A REVISION OF THE GENUS
LEDEBOURIA ROTH (HYACINTHACEAE)
IN SOUTH AFRICA
by
STEPHANUS VENTER
Submitted in fulfillment of the
requirements for the degree of
Master of Science.
Department of Botany, Faculty of Science,
University of Natal,
Pietermaritzburg.
1993 .
Supervisor: Mr. T.J. Edwards
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LEDEBOURlA IN SOUTH AFRICA
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DECLARATION
DECLARATION
The results of the research done for this thesis have not been submitted to any
other University. Except where acknowledged in the text, all data is the result of
my own research carried out in the Department of Botany, Natal University and
Department of Botany, University of the North, Sovenga, under supervision of
Mr. T.J. Edwards.
Stephanus Venter
Signed at:
f,e/-e.fs btNIJ
Date:
LEDEBOURIA IN SOlITH AFRICA
!O/O'8/ /993
III
ABSTRACT
ABSTRACT
Members of the genus Ledebouria Roth (Hyacinthaceae), which occur in South
Africa, are revised. This genus occurs throughout Africa, India and
Madagascar. 33 Species are recognized and placed into nine provisional
infrageneric groups.
A multidisciplinary approach was used in an attempt to provide natural
groupings. The following characters were analysed; morphology,
micromorphology, palynology and caryology. Aspects of ovary structure and
leaf micromorphology proved especially useful in the synthesis of infrageneric
and specific concepts.
Keys, descriptions, illustrations, distributional, ecological, medicinal and
toxicological data are provided. This study is based on plants in their natural
habitat, cultivated specimens and representative herbarium specimens from
herbaria in South Africa and in Europe.
UITTREKSEL
Verteenwoordigers van die genus Ledebouria Roth (Hyacinthaceae), wat in
Suid-Afrika voorkom, is hersien. Hierdie genus kom voor oral op die Afrika
kontinent, in Indie en Madagaskar. 33 Spesies word herken en in nege
voorlopige infrageneriese groepe geplaas.
n Multidisiplinere benadering was gebruik om natuurlike groeperings te verkry.
Die volgende kenmerke is geanaliseer; morfologie, mikromorfologie,
palinologie en kariologie. Aspekte van die vrugbeginsel struktuur en blaar
mikromorfologie was veral behulpsaam met die sintese van infrageneriese en
spesifieke begrippe.
I
Identifikasie sleutels, beskrywings, illustrasies, verspreiding, ekologiese data en
medisinale gebruike word verskaf. Hierdie studie is gebaseer op plante in hulle
natuurlike habitat, gekweekte plante asook van herbarium materiaal in Herbaria
in Suid-Afrika en Europa.
UD£BOURIA IN SOUTH AFRICA
IV
CONTENTS
CONTENTS
Frontispiece - Ledebouria
I.
Declaration
Il.
Abstract
Contents
Ill.
Unpublished name changes in the thesis
VII.
IV.
1.0
INTRODUCTION
1.
2.0
HISTORICAL SYNOPSIS
2.
2.1.
The characters used by the various authors in their
placement of Ledebouria
2.2.
Revisions of Scilla subgenus Ledebouria
2.3.
Dr. EZ. van der Merwe's contribution
2.
4.
4.
3.0
MATERIALS AND METHODS
6.
4.0
GENERIC DELIMITATION
10.
5.0
MORPHOLOGY
14.
5.1.
Habit
14.
5.2.
Roots
14.
5.3.
Bulb
15.
5.4.
Leaves
5.5.
5.6.
5.7.
Infraspecific leaf variation
Leaf anatomy
Inflorescence
5.8.
Floral characters
5.8.1. Perianth
16.
19.
22.
25.
32.
32.
5.8.2. Stamens and pollen
32.
5.8.3. Ovary
35.
5.8.4. Capsule and seed
36.
5.8.5. Caryology
37.
LEDEBOURIA IN SOlJI'H AFRICA
v
CONTENTS
6.0
7.0
8.0
INFRAGENERIC CLASSIFICATION
42.
6.1.
Phenotypic variation
42.
6.2.
Genotypic variation
42.
6.3.
Designation of ranks
43.
6.3.1. Species ranks
43.
6.3.2. Species arrangement and sectional ranks
43.
PHENOLOGY AND LEAF PERIOD
44.
7.1.
Flowering period
44.
7.2.
Pollination biology
46.
SEED DISPERSAL
48.
8.1.
Wind dispersal
48.
8.2.
Water dispersal
48.
9.0
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
49.
10.0
PHENETICS
59.
11.0
PHYTOGEOGRAPHY
66.
11.1. Distribution
66.
11.2. Factors promoting speciation and distribution
70.
TAXONOMY
71.
12.1. Generic description
71.
12.2. Key to the sections
72.
12.3. Key to the species
73.
12.4. Species descriptions
79.
12.0
13.0
NOMINA DUBIA AND NOMINA EXCLUSA
340.
14.0
HYBRIDS
341.
LEDEBOURIA IN SOlITH AFRICA
VI
CONTENTS
15.0
16.0
CULTURAL USES
341.
15.1. Food
341.
15.2. Medicine
341.
15.3. Poisoning
342.
15.4. Vernacular names
344.
CAPTIONS TO ILLUSTRATIONS AND TABLES
345.
ACKNOWLEDGMENTS
364.
REFERENCES
365.
LEDEBOURIA IN SOllTH AFRICA
UNPUBLISHED NAME CHANGES IN THE THESIS
UNPUBLISHED NAME CHANGES IN THE THESIS
(a) New combinations
Ledebouria asperijolia (Van der Merwe) S. Venter
Ledebouria ensijolia (Eclcl.) S. Venter
Ledebouria galpinii (Bale.) S. Venter
Ledebouria lepida (N. E. Br.) S. Venter
Ledebouria minima (Bak.) S. Venter
Ledebouria petiolata (Van der Merwe) S. Venter
Ledebouria rupestris (Van der Merwe) S. Venter
Ledebouria sandersonii (Bak.) S. Venter
(b) New taxa
Ledebouria atro-brunnea S. Venter
Ledebouria coriacea S. Venter
Ledebouria crispa S. Venter
Ledebouria dolomiticola S. Venter
Ledebouria glauca S. Venter
Ledebouria monophylla S. Venter
Ledebouria papillata S. Venter
Ledebouria parvijolia S. Venter
LEDEBOURlA IN sourH AFRICA
VII
1
INTRODUCTION
1.0. INTRODUCTION
The genus Ledebouria Roth belongs to the family Hyacinthaceae (Dahlgren et
al. 1985). The distribution of the genus includes the African Continent, India
and Madagascar. The centre of diversity is in the Transvaal and Natal Provinces
of the Republic of South Africa. Members of this genus occur in desert,
grassland, woodland and even on forest floors (Entabeni Forest on the
Soutpansberg, northern Transvaal).
The genus Ledebouria was described by Roth in 1821 with Ledebouria
hyacinthina L., from India, as the type species. This species was transferred to
the genus Scilla L. and provided the basionym for the largest section of Scilla
namely the section Ledebouria (Baker 1870d). Jessop (1970) reinstated the
genus Ledebouria.
Basically three revisions of Ledebouria have been done. The first was Baker's
monograph of Scilla, section Ledebouria and the genus Drimiopsis published in
January 1870. This was followed in September by a revision of the herbaceous
capsular gamophyllous Liliaceae (Baker 1870b). Baker (1896) also published
the revision of Scilla in Flora Capensis. 74 Years later, Jessop (1970) completed
the last revision for the genus.
The large degree of phenotypic variation in Ledebouria, particularly with
regard to habit and leaf maculation, has resulted in the publication of 102
species in four different genera. This led to numerous combinations and changes
throughout the taxonomic history of the genus. As a result the species concepts
are a source of considerable nomenclatural and taxonomic confusion.
In this revision 33 species are recognized and placed in nine sections. The aim
of this study was to produce a logical classification of the genus Ledebouria.
LEDEBOURIA IN smrrH AFRICA
2
HISTORICAL SYNOPSIS
2.0. HISTORICAL SYNOPSIS
2.1.
The characters used by the various authors in their placement of
Ledebouria.
Various authors described Ledebouria species placing them under several
different genera. A summary of the taxonomic treatments by the various authors
is depicted in Table 1.
Table 1. Summary of the taxonomic treatments of Ledebouria by the various
authors.
AUTHOR
DATE
FAMILIAL
PLACEMENT
GENERIC
PLACEMENT
CHARACTERS USED
Linnaeus!
1782
Liliaceae
Hyacimhus
Jacquin!
1794
Liliaceae
Lachenalia
Andrews
1803
Liliaceae
Lachenalia
Ker-Gawler
1811
Liliaceae
Drimia
Trattinick
1814
Liliaceae
Lachenalia
Von Schrank
1820
Liliaceae
Scilla
Roth
1821
Liliaceae
Ledebouria
Schrader
1827
Liliaceae
Drimia
Hooker!
1866
Liliaceae
Scilla
Baker
1868
Liliaceae
Drimia
Corolla shape; leaf attitude and
shape.
Leaf shape; maculation and attitude; scape, raceme and petal attitude.
Leaf shape and number; petal shape
and colour; ovary shape.
Leaf shape and maculation; raceme
attitude and flower characters.
Leaf maculation and colour; inflorescence attitude; petal and ovary
shape.
Leaf, scape, raceme, perianth,
stamens, ovary and fruit characters.
Bulb, leaves, scape, raceme, flower, ovary, capsule and seed
characters.
Bulb, leaf, scape, raceme and
flower characters.
Bulb, leaf, scape, raceme, flower
and ovary characters.
Bulb, leaf, scape, raceme, flower
and ovary characters.
Baker
1870
Liliaceae
Jessop
1970
Liliaceae
Scilla sub-genus
Ledebouria
Perianth segment shape and filament bases.
Ledebouria
Bulb, leaf, inflorescence, perianth
and ovary characters.
Ledebouria
Roots, blub, leaf, inflorescence,
flower and fruit characters.
Dahlgren et al. 1982
Hyacinthaceae
LEDEBOURIA IN SOlITH AFRICA
3
HISTORICAL SYNOPSIS
Roth (1821) described the genus Ledebouria based on the following: corolla six
- lobed, stamens fused to base of tepals, ovary stipitate, style thin and simple,
locules three with base connected. The stipitate ovary is the diagnostic character
of the genus Ledebouria (Roth 1. c.).
Schrader (1827) ignored Roth s circumscription of Ledebouria, and followed
Ker-Gawler (1803) in his broad concept of Drimia using mostly inflorescence
and floral characters. Hooker (1866) conformed to Linnaeus'and von Schrank's
(1820) circumscription of Scilla (Hooker 1866).
I
Baker (1868) described two new species which he hesitantly placed under
Drimia. His innovative classification was the first to distinguish between Scilla,
Lachenalia and Drimia as sub-genera. Baker transferred Ledebouria
hyacinthina, described by Roth, to the genus Scilla
(Baker 1870c).
The most recent revision of Ledebouria forms part of a study by Jessop (1970)
in the bulbous Liliaceae. He concentrated on the generic concepts of Scilla,
Ledebouria and Drimiopsis and reinstated the genus Ledebouria, based on its
stipitate ovary.
In 1982 Dahlgren et al. subdivided the Liliaceae sensu lato into various families
placing Ledebouria within the Hyacinthaceae.
Linnaeus described the genus Hyacinthus which differs from Ledebouria in the
shoots surrounded by basal, tubular, membranous squamae; leaves linear;
stamens short, epitepalous; perianth tube short; ovary sessile. From the above
characters it is clear that Linnaeusf. placed Hyacinthus revolutus under the
wrong genus. The genus Lachenalia as it is accepted today, was published by
Jacquin (1794). It differs from Ledebouria in the upper flowers of the raceme
being mostly vestigial, the outer perianth lobes being shorter than the inner
lobes and the ovary being sessile. Jacquin (l. c.), Andrews (l. c.) and Trattinick
(l. c.) failed to observe these differences between Ledebouria and Lachenalia.
The genus Drimia, described in 1799, was considered congeneric with
Ledebouria by Ker-Gawler (l.c.), Schrader (l.c.) and Baker (1870a). Drimia
differs from Ledebouria in its: loose imbricate bulb scales; spurred lower
bracts; tubular, caducous perianth, exserted stamens, cohering apically and
flattened seeds.
LEDEBOURIA IN SOlITH AFRICA
4
HISTORICAL SYNOPSIS
Von Schrank (1. c.), Hooker f (1. c.) and Baker did not emphasize the stipitate
ovary of Ledebouria and instead maintained a broad concept of Scilla. Scilla
also differs from Ledebouria in the bulb scales being hard and tightly set,
immaculate leaves; central inflorescence; basally spreading perianth lobes;
sessile ovary and many ovules per locule.
2.2.
Revisions of Scilla sub-genus Ledebouria.
Baker (1870c) described two tribes, the Hyacintheae embracing the genus
Drimia and the Scilleae with the genus Scilla. Baker also described three new
species of the sub-genus Ledebouria and provided a key for all 28 existing Scilla
speCIes.
In 1896 Baker completed his second revision of Scilla. Here he described only
members of the sub-genera Euscilla and Ledebouria covering 48 species of
Ledebouria with 16 new species. Many of the species he recognized were
known from single specimens or sometimes only from illustrations. Baker never
visited South Africa and the living plants he used for the descriptions were all
grown in hothouses in Great Britain.
This revision was the most complete work on the genus Scilla. Jessop's revision
would follow 73 years later and would encompass a multidisciplinary approach
including: morphology, cytology, anatomy, cytochemistry and ontogeny.
Jessop (1970) stresses that only a few species of Ledebouria are defined on
really satisfactory qualitative characters. These species are L. hypoxidioides, L.
viscosa, L. ovatifoiia and perhaps L. luteola.
2.3.
Dr. F. Z. van der Merwe's contribution.
Between 1935 - 1960 Van der Merwe gathered information on the genus Scilla
with the idea of revising it. This information, contained in three files, is housed
at the National Herbarium, Pretoria.
LEDEBOURIA IN SOtJrH AFRICA
5
HISTORICAL SYNOPSIS
Van der Merwe concentrated on the generic concepts of the Scilleae and
recognized three genera within Scilla. Most of the species were retained in
Scilla (Van der Merwe 1943a) and he described the genus Resnova (Van der
Merwe 1946) for a small group of plants with sessile ovaries and ascending
perianth segments, which are never blue or blue-purple. He erected the genus
Schizocarphus (Van der Merwe 1943b) for the plants, very closely related to
Scilla, in which the tops of the bulbs are fibrous. The genus Resnova is regarded
as a synonym of Drimiopsis by Jessop (Jessop 1972a).
On his trips throughout Natal and Transvaal, Van der Merwe made thorough
collections of Ledebouria and Scilla. All these plants were grown for him by the
garden staff of the Division of Botany and Plant Pathology, Department of
Agriculture, Pretoria. Many of these plants were illustrated in "Flowering Plants
of Africa" with descriptions provided by Van der Merwe.
LEDEBOURlA IN SOlITH AFRICA
6
MATERIALS AND METHODS
3.0 MATERIALS AND METHODS
Chromosome counts.
Young flower buds were collected between IlhOO and 14hOO and placed into
Carnoy's solution (6 parts ethanol - 3 parts chloroform - 1 part glacial acetic
acid). Anthers were separated from the flower buds, placed on a clean
microscope slide and macerated with an aluminium rod. A drop of aceto
carmine was placed over the anther material with 0.5 ml ferri-acetate and mixed
together. This was heated over an alcohol lamp, a cover slide placed on the
material and heated again. Pressure was applied to the cover slide after placing
it between blotting paper. The microscope slides were made permanent by
freezing them with carbondioxide. The coverglass was lifted off and the slide
immersed into 100% ethanol for 5 minutes. A small drop of euparal was placed
on the material and a cover glass placed on top. These slides were kept on a
hotplate at 40°C for 24 days. The finished product was studied under a Reichert
Microscope.
Computer programmes.
The programme NTSYS-pc was used to analyse data sets. The resultant output
file from the one programme is used as the input file for the next programme.
The initial data matrix file DATA was prepared with the WS2000+ programme
and incorporated into the NTSYS-pc programme. The following procedures
were followed for cluster analysis.
a) OUTPUT - A:DATA
b) STAND
- input A:DATA
output SDATAC
c) SIMINT
- input SDATAC
output SIMINTC
LEDEBOURlA IN SOlITH AFRICA
7
MATERIALS AND METHODS
d) SAHN
- input SIMINTC
output SAHNC
- input SAHNC
e) COPH
output COPHC
t) MXCOMP - input X = SIMINTC
Y = COPHC
output MXCOMPC
g) TREEG
- input SAHNC
output = Phenogram
The DELTA program was used to give concise representation and manipulation
of taxonomic descriptions. Three basic DELTA format files, SPECS =
specifications file, CHARS = character file and the ITEMS = items file were
generated. The CHARS list was done with the WS2000+ programme and
incorporated into the DELTA programme. The ITEMS file contains the
descriptions of the 33 Ledebouria species followed by the SPECS file with
specifications on the character types, number of states, implicit values and
dependant characters.
DELTA was used to generate identification keys, descriptions in natural
language and summary of the data. INTKEY was used to retrieve information
on descriptions of species, separation characters to separate a particular species
from the other species, to determine the differences between two species and to
display the similarities between two species.
Cultivation.
Plants of the various species of Ledebouria were cultivated in an aridarium at
the University of the North.
LEDEBOURIA IN SOlITH AFRICA
8
MATERIALS AND METHODS
Drawings of the plants.
Drawings of the various species were made from live specimens collected in the
veld, SEM micrographs and using a camera lucida.
Electron microscopy.
Plant material of Ledebouria was fixed in EA.A. (formalin - acetic acid alcohol) at a ratio of 1:18:1 or in 2.5 % glutaraldehyde. After fixation, the
samples were washed for three successive periods of 10 minutes each in 0.2 M
Phosphate buffer (pH 7.2) (Glauert 1980). The samples were then dehydrated
for 10 minutes in 50 % acetone, 10 minutes in 70 % acetone and 30 minutes in
100 % acetone. The samples were then ultrasonically cleaned, and placed in 100
% acetone for 60 minutes. The samples were critical point dried, placed on
stubs with double - sided tape and Gold - Palladium coated for 2 minutes.
Material was viewed with Hitachi 450 and Hitachi S -570 scanning electron
microscopes at an accelerating voltage of 10 KV. Voucher specimens are housed
in the Herbarium of the University of the North (UNIN) and the National
Herbarium (PRE), Pretoria. Measurements of epidermal cells and stomata were
taken from scanning electron micrographs.
Pollination studies.
Populations of various species were visited at various times of the day to
determine the visiting hours of the insects. Insects were collected with a swipe
net, suction tube or directly into plastic vials. The insects were transferred into a
Cyanide Killing Bottle for 20 minutes and then placed in small glass vials.
Rock sampling.
Rock samples (hand specimens) were collected at all the populations visited by
the author and identified with the help of geologists from the office of the
Geological Survey, Pietersburg and the Department of Geology, University of
the North, Sovenga.
LEDEBOURIA IN SOlrrH AFRICA
MATERIALS AND METHODS
9
Seed distribution.
A population of L. revoluta was chosen with individuals spread over a flat area
(0°_ 5°) to individuals growing on a slope (15°- 30°). The soil texture varies
from fine sandy soil to coarse gritty soil. Seeds in the capsules of a specific
individual plant were marked with a red vegetable dye. The distances that the
dyed seed travelled were measured after 10 different rainstorms.
Soil sampling.
A sieve analysis was undertaken on soil samples collected at Ledebouria
localities. Soil samples were screened through a 2 mm mesh and immersed in
100% hydrogen peroxide to oxidize the organic component. The sample was
dried and sieved through two further screens 0.02 and 0.002 mm gauge. Soil
depth was measured with a measuring tool designed and built by the author for
this purpose. It consists of a 7 mm diameter drill bit welded onto a 1300 mm
long shaft of hardened rod iron, 7 mm in diameter. At the top end of the shaft a
300 mm long piece of rod iron was welded to form a cross bar for pressing or
turning the tool into the ground. The long main shaft was marked off in
centimeters. Soil depth was measured at various individual plants in a population
by inserting this tool into the ground next to the plants. At least 30
measurements were taken per population depending on the population size. An
avarage soil depth was calculated from these readings.
LEDEBOURlA IN SOlJTH AFRICA
10
GENERIC DELIMITATION
4.0 GENERIC DELIMITATION
The recent restructuring of the Liliaceae senso lato by Dahlgren and Clifford
(1982), is widely accepted. The classification of Ledebouria accepted by the
author, is as follows:
Superorder
LILIIFLORAE
Order
Asparagales
Family
Hyacinthaceae
Genus
Ledebouria
The following genera constitute the Hyacinthaceae in South Africa (Gibbs
Russell 1985):
Albuca L. (57 spp.), Amphisiphon Schltr. (1 sp.), Androsiphon Schltr. (1 sp.),
Bowiea Harv. ex Hook! (2 spp.), Daubenya Lindl. (1 sp.), Dipcadi Medic. (14
spp.), Drimia Jacq. ex Willd. (14 sPP.), Drimiopsis Lindl. (5 spp.), Eucomis L'
Herit. (13 spp.), Galtonia Decne. (2 spp.), Lachenalia Jacq.j. ex Murray (96
spp.), Ledebouria Roth (33 spp.), Litanthus Harv. (1 sp.), Massonia Thunb. ex
Houtt. (5 spp.), Neobakeria Schltr. (4 spp.), Neopatersonia Schonl. (3 spp.),
Omithogalum L. (62 spp.), Polyxena Kunth (3 spp.), Pseudogaltonia 0. Kuntze
(1 sp.), Rhadamanthus Salisb. (11 spp.), Scilla L. (5 spp.), Schizobasis Bak. (1
sp.), Ienicroa Raf. (5 spp.), Thuranthus C.H. Wright (3 spp.), Urginea Steinh.
(26 spp.), l-Cltheimia Gledisch (2 spp.) and Whiteheadia Harv. (1 sp.).
The generic position of Ledebouria has been in question since its inception. Two
species of Ledebouria were described as Drimiopsis, first D. engleri [ = L.
rautanenii (Schinz) S. Venter ined.] by Krause (1914), and then D. papillosa (
=L. scabrida Jessop ) by Dinter (1921). Jessop (1970) reinstated Roth's genus
Ledebouria basing it mainly on the unique stipitate ovary.
Ledebouria is a well defined genus which is easily distinguished from the other
genera in the Hyacinthaceae both in vegetative and floral characters. The most
significant difference is the shape of the ovary. Drimiopsis and Scilla resemble
UDEBOURLA IN SOurH AFRICA
11
GENERIC DELIMITATION
Ledebouria closely. The vegetative differences between these three genera are
shown in Thble 2.
Table 2. Differences between the genera Ledebouria, Drimiopsis and Scilla.
GENUS
BULBSCALES
LEAVES
Ledebouria
Membranous to fleshy
With or without spots
Drimiopsis
Fleshy and loose
With or without spots
Scilla
Hard, dry and tightly packed
Without spots
The leaves of Ledebouria and Drimiopsis are very similar in shape, texture,
marking and colouration (Figure 2). The leaves of Scilla are firmer, resulting
from the lignified sheath around the vascular bundles (Jessop 1972a), without
any spots and often covered with hairs. Some glabrous forms of S. natalensis
occur in the Olivier's Hoek Pass area (pers. obs.).
In Ledebouria and Drimiopsis the inflorescence is axillary and fleshy but in
Scilla it is terminal and rigid. The inflorescences in Ledebouria and Drimiopsis
are mostly flaccid, whereas they are always erect in Scilla.
Although some species of Ledebouria have cucullate perianth lobes, these are
never as pronounced as in Drimiopsis. In Scilla the apices of the perianth lobes
are always acute to acuminate. The filaments are basally fused in Scilla but are
separate in Drimiopsis and Ledebouria.
LEDEBOURlA IN SOlJTH AFRICA
12
GENERIC DELIMITATION
Figure 1 - 6. 1. The bulbs of A, Ledebouria revoluta (L.t) Jessop (lenter
13,209); B, Drimiopsis sp. novo [lenter 13,459 (UNIN)] and C, Scilla
dracomontana Hilliard [Edwards s.n. (UNIN)] to show the differences in the
bulb scales. Bar = 50 mm.
Figure 2. Leaves of the genera A, Scilla natalensis Planch. [lenter
S. n.
(UNIN)]; B, Ledebouria revoluta (lenter 13,209) and Drimiopsis burkei Bak.
[lenter s.n. (UNIN)] to show the differences in the leaves. Bar
= 100 mm.
Figure 3. Three different types of root in the genus Ledebouria. A, contractile
roots of L. revoluta (l-enter 13,207); B, fusiform roots of L. apenijlora (Bak.)
Jessop (Venter 12,686) and C, fleshy roots of L. cooperi (Hook}:) Jessop (Glen
2295). Bar = 50 mm.
Figure 4. Plants of L. crispa S. Venter showing the gregarious habit ("\.enter
11 ,202). Bar = 10 mm.
Figure 5. Basal stems of L. glauca S. Venter (l-enter 13,368). Bar = 50 mm.
Figure 6. SEM micrograph of the spiral threads in the leaves of L. crispa
(l-enter 13,209a). Bar = 5 JLm.
LEDEBOURIA IN SOlITH AFRICA
13
GENERIC DELIMITATION
LEDEBOURlA IN SmITH AFRICA
14
MORPHOLOGY
5.0. MORPHOLOGY
5.1. Habit.
Ledebouria species are geophytic with annual leaves. Plants vary in size from 3
mm tall, with the leaves appressed to the ground (L. galpinii), to 1 meter tall (L.
zebrina) with erect, apically flaccid leaves. A few species are soboliferous
which results in the formation of colonies consisting of few clones connected by
subterranean stolons up to 200 mm long. The largest colony recorded for a
soboliferous individual was 5 meters in diameter in L. cooperi
retn~(
13,342 ).
Species growing on cliffs and steep moist slopes, have pendulous leaves. In the
cliffs at Serala Peak in the north-eastern Transvaal, L. floribunda grows with
leaves hanging down the cliff face. Plants from this colony cultivated in an
aridarium at the University of the North retained this habit.
5.2. Roots.
Roots vary from white (fleshy roots), pink (fusiform roots) to brown and dark
brown (contractile and 'wire' roots).
Contractile roots are hard, at maturity, with transverse wrinkles caused by
contraction of the root which is accomodated by corrugation of the cortex.
These roots compensate for the pseudomonopodial growth of the bulb by
drawing it firmly downwards, and are formed annually (Figure 3A).
Branching fleshy roots are common in Ledebouria and are up to 10 mm in
diameter. They occur in species growing in moist habitats or in deep sandy soils
i.e. Kalahari sand. L. glauca is common in Kalahari sand near Van Zylsrust
and produces basal stems and fleshy roots (Figure 3C).
Fusiform roots are usually soft and are restricted to L. apertijlora and L.
ensifolia. These species have poorly developed bulbs. The roots assume the
storage function of the bulbs. Roots are spread around the plant near the surface
to enhance the uptake of moisture. This is possibly an adaptation to the habitat
LEDEBOURlA IN SOUTH AFRICA
15
MORPHOLOGY
of these taxa as both occur in sandy soils in dry woodland and shrubland (Figure
3B).
'Wire' roots occur in taxa with large bulbs, growing in compact or stoney soil.
These roots are dark brown and difficult to break (L. marginata). They may be
up to 450 mm long and are rarely branched.
5.3. Bulb.
According to Jessop (1970) the development of the bulbs in Ledebouria is
monopodia!. Miiller-Doblies (1971) showed the development of the bulb in
Galanthus to be sympodial. Ontogenetically the inflorescence has a terminal
position with the structure of the bulb a sympodium. This study shows that there
is no evidence that monopodial bulbs exist.
The bulb is sometimes reduced to a thickening or swelling in the leaves. This is
common in soboliferous species of marshy habitats (L. cooperi). In L. ensifolia
the bulb is small and clothed in many layers of dry, hard, dark brown to
purplish brown bulb scales.
Plants of Ledebouria are usually solitary, however L. revoluta and L. asperi/olia
may produce up to seven daughter bulbs. L. crispa is strongly gregarious with
up to 30 plants in a single clump (Figure 4). L. socialis is also gregarial with
bulbs forming on the basal stem.
Bulb elevation is variable. Usually bulbs are hypogeal, a few species have semi
- epigeal bulbs where 25 - 50 % of the bulb is exposed (L. crispa and L.
concolor). Only two species (L. dolomiticola and L. socialis) have epigeal bulbs
with 80 - 100% exposed. These species occur on rock sheets or in deeper soil
but always retain an epigeal habit.
Basal stems occur in a few taxa. In L. dolomiticola and L. papillata they are up
to 10 mm long. In L. viscosa basal stems are up to 130 mm long and 20 mm in
diameter and in exceptional cases branching occurs (Figure 5).
LEDEBOURIA IN SOlITH AFRICA
16
MORPHOLOGY
Bulb width at flowering time for adult plants varies from 5 mm (L. galpinii) to
220 mm (L. revoluta) with the bulb from 10 mm to 230 mm long. Bulb shape is
variable intraspecifically and is therefore of limited taxonomic use.
The colour of the interior of the bulb is usually white but can be purple ( L.
ovatifoiia), or with the upper part of the bulb purplish and the upper part of the
bulb scales markedly spotted. These markings are diagnostic in L. inquinata,
even when the bulb scales are dry. The dead bulb scales may be membranous or
hard and rigid as in L. atro-brunnea. Colour of the bulb scales varies from a
light honey-brown to blackish-brown, various shades of purple also occur. The
bulb scales are normally tightly clustered but can, in some species, be loosely
arranged (L. ovatifoiia and L. galpinii).
Torn bulb scales produce spiral threads, derived from the xylem, in some
species (Figure 6). These threads are usually sparse, but in L. luteola artd L.
ovatifoiia they are prolific and may be used as a diagnostic character (See
chapter 5.6.).
The occurrence of a subterranean or epigeal neck on bulbs is not common.
When it occurs it.is smooth or, more usually, covered by dry leafbases and
varies from 10 - 70 mm in length and from 5 - 20 mm in diameter.
Some species are soboliferous and form bulblets at the base of the bulb scales
but more commonly bulblets are formed at the base of the bulb or on the basal
stem. Some species form bulblets on the upper surface of broken basal stems
resulting in small colonies above ground.
5.4. Leaves.
The leaves of Ledebouria are rosulate and there is only one hysteranthous
species (L. undulata).
In most of the species the leaves are fully emerged at flowering time. Species in
the subsection Stellatae have partly emerged leaves at flowering time.
LEDEBOURIA IN SOtJrH AFRICA
17
MORPHOLOGY
Leaves are predominantly spreading or in some instances erect (L. atrobrunnea) or appressed to the ground (L. galpinii and L. monophylla). In L.
ovatifolia the leaves are usually appressed to the ground except in certain
populations on the South Coast between Durban and Scottburgh. The leaves of
these plants tend to spread. L. marginata, L. leptophylla and L. atro-brunnea
have spirally twisted leaves. Drier and rockier habitats, have more twisted
leaves.
L. monophylla is the only Ledebouria species with a single leaf. Very rarely,
under optimum conditions, the plants may produce another leaf which always
remains poorly developed. The highest number of leaves counted on a single
plant of Ledebouria is 25 (an old specimen of L. marginata).
In Ledebouria cataphylls are rare. Up to three cataphylls occur and these are
usually exserted above ground level. These cataphylls contain chlorophyll only
in the above ground parts.
Leafshape varies from linear to ovate (Table 3) and with various combinations
between these limits (Figure 7). In some species the leatbase is petiolate (Thble
3). The leatbase is usually canaliculate but can be rolled, terete or flat. Leaf
apices may be obtuse, emarginate, acute or acuminate.
Leaves are mostly fleshy and soft but can be leathery, or leathery with
prominently raised veins. The texture of the lamina margin is smooth, finely
serrate, ciliate or papillate. In addition the margin may be smooth, thickened or
undulate. In L. inquinata the proximal part of the lamina margin becomes
undulate only in adult plants. Margins are mostly concolorous but may be
discolorous green, -red, -white or -purple.
Upper and lower lamina surfaces are usually smooth but may be pitted (L.
galpinii), viscid (L. viscosa) , hairy (L. hypoxidioides) or. with rows of asperities
(L. asperijolia). The occurrence of hairs and cilia on the leaf surface of
Ledebouria is rare but consistent. In L. lepida and L. parvifolia hairs are
unicellular but in L. hYPoxidioides they occur in stellate groups.
LED£BOURlA IN SOlJfH AFRICA
18
MORPHOLOGY
Table 3. Leaf characters for the genus Ledebouria in South Africa.
BASE
TEXTURE
APEX
lanceolate
canaliculate
fleshy
acute
L. asperifolia
lanceolate
canaliculate
fleshy
acute
L. atro-brunnea
lanceolate
canaliculate
fleshy
acute
L. concolor
oblong-lanceolate
flat
fleshy
obtuse
L. cooperi
lanceolate
canaliculate
fleshy
acute
L. coriacea
lanceolate
canaliculate
leathery
acute
linear-lanceolate
canaliculate
fleshy
acute
L. dolomiticola
lanceolate
canaliculate
fleshy
acute
L. ensifolia
ensiform
canaliculate
fleshy
acute/obtuse
L. floribunda
lanceolate
canaliculate
fleshy
acute
L. galpinii
ovate/spathulate
canaliculate
fleshy
acuminate
L. glauca
lanceolate
canaliculate
leathery
acute
ovate-lanceolate
canaliculate
fleshy
acute
L. inquinata
lanceolate
canaliculate
fleshy
acute
L. lepida
lanceolate
canaliculate
fleshy
acute
linear
canaliculate
leathery
acute
L. luteola
linear-lanceolate
canaliculate
fleshy
acute
L. macowanii
linear-lanceolate
canaliculate
fleshy
acute/obtuse
L. marginata
lanceolate
flat
leathery
acuminate
L. minima
linear
canaliculate
fleshy
acute
L. monophylla
ovate
canaliculate
succulent
obtuse/acute
L. ovalifolia
oblanceolate
canaliculate
fleshy
obtuse
L. ovatifolia
ovate/deltate
canaliculate
fleshy
acute
L. papillata
linear/lanceolate
petiolate
fleshy
L. parvifolia
lanceolate/oblong
canaliculate
fleshy
acute
L. petiolata
lanceolate
petiolate
fleshy
acuminate
L. revoluta
lanceolate
canaliculate
fleshy
acute
L. rupestris
lanceolate
petiolate
fleshy
acute
lanceolate/ovate
canaliculate
fleshy
acute
L. socialis
oblanceolate
canaliculate
fleshy
acute
L. undulata
linear-lanceolate
canaliculate
fleshy
acute
flat
fleshy
obtuse
flat
fleshy
acute
SPECIES
SHAPE
L. apertijlora
L. crispa
L. hypoxidioides
L. leptophylla
L. sandersonii
L. viscosa
spathulate
L. zebrina
oblong/lanceolate
,
UDEBOURJA IN SOUTH AFRICA
acute!acuminate
19
MORPHOLOGY
During mist spells, the pitted areas on the leaves of L. galpinii fill with water.
The thick resinous layer on the leaves of L. viscosa appears to be unpalatable
for animals as no plants were found showing signs of herbivory. The hairy
leaves of L. hypoxidioides closely resemble the strong fibrous leaves of Hypoxis
villosa Lf. (Hypoxidaceae). It is easily mistaken for the unpalatable Hypoxis
plant.
The colour of the upper and lower lamina surfaces are variably green, glaucous
or pruinose. When maculated, the surface markings occur as longitudinal
stripes, or combinations of stripes and spots, spots and blotches or tessellation at
the lamina base. These markings are usually dark green or various shades of
purple. The markings on the leaves of Ledebouria species are highly variable
except in L. glauca, L. lepida and L. papillata where the purple cross bands on
the petioles are always present. This cross banding also occurs in other families
and genera, in some Urginea spp., Sypharissa spp. (Obermeyer 1980),
Rhadamanthus fasciatus B. Nord. (Nordenstam 1970), Albuca and four
Ornithogalum spp. (Miiller-Doblies 1981) and Habenaria ciliosa Lindl. (Bolus
1913).
The leaf shape and colour of L. leptophylla are cryptic. In early spring when the
grass is not green, the dull linear leaves and greyish-pink flowers of L. '
leptophylla are difficult to distinguish.
Prominent costae occur in L. marginata. This tends to be a feature of plants
growing in drier habitats.
5.5. Infraspecific leaf variation within the species.
Within all of the species there is a wide range of variation in qualitative as well
as quantitative features.
L. monophylla and L. papillata are the only species where the vegetative and
flowering periods are synchronous, four months in L. monophylla and three
months in L. papillata (Table 4). Plants with a long vegetative period (longer
than six months) are primarily grassveld species. Variation occurs in some
species where the leaves die whilst the plant is in fruit (L. ovalifolia).
LEDEBOURIA IN SOUTH AFRICA
20
MORPHOLOGY
Figure 7. Different leafshapes in Ledebouria. A, L. cooperi (l-enter 13,342); B,
L. revoluta
(~nter
13,207); C, L. revoluta
(~nter
13,203); D, L. luteola
(l-enter 13,217); E, L. revoluta (l-enter 13,362); F, L. revoluta (l-enter 13,363);
G, L. marginata (l-enter 13,358); H, L. asperi/olia (l-enter 13,382); I, L. revoluta (Scott-Shaw s.n.); J, L. cooperi
(~nter
(~nter
13,334); K, L. floribunda
13,315); L, L. zebrina (Cunningham s.n.); M, L. revoluta ( l-enter 13,322); N,
L. cooperi (~nter
(~nter
13,388); 0, L. asperifolia
13,383); Q, L. revoluta
(~nter
(~nter
13,382); P, L. cooperi
13,257); R, L. inquinata
13,335) and S, L. ovatifolia (l-enter 13,337). Bar = 100 mm.
LEDEBOURlA IN SOlITH AFRICA
(~nter
MORPHOLOGY
LEDEBOURlA IN SOUTH AFRICA
22
MORPHOLOGY
There are only a few species of Ledebouria where the shape of the lamina is
stable (L. atro-brunnea, L. crispa, L. galpinii, L. monophylla, L. petiolata and
L. viscosa). The remaining species exhibit a wide range in leaf shape and size,
L. cooperi and L. revoluta are exceptional in this regard.
Lamina colour and markings vary intraspecifically even within populations. To
show the extent of variation in mottling on the lamina, L. cooperi is discussed.
The lamina is green on both sides or with spots and blotches of purple on the
abaxial surface. In some populations the markings on both surfaces comprise of
deep purple, longitudinal stripes and form a stark contrast to the light green
background. This form has been in cultivation in America for the last 15 years,
(E. van Jaarsveld, pers. comm.). Similar longitudinal purple stripes occur in
Natal populations of L. zebrina (Bak.) S. Venter.
5.6. Leaf anatomy.
Jessop (1970) provides descriptions of transverse sections of leaves in
Ledebouria. He mentions that the vascular bundles are arranged in a single row
with the xylem uppermost and without lignification of the bundle sheath.
Lignification is restricted to the tracheary elements. No prominent palisade layer
has been observed, but in L. floribunda the two adaxial rows of mesophyll cells
are almost twice as tall as they are broad.
The presence of crystals especially raphides in the cells is common.
In most species the leaves, when torn, produce spiral threads (Figure 6). These
originate from the xylem thickening. They do not stain with thiobarbituric acid
nor with phloroglucine + HCI, therefore they are not lignified (Badenhuizen
1954). The average diameter of these spirals is 2.7 ]Lm. According to
Badenhuizen (1954) there are two types of spiral threads, the R-type (removable
type) and the NR-type (non-removable type).
R-type spirals occur in the metaxylem and leave distinct impressions on the wall
when removed. The substance in which the impressions are made is pectin
(Badenhuizen 1954). The NR-spirals are protoxylem and can be staine.d by
saffranin, phloroglucine + HCI and thiobarbituric acid, thus showing them to be
lignified.
LED£BOURlA IN SOlITH AfRICA
Table 5. Microsculpturing of the leaf surfaces.
~
Species
Character of adaxial epidennal cells
L. apertiflora
L. asperifolia
L. arro-brullllea
Smooth; long and narrow; apices truncate.
With wax platelets; cells four-sided; apices truncate.
Thick covering of wax platelets; cells long and narrow;
keel roundish; apices truncate.
Covered with waxy layer; cells long and narrow;
four-sided; apices truncate.
Smooth; long and narrow; extended out to stomata;
apices truncate.
Covered with thick waxy layer; cells short and wide;
eight-sided; lengthned out to stomata; apices truncate.
Covered with thin waxy layer; cells very long and narroW;
four-sided; apices forked.
Covered with thick waxy layer; cells very long and narrow
keeled; apices truncate.
Covered with thin waxy layer; cells very long and narrow;
apices truncate.
Smooth; cells very long and narrow; four-sided; apices
truncate.
Smooth; cells rectangular; four-sided; apices truncate.
Covered with thick waxy layer; cells rectangular;
four-sided; apices round.
Surface covered in long hairs; cell surface smooth;
boundaries not prominent.
Covered with thin waxy layer; cells shortly rectangular;
four-sided; apices truncate.
Smooth; scattered rows of papillae; cells short and
broad; four- to eight-sided; apices truncate.
Stomata
Character of abaxial epidennal cells.
In deep crypts.
In shallow crypts.
In deep crypts.
Smooth; nearly as broad as long; apices truncate.
With wax platelets; cells four-sided; apices truncate.
Thick covering of wax platelets; prominent rib on
the keel; apices forked.
Thick layer of wax; cells long and narrow; four-sided;
apices truncate.
Smooth; long and narrow; four-sided; apices truncate.
~
==
o
t""
L. cOllcolor
L. cooperi
~
~
!:Ic
~
51
~
L. coriacea
L. crispa
~
L. dolomiticola
L. e/lsifolia
L. floribunda
L. galpi/lii
L. glauca
L. hypoxidioides
L. i/lquillata
L. lepida
In shallow crypts.
In shallow crypts.
Waxy layer thin; cells longer than narrow; apices
truncate.
Waxy layer thin; cells shortly rectangular; apices
In deep crypts.
truncate.
Waxy layer thick; cells long and narrow; keels round;
In shallow crypts.
apices truncate.
Waxy layer thin; cells long and narrow; four-sided;
In shallow crypts.
apices truncate.
Smooth; cells long and narrow; four-sided; apices
In shallow crypts.
truncate.
Smooth; cells rectangular; four-sided; apices truncate.
In shallow crypts.
Waxy layer thick; cells longer than broad; eight-sided;
In shallow crypts.
apices square.
Protrude above cells. Surface covered in long hairs; cell surface smooth;
boundaries not prominent.
Smooth; cells very long and narrow; four-sided;
In shallow crypts.
apices truncate.
Smooth; cells long and narrow; four-sided; apices
In shallow crypts.
truncate.
g
0<
In deep crypts.
~
Table 5. Continued
L. leplophylla
L. IUleola
L. macowanii
L. marginala
L. minima
L. monophylla
L. ovalifolia
~
~
l:lc
L. ovalifolia
L. papillala
~
!Z
~
~
!S
L. parvifolia
L. peliolala
L. revolula
L. rupeslris
L. sandersonii
L. socialis
L. ulldulala
L. viscosa
L. zebrina
.
Smooth; cells long and narrow; lengthened out to stomata;
apices deeply forked.
Cells smooth; covered with small warts; cells long and
narrow; apices truncate.
Smooth; cells long and narrow, lengthened out to stomata;
apices truncate.
Smooth; cells very long and narrow, apices tapered or
forked.
Smooth; cells short and broad; six- to eigth-sided;
strongly lengthened to stomata; apices truncate.
Smooth; cells narrowly rectangular; apices tapered.
Smooth; cells mostly narrowly rectangular; apices
truncate.
Covered with thin layer of wax; cells branched, long
and narrow; apices rounded.
Smooth; cells rectangular, lengthened out to stomata;
apices truncate; scattered rows of papillae.
Smooth; dense rows of papillae; cells short and broad,
six- to eight-sided; apices truncate.
Smooth; cells rectangular, lengthened out to stomata;
apices tapered.
Covered with thin layer of wax; cells long and narrow;
apices tapered.
Covered with thin layer of wax; cells narrowly rectangular, lengthened out to stomata; apices truncate.
Smooth; cells narrowly rectangular; apices truncate.
Smooth; cells long and narrow; apices truncate.
In deep crypts.
Smooth; cells long and narrow; apices forked.
In deep crypts.
With scattered wax platelets; cells very long and
narrow; apices truncate.
Smooth; cells long and narrow, lengthened out to
stomata; apices truncate.
Smooth; cells long and narrow; lengthened out to
stomata; apices forked.
Smooth; cells very long and narrow; apices truncate.
Very shallow crypts.
In shallow crypts.
In deep crypts.
In deep crypts.
Smooth; cells short and broad, mostly six-sided;
apices truncate to tapered.
Smooth; scattered rows of papillae; cells short and
broad, six-sided; apices truncate.
With thin layer of wax; cells branched, long and
narrow; apices rounded.
Smooth; cells rectangular, lengthened out to stomata;
apice truncate.
Smooth; cells long rectangular; apices truncate.
In shallow crypts.
Smooth; cells long rectangular; apices tapered.
In shallow crypts.
With thin layer of wax; cells long and narrow; apices
tapered.
Smooth; cells very long and narrow; apices truncate.
In shallow crypts.
In shallow crypts.
In deep crypts.
In shallow crypts.
In shallow crypts.
Smooth; cells narrowly rectangular; apices truncate.
In shallow crypts.
Protrude above cells. Smooth; cells short and broad, six-sided; apices
truncate.
Covered with thick layer of wax platelets; cells
With thick layer of wax platelets; cells narrowly
In deep crypts.
rectangular; apices truncate.
rectangular; apices truncate.
With thick layer of resin; cells rectangular; apices
Covered with thick layer of resin; cells rectangular;
In shallow crypts.
apices truncate.
truncate.
Waxy layer thick; cells narrowly rectangular; apices
Covered with thick waxy layer; cells narrowly rectangular; In shallow crypts.
apices truncate.
truncate.
s
~
ot'"
g
0<
to>
~
25
MORPHOLOGY
The microsculpturing of the leaf surfaces is diagnostic for certain species (Table
5). Microsculpturing of leaves is an aid to classification in various families.
Glen & Hardy (pers. comm.) studied the genus Aloe L. making use of epidermal
leaf characters in their final synthesis. Glen & Hardy (1990) used SEM studies
of the adaxial leaf surfaces of Aloe parvibracteata Schonl. and Aloe dumetorum
Mathew & Brandham to show that A. dumetorum does not belong to the Aloe
maculata All. group. Van Jaarsveld (1989) found that the sculpturing of the
outer leaf epidermis is most useful and of diagnostic value in distinguishing taxa
in the genus Gasteria Duval .
In Ledebouria the adaxial and abaxial surfaces of the lamina are usually smooth
but can be lacunose (L. galpinii), papillate (L. papillata) , asperate (L.
asperifolia), ciliate (L. lepida), viscose (L. viscosa) or covered with hairs (L.
hypoxidioides). The main textural features of the leaves are the rows of papillae
and cilia. Some of the species (L. coriacea, L. dolomiticola and L. marginata)
have a thick waxy cuticle. In L. marginata this glaucous waxy layer can be
rubbed off. The cuticle in L. coriacea is so resilient that after repeated washes
in ethanol and acetone it is still visible under the Scanning Electron Microscope.
Epidermal cell measurements for the genus range as follows: adaxial surface,
100 - 200 x 6 - 10 JLm; abaxial surface 30 - 70 x 10 - 20 JLm.
5.7. Inflorescence.
The inflorescence of Ledebouria is a simple unbranched axillary raceme. A
single raceme is produced annually in 10 species but in the other members 2 14 may be produced. In most species inflorescences are declinate but some are
erect. In L. revoluta the inflorescence is initially erect but gradually becomes
declinate. The length of the inflorescence varies from 30 - 300 mm. The scape
varies from 1 - 6 mm in diameter. The base of the scape is normally terete, but
may be compressed, winged or angular. It is green, purplish-brown or purple
and may frequently be spotted or striped basally.
The rachis is either consistently smooth or ridged. Flowers are spirally arranged
and racemes are either dense or lax. The overall shape is dependent on pedicel
length, posture and insertion and varies from cylindric to globose.
LEDEBOURlA IN SOtJrH AFRICA
26
MORPHOLOGY
Figure 8. Flower bracts. A, L. concolor, vestigial bract ('~nter
s.n.); B, L.
floribunda (Venter 13,315); C, L. apertijlora (Mauve et al. 179); D, L. atrobrunnea (Venter 13,483); E, L. ovat.ifolia (Venter 13,376) and F, L. zebrina
(Venter 13,395). Bar = 500 J.Lffi. Bracteoles indicated with an arrow.
LEDEBOURJA IN SOlJfH AFRICA
27
MORPHOLOGY
LEDEBOURJA IN SOlTrH AFRICA
28
MORPHOLOGY
Figure 9. L. revoluta flower. A, lateral view. B, distal view. Both from Venter
13,007 (X 8).
Figure 10. Tepal apices of A, Drimiopsis burkei Bak. (Venter 13,341) and B,
Ledebouria apertiflora (Bak.) Jessop (Mauve et al. 179) (X 16).
Figure 11. Various parts of the ovary (X16).
LEDEBOURIA IN SOtrTH AFRICA
29
MORPHOLOGY
B
-A
10
. .- - - - - - - - - Stigma
1---------- Style
:L.!- J<-~ - - Shoulder
'--'r---\--------Vertical groove
r------Expanded lobe
= c;z:....--------
....,.;;::""'::::::::::>'"............
11
~- - - - Stipe
UDEBOURIA IN SOlTTH AFRICA
Basal lobe
30
MORPHOLOGY
Figure 12. SEM micrographs of A, tepal apex in L. zebrina (Bak.) S. Venter
(Venter 13,395). Bar = 50 JLm. B, anther of L. ensifolia (Eckl.) S. Venter
(Venter 13,521). Bar = 0.38 mm. C, pollen grain of L. revoluta (L..f.) Jessop
(Venter 13,207). Bar = 12.6 JLm. D, reticulate 'exine of L. revoluta (Venter
13,207). Bar = 5 JLm. E, stigma of L. revoluta retn~(
13,257). Bar = 86 JLm.
F, nectaries on the base of the ovary lobes in L. sandersonii (Bak.) S. Venter
retn~(
13,465). Bar = 200 JLm. G, seed surface of L. glauca S. Venter retn~(
13,386). Bar = 1.2 mm. H, detail texture of the testa of L. luteola Jessop. Bar
= 43 JLm.
LEDEBOURIA IN SOlJfH AFRICA
MORPHOLOGY
31
LEDEBOURLA IN SOUTH AFRICA
32
MORPHOLOGY
Pedicels are patent, drooping or curved but are always initially erect, becoming
horizontal when the flower opens and cernuous after fertilization. They vary in
length from 2 - 20 mm and vary from white to white speckled with pink or
purple. Pedicels become green after fertilization.
In L. concolor floral bracts are absent or, at most, vestigial. In other species the
bracts are sometimes accompanied by lateral bracteoles. Flower bracts can
either be membranous or fleshy and vary in length from 0.5 - 6.0 mm. Shape is
of taxonomic importance (Figure 8). Colour varies considerably through pink,
purple, grey, white and green.
5.8. Floral characters.
5.8.1. Perianth.
Perianth shape is variable from almost tubular to stellate or campanulate with
recurved or reflexed lobes. Perianth length varies considerably in some species
(L. cooperi, L. ovatifolia and L. revoluta). Lobes are oblong, linear, lanceolate
or ovate.
Tepal apices are acute, obtuse, or with recurved points. In some species apices
are cucullate however these are very different from the connivent tepals of
Drimiopsis (Figure 10). Papillae on the apex of the perianth lobe help to keep
the lobe apices together while the flower is still in bud (Figure l2A).
Colour is, in most cases, not of taxonomic importance. Tepals are green, green
on white, green on pink, brown on pink or pink to purple.
5.8.2. Stamens and pollen.
Stamens are patent, erect or connivent. Filament apices are usually acute but
may be truncate. The filament bases are cylindrical or occasionally slightly
flattened. Filament colour is usually more intense than that of the perianth lobes,
and varies through maroon, pink, white and green. The distal part of the
filament is more intensely coloured. In some species the filaments are free from
the tepals and exhibit swollen bases.
LEDEBOIJRJA IN SOlITH AFRICA
33
MORPHOLOGY
Figure 13. SEM micrographs of the different ovary shapes. A, L. undulata
B, L. papillata S.
(Jacq.) Jessop (Muller-Doblies 89129). Bar = 500 ~m.
Venter (li?nter 13,186). Bar = 1 mm. C, L. parvifolia S. Venter (li?nter s.n.).
Bar = 1 mm. D, L. ensifolia (Eckl.) S. Venter retn~(
13,278). Bar = 500 .m~
E, L. apertiflora (Bak.) Jessop (Mauve et al. 179). Bar = 500 ~m.
F, L.
leptophylla (Bak.) S. Venter retn~(
13,251). Bar = 500 .m~
G, L. asperifolia
(Van der Merwe) S. Venter (li?nter 13,382). Bar = 0.86 mm. H, L. zebrina
and I, L. hypoxidioides (Sch6nl.) Jessop
(Bak.) S. Venter. Bar = 500 ~m.
(~nter
13.311). Bar = 500 .m~
LEDEBOURIA IN SOUTH AfRICA
34
MORPHOLOGY
LEDEBOURlA IN SOUTH AFRICA
3S
MORPHOLOGY
The anthers in Ledebouria are epipeltate and dehisce longitudinally. They are
normally yellow but can be white, pale yellow, pale violet or deep violet in
colour (Figure 12B).
Only a few publications deal with pollen morphology of the South African
species of the Asparagales. Pollen in Ledebouria is mono-sulcate. The exine is
reticulate (Erdtman 1969; Faegri & Iversen 1975). Morphology of grains
according to Jessop (1970) is as follows: grains 37.7 - 61.5 i'm long,
monosulcate; exine smooth to reticulate with the lumina less than 0.1 i'm long
and decreasing in size towards the poles. The degree of reticulation also varies
intra specifically (Figure 12C & 12D).
Our examination reveals that pollen is consistent throughout the genus. Slight
variation occurs in shape and size of the pollen infra-specifically (Figure 12).
5.8.3. Ovary.
The length of the stigmatic lobes differs slightly between certain species.
Initially the lobes are appressed but part when the stigma reaches maturity.
The length of the style at an thesis varies in different species. As the flower
opens the style curves outwards so that the stigma protrudes away from the
perianth during elongation and eventually becomes erect thus avoiding contact
with the anthers. The stigma can be lower than, at the same level as or held
higher than the anthers when receptive.
The style is terete or triangular in cross-section and is white, green, purple or
with the upper part purple and the lower part white. It varies in length from 4.0
- 6.0 mm and in some species is shorter than the ovary (Thble 6).
Ovary shape is taxonomically useful (Table 6, figure 13). The ovary is 3-lobed
in L. galpinii, L. monophylla and L. ovalifolia and 6-lobed in the remaining
species. If ovary shoulders are present, these are tapered into the style,
rectangular or raised. Basal lobes of the ovary are sometimes present (Figure
11) with nectaries on these in some species (Figure 12F). The stipe is usually as
long as it is broad varying from 0.25 - 2.0 mm.
L£DEBOURlA IN SOIJrH AFRICA
36
MORPHOLOGY
Table 6.
Ovary and fruit characters.
1
= length in mm.
2
= width in mm.
3
= Style length:
ovary length
FRUIT
OVARY
SPECIES
1
Lobe shape
L. apertijlora
L. asperi/olia
L. atro-brunnea
L. con,color
L. cooperi
L. coriacea
L. crispa
L. dolomiticola
L. ensifolia
L. floribunda
L. galpinii
L. glauca
L. hypoxidioide.s
L. inquinata
L. lepida
L. leptophylla
L. luteola
L. macowanii
L. marginata
L. minima
L. monophylla
L. ovalifolia
L. ovatifolia
L. papillata
L. parvifolia
L. petiolata
L. revoluta
L. rupestris
L. sandersonii
L. socialis
L. undulata
L. viscosa
L. zebrina
obtusely deltate
narrowly transversely elliptic
obtusely deltate
narrowly transversely elliptic
narrowly transversely oblong
obtusely deltate
obtusely deltate
narrowly transversely oblong
narrowly transversely oblong
widely trullate
depressed ovate
narrowly transversely oblong
obtusely deltate
obtusely deltate
depressed ovate
narrowly transversely oblong
obtusely deltate
obtusely deltate
narrowly transversely oblong
depressed ovate
depressed ovate
depressed ovate
narrowly transversely elliptic
depressed ovate
narrowly transversely oblong
obtusely deltate
narrowly transversely elliptic
obtusely deltate
narrowly transversely oblong
obtusely deltate
narrowly transversely oblong
obtusely deltate
obtusely deftate
1.0
1.0
1.0 - 1.5
1.0 - 1.5
2.0
1.25
2.5
1.5
1.0
1.75
1.5
0.5
1.5
1.0
0.8
1.0
3.0
1.5
0.75
0.5
1.0
1.5
1.75
1.5
1.0
1.0
0.75 - 1.0
1.0
1.0
2.0
1.5
1.0
2.25
3
2
.,
1.2
2.0
2.0 - 2.5
2.5
4.0
2.0
3.0
2.5
2.5
1.5
1.0
1.0
3.0
1.8
1.25
2.0
4.0
2.0
1.5
1.12
2.0
2.0
3.0
2.5
2.0
2.0
1.75 - 2.0
2.0
2.0
2.5
2.0
2.0
3.0
1.5: 1
5: 1
2: 1
4: 1
1.5:1
3:1
1.5:1
2: 1
1.5: 1
8:1
2: 1
6: 1
3: 1
3: 1
4: 1
3:1
1: 1
1.2:1
4: 1
4: 1
1.5:1
1.3:1
2: 1
2.3: 1
3: 1
2.5: 1
7: 1
1.5:1
3: 1
1.5: 1
3: 1
1.5: 1
2: 1
shape
clavate
globose
clavate
cylindrical
globose
globose
globose
globose
globose
clavate
globose
globose
globose
globose
globose
globose
globose
clavate
globose
clavate
globose
globose
globose
globose
globose
globose
clavate
clavate
globose
globose
globose
clavate
clavate
5.8.4. Capsule and seed.
Jesso? .0975), used diffrrent ,types of seed surface to form five major groups in
the Llhaceae sensu lato:
(\
(
1.
2.
3.
4.
5.
The Massonia Group.
The Ornithogalum Group.
The Undulate Margin Group.
The Angular Cell Group.
The Drimia Group.
LEDEBOURIA IN SOlJfH AFRICA
37
MORPHOLOGY
Ledebouria can be grouped into Jessop's Massonia Group. According to Jessop
(1975), cells are arranged in regular rows, more or less 4-angled and 15.0 25.0 JLm in diameter.
The fruit is a 3-lobed, loculicidal capsule but some of the locules can abort
leaving a 2-lobed or I-lobed fruit. The shape is cylindrical, globose or clavate
with the base either truncate or tapering. The outer and inner surfaces of the
valves are smooth.
Seed shape varies considerably within species. The testa is black, reddishbrown, brown or yellowish-brown and variously wrinkled. The material studied
has mostly 5-angled or 6-angled cells randomly arranged. They are 21.5 - 43.0
x 13.0 - 21.5 JLm. Seed length varies from 1.0 - 5.0 mm.
5.8.5. Caryology.
Only three papers on Ledebouria cytology have been published (GimenezMartin 1959, Fernandes & Neves 1962 and Jessop 1972b).
Materials and methods are outlined on page 6.
Gemenez-Martin (l.c.) gives a somatic number of 12 for L. leptophylla and
Fernandes & Neves (l.c.) give a somatic number of 24 for L. apeniflora. Jessop
(l.c.) reports on meiotic chromosomes from pollen mother cells of Ledebouria.
The chromosomes are depicted in the form of idiograms. The results of his
study are given in table 7.
Table 7. Results of Jessop's chromosome studies (1972).
Meiotic chromosomne count n= 10 - 17
Species
L. cooperi
n = 18
L. concolor
L. marginata
L. revoluta
L. socialis
LEDEBOURlA IN sOurH AFRICA
n
=
27
L. ovatifolia
n = 30
n = 46
L. floribunda L. luteoia
MORPHOLOGY
38
Figure 14. Meiotic chromosomes (metaphase I) L. apertiflora: 2n = 26. A - C
from Venter s.n. and L. ensifolia: 2n = 30. D - F from Venter 13,278. A, C
and E X 1000; B, X 600 and F X 400.
UDEBOURlA IN SOIJTH AFRICA
39
MORPHOLOGY
UDEBOURJA IN SOlITH AFRICA
Map 1.
Pictorialized map showing the variation in leafshape, in L. marginata (Bale.) Jessop. A, ll?nter 13,487. B, ll?nter
13,246. C, ll?nter 13,358 and lenter s.n.
12°
14°
18°
16°
20°
22°
24°
26°
28°
30°
32°
34°
s
jtr
~
t"'"
g
-<
20°
'\
22°f--
____7
I
t1
i
220
~
z
fS
;j
~
\
---132°
32°r
ctttt±tYJn'f I I I]'4
I
12°
l.S.O. Mise. 8091
14°
16°
18°
km I??
20°
I
,
!
?
22°
l?O
2YO
24°
26°
28°
3?O 4?O 5?O 6?O 7?O 8?O km
30°
32°
34°
I
-
0
36°
G.P.·S. 13--0959
~
41
MORPHOLOGY
Jessop (1972b) gives somatic numbers for L. ensijolia ranging from 20 to 30.
Jessop (I.e.) detected no evidence for abnormalities, during meiosis in his study.
It was found that the chromosomes did not fall into natural size categories. No
correlation was found between chromosome numbers and distribution or
chromosome size. Jessop misidentified some of the material he used for
chromosome counts. This led to the great variation in chromosome numbers he
encountered for some of the species.
The present author reports somatic numbers of 26 for L. apertijlora and 30 for
L. ensijolia (Figure 14). No basic number is given here for Ledebouria. This
would only be possible after chromosome counts were done for all the South
African species.
LED£BOURIA IN 50lJfU AFRICA
42
INFRAGENERIC CLASSIFICATION
6.0. INFRAGENERIC CLASSIFICATION.
6.1. Phenotypic variation.
The phenotype results from interactions between the genotype and the
environment. The extent to which the same genotype can give rise to different
phenotypes is known as phenotypic plasticity. The degree of plasticity is thus
under genetic control (Richards 1986). Many phenotypic modifications are also
adaptive (Grant 1963). To distinguish phenotypic from genotypic variation, a
thorough study must be made of populations in the field, as well as growing and
studying plants under uniform conditions.
The first detailed studies of natural populations of Ledebouria were undertaken
by Jessop (1970) who last revised the genus. The long lists of synonyms for
certain species reflect the phenotypic variability in this genus.
In the present study populations of each species were examined in the field to
account for intraspecific variation.
Leaf arrangement and floral structure remain mostly unaffected but bulb size,
leaf dimensions and maculation and flowering time vary. In the case of L.
ensifolia, for example, the shape and colour of the leaves are more plastic than
the flower shape and flower colour.
6.2. Genotypic variation.
The occurrence of many small populations with occasional transfer of genes
may be an antidote for the loss of genetic diversity. If a population is smaller
than approximately 50 effectively breeding individuals there is an increased risk
of fixation of alleles and loss of diversity due to inbreeding (Richards 1986).
LEDEBOURIA IN SOlITH AFRICA
43
INFRAGENERIC CLASSIFICATION
6.3. Designation of ranks.
6.3.1. Species rank.
For pragmatic purposes the rank of species is given to taxa where at least two
stable discontinuous characters are present (abrupt variation). In Ledebouria
three major convergent characters recognized. Fusiform roots occur in both L.
apertijlora and L. ensifolia. In the arid habitats of these species, the storage
function of the bulb is primarily taken over by the roots. Loose bulb scales
occur in L. sandersonii, L. monophylla and certain forms of L. cooperi. This
character commonly occurs in species growing in moist or marshy ground.
Three typical grassland species, L. luteola, L. marginata and L. ovatifolia
possess multiple thread bundles in the bulb scales and leaves. These different
species developed this character independently possibly as an adaptation to
herbivory.
A solitary inflorescence, reflexed tepals, many-lobed ovary and glabrous leaves
are regarded as primitive characters. According to this combination of
characters, L. coriacea, L. crispa, L. dolomiticola and L. undulata are
considered primitive. It is postulated that these primitive features have been
successfully retained as the above-mentioned species occur in dense populations
or grow with individual plants closely packed together.
6.3.2. Species arrangement and sectional ranks.
The objective was to produce a natural classification with taxa in monophyletic
groups. Nine sections and nine subsections were identified. The classification of
sections is based on: mature bulb size, bulbil occurrence; leaf number, texture,
occurrence of helical threads; ovary morphology and capsule shape.
LEDEBOURJA IN SOlITH AFRICA
44
PHENOLOGY AND LEAF PERIOD
7.0. PHENOLOGY AND LEAF PERIOD.
7.1. Flowering period.
Thble 8. Flowering period (.) of the genus Ledebouria in South Africa.
Species
apertijIora
asperijolia
atro-brunnea
concolor
cooperi
coriacea
crispa
dolomiticola
ensifolia
floribunda
L. galpinii
L. glauca
L. graminifolia
L. hypoxidioides
L. inquinata
L. lepida
L. luteola
L. macowanii
L. marginata
L. minima
L. monophylla
L. . ovalifolia
L. ovatifolia
L. papillata
L. parvifolia
L. petiolata
L. revoluta
L. rupestris
L. sandersonii
L. socialis
L. undulata
L. viscosa
L. zebrina
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
J
F
Month
M A M J J A S 0 N D
•
• •
• • • •
• •
•
•
•
•
•
• •
•
•
•
•
•• ••
•
• • • •
• • • •
•
• • • •
•
• •
• •
•
•
•
•
• •
• •
•
•
• •
• •
•
• • •
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
• • •
All Ledebouria species flower either at the beginning of the rainy season or well
into it (September to April).
LEDEBOURlA IN SOlITH AFRICA
Thble 4.
I
Phenology of the genus Ledebouria in South Africa.
0<
~
§
Phenology of Ledebouria in South Africa.
~
g
12
I
,
10 -+ ------..-
m
..... '-"'--
-------_
__ ._._._-_. __ .
__._---_._. __ .._------------------_ .. _-_ .. _----------------_..
c
~
2:
~
~
~
M
8 --1._-_.__ .
o
n
t
h
s
6
4
2
o
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19 20
21
22
23
24
25
26
27
28
29
30
31
32
33
Species
,
I-
I
In leaf
-
In flower!
~
46
PHENOLOGY AND LEAF PERIOD
7.2. Pollination biology.
During mid-morning light intensity and temperatures are high, acting as
stimulants for insect activity (Faegri and Van der Pijl 1979). The beginning of
nectar secretion reflects the activity period of the pollinator class and is in
general one to four hours before the pollinators become active (Bentley &
Thomas 1983). Nectar secretion in the flower can be affected by local climatic
conditions such as temperature and rain (Bentley & Thomas 1983).
There appears to be a close correlation between pollinator needs (nectar, pollen
and oil) and floral attraction (colour, shape and odour) (Rebelo 1987). These
are primary and secondary attractants respectively.
In Ledebouria specific flower colour does not seem to be a determining factor.
Bees were seen visiting flowers ranging from green to purplish-pink. U. V.
reflectance would possibly throw more light on this. The scape and rachis are
brightly coloured in some species and may contribute to pollinator attraction.
In most ecosystems, social bees are the dominant pollinators (Richards 1986).
Bees are associated with all flower types except those with narrow tubes. Pollen
from a flower of a particular species is unlikely to be received by a flower of
the same species. Because of this more energy is required for pollen production
per ovule fertilized (Richards 1986). For short distance attraction the surface
texture of the flower together with odour are important factors (Faegri & Van
der Pijl 1966).
Throughout the distributional range of Ledebouria, the butterfly fauna is very
rich except in the southern and south-western Cape which has a relatively
depauperate butterfly fauna (Cottrell 1978).
Wasps are opportunistic foragers and have a low fidelity to specific flowers
(Rebelo 1987).
In Ledebouria, the flowers are numerous and born on long inflorescences with
unsophisticated shapes and simple pollen presentation and receipt.
LEDEBOURlA IN SOlITH AFRICA
47
PHENOLOGY AND LEAF PERIOD
In the case of Ledebouria, nectar plays a minor role in attraction and was
observed only in L. cooperi and L. revoluta. Extrafloral nectaries are situated on
the flowering axis just above the pedicel. These amorphous nectaries secrete
nectar seen as small shiny drops of liquid. Other nectaries occur as large
papillae at the base of the ovary lobes (Figure 12F).
Pollinators visit the flowers of Ledebouria from mid-morning (lOhOO) to early
afternoon (15hOO).
Lepidoptera: (Psychophily)
The only butterflies recorded visiting Ledebouria flowers, are from the family
Lycaenidae, the largest family of southern African butterflies (Dickson & Kroon
1978; Migdol 1987). In the north-eastern Transvaal, on the high-lying areas,
Euchrysops subpallida Bethune-Baker and Lepidochrysops patricia Trimen (Vari
& Kroon 1986), were recorded visiting the flowers of L. revoluta and L.
luteola.
Hymenoptera:
Wasps. (Vespidophily)
On two occasions wasps were seen visiting flowers of Ledebouria.
Bees. (Mellitophily)
The main bee pollinator of Ledebouria is Apis mellifera adansonii (Honey bee).
Specimens of the stingless bee (Trigona gribodoi) were collected at Tzaneen in
the eastern Transvaal, visiting the inflorescences of L. revoluta (Skaife 1979).
These specimens were covered with L. revoluta pollen. The reward for the
pollinator of Ledebouria is only pollen which is used as an energy source. Long
range attractants for bees include colour and scent. L. sandersonii, L. cooperi,
L. monophylla and L. zebrina have a strong, sweet and fresh odour that attracts
bees.
LEDEBOURIA IN SOlITH AFRICA
48
SEED DISPERSAL
8.0. SEED DISPERSAL.
8.1. Wind dispersal.
In many of the species the erect infructescence and capsules are adapted to seed
dispersal by wind (anemochory). The whole infructescence moves about in the
wind and the seeds fall to the ground where they are sometimes further
dispersed by water (Van der Pijl 1969).
8.2. Water dispersal.
Exposed seeds in the capsules of L. revoluta were marked with red vegetable
dye to make sure that only those seeds were used for distance measuring.
Distances travelled by the marked seed were measured after light and heavy
rain storms.
The longest distance recorded for seed of any Ledebouria species transported by
water, is 5.2 meters from the mother plant.
The most important method of dispersal of Ledebouria seed is by hydrochory
with rainwash (ombrohydrochory).
LEDEBOlfRlA IN SOUfH AFRICA
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
49
9.0. ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES.
Acocks (1988) "Veld Types of South Africa" was used for habitat description.
Vegetation structural groups and formation classes follow the system of
Edwards (1983).
The genus Ledebouria occurs in both winter and summer rainfall areas.
The veld types with the highest species density are predominantly woodland
with large areas of grassveld or pure grassveld types.
Ledebouria species occur in the following vegetation types:
Coastal Tropical Forest
Coastal Forest and Thornveld (10 spp.)
Pondoland Coastal Plateau Sourveld (8 spp.)
Ngongoni Veld (10 spp.)
Eastern Province Thornveld (7 spp.)
Inland Tropical Forest
North-eastern Mountain Sourveld (18 spp.)
Lowveld Sour Bushveld (7 spp.)
Tropical Bush and Savanna
Lowveld (4 spp.)
Arid Lowveld (3 spp.)
Turf Thornveld (3 spp.)
Arid Sweet Bushveld (6 spp.)
Mopani Veld (2 spp.)
Kalahari Thornveld (3 spp.)
Mixed Bushveld (6 spp.)
Sourish Mixed Bushveld (4 spp.)
Sour Bushveld (13 spp.)
LEDEBOURIA IN SOUTH AFRICA
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
so
Karoo
Valley Bushveld (11 spp.)
Karroid Broken Veld (4 spp.)
Central Upper Karoo (2 spp.)
Arid Karoo (2 spp.)
Succulent Karoo (1 sp.)
Namaqualand Broken Veld (1 sp.)
Temperate and Transitional Forest and Scrub
Highland and Dohne Sourveld (21 spp.)
Natal Mist belt 'Ngongoni Veld (12 spp.)
Coastal Fynbos (4 spp.)
Pure Grassveld
Cymbopogon - Themeda Veld (6 spp.)
North-eastern Sandy Highveld (6 spp.)
Themeda - Festuca Alpine Veld (10 spp.)
Stormberg Plateau Sweetveld (6 spp.)
False Grassveld
Natal Sour Sandveld (6 spp.)
Pietersburg Plateau False Grassveld (11 spp.)
L. cooperi, L. monophylla and L. minima are commonly found growing in
seepages, especially in mountain terrain but only L. cooperi is recorded growing
in water (mostly running water).
Twelve species of Ledebouria are recorded from open short grassland which is
common in mountainous and flat areas throughout South Africa. Of these L.
inquinata, L. asperifolia, L. minima, L. parvifolia and L. sandersonii favour
grassy mountain slopes. L. cooperi, L. leptophylla, L. luteola, L. marginata, L.
UDEBOURIA IN SOUTH AFRICA
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
51
ovalifolia, L. ovatifolia and L. revoluta favour flat to gently undulating areas.
Ten species are found in open low grassland to which L. petiolata is restricted.
L. monophylla, L. rupestris, L. leptophylla, L. galpinii, L. minima and L.
sandersonii are typically small grassveld species of high altitude.
L. ovatifolia was found growing in closed tall grassland of the secondary dunes
at Scottburgh on the Natal South Coast. These plants have atypical spreading to
erect leaves in contrast to the usually flat leaves of this species. The long thick
grass cover is possibly the cause of this growth pattern. Plants collected from
this population produced normal leaves the next season.
Ledebouria occur in evergreen shrubland at high altitude. On the Karoo and
Namaqualand Flats L. undulata grow sparingly between short evergreen shrub
in full sun.
27 Species were recorded growing in some form of Woodland with the largest
number of Ledebouria populations in deciduous woodland. There is a
correlation between the diversity of Ledebouria species and the occurrence of
deciduous woodland. Most species occur in low deciduous woodland (19
species).
Evergreen woodland is typical of high rainfall areas in South Africa, on the
mountain tops of the Transvaal and the coastal flats of the eastern Cape and
Natal. Of the five Ledebouria species in the evergreen low woodland, only L.
eoneolor and L. ensifolia grow on the coastal flats with L. eoneolor favouring
the shady areas. In the eastern Transvaal, L. asperifolia grows on rock outcrops
and steep mountain slopes covered with evergreen woodland. L. floribunda
occurs commonly in deep or light shade in evergreen tall woodland.
Most of the species (21 spp.) occur predominantly on mountain or hillslopes
(Table 10). Eight species occur on flatlands along the coast and on plateau
areas on the mountains. Of these, L. viseosa occurs only on inland plateaus near
Thabazimbi. L. minima occurs on plateaus of the Transvaal Drakensberg escarp
with L. eoriaeea and L. ovalifolia on coastal flats in the Cape Province. There
are relatively few species groYiing on hill or mountain tops. Occasionally plants
of L. asperifolia, L. inquinata and L. revoluta are found growing in these areas.
UDEBOURlA IN SOlJfH AFRICA
52
ECOLOGY, HABITAT AND ADAPTIVE SfRATEGIES
Table 10. Aspect, slope and topography of Ledebouria in South Africa
SPECIES
ASPECT
SLOPE
TOPOGRAPHY
(degrees)
L. apertiflora
S,W
0-30
Flats and low hills.
L. asperifolia
E,W
5-90
Flats and mountain slopes.
L. atro-brunnea
L. concolor
S,E
S,E
15-30
0-15
L. cooperi
N,S,E,W,NE,SW
5-45
Flats to mountain slopes.
L. coriacea
E,SE
0- 5
Lowlands near sea.
L. crlspa
N,E
5-10
Hill slopes.
L. dolomiticola
E,ES
30-90
Mountain slopes.
L. ensifolia
L. floribunda
N,W
10-30
Flats and mountain slopes.
W
15-90
Mountain and hill slopes.
Hillslopes.
Lowlands near sea.
L. galpinii
L. glauca
N,W,SW
0-10
Mountain tops.
S,E,W
0-10
Flats.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
hypoxidioides
inquinata
N,E
N,S,E,W
5-10
0-45
Hill slopes.
Flats and hill slopes.
lepida
W
0- 5
Mountain slopes.
leptophylla
N,S,E,W
0-30
Flats to mountain tops.
luteola
W
0- 5
Flats to hillslopes.
macowanii
S,E
0-30
Mountain and hill slopes.
marginata
S,E,SE
5-30
Flats to mountain tops.
minima
W
S,E,SW
5-10
Flats to mountain tops.
0-15
Mountain tops.
E
N,S,E,W,SE
0- 5
Flats.
0-30
Flats to mountain tops.
N,SE
0-15
Flats and mountain slopes.
E
S,W
15-30
0-30
Mountain slopes.
Mountain tops.
N,S,E,W,NE,NW
0-45
5-10
15-30
5-15
Flats to mountain tops.
Mountain tops.
monophylla
ovalifolia
ovatifolia
papillata
parvifolia
petiolata
revoluta
rupestris
sandersonii
L. socialis
L. undulata
L. viscosa
L. zebrina
E
S
N
S,W
Mountain slopes and tops.
Flats to hill slopes.
0-15
Flats.
S
0- 5
Flats.
E
15-45
LEDEBOU1lJA IN SmITH AFRICA
Mountain slopes.
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
S3
Of the 33 species, 25 are commonly found in full sun. Some of them, (19 spp.)
are restricted to this exposure with 18 species found growing in various degrees
of shade. Only four of these (L. concolor, L. floribunda, L. rupestris and L.
socialis) tolerate dense shade. Some species occur in the shade of trees as well
as in full sun (L. asperijolia and L. zebrina), but here exposed plants are more
robust.
The genus is concentrated in the moist eastern part of South Africa with only L.
undulata in the dry Karoo and Namaqualand.
The lowest temperature experienced across the geographical range of
Ledebouria is -7 °C with 47 °C the maximum in the Limpopo Valley and at
Upington. Night temperatures are mostly mild, day temperatures soar to a
maximum in January.
Frost occurs over most of South Africa (0 - 40 days) except in areas near the
coast, Transvaal Lowveld and the northern Transvaal. The Drakensberg in the
eastern Cape, Natal and Lesotho receive more than 150 days of frost per annum
and snowfall in winter (Stamp & Morgan 1972). On the high plateau of the
Drakensberg soil temperatures frequently drop below freezing point. Only L.
sandersonii tolerates these conditions.
The distribution of Ledebouria is closely associated with the annual
precipitation. Approximately 27% of the surface is medial (600 - 1000
mm/annum) and 4 % of the surface is moist receiving 1000+ mm per annum
(Barnard et al. 1972). The highest diversity of species and density in numbers
occurs in the medial and moist areas.
Winter rain predominates in the south-western and southern Cape from April to
September. An intermediate area that receives summer and winter rains extends
as a narrow belt from behind the fold mountains of the Cape Province up to
Fraserburg to Goodhouse on the Orange River (Cowling & Roux 1987). In
summer rainfall areas, rain falls mostly from October to March.
L£DEBOURIA IN SOlITH AFRICA
-
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
S4
Mist occurs over the high lying areas of the great escarp and isolated mountains
scattered throughout South Africa. Most of the endemic Ledebouria species (L.
galpinii, L. monophylla, L. parvijolia, L. petiolata, L. rupestris and L.
sandersonii) occur in this mist belt.
Plants of Ledebouria occur from sea level on the coastal plain up to 3,400
meters above sea level at Njasuthi on the Natal Drakensberg Escarp above
Giant's Castle Game Reserve. However more species occur on the high lying
areas than on the low lying areas of the interior and coastal plain.
Twelve species were found growing on sandstone of the Witteberg, Natal,
Beaufort and Uitenhage Groups (SACS 1980). Of these, only L. ovalifolia
seems to be restricted to sandstone.
Dolomites are frequently favoured by Ledebouria with six species commonly
occurring on this substrate. L. dolomiticola has only been recorded from the
dolomites of the Eccles Formation in the Chuniespoort Group (SACS 1980).
Five species of Ledebouria occur on the chert rocks of the Sheba Formation in
the Fig Tree Group but none of the species seem to be restricted to this rock
type.
Granites occur either as intrusions or as part of the basement structure (Du Toit
1956; SACS 1980). No species of Ledebouria are restricted to granitic soils. Six
species were found on granite with L. revoluta the most common.
Dolerite intrusions in the form of sheets, dykes and sills (Blatt et al. 1972;
Truswell 1970; Thornbury 1969) occur over most of South Africa, associated
with characteristic clay to clay-loam soils. Four species were found in
association with this substrate.
Three species were found on andesitic tuff and lava of the Westonaria
Formation of the Ventersdorp Supergroup. L. ovatifolia and L. revoluta are the
most common species on this rock type. Schists and serpentinite occur in the
Komati Formation of the Onverwacht Group at Barberton and near Pietersburg
in the Mothiba Formation of the Pietersburg Group (SACS 1980). Four species
occur on these rocks with L. crispa restricted to the schists and serpentinites in
the Pietersburg Area.
LEDEBOURIA IN sourH AFRICA
55
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
One species, L. hypoxidioides, occurs on Dwyka tillite, but is not restricted to it
(TruswellI970).
L. coriacea and L. papillata from the Port Elizabeth Area occur on Tertiary
limestones of the Alexandria Limestone Formation (SACS 1980) with L.
coriacea endemic to this area (Map 15).
Only two Ledebouria species show in their distribution to be associated with
specific geological formations. L. crispa is a serpentinite endemic so far only
recorded from the serpentinite and schists of the Mothiba Formation in the
Pietersburg Group. L. dolomiticola is endemic to the dolomites of the Eccles
Formation in the Chuniespoort Group in the Strydpoort Mountains south of
Pietersburg.
The genus Ledebouria occurs on nearly all soil types with the exception of some
species that occur only on certain soil types (Table 11, figure 15a). Thirteen
Ledebouria species occur commonly on quartzite derived soils and L. galpinii,
L. lepida, L. monophylla, L. rupestris and L. viscosa may be restricted to
quartzitic soils.
Thble 11. The major soil types (Buckman & Brady 1969):
Sand
Sand 100%
Loamy sand
Sand 85%
silt 10%
clay 5%
Sandy loam
Loam'
Sand 65%
Sand 45%
silt 25%
silt 40%
clay 10%
Silty loam
Sand 20%
silt 60%
clay 20%
Silty clay loam
Sand 15%
silt 55%
clay 30%
Clay loam
Sand 28%
silt 37%
clay 35%
Clay
Sand 25%
silt 30%
clay 45%
Methods and Materials for soil type evaluation on page 9.
LEDEBOURlA IN SOUTH AFRICA
clay 15%
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
S6
Clay loam and sandy soil seem to be the most frequently inhabited substrate.
Dolerite, granite, chert breccia, dolomite, lava and serpentinite are the mother
material for the clay loam soils. Seven species were found growing in clay loam
with a fine granular to coarse granular structure. Six species were found
growing on various forms of lithosols that vary from 10 mm thick on rock
sheets to 3000 mm thick on scree slopes. L. marginata and L. ovatifolia are the
only species found on gritty sandy loam.
Sandy soils are derived from sandstone, quartzite, conglomerate and granite.
Various structures and a percentage of humus occur in the sandy soil types. Five
species occur on gritty sandy soils especially on small plateau areas on gentle to
medium slopes. These soils are always shallow, 50 - 200 mm deep.
Deep sandy soil with a loose structure is common in the northern and western
Transvaal, western Orange Free State and north-central Cape Province. These
soils are aeolian and occur on limestone or calcrete (Van der Merwe 1962).
Three species were found growing in this substrate and L. viscosa is restricted
to it.
Sandy loams are distributed throughout South Africa. Sandy loam is derived
from granite, quartzite, conglomerate, schist or sandstone. In mountainous
areas, sandy loam lithosols are common and seven species occur on them. L.
atro-brunnea and L. zebrina are endemic to this substrate. On small mountain
plateaus the sandy loam tends to be of a gritty nature harbouring four species of
Ledebouria. Of these, L. lepida is endemic to the substrate. Humus-rich sandy
loam is an uncommon soil type, occurring only in mountainous areas of Natal
and the eastern Transvaal. L. floribunda, L. petiolata and L. rupestris were
collected growing in this soil, often in shady conditions.
Loamy soils are commonly associated with quartzite, chert, ferrous shales and
tillite. Four Ledebouria species were collected in loamy lithosols and three
species in gritty loam. No species were encountered that are restricted to loamy
soils.
Of the seven biomes in South Africa (Rutherford & Westfall 1986), Ledebouria
is represented in the Savanna, Grassland, Nama-Karoo, Succulent-Karoo and
Fynbos biomes. The Savanna and Grassland biomes carry the largest number of
Ledebouria species (Chapter 10).
LEDEBOURJA IN SOlrm AFRICA
ECOLOGY, HABITAT AND ADAPTIVE STRATEGIES
57
Figure 15a. Graphic guide for sandy soil textural classification with A, the
different types of sandy soil for Ledebouria (shaded portion) and B, soil textures
of the habitats (shaded portion).
LEDEBOURIA IN sOurH AFRICA
~
§
.:<
==
~
Note: Velues ere expressed
es percenteges of the
10lal sand.
coarse sand
medium sand
fine sand
2,0-0,5 mm
0.5-0.2 mm
0.2-0.02 mm
100
sand
silt
clay
2,0 -0,02 mm
0,02-0,002 mm
0,002 mm
<
<e.
<....
t:l
c
~
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~
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Si
::
,.~
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V_
\/
-.
-
V
-
\/
......
%COARSE SAND
SAND GRADE CHART
A
_.. ~
\Y.:;.i£.·...'§c~·.-
.. ""
%
~
Cb
~
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~
--
TEXTURE CHART
~
<a
rt>
~
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'~SAND
8
VI
go
S9
PHENETICS AND RELATEDNESS
10.0. PHENETICS AND RELATEDNESS.
Rohlf and Sokal (1981) express the view that, if a classification is intended to
represent a similarity scheme, it is phenetic and if it is intended to show
evolutionary branching sequences it is cladistic. Similarly Cain and Harrison
(1960) "refer to arrangement by overall similarity based on all available
characters without any weighting as phenetic". A phenetic classification is thus
based upon the overall present-day similarities and differences of living
organisms. The only approach to phenetic classification is when the
classification is performed by phenetic similarity clustering. The phenetic
approach to classification uses all available characters of the species being
classified without a priori weighting. The diagnostic feature of phenetic
classification is to provide a convenient general-purpose framework for
accommodating the diversity of plants (McNeil 1978).
The programme NTSYS-pc (Numerical Taxonomy and Multivariate Analysis
System) has been used in this -thesis to analyse data sets for the species of
Ledebouria in South Africa. NTSYS-pc is a system of programmes that is used
to find and display structure in multivariate data. This programme can be run on
an IBM compatible PC (XT or AT) computer with RAM memory capacity of
360 K. To run the programme consult the manual "NTSYS-pc, Numerical
Taxonomy and Multivariate Analysis System" (Rohlf 1988).
To describe the relationship amongst taxa it is necessary to reduce dimensions of
space by using cluster analysis. Cluster analysis produces a hierarchical
classification of species based on a similarity matrix. An interactive file for
cluster analysis and coordinates analysis based on the data matrix of the
Ledebouria species for South Africa is used. To arrive at a phenogram, it is
essential to follow a strict series of steps making use of various programmes in
NTSYS-pc.
Clusters which are identical in the analyses are called "ball clusters". These
would be the only clusters in a strict consensus tree between these two trees
(Rohlf 1988). These are usually the most distinct clusters in a set of data.
The tree matrix is an efficient method for describing a system of nested clusters
where each cluster has a corresponding level (Rohlf 1988).
LEDEBOURlA IN SOIJTH AFRICA
60
PHENETICS AND RELATEDNESS
The DELTA (DEscription Language for TAxonomy) programme is used for
concise representation and manipulation of taxonomic descriptions. To run
DELTA, an IBM compatible PC-XT or AT with at least 512K memory is
needed. The format-conversion programme, CONFOR, converts the coded
description into natural language and into formats required by programmes for
key-generation, interactive identification, and numerical classification (Dallwitz
& Paine 1986).
There are four major types of output from the DELTA-format data: the printed
key, interactive key, natural language descriptions and a summary of the data.
To run the programme DELTA, refer to the" User's Guide to the DELTA
System" (Dallwitz and Paine 1986).
One of the major advantages of DELTA is the output of descriptions into natural
language. The descriptions in chapter 12.0 are all generated by DELTA.
SUMMARY gives information on the character, type of character, number of
states, distribution of states, items coded, standard deviation as well as
t~e
minimum and maximum for that character. This is a great help when two or
more species are compared or differences between species studied.
INTKEY is an interactive programme for identifying a specimen by comparing its
characteristics with stored descriptions of species. The INTKEY programme was
used for various information retrieval computations. The most important
computation is the identification of an unidentified specimen. To identify a
specimen, information about the specimen to be identified, is entered,
eliminating species whose descriptions do not match the description of the
specimen, until only one species remains.
When comparing the data sets of the UPGMA clustering method (Unweighted
pair-group method, arithmetic average) and COMPL (complete-link method)
clustering methods, they show 9 ball clusters in common. These are the clusters
indicating the subsections Acutilobae, Erectifoliae, Stellatae, Zebrinae,
Piliferae, Asperifoliae, Glaucae, Ebractae and Longicollae .
UD£BOURlA IN SOIJTH AFRICA
61
PHENETICS AND RELATEDNESS
In the phenogram, (Figure 15b) labels of the objects being clustered are plotted
to the right and the clustering (splitting levels) is shown along the ordinate. Each
vertical line in the phenogram cut by the phenon line corresponds to a cluster
(section or species) whose members are the species connected to that fragment
of the phenogram (Rohlf 1975). The adequacy of a phenogram in representing
taxonomic structure is indicated by a high cophenetic correlation, and if r
(cophenetic correlation) = 0.8 the phenogram is satisfactory in this respect
(Sneath & Sokal 1973). The closeness of fit for the cluster analysis for
Ledebouria shows that r = 0.81428.
The electronic grouping of the sections in the phenogram corresponds well with
intuitive grouping. In this grouping, L. asperi/olia was placed within the section
Magnibulbae but the globose capsule and smaller bulb places it within section
Globosae. Initially L. undulata was in a group of its own based on the
hysteranthus habit. It is best placed in section Efiliferae due to the torn leaves
lacking threads and the distinct neck of the bulb.
The 42 phenon connates groups (sections) affiliated at no lower than 42 % on the
similarity scale used in the analysis. This phenon level was chosen as it indicates
the nine sections as well formed clusters.
In the phenogram two phenon lines are used. They indicate fixed levels of
similarity. The first line at a similarity value of 42 % indicates the 42 phenons.
These are regarded as sections. They are again divided into various subsections.
The area above the 90% phenon line indicates the different species.
The characters used to distinguish between sections are not influenced by the
environment.
The presence of a single leaf and erect inflorescence in the monotypic (L.
monophylla) Monophyllae is sufficient to distinguish it from other sections. This
section shows an interesting disjunction between the main distribution area at
Graskop and the limited range at Pietermaritzburg.
LEDEBOURlA IN SOlITH AFRICA
63
PHENETICS AND RELATEDNESS
Phenogram from tree matrix: sahnc
ID records:
"Ledebouria. for South Africa"
" STAND: input=data, divide=STD, subt=YBAR, direction=C
" SIMINT: input=sdatac, coeff=EYCLID, direction=C
" SAHN: input=simintc, method=UPGMA, tie=WARN
type=5, size=33 by 2, nc=0
0.05
0.04
0.03
0.02
0.01
re
r--
ape
ene
ean
dol
atr
Humeral;
vIa
'pt
!e~
I
I
F"r"'Iln
rup
..--
Coriaceae
cor
flo
~I '
I
r~
L---.r-
zeb
lut
::J--
O\/t
rr"'Ion
~ ': J-
-
~ r:,
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....--
rr"'I<:lt-
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t-nr:JC
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90
u:DEBOURIA IN sourH AFRICA
}.
Filiferae
Monophyllae
Bulbilia
Globosae
Efiliferae
8C~C
I
42
J-
Magnibulbae
--r-- Paucifol{ae
64
PHENETICS AND RELATEDNESS
Section Coriaceae has only one member, L. coriacea. The leathery leaves with
thickened margins and the erect inflorescence with few flowers is diagnostic to
section Coriaceae. It is postulated that L. coriacea grows in close packed
populations to ensure pollination of the flowers resulting in a good fruit
formation. L. coriacea is known from a single locality on the coast near Port
Elizabeth.
In spite of the fact that threads in torn bulb scales and leaves are a common
feature in some monocot groups, it is felt that the presence or absence of these
threads in a section is stable enough to use as a distinguishing character. Section
Efiliferae consists of members that show no threads in torn bulb scales or
leaves. The small (1.0 - 1.3 mm long) globose ovary readily separates it from
the section Paucifoliae. L. crispa and related L. undulata both have undulate
spreading leaves and solitary erect inflorescences. These species grow in full
sun and occupy gravely areas covered with little vegetation. Section Efiliferae is
widely distributed with most of the species in the Cape.
The section Paucifoliae is the only section with members (L. galpinii and L.
papillata) having less than five leaves, prominent cataphylls and a depressedovate ovary. This combination of characters is sufficient to render sectional
status. L. galpinii is a Transvaal endemic and occurs around Kaapsche Hoop in
short montane grassland. L. papillata occupies a variety of habitats in the
Transvaal and eastern Cape.
There are six species (L. cooperi, L. crispa, L. dolomiticola, L. galpinii, L.
parvifolia and L. socialis) with bulblets. Members of the section Bulbilia (L.
cooperi and L. parvifolia) show a prominent cataphyll. The bulblets and the
cataphyll distinguishes this section. L. cooperi is widely distributed in South
Africa except for the Cape with only a few widely distributed localities. L.
parvifolia is recorded from an isolated valley near Graskop in montane
grassland.
Many species (16 species) show threads when bulb scales and leaves are torn.
Members of section Filiferae (L. luteola and L. ovatifolia) reveal copious
threads in the bulb scales and leaves when torn. It is postulated that these
species are adapted to veld fires and herbivory in their grassland habitat. This
section occupies most of the eastern half of South Africa. The distribution of L.
luteola and L. ovatifolia is similar.
UDEBOURIA IN SOlITH AFRICA
6S
PHENETlCS AND RELATEDNESS
The combination of bulb scales without threads when torn and prominent
shoulders to the apex of the ovary is sufficient to support sectional status to
Humerati. L. apertiflora and related L. ensifolia have fusiform roots, thin
cylindrical bulbs, many dry bulb scales and occur in dry lowland habitat types.
These characters are adaptations to xeric conditions. Section Humerati is widely
distributed with members of subsection Erectifoliae and L. lepida and L.
rupestris of the subsection Stellatae each known from a different locality in the
central and eastern Transvaal.
In section Magnibulbae (four species), species are grouped together on the basis
of mature bulbs (flowering bulbs) 60 - 150 mm wide, inflorescence 80 - 350
-flowered and 8 - 15 mm long pedicel. This combination of characters
circumscribe the section. L. floribunda and L. zebrina have inflorescences
exceeding 200 mm in length carrying between 60 - 150 flowers. They occupy
similar habitats in montane grassland and montane woodland. Section
Magnibulbae is widely distributed in South Africa. L. hypoxidioides is the only
species in this section with a restricted distribution, in and around
Grahamstown.
Mature bulbs being 20 - 40 mm in diameter plus prominently globose capsules
characterises section Globosae. Under exceptional conditions plants in other
sections may have bulbs between 20 - 30 mm diameter but then the capsules are
not globose. L. marginata shows similar thread bundles in the leaves as section
Filiferae but does not correlate to characters such as bulb 25 - 50 mm wide,
copious threads in bulb scales and capsules not globose. The thread bundles in
L. marginata are regarded as convergent. Members of Globosae are widely
distributed in the eastern half of South Africa with L. inquinata restricted to
Transvaal.
LEDEBOURlA IN SOlJfH AFRICA
66
PHYTOGEOGRAPHY
11.0. PHYTOGEOGRAPHY.
11.1 Distribution.
The genus Ledebouria occurs in the south-eastern part of India, the Island
Nossi-Be on the north-western coast of Madagascar (Humbert 1935) and
throughout most of Africa (Map 2). Ledebouria appears to be absent from
Australia and South America, which together with India and Africa, formed part
of greater Gondwana. This is an indication that Ledebouria arose after the
separation of Africa from the rest of the Gondwana masses at the end of the
Triassic (Goldblatt 1978). In the countries north of South Africa, the largest
concentration of species occurs on the mountain ranges that receive high
rainfall. Isolated populations occur in drier areas in desert type vegetation of
Namibia and Angola (pers. obs.).
Ledebouria has its main centre of diversity in the eastern part of southern
Africa, notably Natal and Transvaal. The distribution is correlated to rainfall in
South Africa (Map 3).
The greatest species diversity (Map 4) occurs in the eastern and central
Transvaal, 8 - 14 species per degree square down to Swaziland with 10 species
in the 2631 (Mbabane) degree square to the Natal Midlands with 7 - 10 species
per degree square, whereas the Natal Uplands has 7 - 9 species per degree
square. Only the 3227 (Stutterheim) degree square in the Cape has a high
density (9 spp.). The areas in South Africa with the lowest diversity are the
Karoo, Namaqualand, northern Cape and the Kalahari. All degree squares
covering this area have between one and three species. Map 4 was compiled
from herbarium material and localities recorded in the literature.
Most of the sections in Ledebouria have a wide distribution except the
monotypic sections Coriacea (Port Elizabeth) and Monophyllae (Graskop and
Pietermaritzburg). Section Erectifoliae has three species. L. atro-brunnea
occurs in high altitude woodland on quartzite near Rustenburg. L. dolomiticola
also occurs in high altitude woodland but on dolomitic rock on the Strydpoort
Mountain. L. viscosa occurs in deep red sandy soil in low altitude woodland
near Thabazimbi. These three species are all endemic to the Transvaal.
U1JEBOURlA IN SOurH AFRICA
PIIYTOGEOGRAPIIY
67
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u:DEBOURlA IN SOlJfll AFRICA
o
o
'0
Map 3.
12°
14°
Known distribution of the genus Ledebouria Roth in South
Africa.
16°
18°
22°
20°
24°
26°
28°
34°
32°
30°
JU
18°
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70
PHYTOGEOGRAPHY
11.2. Factors promoting speciation and distribution.
Environmental fluctuations can effectively isolate populations and thereby
accelerate allopatric speciation. Under favourable conditions marginal
populations may be established. When conditions become severe these
populations may be isolated and differentiate (Valentine 1967).
Evolution tends to accelerate in arid regions where species on the margin of
their distribution range become isolated leading to specialization and eventual
speciation (Stebbins 1952, Axelrod 1972).
Mountains act as refugia for many plant taxa, especially during climatic changes
(Van Jaarsveld 1989). It can be hypothesized that during drier periods in the
past, many Ledebouria, Gasteria Duval and Othonna L. species survived on
mountains in the cooler and moister habitats with only a few sPecies that could
survive outside this refuge, in hotter and drier areas (Van Jaarsveld 1989).
Today most Ledebouria species occur on mountain ranges.
LEDEBOURlA IN SOlJfH AFRICA
71
TAXONOMY
12.0. TAXONOMY.
12.1. Generic description.
Ledebouria Roth in Novae Plantarum Species: 194 (1821); Jessop in 11 S. Afr.
Bot. 36(4): 244 (1970); Dyer in Gen. S. Afr. Fl. Plants 2: 937 (1983).
Scilla L. pro pane Baker in Flora Cap. 6: 478 (1896); Phillips in Gen. S. Afr.
Fl. Plants :191 (1951).
Type: L. hyacinthina Roth (=S. indica Bak.), from India, Curtis's Bot. Mag.
60: t.3226 (1833).
Plants solitary or gregarious. Roots wiry, contractile, fleshy or fusiform. Bulb
10 - 200 mm diameter; sometimes necked; bulb scales brown to purple; tips
truncate or elongate; basal stem occasionally present; sometimes with threads
when torn; live bulb scales loosely or tightly arranged; bulblets sometimes
produced; basal stem 0 - 120 mm long; cataphylls 0 - 3. Leaves 1 - 20; partly
or fully developed at flowering; erect to adpressed, sometimes spirally twisted;
sometimes petiolate; lamina linear to ovate; fleshy or leathery, green to purple
with or without darker green or purple markings. Inflorescences 1 - 10,
racemose, 4 - 150 -flowered; longer or shorter than the leaves. Peduncle
smooth or papillate; sometimes marked. Raceme lax or dense; cylindric to
globose. Bracts with or without bracteoles, 0.25 - 1.0 x 0.5 - 5.0 mm, deltate
to linear. Pedicels spreading; 1.0 - 15.5 mm long. Perianth almost tubular to
stellate; segments linear to oblong, green to purple sometimes with a green keel;
apex acute, obtuse or sharply upward curving. Stamens erect or spreading,
filaments white, green, pink to purple, base cylindric to slightly flattened, free
or epipetalous; anthers white, yellow or violet. Ovary stipitate, 3- or 6-lobed,
lobes variously shaped, base glabrous or papillate, distallobes present or absent,
apex of ovary tapering into the style, rectangular or raised. Style triangular to
terete; stigma papillate. Capsule trilocular, cylindrical, globose or clavate. Seed
globose, drop-shaped or disc-shaped, surface strongly wrinkled.
LEDEBOURlA IN SOUTH AFRICA
72
TAXONOMY
12.2. Key to the sections.
1.
· Section MONOPHYLLAE
Leaf single
Leaves 2 or more
2.
2.
Leaves thickly leathery, not spreading,
inflorescence few - flowered
Leaves fleshy, spreading, inflorescence dense
3.
5.
Leaves without threads when tom, more than 4;
ovary globose, 1.0 mm long
Leaves with threads when tom, 2 - 4; ovary
depressed-ovate, 1.5 mm long
5.
3.
····.· 4.
Bulb neck prominent
Bulb neck absent
4.
Section CORIACEAE
Cataphylls present; leaves without threads
when tom, bulblets often present
Section EFILIFERAE
Section PAUCIFOLIAE
Section BULBILIA
Cataphylls absent; leaves with threads when
torn
6.
6.
Bulb scales and leaves with copious threads
when tom
Bulb scales and leaves lack copious threads
Section FILIFERAE
when tom (although a few are often
present)
7.
7.
Bulb scales lacking threads when tom; apex of
ovary forming prominent shoulders
..................................................... Section HUMERATI
Bulb scales producing some threads when tom;
apex of ovary lacks prominent shoulders
LEDEBOURlA IN SOIITH AFRICA
8.
73
TAXONOMY
8.
Mature bulbs 60 - 150 mm wide; raceme cylindric;
capsules variously shaped but not globose
............................................... Section MAGNIBULBAE
Mature bulbs 20 - 60 mm wide; raceme not
cylindric; capsules globose
Section GLOBOSAE
12.3. Key to the species.
1.
2.
Bulb epigeal
2.
Bulb semi-epigeal
3.
Bulb hypogeal.. .
4.
Inflorescence erect; bracts fleshy; apex of
ovary not forming shoulders; seed black;
plants gregarious; eastern Cape
Inflorescences flaccid; bracts membranous;
30. L. socialis
ovary shoulders present; seed brown;
; north-eastern Transvaal
3.
4. L. dolomiticola
Inflorescence erect; bulb scales attenuate,
dead bulb scales brown; ovary shoulders
tapering into the style; north-eastern
Transvaal
Inflorescences flaccid; bulb scales truncate,
26. L. crispa
dead bulb scales purplish-brown; ovary
shoulders raised; eastern Cape
4.
Inflorescences erect
5.
Inflorescences flaccid
5.
25. L. concolor
15.
Ovary 3-lobed
6.
Ovary 6-lobed
8.
LEDEBOURlA IN SOUTH AFRICA
74
TAXONOMY
6.
Bulb scales truncate, with threads when tom;
leaf solitary, margin discolorous; rachis
ridged; base of peduncle compressed;
eastern Transvaal
Bulb scales attenuate, without threads when
tom; leaves more than 1, margin concolorous;
rachis smooth; base of peduncle terete
7.
Upper leaf surface smooth; leaves without
threads when tom, margin smooth; bulb
ovoid, bulblets absent; Cape
Upper leaf surface hairy; leaves with
threads when tom, margin ciliate; bulb
obovoid, bulblets present; eastern
Transvaal
18. L. monophylla
7.
28. L. ovalifolia
20. L. parvifolia
8.
Petiole present
Petiole absent
9.
Bulb scales with threads when tom
10.
Bulb scales without threads when tom
12.
10.
33. L. papillata
9.
Upper leaf surface smooth; leaves with threads
when tom, margin discolorous, texture
leathery; seed brown; widespread
Upper leaf surface hairy; leaves without
threads when tom, margin concolorous,
texture fleshy; seed black; northwestern Transvaal
11.
11.
8. L. lepida
Rachis smooth; bracts fleshy; bulb scales
attenuate; apex of ovary with rounded
shoulders; eastern Cape
Rachis ridged; bracts membranous; bulb scales
11. L. coriacea
truncate; ovary shoulders absent;
Transvaal and northern Cape
LEDEBOURlA IN SOtrrH AFRICA
22. 1. glauca
7S
TAXONOMY
12.
Rachis smooth; inflorescences longer than
leaves; tepals not cucullate
13.
Rachis ridged; inflorescences shorter than
leaves; tepals cucullate
13.
14.
Upper leaf surface viscid; live bulb scales
tightly arranged; bracts membranous; ovary
shoulders absent; north-western Transvaal
6. L. viscosa
Upper leaf surface not viscid; live bulb scales
loose; bracts fleshy; ovary shoulders
rounded; widespread
14.
3. L. sandersonii
Leaves fully developed at anthesis, with threads
when torn; bulb scales attenuate; live
bulb scales tightly arranged; bracts
membranous; widespread
12. L. floribunda
Leaves absent at anthesis, without threads
when torn; bulb scales truncate; live
bulb scales loosely arranged; bracts fleshy;
Cape
15.
27. L. undulata
Petiole present; eastern Transvaal
16.
Petiole absent; not restricted to eastern
Transvaal
16.
17.
Rachis smooth; upper leaf surface hairy,
without threads when torn; seed brown
Rachis ridged; upper leaf surface smooth,
10. L. rupestris
with threads when torn; seed
yellowish-brown
17.
29. L. petiolata
Ovary 3-lobed
18.
Ovary 6-lobed
19.
LED£BOURlA IN SOlITH AFRICA
76
TAXONOMY
18.
Bulb scales truncate, with threads when torn;
upper leaf surface with lacunae; rachis
ridged; ovary shoulders absent; Kaapsche
Hoop area
32. L. galpinii
Bulb scales attenuate, without threads when torn;
upper leaf surface smooth; rachis smooth;
ovary shoulders rounded; Cape
19.
20.
21.
28. L. ovali/olia
Bulb scales with threads when torn
20.
Bulb scales without threads when torn
28.
Leaves spirally twisted
21.
Leaves straight
22.
Ovary shoulders present; bulb obovoid; leaves
leathery, difficult to tear, margin
discolorous; widespread
24. L. marginata
Ovary shoulders absent; bulb cylindrical; leaves
fleshy, easily torn, margin concolorous;
north-western Transvaal
22.
5. L. atro-brunnea
Leaves hairy, margin ciliate; bracts fleshy;
tepals lanceolate; eastern Cape
13. L. hypoxidioides
Leaves glabrous, margin smooth; bracts
membranous; tepals oblong; widespread
23.
24.
23.
Leaves partly emerged at anthesis
24.
Leaves fully developed at anthesis
26.
Bulb scales truncate, live bulb scales loosely
arranged; leaves glossy, lower surface
monochromatic, base canaliculate
Bulb scales attenuate; live bulb scales tightly
17. L. ovatifolia
arranged; leaves lustreless, lower surface
dichromatic, base flat
LEDEBOURIA IN SOlITH AFRICA
25.
77
TAXONOMY
25.
Ovary shoulders present; bulb ovoid, 30 - 35 mm
diameter; bulb scales with prominent purple
blotches; leaf margin discolorous; seed
brown
23. L. inquinata
Ovary shoulders absent; bulb obovoid, 100 - 150 mm
diameter; bulb scales without purple blotches;
26.
leaf margin concolorous; seed black
15. L. zebrina
Seed black; tepals not cucullate; pedicels
green; perianth predominantly green
15. L. zebrina
Seed brown; tepals cucullate; pedicel pink;
perianth pink to purple
27.
27.
Bulb ovoid; leaves fleshy, glossy, margin
concolorous, venation obscure
14. L. revoluta
Bulb obovoid; leaves leathery, lustreless, margin
discolorous, venation prominent
28.
29.
17. L. luteola
Leaves with threads when torn; rachis ridged
29.
Leaves without threads when torn; rachis smooth
31.
Leaves spirally twisted, partly emerged at
anthesis, margin concolorous; tepals
linear; seed black
7. L. leptophylla
Leaves straight, fully developed at anthesis,
margin discolorous; tepals oblong; seed brown
30.
30.
Ovary shoulders tapering into the style; bulb
ovoid; leaf margin smooth; venation
obscure
Ovary shoulders raised; bulb subglobose; leaf
margin papillate; venation prominent
31.
12. L. floribunda
21. L. asperifolia
Bracts fleshy
32.
Bracts membranous
36.
LEDEBOURlA IN SOlTfH AFRICA
78
TAXONOMY
32.
Ovary shoulders present, base of ovary lobes
smooth
33.
Ovary shoulders absent, base of ovary lobes
papillate
33.
35.
Ovary shoulders truncate; leaf apex obtuse,
venation prominent; perianth stellate
31. L. macowanii
Ovary shoulders tapering into the style; leaf
apex acute, venation obscure; perianth
recurved
34.
34.
Live bulb scales loosely arranged; dead bulb scales
brown; lower leaf surface monochromatic;
inflorescences longer than leaves
3. L. sandersonii
Live bulb scales tightly arranged; dead
bulb scales purplish-brown; lower leaf surface
dichromatic; inflorescences same length
1. L. aperliflora
as leaves
35.
Leaves linear to narrowly elliptic; perianth
stellate
Leaves lanceolate to oblong; perianth sharply
2. L. ensi/olia
reflexed
36.
9. L. minima
Ovary shoulders present; bulb obovoid, live
bulb scales tightly arranged, bulblets
absent; leaves elliptic
Ovary shoulders absent; bulb subglobose,
1. L. aperliflora
live bulb scales loose, bulblets present;
leaves lanceolate
LEDEBOURlA IN SOlITH AFRICA
19. L. cooperi
79
TAXONOMY
12.4. Species descriptions.
Section HUMERATI S. lenter
Section Humerati S. Venter, sect. nov., squamis efibrosis; ovarii lobis ad
apicem prominenter rotundatis.
Species typica: Ledebouria apenijlora (Bak.) Jessop.
Typus: Saund. Ref. Bot. 1: t.19., "Cape, Cooper s.n." (1868).
Species: L. apertiflora (Bak.) Jessop, L. ensijolia (Eckl.) S. Venter, L.
sandersonii (Bak.) S. Venter, L. dolomiticola S. Venter, L. atro-brunnea S.
Venter, L. viscosa Jessop, L. leptophylla (Bak.) S. Venter, L. lepida (N.E. Br.)
S. Venter, L. minima (Bak.) S. Venter and L. rupestris (Van der Merwe) S.
Venter.
Plants solitary. Bulbscales without threads when torn. Leaves erect to
spreading. Ovary with prominent shoulders.
Subsectio Acutilobae S. Venter, sub sect. nov., tepalorum apicibussigillatim
acutis.
Species typica: L. ensifolia (Eckl.) S. Venter.
Species: L. apertiflora (Bak.) Jessop, L. ensijolia (Eckl.) S. Venter, L.
sandersonii (Bak.) S. Venter.
Plant without threads when torn. Bracts fleshy. Tepal apices markedly acute.
1. LEDEBOURIA APERTIFLORA (Bak.) Jessop
Ledebouria apertiflora (Bak.) Jessop in 11 S. Afr. Bot. 36(4): 254 (1970).
Drimia apertijlora Bak. in Saund. Ref. Bot. 1: t.19 (1868).
Type: Saund. Ref. Bot. 1: t.19 (1968)!. (lecto. selected here - Art. 9.3, Greuter
et al. 1988).
LEDEBOURLA IN SOlTfH AFRICA
80
TAXONOMY
F
E
c
•
.
. :..'7;:
....
--
.
:;
B
G
Figure 16. Illustration of L. apenijlora (Bak.) Jessop. A, habit X 1. B, lamina
margin X 300. C, bract X 10. D, tepal with stamen X 10. E, apex of tepal X
20. F, section through lamina X 5. G, flower X 10. H, lateral view of ovary X
10 and I, dorsal view of ovary X 10. All from retn~
s.n.
UDE:JJOURlA IN SOUTH AFIlICA
81
TAXONOMY
Scilla lorata Bale. in Saund. Ref. Bot. 3(Append.): 14 (1870).
Iconotype: As for Drimia apenijlora.
Scilla linearijolia Bale. in Saund. Ref. Bot. 3: t.184 (1870).
Iconotype: Saund. Ref. Bot. 3: t.184. (1870)!.
Scilla apenijlora (Bale.) C.A. Sm. in Kew Bull. :250 (1930).
Type: As for Drimia apenijlora.
Plants solitary. Bulb hypogeal, 40 - 50 x 40 - 50 mm, + obovoid; dead
bulb scales purplish-brown, apices attenuate, live bulb scales fleshy, tightly
arranged, without threads when torn, white inside. Leaves fully developed at
anthesis, 5 - 10, spreading, lanceolate, 100 - 225 x 7 - 25 mm, without threads
when torn, fleshy, surfaces maculate to fasciate, green with darker green or
purple spots and blotches in the lower part, venation obscure; margins smooth
but sometimes undulate; leaf base canaliculate; apex acute. Inflorescences 2,
dense, cylindric, 50 - 100 x 15 - 20 mm, flaccid, 30 - 70-flowered, as long or
longer than the leaves; scape basally terete, green-spotted, dark green to purple
in lower part, glabrous; rachis smooth, scape smooth, 80 - 225 mm long.
Bracts semi - fleshy to membranous, 0.5 x 0.2 mm, deltoid to linear, green,
bracteole sometimes absent. Pedicels spreading, 6 - 12 mm long, pink or green.
Perianth 5 - 6 mm long, tepals recurved, equal, oblong, 6 x 1.5 mm, apex
acute, cucullate, pink to purple with a green keel. Stamens erect, 5.0 - 5.5 mm
long, filaments maroon to pink, epitepalous; anthers 1 mm long, yellow. Ovary
ovoid, 6 -lobed, 1 x 1.2 mm, lobes obtusely deltate, glabrous, apex tapering
into the style. Style 1.5 mm, terete, glabrous, white to purple; stigma below the
anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, clavate; base
truncate. Seed drop-shaped, ± 3 mm long, strongly wrinkled, brown.
Chromosome number 2n = 26. (Figure 16).
L. apenijlora differs from L. cooperi in the absence of bulblets and bracteoles;
acute tepal apices (Figure 16E), style 1.5 mm long and ovary 1.2 mm wide. L.
revoluta differs from L. apertiflora in having threads in the leaves and
bulb scales when torn, its ridged rachis, wider tepals (+ 3 mm) with obtuse
apices, longer style (6 mm) and ellipsoidal ovary.
LEDEBOURIA IN SOllTH AFRICA
82
TAXONOMY
Specific epithet etymology.
Refers to the open flowers.
Variation.
A variable species without fixed bulbshape which is influenced by the soil and
rainfall. Plants of L. apenijlora can be placed into two groups according to the
lengthened, closely packed or open and spreading, hardened dead bulb scale
apices (Map 5). In map 5 the area indicated by black dots shows the distribution
of plants with the truncate bulb scales. The area indicated by black squares
shows the recorded distribution of plants with dead bulb scales much lengthened
and hardened. After the leaves have died back, a rosette of dead bulb scales
protrudes up to 20 mm above the soil surface.
Leaf shape is polymorphic. Linear to linear-lanceolate leaved plants were placed
in Scilla linearifolia (Baker 1870a). The colour of the leaves varies from light
green (Jozini, northern Natal) to glaucous green (Messina, northern Transvaal)
and may have or lack markings. Populations in the Lebombo Mountains of
northern Natal and Swaziland tend to have strongly undulate leaf margins.
Flower colour is also variable but predominantly pink to purple with a green
keel. Populations in northern Swaziland, eastern and northern Transvaal tend
towards green with only traces of pink at the base of the tepals. Whitish-green
flowers are recorded from the Pongola Area (Nel103).
Jessop (1970) failed to define this species accurately resulting in specimens of L.
cooperi (Hook.f.) Jessop and L. revoluta (L.f.) Jessop being placed in L.
apenijlora.
Flowering period
From October to December.
UDEBOURlA IN SOlITH AFRICA
Map 5.
12°
18 ~o
Pictorialized map of L. apertiflora with the known distribution of the tapered and truncate bulbscaled plants.
14°
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84
TAXONOMY
Figure 17. A, lectotype of L. apenijlora (Bak.) Jessop under Drimia apenijlora
Bak. in Saunders Refugium Botanicum 1 : t. 19 (1868); B, SEM micrograph of
the adaxial lamina surface. C, SEM micrograph of the abaxial lamina surface.
D, habitat near Phalaborwa, north-eastern Transvaal. The vegetation consists of
closed deciduous tall Combretum apiculatum - C. imberbe - Acacia nigrescens
woodland; E, SEM micrograph of stomata and F, SEM micrograph of the fine
s. n.
texture of the testa. B - C and E - F from retn~
LEDEBOURlA IN SOUTH AFRICA
85
TAXONOMY
LEDEBOURlA IN SOUTH AFRICA
86
TAXONOMY
Distribution (Map 5).
Southern Cape to Zimbabwe border excluding Transkei and Lesotho. Suitable
habitats exist in Transkei and the species may have been overlooked. L.
apertiflora is not a plant of montane grassland and is unlikely to occur in
Lesotho. It is restricted to the drier woodland areas with sandy soil.
Habitat
L. apenijlora is recorded from areas where the underlying rock strata is either
granite or gneiss (Transvaal), limestone (Cape) or rhyolite from the Jozini
Formation, Lebombo Group and feldspathic sandstone from the Mlazi
Formation, Natal Group (SACS 1980).
Soils are mostly sandy to a rocky sandy loam or sometimes clay loam. Some of
the Orange Free State populations occur on brackish clay loam near Fouresmith.
On the floodplains below the Lebombo Mountains, some plants were found at
Mkuze, growing in yellow-brown clay.
In the Transvaal, L. apertiflora is associated with woodland (Figure 17D)
whereas in the Cape, it is mostly associated with shrubland.
Dense populations of L. apertijlora rarely occur. The only dense populations
encountered during the survey are on the foothills of the Blouberg in northern
Transvaal. Here the plants grow as scattered individuals. In some areas plants
are heavily grazed.
Historical background
Baker, in April 1868, described L. apertiflora as Drimia apertiflora from a
living plant that flowered in 1868 with Mr. Wilson Saunders at Reigate. Two
years later, Baker placed D. apenijlora into the genus Scilla under the name S.
[orata, discarding the earlier specific epithet, stating "Placing this in Scilla, the
specific name first given is appropriate no longer" (Baker 1870d; Smith 1930).
LEDEBOURIA IN SOIITH AfRICA
87
TAXONOMY
Specimens examined
VENDA. - 2230 (Messina): Thate Vondo (-CD),
retn~
12,242 (UNIN).
TRANSVAAL. - 2229 (Waterpoort): Messina, farm Fontainbleau,
Venter 12,824 (UNIN); Mopane (-DB), Strey 3475 (PRE). 2230 (Messina):
Messina (-AC), Moss & Rogers 92 and 208 (J). 2327 (Ellisras): Villa Nora (DB), Smook 4228 (PRE). 2328 (Baltimore): Villa Nora, farm Wellust (-CA),
Schmidt 95 (PRU). 2329 (Pietersburg): Louis Trichardt (-BB), Martley s.n. sub
BOL 22,502 (BOL). 2330 (Tzaneen): Giyani, Middle Letaba Dam (-AD),
Venter 13,065 (UNIN). 2427 (Thabazimbi): Thabazimbi, Krantzberg (-BC),
Wzn der Merwe 2024 (PRE). 2428 (Nylstroom): Warmbaths (-CC), Leendertz
6651 (PRE). 2429 (Zebediela): Lulu Mountains, farm Parys (-DB), Barnard &
Mogg 703 (PRE). 2430 (Pilgrim's Rest): Klaserie, Venter 12,686 and 12,708
(UNIN). 2431 (Acornhoek): Timbavati Game Reserve (-CB), Zimbatis 1515
(PRE); Manyeleti Game Reserve, (-DA), Bredenkamp 1187 (PRE). 2531
(Komatipoort): Kruger National Park, Skipberg (-AB), Van der SchYff 3214
(NPB); Malelane (-BC), Lang s.n. sub PRE 30,410 (PRE); Komatipoort (-BD),
Moss & Rogers 540 (J), Rogers 22,210 (1); Kruger National Park, Randspruit,
Wzn "yk 4803 (NPB). 2626 (Klerksdorp): Klerksdorp, farm Drooge Spruit (-
DD), Ubbink 706 (PUC). 2630 (Carolina): Carolina (-AA), Moss & Rogers
1375 (1). 2725 (Bloemhot): Christiana, Kameelpan (-CC), Theron 5477 (PRE).
ORANGE FREE STATE. - 2925 (Jagersfontein): Fauresmith (-CB),
Verdoorn 883 (PRE). 2926 (Bloemfontein): Bloemfontein (-AA), Smith 8751B
(PRE); Griqualand West, Mostertshoek (-CA), Acocks H993 (PRE).
SWAZILAND. - 2631 (Mbabane): Stegi (-BD), Compton 31,216
(NBG), Palata, Compton 31,216A (PRE); Bulunga Poort (-DA), Compton
32,375 (NBG), Karsten s.n. sub PRE 703,802 (PRE). 2632 (Bela Vista):
Nkumbane Valley (-AA), Culverwell 253 (PRE); Abercorn Drift (-CC), Moll &
Pooley 4182 (NH). 2732 (Ubombu): Ingwavuma Poort (-AA), Compton 29,817
(NBG).
LED£BOURIA IN SOlITH AFRICA
TAXONOMY
88
NATAL. - 2731 (Louwsburg): Pongola Plaas (-BC), Nel 103 (PRE);
Jozini Dam (-BD), Strey 5346 (NH), Strey & Moll 3649 (PRE), Scott-Smith 105
(NU). 2732 (Ubombu): Ingwavuma Poort (-AA), lUn der Merwe 2732 (PRE),
Compton 29,460 (PRE); Ndumu Game Reserve (-AB), Pooley 669 (NU);
Mkuze Game Reserve (-CA), Goodman 793 (NH). 2831 (Nkandla): Hlabisa (BB), Harrison 231 (NH); Mtunzini, Umhlatuzi Flats (-DC), Venter 2564
(BLFU) , Umhlatuzi Bridge, Lawn 1862 (NH). 2930 (pietermaritzburg): Inanda
(-DB), WJod 1208 (NBG).
CAPE. - 2821 (Upington): Upington, The Halt (-AC), Glover s.n. sub
BOL 22,500 (BOL). - 2823 (Griekwastad): Campbell (-DC), Power s.n. sub
GRA A7422 (GRA). 2824 (Kimberley): Warrenton (-BB), Adams 212 (BOL);
Barkley West (-DA), Acocks 1466 (PRE); Kimberley (-DB), Oliver 127 (NBG),
Wilman s.n. sub BOL 13,939 (BOL), Kimberley; Platrand, Brueckner 920
(PRE). 2919 (Pofadder): Pella (-AA), Pearson 3596 (BOL). 2922 (Prieska):
Prieska (-DA), Bryant J173 (STE), Moss 10,857 (1), Acocks 2548 (PRE). 3020
(Brandvlei): Brandvlei (-BC), Frandsen s.n. sub NBG 142,872 (NBG). 3024
(De Aar): De Aar (-CA), Moss 10,851 (J), Purcell s.n. sub BOL 50,892 (BOL).
3221 (Merweville): Beaufort West, Layton (-BB), Shearing 1142 and 1174
(PRE). 3224 (Graaff Reinet): Karoo National Park (-AB), Palmer 1104 (PRE).
3225 (Somerset East): Cradock (-BA), Zietsman 1292 (PRE); Mortimer (-BC),
Kersit 9306 (BOL); Somerset East (-DA), Barker 9178 (NBG). 3226 (Fort
Beaufort): Glenthorne (-AC), Acocks 16,273 (PRE). 3227 (Stutterheim):
Keiskamma Hoek, Red Hill (-CA), Steyner 59 (GRA), Hogsback, Grant 3006
(PRE); Kubusie Valley (-DD), Wehmeyer 9 (PRE). 3318 (Cape Town):
Riebeeck West (-BD), 'WIn der Merwe 1745 (PRE).
UDEBOURJA IN SOlITH AFRICA
89
TAXONOMY
2. LEDEBOURIA ENSIFOLIA (Eckl.) S. Venter
Ledebouria ensifolia (Eckl.) S. \enter, comb. novo
Type: As for Drimia ensifolia.
Drimia ensifolia Eckl. in S. Afr. Quart. J. 1: 364 (1830).
Type: CAPE - District Ditenhage, Zwartkops River, Zeyher 10 (K!, lecto.
selected here; PREI, photo.).
Scilla ensifolia (Eckl.) Britten in Journ. Bot. 46: 201 (1908).
Type: As for D. ensifolia.
Drimia ludwigii Miq. in Bull. Sc. Phys. Need. :39 (1839).
Type: CAPE - Cap RSpei, Ecklon & Zeyher 1064 (D, halo.; GRAI; PRE!).
Idothea (?) ludwigii Kunth in Enum. Pl. 4: 681 (1843).
Type: As for Drimia ludwigii.
Scilla prasina Bak. in Saund. Ref. Bot. 3(App.): 10 (1870).
Type: CAPE - Kaffirland, Gill s.n. (Kl, halo.; PREI, photo.).
Scilla ludwigii Bak. in Saund. Ref. Bot. 3(App.): 9 (1870).
Type: CAPE - Cap R Spei, Zeyher 4262 (Kl, halo.; PREI, iso.).
Scilla pusilla Bak. in J. Bot., Land. 5: 183 (1876).
Type: Transkei - Bazeia, Bauer 293 (Kl, halo.; BOLl, drawing; PRE1, photo.).
Scilla ecklonii Bak. in Bot. Jahrb. 15(35): 7 (1892).
Type: Cape, Tambukiland, mountains between Silo and Windvogelberg, Ecklon
& Zeyher No. 12 (Bl, halo.).
LEDEBOURIA IN SOlJfH AFRICA
90
TAXONOMY
Plants solitary. Bulbs hypogeal, 40 - 60 x 20 - 30 mm, cylindrical; dead
bulb scales hard, dark brown to purplish-brown, apices attenuate, live bulb scales
membranous, tightly arranged, without threads when tom, white inside. Leaves
fully developed at anthesis, 5 - 10, spreading, narrowly ovate to ensiform, 80 150 x 15 - 40 mm, without threads when torn, fleshy, surfaces dull glaucous
green, adaxial surface without markings or with darker green or purple
blotches, abaxial surface without markings, venation obscure; margin smooth;
leafbase canaliculate; apex obtuse to acute. Inflorescences 1 - 3, dense,
cylindric, 50 - 120 x 10 - 20 mm, flaccid, 30 - 100 -flowered, longer than the
leaves; scape terete at base, green to brownish-green, glabrous; rachis smooth,
scape smooth, 50 - 200 mm long. Bracts fleshy, 1 x 1 mm, deltoid, green,
without bracteoles. Pedicels spreading, 3 - 4 mm long, green. Perianth 3 mm
long, tepals sharply reflexed, subequal, oblong, 3 x 1.5 mm, apex acute, green
to pink with a dull green keel. Stamens erect, 3 mm long, filaments pink,
epitepalous; anthers 1 mm long, yellow. Ovary ellipsoidal, 6 -lobed, 1.0 x 2.5
mm, lobes narrowly transversely oblong, apex shoulders rectangular, base of
lobes papillate. Style 1.5 mm long, triangular, glabrous, purple; stigma equal
height to anthers; stipe 0.5 x 1.0 mm. Capsule 3 - lobed, symmetrical, globose;
base truncate. Seed drop-shaped, 3 - 5 mm long, surface strongly wrinkled,
brown. Chromosome number 2n = 30. (Figure 18).
L. ensifolia is closely related to L. apertiflora sharing fusiform roots, dark dry
bulb scales, acute tepals and the apex of the ovary with prominent shoulders. L.
ensifolia differs from L. apertifolia in the cylindrical bulbs, ensiform leaves and
inflorescence which are longer than the leaves.
Specific epithet etymology.
Refers to the sword-shaped leaves.
Flowering time
From November to February.
LED£BOURIA IN SOlITH AFRICA
91
TAXONOMY
·G
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J
Figure 18. lllustration of L. ensijolia (Eckl.) S. Venter. A, habit Xl; B, lamina
margin X 300; C, lamina apex X 10; D, bract X 10; E, section through lamina
X 5; F, tepa! apex X 20; G, tepa! with stamen X 10; H, flower X 10; I, ovary
dorsal view X 10; J, ovary lateral view X 10. All from Smith 155.
LEDEBOURlA IN
sourn AFRICA
Map 6.
12°
Known distribution of L. ensifolia (Eckl.) S. Venter
14°
16°
18°
20°
22°
24°
26°
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30°
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G.P.·S. 13--0959
~
93
TAXONOMY
Figure 19. A, lectotype of L. ensifolia (K); B, habitat at the Zwartkops River
estuary. The vegetation consists of tall, closed, evergreen shrubland; C, SEM
micrograph of the adaxial lamina surface. Bar = 50 J.'m; D, SEM micrograph
of the abaxial lamina surface. Bar = 50 J.'m; E, SEM micrograph of stomata.
Bar = 50 J.'m; F, inflorescence. Bar = 10 mm. C - F from l€nter 13,278.
LEDEBOURIA IN SOurH AFRICA
94
TAXONOMY
LEDEBOURJA IN SOlJfH AFRICA
9S
TAXONOMY
Distribution (Map 6).
Eastern Cape with a few scattered localities in the western Cape, northern Cape
and Transkei.
Habitat
Most populations occur on quartzites and sandstones of the Dirkskraal Sandstone
Formation of the Witteberg Group. These are commonly known as the
Witteberg Quartzites.
L. ensijolia occurs commonly on sandy soils derived
from quartzites and sandstone. A few populations were found growing in loamy
to clay-loam soils with one population growing in clay soil.
L. ensijolia occurs most commonly in woodland and shrubveld (Figure 19B).
Plants grow in full sun or light shade.
Variation
Leaves of plants growing near the coast tend to be thinner (probably due to the
higher rainfall) than those of plants growing in drier inland habitats. A few
populations with pure green flowers were found in the vicinity of Grahamstown
("\.enter 13,256). Flowers with a slight sweet scent were recorded in some of the
Transkei populations.
Historical background
Although Ecklon named this species Drimia ensijolia, he noted that together D.
nitida and = D. lanceaefolia Ker. (L. revoluta) differ markedly from most
Drimia species and perhaps represent a distinct genus (Ecklon 1830).
In publishing Idothea ludwigii, Kunth placed a question mark in front of the
species name, indicating his uncertainty about the specific name (Kunth 1843).
LEDEBOURJA IN SOllfH AFRICA
96
TAXONOMY
Baker described Scilla ludwigii from material collected by Zeyher (Zeyher
4262). He collected these specimens at the Zwartkops River Valley and
adjoining hills from Villa Paul Mare to Uitenhage on one of his regular visits to
the area. The description of S. ludwigii and Drimia ludwigii are very similar and
Baker did not mention his reasons for describing S. ludwigii.
Baker (1876) expressed the view that Scilla pusilla is closely allied to Scilla
prasina Bak. A specimen labelled as Bauer 293 sub BOL 23281 is in fact a
collection of Ledebouria sandersonii (Bak.) S. Venter. As the species are very
different, it is assumed that in handling the specimens, labels were misplaced.
The holotype leaves no doubt that S. pusilla is a synonym of L. ensifolia.
As Drimia ensijolia was placed in synonymy under Scilla ludwigii, a name
published 50 years after D. ensijolia , Britten (1908) made a new combination
Scilla ensijolia (Eckl.) Britten, using the oldest description.
Specimens examined
TRANSKEI. - 3128 (Umtata): Umtata Waterfall (-DB), Schonland 3774
(GRA). 3129 (Port St. Johns): Annesville, Sipetu (-AA), Strever 164 (NH).
3028 (Matatiele): Mount Frere (-DD), W!nter 13,325 (UNIN).
CAPE. - 3421 (Riversdale): Gouritz River Mouth (-BD), Schlechter 18
(BOL). 3423 (Knysna): Plettenberg Bay (-AB), Smart s.n. (BOL). 3424
(Humansdorp): Humansdorp (-BB), Jeppe 4946 (BOL). 3318 (Cape Town):
Grey Reservoir (-DC), Schonland 146 (GRA). 3323 (Willowmore):
Willowmore (-AD), Barker 5013 (NBG). 3324 (Steytlerville): Campher's Poort
(-AA), Barker 5007 (NBG); Pienaarspoort (-AD), Barker 5070 (NBG);
Uitenhage, Kleinpoort (-BD), Barker 5071 (NBG); Hankey, along Klein River
(-DD), Venter 13,278 (UNIN). 3325 (Port Elizabeth): Zuurberg (-AD), Repton
5887 (PRE); Kommadagga, Water Tower Hill (-BB), Perry 1626 (NBG);
Sheldon, Hutton 493a (BOL); Humansdorp (-CC), Barker 6900 (NBG); Uitenhage, Zwartkops River Mouth (-CD), Zeyher 4262 (BOL); Ecklon & Zeyher
1064 (BOL); Alexandria, Congo's Kraal (-DB), Archibald 4104 (PRE); Port
LEDEBOURIA IN SOlITH AFRICA
97
TAXONOMY
Elizabeth, Komachs, Paterson 2392 (BOL); Bluewater Bay, Unon 889 (PEU);
Redhouse, Paterson 387 (BOL). 3326 (Grahamstown): Grahamstown, Woest
Hill (-AC), Venter 13,254 (UNIN); Grahamstown , Venter 13,256 (UNIN);
Hepbum s.n. (BOL); Queenstown, Page s.n. (BOL); Jacot Guillarmod 10,089
(GRA); Mayor's Seat, Rogers s.n. (GRA); Sutherland's Farm, Deacon 3
(GRA); Commitee Drift (-BB), liln der Merwe 1879 (PRE); Albany, Manley
Flats (-BC), Rogers 28366 (GRA). 3327 (Peddie): Peddie (-AA), Herre s.n.
(BOL); Leighton 2648 (BOL); East London, Kidd's Beach (-BA), Hall 235
(NBG). 3218 (Clanwilliam): Middelburg, Grootfontein (-BA), Gi1l21 (PRE).
3224 (Graaff Reinett): SW of Aberdeen (-AC) , Barker 7120 (NBG); Graaff
Reinett, Lettskraal Station (-BB), Acocks 16,235 (PRE). 3225 (Somerset East):
Cradock (-BA), Barker 7064 (NBG); Mortimer (-BC), Davison 25 (BOL);
Somerset East, Boschberg (-DC), Venter 13,286 (UNIN); MacOwan s.n.
(GRA); -3226 (Fort Beaufort): Alice (-DD), Venter 13,319 (UNIN). 3227
(Stutterheim): Stutterheim Townlands (-CB), Venter 13,418 (UNIN); King
Williamstown (-CD), Landrey sub PRE 35,336 (PRE); East London, Fort
Jackson (-DC), Atkins 3121 (PRE). 3124 (Hanover): Dwaal Station (-BA),
Acocks 16,315 (PRE); Middelburg, Barend's Kraal (-DB), Thome s.n. (BOL).
3125 (Steynsburg): Middelburg, Farm Bangor (-AC) , Bolus 14,110 (BOL);
Albany, Fish River (-CD), liln der Merwe 1895 (PRE).
Without precise locality
- Cap B. Spei, Ecklon & Zeyher 1064 (GRA, PRE, U); Ecklon & Zeyher 4262
(K, PRE); Kaffirland, Gill s.n. (K, PRE).
LEDEBOURIA IN SOurH AFRICA
98
TAXONOMY
3. LEDEBOURIA SANDERSONII (Bak.) S. Venter
Ledebouria sandersonii (Bak.) S. lenter, cqmb.nov.
Scilla sandersonii Bak. in Saund. Ref. Bot. 3 (App.): 5 (1870).
Type: Transvaal, without precise locality, Sanderson s. n. (K!, holo.; PRE!,
photo.).
Scilla baurii Bale. in Flora Cap. 6: 484 (1896).
~
Type: Cape, Tembuland, Bazeia Mountain, Bauer 550 (K!, holo.; PRE!,
photo.; SAM1).
-f
Scilla tysonii Bak. in Flora Cap. 6: 484 (1896).
Type: Cape, Griqualand East, Tyson s.n. (Kl, holo.; BOLl & GRA! drawing).
Scilla oostachys Bak. in Flora Cap. 6: 487 (1896).
~
Type: Natal, Upper Umkomaas, Wood 4627 (Kl, holo.; NH!; PRE1, photo.;
BOLl, drawing).
\
Scilla diphylla Bak. in Flora Cap. 6: 489 (1896).
'" Type: Transvaal, Barberton, Saddleback Range, Galpin 1182 (Kl, holo.; BOLl;
~
GRA!; NH!; PRE1; SAM1).
Scilla bella Markotter in Ann. Univ. Stell. 8 (Sec. A No. 1.): 13 (1930).
Type: Natal, Oliviers Hoek Pass, Thode s.n. in STE 3372 (STE1, holo.; PRE1,
photo.).
Plants solitary. Bulb hypogeal, 10 - 30 x 10 - 15 mm, ovoid to subglobose;
dead bulb scales brown, membranous, apices attenuate, live bulb scales loosely
arranged, without threads when torn, white inside. Leaves fully developed at
anthesis, 1 - 6, spreading or sometimes appressed, ovate to lanceolate, 15 - 75 x
18 - 30 mm, without threads when torn, fleshy, adaxial surface green sometimes
with dark purple blotches, abaxial surface green suffused purple to wholly
purple, venation obscure; margin smooth; leaf base canaliculate; apex acute.
UDEBOURJA IN SmITH AFRICA
99
TAXONOMY
Inflorescences 1 - 2, dense, oblong, 20 - 25 x 18 - 20 mm, erect to flaccid, 12 20 -flowered, longer than the leaves; scape terete at base, green, glabrous;
rachis smooth, scape smooth. Bracts fleshy, 1 x 0.5 mm, dentate, pink to
purple without bracteoles. Pedicels spreading, 6 - 8 mm long, pink. Perianth 3
mm long, tepals recurved, equal, oblong, 3 x 1 mm, apex acute, pink
sometimes with a green keel. Stamens erect, 2.75 mm long, filament with
upper part purple and lower part white, epitepalous; anthers 0.75 mm long, pale
yellow. Ovary ovoid, 6 -lobed, 1 x 2 mm, lobes narrowly transversely oblong,
apex tapering into style, base of lobes with papillate nectaries. Style 3 mm long,
triangular, glabrous, purple; stigma equal height to anthers; stipe 0.5 x 0.5
mm . Capsule three-lobed, symmetrical, globose; base truncate. Seed dropshaped, 2.5 mm long, surface strongly wrinkled, brown. (Figure 20).
L. sandersonii is related to L. apertiflora and L. ensijolia. Together they form
the subsection Acutilobae. L. sandersonii differs from the abovementioned
species in the thin fleshy roots, loosely arranged live bulb scales, ovate to
lanceolate leaves and dense oblong raceme.
Specific epithet etymology.
Commemorates the journalist, trader and draughtsman John Sanderson (Gunn &
Codd 1981).
Flowering period
From August to March with peak flowering from October to November.
Distribution (Map 7).
Distributed in the eastern part of South Africa. The species is most abundant in
Natal.
LEDEBOURIA IN SOlITH AFRICA
100
TAXONOMY
o
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K
J
B
H
Figure 20. Illustration of L. sandersonii (Bak.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, section through lamina X 2; D, bract X 10; E, tepa!
with stamen X 10; F, flower X 10; G, section through lamina depicted in H, X
2; H, lamina, shade form Xl; I, tepal apex X 20; J, ovary lateral view X 10;
K, ovary dorsal view X 10. A - F from Crouch 7 and H to K from Venter
13,464.
LEDEBOURlA IN SOlITH AFRICA
Known distribution Of L. sandersonii (Bak.) S. Venter
Map 7.
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G.P.-$. 13-0959
102
TAXONOMY
Figure 21. A, type of L. sandersonii (Bak.) S. Venter (K); B, plants of the
spotted-leaf form, near Dullstroom, eastern Transvaal. Bar = 25 mm; C, SEM
micrograph of the adaxial lamina surface. Bar = 50 J..'m; D, SEM micrograph
of the abaxial lamina surface. Bar = 50 J..'m; E, SEM micrograph of stomata.
Bar = 50 J..'m; F, habitat in the basalts at Cathedral Peak. The vegetation
consists of short montane grassland. C - E from Crouch 7.
LEDEBOURlA IN SOIJrH AFRICA
103
TAXONOMY
LEDEBOURJA IN SOlJfH AFRICA.
104
TAXONOMY
Habitat
L. sandersonii occurs in humusrich, shallow (5 - 60 mm deep) to deep (30 - 260
mm), medium grained (0.25 - 1 mm <!» to coarse grained (1 - 3 mm <!», well
drained grey to greyish-brown sandy soil. Most of the plants occur in shallow
soil overlaying the rock sheets. The few populations on the Natal Drakensberg
escarp are all on Drakensberg Basalt of the Drakensberg Group (Figure 21F).
Soils derived from the basalt are blackish, shallow (10 - 160 mm deep), coarse
grained (1 - 3mm <!» and medium drained (Van der Merwe 1962). These soils
are subject to freezing during the winter months.
L. sandersonii is associated with montane grassland. Along the Transvaal
Drakensberg escarp and Steenkampsberg, L. sandersonii commonly grows on
the edges of shallow seepages, sometimes in soil sodden for months.
Variation
Plants growing in seepages have loosely arranged live bulb scales compared with
the thinner and the relatively tightly arranged live bulb scales of plants in drier
habitats. L. sandersonii has two distinct forms, an immaculate form and a
mottled form (Figure 21B). Mixed populations of these forms have not been
recorded. Tepal colour varies from dark pinkish purple to light pink with a
prominent green keel.
Specimens examined
TRANSVAAL - 2329 (Pietersburg): Haenertsburg (-DD), Pot! 4725
(PRE). -2430 (Pilgrim's Rest): Vaalhoek (-DB), Rogers 25,065 (1); Mariepskop
(-DB), Raal 1676 (Transvaal Provincial Administration); Venter 12,771
(UNIN); Morgenzon Nature Reserve (-DC), Kluge 2070 (PRE); Mount Sheba
(-DC), Ker/oot 8168 (PRE); Graskop (-DD), Thorncro/t 18,299 (PRE);
Graskop, Stanley Bush (-DD), Raal1038 (Transvaal Provincial Administration);
Mac Mac (-DD), ltln der Merwe 1594 (PRE); Sabie (-DD), Koeleman S.n. sub
LEDEBOURIA IN SOUTH AFRICA
105
TAXONOMY
PRE 36,634 (PRE). -2530 (Lydenburg): Draaikraal (-AA), Hurling & Nel s.n.
sub BOL 22492 (BOL); Steenkampsberg (-AC), Bloem 128 (PRE); Pole-Evans
2031 (PRE); Dullstroom (-AC), Drews 18 (PRE); Killick & Strey 2552 (PRE);
Dullstroom, Suikerboskop (-AC), Burgoyne 372 (PRU); Verloren Vallei Nature
Reserve (-AC), Bloem 128 (PRU); Hartebeesvlakte (-BA), Mohle 34 (PRE);
Witklip Plantation (-BD), Kluge 278 (PRU); Kluge 105 (PRE); Dullstroom,
Lakenvlei Spruit (-CA), McMurtry 5204 (Johannesburg Botanical Garden);
Belfast (-CA), liln der Merwe 1228 (PRE). -2531 (Komatipoort): Saddleback
Range (-CC), Van der Merwe 1823 (PRE). -2730 (Vryheid): Wakkerstroom,
Hlangapies (-AB), Van der Merwe 2075 (PRE); Kwa-Madlangampisi, farm
Groothoek (-BA), du Toit 95 (PRE).
SWAZILAND -2631 (Mbabane): Forbes Reef (-AA), Compton 30,122
(PRE); Compton 31,727 (NBG, PRE); Compton 26,130 (NBG); Compton
27,803 (NBG, PRE); Mbabane (-AC), Compton 30,781 (NBG); Moss 25,942
(1); Rogers 11,589 (BOL);Mbabane, Abner's Farm (-AC), Compton 26,106
(NBG); Stroma (-AC), Compton 25,196 (NBG). -2632 (Bela Vista): Ndumu (CD), Compton 30,338 (NBG).
ORANGE FREE STATE -2827 (Senekal): Vrede (-DB), O'Meara s.n.
(BOL, NBG). -2829 (Harrismith): Harrismith, Platberg (-AC), Jacobsl 2615
(NBG).
NATAL -2730 (Vryheid): Altemooi (-AD), Thode 3368 (STE); Tendeka
Rock (-DB), Gerstner 5097 (PRE). -2731 (Louwsburg): Ngome (-CD), Strey
9362 (NH, PRE); Ngome State Forest (-CD), Nicholas & liln der Berg 1764
(NH, Killick Herbarium). -2828 (Bethlehem): Bester's Vlei (-DB), Bolus s.n.
sub BOL 22,496 (BOL); Royal National Park (-DB), Van der Merwe 2613
(NU). -2829 (Harrismith): Van Reenen, Nolens Volens (-AD), Jacobsl 1660
(PRE); Oliviers Hoek Pass (-CA), Wilson 1086 (PRE); Thode s.n. sub STE
3372 (STE); Cathedral Peak (-CC), Killick 1607 (PRE, Killick Herbarium);
Killick 1570 (PRE); Killick 1114 (PRE); Fongosi River (-DC), Fisher 265
(NU). -2830 (Dundee): Babanango (-BD), liln der Merwe 2786 (PRE); Griffins
Hill (-CC), Acocks 10,734 (NH). -2929 (Underberg): Cathkin Park (-AB),
Henrici 3526 (PRE); Tabamhlope (-BA), Dowling 222 (NU); Ntabamhlope
LEDEBOURlA IN SOlITH AFRICA
106
TAXONOMY
Peak (-BA), lbn der Merwe 2564 (PRE); Mpendhle, Mulangane (-BC), Billiard
& Bunt 18,611 (NU); Estcourt, farm Lanner Veane (-BC), Manning & Balkwill
338 (NU); Kamberg (-BD), Gordon-Grey 99 (NU); Cobham Nature Reserve
(-CB), Billiard & Burtt 9316 (NU); Underberg, Chameleon Cave (-CB),
Billiard & Bunt 17,740 (NU, PRE); Bamboo Mountain (-CB), Billiard & Bunt
15,632 (NU); Grice s.n. (NU); Billiard & Bunt 15,603 (PRE); Castle View
Farm (-CB), Billiard & Bunt 13,644 (PRE); Mkomazi River (-CB), Billiard &
Bunt 15,861 (PRE); Garden Castle (-CD), lbn der Merwe 2761 (NU); lbn der
Merwe 2762 (NU, PRE). -2930 (Pietermaritzburg): Merrivale (-AC), Moll
1341A (NU); Moll 1341b (NU); Ivanhoe (-AC), Moll 2597 (NU); Howick (-
AC), Wood 3479 (NH); Wood 8136 (NH); Drayton, Nottingham Road (-AC),
Smith 188 (NU); Karkloof (-AD), Stirton 12,251 (UNIN); Greytown (-BA),
Rjlie s.n. sub PRE 34,293 (PRE); Rjlie s.n. sub NH 21,692 (NH); Elandskop,
farm Glazley (-CA), Morris 107 (PRE); Elandskop (-CA), Webb 37 (NU);
Zwartkops (-CB), Lindah152 (NU); Williams 66 (NU); Pietermaritzburg (-CB),
Van der Merwe 2846 (PRE); Pietermaritzburg, Town Hill (-CB), Van der
Merwe 2602 (NU); Bond 1296 (NBG); Richmond (-CD), Vivyan s.n. sub BOL
50,786 (BOL); Umgeni, Kransberg (-DA) , Thode 5400 (STE); Gilletts (-DD),
Wood 11,568 (PRE); Wood 19,249 (PRE). -3029 (Kokstad): Kokstad, Mount
Currie (-AD) Goossens 245 (PRE); Ngeli Mountain (-DA), Van H)7k 7513
(PRU); Ngeli Peak (-DA), MacDevette 1311 (NH). -3030 (Port Shepstone):
Umdoni Park (-BC), Jarman & Grey 394 (NU); Port Shepstone (-CB), Brown
323 (BOL); Port Shepstone, farm The Valleys (-CB), Mogg 13,884 (PRE); Sea
Park (-CB), Elery s.n. sub NBG 73,436 (NBG).
TRANSKEI -2929 (Underberg): Sehlabathebe National Park (-CC),
Hoener 2081 (PRE). -3128 (Umtata): Mhlhlani Forest (-BC), Perry 3415
(NBG); Baziya Forest Station (-CB), Van der Merwe 1881 (PRE). -3228
(Butterworth): Elliotdale, The Haven (-BB), Gordon-Grey 812 (NU); Kentani (CB), Pegler 1141 (BOL).
CAPE -3227 (Stutterheim): Stutterheim (-CB), Rogers 12,796 (GRA);
Gonubie (-DD), Batten 6 (NBG).
LEDEBOURJA IN SOUTH AFRJCA
107
TAXONOMY
Subsectio Erectifoliae S. \-enter, subsect. nov., bulbis cylindricis; foliis erectis.
Species typica: L. viscosa Jessop.
Species: L. dolomiticola S. Venter, L. atro-brunnea S. Venter, L. viscosa
Jessop.
Bulbs cylindrical. Leaves erect, concolorous. Raceme lax.
4. LEDEBOURIA DOLOMITICOLA S. Venter
Ledebouria dolomiticola S. Venter, sp. nov., ad L. socialem (Bak.) Jessop
affinis sed bulbis erectis cylindricis epigaeis; foliis glaucis erectis et
inflorescentia flaccidis differt.
Type: Lebowa, Strydpoort Mountain, Donkerkloof, Farm Rivierplaats 354,
W!nter 13,089a (PRE!, holo.; NU!, UNIN!).
Bulbs epigeal, 40 - 100 X 15 - 30 mm, elliptic to cylindrical, in dense groups;
dead bulb scales thinly membranous, brown, apices attenuate, live bulb scales
tightly arranged, without threads when torn, white inside; bulblets on basal
stem. Leaves fully developed at anthesis, 3 - 5, erect, lanceolate, 80 - 100 x 8 20 mm, without threads when tom, fleshy, surfaces with a dull lustre, glaucous
green without markings, venation obscure; margin smooth; leaf base
canaliculate; apex acute. Inflorescence solitary, lax, cylindric, 40 - 60 x 20 - 30
mm, flaccid, 30 - 40 -flowered, longer than leaves; scape terete at base, green,
glabrous; rachis smooth, scape smooth, 60 - 90 mm long. Bracts membranous,
1.5 x 0.25 mm, linear-lanceolate, grey to white with bracteoles present. Pedicel
spreading horizontally, 8 - 10 mm long, white to purple. Perianth 5 mm long,
tepals recurved, subequal, oblong, 5 x 1.0 - 1.5 mm, apex obtuse, pink to
purple with a green keel. Stamens erect, 3 mm long, filaments maroon,
epitepalous; anthers 0.5 mm long, yellow. Ovary globose, 6 -lobed, 1.5 x 2.5
mm, lobes narrowly transversely oblong, apex shoulders raised, basal lobes
present. Style 3 mm long, triangular, glabrous, purple above and white below;
stigma above anthers; stipe 0.25 - 0.5 mm. Capsule 3 - lobed, symmetrical,
globose; base truncate. Seed globose, 4 mm long, surface strongly wrinkled,
brown. (Figure 22).
LEDEBOURlA IN SOlITH AFRICA
108
TAXONOMY
E
c
C>
H
A
~
J
~
B
Figure 22. Illustration of L. dolomiticola S. Venter. A, habit Xl; B, lamina
margin X 300; C, lamina apex X 20; D, bract and bracteole X 10; E, section
through lamina X 5; F, tepa! apex X 20; G, flower X 10; H, tepa! with stamen
X 10; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from Venter
13,208a.
LEDEBOURJA IN SOUTH AFRICA
Known distribution of L. dolomiticola S. Venter
Map 8.
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G.P.-S. 13--0959
110
TAXONOMY
Figure 23. A, holotype of L. dolomiticola S. Venter (PRE); B, habitat at
Donkerkloof, the type locality. The vegetation consists of closed deciduous low
Kirkia wilmsii - Dombeya autumnalis - Obetia tenax woodland on dolomite; C,
plants of L. dolomiticola growing in rock cracks showing the epigeal bulbs. Bar
= 100 mm; D, SEM micrograph of the adaxial lamina surface. Bar = 100 JLm;
E, SEM micrograph of the abaxial lamina surface. Bar = 100 JLm; F, SEM
micrograph of stomata. Bar = 43 JLm. D - F from W!nter 13,089a.
LED£BOURlA IN SOUTH AFRICA
111
TAXONOMY
LED£BOURIA IN SOlITH AFRICA
TAXONOMY
IU
L. dolomiticola is closely related to L. atro-brunnea and L. viscosa in having
cylindrical bulbs, erect leaves (uncommon in Ledebouria) and the ovary apex
forming prominent shoulders. L. dolomiticola differs from both in its epigeal
bulbs and membranous dead bulb scales.
Specific epithet etymology.
Refers to the rock type upon which this species is localized.
Flowering period
From January to April.
Distribution (Map 8).
Known only from the Strydpoort Mountain near Pietersburg, north eastern
Transvaal.
Habitat
Plants are limited to dolomite of the Eccles Formation of the Chuniespoort
Group (SACS 1980). Where an increase in chert (quartz) content in the dolomite
occurs L. dolomiticola appears to be excluded. The steep slopes and cliffs upon
which L. dolomiticola grows experience high temperatures especially during
summer months. The soil is a dark brown clay loam. Plants tend to grow in
rock cracks (Figure 23C) with some plants in lithosols.
Population structure
Two populations are known, the smaller consists of few (+ 35) individuals on a
90° cliff. The larger population consists of ± 121 individuals, on a steep 30° 45 ° slope covered mostly in large dolomitic rocks. Adult plants predominate
with 5 - 18 epigeal bulbs per plant.
Specimens examined
TRANSVAAL. - (2429): Strydpoort Mountain, Donkerkloof, Farm
Rivierplaats 354 (-BA), l-enter 13,089a (PRE, UN, UNIN).
UDEBOURIA IN SOUTH AFRICA
113
TAXONOMY
5. LEDEBOURIA ATRO-BRUNNEA S. Venter
Ledebouria atro-brunnea S. Venter, sp. nov., ad L. marginatam (Bak.) lessop
affinis sed squamis duris, atro-brunneis et foliis glaucis erecto-tortis clare
differt.
Type: Transvaal, Rustenburg, Kroondal on farm Zuurplaat 337, lenter 13,460
(PRE!, holo.; NU!; UNIN!).
Plants solitary. Bulb hypogeal, 30 - 60 x 20 - 30 mm, cylindrical; dead
bulb scales purplish-brown, very hard, apices attenuate, live bulb scales loosely
arranged, with threads when torn, bulb white spotted heavily purple inside.
Leaves partly emerged at anthesis, 4 - 6, erect, spirally twisted, linearlanceolate, 60 - 80 x 3 - 10 mm, with threads when torn, fleshy, surfaces dull
glaucous green, venation obscure; margin smooth but wavy in lower part; leaf
base canaliculate; apex acute. Inflorescences I - 4, lax, oblong, 20 - 50 x 25 30 mm, flaccid, 30 - 60 -flowered, longer than the leaves; scape compressed at
base, purple with darker spots, glabrous; rachis ridged, 25 - 60 mm long.
Bracts semi-fleshy, I x 0.25 - 0.5 mm, linear to bifurcate, grey-white with
bracteoles always present. Pedicels spreading, 6 - 7 mm long, speckled or plain
coloured pink. Perianth 5 mm long, tepals recurved, subequal, linear-oblong, 5
x 1.5 mm, apex acute, slightly cucullate, pink to purple with a green keel.
Stamens erect, 3 - 4 mm long, filaments pink, base slightly flattened,
epitepalous; anthers 0.75 - 1.0 mm long, violet. Ovary ovoid, 6 -lobed, 1.0 1.5 x 2.0 - 2.5 mm, lobes obtusely deltate, apex shoulders absent, basal lobes
present. Style 2.0 - 2.5 mm long, triangular, glabrous, purple; stigma equal
height to anthers; stipe 0.25 x 0.25 - 0.5 mm. Capsule 2- to 3 - lobed,
symmetrical, clavate; base truncate. Seed drop-shaped, 4 - 5 mm long, surface
strongly wrinkled, reddish-brown. (Figure 24).
L. atro-brunnea is related to L. dolomiticola and L. viscosa in having cylindrical
bulbs ~tiw
~uonarb 8
.Q@atl btllb sCMes-and erect leaves but is distinguished by
the twisted leaves and the very hard bulb scales.
L£DEBOURLA IN SOtITH AFRICA
114
TAXONOMY
B
Figure 24. Illustration of L. atro-brunnea S. Venter. A, habit Xl; B, lamina
margin X 300; C, section through lamina X 5; D, bract X 20; E, tepal with
stamen X 10; F, flower X 10; G, tepal apex X 20; H, ovary lateral view X 10;
I, ovary dorsal view X 10. All from \enter 13,460.
UDEBOURJA IN SOUTH AFRICA
Known distribution of L. atro-brunnea S. Venter
Map 9.
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116
TAXONOMY
Figure 25. A, holotype of L. atro-brunnea S. Venter (PRE); B, habitat near
Kroondal, north-western Transvaal. The vegetation consists of closed deciduous
tall Protea caffra subsp. caflra - Xerophyta retinervis - Parinari capensis
woodland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of wax platelets
forming the cuticle; F, plants of L. atro-brunnea showing the colonial growth
and the erect, twisted leaves. Bar = 30 mm. C - F from \enter 13,460.
LEDEBOURlA IN SOlITH AFRICA
117
TAXONOMY
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LEDEBOURlA IN SOlITH AFRICA
118
TAXONOMY
Specific epithet etymology.
Refers to the dark brown bulb scales.
Flowering period
From October to December.
Distribution (Map 9).
Kroondal near Rustenburg, north-western Transvaal, at the foothills of the
Magaliesberg.
Habitat
Throughout its distribution range, the rocks are from the Magaliesberg Quartzite
Formation of the Pretoria Group. These rocks are coarse, recrystallised
quartzite forming mother material for the shallow reddish-brown sandy lithosol.
The vegetation is Bankenveld (Acocks 1988) and consists of closed deciduous
low Protea caffra var. caffra - Lannea discoIor woodland (Figure 25B).
Population structure
A single population of ± 500 x 1000 m was found consisting of ± 300
individuals. Plants were scattered or occasionally occurred in small clumps
(Figure 25F). Most of the plants were adults. Few seedlings in the two leaf
stage were seen but many seeds were collected next to most of the adult plants.
The leaves of the seedlings are erect from the two leaf stage and purple
underneath, becoming glaucous with age.
LEDEBOURlA IN SOlITH AFRICA
TAXONOMY
119
The only protection afforded plants is from grass tussocks. Veld fires commonly
occur and the very hard dead bulb scales protect the viable inner leaves. The
apices of the dead bulb scales possibly act as a defence mechanism against
herbivores.
Variation
Plants growing in the shade of trees and shrubs have more prominent dead
bulb scales and the leaves of the plants growing in shade tend to be less twisted.
Plants grown under controlled conditions in an aridarium at the University of
the North, Sovenga, showed leaf twisting to be independant of moisture
regimes.
Specimens examined
CD),
~nter
TRANSVAAL. - 2527 (Rustenburg): Kroondal, Farm Zuurplaat 337 (13,460 (NU, PRE, UNIN); ~nter
13,483 (PRE, UNIN).
LEDEBOUlUA IN SOUTH AFRICA
120
TAXONOMY
6. LEDEBOURIA VISCOSA Jessop
Ledebouria viscosa Jessop in Jl S. Afr. Bot. 36(4): 264 (1970).
Type: Transvaal, Thabazimbi, Kransberg, Meeuse 10,493 (PRE!, holo.).
Plants solitary. Bulb hypogeal, 50 - 60 x 12 - 20 mm, cylindrical; dead
bulb scales brown, membranous, apices attenuate, live bulb scales fleshy, tightly
arranged, without threads when torn, white inside; basal stem up to 120 mm
long. Leaves fully developed at anthesis, 1 - 3, erect, oblanceolate to spathulate,
90 - 230 x 17 - 30 mm, without threads when torn, fleshy, surfaces covered
with a thick resinous layer, viscid, immaculate, dark green with sometimes a
purple tinge, venation sometimes prominent; margins smooth; leaf base flat to
very shallowly canaliculate; apex obtuse. Inflorescence solitary, lax, cylindric,
100 x 300 mm, erect, 20 - 30 -flowered, longer than the leaves; scape terete at
base, green, glabrous; rachis smooth, 150 - 170 mm long. Bracts membranous, .
1.5 x 0.5 mm, linear-lanceolate, bifurcate at base of raceme, grey to white,
without bracteoles. Pedicels spreading horizontally, 7 - 8 mm long, white.
Perianth 5 mm long, almost tubular, tepals equal, linear - oblong, 5 x 1.5 mm,
apex acute, pink to purple with a green keel. Stamens erect, 4 - 5 mm long,
filaments maroon, epitepalous; anthers I mm long, pale violet. Ovary ovoid, 6
-lobed, 1 x 2 mm, lobes deltate, apex shoulders prominent. Style 1.5 mm long,
terete, glabrous, white; stigma below anthers; stipe 0.5 x 0.5 mm. Capsule 2 3 lobed, asymmetrical, clavate; base truncate. Seed drop-shaped, 4 mm long,
surface strongly wrinkled, brown. (Figure 26).
L. viscosa is related to L. dolomiticola and L. atro-brunnea. Together they
constitute the subsection Erectifoliae.
L. viscosa cannot be confused with other Ledebouria species in South Africa.
The erect viscid leaves and solitary erect inflorescence are diagnostic (Figure
26A).
LEDEBOURlA IN 50IJTH AFRICA
Ul
TAXONOMY
o
:t,....
yil{.
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G
H
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A
Figure 26. lllustration of L. viscosa Jessop. A, habit X 1; B, lamina margin X
300; C, section through lamina X 2; D, bract X 10; E, flower X 10; F, tepal
with stamen X 10; G, tepal apex X 20; H, ovary lateral view X 10; I, ovary
dorsal view X 10. All from \enter 13,455.
LEDEBOURlA IN SOlJfH AFRICA
. Map 10.
12°
Known distribution of L. viscosa Jessop
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123
TAXONOMY
Figure 27. A, holotype of L. viscosa Jessop (PRE); B, habitat near Thabazimbi.
The vegetation consists of closed deciduous low Terminalia sericea - Acacia
tortilis subsp. heteracantha - Grewia flava - Digitaria eriantha woodland; C,
SEM micrograph of the adaxial lamina surface; D, SEM micrograph of the
abaxial lamina surface; E, leaves of L. viscosa with sand particles adhering to
the surfaces. Bar = 10 mm. F, SEM micrograph of the stomata; G, SEM
micrograph showing the thick resinous covering of the lamina; H, leaf
unrolling, a unique character in Ledebouria. Bar = 20 mm. C - H from l.enter
13,455.
LEDEBOURlA IN SOurH AFRICA
124
TAXONOMY
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LEDEBOURlA IN SOUTH AFRICA
125
TAXONOMY
Specific epithet etymology.
Describes the viscid leaves (Figure 27E).
Flowering period
From January to April. Most plants flower during March and April.
Distribution (Map 10).
Endemic around Thabazimbi in the north-western Transvaal.
Habitat
Known populations of L. viscosa occur on well drained, loose, red to brown,
medium grained (0.25 - 1.0 mm cP) deep sandy soil of the Gordonia Sand
Formation of the Kalahari Group.
The vegetation consists mainly of closed deciduous tall Terminalia sericea -
Acacia tortilis subsp. heteracantha - Digitaria eriantha woodland (Figure 27B).
Specimens examined
TRANSVAAL -2427 (Thabazimbi): Kransberg (-BC), Meeuse 10, 493
(PRE); Kransberg, farm Waterval (-BC), Dyer & Ehrens 4201 (PRE); farm
Buffelshoek 446 (-DA), lknter 13,455 (UNlN).
LEDEBOURlA IN SOUTH AFRICA
126
TAXONOMY
Subsectio Stellatae S. lknter, subsect. nov., perianthio sigillatim stellato.
Species typica: L. rupestris (Van der Merwe) S. Venter.
Species: L. leptophylla (Bak.) S. Venter, L. lepida (N.E. Br.) S. Venter, L.
minima (Bak.) S. Venter, L. rupestris (Van der Merwe) S. Venter.
Leaves with adaxial surface concolorous. Raceme 10 - 25 mm long. Perianth
stellate; tepal apices acute.
7. LEDEBOURIA LEPTOPHYLLA (Bak.) S. Venter
Ledebouria leptophylla (Bak.) S. lknter, comb. novo
Scilla leptophylla Bak. in Flora Cap. 6: 483 (1896).
Type: Transvaal, Near the Devil's Kantoor, Kaapsche Hoop, Bolus 7623 (K !,
holo.; BOL !, drawing and iso.).
Scilla graminijolia Bak. in Bull. Herb. Boiss. ser. 2(4): 1001 (1904).
Type.: Transvaal, Modderfontein, Conrath 703 (K!, holo.; BOL!, drawing;
GRAZ; PRE!, photo.; Z!).
Ledebouria graminijolia (Bak.) Jessop in Jl S. Afr. Bot. 36(4): 259 (1970).
Type: Same as S. graminijolia.
Scilla stenophylla Van der Merwe in Flower. PI. S. Afr. 25: t.959 (1944).
Type: Natal, Paulpietersburg, lfln der Merwe 2655 (PRE!, holo.).
Plants solitary. Bulb hypogeal, 40 - 60
X
20 - 30 mm, obovoid to globose; dead
bulb scales light to dark brown, membranous, apices attenuate, live bulb scales
fleshy, tightly arranged, without threads when tom, white inside. Leaves partly
emerged at anthesis, 8 - 20, erect or sometimes spreading, spirally twisted,
linear, 60 - 150 x 2 - 5 mm, with threads when torn, thinly fleshy to slightly
LEDEBOURIA IN SOlJTH AFRICA
127
TAXONOMY
F
o
H
~
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Figure 28. Illustration of L. leptophylla (Bak.) S. Venter. A, habit X 1; B,
lamina margin X 300; e, section through lamina X 10; D, tepa! apex X 20; E,
tepa! with stamen X 10; F, bract with bracteole X 10; G, flower X 10; H, habit
of nonspiral-leaved form X 1; I, ovary lateral view X 10; J, ovary dorsal view
X 10. All from \enter 13,214 except H from lenter 13,251.
LEDEBOURlA IN SOUTH AFRICA
Map 11.
Known distribution of L. leptophylla (Bak.) S. Venter
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12°
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129
TAXONOMY
Figure 29. A, holotype of L. leptophylla (Bak.) S. Venter (K); B, miniature
form resembling L. minima (Bak.) S. Venter. Bar = 20 mm; C, SEM
micrograph of the adaxial lamina surface; D, SEM micrograph of the abaxial
lamina surface; E, SEM micrograph of stomata; F, habitat at Angle Station,
between Barberton and Havelock. The vegetation consists of open low Protea
roupelliae var. roupelliae - Helichrysum splendidum - Xerophyta retinervis
woodland; G, plants with nonspiral leaves. Bar = 20 mm. Band G from "U?nter
13,251 and C - E from l-enter 13,214.
LEDEBOURJA IN SOUTH AFRICA
130
TAXONOMY
.
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LEDEBOURIA IN SOlITH AFRICA
131
TAXONOMY
leathery, surfaces with a dull lustre, grey-green to glaucous green, abaxial
surface with dull purple spots and cross bars at base, venation mostly
prominent; margin smooth to finely papillate; leaf base canaliculate; apex acute.
Inflorescences 1 - 3, dense, oblong, 25 x 15 mm, erect to flaccid, 25 - 30
-flowered, longer than the leaves; scape terete at base, green, glabrous; rachis
ridged, scape smooth, 18 - 20 mm long. Bracts membranous, 1 x 0.25 mm,
linear - lanceolate, grey to white, with bracteoles. Pedicels spreading, 4 mm
long, pink. Perianth 4 - 6 mm long, stellate, tepals suberect, equal, 4 - 6 x
1.75 mm, apex obtuse, pink with a green keel. Stamens erect, 4 - 6 mm long,
filaments maroon, epitepalous; anthers 1 mm long, pale violet. Ovary ovoid, 6
-lobed, 1 x 2 mm, lobes narrowly transversely oblong, apex shoulders
rectangular, base lobed. Style 3 mm long, terete, glabrous, purple; stigma equal
height to anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical,
globose; base truncate. Seed drop - shaped, 2 - 3 mm long, surface strongly
wrinkled, black. (Figure 28).
L. leptophylla is related to L. minima in having glaucous, linear leaves, stellate
flowers and prominent ovary shoulders but differs in the papillate lamina
margin, leaves with threads when torn, ridged rachis, prominant bracteole and
ovary shoulders.
Specific epithet etymology.
Describes the narrow, linear leaves.
Flowering period
From August to January with peak flowering in November.
Distribution (Map 11).
L. leptophylla occurs mainly in woodlands of the Transvaal with scattered
localities in Swaziland, Natal and the Orange Free State.
LEDEBOURJA IN SOlJfH AFRICA
132
TAXONOMY
Habitat
L. leptophylla grows in full sun in open grassveld areas. In high lying areas it
occurs in montane grassland only.
Variation
There are two distinct forms of L. leptophylla. One form has leaves slightly
twisted, 4 - 5 mm wide, spreading and green. It occurs on the Transvaal
Drakensberg Escarp between Graskop and Sabie (lenter 13,251) (Figure 29G).
The more common form has strongly twisted glaucous green leaves 2 - 4 mm
wide, leathery textured and copious threads when tom (lenter 13,214).
The flower colour varies from green (rare), pink with a green keel (common
form) to pink. Although the flowers of L. leptophylla are stellate, the proximal
half of the tepals tends to bend slightly backwards in some plants.
Specimens examined
TRANSVAAL. - 2229 (Waterpoort): Messina, Dongola (-BC), PoleEvans 4373 (PRE). -2230 (Messina): Messina (-AC), Moss & Rogers 7 (J);
Moss & Rogers 205 (J). -2329 (Pietersburg): Bandelierkop (-BD), Van der
Merwe 1727 (PRE); Pietersburg (-CD), ~nter
13,332 (UNIN); ~nter
13,367
(UNIN). -2330 (Tzaneen): Middle Letaba Dam (-AD), lenter 13,064 (UNIN).
-2427 (Thabazimbi): Kranzberg (-BC), WJn der Merwe 2025 (PRE); WJn der
Merwe 2019 (PRE). -2428 (Nylstroom): Vaalwater, Geelhoutkop (-AD), lenter
13,214 (UNIN); Nylstroom (-CB), Hafstrom & Acocks 233 (PRE); Warmbaths
(-CC), Leendertz 6654 (PRE); Leendertz 6653 (J). -2429 (Zebediela):
Sekoekoeniland (-DB), WJn der Merwe 1768 (PRE). -2430 (Pilgrim's Rest):
Graskop, Kowyn's Pass (-DD), ~nter
12,573 (UNIN); Mac Mac (-DD), lenter
13,251 (UNIN). -2528 (Pretoria): Pretoria (-CA), Moss 8594 (J); Mogg 15,189
(PRE); Pretoria, Faerie Glen (-CD), WJn k~
2379 (PRU); -2529 (Witbank):
Stoftberg (-BD), WJn der Merwe 2025 (PRE). -2530 (Lydenburg): Kaapsche
LEDEBOURlA IN SOlITH AFRICA
133
TAXONOMY
Hoop (-DB), Rogers 21,408 (J); Nelshoogte, Thorncroft Nature Reserve (-DD),
Raal & Raal1602 (Transvaal Provincial Administration). -2626 (Klerksdorp):
Ventersdorp, farm Somerville (-BD), Louw 2292 (PUC). -2627
(Potchefstroom): Potchefstroom (-CA), Venter 13, 220a (UNIN). -2628
(Johannesburg): Widsor Park (-AA), Gilliland 26138 (1); Zoo Koppies (-AA),
Moss 7144 (1); Edenvale, Rietfontein (-AA), Venter 13,340 (UNIN); Alberton
(-AC), Moss 13,902 (J); Palmietfontein (-AC), Lloyd s.n. (J). -2629 (Bethal):
Ermelo (-BD), Scholars 60 (PRE); Between Ermelo and Breyten (-BD), \enter
3540 (PRE). -2630 (Carolina): Carolina (-AA), Rogers 21,300 (J); Amsterdam,
Lichfield (-DD), liln der Merwe 1071 (PRE). -2730 (Vryheid): Piet Retief,
between Wittenberg and Bergen (-BB), Acocks 11,735 (BOL).
SWAZILAND. - 2631 (Mbabane): Bomvu Ridge (-AC), Karsten s.n.
(PRE); Sibebe Hill (-CB), Prior 257 (PRE).
ORANGE FREE STATE. - 2824 (Kimberley): Riverton (-DB), Wilman
s.n. sub BOL 22,488 (BOL). -2926 (Bloemfontein): Bloemfontein (-AA), Moss
10,806 (1); Potts 1340 (BOL).
NATAL. - 2731 (Louwsburg): Ngotshe, Ngome (-CD), Billiard & Bunt
8421 (NU, PRE). -2930 (Pietermaritzburg): Oribi Airfield (-CB), Moll 1861
(NU).
LEDEBOURlA IN SOlJfH AFRICA
134
TAXONOMY
8. LEDEBOURIA LEPIDA (N.E. Br.) S. Venter
Ledebouria lepida (N.E. Br.) S. \-enter, comb. novo
Scilla lepida N.E. Br. in Kew Bull. : 299 (1921).
Type: Transvaal, Palala River, Breyer s.n. in Herb. Rogers 24009 (K!, holo.;
BOLl, drawing; PRE!).
Plants solitary. Bulb hypogeal, 10 - 15 x 10 - 15 mm, ovoid to subglobose;
dead bulb scales brown, membranous, apices attenuate, live bulb scales fleshy,
tightly arranged, with threads when torn, white inside. Leaves fully developed
at anthesis, 2 - 3, spreading or sometimes erect, lanceolate, 30 - 50 x 8 - 20
mm, without threads when torn, fleshy, adaxial surface with rows of cilia,
surfaces glossy dark green with purple cross bars at the leaf base, immaculate
above, venation prominent; margins occasionally revolute, with rows of cilia;
leaf base canaliculate; apex acute. Inflorescences 1 - 2, dense, oblong, 10 - 15
x 10 - 12 mm, erect, 13 - 20 -flowered, longer than the leaves; scape terete at
base, purple, pilose; rachis smooth, 30 - 50 mm long. Bracts fleshy, 0.5 x 0.5
mm, dentate, pink to purple, with bracteoles. Pedicels spreading, 2.0 - 2.5 mm
long, purple. Perianth 3.0 - 3.5 mm long, stellate, tepals suberect, equal,
oblong to linear, 3.0 - 3.5 x 1.5 mm, apex acute, pink to purple. Stamens erect,
3 mm long, filaments maroon, epitepalous; anthers 1 mm long, violet. Ovary
globose, 6 -lobed, 0.75 - 0.80 x 1.25 - 1.50 mm, lobes depressed ovate, apex
tapering into the style. Style 3 mm long, triangular, glabrous, purple; stigma
above anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, globose;
base truncate. Seed globose, 2 mm long, surface strongly wrinkled, black.
(Figure 30).
Together with L. leptophylla (Bak.) S. Venter, L. minima (Bak.) S. Venter and
L. rupestris (Van der Merwe) S. Venter, it comprises the subsection Stellatae.
A distinctive feature of this species is the pilose scape and the small flowers.
L£D£BOURIA IN SOUfH AFRICA
135
TAXONOMY
E
G
H
•
~
J
B
A
Figure 30. Illustration of L. lepida (N.E. Br.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, bract X 10; D, hairs on peduncle X 10; E, se~tion
through lamina X 2; F, flower X 10; G, tepa! with stamen X 10; H, tepa! apex
X 10; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from retn~
13,415.
UDEBOURlA IN SOUTH AFRICA
Map 12.
12°
18
Known distribution of L. lepida (N.E. Br.) S. Venter
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137
TAXONOMY
Figure 31. A, holotype of L. lepida (N.E. Br.) S. Venter (K); B, habitat near
Palala, Waterberg. The vegetation consists of closed deciduous low Combretum
zeyheri - Mimusops zeyheri - Freylinia tropica woodland; C, SEM micrograph
of the adaxial lamina surface. Bar
=
100 J.'m; D, SEM micrograph of the
abaxial lamina surface; E, SEM micrograph of stomata. Bar = 43 J.'m; F,
abaxial surface of the lamina showing the prominent purple zebra stripes on the
petiole. Bar = 10 mm. C - F from l-enter 13,415.
LEDEBOURlA IN SOlITH AFRICA
TAXONOMY
138
- - .•
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LEDEBOURlA IN SOlJTH AFRICA
139
TAXONOMY
Specific epithet etymology.
Refers to the hairs on the leaves and scape.
Flowering period
From November to January. Peak flowering in December.
Distribution (Map 12).
A Transvaal and Waterberg endemic. This species is known only from the
Palala area.
Habitat
The rocks in the Palala area are feldspathic sandstones of the Vaalwater
Formation in the Waterberg Group (SACS 1980). The soils derived from these
rocks are a gritty brown sandy loam with the grain size usually 0.5 - 2.0 mm
diameter. These soils are usually shallow (10 - 45 mm) where the rocks form
outcrops but deeper (100 - 450 mm) along the drainage lines.
L. lepida grows in seepages in Combretum woodland. The vegetation is a closed
deciduous low Combretum nelsonii - Rhoicissus digitata - Heteropyxis natalensis
Woodland (Figure 31B).
Specimens examined
TRANSVAAL. - 2428 (Nylstroom): Vaalwater, Geelhoutkop (-AD),
liln der Merwe 324 (PRE); Vaalwater, Palala (-BC), Breyer s.n. sub Herb EA.
Rogers 24,009 (K, PRE); Venter 13,415 (UNIN).
LEDEBOURIA IN SOlITH AFRICA
140
TAXONOMY
9. LEDEBOURlA MINIMA (Bak.) S. Venter
Ledebouria minima (Bak.) S. \enter, comb. novo
Type: As for Scilla minima.
Scilla minima Bak. in Saund. Ref. Bot. 3(App.): 6 (1870).
Type: Transvaal, on Macalisberg, Burke s. n. (K!, holo.; BOLl, drawing; PRE!,
photo.).
Plants solitary. Bulb hypogeal, 8 - 10 x 4 - 7 mm, ovoid to obovoid; dead
bulb scales brown to whitish, membranous, apices attenuate, live bulb scales
membranous, tightly arranged, without threads when tom, white inside. Leaves
fully developed at anthesis, 2 - 5, spreading, linear, 24 - 36 x 2 - 3 mm, without
threads when tom, fleshy, surfaces dull green with dull purple cross bars at base
of leaf, venation obscure; margin smooth; leaf base canaliculate; apex acute.
Inflorescences 1 - 2, lax, oblong, 18 - 24 x 10 - 12 mm, flaccid, 12 - 20
-flowered, longer than the leaves; scape terete at base, green, glabrous; rachis
smooth, scape smooth, 38 - 50 mm long. Bracts fleshy, 1 x 0.5 mm, dentate,
pink to purple, without bracteoles. Pedicels spreading, mostly horizontal, 3 - 4
mm long, pink. Perianth 2 mm long, stellate, tepals slightly recurved, equal,
linear-oblong, 2 x 1 mm, apex acute, keel green on pink. Stamens erect, 2 mm
long, filaments pink, epitepalous; anthers 0.5 mm long, violet. Ovary
depressed ellipsoidal, 6 -lobed, 0.5 x 1.12 mm, lobes depressed-ovate, glabrous
with base of lobes papillate. Style 2 mm long, triangular, glabrous, purple;
stigma carried above anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed,
symmetrical, clavate; base truncate. Seed drop-shaped, 3 mm long, surface
strongly wrinkled, brown. (Figure 32).
L. minima is closely allied to L. rupestris (Van der Merwe) S. Venter. It differs
in the glabrous, linear leaves and absence of bracteoles. Together with L. lepida
and L. leptophylla they constitute the subsection Stellatae.
LEDEBOURlA IN SOUTH AFRICA
141
TAXONOMY
o
E
G
B
A
H
Figure 32. Illustration of L. minima (Bak.) S. Venter. A, habit X 1; B, section
through lamina X 5; C, lamina margin X 300; D, bract X 10; E, tepal apex X
20; F, flower X 10; G, tepal with stamen X 10; H, ovary lateral view X 10; I,
ovary dorsal view X 10. All from W!nter 13,404.
LEDEBOURlA IN SOUTH AFRICA
Map 13.
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143
TAXONOMY
Figure 33. A, holotype of L.minima (Bak.) S. Venter (K); B, habitat on "The
Downs" near Trichardtsdal, north-eastern Transvaal. The vegetation consists of
Tristachya leucothrix - Panicum natalense - Scilla natalensis montane sour
grassland; C, SEM micrograph of the adaxial lamina -surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata; F,
plants showing filiform leaves. Bar = 20 mm. C - F from W!nter 13,404.
LEDEBOURIA IN SOlJTH AFRICA
144
TAXONOMY
UDEBOURlA IN SOlITH AFRICA
145
TAXONOMY
Specific epithet etymology.
Describing the small stature of the plants.
Flowering period
From October to March with peak flowering from October to December.
Distribution (Map 13).
Throughout South Africa with two centres of distribution. The first centre
around the PWV (Pretoria - Witwatersrand - Vereeniging) area and the second
covering the southern Cape.
Habitat
L. minima grows in medium grained (0.25 - 0.25 mm 4» shallow (l - 5 mm
deep) humusrich sandy to loamy soil. Most of the populations seen occur in
perennial seepages in short grassland (Figure 33B).
Population areas are usually small (
< 200 m2 ). Leaf shape and colour of
seedlings resemble grass making them cryptic.
Variation
Markings on the leaves in the northern populations tend to be restricted to dull
purple cross bands at the leafbase. The Cape populations have plants with
purple-mottled leaves, somewhat broader than their northern counterparts.
Some individuals have plain pink flowers compared to the normal pink flowers
with a green keel on the tepals.
Specimens from Gordonia (Leistner 2034) and Prieska (Bryam 743) are here
referred to as L. minima but differ in their broader leaves and larger flowers.
LEDEBOURIA IN SOlJfH AFRICA
146
TAXONOMY
Specimens examined
TRANSVAAL - 2430 (Pilgrim's Rest): The Downs (-AA), Stalmans 832
(PRE). -2527 (Rustenburg): Magaliesberg, Rietpoort (-DC), Moss & Maguire
s.n. sub J 22362 (1); Rustenburg, farm Uitkomst (-DD), Coetzee 438 (PRU).
-2528 (Pretoria): Irene (-CC), Obermeyer 77 (PRE); Leendertz 923 (PRE).
s.n. sub J
-2627 (Potchefstroom): Modderfontein (-BB), Lloyd s.n. (PRE); ts~
31505 (1); Florida (-BB), Dahlstrand 1015T (PEU); Sterkfontein Caves (-BB),
Mogg 36,275 (1); Potchefstroom, Boskop (-CA), Ubbink 506 (PUC);
Potchefstroom (-CA), Ubbink 742 (PRE). -2628 (Johannesberg): Johannesburg
(-AA), Moss 20,518 (1); Elsburg (-AA), Lloyd s.n. (1); Northcliff (-AA),
Gilliland s.n. (1); Thorntree Kloof (-AA), Moss 5588 (J); Moss 10,813 (J);
Frankenwald (-AA), Van Rensburg 25,411 (1); WIn Rensburg 25,313 (PRE);
Phillips s.n. sub J 25892 (J); Jessop 687 (PRE); Jessop 690 (PRE); Heidelberg,
farm Schoongesicht 302 (-AD), Burtt Davy 15,427 (BOL); Vaaldam, farm
Sandfontein (-CD), Aspoas s.n. sub J 48392 (1).
ORANGE FREE STATE - 2927 (Maseru): Thaba Nchu,
Mensvretersberg (-AA), Peeters et al. 323 (PRE). -2926 (Bloemfontein):
Bloemfontein (-AA), Maller 420 (NBG,PRE); Hillandale (-AA), Vtbssenjall 856
(NBG).
NATAL - 2731 (Louwsburg): Ngome Forestry (-CD), WIn ")1k 7095
(PRU).
CAPE - 2822 (Glen Lyon): Gordonia, Skeurberg Mountains on farm
England (-AD), Leistner 2034 (PRE). 2922 (Prieska): Prieska (-DA), Bryant
743 (BOL); Bryant s. n. sub STE 18328 (STE); Fuller 114 (BOL). 3221
(Merweville): Prince Albert (-DC), Krige s.n. sub BOL 12980 (BOL). -3320
(Montagu): Kareevlakte (-DA), Perry 1240 (NBG). 3323 (Willowmore):
Uniondale, Kouga (-CA), Esterhuisen 4672 (BOL). 3325 (Port Elizabeth):
Bluewater Bay (-DC), Urton 890 (PEU); 3419 (Caledon): Zoetendals Vlei (DB), Fellingham 396 (STE). 3420 (Bredasdorp): Potberg Nature Reserve (-BC),
Burgers 1812 (STE). 3421 (Riversdale): Glen (-AB), Muir 3004 (BOL). 3422
(Mossel Bay): Mossel Bay (-AA), Marloth 7784 (PRE); Belvedere (-BB),
Duthie 506 (STE). 3423 (Knysna): Knysna Heads (-AA), Schonland 3677
(BOL).
LEDEBOURlA IN SOtJrH AFRICA
147
TAXONOMY
10. LEDEBOURIA RUPESTRIS (Van der Merwe) S. Venter
Ledebouria rupestris (llin der Merwe) S.
,retn~
comb. novo
Scilla rupestris Van der Merwe in Flower. PI. S. Afr. 21: t.828 (1941).
Type: Transvaal, near Mac Mac and Sabie, Wln der Merwe 1586 (PRE!, holo.).
Plants mostly gregarious, sometimes solitary. Bulb hypogeal, 20 x 15 mm,
ovoid; dead bulb scales brown, membranous, apices attenuate, live bulb scales
fleshy, tightly arranged, without threads when torn, white inside. Leaves fully
developed at anthesis, 3 - 6, spreading, lanceolate to oblanceolate, 30 - 100 x 5
- 10 mm, without threads when torn, fleshy, surfaces immaculate, glossy green,
adaxial surface covered in rows of asperities, venation obscure; margins ciliate;
leaf base canaliculate, petiolate, petiole 10 - 30 mm long; apex acute.
Inflorescence solitary, lax, cylindric, 20 x 15 mm, flaccid, 8 - 10 -flowered,
longer than the leaves; scape terete at base, green, glabrous; rachis smooth, 50 60 mm long. Bracts fleshy, 1 x 0.25 - 0.50 mm, linear to bifurcate, pink to
purple with bracteoles. Pedicels spreading, 2 mm long, pink. Perianth 4 mm
long, stellate, tepals slightly recurved, equal, 4 x 1.0 - 1.5 mm, apex acute,
pink to pink with a green keel. Stamens spreading, 3.5 mm long, filament
maroon, epitepalous; anthers 0.5 mm long, violet. Ovary ovoid, 6 -lobed, 1 x
1.5 - 2.0 mm, lobes obtusely deltate, glabrous, apex tapering into the style.
Style 1.5 mm long, triangular, glabrous, purple above and white below; stigma
at same height than anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed,
asymmetrical, clavate, base tapering. Seed drop-shaped, 3 - 4 mm long, surface
strongly wrinkled, brown. (Figure 34).
L. rupestris is closely related to L. minima. The leaves and bulb scales lack
threads when torn, leaves are monochromatic, racemes lax with a smooth
rachis, bracts fleshy and the stellate flowers with acute tepals. Together with L.
lepida and L. leptophylla they form the subsection Stellatae.
L. rupestris is easily distinguished by its petiolate leaves with adaxial asperities.
UDEBOURJA IN SOlJfH AFRICA
,},r
~h ..'
rd-> raJ(
148
TAXONOMY
E
F
H
o
G
",
-._B
Figure 34. Illustration of L. rupestris (Van der Merwe) S. Venter. A, habit X 2;
B, lamina margin X 300; C, section of the peduncle X 10; D, section through
lamina X 5; E, bract with bracteole X 10; F, tepal with stamen X 10; G, tepal
apex X 20; H, flower X 10; I, ovary lateral view X 10; J, ovary dorsal view X
13,250.
10. All from retn~
UDEBOURlA IN SOlITH AFRICA
Map 14.
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G.P.·S. 13--0959
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150
TAXONOMY
Figure 35. A, holotype of L. rupestris (Van der Merwe) S. Venter (PRE); B,
habitat near Mac Mac Falls, eastern Transvaal. The vegetation consists of open
short Pteridium aquilinum - Parinari capensis - Digitaria eriantha montane
grassland; C, plants showing globose inflorescences with stellate flowers. Bar =
20 mm; D, SEM micrograph of the adaxial lamina surface. Bar = 100 J.£m; E,
SEM micrograph of the abaxial lamina surface. Bar = 100 J.£m; F, SEM
micrograph of a stomata. Bar
= 43 J.£m. C - F from W?nter 13,250.
LED£BOURIA IN SOlITH AFRICA
TAXONOMY
151
A
LEDEBOURIA
IN SOlITH AFRICA
152
TAXONOMY
Specific epithet etymology.
Describes the rock-dwelling habit of this species.
Flowering period
From September to November.
Distribution (Map 14).
Only known from Mac Mac near Sabie in the eastern Transvaal.
Habitat
The rocks in this area consist of Black Reef Quartzite and conglomorate of the
Wolkberg Group. Soil derived from these rocks is a shallow (5 - 120 mm),
medium grained (0.25 - 1.0 mm </», well drained sandy soil. L. rupestris grows
in peaty humusrich sandy soil pockets in the rock cracks with Selaginella dregei
(Presl) Hieron. covering the surfaces. These pockets are sheltered, receiving full
sunlight only at noon.
The vegetation is interspersed with massive boulders and is an open short
montane grassland flanked by closed evergreen low woodland (Figure 35B).
Variation
Only one specimen could be found that matches the type (lan der Merwe 1,586)
which is probably somewhat aberrant. The type specimen lacks the lanceolate
lamina and long petiole.
Specimens examined
TRANSVAAL - 2430 (Pilgrim's Rest): Mac Mac (-DD), lan der Merwe
1586 (PRE); Mac Mac Falls (-DD), Tknter 13,250 (UNIN).
L£D£BOURIA IN SOUTH AFRICA
153
TAXONOMY
Sectio Coriaceae S. Venter, sect. nov., foliis crassicoriaceis crassimarginatis;
inflorescentia erecta pauciflorifera.
Species typica: Ledebouria coriacea S. Venter.
Typus: Cape, Port Elizabeth, 'The Aloes' Railway Station, Venter
13,307 (PRE).
Plants 40 - 50 mm tall. Bulbs 15 - 20 mm wide. Leaves appressed to ground,
thickly leathery, lamina margin thickened. Inflorescence solitary, erect, few
flowered, rachis smooth.
Distribution and habitat.
Port Elizabeth area, in closed evergreen succulent shrubland.
The only section with thickly leathery leaves, thickened lamina margins and
few-flowered inflorescences.
11.
LEDEBOURIA CORIACEA S. 'Venter
Ledebouria coriacea S. Venter, sp.nov., ad L. cooperi (Hook.f) Jessop arcte
affinis sed foliis coriaceis marginibus incrassatis; inflorescentia erecta solitaria
et ovarii lobis ad apicem conspicue rotundatis prominentibus bene distinguenda.
Type: Cape, Port Elizabeth, 'The Aloes' Railway Station, S. Venter 13,307
(PRE!, holo.; NU!; UNIN!).
Plants gregarious. Bulb hypogeal, 30 - 40 x 15 - 20 mm, cylindrical to obovoid;
dead bulb scales brown, apices attenuate, live bulb scales fleshy, white, tightly
arranged, with threads when torn. Leaves fully developed at anthesis, 2 - 9,
spreading but mostly appressed, lanceolate, 30 - 50 x 8 - 15 mm, with sparse
threads when torn, leathery, surfaces dull, green with faint darker green
blotches, venation obscure; margins thickened; leafbase canaliculate; apex
acute. Inflorescence solitary, lax, oblong to cylindric, 30 - 40 x 15 - 20 mm,
erect, 15 - 20 -flowered, longer than the leaves; scape terete at base, green,
LEDEBOURJA IN SOIJfH AFRICA
154
TAXONOMY
.----------------------_._-----------------,
E
.;
F
G
c
H
8
Figure 36. Illustration of L. coriacea S. Venter. A, habit X 2; B, lamina margin
X 300; C, section through lamina X 5; D, lamina apex X 20; E, bract X 10; F,
flower X 10; G, tepal apex X 10; H, tepal with stamen X 10; I, ovary lateral
view X 10; J, ovary dorsal view X 10. All from lCnter 13,307.
UDEBOURIA IN SOlJTH AFRICA
Known distribution of L. coriacea S. Venter
Map 15.
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156
TAXONOMY
Figure 37. A, holotype of L. coriacea S. Venter (PRE); B, habitat at 'The
Aloes near Port Elizabeth. The vegetation consists of evergreen tall Euclea
I
undulata - Ficus burtt-davyi - Brachylaena floribunda shrubland with open
patches covered in Pteronia incana (Burm.) DC. L. coriacea occurs under the
Pteronia shrublets; C, SEM micrograph of the adaxial lamina surface. Bar =
100 J.!m; D, SEM micrograph of the abaxial lamina surface. Bar = 100 J.!m; E,
SEM micrograph of stomata. Bar = 43 J.!m. A and C - E from lenter 13,307.
LED£BOURIA IN SOlTfH AFRICA
157
TAXONOMY
LEDEBOURlA IN SOUfH AFRICA
158
TAXONOMY
glabrous; rachis smooth, 40 - 50 mm long. Bracts fleshy, 0.75 X 0.75 mm,
deltoid, green with bracteoles. Pedicels cernuous, 5 mm long, deltate, speckled
pink soon turning green. Perianth 5 mm long, tepals recurved, equal, oblong, 5
x 1.5 mm, apex acute, slightly cucullate, green suffused purple. Stamens erect,
3 mm long, upper part of filament purple, lower part white, base slightly
flattened, epitepalous; anthers 1 mm long, violet. Ovary ovoid, 6 -lobed, 1.25 x
2 mm, lobes deltate, apex shoulders raised. Style + 3.5 mm long, triangular,
glabrous, purple; stigma above anthers; stipe 0.5 x 0.5 mm. Capsule threelobed, symmetrical, globose; base truncate. Seed drop-shaped, 2 mm long,
surface strongly wrinkled, brown. (Figure 36).
L. coriacea is not closely allied to other Ledebouria species.
Specific epithet etymology.
Alluding to the leathery leaves.
Flowering period
From October to December.
Distribution (Map 15).
Known only from' The Aloes' railway station near Port Elizabeth where it is
rare.
Habitat
'The Aloes' is situated on alluvial sand and fine gravel underlain by calcrete
(SACS 1980). All the populations grow in brown gritty sand, sometimes
interspersed with large pieces of calcrete.
Most of the plants occur in light shade provided by Pteronia incana (Burm.)
DC. (Figure 37B).
LEDEBOURIA IN SOlrrH AFRICA
159
TAXONOMY
Population structure
Five populations were found in a 5 km radius of 'The Aloes I . The smallest
population consisted of 11 adult plants, and the largest population numbered 158
individuals. Approximately 95 % of the populations consist of closely packed
adult plants with juveniles on the outer perimeter. Only a few seedlings in the
two-leaf stage were found. Even at this stage of development the leaves are
markedly leathery compared with seedlings of other species.
Variation
Plants growing in full sun tend to be more prominently marked on the adaxial
surface of the leaves.
Specimens examined
CAPE. - 3325 (Port Elizabeth): Port Elizabeth, 'The Aloes' Station (DC), lknter 13,307 (PRE, NU, UNIN).
LEDEBOURJA IN SOlJfH AFRICA
160
TAXONOMY
Sectio Magnibulbae S. Venter, sect. nov., bulbis maturis 60 - 150 mm diam.;
inflorescentia floribus plus quam octoginta; pedicellis 8 - 15 mm longis.
Species typica: Ledebouria zebrina (Bak.) S. Venter.
Lecto-typus: Transvaal, Near Barberton, Galpin 1184 (GRA).
Species: L. floribunda (Bak.) Jessop, L. hypoxidioides (Schonl.) Jessop, L.
revoluta (Lf) Jessop and L. zebrina (Bak.) S. Venter.
Plants solitary, 0.25 - 0.7 m tall. Bulb 60 - 150 mm wide, torn bulb scales
producing threads. Leaves 4 - 10, pulling threads when torn. Inflorescence 80 350 -flowered; rachis ridged; raceme cylindric, 20 - 50 mm wide; pedicel 8 - 15
mm long. Perianth 5 - 7 mm long.
Distribution and habitat
From the southern Cape to northern Transvaal. Coastal shrubland to montane
grassland.
12. LEDEBOURIA FLORIBUNDA (Bak.) Jessop
Ledebouria floribunda (Bak.) Jessop in Jl S. Afr. Bot. 36(4): 251 (1970).
Scilla floribunda Bak. in Saund. Ref. Bot. 3: t.188 (1870).
Type: Cap. b. Spei, Cooper s.n. (K!, holo.; PRE!, photo.).
Scilla pendula Bak. in Saund. Ref. Bot. 3 Appendix: 14 (1870).
Type: Cape of Good Hope, Burchell s.n. (K!, holo.; BOLl, drawing).
Scilla princeps Bak. in Saund. Ref. Bot. 3: t.189 (1870).
Type: Cape of Good Hope, Cooper s.n. (K!, holo.; PRE!, photo.).
Scilla polyantha Bak. in Gardnr's Chron. 9: 104 (1878).
Type: Natal, York, Bull s.n. (K!, holo.; PRE!, photo.).
LEDEBOURlA IN SOUTH AFRICA
161
TAXONOMY
Scilla tricolor Bale. in Gardnr's Chron. 14: 230 (1880).
Type: Cape, Port Elizabeth, Elwes s.n. (K!, painting; BOLl, copy of painting;
PRE!, photo. of painting).
Scilla subsecunda Bale. in Gardnr's Chron. 16: 38 (1881).
Type: Cape, Eastern Districts, Bowker 218 (K!, holo.; BOLl, drawing; PRE!,
photo. ).
Scilla lauta N.E. Br. in Kew Bull. :299 (1921).
Type: Transvaal, Pietersburg Div., The Downs, Rogers 23,990 (K!, holo.;
BOLl, drawing; PRE!, photo.).
Plants solitary. Bulb hypogeal, 60 - 100 x 25 - 35 mm, ovoid; dead bulb scales
brown, apices attenuate, live bulb scales fleshy, tightly arranged, lacking threads
when torn, purplish inside. Leaves fully developed at anthesis, 4 - 6, spreading,
lanceolate, 200 - 300 x 40 - 50 mm, with threads when torn, fleshy, glossy,
with spots and blotches, green and basally purple, venation obscure; margins
smooth; leaf base flat to shallowly canaliculate; apex acute. Inflorescences 1 3, dense, cylindric, 150 - 200 x 30 - 50 mm, erect to flaccid, 60 - 100
-flowered, shorter than or as long as the leaves; scape terete at base, green,
glabrous; rachis ridged, 150 - 225 mm long. Bracts membranous, 1.5 x 0.5
mm, linear-lanceolate, pink to purple with bracteoles sometimes present.
Pedicels spreading to horizontal, 12.5 - 15.6 mm long, pink. Perianth 7 - 9 mm
long, stellate, tepals equal, oblong, 7 - 9 x 3 mm, apex obtuse, cucullate, green
to pink with a green keel. Stamens erect, 5 - 6 mm long, filaments purple
above, white below, epitepalous; anthers 0.75 mm long, yellow. Ovary
ellipsoidal, 6 -lobed, 0.75 x 1.5 mm, lobes widely trullate, apex tapering into
style, basal lobes present. Style 6 mm long, triangular, glabrous, purple; stigma
equal height to anthers; stipe 0.25 x 0.5 mm. Capsule three-lobed, symmetrical,
clavate; base truncate. Seed drop-shaped, 4 - 5 mm long, strongly wrinkled,
brown. (Figure 38).
L. floribunda can be confused with L. revoluta but differs in the bulb scales
having threads when torn, tepals 2 - 4 mm longer, stigma and stamens equal
height, ovary widely trullate not narrowly transversely elliptic and the seed 3.0 3.5 mm longer.
LEDEBOURlA IN SOurH AFRICA
162
TAXONOMY
~."
~
c
". " ': ,;
f3/
::.
\
:
.
G
~
.:
.
.'
"
.
;"
J
B
A
Figure 38. Illustration of L. floribunda (Bak.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, lamina apex X 5; D, bract with bracteole X 10; E, section
through lamina Xl; F, flower X 10; G, tepal with stamen X 10; H, tepal apex
X 20; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from Venter
13,475.
LEDEBOURlA IN SOUTH AFRICA
Map 16.
14°
12°
P'\
Known distribution of L. floribunda (Bak.) Jessop
~
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164
TAXONOMY
Figure 39. A, type of L. floribunda (Bak.) Jessop (K); B, habitat at Thabina
Falls in the Wolkberg Wilderness Area near Tzaneen. The vegetation consists of
evergreen low Cliffortia linearifolia - Agapanthus inapertus subsp. pendulus -
Helichrysum splendidum shrubland in a seepage area next to a waterfall; C,
SEM micrograph of the adaxial lamina surface. Bar = 50 ",m; D, SEM
micrograph of the abaxial lamina surface. Bar = 50 ",m; E, SEM micrograph of
stomata. Bar = 50 ",m. C - E from 'knter 13,475.
LEDEBOURlA IN SOlJfH AFRICA
TAXONOMY
165
LEDEBOURlA IN SOlTrH AFRICA
166
TAXONOMY
Specific epithet etymology.
Means flowering profusely.
Flowering period
From October to January
Distribution (Map 16).
Restricted to the moist eastern parts of South Africa. Mostly associated with
mountainous areas.
Habitat
It is clear from herbarium material, literature, and populations visited in the
veld, that L. floribunda occurs mostly on quartzites and sandstones. In the
Nelspruit area, L. floribunda grows on the Kaap Valley Granites (SACS 1980).
L. floribunda grows in humus-rich soils in shade of scrub and montane forest.
Most of the specimens collected were found growing in moist situations (Figure
39B).
Population structure
Plants are either closely packed Clknter 13,4(0) or scattered. Most of the plants
seen in the veld were adult plants. It appears that on steep slopes and cliffs
where the plants grow, seed is washed away from the adult plants.
LEDEBOURIA IN SOUTH AFRICA
167
TAXONOMY
Variation
The leaves tend to be longer and narrower in the Cape and Natal than in any
other area. Intermediate forms occur in Swaziland. In some populations the
purple markings on the leaves are distinctly raised CW!nter 13,400).
Historical background
Burchell (1822) collected the type material of Scilla pendula Bak. around
Kommadagga in the Somerset Division. These bulbs were cultivated by Burchell
at his father's Fulham Nursery (Hutchinson 1946, Gunn and Codd 1980). He
completed a description of this plant naming it Drimia pendula but it was never
published. This description is on the type sheet at Kew. His main plant
collection was presented to Kew by Miss Burchell, presumably his daughter.
There is no note as to the name of the collector on the type sheet or a collector
mentioned in the type description of Scilla polyantha. Mr. Bull in Great Britain
cultivated bulbs he received from York, Natal and flowered these plants in
December 1877. These plants were forwarded to Baker at Kew.
Specimens examined
VENDA. - 2230 (Messina): Dzamba (-CD), WIn Ryk & Theron 4683
(PRU).
TRANSVAAL. - 2230 (Messina): Entabeni State Forest (-CC), Venter
13,400 (UNIN). -2430 (Pilgrim's Rest): The Downs (-AA), Moss & Rigers 291
(J); Rogers 23,990 (K); Ohrigstad, Branddraai, Road (-DA), Young A621b
(PRE); Klaserie (-DB), W!nter 12,708. (UNIN); Pilgrim's Rest (-DD), Rogers
14,955 (J). -2531 (Komatipoort): Komatipoort (-BD), Moss & Rogers 546 0);
Nelspruit, farm Rhenosterkop (-CA), Ondersta1l1314 (PRE). -2630 (Carolina):
Carolina (-AA), Moss & Rogers 1376 (J).
LEDEBOURIA IN SOlJfH AFRICA
168
TAXONOMY
SWAZILAND. - 2631 (Mbabane): Ukutula (-AC) , Compton 24,489
(NBG); Manzini, Bu1unga Poort (-DA), Karsten s.n. (NBG).
LESOTHO. - 2927 (Maseru): Maseru (-BB), Gormley & Barber 19
(PRE).
NATAL. - 2730 (Vryheid): Utrecht, 13 km on Paulpietersburg road (AA), Buthelezi 227 (NH). -2731 (Louwsburg): Itala Nature Reserve (-AC) ,
Brown & Shapiro 140 (PRE); Gwaliweni (-BD), Wm der Merwe 2719 (PRE);
Louwsburg,
± 21
km on road to Nongoma (-CB), Schrire 1116 (NH). -2732
(Ubombo): Ingwavuma (-AA), Wells 2229 (PRE). -2828 (Bethlehem): Mountaux-Sources (-DD), Uln der Merwe 2614 (PRE). -2831 (Nkandla): Nkandhla (CA), Jessop 1062 (GRA); Eshowe (-CD), Lawn 1162 (NH); Mtunzini, Ubisana
Valley (-DC), lenter 1228 (BLFU); Ngoya, lenter 602 (NH).
CAPE. - 3227 (Stutterheim): Pirie (-CC), Sim 619 (NU), Sim 621 (NU);
King Williamstown (-CD), Sim 620 (NU); Komgha (-DB), Flanagan 1112
(BOL). -3326 (Grahamstown): Grahamstown, Bloukrantz (-BC), Uln Jaarsveld
9088 (NBG).
Without precise locality.
- Cap B. Spei, Cooper s.n. (K, PRE); Cape of Good Hope, Cooper s.n. (K,
PRE); Burchell s.n. (BOL, K); Cape, Eastern Districts, Bowker 218 (BOL, K,
PRE).
LEDEBOURlA IN SOlITH AFRICA
169
TAXONOMY
13. LEDEBOURIA HYPOXIDIOIDES (Schonl.) Jessop
Ledebouria hypoxidioides (Schonl.) Jessop in 11 S. Afr. Bot. 36(4): 263
(1970).
Scilla hypoxidioides Schonl. in Rec. Albany Mus. 1: 48 (1903).
Type: Cape, Grahamstown, Daly & Sole 435 (GRA!, holo.; BOLl; SAM!; Z!).
Plants solitary. Bulb hypogeal, 80 - 100 X 40 - 60 mm, ovoid, dead bulb scales
purplish-brown, apices attenuate, live bulb scales fleshy, tightly arranged, with
copious threads when tom, white inside. Leaves fully developed at anthesis, 4 6, spreading, ovate-lanceolate, 100 - 140 x 20 - 40 mm, with threads when tom,
fleshy, pilose, venation obscure, dull green with faint darker green blotches
above; base canaliculate; apex acute. Inflorescences 1 - 2, flaccid, dense, 30 120 -flowered, longer than leaves. Peduncle glabrous, compressed at base,
green spotted purple, 50 - 120 mm long; rachis ridged. Raceme dense,
cylindric; 40 - 120 x 20 - 30 mm. Bracts with bracteoles; fleshy, 0.5 x 0.25,
lanceolate, white to purple. Pedicels spreading; 12 - 15 mm long, green.
Perianth 5 mm long; tepals recurved, equal, lanceolate, 5 x 2 mm, green tinted
pink sometimes with a green keel; apex curving sharply upwards, cucullate.
Stamens erect, 4 - 5 mm long; filaments pink, base slightly flattened,
epitepalous; anthers 1 mm long, pale yellow. Ovary ovoid, 6 -lobed, 1.5 x 3
mm, expanded lobes obtusely deltate; apex shoulders raised. Style 5 mm long,
triangular, glabrous, purple; stigma higher than anthers; stipe 0.5 x 0.25 mm.
Capsule symmetrical, globose, 3 -lobed; base truncate. Seed 4 mm long, dropshaped, surface strongly wrinkled, reddish-brown. (Figure 40).
L. hypoxidioides cannot be confused with any other Ledebouria in being the
only species with pilose leaves (Figure 41F & G).
Specific epithet etymology.
Describe the leaves likened to that of the genus Hypoxis L. (Hypoxidaceae).
LEDEBOURlA IN SOUTH AFRICA
170
TAXONOMY
o
E
F
G
A
H
Figure 40. Illustration of L. hypoxidioides (Schonl.) Jessop. A, habit X 0.25; B,
lamina margin X 110; C, section through lamina X 2; D, bract X 10; E, tepal
with stamen X 10; F, flower X 8; G, tepal apex X 20; H, ovary lateral view X
10; I, ovary dorsal view X 10. All from l.f!nter 13,311.
LEDEBOURlA IN sourH AFRICA
Map 17.
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172
TAXONOMY
Figure 41. A, holotype of L. hypoxidioides (Sch6nl.) Jessop (GRA); B, habitat
at Grahamstown. The vegetation consists of False Fynbos with Passerina rigida
- Ficus burtt-davyi - Euphorbia tetragona shrubveld; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of stomata; F, plants from the type locality. Note the hairy
leaves. Bar
=
10 mm; G, SEM micrograph of the leaf surface to show the
hairs. C - G from l-enter 13,311.
LEDEBOURIA IN SOIJfH AFRICA
173
TAXONOMY
u:DEBOURlA IN SOlITH AFRICA
174
TAXONOMY
Flowering period
From October to December.
Distribution (Map 17).
L. hypoxidioides is known only from the hills around Grahamstown.
Habitat
L. hypoxidioides occurs mainly on Dwyka Tillite of the Dwyka Formation of the
Karoo Sequence (SACS 1980). Soil derived from tillite is a brown loamy soil,
with a fine grained structure, usually 200 - 2000 mm deep. On the northwestern side of Grahamstown, it occurs on the Dirkskraal Sandstone Formation,
Witteberg Group of the Cape Supergroup (SACS 1980). Soils derived from this
sandstone are grey-brown gritty sandy loam, 10 - 250 mm deep.
Plants of L. hypoxidioides occur in open grassy areas between trees and shrubs
(Figure 41B). Although it prefers to grow in short grass in the sandstone areas,
areas with long grass are favoured on tillite.
L. hypoxidioides is a relatively rare plant known from only a few scattered
populations.
Specimens examined
CAPE. - 3326 (Grahamstown): Grahamstown (-AC), Van der Merwe
2138 (PRE); Bayliss 3110 (STE); Grahamstown, behind Fort England (-AC),
Daly & Sole 435 (BOL, GRA, SAM); Venter 13,311 (UNIN); Grahamstown,
Settlers Hill (-AC), Venter 13,424 (UNIN).
LEDEBOURIA IN SOtlTH AFRICA
175
TAXONOMY
14. LEDEBOURIA REVOLUTA (Lj:) Jessop
Ledebouria revoluta (Lj:) Jessop in Jl S. Afr. Bot. 36(4): 255 (1970).
Hyacinthus revolutus Lj: in SuppI. Plant.: 204 (1781).
Type: Cape, Cap. bonae Spei, Thunberg s.n. (UPS, holo.; BOL & PRE1, on
Herb. Thunb. Microfiche no. 8508).
Lachenalia lanceaejolia Jacq. in !cones PI. Rar. 2: t.402 (1794).
Iconotype: Jacquin in Icones PI. Rar. 2: t.402 (1794).
Phalangium revolutum (Lj:) Pers. in Syn. PI. 1: 367 (1805).
Type: Cap. bonae Spei, Thunberg s.n. (UPS, holo.; BOL & PRE1, on Herb.
Thunb. Microfiche no. 8508).
Lachenalia lanceaejolia Sims var. maculata Tratt. in Archiv der Gawachskunde
2: 132, t.168 (1814).
Iconotype: Tratt. in Archiv der Gewachskunde 2: t.168 (1814).
Scilla maculata Schrank in PI. Rar. Hort. Acad., Monac. 2: foI.100, t.100.
"Promontium Bonae Spei". (1820).
Iconotype: As for Drimia lanceaejolia Lodd. var. longipedunculata Schrader.
Drimia acuminata Lodd. in Bot. Cab.: t.1041 (1825).
Iconotype: Lodd. Bot. Cab. : t.1041. "Cape of Good Hope" (1825).
Drimia lanceaejolia Lodd. var. longipedunculata Schrader in Blumenb.: 30
(1827).
Iconotype: PI. Rar. Hort. Acad., Monac. 2.: fo1.100, t.l00 "Promontium
Bonae Spei" (1819).
Scilla revoluta (Lj:) Bak. in Saund. Ref. Bot. 3(app.): 6 (1870).
Type: Cape, Caledon Div., on Donker Hoek mountain, Burchell 7982 (K,
holo.).
l.£DEBOURIA IN SOlJrH AFRICA
176
TAXONOMY
Scilla spathulata Bale in Saund. Ref. Bot. 3: t.187 (1870).
Iconotype: Saund. Ref. Bot. 3: t.187, "Cape of Good Hope, Cooper s.n."
(1870).
Scilla livida Bale in Gdnrs' Chron. 20: 166 (1883).
Type: Cape of Good Hope, Hort. F. Horsman & Co." (K!, holo.; PRE!,
photo.; BOL!, drawing).
Scilla polyantha Bale var. angustifolia Bale in Flora Cap. 6: 488 (1896).
Type: Transvaal, Saddleback Mountain, Barberton, Galpin 1096 (PRE!, holo.).
Scilla moschata Schonl. in Rec. Albany Mus. 3: 60 (1914).
Type: Cape, Stutterheim, Rogers 12,786 (GRA!, holo.; PRE!, photo.).
Scilla carnosula Van der Merwe in Flower. PI. S. Afr. 24: t.958 (1944).
Type: Natal, near Greytown, WIn der Merwe 2592 (PRE!, holo.).
Plant solitary. Bulb hypogeal, 60 - 80 x 40 - 60 mm, ovoid; dead bulb scales
brown to purplish brown, membranous, apices attenuate, live bulb scales
membranous, live bulb scales tightly arranged, with threads when torn, white to
purple inside. Leaves fully developed at anthesis, 5 - 10, spreading, lanceolate,
80 - 130 x 30 - 90 mm, with threads when torn, fleshy, adaxial surface green
with darker green to purple spots and blotches, abaxial surface green or
variously marked purple, venation obscure; margin smooth; leaf base
canaliculate; apex acute. Inflorescences 4 - 10, dense, cylindrical, 70 - 120 x
30 - 40 mm, flaccid, 30 - 70 -flowered, longer than the leaves; scape terete at
base, green spotted purple, glabrous; rachis ridged, 90 - 110 mm long. Bracts
membranous, 1 x 0.5 - 1.0 mm, lanceolate to linear, white to purple with
bracteoles mostly present. Pedicels spreading, 10 - 13 mm long, pink to white
speckled pink. Perianth 5 mm long, tepals recurved, equal, oblong, 5 x 3 mm,
apex obtuse and thinly cucullate, green fused pink to dark pink with a green
keel. Stamens erect, 5 mm long, filaments maroon, base slightly swollen,
epitepalous; anthers 1 mm long, pale yellow. Ovary ellipsoidal, 6 -lobed, 0.75 1.0 x 1.75 - 2.0 mm, lobes narrowly transversely elliptic, distallobes present,
LEDEBOURJA IN SOlITH AFRICA
177
TAXONOMY
H
8
Figure 42. Illustration of L. revoluta (L.f.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina X 2; D, apex of tepal X 20; E, bract
with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H, ovary
lateral view X 10; I, ovary dorsal view X 10. All from retn~
13,430.
LEDEBOURlA IN SOUfH AFRICA
Map 18.
Known distribution of L. revoluta (L.f.) Jessop
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179
TAXONOMY
Figure 43. A, type of L. revoluta (L.f) Jessop under Lachenalia lanceaefolia
Jacq. in Icones Plantarum Rariorum 2: t.402 (1794); B, SEM micrograph of the
adaxial lamina surface. Bar = 100 Jlm; C, habitat near Thabazimbi in the
Waterberg, north-western Transvaal. The vegetation consists of closed
deciduous low Albizia tanganyicensis var. tanganyicensis - Croton gratissimus
var. subgratissimus - Myriothamnus jlabellijolia woodland; D, SEM micrograph
of the abaxial lamina surface. Bar
Bar
= 100 Jlm; E, SEM micrograph of stomata.
= 43 Jlm; F, plant of L. revoluta with thickly packed dry bulb scales for
protection against fire. Bar = 30 mm. Band D - E from If?nter 13,(XJ9.
LEDEBOURlA IN SOlITH AFRICA
180
TAXONOMY
LEDEBOURJA IN SmJfH AFRICA
181
TAXONOMY
apex forming shoulders. Style 6 mm long, triangular, glabrous, purple; stigma
above anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, clavate,
base truncate. Seed drop-shaped, 1.0 - 1.5 mm long, surface strongly wrinkled,
brown. (Figure 42).
L. revoluta is closely related to L. hypoxidioides. Together with L. zebrina
(Bak.) S. Venter and L. floribunda they form the section Magnibulbae. L.
hypoxidioides is easily separated from L. revoluta in that the leaves are covered
with long hairs.
Specific epithet etymology.
Describes the revolute perianth segments.
Flowering period
Flowering extends over most of the year with the plants in the Cape Province
flowering mostly from April to August, the rainy season. In the summer rainfall
areas L. revoluta flowers from October to February with a peak from October to
November.
Distribution (Map 18).
L. revoluta is recorded throughout the eastern half of South Africa except in
Lesotho. The species is concentrated in the Transvaal and Natal.
Habitat
L. revoluta is the most widespread Ledebouria species in South Africa
occurring on many soil types and in most of Acocks s veld types. It is absent
I
however from the Karoo, Namaqualand and the Richtersveld.
LEDEBOURIA IN SOtJrH AFRICA
182
TAXONOMY
Most of the specimens were collected in woodland vegetation (Figure 43C).
Plants occur mainly in full sun but some lightly shaded populations were
encountered.
Variation
Plants of L. revoluta growing in frequently burnt woodlands have dry bulb
scales which do not disentegrate but form a thick layer around the bulb and
presumably insulate it from heat damage (Figure 43F). Plants from moist
habitats tend to have very few dry bulb scales.
The leaves are polymorphic in shape, fleshiness and markings. Leaf shape
varies from linear-lanceolate (Wln der Merwe 2714) to broadly ovate (Venter
13,381). The 'albomarginata' form of L. revoluta has leaves appressed to the
ground similar to L. ovatifolia. The lamina margin is broad, hyaline and
distinctly whitish. These plants may prove to be a separate species. They are
tentatively included within L. revoluta in this revision. Flower colour varies
from green tinged with pink, to a dark pinkish purple. Flowers are sometimes
scented.
Historical background
Trattinick described Lachenalia lanceaejolia Sims var. maculata (1814) and
based it on Curtis's Botanical Magazine t.643. The illustration of L. lanceaejolia
var. maculata is a composite copy of Curtis's Botanical Magazine t.643 (plant)
and the flower, gynoecium and small drawing of a flower from Jacquin' s Icones
Plantarum Rariorum. Trattinick was the first to mention the differences between
Redoute's (1807) plant and the plant in Curtis's Botanical Magazine. C.A.
Smith compared type material at Kew and came to the conclusion that the type
of Scilla livida Bak. agrees in all essential details with the Redoute plate thus he
placed Trattinick's plant in synonomy under S. livida (Smith 1930).
Common names: Wave-leaved Hyacinth, Wave-leaved Cape Hyacinth (Aiton
1789), LanzenbHittrige Lachenalie (Willdenouw 1799), Spotted Copperas-leaved
Lachenalia (Ker-Gawler 1803).
LED£80URIA IN SOlTrH AFRICA
183
TAXONOMY
Specimens examined
VENDA. -2230 (Messina): Thate Vondo (-BD), Hemm 640 (J); Tate
& Theron 4838
Vondo (-CD), Steynberg 640 (PRE); Dzamba (-CD), WIn k~
(PRU); Mutale, Lokokoko (-DB), Hardy 6094 (PRE).
TRANSVAAL. -2328 (Baltimore): Vaalwater, farm Kliphoek (-CD),
Venter 11,198 (UNIN). -2329 (Pietersburg): Schyffontein (-BB), Van der
Merwe 1202 (PRE); Pietersburg (-CD), Venter 13,207 (UNIN); Solomondale (-
DC), Van der Merwe 1188 (PRE); Haenertsburg (-DD), Pott 4737 (PRE);
Venter 13,191 (UNIN). -2330 (Tzaneen); Duiwelskloof (-CA), Venter 13,371
(UNIN); George's Valley (-CC), Venter 13,333 (UNIN); Letsitele Valley (CD), Van der Merwe 1173 (PRE). -2427 (Thabazimbi): Thabazimbi, farm
Groothoek (-BC), Raal1251 (LYD); Thabazimbi (-CB), Jourdan s.n. sub NBG
227/39 (NBG). -2428 (Nylstroom): Sterkstroom (-AB), Van der Merwe 1761
(PRE); Nylstroom (-CB), Strydom 1802 (PRE); Naboomspruit (-DA), Mauve
4315 (PRE); farm Mosdene (-DA), Galpin M363 (PRE). -2429 (Zebediela):
Potgietersrust (-AA), van der Merwe 1210 (PRE); Pietersburg, farm Vrederust
(-AB), Van der Merwe 1959 (PRE). -2430 (Pilgrim's Rest): Wolkberg
Wilderness Area (-AA), Venter 11,073 (UNIN); Dublin Mine (-AA), Fourie
593 (PRE); Lekgalameetsi Nature Reserve (-AA), Stalmans 1439 (PRE,
UNIN); Kromellenboog Mine (-AD), Venter 13,203 (UNIN); Mariepskop (DB), WIn der SchYff 6381 (PRE, PRU); Mount Sheba Nature Reserve (-DC),
Onderstall 1384 (PRE); Lisbon Falls (-DD), Louw 2365 (STE). -2431
(Acornhoek): Hoedspruit (-AC), van der Merwe 2005 (PRE); Manyeleti Game
Reserve (-DA), Bredenkamp 1349 (PRE); Leeupan (-DD), WIn der SchYff 4069
(BOL). -2527 (Rustenburg): Brits, farm Welgevonden (-AD), Mogg 14,505
(PRE); Crocodile River (-DA), Van der Merwe 1833 (PRE); Hartebeespoort
Dam (-DD), Van der Merwe 2173 (PRE). -2528 (Pretoria): Pretoria (-CA),
Theron 175 (PRE); Meyerspark (-CA), Theron 175 (PRU); Jonker 25 (PRE);
Trigaardts Poort (-CA), van der Merwe s.n. sub PRE 35304 (PRE); Fountains
(-CA), Repton 1 (PRE); van der Merwe 2002 (PRE); Klapperkop (-CA), Smith
601 (PRE); Roodeplaat Dam Nature Reserve (-CB), Van Rooyen 3088 (PRU);
WIn Rooyen 2576 (PRU); Rietvlei (-CD), Codd 1755 (PRE). -2529 (Witbank):
LEDEBOURIA IN SOlJTH AFRICA
184
TAXONOMY
Loskop Dam Nature Reserve (-AD), Mogg 30,723 (J); Loskop, farm
Donkerhoek (-AD), Theron 1894 (PRE, PRU); Tautesberg, Diepkloof (-BA),
WIn der Merwe 2226 (PRE); Middelburg (-CB), du Plessis 395 (PRU); Young
A15 (PRE); Middelburg, farm Soetvelden (-CD), WIn der Merwe 2197 (PRE);
Belfast, farm Wonderfontein (-DD), Van der Merwe 1575 (PRE). -2530
(Lydenburg): Steenkampsberg, Boschhoek (-AB), Young A438 (PRE); Nelspruit
(-BD), Van Jaarsveld 20 (NBG); Van Jaarsveld 35 (NBG); Van der Merwe
1791 (PRE); Brondal (-BD), WIn der Merwe 1809 (PRE); Rosehaugh (-BD),
Sim s.n. sub BOL 3640/14 (BOL); Belfast, farm Schoongesicht (-CA), lbn der
Merwe 1239 (PRE); Witkloof (-CC), Van der Merwe Se.3 (PRE);
Coetzeestroom (-DA), WIn der Merwe 1647 (PRE); Berlin Forest Station (-DA),
Fourie 505 (PRE); Kaapsche Hoop (-DB), Van der Merwe 1527 (PRE). -2531
(Komatipoort): Mestelspruit (-AA), Van der SehYff 3906 (PRE); Skipberg (AB), Van der Sehyff 1127 (KNP); Malelane (-AD), Codd 5510 (PRE); Impala
Station (-BC), Kluge 2597 (PRE); Kaapmuiden (-CB), Rogers 24,661 (PRE);
Malelane (-CB), Codd 5510 (PRE); Barberton (-CC), Van der Merwe 1818
(PRE); Galpin 508b (PRE); Barberton, farm Oosterbeek (-CC), l-enter 13,378
(UNIN); Saddleback Range (-CC), Van der Merwe 1816 (PRE). -2627
(Potchefstroom): Ventersdorp, farm Goedgedacht (-AA), Sulton 629 (PUC);
Roodepoort, Ruimsig (-BB), Steel 50 (PRE); Potchefstroom (-CA), WIn der
1025 (PUC); Sasolburg (Westhuizen 431 (PUC); Dassiesrant (-CA), WIn k~
DD), Theron 694 (PRE). -2628 (Johannesburg): Kloof (-AA), Moss 10,812 (1);
Frankenwald (-AA), Gilliland s.n. sub J 25,410 (1); Edenvale (-AA), Venter
13,336 (UNIN); Klipriviersberg (-AC), Gilliland s.n. sub J 26,295 (J); Delmas
(-BA), WIn der Merwe 1041 (PRE); -2629 (Bethal): Ermelo, Nooitgedacht (DB), Balsinhas 2829 (PRE). -2630 (Carolina): Carolina (-AA), Moss & Rogers
1377 (J); Moss & Rogers 1299 (1); Maviriestad (-CA), Port 5190 (BOL);
Amersfoort, Kalkoenkranz (-CC), Turner 319 (PRE); Amsterdam (-DA), WIn
der Merwe 1092 (PRE); Piet Retief, farm Wolwenkop (-DC), Venter 13,365
(UNIN); Piet Retief (-DC), lbn der Merwe 1837 (PRE); Kemp Station (-DD),
Venter 13,362 (UNIN); Venter 13,363 (UNIN). -2730 (Vryheid): Piet Retief,
Hlangapies Mountain (-AA), WIn der Merwe 111 (PRE); Amsterdam (-AB),
Van der Merwe 1055 (PRE); Wakkerstroom, Watervalberge (-AC), Van der
Merwe 2059 (PRE); Wakkerstroom, farm Oshoek (-AD), Devenish 287 (PRE);
Piet Retief (-BB), Steyn s.n. sub PRE 13,397 (PRE).
LEDEBOURIA IN SOlITH AFRICA
185
TAXONOMY
SWAZILAND. -2631 (Mbabane): Malandela (-AB), Compton 32,483
(NBG); Mbabane (-AC) , Compton 23,796 (NBG); Compton 27,043 (NBG;
PRE); Ukutula (-AC), Compton 25,194 (NBG); Highveld (-AC), Bolus s.n. sub
BOL 12,407 (BOL); Dalriach (-AC) , Compton 28,478 (NBG); Black Umbeluzi
River (-AC), Compton 28,066 (PRE); Manzini (-AD), Karsten s.n. sub NBG
71,331 (NBG); Compton 31,077 (PRE); Mankaiana (-CA), Compton 29,062
(PRE); Compton 31,581 (PRE); Mhlambanyati (-DA), Dlamini s.n. sub PRE
703,374 (PRE).
ORANGE FREE STATE. -2828 (Bethlehem): Clarens (-CB), de Leeuw
s.n. (BOL). -2829 (Harrismith): Harrismith Botanical Garden (-AC), Jacobsl
2282 (PRE).
NATAL. -2632 (Bela Vista): Ndumu Game Reserve (-CD), Moll 4165
(DWEST); Tembe Elephant Park (-CD), Ward 1372 (NH). -2729 (Volksrust):
Newcastle (-DD), ltln der Merwe 2081 (PRE). -2730 (Vryheid): Utrecht, farm
Retirement (-AC), Hilliard & Burtt 18,553 (NU); Devenish 1597 (PRE);
Vryheid (-DD), Venter 13,360 (UNIN); ltln der Merwe 2402 (PRE); Zinguin
Mountain (-DB), ltln der Merwe 2745 (NU). -2731 (Louwsburg): Magut (DA), Van der Merwe 2714 (PRE). -2732 (Ubombo): Jozini area (-AC),
Vahrmeyer 1258 (PRE); Sordwana State Forest (-CD), Van ""yk 951 (NH);
Mkuzi (-CA), ltln der Merwe 2410 (PRE); Mkuzi Game Reserve (-CA), Wlrd
3515 (NH). -2828 (Bethlehem): Royal National Park (-DB), Hutchinson et al.
131 (NH). -2829 (Underberg): Oliviers Hoek Pass (-CA), Thode s.n. sub STE
3369 (STE); Estcourt, farm Heartsease (-CD), Du Toit 2 (PRE); Colenso (-DB),
Stirton 12,292 (UNIN); Draycott (-DC), Van der Merwe 2560 (PRE). -2830
(Dundee): Impati Mountain (-AA), Shirley 69 (NU); Weenen, Culvers (-CC),
Rogers 28,271 (GRA); Muden (-CD), West 1215 (PRE). -2831 (Nkandla):
Hlabisa, farm Palm Ridge (-BB), Harrison 283 (NH); Itala Nature Reserve (CA), McDonald 158 (NU); Eshowe (-CD), Lawn 968 (NH); Ngoye Forest (DC), Huntley 716 (NU); Ward 1624 (NH). -2832 (Mtubatuba): Hluhluwe Game
Reserve (-AA), Ward 3348 (NH); Ward 1878 (NR). -2929 (Underberg):
Estcourt (-BB),
ts~
1347 (PRE); Rele Rela (-DC), Strey 9203 (NH); Bulwer (-
DD), ltln der Merwe 2764 (PRE). -2930 (pietermaritzburg): Greytown (-BA),
Schrire 1592 (NH); RYlie 27,969 (NH, PRE); ltln der Merwe 2592 (PRE);
UDEBOURlA IN SOIJfH AFRICA
186
TAXONOMY
Pietermaritzburg (-CB), Carnegie s.n. sub NBG 358/31 (BOL, NBG); Allsopp
951 (NH); Scot! s.n. (NU); Douwes-Dekker 19 (NU); Duncaim (-CB), Wlrren
42 (PRE); Oribi (-CB), Lawson 615 (NH); Fox Hill (-CB), Van der Merwe
2555 (NU); Scottsville (-CB), Frankish 42 (NU); Thomville, Richmond (-CB),
Moll 3422 (NU); Thomville, Tala Farm (-CB), Moll 3422 (PRE); Drummond (DA), Hilden s.n. (BOL); Inanda (-DB), Strey 5337 (NH); Mid Il1ovo (-DC),
Thode s.n. sub STE 3385 (STE); Isipingo North (-DD), Ward 460 (DWEST);
Krantzkloof Nature Reserve (-DD), Palmer 26 (NH); Hayganh s.n. sub STE 83
(STE); Haygarth s.n. sub STE 85 (STE); Si1verglen Nature Reserve (-DD),
Ngwenya 414 (NH); Westville (-DD), Vttllsh 23 (NU); Westville, Atho1 Heights
(-DD), Feldman s.n. (NU). -3029 (Kokstad): Impendhle (-AC), Levet! 9 (NH);
Elandskop (-AC), Pascoe s.n. sub NBG 428/44 (NBG); Harding (-DB), lbn der
Merwe 2756 (NU). -3030 (Port Shepstone): Umkomaas (-BB), lbn der Merwe
2766 (NU); Archbell s.n. sub NBG 252/32 (NBG); Vernon Crookes Nature
Resrve (-BC), Balkwill & Cadman 2254 (1); Umgaye Flats (-BC), Rudatis 222
(STE); Oribi Gorge (-CA), Nicholson 1867 (PRE); Balkwill & Cron 187 (1);
Izotsha (-CB), Rump s.n. (NU); Izingolweni (-CC), WJod 162 (NU); Umzumbi
(-DA) , llin der Merwe 2534 (NU, PRE). -3130 (Port Edward): Port Edward (AA), Moss 19,143 (1); Strey 4934 (NH); O'Conner 40 (NU); Umtamvuna
Nature Reserve (-AA), Abbot! 1311 (NH, PRU); Abbot! 2131 (PRU); Abbott
1493 (NH).
TRANSKEI. -3027 (Lady Grey): Sterkspruit (-CB), Van der Merwe
1790(a) (PRE). -3128 (Umtata): Mahlahlani Forest Reserve (-BC), Perry 2431
(NBG). -3129 (Port St. Johns): Ntsubane Forest Reserve (-BC), Venter &
Vorster 33 (PRE); Mkambathi Nature Reserve (-BD), Shackleton 23 (KEI);
Smook 6113 (PRE); Bizana (-CB), Loughlin s.n. (BOL). -3228 (Butterworth):
Willowvale, Ngqaqini area (-AD), llin Eeden B420 (PRE); Dwessa Forest (BD), Linder 1210 (PRE).
CAPE. -3024 (De Aar): De Aar (-CB), Smook 3550 (PRE). -3118
(Vanrhynsdorp): Clanwilliam, Nardouw Kloof (-DC), Stokoe s. n. sub BOL
64,493 (BOL). -3221 (Merweville): Fraserburg, Layton (-BB), Shearing 750
(PRE). -3222 (Beaufort West): Beaufort West (-BC), de lager s.n. (BOL).
-3225 (Somerset East): Somerset east (-DA), Harris s.n. sub NBG 2605/29
LEDEBOURlA IN sourH AFRICA
187
TAXONOMY
(NBG); Boschberg (-DC), MacOwan 1840 (GRA). -3226 (Fort Beaufort):
Hoggsback Forestry Reserve (-DB), Dahlstrand 1834 (1). -3227 (Stutterheim):
Prospect (-CC), Flanagan 112 (BOL); King Williams Town (-CD), Barker s.n.
sub BOL 891/32 (BOL). -3228 (Butterworth): Butterworth (-CC), Taylor 3654
(NBG). -3319 (Worcester): Klein Drakenstein (-CC), du Plessis s.n. sub STE
19,664 (STE). -3320 (Montagu): Bonnievale (-CC), Compton et al. s.n. sub
NBG 558/41 (NBG); Swellendam, Warmwaterberg (-DD), Van Niekerk 564
(BOL). -3321 (Ladismith): Riversdale, Gouritz Bridge (-DC), Aco~ks
21,632
(PRE). -3322 (Oudshoorn): George (-CD), Bolus s.n. sub BOL 22,497 (BOL).
-3323 (Willowmore): Joubertina (-DD), Fourcade 2404 (BOL). -3324
(Steytlerville): Hankey, Zuurbron (-DD), Fourcade 3216 (BOL). -3325 (Port
Elizabeth): Port Elizabeth (-DC), Copeman s.n. sub PRE 35,301 (PRE); Cruden
377 (PRE); Zwartkops River (-DC), Ecklon & Zeyher s.n. (BOL); Redhouse (DC), Paterson 923 (BOL); Donkin Reserve (-DC), Troughton s.n. (GRA).
-3326 (Grahamstown): Albany, Mitford Park (-AA), Britton s.n. sub BOL
22,498 (BOL); Alicedale (-AC), Cruden 211 (GRA); Howison's Poort (-AD),
Hall 223 (NBG); Albany, Manley Flats (-BC), Leighton 2638 (BOL);
Grahamstown (-BC), Glass s.n. sub BOL 23,299 (BOL); Blaauwkrantz (-BC),
Sinclair 1052 (GRA); Fraser's Camp (-BD), Barker 6995 (NBG). -3419
(Caledon): Greyton (-BA), Van der Merwe 2139 (PRE). -3421 (Riversdale):
Riversdale (-AB), Muir 2708 (BOL). -3420 (Bredasdorp): De Hoop,
Hardevlakte (-AD), Van "yk 2148 (STE); Westfield (-BD), Forrester 312
(NBG); Bredasdorp Flats (-CA), Taylor 7680 (STE); Leighton s.n. sub BOL
50,841 (BOL). -3421 (Riversdale): Riversdale (-AB), Muir 3003 (BOL); Muir
1877 (BOL, PRE); Riversdale, farm Tolange (-AB), Bohnen 7284 (STE);
Stillbay (-AD), Bohnen 5763 (PRE); Albertinia (-BB), Hall 4680 (NBG). -3422
(Mossel Bay): Groot Brak (-AA), Bayer 2321 (NBG). -3423 (Knysna): Knysna
(-AA), Thesin s.n. sub BOL 275/13 (BOL); Duthie s.n. sub BOL 1197/15
(BOL). -3424 (Humansdorp): Jeffrey's Bay (-BB), Fourcade 3937 (BOL).
LEDEBOURIA IN SOlJfH AFRICA
188
TAXONOMY
15. LEDEBOURIA ZEBRINA (Bale) S. Venter
Ledebouria zebrina (Bak.) S. "lknter, comb. novo
Scilla zebrina Bale in Saund. Ref. Bot. 3: t.185 (1870).
Type: "Cape of Good Hope, Cooper s.n." Saund. Ref. Bot. 3: t.185 (1870).
Scilla microscypha Bale in Gdnr's Chron. 16: 102 (1881).
Type: Cape, eastern Provinces, Bowker s.n. (Kl, holo.; PRE1, photo.).
Scilla megaphylla Bale in Flora Cap. 6: 490 (1896).
Type: Transvaal, near Barberton, Galpin 1184 (GRA1, lecto.; NHl; PREl,
photo.; SAM!) (lecto. selected here - Art. 9.3, Greuter et al. 1988).
Scilla grandijolia Schonl. in Rec. Albany Mus. 3: 61 (1914).
Type: Transvaal, Tzaneen, Duiwelskloof, P.A. Rogers s.n. (GRAl, ho10.;
PRE1, photo.).
Plants solitary. Bulb hypogeal, 100 - 150 x 100 - 150 mm, obovoid; dead
bulb scales purplish brown, membranous, apices attenuate, live bulb scales tightly
arranged, with threads when torn, white inside. Leaves partly to fully developed
at anthesis, 4 - 6, spreading, lanceolate to oblong-lanceolate, 300 - 500 x 90 120 mm, with threads when torn, fleshy, dull green, sometimes with
longitudinal purple stripes or with large purple blotches, venation obscure;
margin smooth; leaf base flat to very shallowly canaliculate; apex acute.
Inflorescences 4 - 10, dense, cylindric, 150 - 200 x 30 - 37 mm, flaccid, 100 150 -flowered, longer than leaves; scape base winged to angled, green,
glabrous; rachis ridged, 225 - 310 mm long. Bracts membranous, 1.5 x 0.5
mm, linear-lanceolate, green with bracteoles below bracts. Pedicels spreading,
8.3 - 12.5 mm long, green. Perianth 4.2 - 7.7 mm long; tepals recurved, equal,
oblong, 4.2 - 7.7 x 1.5 mm, apex obtuse, light green with darker green keel.
Stamens erect, 4 mm long, green, epitepalous; anthers 1 mm long, pale yellow.
Ovary ovoid, 6 -lobed, 2.25 x 3 mm, lobes deltate, basal lobes present, apex
taper~ng
into style. Style 3 mm long, triangular, glabrous, white to light green;
stigma above anthers; stipe 0.25 x 0.25 mm. Capsule three-lobed, symmetrical,
clavate, base truncate. Seed drop-shaped, 4 mm long, surface strongly
wrinkled, black. (Figure 44).
LEDEBOURLA IN SOlJfH AFRICA
189
TAXONOMY
A
H
Figure 44. Illustration of L. zebrina (Bak.) S. Venter. A, habit X 0.25; B,
section through lamina X 0.5; C, lamina margin X 300; D, bract with bracteole
X 5; E, tepal with stamen X 5; F, apex of tepal X 10; G, flower X 5; H, ovary
lateral view X 10; I, ovary dorsal view X 10. All from lenter 13,395.
LEDEBOURJA IN sourH AFRICA
Map 19.
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191
TAXONOMY
Figure 45. A, type of L. zebrina (Bak.) S. Venter under Scilla zebrina Bak. in
Saunders Refugium Botanicum 3: t.185 (1870); B, plant with leaves 1000 mm
long showing growth habit. Bar = 60 mm; C, SEM micrograph of the adaxial
lamina surface; D, SEM micrograph of the abaxial lamina surface; E, SEM
micrograph of a stomatum; F, habitat near Barberton. The vegetation consists of
closed evergreen low Acacia sieberana var. woodii - Pavetta edentula - Schotia
brachypetala woodland; G, plant from Noodsberg near Greytown showing
the prominent purple zebra stripes on the leaves. Bar = 60 mm. B - E from
lenter 13,395 and G from Smith 115.
LEDEBOURlA IN SOUTH AFRICA
TAXONOMY
192
LEDEBOURlA IN SOlJfH AFRICA
TAXONOMY
193
L. zebrina is closely related to L. revoluta, L. hypoxidioides and L. floribunda.
Together they constitute the section Magnibulbae. L. zebrina is easily
distinguished from all other species by its size.
Specific epithet etymology.
Describes the purple, parallel longitudinal lines on the leaves.
Flowering period
From August to January with a peak from October to November.
Distribution (Map 19).
Occurs in eastern South Africa to Mooketsi in the north with an isolated locality
in the Magaliesberg.
Habitat
L. zebrina grows in shallow (10 - 120 mm deep), fine grained (0.025 - 0.25 mm
c/» to medium grained (0.25 - 1.0 mm c/», humus-rich sandy soils derived from
quartzites and conglomorates. They usually grow in moist, shaded conditions
often associated with forest vegetation.
In the Magaliesberg the L. zebrina populations grow in humus-rich black sandy
soil derived from Magaliesberg Quartzite. This soil is medium grained (0.25 _
0.5 mm c/» and up to 26 mm deep in places. Plants occur in dense shade in
kloof forest vegetation.
LEDEBOURIA IN SOIJfH AFRICA
194
TAXONOMY
Variation
Plants from Barberton are the largest. The leaves are either immaculate or with
dark green to purple blotches or striae (Figure 45G). Flower colour varies from
near white (Swaziland), to green suffused with pink in the northern Transvaal.
Specimens examined
TRANSVAAL. -2330 (Tzaneen): Duiwelskloof, Westfalia (-CA),
Scheepers 468 (UNIN); Tzaneen (-CC), Rogers 12,504 (1); Rogers 12,527 (1).
-2430 (Pilgrim's Rest): The Downs (-AA), Moss & Rogers 460 (1); Wolkberg,
farm Fertilis (-AA), Van der Merwe 2254 (PRE); Nelspruit (-CB), Van der
Merwe 1792 (PRE); Abel Erasmus Pass, farm California 228 KT (-DA), Raal et
al. 978 (LYD); Pilgrim's Rest, farm Vaalhoek (-DB), Rogers 20,974 (PRE);
Rogers 25,010 (BOL, J). -2431 (Acornhoek): Manyeleti Game Reserve (-DA),
Bredenkamp 1167 (PRU). -2530 (Lydenburg): Nelspruit (-BD), Rogers 16,208
(BOL); Buitendag 263 (NBG); Van Jaarsveld 107 (NBG). -2531 (Komatipoort):
4806 (PRE); Randspruit (-BB), Van der SchYff
Paben Outpost (-BB), Van k~
4022 (KNP); Nelspruit, farm Rhenosterkop (-CA), Onderstall 937 (PRE);
Baberton Townlands (-CC), Venter 13,395 (UNIN); Van der Merwe 1825
(PRE); Thorncrojt 2762 (PRE); Rogers 24,663 (PRE). -2627 (potchefstroom):
Sterkfontein Caves (-BB), Mogg 35,541 (J).
SWAZILAND. -2631 (Mbabane): Komati Old Ferry (-AA), Compton
29,820 (NBG); Mbabane (-AC), Martin s.n. (NBG); Manzini (-AD), Kemp 431
(PRE); Manzini, near Bremersdorp (-AD), Compton 28,105 (NBG); Stegi (BD), Compton 31,239 (NBG); Malboch s.n. (NBG); Mbuluzi, Mlawula Nature
Reserve (-BD), Culverwell 2003 (PRE). -2632 (Bela Vista): Umbuluzi Gorge,
Ndzindza Nature Reserve (-AA), Culverwe1l1256 (PRE).
NATAL. -2731 (Louwsburg): ltala Nature Reserve (-CB), Jordaan 2077
(NH); McDonald 419 (NU); McDonald 344 (NU); Ngotshe, Ngome area (-CD),
Hilliard & Burtt 8479 (NU). -2732 (Ubombo): Tembe Elephant Park (-AB),
Ward 2146 (NH). -2829 (Ladysmith): Ladysmith (-DB), Van der Merwe 2603
LEDEBOURIA IN SOUfH AFRICA
195
TAXONOMY
(PRE). -2831 (Nkandla): Hlabisa, Hluhluwe Valley (-BB), Scott-Smith & Wlrd
1 (NH); Hluhluwe Nature Reserve (-BB), Ward 1559 (NU); Eshowe (-CD),
Lawn 1105 (NH); Gerstner s.n. (NBG); Acocks 12,965 (PRE); Mtunzini,
Ubisana Valley (-DC), Venter 1228 (BLFU); Venter 566 (NH); Mtunzini,
"Twinstreams" (-DD), Hilliard & Burtt 6849 (NU). -2832 (Mtubatuba):
Hlabisa, Dukuduku Farm (-AC), Strey 5557 (NH); St. Lucia Bay (-AD), Henry
s.n. (NBG). -2930 (Pietermaritzburg): Richmond, Hela Hela (-CC), Strey 9234
(NH); Pietermaritzburg (-DB), lbn der Merwe 2094 (PRE); Strey 4360 (NH);
Bothas Hill (-DC), Wood 4,776 (NH). -2931 (Stanger): Darnall (-AD),
Lutchminarain 27 (DWEST); Stanger (-AD), Sewbarun 3 (DWEST); Durban,
Clover Estate (-CC), Nair 16 (DWEST). -3029 (Kokstad): Alfred, Weza (-DA),
Law s.n. (NU). -3130 (Port Edward): Umtamvuna Nature Reserve (-AA),
Abbott 2148 (NH).
CAPE. -3228 (Stutterheim): East London, Gonubie (-BB), Carter s.n.
(NBG).
LEDEBOURlA IN SOUTH AFRICA
196
TAXONOMY
Sectio Filiferae S. '\.enter, sect. nov., squamis longitudinaliter copiose fibrosis.
Species typica: Ledebouria luteola Jessop.
Typus: Transvaal, 6.5 miles south of Hammanskraal, Codd 5625 (PRE).
Species: L. luteola Jessop and L. ovatifolia (Bale) Jessop.
Plants 100 - 200 mm tall. Bulb 25 - 50 mm wide, torn bulb scales produce
copious threads. Leaves pulling threads when torn. Inflorescence a dense
raceme, 30 - 50 mm long; rachis ridged; bracts membranous, bracteoles
present. Capsule globose.
Distribution and habitat
From the eastern Cape to northern Transvaal. Coastal grassland and woodland
to montane grassland.
16. LEDEBOURIA LUTEOU Jessop
L. luteola Jessop in 11 S. Afr. Bot. 36(4): 260 (1970).
Type: Transvaal, 6.5 miles south of Hammanskraal' Codd 5625 (PRE!, holo.).
Plants solitary. Bulb hypogeal, 35 - 50
X
25 - 30 mm, obovoid; dead bulb scales
brown, membranous, apices attenuate, live bulb scales tightly arranged, with
copious threads when torn, white inside. Leaves fully developed at anthesis, 4 10, spreading, linear-lanceolate to lanceolate, 45 - 90 x 5 - 22 mm, with copious
threads when torn, leathery, with a dull lustre, purple spots and blotches with
purple cross bands at base, sometimes with dull purple to darker green blotches,
venation obscure; margin smooth, undulate at base; leaf base canaliculate; apex
acute. Inflorescences 1 - 2, dense, elliptic, 30 - 40 x 15 - 20 mm, flaccid, 30 60 -flowered, longer than leaves; scape base terete, green, glabrous; rachis
ridged, 50 - 70 mm long. Bracts membranous, 1.5 x 0.5 mm, bifurcate to
linear, grey to white, with bracteoles. Pedicels spreading, 4 mm long, pink.
Perianth 4 - 6 mm long, tepals suberect to recurved, equal, oblong, 4 - 6 x 3
mm, apex obtuse, cucullate, pink to purple with green keel. Stamens erect, 4 -
LEDEBOURlA IN SOlITH AFRICA
197
TAXONOMY
E
G
o
H
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Figure 46. Illustration of L. luteola Jessop. A, habit Xl; B, lamina margin X
300; C, apex of lamina X 20; D, section through lamina X 5; E, bract with
bracteole X 10; F, flower X 20; G, tepa! with stamen X 20; H, apex of tepa! X
10; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from Venter
13,458.
LED£BOURlA IN SOlITH AFRlCA
Map 20.
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Figure 47. A, holotype of L. luteola Jessop (PRE); B, habitat at Potchefstroom.
The vegetation consists of open deciduous low Acacia caffra - Aloe greatheadii
var. davyana - Themeda triandra woodland; C, plant with the previous years
leaves still on the plant. Bar = 10 mm; D, SEM micrograph of the adaxial
lamina surface; E, SEM micrograph of the abaxial lamina surface; F, SEM
micrograph of stomata. C - F from If!nter 13,217.
LEDEBOURIA IN SOlITH AFRICA
200
TAXONOMY
NA IK)."\IAL IIEllftAJllU ..
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LEDEBOURJA IN SmITH AFRICA
201
TAXONOMY
5 mm long, filaments maroon, epitepalous; anthers 0.75 mm long, pale violet.
Ovary ovoid, 6 -lobed, 3 x 4 mm, lobes obtusely deltate, basal lobes present,
apex tapering into style. Style 3 mm long, triangular, glabrous, purple; stigma
above anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, globose;
base truncate. Seed drop-shape, 4 mm long, surface strongly wrinkled, brown.
(Figure 46).
L. luteola is closely related to L. ovatifolia (Bak.) Jessop, together comprising
the section Filiferae. L. luteola differs in the apices of the bulb scales not at all
truncate, leaves spreading and linear-lanceolate to lanceolate.
Specific epithet etymology.
Referring to the colour of the live bulb scales.
Flowering period
From August to February with a peak from October to November.
Distribution (Map 20).
Most common in the southern half of the Transvaal and the whole Orange Free
State. A few localities are recorded in the extreme north-eastern Cape with a
single locality at Ladysmith in Natal.
Habitat
L. luteola occurs in woodland and grassland but is principally a Highveld
element (Figure 47B). Jessop (1970) mentions that L. luteola occurs in damp
areas. The opposite was found to be the case during the present study.
Some plants show signs of grazing by the Common Duiker (Sylvicapra grimmea
L.) (Smithers 1983) according to spoor and droppings found.
LEDEBOURIA IN SOlJfH AFRICA
202
TAXONOMY
Variation
The eastern Transvaal populations have more erect inflorescences with longer
and darker perianths.
Specimens examined
TRANSVAAL. - 2426 (Mochudi): Derdepoort, farm Buffelsdooring (DA), Chadwick 286 (PRE). -2427 (Thabazimbi): Rooiberg (-DC), Van der
Merwe s.n. (PRE). -2430 (Pilgrim's Rest): Graskop, Lisbon Falls (-DD), ~nter
13,241 (UNIN). -2524 (Vergelee): Molopo Nature Reserve (-DD), Peeters et
al. 222 (PRE). -2527 (Rustenburg): Rustenburg, Paul Kruger (-DB), Van der
Merwe 1127 (PRE); Rustenburg, farm Uitkomst 499 JQ (-DD), Coetzee 210
(PRE); Zwartkops (-DD), Moss 19,059 (J). -2528 (pretoria): Rust der Winter (AB), Pole-Evans 3892 (PRE); Herzogville (-BA), Greyling 4 (PRU); Pretoria,
Van Riebeeck Nature Reserve (-BC), Kok 220 (PRE); Pretoria (-CA), Moss
10,810 (J); Bon Accord Dam (-CA), Wm der Merwe 1988 (PRE); Irene (-CC),
Leendertz 925 (BOL). -2529 (Witbank): Buffelshoek (-AC) , lfln der Merwe
2018 (PRE); -2530 (Lydenburg): Sabie (-BB), Van der Merwe s.n. (PRE);
Nelspruit (-BD), Buitendag 639 (NBG); Belfast (-CA), Van der Merwe 1938
(PRE); Van der Merwe 1227 (PRE); Starvation Creek Nature Reserve (-DA),
Kluge 1351 (PRE). -2531 (Komatipoort): Kruger National Park, Numbi (-AA),
WIn der Schyff 733 (KNP). -2626 (Klerksdorp): Ventersdorp, farm Somerville (-
BD), Louw 3884 (PUC); Louw 4232 (PUC); Klerksdorp (-DC), Strydom 2259
(PRE); Ventersdorp, farm Goedgedacht (-DD), Louw 1982 (PUC); Sutton 631
(PRE). -2627 (Potchefstroom): Rysmierbult (-AC), Ubbink 730 (PUC);
Carltonville, Abe Bailey Nature Reserve (-AD), WIn RYk 312 (PUC); lfln RYk
271 (PUC); Witpoortjie (-BB), Gilliland s.n. (J); Lloyd s.n. sub J 26,629 (1);
Moss 4133 (1); Potchefstroom (-CA), Bredenkamp & Smith 94 (PUC); Venter
13,226 (UNIN); Dassiesrant (-CA), WIn der Vresthuizen 941 (PUC); Boskop (CA), Louw 481 (PUC); Klington (-CA), Goossens 1660 (PUC). -2628
(Johannesburg): Johannesburg (-AA), Moss 19,048 (J); Klipriviersberg (-AA),
Lloyd s.n. (1); Rietfontein (-AA), Moss 10,807 (1); Bedford (-AA), Gilliland
s.n. (J); Zoo Koppies (-AA), Rogers 19,605 (J); Melville (-AA), Gilliland S.n.
LEDEBOURIA IN SOIITH AFRICA
203
TAXONOMY
(1); MacNae 1166 (1); Suikerbosrand (-CA), Bredenkamp 1000 (PRU);
Bredenkamp 27 (PRU). -2630 (Carolina): Carolina (-AA), Gilliland s.n. (1).
-2725 (Bloemhof): Wolmaranstad, farm Boskuil (-BD), Sutton 465 (PUC).
ORANGE FREE STATE. - 2725 (Bloemhof): Hoopstad (-DD),
Zietsman 1356 (PRE). -2727 (Kroonstad): Heilbron, Vegkop (-BD), WIn der
Merwe 1134 (PRE). -2825 (Boshof): Boshof (-CA), Potts 1058 (BLFU). -2827
(Senekal): Senekal, farm Doornkop (-DA), Sutton 631 (PUC). -2924
(Hopetown): Hopetown, farm Panfontein (-BD), Kok 220 (PRU).
-2925 (1agersfontein): Parys (-CC), Van der Merwe 2188 (PRE). -2926
(Bloemfontein): Bloemfontein (-AA), Potts 2862 (BLFU); Potts 1340 (BLFU);
Maller 379 (BLFU); Bloemfontein, Rhenoster Spruit (-AA), Potts 2801
(BLFU); Bloemfontein, Vaalbank (-CA), Smith 8602b (PRE). -2927 (Maseru):
Exelsior, Mensvreterberg (-AA), Zietsman 24 (PRE).
NATAL. - 2829 (Harrismith): Ladysmith (-DB), WIn der Merwe 2606
(PRE).
CAPE. - 2724 (Taung): Vryburg, farm Zoetvlei (-AA), Speedy 19/21A
(PRE). -2824 (Kimberley): Barkley West, between Gaap & Kneukel (-AC),
Acocks 8501 (PRE).
l.£DEBOURlA IN SOurH AFRICA
TAXONOMY
204
17. LEDEBOURIA OVATIFOLIA (Bak.) Jessop
Ledebouria ovatifolia (Bak.) Jessop in 11 S. Afr. Bot. 36(4): 262 (1970).
Scilla ovatifolia Bak. in Saund. Ref. Bot. 3: t.183 (1870).
Type: Natal, Cooper s.n. (K!, holo.; PRE!, photo.).
Scilla lanceaejolia (Jacq.) Bak. var. ovatifolia Bak. in 11 Linn. Soc. 11(54): 252
(1870).
Iconotype: Saund. Ref. Bot. 3: t.183. "Cap. B. Spei, Cooper s.n."
Scilla lanceaejolia sensu Wood & Evans in Natal Plants 3(4): t.202 (1900), non
Lachenalia lanceaejolia Jacq.
Scilla guttata C.A. Srn. in Kew Bull. : 243 (1930).
Type: Natal, Durban Div., Coopers.n. (K!, holo.).
Scilla cicatricosa C.A. Srn. in Kew Bull. : 246 (1930). Nom. novo only.
Scilla climacocarpha C.A. Srn. in Kew Bull. : 249 (1930).
Type: Orange Free State, Bethlehem, Phillips 3068 (PREl, holo.).
Scilla albomarginata Van der Merwe in Flower. PI. S. Afr. 24: t.947 (1944).
Type: Natal, Umzinto, Wm der Merwe 2669 (PRE!, holo.).
Scilla elevans Van der Merwe in Flower. PI. S. Afr. 24: t.948 (1944).
Type: Natal, Vryheid, liln der Merwe 2677 (PRE!, holo.).
Scilla collina Hutch. in Bot. in S. Afr. :344 (1946).
Type: Transvaal, Soutpansberg, Klein Australe, Smuts & Gillett 4186 (K!,
holo.; PREl, photo.).
LEDEBOURJA IN SOlJTH AFRICA
205
TAXONOMY
Plants solitary. Bulb hypogeal, 35 - 70 x 30 - 50 mm, ovoid; dead bulb scales
brown to purple, apices truncate, live bulb scales fleshy, loosely arranged, with
copious threads when torn, white to purple inside. Leaves partly emerged at
anthesis, 4 - 15, sometimes spreading, mostly appressed, ovate to deltate, 50 120 x 36 - 60 mm, with threads when torn, thickly fleshy, adaxial surface green
with purple or dark green spots and blotches, abaxial surface green or more
commonly purple, venation obscure; margin smooth, white to purple; leaf base
canaliculate; apex acute. Inflorescences 1 - 4, dense, ovate, 30 - 50 x 25 - 30
mm, flaccid, 30 - 40 -flowered, longer than the leaves; scape basally
compressed, green, glabrous; rachis ridged, 50 - 70 mm long. Bracts
membranous, 1 x 0.5 mm, linear to lanceolate, pink to purple with bracteoles.
Pedicels spreading, 7 - 12 mm long, pink to purple. Perianth 5 mm long, tepals
recurved, equal, oblong, 5 x 1.5 mm, apex obtuse, pink to purple with a green
keel. Stamens erect, 5 mm long, filaments maroon, epitepalous; anthers 1 mm
long, yellow. Ovary cylindrical, 6 -lobed, 1.75 x 3 mm, lobes narrowly
transversely elliptic, apex forming shoulders. Style 3.5 mm long, triangular,
glabrous, purple; stigma above anthers; stipe 1 x 1 mm. Capsule three-lobed,
symmetrical, globose; base truncate. Seed globose, 3 - 4 mm long, surface
strongly wrinkled, brown. (Figure 48).
L. ovatifolia is closely related to L. luteola. Together they constitute the section
Filiferae. L. ovatifolia differs from L. luteola in the truncate apices of the
bulb scales (Figure 49B) and the partly emerged, ovate to deltate, mostly
humifuse leaves.
Specific epithet etymology.
Describes the ovate leaves.
Flowering period
From August to March with a peak from August to November. Plants from the
lowveld areas of Transvaal and Natal flower later than plants from high lying
areas.
LEDEBOURIA IN SOlJTH AFRICA
206
TAXONOMY
E
Figure 48. Illustration of L. ovatifolia (Bak.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina X 2; D, apex of lamina X 20; E, bract
with bracteole X 10; F, apex of tepal X 20; G, flower X 5; H, tepal with
stamen X 5; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from
Stinon 11,180.
UD£BOURIA IN SOUTH AFRICA
Map 21.
12°
/: = ~
Known distribution of L. ovatifolia.(Bak.) Jessop
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G.P.·S. 13-0959
208
TAXONOMY
Figure 49. A, holotype of L. ovatifolia (Bak.) Jessop (distalleft) (K); B, plant
of L. ovatifolia showing the truncate apices of the bulb scales; C, SEM
micrograph of the adaxial lamina surface; D, SEM micrograph of the abaxial
lamina surface; E, SEM micrograph of a stomatum; F, habitat near
Trichardtsdal, north-eastern Transvaal. The vegetation consists of open low
evergreen Protea roupelliae var. roupelliae - Themeda triandra woodland. B
from llin der Merwe 2172 and C - E from Venter 13,349.
LEDEBOURIA IN SOlITH AFRICA
TAXONOMY
209
f'\:
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LEDEBOURlA IN SOUTH AFRICA
210
TAXONOMY
Distribution (Map 21).
Widely distributed throughout Transvaal, Swaziland, Orange Free State,
Lesotho , Natal and Transkei with a few scattered localities in the eastern and
northern Cape.
Habitat
L. ovatifolia grows in grassland or woodlands (then in large open grassy areas
between the trees and bushclumps) (Figure 49F).
Leaves are occasionally grazed but the copious threads in the bulb scales protect
it from being eaten by animals except the Porcupine (Hystrix africaeaustralis
Peters).
Variation
Plants of L. ovatifolia growing in shady conditions tend to have more lanceolate
spreading leaves (\enter 13,372) with dull darker green or purplish markings (
sometimes immaculate ). Such plants, collected at Vryheid, led Van der Merwe
to describe a separate species namely Scilla elevans (1944b).
Historical background
The plant described by Wood & Evans (1900) as Scilla lanceaefolia does not
agree with the figure of the nomenclatural type (Jacquin 1794). Wood & Evans
cite no specimen but state that the plant figured was collected around Durban.
The name S. lanceaefolia used by Wood & Evans belongs to Ledebouria
revoluta (L.t) Jessop, consequently C.A. Smith (1930) proposed the name Scilla
cicatricosa for this plant.
LEDEBOURJA IN SOUTH AFRICA
211
TAXONOMY
Smith (1930) correctly states that the plant described by Baker (1870d) as Scilla
lanceaejolia, is not the same as S. lanceaejolia sensu Jacquin. Smith (1930)
failed to recognise Baker's species as Ledebouria ovatifolia as stated by him in
the synonym list under Scilla guttata where he mentions Scilla lanceaefolia
(Jacq.) Bak. var. ovatifolia Bak. as a synonym.
Specimens examined
VENDA. -2230 (Messina): Thate Vondo (-CD), Mugwedi 1312 (PRE).
TRANSVAAL. -2330 (Tzaneen): Westfalia Estate (-CA), Scheepers 468
(PRE). -2430 (Pilgrim's Rest): Mount Sheba Nature Reserve (-DC), Kerfoot 94
(J); Kerjoot 82 (J); Pilgrim's Rest (-DD), Rogers 14,951 (J, PRE); Rogers
14,701 (J); Graskop, Lisbon Falls (-DD), Venter 13,248 (PRE). -2527
(Rustenburg): Rustenburg Nature Reserve (-CA), Jacobsen 1031 (PRE). -2528
(Pretoria): Pretoria (-CA), \Un der Merwe 1989 (PRE); Fountains Valley (CA),
Van der Merwe 2001 (PRE); Wonderboompoort (-CA), Labuschagne s.n.
(PRE); Wonderboom (-CA), Rogers 21,718 (J); Meyerspark (-CA), Theron 176
(PRU); Trigaardts Poort (-CB), Van der Merwe s.n. (PRE); Roodeplaat Dam
Nature Reserve (-CB), Van Rooyen 2007 (PRU); Van Rooyen 2319 (PRU);
Olifantsfontein (-CC), Van der Merwe 1971 (PRE). -2529 (Witbank):
Middelburg, farm Doornkop 273 JS (-CD), du Plessis 727 (PRU); Middelburg,
Soetevelden (-CD), Van der Merwe 2198 (PRE). -2530 (Lydenburg):
Lydenburg, farm Nooitgedacht 266 (-AB), \Un der Merwe 1778 (PRE); Sabie (BB), Rogers 23,030 (PRE); Nelspruit (-BD), \Un Jaarsveld 18 (PRE); Witklip
Plantation (-BD), Kluge 127 (PRE); Kluge 128 (PRE, PRU);
Waterval - Boven (-CB), Rogers 227 (GRA); Rogers 5116 (PRE);
Machadodorp, Bambi Hotel (-CB), Booysen s.n. (BLFU); Machadodorp,
Goedverwagting (-CB), Van der Merwe 1252 (PRE). -2531 (Komatipoort):
Pretorius Kop (-AB), Letty 204 (PRE); \Un der Schyff 3226 (KNP); Barberton
(-CC), \Un der Merwe 1817 (PRE); Calpin 508 (PRE); Barberton, Maid of the
Mist Mountain (-CC), Venter 12,543 (UNIN); Barberton, farm Oosterbeek (CC), Venter 13,376 (UNIN). -2626 (Klerksdorp): Ventersdorp, farm Somerville
(-BD), Louw 3917 (PUC); Bosch & Kruger S.n. (PUC); Ventersdorp, farm
LEDEBOURIA IN SOlITH AFRICA
212
TAXONOMY
Goedgedacht (-DB), Sutton 467 (PUC). -2627 (Potchefstroom): Abe Bailey
Nature Reserve (-AD), Van k~
213 (PUC); Modderfontein (-BB), Van der
Merwe 1983 (PRE); Sterkfontein (-BB), Mogg 34,438 (J); Vereeniging (-DB),
Burtt Davy 17,149 (BOL). -2628 (Johannesburg): Johannesburg (-AA), Moss
7347 (J); Bedford (-AA), Gilliland s.n. (1); Morningside (-AA), Gilliland s.n.
(1); Bryanston (-AA), Dahlstrand 364T (PEU); Limbro Park (-AA), Gilliland
s.n. (J); Houghton Hill (-AA), Cohen s.n. sub J 31,881 (J); Klipriviersberg (AC), Gilliland s.n. (J); Wattles (-AC), Moss 10,811 (J); Rosettenville (-AC),
Moss 18,929 (J). -2629 (Bethal): Bethal (-AD), Leendertz 10,139 (PRE);
Breyton (-BD), Steyn 976 (NBG); Standerton, farm Starkville (-CD), ~nter
2049 (PRE); Bethal, Vermaakskraal (-DD), Turner 1154 (PRE). -2630
(Carolina): Carolina (-AA), Galpin 7 (BOL); Rogers 21,301 (1); Rogers 19,604
(1); Galpin 13,096 (PRE); Lichfield (-DA), lhn der Merwe 1073 (PRE). -2729
(Volksrust): Wakkerstroom, Sandspriut (-BA), Gillispie 12,506A (PRE);
Amersfoort (-BB), Turner 1107 (PRE). -2730 (Vryheid): Wakkerstroom,
Oshoek (-AD), Devenish 175 (PRE).
SWAZILAND. -2631 (Mbabane): Mankaiana, Ntondozi (-CA),
Compton 30,892 (NBG, PRE).
ORANGE FREE STATE. -2727 (Kroonstad): Kroonstad (-CA),
Chesnell124 (1); Vals River (-CB), Pont 298 (PRE).
LESOfHo. -2927 (Maseru): Mafeteng (-CC), Dieterlen 1361 (PRE).
NATAL. -2729 (Volksrust): New Castle, Chelmsford (-DD), Smit
chf. 011 (NU). -2730 (Vryheid): Vryheid, farm Groenvlei (-AD), Strey 9078
(NH); Vryheid (-DD), lhn der Merwe 2399 (PRE); Vryheid Nature Reserve (DD), Schrire 1272 (NU). -2731 (Louwsburg): Hlatikulu, farm Goedgegun (AA), Ross 1470 (NU); Josini (-BD), Strey 4621 (PRE); ltala Nature Reserve (CA), McDonald 184 (NU). -2732 (Ubombo): Mazengwenya (-BD), Wlrd 1558
(NH). -2828 (Bethlehem): Royal National Park (-DB), Schelpe 1747 (NH).
-2829 (Harrismith): Swinburne (-AD), Coetzee s.n. (BOL); Ladysmith, farm
Elandslaagte (-BD), Rendles 54 (NU); Cathedral Peak (-CC), Esterhuysen
15,463 (BOL). -2830 (Dundee): Dundee Airfield (-AA), Shirley 114 (NU);
LEDEBOURIA IN SOlITH AFRICA
213
TAXONOMY
Dundee, Impati Hill (-AA), Shirley 1 (NU); Kranskop (-AA), Johnson 3900
(NH); Schrire 1557 (NH); Muden (-CD), Cromwright s,n. sub NBG 73,448
(NBG). -2831 (Stanger): Hlabisa, farm Palm Ridge (-BB), Harrison 122 (NH);
Mtunzini , Umhlatuzi Flats (-DD),
Venter 2563 (PRE). -2832 (Mtubatuba):
.
Hluhluwe Game Reserve (-AA), Fakude & Bourquin 11 (NU, PRE); Poplela,
Sunset, Petlos Krantz (-CB), Rennie 78 (NU); -2929 (Underberg): Kamberg,
Fourteen Streams (-BC), Cooke 5227 (BOL); Kamberg (-BD), Wright 1855
(NU); Wright 2221 (NU); Ruddock 20 (NU); Sani Pass (-CB), Hilliard & Burrt
18,055 (NU); Cathedral Peak (-CC), Everson 231 (PRE); Underberg (-CD),
WIn der Merwe 2760 (NU, PRE); Garden Castle (-CD), Hilliard 8159 (NU);
Loteni Game Reserve (-DA), Phelan 128 (NU); Cunningham 2131 (NH);
Underberg, Nkonzo State Forest (-DD), Nicholas & Norris 1148 (CPF, NH,
PRE); Polela, Highbury (-DD), Landsdell s.n. sub NH 42772 (NH). -2930
(Pietermaritzburg): Tweedie (-AC), Mogg 1302 (PRE); Lions River (-AC),
Strey 3514 (PRE); Caversham (-AC), Mogg 1491 (PRE); Zwartkop Location (AC), Moll 1142 (PRE); Lidgetton (-AC) , Mogg 1408 (PRE); Nottingham Road
(-AC), Galpin 9445 (PRE); Merrievale (-CA), Smith s. n. (PRE);
Pietermaritzburg (-CA), Sanden-Smith 19 (NH); Clarendon (-CB), Newmark 49
(NU); Hayfield (-CB), Nicholas 1998 (CPF, NH); Howick (-CB), MacDevette
1070 (NH); Foley s.n. (BOL); Scottsville (-CB), Daffey s.n. (NU); Zwartkops (CD), Lawson 237 (NU); New England Road (-CD), Lachman 3 (NU); Town
Hill (-CD), Clarkson 62 (NU); Killick 184 (NU); Oribi Airfield (-CD),
Brayshaw 17 (NU); Richmond (-CD), \yvyan 30 (BOL); Cato Ridge (-DA),
McClean & Ogilvie s.n. (NH); WIn der Merwe 2533 (PRE); Ross 2160 (NH,
NU); Inchanga (-DA), Loubser 15 (NU); Kloof, Daviesville (-DD), Herre s.n.
(BOL). -2931 (Stanger): Durban (-CC), Wood 181 (BOL); Wood 1392 (NH).
-3029 (Kokstad): Glen Garry, Mealiefontein (-BC), Strey 9165 (NH); Kokstad (CB), Tyson 1123 (BOL); Weza (-DA), Strey 10,500 (NH); Harding Road (_
DB), Strey 6977 (NH). -3030 (Port Shepstone): Scottburgh (-BC), Venter
13,195 (UNIN); Port Shepstone (-CB), Wlrren s.n. (PRE); Umtamvuna Nature
Reserve (-CC), Abbot! 2100 (PRU).
TRANSKEI. -3028 (Matatiele): Ramatseliso Gate (-BB), Boardman 147
(PRE); Mount Frere (-DD), Venter 13,324 (UNIN). -3128 (Umtata): Umtata,
UNITRA Campus (-DB), Hutchings 589 (KEI).
LEDE80URIA IN SOlITH AFRICA
214
TAXONOMY
CAPE. -2824 (Kimberley): Warrenton (-BB), Adams 98 (BOL). -3128
(Umtata): MaClear (-AB), Wederman & Oberdieck 1156 (PRE). -3227
(Stutterheim): Stutterheim, Evelyn Valley (-CB), Barker 4228 (NBG); Komgha
(-DB), Flanagan 1301 (BOL). -3318 (Cape Town): Claremont (-CD),
Schlechter 2940 (BOL). -3325 (Port Elizabeth): Port Elizabeth (-DC), Fourcade
s.n. sub NBG 73,449 (NBG).
UDEBOURIA IN SOUTH AFRICA
215
TAXONOMY
Sectio Monophyllae S. ,retn~
sect. nov., folio unico humifuso; inflorescentia
erecta, racemo globoso.
Species typica: Ledebouria monophylla S. Venter.
Typus: Transvaal, Graskop, Paradise Camp, retn~
13,235 (PRE).
Species: L. monophylla S. Venter.
Plants solitary, 30 - 50 mm tall. Bulbs 10 - 20 mm wide; apices of dry
bulb scales truncate. Leaf single, appressed to ground, thickly fleshy.
Inflorescence solitary, erect. Peduncle thick, base compressed. Raceme
globose. Perianth stellate.
Distribution and habitat
Transvaal Drakensberg Escarp and two isolated localities near Pietermaritzburg,
Natal.
Occurs in montane grassland.
18. LEDEBOURIA MONOPHYLLA S. Venter
Ledebouria monophylla S. ,retn~
sp. nov., ad L. cooperi (Hook.f) Jessop
sed folio singulo, late ovato adpresso; inflorescentia solitaria erecta, pedunculo
basi compresso et floribus stellaribus, lobis cucullatis distinctissima.
Type: Transvaal, Graskop, Paradise Camp,
retn~
13,235 (PRE!, holo.; NU!;
UNIN!).
Plants solitary. Bulbs hypogeal, 10 - 20 x 8 - 12 mm, ovoid; dead bulb scales
light brown, apices truncate, live bulb scales fleshy, tightly arranged, with
threads when torn, white inside. Leaf single, sometimes partly emerged but
usually fully developed at anthesis, appressed to ground, broadly ovate, 30 - 40
x 30 - 40 mm, without threads when torn, thickly fleshy, glossy green,
immaculate, venation obscure; margin smooth, red; leaf base canaliculate; apex
LEDEBOURIA IN SOurH AFRICA
216
TAXONOMY
obtuse to acute. Inflorescence solitary, erect, dense, globose, 10 - 15 x 15 mm,
erect, 20 - 30 -flowered, longer than the leaves; scape basally compressed,
green spotted purple, glabrous; rachis ridged, 10 - 30 mm long. Bracts fleshy,
lanceolate to bifurcate, pink with bracteoles. Pedicels spreading, 3 - 5 mm long,
pink. Perianth 4 mm long, stellate, tepals equal, oblong, 4.0 x 1.5 mm, apex
obtuse, cucullate, pink to purple. Stamens spreading, 1.5 - 4.0 mm long,
filament base slightly flattened, upper part purple with lower part white, free;
anthers 0.5 mm long, yellow. Ovary globose, 3 -lobed, 1 x 2 mm, lobes
depressed ovate, shoulders raised. Style 1.5 mm long, triangular, glabrous,
purple; stigma above anthers; stipe 0.25 x 0.25 mm. Capsule three-lobed,
symmetrical, globose; base truncate. Seed drop-shaped, 1.5 mm long, surface
strongly wrinkled, brown. (Figure 50).
L. monophylla is not closely allied to other South African species. It is
characterized by the solitary, appressed fleshy leaf, the solitary inflorescence
with depressed peduncle, globose raceme and stellate flowers (Figure 50).
Specific epithet etymology.
Alludes to the single leaf.
Flowering period
From September to December with a peak from September to October.
Distribution (Map 22).
L. monophylla occurs in the eastern Transvaal with a single record from
Zwartkop near Pietermaritzburg, Natal.
LEDEBOURlA IN SOlITH AFRICA
217
TAXONOMY
D
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Figure 50. Illustration of L. monophylla S. Venter. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, bract with bracteole X 10; E,
flower X 10; F, tepal with stamen X 10; G, apex of tepal X 20; H, ovary lateral
view X 10; I, ovary dorsal view X 10. All from
LEDEBOURlA IN
retn~
soum AFRICA
13,235.
Map 22.
12°
Known distribution of L. monophylla S. Venter
14°
16°
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22°
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G.P.·S. 13-0959
219
TAXONOMY
Figure 51. A, holotype of L. monophylla S. Venter (PRE); B, plants showing
the solitary humifuse leaf. Bar
= 20 mm; C, plant in habitat near Graskop,
showing the solitary leaf and erect globose inflorescence. Bar = 10 mm; D,
SEM micrograph of the adaxial lamina surface; E, SEM micrograph of the
abaxial lamina surface; F, SEM micrograph of stomata; G, habitat at the type
locality near Graskop. The vegetation consists of closed evergreen low Erica
leucopelta - Passerina montana - Pteridium aquilinum shrubland. A - F from
lenter 13,235.
LEDEBOURlA IN SOUTH AFRICA
220
TAXONOMY
HERBARIUM
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LEDEBOURIA IN SOlITH AFRICA
221
TAXONOMY
Habitat
L. monophylla occurs in humusrich, medium grained (0.25 - 1.0 mm eJ» deep
(30 - 300 mm) grey to black peaty sands derived from quartzites and sandstones.
The habitat vegetation is mainly montane grassland and Fynbos next to the
streams (Figure 51 C). Populations are scattered and consist of up to 200
individuals.
Variation
Plants growing in full sun away from moisture tend to have less succulent, redtinged leaves. Flower colour varies from light pink to dark pink.
Historical background
The earliest specimen (Thorncroft 998) was collected in 1917 at Graskop. The
next specimen was collected 50 years later (Rauh & Schlieben 9785).
'\-enter 13,235 shows a developmental series of plants and is thus chosen as the
type.
Specimens examined
TRANSVAAL - 2430 (Pilgrim's Rest): Trichardtsdal, farm Malta (-AA), Junod
4434 (PRE); Mount Sheba Nature Reserve (-DC), Geliffe et al. 93 (1); Graskop
(-DD), Thorncroft 998 (PRE); Paradise Camp (-DD), '\-enter 12,602 (UNIN);
'\-enter 13,235 (NU, PRE, UNIN); Graskop, Stanley Bush Kop (-DD), Raal et
al. 1038 (PRE); God's Window (-DD), Mogg 33,289 (1). -2530 (Lydenburg):
Mt. Anderson (-BA), Mohle 427 (PRE); Long Tom Pass (-BA), Rauh &
Schlieben 9785 (PRE); Witklip (-BD), Kluge 199 (PRU).
NATAL - 2930 (Pietermaritzburg): Zwartkop (-CB), MacDevette 1076
(NH).
LED£BOURJA IN SmITH AFRICA
222
TAXONOMY
Sectio Bulbilia S. lknter, sect. nov., bulbis cataphyllisque frequentibus.
Species typica: Ledebouria parvijolia S. Venter.
Typus: Transvaal, Pilgrim's Rest, lknter s.n. (PRE).
Species: L. cooperi (Hookf) Jessop and L. parvijolia S. Venter.
Plants 25 - 35 mm tall. Bulb 10 - 25 mm long, torn scales lacking threads,
bulblets present. Leaves with 1 cataphyll, leaves spreading, 10 - 20 mm wide.
Inflorescence rachis smooth. Perianth lobe apex obtuse. Ovary lobes narrowly
transversely oblong; style 3 mm long. Capsule globose.
Distribution and habitat.
Coastal bush, woodland to montane grassland.
Subsectio Zebrinae S. lknter, subsect. nov., tepalis ad apicem valde cucullatis.
Species typica: L. cooperi (Hookf) Jessop.
Species: L. cooperi (Hookf) Jessop.
Inflorescence as long or shorter than the leaves, rachis smooth. Bracts with
bracteoles always present. Tepals with apices strongly cucullate. Ovary 2 mm
long.
19. LEDEBOURIA COOPERI (Hook):) Jessop
Ledebouria cooperi (Hook. f.) Jessop in Jl S. Afr. Bot. 36(4): 247 (1970).
Scilla cooperi Hook): in Bot. Mag. 92: t.5580 (1866).
Type: Cape, Cooper s.n. (Kl, holo.; PREI, photo.).
Scilla subglauca Bak. in Saund. Ref. Bot. 3: t.186 (1870).
Iconotype: Saund. Ref. Bot. 3: t.186, "Cape of Good Hope, Cooper s.n. ".
LEDEBOURIA IN SOUTH AFRICA
223
TAXONOMY
Scilla concinna Bak. in Saund. Ref. Bot. 4: t.235 (1870).
Type: Saund. Ref. Bot. 4: t.235. "Cape Colony, Cooper s.n.".
Scilla exigua Bak. in Jl Linn. Soc. (Bot.) 13: 247 (1873).
Type: Natal, Camperdown, Farm Assegai Kraal, Sanderson 670 (feD, holo.).
Scilla barberi Bak. in 11 Linn. Soc. (Bot.) 13: 247 (1873).
Type: Transkei, Ad ripas flu minis Tsomo, Barber 805. Type not found.
Scilla saturata Bak. in J. Bot. 3: 365 (1874).
Type: Orange Free State, Cooper 993 (Kl, holo.; PREl, photo.).
Scilla adlamii Bak. in Gdnrs' Chron. 9(3): 521 (1891).
Type: Natal, Adlam s.n. (K!, holo.; BOLl, drawing; PREl, photo.).
Scillajehri Bak. in Bot. Jahrb. 15(35): 7 (1892).
Type: Transvaal, Pretoria, Fehr s.n. ( Zl, holo.).
Scilla inandensis Bak. in Flora Cap. 6: 483 (1896).
Type: Natal, Inanda, WJod 630 (SAMl, lecto.; BOLl; NH!).
Scilla globosa Bak. in Flora Cap. 6: 484 (1896).
Type: Natal, Griqualand Orientalis, In humilosis prope Kokstad, Tyson 1557.
(SAMl, holo.; BOLl).
Scilla rogersii Bak. in Flora Cap. 6: 486 (1896).
Type: Cape Colony, Rogers s. n. (K!, holo.; PRE!, photo.).
Scilla palustris Wood & Evans in J. Bot. 37: 251 (1899).
Type: Natal, in swamp near Newcastle, Wood 6501 (NHl, holo.; PREl,
photo.).
Scilla rehmannii Bak. in Bull. Herb. Boiss. 2(1): 853 (1901).
Type: Natal, Inanda, Rehmann 8277 (Z!, holo.).
LEDEBOURIA IN SOlITH AFRICA
224
TAXONOMY
Scilla aggregata Bak. in Bull. Herb. Boiss. ser. 2(4): 1001 (1904).
Type: Transvaal, Modderfontein, Conrath 701 (Kl, holo.; GRAZ; PREl,
photo.).
Scilla tristachya Bak. in Bull. Herb. Boiss. ser. 2(4): 1001 (1904).
Type: Transvaal, Modderfontein, Conrath 693 (GRAZ, holo.; BOLl, drawing;
PREl, photo.; Z.).
Scilla conrathii Bak. in Bull. Herb. Boiss. ser. 2(4): 1002 (1904).
Type: Transvaal, Modderfontein, Conrath 699 (Kl, holo.; GRAZ; PREl,
photo.).
Scilla londonensis Bak. in Bull. Herb. Boiss. ser. 2(4): 1002 (1904).
Type: Cape, East London, Conrath 694 (GRAZ; PREl, photo.).
Scilla glaucescens Van der Merwe in Flower. PI. S. Afr. 23: t.912 (1943).
Type: Transvaal, Carolina, on farm Onbekend, 'llin der Merwe 2073 (PREl,
holo.).
Plants solitary or gregarious. Bulb hypogea1, 10 - 25 x 10 - 25 mm, subglobose;
dead bulb scales brown to purple, membranous, apices attenuate, live bulb scales
loosely arranged, without threads when torn, white inside. Cataphyll solitary,
exerted beyond ground level. Leaves fully developed at anthesis, 3 - 8,
spreading, lanceolate to oblanceolate, 200 - 250 X 16 - 18 mm, without threads
when torn, fleshy, glossy, green to glaucous green, upper surface with dark
green or purplish spots and blotches, lower surface with longitudinal stripes and
spots, venation sometimes prominent; margin light green; leaf base canaliculate;
apex acute. Inflorescences 1 - 3, dense (rarely open), cylindric to sub-globose,
50 - 80 x 20 - 30 mm, erect to flaccid, 20 - 60 -flowered, as long or longer than
the leaves; scape terete at base, green, glabrous; rachis smooth, 0 - 150 mm
long. Bracts membranous, 1.5 x 0.5 mm, linear to bifurcate, pink to purple
with bracteoles sometimes present. Pedicels spreading, 6 - 12 mm long, pink.
Perianth 4 - 7 mm long, tepals recurved, equal, oblong, 4 - 5 x 1.5 - 2.0 mm,
apex obtuse, strongly cucullate, pink to deep pink with a green keel. Stamens
LEDEBOURlA IN SOlJfH AFRICA
225
TAXONOMY
erect, 3 - 4 mm long, filaments pink, free; anthers 1 mm long, pale yellow.
Ovary ovoid, 6 -lobed, 2 x 4 mm, lobes narrowly transversely oblong,
shoulders of apex raised. Style 3 mm long, triangular, glabrous, purple; stigma
above anthers; stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, globose;
base truncate. Seed drop-shaped, 4 mm long, surface strongly wrinkled,
yellowish-brown to brown. (Figure 52).
L. cooperi is most closely related to L. parvijolia S. Venter but differs in having
glabrous leaves.
Specific epithet etymology.
Commemorates Mr. Thomas Cooper, who collected the type material.
Flowering period
From October to February.
Distribution (Map 23).
Throughout higher rainfall areas of South Africa mostly Natal and the
Transvaal. Poorly represented in the eastern Cape.
Habitat
L. cooperi usually occurs in moist habitats (Figure 53B), on humic clay soils.
Soils include clay loams, sandy loams and on the mountain ranges a sandy peat.
Soils in the low lying habitats tend to be rich in clay and are inundated with
water for long periods.
This species occurs in grassland. Most of the recorded localities are situated in
marshy grassland areas.
LED£BOURLA IN SOurH AFRICA
226
TAXONOMY
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F
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H
B
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Figure 52. Illustration of L. cooperi (Hook.f.) Jessop. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, flower X 10; E, bract with
bracteole X 10; F, apex of lamina X 20; G, tepal with stamen X 10; H, ovary
lateral view X 10; I, Ovary dorsal view X 10. All from Crouch 97.
LEDEBOURJA IN SOUTH AFRICA
Known distribution of L. cooperi (Hook.f.) Jessop
Map 23.
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228
TAXONOMY
Figure 53. A, holotype of L. cooperi (Hook..f.) Jessop (distal specimen) (K); B,
habitat on slopes of Mount Currie near Kokstad with montane grassland and
swampy areas; C, SEM micrograph of the adaxial lamina surface. Bar = 100
J.'m ; D, SEM micrograph of the abaxial lamina surface. Bar = 100 J.'m; E,
SEM micrograph of a stomatum. Bar = 43 J.'m; F, plants of L. coopen showing
parallel purple markings on the lamina. Bar = 20 mm. C - F from Venter
13,194.
LED£BOURIA IN SOlJfH AFRICA
TAXONOMY
229
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LED£BOIJRIA IN SOUTH AFRICA
230
TAXONOMY
Variation
The number of synonyms reflect the variability of this species. Baker described
most of the species now regarded as synonyms of L. cooperi.
Historical background
Hooker (1866) described Scilla cooperi to commemorate Thomas Cooper who
collected the first specimens. These were cultivated at Kew and flowered in
1863 and the original description was drawn up from this material (Baker
l870d).
Baker (1896) described S. globosa from material collected near Kokstad (Tyson
1557). In his description, Baker does not mention either a collecting number or
the herbarium where the type is housed. The specimen Tyson 1557 was
mentioned by Jessop (1970) but with reservation indicated by a question mark.
From the literature (Gunn & Codd 1981) and herbarium material it is shown that
the above specimen is indeed the type of S. globosa. The Rev. W.M. Rogers
made various collections in the Riversdale and George areas from 1860 - 1862.
The type of S. rogersii (Rogers s.n.) is without locality, but, on the type sheet
there is a note in pencil by Archdeacon P.A. Rogers that his father could have
collected the specimen at George. This may be the case for there is a specimen
in the Moss Herbarium (1) collected by the Rev. Rogers at George, which
matches the type.
Van der Merwe (1943b) described S. snec ~ ualg
from material he collected.
He expresses the view that S. glaucescens resembles S. cooperi in its general
appearance but differs, in its relatively wider and shorter leaves, and the
absence of vertical stripes on the abaxial surfaces. These characters are not
regarded as diagnostic by the present author.
LEDEBOURJA IN SOlJfH AFRICA
231
TAXONOMY
Specimens examined
TRANSVAAL. - 2327 (Ellisras): Waterberg, Farm Elandsbosch (-DD),Page
s.n. sub 1 31,823 (1). - 2329 (Pietersburg): Woodbush (-CD), Moss 15,586 (1);
Haenertsburg (-DD), Pot! 13,681 (PRE). - 2330 (Tzaneen): Westfalia Estate (CA), Scheepers 695 (UNIN, PRE); Tzaneen (-CC), Rogers 12,526 (BOL). 2427 (Thabazimbi): Leeupoort Tin Mine (-BC), Rogers 22,764 (1); Kransberg,
Farm Groothoek (-BC), Jacobsen 3493 (PRE); Waterberg, Farm Bergkrans (BC), Jacobsen 3500 (PRE). - 2429 (Zebediela): Makapansgat (-AA), Leendertz
896 (BOL). - 2430 (Pilgrim's Rest): Serala Wilderness Area (-AA), Venter
11 ,119 (UNIN); Venter 11,038 (UNIN); Venter 10,239 (LYD); Lekgalameetse
Nature Reserve (-AA), Stalmans 668 (UNIN); The Downs (-AA), Moss 11,711
(1); Mount Sheba (-CD), Kerfoot et al. 246 (1); Mariepskop (-DB), WIn der
SchYff 6363 (PRU); WIn der Schijff 6332 (PRU); Venter 12, 732 (UNIN); Raal
1676 (1); Mount Sheba Nature Reserve (-DC), Keifoot K8107 (1); Keifoot 8168
(1); Goodman s.n. sub 1 63,701 (1); Ohrigstad Nature Reserve (-DC), Jacobsen
1580 (PRE); Jacobsen 1803 (LYD); Pilgrim's Rest (-DD), Rogers 14,952 (1);
Rogers 14,953 (1); Rogers 18,619 (1); Rogers 14,956 (1); Graskop (-DD),
Pienaar 756 (PRE); Holland s.n. sub BOL 50,788 (BOL); Graskop, Paradise
Camp (-DD), Venter 8885 (LYD); Kowyns Pass (-DD), Venter 12, 578
(UNIN); Louw 2354 (STE); Prosser 1500 (PRE); God's Window (-DD), Venter
13,239 (UNIN); Belvedere (-DD), Davidson 448 (1); Macmac (-DD), WIn der
Merwe 343 (PRE). - 2527 (Rustenburg): Rustenburg Nature Resrve (-CA),
Jacobsen 1034 (PRE); Rustenburg Kloof (-CA), Lanham 80 (PRE); Rustenburg
(-CD), Pegler 975 (BOL); Rustenburg, Farm Uitkomst (-DD), Coetzee 438
(PRU); Hennops River (-DD), Prosser 1629 (PRE); - 2528 (Pretoria): Pretoria
(-CA), l-bgts s.n. sub NBG 1639/28 (NBG); pretoria, Faery Glen (-CA), Mogg
s.n. sub PRE 36,658 (PRE); Schweikerdt 27,640 (PRE); Wonderboom Reserve
(-CA), Repton 1992 (PRE); Knoppiesfontein (-CA), Van der Merwe 1999
(PRE); Bronkhorstspruit (-CD), Codd & De Winter 3138 (PRE); Premier Mine
(-DA), Rogers 25,250 (PRE); Premier Mine, Farm Kaffirkraal (-DA), Moss
19,098 (1); Pretoria (-CA), Vogts s.n. sub BOL 1639/28 (BOL); Rietvlei
Reserve (-CD), Repton 3132 (PRE). - 2529 (Witbank): Middelburg, Farm
Doornkop (-CB), du Plessis 615 (PRU); du Plessis 928 (PRU); du Plessis 1145
(PRE); Middelburg, Farm Donkerhoek (-CB), Theron 1852 (PRU); .lrn 0.50"
LEDEBOURlA IN SOlJfH AFRICA
232
TAXONOMY
Middelburg, Hoedspruit Vlei (-CD), Van der Merwe 2007 (PRE); Carolina,
Farm Hillside (-CD), van der Merwe Se. 2. (PRE); Middelburg (-CD), Young
A58a (PRE). - 2530 (Lydenburg): Steenkampsberg (-AA), Codd 8059 (PRE);
Sabie, Hartebeesvlakte (-AB), Kluge 2335 (PRE); Verloren Vallei Nature
Reserve (-AC), Bloem 127 (PRU); Dullstroom (-AC), Strey 3435 (PRE); Hunter
55 (Johannesburg Botanical Garden); Sabie (-BB), Moss 7345 (J); Koeleman
s.n. sub PRE 35,461 (PRE); Koeleman s.n. sub PRE 35,485 (PRE);
Wonderkloof Nature Reserve (-BC), Elan-Puttiek 309 (PRE); Houtbosloop (BC), WIn der Merwe 1788 (PRE); Witklip Plantation (-BD), Kluge 105 (PRU);
Rosehaugh (-BD), Sim s.n. sub BOL 50,807 (BOL); Carolina, Farm Bergendal
(-CA), Galpin s.n. sub BOL 22,494 (BOL); Belfast (-CA), WIn der Merwe 1228
(PRE); Franks s.n. sub PRE 9,769 (PRE); Bolus s.n. sub BOL 12,406 (BOL);
Long Tom Pass (-CA), Stirton 232 (NU); Kaapsche Hoop (-DB), retn~
13,390
(UNIN); WIn Jaarsveld 960 (NBG); Kaapsche Hoop, Duiwels Kantoor (-DB),
Thode A1650 (NH, PRE). - 2531 (Komatipoort): Barberton, Reception Siding (BD), Rogers 22,175 (J); Barberton (-CC), Rogers 24,653 (PRE); Saddleback
Range (-CC), WIn der Merwe 1822 (PRE); Maid of the Mist (-CC), ~nter
12,
558 (UNIN); Angle Station (-CC), Venter 13,388 (UNIN); Havelock,
Nottingham Peak (-CC), Saltmarshe 1053 (PRE). - 2627 (Potchefstroom):
Welverdiend (-AD), Louw 88 (PUC); Magaliesberg (-BA), WIn der Merwe 2039
(PRE); Krugersdorp (-BB), Behr 963 (PRE); Johannesburg, Discovery (-BB),
Lueas s.n. sub J 36,311 (J); Mulders Drift (-BB), Gilliland s.n. sub J 26,178
(1); Witpoortjie (-BB), Gilliland s.n. sub J 26,628 (1); Hugo s.n. sub STE
31,923 (STE); Moss 7143 (J); Johannesburg, Jackson's Drift (-BD), Gilliland
s.n. sub PRE 26,273 (PRE); Lloyd s.n. sub J 26,863 (1); Potchefstroom (-CA),
Louw 1613 (PUC); Badenhuizen s.n. sub J 28,588 (J); Boskop Dam (-CA),
Kloof (-CB), Moss 25,860
Ubbink 1279 (PUC); Louw 2295 (PUC); He~dlbrg
(J); Vereeniging (-DB), Leendertz s.n. sub TM 3948 (PRE); Leendertz 3947
(PRE). - 2628 (1 ohannesburg) : Johannesburg (-AA), Moss 2723 (1);
Modderfontein (-AA), WIn der Merwe 1948 (PRE); Lynn Park (-AA), Munday
553 (J); Olifantsvlei (-AA), Dimovie s.n. sub J 28,083 (J); Lueas s.n. sub J
35,244 (1); Bryanston (-AA), Gilliland s.n. sub J 26,272 (J); Gilliland s.n. sub J
26,269 (1); Lueas 325 (J); Florida (-AA), Moss 9757 (J); Florida Marsh (-AA),
Moss 7350 (J); Houghton Estate (-AA), Moss 2161 (J); Killarney Vlei (-AA),
Moss 4136 (1); Heidelberg (-AD), Leendertz 1051 (PRE); Delmas,
LEDEBOURlA IN SOUTH AFRICA
233
TAXONOMY
Olifantsfontein (-BA), Van der Merwe 1972 (PRE); Van der Merwe 1973
(PRE); Heidelberg (-CB), Moss 17,723 (1); Vaaldam, Farm Sandfontein (-CD),
Aspoas s.n. sub J 48392 (J). - 2629 (Bethal): Ermelo, Farm Nooitgedacht (DB), Henrici 1296 (PRE); Dyer 4165 (PRE); Ermelo at Vaal River (-DB), lbn
der Merwe Se.1. (PRE). - 2630 (Carolina): Carolina (-AA), Rogers 19,600 (J);
Rogers 19,112 (1); Rogers 19,109 (J); Rogers 19,601 (J); Rogers 19,602 (1);
Ermelo, The Gem (-BC), Van der Merwe 1097 (PRE); Walker s.n. sub PRE
36,647 (PRE); Maviriestad (-CA), Pott 15,257 (PRE). - 2725 (Bloemhof):
Wolmaranstad, Leeuwfontein (-BB), llin ~k
1385 (PUC). - 2730 (Vryheid):
Wakkerstroom (-AC), Rogers 22,493 (BOL); Wakkerstroom, Farm Groothoek
(-BA), Bahrman 26 (PRE); Hlangapies Mountain (-BA), llin der Merwe 2073
(PRE); Van der Merwe 1115 (PRE); Wakkerstroom, Farm Oshoek (-BA),
Devenish 102 (PRE); Devenish 99 (PRE).
SWAZILAND. - 2631 (Mbabane): Forbes Reef (-AA), Compton 26,130
(PRE); Compton 29,552 (NBG); Black Mbuluzi Valley (-AA), Compton 28,066
(NBG); Usutu Forest (-CA), Mott 455 (PRE); Hlatikulu (-CD), Stewart 2530
(NBG); Stewart 2529 (NBG); Ross 1761 (PRE).
NATAL. - 2729 (Volksrust): Top of Majuba (-BD), Van der Merwe
2776 (PRE); Chelmsford Nature Reserve (-DD), Smit 005 (NU). - 2730
(Vryheid): Hlangapies Mountain (-AB), llin der Merwe 1115 (PRE); Utrecht,
Farm Retirement (-AC), Hilliard & Bunt 18,565 (NU); Vryheid (-BD), llin der
Merwe 2463 (PRE); Germishuizen 2264 (PRE); Kambula Mountain (-DD),
Aeoeks 11,767 (NH). - 2731 (Louwsburg): Ntendeka Wilderness Area (-CD),
Van k~
6983 (PRU). - 2828 (Bethlehem): Royal National Park (-DB), Bews
337 (NU); Mount-aux-Sources (-DD), Edwards 314 (NU); Sehweiekerdt s.n.
sub PRE 34,919 (PRE). - 2829 (Harrismith): Tintwa Spruit (-AC), Van der
Merwe 2702 (PRE); Tintwa Mountain (-AD), llin der Merwe 6298 (PRE); Van
Reenen (-AD), Jaeobsz 1659 (PRE); Oliviers Hoek Pass (-CA), Gower 27
(NU); Bergville (-CB), Killiek 1009 (PRE); Cathedral Peak (-CC), Sehelpe 922
(NU). - 2830 (Dundee): Babanango (-BD), llin der Merwe 2787 (PRE); King
293 (PRE); Weenen, Griffins Hill (-CC), Aeoeks 10,678 (PRE); Umhlamba
Mountain (-CC), Aeoeks 13,867 (PRE). - 2831 (Nkandla): Hlabiza, Monzi (BB), Pooley 1874 (NU); Eshowe (-CD), Thode A1269 (PRE); Empangeni (-
LEDE:BOURIA IN SmITH AFRICA
234
TAXONOMY
DB), Robens 12 (BOL); Mtunzizni, Ngoye Forest (-DC), \enter 2487 (BLFU);
1068 (1); Mtunzini, Himewith (-DD), Mogg 4412 (PRE).
Lowrey & WIn k~
-2832 (Mtubatuba): Simbomvini (-AB), Nicolas & MacDevette 2141 (NH); St.
Lucia Resort (-AD), Pooley 1986 (NU); Pooley 2164 (NU). - 2929
(Underberg): Cathkin Park (-AB), Goossens 1941 (PUC); Giants Castle (-AB),
Symons 26,040 (PRE); Symons 517 (PRE); Estcourt, Ntabamhlope Vlei (-BA),
Downing 255 (NU); Van der Merwe 2576 (PRE); Mauve 4374 (PRE);
Ntabamhlope (-BA), W?st 621 (NBG); W?st 844 (PRE); Broadmoor Vlei (-BB),
Downing Z32 (NU); Downing 173 (NU); Mpenhle, Mulangane Ridge (-BC),
Hilliard & Bunt 16,972 (PRE); Impendhle (-BC), Huntley 72 (NU); Highmoor
Forest (-BC), du Toit 2556 (PRE); Mpendhle, Carters Creek (-BC), Hilliard &
Burtt 18,643 (NU); Hilliard & Burtt 16,972 (NU); Lions River, Kamberg (BD), Wright 1900 (NU); Wright 2259 (NU); Polela (-CB), Rennie 80 (NU);
Underberg, Bamboo Mountain (-CB), Grice s.n. (NU); Sani Pass (-CB), Gillies
42 (NU); Upper Tugela (-CC), Everson & Muller s.n. (CPF); Underberg,
Garden Castle (-CD), Van der Merwe 2763 (PRE); Underberg (-CD), Dyer
3265 (PRE); Himeville (-DC), Arnold 524 (PRE); Scott 32 (NU); Hiercel 73
(NU); Donnybrook (-DD), Van der Merwe 2758 (PRE); WIn der Merwe 2759
(PRE). - 2930 (Pietermaritzburg): Rietvlei (-AB), Fry s.n. sub PRE 5731
(PRE); Lions River, Umgeni Poort (-AC), Moll 1373 (NU); Moll 2428 (NU);
Moll 1388 (NU); Lions River (-AC) , Moll 1181 (NU); Lions River, Balgowan
(-AC), Moll 1197 (NU); Howick, Midmar (-AC), Moll 1110 (NU); Nottingham
Road (-AC) , Galpin 10,265 (PRE); Dargle Road (-AC) , Mogg 5698 (PRE);
Mogg 5701 (NH); Cramond, Blinkwater (-AD), 'WJnmann 3 (NU); Greytown (BA), eil~
s.n. sub BOL 22,516 (BOL); Dalton (-BC), Hardley 46 (NU);
Swartkop (-CB), MacDevette 1404 (NH); Pietermaritzburg (-CB), Allsopp 478
(NH); \enter 13,194 (UNIN); World's View (-CB), Moll 2628 (NU); Fox Hill
(-CB), Fisher 402 (NU); Mgeni Poort (-DA), Moll & Mauve 2428 (PRE);
Umzinyathi (-DB), 'WJod 1356 (NH); Inanda Swamp (-DB), 'WJod 1356 (NH);
Inanda (-DB), Wood 1056 (NH); Eisdumbeni (-DB), Wood 4791 (NH);
Krantzkloof Nature Reserve (-DD), Collocott 23 (NU); Hayganh s.n. sub STE
84 (STE); Isipingo Flats (-DD), Ward 6813 (PRE); Kloof, Highbury (-DD),
Landsdell s.n. sub NH 42,771 (NH). - 2931 (Stanger): Mapumulo, Thring's
Post (-AA), Moll 2316 (NU); Brighton Beach (-CC), Lutchminarain 28
(DWEST); Durban Bluff (-CC), Hennessy s.n. (DWEST); Durban (-CC), Singh
LEDEBOUR1A IN SOlJTH AFRICA
235
TAXONOMY
24 (DWEST); Badri 28 (DWEST). - 3028 (Matatiele): Matatiele (-BD),
McLoughlin s.n. sub BOL 50,774 (BOL). - 3029 (Kokstad): Mt. Currie (-AD),
liln der MelWe 2109 (PRE); Sneezewood Plantation (-BC), Strey 9175 (PRE);
Mvenyani (-CA), Bandert 115 (GRA); Kokstad (-CB), Bayliss 2497 (NBG);
Weza State Forest (-DA), Balkwill et al. 2368 (J). - 3030 (Port Shepstone):
Ixopo (-AA), Mogg 2300 (PRE); Kenterton (-AB), Thode s.n. sub STE 3388
(STE); Dumisa Station (-AD), Rudatis 704 (PRE); Isipingo Beach (-BB), Wlrd
972 (NU); Isipingo (-BB), Jankee 16 (DWEST); Pennington (-BC), "W?intraub
s.n. sub J 31,814 (1); Alexandra (-BC), Rudatis 217 (STE); Umtentwini (-CB),
Vogel s.n. sub NBG 48/61 (NBG); Uvongo (-CD), Mogg 13,540 (PRE);
Southbroom (-CD), Wulff28 (NU); Umzumbi (-DA), Van der Merwe 2534
(PRE). - 3130 (Port Edward): Port Edward (-AA), Stirton 12,141 (UNIN); liln
der MerWe 2664 (PRE); Moss 19,144 (J); Umtamvuna Nature Reserve (-AA),
Abbott 3993 (PRU); Strey 4487 (NH); Stirton 12,085 (UNIN); Leisure Bay (-
AA), Ubbink 80A (PUC).
ORANGE FREE STATE. - 2727 (Kroonstad): Vals River (-AC), Pont
520 (PRE). - 2827 (Senekal): Steynsrust (-AB), Grobler 89 (BLFU); Doornkop
(-DD), Goossens 777 (PRE). - 2828 (Bethlehem): Clarence (-CB), liln Hoepen
18,269 (PRE). - 2829 (Harrismith): (-AC), Compton 22,491 (BOL); Compton
13,218 (NBG); Harrismith, Farm Bloemhof (-AC), liln der Zyde s.n. sub NBG
133,281 (NBG). - 2926 (Bloemfontein): Naval Hill (-AA), du Preez 8 (PRE);
Thaba Nchu (-BB), Roberts 2866 (PRE). - 2927 (Maseru): Mensvretersberg (AA), Peeters et al. 323 (J).
LESOTHQ. - 2828 (Bethlehem): Butha Buthe (-CC), Ashton s.n. sub
BOL 22,525 (BOL); Leribe, Dieterlen 57c (NBG); Dieterlen 57 (NBG);
Dieterlen 662 (STE). - 2927 (Maseru): Roma (-AC), Schmitz 7959 (PRE);
Dieterlen 57a (PRE); Tebetebeng River (-BB), Guillarmod 364 (PRE). - 2928
(Marakabei): Semonkong (-CA), Davidson 3037 (J). - 3028 (Matatiele): Thaba
Tsueu (-AB), Page s.n. sub BOL 22,489 (BOL).
TRANSKEI. - 3128 (Umtata): Nqadu (-BC), Hutchings 832 (KEI);
Mjika, Mahlahlane Forestry (-BC), Hutchings 1313 (KEI). - 3129 (Port St.
Johns): Mgodini River (-BA), Strey 10,083 (PRE); Ntsubane Forest Station (-
LED£BOURlA IN SOlITH AFRICA
236
TAXONOMY
BC), Venter & Vorster 52 (PRE); Magwa Falls, Balkwill et al. 1869 (J);
Hutchings 774 (KEI); Magwa Tea Estate, Hutchings 1183 (KEI); Germishuizen
1182 (PRE); Coffee Bay (-CC), Tyson s.n. sub PRE 57,458 (PRE).
CAPE. - 2824 (Kimberley): Rooipoort (-CA), Wilman s.n. sub BOL
22,513 (BOL); - 3024 (De Aar): Katberg (-CC), Young 15,011 (J). - 3121
(Fraserburg): Fraserburg (-DC), Nel s.n. sub STE 15,814 (STE). - 3227
(Stutterheim): Cathcart, Fairford (-AC), Cotterrell131 (GRA); Pirie (-CC), Sim
618 (PRE); King Williams Town (-CD), Norris s.n. (NBG); Sim 1702 (NU);
Gonubie River (-DA), Carter s.n. sub BOL 22,517 (BOL). - 3228
(Butterworth): Kentani (-CB), Pegler 297 (PRE); Pegler 1141 (PRE); Pegler
1464 (PRE). - 3318 (Cape Town): Claremont (-CD), Schlechter 2959a (PRE). 3326 (Grahamstown): Trappe's Valley (-BD), Daly 582 (GRA). - 3421
(Riversdale): Glen Leith (-AB), Muir 3004 (PRE). - 3423 (Knysna): Knysna
Heads (-AA), Fourcade 2030 (BOL).
Without precise locality
- Orange Free State, Cooper 993 (K, PRE).
- Natal, Adlam s.n. (K, PRE).
- Cape, Cooper s.n. (K, PRE); Cape Colony, Rogers s.n. (K, PRE).
LED£BOURJA IN SOlITH AFRICA
237
TAXONOMY
Subsectio Piliferae S. Venter, subsect. , novo foliis adaxialiter pilosis, pHis
seriatim longitudinaliter dispositis.
Species typica: L. parvijolia S. Venter.
Species: L. parvijolia S. Venter.
Basal stem present. Lamina with rows of hairs on adaxial surface.
Inflorescence solitary, erect; rachis smooth. Bracteoles always absent.
20. LEDEBOURIA PARVIFOLIA S. Venter
Ledebouria parvifolia S. Venter sp.nov., ad L. cooperi (Hook./.) Jessop
cognata sed foliis applanatis, lamina adaxiali pilis longitudinaliter seriatum
instructa et inflorescentia sine bracteolis valde distincta.
Type: Transvaal, Graskop, near Lisbon Falls, farm Lisbon 531 KT, lenter s. n.
(PRE!, holo.).
Plants solitary. Bulb hypogeal, 10 - 25 x 10 - 25 mm, obovoid; dead bulb scales
brown, membranous, apices attenuate, live bulb scales fleshy, tightly arranged,
without threads when torn, white inside; bulblets present on basal stem; basal
stem 2 - 4 mm long. Cataphyll one, exserted above ground level. Leaves fully
developed at flowering, 2 - 3, appressed to ground, lanceolate to oblong, 15 30 x 15 - 20 mm, with threads when torn, fleshy, surfaces dull green, adaxial
surface with rows of hair-like papillae, venation obscure; leaf margin ciliate;
leaf base canaliculate; apex acute. Inflorescence solitary, lax, elliptic, 15 - 20 x
10 - 15 mm, erect, 8 - 12 -flowered, longer than the leaves; scape terete at base,
purple, glabrous; rachis smooth, 10 - 15 mm long. Bracts fleshy, 0.5 x 0.5
mm, deltoid, pink to purple without bracteoles. Pedicels cernuous, 3 mm long,
pink. Perianth 3 mm long, tepals recurved, equal, oblong, 3 x 1 mm, apex
obtuse, pink with a green keel. Stamens erect, 3 mm long, pink, epitepalous;
anthers 0.5 mm long, pale violet. Ovary ovoid, 3 -lobed, 1 x 2 mm, lobes
narrowly transversely oblong, apex shoulders raised. Style 3 mm long,
triangular, glabrous, purple; stigma equal height than anthers; stipe 0.5 x 0.5
mm. Capsule three-lobed, symmetrical, globose; base truncate. Seed dropshaped, 3 mm long, drop-shaped, surface strongly wrinkled, brown. (Figure
54).
UDEBOURJA IN SOlITH AFRICA
238
TAXONOMY
o
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Figure 54. Illustration of L. parvijolia S. Venter. A, habit X 2; B, lamina
margin X 300; C, section through lamina X 3; D, bract X 10; E, flower XIS;
F, tepal with stamen XIS; G, tepal apex X 20; H, ovary lateral view X 10; I,
ovary dorsal view X 10. All from
retn~
s.n.
LEDEBOURlA IN SOlITH AFRICA
Map 24. .Known distribution of L. parvifolia S. Venter.
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240
TAXONOMY
Figure 55. A, holotype of L. parvifolia S. Venter (PRE); B, plant showing the
bulbils, small leaves and the solitary inflorescence. Bar
= 20 mm; C, SEM
micrograph of the adaxial lamina surface. Bar = 50 J.Lm; D, SEM micrograph
of the abaxial lamina surface; E, SEM micrograph of stomata; F, habitat near
Pilgrim s Rest, eastern Transvaal. The vegetation consists of closed deciduous
I
low Dombeya rotundifolia subsp. rotundifolia - Acacia ataxacantha - Scilla
natalensis woodland; G, SEM micrograph of the rows of hairs on the adaxial
lamina surface; A - F from Venter s. n..
UDEBOURlA IN sourH AFRICA
241
TAXONOMY
HERBARIUM
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LEDEBOURlA IN SOlJfH AFRICA
-~.
242
TAXONOMY
L. parvifolia is related to L. cooperi but differs in its humifuse leaves with
longitudinal rows of hair-like papillae, solitary, erect, lax inflorescence and 3
-lobed ovary.
Specific epithet etymology.
Describes the small leaves.
Flowering period
From October to December.
Distribution (Map 24).
Endemic to the eastern Transvaal, known from a single locality.
Habitat
L. parvifolia grows on the Malmani Formation dolomite of the Chuniespoort
Group in the Transvaal Sequence (SACS 1980). Soil derived from these rocks is
a shallow (l0 - 120 mm deep), dark brown, fine grained (0.025 - 0.25 mm <t»
clay loam with many rock particles. Most of the hill slopes are comprised of
dolomite rocks and cliffs and covered in montane grassland with scattered
Dombeya rotundifolia (Hochst.) Planch. var. rotundifolia and Faurea speciosa
(Welw.) Welw. trees (Figure 55G).
Specimens examined
TRANSVAAL. - 2430 (Pilgrim s Rest): Graskop, Lisbon Waterfall,
farm Lisbon 531 KT, Venter s.n. (PRE).
I
LEDEBOURlA IN SOIlfH AFRICA
243
TAXONOMY
Sectio Globosae S. Venter, sect. nov., bulbis maturis 20 - 40 (-60) mm dia.;
capsulis sigallatim globosis.
Species typiCa: Ledebouria inquinata (C.A. Srn.) Jessop.
(K!).
Typus: Transvaal, near Pretoria, along Aapies River, Burke s.n.
Species: L. asperijolia (Van der Merwe) S. Venter, L. glauca S. Venter, L.
inquinata (C.A. Srn.) Jessop and L. marginata (Bale.) Jessop.
Plants solitary. Leaves spreading to erect-spreading, producing threads when
torn. Inflorescences with raceme dense, oblong; rachis ridged; bracts
membranous. Capsule distinctly globose.
Distribution and habitat
Eastern and northern Cape to northern Transvaal. Grassland and woodland.
Subsectio Asperifoliae S. Venter, subsect. nov., foliis scapo dense asperatis.
Species typica: L. asperijolia (Van der Merwe) S. Venter.
Species: L. asperijolia (Van der Merwe) S. Venter.
Leaves and scape with asperities. Rachis ridged; bracteoles absent.
2l.LEDEBOURIA ASPERIFOLIA (Van der Merwe) S. Venter
Ledebouria asperifolia (Van der Merwe) S. Venter comb.nov.
Scilla asperijolia Van der Merwe in Flower. Pi. S. Afr. 24: t.944 (1944).
Type: Natal, hillsides at Ladysmith, Wm der Merwe 2604 (PRE!, holo.; NU!).
Plants solitary. Bulb hypogeal, 40 - 50 X 40 - 50 mm, subglobose; dead
bulb scales purplish-brown, apices attenuate, live bulb scales membranous, tightly
arranged, without threads when torn, white inside, sometimes forming a neck 10
- 20 x 5 - 10 mm. Leaves fully developed at anthesis, 4 - 8, spreading,
lanceolate to ovate, 100 - 170 x 25 - 100 mm, with threads when torn, fleshy,
with a dull lustre, adaxial surface green often with dull dark green to purple
LF.DEBOURlA IN SOurH AFRICA
244
TAXONOMY
blotches, abaxial surface either green, basally purple or the whole under surface
purple, venation mostly prominent, with rows of tongue-shaped asperities,
sometimes only adaxial; margins finely papillate; leaf base canaliculate; apex
acute to obtuse. Inflorescences 1 - 3, dense, elliptic, 50 - 120 x 20 - 30 mm,
flaccid, 30 - 50 -flowered, as long or longer than the leaves; scape terete at
base, green to purple, covered with tongue-shaped papillae; rachis ridged, 50 150 mm long. Bracts membranous, 1 x 0.5 mm, lanceolate, grey-white with
bracteoles absent. Pedicels spreading, 10 - 20 mm long, white to pink. Perianth
5 mm long, tepals recurved, subequal, oblong, 5 x I mm, apex cuspidate,
cucullate, pink to purple with a brown keel. Stamens erect, 5 mm long,
filaments maroon, epitepalous; anthers I mm long, pale violet. Ovary
ellipsoidal, 6 -lobed, 1 x 2 mm, lobes narrowly transversely elliptic, apex
shoulders raised. Style 5 mm long, terete, glabrous, purple; stigma equal height
to anthers; stipe 1 x 0.5 mm. Capsule one- to three-lobed, asymmetrical,
globose; base truncate. Seed globose, 3 mm long, surface strongly wrinkled,
brown. (Figure 56).
The only species with which L. asperijolia can be confused, is L. revoluta. The
tongue-shaped asperities on the leaves and scape are however diagnostic.
Specific epithet etymology.
Referring to the asperity covered leaves.
Flowering period
From September to December.
Distribution (Map 25).
Occurs in Natal, a few localities in Swaziland and the mountainous areas near
Barberton, the Transvaal Drakensberg escarp and as far north as Pietersburg.
LEDEBOURlA IN SOIIfH AFRICA
245
TAXONOMY
B
Figure 56. Illustration of L. asperijolia (Van der Merwe) S. Venter. A, habit X
1; B, lamina margin X 300; C, leaf apex X 10; D, section through lamina X 4;
E, bract X 10; F, tepal apex X 20; G, flower X 10; H, tepal with stamen X 10;
I, ovary lateral view X 10; J, ovary dorsal view X 10. All from l-enter 13,249.
LEDEBOURJA IN SOtTl1l AFRICA
Known distribution of L. asperijolia (Van der Merwe) S. Venter
Map 25.
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247
TAXONOMY
Figure 57. A, holotype of L. asperijolia (Van der Merwe) S. Venter (PRE); B,
habitat on the the Makonjwa Mountain between Barberton and Havelock. Plants
were collected (Venter 13,382) on the cliffs in the foreground; C, plants of L.
asperijolia after a recent veld fire, Lisbon Falls, Graskop. Bar = 100 mm; D,
plant from Barberton showing the rows of asperities on the leaves. Bar = 10
mm; E, SEM micrograph of the adaxial lamina surface. Bar = 100 JLm; F,
SEM micrograph of the abaxial lamina surface. Bar = 100 JLm; G, SEM
micrograph of stomata. Bar = 43 JLm. C from Venter 13,249 and D - G from
Venter 13,382.
LEDEBOURJA IN SOUTH AFRICA
248
TAXONOMY
LEDEBOURlA IN SOlJfH AFRICA
249
TAXONOMY
Habitat
L. asperijolia occurs on dolerite dykes, sandstones, shales, quartzites and
conglomorates, banded ironstone, chert, dolomite and granite. L. asperijolia is
not restricted by soil type, but mOst of the populations occur either on clay
loams or sandy loams.
During this survey, L. asperijolia was encountered growing in various woodland
types. The southern populations are all associated with Acacia woodland.
Population structure
Plants grow mostly as scattered individuals except near Pilgrim's Rest where, at
the foot of dolomite cliffs, isolated colonies were encountered (Figure 57C).
Variation
Leaves of plants growing in the shade tend to be longer and narrower, not as
fleshy, and with less asperities. Markings on the leaves vary considerably.
Mottling of the leaves is uncommon in L. asperijolia, although plants with
prominent purple mottling occur in the mountains around Barberton (Figure
57D).
Historical background
This species was discovered and described by Dr. EZ. van der Merwe from
collections made near Ladysmith, Natal. Jessop (1970) in his studies on the
bulbous Liliaceae, placed S. asperijolia under L. revoluta.
UDEBOURIA IN SOlITH AFRICA
250
TAXONOMY
Specimens examined
TRANSVAAL. - 2329 (Pietersburg): Pietersburg (-CD), Venter s.n.
(UNIN). - 2430 (Pilgrim's Rest): Ohrigstad Nature Reserve (-DC), Jacobsen
1839 (PRE); Graskop, Lisbon Falls (-DD), Venter 13,249 (UNIN). - 2530
(Lydenburg): Between Lydenburg and Houtbosloop (-AB), lbn der Menve 1784
(PRE); Farm Nooitgedacht (-AB), lbn der Menve 1777 (PRE); Belfast, farm
Schoongesicht (-CA), lbn der Menve 1239 (PRE); Carolina, tunnel at Waterval
Boven (-CB), lbn der Menve 1245 (PRE); Starvation Creek Nature Reserve (DA), Kluge 1104 (PRE). - 2531 (Komatipoort): Barberton, farm Oosterbeek (CC), Venter 13,382 (UNIN); Tienie Louw Nature Reserve (-CC), Buitendag
1140 (PRE).
SWAZILAND. - 2631 (Mbabane): Manzini, Malkerns (-CA), Compton
s.n. (PRE).
NATAL. - 2731 (Louwsburg): Pongola, Magut (-DA), WIn der Menve
2715 (PRE). - 2829 (Harrismith): Ladysmith (-DB), Van der Merwe 2604
(PRE). - 2830 (Dundee): Weenen (-CC), Acocks 13,863 (PRE); Muden
Townlands (-CD), l\Ylie s.n. (NH). - 2832 (Mtubatuba): Hluhluwe Game
Reserve (-AA), Ward 3288 (NU). - 2930 (Pietermaritzburg): Otto's Bluff (AD), Feldman s.n. (NH); lOm north of Greytown (-BA), Mogg s.n. (PRE);
Epol Hill (-CB), Noel 2720 (NU).- 2931 (Stanger): Eshowe, Mandini (-AB),
Edwards 1598 (NU). - 3029 (Kokstad): Harding (-DB), lbn der Menve 2756
(PRE).
LEDEBOURIA IN SOlJfH AFRICA
251
TAXONOMY
Subsectio Glaucae S. ,retn~
subsect. nov., foliis glaucis; bracteolis bracteam
aequantibus.
Species typica: L. glauca S. Venter.
Species: L. glauca S. Venter, L. inquinata (C.A. Srn.) Jessop, L. marginata
(Bale) Jessop.
22. LEDEBOURIA GLAUC4. S. Venter
Ledebouria glauca S. retn~
sp.nov., ad L. cooperi (Hook!) Jessop cognata sed
foliis coriaceis, glaucis; inflorescentiis erectis, quam foliis brevioribus; ovario
basi lobato manifeste differt.
Type: Transvaal, Pietersburg, Fauna Park,
~nter
13,368 (PRE! holo.; NU!)
Plants solitary. Bulb hypogeal, 15 - 70 x 15 - 30 mm, cylindrical; dead
bulb scales light brown, apices truncate, attenuate only at the extreme tips, live
bulb scales fleshy, with threads when torn, white inside, neck 3 - 30 x 3 - 10
mm; basal stem 5 - 50 x 10 - 20 mm. Leaves fully developed at anthesis, 2 - 4,
spreading, oblanceolate, 50 - 100 x 8 - 18 mm, with threads when torn,
leathery, dull, glaucous with purple stripes and blotches, purple cross bars at
base of leaf, venation obscure; margins finely papillate; leaf base canaliculate;
apex acute. Inflorescences 1 - 2, dense, oblong to cylindrical, 30 - 50 x 10 - 20
mm, erect, 16 - 60 -flowered, shorter than leaves; scape terete at base, spotted
purple, glabrous; rachis ridged, 20 - 30 mm long. Bracts membranous, 1 - 2 x
0.5 mm, lanceolate to bifurcate, white to pinkish purple, bracteoles present.
Pedicels spreading horizontally; 5 - 8 mm long, pink. Perianth 5 mm long,
tepals recurved, equal, oblong, 5 x 1.5 mm, apex obtuse, cucullate, green and
pink to purple with a green keel. Stamens erect, 3 - 4 mm long, filaments
slightly flattened at base, maroon, epitepalous; anthers 1 mm long, yellow.
Ovary ovoid, 6 -lobed, 0.5 x 1 mm, lobes narrowly transversely o~lng,
distal
lobes present. Style 3 mm long, triangular, glabrous, purple; stigma equal
height to anthers; stipe 0.25 x 0.25 mm. Capsule three-lobed, symmetrical,
globose; base truncate. Seed drop-shaped, 3 - 5 mm long, surface strongly
wrinkled, brown. (Figure 58).
LEDEBOURJA IN SOIITH AFRICA
252
TAXONOMY
G\
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E
D
c
G
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8
Figure 58. Illustration of L. glauca S. Venter. A, habit X 1; B, lamina margin
X 300; C, section through lamina X 5; D, bract with bracteole X 10; E, apex of
tepal X 20; F, tepal with stamen X 10; G, flower X 10; H, ovary lateral view X
10; I, ovary dorsal view X 10. All from \enter 13,368.
LEDEBOURlA IN SOIJTH AFRICA
Known distribution of L. glauca S. Venter
Map 26.
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TAXONOMY
Figure 59. A, holotype of L. glauca S. Venter (PRE); B, habitat in the Kalahari
near Vanzylsrus. The vegetation consists of open low Acacia erioloba - A.
haematoxylon - Stipagrostis uniplumis woodland; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of stomata. A and C - E from Wnter 13,368.
LEDEBOURIA IN SOurH AFRICA
255
TAXONOMY
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LEDEBOURIA IN SOlITH AFRICA
256
TAXONOMY
L. glauca is closely related to L. inquinata (C.A. Srn.) Jessop and L. marginata
(Bale) Jessop, but differs in having a basal stem and thickly leathery leaves.
Specific epithet etymology.
Refering to the glaucous colour of the plant.
Flowering period
From October to December.
Distribution (Map 26).
Known to occur in the Transvaal and the northern Cape.
Habitat
L. glauca occurs on sandy soils of various textures and origins favouring open
grassy and sandy areas in full sun, between the trees and shrubs of various
woodland types (Figure 59B). Various Acacia species and Grewia flava DC.
tend to be the dominant woody members.
Plants occur mostly as scattered individuals in the veld. Groups of three to seven
plants occur occasionaly near Van Zylsrust (northern Cape). This species is
rare. On few occasions seedlings were noted growing around the adult plants
(within a radius of four meters).
Historical background
The first record of L. glauca appears to be Acocks 1427 collected during
December 1936. Since then, few collections have been made mostly from the
northern Transvaal and northern Cape.
LEDEBOURIA IN SOlITH AFRICA
257
TAXONOMY
Specimens examined
TRANSVAAL. - 2229 (Waterpoort): Vivo, Langjan Nature Reserve (CC), Zwanziger 539 (PRE). - 2329 (Pietersburg): Pietersburg, Flora Park (CD), lenter 13,368 (PRE, UN, UNIN). - 2528 (pretoria): La Montagne (-CA),
van
k~
2414 (PRE). - 2626 (Klerksdorp): Ventersdorp, farm Somerville (BD), Louw 2980 (PUC).
CAPE. - 2520 (Mata Mata): Kalahari Gemsbok Park (-CD), WIn Rooyen
2904 (PRU); Leistner 1001 (PRE). - 2624 (Vryburg): Vryburg, farm Zoetvlei (DC), Speedy
91
(NBG). - 2722 (Olifantshoek): Katok (-DC), de Lange 136
(UNIN). - 2823 (Griekwastad): Campbell (-DC), Acocks 1427 (PRE).
LEDEBOURIA IN SOlITH AFRICA
258
TAXONOMY
23. LEDEBOURIA INQUINATA (C.A. Srn.) Jessop
Ledebouria inquinata
rc.A.
Sm.) Jessop in 11 S. Afr. Bot. 36(4): 257 (1970).
Scilla inquinata C.A. Smith in Kew Bull. :248 (1930).
Type: Transvaal, near Pretoria along Aapies River, Burke s. n. (Kl, holo.;
PREl, photo.). Designated here as lectotype (Greuter et al. 1988. Article 7.5).
Plants solitary. Bulb hypogeal, 30 - 35 x 30 - 35 mm, ovoid; dead bulb scales
brown speckled with purple apically, membranous, apices attenuate, live
bulb scales tightly arranged, with sparse threads when tom, purplish inside, neck
8 - 10 x 5 - 8 mm. Leaves partly emerged at anthesis, 6 - 10, spreading, linearlanceolate to lanceolate, 50 - 80 x 8 - 20 mm, with threads when torn, fleshy,
surfaces dull glaucous green with dark green to purple markings, venation
obscure; margins smooth, undulate at base; leaf base shallowly canaliculate;
apex acute. Inflorescences 2 - 3, dense, oblong, 20 - 60 x 12 - 30 mm, flaccid,
40 - 60 -flowered, shorter than the mature leaves; scape terete at base, green
spotted dull purple, rachis ridged, 15 - 30 mm long. Bracts membranous, 1.5 x
1 mm, linear to lanceolate, grey to white with bracteoles. Pedicels spreading, 7
mm long, pink. Perianth 4 mm long, tepals recurved, equal, oblong, 4 x 1.5
mm, apex obtuse, cucullate, pink to purple with a brown keel. Stamens erect, 2
mm long, filaments maroon, epitepalous; anthers 0.5 mm long, yellow. Ovary
depressed globose, 6 -lobed, 1 x 1.8 mm, lobes obtusely deltate, apex shoulders
raised, basal lobes present. Style 3 mm long, triangular, glabrous, purple;
stigma above anthers; stipe 0.2 x 0.4 mm. Capsule three-lobed, symmetrical,
globose; base truncate. Seed globose, 2 - 3 mm long, surface strongly wrinkled,
brown. (Figure 60).
Together with L. glauca and L. marginata (Bak.) Jessop, L. inquinata belongs
to the subsection Glaucae but is distinguished by the purple spots and blotches
on the live bulb scales (Figure 61D), leaves undulate at the base and depressed
globose ovary.
LED£BOURIA IN SOUTH AFRICA
259
TAXONOMY
E
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Figure 60. Illustration of L. inquinata CC.A. Srn.) Jessop. A, habit Xl; B,
lamina margin X 300; C, apex of lamina X 10; D, section through lamina X 5;
E, bract with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H,
apex of tepal X 20; I, ovary lateral view X 10; J, ovary dorsal view X 10. All
from lenter 13,343.
u:DEBOURlA IN SOlTTH AFRICA
Known distribution of L. inquinara CC.A. Srn.) lessop
Map 27.
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TAXONOMY
Figure 61. A, lectotype of L. inquinata (C.A. Srn.) Jessop (K); B, isotype of L.
inquinata designated by Smith as co-type (K); C, habitat at Linksfield Ridge,
Bedfordview, with open evergreen Protea caffra subsp. caffra - Aloe
greatheadii var. davyana - Digitaria eriantha woodland; D, plant showing the
diagnostic moUling on the live bulb scales. Bar = 25 mm; E, SEM micrograph
of the adaxial lamina surface; F, SEM micrograph of the abaxial lamina surface;
G, SEM micrograph of stomata. D - G from W?nter 13,343.
LEDEBOURIA IN SOtJrH AFRICA
262
TAXONOMY
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LEDEBOURIA IN SOurH AFRICA
--
263
TAXONOMY
Specific epithet etymology.
Means stained, describing the prominent purple blotches on the dry bulb scales.
Flowering period
From August to November peaking in September.
Distribution (Map 27).
Transvaal and the southern part of Botswana.
Variation
Plants from near Graskop tend to be larger and more robust ('W?nter 13,241)
than plants from the central and western Transvaal.
Jessop (1970) apparently failed to see the differences between the type
specimens and material he recognized as L. inquinata. This material is in fact an
undescribed species related to L. atro-brunnea .
Specimens examined
TRANSVAAL. - 2429 (Zebediela): Potgietersrust (-AA), Leendertz
1453 (PRE). - 2430 (Pilgrim's Rest): Graskop, Lisbon Waterfall (-DD), lCnter
13,241 (UNIN). - 2527 (Rustenburg): Rustenburg (-CA), Leendertz 10,140
(PRE); Rustenburg Nature Reserve (-CA), Coetzee 1513 (PRE); lCnter 13,231
(UNIN). - 2528 (Pretoria): Hammanskraal, farm Kromdraai (-AD), 'W?nter
13,335 (UNIN); Pretoria (-CA), Burtt Davy 2200 (PRE); McLea 101 (BOL);
Tillwich s.n. (PRU); Moss 10,810 (1); Leendertz 4049 (PRE); Horns Neck (-
CA), Perry 2019 (NBG); Meintjies Kop (-CA), Mogg 14,077 (PRE);
LEDEBOURlA IN SOlITH AFRICA
264
TAXONOMY
Queenswood (-CA), Theron 193 (PRU); Groenkloof (-CA), Mogg 3009 (PRE);
Onderstepoort (-CA), Smith 6035 (PRE); Waverley (-CB), Venter 13,344
(UNIN); Venter 13,345 (UNIN); Pienaars Poort (-CB), Leeman s.n. (PRE);
Roodeplaat Dam (-CB), Van Rooyen 2666 (PRU); Fountains Valley (-CC),
Venter 13,350 (UNIN). - 2529 (Witbank): Middelburg, Tautes Berg (-BB),
Young A148 (PRE); Hartebeeshoek (-DB), Verdoorn & Mogg s.n. (PRE). 2626 (Klerksdorp): Ventersdorp, farm Goedgedacht (-BD), Sutten 630 (PUC);
Ventersdorp, farm Morgenzon 42 (-BD), Mogg 22,209 (J). - 2627
(Potchefstroom): Welverdiend (-AD), Louw 3 (PRU, PUC); Krugersdorp, farm
Zwartkrans (-BA), Franklin 10 (1); Sterkfontein Caves (-BB), Mogg 35,540 (1);
Roodepoort (-BB), Moss 13,418 (1); Potchefstroom (-CA), Ubbink 692 (PUC);
Boskop (-CA), Louw 360 (PUC); Dassiesrant (-CA), Venter 13,222 (UNIN);
Scandinavia Drift (-CC), Venter 13,227 (UNIN). - 2628 (Johannesburg):
Johannesburg (-AA), Norwood Young 786 (J); Moss 18,927 (1); Milner Park (AA), Moss 13,703 (J); Sandton, Buccleuch (-AA), Venter 338 (UNIN);
Witwatersrant (-AA), Moss 19,078 (1); Killarney (-AA), Page s.n. (J);
Westc1iffe (-AA), Moss 12,071 (J); Melville Koppies (-AA), Knep & Higgs s.n.
(1); West Morningside (-AA), Gilliland s.n. (J); Bryanston (-AA), Dahlstrand
363T (PEU); Thorntree Kloof (-AA), Moss s.n. sub J 31,789 (J); Edenvale (AA), Venter 13,338 (UNIN); Alberton, Klipriviersberg (-AC), Gilliland s.n. (1,
PRE); Krugersdorp, Witpoortjie (-DC), Rugo s.n. sub STE 31,924 (STE).
LEDEBOURlA IN SOlJTH AFRICA
265
TAXONOMY
24. LEDEBOURIA MARGINATA (Bale) Jessop
Ledebouria marginata (Bak.) Jessop in 11 S. Afr. Bot. 36(4): 260 (1970).
Scilla marginata Bak. in Bull. Herb. Boiss. ser. 2(4): 1002 (1904).
Type: Transvaal, Modderfontein, Conrath 703b (GRAZ!; PRE!, photo.; Z.).
Scilla neglecta Van der Merwe in Flower. PI. S. Afr. 22: t.865 (1942).
Type: Transvaal, Pretoria, Colbyn, llin der Merwe 2441 (PRE!, holo.).
Plants solitary. Bulb hypogeal, 60 - 80 x 30 - 50 mm, obovoid; dead bulb scales
light to dark brown, membranous, apices attenuate, live bulb scales tightly
arranged, with threads when torn, white inside, neck 50 - 70 x 20 mm. Leaves
partly emerged at anthesis, 4 - 10, erect to erect-spreading, spirally twisted,
lanceolate, 120 - 150 x 10 - 100 mm, with thick bundles of threads when tom,
difficult to tear, leathery, dull glaucous green with dark green to purple spots
and blotches with cross bars at base of lamina on the abaxial surface, venation
prominent; margins smooth, basal part usually undulate; leaf base nearly flat to
shallowly canaliculate; apex acute. Inflorescences 4 - 10, dense, oblong; 20 100 x 20 - 50 mm, flaccid, 80 - 100 -flowered, as long or longer than leaves;
scape winged at base, green spotted darker; rachis ridged, 50 - 100 mm long.
Bracts membranous, 3 x 0.5 mm, linear to lanceolate, grey to white with
bracteoles. Pedicels spreading, 3 - 10 mm long, pink. Perianth 3 - 4 mm long,
tepals recurved, subequal, oblong to lanceolate, 3 - 4 x 1.5 mm, apex obtuse,
cucullate, pink to purple with green to brown keel. Stamens erect, 3 mm long,
filaments maroon, epitepalous; anthers 0.75 mm long, yellow. Ovary globose,
6 -lobed, 0.75 x 1.5 mm, lobes narrowly oblong, apex shoulders raised. Style 3
mm long, triangular, glabrous, upper half purple, lower half white; stigma equal
height to anthers; stipe 0.12 x 0.2 mm. Capsule three-lobed, symmetrical,
globose; base truncate. Seed drop-shaped, 2 - 3 mm long, surface strongly
wrinkled, yellowish brown. (Figure 62).
L. marginata is closely related to L. inquinata and L. glauca but is easily
distinguished by the tough glaucous leaves and many inflorescences.
LEDEBOURlA IN SOlJfH AFRICA
266
TAXONOMY
8
Figure 62. Illustration of L. marginata (Bak.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina Xl; D, bract with bracteole X 10; E.
tepal with stamen X 10; F, flower X 10; G, apex of tepal X 20; H, lamina of
the narrow leaved form from Vryheid X 0.5; I, section through lamina of H. X
2; J, ovary lateral view X 10; K, ovary dorsal view X 10. A - G and J - K from
\enter 13,327 and H - I from \enter 13,364.
LEDEBOURIA IN sourH AFRICA
Map 28.
12°
Known distribution of L. marginara (Bak.) Jessop
14°
16°
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268
TAXONOMY
Figure 63. A, lectotype of L. marginata (Bak.) Jessop (GRAZ); B, SEM
micrograph of the adaxial lamina surface; C, SEM micrograph of the abaxial
lamina surface; D, SEM micrograph of a stomatum; E, plant showing diagnostic
twisted leaves. Bar = 30 mm; F, plants of the narrow-leaved form from near
Vryheid. Bar
= 40 mm.
B - E from lenter 13,230 and F from lenter 13,358.
LEDEBOURlA IN SOlrrH AFRICA
269
TAXONOMY
W~I'ROU: ~ I V L'"~ .:=t I~ :C8
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LEDEBOURIA IN SOUTH AFRICA
"\11
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270
TAXONOMY
Specific epithet etymology.
In reference to the broad membranous lamina margin.
Flowering period
From August to February with a peak from September to November.
Distribution (Map 28 ).
Widely distributed in Transvaal, Natal, Orange Free State with scattered
localities in Lesotho, north-eastern and eastern Cape.
Habitat
Although some plants of L. marginata occur on fine to medium grained (0.025 0.25 mm cP) soils derived from andesitic tuff and schist, most plants occur on
soils derived from either sandstone, conglomorate or quartzite.
L. marginata is associated with grasslands and grassy areas in woodland but
rarely occurs in Acacia woodland on heavy clay soils. It is common in short
grassland but rare in tall grassland, where it occurs on rock outcrops. Only one
population, that on the farm Rhenosterspruit at Lanseria, was found in semishade underneath Acacia caffra (Thunb.) Willd. on dolomite rock outcrops in
closed deciduous low woodland (Venter 13,487).
L. marginata is susceptible to grazing. Leaf apices were eaten leaving the strong
fibres exposed.
LEDEBOURIA IN SOlJTH AFRICA
271
TAXONOMY
Variation
In L. marginata there are two groups with respect to leaf shape. The first
group, the largest, has typical L. marginata leaves but the second group has
long narrow leaves (Map 1., Fig. 62H) (Venter 13,358) and occurs in the
Vryheid - Piet Retief area. The leaves of these plants are all erect with a
characteristic spiral twist. Leaf colour varies. In the Krugersdorp area, plants
are immaculate (Venter 13,487) with those from Waverley, Pretoria, heavily
marked with dull purple (Venter 13,346).
The common flower colour is pink, with green keels to the perianth segments
but plants with entirely green flowers grow at the Lisbon Falls near Graskop
(Venter 13,246).
The spiral twist of the leathery, glaucous green leaves and the undulate lamina
margins are distinctive (Figure 63E).
Specimens examined
TRANSVAAL. - 2229 (Waterpoort): Langjan Nature Reserve (-CC),
Zwanziger 365 (PRE). - 2230 (Messina): Messina (-AC), Rogers 22,967 (1). 2329 (Pietersburg): Pietersburg (-CD), Moss 15,693 (1). - 2330 (Tzaneen):
Duiwelskloof (-CA), liln der Merwe s.n. (PRE). - 2425 (Gaborone): Zeerust,
farm Koedoesrant 9 KO (-DD), Krynauw 192 (LYD). - 2427 (Thabazimbi):
Rooiberg, farm Blaauwbank (-DC), liln der Merwe 1964 (PRE); Rooiberg (DC), Van der Merwe 2020 (PRE); Van der Merwe 2475 (PRE). - 2428
(Nylstroom): Nylstroom (-CB), Jacques s.n. (PRE); 19 Miles north of
Nylstroom (-CB), liln der Merwe 1954 (PRE); 5 Miles east of Warmbaths (_
CD), Collett s.n. (PRE). - 2429 (Zebediela): Potgietersrust (-AA), Leendertz
10,136 (PRE); Arabie (-CD), Ellery 288 (PRE). - 2430 (Pilgrim's Rest):
Graskop, Lisbon Falls (-DD), Venter 13,246 (UNIN). - 2431 (Acornhoek):
Hoedspruit (-AC), Van der Merwe 2004 (PRE). - 2524 (Vergelee): Molopo
Nature Reserve (-DD), Peeters et al. 222 (J). - 2526 (Zeerust): Zeerust (-CA),
WIn der Merwe 1213 (PRE). - 2527 (Rustenburg): Rustenburg (-CA), Nation
LEDEBOURLA IN SOlTfH AFRICA
272
TAXONOMY
351 (BOL); Rustenburg Nature Reserve (-CA), Coetzee 1518 (PRE); Kroondal
(-CB), Louw 3455 (PUC); Brits, farm Welgevonden (-DB), Mogg s.n. (PRE);
Witwatersrantberg, Skeerpoort (-DC), Bidlingmaier 7 (PRU); Magaliesberg,
farm Zeekoeihoek (-DC), Van der Merwe 2038 (PRE); Rustenburg, farm
Uitkomst 499 JQ (-DD), Coetzee 233 (PRE). - 2528 (Pretoria): Pretoria (CA),
Moss 11,712 (J); Knoppiesfontein (-CA), lhn der Merwe 1998 (PRE); Pienaars
River (-AB), U!nter 13,327 (UNIN); Van Riebeeck Nature Reserve (-CA), Kok
220 (PUC); Wonderboompoort (-CA), Repton 1818 (PRE); Roodeplaat Nature
Reserve (-CB), lhn Rooyen 2194 (PRE, PRU); Van Rooyen 3534 (PRU); lhn
Rooyen 2667 (PRU); Van Rooyen 1942 (PRU); Van Rooyen 2865 (PRU);
Brumeria (-CB), Codd 3143 (PRE); Baviaanspoort (-CB), Goossens 31 (PRE). 2529 (Witbank): Middelburg, farm Doornkop 273 JS (-CB), Du Plessis 567
(PRU). - 2530 (Lydenburg): Dullstroom (-AC), Van der Merwe 1273 (PRE);
Farm Verloren Vallei (-AC), Drews 48 (PRE); Sudwalas Kraal (-BC),
Liebenberg 3035 (PRE); Belfast (-CA), Van der Merwe s.n. (PRE). - 2625
(Delareyville): Wolmaranstad, farm Strydpoort (-DD), Sutton 466 (PRE). 2626 (Klerksdorp): Lichtenburg, farm Hakboslaagte (-AC), Kinges 1818 (PRE);
Ventersdorp, farm Goedgedacht (-DB), Sutton 628 (PUC). - 2627
(Potchefstroom): A. Bailey Nature Reserve (-AD), Van k~
272 (PRE);
Witpoortjie (-BB), Gilliland s.n. (PRE); Modderfontein (-BB), lhn der Merwe
1982 (PRE); lhn der Merwe 2010 (PRE); Randparkridge (-BB), Taylor 136 (J);
Krugersdorp (-BB), Codd 3157 (PRE); Roodepoort (-BB), Behr 15 (NBG);
Sterkfontein Caves (-BB), Lucas & Glen sub Mogg 34,650 (J); Mogg 36,694
(J); Mogg 34,564 (J); Potchefstroom (-CA), Botha & Ubbink 1543 (PRE);
Klington (-CA), Goossens 1660 (PRE). - 2628 (Johannesburg): Johannesburg (AA), Watt & Brandwyk 2254 (PRE); Frankenwald (-AA), Gilliland s.n. (1);
Milner Park (-AA), Moss 13,663 (1); Moss 7348 (1); Moss 19,060 (1);
Germiston(-AA), Dryden s.n. (BOL); Germiston, Lombardy Estate (-AA),
Holden 14,151 (PRE); Rietfontein (-AA), Moss 10,808 (1); Melville Koppies (AA), Macnae s.n. (1); Parktown (-AA), Walker s.n. sub J 27,357 (J); Mulder's
Drift road (-AA), Gilliland s.n. (1); Forest Hill (-AA), Moss 10,026 (J);
Houghton Estate (-AA), Moss 2162 (1); Bryanston (-AA), Gilliland s.n. (1);
Lloyd s.n. sub J 26,620 (1); Van Wyk's Rust (-AA), Dimovic s.n. sub J 28,056
(1); Bedford Hill (-AA), Gilliland s.n. (1); Thorntree Kloof (-AA), Moss 6203
(1); Suikerbosrand Nature Reserve (-CA), Bredenkamp 982 (PUC). - 2725
2429 (PRE);
(Bloemhof): Wolmaranstad, farm Vaalbosch (-BB), Van k~
Wolmaranstad, farm Rooipoort (-BB).
LEDEBOURlA IN SOUTH AFRICA
273
TAXONOMY
ORANGE FREE STATE. - 2727 (Kroonstad): Kroonstad (-CA), Pont
624 (BLFU). - 2729 (Volksrust): Memel (-DA), Muller 1034 (PRE). - 2824
(Kimberley): Boshof, 14 miles north of Kimberley (-DB), Leistner 2875 (PRE).
- 2827 (Senekal): Between Paul Roux and Senekal (-BD), Zietsman 70 (PRE). 2828 (Bethlehem): Bethlehem (-AB), Van der Merwe 1143 (PRE). - 2829
(Harrismith): Sterkfontein Dam (-CA), Blom 103 (PRE).
LESOTHO. - 2927 (Maseru): Mamathes, Lenea (-BB), JacotGuillarmod 2744 (PRE); Roma (-BC), Schmitz 8024 (PRE); Schmitz 6745
(PRE).
NATAL. - 2632 (Bela Vista): Tembe Elephant Park (-CD), Ward 1702
(NH). -2730 (Vryheid): Vryheid (-BB), fun der Merwe 2465 (PRE); Vryheid
Hill (-DD), Venter 13,358 (UNIN). - 2732 (Ubombo): Ubombo (-CA), fun der
Merwe 2839 (PRE). - 2828 (Bethlehem): Leribe (-CC), Dieterlen 228 (NBG). 2829 (Harrismith): Ladysmith (-DB), Ditton s.n. sub NBG 390/39 (NBG). 2832 (Mtubatuba): Sibomvini, Eastern shores State Forest (-AB), fun "yk 904
(CPF). - 2929 (Underberg): Estcourt (-BB), fun der Merwe 2359 (PRE); Van
der Merwe 2443 (PRE). - 2930 (Pietermaritzburg): Albert Falls (-AD), fun der
Merwe 2601 (PRE); Camperdown (-DA), fun der Merwe 2419 (PRE). - 3030
(Port Shepstone): Umkomaas (-BB), Archbell s.n. (BOL).
TRANSKEI. - 3028 (Matatiele): Ongeluksnek (-AD), Strever 303 (KEI).
CAPE. - 2624 (Vryburg): Clober (-BB), Lamb s.n. sub NBG 51/26
(NBG). - 2722 (Olifantshoek): Olifantshoek (-DD), Thome s.n. (BOL). - 3026
(Aliwal North): Elandshoek (-DA), Bolus 158 (BOL); Bolus 213 (BOL);
Doctor's Drift (-DA), Gerstner 67 (PRE). - 3027 (Lady Grey): Lady Grey (CA), fun der Merwe 1855 (PRE); Herschel, Sterkspruit (-CB), Hepbum s.n.
sub GRA 7426 (GRA). - 3126 (Queenstown): Queenstown (-DD), Galpin 1659
(GRA). - 3225 (Somerset East): Cookhouse (-DB), Rogers s.n. (BOL). - 3326
(Grahamstown): Grahamstown (-BC), Zeyher 9 (BOL).
UDEBOURIA IN SOUTH AFRICA
274
TAXONOMY
Sectio Efiliferae S. \enter, sect. nov., foliis plus quam quatuor; ovario globoso
usque ad 1 mm longo.
Species typica: Ledebouria ovalifolia (Schrad.) Jessop.
Typus: Lodd. Bot. Cab. :278 (1818) sub Drimia lanceaejolia.
Species: L. concolor (C.A. Srn.) Jessop, L. undulata (Jacq.) Jessop, L. crispa
S. Venter, L. socialis (Bak.) Jessop, L. macowanii (Bak.) S. Venter, L.
ovalifolia (Schrad.) Jessop, L. petiolata (Van der Merwe) S. Venter.
Plants 50 - 120 mm tall. Bulbs 10 - 30 mm wide, neck prominent, torn
bulb scales without threads. Leaves erect, spreading or appressed to ground,
without threads when torn. Stamens erect. Ovary 1 - 1.5 mm long, globose.
Distribution and habitat.
Throughout South Africa, from semi - desert through coastal woodland, to
montane grassland.
The only section with a prominent neck to the bulb and when torn, lacking
threads in the leaves.
Subsectio Ebractae S. Venter, subsect. nov., bracteis carentibus brevissisve
usque ad 0.25 mm.
Species typica: L. concolor (Bak.) Jessop.
Species: L. concolor (Bak.) lessop.
25. LEDEBOURIA CONCOLOR (BaJc) Jessop
Ledebouria concolor (Bak.) Jessop in Jl S. Afr. Bot. 36(4): 254 (1970).
Drimia cooperi Bak. in Saund. Ref. Bot. 1: t.18 (1868).
Type: Cap. B. Spei, Cooper s. n. (K!, holo.; PRE!, photo.).
LEDEBOURlA IN SOlITH AFRICA
275
TAXONOMY
Scilla concolor Bak. in Saund. Ref. Bot. 3, Appen.: 13 (1870).
Type: As for Drimia cooped .
Plants gregarious. Bulbs mostly semi-epigea1, 60 - 120 x 30 - 50 mm, obovoid;
dead bulb scales purplish-brown, membranous with horizontal ridges, inner
bulb scales white, fleshy, apices truncate, without threads when torn; neck of
bulb 3 - 20 x 10 - 20 mm, basal stem present. Leaves fully developed at
anthesis, 4 - 6, spreading, lanceolate to oblong, 100 - 125 x 30 - 40 mm,
without threads when torn, fleshy, glabrous, concolorous, glossy green,
venation prominent; margins smooth; leaf base flat to shallowly canaliculate;
apex obtuse. Inflorescences 1 - 3, dense, oblong to cylindric, 80 - 120 x 30 - 40
mm, longer than the peduncle, erect to flaccid, 30 - 50 -flowered, longer than
the leaves; scape terete at base, green, glabrous; rachis smooth, 50 - 75 mm
long. Bracts absent or vestigeal. Pedicel spreading horizontally then drooping,
4 - 8 mm long, green. Perianth 3.7 - 6.2 mm long, tepals recurved, equal,
oblong, 4 - 7 x 2 - 3 mm, apex obtuse, slightly cucullate, green with a tinge of
pink at base, keel green. Stamens erect, 4 - 5 mm long, filaments green to pink,
epitepalous; anthers 1 mm long, yellow. Ovary globose, 6 -lobed, 1.0 - 1.5 x
2.5 mm, lobes narrowly transversely elliptic, base papillate, apex shoulders
raised. Style 4 mm long, triangular, glabrous, purple; stigma equal height to
anthers; stipe 0.5 x 0.5 mm. Capsule two- to three-lobed, asymmetrical,
cylindrical; base truncate. Seed disc-shaped, 3.5 - 4.0 mm long, surface
strongly wrinkled, brown. (Figure 64).
L. concolor is not closely allied to any other species of Ledebouria in South
Africa.
Specific epithet etymology.
Referring to the immaculate leaves.
Flowering period
From October to March.
LED£BOURlA IN SOUTH AFRICA
276
TAXONOMY
D
E
c
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Figure 64. Illustration of L. concolor (Bak.) Jessop. A, habit X 1; B, lamina
margin X 300; C, vestigial bract X 10; D, section through lamina X 2; E,
flower X 10; F, tepal with stamen X 10; G, apex of tepal X 20; H, ovary lateral
view X 10; I, ovary dorsal view X 10. All from Stirton 12,474.
LEDEBOURlA IN SOlTfH AFRICA
Known distribution of L. concolor (Bak.) Jessop
Map 29.
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G.P.-S. 13-0959
278
TAXONOMY
Figure 65. A, holotype of L. concolor (Bak.) Jessop (lower specimen) (K); B,
habitat at the mouth of the Zwartkops River near Port Elizabeth. The vegetation
consists of closed evergreen tall Sideroxylon inerme - Aloe ferox - Euphorbia
ledenii shrubland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata. C - E
from llin Jaarsveld 9060.
LEDEBOURLA IN sOurH AFRICA
TAXONOMY
279
UDEBOURLA IN SOlJTH AFRICA
280
TAXONOMY
Distribution (Map 29.).
Eastern Cape and two localities in the Ciskei. Most of the localities are in the
Port Elizabeth - Uitenhage area.
Habitat
Most of the populations occur on sandstone or shales of the Sondags River
Formation from the Uitenhage Group (SACS 1980). A few populations occur on
alluvial material from Cenozoic deposits just north- east of Port Elizabeth.
Soils found at the localities are either a brown, humusrich sandy loam or a grey,
humusrich sandy soil.
L. concolor is associated with Acocks veld type 23 - Valley Bushveld (Acocks
1988). The vegetation consists mostly of closed evergreen Dombeya cymosa Dalbergia obovata - Euphorbia triangularis woodland (Figure 65B). At Addo
the populations occur in the shade of closed evergreen tall Sideroxylon inerme Azima tetracantha - Scutia myrtina shrubland. Plants grow in light to dense
shade.
Population structure
Mostly small populations of between 20 - 50 individuals. The populations near
East London consist of 150 - 300 individuals. Most of the young plants are those
formed on the basal stems but seedling recruitment is also high. Most adult
plants flower every season with approximately 60% fruit formation.
Variation
Some populations tend to have plants with dry bulb scales and attenuate apices
showing the characteristic horizontal ridges, whereas populations in the Addo
area tend to have bulb scales with the apices truncate. The leaves are mostly
concolorous but a few individuals have been recorded as having dull dark green
blotches on the adaxial surfaces.
LEDEBOURLA IN SOIJfH AFRICA
281
TAXONOMY
Specimens examined
CAPE. - 3325 (Port Elizabeth): Glenconner (-AC), Long 1320 (PRE);
Port Elizabeth, farm Oudekraal (-BC), lizn Jaarsveld 9060 (NBG); Enon (-BC),
Thode A2779 (PRE); Gamtoos River Drift (-CC), Fourcade 3625 (BOL); Addo
(-DA), Barker 5089 (NBG); Komachs (-DC), Paterson s.n. (BOL); Redhouse (DC), Paterson 2678 (GRA), Paterson 4611 (BOL); Paterson 2603 (GRA); Port
Elizabeth (-DC), Drege 138 (BOL); Zwartkops River (-DC), Zeyher 4261
(BOL). - 3326 (Grahamstown): Bathurst, Kowie West (-DB), without collectors
name (BOL). - 3327 (Peddie): East London (-BB), Rattray s.n. (BOL).
l.EDEBOURIA IN SOlJfH AFRICA
282
TAXONOMY
Subsectio Longicollae S. l-enter, subsect. nov., bulbi collo longissimo, 7 - 15 x
5 - 10 mm;
Species typica: L. crispa S. Venter.
Species: L. crispa S. Venter, L. undulata (Jacq.) Jessop, L. ovalifolia (Schrad.)
Jessop, L. petiolata (Van der Merwe) S. Venter, L. socialis (Bak.) Jessop, L.
macowanii (Bale.) S. Venter.
26.LEDEBOUR/A eR/SPA S. Venter
Ledebouria crispa S. Venter, sp.nov., ab L. undulata (Jacq.) Jessop differt
habitu synantho bulbis semi-epigaeis gregariis; foliis 2 - 3 mm latis;
inflorescentia laxa; ovarii lobis basalis.
Type: Transvaal, Lebowa, Farm Majebes Kraal1002 LS, l-enter 11,202 (PRE!,
holo.; UN!; UNIN!).
Plants gregarious. Bulb semi-epigeal to epigeal, 15 - 30 x 10 - 20 mm, ovoid to
obovoid; dead bulb scales honey-brown, membranous, apices attenuate, live
bulb scales fleshy, tightly arranged, without threads when torn, white inside,
neck of bulb 5 - 10 x 4 - 5 mm, bulblets present, at bases of bulb-leaves.
Leaves fully developed at anthesis, 4 - 6, spreading to erect, linear-lanceolate,
20 - 70 x 2 - 3 mm, without threads when torn, fleshy, dull glaucous green,
without markings, venation obscure; margins undulate to crispate; leaf base
canaliculate; apex acute. Inflorescence solitary, lax, cylindric, 20 - 30 x 13 - 15
mm, erect, 8 - 12 -flowered, longer than the leaves; scape terete at base, green
to purplish-brown, glabrous; rachis smooth, 40 - 50 mm long. Bracts
membranous, 1.2 - 1.5 x 0.5 mm, linear-lanceolate, grey to white with
bracteoles. Pedicels spreading, 2 - 4 mm long, pink speckled white. Perianth 3
- 3.5 mm long, tepals sharply reflexed, equal, oblong, 5 x 1.5 mm, apex acute,
slightly cucullate, pink to purple with a dull green keel. Stamens erect, 3.0 3.5 mm long, filaments pink, epitepalous; anthers 0.5 mm long, mauve. Ovary
depressed ovoid, 6 -lobed, 2.5 x 3 mm, lobes obtusely deltate, apex tapering
into the style, basal lobes present. Style 3.5 mm long, terete, glabrous, white;
stigma equal height to the anthers; stipe 1 x 1 mm. Capsule three-lobed,
symmetrical, globose; base truncate. Seed drop-shaped, 3.0 x 1.5 mm, surface
strongly wrinkled, black. (Figure 66).
LEDEBOURIA IN SOlITH AFRICA
283
TAXONOMY
E
D
~
F
C
G
A
H
8
Figure 66. Illustration of L. crispa S. Venter. A, habit X 1; B, lamina margin X
300; C, section through lamina X 5. D, bract with bracteole X 10; E, tepa! with
stamen X 10; F, flower X 10; G, apex of tepal X 10; H, ovary lateral view X
10; I, ovary dorsal view X 10. All from
retn~
11,202.
LEDEBOURlA IN SOUTH AFRICA
Map 30.
Known distribution of L. crispa S. Venter
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TAXONOMY
Figure 67. A, holotype of L. crispa S. Venter (PRE); B, colony of plants
showing the typical colonial growth. Bar
=
20 mm; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of a stoma. All from l-enter 11,202.
UDEBOURIA IN SOUTH AFRICA
286
TAXONOMY
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LEDEBOURIA IN SOlITH AFRICA
287
TAXONOMY
L. crispa is closely related to L. undulata (Jacq.) Jessop but differs in having
the bulb 15 - 25 mm long whereas with L. undulata the bulb can be 30 - 50 mm
long. In L. crispa the leaf margin is always crisped and 2 - 3 mm wide whereas
with L. undulata the leaf margin is only undulate and 8 - 20 mm wide and rather
firmer in texture.
Specific epithet etymology.
Describes the lamina margin.
Flowering period
From October to December.
Distribution (Map 30).
Known only from around Pietersburg in the north-eastern Transvaal.
Habitat
L. crispa is restricted to the ultramaphic rocks (quartzitic schist and chlorite-talc
schist) of the Mothiba Formation of the Pietersburg Group (SACS 1980). These
rocks result in a series of low hills between Potgietersrust and Houtboschdorp.
The soil at both cited localities is brown sandy loam lithosols of schist overlaid
by white quartz. Some of the plants grow in rock cracks with very little soil but
prefer to grow in the loose quartz rubble. At both localities L. crispa grows in
full sun.
Plants grow in scattered groups of 5 - 20 plants (Figure 67B) or occasionally as
individuals. Ten seeds from different plants were sown and kept under
controlled conditions (Aridarium at the University of the North). Three seeds
germinated after three weeks, six seeds after five weeks and one seed after eight
weeks. All the seeds were ± the same age. This erratic germination was found
to be common with all Ledebouria species.
LEDEBOURlA IN SmITH AFRICA
288
TAXONOMY
Specimens examined
TRANSVAAL. - 2329 (Pietersburg): Pietersburg Game Reserve (-CD),
Venter 13,209a (PRE, UNIN); Lebowa, farm Majebeskraal 1002 LS (-DC),
Venter 11,202 (pRE, UN, UNIN).
LEDEBOURIA IN SOUTH AFRICA
289
TAXONOMY
27. LEDEBOURIA UNDULATA (Jacq.) Jessop
Ledebouria undulata (Jacq.) Jessop in Jl S. Afr. Bot. 36(4): 258 (1970).
Drimia undulata Jacq. in Icones Plantarum Rariorum 2(15): t.376 (1794).
Iconotype: Icones Plantarum Rariorum 2(15): t.376 (1794).
Scilla undulata (Jacq.) Bak. in Saund. Ref. Bot. 3(Append.): 11 (1870). Nom.
illegit., non S. undulata Desf., 1798.
Scilla undulatifolia Von Poelln. in Ber. dt. Ges. 41: 209 (1944).
Type: As for Drimia undulata Jacq.
Plant solitary. Bulb hypogeal, 50 - 60 x 30 - 50 mm, ovoid; dead bulb scales
light brown, membranous, apices truncate, live bulb scales fleshy, loosely
arranged, without threads when torn, white inside, neck 10 - 20 x 10 - 20 mm.
Leaves hysteranthous, 3 - 9, spreading, linear-lanceolate to lanceolate, 70 - 110
x 6 - 10 mm, without threads when torn, fleshy, glabrous, dull green to
glaucous green, venation sometimes prominent; margins undulate; leaf base
canaliculate; apex acute. Inflorescence solitary, dense, cylindric, 25 - 40 x 15 25 mm, erect, 20 - 30 -flowered, shorter than the leaves; scape terete at base,
green, glabrous; rachis ridged, 15 - 20 mm long. Bracts fleshy, 1 x 0.5 mm,
linear to dentate, green to pinkish with bracteoles present. Pedicels spreading
horizontally, 3 - 7 mm long, pink. Perianth 5 - 6 mm long, tepals recurved,
equal, oblong, 5 - 6 x 1.0 - 1.5 mm, apex acute, thickly cucullate, pink with a
brown keel. Stamens spreading, 4 mm long, filaments white, epitepa1ous;
anthers 0.5 mm long, yellow. Ovary globose, 6 -lobed, 1.5 x 2 mm, lobes
narrowly transversely oblong, apex tapering into style. Style 5 mm long,
triangular, glabrous, white; stigma above anthers; stipe 0.25 x 0.25 mm.
Capsule three-lobed, symmetrical, globose, base truncate. Seed drop-shaped, 3
mm long, surface strongly wrinkled, brown. (Figure 68).
LEDEBOURlA IN SOtrfH AFRICA
290
TAXONOMY
E
~
B
~
J
A
Figure 68. Illustration of L. undulata (Jacq.) Jessop. A, habit, leafing period X
1; B; habit, flowering period Xl; C, lamina margin X 300; D, bracts with
bracteoles X 10; E, section through lamina X 4; F, flower X 10; G, apex of
tepal X 20; H, tepal with stamen X 10; I, ovary lateral view X 10; J, ovary
dorsal view X 10. A and C - J from Muller-Doblies 89129 and B from Hall
3200.
LEDEBOURlA IN SOUTH AFRICA
Map 31.
Known distribution of L. undulata (Jacq.) Jessop
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G.P.-S. 13-0959
292
TAXONOMY
Figure 69. A, type of L. undulata (Jacq.) Jessop under Drimia undulata Jacq. in
!cones Plantarum Rariorum 2: t.376 (1794); B, habitat on the Gamsberg Flats,
Bushmenland. The vegetation consists of closed Euphorbia mauritanica - Euclea
undulata shrubland; C, plant of L. undulata with semi-fleshy leaves and the
inflorescence absent. Bar = 10 mm; D, SEM micrograph of the adaxial lamina
surface; E, SEM micrograph of the abaxial lamina surface; F, SEM micrograph
of stomata. C - F from Miiller-Doblies 89129.
UDEBOURlA IN SOtJrH AFRICA
TAXONOMY
293
.(./ )/1/1/1<1
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LEDEBOURJA IN SOlJfH AFRICA
294
TAXONOMY
L. undulata is the only hysteranthous species of Ledebouria. In leaf it is easy to
identify by the fleshy texture and dull green to glaucous colour.
L. undulata is closely related to L. crispa and belongs to the subsection
Longicollae, section Efiliferae. It differs from L. crispa in having the bulb 50 60 mm long whereas with L. crispa the bulb is 15 - 30 mm long. In L. undulata
the leaf margin is undulate with the lamina being 6 - 10 mm wide whereas with.
L. crispa the leaf margin is crispate and 2 - 3 mm wide and rather thinner in
texture.
The absence of leaves at anthesis in L. undulata is the major difference between
L. undulata and L. crispa.
Specific epithet etymology.
Describes the undulate lamina margin.
Flowering period
L. undulata flowers sporadically depending on rainfall. Most flowering
herbarium specimens were collected between June and September.
Distribution (Map 31).
A Cape endemic.
Habitat
L. undulata occurs on well drained, medium grained (0.25 - 1.0 mm c/» to
coarse grained (l - 3 mm c/», shallow (l0 - 50 mm deep) sandy soil derived
from gneiss or granite. The vegetation is typical Namaqualand Broken Veld
(Acocks 1988) (Figure 69B).
LEDEBOURIA IN SOlIrH AFRICA
295
TAXONOMY
Around Clanwilliam populations occur on Ceres Formation sandstone of the
Bokkeveld Group (SACS 1980). Soil derived from these rocks is well drained,
medium grained (0.25 - 1.0 mm 4» and deep (120 - 320 mm). The vegetation is
primarily succulent veld with a few woody shrubs in protected areas.
From Langebaan to the Olifants River on the west coast, L. undulata occurs on
Kalahari Group sand on limestone. This consists of greyish-brown loose sandy
material, varying in depth from 10 - 320 mm (Van der Merwe 1962). The
vegetation consists mostly of woody shrubs with Restionaceae and smaller
Fynbos elements in the open areas.
Variation
Plants growing in moist habitats tend to have thinner live bulb scales, leaves
more fleshy and greener with a glossy surface lacking venation. Plants in xeric
habitats have longer, semi-fleshy, dull glaucous green leaves with prominent
venation and undulate margins.
Specimens examined
CAPE. - 2917 (Springbok): 2 km SE of Steinkopf (-BC), Muller-Doblies
89129 (UNIN). - 3017 (Hondeklipbaai): Karkams (-BD), Snijman 280 (NBG). 3018 (Kamiesberg): Garies (-CA), Leighton 2393 (BOL); Compton 18,851
(NBG). - 3019 (Loeriesfontein): Near Kliprand (-CA), Perry 1966 (NBG); 3022 (Carnarvon): Carnarvon (-CC), Steyner s.n. (NBG). - 3119 (Calvinia):
Clanwilliam, Botterkloof Pass (-CD), Hall 3221 (NBG). - 3218 (Clanwilliam):
Clanwilliam (-BB), Leipoldt s.n. (BOL); Van der Merwe 19 (PRE). - 3219
(Wuppertal): Pakhuis Pass (-AA), Compton 4,826 (BOL). - 3318 (Cape Town):
Langebaan, Donkergat (-AA), Hall 3,200 (PRE); Hopefield (-AB), Bolus s.n.
(BOL); Hall 3131 (NBG); Malmesbury (-DC), Gillett 3639 (BOL).
Without precise locality
Little Namaqualand, Zeyher 11 (BOL).
UDEBOURlA IN SOUTH AFRICA
296
TAXONOMY
28. LEDEBOURIA OVALIFOLIA (Schrad.) Jessop
Ledebouria ovalifolia (Schrad.) Jessop in 11 S. Afr. Bot. 36(4): 246 (1970).
Drimia ovalifolia Schrad. in Blumenb. :28 (1827).
Iconotype: Lodd. Bot. Cab. 3: t.278, sub Drimia lanceaejolia, (1818).
Lachenalia reflexa Andr. in Bot. Rep. 5: t.299 (1803), nom. illegit., non
Thunb.
Drimia lanceolata Schrad. in Blumenb. :28 (1827).
Iconotype: Andr. Bot. Rep. 5: t.299, sub Lachenalia rejlexa Andr.(1803).
Drimia gawleri Schrad. in Blumenb. :30 (1827).
Iconotype: In (1811) Curtis's Bot. Mag. 33: t1380, sub Drimia lanceaejolia (13),
(1811).
Scilla lanceolata (Schrad.) Bak. in Saund. Ref. Bot. 3(Append.): 14 (1870).
nom. illegit., non Viviani (1830).
Scilla revoluta (L.f) Bak. sensu Bak. in Flora Cap. 6: 485 (1896).
Scilla ovalifolia (Schrad.) C.A. Srn. in Kew Bull. :245 (1930).
Iconotype: Andrews Bot. Rep. 5: t.229 (1803).
Scilla doratophylla C.A. Srn. in Kew Bull. :245 (1930).
Type: As for Drimia lanceolata Schrad.
Scilla genadendalensis Von Poelln. in Port. Acta bioI., ser.B, 1: 212 (1945).
Type: Cape, Genadendal, Schlechter 10,327 (PRE!, lecto.; BOLl; Z!). The
PRE specimen designated here as lectotype (Greuter et aI1988).
UDEBOURlA IN SOlITH AFRICA
297
TAXONOMY
Plants solitary. Bulbs hypogeal, 10 - 20
X
10 - 15 mm, ovoid; dead bulb scales
brown, membranous, apices attenuate, live bulb scales tightly arranged, without
threads when torn, white inside, neck ± 5 mm long and wide. Leaves partly
emerged at anthesis, 2 - 4, appressed to ground, sometimes spreading,
oblanceolate, 30 - 50 x 20 - 30 mm, without threads when torn, fleshy, smooth
or with short rows of papillae; dull green, abaxial surface purple, sometimes
purple cross bars at base of lamina, venation obscure; margins finely papillate;
leaf base canaliculate; apex obtuse. Inflorescence solitary, lax, elliptic, 15 - 30
x 15 - 20 mm, flaccid to erect, 4 - 12 -flowered, longer than leaves. Peduncle
glabrous, terete at base, green, glabrous; rachis smooth, 30 - 50 mm long.
Bracts semi-fleshy, 1 x 0.5 mm, deltoid, pink to purple without bracteoles.
Pedicels spreading horizontally, 4 - 7 mm long, pink. Perianth 3.5 - 4.0 mm
long, tepals recurved, equal, oblong, 3.5 - 4.0 x 1 mm, apex obtuse, pink to
purple, keel dull green. Stamens erect, 2.5 - 3.0 mm long, base slightly
flattened, swollen, maroon, epitepalous; anthers 1 mm long, violet. Ovary
ovoid, 3 -lobed, 1.5 x 2 mm, lobes depressed ovate, apex shoulders raised.
Style 2 mm long, terete, glabrous, purple; stigma above anthers; stipe 0.25 x
0.25 mm. Capsule three-lobed, symmetrical, globose; base truncate. Seed dropshaped, 3 mm long, surface strongly wrinkled, brown. (Figure 70).
L. ovalifolia resembles L. petiolata (Van der Merwe) S. Venter but differs in
the leaves partly emerged at anthesis, petiole not as well formed, solitary
inflorescence, rachis smooth, no bracteoles and obtuse tepal apices. Together
with L. socialis (Bak.) Jessop and L. macowanii (Bak.) S. Venter they constitute
the subsection Longicollae.
Specific epithet etymology.
Describes the oval leaf shape.
Flowering period
From January to August with a peak from January to April.
LEDEBOURIA IN SOlJfH AFRICA
298
TAXONOMY
E
o
H
~
..,
. . . . .,'.
.
'."
;.~
~
J
A
B
Figure 70. Illustration of L. ovalifolia (Schrad.) Jessop. A, habit X 1; B, lamina
margin X 300; C, apex of lamina X 20; D, apex of tepal X 20; E, section
through lamina X 5; F, bract X 10; G, flower X 10; H, tepa! with stamen X 10;
I, ovary lateral view X 10; J, ovary dorsal view X 10. All from "Cnter 13,263.
.u:DEBOURJA IN SOUTH AFRICA
Map 32.
12°
Known distribution of L. ovalijo/ia (Schrad.) Jessop
14°
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TAXONOMY
Figure 71. A, type of L. ovalifolia (Schrad.) Jessop under Drimia lanceaefolia
Lodd. in Loddiges Botanical Cabinet 3: t.278 (1818); B, plant of L. ovalifolia
with flatly spreading leaves. Bar
=
20 mm; C, SEM micrograph of the adaxial
lamina surface; D, SEM micrograph of the abaxial lamina surface. Bar
I-'m; E, SEM micrograph of stomata. Bar
=
=
50
20 I-'m; F, SEM micrograph of the
parallel rows of papillae on the abaxial lamina surface; G, habitat at Baakens
River, Port Elizabeth. The vegetation consists of Protea - Watsonia Leucospermum Fynbos. B - F from Venter 13,263.
LEDEBOURlA IN 50tJfH AFRICA
301
TAXONOMY
u:DEBOURlA IN SOurH AFRICA
302
TAXONOMY
Distribution (Map 32).
Endemic to the Cape. Most localities are in the southern Cape with an isolated
locality at Grahamstown.
Habitat
L. ovalifolia grows in open areas between shrubs and grass in Fynbos and
shrubland in sandstone areas (Figure 71G). Soils are medium grained (0.25 1.0 mm c/», well drained acidic sandy soils. These soils are from 30 - 750 mm
deep with the surface horizon containing grit, gravel and small pebbles
consisting of white quartz fragments and fine iron oxide concretions (Van der
Merwe 1962).
Variation
A few populations have plants with leaves larger and not appressed but slightly
spreading, especially those in light shade. The purple or dark green stripes at
the base of the lamina may be indistinct but are always present. The small
papillae on the adaxial lamina surface are absent in some populations.
Specimens examined
CAPE. - 3319 (Worcester): Worcester, De Doorns (-DA), Walters 1837
(NBG). - 3320 (Montagu): Bredasdorp, De Hoop (-AD), Barker 8744 (NBG);
The Baths (-CC), Page s.n. sub BOL 17,407 (BOL); Swellendam, between
Lemoenshoek and Naauwkrantz (-DD), Stokoe s.n. sub NBG 99,678 (NBG). 3322 (Oudtshoorn): George, Lancewood (-DD), de Villiers s.n. sub STE 17,034
(STE). - 3326 (Grahamstown): Mayor's Seat (-BC), Daly 846 (GRA). - 3419
(Caledon): Caledon Hill (-AB), Purcell 80 - 88 (NBG); Bot River Valley (-AC),
Guthrie 2321 (BOL); Between Onrust and Hawston (-AC), Thomas s.n. sub
NBG 92,553 (NBG); Riviersonderend Mountains (-BA), Lewis 2980 (NBG);
LEDEBOURlA IN SOlITH AFRICA
303
TAXONOMY
Genadendal (-BA), Bolus & Gillett 880 (BOL); Schlechter 10,327 (BOL); Near
Lindeshof (-BB), Esterhuysen s.n. (BOL). - 3420 (Bredasdorp): Swellendam, 10
0' Clock Mountain (-AB), Wurts 560 (NBG); De Hoop, Potberg Nature
Reserve (-AD), Burgers 1723 (STE); Swellendam, Potteberg (-BC), Lewis 3224
(BOL). - 3422 (Mossel Bay): George, Great Brak River (-AA), Fourcade 3878
(BOL). - 3423 (Knysna): Plettenberg Bay, Formosa (-AB), Fourcade 597
(BOL); Fourcade 1052 (BOL).
LEDEBOURJA IN SOUTH AFRICA
304
TAXONOMY
29. LEDEBOURIA PETIOLATA (Van der Merwe) S. Venter
Ledebouria petiolata (WIn der Merwe) S. lenter comb.nov.
Scilla petio[ata Van der Merwe in Flower. PI. S. Afr. 21: 1. 832 (1941).
Type: Transvaal, North of Graskop, Strydom s. n. sub PRE 26,397 (PRE!,
holo.; iso. !).
Plants solitary. Bulb hypogeal, 10 - 30 x 10 - 15 mm, obovoid; dead bulb scales
brown, membranous, apices attenuate, live bulb scales tightly arranged, with
sparse threads when torn, white inside, neck 20 x 5 - 8 mm. Leaves fully
developed at anthesis, 5 - 8, spreading, broadly lanceolate, 40 - 60 x 5 - 8 mm,
without threads when torn, fleshy, dull green tinged with purple, or abaxial
surface purple, immaculate, venation obscure; margins finely finely papillate
and thickned; petiole up to 30 x 1 mm; apex acute. Inflorescence 1 - 2, lax,
elliptic, 15 x 30 mm, flaccid, 15 - 20 -flowered, longer than the leaves; scape
terete at base, green, glabrous; rachis ridged, 70 - 100 mm long. Bracts fleshy,
0.5 x 0.5 mm, dentate, pink to purple with bracteoles. Pedicels spreading, 8 - 9
mm long, purple. Perianth 4 - 7 mm long, tepals recurved, equal, oblong, 5 x
1.5 - 2.0 mm, apex acute, thinly cucullate, bright pink with sometimes a green
keel. Stamens erect, 4.5 mm long, filaments maroon, epitepalous; anthers 0.5
mm long, yellow. Ovary ovoid, 6 -lobed, 1 x 2 mm, lobes obtusely deltate,
apex tapering into style. Style 2.5 mm long, triangular, glabrous, purple above
and white below; stigma equal height to anthers; stipe 0.5 x 0.5 mm. Capsule
three-lobed, globose; base truncate. Seed drop-shaped, 2.5 mm long, strongly
wrinkled, yellowish-brown. (Figure 72).
Cannot be confused with any other Ledebouria species. The long, thin petiole is
umque.
Specific epithet etymology.
Describes the prominent petiole.
I.£DEBOURJA IN SOlITH AFRICA
305
TAXONOMY
E
G
o
F
c
~
~
H
A
Figure 72. Illustration of L. petiolata (Van der Merwe) S. Venter. A, habit X 1;
B, lamina margin X 300; C, section through lamina X 5; D, apex of tepal X 20;
E, bract with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H,
ovary lateral view X 10; I, ovary dorsal view X 10. All from Middleton s.n..
LEDEBOURJA IN SOlITH AFRICA
Known distribution of L. petio!ata (Van der Merwe) S. Venter
Map 33.
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TAXONOMY
Figure 73. A, holotype of L. petiolata (Van der Merwe) S. Venter (PRE); B,
habitat near Graskop. The vegetation consists of open short Passerina montana -
Ischyrolepis schoenoides - Panicum natalense grassland. C, SEM micrograph of
the adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface;
E, SEM micrograph of stomata; F, plants of L. petiolata typically growing in
moss filled rock cracks. Bar
= 20 mm. C - F from lenter s.n.
LEDEBOURIA IN SOllfH AFRICA
308
TAXONOMY
A
LEDEBOURIA IN SOUTH AFRICA
309
TAXONOMY
Flowering period
From September to January with most flowering specimens collected during
January.
Distribution (Map 33).
Known only from the Graskop and Kaapsche Hoop areas on the Transvaal
Drakensberg Escarp.
Habitat
L. petiolata occurs on Black Reef Quartzite of the Wolkberg Group (SACS
1980). Soil derived from these rocks is shallow (10 - 50 mm deep), medium
grained (0.25 - 1.0 mm cP) grey sandy soil. Plants grow mostly in rock cracks
and shallow soil associated with the moss Fissidens submarginatus Bruch ex
Krauss (Figure 73F). The vegetation is montane grassland with scattared Aloe
arborescence Mill. and Rhus tumulicola S. Moore var. tumulicola shrubs
(Figure 73B).
Variation
The main variation found in L. petiolata is in the colour of the abaxial lamina
surface. This can either be green or purple depending on the exposure to
sunlight.
Specimens examined
TRANSVAAL. - 2430 (Pilgrim's Rest): Bourke's Luck, De Berg (-DB),
Wm der Merwe 1591 (PRE); Belvedere (-DD), Davidson 483 (J); Pirow's Grave
(-DD), Venter s.n. (UNIN); Graskop (-DD), Strydom s.n. sub PRE 26,397
(PRE).
UDEBOURlA IN SOlJTH AFRICA
TAXONOMY
310
30. LEDEBOURIA SOCIALIS (Bak.) Jessop
Ledebouria socialis (Bak.) Jessop in 11 S. Afr. Bot. 36(4): 253 (1970).
Scilla socialis Bak. in Saund. Ref. Bot. 3: t.180 (1870).
Type: Natal, without precise locality, Cooper 3635 (Kl, holo.; PRE1, photo.).
Scilla paucifolia Bak. in Saund. Ref. Bot. 3: t.181 (1870).
Iconotype: Saund. Ref. Bot. 3: t.181, "Cape, Cooper s.n." 1870.
Scilla laxijlora Bak. in Gdnr's Chron. 9: 668 (1891).
Type: Hortus N.E. Brown, Kew. (Kl, holo.; BOLl, drawing).
Scilla violacea Hutch. in Kew Bull. : 511 (1932).
Type: Cape, Uitenhage Div., Hankey, Hutchinson & Long s. n. (K!, holo.;
BOL!; PRE!).
Ledebouria violacea (Hutch.) W.L. Tjaden in British Cactus and Succ. Journ.
7(1): 11 (1989).
Type: As for Scilla violacea Hutch.
Plants gregarious. Bulb epigeal, 40 - 50 x 30 - 36 mm, ovoid to obovoid; dead
bulb scales brown to purple, membranous, apices attenuate, live bulb scales
fleshy, tightly arranged, without threads when torn, white inside, neck of bulb
10 - 15 x 5 - 10 mm, bulblets on basal stem. Leaves fully emerged at anthesis, 3
- 4, spreading, oblanceolate, 50 - 75 x 20 - 25 mm, without threads when torn,
fleshy, glossy, adaxial surface green to green with a silvery sheen, immaculate
or with darker green to purple blotches, abaxial surface green to purple,
venation obscure; margins smooth; leaf base canaliculate; apex acute.
Inflorescence solitary, lax, cylindric, 30 - 50 x 20 - 30 mm, erect, 10 - 30
-flowered, longer than leaves; scape terete at base, brownish to green; rachis
smooth, 50 - 70 mm long. Bracts fleshy, 0.5 x 1 mm, deltoid, green without
bracteoles. Pedicels spreading, 3 - 4 mm long, pink to green. Perianth 3.5 _
LEDEBOURlA IN SOlJfH AFRICA
311
TAXONOMY
4.0 mm long, tepals recurved, equal, oblong, 3.5 - 4.0 x 1.5 - 2.0 mm, apex
obtuse, lobes whitish green to pink with a green keel. Stamens erect, 3.5 - 4.0
mm long, filaments with upper part purple, white below, epitepalous; anther 0.5
mm long, yellow. Ovary ellipsoidal, 6 -lobed, 2 x 2.5 mm, lobes obtusely
deltate, apex shoulders raised. Style 3 mm long, triangular, glabrous, purple
above and white below; stigma above anthers; stipe 0.25 x 0.25 mm. Capsule
three-lobed, globose; base truncate. Seed drop-shaped, 3 mm long, surface
strongly wrinkled, black. (Figure 74).
L. socialis is closely related to L. macowanii (Bak.) S. Venter but differs in
having epigeal bulbs. Both these species belong to the subsection Longicollae.
L. socialis is the only species with epigeal bulbs, an erect inflorescence and
small flowers with strongly reflexed lobes.
Specific epithet etymology.
Describes the gregarial habit.
Flowering period
From July to December peaking from October to November.
Distribution (Map 34).
Endemic to the Cape Province. Most of the known localities are in the eastern
Cape.
LEDEBOURlA IN SOUTH AFRICA
312
TAXONOMY
~
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F
G
B
Figure 74. Illustration of L. socialis (Bak.) Jessop. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, bract X 10; E, flower X 10;
F, tepal with stamen X 10; G, ovary lateral view X 10; H, ovary dorsal view X
10. All from "\.enter 13,272.
LEDEBOURlA IN sourH AFRICA
Known distribution of L. socialis (Bak.) Jessop
Map 34.
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314
TAXONOMY
Figure 75. A, holotype of L. socialis (Bak.) Jessop (K); B, plant showing the
gregarious habit. Plant growing in deep shade. Bar
showing the pendulous flowers. Bar
=
=
40 mm; C, plants
20 mm; D, SEM micrograph of the
adaxial lamina surface; E, SEM micrograph of the abaxial lamina surface; F,
SEM micrograph of stomata. B - F from
retn~
13,272.
LEDEBOURIA IN SOlITH AFRICA
TAXONOMY
315
(
UDEBOURlA IN SOlITH AFRICA
316
TAXONOMY
Habitat
L. socialis occurs on sandstone in fine to medium grained (0.025 - 1.0 mm cP),
shallow (10 - 24 mm) to deep (50 - 320 mm) well drained humusrich sandy soil.
It usually grows in shade mostly in closed evergreen woodland. A few
populations occur in evergreen scrub forest. Sometimes a thick layer of litter
covers the plants. Most of the localities are in Acocks veld type 23 - Valley
Bushveld. The distribution of L. socialis correlates with river valleys.
Variation
Plants of L. socialis growing in full sun tend to be smaller with ovoid bulbs,
short scapes and pink flowers with dull green tepal keels. The colour of the
bulb scales varies from green to a deep purple.
The leaf colour variation has caused confusion since 1870 in horticultural
circles. Plants in cultivation originated from collections made of green-leaved,
mottle-Ieaved and silvery-leaved forms. The latter being the most popular. This
resulted in Tjaden (1989) making the new combination Ledebouria violacea
(Hutch.) W.L. Tjaden based on leaf colour only.
Historical background
Baker (1870a) in his description of Scilla socialis cites the locality as "Cape of
Good Hope, discovered by Mr. Cooper", but on the type specimen the locality
is given as Natal. It was described from a plant cultivated by Mr. Wilson
Saunders at Reigate about 1870 (Baker 1896). The species has never been recollected in Natal and Cooper's locality in Natal remains a mystery.
LEDEBOURIA IN SOIITH AFRICA
317
TAXONOMY
Specimens examined
CAPE. - 3123 (Victoria West): Victoria West (-CA), Bolus s.n. (NBG).
- 3323 (Willowmore): Albany, Paardekraal (-CC), Dyer 2156 (GRA). - 3324
(Steytlerville): Hankey (-DD), Long s.n. (BOL). - 3325 (port Elizabeth): Addo
(-DA), Barker 4969 (NBG). - 3326 (Grahamstown): Fraser's Camp (-BD),
Barker 6986 (NBG); Blaauwkrantz (-BD), HUner lOO (GRA); Kariega River (DA), Wm ")'k & Kok 5840 (PRE); Kowie (-DB), Tyson 19,250 (PRE); Port
Alfred (-DB), Schonland 1554 (GRA). - 3327 (Peddie): Peddie (-AA), Herre
s.n. (BOL); Line drift (-AA), Leighton s.n. (BOL); East London (-BB), Rattray
234 (GRA); Rattray s.n. (BOL).
LEDEBOURIA IN SOUTH AFRICA
318
TAXONOMY
31. LEDEBOURIA MACOWANII (Bale.) S. Venter
Ledebouria macowanii (Bak.) S. \enter comb.nov.
Scilla macowanii Bale. in Gdnr's Chron. 3: 748 (1875).
Type: Cape, Somerset Division, Boschberg, MacOwan 1841 (GRA!, lecto.;
BOL!; PRE1, photo.; Z). Here designated as lectotype (Greuter et alI988).
Scilla nelsonii Bale. in Flora Cap. 6: 488 (1896).
Type: Cape, Vaal River, Nelson 167 (K!, holo.; PRE1).
Plants solitary. Bulb hypogeal, 15 - 30 x 15 - 30 mm, ovoid; dead bulb scales
brown, membranous, apices attenuate, live bulb scales tightly arranged, without
threads when torn, neck of bulb 5 x 10 mm, bulb white inside. Leaves fully
developed at anthesis, 3 - 4, spreading, linear-lanceolate, 75 - 100 x 6 - 8 mm,
without threads when torn, fleshy, dull green, abaxial surface green with
occasional purple spots and blotches in lower third to half, venation obscure;
margins smooth; leaf base canaliculate; apex obtuse to acute. Inflorescences 1 2, lax, cylindric, 25 - 60 x 17 - 25 mm, flaccid, 20 - 40 -flowered, as long to
longer than the leaves; scape base terete, green spotted purple; rachis smooth,
40 - 100 mm long. Bracts fleshy, 0.5 x 0.5 mm, deltoid, green to pink without
bracteoles. Pedicels spreading, 6 - 7 mm long, pink to purple. Perianth 4 mm
long, stellate, tepals slightly recurved, equal, oblong, 4 x 1.5 mm, apex obtuse,
green tinged pink. Stamens spreading, 3.8 mm long, filaments with upper part
purple and white below, epitepalous; anthers 0.5 mm long, pale violet. Ovary
globose, 6 -lobed, 1.5 x 2 mm, lobes obtusely deltate, apex shoulders raised.
Style 1.8 - 2.0 mm long, triangular, glabrous, purple; stigma exceeding anthers;
stipe 0.5 x 0.5 mm. Capsule three-lobed, symmetrical, clavate; base tapering.
Seed drop-shaped, 3 - 4 mm long, surface strongly wrinkled, brown. (Figure
76).
L. macowanii is closely related to L. socialis with the bulbs hypogeal and not
epigeal.
LEDEBOURlA IN SOlITH AFRICA
319
TAXONOMY
o
E
B
Figure 76. Illustration of L. macowannii (Bak.) S. Venter. A, habit Xl; B,
lamina margin X 300; C, section through lamina X 2.5; D, bract X 10; E, apex
of tepal X 10; F, flower X 5; G, tepal with stamen X 10; H, ovary lateral view
X 10; I, ovary dosal view X 10. All from
retn~
13,413.
LEDEBOURJA IN SOUTH AFRICA
Known distribution of L. macowannii (Bak.) S. Venter
Map 35.
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TAXONOMY
Figure 77. A, lectotype of L. macowanii (Bak.) S. Venter (GRA); B, habitat
near Pafuri in the Kruger National Park. The vegetation consists of closed
deciduous Terminalia sericea - Hymenocardia ulmoides - Holarrhena pubescens
woodland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata; F,
plants of L. macowanii showing the lax racemes. Bar = 50 mm. C - F from
Venter s.n..
LEDEBOURJA IN SmITH AFRICA
TAXONOMY
322
UDEBOURlA IN SOUTH AFRICA
323
TAXONOMY
Specific epithet etymology.
Commemorates Dr. P. MacOwan who collected the type material.
Flowering period
From October to March with a peak from November to December.
Distribution ( Map 35).
Widely scattered throughout the eastern half of South Africa mostly in the
Transvaal and eastern Cape. This species is mostly associated with mountains.
Habitat
L. macowanii occurs in shallow, 10 - 50 mm deep, humusrich sandy soil
derived from quartzite, conglomorate or sandstone. In the western Transvaal,
southern Cape and on the Natal Coast this species grows on damp cliff ledges in
semi shade.
Variation.
Plants from dry lowland areas of Kimberley and the Kruger National Park tend
to be small. The neck of the bulbs are usually above ground level except in
some populations in the Waterberg (Venter 13,413). Plants from the Cape are
prominently marked with purple, whereas the plants from the northern
populations are immaculate.
LEDEBOURJA IN SOlITH AFRICA
324
TAXONOMY
Historical background
Jessop (1970) regarded Scilla macowanii as a synonym of Ledebouria cooperi
without giving a reason. He also placed Scilla nelsonii as a synonym of
Ledebouria undulata with a note that it may be nearer to L. cooperi than to L.
undulata.
Specimens examined
TRANSVAAL. - 2231 (Pafuri): Punda Maria, Dzundwini Hill (-CC),
Codd & Dyer 4598 (KNP, PRE). - 2427 (Thabazimbi): Rooiberg (-DD), WIn
der Merwe 2044 (PRE). - 2329 (Pietersburg): Louis Trichardt (-BB), Koker 3
(PRE). - 2331 (Phalaborwa): Letaba Camp (-DC), Codd 4675 (KNP). - 2431
(Acornhoek): Skukuza (-DC), Codd & De Winter 5076 (PRE); Mabase (-DC),
Lang s.n. (PRE). - 2527 (Rustenburg): Rustenburg (-CA), WIn der Merwe 1558
(PRE).
SWAZILAND. - 2631 (Mbabane): Komati Bridge (-AA), Compton
29,391 (PRE); Usuthu Dam (-CA), Compton 27,120 (PRE). - 2632 (Bela Vista):
Mbuluzi Nature Reserve (-AA), Culverwe1l1389 (PRE).
ORANGE FREE STATE. - 2924 (Fauresmith): Fauresmith, farm
Wanda (-CB), Smook 3045 (PRE). - 2827 (Seneka1): Seneka1, farm Doornkop
(DD), Goossens 777 (PRE). - 2828 (Bethlehem): C1arence (-CB), WIn Hoepen
18,270 (PRE). - 2927 (Maseru): C10co1an, farm Hillcrest (-BA), Crosby 420
(PRE).
NATAL. - 2732 (Ubombo): Phelendaba Crossing (-BA), Germishuizen
3533 (PRE). - 2832 (Mtubatuba): Ense1eni Nature Garden (-CC), Venter 6115
(PRU).
TRANSKEI. - 3027 (Lady Grey): Herbert, Douglas (-DC), Orpen 57
(BOL). - 3127 (Lady Frere): Glen Grey, Nzebanya Mountain (-CC), Galpin
1914 (PRE).
LEDEBOURlA IN SOllTH AFRICA
325
TAXONOMY
CAPE. - 2824 (Kimberley): Riverton Pont (-DB), Leistner 2016 (PRE).
- 3226 (Fort Beaufort): Katberg Mountain, above Seymour (-DB), Bayliss 7442
(NBG). - 3227 (Stutterheim): Pirie (-CC), Taylor 1760 (BOL); King Williams
Town (-CD), Sim 628 (NU); Sim 1075 (BOL); de Victoria 15 (BOL). - 3326
(Grahamstown): Blaauw Krantz (-BD), Daly 1026 (BOL); Peddie, Committee's
Drift (-BB), lfln der Merwe 2132 (PRE). - 3327 (Peddie): East London (-BB),
WJod 2817 (BOL); Rattray s.n. sub BOL 13,689 (BOL); Pamphlett 77 (NBG).
- 3228 (Butterworth): Kwenqura River (-CC), Galpin 5814 (PRE). - 3421
(Riversdale): Riversdale (-AD), Bohnen 7461 (STE); Still Bay (-AD), Bolus s.n.
sub BOL 19,487 (BOL); Ferguson s.n. sub BOL 22,515 (BOL); Albertinia,
Ystervarkpunt (-BC), Willemse 107 (STE). - 3423 (Knysna): Knysna (-AA),
Breyer 23, 711 (PRE).
UDEBOURJA IN SOlITH AFRICA
326
TAXONOMY
Sectio Paucifoliae S. Venter, sect. nov., foliis numquam plus quam quatuor;
cataphylliis duo vel tres; ovario depresse ovato.
Species typica: Ledebouria galpinii (Bak.) S. Venter.
Lectotypus: Transvaal, Kaapsche Hoop, Galpin 672 (PRE).
Species: L. galpinii (Bak.) Jessop, L. papillata S. Venter.
Plants 30 - 70 mm tall. Bulb 8 - 25 mm wide; neck 3 - 7 mm long; basal stem
present; cataphylls 1 - 3. Leaves 2 - 4, adaxial surface unspotted. Inflorescence
rachis ridged; raceme 10 - 20 mm wide; bracts fleshy. Tepal segments slightly
cucullate. Ovary depressed-ovate, 1.5 mm long. Seed 1.5 - 2.0 mm long.
Distribution and habitat.
Transvaal and the eastern Cape, in montane grassland and coastal shrubland.
32. LEDEBOURIA GALPINII (Bak.) S. Venter
Ledebouria galpinii (Bak.) S.
retn~
comb.nov.
Scilla galpinii Bak. in Flora Cap. 6: 487 (1896).
Type: Transvaal, summit of Devil' s Kantoor, Galpin 672 (PRE! 1ecto; BOLl;
GRA!; NH!; PRE!, photo.; SAM!; Z). Designated here as lectotype (Greuter et
al. 1988).
Plants sometimes solitary, usually gregarious. Bulb hypogeal, 20
- 30 X 20 - 25 mm, ovoid to subglobose; dead bulb scales brown, apices
truncate, live bulb scales fleshy, loosely arranged, white with numerous purplish
spots in upper part, with occasional threads when torn, neck 1 - 2 x 3 - 5 mm;
bulblets on basal stem; cataphylls 1 - 3, exerted above ground level. Leaves
fully developed at anthesis, 3 - 4, humifuse, oblong to ovate-spathulate, 50 - 80
x 20 - 25 mm, without threads when torn, thickly fleshy, glossy purple to
purplish-green, adaxial surface distinctly lacunose, venation obscure; margins
smooth; leaf base canaliculate; apex acuminate to mucronate. Inflorescences 1 -
LEDEBOURlA IN SOUTH AFRICA
327
TAXONOMY
3, dense, sub-globose, 10 - 20 x 10 - 20 mm, flaccid, 20 - 30 -flowered, as
long or longer than the leaves, lengthening when in fruit; scape winged at base,
purple; rachis ridge4, 40 - 50!,~_I(mg)Bracts
fleshy, 1 x 0.5 mm, linear-
lanceolate, pink to purple withbiacteoles present. Pedicels spreading, 3.0 - 4.2
mm long, purple. Perianth 4.0 - 4.2 mm long, stellate, tepals equal, oblong, 4 5 x 2 mm, apex obtuse, cucullate, pink to purple sometimes with a dull green
keel. Stamens spreading, 3.8 - 4.0 mm long; filaments maroon, epitepalous;
anthers 0.5 mm long, violet. Ovary globose, 3 -lobed, 1.5 x 1 mm, lobes
obtusely depressed-ovate. Style 3 mm long, triangular, glabrous, purple; stigma
equalling anthers; stipe 0.25 x 0.5 mm. Capsule three-lobed, symmetrical,
globose; base truncate. Seed drop-shaped, 1.5 - 2.0 mm long, surface strongly
wrinkled, reddish-brown. (Figure 78).
This species is closely related to L. papillata but differs in having humifuse
leaves with prominent corrugations (lacunae) on the adaxial surface (Figure
79B) and flaccid inflorescence.
Specific epithet etymology.
Commemorates E.E. Galpin who collected the type material.
Flowering period
From October to November.
Distribution (Map 36).
L. galpinii is known only from Kaapsche Hoop in the eastern Transvaal.
LEDEBOURIA IN SOlJfIl AFRICA
328
TAXONOMY
E
o
G
H
c
B
Figure 78. Illustration of L. galpinii (Bak.) S. Venter. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, apex of lamina X 20; E,
bracts X 10; F, flower X 10; G, apex of tepal X 20; H, tepal with stamen X 10;
I, ovary lateral view X 10; J, ovary dorsal view X 10. All from \enter 13,389.
UDEBOURlA IN SOlJfH AFRICA
Map 36.
140
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TAXONOMY
330
Figure 79. A, lectotype of L. galpinii (Bak.) S. Venter (PRE); B, plants with
prominent lacunae on the leaves. Bar
=
10 mm; C, habitat at Kaapsche Hoop,
eastern Transvaal. The vegetation consists of closed short Panicum natalense -
Eragrostis racemosa - Scabiosa columbaria montane grassland; D, SEM
micrograph of the adaxial lamina surface. Bar = 50 JLm; E, SEM micrograph of
the abaxial lamina surface. Bar = 50 JLm; F, SEM micrograph of stomata. Bar
= 50 JLm. Band D - F from Venter 13,389.
LEDEBOURIA IN SOlITH AFRICA
331
TAXONOMY
~
/
Jr'
A
.~_,
LED£BOURlA IN SOUTH AFRICA
332
TAXONOMY
Habitat
Kaapsche Hoop is situated on quartzites and conglomerates of the Black Reef
Quartzite Formation of the Wolkberg Group (SACS 1980). Soil derived from
these quartzites is a shallow grey sandy soil with the grains 0,25 - 1.0 mm
diameter. Wherever L. galpinii occurs, the soil has a high humus content.
L. galpinii grows in protected soil pockets between and in openings under stones
and boulders. It grows in association with various moss species and
StreplOcarpus galpinii J.D. Hooker (Hilliard & Burtt 1971). Plants receive
dire<:t sunlight in the mornings and are frequently subjected to mist (Figure
79C).
Population structure
The 19 populations surveyed extend over
± lkm 2 and
usually number 5 - 20
individuals but one population that occurs in a vlei consists of 1000+
individuals.
Variation
Plants growing in dense shade tend to have long, green leaves which are not
appressed to the ground. Plants have pure pink to purple flowers and shaded
individuals develop a prominent dull green keel on the tepals.
Specimens examined
TRANSVAAL. - 2530 (Lydenburg): Kaapsche Hoop, Devil's Kantoor (_
DB), Galpin 672 (BOL, GRA, NBG, NH, PRE); WIn der Merwe 2047 (PRE);
Codd 9789 (PRE); Venter 13,389 (UNIN).
LEDEBOURIA IN sOurH AFRICA
333
TAXONOMY
33. LEDEBOURIA PAPILLATA S. Venter
Ledebouria papillata S. \enter, sp.nov., ad L. cooperi (Hook}:) Jessop affinis
sed squamis ad apicem truncatis; petiolo purpureo fasciato; inflorescentiis
erectis, pedunculo pappilato; ovario stipitato, stipite 0.5 mm longo; ovario
dorsaliter lobato differt.
Type: Transvaal, Pietersburg, \enter 13,186 (PREl, holo.; NU,I; UNIN I).
Plants solitary. Bulb hypogeal, 15 - 30 x 8 - 20 mm, obovoid; dead bulb scales
light brown, membranous, apices truncate, live bulb scales fleshy, without
threads when torn, white inside, neck 3 - 27 x 3 - 7 mm; basal stem 5 - 15 mm
long. Cataphylls 2, reaching ground level. Leaves fully developed at anthesis,
2 - 4, spreading, linear-lanceolate to oblanceolate, 30 - 90 x 4 - 10 mm, with
few threads when torn, fleshy, adaxial surface glossy dark green, abaxial
surface dull purplish green with rows of longitudinal papillae, prominent dark
purple cross bars at base, venation prominent; margins papillate; leaf base
petiolate, canaliculate; apex acute to acuminate. Inflorescences 1 - 2, dense,
cylindric, 20 - 60 x 10 - 20 mm, erect, 14 - 45 -flowered, longer than the
leaves; scape terete at base, green, spotted and striped purple, longitudinally
papillate, rachis ridged, 50 - 120 mm long. Bracts fleshy, 0.5 x 0.25 mm,
oblong to bifurcate, pink, white or green without bracteoles. Pedicels spreading·
horizontally, 2.5 - 5.0 mm long, pink. Perianth 2.0 - 3.5 mm long, tepals
reflexed, equal, oblong, 3.0 - 3.5 x 1.0 - 1.5 mm, apex obtuse, thinly cucullate,
pink to purple with a green keel. Stamens erect, 3 mm long, filaments maroon,
epitepalous; anthers 0.5 mm long, violet. Ovary globose, 6 -lobed, 1.5 x 2.5
mm, lobes depressed ovate, apex shoulders raised, distallobes present. Style
3.5 mm long, terete, glabrous, purple; stigma equal height to anthers; stipe 0.5
x 0.5 mm. Capsule three-lobed, symmetrical, globose; base tapering. Seed
globse, 2 mm long, surface strongly wrinkled, brown. (Figure 80).
L. papillata is closely related to L. galpinii and together they constitute the
section Paucifoliae. It differs from L. galpinii in having longitudinal rows of
papillae on the adaxial leaf surface, purple cross bars on the petiole, erect
inflorescences and papillate peduncles.
LEDEBOURlA IN SOlTfH AFRICA
334
TAXONOMY
E
o
F
G
&
c{JJ
J
B
A
Figure 80. Illustration of L. papillata S. Venter. A, habit X 2; B, section
through leaf X 5; C, lamina margin X 300; D, bract X 10; E, papillae on
peduncle X 10; F, flower X 10; G, tepa! with stamen X 10; H, apex of tepa! X
20; I, ovary lateral view X 10; J, ovary dorsal view X 10. All from Venter
13,008.
UDEBOURJA IN SOlJfH AFRICA
Known distribution of L. papillata S. Venter
Map 37.
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336
TAXONOMY
Figure 81. A, holotype of L. papillata S. Venter (PRE); B, habitat near
Pietersburg. The vegetation consists of deciduous open low Aloe marlothii var.
marlothU - Ormocarpum trichocarpum - Themeda triandra woodland; C, SEM
micrograph of the adaxial lamina surface. Bar = 100 JLm; D, SEM micrograph
of the abaxial lamina surface. Bar
=
100 J.tm; E, SEM micrograph of stomata.
Bar = 43 J.tm. A and C - E from lenter 13,186.
LEDEBOURlA IN SOUTH AFRICA
337
TAXONOMY
J'
/
:J,t
HERBARIUM
'l'I.IIIIJnOlT"l'lfOfII'lO
U.O_A .OVTM ,""'CA
LEDEBOURlA IN SOurH AFRICA
338
TAXONOMY
Specific epithet etymology.
Describes the papillate surface of the plant.
Flowering time
From October to January with a peak in November.
Distribution (Map 37).
The distribution is mainly in the Transvaal with a strangely disjuncted extension
in the eastern Cape near Port Elizabeth and Peddie.
Habitat
Plants of L. papillata grow in shallow (10 - 35 mm deep), medium grained
(0.25 - 1.0 mm 4» to fine grained (0.025 - 0.25 mm 4», sandy to sandy loam.
Plants grow in small populations in woodland.
In the eastern Cape, near Grahamstown, populations occur on Dwyka Tillite of
the Karoo Sequence (SACS 1980). The vegetation consists of grassland with
scattered bushc1umps. At 'The Aloes' near Port Elizabeth, L. papillata occurs
on limestone and sand of the Alexandria Formation in the Kalahari Group
(SACS 1980). The vegetation is a closed evergreen tall shrubland.
Variation
The neck of the bulb is usually short (3 - 5 mm) in plants growing in shallow
soil in exposed situations. Plants from the Blouberg grow in deeper soil
resulting in a longer neck (10 - 27 mm). Plants from Port Elizabeth (Venter
13,308) exhibit long linear-lanceolate leaves compared to the short oblanceolate
leaves of the Transvaal populations.
LEDEBOURlA IN SOtJrH AFRICA
TAXONOMY
339
Historical background
Leendertz (from the Transvaal Museum) collected the first specimen,
subsequently only a few collections have been made mainly in the Transvaal.
Jessop (1970) regarded L. papillata merely as a form of L. cooperi but did not
mention the papillate nature of the epidermis.
Specimens examined
TRANSVAAL. - 2329 (Pietersburg): Louis Trichardt (-BB), Breyer 22,714
(PRE); Pietersburg (-CD), Van der Merwe 2244 (PRE); farm Bloedrivier,
Venter 12,216 (UNIN); Flora Park, Venter s.n. (UNIN). - 2428 (Nylstroom):
Naboomspruit, farm Mosdene (-DA), Galpin M362 (PRE). - 2429 (Zebediela):
Potgietersrus (-AA), Leendertz 6627 (PRE); Potgietersrus, farm Riebeeck West,
liln der Merwe 1744 (PRE). - 2527 (Rustenburg): Witwatersberg, Nefdt (-DD),
Prosser P1109 (PRE). - 2627 (Potchefstroom): Potchefstroom (-CA), Ubbink
332 (PUC); Potchefstroom Dam, Ubbink 742 (PUC).
CAPE. - 3325 (Port Elizabeth): Uitenhage (-CD), liln Jaarsveld 9115
(NBG); The Aloes (-DC), Venter 13,308 (UNIN). - 3327 (Peddie): Peddie (AA), lfln der Merwe 2129 (PRE).
LEDEBOURlA IN SOlITH AFRICA
340
NOMINA DUBIA AND NOMINA EXCLUSA
13.0. NOMINA DUBIA AND NOMINA EXCLUSA.
Nomina dubia
Drimia angustifolia Kunth in Enum. PI. 4: 340 (1843).
Type: Cap. b. Spei, Drege 8618b. Not located and description inadequate.
Drimia dregeana Kunth in Enum. PI. 4: 340 (1843).
Type: Cap. b. Spei, Drege 1616c. Not located and description inadequate.
Drimia nitida EcId. in S. Afr. Quart. JOUffi. 1: 364 (1830).
Type: Uitenhage, Addo, Drege 8616b. Not located and de~cripton
inadequate.
Drimia viridiflora EcId. in Topogr. Verzeichn. von Chr. Fr. EcIdon (1827).
Type: No locality, Zeyher s. n. Specimen not found and there is no description.
Drimia viridiflora Kunze in Linnaea 20: 10 (1847).
Type not located and description inadequate.
Scilla leichtlenii Bale in Flora Cap. 6: 486 (1896).
Type: Without locality, cultivated specimen. (K!, holo.).
Described from a living plant that flowered with Max Leichtlin, at Baden-Baden
in June 1878. Baker's description does not fit that of any Ledebouria species and
the type specimen consists of a single inflorescence.
Scilla sphaerocephala Bak. in Flora Cap. 6: 484 (1896).
Type: Little Namaqualand, Bolus s.n. Type not found and the description
inadequate.
Scilla viridiflora (Kunze) Bak. in 1. Linn. Soc. (Bot.) 13: 255 (1873).
Nomina exclusa
-------.
Hyacinthusflexuosus Thunb. in Prodr. FI. Cap. :64 (1794).
Type: Cape, Thunberg s. n. = Ornithogalum sp.
Scillaflexuosa (Thunb.) Bak. in 1. Linn. Soc. (Bot.) 13: 245 (1873).
LEDEBOlfRJA IN SOlTfH AFRICA
341
HYBRIDS AND CULTURAL USES
14.0. HYBRIDS.
Only one putative hybrid in the genus Ledebouria is known. On Settlers Hill at
Grahamstown, plants were found that looked superficially like L. revoluta. On
closer inspection these plants showed characters of both L. hypoxidioides and L.
revoluta both of which occur at this locality.
15.0. CULTURAL USES.
15.1. Food.
Some species of Ledebouria are a popular food source for various animals. L.
ovatifolia bulbs are dug up and eaten by Chacma Baboon, [Papio ursinus (Kerr,
1792)] in the Rustenburg Nature Reserve in the north-western Transvaal (pers.
obs.). Notes to this effect are made on the label of a specimen collected at the
same locality (Jacobsen 1031).
Ledebouria bulbs are utilized by the Bushman in Namibia, the Kalahari and
Botswana. In the central Kalahari, in the Kweneng District, Botswana, the bulbs
of L. apertiflora are baked in warm ashes before eating and are called
"Dinojane" (Barnard 219).
Bushmen in the Sandfontein area, Namibia, eat the
baked bulbs of L. revoluta (Bleek s.n. sub PRE 26,143).
15.2. Medicine.
Many Ledebouria species are used medicinally by local tribes throughout the
distribution range (Watt & Breyer-Brandwijk 1962).
L. apertijlora - Bulbs are used by the Bakone Sotho of Sekhukhuniland for chest
problems (pers. obs.).
L. concolor - Diviners wash their bodies in water with pieces of the bulb so that
the patients will not mind paying a fee for the consultation (pers. obs.).
UDEBOURIA IN SOlITH AFRICA
342
HYBRIDS AND CULTURAL USES
L. cooperi - A preparation is given to a woman in the fourth month of
pregnancy to ease her discomfort. A decoction is given to cows to ensure a
succession of calves of one sex (Jacot Guillarmod 1971).
L. hypoxidioides - Xhosa herbalists use bulbs of this species in and around
Grahamstown as a remedy for stomach problems (pers. obs.).
L. luteola - At Melville Koppies area near Johannesburg, Sotho and Tswana
people use crushed bulbs as an enema for children with meningitis (Notes on
label of Macnae 1166).
L. marginata - In the Melville Koppies area near Johannesburg bulbs are cooked
and the infusion used as enema for children with' meningitis' (Macnae 1166).
Also used in treating lumbago (Jacot Guillarmod 1971). Meat of sheep cooked
with the bulbs can be used as a remedy against insanity (diviner pers. comm.)
L. revoluta - The Tswana use a decoction of the bulb as a cure for sadness. If
the heart beats too fast, the patient must eat bulbs boiled with chicken. To
prevent mishaps, bulbs are mixed with Craterostigma plantagineum Hochst.,
Pittosporum viridiflorum Sims and Brunsvigia radulosa (Jacq.) Aitk. A plant of
this species in a village gives protection and a person who speaks badly of the
village will become ill at his house. L. revoluta is used as muti to assure a
bumper harvest. In Lesotho the natives cultivate this plant outside houses as a
good luck charm and to drive away lightning (Jacot Guillarmod 1971).
Ledebouria sp. - Used by the Nanga of Zimbabwe as Fontanelle - "chipande"
(Gelfand et al 1985).
15.3. Poisoning.
Some species of Ledebouria are toxic. The following species contain cardiac
poisons: L. inquinata, L. revoluta, L. ovatifolia and L. cooperi.
LEDEBOURlA IN SOUTH AFRICA
343
HYBRIDS AND CULTURAL USES
L. cooperi - This plant has the typical action of the digitalis group. It was shown
that the minimal one-hour systolic doses per gram for a frog was 0.0055 - 0.02
cc (Gunn et al. 1924). Toxic to sheep but seldom eaten in quantity (Galpin
5814).
L. ovatifoiia - The results of toxicity tests done at Onderstepoort Research
Station during December 1957 are given in table 10.
Thble 10. Toxicity tests with L. ovatifolia.
Animal
Weight (Kg)
Dosage
Period
Result
Rabbit
Rabbit
Sheep
Sheep
Sheep
2.5
2.75
27.21
26.3
25.8·
20g
20g
30g
50g
50g
3 days
3 days
5 days
2 days
5 days
Died
Recovered
No effect
Died
Very sick
The level where this plant is toxic is 2g per Kg live mass. The bulbs used were
sent from Horse Shoe Farm, Port Shepstone.
L. revoluta - The bulb contains a combination of alkaloids (Willaman &
Schubert 1958). A decoction of these bulbs is used to treat gall-sickness (Smith
1895). Bulbs are inedible and toxic according to the !kun Bushmen (Story
1958). Produces ill effects if eaten by sheep but not usually fatal (Gane 202).
LEDEBOURlA IN SOlTfH AFRICA
344
HYBRIDS AND CULTURAL USES
15.4. Vernacular names.
L. apertijlora - (Afrikaans) Kaffir Ui; (Sotho) Mokxetlepyane.
L. concolor - (Xhosa) iDumu.
L. cooperi - (Sotho) beokho, bookhoe, phetola (Jacot-Guillarmod 1971).
L. luteola - (Tswana) Untlokwana.
L. marginata - (Tswana) Untlokwana, Bogokwe, Bogokgwe, Mositakgomo.
L. ovalifolia - (English) Reflex-flowered Lachenalia (Andrews 1803).
L. revoluta - (!kun Bushman)' lo/ni, //nanna, //khabhe; (English) Wave-leaved
Hyacinth (Aiton 1789), Spotted Copperas-leaved Lachenalia (Aiton 1811);
(German) Lanzenblattrige Lachenalie (Willdenow 1799), Gebogene Hyacinthe
(Willdenow 1799); (Shona) chitupatupa (Gelfand et al. 1985); (Sotho) boakhoe,
boekhoe, bookhoe; (Tswana) Sejabaleki, Mositakgomo, Bogokgwe; (Xhosa)
ubuHlungu; (Zulu) iCubudwane.
L. undulata - (German) Wellenblattrige Giftlilie (Willdenow 1799).
UDEBOURlA IN SOlJrH AFRICA
345
CAPTIONS TO ILLUSTRATIONS AND TABLES
16.0. CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 1 - 6. 1. The bulbs of A, Ledebouria revoluta (L.f.) Jessop
(~nter
13,209); B, Drimiopsis sp. novo [Venter 13,459 (UNIN)] and C, Scilla
dracomontana Hilliard [Edwards
bulb scales. Bar = 50 mm.
S. n.
(UNIN)] to show the differences in the
Figure 2. Leaves of the genera A, Scilla natalensis Planch. [Venter s.n.
(UNIN)]; B, Ledebouria revoluta
(~nter
13,209) and Drimiopsis burkei Bak.
[l-Cnter s.n. (UNIN)] to show the differences in the leaves. Bar = 100 mm.
Figure 3. Three different types of root in the genus Ledebouria. A, contractile
roots of L. revoluta (l-Cnter 13,207); B, fusiform roots of L. apertijlora (Bak.)
Jessop (l-Cnter 12,686) and C, fleshy roots of L. cooperi (Hook!) Jessop (Glen
2295). Bar = 50 mm.
Figure 4. Plants of L. crispa S. Venter showing the gregarious habit (~nter
11 ,202). Bar = 10 mm.
Figure 5. Basal stems of L. glauca S. Venter (l-Cnter 13,368). Bar = 50 mm.
Figure 6. SEM micrograph of the spiral threads in the leaves of L. crispa
(l-Cnter 13,209a). Bar = 5 J-tm.
Figure 7. Different leafshapes in Ledebouria. A, L. cooperi (l-Cnter 13,342); B,
L. revoluta (Venter 13,207); C, L. revoluta (Venter 13,203); D, L. luteola
(l-Cnter 13,217); E, L. revoluta (Venter 13,362); F, L. revoluta (l-Cnter 13,363);
G, L. marginata (Venter 13,358); H, L. asperifolia (Venter 13,382); I, L.
revoluta (Scott-Shaw s.n.); J, L. cooperi (~nter
13,334); K, L.floribunda
(~nter
13,315); L, L. zebrina (Cunningham s.n.); M, L. revoluta ( ~nter
13,322); N, L. cooperi (Venter 13,388); 0, L. asperifolia (~nter
13,382); P,
L. cooperi (Venter 13,383); Q, L. revoluta (~nter
13,257); R, L. inquinata
(l-Cnter 13,335) and S, L. ovatifolia (l-Cnter 13,337). Bar = 100 mm.
LEDEBOURJA IN SOlITH AFRICA
346
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 8. Flower bracts. A, L. concolor, vestigial bract
floribunda
retn~(
(~nter
s.n.); B, L.
13,315); C, L. apertijlora (Mauve et al. 179); D, L. atro-
13,483); E, L. ovatijolia (~nter
13,376) and F, L. zebrina
brunnea (~nter
(~nter
13,395). Bar = 500 j.tm. Bracteoles indicated with an arrow.
Figure 9. L. revoluta flower. A, lateral view. B, distal view. Both from
~nter
13,()(J7 (X 8).
Figure 10. Tepal apices of A, Drimiopsis burkei Bak.
retn~(
13,341) and B,
Ledebouria apertijlora (Bak.) Jessop (Mauve et al. 179) (X 16).
Figure 11. Various parts of the ovary (X16).
Figure 12. SEM micrographs of A, tepal apex in L. zebrina (Bak.) S. Venter
retn~(
13,395). Bar = 50 j.tm. B, anther of L. ensijolia (Eckl.) S. Venter
(Venter 13,521). Bar = 0.38 mm. C, pollen grain of L. revoluta (Lf.) Jessop
(~nter
13,207). Bar = 12.6 j.tm. D, reticulate exine of L. revoluta (~nter
13,207). Bar = 5 j.tm. E, stigma of L. revoluta (Venter 13,257). Bar = 86 j.tm.
F, nectaries on the base of the ovary lobes in L. sandersonii (Bak.) S. Venter
(Venter 13,465). Bar = 200 j.tm. G, seed surface of L. glauca S. Venter (Venter
13,386). Bar = 1.2 mm. H, detail texture of the testa of L. luteola Jessop. Bar
= 43 j.tm.
Figure 13. SEM micrographs of the different ovary shapes. A, L. undulata
(Jacq.) Jessop (Muller-Doblies 89129). Bar = 500 j.tm. B, L. papillata S.
Venter (Venter 13,186). Bar
Bar
=
1 mm. C, L. parvijolia S. Venter (Venter s.n.).
= 1 mm. D, L. ensijolia (Eckl.) S. Venter (Venter 13,278). Bar
= 500 j.tm.
E, L. apertiflora (Bak.) Jessop (Mauve et al. 179). Bar = 500j.tm. F, L.
leptophylla (Bak.) S. Venter (Venter 13,251). Bar = 500 j.tm. G, L. asperijolia
(Van der Merwe) S. Venter (Venter 13,382). Bar = 0.86 mm. H, L. zebrina
(Bak.) S. Venter. Bar = 500 j.tm. and I, L. hypoxidioides (Schonl.) Jessop
(Venter 13.311). Bar = 500 j.tm.
LEDEBOURIA IN SOUTH AFRICA
347
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 14. Meiotic chromosomes (metaphase I) L. apenijlora: 20 = 26. A - C
from Venter s.n. and L. ensifolia: 2n
= 30. D - F from Venter 13,278. A, C
and E X 1000; B, X 600 and F X 400.
Figure 15a. Graphic guide for sandy soil textural classification with A, the
different types of sandy soil for Ledebouria (shaded portion) and B, soil textures
of the habitats (shaded portion).
Figure 15b. Phenogram generated with the program 'NTSYS-pc'. ape = L.
apenijlora; ens = L. ensifoiia; san = L. sandersonii; dol = L. dolomiticola; atr
= L. atro-brunnea; vis = L. viscosa; lpt = L. leptophylla; lep = L. lepida;
min = L. minima; rup = L. rupestris; cor = L. coriacea; flo = L. floribunda;
hyp = L. hypoxidioides; rev = L. revoluta; zeb = L. zebrina; lut = L. luteola;
ovt = L. ovatifolia; mon = L. monophylla; coo = L. cooperi; par = L.
parvifolia; asp = L. asperifolia; gla = L. glauca; inq = L. inquinata; mar =
L. marginata; con = L. concolor; cri = L. crispa; und = L. undulata; ov1 =
L. ovalifoiia; pet = petiolata; soc = L. socialis; mac = L. macowanii; gal =
L. galpinii and pap = L. papillata.
Figure 16. Illustration of L. apenijlora (Bale.) Jessop. A, habit X 1. B, lamina
margin X 300. C, bract X 10. D, tepal with stamen X 10. E, apex of tepal X
20. F, section through lamina X 5. G, flower X 10. H, lateral view of ovary X
10 and I, distal view of ovary X 10. All from Venter s.n.
Figure 17. A, lectotype of L. apenijlora (Bale.) Jessop under Drimia apenijlora
Bale. in Saunders Refugium Botanicum 1 : 1. 19
(1868); B, SEM micrograph of the adaxial lamina surface. C, SEM micrograph
of the abaxial lamina surface. D, habitat near Phalaborwa, north-eastern
Transvaal. The vegetation consists of closed deciduous tall Combretum
apiculatum - C. imberbe - Acacia nigrescens woodland; E, SEM micrograph of
stomata and F, SEM micrograph of the fine texture of the testa. B - C and E - F
from Venter s. n.
LEDEBOURlA IN SOlITH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
348
Figure 18. Illustration of L. ensifolia (Eckl.) S. Venter. A, habit X 1; B, lamina
margin X 300; C, lamina apex X 10; D, bract X 10; E, section through lamina
X 5; F, tepal apex X 20; G, tepal with stamen X 10; H, flower X 10; I, ovary
distal view X 10; J, ovary lateral view X 10. All from Smith 155.
Figure 19. A, lectotype of L. ensifolia (K); B, habitat at the Zwartkops River
estuary. The vegetation consists of tall, closed, evergreen shrubland; C, SEM
micrograph of the adaxial lamina surface. Bar = 50 ~m;
D, SEM micrograph
of the abaxial lamina surface. Bar = 50 ~m;
E, SEM micrograph of stomata.
F, inflorescence. Bar = 10 mm. C - F from '{knter 13,278.
Bar = 50 ~m;
Figure 20. Illustration of L. sandersonii (Bak.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, section through lamina X 2; D, bract X 10; E, tepal
with stamen X 10; F, flower X 10; G, section through lamina depicted in H, X
2; H, lamina, shade form Xl; I, tepal apex X 20; J, ovary lateral view X 10;
K, ovary distal view X 10. A - F from Crouch 7 and H to K from Venter
13,464.
Figure 21. A, type of L. sandersonii (Bak.) S. Venter (K); B, plants of the
spotted-leaf form, near Dullstroom, eastern Transvaal. Bar = 25 mm; C, SEM
D, SEM micrograph
micrograph of the adaxial lamina surface. Bar = 50 ~m;
of the abaxial lamina surface. Bar = 50 ~m;
E, SEM micrograph of stomata.
Bar = 50 ~m;
F, habitat in the basalts at Cathedral Peak. The vegetation
consists of short montane grassland. C - E from Crouch 7.
Figure 22. Illustration of L. dolomiticola S. Venter. A, habit X 1; B, lamina
margin X 300; C, lamina apex X 20; D, bract and bracteole X 10; E, section
through lamina X 5; F, tepal apex X 20; G, flower X 10; H, tepal with stamen
X 10; I, ovary lateral view X 10; J, ovary distal view X 10. All from Venter
13,208a.
Figure 23. A, holotype of L. dolomiticola S. Venter (PRE); B, habitat at
Donkerkloof, the type locality. The vegetation consists of closed deciduous low
Kirkia wilmsii - Dombeya autumnalis - Obetia tenax woodland on dolomite', C,
plants of L. dolomiticola growing in rock cracks showing the epigea1 bulbs. Bar
= 100 mm; D, SEM micrograph of the adaxial lamina surface. Bar = 100 ~m;
E, SEM micrograph of the abaxial lamina surface. Bar = 100 ~m;
F, SEM
micrograph of stomata. Bar = 43 .m~
D - F from lknter 13,089a.
LEDEBOURlA IN SOlITH AFRICA
349
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 24. Illustration of L. atro-brunnea S. Venter. A, habit Xl; B, lamina
margin X 300; C, section through lamina X 5; D, bract X 20; E, tepal with
stamen X 10; F, flower X 10; G, tepal apex X 20; H, ovary lateral view X 10;
I, ovary distal view X 10. All from lknter 13,460.
Figure 25. A, holotype of L. atro-brunnea S. Venter (PRE); B, habitat near
Kroondal, north-western Transvaal. The vegetation consists of closed deciduous
tall Protea caffra subsp. caffra - Xerophyta retinervis - Parinari capensis
woodland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of wax platelets
forming the cuticle; F, plants of L. atro-brunnea showing the colonial growth
and the erect, twisted leaves. Bar
= 30 mm.
C - F from lknter 13,460.
Figure 26. Illustration of L. viscosa Jessop. A, habit Xl; B, lamina margin X
300; C, section through lamina X 2; D, bract X 10; E, flower X 10; F, tepa!
with stamen X 10; G, tepal apex X 20; H, ovary lateral view X 10; I, ovary
distal view X 10. All from Venter 13,455.
Figure 27. A, holotype of L. viscosa Jessop (PRE); B, habitat near Thabazimbi.
The vegetation consists of closed deciduous low Terminalia sericea - Acacia
tortilis subsp. heteracantha - Grewia flava - Digitaria eriantha woodland; C,
SEM micrograph of the adaxial lamina surface; D, SEM micrograph of the
abaxial lamina surface; E, leaves of L. viscosa with sand particles adhering to
the surfaces. Bar
=
10 mm. F, SEM micrograph of the stomata; G, SEM
micrograph showing the thick resinous covering of the lamina; H, leaf
unrolling, a unique character in Ledebouria. Bar = 20 mm. C - H from Venter
13,455.
Figure 28. Illustration of L. leptophylla (Bak.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, section through lamina X 10; D, tepal apex X 20; E,
tepal with stamen X 10; F, bract with bracteole X 10; G, flower X 10; H, habit
of nonspiral-leaved form Xl; I, ovary lateral view X 10; J, ovary distal view
X 10. All from Venter 13,214 except H from Venter 13,251.
LEDEBOURlA IN SOlITH AFRICA
350
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 29. A, holotype of L. leptophylla (Bak.) S. Venter (K); B, miniature
form resembling L. minima (Bak.) S. Venter. Bar = 20 mm; C, SEM
micrograph of the adaxial lamina surface; D, SEM micrograph of the abaxial
lamina surface; E, SEM micrograph of stomata; F, habitat at Angle Station,
between Barberton and Havelock. The vegetation consists of open low Protea
roupelliae var. roupelliae - Helichrysum splendidum - Xerophyta retinervis
woodland; G, plants with nonspiralleaves. Bar = 20 mm. B and G from \enter
13,251 and C - E from Venter 13,214.
Figure 30. Illustration of L. lepida (N.E. Br.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, bract X 10; D, hairs on peduncle X 10; E, section
through lamina X 2; F, flower X 10; G, tepal with stamen X 10; H, tepal apex
X 10; I, ovary lateral view X 10; J, ovary distal view X 10. All from Venter
13,415.
Figure 31. A, holotype of L. lepida (N.E. Br.) S. Venter (K); B, habitat near
Palala, Waterberg. The vegetation consists of closed deciduous low Combretum
zeyheri - Mimusops zeyheri - Freylinia tropica woodland; C, SEM micrograph
of the adaxial lamina surface. Bar
=
100 J.tm; D, SEM micrograph of the
abaxial lamina surface; E, SEM micrograph of stomata. Bar
=
43 J.tm; F,
abaxial surface of the lamina showing the prominent purple zebra stripes on the
petiole. Bar = 10 mm. C - F from Venter 13,415.
Figure 32. Illustration of L. minima (Bak.) S. Venter. A, habit X 1; B, section
through lamina X 5; C, lamina margin X 300; D, bract X 10; E, tepal apex X
20; F, flower X 10; G, tepal with stamen X 10; H, ovary lateral view X 10; I,
ovary distal view X 10. All from Venter 13,404.
Figure 33. A, holotype of L. minima (Bak.) S. Venter (K); B, habitat on "The
Downs" near Trichardtsdal, north-eastern Transvaal. The vegetation consists of
Tristachya leucothrix - Panicum natalense - Scilla natalensis montane sour
grassland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata; F,
plants showing filiform leaves. Bar = 20 mm. C - F from Venter 13,404.
LEDEBOURlA IN SOurH AFRlCA
351
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 34. Illustration of L. rupestris (Van der Merwe) S. Venter. A, habit X 2;
B, lamina margin X 300; C, section of the peduncle X 10; D, section through
lamina X 5; E, bract with bracteole X 10; F, tepal with stamen X 10; G, tepal
apex X 20; H, flower X 10; I, ovary lateral view X 10; J, ovary distal view X
10. All from lknter 13,250.
Figure 35. A, holotype of L. rupestris (Van der Merwe) S. Venter (PRE); B,
habitat near Mac Mac Falls, eastern Transvaal. The vegetation consists of open
short Pteridium aquilinum - Parinari capensis - Digitaria eriantha montane
grassland; C, plants showing globose inflorescences with stellate flowers. Bar
20 mm; D, SEM micrograph of the adaxial lamina surface. Bar
=
= 100 JLm; E,
SEM micrograph of the abaxial lamina surface. Bar = 100 JLm; F, SEM
micrograph of a stomata. Bar
= 43 JLm. C - F from lknter 13,250.
Figure 36. Illustration of L. coriacea S. Venter. A, habit X 2; B, lamina margin
X 300; C, section through lamina X 5; D, lamina apex X 20; E, bract X 10; F,
flower X 10; G, tepal apex X 10; H, tepal with stamen X 10; I, ovary lateral
view X 10; J, ovary distal view X 10. All from lknter 13,307.
Figure 37. A, holotype of L. coriacea S. Venter (PRE); B, habitat at 'The
Aloes near Port Elizabeth. The vegetation consists of evergreen tall Euclea
I
undulata - Ficus burtt-davyi - Brachylaena floribunda shrubland with open
patches covered in Pteronia incana (Burm.) DC. L. coriacea occurs under the
Pteronia shrublets; C, SEM micrograph of the adaxial lamina surface. Bar =
100 JLm; D, SEM micrograph of the abaxial lamina surface. Bar
= 100 JLm; E,
SEM micrograph of stomata. Bar = 43 JLm. A and C - E from lknter 13,307.
Figure 38. Illustration of L. floribunda (Bale) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, lamina apex X 5; D, bract with bracteole X 10; E, section
through lamina X 1; F, flower X 10; G, tepal with stamen X 10; H, tepal apex
X 20; I, ovary lateral view X 10; J, ovary distal view X 10. All from Venter
13,475.
LEDEBOURlA IN SOlITH AFRICA
352
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 39. A, type of L. floribunda (Bak.) Jessop (K); B, habitat at Thabina
Falls in the Wolkberg Wilderness Area near Tzaneen. The vegetation consists of
evergreen low Cliffortia linearifolia - Agapanthus inapertus subsp. pendulus -
Helichrysum splendidum shrubland in a seepage area next to a waterfall; C,
SEM micrograph of the adaxial lamina surface. Bar = 50 JLm; D, SEM
micrograph of the abaxial lamina surface. Bar = 50 JLm; E, SEM micrograph of
stomata. Bar = 50 JLm. C - E from Venter 13,475.
Figure 40. Illustration of L. hypoxidioides (Schonl.) Jessop. A, habit X 0.25; B,
lamina margin X 110; C, section through lamina X 2; D, bract X 10; E, tepal
with stamen X 10; F, flower X 8; G, tepal apex X 20; H, ovary lateral view X
10; I, ovary distal view X 10. All from Venter 13,311.
Figure 41. A, holotype of L. hypoxidioides (Schonl.) Jessop (GRA); B, habitat
at Grahamstown. The vegetation consists of False Fynbos with Passerina rigida
- Ficus burtt-davyi - Euphorbia tetragona shrubveld; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of stomata; F, plants from the type locality. Note the hairy
leaves. Bar = 10 mm; G, SEM micrograph of the leaf surface to show the
hairs. C - G from Venter 13,311.
Figure 42. Illustration of L. revoluta (L.t) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina X 2; D, apex of tepal X 20; E, bract
with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H, ovary
lateral view X 10; I, ovary distal view X 10. All from Venter 13,430.
Figure 43. A, type of L. revoluta (Lf.) Jessop under Lachenalia lanceaejolia
Jacq. in Icones Plantarum Rariorum 2: t.402 (1794); B, SEM micrograph of the
adaxial lamina surface. Bar = 100 JLm; C, habitat near Thabazimbi in the
Waterberg, north-western Transvaal. The vegetation consists of closed
deciduous low Albizia tanganyicensis var. tanganyicensis - Croton gratissimus
var. subgratissimus - Myriothamnus jlabellifolia woodland; D, SEM micrograph
of the abaxial lamina surface. Bar
= 100 JLm; E, SEM micrograph of stomata.
Bar = 43 JLm; F, plant of L. revoluta with thickly packed dry bulb scales for
protection against fire. Bar = 30 mm. Band D - E from Venter 13,009.
LEDEBOURlA IN SOlITH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
353
Figure 44. Illustration of L. zebrina (Bak.) S. Venter. A, habit X 0.25; B,
section through lamina X 0.5; C, lamina margin X 300; D, bract with bracteole
X 5; E, tepal with stamen X 5; F, apex of tepal X 10; G, flower X 5; H, ovary
lateral view X 10; I, ovary distal view X 10. All from l-enter 13,395.
Figure 45. A, type of L. zebrina (Bak.) S. Venter under Scilla zebrina Bak. in
Saunders Refugium Botanicum 3: t.185 (1870); B, plant with leaves 1000 mm
long showing growth habit. Bar = 60 mm; C, SEM micrograph of the adaxial
lamina surface; D, SEM micrograph of the abaxial lamina surface; E, SEM
micrograph of a stomatum; F, habitat near Barberton. The vegetation consists of
closed evergreen low Acacia sieberana var. woodii - Pavetta edentula - Schotia
brachypetala woodland; G, plant from Noodsberg near Greytown showing the
prominent purple zebra stripes on the leaves. Bar = 60 mm. B - E from Wnter
13,395 and G from Smith 115.
Figure 46. Illustration of L. luteola Jessop. A, habit Xl; B, lamina margin X
300; C, apex of lamina X 20; D, section through lamina X 5; E, bract with
bracteole X 10; F, flower X 20; G, tepal with stamen X 20; H, apex of tepal X
10; I, ovary lateral view X 10; J, ovary distal view X 10. All from Venter
13,458.
Figure 47. A, holotype of L. luteola Jessop (PRE); B, habitat at Potchefstroom.
The vegetation consists of open deciduous low Acacia caffra - Aloe greatheadii
var. davyana - Themeda triandra woodland; C, plant with the previous years
leaves still on the plant. Bar = 10 mm; D, SEM micrograph of the adaxial
lamina surface; E, SEM micrograph of the abaxial lamina surface; F, SEM
micrograph of stomata. C - F from l-enter 13,217.
Figure 48. Illustration of L. ovatifolia (Bak.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina X 2; D, apex of lamina X 20; E, bract
with bracteole X 10; F, apex of tepal X 20; G, flower X 5; H, tepal with
stamen X 5; I, ovary lateral view X 10; J, ovary distal view X 10. All from
Stinon 11,180.
LEDEBOURJA IN SOlffH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
354
Figure 49. A, holotype of L. ovatifoiia (Bak.) Jessop (distalleft) (K); B, plant
of L. ovatifolia showing the truncate apices of the bulb scales; C, SEM
micrograph of the adaxial lamina surface; D, SEM micrograph of the abaxial
lamina surface; E, SEM micrograph of a stomatum; F, habitat near
Trichardtsdal, north-eastern Transvaal. The vegetation consists of open low
evergreen Protea roupelliae var. roupelliae - Themeda triandra woodland. B
from llin der Merwe 2172 and C - E from Venter 13,349.
Figure 50. Illustration of L. monophylla S. Venter. A, habit Xl; B, lamina
margin X 300; C, section through lamina X 5; D, bract with bracteole X 10; E,
flower X 10; F, tepal with stamen X 10; G, apex of tepal X 20; H, ovary lateral
view X 10; I, ovary distal view X 10. All from Venter 13,235.
Figure 51. A, holotype of L. monophylla S. Venter (PRE); B, plants showing
the solitary humifuse leaf. Bar = 20 mm; C, plant in habitat near Graskop,
showing the solitary leaf and erect globose inflorescence. Bar = 10 mm; D,
SEM micrograph of the adaxial lamina surface; E, SEM micrograph of the
abaxial lamina surface; F, SEM micrograph of stomata; G, habitat at the type
locality near Graskop. The vegetation consists of closed evergreen low Erica
leucopelta - Passerina montana - Pteridium aquilinum shrubland. A - F from
Venter 13,235.
Figure 52. Illustration of L. cooperi (Hookf) Jessop. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, flower X 10; E, bract with
bracteole X 10; F, apex of lamina X 20; G, tepal with stamen X 10; H, ovary
lateral view X 10; I, Ovary distal view X 10. All from Crouch 97.
Figure 53. A, holotype of L. cooperi (Hookf) Jessop (distal specimen) (K); B,
habitat on slopes of Mount Currie near Kokstad with montane grassland and
swampy areas; C, SEM micrograph of the adaxial lamina surface. Bar = 100
; D, SEM micrograph of the abaxial lamina surface. Bar = 100 ~m;
E,
F, plants of L. cooperi showing
SEM micrograph of a stomatum. Bar = 43 ;m~
~m
parallel purple markings on the lamina. Bar = 20 mm. C - F from Venter
13,194.
LEDEBOURlA IN SOtJrH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
355
Figure 54. Illustration of L. parvifolia S. Venter. A, habit X 2; B, lamina
margin X 300; C, section through lamina X 3; D, bract X 10; E, flower XIS;
F, tepal with stamen XIS; G, tepal apex X 20; H, ovary lateral view X 10; I,
ovary distal view X 10. All from l-enter s.n.
Figure 55. A, holotype of L. parvifolia S. Venter (PRE); B, plant showing the
bulbils, small leaves and the solitary inflorescence. Bar = 20 mm; C, SEM
micrograph of the adaxial lamina surface. Bar = 50 Jlm; D, SEM micrograph
of the abaxial lamina surface; E, SEM micrograph of stomata; F, habitat near
Pilgrim s Rest, eastern Transvaal. The vegetation consists of closed deciduous
I
low Dombeya rotundifolia subsp. rotundifolia - Acacia ataxacantha - Scilla
natalensis woodland; G, SEM micrograph of the rows of hairs on the adaxial
lamina surface; A - F from l-enter s. n..
Figure 56. Illustration of L. asperijolia (Van der Merwe) S. Venter. A, habit X
1; B, lamina margin X 300; C, leaf apex X 10; D, section through lamina X 4;
E, bract X 10; F, tepal apex X 20; G, flower X 10; H, tepal with stamen X 10;
I, ovary lateral view X 10; J, ovary distal view X 10. All from l-enter 13,249.
Figure 57. A, holotype of L. asperijolia (Van der Merwe) S. Venter (PRE); B,
habitat on the the Makonjwa Mountain between Barberton and Havelock. Plants
were collected (Venter 13,382) on the cliffs in the foreground; C, plants of L.
asperijolia after a recent veld fire, Lisbon Falls, Graskop. Bar = 100 mm; D,
plant from Barberton showing the rows of asperities on the leaves. Bar = 10
mm; E, SEM micrograph of the adaxial lamina surface. Bar = 100 J.'m; F,
SEM micrograph of the abaxial lamina surface. Bar = 100 Jlm; G, SEM
micrograph of stomata. Bar
=
43 J.'m. C from Venter 13,249 and D - G from
Venter 13,382.
Figure 58. Illustration of L. glauca S. Venter. A, habit Xl; B, lamina margin
X 300; C, section through lamina X 5; D, bract with bracteole X 10; E, apex of
tepal X 20; F, tepal with stamen X 10; G, flower X 10; H, ovary lateral view X
10; I, ovary distal view X 10. All from l-enter 13,368.
LEDEBOURIA IN SOUTH AFRICA
356
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 59. A, holotype of L. glauca S. Venter (PRE); B, habitat in the Kalahari
near Vanzylsrus. The vegetation consists of open low Acacia erioloba - A.
haematoxylon - Stipagrostis uniplumis woodland; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of stomata. A and C - E from
retn~
13,368.
Figure 60. Illustration of L. inquinata (C.A. Srn.) Jessop. A, habit Xl; B,
lamina margin X 300; C, apex of lamina X 10; D, section through lamina X 5;
E, bract with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H,
apex of tepal X 20; I, ovary lateral view X 10; J, ovary distal view X 10. All
from
retn~
13,343.
Figure 61. A, lectotype of L. inquinata (C.A. Srn.) Jessop (K); B, isotype of L.
inquinata designated by Smith as co-type (K); C, habitat at Linksfield Ridge,
Bedfordview, with open evergreen Protea ca/Ira subsp. caffra - Aloe
greatheadii var. davyana - Digitaria eriantha woodland; D, plant showing the
diagnostic moUling on the live bulb scales. Bar = 25 mm; E, SEM micrograph
of the adaxial lamina surface; F, SEM micrograph of the abaxial lamina surface;
G, SEM micrograph of stomata. D - G from
retn~
13,343.
Figure 62. Illustration of L. marginata (Bak.) Jessop. A, habit X 0.5; B, lamina
margin X 300; C, section through lamina Xl; D, bract with bracteole X 10; E.
tepal with stamen X 10; F, flower X 10; G, apex of tepal X 20; H, lamina of
the narrow leaved form from Vryheid X 0.5; I, section through lamina of H. X
2; J, ovary lateral view X 10; K, ovary distal view X 10. A - G and J - K from
retn~
13,327 and H - I from Venter 13,364.
Figure 63. A, lectotype of L. marginata (Bak.) Jessop (GRAZ); B, SEM
micrograph of the adaxial lamina surface; C, SEM micrograph of the abaxial
lamina surface; D, SEM micrograph of a stomatum; E, plant showing diagnostic
twisted leaves. Bar = 30 mm; F, plants of the narrow-leaved form from near
Vryheid. Bar
= 40 mm.
B - E from Venter 13,230 and F from retn~
LEDEBOlfRJA IN SOlJTH AFRICA
13,358.
357
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 64. Illustration of L. concolor (Bak.) Jessop. A, habit X 1; B, lamina
margin X 300; C, vestigial bract X 10; D, section through lamina X 2; E,
flower X 10; F, tepal with stamen X 10; G, apex of tepal X 20; H, ovary lateral
view X 10; I, ovary distal view X 10. All from Stirton 12,474.
Figure 65. A, holotype of L. concolor (Bak.) Jessop (lower specimen) (K); B,
habitat at the mouth of the Zwartkops River near Port Elizabeth. The vegetation
consists of closed evergreen tall Sideroxylon inerme - Aloe ferox - Euphorbia
ledenii shrubland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata. C - E
from ltln Jaarsveld 9060.
Figure 66. Illustration of L. crispa S. Venter. A, habit X 1; B, lamina margin X
300; C, section through lamina X 5. D, bract with bracteole X 10; E, tepal with
stamen X 10; F, flower X 10; G, apex of tepal X 10; H, ovary lateral view X
10; I, ovary distal view X 10. All from 'W?nter 11,202.
Figure 67. A, holotype of L. crispa S. Venter (PRE); B, colony of plants
showing the typical colonial growth. Bar
=
20 mm; C, SEM micrograph of the
adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface; E,
SEM micrograph of a stomatum. All from 'W?nter 11,202.
Figure 68. Illustration of L. undulata (Jacq.) Jessop. A, habit, leafing period X
1; B, habit, flowering period Xl; C, lamina margin X 300; D, bracts with
bracteoles X 10; E, section through lamina X 4; F, flower X 10; G, apex of
tepal X 20; H, tepal with stamen X 10; I, ovary lateral view X 10; J, ovary
distal view X 10. A and C - J from Muller-Doblies 89129 and B from Hall
3200.
Figure 69. A, type of L. undulata (Jacq.) Jessop under Drimia undulata Jacq. in
!cones Plantarum Rariorum 2: t.376 (1794); B, habitat on the Gamsberg Flats,
Bushmenland. The vegetation consists of closed Euphorbia mauritanica - Euclea
undulata shrubland; C, plant of L. undulata with semi-fleshy leaves and the
inflorescence absent. Bar
=
10 mm; D, SEM micrograph of the adaxial lamina
surface; E, SEM micrograph of the abaxial lamina surface; F, SEM micrograph
of stomata. C - F from Muller-Doblies 89129.
LEDEBOURIA IN SOlJfH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
358
Figure 70. Illustration of L. ovalifolia (Schrad.) Jessop. A, habit X 1; B, lamina
margin X 300; C, apex of lamina X 20; D, apex of tepal X 20; E, section
through lamina X 5; F, bract X 10; G, flower X 10; H, tepal with stamen X 10;
I, ovary lateral view X 10; J, ovary distal view X 10. All from \enter 13,263.
Figure 71. A, type of L. ovalifolia (Schrad.) Jessop under Drimia lanceaejolia
Lodd. in Loddiges Botanical Cabinet 3: t.278 (1818); B, plant of L. ovalifolia
with flatly spreading leaves. Bar = 20 mm; C, SEM micrograph of the adaxial
lamina surface; D, SEM micrograph of the abaxial lamina surface. Bar = 50
J.'m; E, SEM micrograph of stomata. Bar = 20 J.'m; F, SEM micrograph of the
parallel rows of papillae on the abaxial lamina surface; G, habitat at Baakens
Ri ver, Port Elizabeth. The vegetation consists of Protea - Watsonia Leucospermum Fynbos. B - F from Venter 13,263.
Figure 72. Illustration of L. petiolata (Van der Merwe) S. Venter. A, habit X 1;
B, lamina margin X 300; C, section through lamina X 5; D, apex of tepal X 20;
E, bract with bracteole X 10; F, tepal with stamen X 10; G, flower X 10; H,
ovary lateral view X 10; I, ovary distal view X 10. All from Middleton s.n..
Figure 73. A, holotype of L. petiolata (Van der Merwe) S. Venter (PRE); B,
habitat near Graskop. The vegetation consists of open short Passerina montana Ischyrolepis schoenoides - Panicum natalense grassland. C, SEM micrograph of
the adaxial lamina surface; D, SEM micrograph of the abaxial lamina surface;
E, SEM micrograph of stomata; F, plants of L. petiolata typically growing in
moss filled rock cracks. Bar = 20 mm. C - F from \enter s.n.
Figure 74. Illustration of L. socialis (Bak.) Jessop. A, habit Xl; B, lamina
margin X 300; C, section through lamina X 5; D, bract X 10; E, flower X 10;
F, tepal with stamen X 10; G, ovary lateral view X 10; H, ovary distal view X
10. All from Venter 13,272.
Figure 75. A, holotype of L. socialis (Bak.) Jessop (K); B, plant showing the
gregarious habit. Plant growing in deep shade. Bar = 40 mm; C, plants
showing the pendulous flowers. Bar = 20 mm; D, SEM micrograph of the
adaxial lamina surface; E, SEM micrograph of the abaxial lamina surface; F,
SEM micrograph of stomata. B - F from Venter 13,272.
LEDEBOURIA IN SOlJTH AFRICA
359
CAPTIONS TO ILLUSTRATIONS AND TABLES
Figure 76. Illustration of L. macowannii (Bak.) S. Venter. A, habit X 1; B,
lamina margin X 300; C, section through lamina X 2.5; D, bract X 10; E, apex
of tepal X 10; F, flower X 5; G, tepal with stamen X 10; H, ovary lateral view
X 10; I, ovary dosal view X 10. All from lknter 13,413.
Figure 77. A, lectotype of L. macowanii (Bak.) S. Venter (GRA); B, habitat
near Pafuri in the Kruger National Park. The vegetation consists of closed
deciduous Terminalia sericea - Hymenocardia ulmoides - Holarrhena pubescens
woodland; C, SEM micrograph of the adaxial lamina surface; D, SEM
micrograph of the abaxial lamina surface; E, SEM micrograph of stomata; F,
plants of L. macowanii showing the lax racemes. Bar = 50 mm. C - F from
lknter s.n..
Figure 78. Illustration of L. galpinii (Bak.) S. Venter. A, habit X 1; B, lamina
margin X 300; C, section through lamina X 5; D, apex of lamina X 20; E,
bracts X 10; F, flower X 10; G, apex of tepal X 20; H, tepal with stamen X 10;
I, ovary lateral view X 10; J, ovary distal view X 10. All from lknter 13,389.
Figure 79. A, lectotype of L. galpinii (Bak.) S. Venter (PRE); B, plants with
prominent lacunae on the leaves. Bar
=
10 mm; C, habitat at Kaapsche Hoop,
eastern Transvaal. The vegetation consists of closed short Panicum natalense -
Eragrostis racemosa - Scabiosa columbaria montane grassland; D, SEM
micrograph of the adaxial lamina surface. Bar = 50 J.'m; E, SEM micrograph of
the abaxial lamina surface. Bar = 50 J.'m; F, SEM micrograph of stomata. Bar
= 50 J.'m. Band D - F from lknter 13,389.
Figure 80. Illustration of L. papillata S. Venter. A, habit X 2; B, section
through leaf X 5; C, lamina margin X 300; D, bract X 10; E, papillae on
peduncle X 10; F, flower X 10; G, tepal with stamen X 10; H, apex of tepal X
20; I, ovary lateral view X 10; J, ovary distal view X 10. All from Venter
13,008.
LEDEBOURlA IN SOlITH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
360
Figure 81. A, holotype of L. papillata S. Venter (PRE); B, habitat near
Pietersburg. The vegetation consists of deciduous open low Aloe marlothii var.
marlothii - Ormocarpum trichocarpum - Themeda tnandra woodland; C, SEM
micrograph of the adaxial lamina surface. Bar = 100 Ilm; D, SEM micrograph
of the abaxial lamina surface. Bar = 100 Ilm; E, SEM micrograph of stomata.
Bar
= 43 Ilm.
Table 1.
A and C - E from Venter 13,186.
Summary of the taxonomic treatments of Ledebouna by the
various authors.
Thble 2.
Differences between the genera Ledebouria, Drimiopsis
and Scilla.
Table 3.
Leaf characters for the genus Ledebouria in South
Africa.
Table 4.
Phenology of the genus Ledebouria in South Africa.
Table 5.
Microsculpturing of the leaf surfaces.
Table 6.
Ovary and fruit characters.
Table 7.
Results of Jessop's chromosome studies (1972).
Thble 8.
Flowering period (+) for the genus Ledebouria in South
Africa.
Table 9.
The major soil types (Buckman & Brady 1969).
Thble 10. The sectional, subsectional and species groupings of
Ledebouria.
Table 11. Distribution and status of the genus Ledebouria in South Africa.
Table 12. Toxicity tests with L. ovatifolia.
LEDEBOURIA IN SOlJrH AFRICA
CAPTIONS TO ILLUSTRATIONS AND TABLES
Map 1.
Pictorialized map showing the variation in leaf shape, in
L. marginata (Bak.) Jessop. A, 1-Cnter 13,487. B, 1-Cnter
13,246. C, 1-Cnter 13,358 and 1-Cnter s.n.
Map 2.
Known distribution of the genus Ledebouria Roth.
Map 3.
Known distribution of the genus Ledebouria Roth in South
Africa.
Map 4.
Concentration of species of Ledebouria in geographical
degree squares.
Map 5.
Pictorialized map of L. apenijlora with the known
distribution of the tapered and truncate bulbscaled plants.
Map 6.
Known distribution of L. ensifolia (Eckl.) S. Venter
Map 7.
Known distribution of L. sandersonii (Bak.) S. Venter
Map 8.
Known distribution of L. dolomiticola S. Venter
Map 9.
Known distribution of L. atro-brunnea S. Venter
Map 10.
Known distribution of L. viscosa Jessop
Map 11.
Known distribution of L. leptophylla (Bale) S. Venter
Map 12.
Known distribution of L. lepida (N.E. Br.) S. Venter
Map 13.
Known distribution of L. minima (Bale) S. Venter
Map 14.
Known distribution of L. rupestris (Van der Merwe)
S. Venter
Map 15.
Known distribution of L. coriacea S. Venter
LEDEBOURlA IN SOtlTH AFRICA
361
CAPTIONS TO ILLUSTRATIONS AND TABLES
Map 16.
Known distribution of L. floribunda (Bak.) Jessop
Map 17.
Known distribution of L. hypoxidioides (Schonl.) Jessop
Map 18.
Known distribution of L. revoluta (Lj:) Jessop
Map 19.
Known distribution of L. zebrina (Bak.) S. Venter
Map 20.
Known distribution of L. luteola Jessop
Map 21.
Known distribution of L. ovatifolia (Bak.) Jessop
Map 22.
Known distribution of L. monophylla S. Venter
Map 23.
Known distribution of L. cooperi (Hookj:) Jessop
Map 24.
Known distribution of L. parvifolia S. Venter.
Map 25.
Known distribution of L. asperijolia (Van der Merwe)
S. Venter
Map 26.
Known distribution of L. glauca S. Venter
Map 27.
Known distribution of L. inquinata (C.A. Srn.) Jessop
Map 28.
Known distribution of L. marginata (Bak.) Jessop
Map 29.
Known distribution of L. concolor (Bak.) Jessop
Map 30.
Known distribution of L. crispa S. Venter
Map 31.
Known distribution of L. undulata (Jacq.) Jessop
Map 32.
Known distribution of L. ovalifolia (Schrad.) Jessop
LEDEBOURIA IN SOtJTH AFRICA
362
CAPTIONS TO ILLUSTRATIONS AND TABLES
Map 33.
Known distribution of L. petiolata (Van der Merwe)
S. Venter
Map 34.
Known distribution of L. socialis (Bak.) Jessop
Map 35.
Known distribution of L. macowannii (Bak.) S. Venter
Map 36.
Known distribution of L. galpinii (Bak.) S. Venter
Map 37.
Known distribution of L. papillata S. Venter
LEDEBOURIA IN SOlITH AFRICA
363
364
ACKNOWLEDGEMENTS
ACKNOWLEDGEMENTS
"Commit your work to the Lord, and your plans will be established" Proverbs
16:3.
I wish to thank the following persons and institu tions who offered
encouragement, assistance, advice and cooperation in the preparation of this
thesis:
Mr. T.1. Edwards, my supervisor, for his advice and constructive comments on
the thesis.
Prof. 1. van Staden, head of the Department of Botany, Natal University, for
encouragement and permission to use the Department in my studies.
The Foundation for Research Development and Mr. M. Dix for financing part
of the thesis.
The staff of the National Herbarium (PRE) of the National Botanical Institute
and the following herbaria B, BLFU, BM, BOL, CPF, GRA, GRAZ, J, K,
KEI, KNP, LYD, NBG, NH, NU, PEU, PRE, SAM, PRU, PUC, SRGH,
STE, UNIN, UWC, Z, for assistance and permission to examine material.
Mrs. Sandy Burrows, the artist, for preparing the ink drawings.
The staff of the Mary Gunn Library for their assistance in the search for
literature.
Messrs. R. Archer, 1. Burrows, I. & N. Croach, G.1. du Toit, T.J. Edwards,
D.S. Hardy, A. le Roux, T. Middleton, G.F. Smith, C. Stirton, E. van
Jaarsveld, 1.A. Venter and W. T. Vos for the collecting of plant material.
The directors and staff of the Electron Microscope Units, Natal University,
Pietermaritzburg and University of the North, Sovenga for constructive
assistance and the use of their facilities.
My wife Julye-Ann Venter for her patience and support in the veld and at home.
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