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A SYSTEMATIC REVISION OF
DRIMIOPSIS Lindl. & Paxt.
(HYACINTHACEAE)
Pearl Dijeng 'Matlhapi Lebatha
A SYSTEMATIC REVISION OF DRIMZOPSZS Lindl. &
Paxt. (HYACINTHACEAE)
Pearl Dijeng 'Matlhapi Lebatha
BSc. UNZA; MSc. SDSU; MEd. SDSU
Thesis submitted in fulfament of the requirements for the degree
Philosophiae Doctor
School of Environmental Sciences and Development
Botany Division
North-West University
Potchefstroom Campus
Promoter: Dr. M. H. Buys
Co-promoter: Prof. P. D. Theron
November 2004
The beginning of &nowledge
consists of learning to call things
their names.
Tuo-TeChing,
Lao-Tzu
DECLARATION
I, the undersigned, hereby declare that this thesis, submitted for the Doctor of
Philosophy Degree at North-West University, Potchefstroom Campus, is, except where
acknowledged, the result of my own investigations and has not been submitted in any
form to another university for a degree.
Signature. .............................
P. D. Lebatha
Date. ........................
ABSTRACT
Drimiopsis Lindl. & Paxt. is one of many genera within the Hyacinthaceae that have
not been comprehensively reviewed.
In this study, a comprehensive systematic
revision of Drimiopsis is accomplished through examination of the anatomy,
morphology, phytochemistry, DNA and cytology with the express aim of establishing a
phylogenetic classification. Results of the research include amongst others, character
and taxa diversity, identification keys as well as the delimitation of the genus fiom the
other closely related genera of the subtribe Ledebouriinae U. & D Miiller-Doblies.
The phylogenetic species concept, a character-based approach inferring phylogeny via
hierarchical distribution of characters, is preferred in elucidating the phylogenetic
history of Drimiopsis and reviewing its taxonomy. To this end, both a phenetic and a
cladistic analysis were done.
This study also adopts a bifurcatus investigative approach encompassing generic
concepts in the Ledebouriinae to enhance phylogenetic inference with respect to
Drimiopsis. Ambiguities exist in the concept of Drimiopsis that cannot be fully
understood in isolation. An analysis of the most immediate sister taxa namely, Resnova
v. d. Merwe and Ledebouria Roth gives a better resolution of interspecific and
intergeneric variation in characters and states as well as relationships.
The leaf epidermis displays taxonomically significant cell arrangement, morphology
and
stomata
characters. Anatomical characters confirm the morphological
differentiation of epidermal cells. Phytochemical analysis of D. burkei Bak. extracts
revealed six novel structured homoisoflavonoids and a scillascillin. The genus
Drimiopsis has basic chromosome numbers x = 10 and x = 11. The former is
predominant in southern Afiican taxa, the latter in tropical African taxa. There are
distinctive intergeneric variations in the bulb, leaf, inflorescence, flower and pollen
char-i~:-ters-within
theZedebourijgae.
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The phenetic analysis of leaf, flower, pollen and phytochemical characters all
demarcate Resnova, Ledebouria and Drimiopsis, clustering Resnova with Ledebouria
throughout.
The consensus cladogram of the cladistic analysis emphasises the monophyly of the
three genera and supports the subtribe grouping of the Ledebouriinae. The total
evidence analysis of morphological and available DNA data produced a well-supported
and resolved tree similar in topology to the tree based on morphological characters
alone. The nodes that were not resolved in DNA data analysis were resolved with the
addition of morphological data to the matrix.
The genus Drimiopsis, now consisting of 18 species, is endemic to Africa with disjunct
distributions in southern and tropical Africa. Thirteen of these species are endemic to
southern Afiica and five to tropical Africa. This study recognizes the current nine taxa
endemic to southern Afiica. Of these two are elevated to species rank and four new
species are described. The four taxa currently endemic to tropical Afiica are recognized
of which two are elevated to the rank of species. Two species are resurrected.
ACKNOWLEDGEMENTS
This study was undertaken under the sponsorship of the Botswana College of
Agriculture, an Associate Institution of the University of Botswana.
I thank Dr. Matt H. Buys of the A. P. Goossens Herbarium, North-West University,
Potchefstroom Campus, for strong leadership, positive guidance and the lesson: "one
cannot dream oneself into a good scientist; one must hammer and forge oneself into
one".
Also from Potchefstroom Campus, with prolific comments and assistance: Prof P. D.
Theron (co-promoter), Dr. Sandra du Plessis and Dr. Carlos Bezuidenhout (DNA
analysis), Dr. Lourens Tiedt (SEM Laboratory), Santie Pieterse (Graphics) and Alta
Lindeque (botanical drawings). From the University of Botswana: Prof. B.M. Abegaz,
Department of Chemistry
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Natural Products Research, and his research team for
assistance with the chemotaxonomy and Dr. Anine Jordaan of Microscopy, Physics
Department, is thanked for assistance with plant anatomy. Prof. J. J. Spies, Department
of Plant Sciences: Genetics, University of the Free State for guidance with the
cytogenetics. Dr. Nigel Barker, Rhodes University and Di James of the DNA
Sequencing Laboratory, University of Cape Town, are acknowledged for DNA
sequencing.
For fiuitful discussions, I acknowledge Dr. 0. M. Mine of the Botswana Harvard AIDS
Institute Partnership for HIV Research and Education, Gaborone; Dr. John Manning of
the South African National Biodiversity Institute (SANBI), Cape Town; Dr. J. P. Jessop
of the State Herbarium of South Australia (AD), Australia; Prof. Dr. W. Wetschnig of
the Institute of Botany, Graz, Austria.
A very special thanks is extended to Prof. Brita Stedje of the University of Oslo for
pi6vidiiig six=-bulbs, photographs discussions and encouragement, Prof. DryDietrich
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Muller-Doblies and Dr. Ute Muller-Doblies of Technische Universitat Berlin, for their
generosity and support. Assistance is acknowledged from staff of the following
herbaria: PRE, Pretoria; Kew, particularly Dr. Paul Wilkin, Dr. Anna Haig and Dr.
Monika Shaffer-Fehre; Botanischer Garten und Botanisches Museum Berlin-Dahlem,
Berlin, especially Dr. Christoph Oberprieler.
The following people are acknowledged for their support and fiee access to their live
collections: Andrew Hankey of the Walter Sisulu National Botanical Garden, Anne
Hutchings of the University of Zululand, Wally Menne of BOTSOC, Durban and Dr.
Trevor Edwards of Natal University, Pietermaritzburg. Profuse gratitude is extended to
all supportive relatives and fiends.
This work is dedicated to my husband, Khotso, for his fortitude and indelible
patronage; to Kiki, Qeni and Can, my raison d'ghe, for sacrifices endured and my
parents, Cuthbert and Margaret Motsepe, for absolute championship.
The following institutions are thanked for long-term loan of herbarium material and for
some, the supply of plant images: B, BLFU, BOL, BR, BTU, GAB, HNN, J, K, NBG,
NH, 0 , P, PRE, PUC, UNIN, and Z.
ABBREVIATIONS AND ACRONYMS
AD
State Herbarium of South Australia, Adelaide, Australia.
B
Botanischer Garten und Botanisches Museum Berlin-Dahlem,
Germany.
BLFU
Geo-Potts Herbarium, Bloemfontein, South Africa.
BOTSOC
Botanical Society of South Africa.
BOL
Bolus Herbarium, Cape Town, South Afiica.
BR
Jardin Botanique National de Belgique, Meise, Belgium.
BTU
Herbarium Technische Universitiit Berlin, Germany.
DNA
Deoxyribonucleic acid.
GAB
National Herbarium, Botswana.
HN-N
Horniman Museum of Natural History, London, England.
ICBN
International Code of Botanical Nomenclature (Tokyo Code)
IPNI
International Plant Name Index.
J
Charles E. Moss Herbarium, Johannesburg, South Afiica.
K
Royal Botanic Gardens Herbarium, Kew, England.
NBG
Compton Herbarium, Cape Town, South Afiica.
NH
Natal Herbarium, Durban, South Afiica.
0
Botanical Museum, Oslo, Norway.
P
Muskum National d'Histoire Naturelle Paris, France.
PRE
National Herbarium, Pretoria, South Afiica.
PUC
A.P. Goossens Herbarium, Potchefstroom, South Afiica.
SANBI
South African National Biodiversity Institute.
SEM
Scanning Electron Microscopy.
UNIN
University of the North Herbarium, Polokwane, South Afiica.
z
Universitat Zurich, Zurich, Switzerland.
ACCEPTED DRIMZOPSZS NAMES IN THIS THESIS
Drimiopsis atropurpurea N.E. Br.
Drimiops is barteri Bak.
Drimiopsis botyoides Bak.
Drimiopsis burkei Bak.
Drimiopsis carrii Lebatha
Drimiopsis comptonii U. & D. Miiller-Doblies
Drimiopsis davidsoniae U. & D. Miiller-Doblies
Drimiopsis $scheri (Engl.) Stedje
Drimiopsis kikiae Lebatha
Drimiopsis liniopapilla Lebatha
Drimiopsis maculata Lindl. & Paxt.
Drimiopsis perfoliata Bak.
Drimiopsis pusilla U. & D. Muller-Doblies
Drimiopsis queae Lebatha
Drimiopsis reilleyana U. & D. Muller-Doblies
Drimiopsis rosea A. Chev.
Drimiopsis stolonissima (U. & D. Muller-Doblies) Lebatha
Drimiopsis woodii Bak.
CONTENTS
Page
Frontispiece
Declaration
rv
Abstract
v
Acknowledgements
VII
Abbreviations and acronyms
IX
Accepted names in this thesis
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1. INTRODUCTION
1.1 Background
1.2 History
1.3 Habit and Habitat
1.4 Uses
1.5 Abundance
1.6 Objectives
1.7 Modus operandi
1.8 Conclusion
2. SYSTEMATIC CONSIDERATIONS
2.1 Introduction
2.2 Species concepts
2.3 Nomenclature
2.4 Conclusion
3. THE LEAF
3.1 Introduction
3.2 Objectives
3.3 Materials and methods
3.4 Results and discussion
3.4.1 Macromorphology
3A.2 Micromorphology
3.4.3 Anatomy
3.5 Phenetic analysis
3.6 Conclusion
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3.7 Key to species of Drimiopsis based on leaf characters
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4. THE FLOWER
4.1 Introduction
4.2 Objectives
4.3 Materials and methods
4.4 Results and discussion
4.4.1 Inflorescence
4.4.2 Perianth
4.4.3 Androecium
4.4.4 Gynoecium
4.5 Phenetic analysis
4.6 Conclusion
4.7 Key to the species of Drimiopsis based on floral characters
5. PALYNOLOGY
5.1 Introduction
5.2 Objectives
5.3 Materials and methods
5.4 Results and discussion
5.5 Phenetic analysis
5.6 Conclusion
6. CHEMOTAXONOMY
6.1 Introduction
6.2 Objectives
6.3 Materials and methods
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6.3.2 Plant material
6.3.3 Phytochemical screening protocols
6.3.4 Chromatography analysis
6.3.5 Phenetic analysis
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6.4 Results and discussion
6.4.1 Phytochemical screening tests
6.4.2 TLC and HPLC analysis
6.5 Phenetic analysis
6.6 Conclusion
7. KARYOLOGY
7.1 Introduction
7.2 Objectives
7.3 Materials and methods
7.4 Results and discussion
7.5 Conclusion
8. DNA ANALYSIS
8.1 Introduction
8.2 Objectives
8.3 Materials and methods
8.4 Results and discussion
8.4.1 Protocol appraisal
8.4.2 PCR results summary
8.4.3 Sequencing results
8.5 Conclusion
9. PHENETIC ANALYSIS
9.1 Introduction
9.2 Objectives
9.3 Materials and methods
9.4 Results and discussion
9.4.1 Phenetic analysis
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9.4.2 Generic circumscription
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9.5 Conclusion
10. CLADISTIC ANALYSIS
10.1 Introduction
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10.2 Objectives
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10.3 Materials and methods
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10.4 Results and discussion
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10.4.1 Cladistic analysis of morphological data
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10.4.2 Total evidence analysis
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10.4.3 Systematic implications of the character analysis
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10.5 Conclusion
11. PHYTOGEOGRAPHY
11.1 Introduction
11.2 Objectives
11.3 Materials and methods
11.4 Results and discussion
11.4.1 Distribution patterns
11A.2 History, evolution and relationships
11.5 Conclusion
12. FORMAL TAXONOMY
12.1 Generic description
12.2 Key to species of Drimiopsis
12.3 Species descriptions
12.3.1 Drimiopsis stolonissima
12.3.2 Drimiopsis carrii
12.3.3 Drimiopsis perfoliata
12.3.4 Drimiopsis botryoides
12.3.5 Drimiopsis rnaculata
12.3.6 Drimiopsis davidroniae
12.3.7 Drimiopsis comptonii
12.3.8 Drimiopsisfischeri
12.3.9 Drimiopsis woodii
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12.3.10 Drimiopsis reilleyana
12.3.1 1 Drimiopsis burkei
12.3.12 Drimiopsis barteri
12.3.13 Drimiopsis rosea
12.3.14 Drimiopsis pusilla
12.3.15 Drimiopsis atropurpurea
12.3.16 Drimiopsis kikiae
12.3.17 Drimiopsis liniopapilla
12.3.18 Drimiopsis queue
12.4 Excluded names
1 2.4.1 Published names
12.4.2 Nomino dubia
3. GENERAL CONCLUSIONS
13.1 Systematic revision
13.2 Future prospects for research
4.
REFERENCES
*PENDICES
Appendix 1: Rf values tables and chromatograms
Appendix 2: DNA protocols
INDEX TO DRIMIOPSIS NAMES
1. INTRODUCTION
1.1BACKGROUND
The family Hyacinthaceae Agardth (Liliaceae Juss. sensu lato) [synonyms: Scillaceae
Vest (18 18, Anleit. Stud. Bot. : 267, 284), Scilloideae Kosteletzky. (1831, Allg. Med.Pharm. F1. 1: 168), Eucomidaceae Salisb. (1866, Gen. PI.: 16) and Lachenaliaceae
Salisb. (1866, Gen. PI.: 20) Reveal (1999)l is part of the superorder Liliiflorae, order
Asparagales (Dahlgren, 1980). The majority of plants in this family consist of
rhizomatous perennial herbs, with fleshy roots or bulbs as well as a basal aggregation of
leaves. The inflorescence is commonly a raceme; the flowers regular, perfect, trimerous
and tepalous.
The Hyacinthaceae consists of ca. 42 genera with the highest diversity occurring in
southern Afiica, followed by the Mediterranean region and south-westem Asia (Pfosser
et al., 2003). The Hyacinthaceae currently comprises five morphologically similar
subfamilies delimited mostly on phytochernical data (Speta, 1W8a, 1998b; Pfosser et
al., 2003). The Chlorogaloideae Speta is confmed to North America and the
Oziroeoideae Speta to South America. Of relevance to Afiica, are the Urgineoideae
Speta, with bufadeinolids, Ornithogaloideae Speta with cardenolides andlor protein
crystals in their nucleus, and the homoisoflavanol rich Hyacinthoideae Link (Link
1829).
The Massonieae Bak. (1870b, J. Linn. Soc., Bot. 11: '359, one of two (Speta, 1998b)
or three (Manning et al., 2004) tribes of the Hyacinthoideae, comprises around 30-45
genera found in Afiica south of the Sahara and into India (Speta, 1998b). The tribe is
diagnosed through the presence of bulbs, spotted leaves, and seeds without a superficial
cellular pattern as well as connate stamens, introrse anthers and a threecelled ovary
producing a loculicidal capsule. Muller-Doblies & Muller-Doblies (1997) place species
of the Massonieae from southern and eastern Afiica having a "tropical growth-form
with iterative innovation" into subtribe Ledebouriinae U. & D Muller-Doblies, thus
separating Drimiopsis Lindl. & Paxt., Resnova v. d. Menve and Ledebouria Roth from
the other Massonieae endemic to South Afiica with annual innovation.
Miiller-Doblies & Miiller-Doblies (1997) characterize the Ledebouriinae on the
occurrence of two basal ovules in each locule-according
are side by side in Ledebouria-leaves
to Speta (1998b) the ovules
without sheathing cataphylls and several
( inflorescences to a tuft of leaves. Of the three genera in the Ledebouriinae, it is only
Ledebouria that occurs in the Mediterranean region, India, as well as Afiica and
Madagascar. Drimiopsis is endemic to Afiica south of the Sahara while Resnova is
restricted to southern Africa.
Manning et al. (2004) disagree with the Muller-Doblies & Muller-Doblies (1997)
classification of the tribe Massonieae. Their phylogenetic analysis of the Hyacinthaceae
resolves the Massonieae into a polychotomy of five clades, one of which is the genus
Ledebouria. This Ledebouria clade houses Drimiopsis and Resnova. It is characterised
by "lack of bracteoles and by its globose or top-shaped ovary containing two ovules per
locule .... most species have spotted leaves and often produce more than a single
inflorescence per plant in one growing season, and the bulb scales are ofien rather
loosely packed and in many species produce fme threads" (Manning et al., 2004).
Manning et al. (2004) also consider differences between Ledebouria and Resnova
qualitative and dismiss tepal differences in Drimiopsis as a pollination adaptation.
Separate revisions have been done on southern Afiican and tropical Afiican taxa of
Drimiopsis (Baker, 1896 & 1898; Jessop, 1972; Stedje, 1994; Muller-Doblies & Miiller-
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Doblies, 1997) producing varying taxonomic opinions. No monograph of the genus
exists to date. Plant descriptions in these revisions are minimal-to
a lesser extent so in
Baker (1896 & 1898). Character and character state allocation is either inconsequent or
entirely lacking, making taxa determination difficult. I assume this may be because of
journal volume restrictions, as there appears to be a decrease in the volume of
taxonomic descriptive works in journals over the years.
1.2 HISTORY
The history of Drimiopsis is intimately interwoven with that of Scilla L. (Limaeus
175T, -1 758); one of the moN specks-rich h u h t l s genera, as thesynonymy list-to most_
names currently recognised will attest. Many species initially described as belonging to
Scilla have found their way to Drimiopsis, Resnova or Ledebouria (see Taxonomic
Treatment, Chapter 12). Resnova, since its inception in 1946, has produced various
subsequent combinations in Drimiopsis, compounding the taxonomic complexities.
I
Scilla sensu lato is, on account of DNA data, not monophyletic (Stedje, 1998; Pfosser
& Speta, 1999; Wetschnig et al., 2002). The distinct sub-Saharan Scilla clade is now
the genus Merwilla Speta (Pfosser & Speta, 1999; Wetschnig et al., 2002bthere is
consensus that Drimiopsis, Ledebouria and Resnova form their own clade (Wetschnig
& Pfosser, 2003; Pfosser et al., 2003).
Lindley and Paxton (1851-1852) first described Drimiopsis as a monotypic genus
based on Drimiopsis maculata Lindl. & Paxt., a species described from material
received by the Horticultural Society (London) from the Cape, South Africa.
Drimiopsis was characterised by
". ..a white fleshy bulb.. . few broad,
fleshy, oblong
leaves, 6-8 in. long, rolled up at their base... form a kind of channelled petiole;
...abundantly clouded with dark green oblong stains upon a paler ground. The scape
tapers, and is about as long as are the leaves, terminated by a close raceme of half
closed campanulate flowers; the lower of which are pendulous, the upper white and
erect, both sepals and petals are herbaceous, ovate, cucullate, concave and united at the
base; the petals are rather shorter and broader than the sepals. The stamens are six,
equal; their filaments inserted by a broad base upon the sepals and petals; the anthers
are ovate and turned inwards. The ovary is ovate, roundish, undivided, gradually
tapered into a style with a simple minute stigma; in each of its three cells erect a pair of
anatropal collateral ovules" (Lindley & Paxton 1851-52).
Although Lindley 's main interest lay in orchids (Lindley, 1834-3 7; 1838; l852), he coauthored D. maculata with Paxton, who also was a keen botanist and publisher (Paxton,
1834-49; Lindley & Paxton, 1882-1 884). Of note is the nomenclatural inaccuracies
demonstrated in the authorship of Drimiopsis. Baker (1870a, 1896,1898), Van der
Menve (1946a, 1946b), Jessop (1972), Dyer (1976), Arnold & De Wet (1993) and
Miiller-Doblies & Muller-Doblies (1997) recognize Lindley as the sole author.
However, the original description of the genus specifically states "We therefore
propose it [Drimiopsis]as a new genus" (Lindley & Paxt, 1851-52: 73). Stedje (1994),
Speh-(4 998a-& -1998h), K a t i u (2000_), Lebathaet al. (2003), Manning & Goldblatt
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(2003) and Lebatha & Buys (a & b, in press) provide the correct author citation.
However Manning et al. (2004), differing from Manning & Goldblatt (2003), refer to
the new combination as "Ledebouriapetiolata J.C. Manning & Goldblatt, nom. nov.,
pro Drimiopsis maculata Lindl." (Manning et al.,2004: 56 1).
Baker (1 870a, 1874b) initially described tropical African species, namely D. botryoides
Bak. and D. barteri Bak., separating them on the basis of leaf shape. He also described
D. kirkii Bak. (Baker, 1874a) and later D. perfoliata Bak. (Baker, 1878), both from Dr.
Kirk's Zanzibar collection.
During that time, Engler (1895), as the Director of the Berlin School of Botany,
contributed by describing a broad-leaved tropical African species, D. holstii Engl. He
also described Scilla fischeri Engl. that was later transferred to Drimiopsis (Stedje &
Thulin, 1995).
Later, Baker (1896), in what may be considered as the first account of southern African
Drimiopsis, discussed seven species. Six of these species he described as new,
delimiting them on the basis of leaf and perianth dimensions. Baker appears to have
focused on differences (i.e. the splitter approach) between taxa e.g. although he
grouped D. maculata with D. minor Bak. on the basis of distinctly petioled leaves with
a cordate blade, he distinguished them on the basis of leaf and flower size. Leaf length
has been found unreliable, as it is generally a function of plant age. The differences in
perianth size as stipulated by Baker are infinitesimal, 0.4 and 0.3 cm long respectively.
D. woodii Bak. and D. maxima Bak., also possessing distinct petioles, were separated
fiom the former two through the occurrence of a leaf blade narrowed at the base. D.
woodii and D. maxima were in turn delimited on the basis of a raceme being 6 . 6 1 1 cm
or 1.1-2.2 cm long respectively. Baker diagnosed the sessile leaved species using leaf
erect, and small flowered in D. burkei Bak. and
orientation and flower size-leaves
leaves erect and large flowered in D. saundersiae Bak. D. humifusa (Bak.) Bak. was
separated on the basis of spreading leaves, with no reference to flower size.
Baker (1898) described the tropical African D. stuhlmannii Bak. and transferred S.
volkensii Engl. to Drimiopsis. Baker (1904) later reconsidered the status of S. humifusa
Bak. (1881) and transferred it to Drimiopsis. This has recently been transferred to
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Carl Darnmer, a 20th century German botanist at the Botanical Museum in BerlinDahlem, described D. erlangeri Damm. (Damrner, 1905) fiom Ethiopia and D. bussei
Damm. fiom Tanzania (Darnmer 1907).
Augustine Chevalier, a French botanist interested in botanical explorations (Chevalier,
1920, 1934), described three taxa of Drimiopsis from Congo Brazzaville namely D.
aroidastrum var. aroidastrum, D. aroidastrum var kabarum with "lanceolate-linear
leaves" and D. rosea. (Chevalier, 1908). D. aroidastrum var. kabarum was cultivated
for medicinal use in the then Congo Brazzaville.
Emile Auguste Joseph De Wilderman (19 11a, 1911b) described D. sereti fiom Zaire.
He mentioned that it is similar to D. barteri, but: "possibly the variety parvifolia
Perkins"-an
invalid name.
Kurt Krause was a co-worker of Adolf Engler. From 1905 Krause was responsible for
compiling small families in Das Pflanzenreich, describing 124 Aracaceae taxa and coauthoring another 75 with Engler. Krause (1914) described D. engleri Krause fiom
Namibia.
Moritz Dinter's main interest lay with the Namib Desert flora. He described D.
papillosa fiom Namaqualand (Dinter, 1921). This taxon has subsequently been
transferred to Ledebouria and synonymised under L. scabrida Jessop (S Venter, 1993).
Dr. Nicholas Edward Brown worked at Kew in the production of Araceae treatments
for floristic works. In 1921 Brown described D. atropurpurea from material collected
by F.A. Rogers, stating that it was: "...similar to the type of the genus but pubescent
with purple flowers".
Phillips (1926) reports Drimiopsis as comprising 19 species, 7 from South Africa and
12 from tropical Africa. He does not however, list the species but his count fits well
with the species (including the variations) already described thus far.
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Van der Menve (1946a) described D. crenata and D. purpurea, empliasisilng-size
differences as diagnostic for D. crenata namely, "...bigger than D. burkei but smaller
than D. saundersiae". D. purpurea was later transferred to D. atropurpurea (Jessop,
1972).
Jessop (1970), while revising Ledebouria, transferred D. engleri to Ledebouria, and
synonyrnised it under L. undulata (Jacq.) Jessop together with a number of Scilla
species. Later Venter (1993) viewed D. engleri as a synonym of his new combination L.
rautanenii (Schinz) S. Venter, a transferral fiom Scilla (Schinz, 1901-1 908).
Jessop (1972) undertook the second major revision of southern African Drimiopsis
since the inception of the genus. Of note here is that although the title to his paper
evokes the impression that his study was confined to South Africa, he clearly also
investigated specimens from Swaziland and Botswana. As he did not recognize
Resnova as a separate genus he transferred Resnova lachenaliodes (Bak.) v.d. Merwe
(based on S. lachenaliodes Bak.) to Drimiopsis as a new combination, and synonymised
the remaining Resnova species namely R. schlechteri (Bak.) v.d. Merwe (based on S.
schlechteri Bak.), R. transvaalensis v.d. Menve, R. pilosa v.d. Menve, R. minor v.d.
Merwe and R. maxima v.d. Menve under Drimiopsis maxima Bak. Jessop's (1972)
decision to transfer Resnova to Drimiopsis was probably encouraged by his misguided
opinion that Van der Merwe (1946b) did not specify type specimens. However, Van der
Menve (1946b: 46) did affirm them. Jessop might have reached a different conclusion
had he studied these. He had a strong conviction that the Scilleae had very few
qualitative characters that could be used in taxonomic work, and that floral structure
displayed very little intergeneric variation.
Jessop (1972), upon studying vascular bundles and cytology of Drimiopsis sensu lato,
concluded that Drimiopsis was more closely related to Ledebouria than to Scilla. This
supports the subsequent grouping of Drimiopsis and Ledebouria (and Resnova) in the
Ledebouriinae (Muller-Doblies & Muller-Doblies, 1997). Jessop (1972) recognized two
groups within Drimiopsis: D. atropurpurea, D. burkei, D. maculata possessing small
flowers (not more than 5 rnm long) and with tepal apices cucullate; and the larger
flowered D. lachenalioides and D. maxima possessing spreading tepals, D. maxima
with slightly cucullate tepal apices. Dyer (1976), Arnold & De Wet (1993) and
Williams (2000) subsequently accepted these five species. The two larger flowered
Drimiopsis species were later transferred to the reinstated Resnova (Miiller-Doblies &
Miiller-Doblies, 1997).
Stedje (1994) and Stedje & Thulin (1995), confming their work to tropical east Afiica,
recognise other than D. maculata, an additional four tropical AfYcan taxa. By
disregarding leaf dimensions and concentrating more on leaf shape and flower
characters, they place all taxa with narrowly lanceolate leaves and small sessile flowers
with perianth segments connivent in D. barteri. Taxa with lanceolate to cordate leaves
as well as sessile or pedicellate flowers with inner perianth segments connivent and the
outer spreading belong to D. botryoides Bak. subsp. botryoides (now also including the
synonymised D. kirkii, D. holstii, D. stuhlmannii, D. erlangeri and D. bussei). The third
group, with broadly lanceolate leaves appressed to the ground and flowers sessile to
very shortly pedicellate, is described as D. botryoides Bak. subsp. prostrata Stedje. S.
fscheri Engl. (Engler, 1895), transferred to D. fischeri (Engl.) Stedje, forms a unique
group in that the long perianth with stamens inserted just below the apex makes it
different from any other Drimiopsis species described so far. In my opinion, Stedje
(1994) and Stedje & Thulin (1995) can also be labelled lumpers as they emphasise
similarities.
Miiller-Doblies & Muller-Doblies (1997), in their revision of southern African
Drimiopsis recognise three of Jessop's species namely D. atropurpurea, D. burkei and
D. maculata. As mentioned, they transfer D. lachenaliodes and D. maxima to the
reinstated Resnova. In the latter case it is noteworthy that D. maxima reported to
possess 9 mm long tepals (Jessop, 1972), is synonymised under R. humifua, reportedly
possessing tepals 5-6 rnrn long (Muller-Doblies & Muller-Doblies (1997). In addition
they resuscitate D. woodii, which Jessop (1972) earlier placed under D. burkei on
account of small flowers after studying the syntype at Kew. They also synonymise
Drimia petiolata G. Koch & C.P. Bouchd (Koch & Bouchi, 1861) under D. maculata.
This is not surprising as Drimiopsis means "bearing a resemblance to Drimia".
Although the title of the Muller-Doblies & Muller-Doblies (1997) paper implies a
southern Africa study, a new tropical Afiican combination D. fscheri (Engl.) U . & D.
Muller-Doblies is included. One subspecies was described, namely D. burkei Bak.
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subsp. stolonissima, differing e
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m-Dmburkei Sak.subs~;
burkei-in-possessings_t~bns._
Four new species were also described: Drimiopsis reilleyana with "leaves, scape,
pedicels hairy". These aforementioned taxa together with D. maculata and D. woodii,
are grouped on the basis of predominantly green tepals. The remaining three new
species described (all possessing pink to lilac flowers) were D. davidsoniae with "scape
and leaves glabrous, leaf margin crenulate"; D. pusiNa with "scape and leaves hairy,
leaf margin entire" and D. comptonii with "scape glabrous, leaves hairy on upper side".
The new taxa together with D. atropurpurea, bring the total for southern African
Drimiopsis taxa to nine as reported by Manning & Goldblatt (2003).
Recent notes for the Flora Zambesiaca area allude to only three Drimiopsis taxa in this
region (Kativu, 2000), namely D. maculata, D. burkei and D. barteri. The genus is
described as possessing isomorphic tepals. Drimiopsis maculata and D. burkei are
reported as having 2-4 and 1-2 leaves, respectively. D. burkei is reported to have a
greater stature when growing in the shade.
Williams (2000) synonymised Resnova under Drimiopsis which in her opinion consists
of ca. 15 African species, five of them southern Africa-no
mention of species names.
She did not elaborate on the remaining ten, notwithstanding citing Stedje (1994 &
1996).
The treatment of Drimiopsis by all the authors mentioned above is indicative of the
morphological similarities possessed by them. Botanists not specializing in Drimiopsis,
hence without an overall insight into the variability of character states, historically
described most species. The varying degrees of polymorphism within species either
promote lumping--creating more polymorphic taxa--or splitting morphotypes into
separate taxa. Notable is that morphotypes, though evident fiom descriptions of taxa
(eg. D. burkei subsp. burkei), have never been considered for reranking into species.
Drimiopsis maculata also displays variation with lamina margins ranging fiom flat to
undulate, base fiom simple cordate to almost sagittate.
The status of Ledebouria has never been seriously questioned since Jessop's (1970)
revision. Roth (1821) reranked section Ledebouria Baker, one of the sub-sections of
SciNa (Baker 1870a), into the genus Ledebouria. It has previously been classified
render Hyacinthus LL&-eus,
-1782)- and Lachenalia Jacq. (Jacquin, 1 794; Andrews,
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1803). Ker-Gawler (181 1) transferred taxa of Ledebouria to Drimia. Later Trattinick
(1814) grouped Ledebouria with Lachenalia and von Schrank (1820) moved it back to
Scilla. Baker (1870a), in his SciNa monograph, also included Ledebouria under SciNa.
Jessop (1970) reinstated Ledebouria as a separate genus proposing the stipitate ovary as
diagnostic. Venter (1993), in his revision, recognised thirty-three Ledebouria species in
South Africa. There are to date sixty-eight (68) names pertaining to Ledebouria in
literature (IPNI, 2004). DNA studies confirm that Ledebouria is a sister group to
Drimiopsis (Stedje, 1996; Pfosser & Speta, 1999; Wetschnig et al., 2002; Wetschnig &
Pfosser, 2003) and has close affinity with Resnova (Wetschnig & Pfosser, 2003;
Pfosser et al., 2003).
As mentioned above, the status of Resnova was reconsidered by Muller-Doblies &
Muller-Doblies (1997). Van der Merwe (1946b) described Resnova based on S.
schlechteri Bak. (1904). In addition he transferred S. lachenalioides Bak to Resnova as
well as describing an additional four species, namely. R. transvaalensis, R. pilosa, R.
minor and R. maxima.
Philips (195 1) returned Resnova to Scilla. This proposal went largely unheeded.
Jessop's (1972) synonymisation of Resnova under Drimiopsis became the status quo
practiced by herbaria (Dyer, 1976; Meyer and Williams, 1997; Arnold & De Wet,
1993; Kativu, 2000) until Miiller-Doblies & Miiller-Doblies (1997) argued again for the
generic status of Resnova on the basis of flower characters, i e . spreading perianth and
ovoid ovary. Recent molecular data seems to support this scenario (Wetschnig &
Pfosser, 2003; Pfosser et al,, 2003). The National Botanical Institute (Manning &
Goldblatt, 2003) and the International Plant Name Index have followed suite. The
number of taxa has grown with the transference of two SciNa species to Resnova
namely R. humifura (Bak.) U . & D. Muller-Doblies (Miiller-Doblies & Muller-Doblies,
1997) and R. nossibeensis (H. Perrier) Speta (Speta, 1998a).
Manning et al. (2004), diverting from the norm, sunk both Resnova and Drimiopsis into
Ledebouria, hence the 68 species of Ledebouria recognised by them. The new
combinations thereof were made "only for type species of genera that are regarded as
synonymous with others, and for species that are treated in more recent accounts for
sout-h-eK Afj-1ca3'r
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Speta (1998a) described Avonsera Speta typified by A. convallariodes (Perrier) Speta
fiom Madagascar-originally
an Ornithogalum species. Included in this new genus is
A. lachenalioides (Bak.) Speta, a transferral of R. lachenalioides and as mentioned
confmed to southern Africa. The status of Avonsera has been seriously questioned
(Stedje, 2001) and is not recognised by Manning and Goldblatt (2003). Speta (1998a)
lists eight southern and three tropical Afiican Drimiopsis species, excluding subspecies.
1.3 HABIT AND HABITAT
Drimiopsis is a geophyte growing in diverse soil types. The taxa are confmed to Afiica
south of the Sahara, being found in Botswana, South Afiica, Somalia, Swaziland,
Mozambique, Zambia, Zimbabwe as well as Tanzania, Kenya and Ethiopia. They
spread across central Afiica, Democratic Republic of Congo, Chad to Cameroon, Niger
and Nigeria. Drimiopsis favours moist shady areas, under trees or tall grass, where
there is an accumulation of plant litter. Regular habitats also appear to be hill slopes
and the base of rocks or boulders.
The adventitious root system arises from the bulb. The roots are white, fleshy, and
shallow, not more than 10 cm deep (Figure 1.1 B - D, F, 1 and H; Figure 1.2 A - G).
The underground bulbs are generally small (0.2-2 cm in diameter), naked, white to
green with fleshy loose scales truncate at their apices. Exceptions to this description are
found in D. maculata with bulbs of ca. 6 cm in diameter (Figure 1.1 C, Figure 1.2 B).
Some D. burkei subsp. burkei plants growing on sandy soil in high temperature regions
(maximums in the 40" C range) of Botswana have bulbs with a brown, scaly
membranous outer covering (Figure 1.2 D). The stoloniferous habit of D. burkei subsp.
stolonissima is found in only a few other species like D. comptonii (Figure 1.2 F - G).
Bulbs produce bulblets at their bases and are rarely solitary (Figure 1.1 C, F & I; Figure
1.2 B & E). Some bulbs have very short basal stems, an extension of the stem plane
beneath the bulb, that is never more than 0.6 cm long (Figure 1.1 B, C, D, I & H; Figure
1.2 A, C & D).
The vegetative state is the dominant stage of the life cycle. The plants range from dwarf
(less than lOcm high) as in D. queae Lebatha to robust as in D. botryoides Bak. subsp.
Botwdes
(morethan 4Q cmhigh)<The plants have colour camouflage in the form of
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cryptically coloured, dotted or streaked leaves that make them inconspicuous. All
Drimiopsis taxa either possess leaves that appear before the flowers (protoantherous) or
leaves and flowers appearing at the same time (synantherous) except for D. rosea A.
Chev. that is hysteranthous (flowers appearing before the leaves). The racemose
inflorescence makes a brief seasonal appearance, commonly from August to October,
with inconspicuous flowers that either h i t very quickly or not at all.
Resnova taxa are gregarious geophytes with white and round to ovate bulbs with
loosely packed scales (Figure 1.1 A & G, Figure 1.2 I). Resnova possesses a limited,
disjunct distribution across the northern and eastern parts of South Afiica, also
preferring shaded areas.
The Ledebouria hypogenous bulbs are gregarious or solitary and are coloured purple to
brown, with the dead bulb scales persistent (Figure 1.1 E, Figure 1.2 H). The majority
of Ledebouria have a unique character of bulb scales producing threads when tom.
According to Venter (1993), Ledebouria species occur in winter and summer rainfall
areas, 76% of them in full sun with only L. concolor (Bak.) Jessop, L. floribunda (Bak.)
Jessop, L. rupeshis (v.d. Menve) S. Venter and L. socialis (Bak.) Jessop growing in the
shade. They grow in seepages, water, mountain terrains, grasslands and wooded areas.
1.4 USES
Herd boys in some southern parts of Botswana traditionally used the bulb of D. burkei
subsp. burkei -Molora-wa-basimanyana,
Thejane (Turton & Ablomberg-Ermatinger,
1988; Cole, 1995) as soap. This tradition lost favour with the advent of commercial
soaps. The vegetative parts are also used in traditional medicine to treat bereaved
persons after the death of a spouse (informants). Washing with the vegetative parts of
the plant is believed to cleanse and protect the surviving spouse.
The bulb of D. maculata-Sekaname-se-sesweu-is
used in South Africa as a
paediatric enema (Hutchings, 1989a & 1989b). The crushed bulbs are put in water and
used
externally
for
cleansing
to
remove
bad
luck
(informants).
Figure 1 .l: Bulb variation within the Ledebouriinae. A & G, bulbs of a Resnova maxima plant; B, of
Drimiopsis carrii; C, bulb of Drimiopsis maculata plant; D, F & I, bulbs of Drimiopsis burkei plants;
E, bulb of a Ledebouria sp. plant; H , Drimiopsis pusilla bulb. (Drawings enhanced by A. Lindeque).
...:I
Figure 1.2: Bulb morphology variation in the Ledebouriinae: A, aD. pusilla bulb displaying loosely
packed bulb scales; B, D. maculata bulb with bulbils; C, D. burkei bulb with a fundus; 0, a D.
comptonii bulb beginning to form stolons; E, a bulb of a D. burkei plant displaying the unusual
membranous outer scales with a tuberescent fundus (stem plate protruding beneath the bulb); F, the
small bulb of D. liniopapilla; G, a D. burkei bulb with bulbil; H, a Ledebouria species with the
outer brownish coloured membranous scales; I, a typical Resnova bulb.
18
Some Ledebouria are used as food or medicine; others have superstitious connotations.
L. inquinata (C.A. Sm.) Jessop, L. revoluta (LJ) Jessop, L. ovafifolia (Bak.) Jessop and
L. cooperi (Hookf) Jessop contain cardiac toxins (Venter, 1993). There is no
ethnobotanical record on Resnova.
1.5 ABUNDANCE
Drimiopsis plants grow in isolated populations. Populations in turn may be sparse or
abundant. Seven species are known from a single locality only. According to Gibson
(1975), D. maculata grows freely in parts of South Africa and on the salty beaches of
the KwaZulu-Natal coast. Currently, it is scarce on the aforementioned habitat but
common as an ornamental plant. In addition, plants are now absent in many previously
collected localities, yet none are listed in the Red Data book for South Africa (Golding,
2002). There are Red Data records for Swaziland. D. maculata is listed as low risk,
with declining populations but of limited concern. D. maxima is listed as 'data
deficient', with not enough data to classify it otherwise (Dlamini & Dlamini, 2002).
Some species of Ledebouria appear to be widespread but others are of limited
distribution (Venter, 1993). The Namibian L. scabrida Jessop is Red Data listed and
has a conservation status of 'data deficient7(Craven & Loots, 2002).
No records exist of the conservation status of Resnova.
1.6 OBJECTIVES
To conduct a comprehensive systematic revision of Drimiopsis by:
o exploring morphological, anatomical, karyological, molecular, palynological
and phytochemical characters and their states;
o evaluating the importance of these characters and their states through phenetic
and cladistic analyses;
o inferring a phylogenetic history
o providing a classification scheme and keys
o delimiting the three genera of the Ledebouriinae
1.7 MODUS OPERANDI
Chapter two complements the introduction. Chapters three to eight examine
characteristics of the genus while nine to twelve evaluate and synthesise data to propose
relationships via a phylogenetic classification scheme, as well as an identification key
and a taxonomic discourse on Drimiopsis.
Some previously synonymised taxa are treated as independent taxa. Otherwise use of
names is as indicated under 'Accepted names, new names, rankings and combinations'
(page X). To heighten understanding of the taxonomic significance of Drimiopsis
characters, this review encompasses the two other genera in subtribe Ledebouriinae,
namely Resnova and Ledebouria.
The plant material used in investigations was collected from the field, institutional
botanical and private gardens and then cultivated under uniform conditions in the
botanical garden of North-West University, Potchefstroom Campus. The study is also
based on herbarium material including type specimens, from B, BLFU, BOL, BR,
BTU, J, K, NBG, NH, 0, P, PRE, PUC, UNIN, and Z.
Records of morphological characteristics, descriptions of the plants in their natural
habitat and after cultivation were kept. Some characters, especially of sister taxa, were
obtained from appropriate literature. Measurement of quantitative characters used is
average measurements from a minimum of five. Photographs and illustrations, where
not by the author, are dully acknowledged. Characters and their states are compiled and
generated via DELTA (Dallwitz et al., 2000).
1.8 CONCLUSION
"If there is one thing that systematics is about, one key word that best sums us up, it is
diversity. We organize it, name it, catalogue it, and try to understand how it has
developed over time. In so doing, we are probably the single most synthetic scientific
discipline there is" (Lammers, 1999: 495). To this endeavour, this systematic revision
explores the old (orthodox) and the new (molecular) sources of data, re-evaluating the
old and generating 'new' concepts. This process raises debates especially on species
concepts and nomenclature.
2. SYSTEMATIC CONSIDERATIONS
2.1 INTRODUCTION
Thirty-seven names pertaining to Drimiopsir have surfaced to date and eight species
described within Resnova in the literature. Manning et al. (2004), diverting from the
norm, sunk both Resnova and Drimiopsis into Ledebouria, hence the 68 Ledebouria
species (IPNI, 2004). Classification of taxa within the Ledebouriinae raises debates
especially on species concepts, nomenclature and the analytical procedures leading to
the major product of a systematic analysis, a hypothesis of relationships. This group
especially presents a challenge as it lacks 'good' delimiting diagnostic characters
(Jessop, 1972; Speta, 1998a; Stedje, 2001; Wetschnig & Pfosser, 2003; Manning et a].,
2004).
2.2 SPECIES CONCEPTS
Taxa bear characters, with their character states, and are recognisable by the unique
combination of such characters (Schuh, 2000). Polymorphism and the resultant diverse
morphotypes within taxa of Drimiopsis Lindl. & Paxt. demand the exploration of
definitions of a species, as potential modes of speciation are somewhat dependent on
what constitutes a species. Several species concepts have been proposed but of
significance to this study are the biological (BSC), morphological (MSC) and
phylogenetic species concepts (PSC).
Two fundamental considerations are at the core of the vast array of species concepts:
pattern and process (Lidin & Oxelman, 1989). To quote the aforementioned authors,
process species concepts are: ". ..functional (spatio-temporal) units
... species with
pattern-forming (evolutionary) entities". Pattern-based species are perceived as "one of
many taxonomic categories, which may be seen either as historical entities
(monophyletic group including the ancestor) which are recognized (and named) or as
mental constructs (classes) which are defined by attributes".
The BSC has a process conceptual framework. Species are "a reproductive community
of populations, reproductively isolated from others, that occupy a specific niche in
nature" (Mayr, 1942, 1982). Species arise through one or another isolating mechanism,
halting gene flow between populations and isolating one such population that then
acquires a unique genotype, which is often (but not always) expressed in the phenotype.
Radiation fiom an ancestral stock, forming various new lineages adapted to differing
habitats, also produces new species. This scenario embraces the geographical species
concept (Kluge, 1989; Frost & Kluge, 1994). The BSC does not apply well to plants,
including Drimiopsis that reproduce both sexually and vegetatively. Each isolated
population in Drimiopsis is not necessarily a species, although these populations are
prime candidates for speciation in theory, if sexual reproduction is preferred.
Practically, biological species are initially delimited using characters followed by a
demonstration of a reproductive isolation mechanism. The occurrence of sibling species
may result in an underestimation of the total number of species present in a genus.
Sibling species are phenetically alike yet reproductively isolated. Added to which are
the problems associated with breeding experiments in an attempt to resolve biological
species. From a cladistic viewpoint, the ability to reproduce is considered a
plesiomorphic character and therefore inapplicable in determining relationships
(Donoghue, 1985). Although the BSC is to a large extent theoretically sound, the nonoperationalism thereof in especially plants has counted against its favour.
The MSC and the PSC are pattern based. These concepts view taxa as products of
evolution, not its determinants. The MSC is defined according to Cronquist (1988) as
"the smallest groups that are consistently and persistently distinct and distinguishable
by ordinary means." These could independently include molecular, phytochemical,
cytogenetic, or any other characters arising from speciation. Thus, the MSC relies on a
singular emphasis of a group of characters in species definition and is thus a purely
operational concept. It is pre-evolutionary and non-evolutionary. Similarity of
characters can be the result of parallel or convergent evolution. The MSC does not
distinguish among these. Parallelism and divergence are possibilities in Drimiopsis.
Convergent evolution of course provides a false signal in terms of relationships. In a
phenetic analysis variation in characters is dealt with by a fixed or intuitive standard as
to the permissible deviation from the pattern, in other words grouping is arbitrary.
According to Simpson (1951) the only serious modern theoretical support for
typological taxonomy comes from those few students who believe that species arise by
abrupt morphological change from one morphotype to another, that is to say, those who
support spontaneous generation of species.
Numerous PSCs exist, grouping taxa based on the best hypothesis of their phylogeny.
Baum's (1992) monophyletic PSC, based on De Queiroz and Donoghue (1990a,
1990b), identifies species via cladistic analysis. Monophyletic groups could be all
descendents of a common ancestor including the ancestor or groups that are more
closely related to each other than to any other organism. These are identified through
synapomorphic characters (remembering that convergence can result in grouping based
on non-homologous characters and that parallelism causes homoplasy).
Species concepts mostly apply in specific investigative situations. O'Hara (1993) aptly
likens "systematic generalization" to cartography. Drawing a map, especially of a small
area, lacks precision.. .the finest details always get sacrificed for clarity. This becomes
apparent when trying to define species. Systematists (De Queiroz & Donoghue, 1988,
1990a; Nixon & Wheeler, 1990; Vrana & Wheeler, 1992; Liden & Oxelman, 1989;
Baum & Donoghue, 1995; Schuh, 2000) concur on the plural and subjective nature of
species concepts.
Choice of a character based or history based PSC, depends on the goal of the systematic
study (Baurn & Donoghue, 1995). The primary goal of this study is to 'describe the
hierarchical distribution of characters' and infer phylogeny therein. Species in this
instance are "from a cladistic viewpoint, the cladistically indivisible elements to be
analysed, and from a phylogenetic viewpoint,
..."the ultimate products of phylogenetic
history" (Nixon & Wheeler, 1990: 214). Species are regarded diagnosable by "unique
combination of character states in comparable individuals" and analysable by cladistic
methods (Nixon & Wheeler, 1990: 218). Although cluster analysis and cladistics may
result in similar topologies, it is only synapomorphic characters that can resolve the
'history' or phylogeny on a theoretically sound basis. Character distributions in
organisms are history based because they are a product of evolution. However, this
view is strongly criticised by the "evolutionary phylogenetics" school that view this as
sitting "on the fence" (Baum & Donoghue, 1995).
I concur with De Queiroz & Donoghue (1988) that diversification of characters and
states established through speciation are validated through character persistency.
Though speciation is difficult to observe, characters developed thus are persistent in
populations while those accrued as a temporary response to changes in the environment
are not.
2.3 NOMENCLATURE
Linnaeus (1753, 1758), theoretically a creationist, devised a simple scheme for naming
and classifying organisms. From his system stemmed three nomenclature principles:
the binomen, priority, and hierarchical categories. A binomen, character naturalis, the
generic name and the dzferentia, the character that differentiates that species fiom the
others of its kind, vis a vis, the specific epithet after 1753, have been used to name
species. The principle of priority; that the first-published name is the one that is used; is
applied by the International Code of Botanical Nomenclature (ICBN) (Greuter et al.,
2000). The Linnaean notion of an inclusive hierarchy of group names has also been
important-a
hierarchical system whereby organisms are placed in categories made up
of ranks of increasing inclusiveness and given names. Genera are organised into orders,
orders into classes etc., and not always necessarily reflecting phylogeny in the past, but
similarity.
The ICBN is based on the Linnaean system i.e. the Linnaean ranks are still employed
although relatedness is now viewed to be the product of evolution.
A new concept has recently been proposed. The PhyloCode believes classification
systems are not real but that phylogeny is (Benton 2000; Langer, 2001). Briefly, the
PhyloCode is rankless, consists of biological entities called clades and has no types but
specifiers (Cantino & de Queiroz, 2000). The PhyloCode purports to "use of
phylogenetic definitions" thus "liberating taxonomy from a 2000 year old tradition of
basing the definitions of taxon names on characters" (De Queiroz & Gauthier, 1990a).
It is based on common ancestry, cladistic analysis and the naming of clades thus
creating "nomenclatural stability". De Queiroz & Gauthier (1992, 1994) claim that the
principles of phylogenetic nomenclature will produce stability in classification. The
ICBN nomenclature is the outcome of classification.
The PhyloCode and the ICBN both reflect the "hierarchical" nature of life. They have
similar goals but approach this from different directions. The criticism put forward by
De Queiroz & Gauthier (1992) that 'Definitions of taxon names based on organismal
traits are fundamentally non-evolutionary. Such definitions were in use long before the
widespread acceptance of an evolutionary world view, and finthemore, they make no
reference to common descent or any other evolutionary phenomenon,' is not persuasive
enough to discard the well tested ICBN.
Unlike the ICBN the PhyloCode removes predictability from names i.e. hierarchical
relationship and taxonomical information is lost and relationship cannot be inferred, as
is the case with binomial names. Also many formal names are attached to intermediate
clades creating too many names and possible confusion arising from incompatible
definitions. The PhyloCode does not improve the phylogenetic structure of a
classification, neither does it improve its value. Abandonment of the Bio-Code will
produce mayhem, confusion, and distress especially in normal systematic work,
revisions and extension of knowledge about the groups (Benton, 2000). Others like
Jsrgensen (2002, 2004) have suggested informal naming of clades and maintaining the
well-proven ICBN. More recently proposals have been put forward adapting the
International Code of Botanical Nomenclature to phylogenetic classification (Barkley
et al., 2004a; Barkley et al., 2004b).
I believe that phylogeny reflects evolution spearheaded by, amongst others, natural
selection (Darwin, 1859). Classifications should reflect real patterns in nature, the result
of evolution. The nodes, the branching points on cladograms, infer splitting, and clades
common ancestry. However, without paleobotanical data one cannot argue common
ancestry, only put forward hypothesise. Cladistics and molecular techniques greatly
improve the potential of phylogenetic hypotheses. Linnaean based classifications work,
are practical and can be used in phylogenetic inference. It is straightforward, has been
used for decades and serves as a universal reference system, not only for veteran
systematists, but also for aspiring systematists and the novice.
2.4 CONCLUSION
Debates, differences of opinion and interpretations reflect diversity and dynamism
conventional in investigating nature. Systematics should be about recreating the best
phylogenetic inference, and presenting the best classification system possible, with the
tools and data at hand. Putting nature in boxes is not feasible (Vrana & Wheeler, 1992),
one can all but look for the best hypothesis-be
it singular or combinations of views.
After all, "systematic botany" is "an unending synthesis" (Constance, 1964).
This study analyzed diagnostic character groupings phenetically and cladistically, the
clades and terminal taxa thus generated used to establish species. The PSC and the
ICBN are furthermore employed in the interest of nomenclatural stability.
(3.THE LEAF
1 3.1 INTRODUCTION '
Although taxonomists have placed greater importance on reproductive characters due to
their conservative nature, vegetative characters remain equally important in systematic
studies. The delimitation of the traditional two major groups of plants, the
monocotyledons and the dicotyledons were based on floral and vegetative characters.
The vegetative stage is the most prevailing stage in the lifecycle of Drimiopsis Lindl. &
I
Paxt. so that plant identification is often required when no flowers or fruit are present.
The macro and micro morphology as well as anatomy of the leaf consequently solicit
(
investigation.
Macroscopic leaf characters have been used in the past to delimit Ledebouriinae U. Br
)
' '
D. Miiller-Doblies and taxa of Drimiopsis in particular. Baker (1896) predominantly
used leaf size, shape, orientation as well as pseudopetiole presence or absence to
delimit southern African taxa (Key 3.1). D.minor Bak., for example, was primarily
1
based on the possession of leaves smaller than those found in D, rnaculata. Jessop
(1972), demonstrating the unreliability of leaf size in delimitations, sunk D. minor Bak.
in D. maculata Lindl. & Paxt. (a decision subsequently supported by Miiller-Doblies &
Miiller-Doblies, 1997)). Jessop (1972) instead mainly uses flower characters, confining
himself to the use of leaf base in addition to number of flower characters to delimit D.
I
maculata, D. burkei and D. ahopurpurea. Miiller-Doblies & Miiller-Doblies (1997) use
numerous leaf characters (except size) in combination with floral characters in
delimiting taxa of Drimiopsis. In addition they simplify matters by de-emphasizing
Baker's (1897) distinction between erect and spreading leaves in synonymising D.
maxima Bak., D. humijisa Bak. and D. saundersiae Bak. under Resnova humifusa
(Bak.) U . & D. Miiller-Doblies.
I
' Part of this chapter is in prep.,Lebatha & Buys (2005a).
Key 3.1 : Key to southern African Drimiopsis species that Baker (1896) first delimited
on the basis of leaf shape. Compiled from Baker (1870a, 1874a, 1874b, 1878, 1898).
Leaves distinctly :
Blade cordate:
Perianth '1 6 in. long
Perianth '/ 8 in. long
Blade narrowed at the base:
Raceme % - 1 in. long
Raceme 3 - 5 in. long
Leaves sessile:
Leaves erect:
Dwarf; small flowered
Tall; large flowered
Leaves spreading
( 1 ) D. maculata
( 2 ) D. minor
(3) D. woodii
(4) D. maxima
(5) D. burkei
( 6 )D. saundersiae
(7) D. humifusa
Similarly Stedje (1994 & 1998), confining herself to the tropical east Afiican taxa,
expands Baker's (1870a, 1874a, 1874b, 1878, 1898) use of leaf shape and adds leaf
orientation in her description of the two subspecies in D. botryoides Bak. (Key 3.2).
Taxa with narrowly lanceolate leaves were placed in D. barteri Bak., those with
lanceolate to cordate and erect leaves in D. botryoides Bak. (=D. botryoides Bak. subsp.
botryoides) and ones with broadly lanceolate leaves appressed to the ground in D.
perfoliata Bak. (=D. botryoides Bak. subsp. prostrata Stedje).
Key 3.2: Key to tropical African Drimiopsis species delimited initially on the basis of
leaf shape and later with the addition of leaf orientation (Stedje 1994, 1998).
1a. Leaves erect
2a. Lamina narrowly lanceolate.. ......................................D. barteri
2b. Lamina lanceolate to cordate. ...........D. botryoides subsp. bofryoides
1b. Leaves appressed to the ground
3a. Lamina broadly lanceolate.. .............. D. botryoides subsp. prostrata
Varying terminologies in describing leaf character has led to inaccuracies, e.g. Baker
,
(1896) and Muller-Doblies & Muller-Doblies (1997) wrongly referred to leaves with an
- - - - -
extended leaf base as-petiolate; A petiole, t h e leaf stalk connecting the lamina to the
- - - - - - - - - - -
- - - - - - - - - - - - - - - - - - -
stem, is absent in the Ledebouriinae. The extended leaf base, referred to in this thesis as
)
pseudopetiole, is sometimes present.
I
Leaf micromorphology studies have previously revealed underlying similarities and )
differences that are not otherwise easily perceived. Scanning electron microscopy of
I
epidermal surfaces has been routine for almost half a century, beginning with the work
of Bringman. & Kuhn (1955a, 1955b) as well as Amelunxen et al. (1967) and Barthlott
& Ehler (1977). Characters, especially those from the epicuticular wax, have been used
widely in angiosperm classifications (Barthlott, 1981; Behnke & Barthlott, 1983;
Dahlgren, 1989; Barthlott, 1990 & 1994; Barthlott et al, 1998). A host of investigators
use leaf micromorphology as an investigative tool, e.g. Haron & Moore (1996), Darok
et a1 (2000), Kong (2001), Wang et al. (2002) and Ao et al. (2002).
Reports by Stace (1965) and Dilcher (1974) that characters like trichomes, stomata
density and distribution, cuticular thickness and striations, as well as the anticlinal wall
patterns of epidermal cells vary according to the environment, create uncertainty about
their systematic value. Nevertheless, ultra structural and micromorphological
characters, in particular the epidermal cell arrangement, cell wall shape, cuticle and
trichomes have been used in resolving lower level systematic problems as the influence
the environment exerts on them have since been found to be minimal (Behnke &
Barthlott, 1983). Experimental evidence from studies on Aloeae A. Rich. (Aloaceae)
hybrids reveal strong genetic control over trichome, stomata and basic pattern of the
cell wall in the first generation hybrids (Cutler & Brandham, 1976). In contrast, wax
particle shape and size varied greatly in the offspring. Even so, tertiary surface
sculpturing, i.e. epicuticular secretions, has been used to delimit genera in certain
instances (Behnke & Barthlott, 1983).
In terms of the Ledebouriinae, the only attempt at studying the leaf micromorphology
in Drimiopsis has been by Shiva et al. (2001) on D. botryoides. The said authors do not,
however, expound on the interspecific nor intraspecific significance of the leaf surface
I
characters. Venter (1993) studied the micromorphology of South African taxa of
Ledebouria Roth, but no micromorphological studies have been done on Resnova v.d.
Menve to date.
Anatomy has been applied with varying degrees of success in delimiting genera (Smith
& Ayensu, 1974, 1976; Behnke et al., 2002) and for families (Dahlgren & Clifford,
1982; Ayensu, 1968, 1969, 1974). At the species level, taxa delimitation based on
anatomy is usually a matter of degree of variation. Secondary xylem, indumentum, and
the distribution of schlerenchyma and ergastic bodies are often
taxonomic ally^
1
significant, the latter two especially so in the Monocotyledonae (Metcalfe & Gregory,
1964; Metcalfe, 1967; Dahlgren & Clifford, 1982; Mauseth, 1988)-the
secondary
xylem, however, is not of use when dealing with herbaceous monocots.
In terms of the Hyacinthaceae, ergastic substances, mucilage, calcium oxalate crystals,
raphides and "rhexigenetic" lacunae have been accounted to be characteristic (Speta,
1998b; Watson & Dallwitz, 2003). Dahlgren et al. (1985) report on the apparent
absence of raphides in the Hyacinthaceae. Phytochemical screening tests reveal the
presence of tannins in Resnova maxima v.d. Merwe and traces thereof in the leaves of
Drimiopsis kirkii Bak. (Chapter 6). Leaves of Hyacinthaceae have been reported as
being bifacial with vascular bundles arranged adaxially and the presence of
isodiametric palisade cells (Speta, 1998). In terms of sister taxa, the anatomy of
Resnova leaves has not been investigated to date and Venter (1993) studied some leaf
anatomy in Ledebouria. The presence of substances in epidermal cells could be
adaptations to stresskl environmental conditions (Metcalfe & Chalk, 1950; Metcalfe,
1967; Jordaan & Theunissen, 1992).
3.2 OBJECTIVES
To investigate the taxonomic significance of macromorphological, micromorphological
as well as anatomical leaf characters and states in Drimiopsis as well as Ledebouria and
Resnova. To supply a key to species of Drimiopsis based on leaf characters.
3.3 MATERIALS and METHODS
The taxa investigated for leaf surface characters and anatomy are listed in Table 3.1.
Plants were observed in their natural habitats then cultivated in the Botanical Garden of
the North-West University, Potchefstroom Campus. Records were kept of field and
garden observations in terms of leaf shape, surface features and coloration. Herbarium
material was used in the case of some Drimiopsis taxa and Schizocarphus nervosa.
Leaf size consists of length (including pseudopetiole when present), a measure of the
lamina up to the apex, and width, a measure of the widest part of the lamina. Only fresh
material was used for anatomical study (Table 3.1). The extended leaf base is termed
pseudopetiole. Tinted refers to colour differing from that of the general "green" of the
lamina. Colouration may occur in the form of transverse stripes (banded), longitudinal
blotches (streaked), dots (spotted); or it can cover the entire surface. Colouration of the
margins may be in the form of bold colours (bordered) or lighter colours (edged)
(Radford et al., 1974).
Terminology follows Barthlott & Ehler (1977), Behnke & Barthlott (1983) and
Christensen & Hansen (1998) for epidermal cell arrangement (including trichomes,
stomata, glands), cell outline (boundaries of the anticlinal walls and curvature of
periclinal walls) and cell sculpturing, including cuticle striations and epicuticular waxes
(Table 3.1).
Fresh material was preserved in 80% ethanol for micromorphological investigations.
Fresh leaves were cut into about 2 rnrn2and dry leaves fiom herbarium material were
cut to about 1 mrn2. Care was taken to ensure that all the sections were taken fiom the
centre of the leaf. The abaxial and adaxial sections were then processed, using the
standard SEM preparation procedure, and then viewed at 1OkV. The dry leaf sections
were carbon treated and gold plated before viewing them with a Philips SL30DX - 4i
SEM.
Anatomical studies were done on freshly cut material of plants in cultivation. Leaves
and pseudopetioles were fixed for 2-2.5 hours in a mixture of paraformaldehyde and
1% glutaraldehyde in 0.1 M phosphate buffer at pH 7.2 at a temperature of 4°C. To
stabilise mucilage that may interfere with the fixation process, 0.05% alcian blue 8GX
was added to the fixative. Material was post-fixed for one hour in 1% aqueous 0 s 0 4 ,
dehydrated in an ethanol series and embedded in LR White resin. Sections were cut
with a Leica Ultracut R ultramicrotome and stained with 0.05% aqueous toluidine blue
followed by 0.05% neofuchsin. Observations were made with a Zeiss Axioskop I1 Light
Microscope and images were captured digitally.
Data was subjected to a multivariate cluster analysis using STATISTICA 6.1 (Statsoft,
2003) with the following settings: tree clustering: Ward's Method of minimumvariance clustering under the amalgamation rule and percentage disagreement as a
measure of distance. A leaf character based key was generated via DELTA.
Table 3.1: Taxa investigated for leaf macromorphology, micromorphology and
anatomy. Ledebouria data fiom S. Venter (1993) has an N/A (not applicable) status.
Accepted names in
this thesis
D. atropurpurea
D. atropurpurea
D.atropulpurea
D. atropurpurea
-
-
I Accession
D. barteri
D. barteri
I D.
I
botryoides subsp. D. botvoides
I Area
1 Status
South Africa
South Africa
South Africa
South Africa
Herbarium
Herbarium
Fresh
Herbarium
Tanzania
Tanzania
Fresh
Herbarium
Herbarium
Kenya
I
D. botryoides
botrvoides
D. botryoides subsp. D. botryoides
botrvoides
D. botryoides subsp. D. botryoides
Lebatha 098
Tanzania
Fresh
Lebatha 003
Kenya
Fresh
J
D. botryoides
Lebatha 099
Tanzania
Lebatha 003
Kenya
Lebatha 004
Kenya
Reid 1090
Kenya
Fresh
Fresh
I
D, botryoides subsp. D. botryoides
D. botryoides
I
D. botryoides
I botrvoides
I D.
&
voucher numbers
Rogers 18508
Schrierp 1330
Lebatha049
Van der Menve
0266 1
Lebatha 002
Greenway
&
Kaburi 14,782
Greenway 12854
Reid 1984
I
Fresh
Herbarium
I
Kenya
D. perfoliata
I
Herbarium
R.B. & A.J. Faden Kenya
74,505
Tanzania
Lebatha 001
botryoides subsp. D. botryoides
Herbarium
Fresh
rostrata
J
1 D. burkei subs^. burkei
I D. burkei subso. burkei
stolonissima
1 D. cawii
-D,ehae
- - - - - - - - - - -
D. lachenalioides
I D. lachenalioides
I D. liniovawilla
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. stolonissima
Lebatha 009
Lebatha 04 1
Lebatha 096
Lebatha 054
Lebatha 103
I Lebatha 056
I Lebatha 095
Theron 1589
D. carrii
D. burkei
D. comptonii
D. davidsoniae
D. Jischeri
D.-kikiae
D. kikiae
D. liniopapilla
R. lachenalioides
Lebatha 015
Codd 80 18
Lebatha 079
Lebatha 038
Fischer 325
Lebatha 045
LebathaU6
I Lebatha 053
Van der Menve
21 17
I Baut 549
Lebatha 060
Lebatha 005
Lebatha 006
Lebatha 007
- - -
- - -
R. lachenalioides
D. liniopapilla
D. maculata
D. maculata
D. maculata
Lebatha 037
-
Fresh
Fresh
Fresh
Fresh
Fresh
I Fresh
I
I Fresh
I
Herbarium
Fresh
South Africa
South Africa
Botswana,
South Africa
Botswana
I South Africa
I Botswana
South Africa
South Africa
South Africa
Fresh
Herbarium
Locality unknown
Fresh
Swaziland
South Africa
Fresh
Herbarium
Tanzania
South Africa
Fresh
s a u t f i - ~ f ~ i ~ ~'Freshh
I South Africa
1 Fresh
Herbarium
South Africa
- - - - - -
I South Africa
South Africa
Botswana
South Africa
Botswana
-
- - - -
I
I Herbarium I
4
Fresh
Fresh
Fresh
Fresh
1 D. maculata
Lebatha 039
Lebatha 062
Lebatha 102
Lebatha 032
Abbot 643 1
Moss 16777
Van Jaarsveld
6010
Venter s.n.
Lebatha 078
Lebatha 055
Van der Merwe
s.n.
Repton s.n.
Liebenberg. s.n.
Rogers 214 09
Codd 5 126
Lebatha 068
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
R. maxima
D. maculata
D.maxima
R. maxima
D.pusilla
D. queue
1 D. queae
D. queae
D. queue
D. queue
D. queue
D. queue
D. reillevana
D. rosea
1 D.rosea
I D. rosea
[ D. woodii
I R. lachenalioides
I
I
R. maxima
R. maxima
R. maxima
R. maxima.
L. concolor
L. floribunda
D. queue
D. queue
D. queue
D. queue
D. reillevana
Fresh
Fresh
Fresh
Fresh
Herbarium
Herbarium
Herbarium
South Africa
Swaziland
South Africa
Herbarium
Fresh
Fresh
Herbarium
South Africa
South Africa
South Africa
South Africa
South Africa
Swaziland
Democratic
Chevalier 8432
D. rosea
Re
I D. rosea
1 Goossens 43
1 South Africa
I D. rosea
I Codd s.n.
I South Africa
I D. woodii
I Lang 32236
I South Africa
R. humifusa
Schlechter 3 174 )
South Africa.
R. humifusa
Devenish 958
South Africa
R. humifusa
Van der Merwe South Africa
s.n.
I R. lachenalioides I Singh 72
I South Africa
R. maxima
Lebatha 042
South Africa
R. maxima
Swaziland
Lebatha 077
R. maxima
Lebatha 042
South Africa
R. maxima.
Lebatha 047
Swaziland
R. megaphylla
Lebatha 05 1
South Africa
Ledebouria sp.
Lebatha 0 10
Botswana
Ledebouria sp.
Lebatha 050
South Africa
Ledebouria sp.
Lebatha 059
I South Africa
L. asperifolia
Lebatha 057
I Swaziland
L. asverifolia
Lebatha 080
South Africa
L. asperifolia
Lebatha 090
Botswana
I L. concolor
I L. jloribunda
I Re: Venter (1993) I South Africa
Herbarium
Herbarium
Herbarium
Herbarium
Fresh
Herbarium
1 Herbarium I
1 Herbarium I
I Herbarium I
Herbarium
Herbarium
Herbarium
I Herbarium I
Fresh
Fresh
Fresh
Fresh
Fresh
Fresh
Fresh
I Fresh
I Fresh
Fresh
Fresh
I
I
I
I
I
I
I
I
L. ovatifolia
L. ovatifolia
L. revoluta
- - - - - - - - - -
South Africa
Swaziland
South Africa
South Africa
South Africa
South Africa
South Africa
I NIA
I NIA
Re: Venter (1993)
I South Africa
L. znquinata
L. luteola
Lebatha 075
Re: Venter (1993)
South Africa
South Africa
Fresh
NIA
L. ovat$olia
L. ovatifolia
L. revoluta
Lebatha 008
Lebatha 063
Re: Venter (1 993)
Botswana
South Africa
South Africa
Fresh
Fresh
NIA
e a t 0
Re: Venter (1993)
Swaziland
South Africa
Fresh
NIA
Codd 373 1
South Africa
(
L. sandersonii
L. socialis
Schizocarphus nervosa
S. nervosa
- - - -
- - - - - - -
1
I
- - - - - - - - - - - - - -
L. sandersonii
L. socialis
I
I
- - - - - - - - -
-
-
Herbarium
Table 3.2: Illustrations and descriptive terminology of cell shape and sculpturing used
in this report. Adapted from Barthlott & Ehler (1997) and Christensen & Hansen
(1998).
A. Primary cell shape:
isodiametric-tetragonal
elongate-tetragonal
B. Anticlinal wall boundary:
1
Irregularly-sinuate , curved or S-undulate
Straight
C. Anticlinal cell boundary can be:
Undelimited; poorly delimited; depressed, channeled or grooved or simply raised
D. Periclinal wall curvature:
1.
Straight
I
tabular cell
2. Tabular-convex cells:
W
i
t
h papillae
with hair-like papillae
with hairs
3. Non-tabular convex cells:
Convex
hemispherical
domed
conical
with hairs
E. Cuticle sculpturing:
-
1. Degree of folding or striations can be: none; weakly folded or prominent.
2. Arrangement of striae can be: unordered-wrinkled; arranged in parallel to each other;
vertically arranged with respect to lamina cell alignment; horizontally arranged or variously
arranged.
- - - - - - - - -
- - - - - - - - - -
- - - - - - - - - - -
- - - - -
- - - - -
- - - - -
- - - - -
- - - - - - - - - - - -
3.4 RESULTS and DISCUSSION
The data matrix is presented in Table 3.3.
3.4.1 MACROMORPHOLOGY
Leaf shape, dimensions and orientation
The leaves of Drimiopsis are simple, erect or appressed to the ground, unsheathed,
parallel veined and sometimes pseudopetiolate. Length ranges from 2 cm in D. pusifla
U. & D. Miiller-Doblies (Figure 3.1 C) to about 45 cm in D. maculata Lindl. & Paxt.
(Figure 3.1 A). The lamina form varies from cordiform in D. atropurpurea N.E. Brown,
D. perfoliata Bak., D. carrii Lebatha (Figure 3.1 B), D. kikiae Lebatha, D. liniopapilla
Lebatha, D. pusiNa (Figure 3.1 C) and D. woodii Bak., or falciform in D. botryoides
Bak., D. burkei Bak., D. stolonissima U. & D. Muller-Doblies, D. davidsoniae U. & D.
Muller-Doblies and D. queae Lebatha; or spathulate in D. comptonii U. & D. MiillerDoblies. Drimiopsis maculata possesses leaves cordiform to slightly sagittiform.
Drimiopsis barteri, D. jkcheri (Engl.) Stedje, D. reilleyana U. & D. Miiller-Doblies
and D. rosea A. Chev. possess linear leaves.
Leaf margin
Leaf margins are entire in D. atropurpurea, D. barteri, D. botryoides, D. perfoliata, D.
stolonissima, D. comptonii and D. fischeri, or crenate in D. burkei, D. davidsoniae, D.
liniopapilla, D. kikiae and D. queae, to crenulate in D. pusilla. The margins are
undulate in D. maculata, D. woodii, D. maxima and D. reilleyana. D. carrii is unique
with a clefted crenate margin (Figure 3.1 B). Most taxa possess margins that are banded
except in D. barteri, D. burkei, D. comptonii, D. fischeri, D. maculata and D. woodii.
Pseudopetiole
Drimiopsis maculata, D. atropurpurea and D. woodii possess pseudopetiole
considerably longer than the lamina. The pseudopetiole in D. kikiae, D. liniopapilla,
and D. queae are approximately as long as the lamina. The oblanceolate lamina of D.
- - - - - - - - -
- - - -
comptoni tapers 36 3 i e baseforming a b n g narrow lamina extension, a n d i s therefore
- - - - - -
considered to be sessile and not pseudopetiolate. The leaves of D. barteri, D. burkei, D.
stolonissima, D. botryoides, D. perfoliata, D. carrii, D. davidsoniae, D. fscheri, D.
pussila, D. reilleyana and D. rosea are also sessile.
Figure 3.1: Variation in leaf shape and orientation in taxa of the Hyacinthoideae. A, pseudopetiolate
young D. mandata plant with cordate spreading leaves; B, D. carli with cordiform to ovate leaves
appressed to the ground; C, D. pusilla, a sessile dwarf plant with thick lamina tinted purple abaxially;
D, L sandersoni: with leaves slightly appressed to the ground; E, L. asperifolia with sessile, lanceolate
leaves with adaxially spotted lamina; F, R. maxima spreading, sessile leaves.
Leaf orientation
The majority of taxa possess erect leaves. The leaves are however spreading in D.
botryoides, D. fischeri, D. kikiae, D. maculata (Figure 3.\ A), D. rosea and D.
36
-
---
----
----
stolonissima. The leaves of D. perfoliata and D. carrii are appressed to the ground
(Figure 3.1 B).
Leaves vary in terms of shape, colouration, maculation, length and width according to
their age. The proportions of lamina length to pseudopetiole length as well as lamina
length to width are generally constant. This suggests that leaf shape-a
length and width to a degree -is
function of leaf
a good delimiting character. Species also react
differently to cultivation. D. queae possessing leaf lamina 2 cm long and 0.5 cm wide
in its natural habitat increased slightly in length and width in cultivation whereas the
remaining taxa all showed considerable increase. Young D. maculata leaves possess
pseudopetioles as long as or a bit longer than the lamina (Figure 3.1 A; Frontispiece).
This sometimes has led to the incorrect description of new taxa, e.g. D. minor, which is
now viewed to be a young form of D. maculata which, when mature, the ratio of lamina
to pseudopetiole can be as much as 1:10, both in the field and in cultivation.
Leaf colouration
Leaf colouration is variously present in Drimiopsis. The lamina is adaxially spotted in
D. atropurpurea, D. botryoides, D. stolonissima, D. carrii, D. kikiae and D. maculata.
Drimiopsis atropurpurea, D. burkei, D. queae, D. kikiae and D. pusilla (Figure 3.1 C )
are tinted deep purple abaxially. D. burkei may be purple streaked abaxially.
Drimiopsis liniopapilla, D. reilleyana and D. maxima are purple streaked, more so at
the lamina base. Pseudopetiole is banded in D. maculata and D. woodii and purple
streaked in D. atropurpurea and D. liniopapilla. Spots vary in intensity. Plants in full
sun display stronger tinted spots compared to those in the shade.
The leaves of Ledebouria (Figures 3.1 D & E) and Resnova (Figures 3.1 F) are
superficially similar to those of Drimiopsis. They are linear, never pseudopetiolate,
their margins simple and entire. The leaves can be either spreading, as in L. asperijolia
(v.d. Merwe) S. Venter (Figure 3.1 E) and R. maxima v.d. Merwe (Figure 3.1 F), erect
orpessed-te &ehegreundas inL. snnders~nii(Bak.)_S, Venter (Figure 3.1 D).
-
-
-
-
-
-
-
-
-
-
-
-
Table 3.3: Data matrix for Dvimiopsis leaf macromorphology and micromorphology characters. The characters and states are listed at
the bottom of the table. Inapplicable characters are coded as -.
I
S. newosus
D. atropurpurea
D. barteri
I
D. botryoides
D. burkei
I
D. carrii
I
D. comptonii
I
D.davidsoniae
I
D. Jischeri
I
D. kikiae
I
D. 1iniopapilla
D. maculata
D.perfoliata
D. pusilla
I
D.queae
I
D, reilleyana
1
D. rosea
I
D. stolonissima I
D. woodii
I
R, maxima
I
R. megaphylla
R, humifisa
R. lachenalioides
L. mperifolia
L. concolor
L, Joribunda
L. inquinata
1
L. ovatifolia
I
L. revoluta
I
L. sandersonii 1
L. socialis
I
1
1
I
1
3
3
1
3
2
2
2
3
3
3
1
3
2
3
1
3
1
3
3
3
2
2
3
3
3
3
3
3
3
3
3
2
1
1
2
1
3
1
3
1
1
2
2
1
1
1
2
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3
2
1
1
2
2
3
1
1
1
1
1
2
3
1
1
1
2
2
1
2
3
3
2
2
2
2
2
3
2
3
2
4
1
1
3
2
3
1
4
3
3
1
1
1
1
3
1
3
3
3
3
3
1
3
3
3
3
3
3
1
3
3
3
5
1
2
1
1
1
2
3
1
1
1
1
2
2
1
1
1
1
1
2
1
2
1
1
1
1
1
1
2
1
1
1
6
2
2
4
3
2
2
2
2
2
2
2
3
2
1
1
2
1
2
2
3
2
2
3
3
3
4
2
2
3
1
2
7
2
3
1
2
2
2
2
2
1
2
2
3
3
1
1
2
1
2
1
2
2
2
2
2
2
2
1
2
3
1
1
8
1
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
2
1
1
1
1
2
2
9
1
1
2
2
2
2
2
2
2
1
1
1
2
2
1
2
2
2
1
2
2
2
2
2
2
2
2
2
2
2
2
10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41
2 3 1 1 1 1 2 - 2 - 1 1 2 1 1 2 1 2 2 2 2 1 3 3 2 2 2 2 1 2 1 2
3 2 1 2 2 1 1 2 1 1 2 2 1 1 1 1 2 1 1 1 2 2 2 3 2 2 2 3 2 1 1 1
- - 1 2 1 1 2 - 2 - 2 3 2 1 1 1 2 1 1 1 2 2 1 1 1 - - - - - - - - 1 2 2 1 1 2 2 - 2 3 2 2 1 1 2 1 1 1 2 1 1 3 1 - - - - - - - - 2 2 1 1 1 1 1 2 2 3 1 1 1 1 2 1 1 1 2 2 2 1 2 2 2 1 2 - 2 1
- - 2 2 1 1 1 2 2 - 2 3 2 2 1 1 2 1 1 1 2 2 2 1 1 - - - - - - - - 1 2 1 1 2 - 2 - 3 1 2 1 1 1 2 1 1 1 2 2 2 3 2 1 1 3 2 - 2 1
- - 2 2 2 1 2 - 2 - 3 3 2 3 1 1 2 1 1 1 2 2 2 3 2 2 1 2 2 - 1 1
- - 1 2 1 2 2 - 2 - 1 3 2 1 1 1 2 1 1 1 2 2 2 3 1 - - - - - - 1 2 2 2 2 1 1 2 1 1 1 1 1 1 1 1 2 1 1 1 2 2 2 1 1 - - - - - - 2 2 2 2 2 1 2 - 1 2 1 1 1 1 1 2 2 1 1 1 2 2 2 2 2 1 2 3 2 1 1 2
3 1 3 2 1 2 1 2 2 - 1 2 2 1 1 2 2 1 1 1 2 2 2 1 1 - - - - - - - - 1 2 2 1 2 - 2 - 2 3 2 2 1 1 2 1 1 1 2 2 2 3 1 - - - - - - - - 4 2 2 1 2 - 1 1 2 3 1 1 1 2 2 1 1 1 2 2 2 3 2 2 2 2 2 - 1 1
2 2 2 2 2 1 2 - 1 1 1 1 1 2 1 2 2 1 1 1 2 2 2 1 1 - - - - - - - - 3 2 2 1 2 - 1 1 2 3 1 1 1 2 2 1 1 1 2 2 2 3 2 2 2 2 2 - 2 1
- - 3 2 1 2 2 - 1 2 1 3 1 1 1 1 2 1 1 1 2 2 1 1 1 - - - - - - - - 1 2 2 1 1 2 2 - 2 3 2 1 1 1 2 1 1 1 2 2 2 1 1 - - - - - - 2 1 3 2 1 2 2 - 2 - 2 1 2 1 1 1 2 1 1 1 2 2 2 3 2 2 1 2 2 2 2 1
- - 3 2 1 1 2 - 2 - 1 3 2 1 1 2 2 2 2 1 1 1 1 1 2 1 1 2 2 - 2 1
- - 1 2 2 1 1 1 1 1 2 3 1 2 1 2 2 2 2 1 1 1 1 1 2 2 1 3 2 - 1 2
- - 1 2 2 1 1 2 1 1 3 3 2 3 1 2 2 2 2 1 1 1 1 1 2 1 1 2 2 - 2 1
- - 1 2 2 1 1 2 2 - 1 1 2 2 1 2 2 2 2 1 1 1 1 3 2 1 2 2 2 - 2 1
- - 1 2 2 1 1 2 1 1 2 1 2 3 1 2 2 2 2 1 1 1 1 1 2 3 1 2 2 - 2 1
- - 1 2 2 1 2 - 2 - 3 1 2 2 1 2 2 2 2 1 2 1 1 1 1 - - - - - - - - 3 2 1 1 1 1 1 2 2 1 2 1 1 2 2 2 2 1 2 1 1 3 1 - - - - - - - - 3 2 1 1 2 - 1 2 2 1 2 1 1 2 2 2 1 1 2 1 1 2 1 - - - - - - - - 1 2 2 1 1 2 1 1 2 1 2 1 1 2 2 1 2 1 2 1 1 1 1 - - - - - - - - 3 2 1 1 1 2 1 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 1 - - - - - - - - 1 2 2 1 1 2 1 1 2 1 1 1 1 2 2 2 2 1 2 1 1 3 1 - - - - - - - - 1 2 2 1 1 2 1 1 2 1 2 2 1 1 1 2 2 2 2 1 1 1 1 - - - - - - -
#1. Leaves <number> 1. 1 (monophyllous), 2.2 (diphyllous), 3. more than 3 (polyphyllous); #2. Leaves <leaf number variation> 1. number of leaves unvarying,
2. sometimes diphyllous, 3. sometimes polyphyllus; #3. Leaves <posture> 1. erect, 2. spreading, 3. appressed to the ground; #4. Leaves <form> 1. cordiform, 2.
falciform, 3. linear, 4. spathulate; #5. Leaves <shape> 1. lanceolate, 2. ovate, 3. oblanceolate, ; #6. Leaves <length> 1. 5 cm long or shorter, 2. 5.1 to 10 cm
long, 3. 10.1 to 20 cm long, 4. longer than 20 cm; #7. Leaves <width> 1. 2 cm wide or less, 2. 2.1 to 4 cm wide, 3. more than 4.1 cm wide; #8. Leaves when
tom <with threads or not> 1. with threads, 2. without threads; #9. Leaves <pseudopetiolate or not> 1. pseudopetiolate, 2. sessile; # 10. Pseudopetiole <length> 1.
exceedingly shorter than lamina, 2. approximately as long as lamina, 3. exceedingly longer than lamina; #11. Pseudopetiole <colour> 1. banded, 2. tinted, 3.
green; # 12. Leaf margin <shape> 1. entire, 2. crenate, 3. undulate, 4. crenulate; # 13. Leaf margin <cartilaginous or not> 1. cartilaginous, 2. noncartilaginous;
#14. Leaf margin <markings> 1. edged purplehrown, 2. bordered purplehrown; #15, Lamina <thickness> 1. thick, 2. membranous; # 16. Lamina <spotted or
not> 1. spotted, 2. unspotted; #17. Lamina <spotted> 1. abaxially, 2. adaxially; #18. Lamina <tinted or not> 1. tinted, 2. green; #19. Lamina abaxailly <tinted>
1. purple, 2. streaked purplehrown; #20. Lamina apex <apex> 1. acurninate, 2. acute, 3. apex obtuse; #21. Lamina base <base> 1. attenuate, 2. cordate, 3.
cuneate; #22. Lamina base <tinted or not> 1. tinted dark purple, 2. green; #23. Epidermal wax cover <wax cover> 1. thin, 2. thick, 3. particulate; #24. Stomata
<type> 1. anomocytic; #25. Stomata distributed <fkequency distribution> 1. sparsely, 2. densely; #26. Stomata crypts <crypts> 1. raised, 2. shallow; #27.
Stomata subsidiary cells <subsidiary cell H-complex or not> 1. form an H-complex, 2. not in an H-complex; #28. Epidermal cells adaxial <adaxial shape> 1.
short polygonal, 2. elongate tetragonal; #29. Epidermal cells abaxial shape <abaxial shape> 1. elongate tetragonal, 2. short polygonal; #30. Epidermal cells
anticlinal boundaries, 1. undelimited, 2. channelled; #3 1. Epidermal cells anticlinal boundaries, 1. straight, 2. irregular-sinuate; #32. Epidermal cells periclinal
wall curvature, 1. straight tabular, 2. tabular-convex, 3. non-tabular convex; #33. Epidermal cells cuticle striae, 1. smooth, 2. regular, 3. irregular; #34.
Indurnentum <presence or absence> 1. absent, 2. present; #35. Indumentum arranged, 1. in rows, 2. randomly, 3. sparsely on margins; #36. Indurnentum in the
form of <type> 1. papilla, 2. hairs; #37. Indumentum <abundance> 1. sparse, 2. fkequent, 3. dense; #38. Indumentum on lamina <absent or present> 1. absent, 2.
present; #39. Indumentum on pseudopetiole Cabsent or present> 1. present, 2. absent; #40. Indumentum on abaxial leaf surface <present or absent> 1. present,
2. absent; #41. Indumentum on adaxial leaf surface <present or absent> 1. present, 2. absent.
Ledebouria leaves are unique within the Ledebouriinae in producing threads when tom
except in L. concolor (Bak.) Jessop, which, like Drimiopsis and Resnova, does not.
3.4.2 MICROMORPHOLOGY
Epidermal cells
The primary sculpture, i.e. the superficial shape of adaxial and abaxial epidermal cells
in Drimiopsis varies (Figures 3.2 A-L). The adaxial cells possess an overall polygonal
shape and are shorter than those abaxially. The abaxial cells are tetragonal and
elongate. The anticlinal walls are sinuate, mostly S-type undulate. Drimiopsis abaxial
epidermal cells range from 199 to 430 pm in length. The shortest cells, on the adaxial
surface, range from 45 to 164 pm, thus making the overall ratio of adaxial to abaxial
cell length approximately 1:3.
The surface indumentum is unicellular and variously elongate. Drimiops is
atropurpurea (Figure 3.2 H), D. burkei, D. liniopapilla, D. pusilla (Figure 3.1 C) and
D. reilleyana possess hairs. The hairs on the adaxial surface of the lamina of
Drimiopsis burkei (Figure 3.2 I ) are shorter than those of aforementioned taxa.
Drimiopsis maxima and D. crenata v. d. Merwe (Figures 3.2 J-K), together with D.
comptonii, D. davidsoniae and D. woodii, possess papilla.
Indumentum distribution varies from very sparse hairs on the adaxial lamina surface in
D. burkei, to the abundant, randomly distributed hairs on both surfaces of the
pseudopetiole and lamina of D. atropurpurea. The hairs on D. liniopapilla leaves are
arranged abaxially in rows forming continuous straight lines from the pseudopetiole to
the lamina. The hairs on D. pusilla are on both adaxial and abaxial surfaces of the
lamina. Those in D. reilleyana are only on the adaxial surface of the lamina. Adaxial
lamina surfaces of D. comptonii and D. woodii possess papillae arranged in rows and
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
randomly respectively. The papillae on D. davihoniae are onb-othia-a-sarfaces:
-
-
Drimiopsis stomata are ovate, mostly parallel to each other and more abundant on the
abaxial than on the adaxial surface (Figures 3.2 A-L). The subsidiary cells together
with the stomata form the letter 'H', the guard cells being the middle line of the letter
'H' (Figures 3.2 C, D, G). The subsidiary cells of the adaxial cells are isodiametricpolygonal.
Figure 3.2: SEM views of leaf epidermal surfaces. A & B, abaxial and adaxial leaf surfaces of D.
maculata, respectively; C & D, abaxial and adaxial leaf surfaces of D. perfoliata, respectively; E, abaxial
cuticle striations and wax crust on D. davidsoniae leaves; F, adaxial wax platelets and striations on D.
davidsoniae leaves; G, cuticle striations on adaxial leaf surface of D. botryoides; H, unicellular hairs on
adaxial leaf surface of D. atropurpurea; I, hairs on adaxial surface of D. burkei leaf; J & K, Drimiopsis
burkei adaxial papilla at increasing magnification; L, adaxial view of D. maculata leaf margin. A, B, E,
H, J & L, Scale 1 cm = 40 JIDl;C, D, F, G, 1& K, Scale 1 cm = 10 JIDl.
41
I
The leaf epidermal cells of taxa of Resnova (Figure 3.3 A-C) and Ledebouria (Figure
3.3 D-E) studied differ from those of Drimiopsis. Resnova cells are, on average, of the
same length on both surfaces, i.e. an adaxial to abaxial ratio of 1:1 in terms of cell
length. Resnova (Figure 3.3 A-C) possesses shorter abaxial cells than Drimiopsis. The
1 ratio of abaxial to adaxial cells in Ledebouria is also 1 1.
:
' Resnova
species studied all have epidermal indumentum. Resnova humifirsa and R.
lachenalioides (Bak.) v.d. Menve (Figure 3.3 I) possess indumentum similar to that
seen on D. burkei (Figure 3.3 I). Figures 3.3 J-K show R. maxima short hairs. Hairs and
papillae appear in few Ledebouria taxa like L. asperifolia. Young Ledebouriinae
leaves, if papillose, are uniformly so on both adaxial and abaxial surfaces. With age,
papillae tend to become more frequent or confined to the adaxial surface.
Resnova abaxial and adaxial surfaces have uniform ovate stomata (Figure 3.3 A-C, F).
The 'H' complex in this group is indistinct. Ledebouria stomata, in contrast, are ovate
and raised, forming protrusions on the epidermal layer (Figure 3.3 WE). The stomata
are more numerous in Resnova and Ledebouria than in Drimiopsis. As in Drimiopsis,
the stomata distribution is denser on the abaxial than on the adaxial surface in both
Resnova and Ledebouria.
Cell outline and orientation
The outer periclinal cell walls of both abaxial and adaxial surfaces in Drimiopsis are
predominantly non-tabular with a convex shaped periclinal wall curvature. Howeve1
some are domed to conical shaped. The cell boundaries are channelled and the
anticlinal epidermal wall boundaries are for the most part irregularly curved. However,
the epidermal cells along the leaf margins differ from those towards the centre of the
lamina by being more isodiametric. In addition, taxa with undulate or crenulate leal
margins also possess epidermal cells with V-undulate anticlinal wall boundaries as
found in for example and D. maculata (Figure 3.2 L). The cells of the lamina are
longitudinally aligned.
I
-
Figure 3.3: SEM views of leaf epidermal surfaces: stomata and indumentum characters 0: A & B, abaxial
and adaxial surfaces of R. maxima, respectively; C, abaxial leaf margin of R. megaphylla; D & E, abaxial
and adaxial leaf surfaces of L. inquinata, respectively; F, adaxial leaf surface of R. schlechteri also
revealing the cuticle and wax characters; G, L. ovatifolia adaxial leaf surface with smooth wax and raised
stomata; H, L. inquinata adaxial leaf surface with wax rodlets and raised stomata; I, R. lachenalioides
adaxial leaf surface hairs; J & K, adaxial papilla on R. maxima leaf at differing magnifications. Scale bar
A, B, C, D, E, J & L, 1 cm = 40 1JDl;F, G, H, I & K, 1 cm = 10 1JDl.
43
- -
-
In terms of the Ledebouriinae, the outer periclinal cell walls of the adaxial and abaxial
leaf surface in Resnova and Ledebouria are non-tabular with a convex shaped periclinal
wall curvature. In Resnova the cell boundaries are poorly delimited or channeled and in
Ledebouria channeled. The anticlinal epidermal wall boundaries in Resnova are straight
to irregularly sinuate. In Ledebouria the anticlinal wall boundary is obscured to an
extent by a wax cover but can be described as elongate-tetragonal (Figure 3.3, D-E &
G-H). The anticlinal wall boundary of the epidermal leaf margins of Resnova is
predominantly straight, but V-undulate margins have been observed in R. maxima
(Figure 3.3 C ) . In the Ledebouria taxa studied the aforementioned margins are straight
(Figure 3.3 L) or they can also be V-undulate (Venter, 1993).
Cell sculpturing
The wax particles on all the taxa studied exhibit rnicromorphological character
diversity. However, the difficulties in describing these particular characters and
intraspecific variation exclude them from W h e r analysis, although Christensen &
Hansen (1998) maintain that they are genetically fixed and not the result of cell
exudates. The striate cuticle on Drimiopsis epidermal cells is generally irregularly
orientated. In some species it is more transversely striate than longitudinally striate, or
vice versa. Cells around the stomata are either transversely or irregularly striate
(Figures 3.2 WE). Some Drimiopsis davidsoniae leaves possesses a wax crust cover
(Figure 3.2 E) or wax crystals (Figure 3.2 F).
Cell sculpturing in the sister genera exhibit similar characteristics to that found in
Drimiopsis. The cuticle on Resnova epidermal surface is longitudinally striate.
Ledebouria generally possesses irregular striations. The Resnova taxa studied possess a
superficial layer of wax platelets (Figure 3.3 F). The Ledebouria taxa investigated
possess either coiled rodlet wax crystals (Figure 3.3 D, F) or thin wax as in L. ovatifolia
(Bak.) Jess. Venter (1993) also mentions the presence of wax platelets in Ledebouria.
3.4.3 ANATOMY
Drimiopsis atropurpurea (Figure 3.4 A) and D. comptonii (Figure 3.4 D ) with three and
two vascular bundles respectively evidence the typical parallel venation expected in this
group. The adaxial epidermis differs from the abaxial epidermis (Figures 3.4 A, C, E)
in that the former possesses a thicker cuticle and the cells are narrower, more compact
and vertically aligned. Some leaves seem to form a secondary epidermal layer that is
slightly differentiated from the mesophyl (Figure 3.4 A, C and G). The abaxial
epidermal cells are wider, more irregularly shaped and loosely arranged. The sparse
rod-shaped stomata are more numerous on the abaxial surface (Figure 3.5 D-F). The
substomatal chamber varies in size, even on the same leaf.
The chlorenchyma cells between the abaxial and adaxial epidermis are undifferentiated
but consist of uniform mesophyll aerenchyma cells (Figures 3.4 A-H). This situation
exists in vertically oriented leaves that catch sunlight on both surfaces (Metcalfe &
Chalk, 1950). Spongy mesophyll cells, loosely arranged with intercellular spaces, are
the site for photosynthesis and are thus common in horizontally oriented leaves. D.
atropurpurea leaves have numerous crystals (Figures 3.4 A-B). The D. pusilla lamina
trichomes (Figure 3.4 G) are larger than those towards the base of the leaf (Figure 3.4
HI*
Calcium oxalate star shaped crystals and raphides were observed in D. atropurpurea
(Figure 3.4 A) and D. pusilla (Figure 3.4 G and H). Crystals are less concentrated in D.
comptonii. Mucilaginous substances are evident inside the cells in Figures 3.4 A-C; G-
H and Figure 3.5 D. Parenchyma cells are interspersed with mucilage free cells. A
typical example in this group is the presence of a stomata1 antechamber, an adaptation
to water conservation. Anatomical characters were not coded for analysis in the data
matrix, as their reliability is not established.
The anatomy of the two sister genera display the parallel venation (Figure 3.5 G). The
adaxial epidermis is similar to the abaxial epidermis (Figures 3.5 B, C, and G). In
addition, the cuticle is thinner and the epidermal cells are slightly more differentiated
and larger when compared to Drimiopsis. The sparsely spaced stomata are more
concentrated on the abaxial surface. Like those of Drimiopsis, they are rod-shaped with
an-antechamber (Figures 3.5 D and F). No crystals were observed in the mesophyll of
- - -
-
-
-
-
-
-
-
- - - -
-
-
-
-
-
-
- - - -
-
-
-
-
-
-
- - - -
- - - - - -
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
either taxon although Venter (1 993) reports that they are common in Ledeboici.ia.-
- -
-
- - -
8)~
1.,,:,..<r \~"
!
I
Figure 3.4: Leaf anatomy of Drimiopsis taxa. A & B, D. atropurpurea leaf base sections showing
vascular bundles, star shaped calcium oxalate crystals (arrow) and inner tissue of uniform
mesophylI celIs. The adaxial and abaxial epidermis celIs differ from one another; C, adaxial
epidermis of D. comptonii leaf; D, abaxial epidermis of D. comptonii leaf with stomata; E & F, D.
pllsilla leaf sections with uniform mesophylI celIs and rod shaped stomata wit antechamber; G & H,
D. pusilla leaf sections with uniform mesophyII celIs, star shaped calcium oxalate crystals, uniform
mesophyll cells, rod shaped stomata, antechamber and hairs.
46
~ml
"
'v
/
...
f
,'""
120pmr
-s: ~-.....
-~
Figure 3.5: Anatomy of the leaves of Resnova and Ledebouria taxa. A, leaf vascular bundles of a
Resnova species; B, abaxial leaf surface of a Resnova species; C, adaxial leaf surface of a Resnova
species; D, stomata on a leaf of Resnova species; E, vascular bundle of a leaf of a Ledebouria species;
F, stomata of a Ledebouria species; G, vascular bundles and stomata of a Ledebouria species. The
stomata are more frequent than on Drimiopsis leaves in Figure 3.4.
47
.
3.5 PHENETIC ANALYSIS
The phenogram (Figure 3.6) based on 18 Drimiopsis taxa (Table 3.1) produces two
primary clusters. Cluster (A) houses southern African taxa with leaf indumentum. The
exceptions to (A) are D. kikiae and D. queae that have no leaf surface indumentum.
Schizocarphus nervosus is in cluster (Al) of Resnova taxa and L. asperifolia, which
possesses spotted lamina with hairs. Taxa in (A2) possess hairy leaves (except D. kikiae
and D. queae) and are pseudopetiolate. Cluster (A3) consists of taxa with sessile and
papillate leaves, except D. woodii that is pseudopetiolate.
Cluster (B), taxa without leaf indumentum, houses southern African Ledebouria and
tropical African Drimiopsis taxa. The exceptions here are D. carri and D. stolonissima
that possess fleshy leaves. D. maculata is of both southern and tropical African
distribution. The fleshy leaved L. concolor, included in this group, is unique within
Ledebouria in having leaves that are without threads when tom.
The phenogram does not support subspecific ranking currently in use. Drimiopsis
burkei (=D. burkei subsp. burkei) and D. stolonissima (=D. burkei subsp. stolonissima)
are housed separately in clusters (A3) and (Bl) respectively. Similarly, D. botryoides
(=D. botryoides subsp. botryoides) groups separate from D. perfoliata (=D. botryoides
subsp. prostrata).
3.6 CONCLUSION
Observations of plants growing in the field and those growing in the botanical garden
confirm that the overall shape of Drimiopsis leaves is not influenced by the
environment and is thus a good diagnostic character. This is important, as plants in the
field are frequently found without inflorescences.
The leaf epidermis in Drimiopsis, unobscured by wax or indumentum, displays
- - - - - - - - - - - - - - -
- - - - - - -
- - - - - - - -
-
- - - - - - - -
- - - - - - -
- -
taxonomically significanttepideei5a1cell arrangement, epiderrnal cell morphology and
-
stomata characters. Anticlinal walls are mostly S-type sinuate. The convex, tabular cells
possess an overall isodiametric-polygonal shape. The adaxial surface has distinct
isodiametric-polygonal cells, the abaxial tetragonal and elongate. The cell boundary
walls are irregularly curved, and the margins channelled sinuate. In the context of taxa
studied, taxa displaying surface indumentum, i.e. secondary sculpture, seemingly do
not have tertiary structure, i.e. epicuticular secretions.
l lpl
S. nervosus
R . l R.a humc ifusa
~
n
~
~
~
~
L. asperifolia
0.atropurpurea
D. liniopapilla
D. kikiae
D.queae
-
A2
t
A3
D . woodii
D. barteri
D. fischeri
D . rosea
D. b otryoides
D. stolonissima
L. concolor
D . pe rfoliafa
0.carrii
D. maculata
L. floribunda
L. inguinata
L. ovatifolia
L. revoluta
L. sandersonii
L. socialis
-
I
-
-
1
B1
B2
I
.
.
Linkage Distance
Figure 3.6: Phenograrn based on data supplied in Table 3.3 of 18 Drimiopsis, 4 Resnova and 8
Ledebouria species, producing two primary clusters (A) and (B) primarily based on presence or
absence of leaf surface indumentum.
Sparse stomata, more abundant on the abaxial than on the adaxial side, form a
diagnostic 'H' complex with subsidiary cells in Drimiopsis. The stomata in all plants
examined in the field are less frequent than those growing in the uniform environment
of the botanical garden. However, stomata size, and the length and width of the
stomata, has similar ranges in all taxa. Wax and cuticular characters vary even within a
given taxon and are thus not reliable taxonomic characters.
Anatomical characters in the Ledebouriinae, other than to highlight the morphological
range of adaxial and abaxial epidermal cells, are unreliable. The epidermal wall cell
range and the uniform mesophyll disprove the hypothesis that adaxial and abaxial cells
differ according to leaf orientation. This implies the two characters and their states, in
this group, are more genetically than environmentally influenced. The presence of
substances, and variations on epidermal cells, could be adaptations to stressful
environmental conditions
The phenetic groupings of subspecies question the validity of previously recognised
s
botryoides and D. burkei subsp.
subspecies in this genus. Drimiopsis b o ~ o i d e subsp.
burkei are housed in different clusters, separate from their sister taxa. Barring phenetic
limitations in classification, the subspecies deserve a species status as the degree of
variation between subspecies is comparatively similar to that between other taxa.
Keeping the shortcomings of phenetics in mind, the phenetic analysis of leaf characters
clearly demarcates the three genera from one another. The results also reveal that
tropical African Drimiopsis has more in common with Ledebouria than with Resnova.
The southern African Drimiopsis similarly has more in common with Resnova than
with Ledebouria. These results raise questions about views in support of sinking
Resnova under Drimiopsis (Phillips, 1% 1; Jessop, 1970, 1972; Dyer, 1976; Arnold &
De Wet, 1993; Meyer and Williams, 1997) and sinking both Resnova and Drimiopsis
under Ledebouria (Manning et al., 2004).
3.7 Key to species of Drimiopsis based on leaf characters.
Leaf margin entire ..................................................................................................................................... 2
Leaf margin crenate ................................................................................................................................... 6
Leaf margin undulate.. ......................................................................................................................... 10
Leaf margin crenulate ..................................................................................................................D. pusilla
Leaf cordiform...........................................................................................................................................3
Leaf falcifonn ..................................................................................................................... D. botryoides
Leaf linear .................................................................................................................................................4
Leaf spathulate .......................................................................................................................D. comptonii
Diphyllous; leaves sessile, appressed to the ground; lamina unspotted.................................. D. perfoliaia
Polyphyllous; leaves pseudopetiolate, erect; lamina spotted ............................................ D. atropurpurea
Leaves erect, 0.5 to 2 cm wide, margin edged; lamina unspotted ............................................................ 5
Leaves spreading, 3 to 4 cm wide, margin bordered; lamina spotted ................................. D. stolonissima
Monophyllous sometimes diphyllous; leaf more than 20 cm long; lamina thick ........................ D. barteri
Polyphyllous; leaf 6 to 10 cm long; lamina membranous .......................................................... D. facheri
Monophyllous...................................................................................................................................... 7
Diphyllous ............................................................................................................................................. 8
Polyphyllous ..........................................................................................................................................9
Leaf 1 to 5 cm long, 0.5 to 2 cm wide; lamina purple tinted abaxially ........................................ D. queue
Leaf 6 to 10 cm long, 3 to 4 cm wide; lamina purplish streaked abaxially ......................... D. liniopcpilla
Leaves spreading, linear, lanceolate ...........................................................................................D. burkei.
Leaves appressed to the ground, cordiform, ovate ........................................................................ D. carrii
Leaf cordifom, pseudopetiolate; lamina spotted ........................................................................ D. kikiae
Leaf linear, sessile; lamina unspotted ................................................................................. D. davidsoniue
Leaf 1 to 5 cm long.......................................................................................................................D. rosea
Leaf 6 to 10 cm long............................................................................................................................. 11
Leaf 11 to 20 cm long..............................................................................................................D. maculata
1l(10). Leaves lanceolate, 3 to 4 cm wide, sessile, margin bordered ................................................. D. reilleyana
Leaves ovate, 0.5 to 2 cm wide, pseudopetiolate, margin edged ................................................. D. woodii
4. THE FLOWER
Traditional taxonomy has always placed greater importance on reproductive characters
due to their resilience to plasticity. Flower morphology characters have been used in the
past to delimit taxa within the Ledebouriinae U. & D. Miiller-Doblies (Baker, 1896 &
1898; Van der Menve, 1946a, 194613; Jessop, 1970, 1972; Venter, 1993; Stedje, 1994;
Miiller-Doblies & Muller-Doblies, 1997; Speta, 1998b; Williams, 2000).
The unfertilized flowers in the Ledebouriinae, like most South African Hyacinthaceae
(Speta, 1998), fall off shortly after blooming. They are hypogenous, actinomorphic,
sextepalous, and pedicellate, shortly pedicellate or sessile (Speta, 1998, Stedje, 1994,
Williams, 2000). The perianth in Drimiopsis Lindl. & Paxt. is dimorphic because the
tepals of one whorl tend to be wider than the other. The inner perianth segments of, for
example, D. botryoides Bak. (=D. botryoides Bak. subsp. botryoides) (Figure 4.1 A &
B), D. burkei Bak. (=D. burkei Bak. subsp. burkei) (Figure 4.2 A) and D. rnaculata
Lindl. & Paxt. (Figure 4.2 B) have been reported as being wider than the outer ones and
the opposite for D. barteri Bak. (Figure 4.1 C & D). Noteworthy is the labeling of taxa
in Figure 2 of Stedje (1994: 47) that should read 2a & b, D. botryoides and 2c & d D.
barteri. The group L referred to in text and illustration (Stedje, 1994: 46) included D.
botryoides, with inner perianth segments wider than the outer. Group R, D. barteri, was
characterized as with inner segments narrower than the outer (Figure 4.1). The Resnova
v.d. Menve (Figure 4.2C) and Ledebouria Roth (Venter, 1993) perianth segments are
isomorphic.
Venter (1993: 32), in the context of Ledebouria, opines: "Colour is, in most cases, not
of taxonomic importance." Yet, Speta (1998) used, amongst others, perianth colour in
delimiting Drirniopsis from Resnova (whitish or greenish, in the former and pink-purple
or greenish brown in the latter), and Ledebouria (greenish sometimes tinged white or
' This chapter has in part been accepted for publication (Lebatha & Buys, 200%)
'Part of this chapter is in prep., Lebatha, Buys & Smit (2005).
pink). Muller-Doblies & Muller-Doblies (1997) also use colour to separate Drimiopsis
taxa. Tepals are viewed to be predominantly green in D. stolonissima (U. & D. MullerDoblies) Lebatha (=D. burkei Bak. subsp. burkei U. & D. Muller-Doblies), D.
reilleyana U. & D. Muller-Doblies, D. maculata and D. woodii Bak. as opposed to
pinkish to purplish in D. davidsoniae U. & D. Muller-Doblies, D. pusilla U. & D.
Miiller-Doblies, D. comptonii U. & D. Muller-Doblies and D. atropurpurea N.E. Br.
Although a thorough analysis of flower colour in the case of Resnova has not been
done, it is noteworthy that Van der Merwe (1946) delimits Resnova fkom other
Hyacinthaceae on the basis of, amongst others, perianth segments "never being blue or
purple" (translation mine).
Figure 4.1 : Dimorphic perianth segments. A & C, inner, ligulate segments of D. botryoides and
D. barteri Bak, respectively; B & D, elliptical outer ones of D. botryoides and D. barteri
respectively (adapted from Stedje, 1994: 47).
Figure 4.2: The opened flowers showing dimorphic perianth segments and androecium of A, D.
burkzi Bak. subsp. burkzi; B, D.maculata Lindl. & Paxt.; C, Resnova humifusa (Bak.) U . & D.
Miiller-Doblies. (Miiller-Doblies & Miiller-Doblies, 1997: 60)
The androecia in the Ledebouriinae are sexstaminate, commonly filiform to deltoid or
ribbon-shaped as well as basally epitepalous (Speta, 1998). Drimiopsis filaments have
been described as deltoid (Stedje, 1994; Miiller-Doblies & Muller-Doblies, 1997) and
fiee or united at the base (Stedje, 1994). The stamens in Drimiopsis are implanted at the
throat of the perianth tube (Figures 4.14.3). The perianth tube in the flower of D.
fischeri (Engl.) Stedje differs fiom that of other Drimiopsis taxa in being long, hence
the higher insertion in Figure 4.3 (Stedje, 1994; Muller-Doblies & Muller-Doblies,
1997). The filaments in Resnova (Figure 4.2 C ) are lanceolate ('filiform') and biseriate
while those in Ledebouria are filiform and, as in Drimiopsis, uniseriate (Jessop, 1970;
Muller-Doblies & Miiller-Doblies, 1997).
Figure 4.3:Drimiopsis Jischeri flower (form Muller-Doblies & Muller-Doblies, 1997: 63) with
the stamens inserted in the throat of a long perianth tube.
The Ledebouriinae possess tricarpellate, trilocular gynoecia with septa1 nectaries and
two basal ovules per locule (Baker, 1896; Jessop, 1970, 1972; Stedje, 1994; MullerDoblies & Muller-Doblies, 1997; Speta, 1998; Williams, 2000). Baker (1896) described
the ovaries of Drimiopsis as globose-trigonous and Jessop (1972) those of Drimiopsis
and Resnova as oblong. Jessop (1972), only using herbarium material of Resnova,
perceived these states taxonomically insignificant.
The inflorescence in the Hyacinthaceae is usually a simple raceme, rarely a spike
(Baker (1896) described Drimiopsis inflorescences as a subspicate raceme). Scapes are
prominent (Speta 1998). The presence, shape and structure of the bracts and prophylls
are variable in the family. Prophylls are here considered the modified leaves at the base
of pedicels. Bracts, in turn, are any additional modified leaves found on the
inflorescence. In terms of the Hyacinthoideae, Speta (1998) reports bracts to be usually
small, or lacking, or rarely large. Prophylls may be present or absent. Speta (1998)
asserts bracts and prophylls to be lacking in Drimiopsis and Resnova, and either small
or lacking in Ledebouria. Whenever authors refer solely to bracts, one is never sure
whether they distinguish between bracts and prophylls as Speta (1998) has done. In the
light of this, Baker (1896), Miiller-Doblies & Muller-Doblies (1997) and Kativu (2002)
report the presence of vestigial bracts in Drimiopsis. Williams (2000), in turn, reports
the presence of minute bracts in Drimiopsis. Stedje (1994), on the other hand, considers
them lacking. To confuse matters more, Muller-Doblies & Muller-Doblies (1997: 61)
also refer to basal bracts reduced to crescent shaped "gibbosities." These seemingly
incongruent reports on the occurrence of "bracts" are indicative of the above-mentioned
uncertainties and should be interpreted with caution.
Although Jessop (1972) reports the inflorescences of Drimiopsis, Resnova and
Ledebouria to be similar, differences in bracts and pedicel length have been
documented (Speta, 1998; Venter, 1993; Muller-Doblies & Miiller-Doblies, 1997).
4.2 OBJECTIVES
To investigate the systematic value of floral characters and their states in Drimiopsis
and sister taxa and supply a key to species based on floral characters.
4.3 MATERIALS and METHODS
Table 4.1 lists the plants investigated including the species names established in this
thesis. Fresh inflorescences were collected in the field and stored in 70% ethanol and
then examined with either a SEM or light microscope. The flowers were excised fiom
their inflorescences and then subjected to critical point drying. Flowers of D. maculata,
R. lachenaliodes (Bak.) v.d. Merwe and L. ovatifolia (Bak.) Jessop, stained with cresyl
. . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
violet acetate, were wax-embedded and processed thiough standard microtome
- -
-
procedure for anatomical study. Morphological terms follow Radford et al. (1974).
Bracts and prophylls in this study were confined to the mature inflorescence. Some
Ledebouria data was obtained from Venter (1993).
(A data matrix based on Table 4.2 was subjected to a cluster analysis using
S T m s T I c A 6.1 with the following settings: tree clustering, Ward's method of
/minimum-variance clustering under the amalgamation rule
and percentage
disagreement as a measure of distance. A floral character-based key was generated via
I DELTA.
1
4.4 RESULTS and DISCUSSION
The data matrix of flower characters is provided in Table 4.2.
I
4.4.1 INFLORESCENCE
Ledebouriinae young inflorescence is scapose, fleshy, erect, axillary and indeterminate
(Figure 4.4). The majority of species of Drimiopsis possess a solitary inflorescence,
except in D. barteri and D. liniopapilla Lebatha where two inflorescences per bulb can
I
occur. Mature Drimiopsis inflorescences all possess vestigial bracts except in D.
fzscheri where the bracts generally fall off.
1
Contradictory reports on the presence of spikes and racemes in Drimiopsis (Sfedje,
1
1994; Stedje & Thulin 1995) could be explained by the fact that some inflorescences
have minutely pedicellate flowers (Figure 4.4 A, D, F). The terminal and upper flowers
1
are minutely pedicellate and the lower flowers shortly pedicellate. Drimiopsis flowers
range from minutely pedicellate (< 0.1 cm long) in D. barteri Bak., D. botryoides, D.
1
1
pegfoliata Bak., D. burkei, D. stolonissima, D. carrii Lebatha, D davidsoniae U. & D.
Miiller-Doblies, D. liniopapilla Lebatha, D. reilleyana U. & D. Muller-Doblies and D.
rosea A. Chev.; to shortly pedicellate (0.1-0.3 cm long) in D. atropurpurea N.E. Br.,
D. kikiae Lebatha, D. maculata, D. pusilla and D. queue Lebatha; and pedicellate (> 0.3
1
cm long) in D. woodii Bak., D. comptonii U. & D. Miiller-Doblies, and D. fzscheri
I
(Engl.) Stedje. The latter condition also occurs in the Resnova and Ledebouria taxa
studied.
Table 4.1: Taxa whose flowers were dissected and investigated.
Taxa
Accepted name in
this thesis
Accession
number:
D. atropurpurea
D. atropurpurea
D. barteri
D. botryoides
D. botryoides
D. pefloliata
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D. stolonissima
D. cawii
D. comptonii
D. davidoniae
D. jhcheri
Lebatha 048
Lebatha 049
Lebatha 002
Lebatha 098
Lebatha 003
Lebatha 00 1
Lebatha 009
Lebatha 041
Lebatha 095
Lebatha 040
Lebatha 046
Lebatha 054
Lebatha 056
Lebatha 103
Lebatha 037
Lebatha 0 15
Lebatha 079
Lebatha 038
Fischer 325
Locality
R. transvaalensis
Charles Craib, South Africa
Luneberg, South Africa
Iringa Distr., Ruaha, Tanzania
Uzaramo District, Tanzania
Kiarnbo District, Kenya
Iringa District, Tanzania
Potchefstroom, South Africa
Parys Dam, South Africa
Rasesa, Botswana
Kosi Bay, South Africa
Vaal River, South Africa
Reitvlei, South Africa
Waterberg, South Africa
Kgale Hill, Botswana
Strydom Tunnel, South Africa
Durban, South Africa
Mbabane, Swaziland
Pilgrim's Rest, South Africa
Precise
locality
unknown,
Tanzania
Louwsburg, South Africa
Lebatha 045
Lebatha 053
South Africa
Lebatha 0 19
Durban, South Africa
Transkei, South Africa
Hallack s.n.
Bezeia Mts., South Africa
Baur 549
Clydesdale, South Africa
Tyson 2878
Lebatha 060
Roossenekal, South Africa
Lebatha 005
Gaborone, Botswana
Gaborone, Botswana
Lebatha 006
Lebatha 007
Soweto, South Africa
Parys Dam, South Africa
Lebatha 039
Lebatha 033
Mtunzini, South Africa
Lebatha 02 1
Durban, South Africa
Lebatha 03 1
Mtunzini, South Africa
Lebatha 062
Mkanga, Swaziland
Lebatha 102
Eastern Cape, South Africa
Lebatha 047
Mandini, Swaziland
Lebatha 078
Mbabane, Swaziland
Waterberg, South Africa
Cebatha-055Lebatha 068
MkhaJg S w a i j m &
Chevalier 8432 Laboum, Belgian-Congo
Natal, South Africa
Wood 774
Saunders s.n.
Natal, South Africa
Sutherland s.n. Natal, South Africa
Killick, Marais Umtata, South Africa
2063
Lebatha 088
Roossenekal, South Africa
van der Menve Amsterdam, South Africa
1889-1 1-36)
van der Merwe Ermelo, South Africa
R. transvaalensis
L. aspenyolia
L. inquinata
Devenish 958
Lebatha 050
Lebatha 0 10
atropurpurea
atropulpurea
barteri
botryoides subsp. botryoides
botryoides subsp. botryoides
botryoides subspprostata
burkei subsp. burkei
burkei subsp. burkei
burkei subsp. bwkei
burkei subsp. burkei
burkei subsp. burkei
burkei subsp. burkei
burkei subsp. burkei
burkei subsp. burkei
burkei subsp. stolonissima
cawii
comptonii
davidroniae
D. kikiae
D.liniopapilla
D. lachenalioides
D. lachenalioides
D. lachenalioides
D. lachenalioides
D. liniopapilla
D. maculata
D.maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maxima
D. reilleyana
D. rosea
D. saundersiae
D. saundersiae
D. woodii
R. lachenalioides
D. kikiae
D. liniopapilla
R. lachenalioides
R. lachenalioides
R. lachenalioides
R. lachenalioides
D.liniopapilla
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
R. maxima
D. pusilla
D. -qzleaeD.reilleyana
D. rosea
R. humifusa
R. humifusa
D. woodii
R. lachenalioides
R. megaphylla
R. transvaalensis
R. megaphylla
R. humifusa
- - - - - - -
- - - -
9
- - - - - - - -
s.n.
L. inquinata
Piet Retief, South Africa
Piet Retief, South Africa
Molepolole, Botswana
57
-
L. ovatifolia
L. sandersonii
Schizocarphus newosa
L. ovatifolia
L. sandersonii.
S. newosa
Lebatha 008
Lebatha 059
Codd 73 1
Gaborone, Botswana
Pilgrim's Rest, South Africa
South Africa
The number of flowers ranges from eight to fifty per inflorescence and is inconsistent
within species. Age and aestivation is usually accompanied by a lengthening of the
rachis. Drimiopsis comptonii (Figure 4.4 C ) possesses a unique 'simple coryrnb-like'
raceme that in reality represents a compressed rachis. Inflorescences can be tight with
flowers having a conical shaped apical head, as in D. burkei (Figure 4.4 A) and D.
pusiNa (Figure 4.4 D). They may forrn a loosely packed inflorescence with flowers
sparsely distributed on the rachis in D. botryoides (Figure 4.4 B), D. maculata and D.
perfoliata. These latter inflorescences have long scapes with cylindrical racemes. This
type of inflorescence is also common to Resnova species e.g. R. maxima v.d. Menve
(Figure 4.4 E), R. pilosa V.D. Menve (Figure 4.4 F-G) and R. lachenaliodes (Bak.) v.d.
Menve.
Drimiopsis inflorescence is erect except in D. botryoides (Figure 4.4 B) and D.
maculata with spreading inflorescences. Typically the mature scape becomes more than
twice as long as the leaves, resulting in a spreading inflorescence. Drimiopsis
comptonii, D. fscheri, D. queae, D. rosea, D. woodii, D. perfoliata and D. stolonissima
possess inflorescences much longer than the leaves. Inflorescence is approximately as
long as leaves in D. atropurpurea, D. barteri, D. burkei, D. davidsoniae, D. kikiae, D.
liniopapilla, D. pusilla and D. reilleyana.
Inflorescence characters alone are inadequate for species delimitation as some taxa, for
example D. maculata, D. burkei, D. stolonissima, D. perfoliata and D. b o t ~ o i d e scan
sometimes display both patterns. Young Drimiopsis inflorescences usually have buds
arranged densely and tightly in a conical shape as in D. pusifla (Figure 4.4 D). The
- - - - - - -
-
- -
- - - - - - - - - - - - -
peduncle is mostly uncO1Oiiid t h ~ i s it
h -is6iiiiikd-in -D atriipurptcrea and Dcwaodii;
.
spotted in D. burkei and D. kikiae; and purplish coloured in D. liniopapilla, D. pusilla,
D. queae and D. rosea.
#1. Inflorescence <arrangement> 1. one to two per bulb, 2. several per bulb; #2. Inflorescence <type> 1. a simple raceme, 2. a simple-corymb-like raceme; #3. Inflorescence
<posture> 1. erect, 2. spreading; #4. Inflorescence <length> 1. shorter than leaves, 2. more or less as long as leaves, 3. considerably longer than leaves; #5. Inflorescence
with <number of flowers> 1. 15 flowers or less, 2. 16 to 30 flowers, 3. more than 30 flowers; #6. Inflorescence with <flower density> 1. flowers sparsely distributed, 2.
flowers densely distributed; #7. Flower <pedicel length> 1. minutely pedicellate (shorter than 0.1 cm), 2. shortly pedicellate (0.1 to 0.4 cm long), 3. elongated pedicel (more
than 0.4 cm long); #8. Rachis <general length> 1. 10 cm or shorter, 2. 10.1 to 20 cm long, 3. more than 20 cm long; #9. Rachis <shape> 1. cylindrical, 2. conical, 3. ovoidcylindrical; #lo. Peduncle <variegation> 1. banded, 2. coloured (usually purplish), 3. spotted, 4. green; #11. Bracts in mature inflorescence <presence or absent> 1. absent,
2. vestigial, 3. developed; #12. Prophylls <present or not> 1. absent, 2. present; #13. Flowers <size> 1. minute (1-2 mrn long), 2. small (2.1-4 mm long), 3. medium-sized
(4.1-6 mm long), 4. large (more than 6 mm); #14. Flowers <type> 1. actinomorphic; #15. Flowers <number of tepals> 1. sextepalous; #16. Flowers <shape> 1. coronate to
stellate, 2. campanulate, 3. tubular; #17. Flowers hypanthium base <shape> 1. truncate, 2. obtuse, 3. rounded; #18. Tepals <type> 1. isomorphic, 2. dimorphic; #19. Tepals
with hypanthium <hypanthium size> 1. inconspicuous, 2. conspicuous; #20. Tepals <colour> 1. whitish to greenish, 2. purplish green, 3. creamy-brownish, 4. pink, 5.
purplehlue; #21. Outer whorl of tepals <posture> 1. connivent, 2. recurved, 3. drooping; #22. Outer whorl of tepals longitudinally <posture> 1. cucullate, 2. flat; #23. Outer
whorl of tepals apically <apex margin shape> 1. conduplicate, 2. flat; #24. Inner whorl of tepals <posture> 1. connivent, 2. recurved, 3. drooping; #25. Inner whorl of tepals
longitudinally <posture> 1. cucullate, 2. flat; #26. Inner whorl of tepals apically <apex margin shape> 1. conduplicate, 2. flat; #27. Vitta, 1. conspicuous, 2. faint, 3. absent;
#28. Androecium <number> 1. 6, #29. Androecium <colour> 1. greenish to whitish, 2. cream, 3. maroonish/purplish; #30. Androecium <posture> 1. erect, 2. spreading;
#3 1. Androecium <perianth adnation> 1. epitepalous; #32. Androecium <arrangement on tepal> 1. uniseriate, 2. biseriate; #33. Adroecium <insertion on tepab 1. inserted at
throat of perianth tube, #34. Androecium <length> 1. shorter than pistil, 2. as long as pistil, 3. longer than pistil; #35. Filaments <cohesion> 1. free, 2. valvate; #36.
Filament <shape> 1. deltoid to accuminate, 2. lanceolate, 3. filiform; #37. Anther <attachment> 1. dorsifixed; #38. Gynoeciurn <number of carpels> 1. tricarpellate, 2.
polycarpellate; #39. Ovules <number per locule> 1. two per locule, 2. more than two per locule; #40. Stigma <shape> 1. roundish, 2. triangular; #41. Stigma papilla <type>
1. stalked, 2. subsessile, 3. sessile; #42. Stigma papilla <shape> 1. round, 2. trilobal; #43. Style <size>l. shorter than ovary, 2. as long as ovary, 3. longer than ovary; #44.
Style <shape> 1. terete, 2. triangular; #45. Ovary <attachment> 1. sessile, 2. stipitate; #46. Ovary <shape> 1. globose, 2. ovoid to oblong, 3. conical; #47. Ovary
transversely <shape> 1. smooth, 2. with ridge below style, 3. severally lobed; #48. Ovary <colour> 1. whitish, greenish, 2. purplish, 3. bluish; #49. Ovary shoulders <present
or absent, 1. absent, 2. present; #50. Nectaries <absent or present> 1. absent, 2. present; #5 1. Flowering time, 1. March to May, 2. September to December, 3. July to
August.
Q
Figure 4.4: Ledebouriinae inflorescences. A, D. burkei with minute flowers; B, D. botryoides
developing inflorescence; C, D. compton;; corymb-like raceme; D, the coloured and hairy D.
pusilla peduncle; E, shortly pedicellate flowers of R. maxima; F & G, typical isomorphic
spreading flowers of R. pilosa; H, flaccid inflorescence of a Ledebouria sp. (Photos A-D; F-G
by U. Mtiller-Doblies).
61
--------
Ledebouria and Resnova flowers are borne on ascending pedicels. Ledebouria
inflorescences are lax, rarely solitary, commonly more than two per bulb (Figure 4.4
H). Bracts and prophylls are often present in mature flowers. The pedicels are the
longest in Ledebouria, up to 7 cm in L. concolor, L. inquinata and L. sandersonii.
Resnova has solitary inflorescences with flowers possessing pedicels 0.1-0.2 cm long
(Figure 4.4 E).
4.4.2 PERIANTH
Perianth segments in Ledebouriinae are sextepalous and inconspicuous due to their
small stature. They may also be dull coloured in Drimiopsis and Resnova. Drimiopsis
tepals are dimorphic with at least the inner segments permanently connivent (Figure 4.5
& 4.7). The inner tepals are conspicuously cucullate and wider than the moderately
cucullate outer tepals (Figure 4.8 A-G). The tepals of Resnova (Figure 4.4 E-G; Figure
4.6 A) and Ledebouria (Venter, 1993) are isomorphic (Figure 4.6 E) with the inner and
outer segments similar in shape and size. Some Ledebouria taxa, e.g. L. aperfolia
(Bak.) Jessop and L. floribunda (Bak.) Jessop, possess cucullate tepal apices (Venter,
1993), but most are flat (Figure 4.6 E).
The vitta is mostly conspicuous in Drimiopsis and distinctive on Resnova and
Ledebouria tepals. In terms of tepal length, Drimiopsis varies fiom 0.2-1.0 cm,
Resnova 0.5-1.8 cm and Ledebouria 0.5-2.2 cm. Flower size is generally consistent
within species. Only Drimiopsis possesses apical tepal margins that are conduplicate
(Figure 4.4 A, B & D; Figure 4.5 A & F).
It is evident that individual flower and bud colour are not significant taxonomic
characters in the Ledebouriinae. A survey of living and herbarium material of
Drimiopsis demonstrates intraspecific variation in perianth colour. For example, an
analysis of D. maculata records reveals 63% of specimens possessing white flowers, a
further 34% with green flowers and 3% with yellowish or cream flowers. 40% of D.
burkei subsp. burkei specimens possess white flowers, 17% green and 43% include
cream, pink or purple flowers. Records also show R. maxima green, white and redbrown and R. lachenaliodes, pink, mauve or blue.
Figure 4.5: Drimiopsis flower morphology. A, D. maculata sectioned flower with inner connivent and
outer cucullate tepals, the globose and sessile gynoecium; B, D. maculata sessile globose gynoecium with
the style as long as the ovary; C & D, D. burkei stigma showing the corrugated, trilobed and sessile
stigmatic papilla; E & F, D. maculata deltoid filaments attached to the base of the perigone segment, as
broad as the segments.
63
--
-
-
-- -
--
Figure 4.6: Flower morphology. A, isomorphic tepals with biseriate androecium and a lanceolate filament
of R. maxima; B, shortly stipitate ovary of Resnova sp. with ovarian ridges showing just below where the
style and ovary merge; C, the round shortly stalked stigmatic papilla of Resnova; D, L. revoluta stigmatic
papillae; E, the Ledebouria filiform filament and the flat longitudinal posture of the tepal apex; F, the
stipitate conical shaped ovary of Ledebouria revoluta.
64
-
- - - --
-
-----
Figure 4. 7: The globose sessile ovaries of Drimiopsis taxa. A, D. macu/ata flower opened perianth
showing the deltoid filaments; B, C and D, D. macu/ata, D. liniopapilla and D. carrii flowers buds
respectively, with tepals removed to reveal the pistil; E, F and G, the dimorphic inner and outer
segments of D. botryoides, D. queae and D. kikiae (similar to D. liniopapilla) respectively.
65
---
-
---
--
--
-----
.
.
.
.
.
.
,;
Figure 4.8: The stipitate ovaries of Resnova taxa showing variously pronounced ovarian ridges at the
base of the style. A-B, R. maxima; C, R. humifusa (=D. saundersiae); D, R. pilosa; E, R. schlechteri;
F, R. transvaalensis.
66
--
--
-
-
..
.
4
.
~
< ~'~"
~.,
"
.)
."
( ,.
J
.
l ..
*
$.
~
.
-::...
.
-.
.0;"
..'
...
.
-. ~..
:~
~
..
.
'"
.
Figure 4.9: Anatomical sections of D. maculata flower. A, longitudinal section illustrating the rounded
hypanthium base and sessile ovary whose base possesses a protrusion; B, transverse section of the
completely syncarpous trilocular ovary with one aborted ovule; C, longitudinal section of D. maculata
flower showing an inconspicuous perianth tube and androecium attachment; D, transverse section of the
rounded style, style canals and uniseriate filaments.
67
--
-
.
.
! ,(1) }I
~
. J. ~
1I
.
ft
I
II
J '~....
r. -"
)'
;
t
,
\
l
1
~'1I1
~
~
h~~
.~{~.
~
""
...~
...
,.
..
.-
Figure 4.10: Anatomicalsections. A-C, L. ovatifolia:A, longitudinalsection showing a truncate shaped
hypanthium base and stipitate ovary; B, cross section revealing three carpels, septal slits and septal nectarines; C,
cross section revealing the triangular shaped style and uniseriate filaments (also visible in B); D-F, R.
lachenalioides: D, longitudinal section through an obtuse shaped hypanthium base with a shortly stipitate ovary
(arrow) and septal slits; E, cross section revealing the ovoid shaped style and the filaments in two series (also
visible in F); F, three carpels, septal slits with septal nectaries in cross section.
68
--
--
--
-
-
--
-
-
-
-
-
--
---
--
4.4.3 ANDROECIUM
Drimiopsis, Resnova and Ledebouria each possess six stamens with dorsifixed anthers.
In Drimiopsis and Resnova the stamens are exclusively epitepalous (fused to the
tepals). In Ledebouria these may be either epitepalous or free (Venter 1993).
Drimiopsis and Resnova possess erect stamens, whereas they may either be erect,
patent or connivent in Ledebouria. The stamens of Drimiopsis are more or less equal in
length and possess deltoid filaments (Figure 4.5 A, E & F; Figure 4.7 A, E-F). The
base is as broad as the tepal in most of the species. Some of the filament bases, like in
D. maculata, D. burkei and D. botryoides subsp. botryoides, are valvate (Figure 4.7 C).
In Resnova the lanceolate stamens are of unequal lengths (biseriate), the one alternating
with the other (Figure 4.6 A). In Ledebouria the filiform stamens (Figure 4.6 E) appear
to be either equal or unequal in length (Venter, 1993). In all three genera the filaments
are flattened to a greater or lesser degree at the base where the ovary rests (Figure 4.5 A
& E). It is well documented that stamen length relative to pistil length can vary
depending on the developmental stage of the flower in terms of male and female phases
(Weberling, 1989). Fully matured flowers of all three genera possess stamens that are
as long as the pistil.
4.4.4 GYNOECIUM
The pistils of all Ledebouriineae are tricarpellate and conduplicate i.e. possessing a
longitudinal groove (Figure 4.5 A & B; Figure 4.6 B & E). Drimiopsis possesses terete
styles, Resnova and Ledebouria triangular. Drimiopsis taxa possess globose and sessile
ovaries (Figures 4.5 A & B; Figure 4.7 B, D-G). All Resnova ovaries studied are ovoid
to oblong in shape and possess a short stipe (Figure 4.6 B; Figure 4.8 A-G). The
aforementioned observation of a stipe in Resnova is novel (Lebatha & Buys 200%).
The stipes vary in length as seen from dissected fiesh and herbarium material (Figure
4.8). Ledebouria in turn possesses conical and conspicuously stipitate ovaries (Figure
4.6 F). The shoulder of the ovaries (sensu Venter, 1993) vary from exclusively tapering
into the style in Drimiopsis and Resnova to either tapering into, or being rectangular to,
- - - -
- - - -
- - - - - -
- - - - - - - - - -
- - - - -
or raised in relation to the base o f t h e style i n L a i e b ~ ~ Venter's
~ r n ~ 41993) work
contains some errors, e.g. in couplet no. 32 of the key Venter (1993: 78) mentions that
L. ensifolia (Eckl.) S. Venter does not possess shoulders, whereas in the full description
thereof (Venter 1993: 90) he mentions the presence of rectangular shoulders. It is only
Ledebouria taxa that possess expanded basal lobes. Nectaries, when present, occur in
the basal lobes. Resnova in turn, solely possesses ridges on the shoulders (Figure 4.6 B;
Figure 4.8 A-F; Miiller-Doblies & Muller-Doblies, 1997: 60, Fig. 2g).
The style in Drimiopsis is as long as the ovary (Figure 4.5 A-B; Figure 4.7 A-G). In
Resnova (Figure 4.6 B) these are shorter than the ovary, and in Ledebouria (Figure 4.6
F), longer. Exceptions in Resnova are D. saundersiae Bak. (Figure 4.8 C), synonym of
R. humifusa (Bak.) U . & D. Muller-Doblies, and R. transvaalensis v. d. Menve (Figure
4.8 F), previously D. lachenalioides (Bak.) Jessop. The stigma of Drimiopsis taxa
investigated has corrugated, trilobed sessile papillae (Figures 4.5 C & D). The Resnova
stigmatic papillae are rough and round (Figure 4.6 C) and those of Ledebouria smooth
and round (Figure 4.6 D).
The anatomy of the flower accentuates the tricarpellate and syncarpous nature of the
ovaries, and the character states differences in Drimiopsis, Resnova and Ledebouria.
Anatomy of the flower of D. maculata reveals the rounded base of the hypanthium
(Figure 4.9 A). The Drimiopsis ovary has a protrusion (Figure 4.9 A & C) that might be
rudimentary, a remnant of a once stipitate ovary. The completely syncarpous trilocular
ovary has one aborted ovule (Figure 4.9 B). The cross section done just above the ovary
shoulders shows the round shape of the style and style canals (Figure 4.9 D). The
deltoid shape and uniseriate arrangement of the filaments on the perianth is displayed
on Figure 4.9 D.
Ledebouria ovatifolia ovary via longitudinal sectioning, displays the truncate shaped
hypanthium base and a stipitate ovary (Figure 4.10 A). Ledebouria possesses septal
I
slits, where septal nectaries can develop (Figure 4.10 B). The triangular shape of the
style is apparent, together with the uniseriate arrangement of the filiform filaments
(Figure 4.10 C & B). Resnova lachenalioides ovary, with a narrower obtuse shaped
hypanthium base, has a short stipe (Figure 4.10 D), an ovoid shaped style (Figure 4.10
F) and filaments arranged on the perianth in two series (Figure 4.10 E & F). Three
,
1
are lanceolate are displayed in Figure 4.10 F. Ledebouria and Resnova ovaries differ 1
carpels, septal slits, septal nectaries and the ovoid shaped filaments that, at lateral view
from those of Drimiopsis (Figure 4.9 B ) in being partially syncarpous (Figure 4.10 B &
F).
4.5 PHENETIC ANALYSIS
A phenogram (Figure 4.1 1) computed with 5 1 Ledebouriinae floral characters (Table
4.2) produced two primary clusters: a Resnova, Ledebouria cluster and a Drimiopsis
cluster. The Drimiopsis cluster groups southern African taxa and tropical African taxa
that include southern African D. comptonii, D. queue and D. woodii. Drimiopsis
barteri, D.maculata and D. rosea occur in both regions. The tropical African cluster
has plants with inflorescences considerably longer than leaves and greenish-white
tepals, except for D. comptonii that has purplish flowers. Drimiopsisfischeri differs by
having tubular flowers and a stipitate ovary. The southern African taxa cluster divides
into two major groups of shortly pedicellate flowers with pinkish to creamy brown
tepals and basally minutely pedicellate taxa with whitish-greenish flowers respectively.
The phenetic grouping, unlike in Figure 3.5, supports subspecific ranking of D.
botryoides subsp. botryoides and D. botryoides subsp. prostrata. It does not however
support the validity of subspecific ranking of D. burkei subsp. burkei and D. burkei
subsp. stolonissima.
The Ledebouria cluster with Schizocarphus nervosus and Resnova. Thus, Resnova
share more character states with Ledebouria than with Drimiopsis. This result raises
questions about views in support of sinking Resnova under Drimiopsis (Phillips, 1951;
Jessop, 1970, 1972; Dyer, 1976; Arnold & De Wet, 1993; Meyer and Williams, 1997)
or of sinking Drimiopsis under Ledebouria (Manning et al., 2004). Given the fact that
the presence of stipitate ovaries in Resnova was included for the first time, an additional
analysis excluding this character was done with the same results. These results only
show similarity groupings.
S narvosus
L aspanfdra
L ovatndta
L rnqurnaia
L. sandersonri
L flmbunda
L rewluta
L concolor
L. socralrs
R maxrma
R lachenalro~des
R humrfusa
R megaphylla
D atmpupurea
D. davrdsonrae
D pusdla
D burker
D krkrae
D. barten
D. liniopapilla
D. stdonissima
D. cam7
D. reiflayena
D. rosee
D. botryoides
D. pMoIieta
D. comptonri'
D. maculata
D. queee
D. woodii
D. Ifscheri
0
1
2
3
4
5
6
Linkage Distance
Figure 4.1 1: Phenogram based on analysis of 51 floral characters (Table 3.2) in
Drimiopsis, Resnova and Ledebouria using Ward's Method of minimum-variance
clustering and percentage disagreement as a measure of distance.
4.6 CONCLUSION
On their own, interspecific flower characters are not significant. This concurs with
Jessop's (1972) and Manning et al. (2004) assumption that these states are
taxonomically uninformative. The key to species of Drimiopsis based on floral
characters demonstrates this (Section 4.7). However, distinctive intergeneric variation
exists. Tepal colour groupings and not individual colour can assist in delimiting
Drimiopsis.
Investigations into the anatomy of three Ledebouriinae taxa confirm the recorded
morphological characters and their states. The character states of sessile or stipitate
ovaries, shape and arrangements of the filaments and the shape of the style are
established.
Drimiopsis botryoides subsp. prostrata and D. botryoides subsp. botryoides cluster
together supporting their subspecific ranking. The subspecific ranking in D. burkei
subsp. burkei and D. burkei Bak. subsp. stolonissima is not supported.
The phenetic analysis of conventional flower characters in the Ledebouriinae, with the
addition of a stipitate ovary in Resnova, provides sufficient data to demarcate Resnova,
Ledebouria and Drimiopsis. In addition, Resnova clusters with Ledebouria.
4.7 Key to species of Drimiopsis based on floral characters.
Peduncle banded ......................................................................................................................... 2
Peduncle coloured purplish .........................................................................................................
3
Peduncle spotted .........................................................................................................................5
Peduncle not coloured .................................................................................................................7
Inflorescence k as long as leaves, flowers shortly pedicellate;
rachis <10 cm long ............................................................................................D. atropurpurea
Inflorescence considerably longer than leaves, flowers shortly pedicelate; rachis 10-20
cm long .....................................................................................................................D. woodii
Rachis cylindrical .......................................................................................................................4
Rachis conical ...............................................................................................................D. pusilla
Rachis ovoid cylindrical .......................................................................................D. liniopapilla
Flowers sparsely distributed, shortly pedicellate, rachis <10 cm .................................. D. queue
Flowers densely distributed, minutely pedicellate, rachis 10-20 cm long....................... D. rosea
Inflorescence solitary, flowers sparsely distributed, rachis conical ............................................6
Inflorescence one to two per bulb, flowers densely distributed, rachis
ovoid cylindrical ..........................................................................................................D. barteri
Flowers minutely pedicellate; tepals whitish to greenish ............................................. D. burkei
Flowers shortly pedicellate; tepals creamy-brownish ....................................................D. kikiae
Flower minute, 1-2 mm long ...................................................................................................8
Flower small, 3-4 mrn long ................................................................................................10
Flower medium-sized, 5-6 mm long ...................................................................... D. maculata
Flower regular sized, >6 mm long .............................................................................. D. fischeri
Inflorescence shorter than leaves; rachis cylindrical ......................................................D. carrii
Inflorescence more or less as long as leaves; rachis conical.......................................................9
Inflorescence considerably longer than leaves; rachis ovoid cylindrical................ D. comptonii
Flowers sparsely distributed, tepals creamy-brownish ........................................ D. davidsoniae
Flowers densely distributed, tepals whitish to greenish .......................................... D. reilleyana
Flowers sparsely distributed; rachis long, cylindrical; tepals fused at base .............................. 11
Flowers densely distributed; rachis short, conical; tepals free............................ D. stolonissima
1l(10). Inflorescence erect; vitta conspicuous .....................................................................D. perfoliata
Inflorescence spreading; vitta faint .......................................................................D. botryoides
5. PALYNOLOGY
5.1 INTRODUCTION
'
Aa member of the order Asparagales (Dahlgren, 1980), the Hyacinthaceae possesses
monosulcate pollen grains (Dahlgren et al., 1985; Watson & Dallwitz, 2000). Pollen
morphology has been found to be of diagnostic value at all levels of the taxonomic
hierarchy (Stuessy, 1990). Zhang & Anderberg (2002) used palinology at the family
level in the Cyrillaceae while Molina et al. (2002) and Orozco (2001) used it to delimit
species in the Rubiaceae and Cunoniaceae respectively. However, pollen has never
been a popular source of data in the Hyacinthaceae, as it is believed taxonomically
insignificant at lower levels (Speta, 1998b; Pfosser & Speta, 1998). Within the
Ledebouriinae, only Ledebouria Roth pollen has been described to a small degree
(Venter, 1993).
5.2 OBJECTIVES
The aims of this investigation were to assess the interspecific and the intergeneric
variation of pollen morphology in Drimiopsis Lindl. & Paxt., Resnova v.d. Merwe and
Ledebouria.
5.3 MATERIALS and METHODS
Table 5.1 lists the plants investigated. The plant accession numbers are for the NorthWest University, Potchefstroom Campus Botanic Garden. Muri and lumen
measurements are based on averages of four readings.
Unacetylised pollen grains of 21 Drimiopsis, 6 Resnova and 5 Ledebouria taxa were
obtained from fresh flowers, air dried, carbon coated and gold plated for viewing with
Philips SL30DX 4i SEM at 10 kv. Samples were viewed at similar magnification with
the whole grain at 3 000 X and ornamentation at 12 000 X magnification. Terminology
Data was subjected to a cluster analysis using STATISTICA 6.1 with the following
settings: tree clustering; Ward's method of minimum-variance clustering under the
This chapter has been accepted, with modifications, for publication: Lebatha & Buys (2005b).
amalgamation rule and percentage disagreement as a measure of distance. A data
matrix was created based on Table 5.2, but excluding characters that coded
polymorphic for one of the three genera, i.e. 6 characters were analysed.
Table 5.1 : Specimens examined with pollen dimensions.
Average
equatorial
diameter
Taxa
Accession
no.
Locality
D.atropurpurea
D.atropurpurea
Lebatha 049
Lebatha 048
Luneberg, South Africa
Charles Craib, South Africa
Lebatha 003
Kiambo District, Kenya
Lebatha 04 1
Lebatha 054
Lebatha 056
Lebatha 095
Lebatha 103
Parys Dam, South Africa
Rietvlei, South Africa
Waterberg, South Africa
Rasesa, Botswana
Kgale Hill, Botswana
Strydom Tunnel, South
Africa
Mbabane, Swaziland
Pilgrim's Rest, South Africa
Vaal River, South Africa
Durban, South Africa
Mtunzini, South Afi-ica
Durban, South Africa
Mtunzini, South Africa
Mkanga, Swaziland
Graharnstown, South Africa
Mbabane, Swaziland
Roossenekal, South Africa
Mkhaja, Swaziland
South of Piggs Peak, South
Africa
Duiwels-kantoor,
South
Africa
Waterberg, South Africa
Durban, South Africa
Durban, South Africa
Piet Retief, South Africa
Roossenekal, South Africa
Roossenekal, South Africa
Waterberg, South Africa
Kwelera River, South Africa
Mandini. Swaziland
51.4
Average polar
axis
(pm)
(pm)
D. botryoides
botryoides
D. burkei
D. burkei
D. burkei
D. burkei
D. burkei
D.
burkei
stolonissima
D.comptonii
D.davidsoneae
D. kikiae
D. maculata
D. maculata
D.maculata
D. maculata
D. maculata
D. maculata
D. pusilla
D. queae
D.reilleyana
subsp.
subsp.
Lebatha 037
Lebatha 079
Lebatha 038
Lebatha 046
Lebatha 039
Lebatha 033
Lebatha 021
Lebatha 03 1
Lebatha 062
Lebatha 102
Lebatha 078
Lebatha 060
Lebatha 068
L. asperifolia
Lebatha 080
L. inquinata
Lebatha 075
L. ovatifolia
Ledebouria sp.
Ledebouria sp.
Ledebouria sp.
Resnova sp.
Resnova sp.
Resnova sp.
R. maxima
R. maxima
Lebatha 063
Lebatha 0 16
Lebatha 0 18
Lebatha 050
Lebatha 05 1
Lebatha 088
Lebatha 052
Lebatha 042
Lebatha 047
40.2
47.9
16.1
22.1
Table 5.2: Pollen characters and states used in a phenetic analysis within the
Ledebouriinae: A = Drimiopsis, B = Resnova and C = Ledebouria.
Character
Character state
Character coding
A
1 Pollen
grain
shape,
equatorial view
2 Pollen grain shape, polar
view
3 Pollen grain shape, lateral
view
4 Pollen grain types
depressed ovate = 0; ellipsoid = 1
0
elliptic = 0; narrowly elliptic = 1
0
blunt = 0; tapering = 1
0
monosporous = 0; heterosporous 0
=1
0
5 Pollen grain equatorial subequiaxe = 0; breviaxe = 1
diameter
0
straight = 0; curved = 1
6 Pollen distal pole
7 Pollen grain ornamentation punctate = 0; reticulate = 1; 0
punctate-reticulate = 2.
5.4 RESULTS and DISCUSSION
The monosulcate pollen grains of all three genera almost become zonasulcate. Viewed
equatorially, Drimiopsis pollen is depressed ovate (Figure 5.1 A & D) and that of
Resnova and Ledebouria ellipsoid (Figure 5.1 G & J). From a polar view, Drimiopsis
pollen is elliptic (Figure 5.1 B & E) while those of Resnova (Figure 5.1 H) and
Ledebouria are narrowly elliptic-harmomegathy
notwithstanding. Drimiopsis pollen
is subequiaxe and possesses a linear pole opposite to the sulcus as well as blunt lateral
sides (Figure 5.1 A & D). The pollen in Resnova and Ledebouria is breviaxe and
possess a tapered pole opposite to the sulcus as well as tapered lateral sides (Figure 5.1
G, J-L). Heterospory in Ledebouria (Figure 5.1 J-L) is due to the pole opposite the
sulcus tapering to a varying degree (fiom an equatorial view) so that the pollen
sometimes resembles either those of Drimiopsis or Resnova.
In terms of ornamentation, Drimiopsis possesses punctate (pitted) pollen (Figure 5.1 M)
- - - -
- - - -
and R&noi;
alnd-~edibouria-ieti-cu~ate
-(Figare-5.1~ jand
- punctuate-reticulate P O I ~
(Figure 5.1 0)respectively.
The larger Drimiopsis pollen grains are represented by D. rnaculata Lindl. & Paxt.
(Figure 5.1 A-C), with an average equatorial diameter of 45.56 pm and a polar axis of
24.84 pm (ratio 1.8: 1). D. burkei Bak. (Figure 5.1 D-F), representing the smaller
grains, has an average equatorial diameter of 26.6 pm and a polar axis of 18.76 pm
(ratio 1.42:1). Muri in Drimiopsis are (0.3 17-)l. 101-1.7(-1.85) pm wide while lumina
are (0.88-)0.2-0.342(-0.507)
pm across. Resnova pollen (Figure 5.1 G-I) has an
average equatorial diameter of 5 1.7 pm and a polar axis of 23.58 pm (ratio 2.19: 1). R.
maxima (Figure 5.1 N ) has muri of (0.244-)0.3-0.404(-0.837) pm wide and lumen
(0.4-)0.504-0.895(-
1.150) pm across. The equatorial diameter of Ledebouria pollen
studied averages 55.35 pm and the polar axis averages 23.2 pm (ratio 2.39:l). The
Ledebouria lumen is (0.404-)1.1-1.39(-2.45)
pm across. The muri is (0.288-)0.3 17-
O.598(-1.39) pm wide.
5.5 PHENETIC ANALYSIS
A phenetic analysis, run with the parameters as set out above, clustered Resnova to
Ledebouria (Figure 5.2). This result is contrary to current thinking, which sinks
Resnova under Drimiopsis (Phillips, 1951; Jessop, 1970, 1972; Dyer, 1976; Arnold &
De Wet, 1993; Meyer and Williams, 1997; Kativu, 2000).
5.6 CONCLUSION
On the ground of pollen morphology, Resnova has more in common with Ledebouria
than with Drimiopsis.
Lebatha and Buys (2005b) have also obtained the
aforementioned grouping of Resnova and Ledebouria with an analysis of an expanded
matrix consisting of 27 characters pertaining to the flower (Chapter 4). However, a
bigger sampling, especially of Resnova and Ledebouria, is needed to validate this
conclusion.
Figure 5.1: (A-C) Pollen of D. macu/ata (Lebatha 019)-equatorial, polar and lateral view respectively;
(D-F) Pollen of D. burkei (Lebatha 056)-equatorial, polar and lateral view respectively; (G-I) Pollen
of R. maxima (Lebatha 042)-equatorial, polar and lateral view respectively; (J-L) Equitorial view of
heteromorphous pollen in three Ledebouria sp. (Lebatha 050, Lebatha 018, Lebatha 016 respectively);
(M-O) Punctate (pitted) pollen in D. burkei (Lebatha 056), reticulate pollen in R. maxima (Lebatha
042) and punctuate-reticulate pollen in Ledebouria sp. (Lebatha 050) respectively. Scale
= 20
JlIIl
78
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!
i
Resnava
Ledebouria
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1.1
1.2
Linkage Distance
Figure 5.2: Cluster analysis of six (non-polymorphic) pollen characters (Table 5.2) in
Drimiopsis, Resnova and Ledebouria using Ward's method of minimum-variance clustering
under the amalgamation rule and percentage disagreement as a measure of distance.
79
--
6.1 INTRODUCTION
Chemotaxonomic data, even fragmentary, are useful in taxonomy though few have
solved taxonomic problems on their own (Harbome, 1984; Hadacek, 2002). Chemical
data have served mostly as a complementary source of data in support of classical
morphologically-based taxonomy (Quicke, 1996). Phytochemical screening and
structure elucidation are predominantly the domain of researchers in the field of natural
medicinal products. Thin Layer Chromatography (TLC) remains a popular technique
(simple, affordable, sensitive, fast and selective) for separating mixtures of compounds
(Harbome, 1983, 1998; Wagner et al., 1984; Dewick, 2000; Wagner and Bladt, 2001).
Qualitative phytochemical studies, evaluation of chromatograms, together with Rf
values can reveal homologous marker compounds. Rf values though are dependent on
several variables that give a 5 to 10 % error expectation (Randerath, 1966; Hamilton &
Hamilton, 1987).
Although the majority of chemotaxonomic studies entail determining the distribution
patterns of a specific compound (Tadesse & Abegaz, 1990; Abegaz
et
al., 1991;
Buschmann & Spring, 1997; Spring et al., 1999; Viljoen & Van Wyk, 1999; Ahmed et
al., 2001; Viljoen et al., 2001, 2002; Jensen et al., 2002), classes of compounds, with
known or unknown molecular structures, can also be taxonomically significant. For
example, naphthoquinones are confined to the Juglandaceae L.; and hydroxyphenolic
acids are confined to Lycopodium L. (Harborne, 1983, 1984). Many alkaloids and
related compounds occur only in specific groups of plants. Ricinine is an example of a
compound so far found only in the genus Ricinus L. (Budavari, 1996). Flavonoids,
though ubiquitous in plants, have been used successfully in revealing hybridization in,
for example, legumes based on data derived from 2-dimensional chromatography
(Harbome, 1983, 1998).
According to Dewick (2000), secondary plant metabolites are:
". ..an expression of the
individuality of a species.. ." and ". ..are not necessarily produced under all conditions".
Secondary metabolites are important to plant survival as, for example, herbivore
deterrents, anti-microbial agents or even as scents for attracting pollinators. The
Liliaceae Juss., sensu lato, contain among others, "veratrine" alkaloids, which consists
of cevadine, veratridine, devadilline, sabadine and cevine (Wagner & Bladt, 2001).
Convallaria majalis L ., Urginea maritima L ., Drimia maritima (L.) Steam, and Drimia
indica (Roxb) Jess. contain cardiac glycoside cardenolides (Wagner & Bladt, 2001).
The Hyacinthaceae have subsequently been delimited into five subfamilies mainly on
chemotaxonomic characters: Chlorogaloideae Speta (North America); Oziroeoideae
Speta
(Andean
South
America);
southem
African
Urgineoideae
Speta;
Ornithogaloideae Speta; and Hyacinthoideae Speta (Speta, 1998b; Pohl et al., 2001).
Three major groups of compounds viz. homoisoflavones, steroids and cardiac
glycosides (bufadienolides and cardenolides) have been identified in the Hyacinthaceae
to date (Speta, 1998; Pohl et al., 2000, 2001; Koorbanally et al., 2001). The marker
compounds for the Hyacinthoideae are homoisoflavones consisting of four basic
structural types namely : a) 3-benzyl-4-chromanone, b) 3-benzyl-3-hy droxy -4chromanone, c) 3-benzylidene-4-chromanone (Speta, 1998, Pohl et al., 2000) and d) the
scillascilins (Koorbanally et al., 2001) containing an additional fourth ring. Eucosterol
derivatives could possibly also be marker compounds for the Hyacinthoideae (Pohl et
al., 2001). The Hyacinthaceae do not synthesise a-homonojjirimycin (a-HN.J) but
Hyacinthus orientalis L. does, thus questioning its placement in the family (Kite et al.,
1998). Hyacinthaceae subfamily Urgineoideae possesses bufadienolides while
subfamily Omithogaloideae is characterised by the presence of cardenolides.
Chemotaxonomic data on Drimiopsis are wanting. Although Dahlgren et al. (1985)
mention that steroidal saponins are common in the Asparagales, Gibbs (1974) notes
their absence in Drimiopsis. He also reports small quantities of tannins and hydrogen
- - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - - -
cyanide in the leaves and shoot of Drimiopsis kirkii ~ a k . - a%ell
s a s leiidbanthocyanins
in D. kirkii and D. maculata Lindl. & Paxt. Hutchings (1989a, 1989b) recorded the
presence of mucilaginous exudates in D. maculata. Koorbanally et al. (2001) isolated
scillascillins as well as homoisoflavones from D. maculata. Homoisoflavones have also
been recorded for Eucomis bicolor Bak. (Boehler & Tamrn, 1967), Scilla scilloides
Druce (Heller & Tarnm, 1981), Ledebouria graminifolia (Bak.) Jessop (Mutanyatta et
al., 2003), L. cooperi (Pohl et al., 2000) and L. ovatifolia (Bak.) Jess. (Pohl et al.,
2000). Sparg et al. (2002) found no saponins in L. ovatifolia but found the cardiac
glycoside bufadienolides (Urgineoideae chemical markers). Hitherto, nothing has been
recorded for Resnova.
6.2 OBJECTIVES
This chapter reports on the taxonomic significance of secondary metabolites identified
via two different techniques: 1. Phytochemical screening protocols (see below) of
Drimiopsis, Resnova and Ledebouria for the following classes of compounds:
alkaloids, anthocyanins, anthranoids, anthroquinones, cardenolides, coumarins,
flavonoids, leucoanthocyanins, polyphenols, tannins, saponins and steroids. 2. Two
chromatographic methods, viz. TLC and HPLC.
6.3 MATERIALS and METHODS
6.3.1 General
HPLC analyses were performed with a HPl 100 instrument fitted with a Diode ray
detector. A Stuart Scientific Orbital SO1 shaker was used for extraction. HPLC grade
solvents were used for HPLC analyses. Extractions were conducted using redistilled
GPR grade or Analytical Grade solvents used without purification. Solvents from
extracts were recovered with a Buchi Rotorvapour R-114 at 40' C at 320-340 mbar.
TLC samples were spotted on a 0.25 rnm thick layer of silica gel coated on alumunium
foil fiom Merck, Kiselgel 60254and Alugram Sil/GUV254(TLC) plates.
6.3.2 Plant material
All the plants used (Table 6.1) were initially collected in the field, planted in the
- - - - -
- - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - - - - -
Botanical Garden at the North-West University, ~ o t c h e f G o mCampus
and-thenharvested 18 months after cultivation to minimize the potential influence of the
environment. Bulk collection of D. burkei Bak. (Lebatha 095b) was made in March
2003 near Matsieng Foot Prints, Rasesa, Botswana.
Table 6.1: Dry yields obtained from bulbs and leaves harvested fLom the garden.
Voucher specimens are housed in the AP Goossens Herbarium (PUC).
Taxon
Accession No.:
Drimiopsis sp. 1.
Lebatha 048
Drimiopsis sp. 2.
Lebatha 053
D.atropulpurea
Lebatha 049
D. bobyoides
Lebatha 0 13
D. burkei
Lebatha 009
D. burkei
Lebatha 04 1
D. burkei
Lebatha 052
D. burkei
Lebatha 095
D. stolonissima
Lebatha 037
D.comptonii
Lebatha 079
D. maculata
Lebatha 0 12
D. maculata
Lebatha 104
D. maxima
Lebatha 036
D. pusilla
Lebatha 078
Ledebouria coopen'
Lebatha 050
L. ovatifolia
Lebatha 055
L. apert$?oIia
Lebatha 090
Resnova sp. 1.
Lebatha 088
Resnova sp. 2.
Lebatha 072
R. maxima
Lebatha 047
Total weight (g.)
6.3.3 Phytochemical screening protocols
The protocols followed in the investigation were adapted from Abegaz (1995). Fresh
plant material (bulbs and leaves) were cleaned with a brush to remove debris, chopped
into pieces, completely dried in an oven at 40-45" C and ground with a mortar and
pestle. The powdered material was kept refrigerated until required. The combined
protocols require at least 13.5 g of plant material. In some there was a paucity of plant
material and this required that the proportions of the various reagents be
correspondingly adjusted.
1. Alkaloids
Two grams of ground plant material was placed in a test tube and treated with 25 ml of
1% HCl for 15 minutes in a water bath. The suspension was filtered into a test tube and
the filtrate divided into 2 parts: A and B. Five drops of Dragendorff s reagent were
added to filtrate A. The formation of a precipitate was taken as indication for the
presence of alkaloids. To confirm the result, filtrate B was treated with saturated
Na2C03solution until a drop of the solution turned the universal indicator paper (UIP)
blue (pH 8-9). The filtrate was then mixed with 4 ml of chloroform. Two separate
layers formed. The upper layer (aqueous) was removed with a pipette and treated with
acetic acid until a drop gave a yellow-brown colour with IUP (pH 5). Five drops of
Dragendorff s reagent were then added. The formation of a precipitate was taken as
indication for the presence of quaternary alkaloids. The lower layer was treated with 2
ml of HCl. On separation of the latter layers, the upper was pipetted and treated with
three drops of Dragendorff s reagent. The formation of a precipitate was taken as
indication of the presence of tertiary alkaloids.
2. Anthocyanins
One gram of plant material was mixed with 15 ml of cold 1% HCl and boiled. The
colour during boiling was noted. Anthocyanins vary in colour from orange-red to bluered.
3. Anthranoids
A mixture of 5 ml 0.5 N KOH, 0.5 ml 5% H202 solution, together with 0.2 g of well-
ground plant material was boiled for two minutes, cooled and then filtered. The filtrate
was treated with 6 drops of acetic acid until the IUP coloured yellowish red. The
resulting solution was mixed with 5 ml of toluene; the upper layer (toluene) was
separated with a pipette, transferred to a test tube and 2 ml of 0,5 N KOH was added. A
red colour appearing in the aqueous layer indicated the presence of anthranoids.
4. Anthraquinones
One gram of ground plant material was macerated with 5 ml of CHC13 and 5 ml of ether
and then filtered. One ml of this solution was treated with 1 ml of 10% NaOH solution.
A red colouration was taken to indicate the presence of quinines.
5. Cardenolides (Cardiac glycosides)
Two grams thoroughly ground plant material was treated with 20 ml of distilled water
in a test tube and kept at room temperature for two hours. Four drops of Kedde's
reagent (4 ml of 3,5% dinitrobenzoic acid and 0,6 ml of 1 N KOH in methanol, to be
prepared extemporarily) were added to the filtrate. A violet blue colour was taken to
indicate the presence of cardenolides.
6. Coumarins
A small amount of moistened sample was placed on a filter paper moistened with 10%
NaOH solution and covered. The paper was exposed to W light for a few minutes.
Appearance of yellow green fluorescence was taken to indicate the presence of
coumarins.
7. Flavonoids
One gram of well ground plant material was extracted in 10 rnl water and 5 ml
methanol and then filtered. A few magnesium turnings were added to 3 ml of the
filtrate and concentrated. HCl was added drop wise. The development of orange, red or
pink colours were taken to represent flavones, flavonols and flavonones respectively.
8. Leucoanthocyanins
One gram of powdered sample was put in 20 rnl water; 5 ml of this mixture was mixed
with 2 ml of 2N HCl and heated on a water bath (100" C) for about 10 minutes. A slow
development of red to violet colour was taken to indicate the presence of
leucoanthocyanins. Digesting a small sample with 2N HCl in propanol for 15-30
minutes identified leucocytes. The slow development of a red to violet colour was taken
to be a positive reaction.
9. Polyphenols
One gram of plant material in 15 ml of distilled water was heated on a water bath (100"
C) for 15 minutes and the solution filtered. Three drops of fieshly prepared ferric
cyanide solution (1 ml of 1% FeC13 and 1 ml of 1% K3Fe(CN)6) was added to 2 ml of
the filtrate. The formation of the blue green colour was taken to indicate the presence of
polyphenols. The green to black colour, resultant fiom the 1% FeC13,is also positive for
tannins.
10. Tannins
Two grams well ground plant material in 15 ml water was heated on a water bath (100'
C) for 5 minutes. AAer cooling to room temperature, the mixture was filtered and
divided into three 3 ml portions: A, B & C.
To A, 5 ml of 2% NaCl solution was added and any suspension filtered off. Five ml of
1% gelatin was then added. As a reference, the standard solution of tannic acid (0,5% in
water) was used. Precipitation was taken to indicate the presence of tannins. To B, three
drops of 1 % &Fe(CN)6 and three drops of 25% NH3were added and the colour change
noted. To C, three drops of 1% FeCl3 solution were added. A green black coloration in
B and C was taken to indicates the presence of tannins.
NB. The polyphenol test (I) is also positive for tannins hence the separate tests.
11. Saponins
One gram of well ground plant material in 5 rnl water in a test tube was heated to 100°
C in a water bath for 15 minutes. The mixture was left to cool to room temperature then
filtered. 10 ml of the filtrate was poured into 100 x 10 mm test tube and shaken for ten
seconds. The height of the persistent hexagonal honeycomb froth was measured. Froth
higher than 1 cm would confirm the presence of saponins.
12. Steroids
One gram of powdered plant material was covered with 10 ml ether, placed on an
orbital shaker and shaken occasionally for 2 hours or continuously for one hour. The
solution was divided into two 1 ml solutions, A & B. Solution A was put on a porcelain
plate to evaporate. A drop of concentrated sulphuric acid was added and stirred and the
colouration noted. To solution B two drops acetic anhydride were added followed by
two drops concentrated sulphuric acid (Lieberman-Burchard test) and again coloration
was noted. The test was positive when coloration was seen with A or B.
6.3.4 Chromatography analysis
Extraction and isolation:
Two grams of ground plant material, in a 50 ml round bottom flask, was extracted using
25 ml of boiling methanol (in a water bath) for one hour. Evaporation of solvent during
extraction was avoided by attaching a reflux condenser to the extraction round-bottom
flask. The solvent was evaporated completely using a rotary evaporator fiom a tarred
flask. Weighing the evaporating flask and noting the differences obtained, the mass of
organic extract residue was determined. The residue was dissolved in a minimum
amount of methanol, and a portion of this sample was used for thin layer
chromatographic analysis.
Dried and powdered bulbs of D. burkzi (Lebatha 095b), weighing 400 g, were
successively extracted twice (12 hr each) with 200 ml methanol-methylene chloride
- - - - - - - - - -
- - - -
- - - -
- - - - - - - - -
- - - -
mixture and 200 ml of methanol for -threeeliours. me ~esidue-obtained- (20.8 g) b y -
evaporation at reduced pressure (T<40° C) was successively partitioned with hexane,
chloroform, ethyl acetate, butanol and water. The chloroform extract was further
subjected to vacuum liquid chromatography VLC using petroleum ether-ethylacetate
and ethylacetate-methanol mixture of varying concentrations. Fractions collected by
VLC gave mixtures, which were purified by several preparative thin layer
chromatographic procedures, employing chloroform-methanol mixture 9.6:0.4 as
developing solvent yielding compounds 1-7 (Table 6.3). Structure elucidation of these
compounds, five homoisoflavanones, one scillascillin and one xanthone, was based on
'H NMR,HMBC, HMQC, DEPT 135,13cNMR, NOESY, MS and W analyses.
The following solvent systems were used for thin layer chromatographic analysis: ethyl
acetate, methanol and water, 10:1.3:1 (System A); hexane and ethyl acetate 2: 1 (System
B); hexane and ethyl acetate 4: 1 (System C), and chloroform-methanol, 9.4:0.6 (System
D).
The seven compounds (Table 6.3) were isolated and l l l y characterized from a wild
sample of D. burkei (Lebatha 095b). This plant, because of its abundance, was used as
reference compounds in conducting the chromatographic analysis of crude extracts of
17 plants of the three genera (Drimiopsis, Resnova and Ledebouria).
The crude
extracts of various species of the genera (Table 6.1), together with the identified
compounds 1-7
were subjected to TLC, eluted with solvent System D. The
chromatogram obtained was detected by observing in visible and W light (254 nm)
and further visualized by spraying with vanillin sulfuric acid spray and subsequent
heating at 100' C until spots became clearly visible. Based on the retention factor (Rf),
observations were made to ascertain the presence of isolated compounds in other
species of the three genera.
The spot pattern, colour and distribution were noted and Rf values were calculated,
tabulated and compared. The process was repeated several times to ascertain uniformity
and validation.
HPLC
Individual compounds 1-7 and the crude extracts of various species (Table 6.1) of the
genera Drimiopsis, Resnova, and Ledebouria (6 mg dissolved in 200 p1 of methanol)
were analysed by HPLC. An X-Terra RP
18
column (3.0 x 250 mm, 5 pm) was used.
The mobile phase was water + 25 mM formic acid (A) and acetonitrile + 25 mM formic
acid (B) where gradient elution was employed with 25-80% of B at Temp 25". Flow
rate was maintained at 0.6 mllmin. and the wave length (A) used was 254 nrn.
6.3.5 Phenetic analysis
Data was subjected to a cluster analysis using STATISTICA 6.1 with the following
settings: Ward's Method of minimum-variance under the amalgamation rule and
percentage disagreement as a measure of distance. Two data matrices were created
using 13 characters from phytochemical data (1 1 OTUs) and 14 chemical (compounds
1-7) data characters (20 OTUs).
6.4 RESULTS and DISCUSSION
The plants analyzed are tiny in stature (height), ranging fiom 3 cm in D. burkei to 40
cm (5 cm lamina + 35 cm pseudopetiole) in D. maculata, thus the total dry yields were
low. Appendix 1 contains the Rf value chromatographs and tables.
6.4.1 Phytochemical screening tests
All the taxa screened (Table 6.2) show the presence of flavonoids, steroids and
polyphenols. The type of flavonoid however differs: D. maculata, L. apertifolia (Bak.)
Jess., L. ovatifolia (Bak.) Jess. and R. maxima v.d. Merwe contain flavones while the
Drimiopsis sp., D. burkei and Resnova sp 1. contain flavonones. Coumarins are found
in the majority of Drimiopsis and all the Resnova taxa screened, while alkaloids, traces
of anthroquinones and tannins are found only in R. maxima. Only the Ledebouria taxa
studied gave positive results for anthocyanins. Leucoanthocyanins are present in D.
maculata, L. apertifolia, L. ovatifolia and Resnova sp 1. Ledebouria and Resnova, in
turn, have in common the presence of saponins. Anthranoids and cardenolides appear to
be absent in the Ledebouriinae. Of the two cardiac glycosides bufadienolides and
cardenolides, Kedde's reagent (see phytochemical protocols) detected only the latter.
Drimiopsis maculata (Lebatha 104) showed a trace of alkaloids but failed the
confirmation test. The presence of other nitrogenous compounds in the plant can give a
positive result hence the two stage testing. The screened morphotypes of D. maculata
(Lebatha 012 and 104) and D. burkei (Lebatha 009, 04 1, 052 and 095) yielded similar
results.
6.4.2 TLC and HPLC analysis
Table 6.4 shows the retention factor (Rf) and retention time (RJ for compounds 1-7
(Table 6.3) obtained fiom TLC and HPLC analysis respectively. Compound 1, with Rf
of 0.26, appeared in all the extracts with the exception of D. botryoides Bak. (Lebatha
013). The same distribution pattern of the mentioned compound was also detected in
the HPLC chromatograms and appeared with a l& of ca 9.6 min (Figures 6.1 and 6.2).
Table 6.2: Results of phytochemical screening on seven Drimiopsis, two Resnova and
two Ledebouria taxa: 1(alkaloids), 2(anthocyanins), 3(anthranoids), 4(anthroquinones),
S(coumarins), 6(flavonoids), 7(leucoanthocyanins), 8(steroids), 9@olyphenols),
1O(tannins), 1 1(saponins), 12(cardenolides), n(flavone), nn(flavonone), -(absence),
+(presence). The presence of a compound is further qualified as: +(faint), ++(medium)
and +++(strong).
RESULTS
Taxon
Drimiopsis sp. 1
D. burkei
D. burkei
D. burkei
D. burkei
D. maculata
D. maculata
Ledebouria apertifol ia
L. ovatifolia
Resnova sp. 1 .
R. maxima
The five D. burkzi morphotypes (Figure 6.2) collected from five different localities
(Lebatha 009, 037, 041, 052, and 095) show identical results with respect to the TLC
and HPLC, except for compounds 2, 5, 6 and 7 (Table 6.4). Variation in the peak
intensity of some constituents was clearly observed (Figure 6.1a-d and 6.2a-d). The
'weak' detection of compounds 4 , 5 and 6 in the morphotypes of D. burkei is due to the
presence of other more abundant metabolites (not yet identified) with similar R( values.
HPLC generally detects the presence of compounds 1-7 better than TLC. However, the
opposite holds true for compound 6 in D. burkzi (Lebatha 009) and D. comptonii U. &
D. Muller-Doblies (Lebatha 079). It is noteworthy that an homoisoflavone isolated
from D. burkei (Lebatha 095b, Rf of 0.75) appeared as a bright pink spot when the
chromatogram was sprayed with vanillin-sulphuric acid, but was not observed again in
the crude extract, even though there were no other constituents in the region of the Rf
where it was observed.
Variations in the qualitative and quantitative composition of secondary metabolites are
not uncommon, and particularly in the Hyacinthaceae (Speta, 1998a, 1998b). This is
clearly noticed in the present study where compound 1 is the predominant component
in D. burkei (Lebatha 041), whilst conspecifics (Lebatha 095b and 052) contain more
metabolites that elute with low as well as high R, values.
Compound 1was detected in all samples analysed and is therefore a prime candidate as
a marker compound for the Ledebouriinae. Compounds 3 and 4 were detected by both
analytical methods in most of the samples (Table 6.4). The exceptions were extracts of
D. comptonii, D. maculata and D. pusiNa U. & D. Muller-Doblies where they were
absent. Also 3 was not detected by either method in two of the three species of
Ledebouria. Silascillin (5), not unexpectedly (Koorbanally et al., 2001), has a limited
distribution. This strained compound was found in all samples of D. burkei and one of
the two samples of D. maculata (Lebatha 012).
The scillascillin and
homoisoflavonoids isolated in this study differ from those previously isolated by Pohl
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
et al. (2000) and ~oorbanallyet all (2001j.-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
This study reveals that TLC and HPLC can, with caution, be used to profile plant
extracts. It is also possible to use compounds such as 1,3 and 4 as indicators of taxa. At
the same time one should recognize that the composition of secondary metabolites can
vary within species, depending on localities and time of collection. Although xanthones
are found in some of the species, it is not entirely clear if these substances are true
metabolites of the plants or products of other lower organisms (fungi and lichens)/
which may or may not have a defined relationship to the plants.
Table 6.3: Compounds 1-7 isolated from the chloroform extract of D. burkei (Wild
095). Compounds 1, 2, 3, 4 and 6 are homoisoflavonoids. Compound 5 a scillascillin
and compound 7, a xanthone.
CMPD R1
ocy
R2
R3
Table 6.4: Thin layer chromatography (TLC) and High pressure liquid chromatography
(HPLC) results for nine Drimiopsis taxa, three of Resnova and two Ledebouria. + =
observed on TLC chromatogram, d
=
observed on HPLC chromatogram,-is
absent
and R represents obscured.
Compound (STD)
Retention time (min)
Retention factor (cm)
D. burkei
D. burkei
D. burkei
D. burkei
D. stolonissima
D.comptonii
D. pusilla
D. sp.
D.atropurpurea
D. maculata
D. maculata
Drimiopsis sp. 2.
D. botryoides
D. maxima
Lede bouria apertfolia
L. cooperi
L. ovatifolia
Resnova maxima
Resnova sp. 1.
1
9.60
0.26
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5
6
7
11.40 15.20 16.02 16.72 17.21 22.14
0.36 0.40 0.32 0.50 0.52 0.56
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Figure 6.1: HPLC chromatograms for crude extracts of a, Drimiopsis maculata
(Lebatha 012); b, Ledebouria apertifolia (Lebatha 019); c, Drimiopsis botryoides
(Lebatha 013) and d, Resnova megaphylla (Lebatha 088).
96
a
8.488
I
mAU
10
eo
2.458
9.593
I
1.907
20.058
1':;~.~2.011
10 ~
.
,
5.2995.869
11.360
I 2.950
4.393
6.372
32903.9184.71!6
7.1408.003 9.313
.
13.562
16.~.7:k8.122
2O.92k.272
12.601
14.673 16~~
19.382 19.709
27.404
27.766
o ij
10
mAU -1
10~
15
20
25
mn
b
1.750
I
i
10
1.894
50
;
..
19.982
9.566
i
22.646
17.922
30'
20.836
4.656
2.735 4367
'3.932",5.2766.396
,
10"1
22.221
F '~g.748
7:-~~ 8.871
25.7142727~.652
O-L
10
15
2.
20
--
mn
C
mAU
2.435
.. -~
..
30
j
25~
1
20-;
j
151
10
j
.
19.971
4.628
9.550
11.314
20.828
17.879
2.741
22.198 22.646
25.720
23.137
18.678
27271
01
'o
mAU-;
10
20
2.
22.678
9.616
I
30;
I
mn
d
1.895
1.710
Ii
..
15
2.444
i
17.942
20.030
20.304
14.994
19.654 20.880 22.255
16.446
18.875
14.655'6.017
17.207
23.143
14.100
16.718
,
11.37912.247
10
O-t
{
10
"
20
2'
mn
Figure 6.2: HPLC chromatograms for crude extracts of Drimiopsis burkei morphotypes:
a, Lebatha 052; b, Lebatha 037; c, Lebatha 009 and d WILD 095.
97
6.5 Phenetic analysis
Patterns of similarity explored by a cluster analysis of the two data sets produced
conflicting results. The phenogram based on the Rf. values alone was taxonomically
uninformative as taxa tended to form one big cluster. The one based on phytochemical
screening data (Figure 6.3) separates Drimiopsis into two groups: Drimiopsis burkei
taxa with Resnova maxima, and Drimiopsis maculata with Ledebouria taxa and
Resnova sp. 1. Figure 6.4 derived from an analysis of data presented in Table 6.4,
separates Drimiopsis into two groups: one group accommodating all D. burkei
subspecies, and the other clustering with the Resnova and Ledebouria taxa analysed
Ledebouria apertifolia forms a cluster with the Resnova taxa analysed. Drimiopsi:
comptonii clusters with L. ovatifolia and L. cooperi. Of note here is the vegetativc
similarity between Resnova and L. apertifolia and the broad-leaved D. maxima Bak
and D. botryoides that group together.
6.6 CONCLUSION
In summary, it appears that Drimiopsis consists of two groups in respect of secondaq
metabolites: an exclusive Drimiopsis burkei group and a Resnova-Ledebouria cluster
The presence of unidentified metabolites with & values similar to those of 4, 5 and t
needs fkther investigation.
D. burkei subsp. burkei
D. burkei subsp. burkei
D, burkei subsp. burkei
D. burkei subsp. burkei
Drimiopsis sp.
R maxima
1
L. asperifolia
L. ovatifolia
Resnova sp. 1
.
D. maculata
D. maculata
Linkage Distance
gure 6.3: The phenogram computed from phytochemical screening tests results.
D. burkei subsp. burkei
D. burkei subsp. burkei (w)
D. burkei subsp. burkei
D. burkei subsp. burkei
D. burkei subsp. burkei
D.burkei subsp. stolonissima
L. asperifolia
Resnova sp. 1
R. maxima
D. comptonii
L. ovatifolia
L. cooperi
Drimiopsis sp. 2
D. maculata
D. pusilla
D. maculata
Drimiopsis sp.
D. atropurpurea
D maxima
D.
,
botryoides
1.0
0.5
1 .O
1.5
Linkage Distance
2.0
2.:
7. KARYOLOGY
7.1 INTRODUCTION
Cytogenetic data is being used in taxonomy to study relationships. It is especially
useful in groups as challenging as the Ledebouriinae U. & D. Miiller-Doblies.
Cytotaxonomy was used to infer evolutionary relationships within the Liliaceae Juss.
sensu lato (De Wet, 1957; Sen, 1975; Greilhuber & Speta, 1976; Speta, 1979; Stedje &
Nordal, 1987). Chromosome numbers published for Drimiopsis Lindl. & Paxt. to date
(Table 7.1) represent only one third of the genus. Matsuura & Sato (1935) published a
somatic chromosome number of 80 for D. botryoides Bak. subsp. botryoides Stedje,
Sato (1942) a number of 64 for D. maculata Lindl. & Paxt., suggesting x
=
8
(Darlington & Wylie, 1956). Hybridisation and allopolyploidy may have produced
diploids and polyploids with x = 6, 7, 8 and 9 leading to n = 11 and 13 (De Wet, 1957;
Mahalakshmi & Sheriff, 1970).
De Wet (1957) proposed that the chromosome counts in Drimiopsis arose fkom x
=
5,
as is found in the tribe Scilleae sensu Baker. He suggested that hybridization then
produced diploids and polyploids giving rise to x = 6, 7, 8,9, 11, 13. Mahalakshmi &
Sheriff (1970) reported gametic numbers of 32, 33 and 34 but they observed several
univalents and multivalents, as well as bivalents in rings and chains during diakinesis.
They suggested that D. kirkii Bak., 2n
=
22n161 of the gametic chromosomes, 2n
68, is a hybrid of uncertain ancestry. The
=
60 (Table 7.1), might also suggest two
different genomes of uncertain origin (Femandes & Neves, 1962). Drimiopsis kirkii
with 2n
= 60
(Matsuura & Sato) and D. maculata with 2n = 80 (Femandes & Neves)
suggest a basic chromosome number of x
chromosome number of x
=
=
10. Gill (1978) proposes the basic
8. He based this on D. volkensii Bak. somatic number of
64. Drimiopsis volkensii has been synonymised with D. botryoides Bak. subsp.
botryoides Stedje (Stedje, 1994), with somatic chromosome number 80. The basic
chromosome number of x
=
8 is supported by Sato (1942) and Darlington & Wylie
-
(1956) iKD. macu7ata-with 2 n '-64:
More recent investigations suggest-the- b a s i ~
chromosome number of Drimiopsis is x = 11 (Stedje & Nordal, 1987; Stedje, 1994).
' This chapter has been published: Lebatha et a].,2003.
Southern Africa seems the center of endemism with Drimiopsis ploidy levels rising
from the center to the north of Afiica. x = 10 is in the south, x = 11 in the east and x =
12, the most advanced basic number, in the west (Stedje & Nordal, 1987). Aneuploidy
is present in this group and the higher the ploidy the higher the asymmetry of the
chromosomes.
Basic number x
=
10 could be a result of allopolyploidy, aneuploidy or dysploidy.
Aneuploidy could have occurred resulting in the gain or loss of chromosomes thus:
Dysploidy, fusion of x
=
10 chromosomes, results in x = 11. Chittenden (1956)
recognizes close relationships of Scilla L. (x
=
5) and Ornithogalum L. (x
=
3) with
Drimiopsis. This raises many hypotheses that can only be answered through
hybridization experiments.
Sen (1975), in his evolutionary inference in the Liliaceae sensu lato, comments on the
polyploidy in Drimiopsis and supports the hypothesis of its derivation fiom Scilleae.
He is convinced that the order Agavales is derived fiom Drimiopsis based on
morphology of the leaves, inflorescence and chromosome morphology. This
evolutionary line links Eucomis L'Herit., Albuca L. and Ornithogalum L to Drimia
Jacq. and Drimiopsis, then ends with the Agavales L. Drimiopsis has lost primitive
characters of Scilla L. and its allies.
The taxonomic significance of chromosome morphology, as compared to chromosome
number, has been found inadequate as shown in, among others, De Wet (1957), Stedje
& Nordal (1987) and Sen (1975). The belief has always been that "Chromosome
morphology, when carefulZy worked out, perhaps offers greater possibilities in helping
with generic dejnitions than does chromosome number" (Rollins, 1954).
Table 7.1: Chromosome numbers of Drimiopsis and sister taxa obtained from
publications.
Source
Gametic # (In)
Somatic
#
Basic
#
I
(2n)
D. maculata
D.crenata
D. botryoides subsp. prostrata
D. burkei subsp. burkei
De Wet, 1957
De Wet, 1957
Fernandes & Neves, 1962
Sharma, 1970
Mahalakshmi & Sheriff,
1970
Vii. et al.. 1982
Vijayavalli & Mathew,
1988
Jessop, 1972
I
Kootin & Sanwu. 1969
1
Stedje & Nordal, 1987
Ovewole. 1988
I
I
Stedie. 1994
Stedje, 1994
Lebatha et al., 2003
D. pusilla
Lebatha et a1.,2003
D. burkei subsp. stolonissima
Lebatha et a1.,2003
L. apertiyora
L. cooperi
L floribunda
L. revoluta
L. undulata
Fernandes & Nives, 1962
Jessop, 1970
Jessop, 1970
Jessop, 1970
Jessop, 1970
L. concolor
L. cooperii
L. jlorib unda
L. luteola
L. marginata
L. socialis
L. ovat$olia
L. revoluta
Jessop, 1972
Jessop, 1972
Jessop, 1972
Jessop, 1972
Jessop, 1972
Jessop, 1972
Jessop, 1972
Jessop, 1972
L. undulata
L. apertijlora
L. ensijiolia
L. somaliemis
L. urceolata
Stedje, 1996
Jessop, 1972
Venter, 1993
Venter, 1993
Stedje, 1996
Stedje, 1996
D. kirkii
I D.maxima
D.barteri
80
44.55
Matsuura & Sato. 1935
Stedje & Nordal, 1987
Stedje, 1994
Sato. 1942
Fernandes & Neves. 1962
1
1 20
I 10
1 44
I 11
1 20
18
10, 13, 15
10, 11, 17,30
13
13, 15
27
9 -13, 15 - 17,
22
10,13 - 15
1 10
I
Table 7.2: Taxa whose root tips were harvested for karyology studies.
Taron
Accession #
L
Origin
Lebatha 042
D./R. maxima
KZN Hutchings garden, RSA
Lebatha 049
Drim iopsis atropuiprea
Luneberg, KZN, RSA
Lebatha 002
Drimiopsis barteri
Tanzania
Lebatha 001
Drimiopsis boryoides subsp. botryoides
Tanzania
Lebatha 003, IN
Drimiopsis botryoides subsp. botryoides
Kenya
1113
I -Leba&a004, IN I Drimiopsis botyoides subsp. botryoides
773
Lebatha 009
Drimiopsis burkei subsp. burkei
I Lebatha 041
1 Drimiopsis burkei subsp. burkei
Lebatha 046
Drimiopsis burkei subsp. burkei
I Lebatha 095
1 Drimiopsis burkei subsp. burkei
1 Drimiopsis burkei subsp. burkei
Lebatha 096
I Drimiopsis burkei subsp. burkei
I
I
I
Lebatha 103
Lebatha 037
I Lebatha 079
1I Drimiopsis burkei subsp. burkei
(
Drimiopsis burkei subsp. stolonissima
I Drimiopsis comptonii
I P a w Dam, ,M A
Rietvlei ,,RSA
I
I
Tierkloof, Waterberg, ,RSA
I Rasesa, Botswana
I Rasesa, Botswana
I
I
I Kgale Hill, Gaborone, Botswana
( Strydom tunnel, Swaziland
I
Swaziland
Drimiopsis maculata
Lebatha 017
Lebatha 02 1
Drimiopsis maculata
University of Zululand garden,, RSA
Lebatha 033
Drimiopsis maculata
KZN garden,, RSA
Lebatha 039
Drimiopsis maculata
Hitchings garden, KZN,RSA
Lebatha 102
Drimiopsis maculata
Grahamstown, ,RSA
Lebatha 104
Drimiopsis maculata
Potch garden, ,RSA
Tshieng, Botswana
I Emtunzini, KZN,,RSA
I
Lebatha 024
Drimiopsis maxima
KZN Hutchings garden, ,RSA
Lebatha 036
Drimiopsis maxima
KZN Hutchiigs garden, ,RSA
Lebatha 069
Lebatha 022
I
I Drimiopsis sp.
I Roossenekal ,,RSA
Drimiopsis reilleyana
I Drimiopsis sp.
I
Lebatha 052
I
Lebatha 058
I Lebatha 014
Drimiopsis sp.
1 Ledebouria
I Ledebouria apertqolia
I Ledebouria apertvoh'a
1 Ledebouria sandersonii
I
Lebatha 06 1
I
Lebatha 081
I
Lebatha 087
Lebatha 047
I Resnova maxima
I Resnova megaphyla
I
Lebatha 086
I
Lebatha 088
I
II Mbabane, Swaziland
I Carolina, ,RSA
1
I
(
Resnova minor
I
Roossenekal, ,RSA
I Drimiopsis maculata
1I Drimiopsis pusilla
I
I
I
Lebatha 006
1 Lebatha 078
I
Vaal river, ,RSA
Drimiopsis liniopapilla
Lebatha 060
I
Potchefstroom, RSA
Drimiopsis burkei subsp. burkei
Lebatha 054
Lebatha 055
I Tanzania
I Parys Dam,,RSA
I
I S 26 27.677 E 30 58.690
I
I Molepolole, Botswana
I
(
Mkanga, Swaziland
I Barberton, ,RSA
I
I Pigs peak, ,RSA
I
I Mandini, Ngwenya Game Reserve, Swaziland
I
I
1 Roossenekal, ,RSA
I
1
Roossenekal, ,RSA
I
Centromere organization however, can play a significant role in relationships. Jackson
(197 l), and all the authors quoted therein, found that mitotic arrestors affect the shape
and size of chromosomes. Masterson (1994) found a significant relationship between
the size of the stomata and the ploidy level: the higher the ploidy, the larger the
stomata. Higher ploidy levels produce plants with more vigor and loss of a whole
chromosome produces dwarfs (Brandham & Cutler, 1978).
Karyology studies on the Ledebouriinae, Table 7.2, give inconclusive results. There is
no record of Resnova v.d. Merwe chromosome numbers in publications. Wolfgang
Wetschnig (2004, Institute of Botany, Austria, personal communication) reports
somatic chromosome numbers of seven Resnova taxa as 2n=10 with a bimodal
chromosome set. SciNa diploid chromosome numbers vary from 2n=18 in S. firmifolia
Bak. (De Wet, 1957) to 2n=28 in S. natalensis Planch. (Jessop, 1970) and 2n=34 in S.
natalensis (De Wet, 1957).
7.2 OBJECTIVES
To determine the as yet unknown chromosome numbers of Drimiopsis taxa. The results
together with chromosome numbers from publications are used as a taxonomic tool to
establish relationships within the genus and with sister taxa Ledebouria and Resnova.
7.3 MATERIALS and METHODS
General
Fresh root tips of plants listed in Table 7.2 were harvested from bulbs between 8:00 am
and 11:30 am. The root tips were cold treated, placed in water at 4' C in a refrigerator,
for 24 hours to stop cell activity. Cold treatment is the preferred mitotic arrestor (as
opposed to other arrestors like colchiline & mono-bromo-neptheiline) (Kleynhans &
Spies, 1999). Mitotic arrestors destroy spindles stopping the cells segregating to the
poles -(maphase),having sells mstlyia the metaphase- stage. _This-stage is commonly
-
-
-
-
-
-
of about half to 3 hour duration in the morning.
The root tips were then hydrolysed in hot (60' C) 1N HCl for 10 minutes, then stained
with leuco basic fuchsin at 4' C in the dark for 24 hours. Stained root tips were then
squashed in aceto-orcein and left to stain for 20 minutes (Darlington & LaCour, 1976;
Kleynhans & Spies, 1999). Meanwhile, coverslips were prepared with Mayer's albumin
and placed on the squashes. The slides were then placed between three filter papers
folded in half, then squashed. The slides were made permanent using the float off
method in 45% acetic acid, dehydration in a series of alcohol and mounting in euparal
(Darlington & LaCour, 1976).
Carnoy's fixative:
6 ethanol : 3 chloroform : 1 glacial acetic acid. (In the field, ratio used is 6:3:2 as acetic
acid is volatile). Refrigerate. Stops cell processes and chloroform helps in removing
lipids.
Ethanol:
70% for storage of mitotic material, 30 % storage of the meiotic material and 100% &
80% also used in the permanent making of slides.
Mayer's albumin:
25 ml egg white, 25 ml glycerol & 0.5 g sodium salicyclate. Used in the preparation of
coverslips in making permanent slides
Stains:
Aceto-carmine: 100 ml 45% acetic acid, 2 g carmine-for
colouring in meiotic
squashes, used in conjunction with Ferri-acetate stain.
Ferri-acetate is not actually a stain but, used with aceto-carmine, improves the staining.
Aceto-orcine: 2.2g orcine, lOOml warm glacial acetic acid. Used after staining mitotic
root tips with Leuco-basic fuchsin.
Leuco-basic fuchsin: l g basic fuchsin, 200ml boiling water, 30ml 1N NCL, 3g
potassium disulphate + 0.5 g active carbon. Store refrigerated and in darkness
(covered with foil). This stain darkens DNA.
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
7.4 RESULTS and DISCUSSION
The following plant material, out of 29 squashes processed, produced positive results:
1)
Drimiopsis burkei Bak. subsp. burkei (1870). Material used (Lebatha 009)
collected at Potchefstroom (2627CA), North West Province (Figure 7.1).
2)
Drimiopsis burkei Bak. subsp. stolonissima U & D M-D (1997). Material used
in this study (Lebatha 037) was collected from the type locality, Pilgrim's Rest
(2430BC), Limpopo Province (Figure 7.2).
3)
Drimiopsis pusilla U & D M-D (1997). The plant studied (Lebatha 078) was
collected at the type locality, Mbabane (263 IBD), Swaziland (Figure 7.3).
Two basic chromosome numbers are reported in Drimiopsis, i.e. x = 11 for D. burkei
subsp. burkei and D. pusilla, and x = 10 for D. burkei subsp. stolonissima (Table 7.3).
These data, combined with recently published data (Stedje & Nordal, 1987; Stedje,
1994), suggest that the majority of Drimiopsis species appear to have a basic
chromosome number of x
11 (Table 7.1). Tetrapolyploids are common but
=
pentaploids and hexaploids seem confined to tropical Africa.
Table 7.3: New Drimiopsis chromosome numbers
Taxon
Collector no.
D. burkei subsp. Lebatha 009
Chromosome
Basic
number
x=
44
11
Distribution
Southern
burkei
Afiica
44
Lebatha 078
D. pusilla
11
Southern
Africa
D. burkei subsp. Lebatha 037
stodcwzissima
-
-
-
-
~- -
-
-
-
-
-
-
-
-
40
-
-
-
10
-
-
-
-
-
-
-
-
Southern
Afiica
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
---.---..-.---.--..-.--....---.
Figure 7.1: Drimiopsis burkei subsp. burkei somatic chromosome number 2n = 44
Figure 7.2: Drimiopsis burkei subsp. st%nissima 2n = 40
,
-:p.
Figure 7.3: Drimiopsis pusil/a somatic chromosome number 2n = 44
107
The somatic chromosome numbers of D. botryoides subsp. botryoides (2n = 80,
Matsuura & Sato, 1935) and D. macu/ata Lindl. & Paxt. (2n = 64, Sato, 1942),
suggesting x = 8, are the only ones with this karyotype known so far in this group. With
the large numbers of chromosomes involved, it is possible that 80 could have been 77
and less likely 88. The 64 described could have been 66 and 68 - the counting of large
chromosome numbers can be tricky giving misleading results.
Chromosome counts of D. crenata v.d. Merwe and D. saundersiae Bak., 2n = 20 (De
Wet, 1957); D. macu/ata, n = 15 (Jessop, 1972) (Table 7.1), together with latest results
(D. burkei subsp. st%nissima, x = 40) support x = 10. D. crenata and D. saundersiae
have been synonymised under D. burkei subsp. burkei (Jessop, 1972), 2n = 44. D.
vo/kensii (Engl.) Bak., 2n = 64 (Gill, 1978) and D. kirkii Bak., 2n = 60 (Fernandes &
Neves, 1962; Vij et a/., 1982), In = 30 (Vijayavalli & Mathew, 1988, 1990) and 2n =
68 (Mahalakshmi & Sheriff, 1970), are synonyms of D. botryoides subsp. botryoides
(Stedje, 1994) with 2n = 80 ( Matsuura & Sato, 1935), 2n = 44, 55 (Stedje & Nordal,
1987, Stedje, 1994) and 66 (Stedje, 1994).
Present and previous studies do not support the synonymisation of D. crenata and D.
saundersiae, x = 10, under D. burkei subsp. burkei, x = 11. Drimiopsis crenata and D.
saundersiae have a similar basic number than D. burkei subsp. st%nissima.
Chromosome numbers of D. kirkii, x = 10; and D. vo/kensii, 2n = 64 also cast doubts on
the synonymisation under D. botryoides subsp. botryoides, x = 11.
7.5 CONCLUSION
Three results were obtained after processing 29 squashes. Thus, the methodology for
harvesting root tips at their maximum meristematic activity needs optimisation. Bulbs
grown in glyco gel had the least meristematic activity and root development. Future
investigations will involve fIrst forcing bulbs into dormancy, replanting in water, then
root tips harvested at their maximum rate of mitotic cell division, at about 1cm long.
108
The present study suggests the genus Drimiopsis has basic chromosome numbers x
=
10 and x = 11. The fonner is the plesiomorphic character still predominant in southern
African taxa. The x
= 11 plants, some southern African but all tropical African, would
then be derived character. This could have arisen through mutations, via speciation,
reticulation and retention, to spread to tropical Africa. Higher chromosome numbers in
the 80's and 60's are in tropical African plants (Table 7.2 & 7.3). The southern African
plants are diploids and tetraploids.
Additional chromosome counts are needed not only to verify published data but also
clarify issues raised in this investigation. In particular attention needs to be given to,
among others, the case of D. barteri Bak. with 2n = 24 (Oyewole, 1988), suggesting x
= 12; D. macu/ata with 2n = 64 (Sato, 1942), n = 15 (Jessop, 1972) and Resnova with
2n = 20.
Drimiopsis and its sister taxa challenge the hypothesis that the higher the ploidy, the
larger the stomata (Masterson, 1994); higher ploidy levels produce plants with more
vigour and loss of a whole chromosome produces dwarfism (Brandham & Cutler,
1978). This situation has not been observed in this complex.
109
8. DNAANALYSIS
8.1 INTRODUCTION
Molecular studies together with morphology have become one of the focal points of
systematics today, providing new insights into possible relationships, especially in
difficult taxa (Hillis, 1987; Doyle, 1993). Some consider DNA data to be better
phylogenetic indicators than morphological characters (Sytsma et a/., 1990). The study
of Triticum dicoccoides (Koorn. ex Aschers. & Graebn.) Aarons and Hordeum
spontaneum Koch. populations having different forms of rDNA seems to present an
exception to this hypothesis. The two populations, from different ecological areas,
suggest that rDNA variation could be determined by selection (Gupta et a/., 2002; Nevo
et a/., 2002).
Molecular systematics originate from population genetics (Hillis, 1987) and traditional
taxonomy from the Linnean system. Some may say this creates bias because molecular
systematics starts with the phylogeny and deals with the taxonomy later. Disregarding
prior taxonomic delimitations when creating monophyletic groups minimizes bias.
Morphological character variations are, however, taken into account for the groups
being analysed otherwise there would be sampling problems. Molecular systematics,
when used in conjunction with morphological data often results in diagnosable clades.
The concept of homology differs in molecular and morphological studies. Homology in
molecular studies is arrived at statistically whereas in morphology it is through
congruence of features (Patterson, 1988). Expediency dictates that species are
morphologically distinct units whereas molecular systematic concepts support species
as having negligible genetic divergence! The two concepts are not, in my view, in
conflict and separation but create internecine systematics. Advances in systematic
software have made it easier to translate molecular data into phylogenetic
reconstruction. It is expected that molecular phylogenies should concur with
morphological phylogenies.
Homoplasy (reversals) is likely to occur at a higher rate in DNA sequences than in
morphology. Assessment of homoplasy is easier when dealing with morphological
characters, as they are distinctive. Assessment of homoplasy in DNA sequences
110
- - -
involves more sequences. Also, insufficient sampling can make the detection of
paralogous genes difficult and using these genes instead of the orthologous ones will
give well resolved and strongly supported trees not representative of phylogeny (Doyle,
1993). Molecular studies traditionally utilise only a section of the whole genome.
Chloroplast, mitochondria and ribosomal (cytoplasmic genome) DNA are popular in
plant molecular systematic studies. However, the chromosomal genome that contains
the Mendelian genes, i.e. the genes involved in phenotypic expression, is not used in
molecular studies (Grant, 1975). This suggests molecular study and traditional
taxonomy deal with different parts of the overall genome and thus should be applied in
synchronology in phylogenetics deliberation. Doyle & Endress (2000) compared
extensive morphological data and molecular data from rbcL, 18S and atpB. The
combination of chloroplast, nuclear and mitochondrial DNA sequences with
morphological characters produced inconclusive results. Some taxa on the one hand got
better resolution and the combined cladograms more inclined to the molecular data
cladograms. On the other hand, conflicts still prevailed. Reed and Frankbam (2001),
while computing the correlation coefficients between 71 data sets of both molecular and
quantitative phenotypic variation of selected plant groups, found the correlation
generally weak. This confIrms that molecular characters do not usually reflect
accurately the quantitative phenotypic variation in a group.
The uniparentally inherited chloroplast DNA (cDNA) is small in size, circular, ca. 120
-217 kilo bp, and conservative and has a low rate of structural and sequence evolution.
The highly conserved chloroplast trnL (UAA) and trnF (GAA) genes, the trnL intron
and the non-coding intergenic spacer (IGS) between the trnL & trnF genes (Taberlet et
ai, 1991), are popularly used in Hyacinthaceae molecular studies. These regions are
especially reliable and allow low sample diversity. The trnL intron and the trnL-trnF
IGS have sequence divergence rates considerably higher than those in the chloroplast
rbcL (Gielly and Taberlet, 1994). The trnL intron has four distinct conserved regions
and three variable regions, while the IGS has higher substitution rates (Fangan et al.,
1994; Eldenas & Linder, 2000). However, Rieseberg and Soltis (1991) maintain that
chloroplast genes have the potential of becoming dissociated from their genome,
making them unreliable. Hybridisation can also result in chloroplast sharing, as was the
case in the Eucalyptus L'Herit. subgenus Monocalyptus Prior & Johnson (Myrtaceae)
111
of southeastern Australia and Tasmania (McKinnon et al., 1999). In addition, genes
selected for analysis could have different evolution rates from those determining
phenetic traits. The evidence derived from such a scenario would indicate close
relationships where phenetic characters indicate more distant relationships.
Molecular data combine with morphological data to give total evidence, promoting a
better estimation of phylogeny (Doyle & Endress, 2000). Morphological data help
support poorly resolved molecular analysis clades and improve inference of
relationships through determination of ancestral character states.
Systematists working on taxonomic groups related to Drimiopsis Lindl. & Paxt. have
found the chloroplast trnL and trnF genes, the trnL intron and IGS between the trnL &
trnF genes phylogenetically informative (Fangan et al., 1994; Stedje, 1996, 1998;
Pfosser & Speta, 1999; Eldenas & Linder, 2000; Pfosser et al., 2003; Wetschnig &
Pfosser, 2003; Manning et al., 2004). Other cDNA genes, ndhF (+restriction sites),
matK (+ restriction sites) and commonly, rbcL, are also used in systematic studies of
the Hyacinthaceae.
Stedje (1998) sequenced the trnL intron and the trnL IGS of tropical African taxa
namely: D. barteri Bak., D. botryoides Bak. (=D. botryoides Bak. subsp botryoides)
and D. perfoliata Bak. (=D. botryoides subsp prostata Stedje). The spacer sequence
was reported more variable (with 33 phylogenetically informative characters) than the
intron sequence (with 31). The cladogram, representing the first published cladogram of
the Hyacinthaceae, resolved Drimiopsis to be monophyletic and sister to Ledebouria
Roth taxa (Figure 8.1a). The same topology was obtained when adding seven
morphological characters to the data matrix (Figure 8.1b). Leaves spotted 1[1] are
hypothesized to be a synapomorphy for the Drimiopsis and Ledebouria clade and
perianth segments 4[1] a synapomorphy for Drimiopsis. The clear rectangles indicate
homoplasy in Scilla autumnalis L. and Drimiopsis, where the bracts were lost 2[0]
twice. The reduction in ovule number 6[1] occurred both in Ledebouria-Drimiopsis
clade and Scilla autumnalis. The stipitate ovary 5[1], present in Ledebouria and Scilla
L., is reversed to sessile 5[0] in Drimiopsis.
112
The tree produced from molecular data produced an unresolved tree, particularly the
polytomy for D. barteri, D. botryoides and D. perfoUata (Figure 8.la). Inclusion of
morphological data collapsed one weak node, increasing tree length but not resolving
the three Drimiopsis taxa polytomy (Figure 8.1b). Ledebouria including L. revo/uta
(L.f.) Jessop from Zimbabwe, India and Cameroon, is paraphyletic (Figure 8.la). This
is, however, resolved with the addition of morphological characters (Figure 8.lb).
Unfortunately, Resnova v.d. Merwe was excluded in this analysis. Stedje (1998)
considers morphological characters distinct enough to separate the genus Drimiopsis
from Ledebouria.
A
OmlhogalumImolillll
+1
Scillaautumnalis
52i
Scillaperuvians
26
Sci!Jaau1umnalis
Scilla peruviana
SciilalazuGna
Scillalazulina
ScillanlllVOSa
...,
00;;
+1
M
a
...,7
iOOi
OmkhogaJum
tenuifolium
ScillanolVOsa
ledebouriaoordifolia
ledeooulla cordilotia
ledebouriarevOO4a
(Zimbabwe)
ledebouriarevoluta(India)
ledcbowia revoluta(Zimbabwe)
LedebouriarlMilla (Cameroon)
ledebouria revoluta(Incrza)
Ledebouriasomarensis
ledebouria revoIuta(Cameroon)
Ledebouriaurceolata
LedebouriasomallWis
Diimiopslsbaited
Led8bouriaurceolata
DriniopsisboIryoidesssp.bollyoides
Dtimiopsisbarteri
Dliniopsisbotryoidesssp.prostrala
Driniopsis bolryoidesssp.botryoides
Dtima altissima
Drimia81al4
I
b
3
7
Drimiopsisbotryolclesssp.ploSlrata
Drimiaaltissima
Drimiaelata
Figure 8.1: A, Cladogram from sequenced trnL intron and the trnL IGS grouping Drimiopsis with
Ledebouria into a strong clade. The Ledebouria-Drimiopsis clade is not well resolved (Stedje, 1998); B,
Cladogram combining DNA sequencing data with morphological data. The dark rectangle represents
synapomorphies, the clear rectangle homoplasies. (Stedje, 1998).
Pfosser & Speta (1999) analyzed the trnL intron and the trnL-trnF IGS sequences in an
expanded matrix (Figure 8.2). This study included taxa analysed by Stedje (1998): three
east African Drimiopsis species in addition to D. macu/ata Lindt. & Paxt., and four
Ledebouria species in addition to L. socia/is (Bak.) Jessop. Ledebouria taxa form a
poorly resolved monophyletic clade with Drimiopsis. Schizocarphus nervosus (Burch.)
v.d. Merwe is basal to the Ledebouria-Drimiopsis
clade. The results support the
validity of grouping Drimiopsis and Ledebouria in the subtribe Ledebouriinae U. & D.
Miiller-Doblies (Miiller-Doblies & Miiller-Doblies, 1997).
113
Figure 8.2: A section of the Pfosser & Speta (1999) DNA sequencing data cladogram
showing the Ledebouriinae. This cladogram was based on the trnL intron and the trnLtrnF IGS sequences in an expanded matrix. Included in the analysis are four Drimiopsis
species and five of Ledebouria.
Wetschnig & Pfosser (2003) analyzed the trnL intron and the trnL-trnF IGS sequences
in an expanded matrix of the Hyacinthaceae (Figure 8.3). Their study within the
Ledebouriinae included five Drimiopsis taxa one of which, D. kirkii Bak., has been
synonymised under D. botryoides Bak., two Resnova taxa and 17 Ledebouria taxa,
eight of which are unidentified. Schizocarphus nervosus is basal to the Ledebouria,
Resnova and Drimiopsis monophyletic clade. The tree hypothesizes Ledebouria
belonging in two major clades, one basal consisting mostly of Ledebouria unnamed
species, and the other imbedded within a poorly resolved Resnova and Drimiopsis
clade. From these results, recognition of Drimiopsis and Resnova as separate genera
renders Ledebouria paraphyletic.
114
-
--
-
Eucomis zambesiaca
Eucomis vandennerwei
Eucomis montana
Eucomis punctata
Eucomis bicolor
Ledebouria hyacinthina 2
Ledetiouria somalensis
Ledebouria I}yacinthina 1
Resnova humifusa
Resnova maxima
Drimiopsis maculata
Drimiopsis sp.
Drimfopsis klrkii
Drimiopsis botryQides
Drimiopsis barteri
Ledebouria socia/is
Ledebouria socia/is agg.
Ledebouria cf. conc%r
Ledebouria sl? 2
Ledebouiia cordifolia
Ledebouria revoluta
Ledebouria sp. 3
Ledebouria sp. 4
Ledebouria sp. 6
Ledebouria sp. 5
Ledebouria sp. 7
Ledebouria sP. 8
Ledebouria floribunda
-
89/100
84/100
93(100 70/97
100(100
861100I .,..
-/96
-
-
84/100
64(99
-
Merw;lla sp. 2
MerwiHa$p. 1
Merwl1(anatatensis 2
Merwilla natalensis 1
Mlb
Mia
Ledebouriasp. 1
Schizocarp/Jus
nefVOSU$ 2
Schizoca;phus nervosus 1
Merwilla krauss;;
Figure 8.3: A section of the Wetschnig & Pfosser (2003) DNA sequencing data cladogram based on
the tmL intron and the tmL-tmF IGS sequences. The study included within the Ledebouriinae, five
Drimiopsis, two Resnova and 17 Ledebouria taxa.
Manning et a/. (2004), on the basis of cDNA trnL and rbcL analysis, generated a
consensus cladogram of 980 trees (Figure 8.4). The analysis resolved tribe Massonieae
Bak. into clades with a topology that does not support the three Miiller-Doblies &
Miiller-Doblies (1997) subtribes, including subtribe Ledebouriinae. In addition, the
Ledebouria sensu /ato clade forms polytomies with, among others, Schizocarphus
nervosus. With these results Manning et a/. (2004) prematurely, without support of
morphological data, sunk Drimiopsis and Resnova into Ledebouria sensu /ato.
115
- -
68
100
99
82
88
70
AmP.hisiphon st~sa
Neobakeria zevheri
Androsiphon alba1
Androslphon alba2
Dauben~ aul'GS
Neobakeria marglnata
Neobakeria angUStlbJia
Androslphon capense
Neobak8ria cemata
Neobakeria marginate
Neobakeria namaquensis
~taea
lachenaliitlora
DriTliopsis kirkii
Drmlopsls maculala
Driniopsis woodii
Ledebouria moperi
Ledebouria ovatifclia
Ledebouria undulata
Ledebouria hyacinthina
Ledebouria soclalis
Resnova pilosa
Resnova humllUsa
Euconis autumnalis
Eucon'is humilis
Eucoms regia
Lachenalla aloides
Lachenalia pusiDa
Periboea ccrymbosa
PoIY)!8na ensifolla
Lac'benaUa namibensis
Massonla depressa
Massonla echlnam
Massonla heterandra
Massonia pustJlata
Wh iteheadla biblla
Whiteheadla eteslonamibensls
Namophlle uropetala
Velthetmia bractBata
Schlzocarphus nervosus
Merv.flla natalensis1
MenNilia natalensis2
PS8udoprqsp@fO firmifotium
Figure 8.4: A section of the Manning et al. (2004) tree based analysis of cDNA trnL and rbcL
genes. This study included three Drimiopsis, two Resnova and five Ledebouria taxa.
Ideally, molecular data should harmonise with morphological data when creating a
phylogenetic classification. Incongruence should be suspect and regarded as a
procedural problem. Molecular results should be verified by including suites of
homologous traditional morphological characters, although this too can be problematic
due to convergence and divergence. Although incongruency should result in a review of
the data matrix for possible coding errors, the existence of incongruency per se is not
unknown. Taxonomic decisions should not be founded on analyses based on a limited
number of morphological or molecular characters in an analysis. Inadequate character
balancing results in unnecessary explosion of new taxa as seen in recent works on the
Hyacinthaceae where proposal of new taxa may well be premature (Stedje, 2001). On
the other hand, as exemplified in Manning et of. (2004), it can lead to premature
lumping. In my opinion, segregation of genera on the basis of "recognizable
116
morphological synapomorphies with reciprocal monophyly of clades" (Manning et a/.,
2004:535) assigns Drimiopsis, Resnova and Ledebouria as separate genera.
8.2 OBJECTIVES
This study conducts molecular studies on Drimiopsis, performing DNA sequencing in
order to resolve intraspecific and interspecific boundaries within the Ledebouriinae and
create a phylogenetic classification through a combination with morphological data.
8.3.MATERIALS and METHODS
The cDNA trnL (UAA) and trnF (GAA) genes, the trnL intron and the non-coding
region of the trnF intergenic spacer, IGS between the trnL & trnF genes (Taberlet et
a/., 1991) are sequenced in this study. These regions were selected because of their
proven reliability especially with low sample diversity. The chloroplast gene was
selected for primary sequencing because it has shorter run times, enhanced resolution
and gives better supported clades. Also, several Ledebouriinae taxa have been thus
sequenced.
Total genomic DNA was extracted from 49 plants: 40 Drimiopsis, 5 Resnova and 4
Ledebouria and sequenced using commercially available primers and a more amenable
alignment processes can cut costs. The fundamental procedure followed was extraction,
quantification and amplification with volume augmentation (to 300 Ill), sequencing and
sequence alignments (Appendix 2).
117
Table 8.1: Taxa sequenced.
Taxa
Accepted names
Drimiopsis sp.
R. lachenalioides
D. kikiae
Accession
number:
Lebatha 019
Locality
D. atropurpurea
Lebatha 048
D. liniopapilla
Lebatha 053
D. atropurpurea
D. atropurpurea
Lebatha 049
D. burkei subsp. burkei
D. kikiae
Lebatha 046
D. burkei subsp. burkei
D. queae
Lebatha 055
D. burkei subsp. burkei
D. burkei
subsp.stolonissima
D. comptonii
D. davidsoniae
D. burkei
Lebatha 095
Durban, South Africa
South
Houtbosdorp,
Africa
Charles Craib, South
Africa
South
Roossenekal,
Africa
South
Luneberg,
Africa
Vaal River, South
Africa
South
Waterberg,
Africa
Rasesa, Botswana
D. stolonissima
Lebatha 037
Strydom Tunnel, RSA
D. comptonii
D. davidsoniae
Lebatha 079
Lebatha 038
D. maxima
D. pusilla
D. reilleyana
Resnova sp.
R. maxima
D. pusilla
D. reilleyana
R. megaphylla
Lebatha 047
Lebatha 078
Lebatha 068
Lebatha 086
Resnova sp.
R. maxima
R. megaphylla
R. maxima
Lebatha 088
Mbabane, Swaziland
Pilgrim's Rest, South
Africa
Mandini, Swaziland
Mbabane, Swaziland
Mkhaja, Swaziland
South
Roossenekal,
Africa
South
Roossenekal,
Africa
Mbabane, Swaziland
Drimiopsis sp.
Drimiopsis sp.
Drimiopsis sp.
Lebatha 045
Lebatha 077
8.4 RESULTS and DISCUSSION
8.4.1 PROTOCOL APPRAISAL
Out of the 49 plants, the total genome of only 17 of Drimiopsis (Table 8.1) were
successfully amplified. The rate of success was dependent on several factors that are
part of ongoing research. The factors were assessed and logical sequential steps taken
to adjust the extraction procedure. The DNA extracted was good but apparently had an
inhibitor preventing sequencing. Other than re-servicing of equipment, the following is
a synopsis of troubleshooting for the persistent problems encountered:
1. DNA concentration inadequate.
Solution: after incubation before centrifuging or washing, the sample was left in the
CTAB-buffer mixture for one week to allow longer protection from enzymes and
118
secondary metabolites activity and more efficient forming of nucleic acids-CTAB
complexes.
2. Samples contain too much phenols thus interfering with the PCR process.
Solution: decrease phenol concentration by changing the PVP to fresh material weight
ratio to 1:1 when grinding. The freshly ground material was placed in heated CTABbuffer + PVP mixture (65° C for 30 minutes), then similarly incubated.
3. DNA would not amplify because of the apparent presence of inhibitors.
Solution: Equipment and chemicals were autoclaved and fresh plant material washed
with 70% ethanol. The magnesium concentrations was adjusted up to 4 Ill. Extra steps
of washing the DNA pellet with 0.1 volume ammonium acetate in 500 IIIultra pure H20
were included in the protocol. These were further washed once with chloroform and
stored in 100% ethanol overnight; centrifuged then washed twice with 70% ethanol and
stored in TE buffer (not in ultra pure H20as in the previous protocol).
4. Primers seemingly form primer-dimers or concatamers.
Solution: Modify two new protocols from Hillis et al. (1990) and Kopperud & Einset
(1995). Use shorter primers and new extraction protocol.
8.4.2 PCR RESULT SUMMARY
The following photographs (Figures 8.6-8.7) illustrate the PCR process followed in the
amplification of the 17 Drimiopsis taxa as well as the complexities encountered.
Amplification would be ideal as in the case of D. atropurpurea (Figure 8.6a) or fail
even with good quality genomic DNA (Figure 8.6b). Re-amplification using e.g.
different conditions, vary MgCI2, PVP, DNA concentration, aneal temperature and
DMSO, produced little result sometimes forming primer-dimers or concatamers
(Figures 8.6c & 8.6d). Purification of material extracted from the agarose gel yielded
poor results (Figures 8.6e-f). The sequences failed to produce any results (Figures 8.7a,
b, d, e, f and g). Sample 28 cut out (gel extract) band was purified and sequenced
(Figure 8.7c).
119
-.----...-----.-..--.-..--......--
0)
.
'" ..
I
,
........
".
-.
.
.
.
...
.' .'...
.
.
..
...
...
'......
'
.
Figure 8.6: PCR results. A, D. atropurpurea results: the amplification was ideal at all magnesium
concentrations of 0.5, 1.0, 1.5 and 2 ,.drespectively; B, samples 1-19 showing good quality genomic DNA
with little protein contamination and lots of RNA. Amplification produced no bands. Aliquots were Rnased
and re-amplified but no bands were produced; C, re-amplified samples 1-13 using different conditions:
vary MgCI2, PVP, DNA concentrations, aneal temperature and DMSO. Good bands with sample 1, no
bands with sample 13. Note the double banding from the primers -primer-dimers or concatamers; D,
amplified samples 1-19 using optimum conditions from scanB. Faint bands seen on samples 1,4 and 14; E,
Bands 1 & 4 & 14 were extracted from the agarose gel, purified and sequenced. Yield was very poor and
the sequence failed to produce any results; F, amplification without PVP shows single band from sample 28
only, R. maxima (sample 24 showed a smear).
120
Figure 8.7: PCR results. A, Samples 26, 28 and 29 are good though when amplified using varying
concentrations of PVP, only sample 28 produced visible bands; B, re-amplification of sample 28; C, cut
out (gel extract) band from sample 28 (in A), that was purified and sequenced; D, some samples
processed with maximum DNA and masses of MgCl2 and PVP. Many non-specific bands of different
sizes are visible; E, gel obtained using different Taq brands: Roche, Expand High Fidelity, Fermentas
and Bioline; F & G, gel obtained using Bioline X-ACT Taq.
121
8.4.3 SEQUENCING RESULTS
D.aJroJlUlpIll'l1Q
D. I1IUir4i
R. maxima
1
ataaanaaaranncc
1
1
e t e tee
a ( t r ace t
a a e ( t t C t a a g t t tea
naantcnnnnnnnenencaln
r
D. aJroJlUlpUf'8Il101
n a a g g g a a u e g a a
D. I1Il1ir4i
a a D a t t t ( I a e (t
R. maxima
101
101
D. aJroJlUlpIll'l1Q 201
D. I1IUir4i
R. maxima
201
201
g t a cae
r tat
( a t
l
tat
a a I a a t
lecClanaagetn
C g a I g g a ( t t g DaD
t t a ate
a a t t c t a
i
acacnniaa
e ( t c a c c D
a a a a a a a a g
ncagganaCagaDCCCD
c t c t t tat
t c ten
t e (I
t t c a c a a a a a t t e t t t e
D. aJroJlUlpIll'l1Q 601
D. I1Il1ir4i
601
e C
e a c I I
(n
(Iant
R. maxima
501
501
601
D. aJroJlUlpUf'8Il701
D. l1urMi
R. maxima
701
701
cae
a a i
a t c t t t
let
tat
t t
l
a I
c In
t a a
tat
( e n n u ( ace
a cat
i
a
tllni
a I g t a
a a a a t
Daaaaa
caanan
t t t c t a
t t tat
c
( n a a a a a
ggaaaaaccccccnccaccn
a tit
glnngggantaaDcnanan
a a t t a a a c cD t t g g a a a t D
tag
a a c a g ( t t ( cat
t I a C
ceccDcccnCnnnca(naaaegagaganganDnaaac
c c c n a n gnu n c c n ant
c e c t
D. aJroJlUlp1ll'l1Q901
D. blU'kei
R. maxima
ncaaaattnceeeaaangngcnaatnnlrnc(agncg
D. aJroJlu'1!Uf'8Il1001
D. l1urMi
acnnnanlceCI(ceannncancngceteatnacalt
801
t a a a I e a
act
( g a
g act
a t
DDaaaaa
aenaga
t t a ate
n t t t t t c
t t t t t eta
t a a a t
D. aJroJlUlpIll'l1Q 801
D. l1urMi
R. maxima
I a ant
t a n a ant
a n a tat
Il
a a a c r
I a a a I a tat
t I a a t tat
t t r
I t c I a e g I a t t t t c e
e t t a c
D. aJroJlUlpIll'l1Q 501
D. l1urMi
R. maxima
l
a
t t t e t tag
i
l
l l
l
l
l
'
l I
l
l
t n n ( ( a ( (i
401
401
a t ( a (
(
nnnann(naaaealaccgaccccnialtatt
g g a a t t e ( t t g ace
a tat
a t g t tea
t t t g t a
( t C c g a ace
a t a ace
a a t t t t I a g tea
eta
a
a C a I tat
ant
g D a t g
c ( t ( a e gat
t t ( t a a
D. l1urMi
R. maxima
u a a
g t e ( ( C I a
D. /JllrMi
R. maxima
D. aJroJlUlpIll'l1Q 401
t e t t ( ( eat
t t ( e ( a t a e C t (
t a e a g t a e a a a t a a t a e a a a
n ( ace
D. aJroJlUlpIll'l1Q 301
301
301
i
anCI
a a a a
I a t a
n a a a n ( (
ttaaatt
a tat
a I I
t tat
tctnaaanaaaana'ln
C ( ( Dca
t D t Dee
c t c t g c a c c tat
e t t t (
cn ( a c
c e t t t
R. maxima
D. aJroJlUlpIll'l1Q1101
D.l1urlwi
R. maxima
e n e C a ( a c c c n c ( C n g c n can
a g a c
1125
Figure 8.8: Sequencing results. The cDNA trnL-trnF IGS sequences of D. atropurpurea, D.
burkei and R. maxima obtained from this study. The D. atropurpurea and D. burkei sequences
had too many unknowns, 1075 and 582 respectively.
122
Three of the 17 samples amplified were successfully sequenced (Figure 8.8). Resnova
maxima Bak. produced a sequence of molecular weight (kDa): ssDNA: 243.31.
dsDNA: 485.6 with 788 base pairs and the composition: 266 A; 116 C; 124 G; 28 1 T; 1
OTHER. Drimiopsis atropurpurea N.E. Br. Sequence consisted of 1125 base pairs: 10
G, 9 A, 21 T, 10 C and 1075 OTHER. The D. burkei Bak. sequence consisted of 820
base pairs: 85 G, 50 A, 74 T, 29 C and 582 OTHER.
8.5 CONCLUSION
This study could generate no useable DNA data for analysis- one 'good' sequence of
the trnL-F gene of R. maxima produced out of 49 extractions. Investigations are
ongoing, the results of which will be combined with morphological data in a total
evidence approach.
9. PHENETIC ANALYSIS
9.1 INTRODUCTION
Grouping natural variation or biodiversity according to similarity and/or difference is as
old as taxonomy. Whewell (1859) maintains that botanical classification depends on
"the ideas of likeness and differences". Systematic studies always begin with an
assessment of morphology, defining groups for further analysis e.g. molecular analysis
(Jensen, 2003).
Phenetic analysis (Sokal & Sneath, 1963) groups taxa according to overall similarity
and does not necessarily suggest evolutionary relatedness. It may be the result of
convergence, reversals and parallel evolution (homoplasy). Phenetic principles are
essentially applied in all species identification. This orthodox method is chosen for
generic relationship analysis in order to build a broad database essential for taxonomy.
Investigating and recording diverse characters and character states encourages
generation of additional data, thus enhancing evolutionary hypotheses and thus
reinforcing classification criteria.
Phenetic groupings are achieved statistically via algorithms calculating relative distance
or percentage agreement or disagreement. The tree construction process is sequential,
either divisive or agglomerative. Agglomerative clustering, c o ~ e c t i n gat each level
individual taxa into pairs to maximise pairs or group members similarities. Phenetic
algorithms used in this study apply because the rates of change do not differ markedly
across lineages on a tree, thus, the degree of similarity could be a measure of
phylogenetic relatedness (Siddall, 1998).
The Hyacinthaceae has always presented a taxonomic challenge because it is deficient
in good diagnostic characters (Speta, 1998b; Stedje, 2001; Pfosser et al., 2003;
Manning et al., 2004). This is illustrated at subfamily level where five morphologically
similar subfamilies are delimited mostly on phytochemical data (Speta, 1%@a, 1998b;
Stedje, 2001; Pfosser et al., 2003). Phylogenetic analysis of the Hyacinthaceae has also
produced various hypotheses as to generic circumscriptions (Chapter 8, DNA).
Revisions done on Ledebouriinae taxa (Baker, 1896 & 1898; Jessop, 1970, 1972;
Venter, 1993; Stedje, 1994; Miiller-Doblies & Miiller-Doblies, 1997) produced
different taxonomic opinions. Character and character state allocation in revisions are
either inconsequent or entirely lacking, making taxa determination difficult. Recently
Manning et al. (2004) sunk Drimiopsis and Resnova into Ledebouria based on
molecular data. They characterise Ledebouria sensu lato by its "lack of bracteoles and
by its globose or top-shaped ovary containing two ovules per locule.. .. most species
have spotted leaves and ofien produce more than a single inflorescence per plant in one
growing season, and the bulb scales are often rather loosely packed and in many species
produce fine threads". Manning et al. (2004) also consider differences between
Ledebouria and Resnova qualitative and dismiss tepal differences in Drimiopsis as a
pollination adaptation.
Phenetic analysis of conventional leaf (Figure 3.8), flower (Figure 4.9), pollen (Figure
5.2) and compounds 1-7 data (Figure 6.4) characters from this study provide evidence
for demarcating Resnova, Ledebouria and Drimiopsis as separate (Table 9.4). In
addition, Resnova and Ledebouria cluster. This gives credence to Wetschnig &
Pfosser's (2003) results where a cladistic analysis of the trnL intron and the t r L - F
intergenic spacer grouped Resnova with one of the Ledebouria clades.
9.2 OBJECTIVES
This chapter investigates the classificatory significance of Drimiopsis characters and
their states accrued from preceding chapters in conjunction with those of the presumed
sister genera. The chapter also presents a circumscription of the Ledebouriinae genera
based on diagnostic morphological characters.
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
9.3 MATERIALS and METHODS
-
-
-
--
-
-
-
-
-
-
-
--
-
-
-
-
-
-
--
-
-
-
-
-
-
--
-
-
-
-
-
The characters and character states of species investigated (Table 9. l), compiled via
DELTA, are listed in Tables 9.2-9.3 together with the character matrix. Characters are
exported to STATISTICA 6.1 for cluster analysis with the following settings: tree
clustering; Ward's method of minimum-variance clustering under the amalgamation
rule and percentage disagreement as a measure of distance (Ward, 1963), delineating
homogeneous and distinct groups and based on assessment of distances (Davis, 1986).
Characters and character states at generic level (Table 9.4), compiled from diagnosable
characters indicated in Tables 9.2-9.3, are similarly analysed with the exclusion of
characters polymorphic for one of the three genera.
Table 9.1: Taxa whose characters and character states were investigated and analysed
phenetically.
Taxa
D. atropurpurea
D. atropurpurea
D. atropurpurea
D. atropurpurea
D. barteri
D. barteri
D. botryoides subsp.
I botryoides
I D. botryoides subsp.
I
I
botryoides
D. botryoides subsp.
botryoides
D. botryoides subsp.
botryoides
D. botryoides subsp.
botrvoides
D. botryoides subsp.
bohyoides
D. botryoides subsp.
hntrvoides
- - - . - - -.- -
I D. botryoides subsp.
botryoides
D. bottyoides subsp.
I
&
Accepted names Accession
voucher numbers
in this thesis
Rogers 18508
D. atropurpurea
Schrierp 1330
D. atropurpurea
Lebatha 049
D. atropurpurea
Van der Merwe
D. atropurpurea
0266 1
Lebatha 002
D. barteri
D. barteri
Greenway
&
Kaburi l4/782
Greenway 12854
D. botryoides
I D. botryoides
D. botryoides
I
I D. botryoides
D. perfoliata
South Africa
South Africa
South Africa
South Africa
Herbarium
Herbarium
Fresh
Herbarium
Tanzania
Tanzania
Fresh
Herbarium
Kenya
Herbarium
I Tanzania
Lebatha 099
Tanzania
Lebatha 003
Kenya
D. botryoides
I D. botryoides
Status
I Lebatha 098
D. botryoides
D. botryoides
Locality
I
Lebatha 004
I Reid 1090
I
1
1 Fresh
Fresh
Kenya
I Kenya
I Herbarium
Kenya
Herbarium
Tanzania
Fresh
Reid 1984
I R.B. & A.J. Faden
I
741505
Lebatha 00 1
. - - - . -- - --
I
1
I
D. burkei subsp. burkei
D. burkei subsp. burkei
D. burkei subsp. burkei
D. burkei subsp. burkei
D. burkei subsp. burkei
D. burkei subs^. burkei
D. burkei subs^. burkei
stolonissima
D. carrii
D. crenata
D.burkei
D. burkei
D. burkei
D. burkei
I D. burkei
I D. burkei
1 D. burkei
D. burkei
D. stolonissima
Lebatha 009
Lebatha 04 1
Lebatha 096
Lebatha 054
I Lebatha 103
1 Lebatha 056
1 Lebatha 095
Theron 1589
Fresh
1 Fresh
South Africa
I Botswana
1 South Africa
1 Botswana
South Africa
I Fresh
1 Fresh
1 Fresh
Lebatha 037
D. carrii
D. burkei
Lebatha 0 15
Codd 80 18
D. comptonii
D.davidsoniae
D. fischeri
Lebatha 079
Lebatha 038
Fischer 325
Lebatha 045
Lebatha 045
Lebatha 053
locali
1 D. kikiae
I
D. kikiae
D. lachenalioides
I
I
I
D. liniopapilla
D. maculata
D. maculata
D. maculata
D. maculata
D. kikiae
D. kikiae
-DLljniopapilla
R. lachenalioides
21 17
Baut 549
R. lachenalioides
D. liniopapilla
Lebatha 060
D. maculata
Lebatha 005
1 D. maculata
1 Lebatha 006
1 D. maculata
1 Lebatha 007
1 D. maculata
1 Lebatha 039
1 Tanzania
1 South Africa
South Africa
South Africa
Southifrica-
1 Herbarium
1 Fresh
-
-
- -
-~
Fresh
Fresh
-Herb&
I
South Africa
South Africa
Botswana
1 South Africa
I Botswana
I South Africa
I
Herbarium
1 Fresh
I Fresh
I Fresh
1 Fresh
1 Fresh
I
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
D. maculata
Lebatha 102
Lebatha 032
D. maculata
D. maxima
D. maculata
R. maxima
I Van Jaarsveld
D. maxima
D. pusilla
D. queae
R. maxima
D. pusilla
D. queue
D. queae
D. queae
D. queue
D. queae
D. queae
D. queae
D. reilleyana
D. queue
D. queue
D. queae
D. queue
D. reilleyana
D. rosea
D. rosea
D. rosea
D. rosea
D. woodii
R. humifusa
R. humifwa
R. humzjka
D. rosea
D. rosea
I D. woodii
R. humifusa
R. humifusa
R. humfusa
6010
Venter s.n.
Lebatha 078
Lebatha 055
Van der Merwe
s.n.
Repton s.n.
Liebenberg. s.n.
Rogers 2 14 09
Codd 5126
Lebatha 068
Chevalier no.
8432
Goossens 43
Codd s.n.
I Lang 32236
Schlechter 3 174
Devenish no.958
Van der Merwe
I South Africa
I Fresh
Herbarium
Fresh
Fresh
Herbarium
South Africa
Swaziland
South Africa
South Africa
South Africa
South Africa
South Africa
South Africa
Swaziland
Democratic
Republic of Congo
South Africa
South Africa
I South Africa
South Africa.
South Africa
South Africa
I
Herbarium
Herbarium
Herbarium
Herbarium
Fresh
~erba&m-
I
Herbarium
Herbarium
Herbarium
Herbarium
Herbarium
Herbarium
I
s.n.
-
R. lachenalioides
R. maxima
R. maxima
I R. maxima
I R. maxima
I R. rnegaphylla
I Ledebouria sp.
I Ledebouria sp.
Ledebouria sp.
L. asperifolia
L. asperifolia
L. asperifolia
R. lachenalioides
Singh 72
R. maxima
Lebatha 042
Lebatha 077
R. maxima
I R. maxima
I Lebatha 042
I R. maxima
I Lebatha 047
I R. megaphylla
I Lebatha 05 1
I Ledebouria sp. I Lebatha 010
I Ledebouria sp. I Lebatha 050
Ledebouria sp.
Lebatha 059
Lebatha 057
L. asperfolia
Lebatha 080
L. asperrfolia
L. asperijiolia
Lebatha 090
I L. concolor
I L. concolor
South Africa
South Africa
Swaziland
I South Africa
1 Swaziland
I South Africa
1 Botswana
I South Africa
South Africa
Swaziland
South Africa
Botswana
I Re: Venter (1993) I South Africa
Herbarium
Fresh
Fresh
I Fresh
I Fresh
I Fresh
I Fresh
I Fresh
Fresh
Fresh
Fresh
Fresh
1
1
I
I NIA
1
I
I
I
I Re: Venter (1993) I South Africa
IL. inquinata
I L. luteola
I
L. inquinata
I L. luteola
I Lebatha 075
I South Africa
I Re: Venter (1993) I South Africa
Fresh
I NIA
L. ovatifolia
L. ovatifolia
L. revoluta
L. ovatifolia
L. ovatifolia
L. revoluta
Lebatha 008
Lebatha 063
Re: Venter (1 993)
Botswana
South Africa
South Africa
Fresh
Fresh
NIA
L. sandersonii
L. socialis
L. sandersonii
L. socialis
Lebatha 085
Re: Venter (1 993)
Swaziland
South Africa
Fresh
N/A
Table 9.2: Data matrix for Schizocarphus nervosus, Drimiopsis, Resnova and Ledebouria. Characters 1-60 and their states are listedbottom of the table. Characters coded as - represent inapplicable characters.
S,nervosus
D. atropurpurea
D. barteri
D. botryoides
D. perfoliata
D. burkei
D. stolonissima
D. carrii
D. comptonii
D. davidsoniae
D. fischeri
D. kikiae
D. liniopapilla
D. maculata
D.pusilla
D. queae
D. reilleyana
D. rosea
D. woodii
R. maxima
R. megaphylla
R. humifusa
R. lachenalioides
L. asperifolia
L. concolor
L. floribunda
L, inquinata
L. ovatifolia
L, revoluta
L. sandersonii
L. socialis
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 I8 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 525354555657585!6[
3 2 1 I I 1 2 1 3 2 3 2 3 2 1 3 1 2 1 1 2 2 1 1 2 3 1 I I 1 2 - 2 - I 1 2 1 1 2 1 2 2 2 2 1 3 3 2 2 2 2 1 2 1 2 2 1 1 3
#6l. Inflorescence with < n h b e r of flowers> 1. 15 flowers or less, 2. 16 to 30 flowers, 3. more than 30 flowers; #62. Flowers distributed <flower density> 1. sparsely
distributed, 2 densely; #63.
<pedicel length> 1. minutely pedicellate (shorter than 0.1 cm), 2. shortly pedicellate (0.1 to 0.4 cm long), 3. with elongate pedicel (more
than 0.4 cm); #64. Rachis ?general length> 1. 10 cm or shorter, 2. 10.1 to 20 cm long, 3. more than 20 cm long; #65. Rachis crachis shape> 1. cylindrical, 2. conical, 3.
ovoid-cylindrical; #66. Peduncle <variegation> 1. banded, 2. colowed purplish, 3. spotted, 4. green, #67. Bracts in mature inflorescence <presence or absence> 1. absent, 2.
vestigial, 3. developed; #6$. Prophylls <present or not> 1. absent, 2. present; #69. Flower <size> 1. minute (1-2 mm), 2. small (2.1-4 mm), 3. medium-sized (4.1-6 mm), 4.
large (more than 6 rnm); #70. Flower <type> 1. actinomorphic; #71. Flower <number of tepals> 1. sextepalous; #72. Flowers <shape> 1. coronate to stellate, 2.
campanulate, 3. tubular; #73. Flowers with hypanthium base <shape> 1. truncate, 2. obtuse, 3. rounded; #74. Tepals <type> 1. isomorphic, 2. dimorphic; #75. Tepals with
hypanthium Chypanthiwn Size> 1 inconspicuous, 2. hypanthium conspicuous; #76. Tepals <colow> 1. whitish to greenish, 2. purplish green, 3. creamy-brownish, 4. pink, 5.
purple/blue; #77. Outer w@l of tepals <posture> 1. connivent, 2. recurved, 3. drooping; #78. Outer whorl of tepals longitudinal posture <outer> 1. cucullate, 2. flat; #79.
Outer whorl of tepals apicqlly <apex margin shape> 1. conduplicate, 2. flat; #80. Inner whorl of tepals <posture> 1. connivent, 2. recurved, 3. drooping; #8 1. Inner whorl of
tepals longitudinally <post$re> 1. cucullate, 2. flat; #82. Inner whorl of tepals apically <apex margin shape> 1. conduplicate, 2. flat; #83. Vitta, 1. conspicuous, 2. faint, 3.
absent; #84. Androeciuml <number> 1. 6, #85. Androeciwn <colow> 1. greenish to whitish, 2. cream, 3, maroonishfpurplish; #86. Androecium <posture> 1. erect, 2.
spreading; #87. Androeciurn <perianth adnation> 1. epitepalous; #88. Androecium <arrangement on tepals> 1. uniseriate, 2, biseriate; #89. Adroecium <insertion on tepab
1. inserted at throat of peripth tube; #90. Androecium <length> 1. shorter than pistil, 2. as long as pistil, 3. longer than pistil; #91. Filaments <cohesion> 1. free, 2. valvate;
#92. Filaments <shape> 1I. deltoid to accuminate, 2. lanceolate, 3. filiform; #93. Anthers <attachment> 1. dorsifixed; #94. Gynoecium <number of carpels> 1. tricarpellate,
2. polycarpellate; #95. O h l e s <number per locule> 1. two per locule, 2. more than two per locule; #96. Stigma <shape> 1. roundish, 2. triangular; #97. Stigma papilla <
arrangemenv 1. stalked, subsessile, 3. sessile; #98. Stigma papilla shape <stigma papilla shape> 1. round, 2. trilobal; #99. Style <size>l. shorter than ovary, 2. as long as
ovary, 3. longer than ovary; #loo. Style <style shape> 1. terete, 2. triangular; #101. Ovary <attachment> 1. sessile, 2. stipitate; #102. Ovary <ovary shape> 1. globose, 2.
ovoid to oblong, 3. conicial; #103. Ovary transversely <ovary shape> 1. smooth, 2. with ridge below style, 3. severally lobed; #104. Ovary <ovary colow> 1. whitish,
greenish, 2. purplish, 3. b'luish; #105. Ovary shoulders <present or absent> l.absent, 2. present; #106. Nectaries Cabsent or present> 1. absent, 2. present; #107. Pollen
<type> 1. isomorphous, monosporous, 2. heteromorphous, heterosporous; # 108. Pollen equatorial view <shape> 1. depressed ovate, 2. ellipsoid; # 109. Pollen polar view
<shape> 1. elliptic, 2. nwowly elliptic; # 110. Pollen laterally <shape lateral view> 1. blunted, 2. tapered; # I l l . Pollen <equatorial diameter> 1. subequiaxe, 2. breviaxe;
#112. Pollen distal pole $type> 1. straight, 2. curved; #113. Pollen sexine <type> 1. smooth, 2. rough; #114. Pollen ornamentation <type> 1. punctate, 2. reticulate, 3.
punctate-reticulate; #I 1: Flowering <time> 1. March to May, 2. September to December, 3. July to August; #116. Distribution <regions> 1. southern Africa, 2. tropical
Africa, 3. both regions. ;
lower
1
a'.
Table 9.4: A list of diagnostic characters for the genera Drimiopsis (D), Resnova (R) and Ledebouria (L). Characters that coded polymorphic are
indicated in brackets but were excluded in the generic delimitation phenetic analysis (Figure 9.2).
Character coding
Character
1
Plants
Character state
often solitary = 0; gregarious = 1
2
Bulb scale arrangement
loosely packed = 0; compact = 1
3
Bulbs below or above ground
hypogeal = 0; epigeal = 1
4
Bulb vegetative reproduction
stoloniferous= 0; nonstoloniferous= 1
5
Bulb scales colour
whitish, greenish = 0; brownish, purplish = 1
6
Outer bulb scales
rarely present = 0; persistant =1
7
Bulb scale texture
fleshy = 0; papery = 1
8
Bulb size approximation
small = 0; medium = 1;large = 2
9
Bulb scales threads
without threads when torn = 0; with thread when torn = 1
10
Leaves threads
without threads when torn = 0; with thread when torn = 1
11
Pseudopetiole
present
12
Leaf epidermal cell size
adaxial shorter than abaxial = 0; adaxial as long as abaxial = 1
13
Leaf epidermal cell wall outline
sinous = 0; straight = 1
14
Stomata guard cells H-complex
absent = 0; present =1
15
Stomata frequency
frequent = 0; not frequent = 1
16
Stomata crypts
shallow = 0; raised = 1
17
Lamina indumenta
frequent =O; not frequent = 1
18
Inflorescence type
simple raceme = 0; pseudo-corymb raceme = 1
19
Inflorescence number
mostly solitary = 0; rarely solitary = 1
20
Pedicel length
minutely pedicellalte = 0; shortly pedicellate = 1; pedicel longer than 0.4 cm = 2
21
Inflorescence posture
raceme erect = 0; raceme spreading = 1
= 0; absent =
1
I
Inflorescence length
I
I
Bracts, prophylls in mature qorescence
Flower hypanthium base shapp
I
Tepal length
I
I
I
I
Tepal type
Tepal transverse
I
Tepal posture
I
I
I
Tepal apex
Tepal apex margin
I
I
Tepal colour
I
Stamen
I
Stamens posture
I
I
Stamen arrangement
I
I
Stamen length
I
I
Filament shape
I
Filament length
I
Filament base
I
I
Anther mobility
I
I
Thecal arrangement
Stigma shape
I
Stigmatic papilla
I
Papilla shape
I
Papilla surface
I
I
I
Style type
Style size
I
Ovary shape
I
Ovary shoulders
I
I
ovary
I
Ovary attachment
I
Pollen grain type
I
Pollen shape from equatorial view
Pollen shape from polar view
Pollen shape from lateral view
Pollen equatorial diameter (
I
Pollen distal pole
I
Ornamentation
I
Geographic distribution !
I
I
I
I
I
I
I
I
I
*, =, to slightly longer than leaves = 0; rarely considerably longer than leaves = 1; considerably longer than leaves = 2
[Ol]
0
2
vestigial = 0; rarely present = 1; developed = 2
[Ol]
0
2
truncate = 0; obtuse = 1;rounded = 2
2
1
0
<5mm=o;>5<15mm= 1; >15mm=2
0
1
LO21
isomorphic = 0; dimorphic = 1.
cucullate = 0; campanulate = 1.
connivent = 0; recurved = 1; drooping = 2.
cucullate = 0; acute = 1;
conduplicate = 0; flat = 1.
can be blue = 0; never blue = 1
free = 0; valvate = 1; hsed = 2.
erect = 0; spreading =1
uniseriate = 0; biseriate = 1;rarely biseriate = 2
as long as pistil = 0; longer than pistil = 1.
deltoid to acurninate = 0; lanceolate = 1; filiform = 2.
shorter than tepals = 0; longer than tepals = 1.
broad = 0; thin = 1.
versatile = 0; rigid = 1.
parallel = 0; oblique = 1.
roundish = 0; triangular = 1
sessile = 0; subsessile = 1; stalked = 2.
trilobal = 0; round = 1.
rough = 0; smooth = 1.
terete = 0; triangular = 1.
shorter than ovary = 0; as long as ovary = 1; longer than ovary = 2.
globose = 0; ovoid to oblong = 1; conical = 2.
absent = 0; present = 1.
smooth = 0; lobed below style = 1; severally lobed = 2.
sessile = 0; stipitate = 1.
Isomorphous = 0; heteromorphous = 1
depressed ovate = 0; ellipsoid = 1.
elliptic = 0; narrowly elliptic = 1.
blunt = 0; tapering = 1.
subequiaxe = 0; breviaxe = 1.
straight = 0; curved = 1.
punctate = 0; reticulate = 1; punctate-reticulate= 2.
southern Afiica = 0; tropical Africa = 1; rarely both regions = 2; outside of Africa = 3
1
0
0
0
0
1
[oil
0
0
0
0
0
0
1
0
0
0
0
0
0
1
1
1
1
1
0
0
1
1
1
0
1
1
1
1
1
1
0
0
1
1
1
1
1
0
Lo11
1
1
1
1
1
0
0
[oil
2
[oil
1
0
[021
Lo11
[I21
1
2
[oil
1
0
1
1
2
1
1
1
[I21
2
1
2
1
1
1
1
1
1
1
[I21
[013]
0
0
0
0
0
0
0
0
0
0
0
0
0
[Ol]
Tree Diagram for 31 taxa within the Ledebouriinae
S. nervosus
L. asperifolia
L. jloribunda
L. rev0 luta
L. inquinata
L. ovati$olia
L. sandersonii
L. concolor
L. socialis
R. maxima
R. lachenalioides
R. humijiusa
R. m egaphylla
-
'
1
J
-
0
1
3
2
5
4
Linkage Distance
Figure 9.1: A phenogram computed from analysis of 116 characters and their states (Table 9.29.3) in Drimiopsis, Resnova and Ledebouria.
Tree Diagram for Drimiopsis, Resnova and L e u r i a
Drim iopsis
Resnova
Ledeboun'a
.
0.4
0.5
0.7
0.6
0.8
1.O
0.9
Linkage Distance
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Figure 9.2: A phenogram computed from analysis of 36 non-polymorphic characters (Table
9.4) in Drimiopsis, Resnova and Ledebouria show that Resnova has more in common with
Ledebouria than with Drimiopsis.
9.4 RESULTS and DISCUSSION
9.4.1 Phenetic analysis
The phenogram computed from Tables 9.2-9.3 matrix results in two main clusters
(Figure 9.1). Firstly, the Ledebouria and Resnova cluster and secondly, the Drimiopsis
cluster that forms two primary clusters: the uppermost cluster housing southern African
taxa and the bottom one tropical African taxa except the southern Afiican D. carrii and
D. stolonissima. Drimiopsis barteri, D. maculata and D. rosea occur in both regions.
The primary uppermost Drimiopsis cluster separates taxa with abaxially purply tinted
pseudopetiole (D. liniopapilla, D. queae and D. kikiae) from predominantly sessile
leaved taxa (bar D. atropurpurea, D. woodii). Drimiopsis atropurpurea and D. woodii,
possessing pseudopetiolate hairy leaves, and a banded peduncle group with D.
comptonii with spathulate papillate leaves. The cluster housing D. burkei, D.
davidsoniae and D. reilleyana possesses fine papilla, except D. pusilla that is hairy.
The primary lowermost Drimiopsis cluster separates taxa with thick textured large
leaves (D. botryoides, D. perfoliata, D. stolonissima, D. carrii, D. maculata) fiom the
linear and thinner leaved D. barteri, D. rosea and D. fwheri.
The results of the phenogram (Table 9.1) support the generic status of Drimiopsis,
Resnova and Ledebouria. Resnova has more in common with Ledebouria than with
Drimiopsis. The phenetic groupings of subspecies question the validity of creating
subspecies in formerly D. burkei that is housed in a different cluster separate from its
sister taxa. The phenogram does however support the division into subspecies ranking
of Drimiopsis botryoides. The overall results reveal a degree of variation between the
subspecies comparatively
similar to
other taxa. Drimiopsis
botryoides is
morphologically distinct from D. perfoliata in size, leaf form, shape and orientation and
number of inflorescences.
9ALGeaericcircumsc_ription-
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Analysis of 36 diagnostic Drimiopsis, Resnova, and Ledebouria characters (Table 9.4)
generated a phenogram (Figure 9.2) that separate the three genera but group Resnova with
Ledebouria. Twenty-two polymorphic characters (Table 9.4: 1, 3, 8, 12-14, 16, 18, 20-
23, 25,27, 29, 32-34, 37, 46, 52, 57, 58) were excluded in the analysis. This supports
the species level analysis (Figure 9.1).
Drimiopsis and Resnova bulbs are whitish-greenish with bulb scales loosely packed
while those of Ledebouria are purplish brown and compact.
In Resnova and Ledebouria the leaves are never pseudopetiolate. The epidermal cell
shape in Drimiopsis is unique with adaxial cells shorter than those abaxially and
possessing an overall polygonal shape while the abaxial cells are tetragonal and
elongate. Stomata form a diagnostic 'H' complex with subsidiary cells in Drimiopsis.
The stomata are less frequent than in the other Ledebouriinae.
Ledebouria and Resnova inflorescences are flaccid. The Ledebouria inflorescence,
unlike in Drimiopsis and Resnova, is rarely solitary, commonly more than two per bulb.
Bracts and prophylls are often present in the mature Ledebouria inflorescence. The
shape of the hypanthium is rounded in Drimiopsis, obtuse in Resnova and truncate in
Ledebouria. The stamens in Drimiopsis and Resnova are exclusively epitepalous and
either epitepalous or free in Ledebouria. Drimiopsis and Resnova possess erect
stamens, whereas in Ledebouria they may either be erect, patent or connivent. The
stamens of Drimiopsis are more or less equal in length, possess deltoid filaments and
are valvate. Those in Resnova are lanceolate and biseriate. In Ledebouria the stamens
are filiform and appear to be either equal or unequal in length.
1
I
Drimiopsis possesses terete styles, Resnova and Ledebouria triangular. Drimiopsis taxa
possess globose and sessile ovaries, Resnova ovoid to oblong ovaries with a short stipe
and ridges on the shoulders, and Ledebouria conical and conspicuously stipitate ovaries
with shoulders. The style in Drimiopsis is as long as the ovary, in Resnova it is shorter
than, and in Ledebouria, longer than the ovary. In Drimiopsis the stigma is trilobed
with sessile papillae, in Resnova round and shortly stalked, and in Ledebouria round
and conspicuously stalked.
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In equatorial view, pollen grains of Drimiopsis are depressed ovate, those of Resnova
and Ledebouria are ellipsoid; the polar view of Drimiopsis pollen is elliptic while that
of Resnova and Ledebouria are narrowly elliptic. Drimiopsis pollen is subequiaxe and
possesses a linear pole opposite to the sulcus as well as blunt lateral sides. The pollen in
1371
Resnova and Ledebouria is breviaxe and possess a tapered pole opposite to the sulcus
as well as tapered lateral sides. Ledebouria is heterosporous while Drimiopsis and
Resnova are isosporous.
9.5 CONCLUSION
Unambiguous intergeneric differences exist in the Ledebouriinae. At macro level, the
bulb, leaf, inflorescence, flower and pollen characters within the Ledebouriinae
complex are diagnostic. At micro level, epidermal cell arrangement and size, cell wall
shape, stomata and trichomes, are taxonomically significant characters at generic level.
Keeping the limitations of a phenetic analysis in mind, the subspecific status of taxa is
not supported in this study. Drimiopsis botryoides Bak. subsp. botryoides should be
ranked with D. botryoides Bak. and D. botryoides Bak. subsp. prostrata Stedje with D.
perfoliata Bak. Similarly, D. burkei Bak. subsp. burkei is D. burkei Bak. and D. burkei
Bak. subsp. stolonissima U. & D. Muller-Doblies is D. stolonissima (U. & D. MullerDoblies) Lebatha (Chapter 12, Taxonomic treatment).
The result of this analysis raises questions about views in support of sinking Resnova
under Drimiopsis (Phillips, 1951; Jessop, 1970, 1972; Dyer, 1976; Arnold & De Wet,
1993; Meyer and Williams, 1997) and sinking both Resnova and Drimiopsis under
Ledebouria (Manning et al., 2004). However, morphological differences between
Drimiopsis, Resnova and Ledebouria are significant enough to warrant generic status.
The differences cannot be simply dismissed as qualitative and adaptive.
The intergeneric variations within the Ledebouriinae complex can be interpreted in one
of two ways: 1. For the splitter, Resnova, Ledebouria and Drimiopsis are three separate
genera, with Resnova having more in common with Ledebouria than with Drimiopsis.
10. CLADISTIC ANALYSIS
10.1 INTRODUCTION
Cladistics is one of the analytical procedures leading to the major product of a
systematic analysis, a hypothesis of relationships. Unlike phenetics, cladistic analysis
can identify polyphyletic or paraphyletic taxa. Cladistics analyzes data so as to portray
clades (cladograms), an easily verifiable criterion for taxa delimitation and phylogenetic
inference. A cladistic analysis begins with an existing taxonomic system followed by a
selection of taxonomic characters and character states and aims for a fully dichotomous
branching scheme, not because speciation occurs that way, but because such schemes
maximise their information content (Schuh, 2000).
Cladistics (Hemig, 1965, 1966; Wagner, 1969; Scotland, 1992) infers phylogeny
through shared derived characters. One of the assumptions of cladistics is the common
ancestry of all organisms with character lineages formed over time. The two most
popular approaches to cladistic analyses are parsimony and maximum likelihood. This
is due to their character distribution explaining capabilities (Siddall, 1998). The
methods, unlike mathematical clustering algorithms in phenetics, are phylogenetically
defendable.
Parsimony has to do with presuming that there is a single origin of similar structures in
the absence of evidence to the contrary (Schuh, 2000). Parsimony accepts that the most
likely phylogenetic hypothesis is the one that requires the least number of character state
transitions-the
least number of evolutionary assumptions ( e g Felsenstein, 1978 &
1983; Farris, 1983, 1986; Kluge, 1997). According to Felsenstein (198 1), parsimony
assumes that character state modifications are essentially unlikely events, that
homoplasy, though existing in nature, is unlikely or rare Farris (1983) counters by
stating, "procedures that minimize something do not have to presuppose the quantity
minimized is rare".
Character polarity is established through the outgroup method. Relationships are then
hypothesized through studying the distribution pattern of these characters, usually
forming a hierarchy of nested groups/clades.
'Cladogams and dendograms are similar in their assumption that hierarchical patterns
exist in nature. Dendograms, by their very nature, result in a loss of information because
no characters or states are plotted on them. Cladograms, on the other hand, make full
use of character-taxon matrices by plotting relevant characters and states on them.
Maximum likelihood is found more applicable in cases where parsimony fails, where
the hypothesis that change is rare does not apply and evolutionary rates along differing
evolutionary branches varies. Maximum likelihood assumes that evolutionary change
occurs by chance, thus each possible phylogeny must have a certain probability of being
the correct phylogeny. It seeks the most probable explanation, not necessarily the best
(Swofford et al., 1996; Siddall, 1998). This method usually yields trees similar to those
of parsimony.
Homoplasy, polymorphism or incomplete data sets can generate multiple trees, hence
their ideal exclusion in a cladistic analysis. Several consensus techniques exist that
objectively compare multiple trees.
The bone of contention in classification lies in the question of naturalness (Stuessy,
1990 & 1997), and parsimony is adopted in this study as the best way to explain natural
phylogeny. Cladistics defines monophyletic groups as composed of all inferred
descendants of an ancestral species (Hennig, 1966; Mayr & Aslock, 1991; Judd et al.,
2002; Brurnrnitt, 2002). A group including only some of the descendants is paraphyletic
and rejected by cladists. They rearrange the paraphyletic groups into one or several
clades, masking the degree of phenotypic differentiation in taxa circumscription. The
use of only monophyletic groups in classification runs the risk of excluding key
characters related to descent.
This study highlights the opinion that "taxonomy must depend on characters related to
descent, not simply on lines of descent" Brumrnitt (2003: 803). This creates groups
which aTe m o n ~ p k y l efor
t ~ -taxonomists but paraphyletic for-cladists. Incongruences
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between taxa or clade groupings are a product of the cladistic method and not an error in
the taxonomic system. Ebach & Williams (2004) argue on the semantics of paraphyly
and monophyly. They maintain paraphyly and polyphyly are paralogous; paraphyly
refers to the absence of monophyly and is therefore not a state. I believe analysis of
nature can result in well-supported informative (monophyletic) or uninformative
(paraphyletic) groupings.
The selected outgroup, Schizocarphus nervosus (Burch.) v.d. Merwe. has been
hypothesised to be the sister group of the Ledebouriinae U. & D. Miiller-Doblies by
Pfosser & Speta (1999), Wetschnig et al. (2002), Pfosser et al. (2003) and Wetschnig &
Pfosser (2003) based on trnL-F data. The atpB gene indicates Merwilla Speta as a
potential outgroup (Pfosser et al., 2003). The results from analysis of the trnL-F & rbcL
genes also suggest S. nervosus as the outgroup (Manning et al., 2004). Furthermore,
seed characters also support S. nervosus as an immediate Ledebouriinae sister group
(Pfosser et al., 2003). Van der Menve (1943) separated Scilla rigidifolia Bak. that
possessed stellate flowers with greenish tinged apices, subequal segments and fibrous
bulb apices into the new genus Schizocarphus v.d. Merwe. This was later transferred to
Scilla L. (Jessop, 1970; Stedje & Thulin, 1995). Molecular data separates Scilla nervosa
(Burch.) Jessop form other SciNa hence the subsequent transference to Schizocarphus
nervosus (Stedje, 1998; Pfosser & Speta, 1999; Wetschnig & Pfosser, 2003; Manning et
al., 2004).
10.2 OBJECTIVES
To perform a cladistic analysis of Drimiopsis Lindl. & Paxt. and infer phylogeny based
on morphology and a total evidence approach within the Ledebouriinae.
10.3 MATERIALS and METHODS
Taxa analysed are listed in Table 10.2 & 10.3. The matrix of 105 morphological
characters and character states coded for 31 taxa generated through DELTA (using the
TOHEN directive) was analysed cladistically using NONA (Goloboff, 1999) via
WINCLADA (Nixon, 2000). Data were analysed with using the following settings:
heuristics with 2500 maximum trees to keep, 500 replications, five starting trees per
replication, a repeated unconstrained search strategy through tree searching using the
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multiple TBR
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+ TBR (muFmaxx*) search strategy.
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I% striet consensus trge w a s
generated in the event of more than one equally parsimonious tree being present.
Bootstrap support values were calculated via 1000 replications with 10 repeats on all
trees.
In addition, available trnL-F gene sequences for taxa for which there was
morphological data were obtained from National Centre for Biotechnology Information
(NCBI, 2004). Sequences were aligned using the ClustalW Multiple Alignment option
(Thompson et al., 1994) in Bioedit (Hall, 2004). Indels were encoded using Gap
Recorder (Ree, 2004). Cladistically uninformative transition/transversion characters
were excluded a priori from the matrix, while uninformative morphological and indel
data was deactivated for the analysis. The resultant matrix is provided in Table 10.4.
The matrix was analysed variously, first using only the morphological data, then
transition/transversion data and indel data and thirdly a total evidence approach by
combining the two.
10.4 RESULTS and DISCUSSION
10.4.1 Cladistic analysis of morphological data
The 105 characters initially initiated via DELTA were scored for thirty-one taxa (Table
10.2). Eleven characters were uninformative. Of the remaining 94 characters, 38 were
multistate, 14 were coded additive and twenty-four nonadditive (Table 10.1). The
analysis resulted in four most parsimonious trees with tree lengths of 427 steps. Two
nodes, confmed to the Ledebouria clade, collapsed in the 431 steps long strict consensus
tree of a Consistency Index (Ci) of 0.32 and a Retention Index (Ri) of 0.66 (Figure
10.1). The low consistency and retention indices are indicative of the high amounts of
homoplasy and lack of hierarchical characters in the analysis.
Table 10.1: List of 38 multistate characters and their character coding: 24 characters were coded
nonadditive and 14 characters additive. The characters and states are explained in Table 10.2.
Nonadditive
Oro12
2
10[0.12
13[012
15[012
~
24[0123
32[012
~
35[012
43[012
~
61[012
63ro12
142
----
---
The high rates of homoplasy result in the majority of species being defmed by a suite of
homoplasious characters. These characters show discrete distributions, making them
effective in delimitating taxa. Drimiopsis species possess few autapomorphies, these
being confmed to D. botryoides possessing a falciform leaf form 16[1]; D. macu/ata
possessing a rachis 11 to 25 cm long 60[2]; D. comptonii possessing a spathulate leaf
form 16[3] and an oblanceolate leaf shape 17[2];D. fischeri lacking bracts in the mature
inflorescence 63[0]; D. pusilla possessing a crenulate leaf margin 24[3] and D. kikiae
possessing a pseudopetiole conspicuously shorter than the lamina 22[0].
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Figure 10.1: Strict consensus tree of four equally parsimonious trees from a cladistic analysis of data as
presented in Tablel0.2. Values in squares represent bootstrap support values. A-C: the major clades within
Drimiopsis.
143
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--
Table 10.2: Matrix of 105 morphological characters and their states for 30 Ledebouriinae taxa
and Schizocarphus nervosus as outgroup. The characters and states are explained in Table 10.3.
TABLE 1. Characters
S. nervosus
D.
D.
D.
D.
atropurpurea
barteri
botryoides
perfoliata
D. burkei
D. stolonissima
D. carrii
D. comptonii
D. davidsoniae
D. fischeri
D. kikiae
D. liniopapilla
D. maculata
D. pusilla
D. queae
D. reilleyana
D. rosea
D. woodii
R. maxima
R. megaphylla
R. humifusa
R. lachenalioides
L. asperifolia
L. concolor
L. floribunda
L.
L.
L.
L.
L.
inquinata
ovatifolia
revoluta
sandersonii
socialis
TABLE 2.
0-62
o
5
10
15
20
25
30
35
40
45
50
55
60
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2100102121210201001101--00001-1-001010111102211111-010002212103
110110010010120001121021011001001100010001112111210000001001010
1101100110211010203011--01001-1-12100100011000
0001210022
2101110000101201102111--0110011-12110100010020
0012100103
1101100000001102011211--01101-1-12110100011120
0002100103
1101100110101121201111--11000101120001000111011101-100001100012
1101000010101201201111--0110011-12100100011100
0002110013
1101000000101102011111--1100011-12110100011100
0000110003
0101000100101120321111--01001-1-201001000111210021-100102002023
1101100100101200201111--11101-1-221201000111211011-000001000013
1101100100201200201011--01011-1-02100100011120
0002102013
1101110000101210001110011110010000000100011100
0001101012
11011101001010100011101111101-010000110001111101210010001110021
2101110010001201012210202101011-01101100011100
0012101223
0101100100101200200011--31101-00120011000111211111-000001111011
01011001001010100000101111101-0000011100011100
0002101001
1101100110101200201111--21101-00120011000111211111-100001110013
0001110100101011200011--21011-0102000100011000
0002210101
11011000001012002110101021011-1-1010010001112110111100002102110
2101100111101201202111--21001-1-021011110000010011-100002111113
1101110010001102011111--01100000120111110000011021-010012101013
1101110110101102201111--01100100221211110000010011-100002101111
1101110010101201202111--0110011-001111110000210111-100001111001
1101100011211201202101--01100100101211110000012011-101011212121
2110102111211201202111--01101-1-20111111010000
1012202103
2100102121211201203101--2100000110101111010020
1011202101
1100111111210201201001--21001-0110101110010010
1010212112
1100101111210202011101--0110010010101101010000
1012212123
2101101121210201202201--2100010010100101010000
1011202102
0100102111211202200011--0110010010001111010020
0012112113
2111101111211201201011--0110010010110011110000
0012102103
Characters
63
S. nervosus
D.
D.
D.
D.
atropurpurea
barteri
botryoides
perfoliata
D. burkei
D. stolonissima
D. carrii
D.
D.
D.
D.
D.
D.
D.
comptonii
davidsoniae
fischeri
kikiae
liniopapilla
maculata
pusilla
D. queae
D. reilleyana
D. rosea
D. woodii
R. maxima
R. megaphylla
R. humifusa
R. lachenalioides
L.
L.
L.
L.
L.
L.
L.
L.
asperifolia
concolor
floribunda
inquinata
ovatifolia
revoluta
sandersonii
socialis
63-104
68
73
78
83
88
93
98
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211020102112112110120211100111000111011110
101101010000000000110000021100000100000000
101101100000000000110000021100000100000000
101101101000001000111000021100000100000000
101101101000000000111000021100000100000000
100101000000000000111000021100000100000000
101101000000000000111000021100000100000000
100101000000000000110000021100000100000000
100101040000000000110000021100000100000000
100101020000000000110000021100000100000000
003201101000000000010000021101100100000000
100101020000000000111000021100000100000000
100101000000000000110000021000000100000000
102101001000000000111000021100000100000000
100101010000000000110000021100000100000000
100101021000000000110000021100000100000000
100101000000000000110000021100000100000000
100101000000000000110000021100000100000000
101101001000000000110000021100000100000000
103110031111110101120100110001111101111101
100210141111110001120100110001111101111101
101111112111110101120100110001111101111101
103110132111110101120100110001111101111101
200120041111110200100200100211221111111112
101120001111110000120200100211221111111112
211020031111110200110200100211221111111112
211120041111110100100200100211221111111112
212120041111110200110200100211221111111112
211120131111110200110200100211221111111112
202120031111110200110200100211221111111112
201120031111110100120200100211221111111112
103
I
144
Table 10.3: The character list used to create the matrix (Table 10.2) for a cladistic analysis.
Characters and character states are analysed with inference on their evolutionary development
based on the strict consensus cladogram furnished in Figure 10.1. The number in brackets in the
first column is the character number as used in DELTA. Figures 10.4-10.27 illustrate
transformations of selected characters.
Character and character state
Proposed hypothesis on character state transformations
Character transformations are ambiguous from the outgroup to the
0(1). Plants <size>
basal branches in the Ledebouria clade. The outgroup, both the basal
O.dwarfed (less than IOcmhigh)
1. medium-sized (10.1 to 15 cm and terminal entities in Ledebouria as well as the terminal taxa in
Clade A are hypothesized to be robust. The majority of Resnova and
high)
Drimiopsis
are medium sized except R. maxima, which is also robust,
2. robust (more than 15 em high)
and the dwarf D. comotonii, D. rosea, D. ousilla and D. queae.
Uninformative and excluded from the analysis.
1(2). Plants <development or
maturation>
O.hysteranthous
1. protantherous to synantherous
2(5). Bulbs <below or above ground>
Hypogeal bulbs are plesiomorphic, transforming into epigeal in 1.
concolor and 1. socialis (Figure 10.12).
O.hypogeal
1. epigeal
Gregarious bulbs, largely confined to Drimiopsis, but also occurring in
3(6). Bulbs <habit>
1. socialis, 1. revoluta and 1. asperifolia, are derived.
O.solitary
1. gregarious
The non-stoloniferous character state is plesiomorphic but its
4(7). Bulbs <vegetative reproduction>
O.stoloniferous
development in Drimiopsis is ambiguous in D. stolonissima and D.
1. non-stoloniferous
carrii and homoplasious in D. comptonii.
5(8). Bulbs with tuberescent fundus The absence of a tuberescent fundus is plesiomorphic. The possession
<type>
of a tuberescent fundus is a homoplasious character state having
O.absent
developed independently in 1. inquinata, D. rosea and the D.
botryoides and D. maculata clade. Due to the ambiguity in the data,
1. present
either state may be homoplasious in Resnova or the terminal branches
of Drimioosis.
6(9). Bulbs <colour>
Brown bulbs are plesiomorphic and are absent from Resnova and
O.whitish
Drimiopsis, which only possess white bulbs. Both brown and white
bulbs are homoplasious: the former as a result of a hypothesized
1. purplish
2. brown
reversal in 1. floribunda and the latter as a result of parallel evolution
in Clade B & C and 1. asperifolia. Similarly, purplish bulbs are also
homoplasious due to parallel evolution within Ledebouria (Figure
10.13).
Ovoid bulbs are plesiomorphic. Roundish bulbs are homoplasious,
7(10). Bulbs <shape>
O.roundish
having developed in representatives of Ledebouria and Resnova as
1. ovoid
well as D. kikiae, D. woodii, and the basal clade A (Figure 10.14).
8(11). Bulbs <diameter>
Ambiguity exists as to whether the common ancestor to taxa analysed
O.small (2.5 cm wide or less)
possess large or medium sized bulbs. The large bulbs in the terminal
1. medium sized (2.6 to 5 cm wide) clade of Ledebouria represent either a parallel or reversal event. All
Resnova analysed possess medium sized bulbs. The medium sized
2. large (more than 5 cm wide)
bulbs in D. maculata are hypothesized to be due to a reversal. Small
bulbs apparently developed independently in all three clades of
Drimioosis.
9(13). Bulb scales <packing>
Compact bulb scales in S. nervosus and Ledebouria evolved to loosely
O.loosely packed
packed scales in Resnova and Drimiopsis with a reversal to compact in
1. compact
R. maxima (Figure 10.8).
10(14). Bulb outer scales <colour>
Brownish-purple outer bulb scales are plesiomorphic. Initial
O.greenish
transformation is to white in Resnova (advancing to greenish in R.
1. white
megaphylla) and Drimiopsis followed by parallel reversals to brownishpurple in D.fischeri and D. barteri and either a parallel development to
2. brown/purple
greenish in D. peifoliata and D. maculata or a reversal to white in D.
botryoides.
145
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11(15). Bulb outer scales <outer
texture>
O.fleshy
1. membranous
12(16). Bulb scales when tom <with
threads or not>
O.with threads
1. without threads
13(17). Leaves <number>
O. 1 (monophyllous)
1. 2 (diphyllous)
2.3 (polvphvllous)
14(18). Leaves < number variation>
O.number of leaves unvarying
1. sometimes diphyllous
2. sometimes polyphyllous
15(19). Leaves <posture>
O.erect
1. spreading
2. appressed to the ground
16(20). Leaves <form>
O.cordiform
1. falciform
2. linear
3. spathulate
17(21). Leaves <shape>
O.lanceolate
1. ovate
2. oblanceolate
Membranous outer bulb scales is plesiomorphic and evolves to fleshy
in Drimiopsis and Resnova but re-emerges in D. barteri.
There is ambiguity in the data as to whether the presence or absence of
threads is the plesiomorphic state. Similarly, ambiguity exists as to
whether subsequent transformations represent reversals or parallel
developments. The production of states is confined to S. nervosus and
1. inQuinata,1. ovatifolia and 1. revoluta.
PolyphyllOus leaves are plesiomorphic. Diphyllous leaves develop
independently in Resnova and each of the three Drimiopsis clades.
Monophyllous leaves also represent a parallel development in D.
barteri and D. rosea as well as D. liniopapilla and D. queae.
Taxa with no variation in leaf number is plesiomorphic. Monophyllous
taxa that are sometimes diphyllous, as well as diphyllous taxa that are
sometimes polyphyllous, represent parallel developments with
Drimiopsis.
Spreading leaves is plesiomorphic. Appressed leaves have evolved
several times in Ledebouria, Resnova and possibly also Drimiopsis.
Erect leaves are confined to Clade B & C in Drimiopsis.
Ambiguity (whether cordiform or linear leaves are plesiomorphic) at
the base of the tree results in the homoplasious distribution of
cordiform leaves being ascribed to either parallelisms or reversals.
Falciform leaves is an autapomorphy of D. botryoides and spathulate
leaves is an autapomorphy of D. comptonii (Figure 10.15).
Lanceolate leaves are plesiomorphic. Ovate leaves developed parallel
in 1. ovati/olia, R. megaphylla, D. wood;;, D. atropurpurea and in D.
carrii, D. perfoliata and D. maculata with a postulated reversal to
lanceolate in D. botryoides. Oblanceolate leaves is an autapomorphy of
D. comvtonii(Figure 10.16).
18(24). Leaves <length>
0.5cm
1. 5.1 to 10 cm
2. 10.1 to 20 cm
3. longer than 20 cm
Leaves 5.1 to 10 cm long is plesiomorphic. Short leaves (1-5 cm long)
developed three times independently in Drimiopsis Clade C and 1.
sandersonii. Leaves between 10.1 and 20 cm long are found in the
terminal taxa of Drimiopsis Clade A as well as Resnova and
Ledebouria. The longest leaves (more than 20.1 cm) developed parallel
in D. barteri and 1. floribunda.
19(25). Leaves <width>
Leaves between 2.1 and 4 em wide is plesiomorphic. Narrow leaves
O.2 cm or less
(0,5-2 cm wide) are to be found in Drimiopsis Clade B & C as well as
1. 2.1 to 4 cm
Ledebouria. Wide leaves (more than 4.1 cm wide) developed in
2. more than 4 cm
parallel in 1. revoluta, D. perfoliata, D. maculata and D. atropurpurea.
The plesiomorphic state in D. botrvoides could well be due a reversal.
20(26). Leaves when tom <with or The plesiomorphic state is unknown due to ambiguity in the data. The
without threads>
majority of taxa analysed possess leaves without threads when tom.
O.with threads
The occurrence of leaves that do produce threads when tom is confined
1. without threads
the terminal clade in Ledebouria and the outgroup. The aforementioned
may represent a reversal or parallel evolution (Figure 10.17).
Leaves
or
21(27).
<pseudopetiolate
Sessile leaves are plesiomorphic. Pseudopetiolate leaves develop
not>
independently in terminal taxa of Drimiopsis Clades A-C (Figure
O.pseudopetiolate
10.18).
1. sessile
22(28). Pseudopetiole <length>
The presence of inapplicable states for this character limits its cladistic
O.exceedingly shorter than lamina usefulness.
1. approximately as long as lamina
2. exceedingly longer than lamina
23(29). Pseudopetiole <colour>
The presence of inapplicable states for this character limits its cladistic
O.banded
usefulness.
1. tinted
146
---
---
2. green
24(30). Leaf margin <shape>
O.entire
1. crenate
2. undulate
3. crenulate
25(31). Leaf margin <cartilaginous or
not>
O.cartilaginous
1. noncartilaginous
26(32). Leaf margin <markings>
O.edged purple/brown
1. bordered purple/brown
27(33). Lamina <thickness>
O.thick
1. membranous
28(34). Lamina <spotted or not>
O.spotted
1. unspotted
Leaf margins entire is plesiomorphic. Undulate margins developed in
parallel in Ledebouria, Resnova and Drimiopsis. Crenate margins
developed in all three Drimiopsis clades. Crenulate margins is an
autapomorphy of D. pusilla (Figure 10.19).
Uninformative and excluded from the analysis
The plesiomorphic state is unknown due to ambiguity in the data.
Edged margins can be variously interpreted as having developed either
by multiple reversals or in parallel in Ledebouria, Resnova and
Drimiopsis.
Thick laminas are plesiomorphic. Membranous laminae evolved in
parallel in D. maculata, D.fischeri, D. woodii and D. rosea.
The plesiomorphic state is unknown due to ambiguity in the data.
Unspotted leaves can be variously interpreted as having developed
either by multiple reversals or in parallel in Ledebouria, Resnova and
Drimiopsis (Figure 10.6).
Uninformative and excluded from the analysis.
29(35). Lamina <where spotted>
O.abaxially
1. adaxially
30(36). Lamina <tinted or not>
O.tinted
1. green
Green laminae are plesiomorphic. In Ledebouria, the distribution of
states can be variously interpreted due to ambiguity in the data. Green
laminae in L. concolor can be interpreted as a reversal (with tinted
laminae having a single origin here) or tinted laminae could have
developed in parallel (in L. socialis and the rest of the Ledebouria.
Tinted leaves in Resnova and Drimiopsis is ascribed to parallelism and
the green leaves of D. barteri is due to a reversal (Figure 10.20).
The presence of inapplicable states for this character limits its cladistic
usefulness.
31(37). Lamina abaxially <tinted>
O.purple
1. streaked purple/brown
32(38). Leaf apex <type>
O.acuminate
1. acute
2. obtuse
33(39). Leaf base <type>
O.attenuate
1. cordate
2. cuneate
34(40). Leafbase <tinted or not>
O.dark purple
1. green
35(49). Epidermal wax cover <type>
O.thin
1. thick
2. particulate
36(51). Stomata distribution
<frequency>
O.sparse
1. dense
37(52). Stomatal crypts <crypts>
O.raised
The plesiomorphic state is unknown due to ambiguity in the data. All
three states can be variously interpreted as having developed either by
multiple reversals or in parallel in Ledebouria, Resnova and
Drimiopsis.
Attenuate leaf bases are plesiomorphic. Cuneate leaf bases are absent
from Ledebouria. The presence of attenuate leaf bases in R.
lachenalioides and Drimiopsis Clades B-C are ascribed to reversals.
Cordate leaf bases in D. maculata and D. atropurpurea represent
parallel evolution (Figure 10.21).
Green leaf bases are plesiomorphic. Tinted leaf bases developed in
parallel in L. sandersonii, R. megaphylla and Drimiopsis Clade C. The
green leaf base in Drimiopsis barteri is due to a reversal (Figure
10.22).
Filmy wax cover is plesiomorphic. The presence of the aforementioned
state is ascribed to a reversal in D. maculata. The remaining states
developed in parallel in Ledebouria and Drimiopsis. Ambiguity in the
data precludes a decision on whether parallelism and/or reversals
occurred in Resnova.
Densely distributed stomata is plesiomorphic. Their presence in D.
maculata, D. reilleyana, D. pusilla, D. liniopapilla and D. queae is
ascribed to reversals. Sparsely distributed stomata developed in parallel
in L. socialis, L. revoluta and Drimiopsis.
The plesiomorphic state is unknown due to ambiguity in the data.
Raised stomatal crypts in L. socialis can be variously interpreted as
147
--
-
-
1. shallow
38(53). Stomatal subsidiary cells
<subsidiary cell H-complex present or
not>
O.form a H-complex
1. not in a H-comolex
39(54). Epidermal cells adaxial shape
O.short polygonal
1. elongate tetragonal
40(55). Epidermal cells abaxial shape
O.elongate tetragonal
1. abaxial cells short polvgonal
41(56). Epidermal cells anticlinal
boundaries
O.undelimited
1. channelled
42(57). Epidermal cells anticlinal
boundaries
O.straight
1. irregular-sinuate
43(58). Epidermal cells periclinal wall
curvature
O.straight tabular cells
1. tabular-convex cells
2. non-tabular convex cells
44(59). Epidermal cells cuticle striae
O.smooth
1. regular
2. irregular
having developed either via a reversal or in parallel to the outgrouP.
Stomatal subsidiary cells that do not form a H-complex represents the
plesiomorphic state. Although largely confined to Drimiopsis, stomata
with subsidiary cells forming a H-complex developed in parallel in L.
ovatifolia and L. revo/uta.
Elongate tetragonal epidermal adaxial cells represent the plesiomorphic
state. Although largely confined to Drimiopsis, short polygonal adaxial
cells develop in parallel in L. inquinata.
The plesiomorphic state is unknown due to ambiguity in the data.
Abaxial cells short polygonal in L. socia/is can be variously interpreted
as having developed either via a reversal or in parallel to the outgroup.
Channelled anticlinal wall boundaries represent the plesiomorphic
state. Although largely confined to Resnova, undelimited anticlinal
wall boundaries develop in parallel in L. asperifolia.
Straight anticlinal cell boundaries represent the plesiomorphic state.
Irregular anticlinal cell boundaries are a synapomorphy of Drimiopsis.
The straight anticlinal cell boundaries in D. botryoides represent a
reversal.
The plesiomorphic state is unknown due to ambiguity in the data.
However, tabular-convex cells are a synapomorphy for Drimiopsis.
The presence of straight tabular cells is ascribed to reversals in D.
botryoides, D. barteri and D. rosea.
The plesiomorphic state is unknown due to ambiguity in the data.
Smooth cuticle striae arise as a result of independent reversal in
Drimiopsis Clades B & C. The presence of aforementioned in D.
macu/ata can be variously interpreted as having developed either via a
reversal or in parallel due to ambiguity in the data
or
Indumentum
Indumenta presence is plesiomorphic. Loss of indumentum occurred
<presence
45(41).
absence>
more than once in Ledebouria and all three Drimiopsis clades
O.absent
respectively. The absence of indumentum in D. liniopapilla could
possibly be due to a reversal, but ambiguity in the data prevents
1. present
clarification. All the Resnova taxa analysed retain the plesiomorphic
state (Figure 10.23).
46(42). Indumentum arranged
The presence of inapplicable states for this character limits its cladistic
usefulness.
O.in rows
1. randomly
2. sparselv on margins
47(43). Indumentum in the form of The presence of inapplicable states for this character limits its cladistic
usefulness.
<type>
O.papillae
1. hairs
48(44). Indumentum <abundance>
The presence of inapplicable states for this character limits its cladistic
usefulness.
O.sparse
1. frequent
2. dense
49(45). Indumentum on lamina <absent Uninformative and excluded from the analysis.
or present>
O.absent
1. present
50(46). Indumentum on pseudopetiole Uninformative and excluded from the analysis
<absent or present>
O.present
1. absent
51(47). Indumentum on abaxial leaf The presence of inapplicable states for this character limits its cladistic
usefulness.
surface <present or absent>
148
---
-
--
--
O.present
1. absent
52(48). Indumentum on adaxial leaf The presence of indumentum is plesiomorphic. Inapplicable states
limits the cladistic usefulness of this character.
surface <present or absent>
O.present.
1. absent
One to two inflorescences per bulb is plesiomorphic. More than two
53(60). Inflorescence <arrangement>
inflorescences per bulb evolved twice in Ledebouria: once in L.
O.one to two per bulb
concolor and once in the terminal clade (Figure 10.7).
1. several per bulb
54(61). Inflorescence <type>
O.a simple raceme
1. a simple pseudo-corvmb raceme
55(62). Inflorescence <posture>
O.erect
1. spreading
56(63). Inflorescence <length>
O.shorter than leaves
1. more or less as long as leaves
2. considerably longer than leaves
Uninformative and excluded from the analysis, although a simplepseudo-corymb raceme is an autapomorphy of D. comptonii.
An erect inflorescence is plesiomorphic. Spreading inflorescence
developed in parallel in Ledebouria, R. megaphylla and D. botryoides
and D. maculata.
Inflorescence considerably longer than leaves is plesiomorphic. The
presence of the aforementioned state in D. rosea and D. queae is
ascribed to a reversal. Due to ambiguity in the data, it is unclear
whether this state in D. comptonii, D. woodii and D.fischeri is also due
to a reversal. Inflorescence more or less as long as leaves develop in
parallel in the terminal clade of Ledebouria, R. lachenalioides and
Drimiopsis Clades B & C. Inflorescences shorter than leaves develops
in parallel in L. inquinata and D. carrii (Figure 10.24).
57(64). Inflorescence with <number of The plesiomorphic state is unknown due to ambiguity in the data.
flowers>
Inflorescences with 15 flowers or less have developed in parallel in D.
O. 15 flowers or less
davidsoniae, D. comptonii and D. atropurpurea.
1. 16 to 30 flowers
2. more than 30 flowers
58(71). Flowers distributed <flower Inflorescences with dense flowers are plesiomorphic. Sparsely
distributed flowers developed in parallel in Ledebouria, Resnova and a
density>
number
of Drimiopsis taxa.
O.sparsely
1. densely
Pedicels more than 0.4 cm long is plesiomorphic and occurs mainly in
59(73). Flowers <pedicel length>
O. minutely pedicellate (shorter Ledebouria. Elongated pedicels in D. comptonii, D. fischeri and D.
woodii is ascribed to a reversal. Shortly pedicellate flowers occur in
than 0.1 cm)
1. shortly pedicellate (0.1 to 0.4 cm Resnova. Their occurrence in some of the terminal taxa in Drimiopsis
Clade C is also ascribed to a reversal. Apart from D. liniopapilla,
long)
2. pedicels elongated (more than minutely pedicellate flowers occur in the basal taxa of Drimiopsis
0.4 cm long)
Clades A-C (Figure 10.25).
60(65). Rachis <general length>
A rachis 11-20 cm long represents the plesiomorphic state. A rachis
O. 10 cm or shorter
more than 20 cm long is an autapomorphy of D. maculata. Due to
1. 11 to 20 cm long
ambiguity in the data, it is unclear whether states 0 or 1 are
2. more than 20 cm long
homoplasious in Resnova. The majority of Drimiopsis taxa possess a
rachis 10cm or shorter. A rachis 11-20 cm long in D. perfoliata, D.
botrvoides, D. woodii and D. rosea is ascribed to parallel reversals.
The plesiomorphic state is unknown due to ambiguity in the data. In
61(66). Rachis <shape>
Drimiopsis, cylindrical and ovoid-cylindrical rachi are homoplasious.
O.cylindrical
1. conical
2. ovoid-cylindrical
Green peduncles are plesiomorphic. The remaining states are all
62(68). Peduncle <variegation>
O.banded
homoplasious, having developed independently across the whole tree.
All taxa in Drimiopsis Clade A retain the plesiomorphic state. Banded
1. coloured purplish
peduncles are confined to D. woodii and D. atropurpurea. Coloured
2. spotted
3. I!reen
peduncles are confined to Drimiopsis Clade C.
63(69). Bracts in mature inflorescence Well-developed bracts in mature inflorescence are plesiomorphic and
<presence or absence>
by and large confined to the outgroup and Ledebouria. Apart from L.
O.absent
concolor, vestigial bracts occur in Resnova and Drimiopsis. The only
149
1. vestigial
2. well develooed
64(70). Prophylls <present or not>
O.absent
1. present
65(72). Flower <size>
O.minute (1-2 mm long)
1. small (2.1-4 mm long)
2. medium-sized (4.1-6 mm long)
3. large (more than 6 mm long)
66(76). Flowers <shape>
O.coronate to stellate
1. campanulate
2. tubular
67(77). Flowers with hypanthium base
< shape>
O.truncate
1. obtuse
2. rounded
68(78). Tepals <type>
O.isomorphic
1. dimorohic
69(79). Tepals with hypanthium
<hypanthium size>
O.inconspicuous
1. conspicuous
70(80). Tepals <colour>
O.whitish to greenish
1. purplish green
2. creamy-brownish
3. pink
4. purple/blue
71(81). Outer whorl of tepals
<posture>
O.cormivent
I. recurved
2. drooping
72(82). Outer whorl of tepals
longitudinal posture <outer>
O.cucullate
1. flat
73(83). Outer tepals apically <apex
margin shape>
O.conduplicate
1. flat
74(84). Irmer whorl of tepals <irmer
posture>
O.cormivent
1. recurved
2. drooping
75(85). Irmer whorl of tepals
longitudinally <posture>
O.cucullate
1. flat
76(86). Irmer whorl of tepals apically
<apex margin shape>
O.conduplicate
exception in the latter is D. fischeri where bracts are absent (Figure
lOA).
The plesiomorphic state is unknown due to ambiguity in the data.
Presence of prophylls is by and large confmed to the terminal clade in
Ledebouria.
The plesiomorphic state is small flowers. The remaining four states are
all homoplasious. having developed independently in all three genera.
The presence of small flowers in D. woodii, D. barteri and D.
atropurpurea is ascribed to reversals (Figure 10.26).
The plesiomorphic state is unknown due to ambiguity in the data. The
majority of taxa possess campanulate flowers. The tubular flowers of
D. fischeri developed in parallel with R. megaphylla. Coronate to
stellate flowers are an autapomorphy of 1. floribunda.
Uninformative and excluded from the analysis.
Isomorphic tepals are plesiomorphic. Dimorphic flowers develop in
parallel in R. humifusa and all the taxa in Drimiopsis.
The plesiomorphic state is unknown due to ambiguity in the data.
However, conspicuous hypanthiums are homoplasious.
The plesiomorphic state is unknown due to ambiguity in the data. All
four states occur as homoplasies within Drimiopsis.
The plesiomorphic state is unknown due to ambiguity in the data.
However, cormivent outer tepals is a synapomorphy for Drimiopsis.
The presence of recurved outer tepals in the terminal taxa of Clades AC is due to reversals.
Longitudinally flat outer tepals are plesiomorphic. Longitudinally
cucullate outer tepals are a synapomorphy for Drimiopsis (Figure
10.10).
Flat outer tepal apex margins are plesiomorphic. Conduplicate outer
tepal apex margins are a synapomorphy for Drimiopsis.
The plesiomorphic state is unknown due to ambiguity in the data.
However, connivent irmertepals are a synapomorphy for Drimiopsis.
Longitudinally flat irmer tepals are plesiomorphic. Longitudinally
cucullate irmer tepal posture is a synapomorphy for Drimiopsis (Figure
10.11).
Flat irmer tepal apex margins are plesiomorphic. Conduplicate irmer
tepal apex margins are a synapomorphy for Drimiopsis.
150
----
1. flat
77(87). Vitta
O.conspicuous
1. faint
2. absent
78(89). Androecium <colour>
O.greenish to whitish
1. cream
2. maroonishlourolish
79(90). Androecium <posture>
O.erect
1. soreading
80(92). Androecium <arrangement on
tepals>
O.uniseriate
1. biseriate
81(93). Androecium <insertion on
tepals>
O. inserted at throat of perianth
tube
82(94). Androecium <length>
O.shorter than pistil
1. as long as pistil
2. longer than oistil
83(95). Filaments <cohesion>
O.free
1. valvate
84(96). Filaments <shape>
O.deltoid to acuminate
1. lanceolate
2. filifonn
85(98). Gynoecium <number of
carpels>
O. tricarpellate
1. oolvcaroellate
86(99). Ovules <number per locule>
O.two per locule
1. more than two oer locule
87(100). Stigma <shape>
O.roundish
Uninfonnative and excluded from the analysis.
Cream coloured androecium is plesiomorphic. Maroonishlpurplish
androecium is confined to Ledebouria. Although all the Drimiopsis
taxa possess greenish to whitish androecium, this particular state also
developed in parallel in R. megaphylla and L. conc%r.
Uninfonnative and excluded from the analysis.
A uniseriate androecium is plesiomorphic The biseriate androecium is
a synapomorphy for Resnova.
Uninfonnative and excluded from the analysis.
Androecium longer than the pistil is plesiomorphic. Resnova only
possesses androecia longer than the pistil. Although the majority of
Drimiopsis taxa possess androecia as long as the pistil, this state also
occurs independently in some Ledebouria.
Free filaments are plesiomorphic. Valvate filaments represent a
homoplasious character state due to its parallel development in
Drimiopsis Clades A, B and C.
Filifonn filaments are plesiomorphic and occur in the outgroup and
Ledebouria. Lanceolate filaments are a synapomorphy for Resnova.
Deltoid to acuminate filaments are a synapomorphy for Drimiopsis
(Figure 10.27).
Uninfonnative and excluded from the analysis.
Uninfonnative and excluded from the analysis.
Triangular stigmas are plesiomorphic. Roundish stigmas are a
synapomorphy
for Drimiopsis
.
1.triangular
88(101). Stigma papilla <type>
O.stalked
1. subsessile
2. sessile
89(102).
Stigma papilla
<stigmatic papilla shape>
O.spheroid
1. trilobal
90(103). Style <size>
O.shorter than ovary
1. as long as ovary
2. longer than oVary
91(104). Style <style shape>
O.terete
1. triangular
92(105). Ovary <attachment>
O.sessile
Stalked papillae are plesiomorphic. Subsessile papillae are a
synapomorphy for Resnova. Sessile papillae are a synapomorphy for
Drimiopsis.
shape
Round papillae are plesiomorphic.
synapomorphy for Drimiopsis.
Trilobal
papillae
are
a
Styles as long as the ovary in Drimiopsis is a plesiomorphic character,
also present in the outgroup. Styles shorter than the ovary develop in
parallel in Resnova and D. liniopapilla. Styles longer than the ovary
are a synapomorphy for Ledebouria.
Triangular styles are plesiomorphic. Terete styles are a synapomorphy
for Drimiopsis and Resnova.
Stipitate ovaries are plesiomorphic. Sessile ovaries are a
synapomorphy for Drimiopsis except in D. fischeri that possesses a
151
---
1. stiuitate
93(106). Ovary <shape>
O.globose
1. ovoid to oblong
2. conical
94(107). Ovary transversely <shape>
O.smooth
1. with ridge below style
2. several lobed
95(109). Ovary shoulders <shoulders
present or absent>
O.absent
1. shoulders
96(110). Nectaries <absent or present>
O.absent
1. uresent
97(111). Pollen <type>
O.isomorphous/monosporous
1. heteromorohous/heterosporous
98(112). Pollen equatorial view
<shape>
O.depressed ovate
1. ellipsoid
99(113). Pollen polar view <shape>
O.elliptic
1. narrowly elliptic
100(114). Pollen laterally <shape
lateral view>
O.blunted
1. tapered
101(115). Pollen <equatorial diameter>
O.subequiaxe
1. breviaxe
102(116). Pollen distal pole <type>
O.distal pole straight
1. distal Dolecurved
103(117). Pollen sexine <type>
O. smooth
1. rough
104(118). Pollen ornamentation <type>
O.punctate
1. reticulate
2. Dunctate-reticulate
stipitate OVarydue to a reversal.
Globose ovaries are plesiomorphic. Conical ovaries are a
synapomorphy for Ledebouria. Although all Resnova possess an ovoid
to oblong ovary, this state is also present in D.fischeri due to a reversal
(Figure 10.5).
Smooth ovaries are plesiomorphic and occur in Drimiopsis. Ovaries
with a ridge below the style are a synapomorphy for Resnova. Several
lobed ovary is a synapomorphy for Ledebouria.
The plesiomorphic state is unknown due to ambiguity in the data.
Gynoecia without shoulders could either have developed in parallel
with the outgroup or represent the plesiomorphic state.
Uninformative and excluded from the analysis.
Heteromorphous/heterosporous pollen is plesiomorphic.
Isomorphous/monosporous pollen is a synapomorphy for Drimiopsis
and Resnova.
Ellipsoid pollen is plesiomorphic. Depressed ovate pollen is a
synapomorphy for Drimiopsis and Resnova.
The plesiomorphic state is unknown due to ambiguity in the data.
Elliptic pollen could either have developed in parallel with the
outgroup or represent the plesiomorphic state.
Pollen with tapering ends are plesiomorphic. Pollen with blunt ends are
a synapomorphy for Drimiopsis.
Breviaxe pollen are plesiomorphic. Subequiaxe pollen are a
synapomorphy for Drimiopsis.
Pollen with a curved distal pole is plesiomorphic. Pollen with a straight
distal pole represents a synapomorphy for Drimiopsis.
Rough sexine is plesiomorphic. Smooth sexine is a synapomorphy for
Drimiopsis and Resnova.
Punctate ornamentation is plesiomorphic. Reticulate ornamentation is a
synapomorphy for Resnova. Punctate-reticulate ornamentation is a
synapomorphy for Ledebouria
152
-
---
Table 10.4: Matrix of 163 characters used for a total evidence analysis. Taxa included were
those for which both morphological data (characters 0-104) and transition! transversion data
(characters 105-134) and indel data (characters 135-163) were available.
Characters
0-66
o
5
10
15
20
25
30
35
40
45
50
55
60
65
S nervosus
2100102121210201001101--00001-1-001010111102211110-0110022121032110
R maxima
2101100111101201202111--21001-1-021011110000010011-1000021111131031
R humifusa
1101110110101102201111--01100100221211110000010011-1000021011111011
D barteri
1101100110211010203011--01001-1-12100100011000
10012100221011
D maculata
2101110010001201012210202101011-01101100011100
00121012231021
D perfoliata 1101100000001102011211--01101-1-12110100011120
00021001031011
D botryoides 2101110000101201102111--0110011-12110100010020
L socialis
2111101111211201201011--0110010010110011110000
00121001031011
00121021032011
2110102111211201202111--01101-1-20111111010000
20122021031011
L floribunda 2100102121211201203101--2100000110101111010020
20112021012110
2101101121210201202201--2100010010100101010000
20112021022111
L concolor
L revoluta
Characters
67-133
67
72
77
82
87
92
97
102 107 112 117 122 127 132
I
s
nervosus
I
I
R maxima
1003111111010112010011011110110111110121202211103223311311331110311
R humifusa
1111211111010112010011011110110111110121202211103223311311331110311
D barteri
0110000000000011000002100000010000000020220013130001133133-03310333
D maculata
01001000000000111000021000000100000000-0321013130001133133-033-0333
D perfoliata
0110100000000011100002100000010000000000220013130001133133-03310333
D botryoides
01101000001000111000021000000100000000*0320013130001133133-03310233
L socialis
2003111111010012020010021120111111111220220031103223311311031130231
L concolor
2000111111000012020010021121111111111220220031103223311311031130231
L floribunda
2003111111020011020010021121111111111200220011303223311311-31112331
L revoluta
2013111111020011020010021121111111111220220011130001133133-03310331
Characters
I
I
20102112112110120211100011020111011110-0220013303223311311331112331
134-163
134
139
144
149
154
159
S nervosus
10-110000100000000010010011011
R maxima
10001111011000001001101001-101
R humifusa
10001111011000001001101001-101
D barteri
10-11111011111---11-1000011001
D maculata
10-11111011111---11-1011111001
D perfoliata 10-11111001111---1001010001001
D botryoides 10-11111101111---11-1010011001
L socialis
10-1111101100001-0011010011001
L concolor
10-1111101100001-0011010011000
L floribunda 10-101110110001--0011010011001
L revoluta
-1101111011111---11-1110011001
153
---
---
--
The four most parsimonious trees differ solely in the topology of the Ledebouria clade
(Figure 10.2). Ledebouria socialis (Bak.) Jessop, L. conc%r
(Bak.) Jessop and L.
sandersonii (Bak.) S. Venter are basal in all four trees. In addition, L. floribunda (Bak.)
Jessop and L. revo/uta (L.f.) Jessop group in all cases. The conflict in the data is largely
confmed to L. ovatifolia (Bak.) Jessop, L. inquinata (C.A. Sm.) Jessop and L.
asperifolia (v.d. Merwe) S. Venter.
L. conc%r
L. socia/is
L. sandersonii
L. sanderson;;
L. ovatifo/ia
L. asperifo/ia
L. ovatifolia
L. asperifo/ia
L. inquinata
L. inquinata
L. floribunda
L. f/oribunda
L. revo/uta
L. revo/uta
L. sanderson;;
L. sandersonii
L. f/oribunda
L. f/oribunda
L. ovatifo/ia
L. asperifolia
L. asperifolia
L. inquinata
L. inquinata
L. ovatifo/ia
Figure 10.2: Four competing hypotheses of relationships in Ledebouria from a cladistic analysis
of the data matrix as presented in Table 10.2. Conflict in the data is confmed to L. asperifolia, L.
inquinata and L. ovatifolia.
The consensus cladogram (Figure 10.1) hypothesizes two main monophyletic groups
with strong bootstrap support, i.e. Ledebouria Roth on the one hand and Resnova v.d.
Merwe with Drimiopsis on the other. The Ledebouria clade is supported by a
combination of four synapomorphic characters, namely: a style longer than the ovary
90[2]; conical shaped 93[2] and several lobed 94[1] ovary and pollen grains with
154
-
---
---
punctate-reticulate ornamentation 104[2], and two homoplasies: leaves mostly with
indumentum absent 45[0] and a spreading inflorescence 55[1].
Seven synapomorphies support the Resnova-Drimiopsis clade, namely: loose bulb
scales 9[0] with outer ones fleshy 11[0]; a deltoid to acuminate filament 84[1],
subsessile stigmatic papillae 88[1]; a terete style 91[0]; isomorphous/monosporous
pollen grains 97[0] with a smooth sexine 103[0]. In addition, this clade is supported by
six homoplasious characters, namely: a gregarious bulb habit 3[1]; whitish bulbs 6[0]
with white outer scales 10[1]; a predominantly cuneate lamina base 33[2]; shortly
pedicellate flowers 59[1] and inflorescence with vestigial bracts present 63[1].
The Resnova clade is supported by three synapomorphies: an androecium with biseriate
arrangement of filaments on the tepals 80[1]; an ovary with ridges below the style 94[1]
and pollen grain ornamentation that is reticulate 104[1]. The Resnova taxa analysed
resolve into two subclades, namely R. maxima v.d. Merwe and R. lachena/ioides (Bak.)
v. d. Merwe as well as R. humifusa (Bak.) U. & D. Miiller-Doblies and R. megaphylla.
The strongest support exists for the Drimiopsis clade with bootstrap value of 100
(Figure 10.1). The aforementioned is largely due to 17 synapomorphic characters
defining this clade, namely: epidermal cells with irregularly-sinuate anticlinal cell
boundaries 42[1], periclinal wall curvature formed by tabular convex cells 43[1];
connivent 71[0], cucullate 72[0] outer tepals with a conduplicate apex margin 73[0] and
inner tepals that are connivent 74[0], cucullate 75[0] with a conduplicate apex margin
76[0]; deltoid to acuminate filaments 84[0]; a round stigma 87[0] with subsessile 88[2]
trilobal stigmatic papilla 89[1], sessile ovary 92[0]; the depressed ovate pollen grain
(equatorial view) 98[0] with blunt ends at lateral view 100[0], a subequiaxe equatorial
diameter 101[0] and a straight distal pole 102[0]. Sparsely distributed stomata 36[0]
forming an "H-complex" with subsidiary cells 38[0], adaxial epidermal cells that are
shortly polygonal 39[0], a minute pedicel 59[0], dimorphic tepals 68[1] and a greenish
androecium 78[0] that is as long as the pistil 82[1] are seven homoplasies additionally
supporting the Drimiopsis clade.
The Drimiopsis clade separates into three smaller clades supported mainly by
homoplasious character suites (Figure 10.1 A-C). The basal clade A consists of D.
155
-
stolonissima (D. & D. Miiller-Doblies) Lebatha, D. carrii Lebatha, D. perfoliata Bak.,
D. botryoides Bak. and D. maculata Lindt & Paxt. These taxa possess roundish bulbs
7[1] with leaves having indumentum present 45[1]. The former character, however, also
occurs in L. asperifolia, R. megaphylla, R. lachenalioides as well as D. woodii and D.
kikiae while the latter is also to be found in the majority of the Ledebouria analysed as
well as in representatives of the other two Drimiopsis clades, but is absent in Resnova.
Drimiopsis botryoides and D. maculata, terminal entities of this clade, possess
autapomorphies of falciform leaves 16[1] and the largest flowers in the genus 65[3]
respectively.
Clades B and C are supported by the synapomorphy of erect leaves 15[0] plus three
homoplasies, namely unspotted leaves 28[1], cuticle irregularly striated 44[2] and
minute flowers 65[0].
No synapomorphies support or are present in clade B. Two autapomorphies, a
spathulate leaf form 16[3] and oblanceolate shaped leaves 17[2] diagnose D. comptonii
and a single autapomorphy, bracts in mature inflorescence absent 63[0], diagnoses D.
fischeri.
Similarly, no synapomorphies support clade C. Two autapomorphies, namely a
crenulated leaf margin 24[3] and a pseudopetiole exceedingly shorter than lamina 22[0],
diagnose D. pusilla and D. kikiae respectively.
10.4.2 Total evidence analysis
A cladistic analysis confmed to the morphological data in Table 10.4 resulted in two
most parsimonious trees. The strict consensus tree recognises a poorly resolved
Ledebouria and fully resolved Resnova and Drimiopsis as separate (Figure10.3A). A
cladistic analysis of the DNA data only (transitions/transversions and indels) also
resulted in two trees of equal length. The resultant strict consensus tree rendered
Ledebouria polyphyletic (Figure 10.3B). The analysis of the aforementioned
morphological and DNA data combined resulted in one tree similar to Figure 10.3A, but
being fully resolved (Figure 10.3C).
156
--
-- - -
S. nervosus
.-I
I
L. socialis
, - I
L. floribunda
L. revoluta
R. maxima
R. humifusa
- I
D. barteri
D. maculata
D. perfoliata
D. botryoides
S. nervosus
1
L. floribunda
L. socialis
L. concolor
R. maxima
R. humifusa
L. revoluta
D. batteri
D. maculata
D. perfoliata
D. botryoides
S. nervosus
G
7
w
*- I
I
-
h
-
I
G
I
I, - I
I
i
L. floribunda
L. revoluta
L. socialis
L. concolor
R. maxima
R. humifusa
D. batteri
D. maculata
D. perfoliata
D. botryoides
Figure 10.3: Cladograms based on data in Table 10.4. A, strict consensus tree of two
most parsimonious trees based on morphological data alone; B, strict consensus tree of
two most parsimonious trees based on DNA data (transversiodtransition & indels)
alone; C, one and only fully resolved tree based on a total evidence approach.
S netVosus
CHAR 93: Ovary<shape>
L=160 CI=66 RI=90
CHAR 63: Braots in mature intloresoenoe <pl'8senoe or absenoe>
L 160 CI 66 RI 86
L IJorlbunda
..
..
=
=
=
0: absent
1: vestigial
2: developed
Ambiguity
L IIorlbunda
..
..
0: globose
1: ovoid to oblong
2: conical
Ambiguity
L rewlUUI
L fevolufa
D IJnlopapll1a
Oquoae
Figure 10. 4: Well-developed bracts in mature inflorescence are
plesiomorphic and by and large confined to the outgroup and Ledebouria.
Apart from L. conc%r, vestigial bracts occur in Resnova and Drimiopsis.
The only exception in the latter is D.fischeri where bracts are absent.
D
I;nlopapilla
Oqu_
Figure 10.5: Globose ovaries are plesiomorphic. Conical ovaries are a
synapomorphy for Ledebouria. Although all Resnova possess an ovoid to
oblong ovary, this state is also present in D.fischeri due to a reversal.
158
S netVosus
S netVosus
CHAR 28: Lamina <spotted
L 450 CI
11 RI 42
..
.
=
=
=
or not>
CHAR 63: Intloresence
per bulb <arrangement>
L 300 CI
33 RI
71
0: spotted
1: unspotted
Ambiguity
L norlbunda
=
=
0: solitary
1: one to two
2: several
Ambiguity
~ norlbunda
L reva/uta
.. =
..
~ ,evo/uta
D Iiniopapllla
D /lnlopapl/la
D queae
D queae
Figure 10.6: The plesiomorphic state is unknown due to ambiguity in the data.
Unspotted leaves can be variously interpreted as having developed either by multiple
reversals or in parallel in Ledebouria, Resnova and Drimiopsis.
Figure 10.7:One to two inflorescences per bulb is plesiomorphic. More than
two inflorescences per bulb evolved twice in Ledebouria: once in L. concolor
and once in the terminal clade.
159
S nervo8u8
S neIVOaU8
CHAR 9: Bulb scales <packing>
..
.
L
= 100
CI
=50
RI
=88
0: loosely packed
1: compact
Ambiguity
L flonbUnda
L_
CHAR 12: Bulb scales when torn <with or without threads>
L.200 CI. 25 RI = 0
0: produce threads
1: do not produce threads
Ambiguity
..
.
L ,..vo/ula
L revo/uta
D IInlopapllla
D Iiniopapllla
Dqueae
D qUeM
Figure 10.8: Compact bulb scales in S. nervosus and Ledebouria
evolved to loosely packed scales in Resnova and Drimiopsis with a
reversal to compact in R. maxima.
Figure 10.9: There is ambiguity in the data as to whether the presence or absence of
threads is the plesiomorphic state. Similarly, ambiguity exists as to whether subsequent
transformations represent reversals or parallel developments. The production of states is
confined to S. nervosus and L. inquinata, L. ovatifolia and L. revoluta.
160
s. n.fVO.U.
L..,,,,'Ifo/M
L.Inqulnal.
CHAR 75: Tepels longitudinal posture <inner>
L.. 50 CI - 100 RI- 100
CHAR 72: Tepels longitudlnel posture <outer>
L.. 150 CI.. 100 RI.. 100
0: cucull8te
1: flat
Ambiguity
..
.
L. ..".'11011.
L. Inqu.....
L. tlol/bun.
..
.
0: cucullate
1:flet
Ambiguity
L tIorlbunda
D.IIn/op.lII.
D. JlnIolMpllJ.
D.qu...
Figure 10.10: Longitudinally flat outer tepals are plesiomorphic.
Longitudinally cucullate outer tepals are a synapomorphy for Drimiopsis.
D. qUH'
Figure 10.11:Longitudinally flat inner tepals are plesiomorphic.
Longitudinally cucullate inner tepal posture is a synapomorphy for
Drimiopsis.
161
S norvosus
S n.,-v08U.
CHAR 2: Bulbs <below or above ground>
L 60 CI. 100 RI- 100
..
.
...
..
CHAR 6: Bulb <colour>
=
L
0: hypogeal
260 CI. 40 RI- 60
0: whitish
1: eplgeal
Ambiguity
1: purplish
2: brown
Ambiguity
L florlbunda
L flortbunda
LrevolutJI
L reva/ura
o /lniopapllla
Oqueae
Figure 10.12: Hypogeal bulbs are plesiomorphic transforming
into epigeal in L. conc%r and L. socia/is.
o /lniopapllla
D que..
Figure 10.13: Brown bulbs are plesiomorphic and are absent from Resnova and
Drimiopsis, which only possess white bulbs. Both brown and white bulbs are
homoplasious: the former as a result of a hypothesized reversal in L. floribunda and the
latter as a result of parallel evolution in Clade B & C and L. asperifo/ia. Similarly,
purplish bulbs are also homoplasious due to parallel evolution within Ledebouria.
162
S nelVo.u.
S nervo.u.
CHAR 16: Leaves <form>
L-360 CI..42 RI..66
CHAR 7: Bulb <shape>
L - 300 CI- 16 RI- 44
..
.
0: roundish
1: ovoid
Ambiguity
L norl",,"da
L "orlbunda
..
..
.
0: cordiform
1: falciform
2: linear
3: spatulate
Ambiguity
L IVvoIUUI
L nwoluta
o IIniopapllla
o Ilnlop"pllla
o qu...
Figure 10.14:Ovoid shaped bulbs are plesiomorphic. Roundish bulbs are
homoplasious, having developed in representatives of Ledebouria and
Resnova as well as D. kikiae, D. woodii, and the basal clade A.
o qu...
Figure 10.15:Ambiguity (whether cordiform or linear leaves are
plesiomorphic) at the base of the tree results in the homoplasious distribution
of cordiform leaves being ascribed to either parallelisms or reversals. Falciform
leaves is an autapomorphy of D. botryoides and spathulate leaves is an
autapomorphy of D. comptonii.
163
S n8tVOSU8
So.........
CHAR 17: Leaves<shape>
L. 360 CI = 28 RI = 16
I 0: lanceolate
I 1: ovate
I 2: oblanceolate
I Ambiguity
CHAR 20: Leave. wh.n tom <produc.
L .. 100 CI" 50 RI.. 80
..
.
L norlbund.
L_do
L rwolut.
L.......
D IinJop.plll.
thr.ads
or not>
0: produc. thr.ad.
1: do not producethr.ads
Ambiguity
D JlnlofMpHla
Dq_
D qu...
Figure 10.16: Lanceolate leavesare plesiomorphic. Ovate leaves
developed in parallel in L. ovatifolia, R. megaphylla, D. woodii, D.
atropurpurea and in D. cardi, D. peifoliata and D. maculata with a
postulated reversal to lanceolate in D. botryoides. Obianceolate leavesis
an autapomorphy of D. comptonii.
Figure 10.17: The plesiomorphic state is unknown due to ambiguity in the data. The
majority of taxa analysedpossessleaveswithout threads when tom. The occurrence of
leavesthat do produce threads when tom is confined the terminal clade in Ledebouria
and the outgroup. The aforementioned may representa reversal or parallel evolution.
164
S fHlIVOSU8
S
rHlTVOSU8
CHAR 21: Leaves <pseudo petiolate or not>
L = 160 CI = 33 RI. 60
..
.
.
CHAR 24: Leaf margin <shape>
L 600 CI. 26 RI. 26
0: pseudopetlolate
1: s85slle
Ambiguity
L florlbund.
L fIorIbund.
L revolut.
L revolut.
..
..
.
0: entire
1: crenate
2: undulate
3: crenulate
Ambiguity
D IIniopaplll.
D IInloplIpill.
D que..
D queM
Figure 10.18: Sessile leaves are plesiomorphic. Pseudopetiolate leaves
develop independently in terminal taxa of Drimiopsis Clades A-C.
Figure 10.19:Leaf margins entire is plesiomorphic. Undulate margins develop in
parallel in Ledebouria, Resnova and Drimiopsis. Crenate margins develop in all
three Drimiopsis clades. Crenulate margins is an autapomorphy of D. pusilla.
165
S
S nervosus
CHAR
MrvOSU8
CHAR 33: lamina <base>
30: lamina <coloured or not>
L
L - 250 CI. 20 RI - 69
..
.
0: coloured
1: uncoloured
Ambiguity
L tlorlbunda
L flot1bunda
..
..
- 360CI- 28 RI- 64
0: attenuate
1: cordate
2: cuneate
Ambiguity
L IVvoluta
L IVVoJutll
D IInlopapllla
D liniopapilla
D quo.
Dqu...
Figure 10.20: Green laminae are plesiomorphic. In Ledebouria, the distribution of states can
be variously interpreted due to ambiguity in the data. Green laminae in L. concolor can be
interpreted as a reversal (with tinted laminae having a single origin here) or tinted laminae
could have developed in parallel (in L. socia/is and the rest of Ledebouria). Tinted leaves in
Resnova and Drimiopsis is ascribed to parallelism and the green leaves of D. barteri is due
to a reversal.
Figure 10.21:Attenuate leaf bases are plesiomorphic. Cuneate leaf bases are
absent from Ledebouria. The presence of attenuate leaf bases in R. lachenalioides
and Drimiopsis Clades B-C are ascribed to reversals. Cordate leaf bases in D.
maculata and D. atropurpurea represent parallel evolution.
166
S nervosus
S nervosus
CHAR
..
.
L=200
CHAR 46: Indumentum <presence
L 360 Cia 14 RI = 63
34: lamina base <coloured or not>
CI=26
..
.
RI=66
0: dark purple
1: uncoloured
Ambiguity
L Worlbunda
L norlbunda
L.revolura
L revoluta
o liniopilpilla
o queH
Figure 10. 22: Untinted leaf bases are plesiomorphic. Tinted leaf bases
developed in parallel in L. sandersonii, R. megaphylla and Drimiopsis
Clade C. The untinted leaf base in Drimiopsis barteri is due to a reversal.
=
or absence>
0: absent
1: present
Ambiguity
D IInlopapllla
o queae
Figure 10.23: Indumentum presence is plesiomorphic. Loss of
indumentum occurred more than once in Ledebouria and all three
Drimiopsis clades respectively. The absence of indumentum in D.
liniopapilla could possibly be due to a reversal, but ambiguity in the data
prevents clarification. All the Resnova taxa analysed retain the
167
S nervosus
S nelVOsus
CHAR 59: Inflorescence
L-350 CI..28 RI..75
CHAR 66: Inllorescence <length>
L .. 600 CI.. 20 RI.. 42
L trorlbunda
..
..
0: shorter than leaves
1: more or less as long as leaves
2: considerably longer than leaves
Ambiguity
l norlbunda
..
..
<pedicel length>
0: minute 0.6-0.1om
1: short 0.1.0.4 om
2: more than 0.4 om
Ambiguity
L ,..voIuu.
Lrevoluta
D IInlopapllla
D quea.
Figure to. 24: Inflorescence considerably longer than leaves is plesiomorphic.
The presence of the aforementioned state in D. rosea and D. queae is ascribed
to a reversal. Due to ambiguity in the data, it is unclear whether this state in D.
comptonii, D. woodii and D. fischeri is also due to a reversal. Inflorescence
more or less as long as leaves developed in parallel in the terminal clade of
Ledebouria, R. lachenalioides and Drimiopsis Clades B & c. Inflorescences
shorter than leaves developed in parallel in L. inquinata and D. carrii.
D Ilniopap/lla
Dqueae
Figure to.25: Pedicels more than 0.4 cm long is plesiomorphic and occurs
mainly in Ledebouria. Elongated pedicels in D. compton;;, D. fischeri and D.
woodii is ascribed to a reversal. Shortly pedicellate flowers occur developed in
Resnova. Their occurrence in some of the terminal taxa in Drimiopsis Clade C
is also ascribed to a reversal. Apart from D. liniopapilla, minutely pedicellate
flowers occur in the basal taxa of Drimiopsis Clades A-C.
168
S mtlVOSUS
CHAR 84: Filament <shape>
CHAR 66: Flower <size>
..
..
.
L
L trorlbunda
=700
CI
=21
RI
=38
0: minute
1: small
2: medium
3: regular
Ambiguity
.. =
..
L
100
CI
= 100
RI
= 100
0: deltoid to aooumlnate
1: lanoeolate
2: filiform
Ambiguity
L "orlbunda
L revolUUl
L ,evaluta
D Iiniopapilla
D Iiniopapllla
D queae
D queae
Figure 10.26:The plesiomorphic state is small flowers. The remaining four
states are all homoplasious, having developed independently in all three
genera. The presence of this state in D. woodii, D. barted and D.
atropurpurea is ascribed to reversals.
Figure 10.27: Filiform filaments are plesiomorphic and occur in the
outgroup and Ledebouria. Lanceolate filaments are a
synapomorphy for Resnova. Deltoid to acuminate filaments are a
synapomorphy for Drimiopsis.
169
10.4.3 Systematic implications of the character analyses
Manning et al. (2004) circumscribed Ledebouria to include Drimiopsis and Resnova
based on the analysis of two genes. They venture listing six morphological characters
that define Ledebouria sensu lato. Accordingly, Ledebouria is circumscribed by: "... the
lack of bracteoles and by its globose or top-shaped ovary containing two ovules per
locule. In addition most species have spotted leaves and often produce more than a
single inflorescence per plant in one growing season, and the bulb scales are often rather
loosely packed and in many species produce fine threads when tom". With regard to
these characters the following:
Manning et al. (2004) do not clarify whether they distinguish between bracts and
prophylls as has been done in this analysis. In addition, they do not specify
whether this state is present in mature andlor developing inflorescences. And,
furthermore, they do not distinguish between vestigial or well-developed bracts.
In the matrix used above, all the Ledebouria analyzed, bar L. concolor possess
well developed bracts, this being a plesiomorphic character state, also present in
the outgroup (Figure 10.4). Bracts are largely vestigial in Drimiopsis. If, on the
other hand, Manning et al. (2004) by bracts refer to prophylls, then this analysis
hypothesises their presence confined to the terminal taxa in Ledebouria.
Manning et al. (2004) distinguish between "globose" or "top-shaped" ovaries.
While it is true that a conical (top-shaped) ovary a synapomorphy for
Ledebouria sensu stricto, the analysis above indicates that a third state within the
Ledebouriinae is also distinguishable, namely ovoid to oblong ovaries found in
all Resnova and D. fzscheri. Globose ovaries in tum are found in all Drimiopsis
taxa as well as the outgroup, i.e. a symplesiomorphic state (Figure 10.5).
The cladistic analysis hypothesises the presence of spotted and unspotted leaves
as having developed either by multiple reversals or in parallel in Ledebouria,
Resnova and Drimiopsis. That is to say, neither character state is a
synapomorphy and should be used with caution in support of systematic
arguments (Figure 10.6).
While it is true that Ledebouria, Resnova and Drimiopsis can produce more than
one inflorescence per season, it is also so in S. nervosus, i.e. a symplesiomorphic
character state in this analysis (Figure 10.7). A distinction has been made in this
thesis between plants producing 1-2 as opposed to three or more inflorescences
per season. When coded thus, all but two of the Ledebouria possessed the latter
homoplasious state.
I differ fiom Manning et al. (2004) and consider Ledebouria to possess compact
bulb scales when compared with Resnova and Drimiopsis. While the
aforementioned is open to interpretation, what is not is that Ledebouria shares its
state with the outgroup and R. maxima, i.e. it is symplesiomorphic in this
analysis (Figure 10.8).
Bulb scales producing threads when tom is present in S. nervosus and some of
the taxa in the terminal Ledebouria clade. As such its presence in Ledebouria is
ascribed to either a homoplasious reversal or parallel development (Figure 10.9).
An additional comment is warranted on Manning et al. (2004) who, seemingly biased
by their molecular work, attempt to negate observed differences in flower morphology
to a "mere" pollination syndrome. The accepted view in the literature on evolutionary
biology is that "Interactions among species are thought to promote the evolution of
diversity in several ways" (Futuyma, 1998). In other words, a large proportion of
diversity of life and life forms is not just due to adaptation to static environments but
also due to biotic interactions. This analysis reveals a number of synapomorphic flower
characters for Drimiopsis, Resnova and Ledebouria. These characters too have a
phylogenetic story to tell and in all fairness, deserve to be heard.
10.5 CONCLUSION
The strict consensus tree produced from a cladistic analysis of morphological data for
18 Drimiopsis, 4 Resnova and 8 Ledebouria species (and S. nervosus as outgroup)
supports the monophyletic status of all three genera in the ingroup.
A total evidence analysis of combined morphological and DNA data based on a reduced
taxon sample produced a fully resolved tree also hypothesising the monophyletic status
of all three genera.
The aforementioned results directly oppose Manning et a1 's (2004) decision to lump
Resnova and Drimiopsis in Ledebouria based solely on a polytomy produced by a strict
consensus tree of trnL-F & rbcL data. In addition, the characters professed by them to
support their lumping prove to be either homoplasious, symplesiomorphous or confined
to the terminal clade in Ledebouria. .
Although Drimiopsis resolves into three clades, no sections or subgenera are recognised
as these clades lack synapomorphies and instead possess homoplasious characters. In
addition, the subspecies formerly recognised are elevated to the rank of species.
Drimiopsis botryoides (=D. botryoides subsp. botryoides) forms terminal taxa with D.
maculata on the cladogram. Recognising the subspecies D. perfoliata (=D. botryoides
subsp. prostrata) rank creates three subspecies that include D. maculata. Drimiopsis
stolonissima and D. burkei cannot be subspecies because they belong in different
Drimiopsis clades: A and B respectively.
The aforementioned is a hypothesis, developed fiom a comprehensive cladistic analysis
of Drimiopsis and sister taxa, and like all hypotheses is subject to falsification when the
matrix is expanded with new data (characters or taxa) in the fbture.
11. PHYTOGEOGRAPHY
11.1 INTRODUCTION
Phytogeography endeavours to describe, analyse and explain patterns of distribution and
the origin of taxa. This involves ecological influences (ecological geography) and plant
distributions resultant from long-term historical factors (historical biogeography)
(Schuh, 2000). Phytogeography has utilised many paradigms over the years to explain
distributions. In the beginning were the dispersal theories originating from Darwin
(Finchrnan, 1977), then land bridges, plate tectonics and the continental drift theories
(Wegener, 1966) that are still relevant today. A Venezuelan botanist, Leon Croizat
(1894-1982) coined "life and earth evolve togetherv1-that geographic barriers and biota
co-evolve, establishing the foundations of a new synthesis between earth and life
sciences. This was coined panbiogeography (Croizat, 1962). He compared distribution
of some endemic taxa, connected their geographical ranges to form "tracks1'with other
taxa. This combination resulted in "generalized tracks" indicating historical connections
of taxa. Panbiogeography signified the start of vicariance biogeography, a combination
of his works and phylogenetic systematics (cladistic biogeography). Vicariances lead to
phylogeography, which uses cladistic methods in analysis and produces area cladograms
(Platnick, 1981, 1991). The basic assumptions of phylogeography are that speciation is a
result of geographic isolation-the
present geographic ranges have preserved ancient
patterns of geographic separation indicative of historical relationships (Schuh, 2000).
These concepts are still subjects of contentious debates.
Phytochoriological studies enhance classification of distribution patterns based on
geographical range of taxa (van Wyk & Smith, 2001). The mapping of taxa reveals
consistent distribution patterns consigning taxa to particular geographical ranges or
phytogeographical areas. The demarcated ranges are centers of endemism of the taxa.
Phytochoria facilitates taxa distributions and history studies in that its geographic ranges
are already characterised by habitat and history. The phytochoria of Africa consists of
one Archipelago-like centre of endemism, eight regional centres of endernism, seven
regional transitional zones and two regional mosaics (Figure 11.3). Traditional plant
distribution classification is based on growth form, separating regions according to
grassland, savannah etc.
The three major types of plant distribution are cosmopolitan, endemic and disjunct
distributions (Radford et al., 1974; Schuh, 2000; van Wyk & Smith, 2001).
Cosmopolitan worldwide distribution is not further considered in this study, as none of
Drimiopsis Lindl. & Paxt. taxa are cosmopolitan (Table 11.1; Figure 11.1 A & B;
Section 11.4). The criteria used to determine place of origin and of distribution is
subject to interpretation but based on available data. This primary method of
"perception or intuitive discernment" (van Wyk & Smith, 2001: 15) or "the use of
abstractions" (Blij, 1971: I), has long been in use.
The principal factors affecting plant distribution are climatic conditions. Edaphic
factors, physiographic and biotic factors are also barriers to plant migration and
dispersal. In addition, species ranges are influenced by genetically controlled tolerances.
Van Wyk & Smith (2001) succinctly illustrate a centre of origin1 dispersal model. The
model assumes that the history of an individual reflects the history of the taxon. This
applies well to taxa recently evolved at a specific location (neoendemic taxa), that
become successfid and extend to maximum geographical ranges. The success of this
juvenile stage in history is determined primarily by environmental conditions. High
vigour can promote lead to the creation of new taxa in the outmost geographic ranges.
In due course vigour is lost leading to declines in populations and disappearances in the
original geographic ranges, and disjunct distributions result. This relict stage may lead
to taxa having restricted ranges (palaeoendemics) and to eventually becoming extinct.
The graduation between stages is of course influenced by environmental, genetic factors
and reproductive failure.
Disjunct distributions are caused by habitat discontinuities, drastic conditions leading to
differential extinction producing relict populations and jump dispersals. The fracturing
of geographic barriers, through any of drastic events/conditions (vicariant events), leads
to widely separated sites (refuges) and the creation sister taxa (vicarious species) (van
Wyk & Smith, 2001). The resultant taxa split can generate continuous distribution of
species.
A geographic distribution pattern of the Ledebouriinae U. & D. Miiller-Doblies,
Drimiopsis, Resnova v.d. Menve and Ledebouria Roth, has not in its entirety been
assessed. The Ledebouriinae, with the exception of Ledebouria, are exclusively
distributed in sub-Saharan Africa. Resnova is endemic to southern Africa, concentrated
in Mpumalanga and KwaZulu-Natal Provinces. Resnova maxim occurs in the Eastern
Cape Province of South Africa and in Swaziland. Ledebouria possesses a wide
distribution-according
to Venter (1993) it occurs in southern Africa, south-eastern
India, the Mediterranean and the western coast of Madagascar. The most species rich
areas are in the KwaZulu-Natal, Mpumalanga and Gauteng Provinces, with the centre of
endemism in Mpumalanga.
11.2 OBJECTIVES
This study explores geographic distribution of Drimiopsis, speculates on distribution
patterns and evolutionary history, and presents an interpretation of relationships with
sister taxa of the Ledebouriinae based on patterns of distribution.
11.3 MATERIALS and METHODS
Data on taxa localities were based on herbarium material, personal collections, and
published data, especially so Venter (1993) for Ledebouria. ArcView GIs 3.2
(Applegate, 1999) was used to plot data and generate maps. Precise localities, where
uncertain, were investigated and established mainly through the geographic database,
National Geospatial-Intelligence Agency, GEOnet Names Server (GNS), 2004. Taxa
groupings are based on results obtained in the cladistic analysis (Figure 11S).
11.4 RESULTS and DISCUSSION
11.4.1 Distribution patterns
Drimiopsis distribution (Figure 11.1 A & B) spans most of Africa from Ghana into
Togo, Nigeria, Cameroon and southern Chad in the Guinea-CongolianISudanian
regional transitional zone-1
1 phytochorion (Figure 11.3). Species richness is highest in
South Africa's Mpumalanga Province (Table 11.2). Drimiopsis collections made in the
equatorial or central parts of Africa (Central Afiican Republic, Democratic Republic of
Congo and Zambia), occur in several phytochoria. The Central African Republic ones
occur in phytochorion-11
and those of Democratic Republic of Congo in the Guinea-
Congolian centre of endemism-1.
endemism-2,
Zambian taxa are in the Zambezian centre of
together with the southern Afiican taxa of Zimbabwe, northern and
portions of eastern Botswana and the Limpopo Province of South Africa. Distribution in
South Africa extends into the Archipelago-like center of endemism-8
Maputoland-Pondoland-15
and the
regions encompassing the Mozambique collection.
Drimiopsis distribution in the Horn of Africa regions occurs in southern Sudan in
phytochorion-1 1, Ethiopia and Somalia in the Sudanian region center of endemism-
4. Indications from east African Kenyan and Tanzanian collections place distribution in
the phytochorion-4. However, the Tanzania taxa also occur in the Swahilian-13a
phytochorion together with Zanzibar taxa on the one hand, and with Burundi taxa in the
Lake Victoria mosaic-1 2.
Table 11.1: Regional distribution of Drimiopsis in various African phytochoria. (Figure 11.3): 1
= Guinea-Congolian centre of endemism; 2 = Zarnbezian centre of endemism; 3 = Sudanian
centre of endemism; 4 = Somalia-Masai centre of endemism; 8 = Archipelago-like centre of
endemism; 11 = Guinea-CongolianISudaniantransition zone; 12 = Lake Victoria mosaic; 13a =
Swahilian centre of endemism; 14 = Kalahari-Highveld transitional zone and 15 = MaputolandPondoland regional mosaic.
Total
Phytochorion (Figure 1 1.3)
1 2 3 4 8 11 12 13a 14 15 phytochoria
Drimiopsis atropurpurea
e . 3
Taxon
Drim iopsis barteri
m
e
m
5
e
Dr imiopsis botryoides
0
0
3
Drim iopsis burkei
2
Drimiopsis carrii
1
Drimiopsis comptonii
1
Drimiopsis davidsoniae
1
DrimiopsisJscheri
1
Drim iops is kikiae
1
a
Drimiopsis liniopapilla
1
Drimiopsis maculata
a
3
Drimiopsis perfoliata
2
Drimiopsis pus illa
1
Drimiopsis queae
1
Drimiopsis reilleyana
1
Drimiopsis rosea
a
2
Drimiopsis stolonissima
1
Drimiopsis woodii
2
Drin,iopsis Dist,"ibution
.
s
Afiica~ cOlmtries,showing mnnber
of speCIes of Drimiopsis collected.
- - o- 1000-
2000
1000
Kilometers
Figure 11.1: Known number of types of species per country for Drimiopsis based on herbarium
collections (Table 11.2)
178
---
-- . . . . . - . . . . . . . . . . . . . - . . .
.
D. if1$,-hlitist
. D. .JfRhniuilt=
. D.1ngJi
D.71J.J&fI
. D.7f1ik.~
. D. qtt&sII
. D.lIwt'"
. D.litiJ~'"
. D. n=:/tlAist
D.~
. D.~1ItII
. D. ulfflTfRtti
. D.M71'i
. D.lmisf
. D. ilMlI~1trfIA
. D. Igh:rJgiu
· D.lm1s7i
s
2000
o
2000
4000
Kilometers
Figure 11.2: Known species richness for Drimiopsis based on herbarium collections (Table
] 1.2)
179
10"
21]'
40"
30"
20"
I]'
50"
'311'
-20"
II]'.
II]'
O'
_
_
_
_
_
_
I]'
Regional Centres of Endemism
1
2
3
HI"
Guineo-Congolia'
Zambezian
rc==JSudanian
4
5
6
7
13a
Somalia-Masal
Cape
c==JKaroo-Namib
11]'1
c:::J Mediterranean
r\
Swahilian
Archlpelago-llkeCantle of Ende..lsm
c:::J
8
21]'.
Regional
10
11
13b
14
30"-
16
17
18
Afromontane
Transition
-{.'
& Afroalpine
/
Zones
Guinea-(ongolia.'lambezian
IDIJI Guinea-Congolia!5udanian
,
/
/
f'
!
J
,20"
SwahilianIMaputaland
c==J Kalahari-Highveld
~
UlU]
Sahel
Sahara
MediterraneanlSahara
6
1-311'1
t(:
Regional Mosaks
12
LakeVictoria
15
.f
21]'
I]'
,
8
Maputaland-Pordoland
10"
500
".<-
'.0
I
10"
-7
20"
1...1f9l.:
I
'(at Equat>t
I
31]'
--
I
41]'
I
50"
Figure 11.3: Phytochoria of Africa. Drimiopsis occurs (Figure 11.1 A & B; Table 11.1 & 11.2)
in 10 of the phytochoria: Guineo-Congolian centre of endemism-I, Zambezian centre of
endemism-2, Sudanian centre of endemism-3, Somalia-Masai centre of endemism-4,
Swahilian centre of endemism-13a, Archipelago-like centre of endemism-8, GuineaCongolianlSudanian transition zone-II,
Kalahari-Highveld transitional zone-I 4, Lake
Victoria mosaic-I 2 and the Maputoland-Pondoland regional mosaic-15 (van Wyk & Smith,
200 I: 10).
180
nl[STI
DJO,-F
CHAD
SUDAN
I
..
·
A
~;H..." ,
CONGO
. Hr
..
KALA
HARt
.
Bodelt-Uk, CIwI
i
..-AdmA
",/,
IleStll.Rillil
~
:EastaraIi ft
=
Figure 11.4: Africa physical map: A the African basins of Djouf, Chad, Sudan, Congo (now
called the Zaire Basin) and the Kalahari surrounded by plateaus. B = Africa rift valleys. (De
BIij, 1971: 319)
181
D. stolonissima
D. carrii
I
D. perfoliata
D. hotryoides
D. maculata
D. davidsoniae
D. comptonii
D.fischeri
D. woodii
.1
I
.
D. reilleyana
D. hurkei
D. harteri
D. rosea
I
D.pusilla
.
.
D. atropurpurea
southernMrica
D.kikiae
D. liniopapilla
tropical Africa
D. queae
Figure 11.5: Drimiopsis cladogram distinguishing between taxa possessing a southern or
tropical African distribution.
182
Table 11.2: Regional distribution of Drimiopsis and Resllova in countries: Bt, Botswana; Br, Burundi; Ca, Cameroon; Cr, Central African Republic; Cd, Chad; Dc,
Democratic Republic of Congo; Eh, Ethiopia; Ga, Ghana; Kn, Kenya; Mz, Mozambique. Provinces of Nigeria: sk, Sokoto; zr, Zaria; yl, Yola; ng, Niger. Sm, Somalia. South
African Provinces: ec, eastern Cape; gu, Gauteng; kz, KwaZulu-Natal; Ii, Limpopo; mp, Mpumalanga; nw, North West. Sd, Somalia; Sw, Swaziland. Tanzania districts: ir,
Iringa; ky, Kyimbila; mb, Mbulu; mp, Mpanda; rn, Rungwa, mr, Morogoro; ta, Tanga; lu, Lushoto; is, the Islands. Tg, Togo; Za, Zambia; Zn, Zanzibar; Zi, Zimbabwe.
Collections are inadequate, uneven. South African appears the most-collectedregion. Total number of taxa per area is indicative of endemism, the highest (+3) in bold red.
Taxon
Bt
Br
D. atropl/rpl/rea
D.barteri
D. botryoide.f
D.bl/rkei
D.carrii
.
.
Ca
.
Cr
Cd
Dc
Eh
Ga
. . . .. .
Kn
Mz
.
.
SIO
Nigeria
sk
.
zr
vi
.
. . .
South Africa
ec
ng
gU
.
kz
Ii
Sd
.
100
nw
S
W
.
. .
.
D. david.wmille
D.jischeri
.
D.kikiae
.
D. petjiJ/ilita
D.pl/.filla
D. reilleyanll
.
.
D.rosea
D.woodii
Re.fIlova
1
1
2
2
2
1
2
1
1
2
1
1
1
1
1
2
.
.
.
3
6
rn
.
. .
lOr
ta
lu
Tg
Za
Zn
Zi
.
.
..
.
2
1
is
. ..
.
2
1
.
.
.
.
.
100
.
.
D. .ftoloni.uitll/I
Total no. of taxa
. .
.
D.ql/eae
lOb
. . . .. .
.
. . .
.
D. macl/lata
kv
. .. . . .
D. cO/nptonii
D. /iniopapilla
Tanzania
ir
1
9
.
2
1
5
3
1
2
3
2
1
1
1
183
1
The distribution of taxa in White's (1983) phytochoria (Table 11.1) points to disjunct
distributions and regional endemism. Drirniopsis barteri Bak. has the widest distribution
in the genus, occurring in five of the phytochoria. Drimiopsis barteri has been collected
fiom Ghana, Togo, Nigeria, Cameroon, Chad, Somalia, Ethiopia, Central African
Republic, Democratic Republic of Congo, Tanzania and Zambia (Table 11.2). One
herbarium specimen from Mpumalanga possessing D. barteri characteristics needs more
floral investigations to validate its identity. The distribution pattern of D. barteri
suggests a much wider distribution. Disjunction probably by habitat discontinuity
caused it to break into sectional distribution.
Drimiopsis atropurpurea N.E. Br., D. botryoides Bak. and D. rnaculata Lindl. & Paxt.,
each occurs in three different phytochoria. Satellite populations (near endemics) of D.
botryoides and D. atropurpurea, are of marginal presence in ranges. Drirniopsis
botryoides has been widely collected in Kenya, Zanzibar, Tanzania, Pemba Islands and
Burundi (Table 11.2). One of the collections, the holotype for D. erlangeri Damrner
collected by Dr. Ellenberg 2043 (April 1901) (K!), records locality as 'Somaliland:
Borau Tarro-Gumbe'. Tarro-Gumbi, according to latest records is in Ethiopia. This is
just one example of locality uncertainties due to obsolete place names. Drirniopsis
maculata occupies widely separated localities in southern and tropical Afiica (Table
11.2). The plants in the Somalia-Masai phytochorion are probably relicts that have
managed to survive in refuges. To conclude that they could have been the result of jump
dispersals would need assessment of all other taxa, flora and fauna, of the area. If these
were to display the same behaviour as D. maculata, then they would be vicarious taxa.
A similar scenario exists with reference to D. rosea A. Chev. which occurs in two very
distant geographic ranges. Drimiopsis rosea has been collected in Chad, Cameroon and
a single specimen from South Afiica (one specimen, Venter s.n. (NH) (Table 11.2).
Drirniopsis burkei Bak., D. perfoliata Bak. and 0.woodii Bak. occur in two different
geographic ranges. The wide distribution of D. burkei in the southern African region
(Table 11.2) suggests previous wide continuous distribution that was interrupted leading
to disjunction. This has resulted in discontinuous distribution with pockets of
populations successful in isolated but similar habitats or refuges. Drirniopsis perfoliata
and D. woodii possess a similar type of disjucnt distribution but in confined areas. They
are marginally present, near-endemics in Tanzania and South Africa respectively. The
occurrence of D. perfoliata in Somalia is based on one herbarium specimen of L. Friss,
V. Alstrup & A. Michelsen s.n. (K!).
The remaining taxa are confined to specific areas possessing neoendemic
characteristics. Neoendemism is illustrated in the terminal taxa on the cladogram, Figure
11.5. Drimiopsis fzscheri (Eng1.) Stedje is known from only one herbarium specimen,
Fischer 325 (K!) collected in Tanzania. The remaining species, namely D. carrii
Lebatha (KwaZulu-Natal), D. kikiae Lebatha (KwaZulu-Natal) and D. stolonissima U.
& D. Muller-Doblies (Mpumalanga) occur in southern African with highest diversity in
the KwaZulu-Natal and Mpumalanga. This suggests these areas are their centres of
origin (Table 11.2). Drimiopsis comptonii U. & D. Muller-Doblies, D. reiNeyana U. &
D. Miiller-Doblies and D. pusisiNa U. & D. Muller-Doblies are of Swaziland origin. All
these remaining taxa have a restricted distribution; some found in single localities and in
small areas as small as approximately 4 m2 in the case of D. comptonii. The limiting
factors in the establishment of Drimiopsis are primarily external: shade, moisture and
because of their delicate size, some type of security in the form of rocks, boulders, trees
and dead leaves or, rarely, richly dense grassland. The plants prefer north facing hill or
mountain slopes, where there are thickets of shrubs.
Species richness in Drimiopsis reflects two nuclear areas, in South Afiica and in
Tanzania that mirror one another. Drimiopsis shows taxa richness in the Mpumalanga
and KwaZulu-Natal Provinces of South Afiica and the Iringa and Mpanda Districts of
Tanzania (Table 11.2). The Mpumalanga Province appears to be the centre of endemism
of Ledebouria and Resnova. The lowest diversity is in the Cape regions and winter
rainfall regions where only Ledebouria occurs.
11.4.2 Kistory/evolution and relationships
The Gondwanaland hypothesis is well supported in the distribution patterns of the
Ledebouriinae. Wide distribution of Ledebouria in Africa, India, the Mediterranean and
Madagascar, suggests pre-separation origin. When the continents separated, some of the
taxa moved with the Indian continent and spread into the Mediterranean regions.
Ledebouria thus became basal within the Ledebouriinae as illustrated by the cladistic
analysis of the Ledebouriinae.
The African continent, after separation moved northwards, away from the Arctic. The
continent, experiencing separation and movement tension, produced rift valleys (Figure
11.4 B). Madagascar is the result of one such a rift. The presence of Ledebouria,
Avonsera (Chapter 12) and the two Manning et al. (2004) taxa (L. cryptodata (Bak.)
J.C. Manning & Goldblatt and L. nossibeensis (H. Perrier) J.C. Manning & Goldblatt)
on the western coastline of Madagascar support this hypothesis.
The distribution pattern of Drimiopsis may be explained via the African continental
rifts. The western African rift caused new barriers in western parts of Afiica where there
was species continuity. Only one species, D. barteri, still occupies this region. Its
disjunct distribution spreads into the Central AGican Republic and the DRC. This rift
separated D. rosea and has refuges in Chad and Cameroon resulting from the rift. The
Ethiopian rift, Eastern rift and western rifts in eastern Afiica, forming the Rift Valley,
also separated taxa. These are the areas with D. botryoides, D. perfoliata, D. fischeri
and D. barteri as well as D. maculata relicts. The barrier in southern Afiica was
probably the Swaziland-Natal section rift. This separated taxa promoting development
of neoendemics like D. comptonii, D. reilleyana and D. pusilla exclusively distributed
in Swaziland. These drastic occurrences triggered chains of events, spanning over years
creating the species diversity of today. The zoning of the phytochoriological regions of
Africa demonstrates this scenario on a larger scale-notably,
sympatry normally exists
in any form of vegetation demarcation into regions.
The African basins, caused by previous masses of ice Gom the Gondwana era, and the
African plateaus have formed favourable habitats and protection for extant Drimiopsis
endemic species, neoendemics, choloendemics and palaeoendemics (Figure 11.4A).
These species, mostly newly described, survive in very localised and restricted habitats.
The genus Drimiopsis is only lately being noticed and having new species described.
This is the result of fieldwork extending into areas that were previously inaccessible.
The small stature and camouflage mechanisms of spotted, banded or streaked leaves of
these plants also render them cryptic.
The relationships between taxa on the cladogram, Figure 10.1 @age 143), portray
Ledebouria, the basal clade within the Ledebouriinae, as the oldest taxon of the group.
The Gondwana and rift hypothesis supports this, as Drimiopsis and Resnova do not
display the wide range distribution pattern of Ledebouria. Resnova clade primarily
possessing intermediary characters, displays restricted distribution. It appears Resnova
is a 'habitat specialist', a holoendemic species confined to isolated habitats with special
environmental conditions. The basal taxa within the Drimiopsis clade include tropical
African and two wholly southern African taxa (Figure 11.5). The taxa in this basal
clade, the implication being they are the most primitive of Drimiopsis taxa, possess light
green coloured thick textured leaves that allow them to survive harsh climatic
conditions (Figure 11.5).
11.5 CONCLUSION
Ledebouria is the basal taxon of the Ledebouriinae group. It also occurs outside of
Africa, and has centres of endemism in Mpumalanga and KwaZulu-Natal Provinces of
South Africa. It is the oldest in origin from whence Resnova and Drimiopsis ancestors
originated in the Mpumalanga and KwaZulu-Natal Provinces. The Resnova centre of
endemism is in the Mpumalanga Province. Drimiopsis appears to be the most recent
taxon with predominantly tropical Afiican basal taxa and seemingly two nuclear areas
of distribution in the Mpumalanga Province of South Africa and the Iringa District of
Tanzania. The overall centre of endemism of the Ledebouriinae is the MaputolandPondoland regional mosaic- 15.
This study is the first comprehensive geographic distribution analysis of the
Ledebouriinae. The paucity of plant collections-
uneven and inadequate collections-
notwithstanding, this study presents a starting point for the inquiry into the intricacies of
geographic distribution, evolution and development within this group.
12. TAXONOMY
-
Z).
1
Z). ea'lIZii
Z).
2
,--,
3
'------'
r-
Z).
4
Z).
5
Z).
6
Z).
7
Z).
,-8
Z).
9
Z).
10
Z).
-
11
Z).
Z).
12
-"
13
'UJ4ea
Z). ft««tia
14
Z).15
Z).
16
Z).
17
Z). «etU
The new names of species presented in this thesis are not viewed effectively published.
188
------
-
- -
- -
18
12.1 GENERIC DESCRIPTION
Drimiopsis Lindl. & Paxt. in Paxton's Flower Garden 2: 73 (1851-52). -Type:
Drimiopsis maculata [icon in] Paxton's Flower Garden 2: 73 (1851-52) (lectotype, hie
designatus).
Plants medium-sized (10.1 cm to 15 cm high), rarely smaller or larger; protantherous to
synantherous, rarely hysteranthous; annuals, bulbaceous. Bulbs hypogeal; gregarious,
simple tunicated, some with tuberescent h d u s , some stoloniferous, whitish to
greenish, roundish to ovoid, small (2.5 cm across or less) to large (more than 5 cm).
Bulb scales loosely packed, when tom without threads, greenish, outer scales white or
purplehrown; fleshy, rarely membranous. Leaves 1 , 2 or 3 or more; erect, spreading or
rarely appressed to the ground; cordiform, falciform, linear or spathulate; ovate to
lanceolate; 1.1 to more than 20 cm long, 2 cm or less to more than 4 wide; when tom
without threads; pseudopetiolate or sessile. Pseudopetiole absent or present, when
present, exceedingly shorter than, as long as or exceedingly longer than lamina; banded
or tinted. Leaf margins entire, crenate, crenulate to undulate, simple, edged or banded
purplehrown. Lamina thick or membranous, adaxially spotted or unspotted, abaxially
green or tinted, if tinted then abaxially streaked purplehrown; apex mostly acuminate,
rarely acute or obtuse; base mostly cuneate, sometimes attenuate, green to tinted dark
purple, streaked or banded. Indumentum absent or present, in the form of papillae or
hairs, sparse, frequent or dense, arranged in rows or randomly; distributed on
pseudopetiole or lamina or both. Stomata sparsely distributed, anomocytic, with
shallow crypts and subsidiary cells forming an H-complex with stomata. Epidermal
cells adaxially shortly polygonal, abaxially elongately tetragonal. Anticlinal cell
boundaries generally channelled; anticlinal cell boundaries irregularly sinuate to
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undilrate; periclinal wall curvature with tabular-convex cells rarely with non-tabular
convex cells. Cuticle striae regular to irregular. Inflorescence a simple raceme, rarely a
pseudo-corymb, solitary or few (one to two per bulb), erect, spreading, rarely flaccid,
shorter than, equal to, or considerably longer than leaves. Rachis cylindrical, conical or
ovoid cylindrical, generally short (1-10 cm) or 11-25 cm long. Peduncle generally
evenly coloured, rarely banded. Bracts and prophylls vestigial or absent in mature
inflorescence. Flowers 10-15 to more than 30, sparsely or densely spaced, minutely
pedicellate (<0.1 cm long), or shortly pedicellate (0.1 to 0.4 cm long), rarely with long
pedicels (>0.4 cm long); minute, small or medium-sized (0.1-0.6 cm), actinomorphic,
campanulate rarely tubular; hypanthium base rounded. Tepals dimorphic, free, rarely
fused at base, predominantly whitish to greenish, creamish to pinkish or rarely purplish.
Outer tepals comivent, imer comivent to recurved rarely spreading. Outer tepal apex
is cucullate to comivent, imer cucullate-comivent with conduplicate apex margin.
Vitta faint to conspicuous, rarely absent. Androecium epitepalous, white to cream,
erect, uniseriate, inserted at throat of short or rarely long perianth tube, shorter than
pistil to as long as pistil. Filaments free to valvate, deltoid to acuminate, base broad.
Anthers dorsifixed. Gynoecium tricarpellate, ovules two per locule. Stigma roundish,
stigmatic papillae sessile, trilobal, surface corrugated. Style shorter than ovary,
roundish. Ovary sessile, globose, smooth, shoulders and ridges absent. Nectaries
present. Pollen isomorphous/monosporous, equatorial view depressed ovate, polar view
elliptic, laterally blunted; subequiaxe, distal pole curved, ornamentation punctate.
Karyology x = 10, x
=
11. Flowering is commonly around July to August in tropical
African species and from September to November (rarely January to February) in other
taxa.
Diagnostic characters
Drimiopsis species are generally small geophytes with thick, usually erect, spotted
leaves growing in shady areas mostly among rocks. The abaxial leaf surface is usually
purple to brownish or streaked. The epidermal cells possess irregularly sinuate
anticlinal cell boundaries with periclinal wall curvature formed by tabular convex cells.
The short-lived inflorescence bears minutely to shortly pedicellate flowers possessing
dimorphic tepals that are connivent and cucullate with conduplicate apex margins. The
greenish filaments are deltoid to acuminate. The globose sessile ovary possesses a
round stigma with subsessile trilobal stigmatic papillae. Pollen grains are depressed
ovate (equatorial view) with blunted ends laterally, a subequiaxeequatodal diameter
p
p
p
p
p
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p
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and a straight distal pole. Sparsely distributed stomata usually form an "H-complex"
with subsidiary cells, adaxial epidermal cells are shortly polygonal.
Distribution and habitat preference
Drimiopsis comprises 18 species variously distributed in Ghana, Togo, and northern
Nigeria in areas of Sokoto, Zaria and Yola. It also occurs along the Chari-Banguirmi
region of Chad into the uppermost Central African Republic. Distribution spans the
eastern parts of Africa into Sudan, the Gambela region of Ethiopia, Somalia and
Zanzibar. In Kenya it occurs in the Taita District, Kilimanjaro, Tsavo National Park.
Tanzanian collections are from the Lushoto, Tanga, Mbulu, Iringa, Kyimbila, Mpanda,
Morogoro and Rungwa Districts and along the coast. It has been collected in the
Bujumbura areas of Burundi. From the Central African Republic distribution extends
towards southern Africa through Katanga regions of the Democratic Republic of
Congo, Kasama District of northern Zambia into Harare, Zimbabwe. In South Africa it
is widely distributed in the summer rainfall areas around Gauteng, North West, Free
State, Mpumalanga, KwaZulu-Natal and Eastern Cape. It occurs in Swaziland, Maputo
and the Inhaca Island of Mozambique. It also occurs in the eastern parts of Botswana in
Gaborone, Kgatleng and Mahalapye. (Figure 11.1 A & B)
Drimiopsis grows in a wide range of shaded habitats in grasslands, wet marshes,
bushland or woodland. It often grows in mountainous areas, among rocks in shaded
areas and under boulders as well as near rivers and streams, on densely grassy slopes,
areas with bush clumps, along river bushveld and coastal forest fringes. It requires
damp areas where there is plenty of leaf litter. Drimiopsis grows in most soil types
favouring dark clayish or sandy soils, brown sandy clay, dolomitic rock, granitic ridges
or hard, dry ground along footpaths at elevations of up to 1600 m above sea level.
Taxonomic note
The sinking of Resnova under Drimiopsis (Phillips, 1951 ; Jessop, 1970, 1972; Dyer,
1976; Arnold & De Wet, 1993; Meyer and Williams, 1997) or of both Resnova and
Drimiopsis under Ledebouria (Manning et al., 2004) is not supported by my analysis.
The significant number of morphological differences between Drimiopsis, Resnova and
Ledebouria support generic ranking for each of the three taxa.
.
.
12.2 KEY TO SPECIES OF DRIMIOPSIS LINDL & PAXT
Leaves mainly 1 ................................................................................................................................. 2
Leaves mainly 2 .................................................................................................................................5
Leaves 3 or more ................................................................................................................................8
Plants dwarf, less than I Ocm; inflorescence solitary..........................................................................3
Plants medium-sized (10.1 to 15 cm); inflorescence one to two per bulb ..........................................4
.
Leaves erect; cordifonn; leaf margin crenate; lamina base attenuate.............................D queae (18)
Leaves spreading; linear; leaf margin undulate; lamina base cuneate............................ D rosea (13)
.
.
Leaves linear; sessile; lamina base cuneate; leaf margin entire ...................................D barteri (12)
Leaves cordate; pseudopetiolate; margin crenate; lamina base attenuate .............D. liniupapilla (17)
Bulbs stoloniferous ........................................................................................................................... 6
Bulbs non-stoloniferous ..................................................................................................................... 7
.
Plants dwarf(>10 cm); leaves erect; lamina base attenuate; bulbs ovoid .................D comptonii (7)
Plants medium-sized(lO.l to 15 cm); leaves appressed to the ground; lamina base cuneate;
bulbs roundish ............................................................................................................... D carrii (2)
.
.
Bulbs roundish; leaves appressed to the ground; cordifom; ovate...........................D perfoliata (3)
Bulbs ovoid; leaves spreading; linear to lanceolate .................................................... D burkei (11)
.
Lamina base attenuate....................................................................................................................... 9
Lamina base cordate........................................................................................................................ 10
Lamina base cuneate ....................................................................................................................... 11
.
Leaves linear to ovate; leaf margin undulate; pseudopetiole banded ............................. D woodii (9)
Leaves cordifom; to lanceolate; margin crenate; pseudopetiole tinted ........................ D kikiae (16)
.
.
Bulbs roundish; plants robust. 4 5 cm; leaves spreading; pseudopetiole banded ......D maculata (5)
Bulbs ovoid; plants medium.sized. 10.1 to 15 cm; leaves erect; pseudopetiole
tinted ................................................................................................................ D atropurpurea (15)
.
.
Plants dwarf, >10cm ....................................................................................................D pusilla (14)
Plants medium.sized, 10.1 to 15 cm ...............................................................................................12
Plants robust, 4 5 cm ................................................................................................D botryoides ( 4 )
.
.
Lamina apex acuminate ................................................................................................D c h i (8)
Lamina apex acute .......................................................................................................................... 13
Lamina apex obtuse ..............................................................................................D davaoniae (6)
.
.
Bulbs roundish; leaves spreading; margin entire; bulbs stoloniferous.................. D stolonissima (1)
Bulbs ovoid; leaves erect; margin undulate; bulbs non-stoloniferous .....................D reilleyana (10)
.
12.3 SPECIES DESCRIPTIONS
12.3.1 Drirniopsis stolonissima
Drimiopsis stolonissima (U. & D. Miiller-Doblies) Lebatha comb. nov.
Drimiopsis burkei Bak. subsp. stolonissima U. & D. Miiller-Doblies in Feddes Repert.
108: 64 (1 997). Type: Muller-Doblies & MNer-Doblies 770173, Strydom Tunnel,
second parking are south of the tunnel; north west facing slope with Euphorbia-trees,
Pilgrims rest, Mpumalanga (B!, holotype; BTU!, Z! isotype).
Ledebouria burkei subsp. stolonissima (U. & D. Miiller-Doblies) Manning & Goldblatt
in Edinburgh. J. Bot. 60(3): 560 (2004).
Description: Figure 12.2
Habit and bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; stoloniferous; with
tuberescent fbndus absent; whitish; roundish; 2-4 cm across. Bulb scales loosely
packed; when torn without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; spreading; linear; lanceolate; (4-)7-12(-15)
cm long; 2-3 cm wide; when tom without threads; sessile. Leaf margin entire;
noncartilaginous; bordered purple/brown. Lamina thick; spotted adaxially; green. Leaf
apex acute. Leaf base cuneate; green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
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shortly polygonal; abaxiallj. elongately tetragonat;ariikliiid 6 0 ~ d a r i e channelled
s
and
- - - - - - -
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, dense, raceme; erect; considerably
longer than leaves. Rachis conical; 14-20 cm long. Peduncle green. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30, small (2.1-4 mrn long);
minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous; campanulate;
hypanthium base rounded. Tepals dimorphic; whitish to greenish; hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted in throat of perianth tube; as long as pistil. Filaments valvate;
deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile; trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis stolonissima has been collected only in the vicinity of Pilgrim's Rest, South
Africa (Figure 12.1). It grows in the shade of trees on the northwest facing slopes of
Strydom tunnel in the Abel Erasmus pass.
Diagnostic characters
Drimiopsis stolonissima possesses fleshy, light green, adaxially spotted, spreading and
lanceolate leaves that taper to an acute apex. The plants are stoloniferous, a habit shared
with three other Drimiopsis species and possess medium sized bulbs.
Taxonomic note
Drimiopsis burkei subsp. stolonissima U & D Miiller-Doblies is elevated to species
rank. The morphological, geographical and ecological differences between the
subspecies are equal to the differences among all other Drimiopsis species studied. The
cladistic analysis places D. burkei and D. stolonissima in different clades with D.
stolonissima in the basal clade. Seventeen differences exist between D. stolonissima
and D. burkei, notably, D. stolonissima is 3 or more and possesses fleshy lighter green
leaves that are always adaxially spotted. Drimiopsis burkei on the other hand is 2
(sometimes 3 or more) and possesses a lamina that is sometimes adaxially spotted but
always abaxially streaked purplish/brownish with a dark purple tinted base.
Specimens studied
-2430:
Strydom Tunnel, second parking are south of the tunnel; north west facing
slope with Euphorbia-trees, Pilgrims Rest, Mpumalanga (-BC), Miiller-Doblies &
Miiller-Doblies 77017b (B, BTU, Z); Strydom Tunnel, Abel Erasmus pass, Pilgrims
Rest, Mpumalanga, (-BC), Lebatha 037 (PUC, PRE).
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Figure 12.1: Known distribution of D. st%nissima.
195
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Figure 12.2: Drimiopsis st%nissima (Lebatha037)plant (life size). 1, bulb with bulbils; 2, part of
the inflorescence; 3, adaxially spotted leaves; 4, sectioned flower (x40) with cucullate and
connivent tepals and pistil.
196
---
12.3.2Drimiopsis carrii
Drimiopsis carrii Lebatha sp. novo -Type:
South Africa, KwaZulu-Natal Province,
Amanzimtoti, along a stream, south of Durban: 300-03'S 300-53'E Lebatha 015 (PRE
holotype, PUC isotype).
Folia prostratis, D. perfoliata remote affmis sed differt folia dilatus-viridia et guttatus
margine crenutus; floribus albus.
Description: Figure 12 4 -12.6
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; stoloniferous; with
tuberescent fundus absent; whitish; roundish; 1-2 cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 2; appressed to the ground; cordiform; ovate; 6-8(-10)
cm long; 3-5 cm wide; when tom without threads; sessile. Mature leaf margin crenate;
noncartilaginous; edged purple/brown. Lamina thick; spotted adaxially; green. Leaf
apex acute. Leaf base cuneate,;green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thick. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, dense, raceme; erect; shorter than
leaves. Rachis cylindrical; 5-10 cm long. Peduncle green. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 mm long);
minutely pedicellate (shorter than 0.1 cm long); actinomorphic; sextepalous;
campanulate; hypanthium base rounded. Tepals dimorphic; whitish to greenish;
hypanthium inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect;
epitepalous; uniseriate; inserted in throat of perianth tube; as long as pistil. Filaments
free; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile and trilobal. Style shorter than ovary; terete.
197
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis carrii is known only from Amanzimtoti, south of Durban, South Africa
(Figure 12.3) and grows in shaded grassland areas in large amounts of organic matter.
Flowering time September to December.
Diagnostic characters
Drimiopsis carrii is similar to D. perfo/iata. Of the nineteen differences between the
two taxa, the wider, lighter green, semisucculent, adaxially spotted, broadly ovate
leaves with acuminate apices, crenate and edged margins (cleft in some areas) as well
as the minute and white flowers, and the inflorescence shorter than the leaves
characterise D. carrii. Drimiopsis carrii leaf margins resemble those of D. burkei, but
are deeper edged.
Specimens studied
-3030:
South Africa, KwaZulu-Natal Province, along a stream, Amanzimtoti, south of
Durban, 30° 03'S 30° 53'E (-AD), Lebatha 015 (pRE holotype, PUC isotype).
198
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Kilometers
Figure 12.3: Known distribution of Drimiopsis carrii.
199
Figure 12.4: Drimiopsis carrii (Lebatha 015) (life size). 1, a simple tunicated, white,
roundish bulb with a small tuberescent fundus; 2, inflorescence with minutely pedicellate
flowers; 3, sectioned flower exposing the pistil (x70).
200
Figure 12.5: Drimiopsis carrii (Lebatha 015). Leaves spotted adaxially and with
crenate margins (life size to x 0.5).
..
Figure 12.6: Drimiopsis carrii (Lebatha 015): developing inflorescence and young leaves
(life size).
201
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----
12.3.3 Drimiopsis perfolircra
Drimiopsis perfoliata Bak. in Gard. Chron. 10: 364 (1878); in Pflanzenw. Ost-Afi.
Zanzibar, Kirk s. n. sub K HI 525/82 ( K !, lecotype, hic designatus) hic(
(1878). -Type:
restituta.
Drimiopsis botryoides Bak. subsp. prostrata Stedje in Nord. J. Bot. l4(l): 49 (1994).
-Type:
Bjwnstad 564, Tanzania, Iringa District ( 0!,holotype; K! , isotype).
Ledebouria botryoides subsp. prostrata (Stedje) Manning & Goldblatt in Edinburgh J.
Bot. 60(3): 560 (2004).
Description: Figure 12.8
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
(
tuberascent fundus absent; whitish; roundish; 2-2.5(-4) cm across. Bulb scales loosely
packed; when tom without threads; outer scales greenish and fleshy.
Leaf morphology. Leaves 2; appressed to the ground; cordiform; ovate; (6-17-9(14) cm long; 4-8 cm wide; when tom without threads; sessile. Leaf margin entire;
noncartilaginous; bordered purplelbrown. Lamina thick; unspotted; green. Leaf apex
acute. Leaf base cuneate; green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thick. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermis cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; considerably
longer than leaves. Rachis cylindrical; 4-7 cm long. Peduncle green. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30, small (2.1-4 rnm long);
minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous; campanulate;
hypanthium base rounded. Tepals dimorphic; whitish to greenish; hypanthium
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c6ri@icuous. Outer whorl of tepals recurved; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted in throat of perianth tube; as long as pistil. Filaments valvate;
-
-
deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitishlgreenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis perfoliata occurs in the Iringa, Mbulu and Mbeye regions of Tanzania,
Zanzibar and Somalia. It has been collected along the Mara River in Mswisuri, on the
Uluguru Mountains south of Morogoro and in the Mpwapwa region (Figure 12.7). It
usually grows in rocky areas, scattered among the rocks and in open or cleared
woodland. Flowering time, July to August.
Diagnostic characters
D. perfoliata bulb is globose, about 2-2.5(-4) cm in diameter, the outer scales greenish.
The plants generally possess 2 leaves that are broadly lanceolate and appressed to the
ground. The lamina is thick, generally unspotted and uniformly green with a cuneate
base. The flowers are minutely pedicellate, almost sessile. See D. carrii for differences
between D. perfoliata and the aforementioned.
Taxonomic note
Baker (1878) mentions that his new D. perfoliata is based on a plant collected by Dr.
Kirk in Zanzibar and grown at Kew Gardens, but does not cite any specimens. The
plant is described as possessing typical Drimiopsis 'inflorescence and perianth' but
with 2 ovate leaves appressed to the ground and possessing a cuneate base. There is a
Kirk specimen in Kew collected in Zanzibar in 1873. Kirk s.n. sub K H1535/82 in all
probability hales from the same collection that provided the mentioned living material
for Kew. I therefore designate the specimen as the lectotype of D. perfoliata. Kirk
initially detted his collection as "Drimiopsis n. sp." but later changed his mind and
considered it to be D. botryoides.
As far as could be ascertained, Bjerrnstad has been the only person since Baker (1878)
to employ the name D. perfoliata when he detted the collection as such (Bjmnstad 534,
Figure 12.8 A). Stedje (1994), however, considered the name D. perfoliata 'dubious'
and opted to typify her D. botryoides Bak. subsp prostrata Stedje with the mentioned
Bjerrnstad specimen.
Being guided by the cladistic analysis in this thesis, which supports a reranking of D.
botryoides subsp prostrata, D. perfoliata is resuscitated and D. botryoides subsp
prostrata synonimised there under.
Kilometers
Figure 12.7: Known distribution of Drimiopsis perfoliata based on herbarium specimens and
Stedje (1993).
.
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HERB, HORT. KEW.
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,~~
;
I$CJTYfE! COOP)
ORI_IOPSIS
..p.
Diet.
s..n
BO'I'RYOlmal Bek.
1WJ8'I"RA""
Stedje
B. St8dj.
tor
'.T.E.A.
199:l
U I
'-'
~.
~
'~~~.
.
KIf~.UII..n. 011
\.
....
J i" .. / /_
) --
~
1.
I,oa... LI' 1'110'.11
~S,.-bc~of/".,...". , di..~
"""'...-.
A.P~H~~R."iA
'DT;~;.r~i5
b..,!",!
~
~.iOcS
folo<:.l~~'!.
&1::
/
/'
/-
>
'"
I
/I
UIUt
".,-/
t'. //
'-/' Iii
l.-
Figure 12.8: A, Bj(j)rnstad564 (K), isotype of D. botryoides subsp. prostrata; B, Kirk s.n. sub K H/535/82 (K), the designated
lectotype of D. perfoliata.
I
205
Specimens studied
-4144: Shabeellaha Hoose, Somalia (-DD), Friss, Alstrup & Michelsen (K).
-0119:
Mbeye District, Usango flats, Mswisuri (-AA), Leedal 5198 (K); Mbeye
District, Usango flats, Mswisuri (-AA), Leedal5298 (K).
-0234: Mara River Guard Post (-DA), Greenway 10,235 (K).
-0436: Mbulu District, Tarangire National Park (-AA), Richards 24734 (K).
- 4 6 3 6 Mpwapwa, Tanzania (-CA), Hornby 2093 (K); Hornby 7380 (K).
-0637 Turiani near Diwali River (-BA), Milne-Redhead & Taylor 7357 ( K ) .
-0639 Zanzibar, Kirk no. Vl87.3 (K) -BB.
-0734
Rungwa Game Reserve (-AC), Richards 20755 (K).
-0735
T.7, Iringa District, Iringa (-AA), Bjarnstad, I. 564 (K, 0),Bjmnstad, A.B.
2035 (K), Greenway & Kanuri 14963 (K); Iringa District, Ibumu Village (-DD),
Richards 15612 (K).
- 4 8 3 5 Ukam., Tanzania (-DC), Stuhlmann 9308 (K).
-0838 Kingupira (-DA), Vollesen 3025 (K).
12.3.4 Drimiopsis bofryoides
Drimiopsis botryoides Bak. in Saund. Ref. Bot. 3. App. 17 (1870); Bak. in Fl. Trop.
Afr. 7: 543 (1898); Agnew & Hanid Upland Kenya Wild flowers: 697 (1966); Molehill
in E. Afr. Nat. Hits. Soc. Bull. 24: 18 (1962); -Type:
Africa orientalis without precise
locality, Blackburn s. n. sub K 1867 (K!,holotype).
Drimiopsis holstii Bak. in Engler, Pflanzenw. Ost-Afi. C: 143 (1895); Bak. in Fl. Trop.
Afr. 7: 544 (1898). -Type:
Tanzania, Usambara, Mlalo, Holst 619 (B!,holotype).
Scilla volkensii Engl. in Pflanzenw. Ost-Afr. C: 142 (1895). -Type:
Tanzania,
Kilimanjaro below Marangu, Volkens 2164 (B! holotype).
Drimiopsis kirkii Bak. in Gard. Chron. 2: 644 (1874); F1. Trop. Afr. 7: 543 (1898). -
Type: Zanzibar, Kirk s. n. (K!,holotype).
Drimiopsis stuhlmannii Bak. in F1. Trop. Afi. 7: 544 (1898). -Type:
Tanzania,
Usararno, Stuhlmann 9308 (B!, holotype).
Drimiopsis erlangeri Damrner in Bot. Jahrb. 38: 63 (1905). -Type:
Ethiopia, Tarro-
Gumbi, Ellenbeck 2043 (B!, lectotype, hic designatus).
Drimiopsis bussei Darnrner in Bot. Jahrb. 38: 62 (1907). -Type:
Tanzania, Mandandu,
Bussei 526 (B!, holotype; EA, isotype).
Drimiopsis botryoides Bak. subsp. botryoides Stedje in Nord. J. Bot. 14(1): 49 (1994).
Stedje, & Thulin, Nord. J. Bot. 15: 594 (1995).
Ledebouria botryoides (Bak.) Manning & Goldblatt in Edinburgh J. Bot. 60(3): 560
(2004).
Description: Figure 12.10
Habit & bulbs. Plants robust (more than 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus present; whitish; roundish; 2-2.5 cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; spreading; falciform; lanceolate; (8-) 10-20(45) cm long; 3-5 cm wide; when tom without threads; sessile. Leaf margin entire;
noncartilaginous; bordered purplehrown. Lamina thick; spotted adaxially; green. Leaf
apex acute. Leaf base cuneate; green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thick. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
straight; periclinal wall curvature straight tabular; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, sparse raceme; spreading;
considerably longer than leaves. Rachis cylindrical; 20-25 cm long. Peduncle green.
Bracts in mature inflorescence vestigial. Prophylls absent. Flowers 16 to 30; small
(2.1-4 mrn long); minutely pedicellate (shorter than 0.1 cm); actinomorphic;
sextepalous; campanulate; hypanthium base rounded. Tepals dimorphic; whitish to
greenish; hypanthium conspicuous. Outer whorl of tepals recurved; longitudinally
cucullate; apically conduplicate. Inner whorl of tepals connivent; longitudinally
cucullate; apically conduplicate. Vitta faint. Stamens 6; greenish to whitish; erect;
epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments
valvate; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two
per locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitishlgreenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Diagnostic characters
Drimiopsis botryoides is characterized by thick semi-succulent, light green, falciform,
lanceolate, maculated leaves with acute apices in combination with greenish flowers.
Distribution and habitat preference
Drimiopsis botryoides occurs in the highlands of Ethiopia and Somalia spreading into
Zanzibar, Kenya in the Taita District, Kilimanjaro, below Marangu and along the Athi
River, Tsavo National Park and near Fourteen Falls. In Tanzania it is distributed in the
Lushoto districts, west of Usambara Mountains, Tanga District, along the coast and
near Mount Tanganyika. Distribution is also in Turiani, Nkulumuzi Valley near the
Amboni Caves, Mbulu District, and Lake Manyara National Park and on the Bongoyo
Island off Msasani Bay. Distribution extends into Bujumbura areas of Burundi (Figure
12.9). Drimiopsis botryoides has been collected in bushland, woodland, on sandy soils
and often among rocks, up to 1600 m above sea level. Flowering time September to
December.
Taxonomic note
Drimiopsis erlangeri is here synonymised under D. botryoides. Thulin has probably
erred in determining the cyanotypes of D. erlangeri as Drama botryoides instead of
Drimiopsis botryoides. The lectotype and syntype of D. erlangeri do possess some
characters reminding of a young D. maculata, e.g. the pseudopetiole and cordate leaf
base (Figure 12.1 0). For the moment though, I synonimise it under D. botryoides until
new data proves otherwise.
Kilometers
Figure 12.9: Known distribution of D. botryoides based on herbarium specimens and Stedje (1993).
Ill'Ir:
A
)l
II
II' !!~II
rB ---I
I
!
!
.,
b~,'
...
F
E
=1
...;~;...'"J"'"'...,
...~..\
~ a.....
.<
~
,
"""
f-
,....
(~
~
~
Museum botanicum Berolinense.
I
..."
J.., "-r0' r-,...
-
.
. J.
.
M"..um b:~,.nic":..Berolinen...
_')i
~
'4...1'''' ~_..f'.:I"
---
Figure 12,10: Drimiopsis erlangeri. A. syntype; B, designated 1ecotype..
211
Figure 12.11: Drimiopsis botryoides: A, mature plant with inflorescences exceedingly longer than the
leaves (xO.25); B, a young developing leaf (life size) (Lebatha 003 (PUC); C, sessile, spreading and
falciform leaves (xO.25)(Lebatha 098 (PUC).
212
---
-
-
---
-
---
Specimens studied
-4137: Near Fourteen Falls, near Donyo Sabbuk, Central Province, Kenya (-DD),
Mwache, Rayner 440 (K);Lebatha 003 (PUC) cult. Nordal113 (0).
-0237: Tsavo National. Park, Worssera (-DD), Greenway & Kanuri 12854 (K).
-0329: Bujumbura. Plaine de la Rusizi, Burundi (-BA), Lewalle 4839 (K).
-4335: Tanzania Mbulu District, Lake Manyara National Park (-DB), Masasa,
Greenway & Kirrika 11105 (K).
-0338: Taita District, 18km fi-om Nairobi to Taveta, via Taveta Road (-BC), Faden,
E.R.B. & Faden, A.J. 74/505 (K), Drummond & Hemsley 2524 (K).
- 0 0 3 7 : Ngeng-Mathew Road, Kenya (-BB), Newbould 31 77 (K).
-4337: Manolo River, Kenya (-DA), Kassner 319 (K).
-0338: Kilimanjaro, below Marangu 300ft. Kenya (-AA), Volkens 2164 (K).
-0426: Lushoto District, West Usambara Mountains (-AB), Faden, R.B., Phillips,
Muasya & Macha n 96/11 (K).
-0438: Lushoto District, T.3, 3.7 km from Lushoto on Lushoto-Mombo road.
Mswaha, Tanzania (-CD), Archibald 3243 (K).
-0439: Kwale District (-BA), Luke 3849 (K); Tanga District, Bomandani, (-CA),
Drummond & Hemsley 3678 (K).
-4636: Tanganyika Territory, north of Mofwapwa (-BC), Hornby 7380 (K).
-0637: Turiani, Tanga district, Tanganyika (-BA), Drummond & Hemsley 2022 (K);
Morogoro District (-CC), Wallace 5 76 (K).
-0639: Zanzibar (-BB), Blackburn s.n. sub K 1867 (K); Bongoyo Island, Msasani Bay
(-CD), Batty 931 (K), Batty 518 (P).
-4735: Morogoro Road (-DD), Hamid 2890 (NHM).
Without precise locality:
Tanzania, Mbudya Islands, Lebatha 098 (PUC) cult. Nordal 1600, Lebatha 004 (PUC) cult.
Bj0rnstad 773; Vanderyst 12129 (BR); Helen Faneknin 1861 (K); Helen Faneknin
-
-
-
-
-
-
1964 (K);
~'biltkriif
1861(BR).~Faufhm-f 964 -(£%It).-Tanganyika,Mrulumuzi Yalley
near Amboni Caves, Verdcourt & Greenway 253 (K). Tanzania: Schieben 5697 (P).
Gallahochland highland, Borau, Tarro Gumbi, Somaliland, Ellenbeck s.n. 2043, 2081
(B). Quarre 1441 (BR); SalGsiens 992 (BR); Belgian Congo, Schmitz 2.1 14 (BR).
Collectors uncertain
sub BR 893093 (BR); sub BR 899195 (BR); sub BR 899197 (BR); sub BR 899201
(BR); sub BR 899205 (BR); sub BR 899215 (BR); sub BR 899 220 (BR); sub BR
899216 (BR); sub BR 89921 7 (BR); sub BR 899218 (BR);sub BR 899219 (BR).
12.3.5 Drimiopsis maculafa
Drimiopsis maculata Lindl. & Paxt. in Paxt. F1. Gard. 2: 73, fig. 172 (185 1-52); Bak.
in Saund. Ref. Bot. 3, App. 17 sub t. 191 (1870); in J. Linn. Soc. 12: 227 (1870); in F1.
Cap. 6: 473, 478 (1896); Fl. P1. South Africa, Plate 304 (1944); F1. P1. Africa. 25: t.
Plate 957 (1946a); Jessop in J. S. Afr. Bot. 38(3): 159-1 60 (1972); Stedje in Nord. J.
Bot. 14(1): 48 (1994); Miiller-Doblies & Miiller-Doblies in Fedd. Repert. 108: 61.
(1997). -Type:
icon in Paxt. Fl. Gard. 2: 73, fig. 172 (1851-52) lectotype, hic
designatus.
Drimia petiolata Koch & Bouchk in Index seminum Berol: App. 3. (1861). -Type:
ex
horto Kewensi in Berolinensem allata, cult 1863 (B! syntype).
Ledebouriapetiolata Manning & Goldblatt in Edinburgh J. Bot. 60(3): 561 (2004).
Drimiopsis minor Bak. in Saund. Ref. Bot. 3, App. 17 sub t. 192 (1870); Bak in J. Linn.
Soc. 13: 227 (1870); Bak. in F1. Cap. 6: 472, 473 (1896). -Type:
I
KwaZulu-Natal,
Cooper i c (B! ,holotype).
Description: Figures 12.13 & 12.14.
Habit & bulbs. Plants robust (more than 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus present; whitish; roundish; (2-)2.2-3(-10)
cm across. Bulb scales
loosely packed; when tom without threads; outer scales greenish and fleshy.
Leaf morphology. Leaves 3 or more; spreading; cordiform; ovate; 4-9 cm long; 4-
6 cm wide; when tom without threads; pseudopetiolate. Pseudopetiole exceedingly
longer than lamina; banded. Mature leaf margin undulate; noncartilaginous; edged
purplehrown. Lamina membranous; spotted adaxially; green. Leaf apex acuminate.
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Leaf base cordate, almost 'saggitate'; g r e 6 Leavesglaljrous;
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
I
densely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
I
Inflorescence. One to two per bulb; a simple, sparse raceme; spreading;
considerably longer than leaves. Rachis ovoid-cylindrical; 15-60 cm long. Peduncle
green. Bracts in mature inflorescence vestigial. Prophylls absent. Flowers 16 to 30;
medium-sized (4.1-6 rnrn); shortly pedicellate (0.1-4 cm long); actinomorphic;
sextepalous; campanulate; hypanthium base rounded. Tepals dimorphic; whitish to
greenish; hypanthium inconspicuous. Outer whorl of tepals recurved; longitudinally
cucullate; apically conduplicate. Inner whorl of tepals connivent; longitudinally
cucullate; apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish;
erect; epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil.
Filaments valvate; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate.
Ovules two per locule. Stigma roundish; papillae sessile, trilobal. Style shorter than
ovary; terete. Ovary sessile; globose; transversely smooth; whitishlgreenish; shoulders
absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis maculata possesses a disjunct distribution, occurring in southern Africa as
well as Tanzania. Even in southern Africa, populations mostly found east and north east
(Mozambican) of the Drakensberg, rarely occurring in Gauteng. It has been largely
collected in Mpwapwa District of Tanzania but also elsewhere (Figure 12.12). The type
of habitat favoured by D. maculata is grasslands, brown sandy clay, along river
bushveld, dolomitic rock, along forest and coastal forest fringes as well as dune forest.
Flowering time is September to December.
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Diagnostic characters
Drimiopsis maculata possesses medium sized bulbs, is pseudopetiolate with bright
green, slightly membranous leaves, maculated a darker green. The margins vary from
entire in younger leaves to undulate in mature leaves (Figure 12.13). The leaf base is
mainly cordate. Drimiopsis maculata possesses spreading, shortly pedicellate, medium-
sized (4.1-6 mm) flowers. The medium sized flowers are uncommon in this genus
possessing predominantly minute flowers. The mature inflorescence is elongate and
spreading.
'}
.5
:.. .::: :"indsav
.4 'i r T"
del
Figure 12.13: Drimiopsis maculata. I, a plant in flower displaying a round bulb with loosely packed
fleshy scales, typical of Drimiopsis; 2, an undulate leaf; 3 a flower bud; 4, a mature flower (x25); 5, a
tepal with deltoid filament; 6, an outer tepal; 7, distal view of a mature flower; 8, the uniseriate filaments
and a globose ovary with the style shorter than the ovary; 9, a cross section of the ovary. (Van der Merwe
1946a).
218
---
Figure 12.14: Variation in leaf shape and margin in D. maculata (xO.5).A, a plant with an undulating leaf
margin and an almost sagittate leaf base growing in the field in Swaziland(Lebatha 062 (PUC); B, the
same plant growing at the botanical garden of the North West University Potchefstroom Campus with
leaves possessing entire margins and cordate bases; C, a plant from Berlin Botanical Gardens
possessesing undulate margins.
219
--
---
-
Taxonomic and nomenclature note
Nomenclature inaccuracies exist in the authorship of Drimiopsis. Baker (1870a,
l896,1898), Van der Menve (1946a, l946b), Jessop (1972), Dyer (1976), Arnold & De
Wet (1993), and Muller-Doblies & Muller-Doblies (1997) recognize Lindley as the sole
author. The description of the genus specifically states "We therefore propose it
[Drimiopsis] as a new genus" (Paxt. Flow. Gard. 2: 73 (185 1-52), Stedje (1994), Speta
(1998a & 1998b), Kativu (2000), Lebatha et al. (2003), Manning & Goldblatt (2003)
and Lebatha & Buys (in press) provide the correct author citation. However Manning et
al. (2004), differing form Manning & Goldblatt (2003), refer to the new combination as
"Ledebouriapetiolata J.C. Manning & Goldblatt, nom. nov., pro Drimiopsis maculata
Lindl."(Manning et al., 2004: 56 1).
The type specimens of Drimiopsis erlangeri Dammer (Figure 12.10, page 212),
currently synonymised under D. botryoides, possess some characters reminding of a
young Drimiopsis maculata, e.g. the pseudopetiole and cordate leaf base. This study
maintains the status quo until more investigations, preferably using fresh material, are
undertaken.
Specimens studied
- 4 0 3 4 : Boran, Merti, North Frontier region of Kenya (-CC), Adamson no. 372 (K).
-4236: Turkana, near El Molo Bay. Kenya (-BB), Lamprey & Cromwell16470 (K).
- 4 8 3 8 T .8, Selous Game Reserve (-CC), VoNensen 3032 (K).
-1039 Tanzania, Mpwapwa District (-CC), Anderson 568 (EA).
-1613 Mozambique. Maputo (-DA), Balsinhas 630 (K, PRE).
-1832 Nelspruit (-A&
ViZjoens.n. (PTECH).
-2428: Naboomspruit Parys (-DB), Lebatha 039 (PUC).
-2430 Pilgrim's rest, Asbestos mines (-AC), van Jaarsveld 9180 (NBG);4 miles north
ofBrmddraai,-Lydenbmggdistrictt(-DD),Codd & De Winter 3261(PRE).
- - - - - - - - - - - - - -
- - - - - - - -
- - - - - - - - - - -
-2528 Pretoria, Wonderboom (-CA), Repton 6981 (PTECH).
-2532
- - - - - - - - - - - - - - - - - -
Maputo, Marracuene para Praia da Costa do Sol (-CC), Nvunga & ~ o A j o387
(NW.
-2628 KwaZulu-Natal, Inanda (-AA), Swart s.n. (J); Inanda (-AA), Buchanan 6/74
(K); Humbert 1741 7 (P).
-2631
Swaziland, Mbabane, Red Tiger Ranch (-AC), Compton 32432 (NBG);
Compton 32432 (PRE), Lebatha 062 (PUC)..
-2632
Inhaca Island, east of Maputo (-BB), Mogg 27433, 27393, ( J ) ; Inhaca Island,
east of Maputo (-BB), Moss s.n sub J 2079 (J), Moss s.n sub J 16777 (J); Maputo (CC), TinZy & Ward 35 (PRE, NH); Maputo (-CD), MoN 4239 (PRE, K). Maputo,
Polana Beach (-DD), Moss & Ottley 11 769 (J).
-2732
KwaZulu-Natal, Sordwana (-AC), Lawn 2020 (NH); Jozini Dam, Lawn s.n.
sub NH 1768 (NH) Moll & Strey 3648 (K); KwaZulu-Natal (-BC), Lubbe 373 (NH);
Ubombo (-DA), Kluge 2533 (NE3G);KwaZulu-Natal (-DC), MacDevette s.n. sub NH
1865 (NH).
-2831 Richards Bay Caravan Park (-CC), Smook 1285 (PRE); Richards Bay Caravan
Park (-CC), Codd s.n. sub PRE 101 72 (PRE).
-2832
St. Lucia-offices, Mtubatuba (-AD), Nicholas 1628 (NH); Urnhlatuzi Lake
Bluff (-CC), Venter 4082 (UCVS).
-2926 Florida (-CC), Foster s.n. (K).
-2929 KwaZulu-Natal, Umkomas (-AC), ArchbeN s.n. sub NBG 2336/32 (NBG).
-2930
KwaZulu-Natal, Bothas Hill (-CC), Hutchins 4687 (K); Krantzkloof (-DD),
Haygarth 22342 (PRE).
-2931
Glen Hill, Mvoti River Bank (-AC), Moll 2289 (PRE); Amatikulu Nature
Reserve (-BA), Ward 2136 (NH); Zimbali Forest Dune (<A), MacDevette s.n. sub
NH 1946 (NH); Ross & Moll s.n. sub NH 21 71 (NH); Stanger, Berea (-CC), Mbonambi
s.n. sub NH 2 (NH); Wood s.n (GRA); Wood. s.n. (NH); Rabinowitz s.n (BOL);Glen
Mill, Lower Tugela ( ), Moll 2289 (PRE); Moll 24586 (GRA); Durban ( ), van Niekerk
s.n. sub NBG 24586 (NBG);van Niekerk s.n sub SAM 73326 (SAM); Berea ( ), Wood
23243 (SAM); Wood 1013 (K); Wood s.n. sub BOL 99552 (BOL) ( ); KwaZulu-Natal,
sedges of bush ( ), Haygarth 273 (NBG); KwaZulu-Natal. Ford 1/79 (K); KwaZulu1 $k (ICY. te b d h a Q31(PUC).
Natal-( J,XiiGn-de~s~
- - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - -
-3029 KwaZulu-Natal (-DD), Abbott 6431 (NH).
-3030
KwaZulu-Natal, Port Shepstone District, The Valleys Farm (-CC), Mogg
13898 (K); Barker 10007 (NBG).
-3129 Port St. John's. Libode (-CA), De ViNiers s. n. sub BOL 15 72/28 (BOL); Port St
Johns, Third Beach, Isinuka (-CB), CZoete 857 (NH); Theron 1574 (K); Port St. John's
(-DA), Moss 8016 (J); Strey 4322 (K); Strey 4322 (PRE).
-3130 (-AA),Abbot 6463 (NH).
-3228 Cebe mouth, Kentani (-CB), Holnzes. s.n. (NBG).
-3228 Gonubie stream bank, East London (-CC), Batten 6-PL82 (NBG).
-3327 Peddie, East London (-BB), Rattray 576 (GRA); Rattray 7881 (PRE); Rattray,
s.n. (BOL).
Without precise locality:
Cultivated pot plants: from Botswana Lebatha 005 (PUC), from Soweto, RSA, Lebatha
007 (PUC); Wood s.n. sub NH 3872 (NH); Johnson s.n. sub NBG 430 (NBG); Edwards
sub Moss s.n. sub J 1345 (J); Mogg s-n (J); Edkins s.n. sub NH 35295 (NH); Codd s.n.
sub PRE; Jodrell Laboratory Jones s.n (K); Beach terminus Limpopo Province, Thode
3395 (NBG); Charles Creek, Barker 10001 7 (NBG).
Collectors uncertain
sub BR 89.3109 (BR); sub BR 899196 (BR).
12.3.6 Drimiopsis davidsoniae
Drimiopsis davidsoniae U & D Muller-Doblies in Fedd. Repert. 108:64 (1997). -
Type: Muller-Doblies & Davidson 77003k, Blyde River Canyon Nature Reserve, rocky
slope, Pilgrim's Rest, Mpumalanga (B!, holotype; BTU!, Z! isotypes).
Ledebouria davidsoniae (U & D Muller-Doblies) Manning & Goldblatt in Edinburgh J.
Bot. 60(3): 560 (2004).
Description: Figure 12.16
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; 0.5-1.8 cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; erect; linear; lanceolate; (10-)12-15(-20)
long; 3-7
cm
cm wide; when torn without threads; sessile. Leaf margin crenate;
noncartilaginous; bordered purplehrown. Lamina thick; unspotted; green. Leaf apex
obtuse. Leaf base cuneate; green. Indumentum present; arranged randomly; in the form
of papillae; frequent; on lamina present; on abaxial leaf surface present; on adaxial leaf
surface present.
Leaf epidermis. Epidermal wax cover particulate. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; more or less as
long as leaves. Rachis conical; 2-7 cm long. Peduncle green. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 15 or less; minute (1-2 rnrn long);
minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous; campanulate;
hypanthium base rounded. Tepals dimorphic; creamy-brownish; hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments free; deltoid to
acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per locule. Stigma
roundish; papillae sessile, trilobal. Style shorter than ovary; terete. Ovary sessile;
globose; transversely smooth; whitishlgreenish; shoulders absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis davidsoniae possesses a localised distribution in the vicinity of Pilgrim's Rest,
Mpumalanga, South Africa (Figure 12.15). It grows on rocky slopes in the shade. Flowering
time September to December.
Diagnostic characters
Drimiopsis davidsoniae is a medium-sized plant possessing sessile, erect, linear to
broadly lanceolate leaves, margin entire and possessing indumentum. The inflorescence
commonly is more or less as long as the leaves. The flowers are creamy-brown.
Specimens studied
-2430: Blyde River Canyon Nature Reserve, Pilgrim's Rest (-DB), Muller-Doblies &
Davidson 77003k (B, BTU, Z ) ; Blyde River Canyon Nature Reserve (-DB), Lebatha
038 (PUC).
Kilometers
Figure 12.15 : Known distribution of Drimiopsis davidsoniae.
Figure 12.16: Drimiopsis davidsoniae (Lebatha 038 (PUC):. 1 , the enlarged mature leaf with
indurnentum; 2, the inflorescence; 3, sessile, erect, linear to lanceolate leaf; 4, Sectioned flower
illustrating globose ovary and deltoid filaments.
12.3.7 Drimiopsis comptonii
Drimiopsis comptonii U. & D. Miiller-Doblies in Fedd. Repert. 108: 64 (1997). Type: Ukutla, slope below farmhouse of Prof. Compton, south facing granitic
outcrops, and peaty pans, Mbabane, Swaziland. -263 1 AC (1982), Muller-Doblies &
Muller-Doblies 820 18g, (B! holotype; BTU! ,Z!, isotypes)
Ledebouria comptonii (U & D Miiller-Doblies) Manning & Goldblatt in Edinburgh J.
Bot. 60(3): 560 (2004).
Descriptions: Figures 12.18 & 12.19
Habit & bulbs. Plants dwarfed (less than lOcm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; stoloniferous; with
tuberescent fimdus absent; whitish; ovoid; 0.5-1.5 cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 2; sometimes 3 or more; erect; spathulate;
oblanceolate; (4-)6-lo(-15)
cm long; 1-3 cm wide; when tom without threads;
sessile. Leaf margin entire; noncartilaginous; edged purplehrown. Lamina thick;
unspotted; green. Leaf apex obtuse. Leaf base attenuate; green. Indumentum present;
arranged in rows; in the form of papillae; dense; on lamina present; on abaxial leaf
surface absent; on adaxial leaf surface present.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled
and irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a sparse pseudo-corymb; erect; considerably
longer than leaves. Rachis ovoid-cylindrical; 1-1.5 cm long. Peduncle green. Bracts
in Mature- inflorescence-vestigial.-ProphyUs absent- Flowers 1-5 or less; minute (1-2
-
-
-
-
-
-
-
-
-
-
-
-
-
mm long); with an elongate pedicel (more than 0.4 cm long); actinomorphic;
sextepalous; campanulate; hypanthium base rounded. Tepals dimorphic; purplehlue;
hypanthium inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect;
epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments
free; deltoid to acurninate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two
per locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous1monosporous; equatorial view depressed ovate; polar
view elliptic
Distribution and habitat preference
Drimiopsis compronii is known from the vicinity of Mbabane, Swaziland (Figure
12.17). It has a extremely localised distribution, restricted to a single rocky outcrop in
Mbabane where it grows on a hill on a rock boulder full of humus in semi shade.
Diagnostic features
Drirniopsis comptonii is a relatively dwarf, stoloniferous, plant possessing two,
spathulate, bright green and oblanceolate leaves with frequent glossy papillae
arranged in rows. The raceme resembles a pseudo-corymb at maturity and bears
striking purple flowers.
Taxonomic note
Drimiopsis cornptonii does not resemble D. atropurpurea as was noted ("remote
affinis") by Miiller-Doblies & Miiller-Doblies (1997).
Kilometers
Figure 12.17: Known distribution of D. comptonii
Figure 12.18: D. comptonii (Lebatha 079 (PUC).1 , simple tunicated, ovoid bulbs with rudimentary
stolons; 2, spathulate leaves; 3, sectioned flower illustrating the pistil and anthers; 4, sectioned
mature flower illustrating deltoid filaments and pistil.
Figure 12.19: Drimiopsis comptonii (Lebatha 079 (PUC). A, plants growing in the botanical
garden of North West University, Potchefstroom Campus displaying their dwarf stature and
spathulate leaves; B, plants from the type locality with stolons; C, pseudo-corymb
inflorescence.
Specimens studied
-2631:
Ukutla, Mbabane, Swaziland, slope below farmhouse of Prof. Compton,
south facing granitic outcrops, and peaty pans (-AC), Miiller-Doblies & MiillerDoblies 82018g (B); Ukutla, Mbabane below the Compton house, south facing
granitic outcrops (-AC), Lebatha 079 (PUC, PRE).
231
-
-
12.3.8Drimiopsis fischeri
Drimiopsis fischeri (Engl.) St~dje in Nordic 1. Bot. 15(6): 593 (1995); Stedje in Fl.
Trop. E. Afr.: 6 (1996); non Drimiopsis fischeri (Engl.) Miiller-Doblies & MiillerDoblies, Fedd. Repert 108: 64 (1997) nomen superflous.
Scillafischeri Engl. in Die Pflanzenw. Ost-Afr. C: 142 (1895); Bak. in Fl. Trop. Afr.
7: 553 (1898). -Type: Tanzania, without precise locality, Fischer 11 (B!, lectotype).
Ledebouria fischeri (Engl.) Manning & Goldblatt in Edinburgh J. Bot. 60(3): 560
(2004).
Drimia fischeri Bak. in Fl. Trop. Afr. 7: 526 (1898). -Type:
Tanzania, without
precise locality, Fischer 1325(B!, lectotype).
Description: Figures 12.21 & 12.22
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous;
with tuberescent fundus absent; whitish; ovoid; 1-1.2 cm across. Bulb scales loosely
packed; when torn without threads; outer scales brown/purple and fleshy.
Leaf morphology. Leaves 3 or more; erect; linear; lanceolate; (6-)8-10(-20) cm
long; 1-2 cm wide; when torn without threads; sessile. Leaf margin entire;
noncartilaginous; edged purple/brown. Lamina membranous; unspotted; green. Leaf
apex acuminate. Leaf base cuneate; green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled
and irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; considerably
longer than leaves. Rachis conical; 1-1.5 cm long. Peduncle green. Bracts in mature
inflorescence absent. Prophylls absent. Flowers 16 to 30; medium-sized (4.1-6 mm);
with an elongate pedicel (more than 0.4 cm long); actinomorphic; sextepalous;
tubular; hypanthium base rounded. Tepals dimorphic; whitish to greenish;
hypanthium conspicuous. Outer whorl of tepals recurved; longitudinally cucullate;
232
--- --
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect;
epitepalous; uniseriate; inserted at throat of perianth tube; longer than pistil. Filaments
free; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two
per locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary stipitate; ovoid to oblong; transversely smooth; whitish/greenish; shoulders
absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
This taxon is based on two separate herbarium specimens, both referring to Tanzania
as the country of collection with no precise locality (Figure 12.20). Flowering time
September to December.
233
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W+E
1000
I
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1000
S
2000
Kilometers
Figure 12.20:Presumed distribution of D.fischeri.
Diagnostic characters
Drimiopsis jischeri possesses brownish bulbs and no indumentmn. The bracts are
absent on mature inflorescence. The leaves are sessile, erect, and lanceolate with no
indumentmn possessing an entire and edged margin. The flower is large (more than 6
mm long), tubular, and shortly pedicellate. The ovary is ovoid to oblong and shortly
stipitate.
Taxonomic note
The two herbarium sheets of Drimiopsisjischeri in Figure 12.22 are seemingly of the
same collection. Figure 12.22A was designated as the type for Scilla jischeri Eng1..
Figure 12.22B is the Drimia jischeri Bak. type. According to Muller-Doblies &
Muller-Doblies (1997), the two specimens are probably duplicates of the same
234
collection. The combination D. fischeri (Engl.) U. & D. Miiller-Doblies (MiillerDoblies & Miiller-Doblies, 1997) is invalid as it is preceded by D. fischeri (Engl.)
Stedje (Stedje & Thulin, 1995).
Ul1-]).
,!:I-~
5mm
~
2mm
Figure 12.21: Flowers of Drimiopsis fischeri. A, long pedicellate flowers (Fischer 11); B,
sectioned flower illustrating long perianth tube and stipitate ovary (Fischer 1325); C, closed
flower (Fischer 1325). Arrows highlight the cucullate inner tepals. (Drawing adapted from
Miiller-Doblies & Miiller-Doblies, 1997: 63).
235
¥~.,
,
\\t.-.
~~',
"
II
.
I;
::..,~t.
j
Figure 12.22: D. fischeri. A, Fischer 11, lectotype of Scilla flScheri Eng1.; B, Fischer 1325, lectotype of
Drimia fischeri Bak.
Specimens studied
Tanzania, without precise locality, Fischer 11 (B); Tanzania, without precise locality,
Fischer 1325 (B).
236
12.3.9 Drimiopsis woodii.
Drimiopsis woodii. Bak. in Fl. Cap. 6: 473-474 (1896); in Fl. PI. Afr., 25: t. Plate 986,
(1946a);Miiller-Doblies & Miiller-Doblies in Fedd. Repert. 108: 63 (1997); excluding
Sutherland s.n. -Type:
KwaZulu-Natal, Pietermaritzburg, Inanda, Wood 656 (K!,
lectotype; BOL!, NH!, SAM isolectotypes).
Ledebouria woodii (Bak.) Manning & Goldblatt in Edinburgh J. Bot. 60(3): 561
(2004).
Description: Figures 12.24 & 12.25
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous;
with tuberescent fundus absent; whitish; roundish; 1.5-2.2 cm across. Bulb scales
loosely packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; erect; linear; ovate; 6-8 cm long; 2-2.5 cm
wide; when tom without threads; pseudopetiolate. Pseudopetiole approximately as
long as lamina; banded. Leaf margin undulate; noncartilaginous; edged purple/brown.
Lamina membranous; unspotted; green. Leaf apex acute. Leaf base attenuate; green.
Indumentum present; arranged randomly; in the form of papillae; frequent; on lamina
present; on pseudopetiole absent; on abaxial leaf surface absent; on adaxial leaf
surface present.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled
and irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; considerably
longer than leaves. Rachis conical; 20-30 cm long. Peduncle banded. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30; small (2.1-4 mm long);
with an elongate pedicel (more than 0.4 cm long); actinomorphic; sextepalous;
campanulate; hypanthium base rounded. Tepals dimorphic; whitish to greenish;
hypanthium inconspicuous. Outer whorl of tepals recurved; longitudinally cucullate;
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect;
237
epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments
free; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two
per locule. Stigma roundish; papillae sessile; trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis woodii is distributed in the eastern regions of South Africa occurring in
Inanda, Klip River, Letaba, northern banks of Merensky Dam, Pilgrim's Rest, Orrie,
Barragwanath Pass, Mtunzini, Empangeni and in the Kruger National Park (Figure
12.23). Its habitat is in shaded northern slope of hills and dunes. Flowering time
September to December
Diagnostic characters
Drimiopsis woodii possesses roundish bulbs, broadly ovate, pseudopetiolate leaves
with undulate margins. The rachis is about 11 to 20 cm long and the peduncle is
banded. The minute flowers are coloured greenish to white.
238
.,-h
-..,
lrJ
v
\
Zimbabwe
.
'"
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V--
( It...
I..r-
r-, .........
...........
-
Sod
,.......,.
A&ka
V
1\
)
...-
900
I
r-- r t---'
Mozambique
I
( SwuiJaJul
"..../
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r
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.
/
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;\
.
V
,/
<
/
r--Botswana
\
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.
--
-
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V
o
I;
.r
/
/"
V
.J
I
N
W-+-E
S
900
1800
Kilometers
Figure 12.23: Known distribution of Drimiopsis woodii.
Taxonomic note
Jessop (1972) considered D. woodii to be a synonym of D. burkei. He realised that it
differed from D. burkei but nonetheless placed it there on the basis of its minute
greenish flowers. He also noted that D. woodii and D macu/ata possess a similar leaf
form. Drimiopsis woodii leaves differ from those of D. macu/ata in shape--the leaves
are linear to ovate, making their base less cordiform compared to that of D. macu/ata.
Both species have entire leaves, but some have entire margins and others have
undulate margins. The leaves are also unspotted. Miiller-Doblies & Miiller-Doblies
(1997) differed with Jessop's (1972) analysis and resuscitated D. woodii.
Baker (1896) cites two specimens when describing D. woodii as new. The first, Wood
656 collected from Inanda, was designated by Miiller-Doblies & Miiller-Doblies
(1997) as lectotype. The second, Suther/and S.n. from Klip River is excluded here
because it conforms to D. queae (See 12.3.18). The Suther/and s.n. collection differs
239
significantly from the D. woodii protologue in having an ovoid bulb, fleshier leaves
with crenulated margins and having leaves tinted abaxially.
Specimens studied
-2030: Zimbabwe, areas of Harare (-BC), Wild 4710 (K, PRE).
-2231: Punda Maria, Kruger National Park (-CA), Lang s.n. (NH).
-2330: Letaba, northern banks of Merensky Dam (-AD), Scheepers 770 (PRE).
-2430: Pilgrim's Rest, Barragwanath Pass (-DB), VanJaarsveld 9158 (NBG).
-2531: Mozambique, Ressano Garcia (-DB), Schlechter 11918 (Z).
-2831:
KwaZulu-Natal, Empangeni (-DB), Edwards 1444/14 (NBG); Empangeni
(-DB), Edwards s.n. (BOL); Mtunzini (-DD), Lawn 1156 (NH); Mtunzini (-DD),
Lang sub PRE TM 32236 (PRE).
-2930:
KwaZulu-Natal, Pietermaritzburg, Inanda (-DB), Wood 656 (BOL, K, NBG,
NH, SAM).
Without precise locality:
Leendertz (pRE 1455, 1455.2).
240
"
-'
,
,
I,
,
e-
r-
.
i.
.
~
i
\
.
..
,
Figure 12.24: Wood 656. Lectotype of Drimiopsis wood;;.
241
;(
i
Figure 12.25: Drimiopsis woodii. 1, the entire plant including the bulb with a tuberesent fondus; 2, the
flower; 3, sectioned perianth illustrating deltoid filaments and dimorphic tepals; 4, globose ovary and
terete style. 5, cross-section through the ovary. 6, portion of the inflorescence rachis showing the vestigial
bracts. (Van der Merwe, I946a).
242
-
----
---
12.3.10 Drimiopsk reilleyana
Drimiopsis reilleyana U . & D. Muller-Doblies in Fedd. Repert. 1O8:64 (1997). -Type:
Mkhaja Nature Reserve, north of Phuzumoja Station, little waterfall in the treesavannah, Mbabane, Swaziland, 263 1DA (198 1), Miiller-Doblies & Miiller-Doblies
820136 (B!, holotype; BTU!, Z! isotypes).
Ledebouria reilleyana (U & D Muller-Doblies) Manning & Goldblatt in Edinburgh J.
Bot. 60(3): 560 (2004).
Description: Figure 12.27
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; (1-)2-4(-5)
cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; erect; linear; lanceolate; (5-)9-lo(-15)
cm
long; 1-4 cm wide; when tom without threads; sessile. Leaf margin undulate;
noncartilaginous; bordered purplehrown. Lamina thick; unspotted; tinted; abaxially
purple. Leaf apex acute. Leaf base cuneate; tinted dark purple. Indumentum present;
arranged randomly; in the form of hairs; frequent; on lamina present; on abaxial leaf
surface absent; on adaxial leaf surface present.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
densely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, dense raceme; erect; more or less as
long as leaves. Rachis conical; 10-30 cm long. Peduncle green. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 rnrn long);
minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous; campanulate;
hypanthium base rounded. Tepals dimorphic; whitish to greenish; hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments free; deltoid to
acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per locule. Stigma
roundish; papillae sessile, trilobal. Style shorter than ovary; terete. Ovary sessile;
globose; transversely smooth; whitishlgreenish; shoulders absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis reillqana distribution is localised in the Mkhaja Nature Reserve in Mbabane,
Swaziland (Figure 12.26), near a little waterfall in the tree-savannah. Drimiopsis reiZZeyana
grows in the dense grasses in the shade under brush. Flowering time September to
December.
Diagnostic characters
Drimiopsis reilleyana is characterized by thick, erect, sessile, and linear to lanceolate
leaves with a thickly banded undulate margin. The stomata are densely spaced. The
flowers are greenish white.
Figure 12.27: Drimiopsis reilleyana (Lebatha 068 (PDC). A and B, leaves are erect; linear to
broadly lanceolate with margins undulate and bordered (x 0.5).
Specimens studied
-2631:
Mkhaja Nature Reserve, Mbabane,
Swaziland
(-DA), Miiller-Doblies
&
Miiller-Doblies 82013b (B, BTU, Z); Mbabane, Mkhaja Nature Reserve, by a mini
waterfall under trees, (-DA), Lebatha 068 (PUC).
245
12.3.1 1 Drimiopsis burkei
Drirniopsis burkei Bak. in Saund. Ref. Bot 3: App 17 (1870), J. Linn. Soc. 13: 228
(1873), F1. Cap. 6: 474 (1896); Van der Merwe in F1. P1. South Afiica 25: t. 975, t. 988
(1946a); Jessop in J. S. Afr. Bot. 38(3): 159 (1972); Muller-Doblies & Muller-Doblies
in Fedd. Repert. 1O8:64 (1997). -Type:
Apies River, Gauteng Province Burke s . n (K!
holotype).
Ledebouria burkei (Bak.) Manning & Goldblatt in Edinburgh J. Bot. 60(3): 560 (2004).
Drimiopsis crenata v.d. Menve in F1. P1. South Afiica 25: t. 975 (1946a). -Type:
Transvaal, Rooiberg, Van der Menve 2805 (PRE! holotype).
Description: Figure 12.29
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; (1-)2-4(-6) cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 2; sometimes 3 or more; spreading; linear; lanceolate;
(5-)6-lo(-20)
cm long; 2-4 cm wide; when tom without threads; sessile. Leaf margin
crenate; noncartilaginous; edged purplehrown. Lamina thick; spotted adaxially; tinted;
abaxially streaked purplehrown. Leaf apex acute. Leaf base cuneate; tinted dark
purple. Indumentum present; arranged randomly; in the form of hairs; sparse; on lamina
present; on abaxial leaf surface absent; on adaxial leaf surface present.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; more or less as
long as leaves. Rachis conical; 1-3 cm long. Peduncle spotted. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 rnm long);
minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous; campanulate;
hypanthium base rounded. Tepals dimorphic; whitish to greenish; hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments valvate;
deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis burkei is widespread in a wide range of habitats in Africa south of the Cunene
and Zambezi rivers. It occurs in South Afiica primarily in the summer rainfall areas
around Gauteng, Limpopo and Mpumalanga Provinces and rarely in the Eastern Cape
and KwaZulu-Natal Provinces. It is widely distributed in eastern Botswana where the
conditions are not arid, near Gaborone Dam, on Khale Hill and along streams in Maruapula
area. In the Kgatleng District it grows in abundance in and around Mochudi village and Rasesa,
and towards Mahalapye in the central district and the Kasama district of northern Zambia. In
addition, it has been collected in the vicinity of Harare, Zimbabwe (Figure 12.28).
Drimiopsis burkei grows on all types of soils and a wide range of habitats among rocks, in
shaded areas and under rock boulders. It is the most widespread of Drimiopsis species with
a plasticity of features that allows it to adapt to extreme conditions. Drimiopsis bulbs in
general, are without papery outer scales but this feature is found in D. burkei growing
i n the sands of the Kgalagadi desert of Botswana. The plants here differ in size; those
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growing in the heavily shaded have big;& l
~
~27 kiii~Img aria
~ 3 4 c~m wide,
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yet maintaining the same leaf shape. Jessop (1972) noted that some, especially those
from damper areas, had longer leaves. Flowering time September to December.
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Kilometers
Figure 12.28: Known distribution of Drimiopsis burki.
Diagnostic characters
Drimiopsis burkei is 2 (sometimes 3 or more), possesses a crenate leaf margin, a lamina
that is adaxially spotted, abaxially streaked purplish/brownish and a dark purple tinted
base.-Theflowers are-ylars&d&ibuted
on the inflorescence and the filaments are
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Taxonomic note
Drimiopsis burkei Bak together with D. burkei Bak. subsp. stolonissima U & D MiillerDoblies occur in two separate clades in the cladistic analysis (Chapter 10). The
differences between the taxa are equal to the differences, among all other Drirniopsis
taxa.
975
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o
11
"
Huthlit},
Figure 12.29:Drimiopsis burkei. I, the entire plant possessing an ovoid bulb and crenate leaves; 2,
flower; 3, sectioned flower bud illustrating dimorphic tepals and a globose ovary; 4, sectioned mature
flower illustrating the deltoid filaments; 5, globose ovary with the style shorter than the ovary; 6, a cross
section of the ovary. (Van der Merwe, 1946a).
250
Specimens studied
-1225: Kasama District, Masaka Village near Mifundu River, Zambia (-AB), Richards
19362 (K); Kalolo Village, 1.6 krn from Kalolo Chambeshi Flats, Zambia (-AB), Richards
19330 (K).
-2030: Harare, Zimbabwe (-BC), Wil4710 (K, PRE).
-2325: Botswana, Lobatse (-BB), Rogers 776/13 (BOL, NBG); Lobatse (-BB),
Rogers 6232 (BOL).
-2328: Waverly, Pretoria (-CB), Venter 13348 (UNIN).
-2329: Lebowa, Sovenga (-AD), Venter s.n. (UNIN).
-2330: Giyani District, Letaba Dam (-AD), Venter 13063 (UNIN); Tzaneen (-CA),
Rogers12530 (BOL); Tzaneen (-DD), Venter 13202 (UNIN).
-2425: Botswana, Mochudi, on hills past the Deborah Memorial hospital, opposite the
Chiefs graveyard (-AC), Lebatha 095 (PUC); Aedume Park, Gaborone (-DC), Hansen
31 78 (PRE); Aedume Park, Gaborone (-DC), Hansen 3462 (GBR); Hansen 3476(K);
Hansen 3467 (PRE); Kgale Hill, southwest facing side before the steep climb, and on
top of the steep climb (-DC), Lebatha 103 (PUC).
-2426: Derdepoort (Mochudi) (-DA), Codd 8855 (GBR, PRE); Rasesa, at the
Matsieng Footprints site (-DA), Lebatha 096 (PUC).
-2427: Leeupoort Tin Mines, Waterberg (-BD), Rogers 22970 (J); Venter s.n. sub
UNIN 13459 (UNIN); Van der Merwe 2805 (PRE); Waterburg (-DC), Lebatha 056
(PUC).
-2428: Naboomspruit (-DA), Galpini 23/l 4 (NBG);Galpini 23/14 (BOL); Parys
Dam (-DB), Lebatha 041 (PUC).
-2430: Ga-Madits, Kromellenboog (-AD), Venter (UNIN); Vaalhoek, Swadini (DB), Raal, P. & Raal, G. 1691 (PRE); Pilgrim's Rest, Bewedere Valley (-DD), Kerfoot
6463 (J, UNIN).
-2527:
Kloofivater Farm, Rustenburg (-CD), Forbes 871 (PRE).
-2528:
Onderstepoort (-CA), Smith 6286 (PRE); Rietvlei (-CD), Lebatha 054 (PUC).
-2530: Lydenburg (-BB), Strey 3662 (K); Carolina District, Stolzburg (-DD),
Blackwill & Blackwill 10091 (J).
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Botswana, Mahalapye (-DD), Rogers 6732 (BOL); ~ & e @ h 1 ISO7X(BUL];
Mansergh 99555 (BOL); Mansergh 115O/27 (NBG); Mansergh 2681/27 (NBG).
I 2627: West Rand, Gauteng, Witpoortjie (-BB), Mogg s.n (J); Roodepoort National
r
IBotanical Garden Reserve (-BB), Perry s.n. (NBG); Randburg, Gauteng (-BB), Reddy,
Reddy & Reddy 406 ( J ) ; Potchefstroom, on a hill in the botanical garden of North West
University, Potchefstroom Campus (-CA), Lebatha 009 (PUC).
-2628: Windsor Park West Rand, Gilliland s.n. (J); Thorntree Kloof, Sandton,
Johannesburg (-AA), Moss 8064 (3); Morningside, Johannesburg (-AA), Moss s.n. (J);
Kapjie, Johannesburg (-AA), Moss s.n. sub J 0601 75 (J). Klipriviersberg (-AC),
Gilliland s.n. (J).
-2831 :KwaZulu-Natal, Eshowe, Campus of the University of Zululand (-CD),
Venter 32 7 (NH); Mtunzini (-DD), Lawn 1717 (NH);Inanda (-DD), Wood 656 (BOL).
-3030: Oribi Gorge Nature Reserve, Port Shepstone, KwaZulu-Natal (-CA),
Vassilatos & Mantel1 731 (4.
Without precise locality
Reddy, R.A., Reddy, K.B. & Reddy, P. s.n. sub J 653 (J); Reddy, R.A., Reddy, K.B. &
Reddy, P. s.n. sub J 842 (J); Reddy, R.A., Reddy, K.B. & Reddy, P. s.n. sub J 1347 (J);
Reddy, R.A., Reddy, K.B. & Reddy, P. s.n. sub J 1388 (J); Reddy, R.A., Reddy, K.B. &
Reddy, P. s.n. sub J 1498 (J); Reddy, R.A., Reddy, K.B. & Reddy, P. s.n. sub J 1846 (J);
Reddy, R.A., Reddy, K.B. & Reddy, P. s.n. sub J 10091 (J); Lang s.n. sub PRE TM3lO88
(PRE); Lang s.n. sub NH 27185 (PRE);Moss 8016 ( J ) ; Strey 3455 (PRE); Balkwill,
MacCullum & Campbell-Young 11084 (9;Williamson 86 (PRE); Kluge 126 (PRE); Balkwill,
Williamson & Smith 8028 (J); Hobson 2204 (PRE); Culvenvell991 (PRE); Smit 1324 (PRE);
Smith 6286 (PRE); Jacobsen 1 791 (PRE); Zambalis 1290 (PRE); Theron 1589 (PRE); L o w
1514 (PRE); Coetzee 6007 (PRE); Turner 1153 (PRE); Moss & Rogers 1345 (NH);
Leendertz TM6628 (PRE); Behr 530 (PRE); Killick & Strey 2705 (PRE); Van der Schoff
4086 (PRE); Du Plessis 928 (PRE); Codd 9767.1 (PRE); Codd 6767.2 (PRE); Codd 8018
(PRE); Codd 8019 (PRE); Codd 31 78 (PRE); Van der Merwe 2 7,253 (PRE); Van der Merwe
2 7,253, 11 77A (PRE); Van der Merwe 11 77A (PRE).
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Collectors uncertain
sub BOL 23/14 (BOL); sub BOL 2681/27 (BOL), sub BOL 776/13 (BOL).
I
12.3.12 Drimiopsis barteri
Drimiopsis barteri Bak. in Saund. Refug. Bot. 3. App. 18 (1870a); in F1. Trop. Afi. 7:
543 (1898); Hepper in F1. W Trop. Afi. 3(1): 104 (1968); Stedje in Nord. J. Bot. 14(1):
49 (1994); Stedje, & Thulin in Nord. J. Bot. 15: 594 (1995). -Type:
Niger, rocky
plains near Nupe, Barter 1512 (K! lectotype, hic designatus); Niger, rocky plains near
Nupe, Barter 3445 (K, syntypes).
Drimiopsis sereti De Wild. in Ann. Mus. Congo, Ser. 5(3): 350 (1906); in Bull. Jard.
Bot. Brux. 3: 268 (191 1). -Type:
Vicinity of Gumbari (10 March 1906), Sereti 543
(BR! holotype, K! isotype).
Drimiopsis aroidashum A. Chev. [var.aroidastrum] in Bull. Soc. Bot. France IV. Mem.
8: 93/94 (1908). -Type:
Saboum village, NdellC, Cameroon (1903), Chevalier 8231
(P! lectotype, hic designatus, K! isotype); Saboum Village, Chevalier 8431 & 8442 (P!,
syntypes).
Drimiopsis aroidastrum var. kabrum A. Chev. in Bull. Soc. Bot. France IV. Mem 8:
93/94 (1908). -Type:
Central Chad, cultivated in Sarh, near Fort Archambault (12
May l9O3), Chevalier 8545 (P! holotype, K! isotype).
Ledebouria barteri (Bak.) Manning & Goldblatt in Edinburgh J. Bot. 60(3): 560
(2004).
Description: Figure 12.31 & 12.32
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; (2-)2.5(-3)
cm across. Bulb scales loosely
packed; when torn without threads; outer scales browdpurple and membranous.
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lanceolate; (20-)24-30(-
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32) cm long; 2-3 cm wide; when torn without threads; sessile. Leaf margin entire;
noncartilaginous; edged purplelbrown. Lamina thick; unspotted; green. Leaf apex
acute. Leaf base cuneate; green. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature straight tabular; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, dense raceme; erect; more or less as
long as leaves. Rachis ovoid-cylindrical; 3-5 cm long. Peduncle spotted. Bracts in
mature inflorescence vestigial. Prophylls absent. Flowers more than 30, small (2.1-4
mm long); minutely pedicellate (shorter than 0.1 cm long); actinomorphic; sextepalous;
campanulate; hypanthium base rounded. Tepals dimorphic; .whitish to greenish;
hypanthium conspicuous. Outer whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect;
epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments
free; deltoid to acuminate. Anthers dorsifixed. Gynoeciurn tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis barteri spans most of Africa from Yendi in Ghana towards Bassar in Togo,
into northern Nigeria in Sokoto to Zaria province, to the Yola area near the Cameroon
I
border. It is distributed along the Chari River in the Chari-Banguirmi region of Chad
into the uppermost Central African Republic in the St. Floris National Park.
Distribution continues towards the eastern parts of Africa into Sudan, around Zalingei,
that is equidistant to Sudan's borderlines with Chad and Central African Republic, into
the Gambela region of Ethiopia spanning into Jelib, Somalia. From the Central African
Republic towards &e southern parts of Africa, distribution spans into Katanga regions
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of the Democratic Republic of Congo, then firstly south into the Kasama District of
northern Zambia, then east into Tanzania. Drimiopsis barteri has been well collected
mainly in Tanzania. Here D. barteri has more vigour and is abundant in the Iringa,
Kyimbila, Mpanda, Morogoro and Rungwa districts in Rungwa Game Reserve and
Mlala Hills. A collection form KwaZulu-Natal, South Africa, Comins, D.M. 470
(221911952) housed in PRE, is here determined to be D. barteri. This extends the
distribution to southern Africa (Figure 12.30). Flowering time March to May.
Drimiopsis barteri grows in grasslands and wet marshes; in bushland or woodland on
dark clay soils, in damp areas, near rivers and streams or on hard and on dry ground
footpaths.
Kilometers
Figure 12.30: Known distribution of D. barteri.
Diagnostic characters
Drimiopsis barteri possesses brownish/purplish membranous outer bulb scales. The
leaves are very narrowly lanceolate and up to 40 cm long, the lamina and lamina base
are uniformly coloured green. The flowers are small (2.1-4 mm long), greenish and the
hypanthium conspicuous.
Taxonomic note
Stedje (1994) proposes Barter 3449 as the lectotype for D. barteri when in the
protologue Barter 1512 and Barter 3445 are cited as syntypes. In accordance with
article 9.10 of the ICBN (Greuter et al. 2000), I therefore propose Barter 1512 as
lectotype partly because Barter 1512 conforms well with the protologue and because
Barter 3445 could not be found.
Chevalier's description of D. aroidastrum fits D. barteri equally well. Drimiopsis
aroidastrum var. kabarum A. Chev. was apparently a cultivated plant. It differs from D.
aroidastrum only in leaf size. As is common in Drimiopsis, the differences in leaf size
of D. aroidastrum var. aroidastrum (19-20 cm long, 15-20 cm wide) and D.
aroidastrum var. kabarum (10-12 x 20-22 cm) could be the result of size variation
generated by cultivation.
Similarly, D. sereti is similar to D. barteri but for the smaller flowers and shorter
inflorescence (De Wilderman 1906, 1911). However, Wilderman (1906) states that the
flowers are not fully developed (suggesting that both the inflorescence and flowers will
get larger with maturity) and that his species resembles D. barteri Bak. var. parvzjlora.
The aforementioned is an unpublished name and not considered further.
Drimiopsis aroidastrum, D. aroidastrum var. kabarum and D. sereti are synonymised
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880 747
I)
rl
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-Figure 12.31: Drimiopsis barteri. Holotype of D. sereti, Sereti 543 (BR).
257
Ai
,
~I
I
f. .
C8.JIUCII'S18 au'I'Iatl
o.t.. .. - dj.
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..
I
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CMfYAlIfR 'U'HSO~lOU""GUI[TDUCHUI
'i.
UKlOk1D'ULIPUS
~14r
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FLORA
".~I'".......
OF
It...
'i"" ~
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f
r
Figure 12.32: A, the herbarium sheet of D. aroidaslntm lectotypeA.
Chevalier 8131 (1903) determined by Stedje. to be D. barteri. B,
herbarium sheet of D. aroidastntm, showing the size variations in the
taxon. The linear leaves in B are opened while folded in A and Figure
12.30.
258
Specimens studied
-0412: Saboum village, Ndelle, Cameroon (-CC), Chevalier 8231 (K, P); Chevalier
8431 (P); Chevalier 8442 (P); Chevalier 8544 (P); Chevalier 8772 (P).
-4434: Mbosi Zambi (Mbosita Mbuga), Tanganyika Territory (-DC), Davies D. 778
(K).
-0605:
Niger Territory, Nupe, Plains of Ilorin. Nigeria (-AC), Barter 3448; Barter
3449 (K); Niger, rocky plains near Nupe (-AC), Barter 1512 (K).
-0631:
Mpanda District, Kapapa Camp Track (-CB), Richards 11648 (K); Mlala
Hills, Mpanda District (-CC), Barter 11564 (K).
-4636: Tanganyika Territory, Mupwapwa (-CC), Hornby 566 (K).
-4637: Morogoro District, Morogoro-Dodoma road (-DC), Harris 2142 (0).
- 4 7 0 1 :About 4 miles east of Yendi, Ghana (-AC), Morton GC9094 (K).
-0711: Kan Gimi, Zaria Province, Anara Forest Reserve (-AC), Keay FHI 22924 (K);
Sarduana, along Gashaka River to Serti (-AD), Medler 290 (K);
Zaria Province, Igabi District (-AD), Photo FHI 25 770 (K).
-0734: Rungwa Game Reserve, Tanzania (-BA), Richards 20755 (K).
-4735: T.7, Msembi-Mbagi Track, 8.3 miles, Iringa District (-BC), Greenway &
Kanuri I4804 (K); T7, Ruaha National Park, Iringa District, 3.5 km north of Msembe
(-BC), Bj~rnstad1158 (K, 0);
Lebatha 002 (PUC) cult. Bj~rnstad1158 (K, 0).
-4804: Sokoto Province, Northern Nigeria (-CB), Dalziel541 (K).
-0810: Yola Bodewa, Nigeria (-BC), Dalziel236 (K); Yola collection 236 (K).
-4821: Central Afiican Republic, Manovo-Gounda, St. Floris National Park (-AA),
Fay 2 728 (K).
-0834: Ethiopia, Gambela, Illubabore Province (-AA), Ash 3502 (K).
-0900: Bassar, Togo (-BB), Hakki, Leuenberger & Shiers 369 (B, K).
-0905:
Chambeshe flats, 1.6 km on a path from Kalolo village (-CC), Richards 1933
-4908: Sanga River Forest Reserve, Zaria Province, Jemaa Division, north of Dogon
Kurmi sawmill Northern Nigeria (-CD), Jones 108 (K).
-4918: Chad, cultivated in Sarh, near Fort Archambault (-CB) Chevalier 8545 (K).
-0933:
Kyimbila District, North of Lake Malawi (-AB), Stolz 1786 (K); Tanzania,
Rungwe district, Chivanje (-BA), Leedal5203 (K).
-1030:
Zambia. Northern Kasama District, Masuka village (-CB), Richards 19362
(K).
-1106: Mada Hills, Northern Nigeria (-AC), Hepburn 79 (K).
-1129:
Katanga, Democratic Republic of Congo (-AA), Lebrun 7566 (K).
-0229:
Gombari (Gumbari) Democratic Republic of Congo (-CB), Sereti 543 (BR,
K).
-02931 :KwaZulu-Natal, South Africa (-AC), Comins 470 (PRE).
Without precise or unknown locality
Barouma, Cameroon: Favios 2032 (P); Jacques-Felix s.n. (P); Jacques-Felix s.n. sub P
1921 5.70 (P); Chevalier 1.3167 (P); Chevalier 13422 (P); Belgian Congo: Humbert
8785 (P); Bequaert 5495 (BR); Bequaert 5495 (BR). Nigeria: Parsons ,5114 (K); T.7,
Triki Nboga Track, 9.7 mile from Msembi Greenway, & Kanuri 14782 (K); 1 km north
west of Jelib, Somalia Thulin & Warfa (K); Jabal Marra, 30 miles south of Zalingei,
Sudan, Wickens, G. E. 1960 (K).
Collections uncertain
sub BR 893092 (BR); sub BR 893106 (BR); sub BR 893107 (BR); sub BR 899188
(BR); sub BR 899204 (BR); sub BR 899206 (BR); sub BR 899210 (BR); sub BR
899211 (BR); sub BR 899212 (BR).
12.3.13 Drimiopsis rosea
Drirniopsis rosea A. Chev. in Bull. Soc. Bot. France IV. Mem. 8: 93 (1908). -Type:
in
Saboun village in the countries of Ndoukas and Koutiin on dry 'laterite' in Cameroon,
Chevalier 8432 (P! holotype, K! isotype) hic restituta.
Description: Figure 12.34
Habit & bulbs. Plants dwarfed (less than lOcm high); hysteranthous; annual;
bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with tuberescent fundus
present; whitish; ovoid; 1-1.5 cm across. Bulb scales loosely packed; when tom
without threads; outer scales white and fleshy.
Leaf morphology. Leaves 1; sometimes 2; spreading; linear; lanceolate; (2-)3-5(-
8) cm long; 0.5-2 cm wide; when tom without threads; sessile. Leaf margin undulate;
noncartilaginous; edged purplehown. Lamina membranous; unspotted; tinted;
abaxially streaked purplehrown. Leaf apex acuminate. Leaf base cuneate; tinted dark
purple. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature straight tabular; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, dense raceme; erect; considerably
longer than leaves. Rachis cylindrical; 10-30 cm long. Peduncle coloured purplish.
Bracts in mature inflorescence vestigial. Prophylls absent. Flowers than 30; minute (1-
2 rnrn long); minutely pedicellate (shorter than 0.1 cm); actinomorphic; sextepalous;
campanulate; hypanthiurn base rounded. Tepals dimorphic; whitish to greenish;
hypanthium inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate;
apically conduplicate. Inner whorl of tepals connivent; longitudinally cucullate;
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apically conduplicate. Vitta conspicuous. Stamens 6; g r e e n ; ~t ~o whitish; - a c t ;
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epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments
free; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitiswgreenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
The plant displays disjunct distribution, known from one herbarium specimen in
Cameroon, and South Africa in Kaapsche Hoop, Pretoria and Mpumalanga (Figure
12.33). Flowering time September to December.
Diagnostic characters
Drimiopsis rosea is dwarf; some of the small (0.5-2.5 cm diameter) bulbs possess a
tuberescent fundus. The mainly single leaf (sometimes 2 leaves do occur) is, tiny (1-5 cm
long) with an undulate margin and possesses a, membranous lamina with an acuminate apex.
The inflorescence is considerably longer than the leaves, with the rachis 11-20 cm long and a
peduncle usually coloured uniformly purple.
Taxonomic note
The South Afiican material was previously misidentified as D. burkei, or unidentified
Drimiopsis species. The plants are dwarf with lamina as small as 1 cm long; the whitish
to violet flowers are sometimes described as pink to mauve. This taxon has eluded
mention in publications since its description in 1908. Fresh material should validate this
decision to reinstate the species.
Specimens studied
- 4 6 1 1 Chari oriental, LaBoum, in the village of Saboun, Cameroon (-CA), Chevalier
8432 (BR; K, P (x2).
-2530
Mpumalanga, Kaapsche Hoop (- CB), Venter s.n (NH).
-2529
Middelburg,(- CB), Codd s.n. sub PRE.
- - . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
- - - - - - - - - - - -
-2528 Baviaanspoort, Pretoria (-CB), Goossens 43.Z (PRE);
- - - - - - - - - - - - - - - - - - - - - - - -
-----.....-.-
,
f
CHEVALIER.
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Figure 12.34: Chevalier8432 (P). Lecotype of D. rosea
264
12.3.14 Drimiopsis pusilla
Drimiopsis pusiNa U. & D. Muller-Doblies in Fedd. Repert. 108: 64 (1997). -Type:
Red Hill road, 4 km from MbabaneKJsutu road to Lundsi, north facing granitic ridges,
shady ledge, Swaziland, Miiller-Doblies & Davidson 82020b (B!, holotype; BTU!, Z!
isotypes).
Ledebouria pusifla (U & D Muller-Doblies) Manning & Goldblatt in Edinburgh J. Bot.
60(3): 560 (2004).
Description: Figures 12.36 & 12.37
Habit & bulbs. Plants dwarfed (less than lOcm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; 2.5 cm or less across, up to 0.2 cm. Bulb
scales loosely packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; erect; linear; lanceolate; (2-)3-5(-8)
0.5-2
cm long;
cm wide; when tom without threads; sessile. Leaf margin crenulate;
noncartilaginous; bordered purple/brown. Lamina thick; unspotted; tinted; abaxially
purple. Leaf apex acute. Leaf base cuneate; tinted dark purple. Indumentum present;
arranged randomly; in the form of hairs; frequent; on lamina present; on abaxial leaf
surface present; on adaxial leaf surface present.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
densely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
Inflorescence. One to two per bulb; a simple, dense raceme; erect; more or less as
long as leaves. Rachis conical; 2-6 cm long. Peduncle coloured purplish. Bracts in
mature inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 mrn
long); shortly pedicellate (0.1-4 cm long); actinomorphic; sextepalous; campanulate;
hypanthium
base
rounded.
Tepals
dimorphic; purplish
green;
hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments fiee; deltoid to
acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per locule. Stigma
roundish; papillae sessile, trilobal. Style shorter than ovary; terete. Ovary sessile;
globose; transversely smooth; whitish/greenish; shoulders absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis pusilla known only from the vicinity of Mbabane, Swaziland along a hill
near a new development (Figure 12.35). It grows in open areas under rock boulders,
north facing granitic ridges in shady ledges. Flowering time September to December.
Diagnostic characters
Drimiopsis pusilla is a dwarf plant with tiny leaves (1-5 cm long) and 0.5-2 cm wide.
The leaves are linear to broadly lanceolate, sessile, abaxially brownish purplish and
possess a crenulate and bordered leaf margin. Crenulated leaf margin is an
autapomorphy to D. pusilla. The lamina possesses an indumentum, densely spaced
stomata, and a dark purple tinted cuneate base. The flowers are purplish greenish tinged
white.
Specimens studied
-2631
Red Hill road, Mbabane, Swaziland (-BD), MiiNer-Doblies & Davidson
82020b ( B , BTU, 2);Lebatha 078 (PUC).
Kilometers
Figure 12.35 : Known distribution of Drimiopsis pusilla.
Figure 12.36: Drimiopsis pusilla (Lebatha 078, PUC). 1 , an ovoid bulb; 2, the broadly lanceolate
sessile leaves with hairs; 3, the inflorescence with sparsely distributed flowers; 4, the dimorphic
tepals with the globose ovary and deltoid filaments. (1,2 & 3 = lifesize; 4 =. x 25)
Figure 12.37: Figure 12.37: D. pusilla (Lebatha078). A, the plant with a thick lamina, tinted
abaxially; B, broadly lanceolate and erect leaves.
269
~--
12.3.15 Drimiopsis atropurpurea
Drimiopsis atropurpurea N.E. Brown in Kew Bull. 1921: 299 (192 1); in F1. PI. South
Africa, Plate 976 (1946); Jessop in J. S. Afi. Bot. 38(3): 160 (1972). -Type:
Roses
Creek, near Barberton (1EO), Thorncrofr 1083 (K! ,holotype; BOL! ,PRE!, isotypes).
Ledebouria atropurpurea W.E. Br.) Manning & Goldblatt in Edinburgh J. Bot. 60(3):
560 (2004).
Drimiopsis purpurea v.d Merwe in F1. P1. Afi. 25: t. 975 (1946a). -Type:
Paulpietersburg near Pivaan, Van der Merwe 2781 (PRE!, holotype).
Description: Figure 12.39
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus absent; whitish; ovoid; (0.S) 1-2(-2.5)
cm across. Bulb scales
loosely packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; erect; cordiform; ovate; (8-)9-13(-15)
cm
long; 5-7 cm wide; when tom without threads; pseudopetiolate. Pseudopetiole
exceedingly longer than lamina; tinted. Leaf margin entire; noncartilaginous; bordered
purplelbrown. Lamina thick; spotted adaxially; tinted; abaxially purple. Leaf apex
acute. Leaf base cordate; tinted dark purple. Indumentum present; arranged randomly;
in the form of hairs; dense; on lamina present; on pseudopetiole present; on abaxial leaf
surface present; on adaxial leaf surface present.
Leaf epidermis
. Epidermal wax cover thin. Stomata anomocytic; distributed sparsely; crypts
shallow; subsidiary cells form an H-complex. Adaxial epidermal cells shortly
polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae irregular.
-
-
-
-
Idorescence; -iie
to two per bulb; a simple, sparse-raceme;erwt; m o g -or less as
-
-
-
-
-
long as leaves. Rachis conical; 2-7 cm long. Peduncle banded. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 15 or less; small (2.1-4 rnm long);
shortly pedicellate (0.1-4 cm long); actinomorphic; sextepalous; campanulate;
hypanthium
base
rounded.
Tepals dimorphic; purplish
green;
hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments fiee; deltoid to
acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per locule. Stigma
roundish; papillae sessile, trilobal. Style shorter than ovary; terete. Ovary sessile;
globose; transversely smooth; whitisWgreenish; shoulders absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis atropurpurea has localised distribution in South Africa in the Limpopo
Province, in Vryheid, Vryheid Nature Reserve, Paulpietersburg, Pivaan and Luneberg,
in KwaZulu-Natal, in Mpumalanga in Rose's Creek, Barberton in the Waterval areas. In
Mozambique it has been collected in Maputo and on the Inhaca Island (Figure 12.38).
Drimiopsis atropurpurea grows in mountainous areas, mostly south facing densely
grassy or areas with bush clumps. Flowering time September to December.
Kilometers
Figure 12.38: Known distribution of Drimiopsis atropurpurea.
Drimiopsis atropurpurea possesses ovate leaves that are more than 4.1 cm wide and a
raceme of 15 flowers or less. Drimiopsis atropurpurea looks like D. maculata but
differs in the erect leaves that are pubescent, the lamina adaxially tinted purple; cordate
base tinted dark purple and the margin entire. The flowers are purple. Drimiopsis
maculata on the other hand possesses leaves that are pubescent, spreading, abaxially
uniformly coloured green with a pseudopetiole much longer than the lamina, and the
flowers are greenish.
Specimens studied
-2328:
Bochum District near Blouberg Mountain summit (-BB), Balkwill, K.,
Balkwill, M.J., Barloe-Kearsley & Gessel6028 ( J ) .
-2530:
Belfast district, Schoemans Kloof (-CA), Young A297 (PRE); Lydenburg,
Barberton (-CA), Mason (K); Rogers 18508 ( K , PRE); (-CC), Thorncrofr 1083 (BOL,
K , PRE). Waterval Boven (-DB), Rogers 30241 b (PRE).
-2730:
Vryheid Nature Reserve, Paulpietersburg (-BD), Schrire 1330 (NH); Rogers
30241 (K, Z ) ; Van der Merwe 2781 (PRE). Luneburg area, Farm Verkocht, northern
KZN (-CD), Turner 412 (GWIT); Lebatha 049 (PUC) sub GWIT 1999, SLT412
(GW1T);Near Luneburg (-CD), Van der Menve 2779, (PRE).
-2929 Nyininye, Underberg (-BA), Du Toit 251 9 (NH).
-3029
Maputo, Mozambique (-DA),
Mozambique (-CB), Mogg 2 7433 (K).
Precise locality unknown
Moss 1081 7 (PRE).
MacDevette 1303 (NH); Inhaca Island,
976
2
G. Parker
del.
Hu.th lith.
Figure 12.39: Drimiopsis atropurpurea. I & 2, plant i11ustratingthe bulb, leaves and inflorescence; 3, the
minute to smal1 single flower; 4, a portion of the dimorphic perianth revealing the uniseriate filaments
typical of al1Drimiopsis species; 5, the globose ovary; 6, a cross section of the ovary (I & 2 ...,lifesize; 3 =
x 40; 5 = x 60) (Van der Merwe, 1946a).
274
--
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--
- ---
12.3.16 Drimiopsis kikiae
Drimiopsis kikiae Lebatha sp. nov. -Type:
Louwsburg, Itala Nature Reserve under
trees and hidden in grasses, Lebatha 045 (PUC, holotype; PRE isotype).
In facie similes Drimiopsis queae sed differt folium quasi magnus, crispatus, subtus
purpureo coloris et supra guttatus.
Description: Figures 12.41 & 12.42
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent hndus present; whitish; roundish; 1-3 cm across. Bulb scales loosely
packed; when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 3 or more; sometimes 2; erect; cordiform; lanceolate; (4)8-12(-18)
cm long; 2 4 cm wide; when tom without threads; pseudopetiolate.
Pseudopetiole exceedingly shorter than lamina; tinted. Leaf margin crenate;
noncartilaginous; bordered purple/brown. Lamina thick; spotted adaxially; tinted;
abaxially purple. Leaf apex acuminate. Leaf base attenuate; tinted dark purple. Leaves
glabrous.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
sparsely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; more or less as
long as leaves. Rachis conical; 8-10 cm long. Peduncle spotted. Bracts in mature
inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 rnrn long);
shortly pedicellate ( 0 . 1 4 cm long); actinomorphic; sextepalous; campanulate;
hypanthium base rounded.
Tepals dimorphic; creamy-brownish; hypanthium
inconspicuous. Outer whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
-
-
-
-
-
-
-
-
-
-
-
-
- - - - - - - - - - - - - -
-
-
-
-
-
-
-
-
-
-
-
-
-
conduplicate. Vitta conspicuous. Stamens 6;-greenisli ID whitish; -erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments valvate;
deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate. Ovules two per
locule. Stigma roundish; papillae sessile, trilobal. Style shorter than ovary; terete.
Ovary sessile; globose; transversely smooth; whitish/greenish; shoulders absent.
Nectaries present.
Pollen. Pollen isomorphouslmonosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis kikiae is known only from the Itala Nature Reserve, Louwsburg, South
Africa (Figure 12.40), in moist shaded areas under trees and rocks. Flowering time
September to December.
Diagnostic characters
Drimiopsis kikiae possesses roundish bulbs, a pseudopetiole much shorter that the
lamina, a spotted peduncle, and valvate filaments. It is a striking plant possessing a
crisp-like lamina with crenulate purply banded margins. The leaves are a rich deep
green, spotted adaxially, and abaxially deep purple. The purple pseudopetiole is shorter
than the lamina. The flowers are white.
Taxonomic note
Drimiopsis kikiae resembles D. queae only in possessing a pseudopetiole and having
crenulated leaf margins. There are thirteen differences between the two plants, notable
amongst which are the D. kikiae larger leaves that are crisp-like with undulating
margins, abaxially purple tinted and adaxially spotted.
Specimens studied
-2731 : Louwsburg,Itala Nature Reserve (-CA), Lebatha 045 (PUC, PRE).
5 kilometers fiom VaalWater (-CA), Charles Craib sub Lebatha 046 (PUC).
Kilometers
Figure 12.40: Known distribution of D.kikiae.
Figure 12.41: D. kikiae (Lebatha 046 (PUC).1, the roundish bulb with a tuberescent fundus; 2,
erect; cordiforrn, broadly lanceolate leaves with a crenate and undulating leaf margin; 3,
sectioned flower illustrating dimorphic tepals, globose ovary, and deltoid filaments; 4, a
developing (left) and a mature inflorescence (right) with sparse, shortly pedicellate flowers
on a short rachis (1 & 2 -life size; 3 = x 60).
Figure 12.42: Drimiopsis kikiae (Lebatha 046 (POC). A, Sparse raceme and leaves
spotted adaxially, with crenate and undulating margins; B young plant (A = x 1.5; B =
life size
279
-
-
-
- -
--
- -
12.3.17 Drim iopsis Iin iopapilla
Drimiopsis liniopapilla Lebatha sp. nov. -Type:
Roosenekal, South Africa, (25" 14'
207" S 29" 57' 403" E), Lebatha 053 (PUC, holotype; PRE, isotype).
Drimiopsis atropurpurea remote affinis, set abbreviatus, lamina basi quasi liniaris,
indumentum lineolatus et flores viridi-albus.
Description: Figures 12.44 & 12.45
Habit & bulbs. Plants medium-sized (10.1 to 15 cm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fundus present; whitish; ovoid; 0.5-2.5 cm across. Bulb scales loosely
packed; when torn without threads; outer scales white and fleshy.
Leaf morphology. Leaves 1; sometimes 2; erect; cordiform; lanceolate; 3-8 cm
long; 1-4 cm wide; when torn without threads; pseudopetiolate. Pseudopetiole
approximately as long as lamina; tinted. Leaf margin crenate; noncartilaginous;
bordered purple/brown. Lamina thick; unspotted; tinted; abaxially streaked
purple/brown. Leaf apex acuminate. Leaf base attenuate; tinted dark purple.
Indumentum present; arranged in rows; in the form of hairs; dense; on lamina present;
on pseudopetiole present; on abaxial leaf surface present; on adaxial leaf surface absent.
Leaf epidermis. Epidermal wax cover thin. Stomata anomocytic; distributed
densely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae regular.
Inflorescence. One to two per bulb; a simple, dense raceme; erect; more or less as
long as leaves. Rachis ovoid-cylindrical; 20-40 cm long. Peduncle coloured purplish.
Bracts in mature inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (12 rnm long); minutely pedicellate (shorter than 0.1 cm long); actinomorphic;
sextepalous; campanulate; hypanthium base rounded. Tepals dimorphic; whitish to
greenish; hypanthium inconspicuous. Outer whorl of tepals connivent; longitudinally
cucullate; apically conduplicate. Inner whorl of tepals connivent; longitudinally
cucullate; apically conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish;
erect; epitepalous; uniseriate; inserted at throat of perianth tube; as long as pistil.
Filaments free; deltoid to acuminate. Anthers dorsifixed. Gynoecium tricarpellate.
Ovules two per locule. Stigma roundish; papillae sessile, trilobal. Style as long as
ovary; terete. Ovary sessile; globose; transversely smooth; whitisWgreenish; shoulders
absent. Nectaries present.
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
Drimiopsis liniopapilla is currently only known from the vicinity of Roosenekal, South
Africa (Figure 12.43). Grows on hillsides, on bushy and wooded slopes and under
rocks. Flowering time September to December.
Diagnostic characters
Drimiopsis Iiniopapilla is characterised by an abaxially purple/brown streaked lamina.
The minutely pedicellate (shorter than 0.1 cm long) flowers are sparsely distributed on
the rachis; tepals coloured whitisldgreen and the style as long as the ovary. The hairs on
the leaves are arranged in parallel rows and with one or two inflorescences per bulb.
Taxonomic note
Drimiopsis liniopapilla resembles D. atropurpurea in having visible hairs and a
pseudopetiole. It is smaller, the lamina linear but narrower, and finely crenulated. The
hairs are arranged in parallel lines and the flowers whitish green, those of D.
ahopurpurea are purple.
Specimens studied
-2529 Roosenekal, South Afiica (-BA), Lebatha 053 (PUC, PRE).
Figure 12.44: Drimiopsis liniopapilla. 1, an ovoid bulb; 2, the inflorescence with minutely pedicellate
flowers on a short rachis; 3 & 4, the pseudopetiolate, linear leaves with an attenuate base and
indumentum lined on the lamina; 5, sectioned flower illustrating the pistil; 6, sectioned flower
illustrating the deltoid filaments..
Figure 12.45: Drimiopsis liniopapilla (Lebatha 053 (PUC). A, a plant, in its natural habitat,
with two inflorescences per bulb as well as leaves abaxially streaked purplish/brownish; B, a
plant with young leaves possessing a pronounced parallelodromous venation; C, developing
plant, cultivated in the garden, with linear leaves..
284
12.3.18 Drimiopsis queae
Drimiopsis queae Lebatha sp. nov. -Type:
Lebatha 055 (PRE holotype, PUC isotype).
Minutes, proxime affinis D. pusilla sed differt folium pseudopetiolus et glabris, lamina
basi cordata margine crenatus.
Description: Figures 12.47-12.49
Habit & bulbs. Plants dwarfed (less than lOcm high); protantherous to
synantherous; annual; bulbaceous. Bulbs hypogeal; gregarious; non-stoloniferous; with
tuberescent fimdus absent; whitish; ovoid; 1-2 cm across. Bulb scales loosely packed;
when tom without threads; outer scales white and fleshy.
Leaf morphology. Leaves 1; sometimes 2; erect; cordiform; lanceolate; 2 4 cm
long; 0.5- 1 cm wide; when tom without threads; pseudopetiolate. Pseudopetiole
approximately as long as lamina; tinted. Leaf margin crenate; noncartilaginous;
bordered purple/brown. Lamina thick; unspotted; tinted; abaxially purple. Leaf apex
acuminate. Leaf base attenuate; tinted dark purple. Leaves glabrous.
Leaf epidermis. Epidermal wax cover thick. Stomata anomocytic; distributed
densely; crypts shallow; subsidiary cells form an H-complex. Adaxial epidermal cells
shortly polygonal; abaxially elongately tetragonal; anticlinal boundaries channelled and
irregular-sinuate; periclinal wall curvature tabular-convex; cuticle striae smooth.
Inflorescence. One to two per bulb; a simple, sparse raceme; erect; considerably
longer than leaves. Rachis cylindrical; 1-2 cm long. Peduncle coloured purplish. Bracts
in mature inflorescence vestigial. Prophylls absent. Flowers 16 to 30; minute (1-2 rnm
long); shortly pedicellate (0.1-4 cm long); actinomorphic; sextepalous; campanulate;
hypanthiurn base rounded. Tepals dimorphic; creamy-brownish; hypanthium
inconspicuous. Outer whorl of tepals recurved; longitudinally cucullate; apically
conduplicate. Inner whorl of tepals connivent; longitudinally cucullate; apically
conduplicate. Vitta conspicuous. Stamens 6; greenish to whitish; erect; epitepalous;
uniseriate; inserted at throat of perianth tube; as long as pistil. Filaments free; deltoid to
acuminate. Anthers-db;rsifixe&.-Gpoeeium tricqellate. O v u l e t~
- - - - - - - - - - - - - - - - - -
w o 1 ~ ~~Stigma~
roundish; papillae sessile, trilobal. Style shorter than ovary; terete. Ovary sessile;
globose; transversely smooth; whitishlgreenish; shoulders absent. Nectaries present.
~
1
Pollen. Pollen isomorphous/monosporous; equatorial view depressed ovate; polar
view elliptic; laterally blunted; subequiaxe; distal pole straight; sexine smooth;
ornamentation punctate.
Distribution and habitat preference
This miniature plant is distributed in the Gauteng, North West and Mpumalanga and
KwaZulu-Natal regions of South Africa: Waterberg District, Tierkloof, Pretoria,
Wonderboom, Modderfontein and Potchefstroom (Figure 12.46). It grows under rocks
in shaded areas. Flowering time September to December.
Figure 12.46: Known distribution o f Drimiopsis queue.
Diagnostic characteristics
Drimiopsis queae is dwarfed, possessing leaves not longer than 5 cm, 2 cm wide or less
and a thin epidermal wax cover. The inflorescence is considerably longer than the
leaves and the outer whorl of tepals recurved. Drimiopsis queae differs fiom D. pusilla
in possessing a thinner lamina, linear pseudopetiolate leaves with crenulate undulating
margins and no indumenta. The lamina colouring also differs; D. queae is a darker
purple abaxially. The short-lived flowers are creamish purple.
Taxonomic note
Sutherland s.n. (K) (Figure 12.49), one of the specimens cited with the original
description of D. woodii, belongs here under D. queae. It differs considerably from D.
woodii in possessing an ovoid bulb, fleshier leaves tinted abaxially and with margins
crenulate.
Specimens studied
-2427:
Leeupoort, Waterberg (-DC), Rogers 22970 (J), Waterberg (-DC), van der
Merwe 2805 (PRE).
-2528:
Pretoria, Wonderboom Reserve (-CA), Repton 1881 (PTECH).
-2529:
Tierkloof (-AA), Lebatha 055 (PUC, PRE).
-2530:
8 km south of Nelspruit (-BD), Bayliss & Leach 718 (NBG).
-2627:
Vredefort (-CD), Du Preez 481 (BLFU); Wonderboom, Mooi River,
Potchefstroom (-CA), Rogers 21 717 (J).
-2628:
Gauteng, Modderfontein (-AA) Rogers 21409 (J).
-2930:
Klip River, Inanda, KwaZulu-Natal (-DB), Sutherland s.n. (K).
Figure 12.47: Drimiopsis queae (Lebatha 055 (PUC). 1, ovoid bulb; 2, cordiform to lanceolate
young leaves without spots and with margins slightly crenulate; 3, rachis with sparse flowers; 4
dissected flower illustrating stamens and pistil (life size except 4 =enlarged x 40).
Figure 12.48: D. queae: (Lebatha 055 (PDC). A, a plant with pseudopetiolate, erect, lanceolate
leaves (life size); B, a plant with shortly pedicellate flowers (life size); C, mature plant (all life
size).
289
---
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..;<
..............
Figure 12.49: Herbarium sheet of Sutherland s.n. illustrating ovoid bulbs, leaves tinted abaxially
and with crenulated margins.
290
12.4 EXCLUDED NAMES
12.4.1 Published names
Drimiopsis avasmontana Dinter ex Solch. Beltr. 1960. Flowers of South West Afi-ica.
Dissertation, University of Munich. [publication unavailable for verification]
Drimiopsis engleri Krause in. Bot. Jahrb. li: 443 (1914). [now synonymised under
Ledebouria rautanenii (Schinz) S . Venter (1993)l.
Drimiopsis humifusa Bak. in F1. Cap. 6: 474 [now synonymised under Resnova
humifisa (Bak.) U . & D. Muller-Doblies in Fedd. Repert. 108: 64 (1997)l.
Drimiopsis lachenalioides (Bak.) Jessop in J. S. Afi. Bot. 38(3): 157 (1972). [now
synonymised under Resnova lachenalioides (Bak.) v. d. Menve in U. & D.
Miiller-Doblies Fedd. Repert. 108: 64 (199711.
Drimiopsis maxima Bak. in F1. Cap. 6: 474. [now synonymised under Resnova
humifusa (Bak.) U . & D. Muller-Doblies in Fedd. Repert. 108: 64 (1997)l.
Drimiopsis papillosa Dint. in Fedd. Repert. 17: 189 (192 1). [now synonymised under
Ledebouria rautanenii (Schinz) S . Venter (1993)l.
Drimiopsis saundersiae Bak. in F1. Cap. 6: 474 [now synonymised under Resnova
humifua (Bak.) U. & D. Muller-Doblies in Fedd. Repert. 108: 64 (1997)l.
12.4.2 Nomino dubia
D. cordata is an invalid herbarium name used by Peters (B).
D. gracilis is an invalid herbarium name used by Moss (PRE).
D. linearis, is an invalid herbarium name used by Peters (B).
D. maculatum Lindl., is an invalid herbarium name used by Rabinouritz (BOL).
D. rudatisii, is an invalid herbarium name used by Schl. (BR).
D. undulata, is an invalid herbarium name used by Peters (B).
13. GENERAL CONCLUSIONS
13.1 SYSTEMATIC REVISION
Drimiopsis Lindl. & Paxt. is endemic to Africa and 18 species are recognised in this
thesis. This study attempts for the first time a comprehensive systematic analysis of
monographic proportions. Species delimitation was based on terminal entities resultant
from a cladistic analysis. Any systematic study is dependent on data and data may be in
various forms. Previous systematic treatments of Drimiopsis taxa were almost
exclusively based on macro-morphological characters. The present study, although
confining itself to the traditional macro- and micro-morphological characters,
investigated for the first time leaf anatomy, stigma and pollen morphology and
chemical composition. A total of 105 characters were coded for DELTA. The large
number of characters held promise of resolving relationships within the genus.
However, this study has shown that synapomorphies are largely absent from the
terminal entities that correspond to species, which instead possess a preponderance of
homoplasious characters. Two species are resurrected, four subspecies are elevated to
the rank of species creating two new combinations and four new southern African
species are described.
This study affirms that Drimiopsis is poorly collected. In general, plants can be said to
have a vulnerable conservation status-species
are at high risk of extinction in the wild
in the medium term. Drimiopsis in general makes poor herbarium specimens with
especially flower characters difficult to discern.
Drimiopsis species are generally small geophytes with thick, usually erect, spotted
leaves. The abaxial leaf surface is usually purple to brownish tinted or streaked. The
- - - - - -
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- - - - - - - - - - -
- - -
short-lived inflorescence be% minutely t o shoitiy pediceHate flowers that posses-~
-
tepals that are dimorphic, connivent and cucullate with a conduplicate apex margin.
The greenish filaments are deltoid to acuminate. The globose, sessile ovary possesses a
terete style, topped by a spheroid stigma with subsessile trilobal stigmatic papilla.
Pollen grains are depressed ovate (equatorial view) with blunt lateral sides, a
subequiaxe equatorial diameter and a straight distal pole.
'
Drimiopsis grows in a wide range of shaded habitats in grasslands, wet marshes,
bushland or woodland. It often grows in mountainous areas, among rocks. It prefers
damp areas where there is plenty of leaf litter, on densely grassy slopes, areas with bush
clumps, along riverine bushveld and coastal forest fringes. It grows in a wide variety of
soils types favouring dark clayish or sandy soils, but also grows on brown sandy clay,
dolomitic rock, granitic ridges or hard, dry ground along footpaths at elevations up to
1600 m above sea level.
This study has revealed that the overall shape (but not dimensions!) of Drimiopsis
leaves is not influenced by the environment and is therefore a character of some
taxonomic significance. The leaf epidermis in Drimiopsis, unobscured by wax or
indumentum, displays taxonomically significant epidermal cell arrangement,
morphology and stomata1 characters. Anatomical characters, although confirming the
morphological differentiation of epidermal cells, are unreliable. The hypothesis that
adaxial and abaxial cells differ according to leaf orientation is disproved in this
investigation, implying that character in Drimiopsis is more genetically than
environmentally influenced. Phenetic analyses of the leaf characters suggest a reranking of subspecies and clearly demarcate three genera, and groups Resnova with
Ledebouria to the exclusion of Drimiopsis.
Interspecific flower characters and states are continuous and thus not taxonomically
significant. Tepal colour groupings can assist in delimiting groups of species.
Anatomical sections of flowers in this study for the first time demonstrate Resnova
ovaries also to be stipitate. An investigation of pollen characters also revealed novel
states, but here too, intergeneric rather than interpsecific variation was observed. The
phenetic analysis of intergeneric flower and pollen characters demarcates Resnova,
Ledebouria and Drimiopsis. Here too, Resnova clusters with Ledebouria.
-Thg phytochemical analysis isolated six novel structure compounds in extracts of D.
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- - - - - -
- - - - -
- - - - - - -
- - - - - - - - - - - - - - - - - - - - -
- - - - -
burkei Bak. Compounds 1, 2, 3, 4 and 6 are homoisoflavonoidsand~Comp0~~d~5,-ascillascillin. Compound 1 was identified as a possible Ledebouriinae U. & D. MiillerDoblies marker compound and compounds 1, 3 and 4 as taxa indicators. TLC and
HPLC can, with caution, be used to profile plant extracts.
The Drimiopsis taxa investigated revealed a basic chromosome count of x = 10 and x =
11. The former, predominant in southern Afiican taxa, appears to be the plesiomorphic
state. All tropical Afiican plants and a few southern African ones are x = 11. Higher
chromosome numbers occur in tropical African plants. Southern African plants are
diploids and tetraploids.
All attempts to sequence the trnL-F gene failed. Investigations are ongoing, the results
of which will be combined with morphological data in a total evidence approach.
Phenetic analysis results revealed unambiguous intergeneric differences in the
Ledebouriinae, which raises questions about views in support of sinking Resnova under
Drimiopsis (Phillips, 1951; Jessop, 1970, 1972; Dyer, 1976; Arnold & De Wet, 1993;
Meyer and Williams, 1997); and sinking both Resnova and Drimiopsis under
Ledebouria (Manning et al., 2004). Intergeneric variations within the Ledebouriinae
can be interpreted in one of two ways: 1. For the splitter, Resnova, Ledebouria and
Drimiopsis as separate genera, Resnova having more in common with Ledebouria than
with Drimiopsis. If lumping is preferred, then Resnova and Drimiopsis should be
grouped in one taxon. I view the morphological differences between Drimiopsis,
Resnova and Ledebouria to warrant generic status.
Although the strict consensus tree based on an analysis of morphological characters
was fully resolved within Drimiopsis, almost no synapomorphies support species,
implying that the taxonomic arrangement of species as presented in this thesis is
tenuous. The strict consensus tree hypothesises the monophyletic status of all three
genera in the ingroup where each genus in question is supported by numerous
synapomorphies. The total evidence analysis of combined morphological and published
hnL-F DNA data based on a reduced taxon sample produced a fully resolved tree also
supporting the monophyletic status of all three genera. These results differ from the
conclusions of Manning et al. (2004) who lump the three genera in question. In
addition, the characters professed by them to support their lumping, prove to be either
homoplasious, symplesiomorphies or confined to the terminal clade in Ledebouria.
Sections or subgenera within Drimiopsis are not recognised due to the general lack
synapomorphies and predominance of homoplasious characters.
An analysis of the geographic distribution of the Ledebouriinae, the paucity of plant
collections and uneven and inadequate collections notwithstanding, show Ledebouria to
be the basal taxon occurring out of Afiica. Drimiopsis and Ledebouria centres of
diversity are Mpumalanga and KwaZulu-Natal respectively. The Resnova centre of
diversity appears to be Mpumalanga. The basal species in Drirniopsis have a
predominantly tropical African distribution. Thirteen Drimiopsis species are endemic to
southern African and five to tropical Africa.
Until the reliability of all conventional characters of the plant genome have been
evaluated and analysed, morphology and molecular data should always be prerequisites
of a systematic study. It is time systematists end dogmatic idealism and see systematics
for what it is, biodiversity governed by genes and nature-no
single factor explains all
aspects of biodiversity.
"Scientists still do not appear to understand sufficiently that all earth sciences must
contribute evidence towards unveiling the state of our planet.. . It is only by combining
the information furnished by all the earth sciences that we can hope to determine 'truth'
here, that is to say, to find the picture that sets out all the known facts in the best
arrangement and that therefore has the highest degree of probability. Further, we have
to be prepared always for the possibility that each new discovery, no matter what
science furnishes it, may modify the conclusions we draw" -Alfred
Wegener (1966).
Ongoing molecular studies of cpDNA to be expanded to include mtDNA and
nuclear DNA.
The maintenance and expansion of the DELTA list of characters and character
states generated in this study.
The pursuing of unidentified metabolites found during the chemical analysis.
The basic chromosome number x = 5 for Resnova is intriguing and needs
further investigation, and hybridization experiments are proposed.
Determination of the pollinators.
The significance of colouration or maculation relative to soil, availability of
sunlight, etc.
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319
Table 2: Rf values from chromatogram with Solvent System B (Figure 2).
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321
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2
APPENDIX 2
METHODS: DNA EXTRAXTION PROTOCOL
PREPARATION OF SOLUTIONS
a. 100 mMol(0.01 Mol) Tris (Hydroxymethyl l), or arninoethane GR buffer
= C4H11N03,
Molecular weight = 121.14 dm01
Calculate the amount of Tris for a 250 rnl buffer solution:
Molecular weight x Moles x volume to be made(1iters)
121.14 g/M x 0.01 Mol x 250/1000 liters
= 3.029 g Tris
b. 0.02 Mol EDTA (Ethylenediaminetetra - acetic acid or Diaminoethanetetraacetic
Acid) or simply, Disodium salt = [CH2.N(CH2.COOH).CH2.COONaI2.2H20
MW = 372.25 dm01
0.02 mol x 326.27 dm01 x 25011000 1
= 1.631 g EDTA
c. 1.4 mol NaCl (sodium chloride) MW = 58.44 glmol
1.4 mol x 58.44 dm01 x 250/1000 1
= 20.454 g NaCl
d. Preparation of 250 ml DNA isolation (extraction) buffer
Mix a, b, and c in a beaker with 150 ml high purity water. Stir, a stirrer bar inside, on
an electrical stirrer until clear. Adjust pH, using a pH meter, to 9.00 with concentrated
acetic acid and NaOH. Adjust volume to 250 ml as while regulating the pH. Pour
mixture into a volumetric flask to verify volume. Add to a schott bottle and autoclave
then refrigerate.
e. Preparation of 2 % CTAB (Hexadecyl trimethyl ammonium bromide) stock
solution - Molecular weight = 354.45: Weigh 4 g CTAB to lOml high purity water.
Make up a 50 ml solution in a volumetric flask. Add to schott bottle and autoclave.
f. Preparation of 100 ml TAE-buffer: Mix 10 mM Tris (-01x121 . 1 4100/1000)
~
=
0.121 14 g and 1 mm EDTA (.002 mol x 326.27 g/mol x 100/1000). Adjust to pH 7,
then autoclave.
g. Gel preparation: Materials needed: 50ml TAE buffer 1:1; 5g Agarose powder and
15micro 1 Ethediumbromide. Cook mixture in the microwave until powder has
dissolved making sure not to overboil to avoid evaporation. Add the ethidium bromide
as the mixture cools. Cool gel in the refrigerator for about three minutes then place
into a tray, insert comb, then refrigerate.
h. Chemicals also needed are 5 ml of 0.2 % Mercaptoethanol, Chloroform, 100% and
70 % ethanol washing solution.
DNA EXTRACTION
Mark 1.5 ml Eppendorf tubes carefully with sample name or number for
identification.
Mix 1000 pl CTAB + buffer with 2 p1 mercaptoethanol in an Eppendorf tube per
sample. CTAB-buffer protects the DNA from enzymes and secondary
metabolites. CTAB also lyses the ceN membranes andforms complexes with
nucleic acids.
Place approx. 100 mg of fresh leaf and 100 mg of PVP in a mortar (PVP binds
to phenols).
Grind material in liquid nitrogen. When tissue is a fine powder transfer to the
Eppendorf tube with the CTAB-mix. This is the mechanical breaking up of ceN
walls.
Incubate sample tubes at 65°C in a water bath for 60 min. Mix a few times
during incubation by gently shaking the tubes. This step lyses the cell releasing
the CTAB-DNA complex into the aqueous solution.
Leave in the refrigerator overnight.
Incubate sample tubes at 65' C for 30 min. Mix a few times during incubation
by gently shaking the tubes.
Centrifuge at 12000 rpm for 15 rnin. Decant supernatant.
Transfer to fume hood. Add 250-500 p1 wet chloroform to the supernatant. Mix
carefully by turning tubes. Spin for ca. 5 rnin. at 10000- 13000 rpm. This step
removes proteins and carbohydrates.
10. Mark a new set of Ependorf tubes. Transfer the aqueous supernatant fiom each
sample to a new tube carefully, taking care NOT to pipet any of the debris
from the middle layer.
11. In the fume hood, add another 250-500 p1 wet chloroform, mix, and spin as
above. This time the middle phase separating the chloroform and the aqueous
phase should look thin and filmy.
12. Mark a new set of Ependorf tubes. Transfer the aqueous supernatant from each
sample to a new tube carefully, taking care NOT to pipet any of the debris
from the middle layer.
13. Repeat steps 9-1 0 once or twice if there is a gel-like suspension forming -a
top
layer
14. Add 1/10 of supernatant volume of 5M NaCl.
15. Add 1.5-2 times the sample volume of ice cold Et-OH (absolute or 95%). Freeze
the samples at -20'
C for overnight to help precipitate the DNA. Divide the
sample into two flnecessary, the ethanoltsample ratio is important. Freezing can
be for one to several hours ifnecessary.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
16. Spin for 10 min. at 10000-1 3000 rpm. Decant the ethanol. Add ca 1000 ~ 1 7 0 %
ethanol per sample. Decant and add 100% Et-OH. Shake the tubes gently to
dislodge pellets from tube walls, and leave for 30 min-1 hour. The ethanol
removes the last remnants of the CTAB.
17. Spin for 10 min. at 10000-13000 rpm. Decant the ethanol carefblly-don't
loose the DNA-pellet. Let the ethanol evaporate by leaving the tubes on their
sides with their caps off (indesicator). When the pellet is dry (ca. 1 hour or so),
add 200 p1 water and resuspend. Store at 4-43' C (for immediate use) or at -20' C
(for long term storage).
DNA QUANTIFICATION
Mix 4 p1 standard buffer with 5 pl loading buffer
Load 10 p1 DNA combined with 5 p loading buffer
Run the gel for whole genome @ 60 volt constant voltage for 45 minutes
DNA AMPLIFICATION-PCR
Taberlet et al. (1991) universal primers used:
T a b c l = 5'TTT CAA A(CT)T CAG AGA AAC CCT GG 3'
Tabfl
= 5'TAA
CTT GGG TTT ATG TCA ATT 3'
Material for PCR was prepared in 50 p1 Ependorf tubes, the Super-mix in a 100 pl
tube. 1.25 p1 of the each speceis plant DNA in super pure water, referred to here as
the template, was mixed with various magnesium concentrations (Table 1). The total
reaction volume is 2 5 ~ 1 .
Table 1: All measurements are in p1. Sp = tube ID, H 2 0 = ultra pure water, Mg =
magnesium concentrations, Buff = super Taq buffer, P 1, P2 = primers c to f = hnL to
tmF, dNTP's are the deoxyribonucleosidetriphosphate, Enzy = Taq DNA polymerase
and Temp = template of the plant species DNA to be amplified.
Enzy
Temp
Total
volume
0.05
1.25
25.0
Label one 100 p1 tube for the supermix that will contain the constant
ingredients, Mg, Buff, P 1, P2, dNTP, Enzy and template (enzyme added last
as it has to stay frozen).
Mark 4 tubes A, B, C & D, place water and magnesium in the tubes then add
5.5 5 supermix:
Super mix total volume = 22.2 p1
Thus 4 equal portion of this is 22.214 = 5 . 5 ~ 1
Place tubes in PCR thermal cycler using the following running conditions:
95' C for 3 min. x 1 cycle
1
95" C for 30 sec.
50" C for 30 sec. For 35 cycles
J
72" C for 1 min.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
72" C for 10 min. x 1 cycle
4.
- - -
Make up agarose gel and run products on 90 volts for 30 minutes.
Mix 4 p1 standard buffer with 5pl loading buffer
To every 10 p1 DNA add 5p loading buffer
SEQUENCING AND ALIGNING
Sequencing was done on a MegaBACE 500, Molecular Dynamics part of Amersham
Pharmacia Biotech (AmershamBiosciences) Automated Capillary DNA Sequencing
System with MegaBACE 500 Sequence Analyser v2.4 software (University of Cape
Town, Dept of Molecular and Cellular Biology, DNA Sequencing Laboratory).
Alignments were done with BioEdit software.
Taxonomic treatment: Index to Drimiopsis names
Drimjopsis names accepted in this thesis
Drimiopsis aroidastrum A.Chevalier
Drimiopsis aroidastrum A. Chevalier var. kabrum .
Drimiopsis atropurpurea N . E. Brown
Drimiopsis avansmontana Dinter ex Solch.
Drimiopsis barteri Baker
Drimiopsis botryoides Baker.
Dr imiopsis botryoides Baker subsp . botryoides
Drimiopsis botryoides subsp. prostrata B.Stedje
Drimiopsis burkei Baker.
Drimiopsis burkei subsp. burkei
Drimiopsis burkei subsp. stolonissima U.Miiller-Doblies & D.Muller-Doblies
Drimiopsis bussei Damm.
Dr imiopsis comptonii U.Muller-Doblies & D.Miiller-Doblies
Drimiopsis crenata van der Merwe
Drimiopsis davidsoniae U.Muller-Doblies & D .Muller-Doblies
Drimiopsis engleri Krause
Drimiopsis erlangeri Damm.
Drimiopsisfischer i (Engl.) B.Stedje
Drim iopsisfischer i (Engl .) U .Miiller-Doblies & D .Miiller-Doblies
Drimiopsis holstii Engl.
Drimiopsis humifusa Baker
Dr imiopsis kir kii Baker
Drimiopsis lachenalioides (Bak.) Jessop
Drimiopsis maculata Lindl. & Paxt.
Drimiopsis maxima Baker
Drimiopsis minor Baker
Drimiopsis papillosa Dinter
Drimiopsis perfoliata Baker
Drimiopsis purpurea van der Merwe
Drimiopsis pusiZZa U.Muller-Doblies & D.Muller-Doblies
Drimiopsis queue Lebatha
Drimiopsis reilleyana U.Miiller-Doblies & D.Miiller-Doblies
Drimiopsis rosea A.Chevalier
Drim iopsis saundersiae Baker
Drimiopsis sereti De Wild.
Drimiopsis stoZonissima U .Miiller-Doblies & D.Miiller-Doblies
Drimiopsis stuhlmanni Baker
Drimiopsis volkensii Baker
Drimiopsis woodii Baker