vy metals)
heavy 44.4-66.6 29.6-43.2 36.17
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 31

Fig. 1-20 : Stomata of fronds of juvenile sporophytes growing on control medium and on polluted media.
1. Adiantum philippense ; 2 . Ceratopteris thalictroides ; 3 , 4 & 5. Christella dentata ; 6-9. Cyclosorus
interruptus ; 10. Diplazium esculentum ; 11-13. Drynaria quercifolia ; 14 & 15. Microsorum punctatum ; 16-17.
Nephrolepis cordifolia ; 18 & 19. Pteris vittata ; 20. Tectaria polymorpha
1, 3, 6, 12, 19, 20. Control media; 2, 7, 11, 14, 17. 0.001 ppm conc. of Pbcl 2 polluted media;
4. 1 ppm conc. Pbcl2 polluted medium; 5, 10, 18. 0.001ppm conc. of Hgcl2 & 9. 0.01 ppm conc. of Hgcl2 polluted
media; 8, 15, 16. 0.001 ppm conc. of Cdcl 2 polluted medium; 13. 0.01 ppm conc. of Cdcl 2 polluted medium
x206
x206; 2. x103
1, 3-10, 13, 16, 17, 20.x206 x103; 11, 12, 14, 15, 18, 19. x172
x172.
32 Indian Fern Journal Volume XXXI (2014)

TABLE 3 : Name of taxon : Chr istella denta


Christella dentatt a

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 29.6-37 22.2-28.6 30.61


0.01 29.6-37 22.2-28.6 29.70
0.1 29.6-37 22.2-28.6 27.27
0.5 25.8-30.5 20.2-26.6 25
1 25.8-30.5 20.2-26.6 18.18
1.5 25.8-30.5 20.2-26.6 16.66
2 22.2-29.6 18.5-25.6 13.88
3 14.8-18.5 12.8-16.5 12.90
Hgcl2 0.001 29.6-37 22.2-28.6 30.43
0.01 29.6-37 22.2-28.6 23.80
0.1 25.8-30.5 20.2-26.6 21.05
0.5 25.8-30.5 20.2-26.6 20
1 21.2-28.5 18.5-25.6 16.66
1.5 20.2-28.5 14.8-22.2 15.78
2 14.8-18.5 12-16.6 12.50
Cdcl2 0.001 29.6-37 22.2-28.6 32
0.01 29.6-37 22.2-28.6 30
0.1 29.6-37 22.2-28.6 28
0.5 29.6-37 22.2-28.6 26
1 20.2-28.5 18.5-25.6 23.91
1.5 20.2-28.5 14.8-22.2 20.45
2 20.2-28.5 14.8-22.2 19.04
3 20.2-28.5 14.8-22.2 18.60
4 20.2-28.5 14.8-22.2 16.21
5 20.2-28.5 14.8-22.2 15
6 20.2-28.5 14.8-22.2 14.28
7 20.2-28.5 14.5-22.2 12.90
8 14.6-27 12.2-16.6 10.34
Contr ol(W
Control(W ithout hea
ol(Without vy metals)
heavy 29.6-37 22.2-28.6 32.14
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 33

z C e rraa topter is thalictr


topteris oides ( T
thalictroides Taa b le 2)
It grows well in unpolluted environment. But it is very susceptible to Cd and
Pb-salt pollution. On the contrary it can tolerate the Hg–salt pollution at least to some
extent. In unpolluted control medium, the juvenile sporophytes develop luxuriantly and the
stomata are very variable in size varying from 44.4 µm to 66.6 µm in length and 29.43 µm
to 43.2 µm in breadth and their frequency is quite high being 36.17. In 0.001ppm conc. of
Pbcl2 and Cdcl2 pollution the stomatal frequency decreases to a considerable degree. The
length and breadth of stomata also decrease significantly. On Hg-polluted substrate juvenile
sporophytes appear soon after fertilization and they grow well even on 0.1 ppm Hgcl2
polluted medium. The stomatal index and stomatal size however, decline gradually. In 0.5
ppm concentration of Hgcl2 pollution, juvenile sporophytes rarely appear and perish within
two or three days and as such nothing about stomata could be recorded (Table 2).
In all kinds of polluted substrate, the juvenile sporophytes commonly develop
polocytic, copolocytic and pseudopolocytic types of stomata (Fig.2). Few desmocytic
stomata also develop. It is interesting to note that occasionally, with Pbcl2 pollution, a few
stomata may become abortive.
z Chr istella (=T
Christella hel
(=Thel ypter
helypter is) denta
ypteris) ta ( T
dentata Taa b le 3)
This fern is a natural weed and grows everywhere including in polluted
environment. On unpolluted medium, numerous juvenile sporophytes appear. Stomata are
oval to almost circular and commonly polocytic or copolocytic ranging from 29.6 µm to
37 µm in length and 22.2 µm to 28.6 µm in width (Fig.3). Healthy juvenile sporophytes
develop even at 3ppm conc. of Pbcl2 and 2 ppm conc. of Hgcl2 on polluted substrate and
develop generally polocytic and copolocytic stomata (Figs. 4 & 5). Cdcl2 pollutant has little
harmful effect on juvenile sporophytes and even in 8ppm conc. of Cdcl2 pollution in the
substrate, the juvenile sporophytes grow more or less healthy and develop polocytic and
copolocytic types of stomata. However, the stomatal frequency declines gradually (Table 3).
z C y cclosor
losor us inter
losorus interrr uptus ( T
Taa b l e 4 )
This fern is moderately tolerant to polluted environment. When grown in unpolluted
substrate, stomata are oval and variable in size, ranging from 29.6 µm to 44.4 µm in length
and 22.2 µm to 29.6 µm in width (Table 4). The stomatal frequency is quite high being
38.88. Stomata are commonly polocytic, copolocytic and seppolocytic (Fig.6). All these
stomatal types occur also in juvenile sporophytes of this taxon growing on the heavy metal
polluted substrate (Fig.7-9). Young sporophytes growing on medium polluted with 0.001
ppm of Cdcl2 very occasionally develop desmocytic stomata along with other types (Fig.8).
Juvenile sporophyte growing on 0.01 conc. of Hgcl 2 medium some time develops
seppolocytic stomata (Fig.9) in addition to other types. Juvenile sporophytes can endure
only up to the 0.5ppm conc. level of Pbcl2 and Cdcl2 pollution. Interestingly they can
tolerate even up to 1ppm conc. level of Hgcl2 pollution.Stomata are of the same types, but
gradually decline in their size and in frequency.
34 Indian Fern Journal Volume XXXI (2014)

TABLE 4 : Name of taxon : Cyc losor


Cyclosor us inter
losorus interrr uptus

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 22.2-29.6 14.8-25.9 31.06


0.01 22.2-29.6 14.8-22.2 29.78
0.1 22.2-29.6 14.8-22.2 28.57
0.5 22.2-29.6 14.8-18.5 27.16
Hgcl 2 0.001 29.6-37.3 22.2-29.6 33.33
0.01 22.2-33.3 18.5-25.9 30.00
0.1 22.2-29.6 18.5-25.9 29.41
0.5 22.2-29.6 18.5-22.2 27.69
1 18.5-29.6 14.5-22.2 25
Cdcl 2 0.001 22.2-33.3 18.5-29.6 31.32
0.01 22.2-29.6 18.5-25.9 28.91
0.1 22.2-29.6 18.5-25.9 28.72
0.5 22.2-29.6 14.8-22.2 27.38
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 29.6-44.4 22.2-29.6 38.88

TABLE 5 : Name of taxon : Diplazium esculentum

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 29.6-37 22.2-29.6 10.35


Hgcl 2 0.001 29.6-37 22.2-29.6 10.34
Cdcl 2 0.001 29.6-37 22.2-29.6 13.53
0.01 29-36 22.2-29.6 11.40
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 29.6-37 22.2-29.6 20.70

z Diplazium esculentum ( T
Taa b le 5)
This fern cannot tolerate polluted environment. In unpolluted medium, the juvenile
sporophytes grow but in a low frequency. The stomatal index is also very low, only being
20.70. Length and breadth of stomata vary a little ranging only from 29.6 µm to 37 µm in
length and 22.2 µm to 29.6 µm in breadth. When the media are polluted with 0.001ppm of
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 35

TABLE 6 : Name of taxon : D rrynar


ynar ia quer
ynaria cif
quercif olia
cifolia

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 37-51.8 22.2-37 25.00


0.01 37-51.8 22.2-29.6 24.00
Hgcl2 0.001 37-51.8 22.2-29.6 24.07
0.01 29.6-44.4 22.2-27 22.22
Cdcl2 0.001 29.6-45 22.2-37 28.24
0.01 29.6-44.4 22.2-37 23.37
0.1 29.2-44 22.2-37 23.07
0.5 22.2-37 22.2-29.6 22.55
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 37-51.8 22.2-37 30.76

Pbcl2, Hgcl2 & Cdcl2, the juvenile sporophytes develop but these are very late in appearance
and very low in frequency. The stomata are only of polocytic type and remain unchanged in
size in metal polluted environment (Fig.10). However, stomatal index decreases greatly even
in slightly polluted environment (Table 5). The sporophytes do not survive after the
appearance of the first leaf. The tolerance of this fern to Cdcl2 pollutant is also very little
and the juvenile sporophytes develop on 0.01ppm Cdcl2 polluted substrate, but die very shortly.
z D rrynar
ynar ia quer
ynaria quercc i ffolia
olia ( T
Taa b le 6)
A common epiphytic fern in West Bengal, which can endure only moderately
polluted environment. In Laboratory condition, on unpolluted medium the juvenile
sporophytes appear on 125 day old gametophytes. The stomata are polocytic, copolocytic
and seppolocytic (Fig.12). Occasionally, however, desmocytic stomata also occur. The
stomata measure 37 µm to 51.8 µm in length and 22.2 µm to 37 µm in breadth. Upto 0.5
ppm of Cdcl2 and 0.01ppm Pbcl2 and Hgcl2 pollution level, the sporophytes develop even
up to 2 leaf stage in culture media and show various types of stomata (Fig.11). Stomata
are as in control medium, polocytic, copolocytic & seppolocytic. Interestingly, 0.01ppm
Cdcl2 pollution, the outer wall of the guard cells sometimes become thickened (Fig.13).The
heavy metal pollution does not show much effect on stomatal size. Only the stomatal index
decreases slightly (Table 6).
z Micr osor
Microsor um puncta
osorum tum ( T
punctatum Taa b le 7)
This epiphytic fern rather shows very little tolerance to polluted environment in
nature. In culture, and also in control medium, more than 60% of prothalli bear sporophytes
and survive beyond 4th leaf stage. The stomata are frequent and their index is 32. They
vary in size from 51.8 µm to 66.6 µm in length and 29.6 µm to 44.4 µm in breadth. These
36 Indian Fern Journal Volume XXXI (2014)

TABLE 7 : Name of taxon : Micr osor


Microsor um puncta
osorum tum
punctatum

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 44.4-66.6 22.2-37 30.43


0.01 22.2-37 14.8-22.4 19.60
Hgcl 2 0.001 22.2-37 22.2-29.6 22.5
Cdcl 2 0.001 22.2-37 14.8-22.2 24.39
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 51.8-66.6 29.6-44.4 32
are commonly polocytic.
In 0.001 ppm Pbcl2 pollution level, the stomatal size and stomatal index of the
leaves of the juvenile sporophytes decrease a little (Fig.14), but at 0.01 ppm pollution level
of Pbcl2, there is drastic decrease in size in length (22.2 µm to 37 µm) and breadth (14.8
µm to 22.4 µm). The stomatal index also declines greatly being only 19.60 (Table 7). In
Hgcl2 and Cdcl2 pollution, these plants suffer a lot and even at 0.001ppm pollution level of
both Hgcl2 and Cdcl2, there are drastic decline in stomatal size and index (Fig. 15). The
sporophytes are only very few in appearance and they are ill developed and soon die. In all
the cases, the stomata are only polocytic and seppolocytic (Figs. 14 and 15).
z N eephr
phr ole
phrole pis cor
olepis dif
cordif olia ( T
difolia Taa b le 8)
In unpolluted culture, only 16% juvenile sporophytes develop from the zygotes and
rests of the zygotes become necrotic. The sporophytes become healthy and develop several
leaves. The stomata are oval, commonly polocytic and cyclocytic. They measure 37 µm to
51.8 µm in length and 22.2 µm to 37 µm in breadth having high stomatal index 43.18
(Table 8). However, this fern is very much susceptible to the heavy metal pollution,

TABLE 8 : Name of taxon : N eephr


phr ole
phrole pis cor
olepis dif
cordif olia
difolia

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 37-51.8 22.2-29.6 27.69


0.01 29.6-44.4 22-29 25.49
0.1 29.6-44.4 21-27 13.33
Hgcl 2 0.001 29.6-44.4 22.2-29.6 25
Cdcl 2 0.001 29.6-44.4 22.2-29.6 16.94
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 37-51.8 22.2-37 43.18
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 37

particularly in Cd-salt and Hg-salt. In 0.001ppm Cdcl2 and Hgcl2 pollutant in the substrate,
the prothalli can develop sporophytes only rarely (2% to 3%). Stomata are polocytic and
cyclocytic (Fig.16). However, it can tolerate Pb-pollution even up to 0.1 ppm concentration
in the substrate. The stomata in 0.001 conc. of Pbcl2 pollutant are similar to those in control

TABLE 9 : Name of taxon : Pter is vitta


Pteris vittatt a

Name of Concentrations of Range of size of stomatal guard cell Stomatal


heavy metal heavy metal Length Breadth Index
compounds compounds in ppm. (µm) (µm)

Pbcl2 0.001 44.4-58.1 29.6-40.3 26.71


0.01 44.4-51.8 29.6-37 24.31
0.1 44.4-51.8 29.6-37 22.66
0.5 37-48.1 29.6-37 19.78
1 29.6-48.1 22.2-29.6 17.50
1.5 29.6-44.4 22.2-29.6 13.21
2 20.3-42.92 22.2-29.6 12.18
Hgcl2 0.001 45.3-51.8 29.6-33.3 28
0.01 44.4-51.8 29.6-33.3 26
0.1 44.4-51.8 29.6-33.3 24
0.5 40.7-51.8 29.6-33.3 22.15
1 40.4-51.8 22.2-29.6 20.80
1.5 29.6-37 22.2-29.6 17.76
2 29.6-37 22.2-25.9 14.78
3 29.6-37 22.2-25.9 11.50
Cdcl2 0.001 44.4-51.8 25.9-29.6 33
0.01 44.4-51.8 25.9-29.6 32
0.1 40.7-48.1 25.9-29.6 30
0.5 40.6-44.4 22.2-29.6 28
1 40-44.4 22.2-29.6 26.81
1.5 33.3-44.4 22.2-29.6 24.45
2 30.9-44.4 22.2-29.6 22.07
3 29.6-44.4 22.2-29.6 20.66
Contr ol (W
Control ithout hea
(Without vy metal)
heavy 67.6-84.5 50.7-67.6 34.21
38 Indian Fern Journal Volume XXXI (2014)

(Fig.17). In all these cases the stomatal index and size decrease gradually (Table 8).
z Pter is vitta
Pteris ta ( T
vittata Taa b le 9)
This fern grows healthy in unpolluted environment and also can tolerate the polluted
environment to a great extent. In unpolluted culture, the stomatal index is quite high being
34.21. The stomata are very large in size varying from 67.6 µm to 84.5 µm in length and
50.7 µm to 67.6 µm in width. These are more or less oval in outline and commonly
polocytic and copolocytic and a few are also seppolocytic. When the culture medium is
polluted with the salts of the heavy metals like Pb, Hg & Cd, the prothalli can tolerate
even in 2ppm to 3ppm concentration of these pollutants. However, at still higher pollution
level, especially with Pb-salt, the sporophytes become extremely ill-developed and develop
crooked leaves. The stomata are commonly of polocytic types (Figs.18 &19). The stomatal
size and in index reduce greatly when grown in polluted environment (Table 9).
z Tectar ia pol
ectaria ymor
polymor
ymorpha
pha
This fern grows only in unpolluted environment, even in slightest polluted condition
it cannot survive. In unpolluted culture medium the percentage of juvenile sporophytes is
extremely poor. The stomata are commonly polocytic and copolocytic. Interestingly
staurocytic stomata may develop occasionally (Fig.20). The stomatal index is quite high
being 37.5 and the stomata vary in size from 37 µm to 51.8 µm in length and 29.6 µm to
44.4 µm in width. In polluted media, sporophytes may develop at 0.001ppm conc. of Cdcl2
and Pbcl2 but they die only after 2 or 3 days, prior to the expansion of leaves. In Hgcl2
pollution, even the zygotes are not formed.
DISCUSSION

It appears from this study that out of the gametophytes of ten fern taxa studied in
culture condition, the gametophytes of some ferns can tolerate heavy metal pollution and
develop juvenile sporophytes. A few others survive only moderately, and the remaining ones
cannot endure metal –polluted environment. It is particularly true for Adiantum philippense
and Tectaria polymorpha which are totally intolerant to polluted environment. Basing on
this new knowledge, it can be presumed that wherever these two fern taxa grow luxuriantly,
the area must be free from environmental pollution. The occurrence of these two taxa in a
locality, therefore acts as a litmus test of pollution free environment. In nature these two
taxa are available only in restricted unpolluted areas. Pollution tolerance capability of
Ceratopteris thalictroides, Diplazium esculentum, Microsorum punctatum and Nephrolepis
cordifolia is comparatively little, yet these taxa can be placed in the Adiantum group. And
places of their occurrence can be considered as almost environmental pollution free region.
However, it must be remembered that Pteris vittata and Christella (=Thelypteris)
dentata can grow in much polluted environment since these two taxa are capable of
withstanding metal pollution to a great extent.They even accumulate the pollutants and grow
healthy.
Ripan Paul et al. : Effect of Heavy Metal Pollution on Stomata of Juvenile Sporophytes 39

The stomatal characters, such as their size, index and thickening of the walls of the
guard cells can be used to ascertain the condition of environmental pollution.
ACKNOWLEDGEMENTS

The authors are thankful to Prof.S.C.Verma (Chandigarh) for his constructive


suggestions.
REFERENCES

BENNICELLI R, STEPNIEWSKA Z, BANACH A, SZAJNOCHA K & OSTROWSKI J 2004 The ability


o f A z o l l a c a r o l i n i a n a t o r e m o v e h e av y m e t a l s [ H g ( I I ) C r ( I I I ) , C r ( V I ) ] f r o m m u n i c i p a l
waste water Chemosphere 5 5 : 141-146
COTTHEM W VAN 1970 Comparative morphological study of the stomata in the Filicopsida Bull Jard
Bot Natn Belg 4 0 : 81-239
D U T TA K P & S E N T 1 9 9 2 G r ow t h a n d d ev e l o p m e n t o f s o m e f e r n s u n d e r d i ff e r e n t e nv i r o n m e n t a l
conditions Indian Fern Journal 9 : 107-112
G U P TA M & D E V I S 1 9 9 5 U p t a ke a n d t o x i c i t y o f C a d m i u m i n a q u a t i c f e r n s J E nv i r o n B i o l 1 6 :
1 3 1 -1 3 6
K ATO Y 1 9 6 9 P h y s i o l o g i c a l a n d m o r p h o g e n t i c s t u d i e s o f f e r n g a m e t o p h y t e s a n d s p o r o p h y t e s i n
a s p e c t c u l t u r e V I I . E x p e r i m e n t a l m o d i f i c a t i o n s o f d i m e n s i o n a l g r ow t h i n g a m e t o p h y t e s o f
Pteris vittata . Phytomorphology 1 9 : 114-121
PA N C H A NA N B & M O N DA L M S 1 9 9 8 N o t e s o n d i s t r i bu t i o n a n d B i o l o g y o f C e r a t o p t e r i s
thalictroides (L) Bongn (Parkeriaceae) with special reference to spore production and fertility
under pollution stress Bull Bot Surv (India) 4 0 ( 1 - 4 ) : 58-61
PAUL R, SEN T & SEN U 2010 Bioaccumula tion of heavy metals in fern gametophytes and their role
in phytoremediation Indian Fern J 2 7 : 101-107
PAU L R , S E N T, S E N U & R A H A M A N S 2 0 1 1 To x i c e ff e c t o f h e a v y m e t a l p o l l u t i o n o n s p o r e
germination and gametophyte development in some ferns Indian Fern J 2 8 : 25-40
S I N G H J , D E V I S , C H AW L A G , G U P TA M & V I S WA NAT H A N P N 2 0 0 8 U l t r a s t r u c t u r a l a n d
biochemical effects of cadmium on the aquatic fern Marsilea minuta Linn New Phytologist
1 4 5 : 1-11
SEN U & DE B 1992 Structure and ontogeny of stomata in ferns Blumea 3 7 : 239-261
S E N T, C H A K R A B O RT I K , R A H A M A N S , BA N DYO PA D H YAY M & S E N U 2 0 1 0 B i o d iv e r s i t y o f
p t e r i d o p hy t e s o f I n d i a n p a r t o f S u n d a r b a n a n d t h e i r i n s i t u c o n s e r va t i o n P ro c Na t Ac a d Sc i
India Sect B, 8 0 p t ( I I I ) : 201-220
Indian Fern J. 3 1 : 40-45 (2014) ISSN 0970-2741

ADIANTUM HISPIDULUM SW AR
SWAR TZ – A NEW REPOR
ARTZ T
REPORT
FR OM SEN
FROM SENAAPATI DISTRICT
PA DISTRICT,, MANIPUR
TH. SUNITA DEVI1 AND AJIT KUMAR DAS2*
1
Department of Botany, Manipur College, Imphal-795008, India
2
Ethnobotany & Medicinal Plants Conservation Laboratory, Department of Ecology &
Environmental Science, Assam University, Silchar-788011, India

(Received December 4, 2013; Revised Accepted)

ABSTRACT
Adiantum hispidulum (Swartz), a species of maidenhair ferns, is reported for the first
time from Senapati District, Manipur, North-East India. A taxonomic description of this newly
reported species along with photographs and drawings has been provided.
d s : Adiantum hispidulum , Maidenhair fern, Senapati district
K e y W o rrd

INTRODUCTION

The maidenhair ferns, belong to the genus Adiantum L., family Pteridaceae (sensu
Copeland 1947) in the order Polypodiales (Smith et al. 2006). The botanical name Adiantum
refers to the way in which water runs off the surfaces of the leaves without wetting them
(Singh & Panigrahi 2005). There are about 200 species of worldwide distribution but in
India 24 species are reported by Nayar & Kaur (1974), whereas Fraser-Jenkins (2008)
recognized only 14 species and 3 subspecies. Adiantum hispidulum (Swartz) is an attractive
fern species with reddish young fronds commonly known as rosy maidenhair fern or rough
maidenhair fern. The species was first described by Olof Swartz from specimens collected
from Australia. Sinha (1986) reported six species of Adiantum L. from Manipur (Adiantum
aethiopicum, A. capillus-veneris, A. philippense, A. incisum, A. venusetum). Singh (1990)
reported only three species of Adiantum (A. caudatum, A.lunulatum, A. phillipense) species
from Tamenglong district, Manipur. [A. lunulatum and A. philippense are in fact synonyms,
and A. philippense is the correct name which has three subspecies, see Verma & Fraser-
Jenkins 2008]. During recent years Snatombi (2008) reported four species (Adiantum
capilus-veneris, A. caudatum, A. flabellalulatum, A. phillippense) from Thoubal district,
Manipur.
MATERIALS AND METHODS
ATERIALS

The present work is based on the field trips and collection of vascular plants in
various localities of Senapati district of Manipur from September 2012 to August 2013.
Routine herbarium practices were followed as suggested by Jain & Rao (1976). Critical
morphological study of rhizome, scales, stipe, frond, mode of branching, rachis shape, size,
pinnule structures, soral features, number of sporangia etc. and consultation of available

* E-mail : ajitkumardas2009@rediffmail.com
Th. Sunita Devi et al. : Adiantum hispidulum Swartz – A New Report from Senapati District, Manipur 41

literatures and pteridophytic Flora (Beddome 1883, 1892, Borthakur et al. 2000) and reveal
the identity of the fern Adiantum hispidulum Sw. and it forms a new addition to
pteridophytic flora of Manipur. Measurements are given in metric scale.
STUDY AREA
STUDY

Senapati district is one of the nine districts of Manipur, North-East India (93029' -
94015' E longitude and (24037'–25037' N latitude) and about 45 km. from Imphal city
along NH39. It covers an area of 3271 sq.km. with six reserve forests of different areas
elevation ranges from 950 meters to 2560 meters above MSL. The reserve forests are
Kangchup reserve forest (9.60 sq. km.), Kangchup Chiru reserve forest (2.59 sq.km.),
Kanglatongbi Kangpokpi (85.47 sq.km.), Khamenlo-Gwaltabi (85.47 sq.km.), and Khuman-
ching (9.60 sq.km.). These forests belong to sub-tropical mixed type. The reported species
has been collected from the foot-hill of Kanglatongbi Kangpokpi reserve forest.

Adiantum hispidulum Swartz, Hook. Syn. Fil. p.126. Bedd.F, S.I.t.3,


Specimens examined
examined: Kanglatongbi-Kangpokpi reserve forest of Senapati district
1018m, Jan.2013. [Mention Collection/Accsssion No. Thangsam & Das, 0596, locality,
Herbarium where deposited Dept. of Ecology]
Ecology : Uncommon, rare, endemic, terrestrial on forest slope, 960-1500 m.
Distr ib
Distrib ution : India, Madras, FM. Jarrett 534(K); Thailand: Loi, P. Nosiriwong 14
ibution
(BCU); AUSTRALIA : New South Wales, J.R. Boosmau s.n.(P); Papua New Guinea:
Sattelberg, R.J. Johns 086(k). [Sri Lanka]
DESCRIPTION

Herb, ca 39 cm tall ; rhizome-short, sub-erect, ca 0.6cm, thick; fronds erect, tufted;


stipe ridged, clothed towards the base with brown scales ( ca 1mm.x1.3mm., entire,
tapering),covered short stiff multicellular hairs,reddish brown in colour, dichotomous with
the main branches flabellate; lamina ovate to deltate in shape (14-22x13) cm, pinnae
alternate, basal pinnae slightly stalked, upper ones sessile middle pinnae largest, linear
oblong, ca. 14 cm, long, hispid hairs, lateral pinnules up to 14 pairs alternate with short
stalk, middle pinnules mostly sub rhomboidal ca (1.1x 0.6) cm, gradually smaller towards
both ends, brownish green, inner and lower margins straight and entire, upper margins with
narrow sinuses and finely toothed; veins free, forked, visible on both sides, stiff, patent,
lateral pinnae shorter than the middle pinnae; sori at vein tips near margin, 3-8 for each
pinnules, indusial flaps oblong bearing brown hairs, spores yellowish brown, fairy granular,
spores -30x36 µm, exine smooth, tetrahedral, trilete (Fig.1 : A-D)

Adiantum hispidulum Sw
Sw.. can be identif ied bby
identified y using the kkee ys
1a. Rachis dichotomosise into branches, fronds flabellate, fan-shaped lamina
42 Indian Fern Journal Volume XXXI (2014)

PLATE I
PLATE

F i g . 1 . Adiantum hispidulum Swartz - Habit.

F i g . 2 . A. hispidulum Sw. (Young) F i g . 3 . A. hispidulum Sw. (Young)


Th. Sunita Devi et al. : Adiantum hispidulum Swartz – A New Report from Senapati District, Manipur 43

PLATE II
PLATE

F i g . 1 : A . Adiantum hispidulum Sw. A. W hole Plant, B . Pinnules showing Sor i, C . Pinnules


showing venation, D . Rhizone scale
44 Indian Fern Journal Volume XXXI (2014)

2a. Leaflets glabrous, veins not raised ........Adiantum flabellulatum


2b. Leaflets hairy, veins raised on both surfaces ........Adiantum hispidulum
1b. Rachis not dichotomizing, fronds linear, lanceolate ........A. stanochlamys
Adiantum hispidulum Sw. closely resembles A. flabellulatum. However, the former
is clearly distinct from the latter by having multicellular hairs throughout the frond and
tufted at the base, bearing only pinnately pinnules, veins raised on both surfaces, indusial
flaps oblong bearing brown hairs.
DISCUSSION & CONCLUSION

Conservation status
A small population of about eighteen individuals including five ones has been
found. We considered registration of Adiantum hispidulum Sw. as Data Deficient (DD)
according to IUCN conservation status since only one population has been found. It is hoped
that some more populations will be found elsewhere in the other protected reserve forests
of the district.
During the present work it has been revealed that due to over population,
construction of roads, collection of fire wood, deforestation the plant is going to lose the
natural habitat. Lack of proper understanding of the status of this fern make the species
vulnerable and put in the danger of extinction. (Language needs revision)
ACKNOWLEDGEMENTS

The first author is thankful to the principal chief conservator of Forest Government
of Manipur for giving permission to study the reserve forest area of Senapati district and
N. Raghumani Singh, Head of Department of Botany Manipur College to provide the
laboratory facilities for the taxonomic study.
REFERENCES

BEDDOME R H 1883, 1892 Handbook to the Ferns of British India, Ceylon and the Malaya
Peninsula with supplement, Tac hker Spink & Co. Calcutta [Re pr int New Delhi !!]
B O RT H A K U M A R S K , D E K A P & NAT H K K 2 0 0 0 Illustrated Manual of Ferns of Assam Bishen
Singh Mahendra Pal Singh Dehradun
DEVI S YUMKHAM 2008 S p e c i e s D iv e r s i t y a n d P a l y n o l o g y o f P t e r i d o p h y t e s i n T h o u b a l D i s t r i c t
Manipur Ph.D. Thesis submitted to Manipur University Canchipur India
JA I N S K & R AO R R 1 9 7 6 H a n d b o o k o f F i e l d a n d H e r b a r i u m M e t h o d s To d a y a n d To m o r r o w
Publishers New Delhi
NAYAR B K & KAUR S 1974 Companion to R H Beddomes Handbook to the Flora of Br itish India
Ceylon and Malaya Peninsula. The Chronicle Botanica New Dehli
SINGH O K 1990 F l o r i s t i c S t u d y o f Ta m e n g l o n g D i s t ri c t , M a n i p u r w i t h e t h n o b o t a n i c a l n o t e s P h . D
Thesis Submitted to Manipur University Canchipur India
Th. Sunita Devi et al. : Adiantum hispidulum Swartz – A New Report from Senapati District, Manipur 45

S I N G H S & PA N I G A R H I G 2 0 0 5 Ferns and Fern Allies of Arunachal Pradesh Bishen Singh


Mahendra Pal Singh Dehradun
SINHA S C 1986 Ethanobotanical study of Manipur Ph D Thesis submitted to Manipur University
Canchipur India
S M I T H A R , P RY E R K M , S C H U E T T P E L Z E , KO R A L L P, S C H N E I D E R H & W O L F P G 2 0 0 6 A
classifica tion of extant fer ns Taxon 5 5 ( 3 ) : 7 0 5 - 7 3 1
VERMA S C & FRASER-JENKINS C R 2008 Adiantum philippense L The correct name for
Adiantum lunulatum Burm f and its subspecies In Perspectives in Pteridophytes (eds S C
Ve r m a , S P K h u l l a r & H C h e e m a ) B i s h e n S i n g h M a h e n d r a Pa l S i n g h , D e h r a D u n p p 6 5 - 9 2
YO U X I N G L & P R A D O J 2 0 1 2 A d i a n t a c e a e I n : Flora of China ( o n l i n e ) h t t p : / / f o c. b i o - m i r ror. c n /
m s s / vo l u m e 0 2 / A d i a n t a c e a e - M O _ c o a u t h o r. h t m

* * * * *

HEARTIEST CONGRA
HEARTIEST TULA
CONGRATULATIONS T O DR. SA
TULATIONS TISH CHANDER VERMA
SATISH

Members of The Indian Fern Society wish to convey their great


appreciation and heartiest congratulations to Dr. Satish Chander Verma, Professor
Emeritus and Editor of the Indian Fern Journal, for being honoured with the
prestigious INSA Teachers Award (2014), during the 80th Anniversary General
meeting of INSA, to be held at GOA Dec 19-21, 2014. During the 101st Indian
Science Congress Session at Jammu, February 2014, he was honoured with the
prestigious ISCA, Professor Archana Sharma Memorial Award.

S. P
P.. Khullar
Secretary, IFS
Indian Fern J. 3 1 : 46-52 (2014) ISSN 0970-2741

ETHNO-MEDICO-BO
ETHNO-MEDICO-BOTTANICAL STUDIES ON
SOME PTERIDOPHYTES OF RAJ
RAJAASTHAN
R. P. KANTHER1* AND DILIP GENA2
1
Department of Botany, S. D. Government College, Beawar (Rajasthan)
2
Department of Botany, Government College, Ajmer (Rajasthan)

(Received December 20, 2013; Revised Accepted June 10, 2014)

ABSTRACT
The object to investigate the issue of recognizing, documentation and acknowledging
of the oral and traditional information related with the true value to medicinal potential of
some pteridophytes of Rajasthan. The Rural and tribal people of Rajasthan have developed and
follow their own household medicinal practices, Many pteridophytes are being used by them to
cure human and veterinary ailments. Each tribal community has its ancient knowledge and they
treat various diseases by making use of the ferns available around their region. They prepare
pastes, decoctions, aqua extract, powder and juice from pteridophytes. Twenty two
pteridophytes were surveyed for ethno-medicinal potential from Rajasthan, after the field survey
and communications with tribal, Ojhas, Vaidhyas, Bhopas and elderly persons (above 60 years).
The survey revealed that all these documented pteridophytes have great ethno-medicinal
potential. In this present enumeration an attempt has been made to document the oral
indigenous information about the use of pteridophytes growing in Rajasthan. Earlier reports in
published literature on the subject have also been taken into account and an attempt to develop
a prospect in ethno-medico-botany is being made.
K e y Wo rrd
d s : Tribes, Ethno-Medico-Botany, Pteridophytes, Rajasthan

INTRODUCTION

Ancient Indian mythological literature like the Vedas, Ramayan, Mahabharat, the
Bhagwat Gita, Puran, Folklore, Folk talks and legends etc are all replete with the importance
of plants-mainly the flowering plants but there is hardly any mention of pteridophytic plants
however, there is mention of “Sanjeevani” (Selaginella) in the Ramayana and “Hansraj” in
Ayurveda could be some (Adiantum spp.). A Few ferns find mention in ancient Greek and
Latin literature. But the economic value of pteridophytes has been known to man for more
than 2000 years. Several species of pteridophytes have been found to be source of food and
medicine. In India, Pteridophytes have remained a much neglected group of plants so far as
their ethno-botanical importance is concerned (Dixit 1975). As such the pteridophytic flora
of Rajasthan also had not attracted attention regarding their medicinal and other economic
value. Sharma & Vyas (1985) were the first to undertake ethno-botanical studies on the fern
and fern allies of Rajasthan. Rajasthan state is situated in the north western part of India
between 2303'N and 30012'N latitude and 69030'E and 78017'E longitude. Rajasthan has a
hot dry summer and cold bracing winter and the rainy season is comparatively short yet
pteridophytic flora represented almost in all Rajasthan.

* E-mail : rpkanther@gmail.com
R P Kanther & Dilip Gena : Ethno-Medico-Botanical Studies on Some Pteridophytes of Rajasthan 47

Fig.1 : Distribution of Pteridophytes in Rajasthan

REVIEW

Information on economic utility of pteridophytes has been given by various workers


from time to time. Information on the Systematics and Taxonomy of pteridophytes of
Rajasthan has been given by Gena 1998, Parihar and Bohra (2004) reported that the extract
of Adiantum spp. were effective against the growth of Salmonella typhi, the causal organism
of typhoid fever. Manickam et al. (2005) described the antibacterial effect of Christella
parasitica leaf glands against human bacterial pathogens. Parihar and Bohra (2006) published
antibacterial effect of Dryopteris cochleata. Parihar and Parihar (2006) studied antimicrobial
characteristics of the species of Adiantum . Chaudhary and Dulavat (2006) reported
occurrence of 19 species of pteridophytes from Sitamata wild life sanctuary, Rajasthan.
Chaudhary and Dulawat (2008) carried out as extensive survey of various localities of
South-East Rajasthan and reported 27 species of pteridophytes belonging to 10 families of
this part of Rajasthan. Bhardwaj & Gupta (2010) studied the medicinal properties of
pteridophytes of the Hadauti Plateau, Rajasthan.
MATERIALS AND METHODS
ATERIALS

The present work was based on the Ethno-medicinal studies of pteridophytes of


Rajasthan. The several tribes like the Sahariyas, Bhils, Gadia Lohar, Raibaries, Kanjars,
Sansi, Kalbelias, in Rajasthan were the main source of this survey.
48 Indian Fern Journal Volume XXXI (2014)

To achieve the goal, botanical excursions were taken two of three times in a month.
Field trips were arranged in such a way as to cover all pteridophytic localities and its
surrounding tribal’s. Interview were conducted with the tribal people and local Vaidyas,
Ojhas, and medicine men, Generally tribesmen who know about the medicinal plants do
not want to give all in formations because they believe that medicinal plant if disclosed its
medicinal properties will be lost forever. For this reason, the information collection from
the tribal is an important aspect of ethno-medico studies. The information which was
provided us was oral information and it was crossed examined at different places.
Efforts were made to identify the pteridophytic materials. The species were
identified with the help of Khullar’s (1994 & 2000) “An illustrated fern flora of the West-
Himalaya” Vols. I & II and with the help of well-known pteridologist and former vice-
chancellor Dr. C. B. Gena.
OBSER
RVVAT I O N S

Sixty three species belonging to 29 genera of pteridophytes have represented by


various workers from Rajasthan. Out of these 23 species of 16 genera were found being
used as ethno-medicines by tribal’s of Rajasthan. Botanical name, family, plant part useful,
medicinal uses of each species are provided here as follows, arranged alphabetically.
1. Actiniopter is rradia
Actiniopteris adia ta L. (Family : Adiantaceae)
adiata
— Fronds are useful in skin disease, plant is bitter, having properties the styptic,
anthelmintic.
2. Adiantum ca pillus-v
capillus-v
pillus-vener is L (Family : Adiantaceae)
eneris
ener
— Fronds and rhizome is useful in fever and cough. It has hypoglycemic,
aphrodisiac, antibacterial anti fungal and antiviral properties.
3. Adiantum incisum F or
For ssk. (Family : Adiantaceae)
orssk.
— Rhizome extract is used by tribesmen in bronchitis, and in cough of children’s.
4. Adiantum philippense L (Family : Adiantaceae)
— Fronds and rhizome are used in fever and dysentery.
— Fronds extract is used in fever, asthma, bronchitis etc.
— The paste of plant used in the treatment of throat swellings in the cattle.
5. Ceratopteris thalictroides (L.) Brong. (Family : Parkenaceae)
— Fronds paste is used in skin disease for its antifungal properties.
6. Cheilanthus albomanginata Clarke (Family : Cheilanthaceae)
— Whole plants have properties against Tuberculosis.
— Tribal’s use extracts as tonic
— Plant has anti bacterial potential
7. Cheilanthus bicolor (Ro (Roxbxb.) F
xb.) Frr aser -J
aser-J enk. (Family : Cheilanthaceae)
-Jenk.
— Tribal’s use rhizome and root extracts as general tonic and in the cure of urine
problems.
— Mature plant decoction is used for wound healing of cattle’s.
R P Kanther & Dilip Gena : Ethno-Medico-Botanical Studies on Some Pteridophytes of Rajasthan 49

8. Christella dentata (Forrsk.) Brownsey & Jermy (Family : Thelypterdiaceae)


— The paste of plant applied to treat backache.
— Decoction of mature leaf is used for hair treatment.
9. Dryopteris cochleata Buch.-Ham. ex D Don (Family : Dryopterdiaceae)
— Whole plant extract is given twice daily orally in case of snakebite.
— Rhizome has antibacterial and anti epileptic properties.
10. Equisetum rramosissim
amosissim
amosissimum um Desv. (Family : Equisetaceae)
— Strobili use for medicines. Whole plant mixed with mustard oil is use in the
treatment of bone fracture, backache and muscular pain.
— Powdered stem dissolved in water in used for enema during stomach disorder.
11. Hypodematium crenatum (Forrsk.) Kuhn (Family : Dryopteridacease)
— Rhyzome is used as an antibacterial agent.
— Leaves are used to facilitate conception in women.
12. Isoetes reticulata Gena & Bhardwaja (Family : Isoetoceae)
— Tribal’s use rhizome decoction as s lotion for burn.
— It is also known as antibacterial, anticancer us, antiseptic properties.
13. Marsilea minuta L. (Family : Marsileaceae)
— Whole plant is used in cough
— Plant has antibacterial properties.
14. Mar silea cor
Marsilea omandelina Willd. Complex (Family : Marsileaceae)
coromandelina
— Plant is used in skin diseases.
15. Nephrolepis cordifolia (L) Presl. (Family : Nephrolepidaceae)
-Rhizome is antibacterial and use in cough, rheumatism etc.
16. Ophio glossum costa
Ophioglossum tum R. Br
costatum Br.. (Family : Ophioglossaceae)
— Plant is used in wounds, and inflammations.
17. Ophio glossum ggrr amineum Willd
Ophioglossum illd.. (Family : Ophioglossaceae)
— Tribals use rhizome decoction as s lotion for burn. It is also known as
antibacterial, anticancer us, antiseptic properties.
18. Ophioglossum reticulatum L. (Family : Ophioglossaceae)
— Used as cooling agent and in the treatment of inflammation and wounds.
— The warm decoction of rhizome used as a lotion fire burns.
19. Pteris vittata L. (Family : Pteridaceae)
— Extract of fronds used in curing skin disease.
— Plant extracts is used as demulcent, hypertensive tonic antiviral and antibacterial.
20. Salvinia aur icula
auricula ta A ub
iculata let (Family : Salviniaceae)
ublet
— Plant used as antifungal agents.
21. Selaginella rajasthanensis Gena (Family : Selaginellaceae)
— Whole plant grinded and applied on wounds, cuts etc
— Possess antiseptic Properties.
50 Indian Fern Journal Volume XXXI (2014)

Selag
22. Sela g inella rree panda (Desv. Ex Poir.) Spring (Family : Selaginallaceae)
— Fronds are used as antibacterial agent
— Possess antiseptic Properties
23. Tectar
ectariaia macr odonta (J
macrodonta (J.. Smith) C Chr
Chr.. (Family : Dryopteridaceaae)
— Fresh Rhizome extract is used for preventing dysentery in children’s.
— Extraction of dried leaves is used in respiratory disorders like cough, asthma
and bronchitis.
DISCUSSION

Twenty three species were found being used as ethno-medicines by tribal’s of


Rajasthan. This data indicates that pteridophytes have an important role in the utilization of
natural resources as a medicine. Further it has been observed that pteridophytes are the most
frequently utilized plants/plant- parts against various ailments. Preparation of leaf decoction
is the most common traditional formulation prepared followed by paste and application of
fresh juice. The ranking of the parts of plants being used against different is as follows-
Fronds > whole plants > rhizome > corm> Strobili. A few parts were found to be effective
against more than one disease. The data complied was compared with pertinent published
literature. The present investigation proves that ethno medicinal knowledge is also important
from humanitarian point of view.

Fig.2 : Relative contributions of various pteridophytes in different disease as medicine.


R P Kanther & Dilip Gena : Ethno-Medico-Botanical Studies on Some Pteridophytes of Rajasthan 51

PLATE I
PLATE

1. Actiniopteris radiata L. 2. Adiantum capillus-veneris L.


(Family : Adiantaceae) (Family : Adiantaceae)

3. Adiantum incisum Forssk. 4. Cheilanthes albomanginata Clarke


(Family : Adiantaceae) (Family : Cheilanthaceae)

5. Marsilea minuta L 6. Nephrolepis cordifolia (L) Presl


(Family : Marsileaceae) (Family : Nephrolepidaceae)
52 Indian Fern Journal Volume XXXI (2014)

REFERENCES

BHARWAJ A & GUPTA C S 2010 Studies on Some Medicinal Pteridoph ytes of the Hadauti Pla teau
Rajasthan Indian Fern J 2 7 : 332-337
CHAUDHARY B L & DULAWAT C S 2006 Distributation of fern and fern-allies in Sitamata Wild Life
Sanctuary Rajasthan India Indian Fern J 2 3 : 75-82
CHAUDHARY B L & KHICHI Y S 2006 Ferns of Kumbhalgarh Wild life sanctuary in Rajasthan India
Indian Fern J 2 3 : 83-91
DIXIT R D 2000 Conspectus of Pteridophytic Diversity in India Indian J Trad Knowledge 1 7 : 77-91
GENA C B 1998 Systematics and Taxonomy of Pteridophytes of Rajasthan Indian Fern J 1 5 : 139-148
K H U L L A R S P 1 9 9 4 , 2 0 0 0 An illustr a ted fer n f lora of the West-Himala ya Vol I (1994) & II (2000)
International Book Distributers India Dehra Dun
M A N I C K A M V S , B E N N I A M I N A & I RU DAYA R A J V 2 0 0 5 A n t i b a c t e r i a l a c t iv i t y o f l e a f g l a n d s o f
Christella paronitica (L) Lev. Indian Fern J 2 7 : 87-88
MAO A A , Hynniewta T M & SANJA P PA M 2009 Plant w ealth of Nor theast India with refer ence to
ethnobotany Indian J Trad Knowledge 8 ( 1 ) : 96-103
M A S A L V P, M E E NA M & D O N G E R 2 0 1 0 S t u d i e s o n E t h n o - M e d i t a t i o n a l u s e s o f p t e r i d o p hy t e s of
Ratnagiri District (Maharashtra India) Indian Fern J 2 7 : 88-93
PARIHAR P & BOHERA A 2004 Ant salmonella activity of three species of Adiantum from Rajasthan
Western India Indian Fer n J 2 1 : 136-139
PA R I H A R P & PA R I H A R L 2 0 0 6 A n t i m i c ro b i a l c h a r a c t e r i s t i c o f T hr e e S p e c i e s o f A d i a n t u m I n d i a n
Fern J 2 3 : 35-42
SHARMA N K 2002 Ethno medicinal studies on ferns and fern allies Zoo's Print J 1 7 ( 3 ) : 732-734
SHIPI DAS 2003 Usefulness of pteridophytes in India with special reference to medicine and
conservation J Econ Taxon Bot 2 7 ( 1 ) : 7-16
SRIVASTAVA K 2007 Importance of f erns in human medicine Ethnobot Leaflets 1 1 : 231-234
VYAS M S 2008 Photochemistry of some pteridophytes of Rajasthan Indian Fern J 2 5 : 61-65
YUMKHAN S D & SINGH P K 2011 Less Known fern and fern-allies of Manipur with Ethnobotanical
uses Indian J Trad Knowledge 1 0 ( 2 ) : 287-291
Indian Fern J. 3 1 : 53-68 (2014) ISSN 0970-2741

PHENETICS OF SOUTH INDIAN TRICHOMANOID FERNS


P. K. VIDYA VARMA1, K. V. MOHANAN2 AND P. V. MADHUSOODANAN3*
1
Department of Botany, Govt. College Madappally, Kerala-673102
2
Department of Botany, University of Calicut, Kerala-673635
3
Malabar Botanical Garden, Calicut, Kerala-673014

(Received April 21, 2014; Revised Accepted July 5, 2014)

ABSTRACT
Cluster analysis of twenty-two taxa of Trichomanes s. l. of South India was done
using twenty multi-state qualitative morphological characters. Cluster analysis was carried out
adopting UPGMA algorithm and percentage disagreement as the statistical test and a
dendrogram was constructed. The dendrogram obtained through the analysis revealed that the
Trichomanes s. l. of the region falls under three distinct groups. The affinities of the five
newly described taxa are also revealed.
K e y Wo rrd
d s : Trichomanoid ferns, South India, Phenetics, dendrogram

INTRODUCTION

Hymenophyllaceae Link is a leptosporangiate family of filmy ferns with ca 600


species (Iwatsuki, 1990) and wide distribution in the tropical rain forests and some wet
temperate regions. The family is characterized by small plants with single-cell thick lamina
and mariginal sori with sporangia on slender receptacle covered by involucres. Traditionally,
the family is comprised of two large genera i.e., Hymenophyllum Sm. (hymenophylloid
ferns) characterized by bivalvate involucres, and Trichomanes L. (trichomanoid ferns),
characterized by tubular involucres. Trichomanes s. l. is morphologically and ecologically
more diverse than the sister genus, Hymenophyllum s. l. The systematics of Trichomanes
s. l. is highly controversial due to the morphological convergence shown by the members
of the genus in relation to the variations in the habitats they occur.
In South India, filmy ferns are found in the humid tropical rain forests of the
Western Ghats. They are restricted to the deeply shaded high altitude localities of these
forests. They are either epiphytic or lithophytic except Trichomanes obscurum, which is
terrestrial. 15 species of filmy ferns were reported from South India by Beddome (1863,
1883) and Manickam & Irudayaraj (1992) whereas Chandra (2000) reported eighteen species
from the region. The floristic study of Hymenophyllaceae of South India by Hameed et al.
(2003) reported twenty seven species, which include five new species of Trichomanes s. l.
(Hameed & Madhusoodanan 1998, 1999, 2003, Madhusoodanan & Hameed 1998, 1999).
Fraser – Jenkins (2008) has raised doubts on the status and validity of some of these newly
described species.
The present study attempts to find out the affinities of the Trichomaniod ferns and
the newly described taxa of Trichomanes s. l. in the South Indian region. (Figs. a – x)

* E-mail : pvmadhu@gmail.com
54 Indian Fern Journal Volume XXXI (2014)

MATERIALS AND METHODS


ATERIALS

Twenty two taxa of Trichomanes s. l. (Operational Taxonomic Units, of Sokal &


Sneath 1963) are considered in this study with their code numbers given in Table 1. The
data for the analysis were collected from herbarium specimens deposited in the Calicut
University Herbarium (CALI) and also from materials preserved in 4% formaldehyde.
Twenty multi-state qualitative morphological characters of the sporophyte alone were
considered in the present study. The characters (with abbreviated codes) used in the cluster
analysis with their character states are given in Table 2. The characters with two states were
coded as 1 and 2 and those with more than two states were given numbers continuously
TABLE 1 : Species of Trichomanes s.l. (OTUs) selected for the study

Code No. Name of taxa (OTUs)

1 T. (Crepidomanmes) agasthianum Madhus. & Hameed


2 T. bilabiatum (Nees & Blume) Copel.
3 T. bimarginatum (Bosch) Ebihara & K.Iwats.
4 T. bipunctatum (Poir.) Copel.
5 T. christii (Copel) Copel.
6 T. exiguum (Bedd.) Copel.
7 T. henzaianum Parish ex Hook
8 T. indicum C.A.Hameed & Madhus.
9 T. insigne (Bosch) S.H. Fu
10 T. intramarginale (Hook. & Grev.) Copel.
11 T. kurzii (Bedd.) Tagawa & K.Iwats.
12 T. (Crepidomanes) lunulatum Madhus. & C.A. Hameed
13 T. (Crepidomanes) malabaricum C.A. Hameed & Madhus.
14 T. mindorense H.Christ.
15 T. (Crepidomanes) proliferum var. minutum (Blume) C.A.Hameed comb. novo
16 T. obscurum (Blume) Ebihara & K.Iwats.
17 T. plicatum (Bosch) R.C. Ching
18 T. proliferum var. proliferum (Blume) Bostock
19 T. saxifragoides (C. Presl) P.S. Green
20 T. schmidianum (Zenker ex Taschner) K.Iwats.
21 T. sublimbatum Mull. Hal.
22 T. vamana Hameed & Madhus.
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns 55

TABLE 2 : Characters used in the cluster analysis

Sl. No. Code Characters Character States

1 Ha Habit : epiphyte, lithophyte or terrestrial


2 Rhi Rhizome : erect to suberect or creeping
3 FrA Frond arrangement : distant or close
4 FrAr Frond architecture : catadromous or anadromous
5 SH Presence of stipe hair : present or absent
6 SW Stipe wing : winged at apex, winged upto the base or
wing absent
7 LaSh Lamina shape : spatulate, deltoid, orbicular, ovate to
elliptical, obovate, lanceolate or ovate to
oblong
8 LA Lamina apex : acute, acuminate or obtuse
9 LaM Lamina margin : undulate, sinuous, crenate or entire
10 LaMorp Lamina morphology : simple, pinnatifid, bipinnatifid, tripinnatifid,
digitate, flabellate or simple with apical
lobes
11 LaB Lamina base : cuneate, truncate, acute, attenenate or
cordate
12 Ven Venation : pinnate, digitate or flabellate
13 SFV Submarginal false vein lets : continuous, interrupted or absent
14 StP Presence of straie : present or absent
15 LFV Presence of lateral false veinlets : present or absent
16 IS Shape of Indusia : bivalvate, campanulate, obconic or tubular
17 IL Indusial lip : rounded, triangular or absent
18 TT Trichomes on lamina : glandular, linear or absent
19 Re Receptacle at maturity : included or exserted
20 SpO Spore ornamentation : granulate, spinulose, echinate, tuberculate,
verrucate

starting with 1. The results of multiple range tests of characters of the OTUs are given in
the Table 3. For cluster analysis, all the twenty characters were tabulated against the twenty-
two OTUs using the numerical codes given for character states and abbreviated codes of
56
TABLE 3 : Results of multiple range test of characters

Code for Characters Grouping of taxa (represented by code numbers) according to their character states

Characters 1 2 3 4 5 6 7

Ha Habit Epiphyte Lithophyte Terrestrial


1, 3, 4, 5, 7, 2, 6, 9, 10, 11, 16
8, 14, 20, 12, 13, 15, 17,
21, 22 18, 19
Rhi Rhizome Erect to Creeping
sub erect 1, 2, 3, 4, 5, 6,
16 7, 8, 9, 10, 11,
12, 13, 14, 15,
17, 18, 19,
20, 21, 22
FrA Frond Distant Close
arrangement 1, 2, 3, 4, 5, 16
6, 7, 8, 9, 10,
11, 12, 13, 14,
15, 17, 18, 19,
20, 21, 22
FrAr Frond Catadromous Anadromous
architecture 3, 6, 7, 14, 1, 2, 4, 5, 8,
21, 22 9, 10, 11, 12,
Indian Fern Journal Volume XXXI (2014)

13, 15, 16, 17,


18, 19, 20
SH Stipe hair Present Absent
1, 3, 4, 5, 6, 7, 2, 18, 19
8, 9, 10, 11, 12,
13, 14, 15, 16,
17, 20, 21, 22
SW Stipe wing Winged at Winged up Absent
apex to the base 1, 3, 6, 8,
2, 4, 5, 9, 10, 12, 7, 11, 17 13, 14, 15,
19, 20, 21,22 16, 18

Contd.
TABLE 3 : Results of multiple range test of characters

Code for Characters Grouping of taxa (represented by code numbers) according to their character states

Characters 1 2 3 4 5 6 7
LaSh Lamina shape Spatulate Deltoid Orbicular Ovate to Ovate Lanceolate Ovate to
7, 8, 11, 13, 16 1, 15, 19 elliptical 10 5, 18 Oblong
14, 21, 22 2, 6, 20 3, 4, 9,
12, 17
LA Lamina Acute Acuminate Obtuse
apex 2, 4, 5, 6, 9, 16 1, 3, 7, 8,
10, 11, 12, 13, 14, 15, 19,
17, 18, 20 21, 22
LaM Lamina Undulate Sinuous Crenate Entire
margin 1, 3, 9, 10, 22 6
13, 14, 17,
21
LaMorp Lamina Simple Pinnatifid Bipinnatifid Tripinnatifid Digitate Flabellate Simple with
morphology 6, 21 8, 9, 10, 11, 20 2, 4, 5, 1, 15 19 a pical lobe
12, 13, 18 16, 17 3, 7,
14, 22
LaB Lamina base Cuneate Truncate Acute Attenenate Cordate
7, 8, 10, 4, 9, 18 1, 2, 3, 5, 14 15, 19
11; 16, 21, 6, 12, 13,
22 17, 20
Ve n Venation Pinnate Digitate Flabellate
2, 3, 4, 5, 6, 1 15, 19
7, 8, 9, 10, 11,
12, 13, 14, 16,
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns

17, 18, 20,


21, 22
SFV Submarginal Continuous Interrupted Absent
false vein 1, 3, 4, 8, 2, 5 6, 7, 9, 15,
lets 10, 11, 12, 16, 17, 18, 19,
13, 14 20, 21, 22
57

Contd.
58
Contd
Contd.. Ta b le 3

Code for Characters Grouping of taxa (represented by code numbers) according to their character states

Characters 1 2 3 4 5 6 7
StP Presence Present Absent
of straie 2, 4, 5, 9, 17 1, 3, 6, 7, 8,
10, 11, 12,
13, 14, 15, 16,
18, 19, 20, 21,
22
LFV Lateral false Present Absent
veinlets 3, 6, 7, 14, 1, 2, 4, 5,
21, 22 8, 9, 10, 11,
12, 13, 15, 16,
17, 18, 19, 20
IS Shape of Bivalvate Campanulate Obconic Tubular
indusia 2, 9 1, 3, 5, 7, 8, 11, 4, 10, 17 6, 15, 18
12, 13, 14, 16,
19, 20, 21, 22
IL Indusial lip Rounded Triangular Absent
2, 3, 5, 6, 7, 4, 17 1, 8, 12, 13,
9, 10, 11, 14, 15, 16, 18,
19, 21 20, 22
TT Trichomes Glandular Linear Absent
Indian Fern Journal Volume XXXI (2014)

on lamina 1, 5, 8, 10, 3, 6, 7, 9 2, 4, 14, 16,


11, 12, 13, 17, 20, 21, 22
15, 18, 19
Re Receptacle at Included Exserted
maturity 1, 7, 8, 11, 2, 3, 4, 5, 6,
12, 15, 22 9, 10, 13, 14,
16, 17, 18,
19, 20, 21
SpO Spore Granulate Spinulose Echinate Tuberculate Verrucate
ornamentation 9, 16, 20, 22 2, 4, 5, 6, 15 1, 3, 7, 8, 11, 19
12, 14, 17, 10, 13
18, 21
Contd.
TABLE 4 : Character states of the OTUs

Character Codes for OTUs

Code
Codess 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Ha 1 2 1 1 1 2 1 1 2 2 2 2 2 1 2 3 2 2 2 1 1 1

Rhi 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2

FrA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1

FrAr 2 2 1 2 2 1 1 2 2 2 2 2 2 1 2 2 2 2 2 2 1 1

SH 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 1

SW 3 1 3 1 1 3 2 3 1 1 2 1 3 3 3 3 2 3 1 1 1 1

LaSh 3 4 7 7 6 4 1 1 7 5 1 7 1 1 3 2 7 6 3 4 1 1

LA 3 1 3 1 1 1 3 3 2 1 1 1 1 3 3 2 1 1 3 1 3 3

LaM 1 4 1 4 4 3 4 4 1 1 4 4 1 1 4 4 1 4 4 4 1 2

LaMorp 5 4 7 4 4 1 7 2 2 2 2 2 2 7 5 4 4 2 6 3 1 7

LaB 3 3 3 2 3 3 1 1 2 1 1 3 3 4 5 1 3 2 5 3 1 1

Ve n 2 1 1 1 1 1 1 1 1 1 1 1 1 1 3 1 1 1 3 1 1 1

SFV 1 2 1 1 2 3 3 1 3 1 1 1 1 1 3 3 3 3 3 3 3 3

StP 2 1 2 1 1 2 2 2 1 2 2 2 2 2 2 2 1 2 2 2 2 2

LFV 2 2 1 2 2 1 1 2 2 2 2 2 2 1 2 2 2 2 2 2 1 1

IS 2 1 2 3 2 4 2 2 1 3 2 2 2 2 4 2 3 4 2 2 2 2
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns

IL 3 1 1 2 1 1 1 3 1 1 1 3 3 1 3 3 2 3 1 3 1 3

TT 1 3 2 3 1 2 2 1 2 1 1 1 1 3 1 3 3 1 1 3 3 3

Re 1 2 2 2 2 2 1 1 2 2 1 1 2 2 1 2 2 2 2 2 2 1

SpO 4 2 4 2 2 2 4 4 1 4 5 2 4 2 3 1 2 2 5 1 2 1
59
60
TABLE 5 : Percent disagreement between OTUs under study

OTUs arranged as per their numerical codes


OTU's
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

1 0 0.75 0.4 0.65 0.55 0.7 0.55 0.25 0.7 0.5 0.5 0.4 0.3 0.5 0.35 0.65 0.65 0.6 0.55 0.5 0.6 0.55
2 0.75 0 0.7 0.35 0.25 0.5 0.75 0.7 0.45 0.5 0.55 0.45 0.55 0.65 0.7 0.65 0.35 0.45 0.5 0.4 0.6 0.75
3 0.4 0.7 0 0.6 0.55 0.4 0.3 0.45 0.55 0.5 0.6 0.55 0.4 0.2 0.7 0.7 0.6 0.7 0.65 0.55 0.35 0.45
4 0.65 0.35 0.6 0 0.3 0.65 0.7 0.55 0.45 0.45 0.55 0.4 0.55 0.55 0.7 0.6 0.25 0.5 0.65 0.4 0.55 0.65
5 0.55 0.25 0.55 0.3 0 0.55 0.6 0.5 0.5 0.45 0.45 0.35 0.45 0.55 0.65 0.6 0.4 0.45 0.5 0.35 0.5 0.65
6 0.7 0.5 0.4 0.65 0.55 0 0.5 0.7 0.55 0.55 0.6 0.55 0.5 0.45 0.6 0.7 0.5 0.45 0.65 0.5 0.4 0.6
7 0.55 0.75 0.3 0.7 0.6 0.5 0 0.35 0.65 0.6 0.4 0.6 0.6 0.35 0.6 0.65 0.7 0.7 0.6 0.55 0.3 0.25
8 0.25 0.7 0.45 0.55 0.5 0.7 0.35 0 0.65 0.4 0.25 0.3 0.25 0.45 0.4 0.5 0.7 0.45 0.55 0.45 0.5 0.4
9 0.7 0.45 0.55 0.45 0.5 0.55 0.65 0.65 0 0.4 0.55 0.5 0.5 0.65 0.65 0.6 0.4 0.5 0.55 0.5 0.55 0.65
10 0.5 0.5 0.5 0.45 0.45 0.55 0.6 0.4 0.4 0 0.3 0.35 0.25 0.55 0.6 0.65 0.45 0.45 0.5 0.5 0.5 0.65
11 0.5 0.55 0.6 0.55 0.45 0.6 0.4 0.25 0.55 0.3 0 0.25 0.3 0.55 0.5 0.6 0.55 0.45 0.45 0.5 0.55 0.55
12 0.4 0.45 0.55 0.4 0.35 0.55 0.6 0.3 0.5 0.35 0.25 0 0.25 0.6 0.45 0.6 0.45 0.35 0.5 0.35 0.6 0.55
13 0.3 0.55 0.4 0.55 0.45 0.5 0.6 0.25 0.5 0.25 0.3 0.25 0 0.45 0.5 0.55 0.45 0.35 0.55 0.4 0.55 0.6
14 0.5 0.65 0.2 0.55 0.55 0.45 0.35 0.45 0.65 0.55 0.55 0.6 0.45 0 0.7 0.65 0.6 0.65 0.65 0.55 0.2 0.35
Indian Fern Journal Volume XXXI (2014)

15 0.35 0.7 0.7 0.7 0.65 0.6 0.6 0.4 0.65 0.6 0.5 0.45 0.5 0.7 0 0.6 0.65 0.4 0.35 0.55 0.7 0.6
16 0.65 0.65 0.7 0.6 0.6 0.7 0.65 0.5 0.6 0.65 0.6 0.6 0.55 0.65 0.6 0 0.6 0.55 0.65 0.4 0.6 0.55
17 0.65 0.35 0.6 0.25 0.4 0.5 0.7 0.7 0.4 0.45 0.55 0.45 0.45 0.6 0.65 0.6 0 0.5 0.65 0.45 0.55 0.7
18 0.6 0.45 0.7 0.5 0.45 0.45 0.7 0.45 0.5 0.45 0.45 0.35 0.35 0.65 0.4 0.55 0.5 0 0.45 0.45 0.65 0.7
19 0.55 0.5 0.65 0.65 0.5 0.65 0.6 0.55 0.55 0.5 0.45 0.5 0.55 0.65 0.35 0.65 0.65 0.45 0 0.5 0.55 0.65
20 0.5 0.4 0.55 0.4 0.35 0.5 0.55 0.45 0.5 0.5 0.5 0.35 0.4 0.55 0.55 0.4 0.45 0.45 0.5 0 0.45 0.4
21 0.6 0.6 0.35 0.55 0.5 0.4 0.3 0.5 0.55 0.5 0.55 0.6 0.55 0.2 0.7 0.6 0.55 0.65 0.55 0.45 0 0.25
22 0.55 0.75 0.45 0.65 0.65 0.6 0.25 0.4 0.65 0.65 0.55 0.55 0.6 0.35 0.6 0.55 0.7 0.7 0.65 0.4 0.25 0
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns 61

T r ee dia
diagg r am ffor
or 22 O TUs U n w
OTUs eighted pair
weighted pair-- ggrr oup
average Percent disagreement

Linkage Distance
Fig.1 : Dendrogram of twenty-two taxa of Trichomanes s.l. obtained by phenetic analysis.
1. T. agasthianum , 2. T. bilabiatum , 3. T. bimarginatum , 4. T. bipunctatum , 5. T. christii ,
6. T. exiguum , 7. T. henzaianum , 8. T. indicum , 9. T. insigne , 10. T. intramarginale , 11. T. kurzii ,
12. T. lunulatum , 13. T. malabaricum , 14. T. mindorense , 15. T. proliferum var. minutum , 16. T. obscurum ,
17. T. plicatum , 18. T. proliferum var. proliferum , 19. T. saxifragoides , 20. T. schmidianum , 21. T.
sublimbatum , 22. T. vamana

characters (Table 4). This tabulated data were used to generate dendrogram using the
statistical package STATISTICA version 5.0 loaded in a personal computer, adopting
Unpaired Group Method with Arithmetic mean as algorithm (Sokal & Michener 1958) and
percent disagreement (Hill & Lewicki 2006) as the statistical test. The percent disagreement
table of OTUs is given (Table 5).
RESULT S AND DISCUSSION
RESULT

The dendrogram obtained (Fig. 1) through the cluster analysis clearly revealed that
the 22 taxa of Trichomanoid ferns of South India fall into three distinct groups. The taxa
62 Indian Fern Journal Volume XXXI (2014)

F i gg.. a–h. Trichomanes spp. of South India a. T. agasthianum , b . T. bilabiatum , c . T. bimarginatum ,


d . T. bipunctatum , e . T. christii , f . T. exiguum , g . T. henzaianum , h. T. indicum
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns 63

F i gg.. i – p . Trichomanes spp. of South India i . T. insigne , j . T. intramarginale , k . T. kurzii , l . T.


lunulatum , m . T. malabar icum , n . T. mindor ense , o . T. prolif erum var. minutum , p . T. prolif erum var.
proliferum .
64 Indian Fern Journal Volume XXXI (2014)

F i g . q – x . Trichomanes spp. of South India q & rr.. T. obscurum , s & t . T. saxifragnoides ,


u. T. schmidianum , v. T. plicatum , w. T. sublimbatum , x. T. vamana
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns 65

with their respective groups are given below :


Group I : 16
Group II : 3, 6, 7, 14, 21 and 22
Group III : 1, 2, 4, 5, 8, 9, 10, 11, 12, 13, 15, 17, 18, 19 and 20
Taxa under Group II and III were clustered together and show 60% dissimilarity
with T. obscurum (Group I) which is morphologically very much different from all other
species. Terrestrial habit and erect rhizome with closely arranged fronds are obviously
different from the epiphytic and lithophytic habit and creeping rhizome with distantly
arranged fronds of other two groups. T. obscurum is placed under the genus Cephalomanes
by Iwatsuki (1984) and under the genus Abrodictiyum by Ebihara et al. (2006). But in both
classifications and also in Morton’s classification, this species is under the section
Pachychaetum. The taxa under Group II were 58% dissimilar with Group III and are with
fronds that were smaller in size and simple or simple with apical lobed lamina. They also
show lateral false veinlets in the lamina. The taxa under Group III were comparatively
larger sized plants with dissected lamina, and without lateral false veinlets.
In Group II, T. exiguum is clustered distantly from other taxa in the group. T.
exiguum is distinct from other taxa in having marginal stellate setae and simple fronds with
an apical solitary sorus. T. exiguum is placed under section Didymoglossum and subgenus
Didymoglossum by workers such as Morton (1968), Iwatsuki (1984) and Ebihara et al.
(2006). T. vamana clusters with T. henzaianum in this study, but it shows same dissimilarity
percentage with T. henzaianum and T. sublimbatum (25%; Table 5). The number of
dissimilar characters shown by T. vamana with T. henzaianum and with T. sublimbatum is
also same (five characters). Even though, T. vamana is clustered along with T. henzaianum,
further studies are needed to confirm its affinity clearly.
Trichomanes bimarginatum and T. mindorense are clustered together and show the
highest affinity (only 20% dissimilarity) to each other in the analysis and are considered
under the same section Microgonium by various workers (Morton 1968, Iwatsuki 1984,
Ebihara et al. 2006). Both are closely related and are frequently confused with each other
(Croxall 1986). They differ in lamina shape, lamina base, and nature of trichomes in lamina
and spore ornamentation.
Group III consists of two clusters viz., Group III-a and Group III-b. The taxa
clustered together in Group III-a are : T. insigne, T. schmidianum, T. christii, T. plicatum,
T. bipunctatum, and T. bilabiatum. These are situated basally in the cluster. T. insigne was
first described by Beddome (1883) as a variety of T. bipunctatum. Iwatsuki (1985)
considered Crepidomanes insigne (= T. insigne ) and Crepidomanes latealatum (= T.
latealatum) as conspecific whereas Manickam & Irudayaraj (1992) consider it conspecific
with T. plicatum. T. insigne is closely related to T. plicatum and T. bipunctatum and often
confused with T. plicatum, but can be easily distinguished by pinnatifid fronds with
marginal setae in the former. T. schmidianum differs from other taxa in the cluster by the
66 Indian Fern Journal Volume XXXI (2014)

tomentose rhizome, absence of false veins and cupular indusia with rounded mouth.
Trichomanes plicatum and T. bipunctatum show only 25% dissimilarity (Table 5)
and are clustered together in the analysis. Nayar & Kaur (1974) and Chandra & Kaur (1987)
consider T. bipunctatum var. plicatum described by Beddome (1883) as a nomenclatural
equivalent to Crepidomanes plicatum (= T. plicatum). T. plicatum and T. bipunctatum show
close similarity and are distinguishable by the nature of stipe wing, lamina margin, lamina
base and nature of sub marginal false veinlets.
Hameed et al. (2003) consider Trichomanes bilabiatum as closely related to T.
bipunctatum and T. plicatum. Iwatsuki (1985) treated T. bilabiatum as conspecific with T.
bipunctatum. But, in the present analysis, T. bilabiatum is found clustered along with T.
christii, which may be due to the fact that T. bipunctatum and T. plicatum are more closely
related to each other than T. bilabiatum that is evident from the percent disagreement values
(Table 5).
Group III-b 1, consists of three taxa viz ., T. saxifragoides, T. proliferum var.
minutum and T. proliferum var. proliferum. T. proliferum var. minutum recognized by
Hameed et al. (2003) is T. proliferum forma minutum of Manickam & Irudayaraj (1992)
which seems to be T. minutum of Blume (1828). T. proliferum var. proliferum is T.
proliferum of Blume (1828). In this study, T. proliferum var. minutum clusters together
with T. saxifragoides instead of T. proliferum var. proliferum. It shows more similarity with
T. saxifragoides than T. proliferum var. proliferum. Thus, considering this taxon as a variety
of T. proliferum is a debatable matter, which needs further studies. Copeland (1958)
consider, Gonocormus prolifer (= T. proliferum) and G. saxifragoides (= T. saxifragoides)
as seasonal forms of G. minutus (= T. minutum). But, later workers (Tindale 1963, Sledge
1968, Croxall 1975) treated them as separate species. Ebihara et al. (2006) place all these
taxa under section Gonocormus under a single polymorphic species Crepidomanes minutum
(Blume) K. Iwats. They recommended further studies for this taxa as these plants show great
genetic variation in the rbcL sequences when collected from various locations, but which
neither corresponds to morphological features nor to geographical distribution (Ebihara et
al. 2006).
Group III-b2 consists of six taxa. In this group, T. lunulatum clusters together with
T. kurzii but when percent disagreement value is considered, both T. kurzii and T.
malabaricum share same disagreement value with T. lunulatum (0.25). T. malabaricum
shows 75% similarity with T. intramarginale (0.25 percent disagreement value) and 70%
with T. kurzii (0.30 percent disagreement value). Similarity of T. malabaricum with T.
intramarginale was pointed by Hameed et al. (2003). These four taxa show close affinities
with each other and cluster in the basal region of Group III-b2. T. agasthianum and T.
indicum are clustered together sharing a similarity of 75%. The affinity between these two
taxa agrees with the observation by Hameed et al. (2003) who reported that T. lunulatum
shows resemblance with T. agasthianum but the present study shows that T. agasthianum is
P K Vidya Varma et al. : Phenetics of South Indian Trichomanoid Ferns 67

allied more to T. indicum (with 0.25 percent disagreement value).


On the basis of molecular phylogenetic studies Pryer et al. (2001) demonstrated the
occurrence of two lineages in the family viz., Trichomaniod lineage and Hymenophylloid
lineage, the former having with about 325 spp. showing maximum diversity in their
morphology and ecological preferences. Ebihara et al. (2006) recognized eight genera under
trichomaniod lineage. The South Indian trichomaniod ferns fall under three genera
recognized by them viz., Abrodictyum, Crepidomanes and Didymoglossum.
Even though the phenetic analysis gives a picture on the affinities of the taxa of
South India including the newly described taxa from this region, further studies and analysis
with data from other sources such as molecular, cytological and anatomical, etc. are needed
to get a vivid picture about the relationships of Trichomanoid ferns of South India.
ACKNOWLEDGEMENTS

The first author thankfully acknowledges the financial support provided to her by
Council of Scientific and Industrial Research (CSIR), New Delhi for this study.
REFERENCES

BEDDOME R H 1863 T h e F e r n s o f S o u t h e r n I n d i a Vo l u m e - 1 To d a y a n d To m o r r ow ’s P r i n t e r s a n d
Publishers New Delhi
BEDDOME R H 1883 Handbook to the f erns of British India, Ceylon and the Malay Peninsula Today
and Tomorrow ’s Printer s and Publishers New Delhi
BLUME C L 1828 En umeratio Plantar um Javae Van Leeuw en Lugduni Bat avorum (Leiden)
CHANDRA S 2000 The Ferns of India ( Enumeration, Synonyms and Distribution ) International Book
distributors Dehra Dun
CHANDRA S & KAUR S 1987 A nomencla tural guide to R H Beddome’s Ferns of Southern India and
Ferns of British India Today and Tomor row’s Printers and Publishers New Delhi
COPELAND E B 1958 Fer n Flora of the Phillipines-I Institute of Science and Technolo gy Manila
CROXALL J P 1975 The Hymenoph yllaceae of Queensland Aust J Bot 2 3 : 509-547
CROXALL J P1986 Microgonium (Hymenophyllaceae) in Malesia with special reference to Peninsular
Malaysia Kew Bull 4 1 : 519-531
EBIHARA A , DUBUISSON J-Y, IWATSUKI K, HENNEQUIN S & ITO M 2006 A taxonomic revision
of Hymenophyllaceae Blumea 5 1 : 221-280
FRASER-JENKINS C R 2008 Endemics and pseudo-endemics in relation to the distribution patterns
of Indian Pteridophytes Taiwania 5 3 : 264-292
H A M E E D C A & M A D H U S O O DA NA N P V 1 9 9 8 A n ew s p e c i e s o f fi l m y f e r n ( H y m e n o p h y l l a c e a e :
Pteridophyta) from South India Sida 1 8 : 519-522
H A M E E D C A & M A D H U S O O DA NA N P V 1 9 9 9 C r e p i d o m a n e s m a l a b a r i c u m , a n ew fi l m y f e r n
species (Hymenophyllaceae – Pteridophyta) from Southern India Nord J Bot 1 9 : 277-279
H A M E E D C A & M A D H U S O O DA NA N P V 2 0 0 3 Tri c h o m a n e s va m a n a , a n ew f i l m y f e r n s p e c i e s
(Hymenophyllaceae : Pteridophyta) from India Nord J Bot 2 3 : 437-440
H A M E E D C A , R A J E S H K P & M A D H U S O O DA NA N P V 2 0 0 3 Filmy ferns of South India Penta
Book Publishers and Distributors Calicut
HILL T & LEWICKI P 2006 Statistics : Methods and Applications StatSoft Inc U S A
68 Indian Fern Journal Volume XXXI (2014)

I WAT S U K I K 1 9 8 4 S t u d i e s o n t h e s y s t e m a t i c s o f film y f e r n s V I I A s c h e m e o f c l a s s i fi c a t i o n b a s e d
chiefly on the Asiatic species Acta Phytotax Geobot 3 5 : 165-179
I WAT S U K I K 1 9 8 5 T h e H y m e n o p h y l l a c e a e o f A s i a , ex c l u d i n g M a l e s i a J F a c S c i U n iv To k y o 1 3 :
501-551
IWATSUKI K 1990 Hymwnophyllaceae In T he families and genera of Vascular Plants – Pter idophytes
and Gymnosper ms Ed . K Kubitzki Springer Verlag Ber lin Vol 1 Pp 157-163
M A D H U S O O DA NA N P V & H A M E E D C A 1 9 9 8 C r e p i d o m a n e s a g a s t h i a n um , a n ew fi l m y f e r n
species (Hymenophyllaceae) from India Nord J Bot 1 8 : 169-170
M A D H U S O O DA NA N P V & H A M E E D C A 1 9 9 9 C r e p i d o m a n e s l u n u l a t u m ( H y m e n o p h y l l a c e a e :
Pteridophyta) a new filmy fern species from South India Fern Gaz 1 5 : 265-268
MANICKAM V S & IRUDAYARAJ V 1992 Pteridophyte flora of the Western Ghats - South India B I
Publishers Pvt Ltd New Delhi
M O RTO N C V 1 9 6 8 T h e g e n e r a , s u b g e n e r a a n d s e c t i o n s o f H y m e n o p h y l l a c e a e C o n t r U N a t H e r b
3 8 : 153-201
NAYAR B K & KAUR S 1974 Companion to Beddome’s handbook to the ferns of Br itish India Pama
Primlane New Delhi
P RY E R K M, S M I T H A R, H U N T J S & D U BU I S S O N J - Y 2 0 0 1 rbcL da ta reveal two monophyletic
g roups of filmy ferns (Filicopsida : Hyminophyllaceae) Amer J Bot 8 8 : 1118-1130
SLEDGE W A 1968 The Hymenophyllaceae of Ceylon Bot J Linn Soc 6 0 : 289-308
S O K A L R R & M I C H E N E R C D 1 9 5 8 A s t a t i s t i c a l m e t h o d f o r eva l u a t i n g s y s t e m a t i c r e l a t i o n s h i p s
Univ of Kangas Sci Bull 2 8 : 1409-1438
SOKAL R R & SNEATH P H A 1963 Principles of Numerical Taxonomy W H Freeman San F rancisco
U S A
T I N DA L E M D 1 9 6 3 P t e r i d o p hy t a o f S o u t h E a s t e r n A u s t r a l i a C o n t r N S W N a t H e r b F l S e r 2 0 1 :
1-49
Indian Fern J. 3 1 : 69-89 (2014) ISSN 0970-2741

MOLECULAR SYSTEMA
SYSTEMATT I C S O F THE FILMY FERNS
(HYMENOPHYLLA CEAE) OF SOUTH INDIA
(HYMENOPHYLLACEAE)
P. K. VIDYA VARMA1 AND P. V. MADUSOODANAN2*
1
Govt. College, Madappally, Vatakara-673102, Kerala
2
Malabar Botanical Garden, Calicut-673014, Kerala

(Received April 22, 2014; Revised Accepted May 16, 2014)

ABSTRACT
Molecular systematics of the filmy ferns (Hymenophyllaceae) of South India is done
based on rbcL nucleotide sequences. Live materials were collected from Western Ghats of South
India. Fresh and silica dried fronds were used for the extraction of DNA. The rbcL gene was
amplified and sequenced. For phylogenetic analysis 19 sequences (eight newly sequenced and 11
sequences from GenBank) were considered. Multiple sequence alignment was done with
ClustalW2 software. The phylogenetic analysis was done with Neighbor Joining method,
Maximum Likelihood method and Bayesian inference. As a result a conspicuous clustering
pattern was obtained showing the Trichomanoid lineage of South India comprising of three
strongly supported clades which correspond to the three genera recognized by Ebihara et al.
(2006) viz., Abrodictyum, Crepidomanes and Didymoglossum.
d s : Molecular systematics, Trichomanoid lineage, South India, rbcL
K e y W o rrd
gene.

INTRODUCTION

The filmy ferns are unique group of leptosporangiate monilophytes belonging to the
family of Hymenophyllaceae Link comprising around 600 species (Iwatsuki, 1990). They
grow in the interior of dense humid forests either as epipetric forms or as epiphytes (except
certain terrestrial species such as Abrodictyum obscurum) and are easily mistaken for thalloid
bryophytes. The molecular phylogenetic study of the family done by Pryer et al. (2001)
showed that there exist two lineages in the family viz ., Trichomanoid lineage and
Hymenophylloid lineage. In this, the Trichomanoid ferns (about 325 species- Dubuisson
1997) show maximum diversity in their morphology and ecological preferences.
The Trichomanoid ferns are well adapted to the humid rain forests of Western Ghats
region of South India. A detailed study of Hymenophyllaceae of South India done by
Hameed et al . (2003) reported twenty two species of Trichomanoid ferns (including
Vandenboschia radicans ((Sw.) Copel.), which also include five new species under the same
lineage (Hameed & Madhusoodanan 1998, 1999, 2003, Madhusoodanan & Hameed 1998,
1999). But the later studies (Fraser-Jenkins 2008a, b, Varma et al. (2014) on the filmy ferns
of South India raised doubts about species status and relationships of the newly described
taxa of this region.
The present study attempts to determine the relationships exist between the
Trichomanoid taxa of South India by employing the rbcL gene sequences and also to

* E-mail : pvmadhu@gmail.com
70 Indian Fern Journal Volume XXXI (2014)

determine the affinities and status of various newly described endemic species prevailing in
the region.
MATERIALS AND METHODS
ATERIALS

In this study, Trichomanoid ferns (Hymenophyllaceae) distributed all over South


India were selected to deduce the systematic relationship existing between them. The prime
aim of the study was to deduce the relationship between these taxa by employing the
molecular systematic techniques. The specimens for the present study were collected from
Western Ghats forests of South Indian region.
A portion of the collected specimen was used for deposition as voucher specimen
in the Calicut University Herbarium (CALI). The other portion of the collected specimen
was utilized for molecular study. As most of the filmy ferns were epiphytic, they were often
found to grow intermingled with habitually similar bryophytes, to avoid the possible
contamination by bryophytic species, the specimens were examined under Nikon SMZ 800
(Type – 104) stereomicroscope and bryophytes were removed carefully from the fronds. The
photographs were taken using Nikon D 100 digital camera (Figs.1 & 2).The clean fronds
were separated from the rhizome and were used fresh or after silica drying for the extraction
of DNA. DNA extraction was carried out by a modified CTAB method (Doyle & Doyle
1987). Details of the species used for DNA extraction and rbcL amplification was given in

TABLE 1 : Details of taxa used ffor


or the DN
DNAA eex action and rbcL ggene
x t rraction ene amplif ica
icatt i o n
amplifica

S l. Name of the Species Herbarium Nature of


No. accession number the material

1. Crepidomanes bilabiatum (Nees & Blume) Copel. CU.119929 Silica dried


material
2. Crepidomanes indicum C. A. Hameed & Madhus. CU.119958 Silica dried
material
3. Crepidomanes insigne (Bosch) S.H.Fu CU.119903 Silica dried
material
4. Crepidomanes intramarginale (Hook. &G rev.) Copel. CU.119907 Silica dried
material
5. Crepidomanes lunulatum Madhus. & C.A. Hameed CU.119980 Silica dried
material
6. Crepidomanes malabaricum C. A. Hameed & Madhus. CU.119973 Silica dried
material
7. Crepidomanes proliferum (Blume) Bostock var. CU.119985 Fresh material
proliferum
8. Crepidomanes saxifragoides (C. Presl) P.S. Green CU.119989 Fresh material
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 71

the Table 1. The quality of the DNA isolated was checked using agarose gel electrophoresis
(0.8%) using Rice DNA (30ng) as the control. The gels were visualized in a UV
transilluminator (Genei) and the image (Fig.3) was captured under UV light using Gel
documentation system (Bio-Rad).
PCR amplification of rbcL gene was done by using the four primers (two forward
and two reverse). The primer combination aF x JYDS5 and RBCF x 1390R was used
(sequences of aF, JYDS5, 1390R from Pryer et al. (2001) and that of RBCF from (Rajiv
Gandhi Centre for Biotechnology (RGCB) Thiruvananthapuram, Kerala). The PCR
amplifications were carried out in 20 µl reaction volume which contained 1X PCR buffer
(100mM TrisHCl, pH-8.3; 500mM KCl), 0.2mM each of dNTPs (dATP, dGTP, dCTP and
dTTP), 2.0mM MgCl2, 20ng template DNA, 1 unit of AmpliTaq Gold DNA polymerase
enzyme, 0.15 mg/ml BSA and 3% DMSO, 0.5M Betaine, 5pM of forward and reverse
primers.The PCR amplification was carried out in a PCR thermal cycler (GeneAmpPCR
System 9700, Applied Biosystems) with an initial 95 o C denaturation cycle for 5 min,
followed by 40 cycles of 95 o C denaturation for 30 sec, primer annealing at 52o C for 40
sec, elongation at 72o C for 1 min and a final one terminal elongation at 72o C for 7 min.
The primer combinations aF x JYDS5 amplified 912bp and the combination RBCF x 1390R
amplified about 790bp of the rbcL gene. The PCR products were viewed in 1.2% agarose
gel along with100bp DNA ladder (NEB) as the molecular standard (Fig.4). The complete
sequences of the gene were obtained by sequencing each of the amplicon in ABI 3730/3500
Genetic Analyzer (Applied Biosystems). The quality was checked using Sequence Scanner
Software v1 (Applied Biosystems). The sequence fragments were edited and assembled into
contiguous alignment by using Geneious Pro v5.6 (Drummond et al. 2012).
For the phylogenetic analysis of Trichomanoid ferns of South India using rbcL gene
sequences, eight newly sequenced taxa, rbcL sequences of eleven species from GenBank
(including three sequences under the genus Hymenophyllum) were considered (Table 2). The
genus Vandenboschia Copel, which is represented by a single species viz., V. radicans (Sw.)
Copel., is not included in this work as its occurrence in South India is doubtful as it is
first reported by d’Almeida (1926) from the region but later works could not collect this
species from the region. The sequence of Crepidomanes latealatum (Bosch) Copel. was
included in the present study as DNA sequence of C. plicatum (Bosch) R.C. Ching was not
available and C. latealatum (Bosch) Copel. was considered to be conspecific with C.
plicatum (Bosch) R.C. Ching (Hameed et al. 2003).The details of the sequences from
GenBank were given in the Table 2.Multiple sequence alignment of all the 19 species
considered for the analysis was done by the ClustalW2 software (Larkin et al. 2007). The
software Jalview version 2 (Waterhouse et al . 2009) was used to view the SNPs and
conserved regions of the aligned sequences. After the alignment, the sequences of the newly
sequenced taxa were trimmed to 1205bp (which was equal to the length of sequences
retrieved from GenBank) by using the software BioEdit v7.1.11 (Hall 1999). DnaSP v5
72 Indian Fern Journal Volume XXXI (2014)

TABLE 2 : Details of the rbcL sequences of taxa accessed fr om GenBank


from

S l. Name of the Species GenBank


No. accession number

1. Abrodictyum obscurum (Blume) Ebihara & K. Iwats. AB574701


2. Crepidomanes bipunctatum (Poir.) Copel. EU122964
3. Crepidomanes christii (Copel.) Copel. HQ638660
4. Crepidomanes kurzii (Bedd.) Tagawa & K. Iwats. EU122969
5. Crepidomanes latealatum (Bosch) Copel. AB064297
6. Crepidomanes schmidianum (Zenker ex Taschner) K. Iwats. AB378492
7. Didymoglossum exiguum (Bedd.) Copel. AB257488
8. Didymoglossum bimarginatum (Bosch) Ebihara & K. Iwats. AB257494
9. Hymenophyllum acanthoides (Bosch) Rosenst. AB064291
10. Hymenophyllum denticulatum Sw. AB574718
11. Hymenophyllum polyanthos (Sw.) Sw. AB574722

(Librado & Rozas 2009) software was used to identify the polymorphic sites, conserved
DNA regions, mutations and also for finding the haplotype distribution in the sequences.
Estimation of evolutionary divergence between different taxa was done by applying the
distance estimation analysis in MEGA version 5.0 (Tamura et al. 2011) using the Maximum
Composite Likelihood method (MCL method-Tamura et al. 2004) with gamma distribution
for the rate variation among sites.
The phylogenetic analysis was conducted by using Distance, Maximum likelihood
and Bayesian inference approaches. As indels and gaps were not present in the rbcL gene,
all the 1205 positions were considered for the analysis. Before the analysis, evolutionary
model test was done in MEGA version 5.0 (Tamura et al. 2011). Rooting of the tree was
done with the three Hymenophyllum species (specifically using H. polyanthos (Sw.) Sw.).
Bayesian analysis was performed in Mr. Bayes version 3.1 (Huelsenbeck & Ronquist 2001,
Ronquist & Huelsenbeck 2003) with two searches run simultaneously for at least two million
generations. Flat Dirichlet as the prior and General Time Reversible model with a
proportion of invariable sites and a gamma-shaped distribution of rates across sites as the
model of nucleotide substitution were opted. Three heated chains (temperature 0.2) and one
cold chain were used in each search and trees were sampled every 1000 generations. The
parameter was then fixed for a bootstrap analysis with 1000 replicates. Tracer version 1.5
(Rambaut & Drummond 2007) was used to evaluate mixing and convergence, and to estimate
appropriate burn-in period.
For distance methods and maximum likelihood approaches the software MEGA
TABLE 3 : Percent Identity Score generated by ClustalW2 between the 19 taxa using rbcL sequences (1205 bp)

1. A. obscurum, 2. C. bilabiatum , 3. C. bipunctatum , 4. C. christii, 5. C. indicum , 6. C. insigne , 7. C. intramarginale, 8. C. kurzii , 9. C. lunulatum,


10. C.malabaricum, 11. C. latealatum, 12. C. proliferum var. proliferum , 13. C. saxifragoides, 14. C. schmidianum, 15. D. bimarginatum , 16. D. exiguum,
17. H. acanthoides, 18. H. denticulatum, 19. H. polyanthos
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

1 100 91 91 91 91 91 91 91 91 91 91 90 90 90 89 91 91 91 92

2 91 100 98 99 98 99 99 98 99 99 99 94 94 98 90 90 89 89 89

3 91 98 100 99 98 98 98 98 98 98 98 95 94 98 89 90 89 89 89

4 91 99 99 100 98 99 98 98 98 98 98 95 94 98 90 90 89 89 89

5 91 98 98 98 100 98 98 98 98 98 98 94 94 98 90 90 90 89 89

6 91 99 98 99 98 100 99 98 99 99 99 94 94 98 90 90 89 89 89

7 91 99 98 98 98 99 100 98 100 100 98 94 94 98 90 91 89 89 89

8 91 98 98 98 98 98 98 100 98 98 98 94 93 98 89 90 89 89 89

9 91 99 98 98 98 99 100 98 100 100 98 94 94 98 90 91 89 89 89

10 91 99 98 98 98 99 100 98 100 100 98 94 94 98 90 91 89 89 89

11 91 99 98 98 98 99 98 98 98 98 100 95 94 98 90 90 89 89 89

12 90 94 95 95 94 94 94 94 94 94 95 100 96 94 88 89 89 89 89

13 90 94 94 94 94 94 94 93 94 94 94 96 100 94 88 89 89 89 89

14 90 98 98 98 98 98 98 98 98 98 98 94 94 100 89 90 89 89 89

15 89 90 89 90 90 90 90 89 90 90 90 88 88 89 100 95 89 89 89

16 91 90 90 90 90 90 91 90 91 91 90 89 89 90 95 100 89 89 89

17 91 89 89 89 90 89 89 89 89 89 89 89 89 89 89 89 100 99 97

18 91 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 99 100 97
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae)

19 92 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 89 97 100
73
74
TABLE 4 : Estimation of evolutionary divergence between 19 taxa (as distances) using the rbcL sequences (1205 bp). Standard error estimate were
shown above the diagonal

C. saxifragoides
C. proliferum var.
proliferum
C. bilabiatum
C. latealatum
C. intramarginale
C. kurzii
C. bipunctatum
C. christii
C. insigne
C. schmidianum
C. malabaricum
C. lunulatum
C. indicum
D. bimarginatum
D. exiguum
H. acanthoides
H. denticulatum
H. polyanthos
A .obscurum

C. saxifragoides 0.007 0.009 0.009 0.009 0.010 0.009 0.009 0.009 0.010 0.009 0.009 0.010 0.018 0.017 0.017 0.017 0.017 0.015
C. proliferum 0.040 0.009 0.009 0.009 0.009 0.008 0.009 0.009 0.009 0.009 0.009 0.009 0.018 0.017 0.017 0.017 0.017 0.015
var. proliferum
C. bilabiatum 0.059 0.053 0.001 0.003 0.003 0.003 0.003 0.002 0.003 0.003 0.003 0.003 0.016 0.015 0.017 0.017 0.016 0.013
C. plicatum 0.058 0.052 0.002 0.003 0.003 0.003 0.003 0.002 0.004 0.003 0.003 0.004 0.016 0.015 0.017 0.017 0.016 0.014
C. intramarginale 0.054 0.055 0.010 0.011 0.003 0.004 0.003 0.003 0.004 0.000 0.000 0.004 0.015 0.015 0.016 0.016 0.016 0.014
C. kurzii 0.065 0.057 0.012 0.013 0.013 0.004 0.004 0.003 0.004 0.003 0.003 0.004 0.016 0.015 0.017 0.017 0.016 0.014
C. bipunctatum 0.058 0.050 0.012 0.013 0.013 0.015 0.002 0.003 0.004 0.004 0.004 0.004 0.016 0.015 0.016 0.016 0.016 0.014
C. christii 0.058 0.052 0.010 0.011 0.012 0.013 0.003 0.003 0.004 0.003 0.003 0.004 0.016 0.015 0.016 0.016 0.016 0.014
C. insigne 0.058 0.054 0.003 0.004 0.010 0.012 0.012 0.010 0.003 0.003 0.003 0.003 0.016 0.015 0.016 0.016 0.016 0.014
C. schmidianum 0.061 0.058 0.012 0.013 0.017 0.019 0.019 0.017 0.012 0.004 0.004 0.005 0.016 0.016 0.017 0.017 0.017 0.014
C. malabaricum 0.054 0.055 0.010 0.011 0.000 0.013 0.013 0.012 0.010 0.017 0.000 0.004 0.015 0.015 0.016 0.016 0.016 0.014
Indian Fern Journal Volume XXXI (2014)

C. lunulatum 0.054 0.055 0.010 0.011 0.000 0.013 0.013 0.012 0.010 0.017 0.000 0.004 0.015 0.015 0.016 0.016 0.016 0.014
C. indicum 0.062 0.056 0.012 0.013 0.015 0.017 0.017 0.015 0.012 0.019 0.015 0.015 0.016 0.016 0.016 0.016 0.016 0.014
D. bimarginatum 0.125 0.123 0.110 0.110 0.107 0.113 0.111 0.109 0.109 0.114 0.107 0.107 0.110 0.008 0.016 0.016 0.017 0.016
D. exiguum 0.115 0.115 0.101 0.102 0.098 0.103 0.103 0.101 0.101 0.105 0.098 0.098 0.103 0.047 0.017 0.017 0.017 0.014
H. acanthoides 0.120 0.120 0.116 0.117 0.112 0.118 0.115 0.115 0.116 0.120 0.112 0.112 0.115 0.114 0.119 0.001 0.005 0.014
H. denticulatum 0.121 0.121 0.117 0.118 0.113 0.119 0.116 0.116 0.117 0.121 0.113 0.113 0.116 0.114 0.119 0.002 0.005 0.014
H. polyanthos 0.121 0.119 0.112 0.113 0.111 0.115 0.112 0.112 0.112 0.117 0.111 0.111 0.113 0.121 0.120 0.024 0.024 0.013
A. obscurum 0.107 0.105 0.092 0.093 0.098 0.096 0.092 0.092 0.094 0.099 0.098 0.098 0.095 0.114 0.094 0.088 0.089 0.085
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 75

version 5.0 (Tamura et al. 2011) was used. Treatment by distance was performed with
Neighbor Joining method (Saitou & Nei 1987). The evolutionary distances were computed
using the Maximum Composite Likelihood method (MCL method-Tamura et al. 2004) and
were in the units of the number of base substitutions per site.The analysis using Maximum
Likelihood method was based on General Time Reversible model with discrete gamma
distribution of rate differences among sites and proportion of invariable sites.In both NJ
and ML, bootstrap analysis (Felsenstein 1985) with 10000 replicates was considered for the
assessing robustness. Initial tree for the heuristic search was obtained by the option of
BIONJ with MCL distance matrix for ML. A majority rule consensus of the bootstrap
replicates was calculated in Consensus in the PHYLIP 3.69 package (Felsenstein 1989).
RESULTS
LT

The rbcL sequence data included 1205bp for each of 19 taxa and no positional
homology ambiguities were observed. rbcL sequence data of eight species under the genus
Crepidomanes (C. Presl) C. Presl, which were generated in this study, were not reported
previously. ClustalW2 used for multiple sequence alignment calculated the best match for
the selected sequences and lined up the sequences accordingly. The Percent identity score
thus generated by ClustalW2 between the 19 rbcL sequences (1205 bp length) were shown
in the Table 3.The Conserved region analysis of the sequences with DnaSP v5 software
showed that there were three highly conserved regions in the 1205 bp sequence. The
Polymorphic site analysis showed that there were 945 invariable sites (monomorphic sites),
260 variable (polymorphic sites), 188 parsimony informative sites and 72 singleton variable
sites in the 1205 bp length sequence. There were 312 mutations in the 1205 bp region and
all were of the silent type. The haplotype distribution analysis showed that there were only
17 haplotypes out of the 19 sequences analysed. The sequences of Crepidomanes
intramarginale (Hook. & Grev.) Copel., Crepidomanes lunulatum Madhus. & C. A. Hameed
and Crepidomanes malabaricum C. A. Hameed & Madhus. were found to be of the same
haplotypes. The results of the estimation of evolutionary divergence between taxa were
given in the Table 4.
The model test done in the MEGA version 5.0 (Tamura et al. 2011) with Akaike
Information Criterion (AIC) as the model selection criteria, showed that the General Time
Reversible model with discrete gamma distribution of rate differences among sites and
proportion of invariable sites was most suitable for the sequence given.
The optimal tree (the sum of branch length = 0.32995444) showing the relationships
of the 19 taxa inferred from Neighbor Joining method was given in Figure 5. In Maximum
likelihood analysis, the tree with the highest log likelihood (-3936.2865) was shown (Fig.
6) and the tree resulted from the Bayesian analyses was also given (Fig.7).
Regardless of the phylogenetic method used for the analysis, strong support (as
given by the Bootstrap values (BS>95) and Bayesian posterior probabilities as percentage
(PP> 95)) was observed for the four different groups. The clustering pattern of different
taxa showed no difference in the three trees derived.
76 Indian Fern Journal Volume XXXI (2014)

Group-A comprised of the species viz., Hymenophyllum polyanthos, H. acanthoides


and H. denticulatum. In this, H. polyanthos was considered as outgroup for rooting the tree.
Group–B consists of only one species Abrodictyum obscurum and Group-C with two species
viz., Didymoglossum bimarginatum and D. exiguum.
Group-D was the largest group in the tree with thirteen species. Group-D consisted
of a smaller cluster (Group-D1) with two taxa (Crepidomanes proliferum var. proliferum
and C. saxifragoides) and a larger cluster (Group-D2) with eleven species. Both sub groups
of Group-D were with strong support (both BS and PP above 97).
The first split in the Group-D2 led to the cluster consisting of Crepidomanes
bipunctatum and C. christii (PP=100, BS>90). The second split gave comparatively a larger
cluster with very low support (BS<50) both in NJ and ML trees but with PP=63 in the
Bayesian tree. This cluster consisted of C. insigne cluster and the C. intramarginale cluster
and three other species. Most remarkable result was that newly described endemic species,
namely C. lunulatum and C. malabaricum, were nested along with C. intramarginale
without any segregation and with extremely strong support (BS>99, PP=100). The C. insigne
cluster was found to be consisting of three species in which C. bilabiatum and C. latealatum
formed a clade with weak support (BS<70) in ML and NJ tree but with good support in
Bayesian tree (PP=94) and C. insigne appeared sister to this clade.
DISCUSSION

The present analysis has yielded a good clustering pattern that reflects relationships
among the taxa. The two lineages in the family Hymenophyllaceae can be seen as two
separate clusters in tree (Fig.7) as presented by other molecular studies (Dubuisson 1997,
Pryer et al. 2001, Ebihara et al. 2006, 2007, Hennequin et al . 2008). The lineage of
Hymenophylloid ferns is represented by three species (Hymenophyllum polyanthos, H.
acanthoides and H. denticulatum) which are clustered together and denoted in the trees as
Group-A. The Trichomanoid lineage of South India is represented by three well supported
groups (Fig.7) which correspond to the three genera of Ebihara et al. (2006). The genera
are Abrodictyum C. Presl (Group-B), Didymoglossum Desv. (Group-C) and Crepidomanes
(C. Presl) C. Presl (Group-D).
The Group-B corresponds to genus Abrodictyum s.s. Ebhihara et al. (2006) and is
represented by a single species, viz., A. obscurum (Blume) Ebihara & K. Iwats. This is the
only South Indian species under Trichomanoid lineage that is terrestrial. The genus
Abrodictyum C. Presl corresponds to the ‘Pa’ clade (Pachychaetum clade) in Ebihara et al.
(2007), and also to the TE (Terrestrial) clade of Hennequin et al. (2008). The taxa in these
clades (including A. obscurum) have a basal position in the Trichomanoid lineage in the
molecular phylogenetic analysis (Ebihara et al. 2006, 2007, Hennequin et al. 2008). The
basal position of this clade is reasonable as evidenced by the studies conducted by Dubuisson
(1997), Dubuisson et al. (2003b) and Hennequin et al. (2008). The fossil evidence from the
Upper Triassic by Axsmith et al. (2001) suggests the terrestrial habitat (with its associated
morphological characters) as the ancestral state for Trichomanoids. Furthermore, the
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 77

F i gg.. 1 A . Abrodictyum obscurum; B . Crepidomanes bilabiatum, C . C. bipunctatum; D . C. christii;


E. C. indicum; F. C. insigne; G . C. intramarginale
78 Indian Fern Journal Volume XXXI (2014)

ig.. 2 A. Crepidomanes kurzii; B . C. lunulatum; C. C. malabaricum; D . C. proliferum var. proliferum;


F ig
E. C. saxifragoides; F. Didymoglossum bimarginatum; G . C. schmidianum; H. D. exiguum
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 79
80 Indian Fern Journal Volume XXXI (2014)
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae)

Fig.6. Most likely topology (with highest - log likelihood = -3936.2865) resulting from the maximum likelihood analysis
using rbcl. sequences (1205 bp) of 19 taxa
81
82 Indian Fern Journal Volume XXXI (2014)
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 83

divergence of TE clade occurred before the divergence of ‘HE’ clade (chronogram in


Hennequin et al. 2008) and also before the divergence of genera Didymoglossum and
Crepidomanes. This aspect makes it sensible for this terrestrial genus to separate out from
other epiphytic genera in the present study (Figs.5, 6, 7) as in the other previous molecular
studies (Ebihara et al. 2006, 2007, Hennequin et al. 2008). Moreover, the phenetic study
done by Varma et al. (2014) of the taxa under Trichomanoid lineage of South India also
illustrates the basal position of A. obscurum to other Trichomanoid taxa in UPGMA tree.
The genera Didymoglossum and Crepidomanes correspond to the ‘HE’ (Hemi-
epiphytic/Epiphytic) clade of Dubuisson (1997) encompassing taxa that are either epiphytic
or epipetric. The two genera have split out from a single node in the NJ and Bayesian
analysis done here (Figs.5, 7) which is in accordance with the previous molecular studies
(Dubuisson 1997, Pryer et al. 2001, Dubuisson et al. 2003a, Ebihara et al. 2006, 2007,
Hennequin et al. 2008). According to the study related to the evolution of epiphytism and
divergence times in the Hymenophyllaceae done by Hennequin et al. (2008), epiphytism have
evolved during Cretaceous in the family. The beginning of Cretaceous was the time at which
these two genera started their split from their preceding ancestor (chronogram given in
Hennequin et al . 2008). This makes sense with their clubbing up in the phylogenetic
analysis.
The Group-C consists of the two species viz ., Didymoglossum bimarginatum
(Bosch) Ebihara & K. Iwats. and D. exiguum (Bedd.) Copel. They represent the genus
Didymoglossum s.s. of Ebihara et al. (2006) and the ‘Di’ clade (Didymoglossum clade) of
Ebihara et al. (2007). In this, D. exiguum represents subclade ‘Le’ ( Lecanium ) and
D. bimarginatum corresponds to subclade ‘Mg’ (Microgonium) of Ebihara et al. (2007).
Group-D corresponds to the genus Crepidomanes s.s. of Ebihara et al. (2006) and
clade ‘PT’ (Pan-tropical) and subclade ‘Cr’ (Crepidomanes) of Ebihara et al. (2007). The
two prominent clusters (Group-D1 and D2) in this group represent the two sections
described by Ebihara et al . (2006). Group-D1 consists of the taxa under the sect.
Gonocormus s.s. of Ebihara et al. (2006) and Group-D2 consists of the taxa under the sect.
Crepodimanes s.s. of Ebihara et al. (2006).
The two taxa clustered together in the Group-D1 are Crepidomanes proliferum
(Blume) Bostock var. proliferum and C. saxifragoides (C. Presl) P. S. Green. Recent revision
of Hymenophyllaceae done by Ebihara et al. (2006) places all the taxa and several other
obscure names under sect. Gonocormus into one polymorphic species complex viz.,
Crepidomanes minutum (Genus Crepidomanes subg. Crepidomanes sect.Gonocormus) and
advocates further studies on these taxa. The study done by Nitta et al. (2011) reveals that
C. minutum complex is a reticulate network including multiple diploid lineages with
stabilized hybrid crosses. They also advocate further study in the complex for defining the
natural taxa.From the analysis of rbcL sequences done here, the taxa, Crepidomanes
proliferum var. proliferum and C. saxifragoides appear distinct and can be treated as
separate species. The percent identity score (Table 3) between Crepidomanes proliferum var.
84 Indian Fern Journal Volume XXXI (2014)

proliferum and C. saxifragoides is 96. The percent identity score for C. bipunctatum and
C. christii is 99 and these taxa are considered as separate species by the former workers
(Ebihara et al. 2007, Hennequin et al . 2008, Nitta et al. 2011). The estimation of
evolutionary distance between Crepidomanes proliferum var. proliferum and C. saxifragoides
shows that the evolutionary distance between them is 0.040 (Table 4) which is almost
similar to the distance between Didymoglossum bimarginatum and D. exiguum (0.047).
Moreover, Sledge (1968) have definitively shown that Trichomanes saxifragoides C. Presl
(= Crepidomanes saxifragoides (C. Presl) P.S. Green) is distinct from T. proliferum Blume
(= Crepidomanes proliferum (Blume) Bostock). The latter has a more pinnate and less
radiating lamina and frequently bears adventive fronds from the stipes (proliferous) whereas
T. saxifragoides has flabellate lamina and non-proliferous stipes. Haplotype distribution
analysis also shows the distinctiveness of the two taxa viz., Crepidomanes proliferum var.
proliferum and C. saxifragoides.
In addition to this, the status of the C. proilferum (Blume) Bostock var. minutum
(Blume) C. A. Hameed, the variety described by Hameed et al. (2003), has to be clarified.
The taxa described by Hameed et al . (2003) as C. proliferum var. proliferum is
T. proliferum Blume of Manickam and Irudayaraj (1992) whereas, C. proliferum var.
minutum is T. proliferum forma minutum of Manickam and Irudayaraj (1992) which seems
to be T. minutum of Blume (1828). It seems that the above mentioned workers followed
Sledge’s (1968) observation that “Trichomanes minutum Blume and T. proliferum Blume
cannot be regarded as more than individual states of same variable species” and thus treated
T. minutum Blume (=Crepidomanes minutum (Blume) K. Iwats.) as a forma/variety of
T. proliferum Blume (=Crepidomanes proliferum (Blume) Bostock). But the affinity of
C. proliferum var. minutum has been questioned by Varma et al. (2014) as this taxon shows
more affinity towards C. saxifragoides in the phenetic analysis and Hameed et al . (2003)
too agrees with the similarity shown by C. proliferum var. minutum to C. saxifragoides.
The rbcL data of C. proliferum var. minutum taxa is not available and the confusion related
its status has remained unsolved. Further studies (probably cytological, anatomical and data
from other molecular markers) are needed to clarify the relationships between the three taxa
of section Gonocormus of South India.
Group-D2 comprises of eleven species which corresponds to the sect. Crepidomanes
s.s. Ebihara et al. (2006). In this, the cluster of Crepidomanes intramarginale consists of
three taxa viz., Crepidomanes intramarginale (Hook. &Grev.) Copel., C. lunulatum Madhus.
& C. A. Hameed and C. malabaricum C. A. Hameed & Madhus. This clade has good
support (PP=100, BS=99) in the three dendrograms obtained in the analysis. In both
Bayesian and ML tree this group appears sister to other six species whereas in the NJ tree
it is sister to C. kurzii with a very low support (BS=39). In this group, C. lunulatum and
C. malabaricum are newly described species from South India and are endemic to the region
(actually endemic to Kerala and are reported only from the type of locality i.e., from the
Athirapally waterfalls, Thrissur Dt.). C. intramarginale is the type species of the genus
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 85

Crepidomanes and is confined to South India and Srilanka (Iwatsuki 1985, Hameed et al.
2003). The clustering pattern obtained from the analysis of rbcL gene pose doubts on the
species identity of the two newly described taxa which are endemic to South India. The
estimation of evolutionary divergence shows that the distance between Crepidomanes
intramarginale , C. lunulatum and C. malabaricum is 0.00 (Table 4) i.e ., there is no
divergence between the taxa when rbcL sequences is considered. The percent identity score
(Table 3) of the three taxa are 100 which also supports the clustering pattern. The haplotype
distribution analysis also shows that Crepidomanes intramarginale, C. lunulatum and C.
malabaricum consist of one and the same haplotype and are not distinct.
Varma et al. (2014) has reported 75% morphological similarity between C.
intramarginale and C. malabaricum. Hameed et al. (2003) have also commented on the
resemblances of C. malabaricum with C. intramarginale. According to them, C. malabaricum
is larger, clearly pinnatifid or pinnatisect, unistratose fronds with close and slightly
overlapping segments, with wavy margin, campanulate indusia with abruptly dilated mouth
whereas C. intramarginale has bistratose, subpinnatifid fronds, and winged rachis with
bivalvate indusia. In this, more striking character is the unistratose lamina of C.
malabaricum which can be distinguishable from the bistratose lamina of C. intramarginale.
Fraser-Jenkins (2008a, b) considers C. malabaricum as “probable synonym of C. kurzii”,
whereas the study done by Varma et al. (2014) shows 70% similarity between C. kurzii and
C. malabaricum.
The clustering of C. lunulatum with C. malabaricum and C. intramarginale is a
surprising one. Earlier workers have different opinions about the affinity of C. lunulatum.
Hameed et al. (2003) have commented that C. lunulatum shows resemblance with C.
agasthianum Madhus. & C. A. Hameed, another newly described endemic species of South
India. But, Chandra et al. (2008) and Fraser-Jenkins (2008a, b) consider C. lunulatum as
the “probable synonym of C. agasthianum”. The study done by Varma et al. (2014), C.
lunulatum has 75% morphological similarity with C. malabaricum and C. kurzii whereas it
shows only 60% morphological similarity with C. agasthianum. The estimate of evolutionary
divergence by rbcL sequences shows that distance between C. kurzii and C. lunulatum is
0.013 (Table 4) and the percent identity score between them is 98 (Table 3) and both values
are similar to that of C. kurzii and C. malabaricum. As the rbcL sequence of C. agasthianum
is not included in the current analysis the affinity of C. lunulatum with this species cannot
be commented here. To deduce the exact identity of the C. lunulatum and C. malabaricum
further studies using other molecular markers that show relatively more variation than rbcL
is needed (e.g.: matk). In addition to this, evidences from cytology and anatomy may also
be useful. A combined phylogenetic analysis using molecular data and non-molecular data
is advisable along with additional sequences from other taxa such as C. agasthianum in order
to clarify the doubts regarding the status of these two newly described taxa.
In the Group-- D2, another important cluster is the C. insigne cluster. This cluster
consists of three species which are considered to be closely related (Iwatsuki 1985, Hameed
et al. 2003). Here, Crepidomanes insigne appears as sister to a cluster with C. bilabiatum
86 Indian Fern Journal Volume XXXI (2014)

and C. latealatum. Iwatsuki (1985) considers C. insigne as conspecific to C. latealatum along


with C. plicatum. Manickam and Irudayaraj (1992) and Fraser-Jenkins (2008a) consider C.
insigne as synonym of C. latealatum. But in the description, Fraser-Jenkins (2008a) states
that Trichomanes (Crepidomanes) insigne are plants with a single submarginal false veinlet.
Iwatsuki (1985) clearly states that the taxa under Crepidomanes latealatum complex
(including Crepidomanes latealatum, C. insigne and C. plicatum) are without submarginal
false veinlet, but with oblique striae and the plants with submarginal veinlet are those under
C. bipunctatum. So it seems that specimens studied by Fraser-Jenkins (2008a) do not belong
to Trichomanes (Crepidomanes) latealatum complex.
As reported by Hameed et al. (2003) and Iwatsuki (1985), C. insigne have no
submarginal false veinlet but with oblique striae.In the present study, C. insigne is not
clustered together with C. latealatum (Figs. 5, 6, 7) but placed as sister to C. bilabiatum-
C. latealatum cluster. The percent identity score (Table 3), estimates of evolutionary
divergence (Table 4) and the haplotype distribution analysis support the distinctiveness of
C. insigne from other related taxa considered in this study.
Crepidomanes indicum C. A. Hameed & Madhus. is a newly described species from
South India which is endemic to the region. The status and affinity of this endemic species
is a matter of dispute. Hameed et al. (2003) has reported that C. indicum shows
resemblances with C. kurzii and C. agasthianum. The study done by Varma et al. (2014)
has showed that C. indicum is more closely allied to C. agasthianum than C. kurzii. Fraser-
Jenkins (2008a, b) considers C. indicum as “probable synonym of C. kurzii”. In the
Bayesian tree (Fig.7), C. indicum clusters with C. insigne group along with C. schmidianum
and this cluster has good support too (PP=94). But in the ML tree (Fig.6) C. indicum is
placed above to C. kurzii. But the estimate of evolutionary divergence (Table 4) shows that
C. indicum appears to be more close to C. bilabiatum and C. insigne than C. kurzii.
In the present study, the C. indicum appears as a distinct species as revealed by the
haplotype distribution analysis and also by its positioning in the three trees obtained. It
seems that additional rbcL data from taxa such as C. agasthianum will be necessary to
clarify the status and affinities of C. indicum.
Further studies on South Indian Trichomanoid lineage are needed to clarify
ambiguities observed in the present study. Additional rbcL sequences data from species such
as Crepidomanes agasthianum Madhus. & C. A. Hameed, which has not been included in the
present study, is needed to obtain more clarity in the relationships of species such as C.
indicum C. A. Hameed & Madhus. and C. lunulatum Madhus. & C. A. Hameed. A combined
phylogenetic analysis using both molecular and non-molecular data (morphology, cytology,
anatomy etc.) will be advisable to get clearer picture about the relationships and status of the
above mentioned problematic taxa. Preferably, a rbcL sequence based phylogenetic study of
Trichomanoid lineage with a pantropical range will be more useful to discuss the taxonomical
position of every species and to propose historical bio-geographical hypotheses for Indian taxa.
Taxonomy is a ‘never ending synthesis’. Being bereft of own data, Taxonomy has
to depend on other disciplines of biology; as and when new disciplines arise Taxonomy has
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 87

to undergo vicissitudes redefining the phylogenetic aspects and lineages restructuring the
groups and their affinities. Earlier, taxonomists used to depend only the visible
morphological characters for classification and elucidation of evolutionary relationships.
Later, palynological, anatomical, cytological, phytochemical characters, etc. were utilized.
The emergence of Molecular Systematics in which the molecular sequences are considered
to be the most dependable character in the determination of species circumscription and
phylogenetic relationships has overtopped the present evidences used in taxonomy.
There are several advantages in using rbcL for systematic analysis. It is situated in
the single copy region of the chloroplast genome which is 1.4bp in length and has fairly
conservative rate of evolution. The gene is universal among all photosynthetic plants which
makes it a good phylogenetic tool. Introns are absent and positional coding of information
permits the unambiguous alignment of rbcL which is advantageous for among phylogenetic
analysis. The CBOL (Consortium for the Barcode of Life) has approved the two plastid
loci rbcL and matK as the official DNA barcode for all land plants including ferns (Li et
al., 2011) thus highlighting the utility of rbcL phylogenetic interpretations.
The protein coding genes in cpDNA are conserved ones and most exploited in
molecular studies. The genes that code for subunit ATP Synthase (atpB), enzyme maturase
(matK) subunit F of NADP Hydrogenase (ndhF) and ribosomal proteins (rpS2 and rpS4)
are often employed in phylogenetic studies. The atpB genes are without any introns and
found to have a rate of evolution as that of rbcL gene (Hoot et al., 1995) whereas matK
evolves relatively as much at rapid rate than rbcL (Soltis & Soltis, 1998). Now matK along
with rbcL are considered to be the potential barcoding markers in the land plants (CBOL
Plant Working Group, 2009).
The present data on nucleotide sequences evidently help to draw the phylogenetic
relationships of Trichomanoid species of South India. This is partly in agreement with the
results of the phenetic analysis done with morphological features of trichomanoid ferns as
three distinct groups (Varma et al., 2014). While morphological features show distinction
of the newly reported species viz., Crepidomanes lunulatum and Crepidomanes malabaricum,
molecular data categorically deny the validity of these species.
Even though the rbcL nucleotide sequences are almost reliable in determining the
identity at species and subspecific level, the process is cumbersome and time consuming
deterring field taxonomists from its applications and to think that Molecular Systematics is
useful only in the understanding of the phylogenetic relationships and not for primary
taxonomic procedures (description, nomenclature, identification and classification) or it can
be applied only to demystify the ‘taxonomically problematic taxa’
ACKNOWLEDGEMENTS

The first author is thankful to University Grants Commission, Govt. of India for
sanctioning Minor Research Project which provided the financial help for this study.The
authors acknowledge the technical support given by DBT, Distributed Information Sub Centre
(IISR) for the computational analysis of data. Many thanks are also addressed to Mr. Suresh
88 Indian Fern Journal Volume XXXI (2014)

Kumar U., DNA Examiner, RGCB, Thiruvananthapuram for his helps and suggestions during
the molecular study and also for providing the sequence of primer, RBCF. The authors also
thank Dr. Santhosh J. Eapen, Principal Scientist & Co-ordinator (Bioinformatics), and Ms.
Rosana O.B., Senior Research Fellow, DISC, IISR, Calicut for their help and suggestions during
phylogenetic analysis. We are also thankful to Mr. Anoop K.P. for his help in photoshop.
REFERENCES

A X S M I T H B J , K R I N G S M & TAY L O R T N 2 0 0 1 A f i l m y f e r n f r o m t h e U p p e r Tr i a s s i c o f N o r t h
Car olina (USA) Amer J Bot 8 88: 1558-1567
BLUME C L 1828 En umer atio Plantar um Javae Van Leeuw en Lugduni Bat avorum (Leiden)
CHANDRA S, FRASER-JENKINS C R, KUMARI A & SRIVASTAVA A 2008 A summary of the status
of thr ea tened pteridophytes of India Taiwania 5 3 : 170-209
C B O L P l a n t Wo r k i n g G r o u p 2 0 0 9 A D NA b a r c o d e f o r l a n d p l a n t s P r o c N a t l A c a d S c i 1 0 6 :
12794-12797
d’ALMEIDA J F R 1926 The ferns of the High wavy mountain J Indian Bot Soc 5 : 20-30
D OYLE J J& DOYLE J L 1987 A rapid DNA isolation pr ocedure fr om small quantities of fresh leaf
tissue Phytochem Bull 1 9 : 11-15
D RU M M O N D A J A S H TO N B C H E U N G M H E L E D J K E A R S E M M O I R R S TO N E S - H AVA S S
THIERER T & WILSON A 2012 Geneious v5 created by Biomattershttp://www.geneious.com
D U B U I S S O N J - Y 1 9 9 7 r b c L s e q u e n c e s : A p r o m i s i n g t o o l f o r t h e m o l e c u l a r s y s t e m a t i c s o f t h e f er n
genus Trichomanes (Hymenophyllaceae) Mol Phylogenet Evol 8 : 128-137
D U B U I S S O N J - Y, H E N N E QU I N S , D O U Z E RY D J P, C R A N F I L L R B , S M I T H A R & P RY E R K M
2003a rbcL phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special
reference to Neotropical taxa Int J Plant Sci 1 6 4 : 753-761
D U B U I S S O N J - Y, H E N N E QU I N S , R A KOTO N D R A I N I B E S & S C H N E I D E R H 2 0 0 3 b E c o l o g i c a l
diversity and adaptive tendencies in the tropical fern Trichomanes L (Hymenophyllaceae)
with special reference to climbing and epiphytic habits Bot J Linn Soc 1 4 2 : 41-63
EBIHARA A, DUBUISSON J-Y, IWATSUKI K HENNEQUIN S & ITO M 2006 A taxonomic revision of
Hymenophyllaceae Blumea 5 1 : 221-280
E B I H A R A A , I WAT S U K I K , I TO M , H E N N E QU I N S & D U B U I S S O N J - Y 2 0 0 7 A g l o b a l m o l e c u l a r
phylogeny of the fern genus Trichomanes (Hymenophyllaceae) with special reference to stem
anatomy Bot J Linn Soc 1 5 5 : 1-27
FELSENSTEIN J 1985 Confidence limits on phylo g enies : An a pproach using the bootstrap Evolution
3 9 : 783-791
FELSENSTEIN J 1989 PHYLIP-Phylogeny Inference Package Cladistics 5 : 164-166
FRASER-JENKINS C R 2008a Ta x o n o m i c r ev i s i o n o f t h r e e h u n d r e d I n d i a n s u b - c o n t i n e n t a l
Pteridophytes with a revised census-list: a new picture of fern-taxonomy and nomenclature in
the Indian subcontinent Bishen Singh Mahendra Pal Singh Dehra Dun
FRASER-JENKINS C R 2008b Endemics and pseudo-endemics in relation to the distribution patterns
of Indian pter idoph ytes Taiwania 5 3 : 264-292
HALL T A 1999 BioEdit : a user friendly biological sequence editor and analysis programme for
Windows 95/98/NT Nucleic Acids Sym Ser 4 1 : 95-98
H A M E E D C A & M A D H U S O O DA NA N P V 1 9 9 8 A n ew s p e c i e s o f f i l m y f e r n ( H y m e n o p hy l l a c e a e :
Pteridophyta) from South India Sida 1 8 : 519-522
H A M E E D C A & M A D H U S O O DA NA N P V 1 9 9 9 C r e p i d o m a n e s m a l a b a r i c u m , a n ew f i l m y f e r n
species (Hymenophyllaceae – Pteridophyta) from Southern India Nord J Bot 1 9 : 277-279
H A M E E D C A & M A D H U S O O DA NA N P V 2 0 0 3 Tri c h o m a n e s va m a n a , a n ew f i l m y f e r n s p e c i e s
(Hymenophyllaceae : Pteridophyta) from India Nord J Bot 2 3 : 437-440
H A M E E D C A , R A J E S H K P & M A D H U S O O DA NA N P V 2 0 0 3 F i l m y f e r n s o f S o u t h I n d i a P e n t a
P K Vidya Varma & P V Madusoodanan : Molecular Systematics of the Filmy Ferns (Hymenophyllaceae) 89

Book Publishers and Distributors Calicut


H E N N E QU I N S S C H U E T T P E L Z E P RY E R K M E B I H A R A A & D U BU I S S O N J - Y 2 0 0 8 D ive rg e n c e
times and the evolution of epiphytism in filmy ferns (Hymenophyllaceae) revisited Int J Plant
Sci 1 6 9 : 1278-1287
H O OT S B , C U L H A M A & C R A N E P R 1 9 9 5 T h e u t i l i t y o f a t p B g e n e s e q u e n c e s i n r e s o l v i n g
phylogenetic relationships Comparison with rbcL and 18S ribosomal DNA sequences in the
Lardiza balaceae Ann Missouri Bot Gard 8 2 : 194-208.
HUELSENBECK J P & RONQUIST F 2001 MRBAYES : Bayesian inference of phylogeny Bioinformatics
1 7 : 754-755
I WAT S U K I K 1 9 8 5 T h e H y m e n o p h y l l a c e a e o f A s i a , e x c l u d i n g M a l e s i a J Fa c S c i U n iv To k y o 1 3 :
501-551
I WATSUKI K 1990 Hymenophyllaceae In The families and gener a of vascular plants - Pteridoph ytes
and Gymnosperms Ed . K Kubitzki Spring er Ver lag Berlin Vol 1 Pp 157-163
L A R K I N M A , B L AC K S H I E L D S G , B ROW N N P, C H E N NA R , M c G E T T I G A N P, M c W I L L I A M H ,
VALENTIN F, WALLACE I M, WILM ALOPEZ R, THOMPSON J D, GIBSON T J & HIGGINS
D G 2007 ClustalW and ClustalX version 2.0. Bioinformatics 2 3 : 2947-2948
LI F W KUO L Y ROTHFELS C J EBIHARA A CHIOU W L WINDHAM M D AND PRYER K M 2011
rbcL and matK earn two thumbs up as the core DNA barcode for ferns PLoS ONE 6 :
e 26597(1-7)
L I B R A D O P & RO Z A S J 2 0 0 9 D n a S P v 5 : A s o f t wa r e f o r c o m p r e h e n s i v e a n a l y s i s o f D NA
polymorphism data Bioinformatics 2 5 : 1 4 5 1 - 1 4 5 2
MADHUSOODANAN P V & HAMEED C A 1998 Cr epidomanesagasthianum , a new filmy fer n species
(Hymenophyllaceae) from India Nord J Bot 1 8 : 169-170
M A D H U S O O DA NA N P V & H A M E E D C A 1 9 9 9 C r e p i d o m a n e s l u n u l a t u m ( H y m e n o p h y l l a c e a e :
Pteridophyta) a new filmy fern species from South India Fern Gaz 1 5 : 265-268
MANICKAM V S & IRUDAYARAJ V 1992 Pteridophyte flora of the Western Ghats - South India B I
Publishers Pvt Ltd New Delhi
N I T TA J H E B I H A R A A & I TO M 2 0 1 1 R e t i c u l a t e ev o l u t i o n i n t h e C r e p i d o m a n e s m i n u t u m s p e c i e s
complex (Hymenophyllaceae) Amer J Bot 9 8 : 1782–1800
P RY E R K M, S M I T H A R , H U N T J S & D U BU I S S O N J - Y 2 0 0 1 rbcL da ta reveal two monoph yletic
g roups of film y ferns (Filicopsida : Hymenophyllaceae) Amer J Bot 8 8 : 1118-1130
RAMBAUT A & DRUMMOND A J 2007 Tracer v1.5. http://beast.bio.ed .ac.uk/Tracer
RO N QU I S T F & H U E L S E N B E C K J P 2 0 0 3 M R BAY E S 3 . 1 : B a y e s i a n p h y l o g e n e t i c i n f e r e n c e u n d e r
mixed models Bioinformatics 1 9 : 1572-1574
S A I TO U N & N E I M 1 9 8 7 T h e n e i g h b o r - j o i n i n g m e t h o d : A n e w m e t h o d f o r r e c o n s t r u c t i n g
phylogenetic trees Mol Biol Evol 4 : 406-425
SLEDGE W A 1968 The Hymenophyllaceae of Ceylon Bot J Linn Soc 6 0 : 289-308
SOLTIS D E & SOLTIS P S 1998 Choosing an approptrpiate gene foy Molecular systematics in plants
I I D NA s e q u e n c i n g E d s . D E S O LY I S , P S S O LT I S A N D J J D OY L E K l u wa r A c a d e m i c
Publishers USA Pp 01-42
TAMURA K , NEI M & KUMAR S 2004 Pr ospects f or infer ring ver y large phylogenies b y using the
neighbor-joining method Pr oc Natl Acad Sci USA 1 0 1 : 11030-11035
TA M U R A K , P E T E R S O N D , P E T E R S O N N , S T E C H E R G , N E I M & K U M A R S 2 0 1 1 M E G A 5 :
Molecular Evolutionary Genetics Anal ysis using Maximum Likelihood, Evolutionary Distance ,
and Maximum Parsimony Methods Mol Biol Evol 2 8 : 2731-2739
WAT E R H O U S E A M , P RO C T E R J B , M A RT I N D M A , C L A M P M & BA RTO N G J 2 0 0 9 “ J a l v i ew
Ver sion 2-a m ultiple sequence alignment editor and analysis wor kbench” Bioinforma tics 2 5 :
1189-1191
VA R M A V P K , M O H A NA N K V & M A D H U S O O DA NA N P V 2 0 1 4 P h e n e t i c s o f S o u t h I n d i a n
Trichomanoid Ferns Indian Fern J 3 1 : 53- 68
Indian Fern J. 3 1 : 90-101 (2014) ISSN 0970-2741

MICR
ROOSTR UCTURAL OBSER
RUCTURAL VATIONS ON THE DEVELOPMENT OF
OBSERV
GAMET O P H Y T E S AND OOGENESIS IN E Q
TO UISETUM A R
QUISETUM VENSE
RV
GE WANG, XI-LING DAI, QUAN-XI WANG AND JIAN-GUO CAO*
College of Life and Environment Sciences, Shanghai Normal University, Shanghai 200234 P.R. China

(Received May 2, 2014; Revised Accepted July 8, 2014)

ABSTRACT
The development of gametophytes and oogenesis of the fern Equisetum arvense
L. were studied using light microscopy. The features of the gametophyte development of
E. arvense are usual for the genus as described previously. Archegonia are formed at the base
of branches or on the top of the apical meristem of the female gametophyte. Section
observations show that the archegonium of E. arvense develops from a superficial cell (initial
cell). The initial cell divides periclinally to form an outer and an inner cell. The outer cell
finally develops into the neck of the archegonium. The inner cell directly becomes the primary
cell, which forms a neck canal mother cell and a central cell by an asymmetrical division. Then
the former forms two oblique arranged neck canal cells and the latter forms an egg and a ventral
canal cell by another asymmetrical division. However, an additional ventral canal cell can be
seen in the section occasionally. During maturation, a prominent feature of the archegonium is
elongation of the terminal cells of the neck. The canal cells degenerate and produce a large
amount of mucilaginous materials. The egg accumulates conspicuous starch grains in the
plastids. No separation cavity and egg envelope can be identified as the egg matures. The
present investigation shows that the archegonial development in the horsetails differs from the
filicopsida.
K e y W o r d s : Horsetails; Equisetum arvense ; Oogenesis; Gametophyte
development.

INTRODUCTION

Various aspects of gametophytes of Equisetum (horsetail), including gametophyte


development, morphology, structure of the gametangia, sexual expression and
spermatogenesis have been studied for more than a century (Sharp, 1912; Hauke, 1967,
1968, 1969, 1971, 1977, 1979, 1980; Duckett, 1970, 1972, 1973a,b, 1977, 1979a,b; 1985;
Jones & Hook, 1970; Duckett & Bell, 1977; Duckett & Duckett, 1980; Liu et al., 1994;
Matzke, 1941; Mesler & Lu, 1977; Renzaglia et al., 2000, 2002). From these studies, we
know that the features of gametophyte and gametangia of Equisetum differ greatly from
those of the Filicopsida. But the modern molecular data show that the extant horsetails
(Equisetum) form a monophyletic group, which is included within the ferns (Pryer et al.
2001, 2004; Des Marais et al. 2003; Smith et al. 2006). Renzaglia et al (2000, 2002) also
indicated that the spermatozoids of Equisetum share several critical features with other ferns
and support monophyly of a fern–Equisetum assemblage. However, features of oogenesis of
Equisetum are still lacking. The features of oogenesis are considered to be more appropriate
to reflect the phylogenetic relationships in contrast to the relatively conserved

* E-mail : caojianguo101@163.com; cao101@shnu.edu.cn


G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 91

spermatogenesis (Cao et al, 2012). The present investigation provides detailed description
of the gametophyte development, archegonial formation and oogenesis of the
microstructural level. The evolutionary relationship reflected by archegonial development
and oogenesis is discussed.
MATERIALS AND METHODS
ATERIALS

Spores of Equisetum arvense were collected from plants collected in Tonghe


County, Heilongjiang Province, P. R. China. The spores, rinsed three times with sterilized
water, were sown on modified Knop’s solution, which is solidified with 1.5% agar in
culture dishes (Cao et al., 2012). These dishes were placed in an artificial climate chamber
under conditions of 25°C in the light (18 h) and 20°C in the dark (6 h). The materials at
various development stages were checked and photographed using a light microscope (Nikon
E-800) and photographed with DXM1200 digital camera.
About 2 months later, archegonia developed on the base of the branches of the
gametophytes. The archegonial gametophytes were checked with a stereomicroscope to
collect the gametophytes with various developmental stages of archegonia. The specimens
were fixed in 3% glutaraldehyde in 0.1 mol/L phosphate buffer (pH 7.0) at room
temperature for 6 to 12 hrs. Subsequently the specimens were washed three times with the
same buffer, postfixed in 2% aqueous osmium tetroxide for 2 h, rinsed three times in the
buffer, and embedded in Spurr’s resin (SPI-Chem, USA) via a graded acetone series. The
specimens were thick-sectioned for presence of the archegonia. All the sections were stained
with Toluidine blue O; examined using a light microscope (Nikon E-800) and photographed
with DXM1200 digital camera.
RESULTS
LT

1. Development of gametophyte
The spores of E. arvense are spherical, chlorophyllous, thin-walled, alete, with
diameter about 35 μm when the spores are released from the sporangium. Each spore has
two elaters, which are attached to the outer spore wall and form four strap-like arms with
spoon-shaped tips (Fig. 1a). The diameter of spore after imbibition increases to 55-60 μm
within several hours after spore sowing. And the elaters are usually shed (Fig. 1b). Under
present culture condition, the spores germinate readily within 24 hours by dividing
eccentrically to produce a small rhizoid cell and a large green prothallus cell (Fig. 1c). The
unicellular rhizoid contains some green chloroplasts. The green prothallial cell usually
undergoes two perpendicular divisions to form four prothallial cells within 2-3 days (Fig.
1d). Certainly, oblique divisions are seen sometimes (Fig. 1e). Further division of the
prothallial cells produces multicellular branched gametophytes in about a week (Fig. 1f).
The gametophytes are basically unisexual. The male and female gametophytes possess no
obvious difference at early stage. But with further development they tend to become
dimorphic. The male gametophytes are usually small and possess some short and thick
92 Indian Fern Journal Volume XXXI (2014)

branches, on top of which form the antheridia in 7-8 weeks (Fig. 1g). The female
gametophytes, usually larger than the male, develop into numerous thin branches (Fig. 1h).
Archegonia firstly develop at the base of the branches in 9-10 weeks. The neck of
archegonium consist of three tiers of four neck cells each (Fig. 1i). The female
gametophytes can grow continuously and form a thick midrib. The apical meristem of the
midrib also can produce archegonia (Fig. 1j). As the archegonium nearly matures, the
terminal four neck cells elongate to almost three to four times longer than broad and spread
apart in an arching manner (Fig. 1k). Several archegonia in the same gametophyte can be
fertilized and form viable sporophytes (Fig. 1l).
2. Ar
Arcc he
hegg onium ffor
or ma
orma tion and oo
mation oogg enesis
The details of the archegonia formation and oogenesis of Equisetum are scarcely
known. Here, the development of the archegonium and oogenesis, from the initial cell of
the archegonium to the mature egg is described with microscopic observations on semi-thin
sections.
2.1. The initial cell
Archegonium of the fern Equisetum arvense develops from a superficial cell
(the archegonial initial cell, IC) on the base of the branches of the female gametophyte
(Fig. 2a) or develops from the apical meristem of the midrib of the old gametophytes (Fig.
2b). The sections show that the initial cell can be identified by possessing dense cytoplasm,
relatively smaller vacuoles and plastids in contrast to the somatic cells. The initial cell
usually shows a cylindrical shape (Fig. 2a), or sometimes shows a triangular shape in
a longitudinal section (Fig. 2b). The nucleus is usually centrally placed in the initial cell
(Fig. 2a-b).
2.2. The primary cell
The initial cell divides periclinally to form two cells (Fig. 2b). The outer cell is
the neck jacket mother cell (Njm) (Fig. 2b, c), which finally form the neck jacket cells of
the archegonium by 2 vertical (Fig. 2e-f) and 2-4 periclinal divisions (Fig. 2g-i). The inner
cell contains several irregular nucleoli within the nucleus. The inner cell does not divide,
but directly becomes the primary cell (PC) (Fig. 2b).
2.3. The central cell
The primary cell usually divides asymmetrically to form two cells before the neck
jacket mother cell divides (Fig. 2d). The cell towards the archegonial neck is the neck canal
mother cell (Ncm), which contains relatively less cytoplasm (Fig. 2d). The lower cell,
getting more cytoplasm from its mother cell, is called the central cell (CC) (Fig. 2d). Soon
after, the neck jacket mother cell divides anticlinally and forms four cells of the neck jacket
of the archegonium (Fig. 2e). Then, large amount of vacuoles occur in the central cell
cytoplasm (Fig. 2f). At this stage, the neck canal mother cell looks like a flat disk with a
height of about 6-7 μm (Fig. 2 e-f).
G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 93

F i g . 1 - G e m e t o p h y t e d e v e l o p m e n t o f E q u i s e t u m a r v e n s e (aa , i , j , l , with a stereomicroscope;


The others, with LM) a . Spore with two elaters (EL); b . A new spore with spiral elaters (EL) c . Within
2 4 h , S p o r e g e r m i n a t i o n , p r o d u c i n g r h i z o i d ( R ) d . A b o u t 2 - 3 d, 2 - 4 c e l l e d g re e n t h a l l i a r e for m e d
e . Seve ra l - c e l l e d g r e e n t h a l l i a re f o r m e d f . Yo u n g b r a n c h e d g a m e t o p hy t e s a r e f o r m e d a b o u t a we e k
g . Male gametophyte with antheridia (An) h . Female gametophyte i . Archegonium is formed at the base
of the br anches of the f emale gametophyte j . Ar ch eg onia occur in the a pical mer istem of the female
gametophyte k . Matured archegonium with four elongated terminal cells (TC); l . Young sporophytes
94 Indian Fern Journal Volume XXXI (2014)

Fig. 2 - Early stage of archegonium formation and oogenesis of Equisetum arvense (with LM)
a , bb.. Archegonial initial cell (IC); c . The initial cell divides into a neck jacket mother cell (Njm) and a
primary cell (PC) d . The primary cell divides to form a neck canal mother cell (Ncm) and a central cell
( C C ) e . The neck jacket mother cell forms four neck jacket cells (Njc) f Large amount vesicles occur in
the central cell (CC); g . A ventral canal cell ( V C C C) and a young egg (E) are formed Two tiers of neck
jacket cells (Njc) form Arrows indicate the starch grains in the egg and the canal cells h , i . Two neck
canal cells (NCC) are formed Three tiers of neck jacket cells (Njc) are formed N, nucleus; P, plastid
G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 95

F i g . 3 - L aatt e s t a g e o f a r c h eeg
g o n i u m ffo
o rrm
m a t i o n a n d o o g e n e s i s (with LM) aa.. Two ventral canal
cells (VCC) can be formed occasionall y; b . The canal cells (NCC, VCC) begin to deg enerate The neck
jacket cells (Njc) begin to elongate The terminal cell (TC) elongates markedly; c . The terminal cell
(TC) elongates and shows dumbbell-shape; d . Mucilageous materials (Mu) appear; e . Matured egg, the
contents are expelled out of the archegonium and a cavity (C) forms above the egg E egg; N,
nucleus; P, plastid
96 Indian Fern Journal Volume XXXI (2014)

2.4. The young egg


The central cell also divides asymmetrically to produce a small ventral canal cell
(VCC) and a young egg cell (E), which gets more cytoplasm from the central cell (Fig.
2g). The plastids are easily identified in the egg and canal cells by presence of starch grains
at this stage (Fig. 2g, arrows). In previous stages, no starch grains are seen in the plastids
(Fig. 2a-f). The vesicles are abundant in the egg cytoplasm (Fig. 2g). As the egg is just
formed, the neck canal mother cell is still mononuclear. But, its volume increases obviously
and bulges upwards (Fig. 2g). At this time, the two tiers of eight neck jacket cells are
formed by a periclinal division (Fig. 2g).
Thereafter, the mononuclear neck canal mother cell forms two neck canal cells by
an oblique division (Fig. 2h). The volume of the two canal cells increase markedly, but
one of them is larger than another (Fig. 2h-i). The lower neck canal cell is usually broad
and short. But the upper neck canal cell is obviously elongated longitudinally (Fig. 2h-i).
The jacket cells undergo another periclinal division and form three tiers of jacket cells (Fig.
2i). Although one ventral canal cell is the rule during oogenesis of Equisetum arvense, two
ventral canal cells can be seen occasionally (Fig. 3a). However, the neck canal cells are
always two without exception.
2.5. Maturation of the archegonium and the egg
The maturation of the archegonium and the egg includes elongation of the neck,
degeneration of the canal cells and development of the egg. The first visible feature of the
archegonium during maturation is the rapid elongation of the neck of the archegonium.
Although the three tiers of the neck jacket cells has been formed previously, each of them
is short in height and contain dense cytoplasm and large nucleus (Fig. 2h-i; Fig. 3a). As
the archegonium matures, the length of each jacket cell increases conspicuously (Fig. 3b).
Especially, the four terminal neck cells of the archegonium elongate rapidly, and finally
reach almost three to four times longer than broad. Each terminal neck cell of the
archegonium has a dumbbell-shape (Fig. 3c). Dense cytoplasm of the jacket cells fades away
and large vacuoles have been formed in the jacket cells (Fig. 3b-d). At the last stage of the
maturation, the lowest cells of the neck divide and totally form 4-5 tiers of neck jacket
cells (Fig. 3d-e). Sometimes, longitudinal divisions of the neck jacket cells can be seen in
the section (Fig. 3d).
The second feature of the archegonium during maturation is degeneration of the
canal cells. The original well defined cell wall between canal cells (Fig. 3a) now becomes
obscure (Fig. 3b-d). The outline of the canal cells becomes irregular (Fig. 3b-d). The starch
grains, which can be seen in the young stage, diminish gradually during maturation.
Simultaneously, mucilaginous materials outside the canal cells increase (Fig. 3b-d). The
upper neck canal cell grows with elongation of the archegonium neck and the cytoplasm of
G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 97

the upper neck canal cell pushes into the neck cavity of the archegonium (Fig. 3c-d).
At the microscopic level, maturation of the egg seems to be a process of
accumulating starch grains. The amyloplasts become more and more prominent (Fig. 3c-e).
The vesicles, recognizable in the young egg stage, diminish gradually in the egg cytoplasm
(Fig. 3a-e). In the last stage of the archegonial maturation, the terminal four neck cells
open (Fig. 1k) and the contents, including the degenerated canal cells and their secretion
materials, are expelled out of the archegonium (Fig. 3e). An egg, with relatively dense
cytoplasm, lies at the bottom of the cavity of the archegonium (Fig. 3e).
DISCUSSION

Although many works about the gametophyte development of the genus Equisetum
have been done (Duckett, 1973a, 1979b; Hauke, 1968, 1969, 1977, 1979, 1980; Liu et
al., 1994; Matzke, 1941; Mesler & Lu, 1977; Walker, 1921, 1931, 1937), few of them
involve the complete and development of the archegonium and oogenesis. The present
investigation provides detailed microstructural observations of archegonium development and
oogenesis.
1. Features of the early development of the archegonium
Although horsetails (Equisetum) are considered to be a member of ferns according
to modern molecular data (Smith et al., 2006), the early development of the archegonium
of Equisetum reflects some obvious differences of horsetail from the ferns (filicopsida).
The archegonial initial of Equisetum only divides once and forms two cells. But the
archegonial initial cell of filicopsida always undergoes two periclinal divisions to form
three cells, i.e. the upper cell, the middle cell (primary cell) and basal cell (Yang et al.,
2009; Huang et al., 2011; Cao et al., 2010a, 2011a). In this point, Campbell (1895) also
indicated that the initial of archegonia in Equisetum divides periclinally to form an outer
and an inner cell. The inner cell divides to form neck canal cells, a ventral canal cell,
and an egg. However, the detailed formation processes of archegonium and oogenesis
were lacking. Here, we clarify the details of the cell divisions and cytological features of
the cells taking part in oogenesis. The present observations show that the inner cell of
Equisetum directly becomes the primary cell, which divides before divisions of the neck
jacket mother cell. But in filicopsida, the primary cell divides after divisions of the neck
jacket mother cell (Yang et al., 2009; Huang et al ., 2011; Cao et al., 2010a, 2011a). In
addition, the canal cells in Equisetum are discus-shaped and have the same transverse
diameter with their mother cell. The transverse diameter of the canal cells in the
filicopsida are smaller than their mother cell (Yang et al., 2009; Huang et al., 2011; Cao
et al., 2010a, 2011a).
2. Features of the canal cells
The present investigation shows that two neck canal cells are formed during
98 Indian Fern Journal Volume XXXI (2014)

oogenesis in Equisetum arvense. The number of the neck canal cells was not determined
previously. It was reported that the archegonium of Equisetum laevigatum has one neck
canal cell in many cases, but only a few archegonia give rise to two boot-shaped neck canal
cells by a vertical division (Walker, 1937). Phatak (1935) reported that four superimposed
neck canal cells were seen in Equisetum (see literature of Walker, 1937). However, in
present investigation, two neck canal cells are constant without variation. But, the number
of ventral canal cells seems to be variable in some extent. In most cases, there is only one
ventral canal cell in the archegonium, but two ventral canal cells can be seen occasionally
(Fig. 3a). The number of neck canal cells perhaps reflects the evolutionary rank of the
archegoniates. Four or eight neck canal cells are a characteristic of the ancient archegoniates.
The liverwort Marchantia polymorpha produces 4 neck canal cells during oogenesis (Durand,
1908; Cao et al, 2011b). The moss Mnium affine possesses 8 neck canal cells (Haupt, 1953).
In the lycophyte, Lycopodium clavatum, the mature archegonium contains six neck canal
cells (Haupt, 1953). But in macrophyllous ferns (including eusporangiate and
leptosporangiate ferns), the archegonium only has one neck canal cell with two nuclei
(Haupt, 1953; Bell & Duckett, 1976; Cao et al, 2009; 2010b,c; 2011a; 2012a,b). Therefore,
it seems reasonable to regard the horsetails as transitional group between the liverworts, the
mosses, lycophytes and the macrophyllous ferns.
It is interesting that the two neck canal cells in Equisetum arvense are divided by
an oblique wall in present investigation. However, vertical division of the neck canal cell in
the species, E. laevigatum, E. telmateia, E. arvense, and E. kansanum (Walker, 1921, 1931);
longitudinal division in E. arvense, E. hyamale, and E. limosum (Jeffrey, 1899); transverse
division in E. telmateia (Campbell, 1918) were reported in previously. Only the species
Equisetum scirpoides shows an oblique division (Walker, 1937). The present investigation
shows that the oblique division perhaps the only correct division mode of the neck canal
cell. Vertical or transverse divisions of the neck canal cells may be caused by the sectioning
directions.
3. Organelles in the egg and the canal cells
Vesicles and plastids in the canal cells and the egg can be identified clearly with
the light microscope. The vesicles are obvious in the central cell and young egg, but vanish
in the later stages egg development. The disappearance of the vesicles is considered to be
related to the separation cavity in filicopsida (Yang et al., 2009; Huang et al., 2011; Cao et
al., 2010b,c, 2011a). However, no prominent separation cavity is formed during oogenesis
of Equisetum. The biological significance of the vesicles in Equisetum is still obscure. The
plastids in the egg and the canal cell can be identified by presence of the starch grains,
which are deep stained by the toluidine blue. The present investigation shows that the starch
grains in the egg accumulate increasingly with development of the egg. However, the starch
G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 99

grains in the canal cells decompose gradually with maturation of the egg. Simultaneously,
mucilaginous materials increase outside the canal cells. Finally, these disintegrating materials
and the decomposed canal cells are expelled outside the archegonium and form a empty
canal for sperm entrance. Features of other organelles, such as mitochondria, Golgi bodies,
endoplasmic reticula, require observations with a transmission electron microscope.
4. The neck of the archegonium
Microscopic observations indicated that the terminal cells of the neck in Equisetum
elongates prominently as the archegonium near matures (Walker, 1937; Hauke, 1968).
Sections show that all the neck jacket cells elongate more or less besides special elongation
of the terminal cells. Furthermore, the elongation of the jacket cells of the archegonial neck
may be at a cost of the cytoplasm, which is abundant in the young archegonium stage (Fig.
3a), but decreases rapidly as the archegonium matures.
ACKNOWLEDGEMENTS

This research was supported by National Natural Science Foundation of China


(30970267), The key project of Shanghai Municipal Education Commission (12ZZ128).
REFERENCES

BELL PR & DUCKETT JG 1976 Gametogenesis and fertilization in Pteridium Bot J Linn Soc 7 3 :
47–78
CAMPBELL D H 1918 The structure and development of mosses and f erns New York Macmillan Co
CAO JG, YANG N Y & WANG QX 2009 Ultrastructure of the mature egg and fertilization in the fern
Ceratopteris thalictroides (L) Brongn J Integr Plant Biol 5 1 : 243-250
C AO J G , H UA N G W J & WA N G Q X 2 0 1 0 a M i c r o s t r u c t u r a l o b s e r v a t i o n o n t h e d ev e l o p m e n t o f
g a m e t o p h y t e s a n d o o g e n e s i s i n t h e f er n A d i a n t u m f l a b e l l u l a t u m L A c t B o t B o r e a l O c c i d e n t
Sin, 3 0 : 0645-0651
C AO J G , WA N G Q X & BAO W M 2 0 1 0 b F or ma t i o n o f t h e f e r t i l i z a t i o n p o r e d u r i n g o o g e n e s i s o f t h e
fern Ceratopteris thalictroides J Integr Plant Biol 5 2 : 518–527
CAO JG, WANG QX, DAI XL & DUCKETT JG 2010c Ultrastructural observations of oogenesis in the
fer n Adiantum flabellula tum L (Adiantaceae) Amer Fern J 1 0 0 : 93–102
C AO J G , DA I X L & WA N G Q X 2 0 1 1 a A r c h e go n i a l d eve l o p m e n t a n d o o g e n e s i s o f t h e f e r n
Plagio gyr ia euphle bia and their phylogenetic significance Amer F er n J 1 0 1 : 231–240
CAO JG, WANG G & WANG QX 2011b Micr ostructur al observations and cytochemical studies on the
archegonial development and oogenesis of Marchantia polymorpha Plant Sci J 2 9 ( 5 ) : 607-
612
C AO J G , DA I X F & WA N G Q X 2 0 1 2 a C y t o l o g i c a l f e a t u r e s o f o o g e n e s i s a n d t h e i r ev o l u t i o n a r y
significance in the fern Osmunda japonica Sex Plant Repro 2 5 : 61-69
C AO J G , DA I X L & WA N G Q X 2 0 1 2 b U l t ra s t r u c t u r a l a n d cy t o c h e m i c a l s t u d i e s o n o o g e n e s i s o f t h e
fern Pteridium aquilinum Sex Plant Repro 2 5 : 147-156
DES MARAIS D L, SMITH A R, BRITTON DM & PRYER K M 2003 Phylogenetic relationships and
100 Indian Fern Journal Volume XXXI (2014)

evolution of extant horsetails, Equisetum , based on chloroplast DNA sequence data (rbcL and
trnL-F) Int J Pl Sci 1 6 4 : 737-751
DUCKETT JG 1970 Sexual behaviour of the genus Equisetum, subgenus Equisetum Bot J Linn Soc
6 3 : 327-352
DUCKETT JG 1972 Sexual behaviour of the genus Equisetum , subgenus Hippochaete Bot J Linn Soc
6 5 : 87-108
DUCKETT JG 1973a Comparative morphology of the gametophytes of the genus Equisetum , subgenus
Equisetum Bot J Linn Soc 6 6 : 1-22
DUCKETT JG 1973b An Ultrastructur al study of the differ entiation of the spermatozoid of Equisetum
J Cell Sci 1 2 : 95-129
DUCKETT JG 1977 Tow a r d s a n u n d e r s t a n d i n g o f s ex d e t e r m i n a t i o n i n E q u i s e t u m : a n a n a l y s i s o f
regeneration in gametophytes of the subgenus Equisetum Bot J Linn Soc 7 4 : 215-242
DUCKETT JG 1979a A n e x p e r i m e n t a l s t u d y o f t h e r e p r o d u c t iv e b i o l o g y a n d h y b r i d i z a t i o n i n t h e
European and North American species of Equisetum Bot J Linn Soc 7 9 : 205-229
DUCKETT JG 1979b Comparative morphology of the gametophytes of Equisetum subgenus
Hippochaete and the sexual behaviour of E ramosissimum subsp debile, (Roxb) Hauke,
E h yemale var affine (Engelm) AA, and E laevigatum A Br Bot J Linn Soc 79: 179–203
DUCKETT JG 1985 Wild Gametoph ytes of Equisetum sylvaticum Amer Fer n J 75:120-127
DUCKETT JG & Bell PR 1977 An ultrastructural study of the mature spermatozoid of Equisetum
Philosophical Transactions of the Royal Society of London Series B, Biological Sciences,
2 7 7 ( 9 5 3 ) : 131-158
DUCKETT JG & DUCKETT AR 1980 R e p r o d u c t iv e b i o l o g y a n d p o p u l a t i o n d y n a m i c s o f w i l d
gametophytes of Equisetum Bot J Linn Soc 8 0 : 1–40
DURAND E J 1908 The Development of the Sexual Organs and Sporogonium of Marchantia
pol ymorpha Bul Torr ey Bot Club 3 5 ( 7 ) : 321-335
H AU K E R L 1 9 6 7 S ex u a l i t y i n a w i l d p o p u l a t i o n o f E q u i s e t u m a r ve n s e g a m e t o p hy t e s A m e r F e r n J
5 7 : 59-66
HAUKE R L 1968 Gametangia of Equisetum bogotense Bull Torrey Bot Club 9 5 : 341-345
H AUKE R L 1969 Gametophyte development in Latin Amer ican horsetails Bull Torr ey Bot Club 9 6 :
568-577
H AU K E R L 1 9 7 1 T h e e ff e c t o f l i g h t q u a l i t y a n d i n t e n s i t y o n s e x u a l e x p r e s s i o n i n E q u i s e t u m
gametophytes Amer J Bot 5 8 : 373-377
H AU K E R L 1 9 7 7 E x p e r i m e n t a l s t u d i e s o n g r ow t h a n d s e x u a l d e t e r m i n a t i o n i n E q u i s e t u m
gametophytes Amer Fer n J 6 7 : 18-31
HAUKE R L 1979 A taxonomic monograph of Equisetum subgenus Equisetum Nova Hedwigia 30:385-
455
H AUKE R L 1980 Gametophytes of Equisetum diffusum Amer Fern J 70(2): 39-44
H UA N G W J , C AO J G & WA N G Q X 2 0 1 1 M i c r o s t r u c t u r a l O b s e r va t i o n s o n t h e D ev e l o p m e n t o f
G a m e t o p h y t e s a n d O o g e n e s i s i n t h e F e r n Pteridium aquilinum v a r latiusculum B u l l B o t R e s
3 1 : 159-163
H AUPT A W 1953 Plant mor phology New York McGr aw-Hill Book Company, Inc
G E WANG et al. : Microstructural Observations on the Development of Gametophytes & Oogenesis 101

JEFFREY E C 1899 The development, structure, and affinities of the genus Equisetum Mem Boston
Soc Nat Hist 5 : 155-190
JONES L E & HOOK P W 1970 Growth and Development in microculture of gametophytes from
stor ed spores of Equisetum Amer J Bot 5 7 ( 4 ) : 430-435
L I U J X , L I U B D , BAO W M & AO Z W 1 9 9 4 F iv e s p e c i e s s t u d i e s o n t h e d ev e l o p m e n t
gametophytes of Equisetaceae Natural Sci J Harbin Normal Univ 1 0 ( 2 ) : 77-89
MATZKE EB 1941 Gametoph ytes of Equisetum Arvense L Torr ey a 4 1 ( 6 ) : 181-187
MESLER MR & LU KL 1977 Large gametophytes of Equisetum hyemale in nor ther n Calif ornia Amer
Fern J 6 7 : 97-98
SHARP L W 1912 Spermatogenesis in Equisetum Bot Gazette 54(2):89-119
PRYER K M, SCHNEIDER H, SMITH A R, CRANFILL R, WOLF PG, HUNT J S & SIPES S D 2001
Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants
Nature 4 0 9 : 618-622
RENZAGLIA K S, DENGATE S B, SCHMITT S J & DUCKETT J G 2002 Novel features of Equisetum
arvense spermatozoids: insights into pteridophyte evolution New Phytologist 1 5 4 ( 1 ) : 159-174
R E N Z AG L I A K S , D U F F R J, N I C K R E N T D L & G A R BA RY D J 2 0 0 0 Veg e t a tive a n d r e pr o d u c t ive
i n n o va t i o n s o f e a r l y l a n d p l a n t s ; i m p l i c a t i o n s f o r a u n i fi e d p h y l o g e n y P h i l o s Ta n s s S e r B ,
3 5 5 : 769-793
S M I T H A R , P RY E R K M , S C H U E T T P E LT E , KO R A L L P, S C H N E I D E R H & WO L F P G 2 0 0 6
A classification for extant f erns Taxon 5 5 : 705-731
WALKER E R 1921 The Gametophytes of Equisetum laevigatum Bot Gazette 7 1 : 378-391
WALKER E R 1931 The Gametophytes of Three Species of Equisetum Bot Gazette 9 2 : 1-22
WALKER E R 1937 The Gametophyte of Equisetum Scirpoides Amer J Bot 2 4 : 40-43
YA N G N Y, C AO J G & WA N G Q X 2 0 0 9 For ma t i o n a n d D ev e l o p m e n t o f A r c h eg o n i u m i n t h e f e rn
Ceratopteris thalictroides (L) Br ongn J Wuhan Bot Res 2 7 : 569-573
Indian Fern J. 3 1 : 102-111 (2014) ISSN 0970-2741

S O M E ADDITIONS T O THE PTERIDOPHYTE FLORA OF


HIMA
ACC H A L PRADESH (W
(W.. H I M A L AYA )
AY
S. P. KHULLAR* AND SUNIL VERMA
Department of Botany, Panjab University, Chandigarh-160014

(Received May 5, 2014; Revised Accepted July 9, 2014)

ABSTRACT
Extensive explorations undertaken for the preparation of a Pteridophyte Flora of
Himachal Pradesh have resulted in the collection of four fern species not previously recorded
from Himachal Pradesh. These are : Leptochilus ellipticus (Thunb.) Noot., Tomophyllum
donianum (Spreng.) Fraser-Jenk. & Parris, Vittaria linearifolia Ching and Cryptogramma stelleri
(S. G. Gmel.) Prantl. New localities have been found for Microlepia setosa (Sm.) Alston and
Oleandra walichii (Hook.) C. Presl. A correction has been made for the report of C stelleri by
Alka et al. 2010.
K e y W o rrd
d s : Himachal Pradesh, India, First, Records, three, ferns.

INTRODUCTION

Ever since [the Europeans] botanists started investigating the flora of India,
especially the Himalaya, Himachal Pradesh has been a centre of great interest and attraction.
Collections of the early European collectors in the Government Herbarium, Saharanpur, now
housed at FRI, Dehra Dun, were catalogued by Trotter and Hope (1890). Some of these
collectors were Clarke, Trotter, Hope, Herschel, Blanford, Duthie, Davidson, King,
McDonell, the Mackinnon brothers and others. Pteridophytes of Himachal Pradesh find
mention in the works of Beddome, Clarke, Blanford and Hope (for details see Khullar,
1994). After Independence in 1947 interest in Pteridophytes was kept alive by Mehra, Bir,
Verma and Loyal at first and was later carried forward by Khullar, Dhir, Vasudeva, Seth
and others. Monumental contributions to the Fern Flora of the West Himalaya (indeed for
the entire country) have been made by Fraser-Jenkins. The first named author (SPK) is in
the process of compiling an Illustrated Pteridophyte Flora of Himachal Pradesh. For this
purpose fresh collections have been made from a number of places not visited earlier and
the results have been very interesting. Some of these findings are enumerated here.
Specimens have been deposited in Herbarium, Botany Department, Panjab University,
Chandigarh (PAN).
1. Le ptoc
Leptoc hilus ellipticus (Thunb.) Noot.
ptochilus
Collected from: Himachal Pradesh, MANDI, Durgapur, c. 1500 m altitude (Coll nos.
SV 5402, SV 5454, SV 5458, July 2008). (PLATES 1 & 2)
Earlier reported by Khullar et al. (2014) as L. pedunculatus which was merely a
lapsus mentis, by which was intended ‘ellipticus’.
Previously reported from the West Himalaya, from Dehra Dun, by Hope (1903) and

* E-mail : sp.khullar@gmail.com
S P Khullar & Sunil Verma : Some Additions to the Pteridophyte Flora of Himachal Pradesh 103

PLATE 1. A Field photograph of Leptochilus ellipticus (Thunb.) Noot.


PLATE
104 Indian Fern Journal Volume XXXI (2014)

PLATE 2. Leptochilus ellipticus (Thunb.) Noot.


PLATE
S P Khullar & Sunil Verma : Some Additions to the Pteridophyte Flora of Himachal Pradesh 105

PLATE 3. Tomophyllum donianum (Spreng.) Fraser-Jenk. & Parris


PLATE
Figs.1. Field photograph, 2, 3. Individual plants
106 Indian Fern Journal Volume XXXI (2014)

PLA TE 4. Vittaria linearifolia Ching.


PLATE
F ig 1. Field photograph, 2. A single plant
S P Khullar & Sunil Verma : Some Additions to the Pteridophyte Flora of Himachal Pradesh 107

from Nepal to Bhutan by Beddome (1883). Fraser-Jenkins (2008) mentions that the name
L. pothifolius (D. Don) Fraser.-Jenk. has been misunderstood and had been frequently
misapplied to L. ellipticus. The species, L. ellipticus has been referred to as L. (Colysis)
pothifolia by many Indian workers (see Fraser-Jenkins 2008).
Khullar (2000) mentioned the distribution of this fern (under the name Colysis
pothifolia (D.Don) C.Presl) from Uttarakhand (Chakrata, Deoban, Mussoorie, Garhwal
Himalaya etc., but not from Himachal Pradesh). No distribution data has been given for
this fern in Fraser-Jenkins (2008: 60-61, 539). However in an email to SPK of 16.4.2014,
Fraser-Jenkins says that L. pothifolius occurs from Nepal eastwards.
L. ellipticus has been mentioned in earlier literature under various genera
(Hemionitis, Selliguea, Polypodium, Gymnogramma).
L. pothifolius has a very long, straight, slightly thicker rhizome than L. ellipticus
(commonly 30 or 40 cm long), often with only one or a couple of fronds throughout the
length of the rhizome. It has a much more glossy, dark green top surface than L. ellipticus
and it has few lobes in the frond (2-4). In sterile fronds the lobes are often very wide,
rounded and widely fused into each other and the lobes are more sloping upwards than in
L. ellipticus.
L. ellipticus has more side lobes, varying considerably in width, but narrower than
L. pothifolius, widely subtended and more deeply cut to their bases (Fraser-Jenkins pers.
comm. to SPK e-mails 16.4.2014; 4.5.2014).
2. Tomoph yllum donian
omophyllum um (Spreng.) Fraser-Jenk. & Parris
donianum
This species was previously known as Ctenopteris subfalcata (Blume) Kunze in
Indian Fern literature but Parris (2000) found that the type species of Ctenopteris belongs
to another genus and Parris & Fraser-Jenkins (in Fraser-Jenkins 2008) found that T.
subfalcatum (Blume) Parris is a distinct species, which they transferred to the genus
Tomophyllum as T. donianum. Presently collected from: KULU: Kasol, c. 2700 m altitude
on damp, mossy rocks (SV 240003; SV 24007). This is the first record of this fern from
Himachal Pradesh. (PLATE 3)
A very rare fern in the West Himalaya it has been recorded by Hope (1903) as
eastwards of Garhwal (Mackinnon); Kumaun, Kalimundi (Duthie). In recent years it has
been reported by Pande & Pande (2003) from Almora: Sunderdhunga Glacier, 4100 m
altitude (YPS Pangtey & SS Samant); Pithoragarh: Kalimuni (PC Pande & HC Pande);
Sikkim; Darjeeling Hills; Arunachal Pradesh; Meghalaya; Manipur; Khasi Hills; Nepal;
Bhutan; China; Myanmar.
3. Vittar ia linear
ittaria if
linearif olia Ching
ifolia
Collected from: KULU near Kasol, 2500 m, lithophytic (SV 10018; SV 10020).
(PLATE 4) This species also occurs in Uttarakhand (BM, BSD) (Fraser-Jenkins, email,
4.5.2014).
108 Indian Fern Journal Volume XXXI (2014)

In the Catalogue of Ferns in the Government Herbarium, Saharanpur (Trotter &


Hope 1890; 115) there is no mention of any Vittaria from Himachal Pradesh, but “V. lineata
(L.) Sm., by which was meant V. flexuosa Fée, has been recorded from a few places in
Kumaun: near Askot; above Gini, near Sosa; and from Landour. Mussoorie (Vicary) and
Garhwal (Duthie). Hope (1903) also recorded V. lineata from Kangra Valley, Kulu (collected
by McDonell and reported in Hope 1903 p 280Trotter’s list. Fraser-Jenkins adds (email
4.5.2014) “V. lineata is exclusively a C. and S. American species, not present in Asia. Any
mention from India is in error for the common and widespread V. flexuosa and V.
taeniophylla (syn.: V. himalayensis), which both occur in the W. Himalaya (taeniophylla
from Uttarakhand)”.
The present specimens from Kulu have been identified by Fraser-Jenkins (Nov.
2013) when he was visiting Chandigarh. Fraser-Jenkins (2008 and email 4.5.2014) adds :
In V. linearifolia the fronds are narrower and smaller than most V. flexuosa - (a more
common species) - not entirely indistinguishable, though the caricina and ophiopogonoides
forms of flexuosa can be as narrow; but most importantly V. flexuosa has a raised inner
lip to the sori. The raised inner lip means the sorus is not superficial in V. flexuosa, but it
is superficial in linearifolia and is actually positioned slightly in from the margin, clear in
younger fronds. Upon maturity the sori fill up all the space on either side of the midrib,
sori are yellowish. V. linearifolia has a characteristically wide and rectangular midrib
[beneath] with angled corners and a flat top surface [to the midrib]
Vittaria linearifolia was first reported from Arunachal Pradesh by Dixit (1981)
and is also known from Nepal, Sikkim, Uttarakhand, Bhutan, and is perhaps scattered
throughout the eastern Himalaya. Fraser-Jenkins (2008) recognized 10 species of this genus
from India. In his list of W. Himalayan fern Fraser-Jenkins lists four species of this genus
(mostly from Uttarakhand eastwards) The present record is the first record of this fern from
Himachal Pradesh.
A CORRECTION

Cryptogramma stelleri (S.G.Gmel.) Prantl


First collected by McDonell from Chamba (see Hope p 107 under Pellaea gracilis
Hook).Presently collected from CHAMBA: Sundarasi, above 3000 m altitude (SV 5399; SV
1867, SV 1887, SV 1888). Also known from Afghanistan, Pakistan, Jammu & Kashmir,
Uttarakhand and further east.
The specimens of Alka Kumari et al. (2012) from KANGRA: Dhauladhar range,
were on examination found to be the more common C. brunoniana Wall. ex Hook. & Grev.
The two species can easily be distinguished by the terminal fertile lobe (pinnule) being much
elongated (almost twice as long as the immediate lateral pinnules) in C. stelleri. PLATES
5 & 6
S P Khullar & Sunil Verma : Some Additions to the Pteridophyte Flora of Himachal Pradesh 109

PLATE 5. F
PLATE igs.1. & 2. Cryptogramma stelleri (S. G. Gmel.) Prantl. 3. A portion enlarged
Figs.1.
110 Indian Fern Journal Volume XXXI (2014)

PLATE 6. Cryptogramma brunoniana Wall. ex Hook. & Grev.


PLATE
S P Khullar & Sunil Verma : Some Additions to the Pteridophyte Flora of Himachal Pradesh 111

EXTENDED DISTRIBUTION

1. Micr ole
Microle pia setosa (Sm.) Alston
olepia
Reported from Himachal Pradesh by Alka Kumari et al. (2010) first from adjacent
to the fernery and garden of the IHBT, Palampur, and later from Banuri . The same species
has also been collected by us from KULU: Parvati Valley, Kasol forest, 2100 m. (SV 7172).
2. Oleandra walichii (Hook.) C.Presl
Earlier known only from SHIMLA District (Glen, Chadwick Falls, Narkanda and
Baghi) has now been collected from MANDI: 4 km from Jhatingri, 2000 m altitude (SV
11120, SV 11115, SV 12119).
ACKNOWLEDGEMENTS

I am highly indebted to my dear friend Mr. C.R. Fraser-Jenkins, Kathmandu, who


identified or confirmed the identity of the specimens besides giving some very useful inputs
and editing the typescript.
REFERENCES

ALKA KUMARI, BRIJ LAL & FRASER-JENKINS C R 2010 Microlepia setosa (Sm.) Alston – A
New Generic Record in the Pteridophyte Flora of Himachal Pradesh, India Indian Fern J 2 7 :
376-382
A L K A K U M A R I , A RU NAVA DAT TA & S A N JAY K R U N I YA L 2 0 1 2 C r y p t o g r a m m a s t e l l e r i ( G m e l )
Prantl. - A new fern r ecord for Himachal Pr adesh Indian J Forestry 3 5 ( 3 ) : 393-395
BEDDOME 1883 Handbook to the Ferns of British India, Ceylon and the Malay Peninsula Thacker
Spink & Co
F R A S E R - J E N K I N S 2 0 0 8 Ta x o n o m i c R ev i s i o n o f Thr ee Hundr ed Indian Subcontinental Pter idoph ytes
With a Revised Census-List Bishen Singh Mahendra Pal Singh Dehra Dun
H O P E C W 1 8 9 9 - 1 9 0 4 Th e Fer n s o f N o r t h We s t e r n I n d i a R ep r i n t B i s h e n S i n g h M a h e n d ra Pa l S i n g h
Dehra Dun 19
K H U L L A R S P 1 9 9 4 , 2 0 0 0 A n I l l u s t r at e d F e r n F l o r a o f We s t H i m a l a y a Vo l 1 ( 1 9 9 4 ) Vo l 2 ( 2 0 0 0 )
International Book Distributors Dehra Dun India
K H U L L A R S P, S U N I L V E R M A & I B P R A S H E R 2 0 1 4 P t r e r i d o p hy t i c F l o r a o f H i m a c h a l P r a d e s h ,
101st Indian Science Cong ress, J ammu Abs. No.086(p 224-225)
PANDE H C & PANDE P C 2003 Fern Flora of the Kumaon Himala ya Vol 1 Bishen Singh Mahendra
Pal Singh, Dehra Dun India
PARRIS B S 2007 F ive new g enera and three new species of Grammitidaceae (Filicales) and the Re-
estab lishment of Oreo gr ammatis Gard’s Bull Singapor e 5 8 ( 2 ) : 233-274
TROTTER EW & HOPE CW 1890 The Catalogue of Ferns in the Government Herbarium, Saharanpur
Govt Pr ess Allahabad
Indian Fern J. 3 1 : 112-117 (2014) ISSN 0970-2741

GC-MS PR OFILE OF LYGODIUM FLEXUOSUM L . ( L


ROFILE YGODIA
LY ACCEAE)
A MEDICIN
MEDICINA AL FERN FR
ROOM NOR
RTT H WESTERN GHA TS
GHAT S..
KANCHAN DUBAL1 AND MANISHA KALE2*
1
Research student, Department of Botany, Jaysingpur College, Jaysingpur.
2
Associate Professor, Department of Botany, Jaysingpur College, Jaysingpur

(Received July 5, 2014; Revised accepted July 31, 2014)

ABSTRACT
Lygodium is an abundant genus comprising of around 6 species in India. Analysis of
the bioactive components present in the whole plant ethanol extract of species Lygodium
flexuosum L. from Northern Western Ghats has been carried out. The maximum number of
compounds recorded is seven, some of which shows positive medicinal properties hindering the
harmful causative agents. The presence of compounds like Butylated Hydroxytoluene (BHT) and
esters of Hexadecanoic acid (Palmitic Acid) are responsible for being the plant as a major potent
medicinal fern.
d s : Lygodium flexuosum (L.) Sw. GC-MS Analysis, Bioactive
K e y W o rrd
components, BHT.

INTRODUCTION

India has an ancient heritage of traditional medicines. However there is no authentic


record of medicines used by the primitive man but the Rig Veda which is the oldest book
supplies curious information on this subject (Kirtikar & Basu 2005). It is well known that
ferns are a potent source of herbal medicines and are very effective against numerous
harmful diseases as they never show the appearance of any such symptoms. Also, nature
has provided a complete remedy to cure all ailments of mankind. The history of herbal
medicines is as old as human civilization (Kokate et al. 2006).
Lygodium flexuosum (L.) Sw. is a large terrestrial climbing rhizomatous perennial
fern commonly known as ‘Maiden hair creeper’. It belongs to the family Lygodiaceae
(Manickam & Irudayaraj 1992). Lygodium is an abundant genus comprising of 10 species
(Dixit & Vohra 1984); six species (Fraser-Jenkins 2008). It is widely distributed in the
India up to an elevation of 1500 m. The fern, L. flexuosum has been very commonly used
in treating various ailments like jaundice, dysmenorrheal, wound healing and eczema. It is
the rich source of alkaloids, flavonoids, saponins and cumarin. Rachis of the plant is often
tied over forehead to reduce headache. The same when tied on hand secures it from evil
spirit (Shil & Dutta Choudhary 2009). Rhizome powder is useful in skin diseases. Plants
are used as expectorant, rheumatism, sprains, scabies, eczema and cut wounds. Fresh roots
boiled with mustered oil used in casbundes and rheumatism (Upreti et al. 2009). The
aqueous extract of the rhizome is used for the treatment of gonorrhea. The part of rhizome
is also tied on the waist (Singh 1999). The plant is used to pleuicy ((Jain S K 1991)) Extract
of stem & rhizome is taken orally twice a day for a week for curing sexual disease like

* E-mail : manishavkale@gmail.com
Kanchal Dubal & Manisha Kale : GC-MS Profile of Lygodium flexuosum L. (Lygodiaceae) 113

gonorrheoa and spermatorrhoea. Plant is considered to have antibacterial properties; spores


cure high fever (Singh et al. 2005).
The GC-MS is a tool of choice widely adopted for identification of different
substances present in the natural raw materials. It is extensively used for the analysis of
these compounds which include esters, fatty acids, alcohols, aldehydes, terpenes etc. It can
analyze compounds even in minor concentrations. The objective of the present study is to
evaluate the phytochemical constitution of L. flexuosum.
MATERIAL AND METHODS
ATERIAL

1. Plant material
Plant material of Lygodium flexuosum (L.) Sw. was collected from wild environs
of Castle Rock, Anmode during the month of September to October in its reproductive
stage. The plant specimen was identified with help of Pteridophyte flora of Western Ghats,
South India (Manickam and Irudayaraj 1992). The voucher specimen is deposited in the
Department of Botany, Jaysingpur College.
2. Preparation of Extract
Whole plant material was washed with distilled water and shade dried. The dried
sample pulverized to fine powder manually. The required quantity of whole plant powder
(5 gm) treated with subsequent quantity of ethanol (50 ml) and were exhaustively extracted
using Soxhlet apparatus. The extract is then concentrated to 1ml and employed in GC-MS
analysis of different compounds.
3. GC-MS Anal ysis
Analysis
GC-MS analysis of ethanol extract of L. flexuosum was performed using a Perkin-
Elmer GC Clarus 500 system and Gas chromatograph interfaced to a Mass spectrometer (GC-
MS) equipped with a Elite-I, fused silica capillary column (30mmX0.25m).
4. Identification of Compound
Identification was based on the molecular structure, molecular mass and calculated
fragments. Interpretation on mass spectrum GC-MS was conducted using the database of
National Institute Standard and Technology (NIST) having more than 62,000 patterns. The
spectrum of the unknown component was compared with the spectrum of the known
components stored in the NIST library. The relative percentage amount of each component
was calculated by comparing its average peak area to the total areas.
RESULT AND DISCUSSION
RESULT

The GC-MS chromatogram of the seven compound peaks was shown in Figure 1.
About seven compounds were identified. These active principles with their retention time
(RT), molecular formula and molecular weight (MW), Structural formula and concentration
(peak area %) in the ethanol extract of whole plant of Lygodium flexuosum (L.) Sw. are
114 Indian Fern Journal Volume XXXI (2014)

F i g u rree 1 : GC-MS Chromatogram of ethanolic extract of whole plant of L. flexuosum (L.) Sw.

depicted in Table1. The prevailing compounds identified are Benzene, 1-chloro-4-methoxy-


2-methyl-; Butylated Hydroxytoluene; 2-Pentadecanone, 6,10,14-trimethyl-; n-Hexadecanoic
acid; Hexadecanoic acid, ethyl ester; Hexadecanoic acid, butyl ester; Octadecanoic acid,
ethyl ester.
T A B L E 1 : G C - M S A n a lly
y zzed
ed Bioacti
Bioactivv e compounds fr
froo m L y ggodium
odium Fle xuosum ( L . ) S w
Flexuosum w..

Sr
Sr.. Bioactive Molecular Molecular Retention Area Structure
N o . Compounds Weight Formula time %

1. Benzene, 1- 156 gms C8H9ClO 7.66 39.94


chloro-4-
methoxy-2-
methyl- $$

2. Butylated 220 gms C15H24O 11.81 3.76


Hydroxytoluene
$$ Phenol, 2,6-
bis(1,1-
dimethylethyl)-
4-methyl- $$ p-
Cresol, 2,6-di-
tert-butyl- $$
Advastab 401 $$
Antioxidant
DBPC
Kanchal Dubal & Manisha Kale : GC-MS Profile of Lygodium flexuosum L. (Lygodiaceae) 115

Contd. Table 1

3. 2-Pentadecanone, 268 gms C18H36O 15.59 3.19


6,10,14-
trimethyl- $$
Hexahydrofarnesyl
acetone $$ 6,10,
14-Trimethyl-2-
pentadecanone

4. n-Hexadecanoic 256 gms C16H32O2 16.74 31.14


acid

5. Hexadecanoic
acid, ethyl ester
$$ Palmitic acid,
ethyl ester $$
Ethyl
hexadecanoate $$
Ethyl palmitate
$$ 284 gms C18H36O2 17.05 15.20
6. Hexadecanoic 312 gms C20H40O2 18.82 2.60
acid, butyl ester
$$ Palmitic acid,
butyl ester $$ n-
Butyl
hexadecanoate
$$ n-Butyl
palmitate $$
Butyl palmitate

7. Octadecanoic
acid, ethyl ester 312 gms C20H40O2 18.89 4.16

Hexadecanoic acid (Palmitic Acid) is having antioxidants hypocholesterolenic


nematicide, pesticide, lubricant antiandrogenic flavor, hemolytic 5 alpha reductase inhibitor
(Sermakkani and Thangapandian 2012). Antifibronolytic and antialopecic.
BHT is a hindered phenolic compound acting as antioxidant and prevents lipid
oxidation by donation of hydrogen to free radical. It is considered generally recognized as
116 Indian Fern Journal Volume XXXI (2014)

safe for addition to food products (Branen 1975). Butylated hydroxytoluene is primarily
used as a food additive that exploits its antioxidant properties. It is also documented as an
antioxidant additive in such diverse products as cosmetics and pharmaceuticals. It has been
reported to have anti-viral effects, particularly in use against herpes family viruses and in
combination with L-lysine and vitamin C.(Snipes W. et.al 1975; Brugh M JR 1977; Richards
J T et.al 1985; Kim K S et.al 1978; Pirtle E C et.al 1986 ; Chetverikova L K et.al 1989;
Chetverikova L K & Inozemtseva L I 1996). BHT(butylated hydroxytoluene) are closely
related synthetic antioxidants used as preservatives in lipsticks and moisturizers, among other
cosmetics. Although, BHT can induce allergic reactions in the skin (http://
hazmap.nlm.nih.gov.)
CONCLUSION

Ethanol extract of the whole plant of Lygodium flexuosum (L.) Sw. possesses
various potent phytochemicals with diverse economical and medicinal uses. The data revealed
through this study also verifies active principles of medicinal value from L. flexuosum. This
information marks importance of the plant in traditional medical system & its
pharmaceutical relevance. This type of study is the gateway towards understanding the
bioactivity basis of the medicinal plants and it may help for further detailed investigations.
ACKNOWLEDGEMENT

The authors are grateful to UGC, New Delhi for financial assistance and the
Principal, Jaysingpur College, Jaysingpur for providing laboratory facilities.
REFERENCES

BRANEN A L 1975 Toxicology and Biochemistry of Butylated Hydroxyanisol and Butylated Hydroxytoulene
J American Oil Chemists Society Feb 1975 Vol 5 2 (2) : 59-63
BRUGH M J R 1977 Butylated hydroxytoluene protects chickens exposed to Newcastle disease virus
Science 1 9 7 ( 4 3 1 0 ) : 1 2 9 1 - 2
CHETVERIKOVA L K, KI’LDIVATOV I I, INOZEMTSEVA L I, KRAMSKAIA T A & FILIPPOV V K
1989 Factors of antiviral resistance in the pathog enesis of influenza in mice Vestn Akad Med
Nauk SSSR 1 1 : 63-8 [in Russian]
CHETVERIKOVA L K & INOZEMTSEVA L I 1996 Role of lipid peroxida tion in the patho genesis of
inf luenza and search for antiviral protective ag ents Vestn Ross Akad Med Nauk 3 : 37-40 [in
Russian]
D I X I T R D & VO H R A J N ( 1 9 8 4 ) A dictionary of the pteridophytes of India Bot Surv India Publ
Dept of Environment Govt of India
F R A S E R - J E N K I N S C R F J 2 0 0 8 Ta x o n o m i c R e v i s i o n o f I n d i a n s u b c o n t i n e n t a l P t e r i d o p h y t e s B i s h e n
Singh Mahendra Pal Singh Dehra Dun
JAIN S K 1991 Dictionar y of Indian Folk Medicine and Ethnobotany (Dee p Publica tions New Delhi)
K I RT I K A R K R & BA S U B D 2 0 0 5 I n d i a n M e d i c i n a l P l a n t s D e h ra d u n : N at i o n a l B o o k D i s t r i bu t i o n s
p 1
Kanchal Dubal & Manisha Kale : GC-MS Profile of Lygodium flexuosum L. (Lygodiaceae) 117

KO K AT E C K , P U R H O I T A P & G O K H A L E S B 2 0 0 6 Tex t b o o k o f P h a rm a c o g n o s y 3 6 t h e d P u n e :
New Delhi Nirali Prakashan p 1
K I M K S , M O O N H M , S A P I E N Z A V, C A R P R I & P U L L A R K AT R 1 9 7 8 I n a c t iva t i o n o f
Cytomegalovirus and Semliki Forest virus by butylated hydroxytoluene”, J Infect Dis
1 3 8 (1) : 91-4
M A N I C K A M V S & I RU DAYA R A J V 1 9 9 2 P t e r i d o p h y t e F l o r a o f We s t e r n G h a t s - S o u t h I n d i a
B.I Publications Pvt Ltd 2012
PIRTLE E C, SACKS J M & NACHMAN R J 1986 Antiviral effectiveness of butylated hydroxytoluene
a g a i n s t p s e u d o r a b i e s ( A u j e s z ky ’s d i s e a s e ) v i r u s i n c e l l c u l t u r e , m i c e , a n d s w i n e, A m e r i c a n
J Vet Res 4 77(9) : 1892-5.
R I C H A R D S J T, K AT Z M E & K E R N E R 1 9 8 5 To p i c a l bu t y l a t e d h y d r o x y t o l u e n e t r e a t m e n t o f
Genital herpes simplex virus infections of guinea pigs Antiviral Res 5 (5) : 281-90
S E R M A K K A N I M & T H A N G A PA N D I A N V 2 0 1 2 GC-MS Analysis of Cassia Italica Leaf Methanol
Extract Asian J of Pharmaceutical & Clinical Research Vol 5 (2) : 90-94
SHIL S & M DUTTA CHOUDHARY 2009 Ethnomedicinal impor tance of pteridophytes used by reang
tribe of Tripura North East India Ethnobotanical Leaflets 1 3 : 634-43
SINGH H B 1999 Potential medicinal pteridophytes of India & their chemical constituents J Econ Tax
Bot 2 3 (1) : 63-78
SINGH S, DIXIT R D & SAHU T R 2005 Ethnomedicinal uses of pter idophytes of Amarkantak M P
Indian J ournal of tr aditional knowledge Vol 4 (4), October 2005, pp 392-395
S N I P E S W, P E R S O N S , K E I T H A & C U P P J 1 9 7 5 B u t y l a t e d h y d r o x y t o l u e n e i n a c t iva t e s l i p i d -
containing viruses, Science 1 8 8 (4183) : 64-6
UPRETI K, JALAL J S, TEWARI L M, JOSHI G C, PANGTEY Y P S & TEWARI G 2009 Ethnomedicinal
u s e s o f p t e r i d o p hy t e s o f K u m a u n H i m a l a y a , U t t a r a k h a n d , I n d i a J A m e r i c a n S c i e n c e 5 ( 4 ) :
167-170
U S National Library of Medicine, in Haz-Map : Occupational Exposure to Hazardous Agents
2010, http://hazmap.nlm.nih.gov
Indian Fern J. 3 1 : 118-123 (2014) ISSN 0970-2741

PHO
OTT OSYNTHETIC PIGMENTS A N D THEIR DEGRAD
DEGRADA AT I O N
UNDER HYPER THERMIA IN T W
HYPERTHERMIA WOO SPECIES OF S E L A GINELLA
AGINELLA
BEA
AUUV
V.. F R OM RAJ
ROM ASTHAN
RAJA
KANTA MEENA*, SHAHDAB HUSSAIN** AND B. L. YADAV***
Department of Botany, M.L.V. Government College, Bhilwara - 311 001, Rajasthan, India

(Received April 5, 2014; Revised Accepted)

ABSTRACT
Photosyntheic pigments and their behavior under hyperthermia in two species of
Selaginella Beauv. namely Selaginella ciliaris (Retz.) Spring and Selaginella repanda (Desv.)
Spring occurring in Rajasthan have been studied to find out their differential tolerance to the
heat and drought conditions of the state. Selaginella ciliaris confined to Sitamata Wildlife
Sanctuary (Pratapgarh) is occurring at the damp margins of water streams while Selaginella
repanda a widely distributed species grows on rocks as well as on moist margins of river.
Selaginella repanda , a species growing on rocky substratum with maximum amount of
photosynthetic pigments, minimum degradation in chlorophyll and maximum decay in
carotenoids at 350C temperature has been found to be the resistant species to drought conditions
while Selaginella ciliaris with minimum content of photosynthetic pigments, maximum
degradation in chlorophylls and minimum in carotenoids at 350C temperature has been found to
be the least resistant species. These data are in confirmation with the survival and growth
period of these species in nature.
d s : Heat and drought resistance, differential tolerance, hyperthermia,
K e y Wo rrd
photosynthetic pigments

INTRODUCTION

Higher plants are known to manifest a wide spectrum of thermal tolerance. Many
of the metabolic changes triggered are concerned with heat injury while a few are associated
with heat tolerance. Ferns in general require low temperature for their optimum growth.
Several workers have carried out phytochemical investigations such as chlorophyll a & b,
carotenoids, carbohydrates, proline, polyphenols, nitrogen and proteins, to find out the
survival potential of different pteridophytes under stress conditions (Rathore & Sharma
1998, Vyas et al. 1989, Rathore & Sharma 1990, John De Britto & Manickam 1994, Kale
& Upadhye 2003, 2005, Kale 2007, Shaikh & Dongare 2008, Vyas 2008, Bhattacharya &
Halder 2009, 2011, Chunari et al. 2012). Effect of hyperthermia on cell memberane
permeability of pteridophytes of Rajasthan has been investigated by Yadav and Bhardwaja
(1994), Kothari and Yadav (2003), Pareek et al. (2005) and Tripathi et al. (2005 a,b).
Efforts have also been made to investigate the variability in thermal tolerance of fern species
(Sharma et al. 1977, Yadav et al. 1978, Kaur and Yadav 1985, Vyas and Sharma 1988,
Sharma 1989, Rathore and Sharma 1991, Yadav 2011). Unfortunately the fern allies have
largely been ignored from such types of studies except a brief report on ecophysiological
aspect of two species of Selaginella (Sharma 1992).
*Email : kantameenalectbotany@gmail.com; **Email : hshahdab@yahoo.in, ***Email : drblyadav@yahoo.com
Kanta Meena et al. : Photosynthetic Pigments and Their Degradation Under Hyperthermia 119

Genus Selaginella Beauv. is represented by four species in Rajasthan (Gena et al.


1979, Sharma 1992, Dulawat & Chaudhary 2008, Yadav et al. 2011), namely Selaginella
ciliaris (Retz.) Spring, Selaginella rajasthanensis Gena, Bharadwaja & Yadav, Selaginella
repanda (Desv.) Spring and Selaginella reticulata (Hook. & Grev.) Spring. There is a
definite sequence of drying up of these species in nature with the onset of adverse
conditions. Photosynthetic pigments and their behaviour under hyperthermia have been
studied to find out the differential tolerance of Selaginella species to the heat and drought
conditions of Rajasthan.
MA
ATTERIALS & METHODS

Fresh mature plants of S. ciliaris and S. repanda were taken as experimental


material. Selaginella repanda is the widely distributed species and therefore two populations
of this species, one growing on rocky substratum at Gwaparnath (Kota) and the second on
moist river bank at Sitamata Wildlife Sanctuary (Pratapgarh). The population of Selaginella
ciliaris occurring only at Sitamata Wildlife sanctuary were taken for this study. Thermal
treatment of 350C was given to plants following Sullivan (1967) for 1hour, 2 hour and 4
hour time period along with controls at room temperature. The plants were then
homogenized in Acetone and photosynthetic pigments (total chlorophylls and total
carotenoids) were measured according to Robbelen’s method (1957).
RESULTS
LT

The observations on photosynthetic pigments in mature plants have been presented


in Table 1. The total chlorophyll content was recorded to be highest (0.91 mg/gfw) in
Selaginella repanda (Gwaparnath population) and the lowest (0.36 mg/gfw) in Selaginella
ciliaris. The total chlorophyll content was recorded to be higher in both the populations of
Selaginella repanda than Selaginella ciliaris. Further a perusal of Table 1 indicates that the
content of chlorophyll a is almost equal to the chlorophyll b in Selaginella ciliaris .
Whereas chlorophyll b is slightly more than chlorophyll a in both the populations of

TA B L E 1 : P h o t o s y n t h e t i c p i g m e n t s i n m a t u r e p l a n t s o f t w
woo s p e c i e s o f S e l aag
g i n e l l a f rro
om Rajasthan
(Cholorophyll & Carotenoid contents mg/g fresh weight)

S. Species Locality Chl-a Chl-b Total Chl Total


No. Carotenoids

1. Selaginella ciliaris Sitamata wild 0.18 0.17 0.36 0.69


life sanctuary
2. Selaginella repanda Sitamata wild 0.18 0.20 0.40 0.32
life sanctuary
3. Selaginella repanda Gwaparnath 0.41 0.46 0.91 0.80
120 Indian Fern Journal Volume XXXI (2014)

TA B L E 2 : E ffff e c t o f hhy
y p e r t h e r m i a o n p h o t o s y n t h e t i c p i g m e n t s ( t o t a l cch
h l o r o p hhy
ylls and carotenoids)
in two species of Selaginella from Rajasthan at various temperature regims for a specified
p e rriod
iod of time (V
(Vaa l u e s rree p rresented
esented as % de
degg r a d aatt i o n )

S. Species Locality Chlorophylls Carotenoids


No. 3 5 0C 3 5 0C

1h 2h 4h 1h 2h 4h

1. S. ciliaris Sitamata wildlife 23.04 21.02 30.15 1.07 14.05 25.98


sanctuary
2. S. repanda Sitamata wildlife 7.03 20.09 27.98 17.24 48.60 52.91
sanctuary
3. S. repanda Gwaparnath 18.15 22.38 22.80 4.16 35.83 56.71

Selaginella repanda which has comparatively broad laminated leaves than Selaginella
ciliaris . Carotenoid contents were recorded to be higher in Selaginella ciliaris
(0.69 mg/gfw) and lower (0.32 mg/gfw) in Selaginella repanda of Sitamata Wildlife
Sanctuary’s population. However, the carotenoids were found to be maximum in the
Selaginella repanda of Gwaparnath population.
Data pertaining to the effect of hyperthermia on chlorophyll and carotenoid
pigments have been presented in Table 2 which indicates a maximum degradation in total
chlorophyll pigments at 35 0C for 1hour temperature treatment in Selaginella ciliaris
(23.04%) whereas the degradation in chlorophyll pigments was minimum in Selaginella
repanda of Sitamata wildlife sanctuary population (7.03%). At 4 hour temperature
treatment of 35 0C it is Selaginella ciliaris showing maximum degradation (30.15%)
followed by Selaginella repanda of Sitamata Wildlife sanctuary population while it is
maximum at 2 hour temperature treatment in Selaginell repanda (22.38%) population of
Gwapernath.
Maximum degradation (17.24%) in the carotenoid contents have been observed in
Selaginella repanda population Sitamata Wildlife Sanctuary (Pratapgarh) while minimum
(1.07%) was recorded in Selaginella ciliaris at 350C temperature for 1hour treatment. The
degradation of carotenoids has been recorded to be higher in both the populations of
Selaginella repanda at 2 hours and 4 hours temperature treatment. Thus, an inverse
relationship has been found between these two pigments. In Selaginella ciliaris there is more
degradation of chlorophyll pigments but less in carotenoids whereas in Selaginella repanda
less degradation was observed in chlorophyll pigments and higher in carotenoids.
DISCUSSION

Ludlow and Wolf (1975) reported that greater quantity of chlorophylls has been
Kanta Meena et al. : Photosynthetic Pigments and Their Degradation Under Hyperthermia 121

found to be present in shade loving species than the sun loving taxa. Herbaceous and well
laminated forms like Adiantum lunulatum and Marsilea minuta were found to possess more
chlorophyll in comparison to narrow, coriaceous and reduced laminated forms like
Actinopteris radiata and Pteris vittata (Vyas and Sharma 1988). They have reported higher
content of chlorophyll a than chlorophyll b in reduced, coriaceous and laminated forms. On
the other hand chlorophyll b occurs in more quantity in broad laminated forms. In soft,
wide laminated forms carotenoids occur in higher amount than hard laminated forms.
The results of the present investigation are in agreement with the reports made by
earlier workers (Ludlow and Wolf 1975, John De Britto and Manickam 1994, Rathore and
Sharma 1991, Shaikh & Dongare 2008). There is a greater quantity of chlorophylls in
Selaginella repanda which is a shade and partially exposed species than Selaginella ciliaris
which is sun loving.
There exist a direct relationship between degradation of pigments and drought
resistance. Drought resistant ferns show higher degradation in carotenoid contents and lower
degradation of chlorophylls (Bohra et al. 1979, Yadav & Bhardwaja 1983, Sharma 1989).
This fits in well with the known function of carotenoids which provide protection to the
chlorophyll pigments from photosensitized oxidation (Krinsky 1966).
ACKNOWLEDGEMENTS

Authors are grateful to Prof. T. N. Bhardwaja, Ex Vice-Chancellor, Kota Open


University, Kota and Prof. C. B. Gena, Ex Vice-Chancellor, Maharaja Ganga Singh
University, Bikaner for critical evaluation of manuscript and valuable suggestions. Authors
are also thankful to forest department of Sitamata Wildlife Sanctuary for their cooperation
during field studies. Financial assistance provided by the UGC to KM and BLY under MRP
is gratefully acknowledged. SH is thankful to UGC for award of MANF, SRF.
REFERENCES

B H AT TAC H A RYA ( G O S WA M I ) N & H A L D E R S 2 0 0 9 Ph y t o c h e m i c a l s t u d y o f s o m e f e r n s o f s e m i -


arid habitats of District Burdwan West Bengal Indian Fern J 2 6 : 37 45
B H AT TAC H A RYA ( G O S WA M I ) N & H A L D E R S 2 0 1 1 Phytochemical analysis of fern plants of
different ecosystems Indian Fern J 2 8 : 120-128
BOHRA D R, SONI S R & SHARMA B D 1979 Ferns of Rajasthan Behaviour of chlorophyll and
carotenoids in drought resistance Experientia 3 5 : 332
CHUNARI S S N, MONDAL S & GHOSH M 2012 Phytochemical analysis and screening for active
compounds in Lycopodium cer nuum Indian F er n J 2 9 : 39-43
D U L AWAT C S & C H AU D H A RY B L 2 0 0 8 Selaginella ciliaris (Retz.) Spring (Selaginellaceae
Pteridophyta) A new record for the Rajasthan India Indian Fer n J 2 5 : 106-109
G E NA C B , B H A R DWA JA T N & YA DAV A K 1 9 7 9 A n ew s p e c i e s o f S e l a g i n e l l a fr o m I n d i a A m e r
Fern J 6 8 : 119-122
122 Indian Fern Journal Volume XXXI (2014)

JOHN DE BRITTO A & MANICKAM S 1994 Phytochemical studies on members of Thelypteridaceae


of the Western Gha ts of South India Indian Fer n J 1 1 : 116-123
KALE M V 2007 P h y t o c h e m i c a l a n a l y s i s o f B o l b i t i s a p p e n d i c u l a t a ( Wi l l d ) I wa t s : R a r e a n d
endangered species from Castlerock and Anmode, Maharashtra, South India Indian Fern J
2 4 : 148-150
KALE M V & UPADHYE A B 2003 Study of chlorophylls in the dimorphic leaves of some homosporous
ferns Indian Fern J 2 0 : 67-70
K A L E M V & U PA D H Y E A B 2 0 0 5 S t u d y o f p r o t e i n i n t h e d i m o r p h i c l e av e s o f t h r e e f e r n s o f
Western Ghats South India Indian F ern J 2 2 : 89-91
K AU R R & YA DAV B L 1 9 8 5 Mechanism of drought endurance of some Rajasthan Ferns Indian
Fern J 2 : 90-92
KOTHARI R & YA DAV B L 2003 Comparative effect of h yperthermia on cell membrane permeability
of some ferns Bionature 2 3 (2) : 65-67
KRINSKY N 1966 Biochemistr y of chlor oplast In Goodwin (Ed). Academic press, London 1
LUDLOW C J & WOLF T F 1975 Photosynthesis and respir ation r ates of ferns Amer Fern J 6 5 (2) :
43-48
PAREEK P, TRIPATHI M K & YADAV B L 2005 Effect of hyperthermia on cell membrane permeability
of some fern taxa of Rajasthan. Indian Fern J 2 2 : 121-123
R ATHORE D & SHARMA B D 1988 Fer ns of Mount Abu (Rajasthan) - Study of pr oline in r ela tion
to stress Phytomorphology J 3 8 (2, 3) : 259-261
R AT H O R E D & S H A R M A B D 1 9 9 0 P t e ri d o p hy t e s o f R a j a s t h a n - P h y t o c h e m i s t r y o f I s o e t e s I n d i a n
Fern J 7 : 121-122
R AT H O R E D & S H A R M A B D 1 9 9 1 Phytochemistry of Rajasthan pteridophytes - Study of leaf
pigments in relation to stress Indian Fern J 8 : 9-13
ROBBELEN G 1957 Ztschr unter suchungen an strahlenin duzier ten Blatt arbrnuta ten von Arabidopsin
thaliana (L ) Heynh Z Indulet Absbamn u Verbungaleher e 8 8 : 189-202
SHAIKH S D & DONGARE M 2008 Analysis of photosynthetic pigments in Adiantum lunulatum
Barm. at different localities of Sindhudurg district (Maharashtra ) Indian Fern J 2 5 : 83-86
SHARMA B D 1989 Ferns and fern allies of Rajasthan - Experimental and phytocemical studies
Indian Fern J 6 : 195-203
SHARMA P, YADAV A K & BHARDWAJA T N 1977 Heat and drought resistance in Actiniopteris and
Adiantum Geobios 4 : 256-258
SHARMA S N 1992 Eco-physiological studies of Rajasthan Selaginellas In T N BHARDWAJA & C B
GENA (Eds) Aspects of Plant Sciences. Toda y and Tommor ow’s Printer s and Publisher s New
Delhi (India) 1 4 : 417-422
S U L L I VA N C Y 1 9 6 7 M e c h a n i s m o f h e a t a n d d r o u g h t r e s i s t a n c e i n g r a m s o rg h u m a n d m e t h o d s o f
measurement In Sorghum in seventies (Eds Rao NCP & LR House) Chap 16 pp 247-264 New
Delhi Oxford IBH 1972
T R I PAT H I M K , M E E NA D K , PA R E E K P & YA DAV B L 2 0 0 5a Isoetes L I n R a j a s t h a n – S t u d y o f
effect of hyperthermia on cell membrane permeability Indian Fern J 2 2 : 157-159
T R I PAT H I M K , M E E NA D K & YA DAV B L 2 0 0 5 b H y p e r t h e rm i a a n d c e l l m e m b r a n e p e r m e a b i l i t y
in some common ferns of Rajasthan J Indian Bot Soc 8 4 : 54-56
Kanta Meena et al. : Photosynthetic Pigments and Their Degradation Under Hyperthermia 123

VYAS M S 2008 Phytochemistr y of some pteridophytes of Rajasthan Indian Fer n J 2 5 : 61-65


VYAS M S & SHARMA B D 1988 Phytochemistry of Rajasthan Pter idophytes Indian Fern J 5 : 143-
149
VYAS M S, RATHORE D & SHARMA B D 1989 Phytochemistry of Rajasthan pteridophytes Study of
Phenols in relation to stress Indian Fern J 6 : 244-246
YA DAV A K , S H A R M A P & B H A R DWA JA T N 1 9 7 8 P h o t o s y n t h e s i s a n d R e s p i r at i o n ra t e s o f s o m e
Rajasthan ferns Geobios 5 : 185-187
YADAV B L & BHARDWAJA T N 1983 Temperature and pigment degradation in Rajasthan species of
Ophioglossum L Sci & Cult 4 9 : 246-247
YA DAV B L & B H A R DWA JA T N 1 9 9 4 E ff e c t o f h y p e rt h e r m i a o n c e l l m e m b e r a n e p e r m e a b i l i t y o f
Ophioglossum L Indian Fern J 1 1 : 94-96
YA DAV B L , M E E NA K L , M E E NA K & H U S S A I N S 2 0 1 1 S e l a g i n e l l a r e t i c u l a t a ( H o o k & G r ev )
S p r i n g ( S e l a g i n e l l a c e a e ) - A n ew r e c o r d t o t h e p t e r i d o p h y t i c f l o r a o f R a j a s t h a n , N o r t h -
Western India J Bombay Nat Hist Soc 1 0 8 (3) : 239-241
YADAV M K 2011 Effect of hyperthermia on photosynthetic pigments of Adiantum incisum occurr ing
in Rajasthan Indian Fern J 2 8 : 1-6
Indian Fern J. 3 1 : 124-131 (2014) ISSN 0970-2741

DISCOVER
DISCOVER Y OF DIPLOID CYT
VERY CYTOO TYPE OF HELMINTHOST
HELMINTHOSTAACHYS
ZEYLANICA (L.) HOOK. (HELMINTHOST
(HELMINTHOSTAA CHYACEAE - PTERIDOPHYT
CHYA A)
PTERIDOPHYTA)
SHASHANK KUMAR SINGH*, S. DOMINIC RAJKUMAR**,
SHOBHIT KUMAR SRIVASTAVA*** & RAVI PRATAP GAUTAM****
Department of Botany, St. Andrew's College (PG), Gorakhpur, UP.

(Received November 4, 2013; Revised Accepted 16 May 2014)

ABSTRACT
Helminthostachys zeylanica (L.) Hook. is a tropical and sub- tropical species. The
widely reported meiotic chromosome number of H. zeylanica is n = 94, interpreted as tetraploid
based on presumptive base number, x = 47. Here we report the discovery of the diploid
cytotype, showing n =47, from Kusmi forests, Gorakhpur (U.P., India), which incidentally
establishes the basic chromosome number for Helminthostachys as x = 47.
d s : Diploid cytotype, Helminthostachys zeylanica, Helminthostachyaceae.
K e y Wo rrd

INTRODUCTION

Helminthostachyaceae is a monotypic fern family found in tropical and subtropical


regions. The family consists of a single genus Helminthostachys and it was previously
placed in the family Ophioglossaceae (Clausen 1938) along with the genus Ophioglossum
and Botrychium. Pichi Sermolli (1977) maintained it as a separate family. Helminthostachys
zeylanica is reported to be present in many localities in India like Haridwar, Moradabad,
Gorakhpur, Darjeeling hills, Bengal, Manipur, Assam, Kerala and Tamil Nadu (Beddome
1883, Roy & Kumar 1959, Dixit & Tripathi 1966, Sharma et al., 1969, Chandra 2000). In
Gorakhpur, Uttar Pradesh, this species has been reported from Kusmi forest (Roy & Kumar
1959) and Lakshmipur forest (Dixit & Tripathi 1966). The Present study area (Fig. 1) –
Uttar Pradesh (Gorakhpur) is bounded by Nepal on the North, Uttarakhand on the North-
East, Himachal Pradesh on the North–West and Bihar on the East. These are situated between
23 0 52'N and 31 0 28'N latitudes and 77 0 30'E and 84? 39'E longitude. The present
cytological study of the material from Gorakhpur revealed the existence of a diploid form
within H. zeylanica, which is intended to be recorded here.
MATERIALS AND METHOD
ATERIALS

For cytology, the methodology of Manton (1950), and conventional method of


preparation of herbarium was followed. The collected specimens are preserved and deposited
in Department of Botany, St. Andrew's College, Gorakhpur, Uttar Pradesh, bearing the
accession numbers : 284, 288.
Specimens Examined : Kusmi forest - Gorakhpur, 12.08.2012, 90m, 284, 288 (Fig.
11 & 12).

* E-mail : shashank2771986@gmail.com, ** dominicrajkumar1@gmail.com, *** sshobhit008@gmail.com,


**** rpgautam0288@gmail.com
Shashank K Singh et al. : Discovery of Diploid Cytotype of Helminthostachys zeylanica (L.) Hook. 125

PLATE 1
PLATE

F i gg.. 2 . Habitat photos of Plant (2 A, 2 B) F i gg.. 1 1 . Voucher specimen - SAC 284 F i gg.. 1 2 . Voucher
specimen - SAC 288
126 Indian Fern Journal Volume XXXI (2014)

PLATE 3
PLATE

Fig. 4. Helminthostachys zeylanica (n = 47), F i g . 5 . Helminthostachys zelaynica (n = 47) Line


diagram, Fig. 6. Helminthostachys zeylanica Spores (40x), Fig. 7. Helminthostachys zelaynic a (40x)

OBSER
OBSERVVATIONS AND DISCUSSION

During the recent exploration of Kusmi forest range of Gorakhpur, Uttar Pradesh
Helminthostachys zeylanica was collected for chromosome count. Helminthostachys
zeylanica is a very common species in the study area (Kusmi forest). The present collection
from Kusmi forests (Figs. 2, 3) turned out to be diploid showing n = 47 (Figs. 4, 5–
voucher 284; Figs. 8 & 9 – voucher 288) and this record of a diploid cytotype is the first
report for the species. This is in contrast to all the earlier records of n-chromosome number
in Helminthostachys zeylanica which refer the species to be tetraploid with n = 94 (see Bir
& Verma 2010), reported widely from several localities in India: Western Ghats –
Trivandrum, Kerala, (Ninan 1956,1958; Mahabale & Nair 1972, Sankari Ammal 1990,
Shashank K Singh et al. : Discovery of Diploid Cytotype of Helminthostachys zeylanica (L.) Hook. 127

PLATE 2
PLATE

Fig. 1. 1: Map of the study area - Uttar Pradesh. Fig. 3 a. Frond of Helminthostachys zeylanica
b . Portion of the pinnule showing venation c. Enlarged portion of Sori
128 Indian Fern Journal Volume XXXI (2014)

PLATE 4
PLATE

Fig. 8. Spore mother cell of (SAC 288) Helminthostachys zelaynica (n = 47) Fig. 9. Line diagram
of Spore mother cell of (SAC 288) H. zelaynica (n = 47) Fig. 10. Helminthostachys zeylanica Stomata (40x)
Shashank K Singh et al. : Discovery of Diploid Cytotype of Helminthostachys zeylanica (L.) Hook. 129

Goswami & Khandelwal 1980: Trivandrum and Gorakhpur, n = 92 + 3-4 univalents),


Darbhanga, North Bihar (Sinha et al., 1979), Bihar (Goswami 1987).
Cytology of the species was first reported by Manton & Sledge (1954) from
Ceylon with n = 92-94. From this and other subsequent reports of n = 94 and 2n = 188
(Ninan 1956,1958), Löve et al . (1977) postulated the basic number of Helminthostachys
to be x = 47, which referred the earlier reports of n = 94 as tetraploid. It is surprising to
notice that Sporne (1970, 1976) stated for Helminthostachys “chromosome counts show a
surprising range within the group (Ophioglossales) for Botrychium has a haploid number
n = 45, Helminthostachys n = 46 or 47... (p.140)”, when there was no published report
of Helminthostachys with n = 47. At best the mention of a haploid count of n = 46 or 47
by Sporne (l.c.) was a presumption of its basic number derived from the report of n =
92-94 from Ceylon by Manton & Sledge (1954). The present report of the existence
diploid form (n = 47) of Helminthostachys zeylanica from Kusmi forest, Gorakhpur (Figs.
4, 5– voucher 284; Figs. 8 & 9 – voucher 288) is the first one, and it confirms the base
number as x = 47, earlier only derived by Löve et al ., (1977) from reports of n = 94.
Gorakhpur in Uttar Pradesh (India) is the first distributional area for the diploid cytotype,
which apparently coexists with the tetraploid cytotype, assuming Goswami & Khandelwal's
(1980) report concerned material from both Trivandrum and Gorakhpur. Obviously, the
Kusmi forest of Gorakhpur is in need of further exploration and cytotaxonomic
evaluation.
The structure of sporangia (Fig. 6) and spore (Fig. 7) were studied and were found
to be normal in appearance. The mean diameter of the spores is 36.2 µm and length of the
stomata is 71.6 µm (Fig. 10). Panigrahi & Dixit (1969) and Pant & Khare (1971) reported
spore size as 32-38µm. Devi (1977) recorded the spore size in H. zeylanica as 33 x 42 µm.
The present observation on spore diameter seems to correspond to all these reports. In this
context, the observation of Goswami & Khandelwal (1980) on spore size is 19-24µm,
distinctly smaller than all the earlier reports. It is very likely that the spore size measures
made by Goswami & Nair (1980) and in the present study were based on spores derived
from plants other than those used for cytological studies. It is, therefore, suggested that the
populations of H. zeylanica in the Gorakhpur forests be examined afresh for chromosome
counts, spore size and other features, and vouchers preserved, for taxonomic evaluation.
Assuming spore size related to ploidy level, in general, we suspect that the large-sized spores
belong to the 4x form and the small-sized spores (reported by Goswami & Nair 1980)
would belong to the 2x form. It needs to be confirmed.
The need to explore the taxonomy of 2x and 4x cytotypes assumes importance,
because H. zeylanica is considered to be highly medicinal by the local people around the
Kusmi forest range of Gorakhpur. The rhizome of H . zeylanica is used to treat
cytotoxicity, anti-inflammatory and pulmonary diseases. In Chinese herbal medicine the
rhizome, is used as antipyretic and antiphlogistic agent (Fitrya et al. 2010). In India, the
130 Indian Fern Journal Volume XXXI (2014)

rhizome is used for curing impotency (Singh et al., 1996). In Malaysia, the rhizome used
as anti-diarrheal agent and chewed with areca for whooping cough relief (Jalil et al. ,
1986). Suja et al., (2004) have reported the aphrodisiac properties of the ethanol extract
of rhizome of H. zeylanica.
ACKNOWLEGEMENTS

The authors are thankful to the Principal, St. Andrew's college (PG), Gorakhpur,
Uttar Pradesh for the facilities and the encouragement given to us. The authors are grateful
to Prof. S.C. Verma for the critical review of the paper. Inspite of his eye problem he had
shown immense interest in critically reviewing and enriching this manuscript. One of the
authors (SDR) is thankful to CSIR, India (CSIR Sanction No. 38 (1282) 11/EMR -II) for
the financial assistance.
REFERENCES
BEDDOME R H (1883) Ferns of British India Thacker Spink & Co Calcutta
BIR SS & VERMA SC 2010 Chromosome Atlas of the Indian Pteridophytes (1951-2009), Bishen Singh
Mahendra Pal Singh, Dehra Dun, pp xx 346. 4º [ISBN 978-81-211-0772-3]
CHANDRA S (2000) The Ferns of India (Enumeration, Synonyms and Distribution), International Book
Distributors Dehra Dun
CLAUSEN R T 1938 A monograph of the Ophioglossaceae MEM TORREY BOT CLUB 19 : 1 – 177
DEVI S 1977 Spores of Indian Ferns Today & Tomorrow's Printers & Publishers, New Delhi Pp 1-228
DIXIT S N & TRIPATHI S M 1966 Occurrence of Helminthostachys zeylanica Hook in Lakshmipur forest
(Gorakhpur) Indian For 9 2 : 275-277
FITRYA, ANWAR L, ELIZA & MUHARNI 2010 Ugonin J Flavonoid from Tunjuk langit ( Helminithostachys
zeylanica Linn) root extract Indo J Chem 10 (2) 233-238
GOSWAMI H K & KHANDELWAL S 1980 Observations on Helminthostachys Kaulfuss (Ohioglosassles) Act
Bot Neerl 2 9 : 199-212
GOSWAMI H K 1987 Ohioglosassles: An overview Bionature 17 : 47-89
JALIL J, BIDIN A A & CHYE T S 1986 Phytochemical study of Ophyoglosaceae family species Proc Malays
Biochemistry society conference Fakultas Sains HayatiUniversitas Kebangsaan Malaysia, Bangi
Malaysia 1 2 : 160-164
LÖVE A, LÖVE D & PICHI SERMOLLI REG 1977 Cytotaxonomical Atlas of the Pteridophyta J Cramer,
Vaduz, Pp i-xviii +398
MANTON I & SLEDGE WA 1954 Observations on the cytology and taxonomy of the Pteridophyte flora of
Ceylon Phil Trans Roy Soc, Londo (Biol Sci) 1238: 127-185
MAHABALE TS & NAIR LN 1972 Cytology of some species of the Ophioglossaceae in India Proc Indian
AcadSci 7 55B: 201-214
NINAN C A 1956 Cytology of Ophioglossaceae Curr Sci 25
25: 161-162
NINAN C A 1958 Studies on the cytology and phylogeny of Pteridophytes VI Observations on the
Ophioglossaceae Cytologia 2 33: 291-316 (315)
Shashank K Singh et al. : Discovery of Diploid Cytotype of Helminthostachys zeylanica (L.) Hook. 131

PANIGRAHI G & DIXIT R D 1969 Studies on Indian Pteridophytes IV The family Ophioglossaceae in India
Proc Natn Inst India 3 55B(3) : 230-266
PANT D D & KHARE P K 1971 Notes on the spore morphology of Ophioglossaceae and the occurrence of
Ophioglossum and its gametophyte in the Gangetic Valley Geophytology 1 (1) : 48-53
PICHI SERMOLLI, R E G 1977 Fragmenta Pteridologiae 6 Webbia 31 : 237 – 259
ROY S K & KUMAR H D (1959) Occurrence of Helminthostachys zeylanica Hookin Gorakhpur Curr Sci,
2 8 : 375
SANKARI – AMMAL L 1990 Studies on the cytology and spore morphology of ferns PhD Thesis, University
of Kerala, Trivandrum, India
SHARMA D N, TRIPATHI S M & SRIVASTAVA A K (1969) Pteridophytic flora of the forests of Gorakhpur
district Indian For 9 5 (8) : 526-531
SINGH V, ALI Z A, ZAIDI S T H & SIDDQUI M K 1996 Fitoterapia, Department of Botany Aligarh Muslim
University Aligarh, India
SINHA B M B SRIVASTAVA D P & VERMA A K 1979 Helminthostachys zeylanica (L) Hook- Meiotic
analysis in a population from N Bihar J Cytol Genet 14 : 88-90
SPORNE KR (1970) 1976 The Morphology of Pteridophytes (1976, 4th ed) Hutchinson & Co Ltd, London
SUJA S R, LATHA P G PUSHPANGADAN P & RAASEKHARAN S 2004 Antihepatotoxic activity of
Spilanthes ciliata on paracetamol induced liver damage in rats Pharmaceutical Biology 4 : 536-541
Indian Fern J. 3 1 : 132-135 (2014) ISSN 0970-2741

HELMINTHOST
HELMINTHOSTA A CHYS ZEYLANICA POPULA TIONS NEED SERIOUS
POPULATIONS
ATTENTION : DISCLOSURES F R
ROO M UNPUBLISHED OBSERVATIONS
OBSERV
HIT KISHORE GOSWAMI*
Retired &Visiting Professor Botany and Genetics, Bhopal

(Received August 10, 2014, Accepted August 30, 2014)

ABSTRACT
This short note is the outcome of an enquiry from Professor S. C. Verma regarding
my earlier studies on the cytology of Helminthostachys zeylanica. Besides referring to the
published reports, I had to reveal my unpublished observations of 2n = 94 (root tip count)
from a Gorakhpur material, and explain that since I had no clear photographable preparation of
mitotic chromosomes this observation was not published. Because Professor Verma had known
about it, this has made him ask me to prepare a brief note as it might inspire someone having
access to Helminthostachys populations to investigate for cytotypic variations. I had also
observed 2n = 94 in a collection from Burdwan (W. Bengal). We believe that both diploid and
tetraploid forms might be co-existing in nature, awaiting to be well-documented and cyto-
taxonomically evaluated.
d s : Helminthostachys , chromosome number
K e y W o rrd

INTRODUCTION

The genus Helminthostachys Kaulfuss is one of the most fascinating and structurally
very complicated; the fertile spike region is non-comparable with any living species among
the pteridophytes (Goswami 1987). This is unfortunate that the monotypic genus has not
attracted many workers and one of the reasons could be that it is not as commonly found
as is Ophioglossum. But even then Helminthostachys zeylanica is found at many places in
India (Goswami 1987) among which reports from Trivandrum and Gorakhpur have been
commonly known (Bir & Verma 2010). Ninan (1956, 1958) observed meiotic as well as
mitotic chromosomes from the plants growing in a locality at Trivandrum (n = 94, 2n =
188), which essentially confirmed the report of n = 92-94 from Ceylon by Manton & Sledge
(1954). Later, this count of n = 94 was confirmed by Mahabale & Nair (1972) from the
plants collected from Trivandrum. Löve et al. (1977) assumed the basic number for the
genus as x = 47, rendering the species to be labeled as tetraploid. The present note intends
to reveal the unpublished observations on chromosome counts from root-tips in material
from Gorakhpur (U.P.) as well as from Burdwan / Bardhman / (W. Bengal), which support
the recent discovery by Singh et al. (2014, IFJ 31,this issue) of a diploid cytotype from
Gorakhpur area with n = 47, at meiosis.
OBSERVAT I O N S
RV

During 1970 we had initiated chromosome studies in Ophioglossaceae at Gwalior


primarily due to a find of very interestingly abnormal plants of Ophioglossum. Many field
trips were made and students helped in bringing plants of different species; we also got a
* 24, Kaushalnagar, P.O. Misrod,Bhopal (MP) 462047, India 91-755 2807950 Email : goswamihk@yahoo.com
Hit Kishore Goswami : Helminthostachys Zeylanica Populations Need Serious Attention 133

few plants of Helminthostachys zeylanica from Gorakhpur forest (exact locality was not
recorded at that time) along with fixed young roots and spikes. Squash preparations were
stained with 1.5% acetic-orcein but unfortunately, we could not find good preparations from
spikes-sporangia for meiosis of Helminthostahcys except a few feebly stained cells at
diakinesis which revealed 92 bivalents plus 3-4 univalents (Goswami & Khandelwal 1980).
Subsequent studies did not confirm this from other plants. Root tips from other collections,
brought fixed in the fixative, from Gorakhpur area showed only 94 chromosomes; the root
tip count of 2n = 94 was further confirmed by the potted plant maintained in Gwalior
(1974-1975), but each time, may be due to improper pretreatment or fixation, the staining
schedule did not give good photographable preparations. Obviously, after having reported
n = 92 II+3-4 Is, based on our preliminary study (Goswami & Khandelwal 1980) I had to
realize that the root tip count in Helminthostachys zeylanica , from different plants,
corresponded to the diploid number (2n = 94), while the commonly recorded number of
n = 94 and 2n = 188 from Trivandrum by Ninan (1956, 1958), and by Mahabale & Nair
(1972); (see also Bir & Verma 2010, for more reports) suggested the species to be tetraploid
based on the presumptive base number of x = 47 (Löve et al. 1977).
The observation of 2n = 94 from root tips was indeed very important requiring
photographic documentation, and calling for in-depth reinvestigation of populations around
Gorakhpur. However, because I had no photographic evidence of mitotic chromosomes
(showing 2n = 94) worthy of presentation to support such a startling, observation, and also
that I could not find an opportunity to visit Gorakhpur-area in person to collect the plants,
the observations were shelved from publication.
The issues raised by Professor Verma were serious: coexistence of two cytological
forms in Gorakhpur, and alternatively the presumptive occurrence of apogamous
reproduction in case the same plant (?, if at all) had shown both n = 94 as well as 2n =
94 (root tip)! How could my observations be explained? Meiotic study of a Gorakhpur
fixation had revealed n = c.94, (reported as 92II + 3-4I, Goswami & Khandelwal 1980),
that corroborated earlier reports from S. India (Trivandrum; see Bir & Verma 2010). But,
the spore size of Gorakhpur material, presumably from a different plant, was distinctly
different (19-24 µm) from earlier reports (32-38 µm) [see Goswami & Khandelwal 1980].
I had myself developed a doubt when root tip count revealed 2n = 94. This was a question
to be resolved because these observations suggested differences among different plants from
Gorakhpur. Unfortunately, exact locality of collections was not recorded by students, hence
this unusual count of 2n = 94 from root tips was neither published, nor even casually
mentioned. That, there might be plants within the same locality or at a different one in
Gorakhpur area, with variable chromosome numbers became a distinct possibility. At any
rate, the lack of photographic proof of the demonstration of 2n = 94 restrained me from
publishing it. Nevertheless, I did carry a belief that the monotypic genus
Helminthostachys (H. zeylanica ) may reveal chromosome variations among/within its
134 Indian Fern Journal Volume XXXI (2014)

populations as it is so common in Ophioglossum and Botrychium and several other ancient


polyploids (Goswami 2013).
Stud
Studyy of ma ter
mater ial fr
terial om Bur
from Burdd w an/ Bar dhman (W
Bardhman (W.. Beng al) :
Bengal)
Chances of studying Helminthostachs zeylanica came up during my short stay at the
University of Burdwan/Bardhman (West Bemngal) as a visiting Professor in 2011, as I was
lucky to get collections from two localities : (1) Baloor Ghat in north Bengal, and
(2) Village area, Devanandapur near Bandyl Rly station, 70 kms. away from Bardhman.
The plants of Helminthostachys from Baloor Ghat are growing in Botanical garden
of Botany Department at Bardhman. It was possible to take out a few roots (not too young,
though) and to attempt their cytology, following pre-treated with 8-hydroxyquinolene and
processed for mitotic squashes. Preparations were poorly stained; a few cells had yielded
nearly 94 chromosomes (2n = c. 94), but this observation again had to be shelved from
publication in the absence of properly photographable count. My observations from
Bardhman area raises the possibility of the occurrence of plants with 2n = 94, as also in
Gorakhpur area. The diploid form seems to be widespread. In fact we need a systematic
cytotaxonomic study on Helminthostachys zeylanica growing in Bengal, and also at higher
altitudes in Assam, besides the populations at Gorakhpur.
INFERENCE

The intention of this short note is to invite the attention of younger colleagues
having interest in chromosome studies and to search for variations in the genome of
Helminthostachys zeylaica. Population must be systematically explored; many individuals
from different areas in the same locality and if possible in different localities should be
studied both for mitotic and meiotic divisions. In passing, this must be mentioned here that
Sporne (1974) had also mentioned a haploid chromosome number of n = 47 for
Helminthostachys but this was not supported by any specific reference, and reference to
n = 92-94 by Manton & Sledge (1954) and n=94 & 2n=188 by Ninan (1956,1958) was not
mentioned. The most reliable meiotic chromosome count for this species accepted world over
has been n = 94, and the basic number suggested by Löve et al . (1977) as x = 47.
Obviously, the report of a diploid with n = 47 from Gorakhpur by Singh et al. (2014, IFJ
31-this issue) must encourage further studies on Helminthostachys zeylanica. Discovery of
cytotypes within fern species will always be a possibility.
There is one more important aspect of Helminthostachys ignored by most of us
interested in Ophioglossales. In Helminthostachys, the axis of the fertile spike bears
numerous sporangiophores in many rows, each bearing several sporangia and a few green
lobes at the tip (see, Fig.3, Goswami 1987; Goswami et al 2007). Such structures are only
comparable to growths found in the Carboniferous fern Botryopyeris. Investigators must also
study this unique morphological feat and record variables as observed in earlier studies
(Goswami & Khandelwal 1980, Goswami 1987) along with mitotic and meiotic
Hit Kishore Goswami : Helminthostachys Zeylanica Populations Need Serious Attention 135

chromosomes. Efforts will not go waste !!


ACKNOWLEDGEMENTS

I am deeply indebted to Professor S. C. Verma for exposing unpublished work by


accelerating my silently kept experience with Helminthostachys chromosomes. I am very
much grateful to Burdwan University Department of Botany for offering me a short term
visiting opportunity, to Professor Radhanath Mukhopadhyay and research scholars of that
department for their manifold help. Here, I also wish to record my grateful thanks to several
old students of 1970s at Science College, Gwalior who had helped me in various ways.
REFERENCES

BIR S S & VERMA S C 2010. Chromosome atlas of the Indian pteridophytes (1951-2009) Bishen Singh
Mahendra Pal Singh Dehradun pp 346
GOSWAMI H K & KHANDELWAL S 1980 Observations on Helminthostachys Kaulfuss (Ophioglossales)
Acta Bot Neerl 29 : 199-202
GOSWAMI H K, VERMA S C & SHARMA B D (eds) 2007 Biology of Pteridophytes I : Ophioglossum
Linnaeus Bionature Monograph Catholic Press Ranchi
GOSWAMI H K 1987 Ophioglossales I : An Overview Bionature 7 : 47-89
GOSWAMI H K 2013 Palaeoploidization and adaptation : An Evolutionary strategy among pteridophytes with
a reference to Ophioglossum L The Nucleus (Springer – Verlag) 5 66(2) : 69-80 (published on line—
14 th July 2013)
LÖVE A, LÖVE D & PICHI-SEMOLLI R E G 1977 Cytotaxonomical Atlas of the Pteridophyta J CRAMER
FI 9490 Vaduz
MAHABALE T S & NAIR L N 1972 Cytology of some species of the Ophioglossaceae in India Proc Indian
Acad Sciences , 55 Sec B 201-214
MANTON I & SLEDGE WA 1954 Observations on the cytology and taxonomy of the Pteridophyte flora of
Ceylon Phil Trans Roy Soc Londo (Biol Sci) 1238 : 127-185
NINAN C A 1956 Cytology of Ophioglossaceae Curr Sci 25 : 161-162
NINAN C A 1958 Studies on the cytology and phylogeny of the pterdophytes VI Observationon the
Ophioglossaceae Cytologia 2 3 : 291-300
SINGH S S, DOMINIC RAJKUMAR S, SRIVASTAVA S K & GAUTAM R P 2014 Discovery of diploid
cytotype of Helminthostachys zeylanica (L) Hook (Helminthostachyaceae - Pteridophyta) Indian Fern
J 31
SPORNE K R 1974 The Morphology of Pteridophytes Hutchinson & Co (publishers) Ltd London 3 rd Edition
(reprint 1974) pp 192
Indian Fern J. 3 1 : 136-138 (2014) ISSN 0970-2741

CYT OLOGY OF THREE ASPLENIUM SPECIES FR


CYTOLOGY OM UTT
FROM ARAKHAND
UTTARAKHAND
ANURITA SHARMA
Department of Botany, P.G.Govt. College For Girls-11, Chandigarh

(Received 20 August, 2014; Revised accepted 1 Sept., 2014)

ABSTRACT
Cytological studies of three Asplenium species : Asplenium nesii Christ., Asplenium
laciniatum D. Don subsp. kukkonenii (Reichst.) Fraser-Jenk. (2008) and Asplenium khullarii
Reichstein & Rasbach ex Fraser-Jenk. (2008) have been studied. All three are tetraploids
with n = 72. Photographs of the chromosome plates are given.
d s : Cytology, three Aspleniums .
K e y W o rrd

INTRODUCTION

The West Himalaya includes the region east of the River Indus (in Pakistan) and
west of River Sharda (on the border of west Nepal). Through her pioneering work entitled
“Problems of Cytology and Evolution in the Pteridophytes, ” Manton (1950), laid the
foundation for cytological studies. Later similar studies were taken up from different parts
of the world. Results on chromosome studies on Indian Pteridophytes from 1951-2009 have
been compiled by Bir and Verma (2010). Although the chromosome numbers of the three
species : Asplenium nesii, A. laciniatum subsp. kukkonenii and A. khullarii have been
recorded in Bir and Verma (2010), it is intended to publish the photographs of the
chromosomes of these three species since these have not been published so far. A very
exhaustive study on the Asplenium laciniatum complex was made by Fraser- Jenkins, Pangtey
and Khullar (2010). This report will make the work of Fraser-Jenkins et al. (2010) more
meaningful with the photographs of the chromosome preparations of these three species.
These ferns were collected from Dehra Dun and Uttarkashi districts of Uttarakhand.
Present collections have been deposited in the Herbarium, Panjab University, Chandigarh.
OBSERVAT I O N S
RV

Asplenium nesii Christ, Nu Giorn. Bot.It. n.s.4: 90 (1897).


Rhizome short, erect or ascending. Stipes short, green, lower half invariably black
with a violet tinge, scaly, rachis green, scantly scaly.Lamina pinnate or partly 2-pinnate (in
lower part), usually widest a little above the middle or lanceolate; pinnate 8-12 pairs beside
the lamina apex, which is more or less linear, incised on both sides to produce 3-5 blunt
teeth on each side directed upwards; lower 4-6 pairs of pinnae much reduced the lowest
much smaller rounded and flabellulate (about half as long as the largest pinnae), pinnae
slightly closer higher up but not crowded although at times touching the next pair. It is
somewhat intermediate between A. exiguum and A. laciniatum complex. Spores dark-brown
minutely spinulose. Chromosome no. n = 72 (Fig. A).

* E-mail : anurita_b@yahoo.com
Anurita Sharma : Cytology of Three Asplenium Species from Uttarakhand 137

F i g 1 . Chromosome plates of : A . Asplenium nesii n = 72(4x) B . Asplenium laciniatum subsp


kukkonenii n = 72 (4x); C . Asplenium khullarii n = 72 (4x). (all x 1500)

Rare, grows in crevices of dry rocks in shade between 2700-3300m altitude.


Present collection : Uttarakhand (Dehra Dun: Chakrata hills, Deoban);
Fraser- Jenkin et al. (2010) report it from Pakistan, W and C. Nepal, Tibet, China
Asplenium laciniatum D.Don subsp . kukkonenii (Reichstein) Fraser-Jenkins (2008),
Asplenium kukkonenii Reichstein (in Viane & Reicheistein 2004)
Rhizome erect; stipes black at the base, remainder green, thin, sparsely scaly; rachis
138 Indian Fern Journal Volume XXXI (2014)

green, glabrous. Lamina 2- pinnate, broadest about the middle, narrowly lanceolate,
herbaceous, glabrous; pinnae many, 8-10 pairs, ovate or broadly lanceolate, fourth pair or
pinnae usually the largest. Mature fronds are narrow with small pinnules which are not
prominently toothed and are not lobed. Spores dark-brown with a broad perine.
Chromosome no. n = 72 (Fig. B).
Grows on moist shaded rocks between 1600-2500 m altitude.
Pr esent collection : Uttarakhand (Uttarkashi, Yamunotri);
Present
Fraser-Jenkin et al. (2010) report it from: India (Jammu & Kashmir, Himachal
Pradesh, Uttarakhand, Arunachal Pradesh); Pakistan, W. Nepal, Bhutan, Tibet and China.
Asplenium khullarii Reichstein & Rasbach ex Fraser- Jenk. (2008)
Rhizome erect, apex scaly, scales dark- brown, lanceolate, apex long acuminate.
Stipes almost as long as the lamina, dark- brown, becoming greenish in the distal region
just below the lamina ; scaly at extreme base, rest is glabrous; rachis glabrous with or
without any vegetative buds. Lamina 2-3 pinnate, narrowly triangular – lanceolate, pinnae
up to 7 pairs, first and second pair of pinnae longest.In morphology it is intermediate
between Asplenium capillipes and A. laciniatum subsp.tenuicaule. The lamina is longer and
larger than in A. capillipes, the dissected segments are bi-ovate in shape but are larger with
slightly more acute teeth than in A. capillipes. Spores dark – brown with a broad perine.
Chromosome no. n = 72 (Fig. C).
A. khullarii grows on shaded and humid rocks found between altitude of 2700 –
3700m.
Pr esent collection : Uttarakhand (Yamunotri about 8 km from Hanumanchatti about
Present
50 m from bridge.
Fraser-Jenkin et al. (2010) report it from: India (Jammu & Kashmir, Uttarakhand),
Pakistan, China, Nepal and Bhutan.
ACKNOWLEDGEMENTS

I am most grateful to Prof. S.P. Khullar, Panjab University Chandigarh for his
valuable guidance and encouragement for the past 25 years.
REFERENCES

BIR S S &VERMA S C 2010 Chromosome Atlas of the Indian Pteridophytes (1951-2009) Bishen
Singh Mahendra Pal Singh Dehra Dun
FRASER-JENKINS C R 2008 Taxonomic Revision of three hundred Indian Subcontinental Pteridophytes
with a revised census list a new picture of fern taxonomy and nomenclature in the Indian
subcontinent pp 685 tt 255 Bishen Singh Mahendra Pal Singh Dehra Dun
F R A S E R - J E N K I N S C R , PA N G T E Y Y P S & K H U L L A R S P 2 0 1 0 Asplenium laciniatum D. Don
(Aspleniaceae) a critical complex and confused species in the Indian subcontinent Indian Fern
J 2 7 : 178-215
MANTON I 1950 Problems of cytology and evolution in the Pteridophyta -Cambridge
Indian Fern J. 3 1 : 139-142 (2014) ISSN 0970-2741

TECTARIA PUBER
TECTARIA ULA (DESV
PUBERULA .) C. CHR. (DR
(DESV.) Y OPTERID
(DRY OPTERIDAA CEAE :
PTERIDOPHYT A), A NEW RECORD FOR ASIA
PTERIDOPHYTA),
RAJU ANTONY1*, C. R. FRASER-JENKINS2**, N. MOHANAN3 AND CHERIYAN P. KOSHY4
1, 3 & 4
Jawaharlal Nehru Tropical Botanic Garden and Research Institute (JNTBGRI),
Palode, Thiruvananthapuram, Kerala, India.
2
Student Guest House, Thamel, P. O. Box no. 5555, Kathmandu, Nepal.

(Received 26 August, 2014; Accepted Sept. 11, 2014)

ABSTRACT
Tectaria puberula (Desv.) C. Chr. is reported for the first time from Asia, having
previously been known only from East Africa, Madagascar and the Mascarene Islands. A detailed
description and photographsare providedto assist initsidentification. It is one of a considerable
number of generally overlooked African phytogeographical connections that occur in India
mainly in the peninsula of C. and S. India.
K e y Wo rrd
d s : Tectaria puberula (Desv.) C. Chr., New record, Asia.

INTRODUCTION

Tectaria Cav. is a large fern genus well represented in tropical and subtropical
regions with most species growing terrestrially in rainforests. The estimated number of
species ranges from 150 (Tryon & Tryon 1982; Kramer 1990) to 210 (Holttum 1991).
Holttum (1991) recognized 105 species of Tectaria from Malesia and deduced that South
East Asiais itscenter of origin. Dixit (1984) listed 21 species and one variety from India
and this has been revised and amended to 22 species by Fraser-Jenkins (2008). The genus
is dryopteroid in its morphology and has usually been placed within Dryopteridaceae, but
in order to avoid paraphyly has recently been placed in a separate family Tectariaceae as a
result of molecular cladonomic studies (Smith et al. 2006, 2008). But as allied genera such
as Ctenitis and even Pleocnemia have been retained in Dryopteridaceae, and morphological
differences are small and insufficient we maintain Tectaria within Dryopteridaceae.
During the course of pteridophytic exploration of the Western Ghats, the first
named author collected an interesting specimen of Tectariagrowing terrestrially along an
earth bank in evergreen forest at Moozhiyar, Pathanamthittta Dist., Kerala, South India. It
did not correspond with Indian species. The specimen was therefore sent to the second
author, who after some difficulty, as it also did not appear to be any Asian species,
identified it as Tectariapuberula (Lam.) Baker, following study and careful comparison with
theTectaria collections at K and BM. The discovery of this species from South India has
added an interesting African species to the Asian fern-floraas well as to India, as it was
previously known only from East Africa (Kenya) and Madagascar.
The African element in the peninsular Indian fern-flora is often overlooked, but is
a small but quite well represented group. Some African species have even been redescribed
as if new species in India due to lack of familiarity with non-Asian ferns. A few more
* E-mail : rajuantonytbgri@rediffmail.com ** E-mail : chrisophilus@yahoo.co.uk
140 Indian Fern Journal Volume XXXI (2014)

PLATE 1. Tectaria puberula (Desv.) C. Chr. A. Habit, B . Rhizome with stipes, C. Frond, D . Terminal
PLATE
pinna, E. Costa showing setose-hairs, F. Portion of pinnae showing sori.
Raju Antony et al. : Tectaria Puberula (Desv.) C. Chr. (Dryopteridaceae : Pteridophyta) 141

examples of African ferns naturally occurring in peninsular India are: Selaginella bryopteris
(L.) Baker (syn.: S. imbricate (Forssk.) Spring); Osmunda huegeliana C. Presl (very closely
related to and probably a synonym of the African O. obtusifolia Willd. ex Kaulf., or O.
capensis C. Presl, non L.); Aneimia schimperiana C. Presl subsp. wightiana (Gardner)
Fraser-Jenk. (a close vicariant of the African subsp. schimperiana); Aleuritopteris scioana
(Chiov.) Fraser-Jenk.; Pellaea longipilosa Bonap. (syn.: P. malabarica Geev. ex Madhus. &
Jyothi); Asplenium aethiopicum (Burm.f.) Bech. Subsp .aethiopicum; Asplenium falcatum
Lam.; Asplenium hymenophylloides Fée and several other species.
That this species is of natural occurrence in India is indicated by its occurrence in
natural, undisturbed forest far from human habitation and in its never having been cultivated
in the well known gardens in India, unlike several other adventive species in India that were
introduced and escaped (see Fraser-Jenkins 2008: 359-366). It is also oflimited occurrence
in Africa and is an uncommon and little known species.
A detailed description and photographs are provided here to help makeits occurrence
in S. India known and to assist botanists to detect and identify it if it should turn up in
other localities.
DESCRIPTION

Tectar ia puber
ectaria ula (Desv.) C.Chr., Dansk Bot. Ark. 7: 67. 1932.
puberula
Polypodium puberulum Desv., Mém. Soc. Linn. Paris 6: 245. 1827, nom. nov. for P.
triphyllum Desv.
Polypodium triphyllum Desv., Mag. Nat. Ges. Nat. Freunde Berlin 5: 315. 1811, non
Jacq. 1788.
Type : from La Réunion (Mascarene Islands).
Plants terrestrial. Rhizome thick, woody, long creeping, densely scaly at apex; scales
upto 5 x 1 mm, lanceolate, light brown with dark brown cells present in the central part of
the scale towards the apex, cordate at base, attenuate at apex, margin with small finger like
outgrowths.Fronds 36-69 x 20-44 cm, widely spaced; stipe in fertile frond longer than the
sterile frond, purplish brown, densely scaly at base, setose-hairy throughout; stipe scales
more or less similar to rhizome scales; lamina 20-45 x 20-44 cm, deltate, pinnate to
bipinnatifid, herbaceous, underside (abaxial surface) rather densely setose-hairy, upperside
(adaxial surface) sparsely hairy; rachis densely covered with setose-hairs; pinnae upto 5
pairs, pale green, opposite, basalmost pinnae much larger, stalked, lobed at margins with
larger and deeper lobes on their basiscopic side than on their basiscopic side; middle pinnae
shortly stalked or sessile, oblong-lanceolate, unequal at base, acuminate at apex, margin
lobed or undulate; terminal pinna dissimilar to lateral pinnae, deltate, pinnatifid, crenate to
decurrent at its base, acuminate at apex; veins distinct, anastomosing with or without
included veinlets in areoles. Sori small, round in irregular rows, indusiate; indusia small,
brown, reniform, deciduous.
142 Indian Fern Journal Volume XXXI (2014)

DISTRIBUTION

Africa : The islands of Tanzania (Zanzibar, Pemba), Kenya, the Comoro Islands,
Madagascar and the Mascarene Islands of La Réunion and Mauritius (the erstwhile “Île
Bourbon” and “Île de France” respectively) (see Roux 2009); S. India: Kerala.
Ecolo
Ecologg y : Very rare species, terrestrial on earth banks in evergreen forest.
Specimen examined : India, Kerala, Pathanamthitta District, Moozhiyar forest, 1010
m alt., Raju Antony 69498, 18 March 2010 (TBGT); ibid. Raju Antony 70191,9 Oct.
2013(TBGT).
Although Roux (2009) treated the similar Asian species, T. subtriphylla(Hook.
&Arn.)Copel.as a synonym of T. puberula, the two are distinct species.
ACKNOWLEDGEMENTS

The authors are thankful to Dr. P.G. Latha, Director, Jawaharlal Nehru Tropical
Botanic Garden and Research Institute (JNTBGRI), Palode, Thiruvananthapuram, Kerala,
India, for encouragement and facilities provided for the study. Thanks are also due to Mr.
S. Suresh, Technical Officer, Conservation Biology Division, JNTBGRI for the preparation
of photo plates and Mr. Harilal Kumar, Gardener, JNTBGRI, for his assistance with our
field study.
REFERENCES

DIXIT RD 1984 A census of the Indian Pter idophyte s pp 177 Botanical Survey of India, How rah
H O LT T U M R E 1 9 9 1 Flora Malesiana , s e ri e s I I , P t e r i d o p hy t a , 2 ( 1 ) , Te c t a r i a gr o u p R i j k s h e r b a r i u m /
Hortus Botanicus, Leiden, 132 pp
FRASER-JENKINS C R 2008 Taxonomic Revision of Three Hundred Indian Subcontinental Pteridophytes
with a Revised Census-List pp 685 Bishen Singh Mahendra Pal Singh Dehra Dun
KRAMER K U 1990 D r y o p t e r i d a c e a e I n K U B I T Z I K ( e d ) T h e F a m i l i e s a n d G e n e r a o f Va s c u l a r
Plants , 1 , Pteridophytes and Gymnosperms 101-144 Spr inger Verla g, Berlin Heidelberg
RO U X J P 2 0 0 9 S y n o p s i s o f t h e Ly c o p o d i o p h y t a a n d P t e r i d o p h y t a o f A f r i c a , M a d a g a s c a r a n d
neighbouring islands Strelitzia 2 3 : 1-296 South African National Biodiversity Institute Pretoria
S M I T H A R , P RY E R K M , S C H U E T T P E L Z E , KO R A L L P, S C H N E I D E R H & WO L F F P G 2 0 0 6 A
classification f or extant fer ns Taxon 5 5 : 705-731
SMITH A R, PRYER K M, SCHUETTPELZ E, KORALL P, SCHNEIDER H & WOLFF P G 2008 Fer n
Classifica tion In KNAKER, T A & HAUFLER CH (eds) The Biology and Evolution of F ern s
and Lycophytes 419-469 Cambr idg e University Press Cambr idge
T RYON R M & T RYON A F 1982 Fer ns and allied plants with special refer ence to Tropical America
pp 857 Spr ing er Verla g, Ber lin, Heidelberg, New York
Indian Fern J. 3 1 : 143-151 (2014) ISSN 0970-2741

FERNS OF THE AABSHAR FOREST AREA (KAND


(KANDAA GHAT,
GHAT
DIST
DIST.. SOLAN), HIMA CHAL PRADESH, WEST HIMALA
HIMACHAL HIMALAYYA .
ANURITA SHARMA*
Department of Botany, P. G. Govt. College For Girls-11, Chandigarh

(Received August 31, 2014; Accepted Sept. 9, 2014)

ABSTRACT
The Aabshar forest area near Kandaghat, Dist. Solan Himachal Pradesh (West
Himalaya) was visited several times for collection of ferns since there is no comprehensive
account of the ferns of this area. The forest here is basically composed of oak ( Quercus
leucotrichophora A. Camus). Forty two fern species have been collected. These are listed
alphabetically in the list provided.
K e y W o rrd
d s : Aabshar, forty two fern species.

INTRODUCTION

The Aabshar forest area (Plate I Fig 1) is a beautiful place for fern collection. It is
located on the Kalka-Shimla highway in Solan district, Himachal Pradesh, two kms before
Kandaghat and 12 kms beyond Solan towards Shimla with a maximum altitude of around
2,000 m. The highest peak of this area is Mount Karol also known as Karol Tibba. The
forest here is basically composed of Oak (Quercus leucotrichophora A. Camus) which form
almost pure stretches of dense and evergreen forests between 1,400-2,000m. Some common
shrubs and trees here are - Berberis asiatica (Roxb.), D.C., Rubus ellipticus J. Smith,
Rhododendron arboretum Sm. Aesculus indica Colebr. ex Camb. and Emblica officinalis
Gaertn. etc. Gymnosperms here are Pinus roxburghii Sarg., and Cedrus deodara (Roxb. ex
Lamb.) D. Don (Plate I Fig 2). Fruit trees like Malus domestica Borkh., nom. cons. (apple),
Prunus domestica subsp. insititia (L.) Schneid. (plum), Prunus persica Stokes (peach), Pyrus
pyrifolia var. culta Nakai. (pear), Punica granatum L. (pomegranate) and Juglans regia L.
(walnut) etc. are also cultivated here.
Water here is plenty with a waterfall (Plate I Fig.3) and a number of connecting
waterways. In earlier times this place was popularly known by the locals as “Dedh Gharat”
(one and a half water mill). This forest is quite rich in ferns due to water being plenty.
The fern flora here is mainly confined to the elevations above 1,500m. It shows a great
deal of diversity and variation. Ferns grow here on the forest floor, on forest slopes, banks
of waterways, in open or in shaded places etc . Epiphytic (PLATE II Figs 4 & 5) and
lithophytic ferns clothe tree trunks and branches, boulders and walls. There are a number
of hardy terrestrial taxa which grow throughout the year.
No comprehensive account of the ferns of this forest is available. It was therefore
thought proper to survey this forest and collect ferns from here. A few reports of ferns
from this area find mention in Khullar (1994, 2000), and Khullar et al. (2008). A
comprehensive and a thorough survey of this forest for ferns resulted in the collection of
* E-mail : anurita_b@yahoo.com
144 Indian Fern Journal Volume XXXI (2014)

42 fern species. In the list provided these names have been arranged alphabetically for the
sake of convenience. All collections are deposited in the Herbarium, Panjab University,
Chandigarh. The PAN numbers are provided in the list.
Generally the nomenclature as suggested by Fraser-Jenkins (2008) has been followed with
some exceptions. Members of Thelypteridaceae have been placed under a single genus
Thelypteris (as suggested in Khullar 2000; Fraser- Jenkins 2008). The Cheilanthoid ferns
are placed in the genus Cheilanthes rather than shifting them under Aleuritopteris as
suggested by Fraser-Jenkins. The name Onychium contiguum (Wall. ex Hope) has been
retained and Dryopteris nigropaleacea Fraser-Jenk. has been named as Dryopteris juxtaposita
Christ., subsp nigropaleacea (Fraser- Jenk.) Khullar (Khullar et al. 2009) rather than
following Fraser-Jenkins in treating it as an independent species. Asplenium yunnanense
Franch. Is retained as a species rather than reduce it as a subspecies (Asplenium exiguum
Bedd. subsp. yunnanense (Franch.) Fraser- Jenk., Pangtey & Khullar (2010)
OBSER
RVVAT I O N S

List of ffer
er ns of the Aa
erns bshar ffor
Aabshar or est.
orest.
Adiantum capillus-veneris L., Sp. Pl. 2 : 1095 (1753).
Common in wet places along banks of streamlets and walls up to 1600 m. PAN
8980.
Adiantum edgeworthii Hook., Sp. 2 : 14. t. 81B (1851).
Grows on shaded rocks between 1,500-1,850 m. PAN 8981.
Adiantum incisum Forssk., Fl. Aeg. Ar. : 187 (1775).
A common low altitude fern that grows in open dry, exposed slopes between 900-
1,200 m. PAN 8983. (PLATE IV, Fig.13).
Adiantum philippense L., Sp. Pl. 2: 1094 (1753).
Found on shaded slopes around 1,500 m. PAN 8984.
Adiantum vven
en ustum D.Don, Prodr. Fl.. Nepal: 17 (1825) subsp. ven
enustum ustum
enustum
Grows from 1,800 m and above. PAN 8982.
Araiostegia pseudocystopteris (Kunze) Copeland, Philip, J. Sci. 34 : 241 (1927).
An epiphyte grows around 1,800 m and above. PAN 8993
Asplenium dalhousiae Hook., Icon. Pl.: t. 105 (1837). (PLATE IV, Fig 12).
A very common fern on exposed rocks in open between 1,500-2,000 m. PAN 8985.
Asplenium yunnanense Franch. Bull. Soc Bot France 32 32:28 (1985)
Occasional on humid rocks in shade, above 1,700 m. altitude. PAN 8987. (PLATE
III. Fig 7).
Asplenium lacinia tum D. Don, Prodr. Fl. Nepal : 8 (1825) subsp. lacinia
laciniatum tum.
laciniatum.
Grows in humid places in the forest between an altitude of 1,600 - 2,000 m. PAN
8986.
Anurita Sharma : Ferns of the Aabshar Forest Area (Kandaghat, Dist. Solan) 145

PLATE I. F
PLATE igs. 1-2. The Aabshar Forest. F i gg.. 3. Waterfall at Aabshar.
Figs.
146 Indian Fern Journal Volume XXXI (2014)

PLATE II. F
AT Fii g s . 4 - 5 . Epiphytes covering tree trunks. F i gg.. 6 . Sphenomeris chinensis (L.) Maxon
Anurita Sharma : Ferns of the Aabshar Forest Area (Kandaghat, Dist. Solan) 147

PLA TE III. F
ATE Fii gg.. 7 . Asplenium yunnanense Franch F i gg.. 8 . Pyrossia porosa (C Presl) Hovenkamp.
F i gg.. 9 . Polystichum obliquum (D. Don) T. Moore
148 Indian Fern Journal Volume XXXI (2014)

PLAATTE IV V.. F i g . 1 0 . Cheilanthes albomarginat a Clarke. F i g . 1 1 . Cheilanthes bicolor (Roxb.)


Fraser-Jenk. F i g . 1 2 . Asplenium dalhousiae Hook. F i g . 1 3 . Adiantum incisum Forssk.
Anurita Sharma : Ferns of the Aabshar Forest Area (Kandaghat, Dist. Solan) 149

Ath yr
Athyr ium pectina
yrium tum (Wall. ex Mett.) Moore, Index Fil. : 152 (1859).
pectinatum
A comparatively low altitude fern found between 1,500- 1,700 m. altitude. PAN
8988
Athyrium schimperi Moug. ex Fee, Mem. Foug. 5 Gen Fil. : 187 (1850-52).
A common fern of the forest floor and forest slopes, found between the altitude
range of 1,200- 2,000 m. PAN 8989.
Cheilanthes albomarginata Clarke, Trans. Linn. Soc. Lond. 2. Bot. 1 : 456 (1880).
Commonly grows on slopes around 1,400-2,000m. PAN 9014 (PLATE IV. Fig. 10.)
Cheilanthes anceps Blanf. Simla, Nat. Hist. Soc. Leaflet 25th June (1888).
Grows on dry slopes between 1,000- 1,800m. PAN 9015.
Cheilanthes bicolor (Roxb.) Griff. ex Fraser-Jenk. Pak Syst 5 (1-20 94 (1991 [1992].
Grows on dry exposed slopes. PAN 9016 (PLATE IV. Fig. 11).
Cheilanthes br breevifr ons (Khullar) Khullar, Indian Fern J. 1: 90 (1984).
vifrons
Quite common on dry exposed rocks between 1,500-1,800m. PAN 9017.
Cyr tomium car
Cyrtomium caryy otideum (Wall. ex Hook. et Grev.) Presl, Tent. Pterid. : 86 (1836).
This fern was found growing on the banks of the rivulets in thick forest from 1,500
m altitude upwards. PAN 8994.
Diplazium maxim
maximum um (D.Don) C.Chr., Index Fil. 1 :235 (1905).
Grows besides streamlets and ravines around 1,200-2,000 m. altitude. PAN 8990.
Dryopteris caroli-hopei Fraser-Jenk., Bull. Brit. Mus. Nat. Hist. Bot. 14 14: 194 (1986).
A fern of the forest floor from 1,500-2,000 m. altitude. PAN 8995.
Dr
Dryy opter is juxta
opteris posita Christ. Subsp. nig
juxtaposita nigrr opaleacea (Fraser– Jenk.) Khullar, Indian Fern J
26 : 91 (2009).
Found growing on the forest floor around 2,000m. altitude. PAN 8996.
Hypodema
Hypodematium tium cr ena
crena tum (Forsk.) Kuhn, in Deck., subsp. lo
enatum y alii Fraser-Jenk. & Khullar,
loy
Bot. Helv. 102 : 146 (1992).
Rare, grows in crevices of dry exposed rocks at 1,500m and above. PAN 9000.
Hypolepis polypodiodes (Blume) Hook. Sp. Fil. 2 : 64 (1852).
Hypolepis punctata (Thub.) Mett., ex Kuhn, Fil. Afr. : 120 (1868).
Commonly grows around 1,500m altitude on the forest floor in moist places or
along open road-sides. PAN 9001.
Le pisor
Lepisor us scolopendr
pisorus scolopendrium ium (Ham. ex D.Don) Mehra & Bir, Res. Bull. Panjab Univ. (n.s.)
15 : 168 (1964).
A common epiphyte on trees or as a lithophyte around an altitude of 1,500m and
above. PAN 9005.
Lepisorus nudus (Hook.) Ching, Bull. Fan Mem. Inst. Biol. (Bot.) 4 : 83 (1933).
Another common epiphyte around 1,800 m altitude. PAN7095
Onychium contiguum Hope, J. Bombay Nat. Hist. Soc. 13 : 444 (1901).
Onychium cryptogrammoides Christ.
150 Indian Fern Journal Volume XXXI (2014)

Grows on the forest floor and open places, forming colonies above 1,700 m. PAN
8992.
Ophioglossum reticulatum L., Sp. Pl.. 2 : 1063 (1763).
Rare, on exposed slopes around 1,800m.altitude. PAN 9003.
P ol ypodiodes amoena (Wall. ex Mett) Ching, Acta Phytotax. Sinica 16(4
olypodiodes 16(4) : 27 (1978).
Occasionally grows on rocks at an altitude of about 1,700m and above. PAN 9007.
P ol ypodiodes micr
olypodiodes orhiz
microrhiz oma (Clarke ex Baker) Ching, Acta Phytotax. Sinica 16 (4) : 27
orhizoma
(1978).
Grows around 1,600m.altitude. PAN 9008.
Polystichum discretum (D. Don) J. Smith, J. Bot.. 3 : 413 (1841).
Grows around elevations of 1,500 in wet places in the forest. PAN 8997.
Polystichum obliquum (D. Don) Moore, Index Fil. : 87 (1858).
In wet places along streamlets at altitudes of 1,500m upwards. PAN 8998. (PLATE
III. Fig.9)
Polystichum squarrosum (D. Don) Fee, Gen. Fil. : 278 (1852).
A fern of the forest floor upwards of 1,800 m altitude. PAN 8999.
Pteris cretica L., Mant. Pl.: 130 (1767).
A Fern of the forest floor commonly growing from altitudes of 1,800m upwards.
PAN 9010.
Pter
Pterisis pseudoquadr
pseudoquadriaur iaur ita Khullar, Illustrared Fern Fl. West Him.. 1 : 272 (1994).
iaurita
Common fern of the forest floor from 1,800m upwards. PAN 9011.
Pter
Pterisis stenoph ylla Wall. ex Hook. et Grev., Ic. Fil. : t. 130 (1829).
stenophylla
A common fern of the forest floor from 1,400-1,600m. PAN 9012.
Pteris vittata L., Sp. Pl.. 2 : 1074 (1753) subsp. vittata
Grows near wet places 1,500m and above. PAN 9013.
Pyrrosia porosa Hovenkemp, Blumea 30 : 208 (1984).
A rare, epiphyte on oak trees here between 1,500-1,800m. PAN 9009 (PLATE III
Fig. 8.)
Sphenomer
Sphenomeris is cchinensis
hinensis (L.) Maxon, J. Wash. Acad. Sci. 3 : 144 (1913).
Very rare grows in cervices of dry rocks from 1,500-2,000 m. PAN 9002. PLATE
II Fig.6)
T hel ypter
helypter is denta
ypteris dentata ta (Forssk.) E.St. John., Amer. Fern J. 26 : 272 (1936).
Very common in wet places along the ravines found up to 2000m. PAN 9018.
T hel ypter
helypter is er
ypteris ubescens (Wall ex Hook.) Ching, Bull. Fan Mem. Inst. Biol. (Bot) 6 : 293
erubescens
(1936).
Grows near waterfalls and ravines at the altitude range of 900-2,000m..PAN 9020.
T hel ypter
helypter is pa
ypteris pilio (Hope) K. Iwatsuki, Mem. Coll. Sci. Univ. Kyoto B 31 : 175 (1965).
papilio
Grows in swampy wet places around around 1,600m. altitude PAN 9019.
Thelypteris penangianum (Hook.) C. Reed, Phytologia 17 : 303 (1968).
Grows in wet areas up to 1,600m. PAN 9021.
Anurita Sharma : Ferns of the Aabshar Forest Area (Kandaghat, Dist. Solan) 151

T hel ypter
helypter is ppyr
ypteris yr rhorhac
yrrhorhac his (Kunze) C M Kuo, Taiwania 30
rhorhachis 30: 60 (1985) subsp. laterepens
(Trotter).Fraser-Jenk. Tax review Indian Subcontinental Pteridophytes 199 (2008).
Rare, grows in the forest along the river banks, or in open wet places at altitudes
between 1,600-2,000m. PAN 9022.
Wood
oodw w ar dia unig
ardia emma
unigemma
emmatata (Makino) Nakai, Bot. Mag. Tokyo 39
39: 103 (1925).
Grows in wet places along ravines and waterfalls from 1,500-2,000 m.altutude.
PAN 8991.
ACKNOWLEDGEMENTS

Sincere thanks to Prof. S. P. Khullar, Department of Botany, Panjab University


Chandigarh for his valuable suggestions concerning the latest nomenclature and correcting
of the manuscript.
REFERENCES

FRASER-JENKINS C R 2008 Ta x o n o m i c R ev i s i o n o f t h r e e h u n d r e d I n d i a n S u b c o n t i n e n t a l
Pteridophytes with a revised census list, a new picture of fern taxonomy and nomenclature
in the Indian subcontinent Bishen Singh Mahendra Pal Singh, Dehra Dun
F R A S E R - J E N K I N S C R , PA N G T E Y Y P S & K H U L L A R S P 2 0 1 0 Asplenium laciniatum D. Don
( A s p l e n i a c e a e ) , A c r i t i c a l C o m p l ex a n d C o n f u s e d S p e c i e s i n t h e I n d i a n s u b c o n t i n e n t I n d i a n
Fern J 2 7 : 178-252
KHULLAR S P 1994, 2000 An Illustrated Fer n Flora of the West Himalaya Vols I (1994) & I I (2000)
International Book Distributors, Dehra Dun (India) pp 506 pp 538
KHULLAR S P, CHADHA J, BAGHLA A & VERMA S 2009 Annotated Inventory of the Pteridophytes
of District Sirmaur (Himachal Pr adesh), West Himalay a Indian Fern J 2 6 : 79-106
KHULLAR S P, SHARMA S & PRASHER I B 2008 Diversity in the pteridophytes of Kangr a district
(Himachal Pradesh) Proc Nationl Acad Sci India 7 8 (sect B, 1) : 1-36
Indian Fern J. 3 1 : 152-153 (2014) ISSN 0970-2741

SOME COMMENTS ON THE P APER "N


PAPER N ATURAL APOSPOR
APOSPORYY IN PTERIS
ARGYRAEA T. MOORE FR OM SOUTH INDIA
FROM INDIA".. BY RAJU A N T ONY
ONY,, S M
TONY
SHAREEF AND N MOHAN AN IN INDIAN FERN JOURN
MOHANAN A L 29 : 148-152
JOURNA
NO 1. COMMENTS BY :

HARVINDER K. CHEEMA*
Department of Botany, Panjab University, Chandigarh -160014

My view point on the observations of their results is as follows :-


The authors have referred aposporous gametophytes to the ones produced on the
abaxial surface of the pinnules. Although they have mentioned that microscopical
observations were carried out but no record or photographs of the same have been shown.
What is the evidence that the unusual outgrowths on the frond as referred to Fig. 1A are
the gametophytes? The filamentous structures are dichotomously branched and these may
be gametophytic or sporophytic or intermediate structures. What are the criteria to call them
as aposporous gametophytes? Since they are arising from the junction of costa and costule
at the tip of the lateral veins and from the costules tip, they may be the outgrowths arising
by the regeneration of the bud and thus can be the case of induction and regeneration of
vegetative bud rather than apospory. It is further supported in Fig. 1B where the
regeneration of complete sporophytes is noticed from the bud and not from the aposporous
gametophyte. Such results have earlier been reported by Mehra and Sulklyan (1969), Loyal
and Chopra (1977), Cheema (1979,1984,1997) and Cheema and Sharma (1994) . Without
looking into the gametophytic characters and cytological confirmation, it is not appropriate
to refer as "aposporous gametophyte". The fact seems to be that it is the case of
regeneration of vegetative bud leading to the development of sporophyte (fig B) reaching
to maturity to bear the sori (Fig. C) . Development of vascular bundles (Fig. D) is a
sporophytic character and hence cannot be an aposporous gametophyte. To state "the
presence of both sori and aposporous gametophytes in the same frond is a rare phenomena"
is not logical as it is not the case of apospory.
REFERENCES

CHEEMA H K 1979 R e g e n e r a t i o n o f r h i z o m e s e g m e n t s o f Regnelidium diphyllum L i n n . i n a s c e p t i c


culture. Curr Sci 4 8 : 640
CHEEMA H K 1984 In vitro studies on reproductive biology and regeneration of the fern
Ceratopteris pterdioides (Hook) Hieron Res Bull Panjab Univ (Sci) 3 5 : 13-17
CHEEMA H K 1997 F e r n s a s a n ex c e l l e n t e x p e r i m e n t a l s y s t e m f o r M o r p h o g e n e s i s : A n O v e r v i ew
Indian Fern J 1 4 : 1-9
CHEEMA H K and SHARMA M B 1994 Induction of multiple shoots from adventitious buds and leaf
callus in Ceratopteris thalictroides Indian Fern J 1 1 : 63-67

* Correspondence Address: 1625, Sector 44 B, Chandigarh, 160047


Harvinder K Cheema : Some Comments on the paper "Natural Apospory in Pteris Argyraea" 153

L OYA L D S a n d C H O P R A H K 1 9 7 7 I n v i t r o l i f e cy cl e , r eg e n e r at i o n a n d a p o s p o r y i n C e r at o p t e ris
pteridoides (Hook.) Heiron Curr Sci 4 6 8 : 89-91
M E H R A P N a n d S U L K LYA N D S 1 9 6 9 I n v i t r o s t u d i e s o n , a p o g a m y, a p o s p o r y a n d c o n t r o l l e d
d i ff e r e n t i a t i o n o f r h i z o m e s e g m e n t s o f t h e f e r n A m p e l o p t e r i s p r o l i f e r a ( R e t z ) C o p e l B o t J
Linn Soc 6 2 : 431-443

No 2. REPLY BY RAJU A N T
REPLY TOONY
Y,, one of the author
authorss :

Dear Dr. Harvinder Cheema,


I am Dr. Raju Antony, working in Tropical Botanic Garden and Research Institute,
Trivandrum. I am writing this mail as per the instruction of Verma Sir. I have been working
on conservation and systematic studies on pteridophytes of Western Ghats for last 22 years
and published more than 25 research papers in both international and national journals and
developed a fernery which holds 250 species of ferns and fern allies, one of the largest
collections in India. I prepared the above research paper in similar line with Natural
apospory in Arachniodes aristata (Forst. f.) Tindale, from South India, published by
Manickam and Irudayaraj in IFJ and Incidence of apospory in Pteris confusa T. G. Walker,
published in Phytomorphology by Manickam et al. At the time of revising this paper the
Editor, IFJ, had asked me to show by sectioning the connection between the branched
gametophyte and parent tissue. So we did the anatomical studies. It is a basic thing that
gametophytes having no vascular bundles so we put aposporous gametophytes in apostrophe
(aposporous 'gametophytes'). After publishing this paper, Fraser-Jenkins who is in good
touch with me, wrote to me that earlier reports of apospory in Arachniodes aristata and
Pteris confusa including Pteris argyraea are due to the infection of Taphrina funcus. He has
also asked me to prepare a research paper on Taphrina infection in ferns. We are thinking
to prepare such a paper and if you wish you can also kindly contribute and we can jointly
publish the same in IFJ.
Yours faithfully

Dr. Raju Antony


Ferns Section TBGRI.
Indian Fern J. 3 1 : 154-161 (2014) ISSN 0970-2741

IS HUPERZIA HAMIL
HAMILT
T ONII (SPRENG .) TREVIS
(SPRENG.) TREVIS.. A HIMALA
HIMALAYYAN ENDEMIC?
AN EMPIRICAL EVALU
EVA ATION USING SPECIES DISTRIB
UA UTION MODELING
DISTRIBUTION
NAWAL SHRESTHA1, 2* and XIAN-CHUN ZHANG1**
1
State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany,
Chinese Academy of Sciences, Beijing 100093, China
2
University of Chinese Academy of Sciences, Beijing 100049, China

(Received September 12, 2014; Revised Accepted)

ABSTRACT
The distribution range of Huperzia hamiltonii (Spreng.) Trevis. has been disputed
among taxonomists since its discovery. In the past it has been thought to be widely distributed
in the Himalaya as well as in parts of China and S.E. Asia, while some recent taxonomists
think it is only a Sino-Himalayan species. Whether this species occurs only in the Himalayan
region or is distributed throughout the Indian subcontinent and Indochina has not been fully
understood, though in Indian literature it has nearly always been reported from both the
Himalaya and peninsular India. There exist inconsistencies among taxonomists regarding its
distribution range to date. The present work is therefore an attempt to evaluate the distribution
of H. hamiltonii using evidence from field-based observations, herbarium specimens and species
distribution modeling. Based on our results, H. hamiltonii is distributed in the western, central
and eastern Indo-Himalaya of India, Nepal and Bhutan and in the Chinese Himalaya, as well
as in south-west India, Sri Lanka, Myanmar, Vietnam and northern Thailand.
d s : Ecological modeling, Himalaya, Huperzia, Maxent.
K e y Wo rrd

INTRODUCTION

Huperzia hamiltonii (Spreng.) Trevis. is an epiphytic club-moss first discovered in


the Kathmandu valley of central Nepal by the Scottish botanist, zoologist and surveyor, Dr.
Francis Buchanan, later Hamilton, in 1805. Since its discovery, there have been many
reports of this species from the Indo-Himalaya as well as from S.W., S., and E. China,
Cambodia, Japan, Laos, Malaysia, Thailand and Vietnam (Tagawa & Iwatsuki, 1979; Dixit,
1984; Iwatsuki, 1988; Boonkerd et al., 2008; Khwaiphan & Boonkerd, 2008; Zhang &
Iwatsuki, 2013; Fraser-Jenkins, 2008, 2010, Fraser-Jenkins & Gandhi, 2015, in prep.). Dixit
(1984, 1987) and Chandra et al. (2008) followed previous Indian literature and reported its
distribution in southern India, Sri Lanka, Myanmar, Indochina and the Malesian islands.
Contrary to previous literature, Fraser-Jenkins (2008) separated the south Indian plant as H.
aloifolia (Wall. ex Grev. & Hook.) Trevis., but has since united them (Fraser-Jenkins &
Gandhi 2015, in prep.). Zhang & Iwatsuki (2013), in the Flora of China, considered H.
hamiltonii to be only a Sino-Himalayan species and believed that all other distributions are
based on misidentification. Thus whether H. hamiltonii is only a Sino-Himalayan species as
treated by Zhang & Iwatsuki (2013), or whether it occurs throughout the Indian
subcontinent and Indochina remains unclear.
Discrepancies in distribution records result partly from its resemblance to other

* Email : nawalshrestha@gmail.com ** Email : zhangxc@ibcas.ac.cn


Nawal Shrestha & Xian-Chun Zhang : Is Huperzia hamiltonii (Spreng.) Trevis. A Himalayan Endemic? 155

closely related taxa. For example, specimens of two Chinese species H. fordii (Baker) R.D.
Dixit and H. henryi (Baker) Holub have often been mistakenly identified as H. hamiltonii.
However the leaves of H. hamiltonii are elliptic, glossy, usually angled upward and have an
obtuse apex (FIGURE 1), while H. fordii has elliptic-lanceolate, non-lustrous and
amplexicaul leaves and H. henryi has elliptic, non-lustrous leaves which are attached at right
angles to the stem (Zhang & Iwatsuki, 2013).
The present work is an attempt to evaluate the distribution range of H. hamiltonii
using the GIS based modeling tool in conjunction with field based data, specimen records
and critical taxonomic study. The study has been conducted to fulfill the following
objectives: (1) does H. hamiltonii occur only in the Himalayan region? (2) what is its actual
distribution range? and (3) what is its potential range of occurrence?

iguree 1. Morphology of Huperzia hamiltonii . A. Habit of the plant; B . Leaf; C. Sporangium


F igur
156
Indian Fern Journal Volume XXXI (2014)

F i g u rree 2 . Potential distribution rang e of Huperzia hamiltonii. AUC value of this model is 0.868 ± 0.070. Black dots represent
occurrence records. Warmer color represents higher probability.
Nawal Shrestha & Xian-Chun Zhang : Is Huperzia hamiltonii (Spreng.) Trevis. A Himalayan Endemic? 157

MATERIALS AND METHODS


ATERIALS

Herbarium Specimens

Specimens deposited at BM, CAL, K, KATH, KUN, PE, PYU and TUCH herbaria
(acronyms from Index Herbariorum , http://sweetgum.nybg.org/ih/) were examined,
including photographs of specimens as well as studying the dried collections. Photographs
of living plants were also studied. The morphology of the species was carefully studied from
herbarium specimens under light microscopy. Wherever possible, high resolution
photographs were consulted for specimens we could not have direct access to.
Occurence and Climate Data

Occurrence records were mainly collected by geo-referencing localities from


herbarium specimens. The geo-referenced data were verified for accuracy by plotting them
against the background map in DIVA-GIS v 7.5 (www.diva-gis.org). In addition to

TA B L E 1 . E nnv
v i rro
o n m e n t vvaa rrii aab
b l e s u s e d i n t h e s p e c i e s d i s t rrii bbu
ution modeling

Bioclimatic variables Symbol

Annual mean temperature Bio1


Mean diurnal range (mean of monthly (max temp- min temp)) Bio2
Isothermality (Bio2/Bio7)(*100) Bio3
Temperature seasonality (standard deviation * 100) Bio4
Max temperature of warmest month Bio5
Min temperature of coldest month Bio6
Temperature annual range (Bio5-Bio6) Bio7
Mean temperature of wettest quarter Bio8
Mean temperature of driest quarter Bio9
Mean temperature of warmest quarter Bio10
Mean temperature of coldest quarter Bio11
Annual precipitation Bio12
Precipitation of wettest month Bio13
Precipitation of driest month Bio14
Precipitation seasonality (coefficient of variation) Bio15
Precipitation of wettest quarter Bio16
Precipitation of driest quarter Bio17
Precipitation of warmest quarter Bio18
Precipitation of coldest quarter Bio19
Actual evapo-transpiration AET
Potential evapo-transpiration PET
158 Indian Fern Journal Volume XXXI (2014)

herbarium specimens, data points were also collected from literature, field collections and
GBIF data portal. Altogether 61 points were collected from the herbarium specimens,
literature, GBIF data portal as well as field collections. In order to remove bias data points
closer than 1 km were removed leaving a total of 43 points for our analysis. Nineteen
bioclimatic variables (TABLE 1) were downloaded from the WorldClim database
(www.worldclim.org) and clipped to the geographical area of our interest using ArcGIS v
9.3. Since we did not know the true environmental conditions required for the growth of
H. hamiltonii, we used all the environment variables in our analysis. In addition, two water
related variables, mean annual potential evapotranspiration (PET) and actual
evapotranspiration (AET) were downloaded from the CGIAR consortium for spatial
information (www.cgiar-csi.org). These variables have been found to have significant effects
on the growth and reproduction of fern and fern allies (Kreft et al., 2010; Mehltreter et
al., 2010) and were included in our analysis.
Species Distribution Modeling

Maxent version 3.3.3k was used to create the distribution model for H. hamiltonii.
Although different modeling algorithms exist, Maxent was used in the present analysis
because it has been found to consistently outperform all others (Elith et al., 2006) even at
lower sample size (Hernandez et al., 2006; Pearson et al., 2007). The default settings of
Maxent (regularization multiplier = 1, maximum number of background points = 10000,
maximum iterations = 500, convergence threshold = 0.00001, regularization parameter = 1,
default prevalence = 0.5) were used for the analysis. In order to evaluate model
performance, each model was run with 10-fold cross validation. Area under the receiving
operator curve (AUC) was used to evaluate the model’s goodness-of-fit.
The model was run twice, first using only the Himalayan records and secondly using
all records. This two step modeling was carried out to check if the model created from
Himalayan records would be able to predict occurrence in other regions as well. We assume
that if a species occurs in adjacent non-Himalayan regions as well, the model developed
from Himalayan records only, must partly be able to predict occurrences in non-Himalayan
regions.
RESULT S AND DISCUSSION
RESULT

Herbarium Specimens

Critical study of herbarium specimens collected from throughout the distribution


range of H. hamiltonii confirmed its presence in Arunachal Pradesh, Goa, Jharkhand,
Karnataka, Kerala, Madhya Pradesh, Maharashtra, Manipur, Meghalaya, Odisha, Sikkim,
Tamil Nadu, Uttarakhand, West Bengal, Bhutan, China, Myanmar, Nepal, Sri Lanka,
Thailand and Vietnam. Unfortunately specimens from Sri Lanka, Malaysia and Vietnam were
not seen by us and the presence of this species there could not be verified, though material
has been verified from Sri Lanka and Vietnam by Fraser-Jenkins & Gandhi (2015, in prep.)
Nawal Shrestha & Xian-Chun Zhang : Is Huperzia hamiltonii (Spreng.) Trevis. A Himalayan Endemic? 159

[pers. comm. 9.2014]. Although H. hamiltonii has been reported from Southern Thailand
(Khwaiphan & Boonkerd, 2008), examination of photographs received from Dr.
Thaweesakdi Boonkerd revealed that the specimens from southern Thailand belong to an
allied S.E. Asian species whose nomenclature is unclear. However specimens collected from
northern Thailand correspond with Himalayan H. hamiltonii and are accepted here. Our
results show that the range of H. hamiltonii extends beyond the Himalayan region, as has
long been suggested in Indian botanical literature. The assumption of Zhang & Iwatsuki
(2013) that this species occurs only in the Himalayan region is therefore erroneous.
Species Distribution Models

The first model created using only the Himalayan records (data points from
Uttarakhand, Nepal, Sikkim, Arunachal, Bhutan, S. Yunnan and N. Myanmar) heavily
predicted its occurrences in these regions with high AUC values (0.914 ± 0.097). In
addition, the model also predicted the occurrence of H. hamiltonii in the south western states
of India (data not shown). Although no data points were used from these states, the model
was able to predict occurrences in these regions too. Since, there are records of collection
from these states and we have already verified its occurrence, we can ascertain that the
accuracy of our model is more than just a random result.
Similarly the model developed using all the occurrence data, showed its potential
distribution in all the Himalayan regions, south western states of India, western Yunnan,
northern Laos and northern Vietnam (Figure 2). Although H. hamiltonii is not yet known
from Laos, our model showed its potential distribution there. Therefore if this species
occurs in Laos as predicted by our model needs further verification. The model was unable
to predict its occurrence in Tibet, parts of central and eastern India and parts of Thailand,
in all of which places it is known to occur. Since distribution records from these regions
were based on literature and were not personally verified by us, the accuracy of records of
its occurrence there could not be ascertained by us. An additional model created by omitting
these points greatly increased the performance of the model which shows that even if it
occurs in these regions, the probability is very low.
CONCLUSIONS

Based on our observations and the results of species distribution modeling, it is


evident that H. hamiltonii is not only a Himalayan endemic as stated by Zhang & Iwatsuki
(2013), but occurs well beyond the Himalayan region, too. The result of species distribution
modeling as well as field and herbarium based observation shows that its occurrence is
heavy in the central and eastern Himalaya as well as in the south-west Indian states of Goa,
Karnataka and Maharashtra. The probability of its occurrence in these regions is much
higher (FIGURE 2) than in other parts in the Indian subcontinent. Its occurrence in some
other Indian states (Madhya Pradesh, Odisha and Jharkhand) were not fully supported by
our species distribution models. However Dixit (1987) reported its occurrences in these
160 Indian Fern Journal Volume XXXI (2014)

regions and Fraser-Jenkins & Gandhi (2015, in prep.) have verified specimens from there
as well as from W. Bengal, Manipur, Kerala and Tamil Nadu. Although, specimens from
Sri Lanka were not seen by us, the model heavily predicted its occurrences in the higher
and moister part of the island.. The region is therefore more likely to harbor H. hamiltonii
and it does (Fraser-Jenkins & Gandhi, 2015, in prep.) Occurrence throughout Thailand, as
stated by Khwaiphan & Boonkerd (2008), was not corroborated by either our model, or
our herbarium study apart from its occurrence in the north. Based on our climate model as
well as field based observation and herbarium specimens, the accepted range of H.
hamiltonii would therefore be Arunachal Pradesh, Goa, Jharkhand, Karnataka, Kerala,
Madhya Pradesh, Maharashtra, Manipur, Meghalaya, Odisha, Sikkim, Tamil Nadu,
Uttarakhand, West Bengal, Bhutan, China, Myanmar, Nepal, Sri Lanka, Thailand and
Vietnam.
ACKNOWLEDGEMENTS

We thank the curators and supporting staff of BM, CAL, K, KATH, KUN, PE,
PYU and TUCH herbaria for allowing access to material (photographs and/or herbarium
specimens) for study. We would also like to thank Dr. Thaweesakdi Boonkerd, Bangkok,
for sending us photographs of specimens from Thailand. We are also grateful to Mr. Chris
Fraser-Jenkins for his help in identification of the collected materials as well as providing
relevant literature and other information. We would also like to express our gratitude to
Dr. Sachin Patil, Kolhapur, for providing pictures of specimens from his collections as well
as sharing his collection localities. Lastly we express our appreciation of the encouragement
and support received from Prof. S.C. Verma, Chandigarh. This work was supported by
Research Grants in Plant Systematics 2012 (received by the first author from the
International Association for Plant Taxonomy), the Chinese Academy of Sciences and the
National Natural Science Foundation of China (Grant Nos. 31070196, 31170199,
31370260).
REFERENCES

B O O N K E R D T, P O L L AWAT N R & DA RU M A S P 2 0 0 8 Taxonomic Revision of the Lycopodiaceae in


Southeast Asia : Final Report Chulalongkorn University, Thailand
CHANDRA S , FRASER-JENKINS C R, KUMARI A & SRIVASTAVA A 2008 A Summary of the status
of thr ea tened Pteridoph ytes of India Taiwania 5 33(2) : 170-209
DIXIT R D 1984 A census of the Indian Pteridophytes Botanical Survey of India, Howrah Kolka ta
DIXIT R D 1987 Lycopodiaceae of India Bishen Singh Mahendra Pal Singh, Dehra Dun India
E L I T H J , G R A H A M C H , A N D E R S O N R P, D U D Í K M , F E R R I E R S , G U I S A N A , H I J M A N S R J ,
H U E T T M A N N F, L E AT H W I C K J , L E H M A N N A , L I J , L O H M A N N L G , L O I S E L L E B A ,
MANION G, MORITZ C, NAKAMURA M, NAKAZAWA Y, OVERTON JMM, PETERSON A T,
PHILLIPS S J, RICHARDSON K, SCACHETTI-PEREIRA R, SCHAPIRE R E, SOBERÓN J,
WILLIAMS S, WISZ M S & ZIMMERMANN N E 2006 Novel methods improve prediction
of species’ distributions from occurrence data Ecography 2 9 : 129-151
Nawal Shrestha & Xian-Chun Zhang : Is Huperzia hamiltonii (Spreng.) Trevis. A Himalayan Endemic? 161

FRASER-JENKINS C R 2008 Ta x o n o m i c R ev i s i o n o f t h r e e h u n d r e d I n d i a n S u b c o n t i n e n t a l
Pteridophytes with a revised Census list Bishen Singh Mahendra Pal Singh, Dehra Dun India
FRASER-JENKINS C R 2010 Ne pal’s little known pter idophytes, the hid den wor k of David Don and
the geography and distribution of Indo-Himalayan ferns with State lists, website version,
updated 4th Oct 2011, on www groups yahoo com/group/Indian-Ferns
FRASER-JENKINS C R & GANDHI K 2015, in prep A revised checklist of Indian pteridophytes
HERNANDEZ P A, GRAHAM C H, MASTER L L & ALBERT D L 2006 The effect of sample size and
species characteristics on performance of different species distribution modeling methods
Ecography 2 9 : 773-785
I WAT S U K I K 1 9 8 8 T h e H i m a l a y a n p l a n t s B u l l e t i n N o 3 1 T h e U n iv e r s i t y M u s e u m , U n iv e r s i t y o f
To k y o J a p a n
K H WA I P H A N W & B O O N K E R D T 2 0 0 8 T h e P t e r i d o p hy t e F l o r a o f K h a o K h i a o , K h a o Ya i N a t i o n a l
Park, Thailand The Natural History Journal of Chulalongkorn University 8 (2) : 69-82
K R E F T H , J E T Z W, M U T K E J & BA RT H L OT T W 2 0 1 0 C o n t r a s t i n g e nv i r o n m e n t a l a n d r eg i o n a l
effects on global pteridophyte and seed plant diversity Ecography 3 3 : 408-419
M E H LT R E T E R K , WA L K E R L R & S H A R P E J M 2 0 1 0 F e r n E c o l o g y C a m b r i d g e U n iv e r s i t y P r e s s ,
Cambridge
PEARSON R G, RAXWO RTHY C J , NA K A M U R A M & TOW N S E N D P E T E R S O N A 2007 Predicting
species distributions from small numbers of occurrence records : a test case using cryptic
geckos in Madagascar Journal of Biogeography 3 4 : 102-117
TAGAWA M & IWATSUKI K 1979 Lycopodiaceae In " Flor a of T hailand 3(1) " Eds T Smitinand & K
Larsen Publisher : Phonphan Printing Bangkok
ZHANG L B & IWATSUKI K 2013 Lycopodiaceae In " Flora of China 2-3 " Eds ZY Wu, PH Raven &
DY Hong Publishers: Beijing Science Press, Beijing and Missour i Botanical Garden Pr ess, St
Louis
Indian Fern J. 3 1 : 162-172 (2014) ISSN 0970-2741

INV
VASION VINIA MOLEST
SALVINIA
ASION OF SAL A D.S
MOLESTA S.. MITCHELL (AFRICAN PAYA L )
PA
IN KERALA AND ITS MAN
MANA A GEMENT
MADHUSOODANAN P. V.1*, SREERENJINI V. K.1**, SMITHA R. B.1*** and PRAKASHKUMAR R.1****
1
Malabar Botanical Garden, G.A. College P.O., Kozhikode, Kerala - 673014

(Received November 05, 2014)

ABSTRACT
Salvinia molesta D.S. Mitchell. (Salviniaceae) a free floating fern, from South
America introduced to Kerala by 1955 for botanical studies became a pernicious aquatic weed
infesting almost all fresh water ecosystems of Kerala afflicting the life of people and
environment in various ways. It affected paddy cultivation, irrigation systems, inland water
transport, fisheries, electricity generation of hydroelectric projects, aquatic flora and fauna and
cause pollution affecting public health. However, this exotic weed was effectively controlled by
using a natural pest (biological control agent), a weevil, viz., Cyrtobagous salviniae introduced
from its original habitat.
d s : Salvinia molesta, weed, problems, biological control, Cyrtobagous
K e y Wo rrd
salviniae

INTRODUCTION

S. molesta D.S. Mitchell (‘African Payal’) is an elegant free floating, heterosporous


fresh water fern which has become a pernicious weed causing great menace to agriculture,
water transport, public health, hydroelectric projects and fish farming throughout Kerala
State (South west coast of Peninsular India). During the past 50 years it has become a
major weed in the state and is reported earlier from Africa and Sri Lanka. The extent of
damage it has caused and tremendous economic loss it is causing are brought out in several
reports (Cook and Gut, 1971; Menon, 1971) and considered as the world’s worst weed
(Nayar and Madhusoodanan, 1979; Rajagopalan 2014).
Salvinia Seguier (named after Antonia Maria Salvini, an Italian Greek Scholar) is a
genus of ca. 11 extant species of which six are restricted to Tropical America, S. hastata
Desv. occurs in tropical and subtropical East Africa (Alston, 1959). S. cucullata Roxb. Ex
Bory is a widely distributed species though not frequent, in North Eastern India, Malaya,
Sumatra and Western Australia (Holttum, 1968). The only temperate species, S. natans (L.)
All. has a wider distribution ranging from British Isles in the west to all over Europe,
Northern India (Kashmir), Northern China up to Japan in the east. Of the seven American
species (the origin of S. molesta D.S. Mitchell still arbitrary), six are restricted to tropical
America and one species, S.molesta D.S. Mitchell has spread to Southern Africa, Southern
Asia, Australia, Papua New Guinea and some Pacific Islands as an aggressive weed posing
grave problems for the economy and environment of the infested area.
The genus Salvinia has not been represented in S. India until the introduction of

* Email : pvmadhu@gmail.com ** Email : vk.sreerenjini@gmail.com *** Email : rbsmitha@gmail.com


**** Email : rprak62@gmail.com
Madhusoodanan et al. : Invasion of Salvinia molesta D.S. Mitchell. (African Payal) in Kerala 163

Salvinia molesta ca. 50 years ago which made its first appearance in Trivandrum, the capital
city of Kerala in 1955. Two species of the genus exist in the N. India viz., S. cucullata
Roxb. Ex Bory with sparse distribution in the north eastern regions (Assam and Bengal)
and S.natans (L.) All. restricted to some lakes of Kashmir. But, both species (and all other
species) are never reported as troublesome weeds though the latter may become a thin carpet
over the lakes during certain part of the season. For long, African payal was confused with
S. auriculata Aubl. – a new world species. Mitchell (1972) has shown that it is a different
and new taxon and named it S.molesta D.S. Mitchell.
The first authentic report of the existence of S. molesta as a weed is from Sri Lanka
where it was observed in 1942 forming small colonies near Kolonnawa, a suburb of
Colombo by an agricultural officer of Sri Lankan Govt. Service misidentifying it as S.
auriculata (Sculthorpe, 1971). Moderately heavy infestations were then reported in 1943 at
Negumbo, nearly 50 Km from Colombo. In the same year it was reported that S.molesta
has infested and covered an area of ca. 95 Km 2 near the Stanley Power Station at
Kolonnawa (Senaratna, 1952; Williams, 1956). It is assumed that it has come as an escape
from the sample of living material obtained by the Colombo University from the Indian
Botanic Gardens, Calcutta. A survey undertaken by Senaratna (1943) revealed that the weed
was restricted to the Western Province in 1943. Another survey conducted by him in 1947
showed that it was spreading fast and by 1952 it has infested the North Western and
Southern Provinces of Sri Lanka (Williams, 1956), but the infestation was restricted to the
coastal belt on the western side of the island. By 1954, ie., 12 years after its first
appearance, it has spread over 850 ha of waterways and 9000 ha of paddy fields (Williams,
1956) all over Sri Lanka and for the first time exhibiting its incredible potentialities as a
gregarious weed. However, Salvinia molesta attracted the attention of the scientific world
when it appeared in the man-made Kariba Lake in Southern Africa. In May, 1959 when the
newly created reservoir in the Zambesi valley (Zimbawe) was filling up, small patches of
S. molesta were observed on the rising waters nearly six months after the closing of the
dam (Hatting, 1961; Schelpe, 1961). Earlier the species was reported on the Zambesi river
as early as 1948, ca. 160 Km upstream from Lake Kariba. It is not clear where the original
infestation of S. molesta happened. But it seems likely that Salvinia existed much earlier to
its detection in the Chobe Swamps of Botswana Land, the swamp formed by the Kwando
river which flows into the Zambesi river beyond the swamps at Kazungula in Zambia. The
building of the Kariba Lake in 1959, providing a vast expanse of lentic water enriched with
decayed plant debris created a highly suited habitat for S. molesta and the plant underwent
an explosive growth and colonization (Jackson, 1960). Within eleven months of its first
appearance in the lake, (by April 1962) it has spread over an area of 1003 Km2 (21.5% of
total lake surface) forming dense floating mats (Mitchell and Tur, 1975). The history of
infestation of Lake Kariba provides a remarkable testimony to the astonishing power of S.
molesta for explosive colonization. The prodigious capacity of the weed to survive and
164 Indian Fern Journal Volume XXXI (2014)

regenerate under formidable conditions is also well demonstrated by the spreading to the
Lake Kariba from the Chobe Swamp area. The very swift current of gorges beyond the
falls also provide a natural hazard which very few floating plants could survive. From the
Victoria Falls to Lake Kariba it is a distance over 150 Km and no Salvinia grows in this
part of river. The heaviest infestation of this weed occurs along the course of the Zambesi
river and some of its tributaries (Guillarmod, 1979). The weed threatened the Okovango
swamp areas. A rigid cordon sanitaire was being maintained to keep the weed out of
Okovango swamps because once this area is infested; it can never be eradicated and will
completely upset the very delicately balanced environment (Guillarmod, 1979). It was owing
to its notoriety as an aggressive weed (World’s worst weed) as reported in Kariba Lake,
Africa. Salvinia molesta is known as ‘African Payal’ (Payal means weed in Malayalam) in
Kerala confusing its original home, S. America (Nayar and Madhusoodan, 1979).
It was originally hoped that the invasion of Salvinia in Lake Kariba was a transient
phenomenon and would get itself controlled as the lake filled up and years go by, especially
when the submerged trees in the area perished so that natural mooring of the colonies will
no longer possible. However, this has proved to be wrong as also a similar hope entertained
in Sri Lanka and S. India. In all these areas, Salvinia infestation grows almost undiminished
till today nearly 50 years after its colonization. Moreover, it has spread to Australia (Usher,
1971) Indonesia (Soerjani, 1976) and Thailand (Junk, 1977).
Salvinia molesta in Kerala
S. molesta probably reached in Southern Kerala somewhere around 1955-60. By
1960-63, there are records of its having been cytologically investigated in the Botany
Department of the Kerala University, Trivandrum. Cook and Gut (1971) reported that it
was first noticed as a weed in Kerala in 1964. However, collections made by Prof. Abraham
and sent to Royal Botanic Gardens, Kew during 1958-60 (now deposited at Kew Herbarium)
reveal its presence in this State much earlier. This has been confirmed by some teachers of
Kerala Govt. Collegiate Service who were students of the Kerala University Botany
Department at Trivandrum during 1955-57 as they were provided with live materials of
S.molesta for practical studies. Misidentifying the taxon as S.auriculata it is presumed that
the weed was inadvertently introduced into the State for teaching of Botany in Kerala
University, Trivandrum from the Indian Botanic Garden, Calcutta. It is documented that the
Indian Botanic garden (formerly Royal Botanic Garden) had S.auriculata (= S.molesta ?) in
cultivation since 1933, the original specimens having been obtained by the Garden from
Germany (Cook and Gut, 1971). The German Botanic Garden appears to have got the
specimens from Rio de Janiero, South America, and had it in cultivation for long. It is
interesting to note that though S.auriculata (?) has long been in the Botanic Gardens of the
University of Singapore, it has not spread as a weed as S.molesta has spread in Kerala, Sri
Lanka and Africa. At the Indian Botanic Gardens it covers extensive areas in all the lakes
and it is regularly removed manually. No preventive measure is taken to check the
Madhusoodanan et al. : Invasion of Salvinia molesta D.S. Mitchell. (African Payal) in Kerala 165

spreading and it is possible that the plant has got washed out into the swampy areas during
rainy season. But still it has not become an aggressive weed. Similarly in Singapore, during
rainy season colonies of S.auriculata (?) washed away from the University Botanic Garden
have been found in several nearby areas (Sculthorpe, 1971). Even so, it has not become
established in the water ways of Singapore Island. It is also to be mentioned that S.molesta
is cultivated since long in the Lalbagh Garden, Bangalore, S. India where a large tank is
seen fully covered by this weed. However, there is no spreading of this weed to other parts
of Karnataka. Cook and Gut (1971) presume that S. molesta reached Kerala from Sri Lanka
though there is no direct evidence, having been inadvertently used as a packing material by
someone. They have stated that Salvinia was supplied from the Indian Botanic Gardens,
Calcutta to be Colombo University in 1939 for botanic studies. Subsequently unwanted
specimens somehow got into nearby waters where they survived beyond all expectations
(Senaratna, 1952). All these presumptions are based on the assumption that S. molesta of
Kerala and Sri Lanka is the same taxon and the material from Indian Botanic Gardens is
the same species. Mitchell (1973) has made a comparative study of the Kerala, Sri Lanka
and African materials and concluded that they are conspecific. The Calcutta material and
the specimen from Lalbagh, Bangalore are verified with Kerala material and confirmed that
they are conspecific.
S. molesta D.S. Mitchell. varies from other specis of this genus, with their elongate
sporocarp bunches having pedunculate megasporocarp and subsessile microsporocarps
arranged distichously in a helical manner (Madhusoodanan,1987). It is a sterile pentaploid
hybrid with parents S.auriculata (hexaploid) and S.biloba (tetraploid) (Mitchell, 1972,
Kuriachan, 1979). It possesses a delicate, spongy, profusely branched, cylindrical,
plagiotropic rhizome with prominent nodes and internodes. At each node, there are two
aerial leaves and a submerged organ (shoot) bearing a tuft of root-like branches at its tip
(Figs.1A-F). Roots are absent .The sporocarps are produced from the centre of the tuft of
root-like branches at its tip. On the upper surface of the aerial leaves there is characteristic
egg-beater like hairs (Fig. 1D). The rhizome cortex is provided with many air chambers,
characteristic of hydrophytes and possesses solenostele in the internode which becomes
siphonostelic at the nodal region (Madhusoodanan and Nampy, 1994).
In 1975, a scientific team was constituted by Kerala Government (Agriculture
Department) to investigate the possibilities of the management of this weed in Kerala
collaborating scientists from Calicut, Kerala and Kerala Agricultural Universities. Calicut
University was entrusted with ecological studies, chemical control was entrusted with
Applied Chemistry Department of Kerala University and the KAU was given the
responsibility of biological control of S. molesta.
MATERALS AND METHODS
ATERALS

Specimens of Salvinia molesta were collected from all over Kerala (149 sites in 14
districts) and the impact of S. molesta infestation on the environment and public life of
166 Indian Fern Journal Volume XXXI (2014)

Kerala was recorded in a Questionnaire prepared for this purpose, by direct observation and
in conversation with local people and scientists of regional research institutions. Data were
collected on the time and ways of introduction of the weed in that area, morphological
variations, impacts on agriculture, water transport, fisheries, etc., associated flora and fauna,
pathological symptoms, natural pests feeding on the weed and utilization prospects of the
weed as practiced by local people. Morphological observations were conducted in the field
and laboratory using standard methods. The collected data were analyzed statistically and
conclusions were derived.
The preliminary biological control experiments were conducted at the Entomology
Division of Kerala Agricultural University, Vellanikkara under the guidance of Prof. P.J.
Joy. The biological control agent was obtained from the Commonwealth Institute of
Biological Control (CIBC), Bangalore. Precautions were taken to prevent the biological
control agent in escaping from the experimental tanks (Joy, 1978). After satisfactory tests
on the feasibility of application and safety of the insect’s infection on other plant species,
the weevil was released on S. molesta population in an open water area in the Pamba River.
RESULTS
LT

The survey has revealed that about 7500 sq.km (30%) and 80 % of the low land
area of Kerala has been infested with S. molesta by 1977 (Madhusoodanan, 1987). Only
one species of Salvinia, i.e., S. molesta D.S. Mitchell occurs in Kerala and exhibits two
growth forms: i.e., primary with small flat leaves and secondary with larger conduplicately
folded velvety leaves bearing sporocarps (Fig. 1 A-E). Since the low lands of Kerala is
interconnected with numerous waterways like canals, rivers, lagoons, back waters with
frequent water transportation via country boats, etc. the spread of the weed was easy and
fast. The application of excess chemical fertilizers in the paddy fields and leaching to
nearby water bodies augmented the explosive growth of this weed. Moreover, the tropical
humid climate of Kerala, equivalent to the climate of its original habitat, was ideal for the
luxuriant growth of S. molesta. From Thrivananthapuram to Kasaragod, about 500 km
distance was occupied by the weed by within five years. In the upland area like Idukki
and Wayanad, the weed was introduced by human beings for the beauty of the velvet-like
the aerial leaves, out of curiosity, to their garden ponds from where it escaped to nearby
waterways during monsoon season. All over Kerala, the plants were found sterile and the
propagation is via vegetative fragmentation only.
Through some natural pests are found feeding on S. molesta, the quantity of the
plant consumed by them was too small to utilize them as biocontrol agents. In Kerala, S.
molesta affected agriculture, irrigation, water transport, inland fisheries, and loss of native
plant species and caused pollution through decay. It also affected the electricity generation
at Sabarimala project (at Moolamattam Power House) by infesting in the Kakki reservoir
and entangling with the turbines of generators chocking the electricity production making it
stand still. Kakki reservoir is 700m above sea level and how S. molesta reached in the
reservoir is still dubious.
Madhusoodanan et al. : Invasion of Salvinia molesta D.S. Mitchell. (African Payal) in Kerala 167

F i g . 1 . A – E . Salvinia molesta DS. Mitchell - morphological features. A . Primary growth form.


B . Intermediate stage showing beginning of folding of aerial leaves, C . Secondary stage with condumplicately
folded aerial leaves, D . Egg beater - like hairs on upper surface of aerial leaves, E. Submerged branch (organ)
showing root-like branches and bunches of sporocarp and F. Sporocarp bunch showing arrangement of sporocarps.
168 Indian Fern Journal Volume XXXI (2014)

F i gg.2. F.. Salvinia molesta infestation and biological control. A . A country boat being trapped in
.2. A – F
Salvinia mats. B . A canal showing thick and complete choking by Salvinia in Aleppy Dt. C. Biological control
using Cyrtobagous salviniae (Inset the weevil Cyrtobagous salviniae – Courtsey : Wikipedia), D. Salvinia mats
showing signs of death and decay after 2 weeds of introduction of the weevil, E. After 4 weeks showing almost
complete destruction and F. A pond showing the destruction of S. molesta .
Madhusoodanan et al. : Invasion of Salvinia molesta D.S. Mitchell. (African Payal) in Kerala 169

However, it is believed that S.molesta probably reached there through the boats
transported to the reservoir area during the construction of the dams either as a packing
material or somehow attached to it. In paddy fields, the weeds fall on the seedlings when
the flood water recedes and suppress them to soil. Attempts to manually remove the weed
from the paddy field, etc. were failed owing to the magnificent capacity of these plants for
regeneration from the left over plant parts, especially the apical and lateral buds, which can
cover the entire area within 10-15 days again. In stagnant ponds, S. molesta forms mats
up to 1m thick and harbors other plants called “sudd” vegetation, mainly grasses, sedges,
herbs and seedlings of trees. It has also affected the populations of the sought after fishes
of Kerala like the ‘Karimeen’ (Atroplus suratensis, the Pearl spot),’bral’ (Channa channa)
and the giant prawn (Macrobrachuium sp.).
The preliminary Biological control experiments were conducted at the KAU campus,
Vellanikkara, using the agent (Cyrtobagous salviniae), a weevil, originally from Brazil,
obtained through the Commonwealth Institute of Biological Control, Bangalore (Now known
as The Bureau of Agriculturally Important Insects). After rearing and feeding tests on the
local crops such as rice, coconut, vegetables, etc. and found safe, the weevil was first trialed
at Athirampuzha in the Pamba River near the MG University Campus, Kottayam. The
results were very promising. The Salvinia population of that area was destroyed within 30
days. The effect was evident from the first week itself showing browning of leaf margin
and yellowing of leaves of the apical regions (Fig. 2 C-F). Later, the agent was introduced
in various affected areas of Kerala and the weed was effectively managed. At present
though present in some localities with depleted vigor, S. molesta is not a serious aquatic
weed in Kerala.
DISCUSSION

The most extensive work on the autecology of Salvinia has been done on S.molesta
of Kariba lake, popularly known as the ‘Kariba weed’ by Mitchell (1970). The nutritional
requirements, rates and factors of growth of S.molesta under natural as well as laboratory
conditions have been investigated by him. These studies are of particular interest considering
the potentiality of the plant for explosive growth only by vegetative means in the context
of controlling or eradicating it from unwanted areas.
As remarked by Bates and Hentges (1976), ‘adverse effects of aquatic vegetation
on the environment are increasingly serious worldwide problem challenging the international
community. The development of control measures will require innovative thinking and
creative research’. Formerly people used to think in terms of eradication i.e., total
devastation of the undesirable species. But nowadays, the adverse effects of such an
indiscriminate deed is considered first before commencing any attempt and started thinking
in the lines of management instead of eradication as it will affect the edaphic stability and
depletion of oxygen and nutrients of the water body. So, now it is considered that how
170 Indian Fern Journal Volume XXXI (2014)

much weed may be efficiently maintained at the selected level at an economic cost
(Sculthorpe, 1971).
Herbicides (weedicides) are often highly effective for eradication of weeds especially
in the early stages of development but are frequently indiscriminative in their devastation
of associated flora and fauna. The selection of appropriate herbicide is very difficult since
it is influenced by several factors such as the morphology and physiology of the plant and
location, nature and uses of the habitat. It is more theoretical than practical to control the
aquatic vegetation by removing essential trace elements from the substratum using specific
chemical techniques as suggested by Martin et al. (1971). This method is however, almost
impossible considering the diversity and complexities of aquatic ecosystems. But it may be
possible to manipulate the environment by addition of internal plant growth regulators such
as gibberellins, if the biochemistry of the weed is adequately understood.
Now a days, attention is much being centered on the possibilities of using a natural
pest or pathogen for the efficient management of weeds. Biological control is considered
to be the most effective and safe method for the control of many weeds as evidenced by
the successful control of the prickly pear (Opuntia sp.) in Australia and Lentana camera of
Hawaii islands (Robins et al., 1942). Biological control has several advantages over other
methods of weed control such as, i) low application cost and persistence, ii) pollution free
and iii) high affectivity due to extreme reachability of the agent to even inaccessible sites
where the weeds grow (Bates and Hentges, 1976). The control agents generally suggested
are phytophagous animals, insects and fungal and microbial plant pathogens. The
disadvantages are: i) it is a slow process and therefore takes a long time ii) if the control
agent is not specific the pest may not have the desired effect, iii) the effect and progress of
the work of the natural enemy will be considerably affected if it is attacked by a hyper
parasite and iv) replacement will become necessary every year if certain seasons of the year
are unfavorable for the agent (Debauch, 1974). The attributes of a natural enemy to be an
effective biological control agent are: i) it should be highly adaptable to the varying
physical conditions of the new environment occupies by the weed possessing high host
reaching capacity; ii) it should multiply faster than the host with high female-male ratio (in
the case of animals) and short life cycle, iii) it should be a voracious consumer of the host,
iv) it should be specific to the host or the range of hosts attacked by the agent should be
narrow, v) the biology of the agent must be synchronized with the host and vi) it should
be able to breed easily under laboratory conditions. On the other hand, when the agents are
indiscriminate, affecting associated flora and fauna including desirable species, an
irreversible damage to the environment could occur. Thus one should be highly cautious to
introduce an exotic pest as a biological control agent and its specificity to the weed should
be convincingly confirmed. At present, the subtle and poorly understood biological
relationship between host, agent and environment place most methods for biological control
of aquatic weeds in a promising experimental, but not commercially useful stage of
Madhusoodanan et al. : Invasion of Salvinia molesta D.S. Mitchell. (African Payal) in Kerala 171

development (Bates and Hentges, 1976). However, a search for pathogens and insects which
possess the virtues of high specificity and vitality, without adverse after effects could be
rewarding. The prospect of such predators seemed rather remote for ‘African Payal’ until
the successful report on biological control came out from Australia. In Lake Moondarra,
Queensland 8000 tones fresh weight of Salvinia molesta was destroyed by the weevil,
Cyrtobagous salviniae Calder and Sands, introduced from Brazil (Room et al., 1981). It is
interesting to note that in Brazil, the native land of S. molesta, this plant is not a weed. It
was discovered that this status is maintained owing to the continuous feeding by natural
pests such as C. salviniae. This is how this weevil was identified as an ideal bicontrol
agent for S.molesta and successfully controlled in Australia. Since then, efficient control of
S.molesta has been achieved in Papua New Guinea, where a subtropical climate exists.
Forno (1985) suggested that C. salviniae should be a successful control agent in tropical
countries. Total eradication of a species, irrespective of the quantum of menace it creates to
human race is unethical and hence effective management of such organisms to affordable
level is logical and rational. The report of collection of C. salviniae from Thrissur for
introduction and in Puttanapalli Lake, Banglore, indicates the persistence of this weevil even
after 30 years of its introduction in Kerala (Rajagopalan, 2014).
REFERENCES

ALSTON A H G 1959 Fer ns and F er n allies of West Tropical Afr ica Suppl : Flora of West Tropical
Africa Ed 2 Crown a gents f or oversea Govts and Adminstr ation London
BATES P & HENTGES Jr J F 1976 Aquatic weeds-eradicate or cultivate? Econ Bot 3 0 : 39-50
COOK C D K & GUT B J 1971 Salvinia in the State of Kerala India Pans 1 7 : 438-447
DEBAUCH P 1974 Biological Control of Natural Enemies Cambridge Univ Press
FORNO I W 1985 H ow q u i c k l y c a n i n s e c t s c o n t r o l S a l v i n i a i n t h e t ro p i c s ? P r o c 1 0 t h A s i a n Pa c i fi c
Weed Sceince Soceity Conf T hailand Nov 1985 pp 271-276
GUILLARMOD J 1979 Water weeds in Souther n Africa Aqua tic Bot 6 : 377-391
H AT T I N G E R 1 9 6 1 Th e p r ob l e m o f Salvinia auricula ta A u bl a n d a s s o c i a t e d a q u a t i c w e e d s i n L a ke
Kariba Weed Res 1 : 303-306
H O LT T U M R E 1 9 6 8 A R ev i s e d F l o r a o f M a l a y a Vo l I I F e r n s o f M a l a y a G ov t P r i n t i n g O ffi c e
Singapore
JACKSON R N B 1960 Hydrobiological research at Kariba New Scientist 8 7 7 : 80
J OY P J 1 9 7 8 E c o l o g y a n d c o n t r o l o f S a l v i n i a ( A f r i c a n p a y a l ) , t h e m o l e s t i n g w e e d o f Ker a l a Tec h
Bull No 2 Ker Agic Univ India
JUNK W J 1977 Note on aquatic weeds in some reservoirs in Thailand Aquatic Bot 3 : 85–90
KURIACHAN P I 1979 I n t e r s p e c i f i c o r i g i n o f Salvinia molesta : E v i d e n c e f r o m k a r y o t y p e I n d i a n J
Bot 2 : 51–54
MADHUSOODA NAN P V 1987 Studies on ‘African Pa yal’ ( Salvinia molesta D S Mitchell) in Kerala
State, Ph D thesis, University of Calicut
M A D H U S O O DA NA N P V & S A N T H O S H NA M P Y 1 9 9 4 M o r p h o g e n e s i s o f s h o r t a n d va s c u l a r
differentiation in the water fern, Salvinia molesta Mitchell (Salviniaceae-Pteridophyta) J
Taiwan Mus 4 7 : 61-68
172 Indian Fern Journal Volume XXXI (2014)

MADHUSOODA NAN P V 1988 Taxonomy and distribution of Salvinia Seguier in the Old Wor ld Pr oc
Internatl Symp In Syst Pterio Beiging, China pp 55–68
M A RT I N D F D O I G M T & M I L L A R D D K 1 9 7 1 Potential control of Florida Elodea by natural
control agents Hyacinth control J 9 : 36–39
MENON M V K 1971 The threatening weed Science Toda y 6 : 33–38
MITCHELL D S 1970 A utecological studies of Salvinia auriculata Ph D Thesis Univ Loondon
MITCHELL D S 1972 The Kariba weed; Salvinia molesta Brit Fern Gaz 1 0 : 251–252
MITCHELL D S 1973 A q u a t i c we e d s i n m a n - m a d e l a ke s G e o - p hy s i c a l M o n o g ra p h S e ries 1 7 : 6 0 6 –
611
MITCHELL D S & TUR N M 1975 The rate of growth of Salvinia molesta ( Salvinia auriculata auct)
in labor ator y and natur al conditions J Appl Ecol 1 2 : 213 –225
NAYAR B K & MADHUSOODANAN P V 1979 The wor ld’s worst weed Science Reporter (CSIR) 1 6 :
15–21
RAJAGOPALAN U 2014 Salvinia weevil released blo g : m bangalore citizen
ROBINS W W CRAFTS A S & RAYNER R N 1942 Weed control – A text book and Manual McGraw
Hill Book Co Ltd New York
RO O M P M H A R L E Y K L S F O R N O I W & S A N D S D P A 1 9 8 1 Successful biological control of
floating weed Salvinia Nature 1 9 4 : 78–80
SCHELPE E A C L E 1961 T h e e c o l o g y o f S a l v i n i a a u r i c u l a t a a n d a s s o c i a t e d v eg e t at i o n i n K a r i b a
Lake JS Afri Bot 2 7 : 481–486
SCULTHORPE C D 1971 T he Biology of Aquatic Vascular Plants Ed ward Arnold London
S E NA R AT NA J E 1 9 4 3 Salvinia auriculata Aubl – a recently introduced free-floating water weed
Trop A gri Ma g Ceylon 9 9 : 1 4 6 - 1 4 9
SENARATNA J E 1952 Preliminary trials for the control of water fern Salvinia Trop Agr Mag Ceylon
1 0 8 : 49-50
SOERJANI M 1976 Aqua tic weed problems in Indonesia with special refer ence to the construction of
m a n - m a d e l a ke s I n : C K Var s h n e y a n d R zo s k a , J E d A q u at i c w e e d s i n S o u t h A s i a p p 6 3 – 7 8
Junk The Hague
USHER J F 1971 Salvinia -a r ival for water hyacinth Cane Grower’s Quart Bull 34: 137 – 138
WILLIAMS R H 1956 Salvinia auriculata Aub l -the chemical eradication of a serious aquatic weed in
C ey l o n Tr o p A gr Train 3 3 : 1 4 5 - 1 5 7
Indian Fern J. 3 1 : 173-186 (2014) ISSN 0970-2741

ECOLOGY OF SOUTH INDIAN PTERIDOPHYTES


MADHUSOODANAN P. V.1*, PRAKASHKUMAR R.1**, SREERENJINI V. K.1*** and SMITHA R. B.1****
1
Malabar Botanical Garden, Kozhikode, Kerala-673 014

(Received December 2, 2014; Revised Accepted December 12, 2014)

ABSTRACT
In South India, pteridophytes are adapted to almost all ecosystems and micro
habitats to represent all ecological types such as hydrophytes, xerophytes, sciophytes,
rheophytes, etc., except parasites and marine plants. Many species are located in high altitudes
above 700m and few only above 900m. They include the small aquatic mosquito ferns (Azolla
pinnata R.Br.) and the arborescent tree ferns like Cyathea crinita. Salvinia molesta Mitch. is a
notorious aquatic weed and Azolla spp. is an effective biofertilizer in paddy fields. Being
sensitive to ecological perturbations, many species have become endangered or extinct from
their original habitat. Unless effective and immediate conservation steps are being taken, many
Pteridophyte species will soon disappear from this area.
K e y W o rrd
d s : Ecology, Pteridophyte, South India

INTRODUCTION

Being the most primitive vascular plants originated some 400 million years ago, but
survived the inimical geological and climatic catastrophes occurred from time to time, the
pteridophytes must have adapted to the then unfavorable environmental factors despite their
less developed vascular system and simple structure. Today the pteridophytes represent
almost all ecological types from Hydrophytes to Xerophytes and Rheophytes to Halophytes.
In size they represent the small mosquito fern (Azolla spp.) to arborescent tree fern
(Cyathea, Alsophila, etc.). They exhibit an independent alternation of generations in their
life cycle. Being bereft of seeds, they propagate through haploid spores produced on leaf.
The myriads of spores, dispersed through wind find a wet place to lodge and germinate to
develop into a prothallus which requires water essential for fertilization. It is this site of
germination which becomes the home of the sporophyte in future. The selection of the site
is a matter of destiny, the natural selection by survival and adaptation becomes a matter of
existence.
In South India (Peninsular India) the pteridophytes occur in almost all eco habitats
but for marine ecosystems. Page (1979) has considered the tropical ferns under five major
categories and 23 subcategories. Mehltreter et al. (2010) discussed various aspects of fern
ecology. Sharpe et al. (2010) discussed the ecological importance of ferns.

THE STUDY AREA


STUDY

South India includes six political states of India viz. Kerala,, Karnataka, Tamil
Nadu, Andhra Pradesh, Telangana, Goa and Union Territories of Mahe and Pondicherry.

* Email : pvmadhu@gmail.com ** Email : rprak62@gmail.com *** Email : vk.sreerenjini@gmail.com


**** Email : rbsmitha@gmail.com
174 Indian Fern Journal Volume XXXI (2014)

Geographically the area falls between 80-200 N latitudes and 740-850 E longitude. It covers
an area of 4,67,186 km2 forming the Peninsular India, with Bay of Bengal on the East,
Arabian Sea on the West and Indian Ocean on the South. On the North it is bordered with
Vindhya – Satpura Mountains of Central India.
South India is essentially a triangular table land (plateau) bordered with mountains
of Eastern Ghats and Western Ghats. The W. Ghats slope rather abruptly to the coastal
region from to 2700 to 0 MSL to provide quick altitudinal gradients. The Western Ghats
with abundant rainfall from the monsoon (June- September) house a variety of plants and
animals in Tropical evergreen forests and other types of vegetation (Fig. 1 A-F). Western
Ghats had one of the ancient civilization and human settlement (Stone age) as depicted at
Edakkal caves in Wayanad.
The pioneer studies on South Indian ferns were done by Beddome (1863), an
amateur pteridologist who undertook the floristic survey of ferns of British India, Ceylon
(Srilanka) and Malay Peninsula. The Malayan and Srilankan pteridophytic flora by Beddome
had been repeatedly revised and updated by Holttum (1968) and Sledge (1982) respectively.
During the last half century, considerable work has been done on pteridophytes of South
India (Madhusoodanan et al. 2001). The ecology of ferns of Palani Hills of South India
has been studied by Manickam and Ninan (1984). Ecology of Lomariopsid ferns of South
India has been done by Majeed et al. (1994). Ecology and distribution of Thelypteriod ferns
of South India was studied by Leena and Madhusoodanan (1992). The ecological impact of
Salvinia molesta Mitch. on the wetland ecosystems of Kerala has been discussed by
Madhusoodanan et al. (2014). Rajagopal and Bhat (1998) have surveyed the pteridophyte
flora of Karnataka. Pullaiah et al. (2003) surveyed the pteridophytes of Andhra Pradesh.
Pteridophytes occupy almost all types of habitat in South India except marine
ecosystems (Table 1).
Hydrophytes
In South India the pteridophytes represent all the three aquatic forms of life i.e.,
submerged, emergent and free-floating. Salvinia molesta D. S. Mitchell and Azolla pinnata
R.Br. are free floating. Salvinia molesta Mitch. is notoriously known as “the world’s worst
weed” (Nayar and Madhusoodanan, 1978) for its gregarious growth and aggressive nature
in competing with native flora, was originally introduced to Kerala some 50 years ago from
South America (Fig. 2B) via Sri Lanka. It became a notorious weed in the Kariba lake of
South Africa (Kariba lake, Gambia – hence also known as Kariba weed). In Kerala, this
weed caused problems during 1955-’85 invading almost all fresh water bodies of the State,
menacing agriculture, water transport, fisheries and hydro-electric projects by tangling on
turbines of generators (in Kakki reservoir of Sabarigiri Project, Pathanamthitta Dt.). This
was later effectively managed using biological control method by introducing a native pest
(weevil- Cyrtobagous salviniae) from Brazil and releasing on the Salvinia populations of
Kerala. In addition to Salvinia molesta Mitch., Salvinia auriculata Aubl. is reported from
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 175

F i gg.. 1 : A–H. Diversity centres of pteridophytes of South India. A. Agasthiamala (Western Ghats of
Kerala) Tropical Rain Forest, B. Eravikulam National Park, Kerala Shola forest, C. Kudajadri Karnataka State
shola forest, D . Westerns Ghats of Goa, E . Palni Hills, Tamil Nadu, F. Silent Valley, National Park Virgin
Evergreen forests, Kerala G . WayanadPass-Mixed forest, Kerala and H . Ercaud (Tamil Nadu) Shevaroy Hills
montane vegetation.
176 Indian Fern Journal Volume XXXI (2014)

Fi g. 2 : A–F F.. Aquatic/wetland peteridophytes of South India. A . Azolla pinnata (free floating –
biofertilizer), B. Salvinia molesta (floating weed-primary stage), C. Marsilea minuta (Emergent), D. Regnellidium
diphyllum (Emergent exotic), E . Osmunda hugeliana (Rheophyte) and F. Acrostichum aureum (Halophyte).
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 177

F.. Adaptation of South Indian pteridophytes to various habitats. A . Actinipteris radiata -


F i gg.. 3 : A – F
xerophyte, B . Pteridium revolutum – an exotic terrestrial weed in open areas, C . Pyrrosia heterophylla-
epiphyte, sensitive to air pollution, D . Botrychium daucifolium -riverine ecosystem, E . Asplenium unilaterale
var. unilaterale- lithophytic sciophyte and F. Angiopteris helferiana, a large mesophyte sensitive to dessication
178 Indian Fern Journal Volume XXXI (2014)

Andhra Pradesh and North India (Madhusoodanan, 1989; Madhusoodanan et al. 2014).
Azolla pinnata R.Br. is native to India, which is popular as an effective biofertilizer
in the inundated paddy fields (Sevichan and Madhusoodanan, 1995). Azolla is free floating,
belonging to the same order, Salviniales but differs from Salvinia in its smaller size, absence
of egg-beater hairs and heteromorphic sporocarps. Azolla sporocarps are unisporangiate and
heteromorphic, the female (macro) sporocarp being somewhat oblong cylindrical, with
pointed tip with a single megasporamgium bearing a single megaspore, while the male
(micro) sporocarps are globose and multisporangiate the sporangia born on a branched
receptacle. On the upper lobe of the bilobed leaf, the Azolla bears mucilaginous cavities
housing the cyanobacterium (BGA), Anabaena azollae. This cyanobacterium is capable of
nitrogen fixation, converts the atmospheric nitrogen into ammonia which is usually leached
to the substratum enriching the fertility of the paddy field. The Chinese and Vietnamese
farmers were utilizing Azolla spp. as a bio-fertilizer since 14th century. Azolla spp. can
provide 40% of the nitrogenous requirement of the paddy thus alleviating the detrimental
effects of the application of chemical fertilizers (Madhusoodanan and Sevichan, 1995) (Fig.
2A)
In addition to Azolla pinnata R.Br., the Chinese species, Azolla rubra R.Br. and
A. filiculoides Lam. have been introduced to South India. While the former seemed to be
ideal for the paddy fields of Kerala, the latter is not viable in this area being a temperate
species.
Ceratopteris thalictroides (L.) Brongn. thrives in all the three aquatic forms. During
the first rain of monsoon itself the spores germinate and gametophytes and young
sporophytes are produced and the young sporophyte within 30 days starts growing under
water. In canals they will be uprooted by the flowing water current and floats (the whole
plant body is spongy with air spaces) to be distributed in various places. Later when the
water recedes, they get anchored in shallow water and settle as emergent ferns. They also
propagate through aerial bulbils produced at the axil of rachis and rachilla. Bulbils are
usually produced on vegetative leaves and seldom on sporophylls. Ceratopteris spores also
germinate on wet places, produce gametophytes and sporophytes but perish during summer
succumbing to drought and desiccation. Ceratopteris populations are breeding centers of
fresh water prawns and fishes.
Cyclosorus interruptus (Willd.) H. Ito grows in wet places, paddy fields and semi
wet lands, sometimes gregariously as a weed. It occupies totally covering the abandoned
paddy fields, with its fleshy long creeping rhizome; it grows fast and reticulates the whole
area. It prefers shallow waters with muddy soil. The long creeping cylindrical rhizome, light
green pinnate leaf having dentate margin, and coriaceous texture distinguishes it from other
common ferns of low land.
Marsilea minuta L. is semi-aquatic but can tolerate the scorching summer heat to a
great extent. M. minuta L. produces sporocarps only when the land is dry in summer.
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 179

During rainy season, whenever the water level increases, the petiole length also increases
accordingly sometimes, up to 2m long. The four leaflets at its tip lie flat on the
water surface. Until recently this species was named Marsilea quadrifolia L., but Bhradwaja
(1980) clarified the identity and declared the absence of Marsilea quadrifolia L. in
India (Fig. 2C).
A relative of Marsilea viz., Regenellidium diphyllum Lindman has been introduced
to South India recently as a horticultural plant. The leaf has only two dark green, semi
circular leaflets and the petiole is sturdier. Usually it does not produce sporocarps. The
sporocarps of Marsilea are bean-shaped and psilose but the sporocarps of Regnellidium are
globose and hairy (Fig. 2D).
Isoetes coromandelina L. has rather frequent distribution in Kerala and reported from
Karnataka and Andhra Pradesh. It was found growing in small seasonal ditches of lateritic
hills of North Kerala and also in the sandy paddy fields of Central and Northern Kerala.
But now this has become very rare and is feared extinct from these localities. On laterite
hills, Isoetes grows only up to 10 cm. but in the inundated low land ditches in coastal areas,
it grows up to 70 cm tall. Isoetes also survives the summer with its dormant ‘rhizomorph
‘(corm) which can resurrect by next rainy season. It is too difficult to distinguish this
lycopod when it grows mixed with grasses and sedges. The sedge, Fuirina umbellata Kunth
(Cyperaceae) mimic them when young and deceives the observer.
Isoetes produces narrow cylindrical leaves with spatulate base and pointed tip and
resembles an onion plant or young Cryptocryne (no wonder the plant is wrongly included
among the Aroideae by Adanson).
Forest Streams – Rheophytes
Rheophytes are plants growing in forest streams tenaciously attached to rocks
braving the swift current of water. During monsoon, they submerge under water and emerge
as and when the water level decreases after the rainy season.
Osmunda hugeliana C. Presl (Syn: Osmunda regalis), the ‘Royal Fern’, known for
its elegant parrot green foliage and crown-like fertile portion (tassel) on leaf tips, was very
common in the forest streams of W. Ghats (above 700m altitude), but now restricted to
interior forests showing its sensitivity to anthropogenic activities(Fig2E).
Trigonospora (with two spp. T.ciliata (Benth.) Holttum and T. caudipinna (Ching)
Sledge) is another genus which grows on rocks of forest streams in S. India. They also
remain submerged during rainy season at least for 2-3 months. Filmy ferns like
Didymoglossum bimarginatum (Bosch) Ebihara & K. Iwats. are found associated in this
habitat. Ferns like Thelypteris spp. ( Pseudocyclosorus octhodes (Kunze) Holttum),
Sphaerostephanos subtruncatus (Borz) Holttum. Sphaerostephanos unitus (L.) Holttum. are
larger rheophytes found in S. India. Blechnum colensoi (Hook.) Wakef. is another rare plant
which is found in rheophytic nature in the Western Ghats of Tamil Nadu (Kothayar Hills)
and at Vellaimala above 1000m alt. in well shaded forest streams on granite rocks. This is
180 Indian Fern Journal Volume XXXI (2014)

one species of Blechnum showing heterophylly with the fertile pinnae reduced to thin
cylindrical leaflets showing similarity to Lomariopsid ferns.
On wet rocks
In the wet evergreen forests big granite boulders are usually covered with thin or
thick layer of humus derived from the fallen leaves putrefied during the rainy season. This
provides a special habitat for some specific fern species such as Asplenium formosum Willd.,
A. unilaterale var unilaterale (Lam.) Clarke (Fig. 3E), Araiostegia pulchra (D.Don) Copel.,
Grammitis spp. and filmy ferns (Trichomanes spp. and Hymenophyllum spp.) giving an
elegant and vibrant look covering the whole boulder. Asplenium grevillei Wall. ex Hook.
& Grev. prefers the woody root bases of Myristica trees in ‘myristica swamps’.
Riverbanks
Primitive ferns like Botrychium daucifolium Wall. ex Hook. & Grev. and
Ophioglossum costatum R.Br. share a unique type of habitat. They perennate through the
rhizome during summer and rainy season in the loamy/sandy soils of forest stream banks
and emerge after the retreat of monsoon during Nov-Dec (winter season) B.daucifolium
Wall. ex Hook. & Grev. usually produces one or two leaves with adnate fertile portion
(spike) and live for 2-3 months and by March wither away except the perennial rhizomatous
portion which remains subterranean in a dormant stage until the onset of the next winter
season which is comparatively mild in the W. Ghats (Fig 3D)
Halophytes
Acrostichum aureum Linn. is the only fern which can grow in brackish waters in
South India. This fern is found all over the West Coast and East Coast in backwaters,
mangroves in shallow waters and muddy banks. Of the two species of Acrostichum only A.
aureum L. is found in India, the other species viz., A. speciosum Willd. which has
acuminate apex for the leaflets is found in Malysia and Australia and can tolerate more
inundation than A.aureum L. (Fig. 2F)
There are no marine pteridophytes.
Ter
errr estr ial Pter
estrial idoph
Pteridoph ytes
idophytes
Wet land plants:
Ferns like Lygodium microphyllum (Cav.) R. Br. and Cyclosorus interruptus (Willd.)
H. Ito prefer wet muddy soil rich in humus. While, L. microphyllum (Cav.) R. Br. prefer
semi-shaded areas with supporting vegetation, C. interruptus (Willd.) H. Ito prefers open
fields with bright sunlight. Lygodium is a leaf climber, goes up to 10 m in some areas.
They prefer banks of forest ponds or muddy stream banks.
Sciophytes – the shade loving ferns
Epiphytes
In fact, most pteridophytes prefer wet shaded habitats which cover the forest floor-
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 181

TABLE 1 : Ecolo
Ecologg ical ca
catt eeg
g o rries
ies of Pter idoph
Pteridoph ytes of S
idophytes S.. India

Sl. Type Candida te/T


Candidate/T ypical Species
te/Typical
No.
1. Hydrophytes Fresh Water Salvinia molesta Mitch.
a) Free Floating Azolla pinnata R. Br.
b) Submerged Isoetes corromandelina L.
(exposed during summer)
Microsorum pteropus (Bl.) Copeland
(exposed during summer)
c) Emergent Ceratopteris thalictroides (L.) Brongn.
(also grow submerged and free floating and as ‘sudd’)
Marsilea minuta L.
2. Halophytes (Brackish water) Acrostichum aureum L.
3. Wetland plants Lygodium microphyllum (Cav.) R. Br.
a) Muddy soil Cyclosorus interruptus (Willd.) H. Ito
Stenochlaena palustris (burm. f.) Bedd. (climber)
Asplenium formosum Willd.
b) Wet rocks Hymenophyllum spp.
Trichomanes spp.
Araiostegia spp.
Grammitis spp.
4. Rheophytes Osmunda hugeliana C. Presl
(rocky forest streams) Thelypteris (Spherostephanos) arbuscula
Thelypteris (Trigonopora) ciliata (Wall. ex Benth.) Ching
Bolbitis presliana (Fee’) Ching
5. Loamy stream beds Botrychium daucifolium Wall. ex Hook. & Grev.
6. Sciophytes
a) Epiphytes Drynaria quercifolia(L.) Js. Sm.
Pyrrossia heterophylla (L.) M.G. Price
Pyrrossia lanceolata (L.) Farw.
Vittaria elongata Sw.
Microsorum nigrescens
Microsorum punctatum (L.) Copeland
Microsorum zipelli (Bl.) Ching
Contd.
182 Indian Fern Journal Volume XXXI (2014)

Contd
Contd.. Ta b le 1

b) Lithophyte Bolbitis (Egenolfia) appendiculata (Willd.) K. Iwats


Bolbitis (Egenolfia) asplenifolia (Bory) K. Iwats
Bolbitis (Egenolfia) virens (Hooker & Grev.) Schott
c) Oxalophytes Macrothelypteris torresiana (Gaud.) Ching
(humus rich soil) Athyrium polypodoides
Athyrium hohenackerianum (Kunze) T. Moore
Deparia boryana (Willd.) M. Kato
7. Heliophytes (open sun) Pityrogramma calomelanos (L.) Link
Christella (Thelypteris) dentata (Forssk.) Brownsey &
Jermy
Cheilanthes tenuifolia (Burm. f.) Sw.
Pteridium revolutum (Blume) Nakai
Blechnum orientale L.
Dicranopteris dichotoma (Thunb.) Bernh.
8. Xerophytes Actiniopteris radiata (Sw.) Link
(arid rocks and soil) Selaginella wightii Hieron
Parahemionitis cordata (Roxb. ex Hook&Grev.) Fraser-
Jenk.
Doryopteris concolor (Langsd. et Fisch.) Kuhn
Chielanthes opposita Kaulf.
No parasites and no marine pteridophytes.
soil rockys, trees trunks, fallen logs, etc. They also make large populations. Probably,
Drynaria quercifolia (L.) Js. Sm. is the most common fern in the West Coast of Peninsular
India found on tree trunks, boles, walls and tiled roof tops, etc. They are conspicuously
heterophyllous with sterile ‘pocket leaves’ and large pinnatified fertile leaves on thick
fleshy rhizome covered with golden paleae. The fertile leaves fall during summer but the
pocket leaves remain even after dried and dead attached on the rhizome till the next
season providing food and moisture to the perennating rhizome. The other epiphytic ferns
in the low land coastal area are Pyrrosia lanceolata (L.) Farw., Vittaria elongata Sw.,
Pyrrosia heterophylla ( L.) M.G. Price (syn: Drymoglossum piloselloides(Linn.) Presl),
etc. (Fig. 3C).
The major epiphytes of higher elevations (above 700 m) include lycophytes like
Huperzia phlegmaria Rothmaler, H. macrostachys (Hook. ex Spring) Holub ferns like,
Microsorum punctatum (L.) Cpoeland, M. membranaceum (D.Don) Ching, Asplenium
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 183

phyllitidis D.Don Lepisorus nudus (Hook.) Ching, Antrophyum reticulatum (G.Forst.)


Kaulf. and filmy ferns such as Crepidomanes bilabiatum (Nees & Blume) Copel., C.
saxifragoides (C.Presl) P.S. Green etc. Phymatopteris montana (Sledge) Pic. Ser. is found
only above 800 m alt., widely distributed in the W. Ghats of Kerala, Karnataka and Tamil
Nadu (Nampy and Madhusoodanan, 1998). Many epiphytes such as Microsorum spp.
Elaphoglossum spp. Vittaria spp. Pyrrosia spp. also grow on wet humus rich rocks.
On forest floor
Members of Thelypteridaceae such as Christella spp. and Thelypteris spp.
(Macrothelypteris, Pronephrium, Ampelopteris, and Amphineuron), members of Athyriaceae
(Athyrium spp., Diplazium spp., Deparia spp.) and Lomariopsidaceae (Bolbitis spp.) are
shade loving ferns. Pteris spp. occupy the forest floor of the forests of Western Ghats.
Whereas Pteris spp. are also found in semi evergreen forests. The members of
Thelypteridaceae and Athyriaceae are found in evergreen forests though some species such
as Athyrium hohneckerianum (Kunze) T. Moore and Thelypteris ( Christella) dentata
(Forssk.) Brownsey & Jermy are found in the lower elevations in semi exposed areas
Heliophytes
Though pteridophytes usually prefer shaded or partially shaded areas, some species
are adapted to semi arid conditions prevailing at least part of the year which can tolerate
high temperature to a great extent. Christella dentata (Forssk.) Brownsey & Jermy is one
fern in open areas in lower elevations but they prefer wet soil forming small colonies. The
Silver fern, Pityrogramma calomelanos (L.) Link an exotic fern naturalized in S. India and
elsewhere, is widespread all over S. India in the lower elevations in open areas, margins of
paddy fields vertical muddy walls and lawns as a weed. Another species called Golden fern,
P. chrysophylla (Sw.) Link is found only above 800m, common in hill stations such as
Kodai, Ooty, Munnar and Mercara. Cheilanthes tenuifolia (Burm F.) Sw. and Parahemionitis
cordata (Roxb. ex Hook. & Grev.) Frase-Jenk. are other ferns found in open grass lands of
lateritic hills. Pteidium revolutum (Blume) Nakai, the notorious ‘Bracken fern’ introduced
from Europe, has become a serious weed in high land plantations (tea, coffee, etc.) and
invaded the grass lands of ‘shola’ vegetation (Fig. 3B). They grow in open sun with the
creeping rhizome deep in the soil and the large leaves covering the whole area and can
withstand forest fires. In Eravikulam National Park, Kerala, this aggressive weed has
invaded the grass lands posing problems for grazing by the ‘Nilgiri Tahr’,( Nilgiritragus
hylocrius), an endangered wild goat species and other herbivores.
Xerophytic pteridophytes
The rain shadow areas on the eastern slopes of W. Ghats, such as Chinnar, some
slopes of Palni Hills and Kudachadri Hills receive little rain annually and the climate is
montane and the terrain is rocky with patches of thin layered soil. This habitat
accommodates special type of vegetation adapted to tolerate xeric conditions (Fig 3A). The
184 Indian Fern Journal Volume XXXI (2014)

common ferns of these areas are Actiniopteris radiata (Sw.) Link and Cheilanthes opposite
Kaulf. These are beautiful and curious ferns found attached on granite rocks with a thin
film of soil. Selaginella wightii Hieron, which looks like a Lycopodium spp. is also found
in this habitat as small carpets.
Biolo
Biologg ical Associa tions
Associations
Some ferns share the same ecological niche and form distinct communities. In
S.India, the Dicranopteris spp. which prefers vertical earth cuttings and forming thicket -
like colonies often share the habitat with Lycopodiella cernuua (L.) Pi. Ser. and Blechnum
orientale L. They usually share the same habitat and grow together. However, what factors
make them share the same microhabitat to form a conspicuous community is unknown.
Ecological Indicators
Some ferns are being used effectively as ecological indicators by local people.
Acrostichum aureum L. indicates the presence of salinity and unpalatable hard water. This
place is usually avoided by people for digging drinking water wells and house constructions.
This area is also not good for usual paddy cultivation. But found to be good for coconut
plantations. However, a special variety of rice called ‘Pokkali’ and ‘Kaipad’ which can
tolerate salinity are cultivated here. Pyrrosia heterophylla (L.) M.G. Price (popularly known
as ‘seethathaali’ in Malayalam) is very sensitive to air pollution, especially to the smoke
emitted from motor vehicles. Hence the presence of this epiphytic fern is considered to be
a reliable indication for fresh, pollution free air. Presence of Ceratopteris thalictroides (L.)
Brongn. in wet land areas is considered to be an indication for the suitability of inundated
paddy cultivation. Presence of filmy ferns shows high humidity in the air. The xerophytic
ferns indicate an arid climate, less humidity and high day temperature. Angiopteris
halferiana C. Presl is very sensitive to desiccation and high temperature and shows lodging
of leaves as if the plant is totally exhausted. If watered, within 30 min all leaves will return
to the previous healthy position and this acts as an indicator of the indigence of water in
ferneries and nature. (Fig.3F).
DISCUSSION

Until some fifty years ago, the Southern India was very rich in Pteridophytes,
especially in the Western Ghats area, in frequency and abundance. However, the post IInd
World War famine necessitated the “grow more food” policy which inspired the people to
migrate from low land to the forests of high land for cultivation. This has resulted
unprecedented and indiscriminate destruction of forests of this area resulting the extinction
of many ferns from the locality and led many to the rare and endangered category.
Some ferns of South India have become very rare and are restricted to one or two
localities at present. Schizaea digitata (L.)Sw. which is reported only from Palode
(Madhusoodanan and Sathish, 1986) Thiruvananthapuram Dt, Kerala. Helminthostachys
zeylanica (L.)Hook. has been reported from several parts of Kerala, but recent survey shows
Madhusoodanan et al. : Ecology of South Indian Pteridophytes 185

that it is restricted to a protected area (oldest Teak Plantation) at Nilambur, Kerala. This
fern prefers loamy soil with rich humus. Blechnum colensoi (Hook.) Wakef. is reported
from Kothayar (Tamil Nadu) and is available at Vellarimala, Kozhikode Dt, Kerala but, very
little in number.
Ampelopteris prolifera (Retz.) Copel. is reported only from Karnataka (Rajagopal
and Bhat, 1998) and Chanthanathode, Wayanad, Kerala (Leena and Madhusoodanan,1992).
From Wayanad now this fern has disappeared. Humata repens (L.f.) Diels, a curious
epiphytic fern with restricted distribution at Sabarimala, Kerala though reported earlier from
Wayand, Kerala (Nayar and Gheevarghese, 1993) has disappeared from that locality.
The destruction of natural vegetation is relentlessly going on. Unless immediate
steps like creating protected areas for the in situ conservation are taken, these ecologically
sensitive plants will face total devastation from this area in near future. Regeneration of
such vegetation and establishment of lost species are cumbersome and almost impossible.
REFERENCES

BEDDOME R H 1863 The Fer ns of Southern India Gantz Bros Madras (Re print Ed 1970) Today and
Tomorr ow’s Printers & Publishers New Delhi
H O LT T U M R E 1 9 6 8 A R ev i s e d F l o r a o f M a l a y a Vo l I I F e r n s o f M a l a y a G ov t P r i n t i n g O ffi c e
Singapore
LEENA K R & MADHUSOODANAN P V 1992 Ecolo gy and Distribution of Thelypter oid Fer ns of S
India Indian Fern J 9 : 174-183
MADHUSOODA NAN P V 1989 Taxonom y and distribution of the g enus Salvinia Senguier in the Old
Wo r l d I n : K H S h i n g a n d K U K r a m e r ( E d s . ) P r o c I n t S y m p S y s t P t e r i d C h i n a ( 1 9 8 8 )
Science and Technol Press Beijing (China) Pp 55-68
MADHUSOODA NAN P V & NAMPY S 1998 Biodiversity of F erns and Fer n Allies of Kerala (India)
J Econ Tax Bot 2 2 : 183-189
M A D H U S O O DA NA N P V A N D S AT H E E S H C 1 9 8 6 Sc hizaea dig itat a (L.)Sw. from South India Ind J
Bot 9 : 144-146
M A D H U S O O DA NA N P V S I J I M O L P S & R A J E S H K P 2 0 0 1 P t e r i d o l o g y i n S o u t h I n d i a - A
Retrospection Indian Fern J 1 8 : 18-34
MADHUSOODANAN P V SREERENJINI V K SMITHA R B & PRAKASHKUMAR R 2014 Invasion of]
Salvinia molesta D.S. Mitchell. (African Payal) in Kerala and its Management Indian Fern
J 3 1 : 156-166
M A J E E D K A L E E NA K R & M A D H U S O O DA NA N P V 1 9 9 4 E c o l o g y a n d di s t r i bu t i o n o f
Lomariopsid Ferns of Kerala Res J Pl Environ 1 0 : 71-74
MANICKAM V S & NINAN C A 1984 Ecolog ical Studies on the Fern Flor a of Palani Hills (S. India)
Today and Tomorr ow’s Printers & Publishers New Delhi
MEHLTRETER K WALKER L R & SHARPE J M Fern Ecolog y Cambridge Univ Press UK
NAYAR BK AND GEEVERGHESE K K 1993 F ern Flora of Malabar Indus publ Co New Delhi
NAYAR B K & MADHUSOODANAN P V 1979 The World’s Worst Weed Science Reporter (CSIR) 1 6 :
15-21
186 Indian Fern Journal Volume XXXI (2014)

PAGE C N 1979 The diver sity of fer ns An Ecolog ical Per spective In : A F Dyer Ed The Experimental
Biology of Ferns Pp 551-589 Academic Pr ess London
P U L L A I A H T A M E E R A H M E D & A M RU T H A L A K S H M I P 2 0 0 3 P t e r i d o p h y t e s i n A n d h r a P r a d e s h
India Regency Publications New Delhi
RAJAGOPAL P K & BHAT K G 1998 Pteridophyte Flor a of Karnataka Sta te India Indian F ern J 15 :
1-28
S E V I C H A N P J & M A D H U S O O DA NA N P V 1 9 9 6 E c o l og i c a l a n d E c o n o m i c I m p a c t s o f A z olla rubr a
as Biofertilizer in the acidic paddy fields of Kerala J Ecobiol 8 : 61-65
S H A R P E M J M E H LT R E T E R K & WA L K E R R L 2 0 1 0 E c o l o g i c a l I m p o r t a n c e o f F e r n s I n : F e r n
Ecology Ed Mehltreter et al Pp 1-21 Cambridge University Press UK
SLEDGE W A 1982 An annotated Check List of the Pteridophyta of Ceylon Bot J Linn Soc 8 4 : 1-30
Indian Fern J. 3 1 : 187-189 (2014) ISSN 0970-2741

LIFE MEMBERS OF INDIAN FERN SOCIETY


2009-2014

2009 7. Dr. M V KALE


1. PROF A S AHLUWALLIA Jaysingpur College, JAYSINGPUR 416101
Depatment of Botany, MAHARASHTRA
Panjab University, manishavkale@gmail.com
CHANDIGARH-160014 8. MR RAJU ANTHONY
phykos@pu.ac.in Technical Officer, TBGRI,
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DELHI 110085 9. DR. BASHA, SHAIK KASIM MUNIR
anjulisood@gmail.com Associate Professor, NBKR, Research Centre,
3. MR RAHUL JAGANNATH MAHAMUNI Affiliated to S V University
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47- Shakti Vihar, Majra H. No. 87, Phase VI,
DEHRADUN -248171 S A S Nagar (Mohali) 160055 Punjab
(Uttarakhand). 12. DR NILIMA BHARDWAJA,
singh_june07@rediffmail.com Lecturer in Botany, Govt. College,
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LIFE MEMBER Paid Rs 5300/- Vill. Pritinagar Co-operative Colony
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balkararya@gmail.com
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14. PROF. ANNETTE SCHÖLCH 20. Dr. MAAJEED HUSSAIN WANI,


Langgewann 22 D-69121 Wagoora, District Baramulla
Heidelberg (Germany) Jammu & Kashmir-193103
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16. Dr. HEER CHANDRA NEGI
Assiatant Professor, Department of Botany,
Assistant Professor, Botany Department,
University of Kalyani,
Govt. Degree College,
Kalyani-741235 (West Bengal)
Rekong/Peo - 172107.
sudhaguptain@gmail.com
District Kinnaur (H. P.)
23. Dr. M JOHNSON,
hcn0918@gmail.com
Assistant Professor,
17. DR. VINEETH KUMAR RAWAT
Department of Plant Biology and Plant
Scientist C
Biotechnology,
Botanical Survey of India,
St. Xavier's College (Autonomous)
Arunachal Pradesh Regional Centre,
Palayamkotai-627002 (Tamil Nadu)
ITANAGAR - 791111
ptcjohnson@gmail.com
rawat_vk2107@rediffmail.com
24. Dr. AJIT KR DAS
18. DR. MALAY BHARTI,
Dept of Ecology & Environment Science,
Assistant Professor,
Assam University, Silchar-788011
Dept. of Botany, Doranda College,
ajitkumardas2009@rediffmail.com
Ranchi-834002 (Jharkhand)
ajitkumar2009@rediffmail.com
malay_bharti@rediffmail.com
25. Dr. MRS M VANDANA E KISHORE
19. Dr. SANJAY KUMAR RAMLAL
‘Shri’ Opp SGM College, Vidyanagar,
RAHANGDALE
Behind Yaser Complex, Karad-415124
Assistant Prof, Botany
(Maharashtra)
Vishal Kranti Nagar, Ale
26. Mr. SHAKOOR AHMED MIR
Tah Junnar, Dist. Pune-412411
Dept of Botany, Jamia Hamdard,
sanjay2@hitmail.com
Hamdard Nagar, New Delhi-110062
soy_sanjay@yahoo.com
shakoorsam@gmail.com
Indian Fern Journal Volume XXXI (2014) 189

2013 2014
27. Dr PREM LAL UNIYAL 34. Mr. DEVENDRA TRIPATHI
Dept. of Botany, University of Delhi, JRF
Delhi-110007 Western Regional Centre
BSI PUNE 07875997263
uniyalpl@rediffmail.com
botanydevendra@gmail.com
28. DR RUCHI SRIVASTAV,
35. PROF. PURSHOTAM KAUSHIK
395/31 Raj Bhawan, Kashmiri Mohalla,
Dept of Botany & Microbiology,
Lucknow-226003
Gurukul Kangri Vishwavidyala,
contactrsri@gmail.com
P & T Gurukul Kangri, Haridwar-249404
29. Ms. KANTA MEENA
purshotam.kaushik@gmail.com
Lecturer Dept of Botany, 36. Dr SHREEKAR PANT
MLV Govt College, Asst Prof.
Bhilwara-311001 (Rajasthan) Centre for Biodiversity Studies,
kantameenalectbotany@gmail.com BGSB University, Rajori-185131 (J & K)
30. Prof. R P KANTHER, shreekarpant.2@rediffmail.com
Pteridophyte Research Lab, 37. Mr. RAVI PRATAP GAUTAM
Deptt. of Botany, S. D. Govt. College, Senior Lecturer
Beawar-305901 (Rajasthan) Department of Botany, St. Andrew’s College,
rpkanther@gmail.com Gorakhpur- 273001 (Uttar Pradesh)

31. PROF. I B PRASHER rpgautam0288@gmail.com


38. MR. SHASHANK KUMAR SINGH
Botany Department, Panjab University,
(Research Scholar)
Chandigarh.
Department of Botany, St. Andrew’s College,
chromista@yahoo.co.in
Gorakhpur- 273001 (Uttar Pradesh)
32. DR. RICHA PUNETHA,
shashank2771986@gmail.com
157, Bajethi Ward,
39. DR. O P SUTHAR
Near Post Graduate College, Senior Lecturer,
Pithoragarh-262502 D R J Govt. Girls College,
punethan_bot@yahoo.co.uk BALOTRA- 342008 (Ajmer) Rajasthan
33. DR. KAMLESH BHAKUNI opsuthar59@yahoo.co.in
c/o Sh. RAMESH SINGH BISHT, 40. DR. RAKESH HARSH
Near Har LAl Petrol Pump, DWARIKA, Dharam Nagar Dwar,
Tanakpur Road, Pithoragarh- 262501 Near Periwal Gas Agency, Bikaner (Raj.)
kammubhakuni@yahoo.com
Indian Fern J. 3 1 : 190-200 (2014) ISSN 0970-2741

BIBLIOGRAPHY OF INDIAN PTERIDOLOGY (201


(20133 -201
-20144)
[Compiled by Dr. R. Mukhopadhyay & Prof. S. P. Khullar]

A L O K M A H E S H P R A S A D , C H AU H A N D K & K H A R E E G 2 0 1 4 Pa l y n o l o gi c a l I nve s t i g a t i o n i n t h e
Middle Miocene Sediments of Tanakpur, Champawat Distr ict, Uttar akhand . Proceedings 101 st
Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 217-218.
ANJUM S , SHREEKAR PANT, HC PANDE & TSEWANG RINCHEN 2014 Ecolo g ical Observations on
the Fern Flora of Thannamandi District Rajouri, Jammu & Kashmir, India. Indian Forester,
1 4 0 ( 1 2 ) : 1 2 3 5 - 1 2 4 4.
A N J U M S , PA N T S & K H U L L A R S P 2 0 1 4 D ive r s i t y a n d d i s t r i b u t i o n p a t t e r n o f p t e r i d o p h y t e s o f
Rajour i district: a pr eliminary sur vey [Abstr act] National Confer ence on Modern A pproac hes
to Pteridophytes: Biolog y, Biodiversity and Bioresource , 20-21 st Dec . , CSIR-IHBT, Palampur,
H.P. , India Abstr act Vol. pages 78-79.
BA N DYO PA D H YAY M 2 0 1 4 L o s s o f f e r n d ive r s i t y b y p r ove r b i a l d i c t u m o f ‘ K h a n a ’ [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresour ce, 20-21 st Dec . , CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 81.
BA N E R J E E A , K U M A R I A , L A L B & K U M A R A 2 0 1 4 U s i n g g e o s p a t i a l t e c h n i q u e s f o r ma p p i n g o f
p o t e n t i a l d i s t r i bu t i o n o f D i p l a z i u m e s c u l e n t u m ( R e t z . ) S w. i n K a n g r a r eg i o n o f H i m a c h a l
P r a d e s h [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l og y,
B i o d iv er s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
page 41.
BEHERA SK & KHARE PB 2014 Pteridophytes of Kendujhar district, Odisha [Abstract] National
C o n fe re n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l og y, B i o d ive rs i t y a n d B i o r e s o u r c e ,
20-21 st Dec ., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. page 29.
B E N N I A M I N A 2 0 1 4 Ta x o n o m i c s t u d y o f fa m i l y D r y o p t e r i d a c e a e i n n o r t h - e a s t I n d i a [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec. , CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pa g es 25-26.
BENNIAMIN A 2014 Pteridophytes of Manipur [Abstract] Na tional Conference on Moder n Appr oaches
to Pteridophytes : Biology, Biodiver sity and Bioresource, 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. page 32.
B E N N I A M I N A , R AWAT V K & T R I PAT H I D 2 0 1 4 F e r n s a n d f e r n a l l i e s o f M a h a r a s h t r a [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresour ce, 20-21 st Dec ., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pag e 34.
BENNIAMIN A & TRIPATHI D 2014 A preliminary sur vey of pteridophytes fr om Biligirir anga S wam y
Temple Wildlife Sanctuar y, Karna taka [Abstract] National Conf erence on Modern Approac hes
to Pteridophytes: Biolo gy, Biodiver sity and Bioresourc e, 20-21 st Dec . , CSIR-IHBT, Palampur,
H.P. , India Abstr act Vol. pages 30-31.
B E R A S , G U P TA S , K H A N M A , D E A & M U K H O PA D H YAY R 2 0 1 4 F i r s t m e g a f o s s i l ev i d e n c e o f
C ya thaceous tr ee f er n from the Indian Cenozoic. J. Ear th Syst. Sci. 1 2 3 no.6 A ugust, 1433-
1438.
BHAKUNI K & PUNETHA R 2014 An updated list of Pteridophytes of Gori valley (Central Himala ya)
[Abstract] National Conference on Modern Approaches to Pteridophytes : B i o l o g y,
st
B i o d iv er s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
page 30.
Bibliography of Indian Pteridology (2013-2014) 191

B H A N DAWAT A , S I N G H G , S H A R M A V, S WA R N K A R M K , S I N G H A K , S I N G H P, S I N G H P S &
SHARMA RK 2014 Mining of transcriptome sequencing of Equisetum ramosissimum for
development of functionally relevant SSR marker resources [Abstract] National Conference
on Modern Appr oaches to Pteridophytes: Biolo gy, Biodiversity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 69-70.
B H A R DWA JA T N 2 0 1 4 M a r s i l a c e o u s f e r n s : p h y l o g e n y a n d i n t e r g e n e r i c r e l a t i o n s h i p s - b i o g e o g r a p h i c
dimension [Abstr act] National Confer ence on Modern A pproac hes to Pteridophytes : Biolo g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 13-14.
BHARTI MALAY & RAM PRAVESH 2014 Phytochemical analysis of Lygodium flexuosum (L.) Sw. and
Ampelopteris prolifera (Retz.) Copel. of Ranchi district of Jharkhand, India. Proceedings
101 st Session Indian Science Congress Jammu Feb 3-7,
2014, Part II Plant Sciences: 2312-234.
C H A K R A B O RT I K , S A D H U K H A N R & BA N DYO PA D H YAY M 2 0 1 4 P o t e n t i a l o r n a m e n t a l f e r n s o f
Gangetic West Bengal [Abstract] National Conference on Modern Approaches to Pteridophytes :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a
Abstr act Vol. pages 85-86.
C H AW L A K D , YA DAV B L & B H A R DWA JA T N 2 0 1 4 E x p e r i m e n t a l i n d u c t i o n o f p a r t h e n o g e n e s i s i n
some xeromorphic species os Marsilea Linn. [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P. , India Abstract Vol. pag e 65.
C H E E M A H 2 0 1 4 H o m o a n d H e t e r o p h a s i s i n t h e in vitro D i f f e r e n t i a t i o n o f F e r n s . P r o c e e d i n g s 1 0 1 s t
Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 118-119.
CHEEMA H 2014 Micropropagation of some aquatic homo and heterosporous ferns [Abstract]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pag es 17-18.
DEEPA J & PARASHURAMA TR 2014 Pteridophytic Flora of Mullaiy anag iri, Kar nataka, South India.
P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b . 3 - 7 , 2 0 1 4 , P a r t I I P l a n t
Sciences : 222-223.
D O G R A D , S H A R M A S & S H A R M A S K 2 0 1 4 Ay u r v e d i c p r o s p e c t iv e o f f e r n s - t h e i r c h e m i c a l ,
pharmacological and medical descriptions [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstr act Vol. pages 45-46.
D O L M A S K , K U M A R I A & E S WA R A R E D DY S G 2 0 1 4 R e p e l l e n t a c t iv i t y o f f e r n ex t r a c t s a ga i n s t
tobacco ca terpillar, Spodoptera litura [Abstr act] National Conf erence on Moder n Appr oaches
to Pter idophytes: Biology, Biodiversity and Bioresour c e, 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. pag es 91-92.
DUBAL K & KALE M 2014 Ethnomedicinal and phytochemical screening of Tectaria coadunata (Wall.
ex Hook. & Gr ev.) C. Chr. from nor th Wester n Ghats (India) [Abstract] Na tional Conf erence
on Modern Appr oaches to Pteridophytes: Biolo gy, Biodiversity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 49.
D U D A N I S N , M A H E S H M K , S U B A S H C H A N D R A N M D & R A M A C H A N D R A T V 2014 Conservation
of endemic pteridophyte habitats in the wet evergreen forests of Uttara Kannada [Abstract]
192 Indian Fern Journal Volume XXXI (2014)

N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o gy, B i o d ive r s i t y a n d
Bior esource, 20-21 st Dec., CSIR-IHBT, Palampur, H.P., India Abstract Vol. pag es 78.
G A N G U LY G , T I WA R I B K & M U K H O PA D H YAY R 2 0 1 3 P hy t o c h e m i c a l a m a l y s i s a n d a n t i m i c r o b i a l
a c t i v i t y o f c r u d e ex t r a c t s a n d ex t r a c t e d p h e n o l s f r o m a l i t h o p h y t i c f e r n A r t h r o m e r i s
wallichiana (Spreng.) Ching. Bionature 3 3 (1) : 1-7.
G A N G U LY G & M U K H O PA D H YAY R 2 0 1 4 R e p r o d u c t iv e s u c c e s s o f s o m e p o l y p o d i a c e o u s f e r n s o f
south Sikkim in relation to venation architecture, soral position and soral size [Abstract]
National Symposium on Plant Diversity: Structure, Function, Utilization and Conservation
D e c e m b e r 4 - 6 , B o t . S o c . B e n g a l , C A S , D e p t . o f B o t a ny, U n ive r s i t y o f C a l c u t t a , Ko l k a t a ,
700019 Abstract Vol. pa ge 4.
G AU TA M R P, D O M I N I C R A J K U M A R S , S R I VA S TAVA S K & S I N G H S K 2 0 1 4 P a l y n o l o g i c a l a n d
stomatal studies on a few selected pter idophytes of Almora [Abstract] National Confer ence on
M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . ,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. pages 57-58.
G H A DAG E D M , G H O R PA D E P N , D U BA L K N & K A L E M V 2 0 1 4 G C - M S P r o fi l e o f Ly g o d i u m
f l e x u o s u m ( L . ) S w ( Ly g o d i a c e a e ) a M e d i c i n a l F e r n f r o m N o r t h We s t e r n G h a t s . P r o c e e d i n g s
101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 233-234.
G H O S H D K & C H A K R A B O RT Y K 2 0 1 4 P e r f o r m a n c e o f A z o l l a p i n n a t a a n d i t s e ff e c t o n t u r m e r i c
c u l t ivat i o n i n r e d l a t e r i t i c z o n e o f We s t B e n ga l [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. pages 90-91.
G O S WA M I B H AT TAC H A RY YA N , H A L D E R S & PA L P K 2 0 1 4 A d a p t iv e s i g n i f i c a n c e o f s t o m a t a
among aquatic Pteridophytes in different ecological settings [Abstract] National Conference
on Moder n A pproaches to Pteridophytes: Biolo g y, Biodiver sity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 37.
G O S WA M I H K 2 0 1 4 Po p u l a t i o n g e n e t i c s o f va r i a b l e s t o u n d e rs t a n d a d a p t ive s t r a t eg i e s o f s p e c i e s i n
n a t u r e [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 14-15.
G O S WA M I N I RU PA M A 2 0 1 4 P h y t o c h e m i c a l s t u d y a n d eva l u a t i o n o f a n t i f u n g a l a c t iv i t y o f
A c r o s t i c h u n a u r e u m L . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 ,
2014, Part II Plant Sciences : 232
G U P TA C S & N I L I M A B H A R DWA J 2 0 1 4 R e p r o d u c t ive B i o l o gy o f a m e d i c i n a l l y I m p o r t a n t F e r n :
A c t i n i o p t e r i s ra d i a t a ( S w. ) L i n k . P ro c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u
Feb 3-7, 2014, Part II Plant Sciences : 236.
G U P TA N , BA L A M , S I N G H B , L A L B & K U M A R I A 2 0 1 4 S e c o n d a r y m e t a b o l i t e s p r o fi l i n g o f
Diplazium s p e c i e s [ A b s t ra c t ] N at i o n a l C o n fer e n c e o n M o d e r n A p p r o a ch e s t o P t e r i d o p hy t e s :
B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstr act Vol. pag es 73-74.
H A R S H R & S H E K H AWAT S 2 0 1 4 L i g n i t e s a m p l e s f ro m R a j a s t h a n - A S t u d y o f P l a n t M i c rof o s s i l s .
P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I P l a n t
Sciences : 217-218.
HARSH R, PUROHIT SN & SHARMA BD 2014 Reproductive biology of ferns of Rajasthan –
Gametophytes [Abstract] National Conference on Modern Approaches to Pteridophytes:
Bibliography of Indian Pteridology (2013-2014) 193

B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstract Vol. page 61.
H E G D E S , 2 0 1 4 T i s s u e c u l t u r e o f F e r n s P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u
Feb 3-7, 2014, Part II Plant Sciences : 119-120.
H E G D E S , M O R A J K A R S , S A J E E V S, H A L A N K A R N & F E R NA N D E S H 2 0 1 4 A D NA B a r c o d e f or
Stenochlaena palustris: an edible medicinal fern [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. pag e 69.
H U S S A I N S , YA DAV B L & M E E NA K A N TA 2 0 1 4 I m p a c t o f h a b i t a t c o n d i t i o n s o n p h o t o s y n t h e t i c
pigmwnts in some pteridophytic soecies from Kumbhalgarh wildlife sanctuary of Rajasthan
[Abstract] National Conference on Modern Approaches to Pteridophytes : B i o l o g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
page 40.
JA I S H E E N & C H A K R A B O RT Y U 2 0 1 4 A n a l y s i s o f a n t i o x i d a n t a c t iv i t i e s , b i o a c t ive c o m p o u n d s a n d
phenolics from three ferns growing in North Bengal region [Abstract] National Conference
on Modern Appr oaches to Pteridophytes: Biolo gy, Biodiversity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 71.
JANGIR BL, BIND RB , LAUREN DR & SOMVANSHI R 2014 Survey, identification, to xin estina tion
and toxicopathological effects of Pteris cretica feeding in Syrian Golden Hamsters
(Mesocricetus auratus) [Abstract] National Conference on Modern Approaches to
P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r,
H.P. , India Abstract Vol. pag es 88-89.
J O H A R I D , S I N G H A P & K H A R E P B 2 0 1 4 S t u d i e s o n r e p r o d u c t iv e b e h av i o r a n d g a m e t o p h y t e
development in Pteris biaurita Limm. and Microlepia speluncae (L.) Moore [Abstract]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pag es 64-65.
K H A R E P B 2 0 1 4 W h a t i s S a n j e eva n i ? [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o
P t e r i d o p h y t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r,
H.P. , India Abstract Vol. pag e 15.
KHOLIA BS 2014 Maidenhair ferns of Sikkim Himalaya : Diversity and future directions for
domestication. Proceedings 101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part
II Plant Sciences : 224.
KHOLIA BS 2014 On the recollection of a rare felt fern Pyrrosia boothii in Sikkim Himalaya
[Abstract] National Conf erence on Modern Appr oaches to Pteridophytes: Biology, Biodiversity
and Bior esource, 20-21 st Dec ., CSIR-IHBT, Palampur, H.P. , India Abstract Vol. pag e 25.
K H O L I A B S 2 0 1 4 A r a r e a n d e n d a n g e r e d f e r n , O p h i o g l o s s u m p e n d u l u m L . - n ew r e c o r d t o S i k k i m
w i t h a n o t e o n i t s r e d i s c o v e r y, d i s t r i bu t i o n a n d h i s t o r i c a l b a c k g r o u n d [ A b s t r a c t ] N a t i o n a l
C o n fe r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u rc e ,
20-21 st Dec . , CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pa ge 79.
KHULLAR SP 2014 I n n o va t ive Te c h n o l o g i e s and B i o d iv e r s i t y, Ta x o n o m y, C o n s e r va t i o n and
D eve l o p m e n t s i n P t e r i d o p hy t e s . P r e s i d e n t i a l A d d r e s s . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n
Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 06-90.
KHULLAR SP 2014 Fern taxonomy and some interesting findings in the Pteridophyte flora of
Himachal Pr adesh, Western Himalaya [Absatract] National Confer ence on Moder n Appr oaches
194 Indian Fern Journal Volume XXXI (2014)

to Pter idophytes: Biolo gy, Biodiver sity and Bioresource , 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. pag es 7-11.
K H U L L A R S P, V E R M A S U N I L K U M A R & P R A S H E R I B 2 0 1 4 P t e r i d o p hy t i c F l o r a o f H i m a c h a l
P r a d e s h . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I
Plant Sciences : 224-225
K U M A R A 2 0 1 4 B i o t e c h n o l o g i c a l a p p l i c a t i o n s i n c o n s e r va t i o n a n d i m p rove m e n t o f f e r n s [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec . , CSIR-IHBT, Palampur, H.P., India Abstr act Vol. pa ge 19.
KUMAR R 2014 Environmental impact and management of a red water fern (Azolla filiculoides)
[Abstract] National Conference on Modern Approaches to P t e r i d o p hy t e s : B i o l o g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t ra c t Vo l .
pages 37-38.
KUMAR S 2014 A study of d i s t r i bu t i o n , status and ethnobotanical importance of selected
P t e r i d o p hy t e s i n J h a r k h a n d , I n d i a [ A b s t r a c t ] N a t i o n a l C o n f e re n c e o n M o d e r n A p p r o a c h e s t o
P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r,
H.P. , India Abstr act Vol. pag es 46-47.
K U M A R I A , G U P TA P, K U M A R I A , NA D DA G , L A L B & S I N G H B 2 0 1 4 I n s e c t i c i d a l a c t iv i t i e s o f
f e r n s o f We s t e r n H i m a l a y a s [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o
P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r,
H.P. , India Abstract Vol. page 91.
K U M A R I A , P R A K A S H O & L A L B 2 0 1 4 E p i p h y t i c F e r n s o f K a n g r a Va l l e y, H i m a c h a l P r a d e s h .
P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I P l a n t
Sciences : 221.
K U M A R I A , P R A K A S H O & L A L B 2 0 1 4 F e rn d iver sity of Kalatop Wildlif e S a n c t u a r y o f H i m a ch a l
P r a d e s h [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 26-27.
KUMARI A, SHARMA P & LAL B 2014 Phenological studies of Adiantum species under ex-situ
condition [Abstr act] National Conf er ence on Moder n Appr oaches to Pter idophytes : Biolo g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 41-42.
KUMARI B & SHARMA V 2014 Evaluation of temper ature and desiccation on ex situ conser vation of
fer n spores [Abstract] National Confer ence on Modern A pproaches to Pteridophytes : Biology,
B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t r a c t
Vo l . p ag e s 7 9 - 8 0 .
LAL B, KUMARI A, SHARMA P, PARKASH O, KUMAR A, GOPICHAND & SINGH RD 2014 Lung ru
[ D i p l a z i u m m a x i m u m ( D . D o n ) C . C h r. ] – A l iv e l i h o o d o f t h e n a t iv e P e o p l e o f w e s t e r n
H i m a l a y a [ A b s t r a c t ] N at i o n a l C o n f e r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y,
B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t r a c t
Vo l . p ag e s 5 0 - 5 1 .
MANDAL SUBHAJIT & SUKUL NEE CHUNARI S 2014 Diversity of Pteridophytes at laterite zone of
We s t B e n g a l [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s :
B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstr act Vol. pag es 39-40.
Bibliography of Indian Pteridology (2013-2014) 195

MAZUMDAR J 2013 A new combination in Thelypteridaceae Phytotaxa 1 3 2 (1): 64.


MAZUMDAR J 2013 New combina tions f or some hybrids in the f ern family Thelypteridaceae Annales
Botanici Fennici 5 00(6) : 398-400.
MAZUMDAR J 2014 (033) Proposal to ad d a new Note with an example after A rt. 11.4 Taxon
6 3 (6) December : 1385.
MAZUMDAR J 2014 Nomenclatural note on three ferns of India Phytotaxa 1 5 8 (3): 297-298.
M A Z U M DA R J 2 0 1 4 A n ew c o m b i n a t i o n i n A r a c h n i o d e s ( D r y o p t e r i d a c e a e ) Ph y t o t a x a 1 5 9 ( 1 ) : 0 2 6 -
028.
MAZUMDAR J & MUKHOPADHYAY R 2013 Notes on some f erns Bionature 3 3 (2) : 45-48.
M A Z U M DA R J & M U K H O PA D H YAY R 2013 Nomenclatural notes on some members of
Thelypteridaceae J. Botan. Soc. Bengal 6 7 (1) : 43-47.
M A Z U M DA R J & M U K H O PA D H YAY R 2013 Nomenclatural notes on some members of
Thelypteridaceae II Bionature 3 33(1) : 13-34.
M A Z U M DA R J & M U K H O PA D H YAY R 2 0 1 4 O n t h e o c c u r r e n c e o f L o x o g ra m m e d u c l o u x i i i n I n d i a
Science & Culture 8 00(7-8) : 228.
M E E NA K , YA DAV B L & H U S S A I N S 2 0 1 4 E ff e c t o f h y p e r t h e r m i a o n c e l l m e m b r a n e p e r m e a b i l i t y
o f I n d i a n s p e c i e s o f S e l a g i n e l l a B e a u v. f r o m R a j a s t h a n [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n
M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . ,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 61-62.
M E H LT R E T E R K 2 0 1 4 N ew F r o n t i e r s o f F e r n E c o l o g y. P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e
Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 106.
M O H A N DA S S & S O M VA N S H I R 2 0 1 4 E va l u at i o n o f t o x i c o - p at h o l o g i c a l e ffe c t s o f Pter is cr etica i n
l a b o r a t o r y r a t s [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstract Vol. page 85.
MIR SA, MISHRA AK, RESHI ZA & SHARMA M P 2014 Preliminary Phytochemical Screening of
Some Pteridophyte Species. Proceedings 101 st Session Indian Science Congress Jammu Feb 3-
7, 2014, Part II Plant Sciences : 231.
MIR SA, MISHRA A K , RESHI ZA & SHARMA MP 2014 Preliminar y phytochemical scr eening of 34
p t e ri d o p h y t e s p e c i e s f r o m K a s h m i r va l l ey, I n d i a [ A b s t r a c t ] N a t i o n a l C o n f er e n c e o n M o d e r n
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. pag e 72.
MIR SA, MISHRA AK, RESHI ZA & SHARMA MP 2014 Elemental analysis of Pteridophytes from
Kashmir valley , India: nutritional and phytoremediation perspectives [Abstract] National
C o n fe r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u rc e ,
20-21 st Dec . , CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pa ge 90.
MUKHERJEE M, MUKHERJEE B, BISWAS O & BERA S 2014 Phytolith spectra of some selected fern
a n d f er n a l l i e s o f A r u n a c h a l H i m a l a y a s [ A b s t ra c t ] N a t i o n a l S y m p o s i u m o n P l a n t D iver s i t y :
Structure, Function, Utilization and Conservation December 4-6, Bot. Soc. Bengal, CAS,
Dept. of Botany, University of Calcutta, Kolkata,700019 Abstr act Vol. page 4.
MUKHOPADHYAY R 2014 Research Tr ends in Pter idology. Pr oceedings 101 st Session Indian Science
Congress Jammu Feb 3-7, 2014, Part II Plant Sciences: 127-128.
PA N D E Y R & T H A K U R S 2 0 1 4 M a r s i l e a a n a p h r o d i s i a c f e r n [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n
M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . ,
196 Indian Fern Journal Volume XXXI (2014)

CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 51.


PANDEY R & THAKUR S 2014 Compar a tiv e study of A zolla & Mar silea at Patna. Proceedings 101 st
Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences: 228-229.
PA N D E Y R & T H A K U R S 2 0 1 4 E ff e c t o f C a l c i u m d i hy d r og e n p h o s p h a t e o n A z o l l a [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresour ce, 20-21 st Dec . , CSIR-IHBT, Palampur, H.P., India Abstr act Vol. page 92.
PAT I L S , & M E E NA D O N G A R E 2 0 1 4 T h e G e n u s B o l b i t i s S c h o t t . f r o m We s t e r n G h a t s o f I n d i a .
P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I P l a n t
Sciences : 228.
PATIL S, SAVLARAM GHANE S & MEENA DONGARE 2014The Gen us Ophioglossum from Wester n
Ghats. Proceedings 101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant
Sciences : 227-228.
PATIL S 2014 Fer n and f er n allies of north Western Gha ts (India) [Abstract] Na tional Confer ence on
M o d e r n A p p r o a ch e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . ,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. page 28.
P R A K A S H A , S I N G H A P & K H A R E P B 2 0 1 4 S t u d i e s o n s o m e u s e f u l P t e ri d o p h y t i c p l a n t s o f tr i b a l
tracts in Uttar Pradesh, India [Abstract] National Conference on Modern Approaches to
P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c. , C S I R - I H B T, P a l a m p u r,
H.P. , India Abstract Vol. page 47.
PUNETHA N 2014 Phenological studies on a Polypodiaceous Fern Drynaria propinqua. Proceedings
101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 132-133.
PUNETHA N & PUNETHA R 2014 Habitat specificity in Central Himalayan Species of Selaginella
( Ly c o p h y t a ) . P ro c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n gr e s s J a m mu F eb 3 - 7 , 2 0 1 4 , Pa r t
II Plant Sciences: 238-239.
P U N E T H A N 2 0 1 4 P h e n o l o g y o f f er n l e a f [ A b s t r a c t ] N a t i o n a l C o n f er e n c e o n M o d e r n A p p r o a c h e s t o
P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c. , C S I R - I H B T, P a l a m p u r,
H.P., India Abstract Vol. page 5.
P U N E T H A N , PA N T J N & P U N E T H A R 2 0 1 4 P h e n o l o g y o f a t e m p e r a t e f e r n Dr yo p t e r is w a l l i c h i n a
( S p r e n g. ) H y l a n d e r [ A b s t r a c t ] N a t i o n a l C o n f er e n c e o n M o d e r n A p p r o a ch e s t o P t e r i d o p hy t e s :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstr act Vol. pag es 40-41.
R AT H O D V I T T H A L N , & BA L D E G OV I N D H 2 0 1 4 B i o d ive r s i t y o f P t e r i d o p h y t e s f r o m To r a n m a l
H i l l s , N a n d u r b a r D i s t r i c t , M a h a r a s h t r a , I n d i a . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e
Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 223.
RAWAT VK & BENNIAMIN A 2014 Contribution to the Pteridoph ytic f lora of India : Ecological and
taxonomical studies on the Pteridophytic flora of Maharashtra [Abstract] National Conference
on Moder n A pproaches to Pteridophytes: Biolo g y, Biodiver sity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. pages 32-33.
R AWAT V K & S AT YA NA R AYA NA P 2 0 1 4 Fa m i l y P o l y p o d i a c e a e i n M e h a o Wi l d l i f e S a n c t u a r y,
A r u n a ch a l P r a d e s h [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A ppr o a c h e s t o P t e r i d o p hy t e s :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstr act Vol. page 27.
R AWAT V K , S I N G H A P & S AT YA NA R AY NA P 2 0 1 4 P e r s p e c t iv e s o f b i o d iv e r s i t y : P t e r i d o p h y t e s a
r e s o u r c e f o r f u t u r e Vi n e e t [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o
Bibliography of Indian Pteridology (2013-2014) 197

P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r,
H.P. , India Abstract Vol. pag e 80.
S AGGO MIS & AKHTER A 2014 Explor ation of cytomor pholog ical diversity in medicinally important
fern : Asplenium trichomanes Linn. From Kashmir [Abstract] Conference on Modern Appr oaches
to Pter idophytes: Biology, Biodiver sity and Bior esourc e, 20-21 st Dec . , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. pag e 63.
S AGGO MIS , KAUR M & KAUR K 2014 Chromosomal studies on some economically important ferns
from north-west India [Abstract] National Conference on Modern Approaches to Pteridophytes:
Biology, Biodiversity and Bioresource, 20-21 st Dec., CSIR-IHBT, Palampur, H.P., India Abstract
Vol. pages 65-66.
SAMANT SS 2014 Research Needs for the Conservation and Management of Biodiversity in Trans,
N o r t h We s t e r n a n d We s t e r n H i m a l a y a . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s
Jammu Feb 3-7, 2014, Part II Plant Sciences : 133-136.
S A R E E N B , S O O D A & B H AT TAC H A RYA A I n v i t r o d ev e l o p m e n t a n d s e x o n t o g e n y o f t wo
medicinally important species of Adiantum [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. pag e 66.
SEN K & MUKHOPADHYAY R 2014 New repor t of vessel elements in Aleuritopteris and Cheilanthes .
Taiwania 5 9 ( 3 ) : 231-239.
SETH A 2014 Role of fer ns on bior emediation [Abstr act] National Conference on Modern Appr oaches
to Pter idophytes: Biology, Biodiversity and Bioresour c e, 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. pag e 87.
S E T H A & S E T H M K 2 0 1 4 P t e r i d o p h y t e s a n d t h e i r e c o n o m i c ex p l o i t a t i o n s [ A b s t r a c t ] N a t i o n a l
C o n fe r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u rc e ,
20-21 st Dec . , CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pa ge 17.
SETH MK & NEGI HC 2014 Asplenium species of district Kinnaur of Himachal Pradesh [Abstract]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d ive r s i t y a n d
Bioresource, 20-21 st Dec., CSIR-IHBT, Palampur, H.P., India Abstr act Vol. pag e 28.
SETH MK & SINGH R 2014 Polystichum Species of District Kangra of Himachal Pradesh. Proceedings
101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 237.
SETH MK & SINGH R 2014 High Altitude Fer ns of District Kangra of Himachal Pradesh. Proceedings
101 st Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 238.
SETH MK & SINGH R 2014 Asplenium species of district Kangra of Himachal Pradesh [Abstract]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d ive r s i t y a n d
Bioresource, 20-21 st Dec., CSIR-IHBT, Palampur, H.P., India Abstr act Vol. pag e 31.
SHARMA BD 2014 Present status of Mesozoic Pteridophytes and associated flora [Abstract] National
Confer ence on Moder n Appr oac hes to Pteridoph ytes : Biology, Biodiver sity and Bioresour ce ,
20-21 st Dec. , CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 11-13.
SHARMA BD 2014 Multimorphic spores in Equisetum ramosissimum Desf. [Abstract] National
C o n fe r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u rc e ,
20-21 st Dec. , CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 55-56.
SHARMA BD & BOHRA DR 2014 Ferns of Rajasthan–Apogamy in Adiantum lunulatum Burm.
P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I P l a n t
Sciences : 221.
198 Indian Fern Journal Volume XXXI (2014)

SHARMA BD & BOHRA DR 2014 Studies on plant fossils of the Rhynie Chert (Lower Devonian
Scotland) [Abstr act] National Confer ence on Modern A pproac hes to Pteridophytes : Biolo gy,
B i o d ive r s i t y a n d B i o r e s o u rc e , 2 0 - 2 1 st D e c. , C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 16-17.
S H A R M A B D & P U RO H I T S N 2 0 1 4 A n a c c o u n t o n t h e s y s t e m a t i c s o f I n d i a n s p e c i e s o f I s o e t e s
L. (Lycopod). Proceedings 101 st Session Indian Science Cong ress J ammu Feb 3-7, 2014, Part
II Plant Sciences : 220.
S H A R M A B D & S U T H A R O P 2 0 1 4 M e s o z o i c F o r e s t s o f J h a r k h a n d S t a t e , I n d i a . P r o c e e d i n g s 1 0 1 st
Session Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 216-217.
S H A R M A O P 2 0 1 4 A p e r s p e c t iv e o f t h e r e s e a rc h o n f e r n i n d u c e d u r i n a r y b l a d d e r c a n c e r i n a n i m a l s
in upland ar eas of Himac hal Pradesh [Abstr act] Na tional Confer ence on Moder n Approac hes
to Pter idophytes: Biolo gy, Biodiver sity and Bioresource , 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstr act Vol. pag es 15-16.
S H A R M A P & B H A R DWA J N 2 0 1 4 Ti s s u e C u l t u r e a p p l i c a t i o n f o r B i o d iv e r s i t y C o n s e r va t i o n o f t h e
E n d e m i c S p e c i e s o f M a r s i l e a i n H a d a u t i R e g i o n . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e
Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 235.
S H A R M A P & B H A R DWA J N 2 0 1 4 S t u d i e s o n M a r s i l e a c o r o m a n d e l i n a c o m p l e x p o p u l a t i o n f r o m
Borawas village of Kota, Rajasthan (India) [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. page 77.
SHARMA R 2014 Enzootic bovine hematuria, a fern-induced neoplastic disease of cattle : update o n
work done in India and future road-map [Abstract] National Conference on Modern
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. pages 86-87.
S H A R M A S K & S H A R M A S 2 0 1 4 M e d i c i n a l a n d N u t r i t i o n a l va l u e o f I n d i a n F e r n s i n Ay u r v e d i c
Li t e r a t u r e [ Ab s t r a c t ] Nat i o n a l C o n fe r en ce o n Mo d er n A p p r o ach es t o P t er i d o p hyt es: B i o l o g y,
B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a A b s t r a c t Vo l .
pages 49-50.
S H A R M A S S 2 0 1 4 F e r n s o f H i m a c h a l P r a d e s h . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s
Jammu Feb 3-7, 2014, Part II Plant Sciences : 148-149.
SHARMA V, SINGH G, SWARNKAR MK, PURIA M, KUMARI A, SHARMA RK, AHUJA PS , SINGH
AK 2014 De novo transcriptome of Equisetum ramosissimum under dehydration stress
[Abstract] National Conference on Modern Approaches to Pteridophytes : Biology, Biodiversity
and Bioresour ce, 20-21 st Dec., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pag es 70-71.
S H I J I T H P P & S R E E N I VA S V K 2 0 1 4 P r e l i m i n a r y s t u d i e s o n t h e p t e r i d o p h y t e s o f A r a l a m Wi l d l i f e
S a n c t u a r y, We s t e r n Ghats [Abstract] National Conference on Modern Approaches to
P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r,
H.P. , India Abstr act Vol. pag es 33-34.
S I N G H A P, J O H A R I D & K H A R E P B 2 0 1 4 B i o - g e o g r a p h i c a l D i s t r i bu t i o n a n d S p e c i e s D iv e r s i t y o f
Pteridophytes in Dudhwa National Park, Uttar Pradesh, India [Abstract] National Conference
on Moder n A pproaches to Pteridophytes: Biolo g y, Biodiver sity and Bioresource , 20-21 st Dec.,
CSIR-IHBT, Palampur, H.P., India Abstract Vol. pages 38-39.
SINGH B 2014 Chemical diversity of western Himalayan Pteridophytes [Abstract] National Conference
on Moder n A pproaches to Pteridophytes: Biolo g y, Biodiver sity and Bioresource , 20-21 st Dec.,
Bibliography of Indian Pteridology (2013-2014) 199

CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 18-19.


SINGH P 2014 Legacy of pteridological research in India and contribution of Botanical Survey of
I n d i a [ A b s t r a c t ] , N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y,
B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c C S I R - I H B T, Pa l a m p u r, H . P. , I n d i a A b s t r a c t
Vo l . p a g e s 3 - 5 .
SINGH SHWETA 2014 Traditional uses of Pteridoph ytes for women’s health care Practices in Madh ya
P r a d e s h . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I
Plant Sciences : 230.
SINGH SHWETA & RITA SINGH 2014 Sur vey, Diver sity, Taxonom y and Conservation status of Tr e e
f e r n s o f P a c h m a r h i B i o s p h e r e R e s e r v e , M a d h y a P r a d e s h . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n
Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 225-226.
S I N G H S H W E TA & R I TA S I N G H 2 0 1 4 P hy t o c h e m i s t r y a n d i d e n t i fi c at i o n o f b i o a c t ive c o m p o u n d s o f
Ethnobotanically important species Selaginella bryopteris (L.) Bak. Proceedings 101 st Session
Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 234-235.
SINGH S & SINGH R 2014 Tree f erns of Panchmar i Biospher e Reserve, Madhya Pradesh : Taxonomy
a n d c o n s e r va t i o n [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstract Vol. page 77.
SINGH SK, DOMINIC RAJKUMAR S, SRIVASTAVA SK & GAUTAM RP 2014 Cheilanthes bicolor L.
(Pteridaceae): an addition to the fern flora of Uttar Pradesh [Abstract] National Conference
o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 s t
Dec., CSIR-IHBT, Palampur, H.P., India Abstract Vol. pag es 29-30.
SINGH VJ, JOHARI D, SINGH AP & KHARE PB 2014 Regeneration potentiality and mass
m u l t i p l i c a t i o n o f g a m e t o p h y t e s i n t r e e f e r n C y a t h e a s p i n u l o s a Wa l l . e x H o o k . [ A b s t r a c t ]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec., CSIR-IHBT, Palampur, H.P. , India Abstr act Vol. pag es 63-64.
S I TA & S R I VA S TAVA M 2 0 1 4 A n a t o m y o f l e ave s o f I s o e t e s c o r o m a n d e l i n a L . ( 1 2 l o c a l i t i e s )
[Abstract] National Conference on Modern A pproaches to Pteridophytes: Biology, Biodiversity
and Bior esource, 20-21 st Dec ., CSIR-IHBT, Palampur, H.P. , India Abstract Vol. pag e 58.
SRIVA S TAVA G 2014 Effect of the pesticide Mala thion on gr owth and antio xidant enzyme pr ofile of
d i ff e r e n t s p e c i e s o f A z o l l a [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o
P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, Pa l a m p u r,
H.P. , India Abstract Vol. pag es 87-88.
S R I VA S TAVA R & U N I YA L P L 2 0 1 4 A c o m p a r a t ive d eve l o p m e n t a l s t u d y o f O ny c h i u m C o n t i g u u m
Wa l l . ex H o p e . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 ,
Part II Plant Sciences : 229.
S R I VA S TAVA R , P R I T I & U N I YA L P L 2 0 1 4 I n - v i t r o d ev e l o p m e n t a l s t u d y o f C h e i l a n t h e s t e n u i f o l i a
( B u r m . ) S w. [ A b s t r a c t ] N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p hy t e s :
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstract Vol. page 62.
S R I VA S TAVA S C & S A X E NA R 2 0 1 4 P t e r i d o p h y t i c d iv e r s i t y i n g e o l o g i c p a s t [ A b s t r a c t ] N a t i o n a l
C o n fe r e n c e o n M o d e rn A p p r o a c h e s t o P t e r i d o p hy t e s : B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u rc e ,
20-21 st Dec. , CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ges 20-22.
SUKUL NEE CHUNARI S & MONDAL S 2014 Evaluating the quantitative ethnobotanical importance
200 Indian Fern Journal Volume XXXI (2014)

o f p t e r i d o p hy t e s i n R a r h r eg i o n o f We s t B e n g a l [ A b s t r a c t ] N at i o n a l C o n fe r e n c e o n M o d e r n
A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R -
IHBT, Palampur, H.P., India Abstract Vol. page 48.
SUKUL NEE CHUNARI S & MONDAL S 2014 Diversity of Pteridophytes and its traditional uses in
Rarh Bengal [Abstract] National Symposium on Plant Diversity : Structure, Function,
U t i l i z a t i o n a n d C o n s e r va t i o n D e c 4 - 6 B o t . S o c . B e n g a l , C A S , D e p t . o f B o t a n y, C a l c u t t a
university, Kolkata 700019 Abstr act Vol. page 16.
S U T H A R O P, H A R S H R & S h a r m a B D 2 0 1 4 A n a t o m y o f f o s s i l O p h i o g l o s s a c e o u s r h i z o m e fr o m t h e
Mesoz oic of the Rajmahal hills, India [Abstract] National Conf erence on Moder n Appr oaches
to Pter idophytes: Biolo gy, Biodiver sity and Bioresource , 20-21 st Dec. , CSIR-IHBT, Palampur,
H.P. , India Abstract Vol. page 55.
V E R M A M , WA L L I A , D H A DWA L S , L A L B , K U M A R I A , S I N H A A K & AG N I H OT R I V K 2 0 1 4
Comparative estimation of natural colors and dyes from some selected Pteridophytes growing
in eastern Himala yas [Abstract] National Conference on Moder n Approaches to Pter idophytes:
B i o l o g y, B i o d ive r s i t y a n d B i o r e s o u r c e , 2 0 - 2 1 st D e c . , C S I R - I H B T, P a l a m p u r, H . P. , I n d i a
Abstr act Vol. pag es 72-73.
V E R M A S C 2 0 1 4 E x p l o r i n g F e r n s i n t h e G e n o m i c A g e . P r o f A r c h a n a S h a r m a M e m o r i a l Aw a r d
L e c t u r e . P r o c e e d i n g s 1 0 1 st S e s s i o n I n d i a n S c i e n c e C o n g r e s s J a m m u F e b 3 - 7 , 2 0 1 4 , P a r t I I
Plant Sciences : 99-100.
V E R M A S U N I L K & K H U L L A R S . P. T h e P t e r i d o p h y t e s o f M a n d i D i s t r i c t , Pa r va t i Va l l e y ( K u l u
District) and Mani Mahesh in Chamba District (Himachal Pradesh). Proceedings 101 st Session
Indian Science Congress Jammu Feb 3-7, 2014, Part II Plant Sciences : 227.
VERMA SC 2014 Exploring Ferns in the Genomics age : Opportunities and challenges [Abstract],
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresource , 20-21 st Dec CSIR-IHBT, Palampur, H.P., India Abstract Vol. pa ge 6.
YA DAV BL 2014 Obser vations on some Indian species of Ophio glossum L. fr om Rajasthan [Abstract]
N a t i o n a l C o n f e r e n c e o n M o d e r n A p p r o a c h e s t o P t e r i d o p h y t e s : B i o l o g y, B i o d iv e r s i t y a n d
Bioresour ce, 20-21 st Dec . , CSIR-IHBT, Palampur, H.P., India Abstr act Vol. page 25.
Indian Fern Journal Volume XXXI (2014) 201

THE INDIAN FERN SOCIETY (NO (NOWW AT CHANDIGARH)


Registered under the Societies Registration Act XXI of 1860 as Amended
by Punjab Government Act 1957.

INCOME & EXPENDITURE ACCOUNT


FOR THE YEAR ENDED 31-03-2014
Particulars Amount Particulars Amount
(Rs.) (Rs.)
To Bank Charges 1,861.00 By Annual Membership 12,316.00
To Mailing Charges 21,124.00 By Life Membership Fees 40,300.00
To Contingency Expneses 5,000.00 By Subscription Foreign 12,404.00
To Printing Expenses 98,781.00 By Subscription Foreign 23,300.00
To Audit Fees 1,124.00 By Collection of Printing & 80,012.00
Mailing Charges
To Excess of Income By Sales / Subscription of Journal 26,150.00
Over Expenditure 1,12,805.00 By Bank Interest 46,214.00

2,40,696.00 2,40,696.00

Auditor's Report
As per our separate report
of even date attached.

F or S
S.. K. Bhasin & Associa tes
Associates
Charter
Char ed Accountants
tered
ter

Sunil Kumar Bhasin


Partner
M.No. 085976
Date : 11.09-2014
Place : Chandigarh
202 Indian Fern Journal Volume XXXI (2014)

AUDIT OR'S REPOR


UDITOR'S REPORTT
We have examined the Receipts & Payments Account, Income & Expenditure Account and
Balance Sheet of the Indian Fern Society, Patiala (Now at Chandigarh) for the year ended 31st
March, 2014.
We have obtained all the information & explanations which to the best of our knowledge
and belief were necessary for the purpose of the Audit.
In our opinion and to the best of our knowledge and according to the explanations given
to us thereto, the said accounts give true & fair view and the payments have been made in
accordance with the by-laws of the society.

F or S
S.. K.Bhasin & Associa tes
Associates
Charter
Char ed Accountants
tered
ter

Sunil Kumar Bhasin Date: 11.09.2014


Partner Place: Chandigarh
M.No. 085976
Indian Fern Journal Volume XXXI (2014) 203

THE INDIAN FERN SOCIETY


(Established 1983)
[Registered under the Societies Registration Act XXI of 1860 as amended by Punjab Amendment Act 1957]
#1633, Sector 7-C, CHANDIGARH-1600 19, INDIA
[Formerly at the Department of Botany, Punjabi University, Patiala-147002, India]
ENROLLMENT PROFORMA FOR FRESH MEMBERSHIP (INDIA)
I desire to enroll myself as an ordinary/ life member* of the Indian Fern Society. I shall abide by the constitution
of the Society, and the decisions taken from time to time.

Name (in BLOCK LETTERS) .........................................................................................................................


Place and Date of Birth ....................................................................................................................................
Qualifications .....................................................................................................................................................
Field /s of Specialisation .................................................................................................................................
.................................................................................................................................................................................
Professional details ...........................................................................................................................................
................................................................................................................................................................................
Address for Correspondence ............................................................................................................................
.................................................................................................................................................................................
........................................................................................................................... PIN............................................
Telephone (Office) ................................................................. (Res.) ...............................................................
Email address .....................................................................................................................................................
The Ann
Annualual Subscr iption* ffor
Subscription* or member
memberss in India with ef
efff ect fr om JJan
from an uar
anuar
uaryy 2012 is Rs. 750/- (Rs. 500/- Register
Annual Fee plus Rs. 50/- as one-time Enrollment Fee), Or Life Membership Subscription* of Rs. 7000/- [For
Annual Members of three year standing, Life Membership is Rs. 55000/-]000/-] is enclosed as DD No................
dated................, + payable at CHANDIGARH, made In favour of 'The Treasurer/Secretary, INDIAN FERN
SOCIETY', CHANDIGARH, India'.

*Please strike out whatever is not applicable.


+The annual/life membership subscription can be remitted also directly to the Bank Account of INDIAN
FERN SOCIETY : [STATE BANK OF PATIALA, MADHYA MARG, SECTOR 7, CHANDIGARH- 160019,
INDIA, SAVINGS BANK ACCOUNT NO. 65055644880]. The IFSC Code of the said Bank is stbp0000202,
and MICR Code is 160007002. You may enquire for any other requirement from your local bank for
direct remittance. Intimation of direct remittance to the Bank Account of Indian Fern Society must be sent
to Prof. S. P. KHULLAR, Secretary, The Indian Fern Society, by email : <sp.khullar@gmail.com>.
Note : The Indian Fern Journal, of the Society, is sent by Registered Post at the given address of all members. However,
to Life Members the supply of the Journal by Registered Post is made on one time payment of Rs. 400/- only, which
may be added to Life Membership Fee. The completed Membership Form along with the DD payable at Chandigarh is to
be sent to Prof. S. P. Khullar, The Secretary, Indian Fern Society, House No.1633, Sector 7-C, Chandigarh-160019, India.

Place :

Date : Signature
204 Indian Fern Journal Volume XXXI (2014)

THE INDIAN FERN SOCIETY


(Established 1983)
[Registered under the Societies Registration Act XXI of 1860 as amended by Punjab Amendment Act 1957]
1633, Sector 7-C, CHANDIGARH-160019, INDIA
[Formerly at the Department of Botany, Punjabi University, Patiala-147002, India]
ENROLLMENT PROFORMA FOR FRESH MEMBERSHIP (FOREIGN)
I desire to enroll myself as an ordinary/ life member* of the Indian Fern Society. I shall abide by the constitution
of the Society, and the decisions taken from time to time.

Name (in BLOCK LETTERS) .........................................................................................................................


Place and Date of Birth ....................................................................................................................................
Qualifications .....................................................................................................................................................
Field /s of Specialisation .................................................................................................................................
...............................................................................................................................................................................
Professional details ............................................................................................................................................
...............................................................................................................................................................................
Address for Correspondence ............................................................................................................................
...............................................................................................................................................................................
..................................................................................................................... PIN..................................................
Telephone (Office) ............................................................. (Res.) ...................................................................
Email address .....................................................................................................................................................

The Annual Subscription* (for members from abroad) of US $ 75 plus US $ 5 as one-time Enrollment
Fee, Or Life Membership Subscription* of US $ 600 is enclosed as DD+ No................................
dated......................... payable at CHANDIGARH, made In favour of 'The Secretary Treasurer/, INDIAN FERN
SOCIETY,, Chandig
SOCIETY Chandigarh, arh, India'.

*Please strike out whatever is not applicable.


Note : The completed Membership Form along with the DD payable to ‘Treasurer/Secretary, Indian Fern Society,
Chandigarh’ is to be sent to Prof. S.P. Khullar, The Secretary, Indian fern Society, House No.1633, Sector 7-C,
Chandigarh-160019, India.
+ The annual/life membership subscription can be remitted directly to the Bank Account of Indian Fern Society:
[STATE BANK OF PATIALA, Madhya Marg, Sector 7, CHANDIGARH- 160019, INDIA, Savings Bank Account
No. 65055644880 ]. The IFSC Code of the said Bank is stbp0000202, and MICR Code is 160007002. You may
enquire for any other requirement from your local bank for direct remittance. Intimation of direct remittance to
the Bank Account of Indian Fern Society must be sent to Prof. S.P. KHULLAR, Secretary, The Indian Fern
Society, by email : <sp.khullar@gmail.com>.

Place :

Date : Signature
205 Indian Fern Journal Volume XXXI (2014)

STATEMENT ABOUT O
STA WNERSHIP AND O
OWNERSHIP THER P
OTHER AR
PARTICULARS OF
ARTICULARS
INDIAN FERN JOURNAL
Pub lished in accor
Published dance with ffor
accordance or
ormm IV
IV,, Rule 8 of the Re
Reg istraa tion of
g istr
Newspaper (Central) Rules, 1956

1. Place of Publication : Department of Botany, Punjabi University, Patiala-147002, India


2. Periodicity of its publication : Yearly Volume (May be issued in two numbers)
3. Printer's Name : Professor S. S. Bir, Editor of Publications
Whether Citizen of India : Citizen of India
Address : Department of Botany,
Punjabi University, Patiala-147002, India
#33, Manauli House, Yadvindra Colony,
The Mall, Patiala-147001
4. Publisher's Name : Professor S. S. Bir, Editor of Publications
Whether Citizen of India : Citizen of India
Address : Department of Botany,
Punjabi University, Patiala-147002, India
#33, Manauli House, Yadvindra Colony,
The Mall, Patiala-147001
5. Editor-in-Chief's Name : Professor S. C. Verma
Whether Citizen of India : Citizen of India
Address : [Department of Botany, Panjab University, Chandigarh, India]
#5452/1, Modern Complex, Manimajra,
Chandigarh-160101
6a. Name and Address of Individuals : The Indian Fern Society (Regd.)
who own the newspapers
6b. Partners or Shareholders holding : Not Applicable
more than one per cent of the
total capital.

I, Professor S.S. Bir, hereby declare that the particulars given are true to the best of my knowledge and
belief.

Sd/-
(S.S. Bir)
Signature of Publisher
Dated : January 20, 2015 Editor of Publications

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