Systematic Studies of the Genus Lebeckia and Related Genera of the Tribe Crotalarieae (Fabaceae)

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Systematic studies of the genus Lebeckia and related genera of the tribe Crotalarieae (Fabaceae)

James Stephen Boatwright

Thesis submitted in fulfilment of the requirements for the degree

PHILOSOPHIAE DOCTOR

BOTANY

in the

FACULTY OF SCIENCE

at the

UNIVERSITY OF JOHANNESBURG

SUPERVISOR: PROF. BEN-ERIK VAN WYK CO-SUPERVISOR: PROF. MICHAEL WINK

APRIL 2009 AFFIDAVIT: MASTER'S AND DOCTORAL STUDENTS TO WHOM IT MAY CONCERN

This serves to confirm that I James Stephen Boatwright Full Name(s) and Surname

ID Number 8204225029081

Student number 920103313 enrolled for the

Qualification PhD (Botany)

Faculty _Science Herewith declare that my academic work is in line with the Plagiarism Policy of the University of Johannesburg which I am familiar with.

I further declare that the work presented in the Systematic studies of the genus Lebeckia and related genera of the tribe Crotalarieae (Fabaceae) (thesis) is authentic and original unless clearly indicated otherwise and in such instances full reference to the source is acknowledged and I do not pretend to receive any credit for such acknowledged quotations, and that there is no copyright infringement in my work. I declare that no unethical research practices were used or material gained through dishonesty. I understand that plagiarism is a serious offence and that should I contravene the Plagiarism Policy notwithstanding signing this affidavit, I may be found guilty of a serious criminal offence (perjury) that would amongst other consequences compel the UJ to inform all other tertiary institutions of the offence and to issue a corresponding certificate of reprehensible academic conduct to whomever request such a certificate from the institution.

Signed at _Johannesburg on this _day of 2009

Signature Print name

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j; "Success is not the key to happiness. Happiness is the key to success. If you love what you are doing, you will be successful."

Albert Schweitzer TABLE OF CONTENTS

SUMMARY . i

ACKNOWLEDGEMENTS V

CHAPTER 1: General introduction and objectives of the study 1 1.1 General introduction 1 1.2 Objectives of the study 3

CHAPTER 2: Material and methods 5 2.1 Morphology 5 2.2 Anatomy 7 2.3 DNA sequencing and phylogenetic analyses 8

CHAPTER 3: Phylogenetic relationships of the tribe Crotalarieae (Fabaceae) inferred from DNA sequences and morphology 15 3.1 Introduction 15 3.2 Results 17 3.3 Discussion 29

CHAPTER 4: Alkaloids and flavonoids of the tribe Crotalarieae 39 4.1 Introduction 39 4.2 Alkaloids of the tribe Crotalarieae 39 4.3 Flavonoids of the tribe Crotalarieae 42 4.4 Other useful compounds 43 4.5 The use of Cape legumes as sources of tea 44

CHAPTER 5: Evidence for the polyphyly of Lebeckia 52 5.1 Introduction 52 5.2 Morphological and anatomical evidence for the paraphyly of Lebeckia 55 5.3 Molecular evidence for the paraphyly of Lebeckia 63 5.4 Taxonomic implications 64 CHAPTER 6: Taxonomy of the genus Calobota 68 6.1 Introduction 68 6.2 Discussion of characters 69 6.3 Taxonomic treatment 86

CHAPTER 7: Taxonomy of the genus Wiborgiella 187 7.1 Introduction 187 7.2 Discussion of characters 189 7.3 Taxonomic treatment 199

CHAPTER 8: The generic concept of Lotononis, reinstatement of Euchlora, Leobordea and Listia and a new genus of the tribe Crotalarieae (Fabaceae) 241 8.1 Introduction 241 8.2 Results 243 8.3 Discussion 247 8.4 Taxonomic treatment 270

CHAPTER 9: Taxonomy of the genus Robynsiophyton 297 9.1 Introduction 297 9.2 Discussion of characters 298 9.3 Taxonomic treatment 300

CHAPTER 10: Taxonomy of the genus Rothia 306 10.1 Introduction 306 10.2 Discussion of characters 308 10.3 Taxonomic treatment 313

CHAPTER 11: General conclusions 328

LITERATURE CITED 343

APPENDICES Appendix A: Voucher specimen information, morphological characters and characters used for parsimony reconstructions 365 Appendix Al 365 Appendix A2 368 Appendix A3 371 Appendix A4 383 Appendix A5 385 Appendix B: Publications .388 SUMMARY

The tribe Crotalarieae (Fabaceae) is a large and diverse group of papilionoid

legumes that largely occur in Africa. A systematic study of generic relationships within

the tribe was undertaken using nucleotide sequences from the internal transcribed

spacer (ITS) of nuclear ribosomal DNA, the plastid gene rbcL and morphological data.

The Crotalarieae are strongly supported to be monophyletic and sister to the tribe

Genisteae. Lebeckia, Lotononis and Wiborgia are all polyphyletic in the molecular

analyses and morphological data support the division of Lebeckia into three more

natural genera (one of which includes the monotypic North African Spartidium). Four

major lineages were identified within the tribe based on sequence data: the "Cape

group", comprising Aspalathus, Lebeckia, Rafnia, Spartidium and Wiborgia; the

Lotononis group, comprising Lotononis pro parte, Pearsonia, Robynsiophyton and

Rothia; a group comprising Lotononis section Leobordea, L. section Listia, and allies;

and the Crotalaria group, comprising Bolusia, Crotalaria and Lotononis hirsuta

(Lotononis section Euchlora). Morphological analysis yielded a similar topology, except

that Lotononis is monophyletic if L. hirsuta is excluded. When the molecular and

morphological data sets are combined, the same major clades are retrieved as in the

molecular analysis, with the notable exception that Lotononis and Lebeckia senso

stricto are supported as monophyletic.

The study showed the need to reinstate the genus Calobota and to describe a new genus (Wiborgiefia). Evidence from ITS and rbcL data clearly showed that the genus Lebeckia is not monophyletic. These analyses, along with morphological and

i SUMMARY

anatomical data, showed that the genus should be divided into three genera: (1)

Lebeckia senso stricto (L. section Lebeckia); (2) Calobota [L. section Calobota and L. section Stiza] including the monotypic, North African genus Spartidium; (3) a new genus, Wiborgiella [L. section Viborgioides, together with L. inflata, L. mucronata and

Wiborgia humilis]. Taxonomic revisions of the genera Calobota and Wiborgiella are presented. Sixteen species are recognized in Calobota, one of which is described as

new, and nine species in Wiborgiella, two of which are described as new. The anatomy of the leaves, stems and fruit walls of the two genera revealed important characters to distinguish these genera from each other and from other genera of the tribe

Crotalarieae. Detailed taxonomic studies of these genera are presented, including keys to the species, descriptions, known geographical distributions, phylogenetic relationships and illustrations.

Lotononis is a large and taxonomically complex genus of the Crotalarieae, with the majority of its species occurring within South Africa. In this part of the study, sequence and morphological data of the rare Lotononis macrocarpa was added to existing data matrices and these were re-analyzed. Molecular systematic data (nrITS and rbcL) indicated that Lotononis is polyphyletic, with Lotononis section Euchlora as sister to Bolusia and Crotalaria and with L. macrocarpa close to the "Cape group" of the

Crotalarieae. As a result, the genus Euchlora is here reinstated and the new genus

Ezoloba is described herein to accommodate the anomalous L. macrocarpa. Ezoloba is distinct in its 5+5 anther configuration, exceptionally large fruit, paired stipules and presence of bracteoles. Within the remaining species of Lotononis, both combined and separate analyses of the morphological and sequence data sets revealed three strongly

ii SUMMARY

supported clades corresponding largely to clades recovered in previous cladistic

analyses based on morphological, chemical and cytological data. These are here

recognised at the generic level as Lotononis s.s., Leobordea and Listia. Characters

supporting these generic re-alignments are discussed and a key to the 16 genera now

recognized in the tribe Crotalarieae is presented.

Pearsonia and Rothia and sister to the latter based on morphological and DNA sequence data. A revision of the genus is presented, including illustrations of vegetative and reproductive features and distributional information.

Rothia is a genus of papilionoid legumes that consists of two species, R. indica and R. hirsuta. The genus is a member of the tribe Crotalarieae and is widely distributed in Africa, Asia and Australia. Molecular systematic studies have shown the genus to be closely related to Robynsiophyton and Pearsonia, but it is easily distinguished from these by its 10 small, rounded anthers and sub-equally lobed calyx

(as opposed to the large, elongate anthers of Pearsonia or the nine stamens of

Robynsiophyton). These three genera share characters such as uniform anthers, straight styles and the presence of angelate esters of lupanine-type alkaloids. The leaf and fruit anatomy of Rothia and Robynsiophyton were also studied but revealed no informative differences. Both genera have dorsiventral leaves (i.e. clearly differentiated into palisade parenchyma adaxially and spongy parenchyma abaxially) with mucilage cells in the epidermis and thin-walled fruit. A taxonomic revision of Rothia is presented,

iii SUMMARY including a key to the species, correct nomenclature, descriptions, illustrations and distribution maps.

The patterns of evolution of 18 morphological, chemical and cytological characters within the Crotalarieae were investigated by reconstructing these onto the minimal length trees from the combined molecular and combined molecular/morphological analyses. This highlighted the taxonomic complexity of the crotalarioid genera, the extreme overlap of characters between genera, convergent patterns in some characters and the value of informative combinations of characters to circumscribe genera in the tribe in the absence of diagnostically useful generic apomorphies.

iv ACKNOWLEDGEMENTS

I would like to express my gratitude to the following persons or institutions:

My promotor, Prof. Ben-Erik van Wyk, for mentorship and guidance

throughout this project. His enthusiasm and motivation is always inspiring and

it was a great experience to work with him. I would also like to thank him for

all the travel opportunities and the opportunity to attend various conferences.

Prof. Michael Wink, my co-promotor, for his advice and major contribution

towards the phylogenetic study.

Prof. Patricia M. Tilney for her patient assistance, guidance and advice with

the anatomical studies.

Dr Michelle van der Bank, Mr Olivier Maurin and Ms Cynthia Motsi from the

Molecular Systematics Laboratory at the University of Johannesburg for the

use of their facilities and the sequencing of samples.

The National Research Foundation (NRF) and the University of

Johannesburg for funding. The NRF and University Research Council are

also thanked for the travel grants to attend the Botany 2008 conference in

Vancouver, Canada.

Curators and staff of the listed herbaria are thanked for their kind hospitality

during study visits (BM, BOL, GRA, J, K, NBG, P, PRE, S and UPS) or for

making specimens available on loan (BOL, GRA, K, MEL, NBG, P, PRE, S

and WIND).

Kew Herbarium and Ms Amida Johns for plant samples and the Jodrell

Laboratory for extraction of the Kew material. ACKNOWLEDGEMENTS

Dr Hugh Glen (National Biodiversity Institute, Durban) is thanked for translating the Latin diagnoses.

Cape Nature for the collecting permit for the Western Cape Province.

The staff of the Mary Gunn Library at the National Herbarium, Pretoria, for their kind assistance during visits.

The Curator of the University of Johannesburg Herbarium, Dr Annah N.

Moteetee, for the management of loans from various herbaria.

I am indebted to Dr Dee Snijman (NBG) for bringing the only flowering material of Wiborgiella dahlgrenii to my attention.

Dr John Manning (NBG) for suggesting the name Wiborgiella for the new genus.

Drs Sarah K. Gess and Friedrich W. Gess for supplying reprints of their publications on pollinators in the Crotalarieae.

The Department of Botany and Plant Biotechnology at the University of

Johannesburg for the use of their facilities.

My fellow students in the Botany Department for their company and support.

Ms Marianne le Roux for her friendship and contribution towards the phylogenetic study.

Mr Anthony R. Magee for his encouragement, inspiration, companionship during field expeditions and support throughout my studies.

My family, especially my mother Suzette, for their love, support and motivation during my studies and always.

The Magee and Ackermann families for their support and encouragement.

vi CHAPTER 1: GENERAL INTRODUCTION AND OBJECTIVES OF THE STUDY

1.1 GENERAL INTRODUCTION

Fabaceae or Leguminosae, the third largest flowering plant family, comprises ca.

727 genera and 19,235 species (Lewis et al. 2005). Its members have a cosmopolitan distribution and form an important component of various vegetation types across the world (Lewis et al. 2005). Lewis et al. (2005) presents a useful synthesis of the literature on the family and also the currently recognized genera. Molecular systematic studies over many years have added to our understanding of relationships within the family and especially within the Papilionoideae (e.g. Kass and Wink 1995, 1996, 1997;

Crisp et al. 2000; Doyle et al. 2000; Wojciechowski et al. 2000; Kajita et al. 2001;

Pennington et al. 2001; Wink 2003; Wink and Mohamed 2003; Wojciechowski et al.

2004).

In Africa the Fabaceae are represented by several tribes. The Papilionoideae are especially well-represented, in particular the genistoid legumes (sensu Polhill 1981; redefined by Crisp et al. 2000), the majority of which are contained in the tribes

Crotalarieae (Benth.) Hutch., Genisteae (Bronn) Dumort. and Podalyrieae Benth. The

Genisteae are widespread with species in Africa, Europe, the Americas, the

Mediterranean and Macaronesia (Polhill and Van Wyk 2005). The Podalyrieae (with the exception of most species of Calpurnia and Podalyria velutina Burch. ex Benth.) are endemic to the Cape Floristic Region of South Africa (CFR; Schutte and Van Wyk

1 CHAPTER 1: GENERAL INTRODUCTION AND OBJECTIVES

1998) and the Crotalarieae are sub-endemic to Africa, with species of Crotalaria,

Lotononis, and Rothia extending to other continents (Van Wyk 1991a).

The tribe Crotalarieae is a large and diverse group of 11 genera and ca. 1,204 species (Van Wyk 2005). Polhill (1976, 1981) in his treatment of the tribe included 16 genera. However, in the latest treatment of the group by Van Wyk and Schutte (1995) the genera of the Argyrolobium group (Dichilus, Melolobium and Polhillia) were transferred to Genisteae and only 11 genera included in the Crotalarieae. To date a comprehensive molecular systematic study of the Crotalarieae has not been completed. Previous molecular studies confirmed the sister relationship of the tribe to the Genisteae, but no complete generic-level molecular phylogeny for the tribe exists.

Such a phylogeny would provide insight into the monophyly of the genera in the tribe as well as their relationships to each other. Most of the genera have been revised in recent years (Aspalathus, Dahlgren 1988; Bolusia, Van Wyk 2003a, Van Wyk et al. submitted;

Crotalaria, Polhill 1982; Lotononis, Van Wyk 1991b; Pearsonia, Polhill 1974; Rafnia,

Campbell and Van Wyk 2001; Wiborgia, Dahlgren 1975), but some uncertainties still remain regarding the monophyly of some of the genera, especially Lebeckia and

Lotononis. Lebeckia, Robynsiophyton, Rothia and Spartidium are the only genera that lack formal revisions. Spartidium has been included in the tribe Crotalarieae, but its tribal affinity and relationships have been uncertain for some time (Polhill 1976; Van

Wyk 1991a).

Lebeckia was last revised in 1862 by Harvey in the Flora Capensis. It is a diverse group of species with high levels of morphological variation and the generic concept (broadened by Bentham in 1844 to include several crotalarioid genera) needs

2 CHAPTER 1: GENERAL INTRODUCTION AND OBJECTIVES to be re-evaluated. Taxonomic and systematic studies of the type section of Lebeckia have recently been completed (Le Roux, 2006; Le Roux and Van Wyk 2007; Le Roux and Van Wyk 2008; Le Roux and Van Wyk 2009), but the other sections of the genus and their relationship to other genera in the tribe have yet to be ascertained. When this has been completed, a treatment of all the genera in the tribe Crotalarieae will have been completed and formally implemented taxonomically.

1.2 OBJECTIVES OF THE STUDY

This project was aimed at a multi-disciplinary study of the genus Lebeckia and related genera of the tribe Crotalarieae. Relationships within the Crotalarieae are explored using DNA sequences (ITS and rbcL) and morphological data and the monophyly of the various genera is assessed. A literature review of chemical characters (mainly alkaloids) was needed in order to evaluate the congruence between molecular, morphological and chemical data and how these patterns should be interpreted within the context of new generic and suprageneric concepts. The DNA study was also aimed at confirming the suspected polyphyly of Lebeckia and Lotononis

(as currently circumscribed) and at improving generic circumscriptions. The aim was to propose and formalize new generic concepts for Lebeckia s.l. and Lotononis s.l. in order to ensure that all of the resultant genera are monophyletic. Part of the objectives was also to explore infrageneric relationships within the newly circumscribed lebeckioid genera through cladistic analyses based on morphological and anatomical data in addition to the DNA sequence data. The project aims included formal taxonomic

3 CHAPTER 1: GENERAL INTRODUCTION AND OBJECTIVES revisions of the new genera derived from Lebeckia s.I., as well as the largely unstudied

Spartidium, Robynsiophyton and Rothia. The overall objective was therefore to not only propose a new generic classification system for the tribe but to actually formalize all the proposed changes in order to eliminate uncertainty and to ensure long term nomenclatural stability.

4 CHAPTER 2: MATERIALS AND METHODS

Authorities for scientific plant names (according to Brummit and Powell 1992) are given in Appendix A3 or in the taxonomic revisions (Chapters 6, 7, 9, 10) and are not repeated elsewhere.

2.1 MORPHOLOGY

Morphological data were obtained from herbarium specimens as well as fresh material of various taxa of the Crotalarieae collected during field excursions. During these collecting trips taxa were studied in their natural environment and fresh material collected. These were preserved in FAA (Sass 1958; formaldehyde: acetic acid: 96% alcohol: water; 10:5:50:35) and silica (Chase and Hill 1991). Observations made in the field provided invaluable insight into species and generic concepts. Photographs of habit and flowers or fruit were taken and all specimens collected are housed in the

University of Johannesburg Herbarium (JRAU). Specimens of Calobota and Wiborgiella from the following herbaria were studied either on site or on loan: BM, BOL, GRA, J,

JRAU, K, NBG (including SAM and STE), P, PRE, S, SBT, UPS and WIND

(abbreviations according to Holmgren et al., 1990). Online photographs of the collections of B, M and Z were also studied. Specimens from BM, K and PRE along with digital images of critical specimens obtained from BR and LISC were studied for the revision of Robynsiophyton. Morphological characters of Rothia were assessed using specimens from BM, BOL, K, MEL, NBG (including SAM), PRE, S and UPS. The

5 CHAPTER 2: MATERIALS AND METHODS material studied for the revisions of Calobota, Robynsiophyton, Rothia and Wiborgiella were arranged according to geographical distribution so that the relevant measurements and observations could be done within several geographical populations. At least three measurements per specimen were done where possible.

Data on the distribution of the various species were gathered from the herbarium material, field notes and from Leistner and Morris (1976; for southern African taxa).

These data are presented as maps. The specimen citation system used in the

"specimens examined" sections of the revisions is that of Edwards and Leistner (1971), i.e. the quarter degree reference system. Additional information on flowering and fruiting phenology, habitat and habit were also noted.

Mature flowers from material fixed in FAA or flowers re-hydrated from rich herbarium specimens were dissected and mounted in glycerol. Specimens from which dissections were made are listed in Appendix A1. Floral characters recorded were: pedicel length, length of the bracts and bracteoles, flower length, length of the calyx, length of the calyx tube, depth of the upper, lateral and lower sinuses of the calyx, width of the upper, lower and lateral calyx lobes, length and width of the standard-, wing- and keel petals and the length of the claws of each petal. The measurements done on the vegetative and reproductive parts are illustrated in Fig. 2.1. Drawings of vegetative and reproductive structures were done using a stereoscope (WILD M3Z) with a camera lucida attachment. All the drawings were done by myself.

6 CHAPTER 2: MATERIALS AND METHODS

2.2 ANATOMY

For anatomical studies, material was fixed in FAA or dried material was rehydrated by briefly boiling in distilled water and then fixed in FAA for 24 hours. The method of Feder and O'Brien (1968) for embedding in glycol methacrylate (GMA) was used, except that the final infiltration in GMA was done for a minimum of five days.

Sections were stained according to the periodic acid Schiff/toluidine blue (PAS/TB) staining method (Feder and O'Brien 1968) and mounted. Voucher information for the material used in the anatomical studies are listed in Appendix A2. Photographs were taken with a JVC KY-F1030 digital camera.

Fruit length Petal length Petal length

Claw length Lamina length /AIIIII"!IUIIIII.rtht Fruit width

IP" Seed length Lamina width Lamina width Keel petal Seed width

Seed

Standard petal Lateral lobe length Wing petal Upper lobe length \ Carina! lobe length

Terminal leaflet length Width idth

Ovary width Lateral leaflet length Calyx length

Calyx Ovary length Pistil Petiole length Bract length Length Width Trifoliolate leaf Bracteole length

Bract and bracteoles Unifoliolate or simple leaf

FIG. 2.1 Measurements of leaves and floral parts used for morphological descriptions.

7 CHAPTER 2: MATERIALS AND METHODS

2.3 DNA SEQUENCING AND PHYLOGENETIC ANALYSES

A phylogenetic study of the Crotalarieae was undertaken in collaboration with my supervisors and other co-workers (Chapter 3; Boatwright et al. 2008a; see Appendix

B3). The methods followed in this study are discussed below.

2.3.1 Plant accessions and choice of outgroups—A total of 175 sequences of

ITS (88% of the total ITS matrix) and 207 sequences for rbcL (91% of the total rbcL matrix) were produced from taxa of the Crotalarieae, representing a total of 135 out of ca. 1,204 species, and combined with previously published sequences of taxa from the tribes Genisteae and Podalyrieae as outgroups (Van Wyk and Schutte 1995; Crisp et al. 2000; Boatwright et al. 2008b). The gene regions ITS and rbcL were selected based on previous studies where they provided sufficient informative characters for analyses at the generic level. Sequences of both the ITS and rbcL regions were available for 161 taxa of the Crotalarieae and these were used for the combined molecular analysis. A morphological matrix was compiled for those species included in the combined molecular analysis, based on 31 characters (including a few chemical and cytological characters) that were scored from examining specimens from BM, BOL, GRA, JRAU,

K, NBG (including SAM and STE), P, PRE, S, UPS and WIND for Lebeckia,

Robynsiophyton, Rothia and Spartidium and from literature for the other genera (Polhill

1974, 1976, 1982; Dahlgren 1975, 1988; Schutte and Van Wyk 1988; Van Wyk 1991b;

Van Wyk and Schutte 1995; Campbell and Van Wyk 2001; Van Wyk 2003a; Van Wyk et al. submitted). These data, along with sequences of the same species, were used to perform a combined molecular/morphological analysis. Subsequent to the study

8 CHAPTER 2: MATERIALS AND METHODS

presented in Chapter 3 (Boatwright et al. 2008a), material of the rare and unusual

Lotononis macrocarpa, a species which is pivotal in studying relationships within the

genus Lotononis, was obtained from the Compton Herbarium (Chapter 8). The

combined rbcL/ITS/morphology data set of Boatwright et al. (2008a) was used to

evaluate relationships within Lotononis and assess the position of L. macrocarpa. DNA

of this anomalous species was extracted using a DNeasy Plant Minikit (Qiagen Inc.,

Hilden, Germany) and sequenced following the procedures outlined below. These sequences of ITS and rbcL were added to the matrix of Boatwright et al. (2008a), which required minimal adjustments to the original alignment, and the matrix was re-analyzed with and without the morphological data. Parsimony (MP) analyses were conducted as described below (see Chapter 8).

2.3.2 DNA extraction, amplification and sequencing—Sequencing of the gene regions was carried out in two independent laboratories at the University of

Johannesburg and University of Heidelberg. Therefore the strategies employed differed slightly. The gene sequencing methods of Kass and Wink (1997) were followed at

University of Heidelberg, whereas sequences generated at the University of

Johannesburg were gathered according to the methods described below. Sequences that were generated at the University of Johannesburg (by J.S. Boatwright) are marked with an asterix in Appendix A3.

DNA was extracted from silica-dried or herbarium leaf material using the 2x hexadecyltrimethylammonium bromide (CTAB) method of Doyle and Doyle (1987) and purified through QlAquick silica columns (Qiagen Inc., Hilden, Germany). Sources of material used in the study are listed in Appendix A3 (for species of Lebeckia,

9 CHAPTER 2: MATERIALS AND METHODS

Spartidium and Wiborgia synonymous names in Calobota and Wiborgiella are provided

in brackets). The internal transcribed spacers (ITS) of nuclear rDNA were amplified

using the primers of White et al. (1990) and Sun et al. (1994), while for rbcL those of

Fay et al. (1997) were used. Amplification reactions were carried out using polymerase

chain reactions (PCR), in 25 pl reactions containing: 22.5 pl ABgene 1.1x PCR

Mastermix (ABgene House, Blenheim Road, Epsom, Surrey, KT19 9AP, UK) consisting

of 1.25 units Thermoprime Plus DNA Polymerase, 75 mM Tris-HCI (pH 8.8), 20 mM

(NH4)2SO4, 1.5 mM MgCl2, 0.01% (v/v) Tween® 20, 0.2 mM each of

deoxyribonucleotide triphosphates (dNTPs); 0.3 pl of both forward and reverse primers

(0.1 ng/pl); 0.8 pl 0.004% bovine serum albumin (BSA); 1.2% dimethyl sulfoxide

(DMSO; for ITS only); 20-50 ng DNA template; and sterile distilled water to make up a

final volume of 25 pl. The DMSO was added to the amplification reactions as this may

improve sequencing of ITS due to relaxation of the template secondary structure during

amplification (Alvarez and Wendel 2003). The PCR cycles used for ITS consisted of 26

cycles of 1 min denaturation at 94°C, 1 min annealing at 48°C, 3 min extension at 72°C

and 7 min final extension at 72°C; and for rbcL, there were 28 cycles of 1 min denaturation at 94°C, 1 min annealing at 48°C, 1:30 min extension at 72°C and 7 min final extension at 72°C. Sequencing reactions were performed using the Big Dye

Terminator version 3.1 Cycle Sequencing Kit (Applied Biosystems Inc., Foster City,

California, USA) and sequenced on a 3130 x/ Genetic Analyzer (Applied Biosystems

Inc.), with the addition of 5% DMSO for the ITS reactions. The cycle sequencing thermal profile consisted of 26 cycles of 10 sec denaturation at 96°C, 5 sec annealing

10 CHAPTER 2: MATERIALS AND METHODS

at 50°C and 4 min at 60°C in a thermal cycler (PCR and cycle sequencing reactions

were performed in a GeneAmp PCR system 9700 or 9800).

2.3.3 Sequence alignment and phylogenetic analyses— Complementary

strands of the sequenced genes were edited in Sequencher version 3.1.2 (Gene Codes

Corporation), and aligned manually in PAUP* version 4.0b10 (Swofford 2002).

Alignments used in this study are available from TreeBASE (study accession number

S2070). Insertions and deletions (indels) of nucleotides were scored as missing data

and thus did not contribute to the combined analysis, but an additional search with gaps from the ITS data set coded as binary characters was performed (no gaps were present

in the rbcL dataset). Gaps were coded in SeqState version 1.32 (Muller 2005) using simple indel coding as described by Simmons and Ochoterena (2000). Maximum parsimony analyses (MP; Fitch 1971) and Bayesian MCMC analysis (BI; Yang and

Rannala 1997; Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003;

MRBAYES version 3.1.2) were performed as described in Boatwright et al. (2008b).

Only the combined ITS/rbcL data set was analysed using BI. Internal support for MP was estimated with 1,000 bootstrap replicates (Felsenstein 1985) using tree bisection- reconnection (TBR) and holding 10 trees per replicate. The following scale for bootstrap support percentages (BP) was used: 50-74%, low; 75-84%, moderate; 85-100%, strong. Congruence of the separate datasets was assessed by examining the individual bootstrap consensus trees in order to compile a combined ITS and rbcL matrix for the

161 taxa where both sequenced regions were available. The bootstrap trees were considered incongruent only if they displayed 'hard' (i.e. strong bootstrap support) rather than `soft' (i.e. low bootstrap support) incongruence (Seelanan et al. 1997; Wiens

11 CHAPTER 2: MATERIALS AND METHODS

1998). In addition, incongruence length difference tests (ILD, Farris et al. 1995) were performed using the partition-homogeneity test of PAUP*. The test was implemented with 1,000 replicate analyses, using the heuristic search option with simple addition of taxa, TBR and the MULTREES option in effect.

The GTR+I+G model [selected by MODELTEST v. 3.06 (Posada and Crandall

1998) using the corrected Akaike information criterion (AICc)] was implemented for the

BI analysis and a total of three million generations were performed with a sampling frequency of 10. A majority rule consensus tree was produced to determine the posterior probabilities (PP) of all observed bi-partitions (only PPs above 0.5 are reported on the tree). The following scale was used to evaluate the PPs: 0.50-0.94, low; 0.95-1.0, strong.

Despite repeated attempts, it was only possible to amplify ITS1 for some samples of Lotononis marlothii, L. pentaphylla, L. platycarpa and L. rostrata. Only the first half of rbcL for some samples of Aspalathus laricifolia subsp. laricifolia, A. shawii subsp. shawii, Lotononis falcate, L. platycarpa, Robynsiophyton vanderystii, Rothia hirsuta, Spartidium saharae and Wiborgia humilis could be amplified. Missing data represented 4.3% of the entire combined molecular matrix due to the fact that both ITS and rbcL sequences were not available for some of the Genisteae included as outgroups.

2.3.4 Morphological analyses— Much debate currently surrounds the inclusion of morphological characters in phylogenetic analyses (e.g., Scotland et al. 2003).

However, Wiens (2004) discusses the importance of morphological characters in phylogenetics, suggesting that their implementation in separate and combined analyses

12 CHAPTER 2: MATERIALS AND METHODS may provide more rigorous phylogenies, while improved resolution has been shown by

Wortley and Scotland (2006) when morphology is combined with molecular data.

Characters and character states used for the cladistic analysis are given in Appendix

A4. The character states were polarised using the method of outgroup comparison.

Phylogenetic analyses were performed in PAUP* (Swofford 2002) with the characters treated as unordered and equally weighted (Fitch 1971). A heuristic search with 1,000 random addition sequences, TBR and the MULTREES option off was performed with a limit of 10 trees held per replicate. Internal support was assessed using 1,000 bootstrap replicates (Felsenstein 1985) as described above. Dichilus gracilis, a suffrutex, was selected as the outgroup for the analysis. A second analysis was performed where

Genista tinctoria L., a shrub, was used as outgroup to test whether polarising the shrubby habit as ancestral had an effect on the topology, but no difference was found in this analysis (tree not shown).

Morphological analyses were performed to study infrageneric relationships in

Calobota (Chapter 6) and Wiborgiella (Chapter 7). All the species within each of these genera were included and thus the sampling was different to that of the combined molecular/morphological analyses. Character states were scored for 17 morphological and anatomical characters for Calobota (see Table 6.1) and 13 characters for

Wiborgiella (see Table 7.1). Where two states were found to co-occur in a single taxon they were scored as polymorphic and autapomorphies were omitted from the analyses.

The morphological data sets were analysed using parsimony (Fitch 1971) in PAUP*. All characters were treated as unordered and equally weighted. Tree searches were performed using the branch-and-bound algorithm for Wiborgiella (random sequence

13 CHAPTER 2: MATERIALS AND METHODS addition and MULTREES on) and a heuristic search for Calobota (1,000 random addition sequences, TBR and MULTREES on). Internal support was assessed using

1,000 bootstrap replicates (Felsenstein 1985) with TBR swapping and MULTREES off.

2.3.5 Evolution of morphological characters.—The patterns of evolution of 12 characters (Table 8.1) traditionally regarded as apomorphies for Lotononis s.l. or groups therein were examined by reconstructing these onto the minimal length trees of both the analysis with and without the morphological data included (presented in

Chapter 8). An additional 6 characters within the Crotalarieae were also included in addition to the 12 mentioned before to examine their patterns of evolution within the tribe (see Appendix A5 and Chapter 11). The character reconstructions were done using parsimony in Mesquite version 2.58 (Maddison and Maddison 2008).

14 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE (FABACEAE)

INFERRED FROM DNA SEQUENCES AND MORPHOLOGY

3.1 INTRODUCTION

Crotalarieae is a tribe of legumes that currently comprises 11 genera and ca.

1,204 species (Van Wyk 2005). It represents the largest tribe of papilionoid legumes in

Africa and also within the genistoid alliance, comprising about 51% of genistoid legumes (species totals in Lewis et al. 2005 used for calculation). The size of the tribe is mainly due to the fact that the genus Crotalaria contains ca. 690 species (Polhill

1982). Some of the species are important commercially such as Aspalathus linearis, which is used for the production of rooibos tea (Van Wyk et al. 1997), and Lotononis bainesii Bak., an important fodder plant (Bryan 1961). Some Crotalaria and Lotononis species have been reported to have medicinal properties (Van Wyk 2005) and a few

are used as what is called Musa -pelo in Lesotho traditional medicine to cure or ease a broken heart (Moteetee and Van Wyk 2007), while others from the same genera are poisonous (Van Wyk et al. 2002).

The Crotalarieae are subendemic to Africa, with species of Crotalaria, Lotononis and Rothia occurring on other continents. Aspalathus, Rafnia and Wiborgia are endemic to the Cape Floristic Region, while Lebeckia (as currently circumscribed) is distributed throughout the Cape and extends into the south-western and central parts of

Namibia. Generic circumscriptions and relationships within the tribe are known to be complicated, with evidence of reticulation and convergence (Dahlgren 1970a; Polhill

15 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

1976, 1981; Van Wyk 1991a). The most recent evaluations of generic relationships in the tribe are those of Polhill (1976, 1981), Van Wyk (1991a) and Van Wyk and Schutte

(1989, 1995). According to Van Wyk (1991a), two main groups can be identified within the tribe, namely the "Cape group" comprising Aspalathus, Lebeckia, Rafnia and

Wiborgia, and the Lotononis group comprising Lotononis, Pearsonia, Robynsiophyton and Rothia. The placement of Bolusia, Crotalaria and Spartidium within these two groups is not clear, but a close relationship between the former two genera has been suggested. Recent revisionary studies of Lebeckia (the first since Harvey's treatment of

1862) have shown that the genus is unlikely to be monophyletic (Le Roux 2006; Le

Roux and Van Wyk 2007; Boatwright, in prep.; Chapters 6 and 7).

Several molecular systematic studies of Fabaceae and specifically the genistoid legumes (sensu Polhill 1976, 1981) have been conducted in recent years (Kass and

Wink 1995, 1996, 1997; Crisp et al. 2000; Doyle et al. 2000; Kajita et al. 2001; Wink and Mohamed 2003; Wojciechowski et al. 2004; Boatwright et al., 2008b). These studies indicate the placement of Crotalarieae within the "core" genistoid clade together with, among others, the South African tribes Podalyrieae and Genisteae, and confirm the exclusion of the Argyrolobium group (Argyrolobium, Dichilus, Melolobium, Polhillia) and the rest of the Genisteae from Crotalarieae (Van Wyk and Schutte 1995). A sister relationship between Crotalarieae and Genisteae, with Podalyrieae successively sister to these, is shown by the molecular studies cited above, but sampling limitations did not allow detailed evaluations at the generic level.

This chapter is aimed at exploring generic circumscriptions and relationships within Crotalarieae, using nrDNA ITS sequences, plastid rbcL sequences and

16 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE morphological data, based on a sample of 135 species representing all of the 11 genera currently recognized, as well as major infrageneric groups within some of the genera. The aim was to assess the monophyly and relationships of the individual genera.

3.2 RESULTS

3.2.1 rbcL data set—The rbcL matrix comprised 1,296 aligned positions with

222 variable and 167 parsimony informative characters. Tree searches produced 2,650 equally parsimonious trees of 527 steps, a consistency index (CI) of 0.50 and a retention index (RI) of 0.92. Overall the strict consensus tree (Fig. 3.1) is poorly resolved with a few supported clades. A Glade consisting of Lebeckia section Calobota

(Eckl. & Zeyh.) Benth., L. section Stiza (E.Mey) Benth. and Spartidium saharae is weakly supported (60 BP). Lotononis s.s. (L. section Lotononis and allies) is moderately supported (74 BP), while the rest of the genus is unresolved except for L. section Listia

(E.Mey.) B.-E.van Wyk, which has high support (89 BP). The Crotalaria Glade is strongly supported (88 BP) and the sister relationship between Bolusia and Crotalaria is also strongly supported with a BP of 90. Both these genera are well-supported to be monophyletic (Bolusia 98 BP; Crotalaria 90 BP) and they are successively sister to

Lotononis hirsuta [L. section Euchlora (Eckl. & Zeyh.) B.-E.van Wyk].

17 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Willa diffuse Rafrria angulata Rafnia swats Rafnia racemosa Rafnia acuminate R&M, globose Lebeckm samosa 1 98 Lebecka inflate 1 Lebeclda inflate 2 79 Lebeckia pauciflora 1 1 Letedda pairciliora Le bob= cfillfioides 1 Le sencea 3 Lebeckla cytwoolessoncea 2 Lebeckia 1 71 1.E Lebetlda saricea 2 Lebeckia halenbergensis 1 65 61 2 Lebeckis multiflora 72 Lebedda lofiononcides 60 68 LI.8 =Tarpons's 2 Lebeckia spinescens Sparlidium sesame 6 Lebeckia macrantha 1 Le Lebeddabeck' =ides 1 LebeUria gi=s 2 Lebeckia psiloloba Lebeckia meyeriana 1 60 Lebeckia meyeriana 2 Lebedda samosa 2 71 LebecJdaer l .aletiana 2 = felbeb%?.9 " ag 2 86 Lebeckia wrighttit 3 Lebeckia might& 1 Lebeckia wrighttii 2 Lebeckia lei 'am 2 Lebecka ambigua 1 Lebeclda sepiarie 2 Lebeckia plukenetiana 1 Lebeckia µbe Lebeclfia ambigua 2 iercukala sseePsfigesj 57 E Viborgia t1SapabzgggsnaiaPrifirr 1 Pearsonw arista 90 E AZOV gia.c i sgg gre "le prsonta sewI ;a f poirsooszwnorora.m. .. Rodle hirsdra'n rYsb' Sg E rigi: rsracer 1 Il iorz tetzgrtdeF, 21

Wiborgia sericea trigg re WagaT 21 Wiborpia monoptera 2 Wiborpia incurvata niPer is adasubsp . namachrensis 89 Lotononis &obi Lotononls sparslflota Lotononis lenticula Lotononis bmchyantha Lotononis falcata 2 63 E tLotononsi4zpsvozn49ab Lotononis —E= Ea22;sgl Vi! subsp. peduncularis 72 = EggaZi °. ie"c11clada iTrani Trgffetri f 2 Lotononis umbefiata to3frogg Peacrhalisa 2 Lotononis involucrata subsp. digitate Lotononis leptoloba 2 Lotononis maximiliani 1 Lotononis lomicaulis 74 Lotononls samosa Lotononls eli0C8IP8 Lotononis elongate. igirag Olferrls Lolononls stricta 66 Lotononis divaricata Lotononis sencophylla 1 Lotononis sericophylla 3 Lotononis meyeri 2 Lotononis mayor, 1 Lotononis pungens 1 Lotononis nyaromiliani 2 Ernorrng ifgoie 1 Lotononis g abra Lotononls micrantha Lotononis macrosepala Lotononis lane 1 E Lotononis lane 2 gi=gif grale

FIG. 3.1 Strict consensus tree of 2,650 equally parsimonious trees from the analysis of the rbcL data set (TL = 527; CI = 0.50; RI = 0.92). Numbers above the branches are bootstrap percentages above 50% from the parsimony analysis.

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Lotononis polycephala 2 Lotononis quinata Lotononis pentaphylla Lotononis bolus!! Lotononis mollis Lotononis moths Lotononis plicata Lotononis polycephala 1 Lotononisbenthamiana Lotononis digitata 2 Lotononis digitata 1 Lotononis ma rutica 57 Lotononis pla carpe 5 Lotononis pia ca $3.1 Lotononis pia Lotononis pia rpa 2 Lotononis pla fpa 3 89 Lotononis sot Lotononis subulata Lotononis marlothil E Lotononis Usti' 71 Aspalathus pendula Aspalathus kneads 2 E Aspalathus (inearis 1 Aspalathus nivea Aspalathus vermiculata Aspalathus camosa , E Aspalathus cronata Aspalathus perfoliata subsp. phillipsii a) co 61 lgliSign hltht rtbsp. hirta Aspalathus laricifolia subsp. laricifolia Aspalathus shawl! subsp. shawl! co Aspalathus willdenowiana Aspalathus macrantha Aspalathus confusa Aspalathus galeata 2 Aspalathus pachyloba 0 Lotononis acuticarpa Lotononis calycina Lotononis decumbens Lotononis platycarpa 6 Lotononis foliose i Lotononis globulosa 100 Lotononis eriantha Lotononis adpressa f__=95 Bolusra amboensls 1 98 Bolusra amboensis 2 &alum acuminata 50 61 Crotalaria capensis 1 61 Crotalaria capensis 2 90 Crotalaria lebeckioides Crotalaria lanceolata Crototaria totoides 71 crotararia virgultalis 2 Crotalana virgultatis 1 88 90 I-- Crotalaria humilis 1 .— Crotalana humilis 2 88 Crotalana distans Crotalana griquensis Crotalana sp. 100 Lotononis hirsuta Lotononis hirsuta 2 L Lotononis hirsuta 3 72 1 Lupinus polyphyllus .-- Ulex europaeus 100 Polhillia brevicalyx Polhillia obsoleta Polhillia involucrata Polhillia canescens Polhillia pallens 91 1 Argyrolobrum !unary subsp. senceum 6 I— Mgifgrrn ma Igirm 1— A/gyro/06/pm sp. Dichilus pilosus 99 oichitusstrictus uichrius renexus fa Dichilus gracilis Dichilus lebeckioides Melolobium adenodes 2 Melolobium aethiopicum 1 Melolobium alpinum a) Melolobium candicans 2 Melolobium canescens \ &Ci= Melolobium humile 61 Melolobium candicans 3 Melolobium microphylium 1 Melolobium subspicatum Melolobium wilmsi! 100 Melolobium adenodes 3 Melolobium aethiopicum 2 kiggstr, Atiecigtg, \ \ Melolobium exudans Melolobium microphyllum 2 Amphithalea alba Amphithalea michrantha Amphithalea williamsonii 0) 75 t$Sgging/satiMia to Podare buxifolio co 57 ip'cirograkaggt? Xiphotheca rellexa 78 Stirtonanthus chrysanthus Caloumia sertcea . 0 88 = ieJopia gerustordes Cydopia subtemata tfirgika divaricata Cadia purpurea

FIG. 3.1 Continued.

19 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Tribe Crotalarieae as a whole is strongly supported as monophyletic (94 BP) and sister to the Genisteae (100 BP). Genisteae and Podalyrieae are both supported as monophyletic (89 BP and 78 BP, respectively).

3.2.2 ITS data set—The analysis of the ITS data set included 560 characters, of which 353 were variable and 256 parsimony informative. Analysis resulted in 5,970 equally parsimonious trees with a tree length (TL) of 1,031 steps, a CI of 0.51 and a RI of 0.85. The analysis in which gaps were coded resulted in identical Glade resolution as the analysis without coded gaps, although some clades had stronger or weaker support

(not shown). The analysis with coded gaps resulted in 2,540 trees (TL = 1,340; CI =

0.51; RI = 0.84). The strict consensus tree based on ITS data (Fig. 3.2) is better resolved than that of rbcL, with several well-supported clades. Aspalathus was weakly supported as monophyletic (64 BP), although this support percentage is higher when the gap characters are included (78 BP). Lebeckia and Wiborgia are both polyphyletic.

Lebeckia section Viborgioides Benth. including Wiborgia humilis is moderately supported (80 BP), while the positions of L. inflata and L. mucronata are unresolved.

The remaining species of Wiborgia form a weakly supported Glade (61 BP; 79 BP with gap coding). Rafnia is weakly supported as monophyletic (75 BP) and Lebeckia section

Calobota along with L. section Stiza and Spartidium saharae also has weak support (59

BP). Lebeckia section Lebeckia is unresolved. Lotononis is left unresolved and L. hirsuta is well separated from the rest of the genus. Three main clades can be distinguished.

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

1— Aspetathus !Reads 1 L— lsp'Prlthrsrelrth 2 7113 camas. sa A.P.10 7E.... Aspalatlfs rnag'rranthe 62 71 _ Aspalathos willdenowisna ... Aspalathus confuse 55 ,-- Aspalathus perfolleta subsp. plullips. 59 .— Aspalathm crenate rAspalathen vemladata 1 53 Aspalathm shawii subw. th aw* 60 Aspelathus laricdolla subs'. laricffolk 78 ASp81811,1111 hes subsp. hute j7t Aspalathus hystrix Aspired!. pe..17= Aspelathus Aspalathus MAR 424— Monne hate 1 ilE Wlborgla ;re 2 ; 11 jii,..52= Magi. obcordata 1 7, 62 Meru!, obcordata 2 Mbar*, manoptera 1 r borgia monoptera 2 iEtinWrborgie incurvata 90 r— 1.2=1:11:1 Lebecida Iniptai 92 Lebeckia 1179643 2 Klboujia hunglis 1 55 91 W.= hunt.4.231. I Lebeckia sena fella I I ! 55 Lebeckia bovereana 1 74 87 Labeckia bowieena 2 Lebeckk! letpoldtrana 2 1 S 1 Lebeckia microns!. Leber-Ada much:nets Rafnla acuminate 74 Rafnla spicata 75 Refute wends 73 78 Rah... globose Rafnia racemosa Rafnia diffuse I I 60 Lebeckia pungens 1 Lebeckie pungens 2 98 Lebeckia macranthe 1 18 i 54 99 54 Lebeckia Inacranthe 2 LebeekR psilobbe i t 59 P Lebeckia rnelilotoides 1 Lebeckia s00000 1 a * 69 iii,-,E,Leb.abeclde,...... sedate 32 g • 7 1 Lebeckia cyllsoldes 1 i I 59 00 Lebeckia cylisoides 2 77 5 —Lebecie mutfon e— Leberida lolononordes Lebeckia hatenbergensis 1 Lebeckia halenbergensis 2 I i Lebeckla spinescens Lebeckia comma Spertidtum wham, Lebeckia carnosa 1 c Lebeckia comma 2 61 Lebeckiapl ukenedene 2 63 Lebeckia a mbigua I Lebeckia 00011800 3 Lebeckia pukonetiana 1 I Lebeckie brevrcarpe Lebeckie amnion! 2 i Lebeckia separia 1 Lebeckia sepiana 2 Lebeckia greets Lebeckia meyenana 1 Lebeckia mosnia09 3 Lebeckia wrchttii 1 i 63 Lebeckia AngtrItii 2 I 88 Lebeckia wrightlii 3 93 I— Lebeckia peucif bra 1 94 .— Lebeckia par...ors 2 Lotononis falcate 1 95 Lotononis pandit... Lotononis fruticoldes Lotononis brachyanthe 1060000)0 falcate 2 Lotononis rabenaviana 57 Lotononis spardtbre lotononis rostrata Lotononis oxyptere Lotononis lenticula Lotonords rostrata subsp . flamer:penes Lotononis taxa 1 r— Lotononis taxa 3 .— Lotononts glabra r— Inforronis rnacranthe .-•■•- Lotononis macrosepala ,-- Lotononis macriate 5868 L—Lotonanis curtii Lotanonis sabulose Lofononis lapIcLortz ni1 I 62 92 8090 72 1-1_°k"c•niiss rpf 2 11111 96 Lotononis umbellate 2 68 Lotononts rigida 2 • 85 Lotononis thanse subsp. NRICOCI.49 92.3 8960 Loton.nr:Lis einvomslipd.,..: ... dig. Lotononis 100010 02 97 Lotononks involucrate sully. Pedunculads Lotonorus prostn3ta 2 ,a, 95 r—L0f011004 filitOrMis 99 '—' 72 60 E Li. °18ofonon44 ;tare 1 Lamont., meyed 1 60 Lotonorm Lotononu =Zeta Lotonona sericophytte 1 70 91= Lotononls settcophylla 2 Lofonona stride se Lotonona camosa 6 ,—. Lotononis pungens 1 so , Lotonorus pungens 2 Lotononrs lototronottles

FIG. 3.2 Strict consensus tree of 5,970 equally parsimonious trees from the analysis of the ITS

data set (TL = 1,031; CI = 0.51; RI = 0.85). Numbers above the branches are bootstrap

percentages above 50% from the parsimony analysis.

21 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Lotononis platycarpa 1 Lotononis calycina 67 71 Lotononis fohosa 1 69 Lotononis foliosa 2 6 66 65 Lotononis eriantha 57 Lotononis adpressa 73 Lotononis acuticarpa Lotononis decumbens 65 C Lotononis platycarpa 2 63 Lotononis platycarpa 5 Lotononis platycarpa 3 Lotononis platycarpa 4 Lotononis bolusii 93 Lotononis mollis 1 81 94 Lotononis mollis 2 96 Lotononis platycarpa 6 92 97 Lotononis benthamiana Lotononis digitata 1 Lotononis digitate 2 Lotononis quinata 64 Lotononis plicata Lotononis globulosa 58 Lotononis magnifica a) Lotononis polycephala 1 co Lotononis polycephala 2 a) Lotononis pentaphylla C. Lotononis solitudinis co 99 90 Lotononis subulata To 100 o Lotononis listii 94 Lotononis marlothii Pearsonia sessilifolia 1 95 Pearsonia sessilifolia 2 97 97 Pearsonia aristata Pearsonia sessilifolia 3 98 82 Pearsonia grandifolia 98 Pearsonia cajanifolia 1 89 1 100 99 97 10 Pearsonia cajanifolia 2 100 uf Rothia hirsute 99 C Robynsiophyton vanderystii 100 Lotononis hirsute 1 E Lotononis hirsute 2 100 Lotononis hirsute 3 99 Crotalaria distans 99 98 Crotalaria griquensis 87 99 Crotalaria sp. 1 94 100 r— Crotalaria capensis 1 99 10 93 I— Crotalaria capensis 2 86 Crotalaria virgultalis 1 64 10010 Crotalaria humilis 1 89 10 Crotalaria humilis 2 67 Crotalaria lanceolata Bolusia amboensis Dichilus pilosus 66 c Dichilus lebeckioides a) 10 62 Dichilus &rictus co 51 00 Dichilus gracilis 54 100 Melolobium adenodes 1 63 100 1-- Melolobium candicans 2 59 51 Polhillia pallens Argyrolobium lunare 0 c Genista teretifolia 89 I Argyrolobium harmsianum 52 Amphithalea alba 99 Amphithalea michrantha 80 100 Amphithalea williamsonii 67 96 r— Xiphotheca reflexa a) CO 99 Xiphotheca guthriei CD 09 Liparia angustifolia 87 90 Liparia hirsuta 86 Podalyria buxifolia CO Podalyria oleaefolia Stirtonanthus chrysanthus 0 Virgilia divaricata CL 60 Calpumia sericea 100 Cyclopia genistoides 100 Cyclopia subtemata Cadia purpurea

FIG. 3.2 Continued.

22 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

The first consists of Lotononis s.s. (also supported in the rbcL tree) with moderate support (84 BP). The second clade, comprising taxa from Lotononis section Leptis

(E.Mey. ex Eckl. & Zeyh.) Benth., L. section Listia and their allies also has low support

(64 BP). The third Glade of Lotononis groups with Bolusia and Crotalaria with strong support (87 BP) and comprises one anomalous species, L. hirsuta, which makes up the monotypic section Euchlora. Crotalaria and Bolusia are weakly supported as sister taxa

(64 BP) and Crotalaria is strongly supported to be monophyletic (99 BP). Pearsonia,

Robynsiophyton and Rothia form a strongly supported Glade (98 BP) and the latter two are strongly supported to be sister genera (97 BP). The taxa of this Glade all possess a

17 base-pair deletion at positions 179-196 in the aligned ITS matrix. The Crotalarieae are strongly supported to be monophyletic (81 BP) and sister to Genisteae (100 BP).

3.2.3 Combined ITS/rbcL data set—Visual inspection of the bootstrap consensus trees resulting from separate analyses of ITS and rbcL sequences presented no strongly supported incongruent patterns. The ILD test indicated significant difference between the two datasets (p=0.001). Following the suggestions of Seelanan et al. (1997) and Wiens (1998), together with suggestions that the ILD test may be unreliable (Reeves et al. 2001; Yoder et al. 2001) these datasets were combined directly. The combined ITS and rbcL matrix consisted of 1,854 included positions, of which 540 were variable and 404 parsimony informative. The MP analysis produced

560 equally parsimonious trees (Fig. 3.3; TL = 1,473; CI = 0.50; RI = 0.86).

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

100 c Aspalathus linearis 1 Aspalathus pendula 1.0 Aspalathus linearis 2 Aspalathus camosa Aspalathus galeata Aspalathus macrantha Aspalathus willdenowiana Aspalathus confusa 68I-- Aspalathus perfoliata subsp. phillipsii 0.99 I-- Aspalathus crenate Aspalathus vermiculata 63 Aspalathus shawii subsp. shawii 1.0 64 Aspalathus laricifolia subsp. laricifolia Aspalathus hirta subsp. hirta Aspalathus hystrix Aspalathus nivea Aspalathus pachyloba 51 Wiborgia fusca 1 80 Wiborgia tetraptera 1 0.99 Wiborgia tetraptera 2 Wiborgia sericea 65 Wiborgia obcordata 1 0.99 1— Wiborgia obcordata 2 96 E Wiborgia monoptera 1 Wiborgia monoptera 2 1.0 Wiborgia incutvata Wiborgia fusca 2 Lebeckia inflata 1 Lebeckia inflata Lebeckia inflata 2 .o Lebeckla mucronata .0 Lebeckia mucronata 60 Lebeckia pungens 1 Lebeckia pungens 2 10 55r Lebeckia macrantha 1 Lebeckia macrantha 2 Lebeckia psiloloba " Lebeckia sericea 1 ..9: Lebeckia sericea 2 Lebeckia sericea 3 roup 100 g 93 Lebeckia cytisoides 1 e Lebeckia cytisoides 2 1.0 89 Lebeckia multitlora

Lebeckia lotononoides ''Cap 55 Lebeckia halenbergensis 1 Lebeckia halenbergensis 2 Lebeckia spinescens Lebeckia cinerea Spartidium saharae 54 Rafnia acuminate Rafnia racemosa 97 Rafnia spicata 1.0 1.0 Rafnia globosa Rafnia diffuse 63 Wiborgia humilis 1 94 9 Wiborgia humilis 2 52 Lebeckia leipoldtiana 1 Lebeckia bowieana 1 64 98r— 1.0 1— Lebeckia bowieana 2 1.0 Lebeckia leipoldtiana 2 0.98 C1— Lebeckia sessilifolia Lebeckia camosa 1

1 Lebeckia camosa 2 la

1— Lebeckia meyeriana 1 k Lebeckia plukenetiana 1 MIIMID

100I— Lebeckia pauciflora 1 bec

1.0 1— Lebeckia pauciflora 2 Le

f.62= Lebeckia wrighttii 1 n

53 100 io 0 Lebeckia wn;ghttii 2 t 1.0 1.0 Lebeckia wnghttii 3 sec I— Lebeckia plukenetiana 2 I— Lebeckia ambigua 1 la k Lebeckia brevicarpa

Lebeckia ambigua 2 bec

Lebeckia sepiaria 1 Le Lebeckia sepiaria 2 I

FIG. 3.3 Strict consensus tree of 560 shortest length trees from the combined analysis of the

ITS and rbcL data sets (TL = 1,473; CI = 0.50; RI = 0.86). Numbers above the branches are

bootstrap percentages above 50% from the maximum parsimony analysis and numbers below

the branches are posterior probabilities above 0.5 from the Bayesian analysis.

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Lotononis falcate 1 98 Lotononis parviflore Lotononis fruticoides 1.0 Lotononis brechyantha Lotononis falcata 2 Lotononis spersiflora 73 r— Lotononis oxyptera to 1 Lotononis rostrata subsp. namaquensis Lotononis rostrata o. 94 Lotononis lanticula 1.0 Lotononis taxa 91 911— Lotononis micrantha to tol Lotononis macrosepala Lotononis glabra 99 r— Lotononis maculate 1.0 "••• Lotononis curtii Lotononis sabulosa 90 Lotononis leptoloba 1 100 , 68 Lotononis maximiliani 1 — 1.0 0,99 Lotononis leptoloba 2 Lotononis pungens 1 0.61 Lotononis pungens 2 Lotononis divancata Lotononis sericophylla 1 0.99 0.521 Lotononis sericophylla 2 73 Lotononis pulchella 1 1.0 Lotononis meyeri 1 100 Lotononis dense subsp. leucoclada 97 54 1.o Lotononis exsfipulata 1.0 9 Lotononis involucrata subsp. digitata 0.94 E Lotononis involucrata subsp. pedunculatis Lotononis involucrata

95 Lotononis filiformis 1.0 Lotononis elongate s Lotononis stricta

0 Lotononis camosa roup g

Lotononis lotononoides

Pearsonia sessilifolia 1 is 98 Pearsonia aristata .o 9 Pearsonia grandifolia 100 Pearsonia cajanifolie 1 1.0 1 Pearsonia cajanifolia 2 97 Roth/a hirsuta Lotonon .0 C Robynsiophyton vanderystii 87 Lotononis platycarpa 1 Lotononis platycarpa 2 1.0 65 Lotononis platycarpa 3 I Lotononis platycarpa 4 , Lotononis platycarpa 5 Lotononis bolusii Lotononis calycina 63 66 81 68 Lotononis foliosa 1 1.0 Lotononis foliosa 2 o 0.85 97 Lotononis ertantha Lotononis adpressa 10 Lotononis acuticarpa Lotononis decumbens 94 Lotononis molris 1 1.0 1.0 Lotononis mollis 2 Lotononis platycarpa 8 Lotononis benthamiana Lotononis digitate 1 Lotononis digitate 2 77 Lotononis quinata 1.0 Lotononis plicate Lotononis globulosa Lotononis magnifica Lotononis polycephala 1 Lotononis polycephala 2 Lotononis pentaphylla Lotononis solitudinis 100 Lotononis subulata 1.0 00 100 Lotononis listit 1.o 91 Lotononis marlothfi 100 E Lotononis hirsute 1 Lotononis hirsuta 2 0 1.0

Lotononis hirsute 3 up v1= Crotalaria distans 99 f ro

Crotaleria griquensis g 99 1 0 Crotalada sp. 1.0 100 r— Crotalaria capensis 1 ia

00 98 1.0 Crotalaria capensis 2 lar ■

1.0 to Crotalada virgultalis 1 ta 92 1 Crotalarla humllis 1 mii to 1 .0 Crotalaria humllis 2 Cro Crotalaria lanceolate

Bolusia amboensis nimi Polhilria pollens Polhillia brevicalyx Dichilus pilosus 44= Dichllus lebeckioides 85 Dichilus gracilis Melolobium candicans 1 1.0 Melolobium adenodes 1 Argyrolobium lunare Argyrolobium harmsianum Argyrolobium lanceolatum Argyrolobium meganhizum 61 Amphithalea alba Amphithalea micluantha Amphithalea williamsonii 97 I Xiphotheca refiexa 1.0 t X/phothece guthdei N Liparia angustifolia 97 Ltparia hirsute 54I 1 O0 Podalyria buxlfolia 93 to Podalyria oleaefolia 1.0 Stirtonanthus chrysanthus Cyclopia genistoides Cyclopie subicatergata Virgule Calpumia sericea Cadia purpurea

FIG. 3.3 (Continued).

25 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

In both the MP and BI analyses, the same major clades could be observed: the

"Cape group" consisting of Aspalathus, Lebeckia, Rafnia, Spartidium and Wiborgia (53

BP; PP 1.0); the Lotononis group consisting of Lotononis s.s., Pearsonia,

Robynsiophyton and Rothia (PP 0.94); a second Lotononis Glade consisting of

Lotononis section Leptis, L. section Listia and allies (77 BP; PP 1.0); and the Crotalaria group consisting of Bolusia, Crotalaria and Lotononis hirsuta/L. section Euchlora (99

BP; PP 1.0). Lebeckia, Lotononis and Wiborgia are all polyphyletic and Lebeckia section Lebeckia is unresolved in both the analyses.

Generic relationships are unresolved within the "Cape group" and represented as a polytomy at the base of this Glade in the bootstrap consensus tree of the parsimony analysis (not shown). In the BI tree, Aspalathus (88 BP; PP 1.0) and

Wiborgia (70 BP; PP 1.0) are sister to each other, with L. mucronata (PP 0.96) and

Lebeckia inflata included in this Glade. These positions are unresolved in the MP analysis. Lebeckia section Calobota (including L. section Stiza and Spartidium saharae;

93 BP; PP 1.0) is sister to Aspalathus and Wiborgia, followed by L. section Viborgioides

(including Wiborgia humilis; 64 BP; PP 1.0) and Rafnia (94 BP; PP 1.0). Lebeckia section Lebeckia is unresolved in both the MP and BI trees.

Lotononis is polyphyletic in both the MP and BI trees. The Lotononis group is moderately supported in the BI tree (PP 0.94) and consists of those species of

Lotononis from the sections Aulacinthus (E.Mey.) Benth., Buchenroedera (Eckl. &

Zeyh.) B.-E.van Wyk, Cleistogama B.-E.van Wyk, Krebsia (Eckl. & Zeyh.) Benth.,

Lotononis, Monocarpa B.-E.van Wyk, Oxydium Benth. and Polylobium (Eckl. & Zeyh.)

Benth. (100 BP; PP 1.0). These are sister to a strongly supported Glade (BP 100; PP

26 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

1.0) consisting of Pearsonia (98 BP; PP 1.0), Robynsiophyton and Rothia. The sister relationship of the latter two genera has high bootstrap support (97%) and PP (1.0).

The next Glade consists of species of Lotononis section Listia (100 BP; PP 1.0) and sections Digitata B.-E.van Wyk, Leobordea (Del.) Benth., Leptis, Lipozygis

(E.Mey.) Benth. and Synclistus B.-E.van Wyk (99 BP; PP 1.0) and these are moderately to strongly supported as sister (77 BP; 1.0 PP).

The Crotalaria group consists of Bolusia and Crotalaria (100 BP; PP 1.0), strongly supported as sister groups (92 BP; PP 1.0), and Lotononis hirsuta

(corresponding to Lotononis section Euchlora). This Glade receives high support in both the MP and BI analyses (99 BP; PP 1.0).

The Crotalarieae are strongly supported as being monophyletic (97 BP; PP 1.0) and sister to Genisteae (100 BP; PP 1.0). The monophyly of the latter tribe receives low to strong support (65 BP; PP 1.0). The Podalyrieae s.s. are strongly supported as monophyletic (93 BP; PP 1.0) and the relationships within the tribe conform to those of previous studies (Boatwright et al., 2008b).

3.2.4 Morphological analysis—The morphological matrix included 31 polarised characters, 29 of which were parsimony informative. Parsimony analysis yielded 261 trees (not shown) of 101 steps (CI = 0.36, RI = 0.93). In the strict consensus tree (not shown), Aspalathus and Rafnia group together, albeit without support. Aspalathus

(excluding A. crenata and A. perfoliata) and Rafnia are weakly supported as monophyletic (55 BP and 71 BP, respectively). Lebeckia section Viborgioides, L. mucronata and Wiborgia s.s. group together, while L. inflata groups with Lebeckia section Lebeckia, but without support. Only Wiborgia is weakly supported as

27 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE monophyletic (78 BP). Lebeckia section Lebeckia is moderately supported as monophyletic (84 BP), but there is no resolution within this group. Lotononis hirsuta is not included in Lotononis, but groups with Crotalaria (63 BP) and Bolusia as in the molecular analysis. The rest of Lotononis is monophyletic (56 BP) in the morphological analysis as opposed to polyphyletic in the combined molecular analysis, but relationships within the genus are largely unresolved. Pearsonia, Rothia and

Robynsiophyton form a strongly supported Glade (87 BP) and a sister relationship between Robynsiophyton and Rothia is strongly supported (88 BP).

3.2.5 Combined ITS/rbcUmorphological data set—The bootstrap consensus trees from the combined ITS/rbcL analysis and the morphological analysis showed no strongly supported incongruent patterns, although an ILD test indicated significant difference between the data sets (p=0.002). The suggestions of Seelanan et al. (1997) and Wiens (1998) were followed and the data sets combined directly. The combined matrix consisted of 1,885 characters, 1,405 of which were constant, 480 variable and

303 parsimony informative. The MP analysis resulted in 370 trees (TL = 1,166, CI =

0.53, RI = 0.84; Fig. 3.4).

The trees resulting from this analysis are similar to those from the combined molecular analysis, except that Lotononis and Lebeckia section Lebeckia are monophyletic. The "Cape group" (73 BP), Lotononis group (including Pearsonia,

Robynsionphyton and Rothia), and Crotalaria group (100 BP) found in the molecular analysis were also retrieved in the combined data set of molecular plus morphological characters. Within the "Cape group", Aspalathus, Lebeckia section Calobota (including

L. section Stiza and Spartidium saharae), L. section Lebeckia, L. section Viborgioides

28 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

(including Wiborgia humilis), Rafnia and Wiborgia (excluding Wiborgia humilis) received moderate to strong support to be monophyletic (99 BP; 95 BP; 77 BP; 87 BP; 98 BP;

94 BP, respectively). Lotononis is monophyletic (75 BP; excluding L. hirsUta) and the groups retrieved within the genus are identical to the separate clades found in the molecular analysis, namely Lotononis s.s. (100 BP), Lotononis section Leptis and allies

(100 BP) and Lotononis section Listia (100 BP). The sister relationship between the latter two is strongly supported with a BP of 88. Pearsonia is strongly supported as monophyletic (99 BP) and sister to Robynsiophyton and Rothia (100 BP). The sister relationship between Robynsiophyton and Rothia is strongly supported (99 BP). The

Crotalaria group is strongly supported (100 BP) and consists of Bolusia, Crotalaria and

Lotononis hirsuta. Crotalaria is strongly supported to be monophyletic (100 BP) and is sister to Bolusia (94 BP).

3.3 DISCUSSION

3.3.1 Phylogenetic relationships—The phylogenetic hypotheses presented in this study are based on a complete sample of Crotalarieae at the generic level and a representative sample of most of the taxonomic and morphological variation within these genera. The data were based not only on DNA sequences but also on salient morphological characters that were carefully polarised and proved to be informative at the generic level.

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

100 r—Aspalathuslinearis 1 Aspalathus pendula Aspalathus camosa Aspalathus vermiculata 99 67 Aspalathus shawii subsp. shawii 63 —Aspalathus laricifolia subsp. laricifolia Aspalathus hirta subsp. hirta Aspalathus hystrix 52 Aspalathus galeata 87 r62 Aspalathus macrantha 94 ` Aspalathus willdenowiana Aspalathus confusa 96 1—Aspalathus perfoliata subsp. phillipsii Aspalathus crenata Aspalathus pachyloba Aspalathus nivea _FEWiborgia fusca 1 Wiborgia tetraptera 1 Wiborgia sericea Wiborgia obcordata 1 93 I— Wiborgia monoptera 1 I— Wiborgia incurvata Wiborgia humilis 1 87 93 Lebeckia leipoldtiana 1 Lebeckia sessilifolia Lebeckia bowieana 1 Lebeckia inflate 2 Lebeckia mucronata Lebeckia pungens 2 100 Lebeckia macrantha 2 Lebeckia psiloloba 73 9 1— Lebeckia sericea 2 I— Lebeckia cytisoides 2 95 90 Lebeckia multiflora Lebeckia lotononoides 81 Lebeckia halenbergensis 2 Lebeckia spinescens Lebeckia cinema Spartidium saharae _TE62 E Lebeckia camosa 2 Lebeckia meyeriana 1 Lebeckia plukenetiana 1 77 Lebeckia pauciflora 2 Lebeckia wrightii 1 Lebeckia brevicatpa Lebeckia ambigua 1 E Lebeckia sepiatia 1 9 r Rafnia acuminata I — Rafnia racemosa 98 99 Rafnia spicata 71 Rafnia globosa Rafnia diffusa

FIG.3.4 Strict consensus tree from the combined molecular (ITS and rbcL) and morphological

analysis. Numbers above the branches are bootstrap percentages above 50% (no. of trees =

370; TL = 1,166; CI = 0.53; RI = 0.84).

CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

Lotononis falcate 1 Lotononis parvifiora Lotononis fruticoides Lotononis brachyantha Lotononis sparsifiora 73 58 Lotononis oxyptera 54 Lotononis rostrata subsp. namaquensis 64 Lotononis lenticula Lotononis laxa 93 731— Lotononis macrosepala 98 I Lotononis micrantha Lotononis glabra

99 Lotononis maculata s.s. Lotononis curtii C is Lotononis sabulosa

99 Lotononis leptoloba 2 tonon Lo Lotononis maximiliani 1 100 Lotononis pungens 2

80 Lotononis divaricata Lotononis sericophylla 1 56 74 Lotononis pulchella Lotononis meyeri 1 Lotononis lotononoides 97 6 I— Lotononis stricta

"— Lotononis camosa

100 Lotononis densa subsp. laucoclada roup g

70 Lotononis exstipulata is

97 Lotononis involucrata a e Lotononis involucrata subsp. peduncularis Lotonon 96 Lotononis filiformis iea

75 Lotononis elongate lar ta Lotononis platycarpa 5

Lotononis calycina Cro 68 85 88 Lotononis foliose 2 65

Lotononis enantha llies a

Lotononis edpresse

59 ♦

Lotononis decumbens t Lotononis acuticarpa Lep

Lotononis bolusii ion

100 t Lotononis motifs 1 sec Lotononis plicate Lotononis quinata

Lotononis polycephala 2 Lotononis Lotononis pentaphylla Lotononis benthamiana Lotononis digitata 1 Lotononis magnifica Lotononis globulosa Lotononis solitudinis 100 H 90 Lotononis subulata Lotononis listii Lotononis marlothil 100 Pearsonia sessilifolia 1 99 Pearsonia aristata 97 Pearsonia grandifolia 100 Pearsonia cajanifolia 2 9 I— Rothia hirsuta "— Robynsiophyton vanderystii Lotononis hirsuta 1 100 1—• Crotalaria dislans Crotalaria griquensis 100 52 2 Crotalaria capensis 1 co 100 iLLE90 Crotalaria virgultalis 1 94 Crotalaria humilis 1 Cmtalaria lanceolata C3 Bolusia amboensis 1 Dichilus gracilis Genisteae

FIG. 3.4 (Continued).

31 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

The low resolution within the Crotalarieae based on morphology and some chemical and cytological characters reflects the somewhat reticulate relationships within the tribe (also mentioned by Van Wyk 1991a; Van Wyk 2005) with remarkable examples of parallelism, convergence and analogy with regard to vegetative and reproductive morphology as outlined by Dahlgren (1970a). The presence of seemingly identical apomorphic states in unrelated genera (e.g. hairy petals in Lebeckia section

Calobota and Lotononis section Leptis) complicates the cladistic analysis. In the absence of single apomorphies with diagnostic value, natural groups can only be delimited using combinations of characters. The results presented here (both molecular and morphological) support some of the current generic concepts within the tribe, but provide novel insights into as yet phylogenetically unstudied groups such as Lebeckia and the smaller genera of the Crotalarieae, namely Robynsiophyton, Rothia and

Spartidium.

Previous studies on relationships within the Crotalarieae have all suggested a close relationship between the "Cape" genera (viz., Aspalathus, Lebeckia, Rafnia and

Wiborgia), as well as the genera that have a zygomorphic calyx (viz., Lotononis,

Pearsonia, Robynsiophyton and Rothia; Dahlgren 1963, 1967a, 1970a; Polhill 1981;

Van Wyk 1991a). The placement of Bolusia, Crotalaria and Spartidium has been uncertain. A sister relationship between the former two genera was suggested by Polhill

(1976) and Van Wyk (1991a), while Spartidium was thought to either be closely allied to

Genisteae or to Lebeckia, but the affinities remained unclear (Polhill 1976). The results of the current study indicate four major lineages within the Crotalarieae, namely the

"Cape group" (Aspalathus, Lebeckia, Rafnia, Wiborgia, including Spartidium saharae),

32 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE two lineages comprising genera with zygomorphic calyces (Lotononis, Pearsonia,

Robynsiophyton and Rothia), plus Bolusia, Crotalaria and Lotononis section Euchlora.

3.3.2 The "Cape group"—The lack of resolution between genera in this group is caused by the low sequence divergence among these taxa. Aspalathus and Rafnia are both strongly supported as monophyletic and indicated as sister taxa by the analysis of morphological data alone because they share sessile leaves and an asymmetrical upper suture of the pod (Campbell and Van Wyk 2001). Aspalathus is placed closer to Wiborgia by both the MP analyses (combined ITS/rbcL and combined

ITS/rbcUmorphology) and BI analysis (combined ITS/rbcL). Although the sample of

Aspalathus is not at all comprehensive, some clades of interest can be noted.

Aspalathus linearis and A. pendula both have simple, terete leaves and this Glade corresponds to the Lebeckiiformes group (Dahlgren 1988), named for its similarity to

Lebeckia section Lebeckia. Aspalathus laricifolia, A. hirta, A. hystrix and A. shawii all have spine-tipped leaves (corresponding to the Laterales group of Dahlgren 1988) and form a Glade, which is well-supported in the BI tree (combined ITS/rbcL). The

Borboniae group (Dahlgren 1988) represented by A. perfoliata and A. crenata form a weak to strongly supported Glade. Within Rafnia, two clades can be identified that correspond to the two section of the genus described by Campbell and Van Wyk

(2001): the R. acuminata Glade representing R. section Rafnia, and the R. spicata Glade representing R. section Colobotropis E.Mey. These two sections differ mainly in the rostrate keel petals and unequal calyx lobes of R. section Colobotropis.

It is clear from both the MP (combined ITS/rbcL and combined

ITS/rbcUmorphology) and BI analysis (combined ITS/rbcL) that Lebeckia and Wiborgia

33 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE are not monophyletic. Lebeckia is represented by several smaller clades within the

"Cape group", while Wiborgia humilis is nested within the Lebeckia leipoldtiana Glade and well-separated from the main Wiborgia clade.

The species of Lebeckia section Calobota form a well-supported Glade with those of L. section Stiza and the monotypic genus Spartidium. These species are all shrubs with trifoliolate or unifoliolate leaves and green mature stems. The petals are usually pubescent and they possess a 4+1+5 anther configuration (five short dorsifixed anthers, four long basifixed anthers and an intermediate carinal anther). Some of these character states also occur in Lotononis species and are thus not informative in the morphological analysis. The species of L. section Stiza and L. section Calobota occur in the Cape and those of the latter also extend northwards into Namibia. Spartidium saharae is a North African plant that has had uncertain affinities within the tribe. The close relationship of this species to L. section Calobota has also been demonstrated independently by Edwards and Hawkins (2007) based on ITS data.

Wiborgia and species of Lebeckia section Viborgioides are superficially very similar in their rigid, woody habit, glabrous petals and highly reflexed standard petals in most species. However, they differ markedly in the winged, few-seeded samaras that are typical of Wiborgia species (Dahlgren 1975). However, Wiborgia humilis has thin- walled, inflated pods that lack the dorsal wing characteristic of Wiborgia (Dahlgren

1970b). The fruits of Lebeckia section Viborgioides are typically turgid, thin-walled, many-seeded and wingless. Wiborgia humilis is the only species of Wiborgia with a 4+6 anther arrangement (6 short, dorsifixed and 4 long basifixed anthers) instead of the

34 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

4+1+5 arrangement found in all other species. These characters therefore support the placement of W. humilis in the L. section Viborgioides clade.

Lebeckia section Lebeckia is unresolved in the BI and MP analyses (combined

ITS/rbcL). This section is morphologically very distinct from the other sections in the genus and its monophyly is supported by four apomorphies, namely the suffrutescent habit, acicular (terete) leaves, a 5+5 anther arrangement and rugose seeds (Le Roux

2006; Le Roux and Van Wyk 2007). This is clear from the moderate support this group received in the morphological analysis as well as the combined morphological/molecular analysis. Lebeckia lotononoides and L. inflata are the only other species of Lebeckia with rugose seeds, but these species share characters with

L. section Calobota and L. section Viborgioides, respectively (Boatwright and Van Wyk

2007; Boatwright, pers. obs.).

The positions of both L. inflata and L. mucronata are unresolved in the combined

ITS/rbcL analysis. Lebeckia mucronata is currently placed within L. section Calobota.

Lebeckia inflata was described by Bolus (1887) without any reference to its position in the infrageneric systems of Bentham (1844) and Harvey (1862). Both of these species, however, share several anatomical and morphological characters with species of L. section Viborgioides, such as a 4+6 anther arrangement, glabrous petals and the dorsiventral leaves with mucilage cells. When morphological characters were combined with the molecular ITS and rbcL data, L. inflata was included in a Glade along with L. section Viborgioides and Wiborgia humilis, while the position of L. mucronata remained unresolved.

35 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

3.3.3 The Lotononis clades—Based on the molecular data, Lotononis is polyphyletic and consists of three clades. Lotononis s.s. groups with the Pearsonia clade (albeit without support in the combined ITS/rbcL MP analysis) and is strongly supported as monophyletic. Two groups were noted within this Glade. The first consists of representatives of L. section Oxydium and the second of representatives of L. sections Aulacinthus, Buchenroedera, Cleistogama, Krebsia, Lotononis, Monocarpa and Polylobium. Lotononis section Oxydium is strongly supported to be monophyletic in the molecular analyses and currently consists of 35 species (accommodated in 14 subsections as described by Van Wyk 1991b) distinguished from other sections by several characters, including single stipules at each leaf base and glabrous wing and keel petals. In the second group, L. section Cleistogama is sister to L. section

Monocarpa, while L. sections Aulacinthus, Lotononis and Polylobium form a Glade without support. The resolution in the rest of the group is low and relationships not clear. In the molecular analysis, Lotononis s.s. and the Pearsonia Glade clearly share similarities in their sequences but morphologically Lotononis s.s. allies with the rest of the genus to form a monophyletic assemblage. When the molecular data were combined with the morphology, Lotononis remained monophyletic, although with moderate support. It is interesting to note the close agreement between the sectional classification proposed by Van Wyk (1991b) and the results of this study.

Pearsonia, Robynsiophyton and Rothia share several morphological characters that are unique among the genera of Crotalarieae, such as the uniform anthers, straight styles and presence of angelate esters of hydroxylupanine. The generic concepts of

Robynsiophyton and Rothia, based mainly on anther characters, have been doubtful

36 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE and Robynsiophyton was thought to be a local derivative of Pearsonia (Van Wyk

1991a). Taxonomic studies of the former two genera and a thorough examination of their morphology and anatomy revealed that these generic concepts are indeed sound and emphasizes the value of androecial characters in the Crotalarieae (Chapter 10;

Boatwright et al. 2008c; Chapter 9; Boatwright and Van Wyk 2009; see Appendices B4 and B5).

The second Glade comprises those species from Lotononis section Leptis,

Lotononis section Listia and allies. This Glade is moderately to strongly supported in the molecular analyses but lacks resolution in the morphological analysis. The concept of a monotypic Listia was broadened by Van Wyk (1991b) and treated as a section of

Lotononis with several distinct characters. This broadened concept is supported in the analyses presented in this study. The absence of suitable material of the rare Lotononis macrocarpa Eckl. & Zeyh. (an anomalous species within this group) prevented its inclusion in this study. See Chapter 8, where L. macrocarpa has indeed been included in a reassessment of Lotononis s.l. Van Wyk (1991b) mentions that no apomorphies exist for L. section Leptis, which is clear from the morphological analysis where this section is left unresolved. The molecular results, however, indicate a close relationship between this section and L. sections Digitata, Leobordea, Lipozygis and Synclistus, all of which are successively sister to L. section Listia. The third Glade comprises L. hirsuta, which groups with the Crotalaria clade.

3.3.4 The Crotalaria Glade—The close relationship between Crotalaria and

Bolusia has been mentioned by both Polhill (1982) and Van Wyk (1991a) and these authors suggested that Bolusia could merely be a local derivative of Crotalaria. Data

37 CHAPTER 3: PHYLOGENETIC RELATIONSHIPS WITHIN THE TRIBE CROTALARIEAE

from the present study indicates that Bolusia is sister to and not embedded within

Crotalaria (rbcL analysis with multiple representatives), so that its generic status seems

justified. Bolusia differs from Crotalaria mainly in its glabrous style and strongly coiled

keel petals. Also included in this Glade is Lotononis hirsuta, representing the monotypic

section Euchlora. This species differs from all other Lotononis species in its peculiar

tuberous habit and from most others in the very large, inflated pods, equally lobed calyx

and very large seeds with smooth surfaces. It shares with Crotalaria and Bolusia the

strongly inflated pods.

3.3.5 Implications for generic classification —The results generated in this

study have very important implications for the generic classification system of the

Crotalarieae. Detailed studies of character variation in the tribe over a period of more

than 23 years are now nearing completion, allowing an informed evaluation of the

congruence between morphological and molecular patterns. The results clearly show

that the genera Calobota Eckl. & Zeyh. and Euchlora Eckl. & Zeyh. should be

reinstated. A new genus should be described that will include Lebeckia sect.

Viborgioides, as well as L. inflata and L. mucronata. Spartidium will be transferred to

Calobota and Wiborgia humilis to the new genus (Chapters 5, 6 and 7; Boatwright et al.

submitted). A new generic classification should be devised for Lotononis given the

polyphyly of the genus based on molecular evidence and also the lack of generic

apomorphies for Lotononis s.l. if L. section Euchlora is excluded (many of these co- occur in the latter). These changes are discussed in Chapter 5 and Chapter 8 and will clearly result in a more practical and predictable generic classification system for the tribe Crotalarieae.

38 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

4.1 INTRODUCTION

The distribution patterns of secondary metabolites in plants, although sometimes erratic, have the potential to provide valuable insight into phylogenetic

relationships. This is especially true for the Fabaceae and more specifically the

Papilionoideae, where chemical patterns are often in agreement with other evidence

(Gomes et al. 1981; Van Wyk 2003b; Wink 2003). Some classes of compounds have been found to be more useful markers than others, such as alkaloids and flavonoids in legumes. Several syntheses of the available data and literature on alkaloids and flavonoids of papilionoid legumes are available (e.g. Harborne 1969,

1971; Mears and Mabry 1971; Salatino and Gottlieb 1980; Gomes et al. 1981;

Kinghorn and Smolenski 1981; Kinghorn et al. 1982; Hegnauer and Grayer-

Barkmeijer 1993; Southon 1994; Hegnauer and Hegnauer 1992-2001). This chapter aims to briefly review the chemotaxonomic value of alkaloids, flavonoids and other taxonomically useful compounds (cyanogenic glucosides and non-protein amino acids) in the Crotalarieae.

4.2 ALKALOIDS OF THE TRIBE CROTALARIEAE

Detailed studies of alkaloids in the Crotalarieae have made it possible to thoroughly evaluate their distribution patterns (e.g. Aspalathus — Van Wyk and

Verdoorn 1989a; Lebeckia — Van Wyk and Verdoorn 1989b; Lotononis — Van Wyk and Verdoorn 1988, Van Wyk and Verdoorn 1989c, Verdoorn and Van Wyk 1992;

39 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

Pearsonia — Van Wyk and Verdoorn 1989d, Verdoorn and Van Wyk, 1990; Van Wyk and Verdoorn 1991a; Rafnia — Van Wyk and Verdoorn 1989a; Robynsiophyton —

Van Wyk and Verdoorn 1991b; Rothia — Hussain et al. 1988; Spartidium — Van Wyk et al. 1989; Wiborgia — Van Wyk and Verdoorn 1989a) and their taxonomic value was discussed by Van Wyk et al. (1988) and Van Wyk and Verdoorn (1990). The chemical structures of all the various alkaloids are presented in Van Wyk and

Verdoorn (1990). The Podalyrieae and Crotalarieae differ from all other genistoids in the absence of a-pyridone alkaloids and from most other tribes in the accumulation of hydroxylated lupanines and their esters, especially in their seeds (Van Wyk

2003b). Within the Crotalarieae, three groups can be distinguished based on alkaloid data (Van Wyk and Verdoorn 1990; Van Wyk 1991a):

An unspecialised group without a-pyridone alkaloids and without esters of alkaloids (Aspalathus, Lebeckia, Rafnia, Wiborgia and Spartidium, the so-called

"Cape group").

A group specializing in lupanine-type esters of alkaloids (Pearsonia,

Robynsiophyton and Rothia, i.e. the Pearsonia Glade)

A group with macrocyclic pyrrolizidine alkaloids (Crotalaria and Lotononis).

These groups are largely consistent with those retrieved by phylogenetic studies on the Crotalarieae based on molecular (nrITS and rbcL) and morphological data. The genera of the "Cape group" form a monophyletic, although moderately supported Glade while the Pearsonia Glade receives strong support (Chapter 3;

Boatwright et al. 2008a), as is also indicated by the chemical data.

The presence of macrocyclic pyrrolizidine alkaloids in Crotalaria and

Lotononis is unique within the Crotalarieae and was thought to support a sister relationship between these genera (Van Wyk and Verdoorn 1990; Van Wyk 2003b).

40 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

It is now clear, however, that the occurrence of these alkaloids are rather the result of convergence as these genera do not share a sister relationship (see Chapters 3 and 8; Boatwright et al. 2008a). Quinolizidine alkaloids, which are present in most of the other genera of the Crotalarieae, are absent from Crotalaria, but present in some species of Lotononis, although they do not co-occur with pyrrolizidine alkaloids in the same species (Van Wyk and Verdoorn 1990). The pattern of alkaloids generally supports the new generic classification system for Lotononis as discussed in

Chapter 8. Lotononis s.s. appears to accumulate only pyrrolizidine alkaloids, while the sections Listia, Leptis and related groups have quinolizidine alkaloids, albeit often only in trace amounts. The presence of pyrrolizidine alkaloids also supports the sister relationship between the Crotalarieae and Genisteae as these are the only tribes that have genera bearing pyrrolizidine alkaloids (Van Wyk 2003b).

Van Wyk and Verdoorn (unpublished) studied alkaloids in the leaves of some

Crotalarieae genera and compared them to alkaloids extracted from the seed of the same species. They observed large quantitative differences in the alkaloid profiles of the leaves and seeds and found that seeds usually contain only a single or a few major alkaloids and are therefore conservative taxonomic characters. The African

Genisteae genera Argyrolobium, Dichilus, Melolobium and Polhillia differ from the

Crotalarieae in accumulating ammodendrine, lusitanine, cytisine, N-methylcytisine and camoensine in their seeds, as opposed to lebeckianine, 3p-hydroxylupanine and 13a-hydroxylupanine accumulated by Aspalathus, Lebeckia, Lotononis and

Pearsonia. Species of Crotalaria and Lotononis s.s. were again unique in accumulating macrocyclic pyrrolizidine alkaloids. A detailed and rigorous study of seed alkaloids of all genera of the Crotalarieae will clearly yield interesting information and will contribute towards a better understanding of how alkaloids

41 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE evolved within the tribe. It is clear that the extreme alkaloidal diversity found in

Lotononis s.l. by Van Wyk and Verdoorn (1988, 1989c) and Verdoorn and Van Wyk

(1992) can now be newly interpreted in relation to the phylogeny as revealed by molecular evidence (see Chapter 8).

4.3 FLAVONOIDS OF THE TRIBE CROTALARIEAE

Flavonoids in legumes and within the Crotalarieae, have not been as well studied as alkaloids, possibly due to the somewhat erratic patterns found in flavonoid data which limits their utility (Harborne 1971; Van Wyk 2003b). The occurrence of isoflavonoids in the Papilionoideae and the absence of a hydroxyl group in position five in all legumes are useful characters from flavonoid data

(Hegnauer and Grayer-Barkmeijer 1993; Van Wyk 2003b). Again seed flavonoids appear to be more conservative than those of the leaves (Hegnauer and Grayer-

Barkmeijer, 1993; De Nysschen et al. 1998).

De Nysschen et al. (1998) showed that the Crotalarieae and the Podalyrieae have the same four major seed flavonoids (butin, 3'-hydroxydaidzein, orobol and vicenin-2; chemical structures presented in De Nysschen et al. 1998), but none of these compounds are present in African Genisteae. No other study has explored flavonoid patterns within the Crotalarieae, which represents an obvious topic for future investigation.

Information on anthocyanins in legumes is rather poor (Harborne 1971). Van

Wyk et al. (1995) showed in the Podalyrieae that anthocyanins were taxonomically informative, with each of the pink- or purple-flowered genera of the tribe having a diagnostic combination of floral anthocyanins (cyanidin and/or peonidin and esters

42 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE of these pigments). The Cape genus Hypocalyptus Thunb. was shown to be unrelated, as it accumulates methylated anthocyanins in its flowers (Van Wyk et al.

1995). However, no information is available for the Crotalarieae. Notable variation in flower colour, with the sporadic occurrence of pink, blue, white or purple flowers, is seen only in Aspalathus, Crotalaria, Pearsonia and Lotononis. However, pink and blue flowers are rather rare in these genera, so that anthocyanins have a limited potential from a chemosystematic point of view. It may nevertheless be interesting to investigate the homology of blue/white/purple/pink flowers in Aspalathus, Pearsonia,

Lotononis and Crotalaria, as these four genera are now known to represent four different lineages within the tribe (see Chapter 3).

4.4 OTHER USEFUL COMPOUNDS

In addition to alkaloids and flavonoids other classes of compounds also provide useful systematic information. Two of these are cyanogenic glucosides and non-protein amino acids. The occurrence of cyanogenic glucosides in legumes was summarized by Seigler et al. (1989). These compounds occur widely in various tribes of legumes, but in the tribe Crotalarieae they are restricted to the genus

Lotononis (Van Wyk 1989; Van Wyk and Whitehead 1990). In Lotononis the cyanogenesis is due to the glucoside prunasin (Van Wyk 1989) and esters thereof

(Seigler et al. 1989). Variation in this character within Lotononis is apparent as some infrageneric groups are acyanogenic (Van Wyk 1991b). This is, however, generally a useful character to distinguish Lotononis from morphologically similar genera, such as Pearsonia and Rothia. It is noteworthy that cyanogenesis is restricted to the one major Glade in Lotononis (sections Oxydium, Monocarpa, Cleistogama,

43 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

Polylobium, Lotononis, Aulacinthus and Buchenroedera) and that it is absent from the sections Listia, Leptis and related sections. This was an important character to support the infrageneric classification system proposed by Van Wyk (1991b) and now becomes highly relevant in the context of new generic delimitations in

Lotononis. Cyanogenesis is a useful diagnostic character for a narrowed generic circumscription of Lotononis (see Chapter 8).

Non-protein amino acids are present in Crotalaria (Pilbeam and Bell 1979).

This character appears to be unique to Crotalaria within the Crotalarieae but the compounds are widely distributed in other papilionoid tribes as well as in the

Mimosoideae and Caesalpinioideae. Wink (2003) and Wink and Mohamed (2003) mapped the distribution of these and other chemical compounds onto phylogenetic trees derived from DNA sequence data to demonstrate their systematic significance.

The high congruence between molecular systematic data and chemical data is noteworthy. The authors speculate that the presence of these compounds as chemical defence or signal compounds are important for plant survival, but are subject to evolution and selection, thereby complicating their interpretation in a phylogenetic context.

4.5 THE USE OF CAPE LEGUMES AS SOURCES OF TEA

The presence of several beneficial compounds in the leaves of legume species has made them popular as traditional sources of tea. These teas have historically been well-documented by several authors under numerous common names, usually describing a property of the tea or the distribution of the source plant. Some of the earliest accounts are those of Thunberg during his travels from

44 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

1772-1775 (Forbes 1986), Pappe (1847, 1857) and Marloth (1925). The species utilized are mainly from three genera of the Crotalarieae and Podalyrieae, namely

Aspalathus, Cyclopia and Rafnia. Some of these herbal teas are already produced commercially such as rooibos- and honeybush tea (Van Wyk et al. 1997; Van Wyk and Gericke 2000; Van Wyk and Wink 2004). These teas are popular mainly because they do not contain caffeine or harmful stimulants and the phenolic constituents are thought to have beneficial properties (Van Wyk et al. 1997; Joubert et al. 2008). The major phenolic compounds of rooibos- and honeybush tea have already been the focus of several studies (e.g. De Nysschen et al. 1996; Van

Heerden et al. 2003). A detailed review of rooibos tea (Aspalathus linearis) and honeybush tea (Cyclopia spp.) was done by Joubert et al. (2008). Teas obtained from Rafnia are tannin-rich and black in colour due to the high concentration of unknown phenolic compounds (Van Wyk and Gericke, 2000). Unlike the related

Aspalathus, there is very little known about its flavonoid chemistry. These legumes are important not only as beverages, but also for various medicinal applications. A short review of the species traditionally used and their common names, medicinal uses and correct taxonomic identities is presented here.

The common- and scientific names of the sources of tea are listed in Table 1, along with the various authors who documented their uses. Thunberg during his travels of the Cape in 1770-1775 mentioned the use of the leaves of Borbonia cordata as a source of tea. This plant is an old Cape remedy and was used for respiratory ailments such as respiratory catarrh, whooping cough and bronchitis

(Pappe 1847; Kling 1923; Watt and Brandwijk 1962). A decoction of the leaves of

Rafnia acuminata (recorded as Crotalaria perfoliata, Vascoa perfoliata or Rafnia perfoliata) was claimed to be a powerful diuretic and was considered a cure for

45 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE

dropsy (Pappe 1847, 1857; Watt and Breyer-Brandwijk 1962; Forbes 1986). Kling

(1923) recommended various teas for an array of respiratory ailments (including

bronchitis and coughs), among these 'heuningtee' and 'hottentotstee' (both teas

from species of Cyclopia), `Stekeltee' (made from Aspalathus species) and

`boesmanstee' (a tea brewed from Rafnia amplexicaulis; Anonymous 2001).

`Stekelthee' was also said to have diuretic properties, but was primarily used to treat

asthma and hydrothorax, whereas 'honigthee/heuningtee' was considered to be a

restorative and was used in the treatment of chronic catarrh or consumption (Pappe

1847, 1857; Speight 1931).

Marloth (1925) in his The Flora of South Africa' applied the term 'bush tea' to

the tea yielded by Cyclopia species. There is a difference in the teas produced in

various districts as different species of Cyclopia are used (Hofmeyr and Phillips

1922; Marloth 1925; Kies 1951). Other tea sources mentioned by Marloth (1925) is

the `stekeltee' (Borbonia cordata) and `rooibostee' (naaldtee) which are produced

from Aspalathus corymbosa (now Aspalathus linearis) that below a certain altitude

turns black when fermented, thus yielding 'black tea'.

The leaves of some Psoralea species, another largely Cape legume genus,

are aromatic due to pellucid glands on the leaves and yielded what has been called

`schaapbos tee'. Marloth (1925) and Smith (1966) mentioned P. bracteata as the

species utilized, but other authors attributed it to P. decumbens (Watt and Breyer-

Brandwijk 1962; Rood 1994). No apparent medicinal properties were mentioned for this tea.

Marloth (1925) gave an account of the process by which 'bush teas' are

prepared: "The young twigs are gathered when in flower and submitted to a process of fermentation or sweating by piling them up in a heap. They are then dried in the

46 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE sun, and if this is done with care and expedition, the tea possesses a sweet aroma".

This process is similar to the commercial process applied to rooibos and honeybush tea. Hofmeyr and Phillips (1922) described a second method used in the preparation of 'bush teas': "...for local consumption is to heat the leaves in an oven for about an hour to produce 'sweating,' after which they are dried in the sun. When thoroughly dry the leaves and smaller twigs are removed from the stem and larger branches, packed, and sold under various names." They also gave a summary of the commercial varieties of the Cyclopia species, which was slightly modified by Kies

(1951) and is summarised in Table 1. Kies (1951) in her revision of Cyclopia mentioned Aspalathus contaminatus and A. tenuifolia as alternative sources of 'bush tea'. Both these species are synonyms of Aspalathus linearis, which is today used to produce rooibos tea (Dahlgren 1988). Dahlgren (1968) described the processing of cultivated rooibostee as follows: "The leafy branch ends are harvested by cutting and when brought to the fermentation yard cut again finely to a length of ca. 0.5 cm, moistened, bruised (e.g. with wooden hammers), and placed in heaps to ferment.

Then the material is spread open in the sun and stirred intermittently until dry. After drying it is sifted, classified, and packed." The preparation of honeybush tea is much the same and Cyclopia genistoides and C. intermedia are the main species used

(Van Wyk et al. 1997; Van Wyk and Gericke 2000; Van Wyk and Wink 2004;

Joubert et al. 2008).

The chemical compounds in rooibos tea and other herbal teas were reviewed by Joubert et al. (2008). The presence of aspalathin (a chalcone C-glycoside) as the dominant phenolic compound in commercial rooibos tea is well known (Koeppen and Roux 1965). However, Van Heerden et al. (2003) showed that the species is chemically complex, with several other phenolic compounds present as major

47 CHAPTER 4: ALKALOIDS AND FLAVONOIDS OF THE TRIBE CROTALARIEAE compounds in various wild populations of the species. The closely related A. pendula R.Dahlgren had rutin as major constituent. Preliminary work by Campbell

(1998) showed that Rafnia species accumulate C-glycosides, apparently similar to those of Aspalathus linearis. It may be of considerable chemotaxonomic interest to explore the main patterns of flavonoids (especially in the "Cape group" of genera) since, apart from Aspalathus linearis, almost nothing appears to be known for any of the genera and species in the tribe Crotalarieae.

48 • • •

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Vle z zzzzz z CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

5.1 INTRODUCTION

The current broad generic concept of Lebeckia Thunb. dates back to

Bentham (1844) and Harvey (1862) and refers to a group of ca. 36 species of papilionoid legumes that occur mainly in the southern and western parts of South

Africa, with some extending into Namibia. The group is particularly common in the

Cape Floristic Region (CFR). Bentham (1844) reduced several genera described by

Meyer (1836) and Ecklon and Zeyher (1836) to the synonymy of an expanded

Lebeckia. This broadened concept included Meyer's Stiza and Sarcophyllum, together with Ecklon and Zeyher's Acanthobotlya and Calobota. A new sectional classification was proposed, based mainly on the shape of the keel and the morphology of the fruit. This comprised five sections, viz. section Calobota, section

Eulebeckia Benth., section Phyllodiastrum Benth., section Stiza and section

Viborgioides. Harvey (1862) followed this sectional classification of Lebeckia for his treatment in the Flora Capensis. For nearly 150 years, the generic concept and relationships between the morphologically rather diverse species were never studied in depth. A revision of Lebeckia s.s. (sections Eulebeckia and Phyllodiastrum) was recently completed by Le Roux and Van Wyk (2007, 2008, 2009). Their results show that the 14 species of Lebeckia s.s. can easily be distinguished by their acicular leaves and 5+5 anther arrangement, not only from all other species of the so-called

"Cape group" of the tribe Crotalarieae (i.e. Aspalathus, Rafnia and Wiborgia) but also from all other species hitherto included in Lebeckia.

52 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

Spartidium is a monotypic genus that occurs in North Africa. The affinities of the genus within the genistoid legumes have been unclear for some time as a possible relationship with both Retama Raf. and Lebeckia has been suggested

(Polhill, 1976). The genus is currently placed within the Crotalarieae based on the open androecial sheath, but an evaluation of its systematic position is clearly desirable. Polhill (1981) considered Spartidium to be "virtually indistinguishable from

Lebeckia". He used the orientation of the seeds in the fruit and the North African distribution as the only key characters to distinguish Spartidium from other genera.

In a broad study of molecular (ITS, rbcL) and morphological data of 117 species representing all the genera of the Crotalarieae, Boatwright et al. (2008a;

Chapter 3) discovered that Lebeckia is polyphyletic (Fig. 5.1a and 5.1b). This study revealed important new relationships within the Crotalarieae as well as Lebeckia and showed the need for new generic circumscriptions. The species can be readily accommodated in three easily recognizable morphological groups (genera) as was proposed by Boatwright et al. (submitted; see Chapters 6 and 7): (1) Lebeckia s.s.

(L. section Lebeckia, including sections Phyllodiastrum and Eulebeckia); (2)

Calobota [L. section Calobota and L. section Stiza, together with the monotypic

North African Spartidium] and (3) Wiborgiella [L. section Viborgioides, together with

L. inflata, L. mucronata and Wiborgia humilis]. Although the position of Lebeckia mucronata is unresolved it shares many morphological and anatomical characters with the other species placed within Wiborgiella and is thus included in this group.

In this chapter, new morphological, anatomical and molecular systematic evidence is presented to show that Bentham's (1844) broad concept of Lebeckia is polyphyletic.

CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

100 E Aspatathus linens 1 00 Aspalathus iinearis 1 Aspalathus pendula CAspakehus pendula 1 0 Aspalathus &wrens 2 Aspalathus samosa Aspalaaws samosa 84 Aspalathus galeata Aspalathia

FIG. 5.1 The "Cape group" taken from (a) a strict consensus tree of the combined analysis of

ITS and rbcL data (no. trees = 560; tree length = 1473 steps; consistency index = 0.50;

retention index = 0.86); and (b) a strict consensus tree of the combined analysis of

molecular (ITS and rbcL) and morphological data for the tribe Crotalarieae from Chapter 3

(Boatwright et al. 2008a; no. trees = 370; tree length = 1166 steps; consistency index =

0.53; retention index = 0.84). Bootstrap percentages are given above the branches and

Bayesian posterior probabilities below the branches. Grey dotted lines indicate alternative

topologies in the Bayesian analysis.

54 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

5.2 MORPHOLOGICAL AND ANATOMICAL EVIDENCE FOR THE PARAPHYLY OF LEBECKIA

5.2.1 Habit and branches—Most genera of the Crotalarieae have a shrubby habit

(Polhill 1976), but variation in habit is quite pronounced within Lebeckia. Species of

Lebeckia section Lebeckia are predominantly suffrutescent plants that branch mainly at ground level, whereas the species of the remaining sections are almost invariably woody shrubs. A strongly spinescent habit is characteristic of Lebeckia section Stiza but also of two species of section Calobota, viz. L. acanthoclada and L. spinescens. Both sections Calobota and Stiza have green young branches (except

L. acanthoclada), as opposed to the woody, rigid, ramified brown to greyish branches of section Viborgioides (similar to the genus Wiborgia). In species with green stems [sections Calobota, Lebeckia and Stiza, and Spartidium saharae] a large part of the cortex is composed of chlorenchyma which serves a photosynthetic function (Metcalfe and Chalk 1950). This layer is absent in species of section

Viborgioides and also L. mucronata.

5.2.2 Leaves—Leaves are extremely variable, ranging from trifoliolate and petiolate (sections Calobota and Viborgioides), to unifoliolate (section Stiza) or simple (section Calobota and Spartidium) to completely phyllodinous (and acicular) in section Lebeckia. Cultivated plants of the section Stiza showed that the juvenile leaves are petiolate and trifoliolate, but a loss of the lateral leaflets and shortening of the petioles result in unifoliolate leaves on the older branches, as was mentioned by

Polhill (1976) and Dahlgren (1970a). This transition is also seen in L. obovata

Schinz. Stipules are absent in all but two species of Lebeckia s.l.: L. wrightii and L. uniflora M.M.le Roux and B.-E.van Wyk (Le Roux and Van Wyk 2009).

55 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

Transverse sections through the leaves of Lebeckia species (Fig. 5.2) revealed a remarkable difference between the species of Calobota/Stiza/Spartidium on the one hand (hereafter called the Calobota group) and Viborgioides/L. inflatalL. mucronata/Wiborgia humilis on the other hand (hereafter called the Viborgioides group), namely that the former group has isobilateral leaves, while the latter has dorsiventral leaves. This difference was unexpected, as anatomical characters are generally regarded as conservative. Even more remarkable was the exact congruence between the anther arrangements of the two groups (respectively

4+1+5 and 4+6 — see later). Leaves of Lebeckia s.s. are completely different from the species mentioned above, as they are phyllodinous, acicular and terete in transverse section.

Another interesting difference between the Calobota group and the

Viborgioides group is the presence of mucilage cells in the latter but not the former

(Fig. 5.2). In this character, the Viborgioides group agrees with other Cape genera

(Aspalathus, Rafnia and Wiborgia, as well as the predominantly Cape Lebeckia s.s.). In contrast, the Calobota group, which extends into arid regions (Northern

Cape and Namibia), does not have mucilage cells in mature leaves.

56 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

FIG. 5.2 Transverse sections through the leaves of Lebeckia species, showing isobilateral

(a, b, Calobota group) and dorsiventral (c, e, f, Viborgioides group) leaflet laminas and an

acicular, terete, phyllode with a circular arrangement of palisade cells (d, Lebeckia s.s.).

Note the presence of mucilage cells in the epidermis. (a) L. cytisoides; (b) L. pungens; (c) L.

sessilifolia; (d) L. sepiaria; (e) L. inflata; (f) L. mucronata.

Voucher specimens: (a) Boatwright et al. 114 (JRAU); (b) Boatwright et al. 106

(JRAU); (c) Van Wyk 2120 (JRAU); (d) Le Roux et al. 10 (JRAU); (e) Johns 162 (JRAU); (f)

Vlok 1726 (JRAU). Scale bar = 0.2 mm.

57 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

The North African Spartidium, however, has mucilage cells in the epidermis, as do immature leaves of L. pungens, suggesting that there is a loss of these cells as the leaves mature (Boatwright, pers. obs.). Mucilage cells are widely distributed throughout the various plant families and several authors have speculated about their function (Gregory and Baas, 1989). Although experimental evidence is lacking, mucilage cells are thought to aid in water storage and reduction of transpiration, protection against intensive radiation and also against herbivory (Gregory and Baas,

1989). Mucilage cells are often associated with plants that occur in Mediterranean climates (Van der Merwe et al. 1994). Bredenkamp and Van Wyk (1999) speculated that in Passerina L. (Thymelaeaceae) the mucilage serves as a regulator of hydration in the leaves, protecting them against water loss, but also helping to accumulate reserve water in the leaves. Lebeckia s.s. and the Viborgioides group are restricted to the CFR and therefore diversifying in a Mediterranean climate could explain the presence of the mucilage cells in these taxa. It is interesting to note the presence of mucilage cells in Spartidium saharae, which also occurs in a

Mediterranean climate. The Calobota group (excluding Spartidium) extend out of the

CFR into summer rainfall, more arid regions and show a different adaptation to drought, viz. isobilateral leaves (Fig. 5.2 a, b; also in Spartidium). Van der Merwe et al. (1994) mention that the presence of more layers of palisade parenchyma improves the transport of water through the leaves and also offers increased protection to the chloroplasts. In contrast, the leaves of species within the

Viborgioides group are dorsiventral (Fig. 5.2 c, e, f). In Lebeckia s.s., the acicular leaves have a complete circular zone of palisade cells, with no spongy parenchyma

(Fig. 5.2 d).

58 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

The petioles are always shorter than the leaflets in the Viborgioides group. In this group the persistent petioles become hard and woody when the leaflets are shed, contributing to the rigid and somewhat thorny appearance of the branches.

These characters are also found in some species of Wiborgia s.s. (Dahlgren 1975).

In the Calobota group, the petioles are either longer or shorter than the leaflets and sometimes persistent, but never rigid and spinescent. The acicular leaves of most species of Lebeckia s.s. are articulated or "jointed" near the middle or reduced to petioles in species with unarticulated leaves, i.e. the leaves are phyllodinous

(Dahlgren 1970a). This distinctive character serves as a convincing synapomorphy for this genus in its narrow circumscription.

5.2.3 Inflorescences—As in most Crotalarieae, the inflorescences in

Lebeckia are all terminal, multi-flowered racemes, varying in length and number of flowers. In Lebeckia s.s., the inflorescences may be relatively long and are often densely flowered with up to 93 flowers per raceme, for example in L. brevicarpa (Le

Roux and Van Wyk 2007). Species of section Calobota generally have fewer flowers per inflorescence, except in L. melilotoides Dahlgr., where more than 100 flowers are found on the elongated spikes (Dahlgren 1967b). The racemes of the three species of section Stiza are characteristically spine-tipped. Very short and few- flowered inflorescences are found in L. section Viborgioides, with the flowers often solitary in L. bowieana Benth.

5.2.4 Flowers—The calyx structure is often an important generic character in the Crotalarieae. Lebeckia species normally have equally lobed or "lebeckioid" calyces (Polhill 1976), as opposed to the zygomorphic calyx ("lotononoid") in

Lotononis (DC.) Eckl. and Zeyh. and Pearsonia Dlimmer or the bilabiate calyx typical of all members of the tribe Genisteae (Van Wyk 1991b; Van Wyk and Schutte

59 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

1995). In the Calobota and Viborgioides groups, the carinal lobe is slightly narrower than the upper and lateral lobes, whereas in Lebeckia s.s. it is equal to the other lobes (Fig. 5.3). In the former two groups, the calyces are pubescent or at least glabrescent as opposed to the usually glabrous calyces found in section Lebeckia

(L. wrightii and L. uniflora are exceptions in having a sparsely pubescent calyx).

The pubescence of the petals, the shape of the keel petal and the arrangement of the anthers closely follow the three major groups within Lebeckia s.l.

Glabrous petals are found in Lebeckia s.s. and the Viborgioides group, whereas species of the Calobota group generally have pubescent petals (or at least pilose along the dorsal midrib of the standard petal). Lebeckia macrantha and L. psiloloba are the only exceptions and have totally glabrous petals. The keel petals in Lebeckia s.s. are characteristically rostrate as opposed to the obtuse keel petals found in the other groups (Fig. 5.3).

5.2.5 Anthers—Surprisingly, it was found that the size and shape of the carinal anther are diagnostic for each of the three groups in Lebeckia s.l. In

Lebeckia s.s., the carinal anther resembles the long, basifixed anthers (Fig. 5.3 h3) resulting in a 5+5 arrangement. In the Calobota group (including Spartidium saharae), the carinal anther is intermediate between the dorsifixed and basifixed anthers (Fig. 5.3 e3, f3, g3), resulting in a 4+1+5 anther configuration (i.e., four long basifixed anthers, an intermediate carinal anther and five short dorsifixed anthers).

The carinal anther is usually attached a little higher up. In the Viborgioides group, the carinal anther resembles the short, dorsifixed anthers (Fig. 5.3 a3, b3, c3, d3), resulting in a 4+6 anther arrangement (i.e., four long anthers and six short anthers).

The anther arrangement is therefore in perfect agreement with the leaf anatomy of the three groups.

60 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

5.2.6 Fruit and seeds—Pods in the Crotalarieae are an important source of systematic information, as specialisations for seed protection and dispersal may result in structural differences. Lebeckia s.l. displays a great diversity of fruit structure, including dehiscent and indehiscent fruits that are either inflated or laterally compressed, and with or without wings (Polhill 1976). Fruits of Lebeckia s.s. are terete to semi-terete and thick- or thin-walled, with wings on the upper suture in some species, e.g. L. meyeriana. Species of the Calobota group generally have terete or semi-terete pods that are thick-walled and spongy or the fruits are thin- walled (membranous), laterally compressed and pubescent or glabrous, as is also found in Spartidium saharae. In contrast, the fruits of section Viborgioides (and L. inflata) are inflated and always glabrous, with highly sclerified, thin walls. The placement of Wiborgia humilis within the Viborgioides group is supported by fruit structure. Wiborgia humilis has inflated pods that lack wings on the upper suture and do not have highly sclerified fruit walls. In contrast, the winged samaras of the rest of

Wiborgia are laterally compressed and have highly sclerified fruit walls in most species (Dahlgren 1975).

61 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA al bl dl Oki a2 b2 c2

a3 c3 @AA b3 d3

g3 h3 f3 iJ k5)

FIG. 5.3. Drawings of the calyces (vestiture not shown), keel petals and anthers of selected species of Lebeckia Spartidium saharae and Wiborgia humilis. Note the relative size of the carinal lobe of the calyx, the apex and vestiture of the keel petal and the relative size of the carinal anther. (al—a3) W. humilis; (b1—b3) L. mucronata; (cl—c3) L. inflata; (dl—d3) L. leipoldtiana; (el—e3) L. cytisoides; (fl—f3) L. pungens; (gl—g3) S. saharae; (hl—h3) L. sepiaria.

Voucher specimens: (al, a3) Van Wyk 3530 (JRAU); (a2) Boatwright et al., 216

(JRAU); (bl—b3) Stirton 10880 (JRAU); (cl—c3) Vlok et al. 2 (JRAU); (dl—d3) Boatwright et al. 123 (JRAU); (el—e3) Boatwright et al. 114 (JRAU); (fl ) Taylor 9386 (NBG); (f2)

Boatwright et al. 106 (JRAU); (f3) Van Wyk 3252 (JRAU); (gl—g3) Hill 1910 (K); (h1, h3)

Barker 6515 (NBG); (h2) Le Roux et al. 24 (JRAU). Scale bars (for all calyces, all keel petals and all anthers, respectively) = 1.0 mm.

62 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

Polhill (1976, 1981) used the orientation of the seed in Spartidium as the only diagnostic character to separate this genus from Lebeckia s.l. A study of most of the species showed that at least three species of the Calobota group (L. macrantha, L. psiloloba and L. pungens) have the seed oriented at right angles to the placenta, exactly as in Spartidum saharae.

The seed surface of species in Lebeckia s.s. is invariably rugose, while the seeds of only one species in the Calobota group (L. lotononoides) and one species in the Viborgioides group (L. inflata) have rugose seeds (Le Roux and Van Wyk

2007, 2008, 2009; Boatwright and Van Wyk 2007).

5.3 MOLECULAR EVIDENCE FOR THE PARAPHYLY OF LEBECKIA

5.3.1 Combined molecular analyses—In the combined molecular analyses

(Parsimony and Bayesian analyses; Fig. 5.1a) presented in Chapter 3 (Boatwright et al. 2008a), the Calobota group is clearly monophyletic with very strong support. In the Bayesian analysis, Spartidium groups with the Lebeckia multiflora group with strong support (Fig. 5.1a). Lebeckia section Viborgioides (including Wiborgia humilis) is monophyletic with weak to strong support, but the positions of L. inflata and L. mucronata are unresolved. It is interesting to note that molecular evidence strongly supports the exclusion of Wiborgia humilis from Wiborgia and the transfer of this species to the Viborgioides group. Surprisingly, Lebeckia s.s. is unresolved in the molecular analyses. Morphologically this group is very distinct from the other groups of Lebeckia s.l. and all other Cape Crotalarieae.

5.3.2 Combined molecular and morphological analysis—When the molecular data were combined with morphological data (Fig. 5.1b), the resolution

63 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA within the "Cape group" and among the three groups of Lebeckia s.l. was much improved. The Calobota group is again strongly supported as monophyletic. Within the Viborgioides group, L. section Viborgioides including Wiborgia humilis is monophyletic with strong support and L. inflate is sister to this group, albeit without support. The position of L. mucronata, however, remains unresolved in this analysis, although abundant morphological and anatomical evidence suggests its placement within this group. The inclusion of W. humilis in the Viborgioides group again receives strong support. With the addition of morphological data Lebeckia s.s. is moderately supported as monophyletic as opposed to being unresolved in the molecular analyses (Chapter 3; Boatwright et al. 2008a).

5.4 TAXONOMIC IMPLICATIONS

Major continuities and discontinuities in morphological characters among the three groups discussed above closely agree with the three main clades revealed by genetic analysis (Fig. 5.1a and 5.1b). The new system proposed by Boatwright et al.

(submitted) is based on a wider consideration of the intricate relationships amongst all genera of the tribe Crotalarieae, all of which have been revised in recent years

(Pearsonia — Polhill 1974; Wiborgia — Dahlgren 1975; Crotalaria — Polhill 1982;

Aspalathus — Dahlgren 1988; Lotononis — Van Wyk 1991b; Rafnia — Campbell and

Van Wyk 2001; Bolusia — Van Wyk 2003a, Van Wyk et al. submitted; Lebeckia s.s. —

Le Roux and Van Wyk 2007, 2008, 2009; Robynsiophyton — Chapter 9; Boatwright and Van Wyk 2009; Rothia — Chapter 10; Boatwright et al. 2008c). Dahlgren (1970a) and Polhill (1976) both discussed the high incidence of convergence in the tribe

Crotalarieae and the intricate relationships between the genera. The difficulty in

64 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA determining generic limits within the Crotalarieae is discussed by Polhill (1976), who emphasized that it would be unwise to propose modifications to the system without a clear understanding of the patterns of character state distributions. Furthermore, alterations to the system should result in a more predictive and useful system without running the risk of instability of circumscriptions and nomenclature. In existing keys to the genera of the Crotalarieae (Polhill 1981; Van Wyk and Schutte

1995), the lack of uniformity and clear-cut diagnostic characters for Lebeckia s.l. is clearly reflected in the fact that the genus keys out no less than three times (in both keys). Unique combinations of morphological characters have now been identified for the three main groups that clearly should be given generic status. The improvement in generic delimitations is also reflected in the following key.

5.4.1. Key to the genera of the Crotalarieae

1. Leaves acicular, terete:

2. Ovary 2 to 4 ovulate, pods few-seeded (±1 to 2-seeded) Aspalathus

Ovary with more than 6 ovules, pods many-seeded (if rarely few-seeded,

then the pods strongly inflated with spongy walls) Lebeckia s.s.

1. Leaves digitate, unifoliolate or simple (flat, never terete) 3

Stipules present 4

Style curved upwards; anthers dimorphic .5

Stipules asymmetrical or single; style glabrous and not

helically coiled; keel obtuse or rostrate; anther arrangement

4+6, 4+1+5 or very rarely 5+5 Lotononis

65 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

5. Stipules symmetrical when present; style with 1 or 2 lines of

hairs or glabrous and helically coiled; keel strongly rostrate

(often at right angles) or helically coiled; anther arrangement

5+5 6

6. Stipules dentate; beak of keel and style helically

coiled, style glabrous Bolusia

6. Stipules entire; beak of keel and style not helically

coiled, style with 1 or 2 lines of hairs Crotalaria

4. Style straight or rarely down-curved; anthers similar in size and

shape 7

7. Stamens nine (five fertile and four lacking

anthers) Robynsiophyton

7. Stamens 10 (all fertile) 8

8. Anthers all rounded; prostrate annuals Rothia

8. Four anthers basifixed, six attached slightly higher up,

all elongate; perennial herbs or shrubs Pearsonia

3. Stipules absent:

9. Calyx with upper and lateral lobes fused on either side Lotononis

9. Calyx sub-equal (upper and lateral lobes not fused) 10

10. Leaves sessile; upper suture of pod asymmetrically

convex 11

11. Plants glabrous except occasionally on bracts and

bracteoles; usually turning black when dried Rafnia

11. Plants usually pubescent on all parts, if leaves

glabrous then standard petal hairy and inner surface of

66 CHAPTER 5: EVIDENCE FOR THE POLYPHYLY OF LEBECKIA

calyx glabrous; not turning black when

dried Aspalathus

10. Leaves usually petiolate, if leaves sessile then plants with many-seeded pods and at least some hairs on leaves or stems; upper suture of pod symmetrically convex 12

12. Petals pubescent; at least on the dorsal midrib of the

standard petal (if glabrous then plants strongly

spinescent, woody, practically leafless shrubs); twigs

green (bark formation late, chlorenchyma present);

leaves isobilateral Calobota

12. Petals glabrous; twigs brown (bark formation early,

chlorenchyma absent; if twigs rarely green then plant a

short-lived fireweed); leaves dorsiventral 13

13. Fruits winged, indehiscent; carinal anther

intermediate (anthers 4+1+5) Wiborgia

13. Fruits without wings, dehiscent (if rarely

indehiscent then ovary and fruit distinctly stalked);

carinal anther resembles short anthers (anthers

4+6) Wiborgiella

67 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

6.1 INTRODUCTION

The genus Calobota should clearly be reinstated to accommodate the 16

species of the Calobota group (Chapter 5; Boatwright et al. submitted). Calobota

was originally described by Ecklon and Zeyher (1836) and consisted of two species,

C. cytisoides and C. pulchella, which are actually the same species (now C.

cytisoides). The genus was later reduced to a section of an expanded concept of

Lebeckia by Bentham (1844). Several species which he included in the section were

previously placed in other genera by Ecklon and Zeyher (1836). Harvey (1862)

followed the sectional classification of Bentham in the last revision of the genus

Lebeckia. In Chapter 5 and Boatwright et al. (submitted) the reinstatement of

Calobota is proposed which now also includes Lebeckia section Stiza and the

monotypic genus Spartidium, based on the results of a phylogenetic study of the

tribe Crotalarieae (Chapters 3 and 5; Boatwright et al. 2008a). These studies, based

on molecular (nrITS and rbcL) and morphological data, revealed that the genus

Lebeckia is not monophyletic and a division into three genera is therefore proposed

by Boatwright et al. (submitted), viz. Lebeckia s.s., Calobota and Wiborgiella. The species of Calobota occur throughout the Cape and extend into Namibia. Some of the species are important components of the Sperrgebiet, a species-rich area of south-western Namibia (Burke and Mannheimer 2004), while two other species, C. namibiensis and C. obovata are endemic to Namibia (Craven and Vorster 2006).

Calobota saharae, however, is endemic to North Africa and occurs in Algeria,

Morocco and Libya (Polhill 1976).

68 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Calobota can be distinguished from Lebeckia and Wiborgiella by a combination of the following characters: green young twigs, uni- or trifoliolate to simple, laminar leaves, hairy petals (C. cuspidosa and C. psiloloba are exceptions), anther configuration of 4+5+1 and the laterally compressed or semi-terete, usually pubescent pods. The species within Calobota are often difficult to distinguish from one another due to the close similarity between related species. This motivated the need for a taxonomic revision that would provide clarity on the number of species within Calobota and a key to these species.

In this chapter, a detailed treatment of the genus Calobota is presented including a key to the species, complete nomenclature, typification, formal descriptions, the known geographical distribution and illustrations of all the species.

6.2 DISCUSSION OF CHARACTERS

6.2.1 Vegetative morphology and anatomy—HABIT AND BRANCHES—All the species of Calobota are perennial, multi-stemmed shrubs (Fig. 6.1) usually about

1.0-1.5 m in height, but C. cuspidosa can reach heights of up to 4 m. Calobota lotononoides is a decumbent shrublet that reaches heights of up to 0.5 m and has extensive underground branches. This species appears to be adapted to sandy habitats as the underground branches allow the stems and leaves to emerge above ground when they become buried (Boatwright and Van Wyk, 2007). Calobota acanthoclada, C. cuspidosa, C. psiloloba, C. pungens and C. spinescens are all spinescent shrubs, with the branch ends and inflorescences terminating in spines.

The typical form of C. sericea sometimes also has spinescent branches, but they are more often unarmed.

69 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Fig. 6.1 Habit of Calobota species. (a) C. cinerea; (b) C. cytisoides; (c) C. halenbergensis;

(d) C. lotononoides; (e) C. pungens; (f) C. sericea; (g) C. spinescens; (h) C. thunbergii.

70 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Fig. 6.2 Transverse sections through the stems and leaves of Calobota species. (a) Portion of the stem of C. spinescens; (b) Portion of stem the of C. sericea; (c) Petiole of C. lotononoides; (d) Petiole of C. elongata; (e) Portion of the juvenile leaf of C. pungens; (f)

Portion of the leaf of C. thunbergii; (g) Potion of the leaf of C. linearifolia; (h) Portion of the leaf of C. obovata. Voucher specimens: (a) Boatwright et al. 158 (JRAU); (b) Boatwright et al. 138 (JRAU); (c) Boatwright et al. 142 (JRAU); (d) Van Wyk 2562b (JRAU); (e) Boatwright et al. 106 (JRAU); (f) Boatwright et al. 151 (JRAU); (g) Dean 673 (JRAU); (h) Kers 152

(WIND). Scale bars: (a, b, e) 0.1 mm; (c, d, f, g, h) 0.2 mm.

71 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

The young branches in species of Calobota, with the exception of C. acanthoclada, are green as opposed to the brown branches found in most species of Wiborgiella. This is due to the presence of several chlorenchyma layers below the epidermis (Fig. 6.2), a feature usually found in papilionoid legumes where the leaves are reduced or absent (Metcalfe and Chalk 1950). Although green stems are also found in other genera of the Crotalarieae, notably Crotalaria, Lebeckia, Lotononis and Rafnia, it is a useful character to distinguish Calobota from Wiborgiella and

Wiborgia. Some species of Calobota lack leaves on their mature branches and in these it is likely that the stem performs the main photosynthetic function. The epidermis in most species is covered by a thick cuticle and the epidermal cells are bottle-shaped.

LEAVES—Leaves in Calobota vary from trifoliolate or rarely 5-foliolate to unifoliolate or simple. In three species with unifoliolate leaves, C. cuspidosa, C. psiloloba and C. pungens, the juvenile leaves are trifoliolate and become unifoliolate on the mature branches due to a loss of the lateral leaflets and shortening of the petioles (Dahlgren 1970; Polhill 1976; Boatwright et al. submitted). A similar reduction has been observed in C. obovata, which also has unifoliolate leaves on the mature branches. The leaves of Calobota species are invariably laminar as opposed to the acicular, phyllodinous leaves of Lebeckia s.s. and usually have long, slender petioles. The leaf margins are entire and stipules are lacking. The leaves are always pubescent or at least pilose.

The leaf arrangement in Calobota is alternate and widely spaced, but in C. acanthoclada and C. spinescens the leaves are fasciculate. This is a useful character to distinguish especially C. spinescens from C. sericea with which it is often confused.

72 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Leaflet shape varies from oblanceolate to elliptic or obovate to suborbicular and less often spathulate. The apices may be acute, obtuse, retuse, truncate, mucronulate or recurved-mucronate and the base of the leaves angustate, attenuate or cuneate. Calobota sericea is a highly variable species and four regional forms can be distinguished based mainly on leaf size, shape and especially the width of the leaflets.

The anatomy of the leaves of Calobota species (Fig. 6.2) provided some interesting characters to distinguish it from Lebeckia and Wiborgiella. In Calobota, the leaves are isobilateral and lack mucilage cells in the epidermis. Transverse sections through juvenile leaves of C. pungens do show mucilage cells in the epidermis, suggesting that there is a loss of these cells in Calobota. Calobota saharae is an exception in that mucilage cells are present in mature leaves. Gregory and Baas (1989) speculate that these cells are usually associated with plants that occur in Mediterranean climates. Calobota saharae is endemic to Mediterranean

North Africa and this could explain the retention of these cells. In Lebeckia the leaves are acicular with palisade parenchyma adjacent to the epidermis and mucilage cells present. Wiborgiella species have dorsiventral leaves (cleary differentiated into palisade and spongy parenchyma) with mucilage cells present in the epidermis.

6.2.2 Reproductive morphology and anatomy—INFLORESCENCE—The structure of the inflorescence in Calobota is fairly uniform and all species have terminal racemes which only vary in length and in the number of flowers per raceme.

This type of inflorescence is rather common in Papilionoideae and also in the

Crotalarieae (Polhill 1976). In one species, C. elongata, the inflorescence is a spike

73 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA rather than a raceme and very densely flowered, with more than 100 sessile flowers

(Dahlgren 1967b).

FLOWERS—Flower size is somewhat diagnostic at species level. The largest flowers are found in C. cytisoides and C. thunbergii (up to 35 mm and 24 mm long, respectively) and C. cuspidosa has flowers of up to 20 mm. The rest of the species range from up to 17 mm in C. linearifolia and C. saharae, to below 12 mm in C. elongata, C. halenbergensis, C. namibiensis, C. psiloloba and C. sericea. The petals are usually pubescent in Calobota (at least pilose along the dorsal midrib) with the exception of C. cuspidosa and C. psiloloba that have glabrous petals. The petals of all the species are yellow. The pedicel is short in most species, but may be long in

C. cytisoides (up to 11 mm) and C. psiloloba (up to 6 mm) or absent in C. elongata.

Bracts and bracteoles vary little between species. A single bract is located at the base of the pedicel, with two bracteoles further up near the middle of the pedicel.

In Calobota the bracts and bracteoles are small, linear and pubescent on the upper surface, except in C. cytisoides and C. obovata where they are often ovate.

The calyces of Calobota species are typically of the "lebeckioid" type (Polhill

1976). They are sub-equally lobed, with the upper sinus often deeper than the lateral or lower sinuses and the carinal lobe consistently narrower than the others.

The outer surface of the calyx is usually pubescent, pilose or at least glabrescent.

The tips of the calyx lobes are minutely pubescent on the inner surface which is found in most of the genera of the "Cape group" of the Crotalarieae, with the notable exception of Aspalathus.

The standard petal is mostly widely ovate in the species of Calobota, but is elliptic in C. elongata, orbicular in C. halenbergensis and transversely oblong in C. lotononoides. The apices are either acute or obtuse, except in C. psiloloba and C.

74 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA lotononoides which have emarginated apices. The claws are relatively short and less than 6 mm, except in C. cuspidosa, C. cytisoides and C. lotononoides where the claws are longer than 6 mm long. The standard petal is usually pubescent in

Calobota species or at least has pilose hairs along the dorsal midrib, except in C. cuspidosa and C. psiloloba which have totally glabrous standard petals.

The shape of the wing petals is fairly uniform in the genus and mostly oblong with obtuse apices. They are either shorter than the keel petals (C. cinerea, C. cuspidosa, C. linearifolia, C. namibiensis, C. obovata, C. psiloloba, C. pungens), as long as or shorter than the keel petals (C. acanthoclada, C. cytisoides, C. spinescens, C. thunbergii), as long as or longer than the keel petals (C. elongata, C. saharae, C. sericea) or markedly longer than the keel petals (C. halenbergensis, C. lotononoides). C. lotononoides is unique within the genus in having a wing- to keel lamina ratio of 2:1; the wing petal laminas of the other species are never more than

1.3 times longer than those of the keel petals (Boatwright and Van Wyk 2007). The wing petals are glabrous in most species, but pilose in C. acanthoclada, C. cinerea,

C. linearifolia, C. obovata and C. pungens. Sculpturing is present in all species on the upper basal and upper central parts of the wing petals. In some species, e.g. C. cytisoides, C. elongata and C. thunbergii, the sculpturing extends to the lower basal, lower central and upper/lower distal parts of the wing petals (see Stirton 1981 for terminology).

The keel petals of all species are boat-shaped with obtuse apices and pockets are invariably present in all species. The petals are either pubescent (C. acanthoclada, C. cinerea, C. linearifolia, C. cytisoides, C. namibiensis, C. obovata,

C. pungens and C. thunbergii) or glabrous (C. psiloloba, C. cuspidosa, C. elongata,

75 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

C. halenbergensis, C. lotononoides, C. saharae, C. sericea). In C. spinescens the keel petals may be either pilose or glabrous.

The value of stamina! characters in the genistoid legumes is well known

(Polhill 1976; Schutte and Van Wyk 1995; Boatwright et al. submitted) and provided very reliable characters to segregate Calobota and Wiborgiella from Lebeckia s.s

(see Chapter 5). The species of Calobota have a 4+5+1 anther arrangement (four long basifixed anthers, five short dorsifixed anthers and an intermediate carinal anther), while those of Lebeckia have a 5+5 arrangement (the carinal anther resembles the long basifixed anthers) and species of Wiborgiella have a 4+6 arrangement (the carinal anther resembles the short dorsifixed anthers). The anther arrangement was found to be in agreement with other evidence (morphological, anatomical and molecular) that support these generic concepts.

The pistil is subsessile or shortly stipitate, usually with a linear ovary and upcurved style. The ovary may be either pubescent or glabrous, while the style is always glabrous. The ovary may contain as few as 3 to 6 ovules (C. elongata, C. namibiensis) or as many as 34 ovules (C. cytisoides). The style is longer than the ovary in C. namibiensis.

The nectar of Calobota species is generally sucrose-rich (Van Wyk 1993), which is typical of most bee-pollinated legumes. The potential pollinators of some of the Cape genera of the Crotalarieae were studied by Gess and Gess (1994). They found that the flowers of C. spinescens and C. thunbergii were visited by solitary aculeate Hymenoptera, specifically Masarinae and Megachilinae of the tribes

Megachilini and Anthidiini. Some species of Xylocopinae also occasionally visited C. thunbergii and Apis mellifera was also often abundant. The megachiline and masarine visitors were able to trip (displacement of the keel petals to come into

76 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA contact with the pollen) the flowers of C. spinescens, but not those of C. thunbergii suggesting that they are potential pollinators of only the former. Apis mellifera, however, was the only species that was able to trip the flowers of Calobota thunbergii making it a potential pollinator of this species (Gess and Gess 1994,

2004, 2006). The diversity of flower visits to other species of Calobota and the

Crotalarieae has been recorded by Gess and Gess (2003).

FRUIT—The pods of Calobota species vary from laterally compressed and membraneous to semi-terete and glabrous or pubescent. The pods are generally many-seeded, with the exception of C. elongata. In this species, the pods are small, light and usually single-seeded. The petals dry out but are persistent and serve as wings for the disperal of the tiny pod by wind, through a rolling action in the open sandy places where these plants typically grow. A similar adaptation to wind dispersal is found in Lotononis section Synclistus (Van Wyk 1991b).

The anatomy of the fruit revealed two types of fruit walls in species of

Calobota, viz. thick- and thin-walled. Those with thick-walled pods usually have a thick mesocarp that may have several layers of sclerenchyma just below the endocarp. The endocarp in most species is composed of a loosely arranged mass of cells which have the appearance of white sponge to the naked eye and seem to envelope the seed, perhaps to serve some protective function. In the species with thin-walled pods, the mesocarp is thinner and has only one or two rows of sclereids adjacent to the endocarp. One or rarely two layers of cells make up the endocarp and the spongy, loosely arranged tissue is absent. In most other genera of the

Crotalarieae, the endocarp is also composed of one or two layers of periclinally elongated or round cells (Fig 6.3). In Lebeckia, however, the pods may be either thin-walled as in L. contaminata, or extremely thick-walled as in L. brevicarpa. The

77 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA spongy tissue present in some Calobota species is also found in Lebeckia, e.g. L sepiaria. Most of the other genera investigated had thin-walled pods. In Wiborgia the pods may be either highly sclerified as in W. sericea or thin-walled with only a few layers of sclerenchyma as in W. incurvata. Crotalaria and Rafnia lack sclerenchyma entirely, while Wiborgiella species have several layers of sclerenchyma adjacent to the endocarp. Melolobium was included to represent a member of the Genisteae with thin-walled pods. The structure of the fruit wall is practically identical to that of

Wiborgiella (Fig. 6.3).

SEEDS—In Calobota the seeds are reniform or oblong-reniform and vary from light brown to pink, sometimes with brown mottling, and generally have a brown hilum. The surface is smooth in all but one species, namely C. lotononoides that has rugose seeds (Boatwright and Van Wyk 2007).

78 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Fig. 6.3 Transverse sections through the fruit walls of Calobota species. (a) C. sericea; (b)

C. namibiensis; (c) C. thunbergii; (d) C. spinescens; (e) C. cinerea; (f) C. elongata. Voucher specimens: (a) Boatwright et al. 138 (JRAU); (b) De Winter and Hardy 7919 (WIND); (c)

Boatwright et al. 151 (JRAU); (d) Boatwright et al. 158 (JRAU); (e) Boatwright et al. 150

(JRAU); (f) Van Wyk 2562b (JRAU). Scale bar represents 0.2 mm.

79 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Fig. 6.4 Transverse sections through the fruit walls of representative genera of the

Crotalarieae. (a) Lebeckia wrightii; (b) Wiborgia sericea; (c) W. monoptera; (d) Melolobium alpinum; (e) Lebeckia ambigua; (f) Wiborgiella sessilifolia; (g) Crotalaria sp.; (h) Rafnia amplexicaulis; (i) Lebeckia brevicarpa; (j) L. pauciflora; (k) L. contaminata. Voucher specimens: (a) Johns 163 (JRAU); (b) Boatwright et al. 124 (JRAU); (c) Boatwright et al.

152 (JRAU); (d) Schutte 160 (JRAU); (e) Boatwright et al. 132 (JRAU); (f) Taylor 4329

(PRE); (g) De Villiers 106 (JRAU); (h) Campbell and Van Wyk 40 (JRAU); (i) Le Roux et al.

4 (JRAU); (j) Le Roux et al. 12 (JRAU); (k) Le Roux et al. 16 (JRAU). Scale bars: all 0.2 mm as in (k); (i) 0.8 mm.

80 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

6.2.3 Phylogenetic Relationships—MORPHOLOGICAL DATA—TO assess infrageneric relationships within Calobota, 17 morphological and anatomical characters were scored for a total of 21 taxa (Table 6.1). Wiborgiella was used as outgroup and Lebeckia and Rafnia included as ingroups based on the results in

Chapter 3 (Boatwright et al. 2008a). All 17 characters are parsimony informative and the analysis resulted in six equally parsimonious trees with a length of 36 steps, CI of 0.64 and RI of 0.81 (Fig. 6.5). The Crotalarieae are known for high incidences of convergence and parallelisms (Dahlgren 1970a, Boatwright et al. 2008a) and the co- occurrence of seemingly identical apomorphic states in distantly related genera complicate morphological analysis. Unique combinations of characters are important to circumscribe genera in the tribe (Boatwright et al. submitted), but these are not reflected in the analysis.

Calobota is weakly supported as monophyletic (63 BP). This is supported by the isobilateral leaves, pubescent petals (although this is reversed in C. cuspidosa and C. psiloloba), the 4+5+1 anther arrangement which is shared with Rafnia and pubescent pods (reversed in C. cuspidosa, C. cytisoides, C. halenbergensis, C. psiloloba and C. saharae). Two clades are present within Calobota, albeit without support, the C. spinescens Glade and the C. cytisoides Glade. Calobota saharae is sister to both these clades. The C. spinescens Glade is supported by the spinescent habit and the laterally compressed fruit. The latter is also present in C. saharae, which differs from the rest of Calobota in having mucilage cells present in the epidermis (these have been lost in the mature leaves of the other species). The close relationship between C. cuspidosa, C. psiloloba and C. pungens receives weak bootstrap support (56 BP), but is supported by the virtual absence of leaves on mature plants (shared with C. namibiensis and C. saharae) and unifoliolate

81 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

leaves (shared with C. obovata). The sister relationship between C. cuspidosa and

C. psiloloba (74 BP) is furthermore supported by the presence of glabrous petals

and pods. Calobota acanthoclada and C. spinescens share the fasciculate

trifoliolate or simple leaves and are weakly supported as sister (51 BP).

The C. cytisoides group is supported by trifoliolate leaves (also present in C.

spinescens), semi-terete and thick-walled fruit (reversed in C. elongata). Calobota

cytisoides and C. thunbergii are sister taxa (73 BP) based on the large flowers

(more than 20 mm long) and the wing petal sculpturing present on the upper basal,

upper central, lower basal, lower central and upper/lower distal parts. The C. sericea

Glade all have wing petals that are longer than the keel petals, which they share with

C. saharae and both Rafnia species.

Lebeckia and Rafnia are supported as sister by the glabrous vegetative parts

(55 BP). Lebeckia is strongly supported by the suffrutescent habit, acicular,

phyllodinous leaves and the 5+5 anther arrangement (100 BP), while Rafnia

received weak support (50 BP).

MOLECULAR DATA—In Chapter 3 (see also Boatwright et al. 2008a), 11 species of Calobota were included in the phylogenetic study of the tribe Crotalarieae

(see Figure 3.4). Calobota was weakly supported as monophyletic by separate analyses of rbcL and ITS, but strongly supported by combined rbcL/ITS and combined rbcL/ITS/morphological data. Although the sampling was not comprehensive, some strongly supported groups were noted. Calobota cuspidosa,

C. psiloloba and C. pungens are strongly supported as being closely related.

Calobota cytisoides and C. thunbergii are strongly supported as sister taxa, as are

C. sericea and C. lotononoides. The expansion of Calobota to include C. saharae

82 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(previously the monotypic genus Spartidium) is also supported by molecular and morphological data.

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CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

51 Calobota acanthoclada Calobota spinescens 4 1 Calobota cuspidosa 56 I Calobota psiloloba Calobota pungens 3 Calobota cytisoides Calobota thunbergii 1 Calobota elongata Calobota halenbergensis Calobota lotononoides Calobota sericea 63 Calobota cinerea Calobota obovata Calobota linearifolia 100 Calobota namibiensis Calobota saharae 100 t Lebeckia pauciflora 55 Lebeckia sepiaria 50 I Rafnia amplexicaulis Rafnia globosa Wiborgiella /eipoldtiana

FIG. 6.5. Strict consensus of six trees produced by parsimony analysis of 17 morphological

characters for the genera Calobota, Lebeckia, Rafnia and Wiborgiella. Values above the

branches are bootstrap percentages above 50%.

85 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

6.3 TAXONOMIC TREATMENT

CALOBOTA Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 192. Jan. 1836, emend. Boatwr.

and B.-E.van Wyk, emend. nov. Lebeckia section Calobota (Eckl. and Zeyh.)

Benth. in Hook. Lond. J. Bot. 3: 358-361. 1844. pro parte majore.—

LECTOTYPE species (here designated): Calobota cytisoides (Berg.) Boatwr.

and B.-E.van Wyk. [Note: This species is chosen as lectotype as it is the only

species included in Ecklon and Zeyher's original concept of Calobota.]

Acanthobotrya Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 192. Jan 1836, pro parte.

Lectotype species (here designated): A. pungens sensu Eckl. and Zeyh. [now

Calobota psiloloba (E.Mey.) Boatwr. and B.-E.van Wyk]. [Note: As mentioned

by Bentham (1844), Acanthobotrya is a mixed concept representing at least

four different genera. However, the diagnosis agrees with the concept of

Stiza E.Mey. (e.g. linear-oblong, compressed fruits); furthermore, C. psiloloba

(at the time still part of the concept of A. pungens), which has all the

diagnostic characters mentioned in the diagnosis, is listed first.]

Stiza E.Mey., Comm. Pl. Afr. Austr. 1: 31. Feb. 1836, synon. nov. Lebeckia section

Stiza (E.Mey.) Benth. in Hook. Lond. J. Bot 3: 355-356, 1844; Harv. in Harv.

in Harv. and Sond., Fl. Cap. 2: 83. 1862.; Bentham and Hooker, Gen. Pl.:

447. 1865.—LECTOTYPE species (designated here): Stiza erioloba E.Mey.

(now Calobota pungens) [Note: Meyer (1836) included only Stiza erioloba

and S. psiloloba E.Mey. (now Calobota psiloloba) in Stiza. The original

description of Stiza is too general to allow for a considered choice of

lectotype, so we here choose C. pungens simply as it is the first-mentioned

species.]

86 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Spartidium Pomel, Nouv. Mat. FL Atl.: 173. 1874, synon. nov. Type species:

Spartidium saharae (Coss. and Durr.) Pomel. [Note: Spartidium is

monotypic.]

Woody, sometimes spinescent shrubs or shrublets. Branches thick and

woody; young branches green, with chlorenchyma and lacking bark (except in C.

acanthoclada), pubescent, often sericeous. Stipules absent. Leaves unifoliolate or

digitately trifoliolate (rarely 5-foliolate); petioles shorter or longer than leaflets,

pubescent; leaflets oblanceolate to elliptic or obovate to suborbicular, less often

spathulate, pubescent. Inflorescence terminal, few to multi-flowered racemes or

rarely a spike. Pedicel pubescent. Bracts linear to obovate, pubescent, caducous.

Bracteoles linear, pubescent, caducous. Corolla yellow, usually pubescent or at

least pilose along the median section of the standard petal (except in C. cuspidosa

and C. psiloloba). Calyx subequally lobed, upper sinus often deeper than the lateral

or lower sinuses, carinal lobe narrower than the others, pubescent or at least

glabrescent, tips of lobes minutely pubescent on inner surface. Standard linear to

widely ovate, with basal callosities in C. cytisoides. Wing petals narrowly oblong to

oblong or slightly ovate, longer or shorter than the keel, terminal parts sometimes

pubescent; apex obtuse. Keel petals oblong, pockets invariably present, terminal

parts sometimes pubescent; apex obtuse. Anthers dimorphic, four long, basifixed

anthers alternating with five ovate, dorsifixed anthers, carinal anther intermediate in

size and shape between the basifixed and dorsifixed anthers. Pistil sub-sessile to very shortly stipitate, ovary linear to slightly elliptic, with 4 to many ovules,

pubescent or glabrous; style longer or shorter than the ovary, curved upwards, glabrous. Pods laterally compressed or semi-terete, linear to oblong, few- to many-

87 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA seeded, glabrous or pubescent, dehiscent or indehiscent. Seeds reniform to oblong- reniform, or less often suborbicular; colour variable, light pink to pink, sometimes mottled with brown; hilum round, brown or black; surface smooth (rugose only in C. lotononoides; seeds of C. namibiensis, C. obovata and C. saharae not seen).

Diagnostic characters— Bark formation is late, so that the twigs remain green and photosynthesizing, while bark formation is early in most species of

Wiborgiella, so that even the young twigs are not green but covered in brown bark

(this is also true of Wiborgia species). The green twigs are a useful diagnostic character, visible even in sterile herbarium specimens. Calobota also differs from

Wiborgiella in the hairy petals (C. cupidosa and C. psiloloba are exceptions) seeing that the petals are glabrous in Wiborgiella. The most reliable diagnostic character to distinguish Calobota from Wiborgiella is the anther configuration of 4+1+5 (4+6 in the latter). The pods are semi-terete or laterally compressed in Calobota and usually pubescent (pods terete or rarely laterally compressed in species of Wiborgiella and always glabrous).

Notes on distribution— The species of Calobota occur in the Eastern,

Northern and Western Cape Provinces, with some restricted to Namibia. One species, Calobota saharae, is endemic to North Africa where it occurs on sand dunes from Libya to Algeria and Morocco (Polhill 1976).

KEY TO THE SPECIES OF CALOBOTA

1. Plants strongly spinescent shrubs, pods laterally compressed, often falcate .2

2. Plants leafless shrubs, if leaves present then alternate and widely

seperated 3

88 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

3. Petals pilose, ovary pubescent and pods densely

tomentose C. pungens

3. Petals glabrous, pods glabrous 4

4. Flowers usually (15—) 16-18 (-20) mm long; pods (21—) 28-

49 (-53) mm long; plants restricted to the Northern

Cape C. cuspidosa

4. Flowers usually 8-11 mm long; pods 14-17 (-33) mm long;

plants restricted to the Eastern Cape C. psiloloba

2. Plants densely leafy, leaves alternate and fasciculate 5

5. Leaves trifoliolate, young stems green, pods constricted between

the seeds C. spinescens

5. Leaves unifoliolate, young stems brown, pods not constricted

between the seeds C. acanthoclada

1. Plants not spinescent, pods semi-terete, if laterally compressed then plants unarmed, erect or virgate, leafless shrubs restricted to North Africa 6

6. Leaves widely obovate or orbicular (8-25 mm wide), short-sericeous 7

7. Leaves trifoliolate, inflorescences long, spicate and densely

flowered (more than 100 flowers), pods ovate to elliptic, 1 to 2-

seeded C. elongata

7. Leaves unifoliolate (rarely trifoliolate), inflorescences short,

racemose and 7- to 16-flowered, pods linear, 5 to 7-

seeded C. obovata

6. Leaves elliptic to oblanceolate, linear, spathulate, or rarely ovate or

obovate and generally less than 10 mm wide, pubescent or sericeous 8

89 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

8. Wing petals longer than keel, leaflets narrow and much shorter than

the petioles 9

9. Plants pilose and not sericeous, pods and pistil

glabrous C. halenbergensis

9. Plants sericeous, pods and pistil pubescent or sericeous...10

10. Leaves not folded inward and directed to the same

side, flowers 7-11 mm long, wing petals not extremely

long (wing to keel lamina ratio of less than

1:1.3) C. sericea

10. Leaves conduplicate and secund, flowers 12-15 mm

long, wing petals extremely long (wing to keel lamina

ratio of 2:1) C. lotononoides

8. Wing petals shorter than the keel, leaflets broad and as long as or

slightly shorter than the petioles or petioles absent 11

11. Plants densely leafy and sericeous at least on the young branches 12

12. Flowers generally large (between 13 and 35 mm long) large, wing petals

with sculpturing extending to the lower basal, lower central and upper or

lower distal parts 13

13. Leaf apices mucronulate; calyx, ovary and pods

glabrous C cytisoides

13. Leaf apices acute, calyx, ovary and pods

sericeous C. thunbergii

12. Flowers medium-sized (between 11 and 17 mm long), wing petals with

sculpturing restricted to the upper basal and upper central parts 14

90 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

14. Leaves trifoliolate with a distinct petiole, pods tomentose and not

constricted between the seeds C. cinerea

14. Leaves simple and sessile, pods short-sericeous and somewhat

constricted between the seeds C. linearifolia

11. Plants practically leafless and not sericeous 15

15. Leaves if present elliptic to ovate, keel petals, pistil and pods glabrous,

restricted to North Africa C saharae

15. Leaves if present linear, keel petals, pistil and pods densely pubescent,

restricted to Namibia C. namibiensis

1. CALOBOTA ACANTHOCLADA (Dinter) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr.

J. Bot.: submitted. 2008). Lebeckia acanthoclada Dinter, Feddes Repert. 30:

196. 1932; Merxm. and Schreiber, FSWA 60: 66. 1970.—TYPE: SOUTH

AFRICA, Namibia, Granietberge [2715 AD], Dinter 6694 (lectotype: B, photo!,

here designated). [Note: The Berlin specimen cited by Dinter is still extant

and is here chosen as lectotype.]

Lebeckia spathulifolia Dinter, Feddes Repert. 30: 197. 1932.—TYPE: SOUTH

AFRICA, Namibia, Kleinfonteiner Flache [2516 BB], Dinter 3735 (lectotype: B,

photo!, here designated; isolectotypes: BM!, PRE!, SAM!, Z, photo! 2 sheets).

[Note: This collection is the only one cited by Dinter in the original description

of this species. The specimen in B is still extant and is designated as

lectotype.]

Lebeckia candicans Dinter, Feddes Repert. 30: 198. 1932.—TYPE: SOUTH

AFRICA, Namibia, Kleinfonteiner Flache [2516 BB], Dinter 3737 (lectotype: B,

photo!, designated here; isolectotypes: BM!, BOL!, NBG!, PRE!, S!, Z,

91 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

photo!). [Note: This single specimen cited by Dinter is still extant in B and is

chosen as lectotype.]

Small, erect or decumbent, multi-stemmed, spinescent shrub up to 1 m in

height. Branches brown; young branches slightly sericeous to pilose; older branches

covered with brown bark, pilose. Leaves simple; petiole absent; fasciculate,

sericeous on both surfaces, spathulate, sessile, 6-20 mm long, 1-2 mm wide, apex

recurved-mucronate, base angustate. Inflorescence 12-25 mm long, racemose, with

3 to 5 flowers; pedicel 2-3 mm long; bract 1.5-2.5 mm long, linear, sericeous;

bracteoles 0.5-1.0 mm long, linear, sericeous. Flowers 9-12 mm long, bright yellow.

Calyx 5-7 mm long, densely sericeous; tube 2.5-4.0 mm long; lobes 2-4 mm long,

subulate. Standard 9.5-14.0 mm long; claw linear, 2-5 mm long; lamina ovate, 6.5-

8.5 mm long, 7-8 mm wide; apex acute; dorsal surface densely sericeous. Wings

9.5-11.0 mm long; claw 2.5-3.5 mm long; lamina oblong, as long as or shorter than

keel, 6.0-7.5 mm long, 2.5-3.0 mm wide, pilose, with 5-7 rows of sculpturing. Keel

10.0-12.5 mm long; claw 3-4 mm long; lamina boat-shaped, 6.5-8.5 mm long, 3.5-

4.5 mm wide, pilose to sericeous on terminal parts. Pistil subsessile to shortly stipitate, pubescent; ovary linear, 7.0-9.5 mm long, 0.7-1.0 mm wide with 9 to 15 ovules; style longer than ovary, 3.5-5.5 mm long. Pods linear, laterally compressed, subsessile to shortly stipitate, 12-19 mm long, 1.5-3.0 mm wide, ±4 to 5-seeded, dehiscent. Seeds oblong-reniform to sub-orbicular, 1.5-2.5 mm long, 1.0-2.5 mm wide, mature seeds light brown to orange, surface smooth (Fig. 6.6). Flowering time:

Flowering specimens have been collected from February through to October.

92 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

d a

i1 i2 i3

kl j2 g1 g2 k2

FIG. 6.6 Morphology of Calobota acanthoclada: (a) flower in lateral view; (b) standard petal;

(g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) intermediate carinal anther; (i3) short, dorsifixed anther; (j1—j2) leaves in abaxial view; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a—g) Mannheimer CM2012a (WIND); (h—i) Williamson 3376

(BOL); (j1—j2) Mannheimer et al. CM 899 (WIND); (k1—k2) Pillans 5149 (BOL). Scale bars:

(a—i) 1 mm; (j—k) 1 cm.

93 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—The species is similar to Calobota spinescens in the fasciculate leaves, ovate and sericeous standard petal and spinescent habit but the branches are brown and not green as in C. spinescens. It also differs in the simple leaves (trifoliolate leaves in C. spinescens), smaller flowers and shorter calyx lobes. The pods are not constricted between the seeds as is the case in C. spinescens.

Distribution and habitat—Calobota acanthoclada is largely restricted to

Namibia but extends into the Richtersveld of South Africa in the south (Fig. 6.7). It grows on sandy loam, rocky soil, limestone or dolomitic limestone at altitudes of between 170 m and 1300 m.

Additional specimens examined

2516 (Helmeringhausen): Kleinfonteiner Flache (—BB), Dinter 6269 (B, BOL).

—2518 (Tses): Kovisberge (—BB), Dinter 6293 (B, BM), Men(meiller and Giess 28445 (PRE,

WIND).

2615 (Luderitz): Klinghardt Mountains (—CA), Clark and Muller 345 (WIND); 1.0 ml. [1.61 km] south of Lagoon, Luderitz Bay (—CA), Giess and Van Vuuren 669 (K, PRE, WIND); between Elizabeth Bay and Kolmanskop, towards coast along old mine road (—CA),

Mannheimer CM2012a (WIND); Luderitz (—CA), MerxmOiler and Giess 3076 (PRE, WIND);

Luderitz, on road to Diazpunt (—CA), Mailer and Jankowitz 270 (WIND); proposed wind farm site, east of Grosse Bucht (—CC), Mannheimer and Burke CM 1700 (WIND); Grillenthal, blue dolomite hills on the road south to Oranjemund from Luderitz in the Diamond Area no. 1 (-

CD), Bean and Oliver 2425 (BOL, NBG); Sperrgebiet South, 1.5 km north of Grillenthal (-

CD), JOrgens 28103 (PRE).

2616 (Aus): 56 mls [90.1 km] west of Aus, Schotterhang (—CB), Leippert 4080 (WIND).

—2715 (Bogenfels): ca. 5 km north of Pomona (—AB), Burgoyne 8270 (PRE); Pomona (—AB),

Dinter 6366 (BM, BOL, NBG 2 sheets, K, PRE, S, WIND); along road to Pomona (—AB),

Mannheimer et al. CM899 (WIND); Rote Kuppe, Chamais road (—BC), Bartsch SB 1043

94 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(WIND); 13 km west of Bogenfels houses, at ridge running south of road (—BC),

Mannheimer CM961 (WIND); Klinghardt Mountains (—BD), Dinter 3980 (BOL, PRE);

Boegoeberg (—DD), Dinter 6574 (B, photo!, BM!), Mannheimer CM1015 (WIND); summit of the Boegoeberg (—DD), Williamson 2594a (BOL 2 sheets).

—2816 (Oranjemund): Brandkaros (—BC), Venter 8863 (PRE); 0.5 km east of Brandkaros on road to Anniesfontein (—DA), Botha 3201 (PRE); Richtersveld, Kortdoorn (—DA), Jurgens

22585 (PRE); Witbank (—DC), Pillans 5149 (BOL, K).

Precise locality unknown: Nautilus, Kinges 2580 (PRE); Summit of Kortderm, Williamson

3376 (BOL).

12 14 16 18 20 22_24_218 30 32 ,_, AllialirlarilE 1111MISITIMIIM 11111ammi j l8 1811111111111111111 1117111 IIVIII imi 1111211111111111■ 111111111111111■ IL ' i IMO WINK"MEI mommo •■ 1111611, ■111 ' isainiul Ilidi il aim111 VIII111111r1111 Mill II II MUM IN ipmji ' " 1111111111111111i lir LI Fans itimmadom ' 11111111111111 Iffell 5111 111111111111111 nal • -MI 11111111/41111 lissalmareLeIIII li NEE orinin28 1111115111111 II NI VII 1111 1 Rilm El MEM II 11111 "11111111111111 v 4 110 ; TN, ■im A - 1 I l 4 :7°71 TPrlit lli oi lI 21 171r7716.1 1 18 m 20mmm. 22 24 26 28 30 32 34 36

FIG. 6.7. Known geographical distribution of Calobota acanthoclada.

95 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

2. CALOBOTA CINEREA (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia cinerea E.Mey., Comm. Pl. Afr. Austr. 1: 35.

Feb. 1836; Benth. in Hook. Lond. J. Bot. 3: 359. 1844.; Walp., Rep. Bot.

Syst.: 453. 1845.; Harv. in Harv. and Sond., Fl. Cap. 2: 87. 1862.; Merxm

and Schreiber, FSWA 60: 67. 1970.; Goldblatt and Manning, Cape Pl.: 493.

2000.—TYPE: SOUTH AFRICA, hills near Noagas [grid unknown], DrOge an

"III, B" (lectotype: P!, flowering specimen on the left, here designated;

isolectotype: BM!, K!, P!). [Note: This specimen was annotated by Meyer

himself and is chosen as lectotype.]

Lebeckia cinerea var. schaferi Dinter, Plantae novae Schaferianae: 342. 1920.—

TYPE: SOUTH AFRICA, "Granitberg zwischen Bogenfels and Prinzenbucht"

[2715 AD], Schafer 598 (Bt?, specimen not located).

Erect, multi-stemmed, unarmed shrub up to 1.5 m in height. Branches green; young branches sericeous or silky to tomentose; older branches sericeous or silky with light brown bark. Leaves digitately trifoliolate; petiole 3-14 mm long, longer or shorter than leaflets; leaflets widely obovate to elliptic or oblanceolate, alternate, sericeous on both surfaces, subsessile, terminal leaflet 5-21 mm long, 1.5-6.0 mm wide, lateral leaflets 3-17 mm long, 1.5-5.0 mm wide; apex recurved-mucronulate to acute; base cuneate. Inflorescence 30-200 mm long, racemose, with 5 to 30 flowers; pedicel 2-3 mm long; bract 1-4 mm long, linear, sericeous; bracteoles 1-4 mm long, linear, sericeous. Flowers 11-16 mm long, pale to bright yellow. Calyx 5-8 mm long, sericeous; tube 3.5-5.0 mm long; lobes 1.5-3.5 mm long, deltoid.

Standard 11.0-13.5 mm long; claw linear, 3.5-4.5 mm long; lamina ovate, 8-10 mm long, 7-11 mm wide; apex obtuse; dorsal surface densely pubescent. Wings 9-13

96 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

mm long; claw 2.5-3.5 mm long; lamina oblong, shorter than keel, 5.5-8.5 mm long,

3-4 mm wide, glabrous or rarely pilose, with 5-8 rows of sculpturing. Keel 11-14

mm long; claw 3.5-4.5 mm long; lamina boat-shaped, 7.5-10.0 mm long, 4.5-5.5

mm wide, pilose. Pistil subsessile to shortly stipitate, pubescent; ovary linear, 6-8

mm long, 0.8-1.2 mm wide with 10 to 16 ovules; style longer than ovary, 5.0-8.5

mm long. Pods linear, semi-terete, subsessile to shortly stipitate, 15-40 mm long, 2-

4 mm wide, ±5 to 6-seeded, tomentose, dehiscent. Seeds oblong-reniform to

reniform, 2-3 mm long, 1.5-2.0 mm wide, mature seeds light brown to orange, often

mottled with brown, surface smooth (Fig. 6.8). Flowering time: Mainly flowering in spring (August to November) and fruiting within the same period. Some specimens collected in Namibia were flowing and fruiting in April and May.

Diagnostic characters—Calobota cinerea is similar to Calobota linearifolia,

but differs in the low, spreading habit as opposed to the virgate or erect habit of C. linearifolia. The leaves are trifoliolate and obovate to elliptic or oblanceolate as opposed to the simple linear to oblanceolate or spathulate leaves of C. linearifolia. In

C. cinerea the inflorescences are more densely flowered with up to 30 flowers (up to

nine flowers in C. linearifolia), the pods are tomentose and not constricted between the seeds, and the seeds light brown to orange or mottled with brown (pods sericeous, constricted between the seeds and the seeds light brown or grey in C. linearifolia).

Distribution and habitat—Calobota cinerea occurs from south-western

Namibia southwards to South Africa as far as Clanwilliam (Fig. 6.9). It occurs on well-drained, sandy, loamy soil or red sand often next to rivers, at altitudes of between ca. 100 m and 500 m. It is recorded to be grazed by livestock or gemsbok.

According to Mucina et al. (2006), C. cinerea is an important component of

97 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Richtersveld (SKr 6 Stinkfontein Eastern Apron Shrubland) and Namaqualand

Sandveld (SKs 2 Northern Richtersveld Yellow Duneveld, SKs 5 Richtersveld Red

Duneveld, Sks 6 Oograbies Plains Sandy Grassland, SKs 8 Namaqualand Coastal

Duneveld).

1•11■111

FIG. 6.8 Morphology of Calobota cinerea: (al—a2) leaves in abaxial view; (b1) bract; (b2) bracteoles; (c) flower in lateral view; (d) standard petal; (e) outer surface of the calyx (upper lobes to the left); (f) wing petal; (g) keel petal; (h) androecium; (0) long, basifixed anther;

(i2) intermediate carinal anther; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view;

(k2) pod in dorsal view.

Voucher specimens: (al) Boatwright et al. 136 (JRAU); (a2—k2) Boatwright et al. 150

(JRAU). Scale bars: (al, a2, kl, k2) 1 cm; (bl—j) 1 mm.

98 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Additional Specimens Examined

—2518 (Tses): Kovisberge (—BB), Wa// 2 (S).

—2615 (Luderitz): Angra Pequena (—BA), Marloth 4809 (PRE), Marloth 14109 (K);

Kovisberge (—CB), Dinter 6292 (BM, BOL, K, NBG); Halenberg (—CB), Merxmiiller and

Giess 3089 (WIND).

—2616 (Aus): near Aris on the Gariep (—BD), Di-0e s.n. "III, B"(BM, P); Farm Klein-Aus (-

CA), Merxm011er and Giess 2998 (PRE, WIND); 15 mls [24.14 km] west of Aus (—CB), Giess and Van Vuuren 833 (K, WIND).

—2715 (Bogenfels): Pomona (—AB), Dinter 6346 (BM 2 sheets, BOL, K, NBG 2 sheets,

PRE), Wall s.n. (S); Pomona area, west facing slope ±15 km from houses going east (—AB),

Mannheimer et al. CM889 (WIND); Prince of Wales Bay (—AB), Marloth 5248 (PRE); Granite

Mountain (Granietberg) near Bogenfels (—AD), Schafer 13 (NBG).

—2716 (Witputz): Diamond Area 1, Inselberg, at beacon 992, end of Uguchab River (—CA),

Germishuizen 10170 (PRE).

—2815 (Diamond Area): Klinghardt Mountains (—BB), Dinter 3918 (BM, BOL, NBG, PRE, S).

—2816 (Oranjemund): ±40 km from Brandkaros (red dune area) near farm fence (—BC),

Petersen s.n. (NBG); Oranjemund (—CB), Clark 8 (WIND); sandy depressions north of

Witbank (—DA), Pillans 5221 (BOL, K); 38 mls [61.14 km] north of Port Nolloth on way to

Alexander Bay (—DA), Werger 517 (K, PRE); Goariepvlakte (—DB), JOrgens 22334 (PRE);

35 km from Port Nolloth on road to Alexander Bay at Holgat River (—DD), Germishuizen

4772 (PRE, WIND); Holgat (—DD), Jurgens 22734, 28804 (PRE).

—2820 (Kakamas): Between Augrabies and Anenous (—CB), Bolus 428 (BM, BOL, K, NBG,

PRE), Bolus 6548 (BOL, K).

—2917 (Springbok): Anenous flats, ca. 8 km west of Farm Grasvlakte (—AB), Goldblatt and

Manning 9461 (K, NBG); on road to Port Nolloth from Steinkopf (—AC), Boatwright et al. 150

(JRAU); sandy slope 40 km east of Port Nolloth (—AC), Goldblatt and Manning 9289 (NBG);

15 mls [24.14 km] north of Port Nolloth (—AC), Herre s.n. (NBG); between Kaus, Natvoet

99 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

th and Doornpoort (—AD), DrOge s.n. (S); Farm Nuwefontein 147 north-west of windmill at 28 mile (—DB), Le Roux 4038 (NBG).

—3017 (Hondeklipbaai): Taaibosvlei in dunes (—BC), Marloth 12418 (NBG, PRE).

—3118 (Vanrhynsdorp): Hardeveld south-west of Nuwerus (—AB), Marloth 8233 (NBG, PRE); sanddunes near Doorn River (—BB), Marloth 2637 (NBG, PRE); Farm Quaggaskop, 16 km on Douse the Glim turn-off, 24 km north of Vanrhynsdorp (—BC), Le Roux 2066 (NBG); towards "Douse the Glim", Knersvlakte (—BD), Stirton 9373 (PRE); Lutzville (—CB), Stirton

6062 (PRE); Vredendal road (—DA), Barker 5692 (BOL, NBG 2 sheets); Farm Liebendal (-

DA), Hall 3700 (NBG 2 sheets); between Vredendal and Lutzville (—DA), Liickhoff s.n.

(NBG); 53 km from Clanwilliam to Vanrhynsdorp (—DA), Stirton 5944 (K, PRE); Farm Spes

Bona, Inedendahl (—DC), Department of Agriculture 29 (PRE); Klawer (—DC), Henrici 3344

(PRE); western aspect of koppie, Vanrhynsdorp road near Klawer (—DC), Lavis 20235

(BOL).

—3218 (Clanwilliam): 8 km from Clanwilliam on road to Vanrhynsdorp (—BB), Boatwright et al. 136, 210 (JRAU); 5 km north of Clanwilliam (—BB), Grobbelaar 2004 (PRE); 10 km before

Clanwilliam from Vanrhynsdorp (—BB), Van Wyk 2598 (JRAU 2 sheets, PRE).

Precise locality unknown: without locality, DrOge s.n. (K); Sandveld, Richtersveld, Herre s.n. sub STE 11879 (NBG); Andaus Poort, Marloth 12250 (NBG, PRE).

100 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

12 14 16 18 20 22 24 26 28 30 32 Alianosal. 11111111 WNW in la era Immo manu18 1 Ramoul Mill MI IMO 111111011111111 0 MIL! ' 11111111111111 I SPA OM ' 11111111111111111 • Fiiii5O11111111 1111111111111 Op min art 111111111111 II IN mitill1111111911111111014 , Is RIM II 11 _MID 1111111ffinito pm orza CIAM HE Inlyilin ■111.31 MOE 111116111111111111 111111111111011113.11111111 Ill jess■11111111111111 # ■ 1111111•25 . 4 ■VI " 1 111o1E1 MI LM 11m111 ii1 2 111II14III11111 MI .1in AMUi1samorin mu ma u t 1012a ir1116 2sill0 z / 26 28 30 Pi3 4 Pric

FIG. 6.9. Known geographical distribution of Calobota cinerea.

3. CALOBOTA CUSPIDOSA (Burch.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Spartium cuspidosum Burch., Tray. S. Africa 1: 348.

1822. Genista cuspidosa (Burch.) DC., Prodr. 2: 147. 1825. Lebeckia

cuspidosa (Burch.) Skeels, U.S. Dept. Agric. Bur. Pl. Industry, Bull. 233: 38

(1911).—TYPE: SOUTH AFRICA, Between `Gatikamma' and `Klaarwater

[now Griquatown; 2823 CC], Burchell 1697 (holotype: K!).

Lebeckia cuspidosa (Burch.) Druce, Rep. Bot. Exch. Cl. Brit. Isles 1916: 631 (1917)

nom Neg.

Lebeckia macrantha Harv. in Harv. and Sond., Fl. Cap. 2: 83-84. 1862; Wilman,

Preliminary checklist of the flowering plants and ferns of Griqualand West

101 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(southern Africa): 53. 1946.—TYPE: SOUTH AFRICA, 'Zooloo country' [grid

unknown], Miss Owen s.n. (lectotype: TCD, photo!, designated here). [Note:

This is the only specimen cited by Harvey in the original description of this

species and was also seen by him. It is therefore designated as lectotype.]

Erect, multi-stemmed spinescent shrub up to 4.0 m tall. Branches green; young branches sparsely pubescent; older branches pubescent to glabrous with light brown bark. Leaves unifoliolate or rarely trifoliolate; petiole short, 1-2 mm long

(up to 5 mm long when trifoliolate); alternate, upper and lower surfaces pilose, linear to slightly obovate, 6-9 (-15) mm long, 2-3 (-6) mm wide; apex obtuse to shallowly retuse; base attenuate. Inflorescences (25–) 40-60 (-90) mm long, racemose, with

(4–) 6 to 12 (-24) flowers; pedicel 2-4 (-5) mm long; bract 0.9-1.2 mm long, triangular to slightly linear; bracteoles 0.5-0.8 (-1.0) mm long, slightly ovate.

Flowers (15–) 16-18 (-20) mm long, bright yellow. Calyx (3.8–) 4.4-5.4 mm long, pubescent on the outer surface; tube (2.5–) 3-4 mm long; lobes narrowly triangular,

(1.2–) 1.8-2.4 (-2.9) mm long, deltoid. Standard 11.0-18.5 mm long; claw linear, 3-

6 mm long; lamina ovate, 8.5-12.5 mm long, (7.5–) 8.0-10.0 mm wide, glabrous; apex obtuse. Wings 10-16 mm long; claw (3–) 5-6 mm long; lamina oblong, shorter than the keel, (7.5–) 9.0-10.5 mm long, (3.0–) 4.0-5.5 mm wide, glabrous, with 5 rows sculpturing. Keel 13.5-19.0 mm long; claw (3.5–) 5.0-7.0 mm long; lamina oblong, 10.0-12.5 mm long, 5-6 mm wide, glabrous. Pistil shortly stipitate, glabrous; ovary linear, 7-14 mm long, 1.0-1.6 mm wide with 9 to 11 ovules; style longer than ovary, 5.5-9.0 mm long. Pods oblong to slightly falcate, laterally compressed, shortly stipitate, (21–) 28-49 (-53) mm long, (5–) 6-10 mm wide, ±4 to 6-seeded, glabrous, indehiscent. Seeds reniform, 3-4 mm long, 2.5-3.0 mm wide, mature

102

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA seeds light brown, surface smooth (Fig. 6.10). Flowering time: This species is mainly spring and summer flowering (October to January), but flowering also sometimes occurs in winter (April to May).

12 i3

FIG. 6.10 Morphology of Calobota cuspidosa: (al—a2) leaves in abaxial view; (b) outer surface of the calyx (upper lobes to the left); (c) flower in lateral view; (d) standard petal; (e) wing petal; (f) keel petal; (g) pistil; (h) androecium; (i1) long, basifixed anther; (i2) intermediate carinal anther; (i3) short, dorsifixed anther; 01) bract; (j2) bracteoles; (kl) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al) MacDonald 76/3 (NBG); (a2) Ethel Anderson 591 (BOL);

(b, i) Gubb 12584 (PRE); (c—h) Van Wyk 3055 (JRAU); (k1—k2) Boatwright et al. 92 (JRAU).

Scale bars: (al, a2, kl, k2) 1 cm; (b—j2) 1 mm.

103 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—Calobota cuspidosa is similar to C. psiloloba, but the large flowers, pods and seeds are very distinctive for the former (in C. psiloloba the flowers, pods and seeds are much smaller). Calobota cuspidosa reaches heights of up to 4 m, while C. psiloloba only reaches heights of 1.2 m. The young stems in this species are normally densely tomentose and the older ones remain at least slightly pubescent (in C. psiloloba the branches are glarbous or only slightly pubescent).

Distibution and habitat—Calobota cuspidosa occurs in the Northern Cape

Province and is centred around the towns of Kuruman and Griquatown at altitudes of between 975 m and 1700 m. It is mainly found in rocky, loamy or sandy soils (Fig.

6.11). According to Rutherford et al. (2006) this species is an important, prominent component of Kuruman Mountain Bushveld (SVk 10 Eastern Kalahari Bushveld) where it is common in low, open bushveld.

Additional specimens examined

—2723 (Kuruman): Batlaros, Bosjesmans doom (—AC), SiIke 16785 (BOL); ca. 10 km west of Kuruman on the road to Olifantshoek (—AD), Dlamini, Nkuna and Van Wyk EvW577 (K);

Kuruman (—AD); Pole-Evans 2108, 2480 (K, PRE), Van Son s.n. sub TRV 31766 (PRE); On road to Olifantshoek (—AD), Germishuizen 5614 (BOL, PRE); 3 mls [4.8 km] east of

Kuruman (—AD), Grobbelaar 1112 (PRE); Kuruman Municipal Reserve (—AD), MacDonald

76/3 (J, NBG, PRE); Buta Asbestos Mine (—AD), Peeters, Gericke and Burelli 264 (J, PRE); ca. 12 km from Kuruman to Vryburg (—AD), Van Wyk 3053 (JRAU); 11 km from Kuruman to

Vryburg (—AD), Van Wyk 3055 (JRAU); Kuruman Hills (—AD), Verdoorn and Dyer 1763

(PRE, K); Vredebron, Blikfontein (—CA), Gubb KMG 10989 (PRE); 10 km west of Kuruman on road to Upington (—CA), Van Wyk BSA1241 (PRE, WIND); 30 km from Kuruman to

Billingshurst (—CD), Arnold and Musil 500 (PRE); Hillsides from Griquatown to Khosis (—

104 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

CD), Bryant J379 (PRE); 30 km south-east of Kuruman (—DA), Jordaan CBK 26 (PRE);

Road from Reivilo to Kuruman, farm Mattana (—DB), Joffe 650 (PRE).

—2724 (Taung): Blesmanspost (—CA), Burtt-Davy 9665 (K).

—2820 (Kakamas): Cape Plateau, Baviaans "Kloor [= Baviaans Krantz], (—DA), Hutchinson

3001 (BOL, K, PRE).

—2822 (Glen Lyon): Hills at Vaalwater (—DA), Acocks 2455 (BOL 2 sheets, K 2 sheets); between Griquatown and Volop, Bakenkop, north of tarred road (—DB), Gubb KMG 10988

(PRE).

—2823 (Griquatown): Postmasburg (—AC), Wilman s.n. (K); 19.5 km from Griquatown to

Campbell (—AD), Boatwright et al. 92 (JRAU); between Postmasburg and Griquatown (—CA),

Hutchinson 3039 (BM, BOL, K, PRE); Jasper veld (—CB), Wilman 4685 (K); Griquatown , commonage, the Asbestos Hills (—CC), Acocks H1049 (PRE); Griquatown (—CC), Barret-

Hamilton s.n. (BM), Botha 2996 (PRE), Burchell 1697, Pole-Evans 22 (K, PRE); 1 ml [1.6 km] east of Griquatown (—CC), Leistner 890 (K, PRE); ca. 5 km from Griquatown on R64 on the road to Upington (—CC), Van Wyk 2534 (JRAU 3 sheets); Broken Hill, north of Campbell

(—DA), Germishuizen 8523 (PRE).

—2824 (Kimberly): Koopmansfontein, Farm Geluk (—AA), Hanekom 2214 (K, PRE);

Schmidtsdrift, Ruigtefontein (—CA), Gubb KMG 12584 (PRE); 9 mls [14.5 km] west of

Schmidtsdrift (—CA), Leistner and Joynt 2690 (K, PRE); Schmidtsdrif (—CA), Wall s.n. (S);

Barkly West, Tierfontein (—DA), Acocks 719 (K 2 sheets); 14 mls [22.5 km] from Vryburg on

Schweizer Reneke road (—DD), Carr 5 (PRE).

—2922 (Prieska): 6 mls [9.6 km] west of Abrahmsdam (—BA), Codd 1263 (K, PRE).

—2923 (Douglas): Mazelsfontein Griqualand West (—BA), Anderson 591 (BOL, PRE); near

Douglas (—BB), Kotze 783 (PRE).

Precise locality unknown: Namaland, Marloth 1042 (PRE, 2 sheets).

105 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

12 14 16 18 20 22 24 26 28 30 32in p 1812sammaymmErammi in ii. nommol_■wiliaromi voli ■ 11111111111% MIN It"0 IMMO 1 ' 1111113111111■■■1 112111 Malt ' 1111161 ■ min■ ow 111111111111111111 K2 ILIIIIIIIN■ , '' 11111111111111111 I "51111111111-- m nal a um 11 la 1170M IIIAMINIF211111 ' 1 MUNI ICEN1 MN 1/11111111111 ■•116 III IE 3 nairM1101111 111 l • ■C:731 MEI 111■1111111128 1111111111111*Mi MA VIII In IM11111111111 111 -MI S ■ ■■ " 11111111 all ■1111 aal l I" ' - ' 111 gill • ■ 4 mos 11111M, OM §11 III n ■ 200E11 PM NM .1111111111 ,Finorairim 0 ,0011a simi 101 ,110111 10 12 14 16 18 116"111 22 24 26 28 30 3-2 --- irri

FIG. 6.11. Known geographical distribution of Calobota cuspidosa.

4. CALOBOTA cy-risoi DEs (Berg.) Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 191 Jan.

1836. Spartium cytisoides Berg., Descr. Pl.: 199. 1767; L.f., Suppl. Pl. 320

1781. Lebeckia cytisoides (L.f.) Thunb., Nov. Gen.: 143. 1800; Prodr. Pl.

Cap.: 122. 1800; Willd. Sp. Pl.: 948. 1800; Thunb., Fl. Cap.: 562. 1823; DC.,

Prodr. 2: 137. 1825; E.Mey., Comm. Pl. Afr. Austr. 1: 36. Feb. 1836; Benth. in

Hook. Lond. J. Bot. 3: 359. 1844; Harv. in Harv. and Sond., Fl. Cap. 2: 87.

1862; Bond and Goldblatt, J. S. Afr. Bot. 13: 289. 1984; Goldblatt and

Manning, Cape Pl.: 493. 2000.—TYPE: SOUTH AFRICA, e Cap. B. Spei [grid

unknown], Grubb s.n. sub Bergius 236.57 (holotype: SBT!).

106 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Ebenus capensis L., Mant II: 264. 1771. Cytisus capensis Lam., Encycl. Meth. 2:

249 (1786); Lebeckia capensis (L.) Druce Rep. Bot. Exch. Cl. Brit. Isles 1916:

631. 1917, nom. illeg.

Crotalaria pulchella Andr., Bot. Repos. 6: plate 417 (Jan–Mar 1805); Sims, Bot.

Mag. t. 1699 (1824); DC., Prodr. 2: 134 (1825). Lebeckia pulchella (Andr.)

Walp. in Linnaea 13: 477 (1839). Calobota pulchella (Andr.) Eckl. and Zeyh.,

Enum. Pl. Afr. Austr. 2: 191 (Jan. 1836).—TYPE: Andr. Bot. Repos. 6: plate

417. 1805. (iconotype). [Note: According to Stafleu and Cowan (1976) Henry

Andrews was a botanical painter and engraver. His artworks were based on

living specimens, but no herbarium specimens are known to exist. Due to the

fact that the painting leaves no doubt as to its identity we here choose it as

iconotype.]

?Lebeckia marginata E.Mey., Comm. Pl. Afr. Austr. 1: 35. Feb.1836; Walp. in

Linnaea 13: 478. 1839; Benth. in Hook. Lond. J. Bot. 3: 359 1844; Harv. in

Harv. and Sond., Fl. Cap. 2: 89 1862.—TYPE: without locality, Drege s.n.

(S!). [Note: This species was described by Meyer (1836) without listing

locality information. The only specimen located is in S and is sterile, but the

mucronulate apices of the leaves look similar to those of Calobota cytisoides.

This taxon might merely represent a synonym of the latter as suggested by

Ecklon and Zeyher (1836), but its identity remains unclear.]

Erect, diffuse, multi-stemmed, unarmed shrub up to 2 m in height. Branches green; young branches sericeous; older branches sericeous to pilose with brown bark. Leaves digitately trifoliolate to rarely 5-foliolate; petiole (8–) 10-40 mm long, as long as or shorter than leaflets; leaflets elliptic to oblanceolate, alternate, sericeous,

107 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA subsessile, terminal leaflet (12–) 14-55 mm long, 3-10 (-17) mm wide, lateral leaflets (11–) 13-45 mm long, 3-9 (-15) mm wide, apex mucronulate to somewhat truncate, base angustate. Stipules absent. Inflorescence 70-255 (-300) mm long, racemose, with 7 to 25 flowers; pedicel relatively long, 3-11 mm long; bract 3-6 mm long, elliptic to ovate, pubescent on outer surface; bracteoles 1-4 (-5) mm long, narrowly ovate to linear, pubescent on outer surface. Flowers 18-35 mm long, bright yellow. Calyx 9-13 mm long, glabrous or very rarely pilose; tube 6.5-10.5 mm long; lobes 1.5-4.0 mm long, deltoid. Standard 18.5-27.0 mm long; claw linear, 4-9 mm long; lamina widely ovate, 13-19 mm long, 14.5-26.5 mm wide; sericeous along dorsal midrib and apex; apex obtuse. Wings 18-28 mm long; claw 5.5-9.0 mm long; lamina oblong to ovate, shorter or ± as long as keel, 12.5-17.5 mm long, 7-12 mm wide, glabrous, with 9-16 rows of sculpturing; apex obtuse. Keel 19.0-29.5 mm long; claw 6.0-12.5 mm long; lamina boat-shaped, 13-20 mm long, 6-10 mm wide, sericeous on terminal parts; apex obtuse. Pistil subsessile to shortly stipitate, glabrous; ovary linear, 14-22 mm long, 1.2-2.0 mm wide with 20 to 34 ovules; style longer than ovary, 7.5-13.5 mm long, glabrous. Pods linear, sometimes somewhat clavate, semi-terete, subsessile to shortly stipitate, 45-70 mm long, 3-5 mm wide,

±5 to 18-seeded, glabrous, dehiscent. Seeds oblong-reniform, 2.8-3.5 mm long,

2.0-2.6 mm wide, mature seeds light brown, surface smooth (Fig. 6.12). Flowering time: Mainly flowering and fruiting in spring and late winter (June to November). One flowering specimen was collected in April.

Diagnostic characters—Calobota cytisoides is similar to C. thunbergii, but differs in the mucronulate leaf apices, long pedicels, larger flowers, sometimes ovate bracts and bracteoles, glabrous or pilose calyx, larger and glabrous pistil and pods

(in C. thunbergii the leaf apices are acute, the pedicels shorter, the flowers smaller,

108 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA the bracts and bracteoles always linear, the calyx sericeous and the pistil and pods shorter and pubescent).

FIG. 6.12 Morphology of Calobota cytisoides: (a, b) leaves in abaxial view; (c) flower in lateral view; (d) outer surface of the calyx (upper lobes to the left); (e) standard petal; (f) wing petal; (g) keel petal; (h) androecium; (i1) long, basifixed anther; (i2) intermediate carinal anther; (i3) short, dorsifixed anther; (j) pistil; (k1, k2) bracts; (k3—k5) bracteoles; (11) pod in lateral view; (12) pod in dorsal view.

Voucher specimens: (a) Van Wyk 3117 (JRAU); (b) Le Roux 2 (JRAU); (c—k1, k4, k5) Boatwright et al. 114 (JRAU); (k2, k3) Barker 9694 (NBG); (11-12) Van Wyk 2439

(JRAU). Scale bars: (a, b, 11, 12) 1 cm; (c—k5) 1 mm.

109 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Distribution and habitat—Calobota cytisoides can be found from

Oudtshoorn in the Eastern Cape through to the Western Cape around Montagu and

Worcester and south through Wupperthal and Clanwilliam as far as Vanrhynsdorp

(Fig. 6.13). It has a more southern distribution than the closely related C. thunbergii and has been collected at altitudes of between 20 m and 1000 m in renosterveld vegetation on sandstone, shale, well-drained sand and clay. Calobota cytisoides is often found on western, southern or south-western slopes, along rivers or in riverbeds and also along disturbed roadsides. Rebelo et al. (2006) and Mucina et al.

(2006) list this species as an important component of Shale Fynbos (FFh 8 Montagu

Shale Fynbos), Shale Renosterveld (FRs 9 Swartland Shale Renosterveld, listed as

C. cinerea), and Rainshadow Valley Karoo (SKv 7 Robertson Karoo).

Additional specimens examined

—3118 (Vanrhynsdorp): "Pillar se punt", northern point of Bokkeveld Mountains, plateau on farm Paardekraal (—BB), Van Wyk 1248 (NBG, PRE, UPS); Vleikraal, east of Klawer (—DA),

Walters 178 (K, NBG); 17.2 km from caravan park, Clanwilliam on old road to Citrusdal (-

DC), Gess 91/92/1 (GRA); 23.9 km from Clanwilliam (—DC), Gess 91/92/10 (GRA); Klawer

(—DC), Lavis 29233 (BOL); 29 km from Clanwilliam to Vanrhynsdorp (—DC), Stirton 5943 (K,

PRE).

—3119 (Calvinia): Vanrhyns Pass (—AC), Acocks 14717 (K, PRE 2 sheets), Bond 1147

(NBG), Esterhuysen 5312 (BOL), Schutte 286 (JRAU 2 sheets), Van Wyk 3117 (JRAU);

Skuinshoogte Pass (—AC), Niemand 1 (JRAU); Niewoudtville, Oorlogskloof Nature Reserve

(—AC), Pretorius 105 (NBG, PRE, WIND); Nieuwoudtville (—AC), Schmidt 636 (K);

Meulsteenvlei near Nieuwoudtville (—AC), Taylor 3942 (NBG); Lokenberg, Stinkfontein Hills

(—CA), Leistner 451 (NBG, PRE).

110 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

—3217 (Vredenburg): Vredenburg (—DB), Grobbelaar 2549 (PRE); above Stompneus (—DB),

Booysen 141 (NBG); 1 ml [1.61 km] east of Paternoster (—DD), Barker 9694 (NBG 2 sheets); Kasteelberg (—DD), Strid and Strid 38078 (NBG).

—3218 (Clanwilliam): Graafwater (—BA), Compton 24209 (NBG); Bulshoek Barrage, Olifants

River Valley (—BB), Barker 6426 (NBG 2 sheets); Clanwilliam (—BB), Bayliss BR1593 (GRA,

K, PRE, S), Davidson s.n. (J), Ecklon and Zeyher 17 (NBG), Eliovson 161 (J), Henrici 2105

(PRE), Levyns 1282 (BOL); along road from Clanwilliam to Algeria (—BB), Boatwright et al.

207 (JRAU); Olifants Dam (—BB), Compton 20015 (BOL, NBG 2 sheets); Clanwilliam south of town next to Olifants River (—BB), De Vos 1717 (NBG, PRE); near Clanwilliam on road to

Biedouw Valley (—BB), Joffe 959 (PRE); Ramskop Wild Flower Garden (—BB), Le Roux

2476 (BOL, NBG); rocky hills near Clanwilliam (—BB), Parker 3589 (BOL, K, NBG); Pakhuis

Pass (—BB), Schulte 262, 264 (JRAU); Bakhoogte near Clanwilliam (—BB), Stirton 9195

(PRE); Nardouwskloof Farm (—BB), Stirton 9364 (PRE); Boschkloof (—BB), Thorne s.n.

(NBG); between Koperfontein and Malmesbury (—BC), Barker 4067 (BOL, NBG); on

Citrusdal road 8 mls [12.87 km] from Clanwilliam (—BD), Gillett 4082 (BOL, PRE); Olifants

River Valley, 23 km north of Citrusdal (—BD), Hugo 426 (K, NBG, PRE); Rondegat River, 10 km from Algeria to Clanwilliam (—BD), Stirton 5926a (K, PRE); roadside between Citrusdal and Clanwilliam (—BD), Taylor 10630 (NBG, PRE), Van Wyk 2313 (JRAU 2 sheets); halfway between Clanwilliam and Citrusdal (—BD), Van Wyk 2439 (JRAU 2 sheets), Van Wyk 2441

(JRAU); Olifants River Valley between Klawer and Citrusdal (—BD), Wilman 828 (BOL); St

Helena Bay, Steenberg Cove (—CA), Wisaru 1710 (NBG); on road between Misverstand

Dam and Velddrif (—CC), Le Roux 2 (JRAU 2 sheets); 9 mls [14.48 km] from Velddrif to St.

Helena Bay (—CC), Marsch 1272 (K, NBG); 2 mls [3.22 km] north-west of Sauer Post Office

(—CD), Acocks 14534 (K, PRE 3 sheets); Moutonsvlei, west of Piquetberg Mountain complex (—DA), Goldblatt 7197 (NBG, PRE, S); base of north slopes below nek at head of

Kapteinskloof, Piquetberg (—DA), Pillans 7701 (BOL); Het Kruis (—DA), Stephens and Glover

8686 (BM, NBG 2 sheets, K, PRE); Kapteinskloof (—DA), Stirton 6133a (K, PRE); near

Warmbad (—DB), Hutchinson 1132 (BM, BOL, K, PRE); Olifants River Valley (—DB), Marloth

111 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

7121 (PRE); Piekenierskloof Pass (—DB), Van Wyk 3219 (JRAU); 3 km south of Piquetberg

(—DC), Ihlenfeldt 1023 (PRE); 3 mls [4.83 km] from Piquetberg up Versfeld Pass (—DC),

Thompson 762 (K, NBG, PRE); outside Piquetberg (—DC), Thorne s.n. (NBG); Piquetberg (-

DD), Bolus 13536 (BOL, PRE 2 sheets), Grant 4714 (PRE), Guthrie 2584 (NBG), Schlechter

5233 (BM, BOL, K), Schlechter s.n. sub TRV 3377 (PRE); roadside near Piquetberg (—DD),

Hafstrom and Acocks 2312 (PRE 2 sheets, S); 3 km south of Piquetberg (—DD), Ihlenfeldt

1023 (PRE); 2 mls [3.22 km] along Piquetberg to Velddrif road (—DD), Marsh 795 (NBG,

PRE).

—3219 (Wuppertal): pass into Biedouw Valley, 6.2 mls [9.98 km] from turnoff from

Clanwilliam-Calvinia road (—AA), Marsh 388 (K, NBG, PRE); Bottom of Biedouw Pass (-

AA), Mauve and Oliver 89 (NBG, PRE); Pakhuis (—AA), Salter 1660 (BM, BOL, K);

Pakhuisberg (—AA), Schlechter 139 (PRE), Schlechter 8657 (PRE), Schlechter 8659 (BM,

K, S); Pakhuis Pass (—AA), Stokoe s.n. sub SAM 52520 (NBG); Pakhuis Pass, east side near Big Bends and gravel pit (—AA), Taylor 11560 (NBG, PRE); Uitkyk Pass, into Biedouw valley (—AA), Thompson 2926 (K, NBG); Cederberg State Forest, Grootkloof (—AC), Andrag

272 (NBG, PRE); Nieuwoudt Pass (—AC), Barnes s.n. (BOL); Olifants River near Algeria (-

AC), Boatwright et al. 119 (JRAU) Cedarberg Reserve (—AC), Hubbard 34 (NBG);

Wupperthal, just outside town on road to Clanwilliam (—AC), Grobbelaar 2975 (PRE);

Wupperthal (—AC), Stirton 6399 (K, PRE); Garskloof, near Algeria (—AC), Stirton 9192

(PRE); near turn-off to Algeria on old National road (—AC), Van der Walt s.n. (NBG, PRE); ca. 8 km from Algeria on road to Clanwilliam (—AC), Van Wyk 2568 (JRAU 2 sheets, PRE);

Cedarberg, Forest Reserve, Langrug (—AC), Viviers 510 (NBG, PRE); Brakfontein (—AD),

Ecklon and Zeyher 1332 (K, PRE, S); south of Trekkloof Pass (—AD), Low 2942 (NBG);

Wuppertal (—AD), Thode A1987 (K, PRE), Thorne s.n. (NBG), Van Rooyen et al. 696 (NBG);

Warm Baths, Clanwilliam division (—CA), Edwards 252 (PRE); near Citrusdal (—CA), Steyn

391 (NBG 2 sheets); just north-east of Citrusdal (—CA), Strid and Strid 37340 (NBG).

—3318 (Cape Town): Riebeek Wes (—BD), Coffer s.n. (NBG); Riebeeck Castle (—BD), DrOge s.n. "III, D, a" (BM, K, S), Pillans 9788 (BOL), Stark and Co. s.n. (BOL).

112 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

—3319 (Worcester): Farm Welbedacht, Biedouw Vally (—AA), Johannesburg Botanical

Garden 3876 (PRE); Hex River (—BC), Bolus 5152 (K), Marloth 1605 (PRE), MacOwen 1605

(BM, K, NBG, UPS); main road passing De Dooms, east of town (—BC), Mauve and Oliver

136 (NBG, PRE); Hex River Pass (—BD), Eliovson 318 (J), Gray s.n. sub BOL 26052 (BOL,

K); 5 mls [8.05 km] from Worcester on Bains Kloof road (—CA), Story 2902 (PRE); Fairy

Glen Hiking trail, Worcester (—CB), Cupido 146 (NBG); 3.5 mls [5.63 km] from Worcester on road to Rawsonville (—CB), Grobbelaar 1183 (PRE); 6 km from Worcester on road to Ceres

(—CB), Grobbelaar 2796 (K, PRE); Worcester Veld Reserve (—CB), Henrici 3807 (BOL,

NBG), Midgley and Bosenberg 7 (NBG 2 sheets, PRE), Olivier 101 (NBG), Van Breda and

Joubert 2177 (K); Farm Vrolikheid, McGregor (—CB), Jooste 192 (NBG); Blinkberg Pass,

Tuinkloof (—CB), Stirton 5907 (K); National road bridge at Breede River (—CB), Taylor 3729

(BOL, K, NBG); Karoo Botanical Garden, Worcester (—CB), Barker 5944 (BOL, NBG 3 sheets), Dobay 52 (NBG), Niemand 14 (JRAU), Van Wyk 1408a (JRAU); Fairy Glen, lower slopes of Brandwag Mountains (—CB), Stirton 9143 (PRE); Brandwagt, in Fairy Glen Kloof

(—CB), Van Breda SKF574 (K, PRE); Robertson veld (—CB) Walters 380 (NBG); roadside

Worcester west en route Brandwacht (—CB), Walters 588 (NBG); Brandwacht (—CB),

Walters 1237 (NBG); hillside Voordenberg, near De Wet station (—CB), Walters 2481 (NBG); between Worcester and Stettyns (—CD), Rycraft 1732 (NBG); De Doorns (—CD), Moss 6798

(BM, J), Schmidt s.n. (NBG); Stettyns (—CD), Van Rensburg 386 (K, NBG 2 sheets, PRE);

Farm Doringkloof, southern foothills of Voetpadsberg (—DA), Morley 506 (NBG); Hex River

Kloof (—DA), Sidey 1878 (PRE, S 2 sheets); Farm Doringkloof, foothills of Voetpadsberg (-

DA), Van Wyk 66 (NBG, PRE, UPS); Jonaskop (—DC), Bayliss BR11228 (K, PRE); on plains between Assegaaikloof and Breede River (—DC), Bowie s.n. (BM); Robertson (—DD), Levyns

2812 (BOL); Vrolykheid Nature Reserve (—DD), Van der Merwe 2969a (K, PRE).

—3320 (Montagu): Kogmanskloof (—CC), Court 399 (GRA), Gillett 4521 (BOL), Mundt s.n.

(K); Bonnievale (—CC), Du Preez 321 (NBG); Rothman NBG221/34 (NBG), Van Breda and

Joubert 1948 (K); near Hassaquaskloof and Kogmanskloof (—CC), Ecklon and Zeyher 1333

(K, S); 15 mls [24.14 km] beyond Montagu on Barrydale road (—CC), Marais NBG525/62

113 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(NBG 2 sheets); Bonnievale, hills of Bokkeveld beds (—CC), Marloth 11811 (NBG 2 sheets,

PRE); 5 km before Montagu (—CC), Van Wyk 544 (NBG, PRE); Goudmyn, between

Bonnievale and Robertson (—CC), Van Wyk 2705 (JRAU); Olifantsberg (—CC), Viviers 422

(NBG 2 sheets, PRE); Scheepersrus (—CD), Johnson 147 (NBG 2 sheets); Barrydale (—DC),

Galpin 3922 (GRA, PRE), Morris 161 (BOL, NBG 2 sheets), Wall 3 (S 2 sheets); 15 mls

[24.14 km] west of Barrydale (—DC), Gillett 1897 (BOL, K); 20 km from Barrydale on road to

Montagu (—DC), Grobbelaar 2246 (PRE); 47 km from Montagu on road to Barrydale (—DC),

Grobbelaar 2782 (K, PRE); shaly hills at Barrydale (—DC), Hafstrom and Acocks 2314

(PRE); 19 mls [30.57 km] south of Barrydale (—DC), Martin 403 (NBG 2 sheets); alongside road up to Wildehondskloof Pass between Montagu and Barrydale (—DC), McDonald 1568

(NBG, PRE); crossroad at Barrydale (—DC), Nel s.n. sub STE 18105 (NBG); between

Barrydale and Montagu (—DC), Rycraft 2627 (NBG 3 sheets), Van Niekerk 582 (K); top of

Wildehondskloof Pass (—DC), Van Wyk 2651b (JRAU).

—3321 (Ladismith): Gamka Mountain Reserve, alongside road to Paardebond (—CB), Cattell and Cattell 169 (K, NBG, PRE).

—3322 (Oudtshoorn): 7 mls [11.26 km] south-west of De Rust (—BC), Theron 2071 (BOL, K,

PRE, UPS); Oudtshoorn (—CA), Besselaar s.n. (NBG); along roadside from Oudtshoorn to

De Rust (—CA), Boatwright et al. 114 (JRAU); Farm Die Krans (—CB), Dahlstrand 2004

(NBG, PRE), Dahlstrand 2312 (NBG, PRE); Oudtshoorn, near Farm Oude Muragie (—CB),

Vlok 2150 (PRE); Farm Doornkraal, 3 mls [4.83 km] east of De Rust (—DA), Dahlstrand 1469

(J, PRE).

—3418 (Simonstown): near Steenberg's Cove, Vredenburg (—BB), Taylor 1535 (NBG 2 sheets).

—3419 (Caledon): Brakfontein (—BB), Schechter 5279 (BM, BOL, S 2 sheets, PRE);

Krommerivier (—BB), Shaw 5624 (BOL, K).

—3420 (Bredasdorp): Breede River (—AA), Mundt 37 (K); 8 km from Stormsvlei to

Swellendam (—AA), Stirton 6147 (K, PRE); Stormsvlei (—AA), Taylor 4048 (NBG 2 sheets);

114 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Swellendam (—AB), Burchell 7475 (K); barren mountain ridges on the Riviersonderend at

Storm Valley, Hassaquaskloof and Breede River (—AB), Zeyher 2320 (PRE 3 sheets).

—3421 (Riversdale): near Platte Kloof (—AA), Muir 671 (PRE); Roadside near Riversdale (-

AB), Bayliss BR1B422 (PRE), Martin 9649 (GRA).

Precise locality unknown: Cape, Bowie s.n. (K); without locality, Drege s.n. (K, PRE); e

Cap. b. Spei, Sparrman s.n. (S), Thunberg s.n. (S), Thunberg s.n. sub UPS 16413, 16414

(UPS); without locality, Zeyher 2320 (K, PRE 2 sheets).

FIG. 6.13. Known geographical distribution of Calobota cytisoides.

5. CALOBOTA ELONGATA (Thunb.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Crotalaria elongate Thunb., Fl. Cap.: 571. 1823.—

TYPE: SOUTH AFRICA, 'Cam prope Bockeveld' [grid unknown], Thunberg

s.n. sub THUNB-UPS 16544 (lectotype: UPS!, designated here). [Note: This

115 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

is the only specimen in Thunberg's herbarium; it is annotated on the reverse

side of the specimen by him and so is here designated as lectotype.]

Lebeckia melilotoides R.Dahlgren, Bot. Notiser 120: 268-271. 1967; Bond.and

Goldblatt, J. S. Afr. Bot. 13: 290. 1984; Goldblatt and Manning, Cape Pl.: 493

2000.—TYPE: SOUTH AFRICA, Platfontein, east of Hottentots Holland Kloof

[3319 BB], H. Hall 177 (holotype: NBG!; isotypes: NBG!, PRE!, S!).

Erect to procumbent, multi-stemmed, unarmed shrub up to 1 m in height.

Branches green or light brown, young and older branches densely sericeous.

Leaves digitately trifoliolate; petiole longer or shorter than leaflets, 8-29 mm long;

leaflets widely obovate, alternate, sericeous, subsessile, terminal leaflet 13-32 mm

long, 8-20 mm wide, lateral leaflets 10-32 mm long, 6-16 mm wide, apex obtuse or

retuse, base cuneate. Inflorescence 80-432 mm long, spicate, with more than 100

flowers; flowers sessile, bract 5-7 mm long, linear, densely tomentose; bracteoles

2-5 mm long, linear, densely tomentose. Flowers 8-11 mm long, pale yellow. Calyx

4.5-6.0 mm long, tomentose; tube 3.5-5.0 mm long; lobes 1.1-1.7 mm long, deltoid.

Standard 10-12 mm long; claw linear, 2.5-3.5 mm long; lamina narrowly ovate to

elliptic, 7.5-9.0 mm long, 5-6 mm wide, pilose along dorsal midrib; apex obtuse.

Wings 10.5-12.0 mm long, claw 3.5-4.0 mm long, lamina oblong, as long as or

longer than keel, 6-8 mm long, 2.5-3.5 mm wide, glabrous, with 5-6 rows of sculpturing. Keel 9.5-12.0 mm long, claw 3.0-4.5 mm long, lamina boat-shaped,

6.5-8.0 mm long, 3.5-4.5 mm wide, glabrous. Pistil subsessile to shortly stipitate, densely pubescent, ovary linear to elliptic, 5.5-6.5 mm long, 0.6-1.0 mm wide with 3 to 6 ovules; style longer than ovary 6-7 mm long. Pods ovate to elliptic, semi-terete, subsessile, 5-8 mm long, 1.5-4.0 mm wide, 1 to 2-seeded, indehiscent. Seeds

116 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA reniform, 2.4-3.0 mm long, 1.3-2.0 mm wide, mature seeds light brown to light pink, often mottled with brown or grey, surface smooth (Fig. 6.14). Flowering time: Late spring to summer (October to November).

Diagnostic Characters—This species is similar to Calobota obovata, but differs in the trifoliolate leaves, dense, spicate inflorescences with smaller, sessile flowers, long, narrow bracts and bracteoles, very short calyx lobes, ovate to elliptic standard petal with a pilose midrib, wing petals that are longer than keel, fewer ovules in the ovary and shorter, tomentose pods with less seeds (in C. obovata the leaves are unifoliolate, the inflorescences racemose with fewer, the flowers pedicellate, the bracts and bracteoles ovate, the calyx lobes longer, the standard petal densely pubescent and widely ovate and the wing petals shorter than the keel.

Furthermore numerous ovules are present in the ovary and the pods are short- sericeous and many-seeded).

Distribution and Habitat—This species occurs between the Touws River and Ceres on sandy soil or dunes at ca. 600 m and has been recorded to be both avoided and grazed by livestock (Fig. 6.15).

Note—The conservation status of this species was assessed as vulnerable

(VU D1+2) by Raimondo et al. (in press). Calobota elongate is largely restricted to the Tanqua Karoo and it is estimated that less than 1,000 individuals are left. It is threatened by overgrazing and building activies.

117

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

j2 j3

FIG. 6.14 Morphology of Calobota elongata: (a) leaf in abaxial view; (b1) bracts; (b2) bracteoles (c) flower in lateral view; (d) outer surface of the calyx (upper lobes to the left);

(e) standard petal; (f) wing petal; (g) keel petal; (h) androecium; (i) pistil; (j1) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view;

(k2) pod in dorsal view.

Voucher specimens: (a—b, d—j) Van Wyk 2229 (JRAU); (c) Hall 177 (NBG); (k1—k2)

Van Breda 4486 (PRE). Scale bars: (a, k1, k2) 1 cm; (b—j3) 1 mm.

118 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Additional specimen examined

—3219 (Wuppertal): Ceres Karoo, near Beukes Fountain (—DC), Schlieben and Van Breda

9811 (K, NBG, PRE).

—3319 (Worcester): Verkeerdevlei (—BD), Van Wyk 2229 (2 sheets), 2230, 2562b (2 sheets),

2710 (JRAU); Farm Littouw (—CB), Van Breda 4461 (PRE).

—3320 (Montagu): near Railway line ca. 5 km south of Touws River (—AC), Van Breda 4486

(PRE).

Precise locality unknown: south end of Tanqua Karroo, Story 3606 (K, PRE).

HEIGHT ABOVE SEA LEVEL CD Over 1500 m CD 900 - 1500 m 300 - 900 m =Undo, 300 m

FIG. 6.15 Known geographical distribution of Calobota elongate.

6. CALOBOTA HALENBERGENSIS (Merxm. and Schreib.) Boatwr. and B.-E.van Wyk,

comb. nov. (S. Afr. J. Bot.: submitted. 2008). Lebeckia halenbergensis

Merxm. and Schreib., Bull. Jard. Bot. Bruxelles 27: 276. 1957; Merxm. and

Schreib., FSWA 60: 67. 1970.—TYPE: SOUTH AFRICA, Namibia, Luderitz

119 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

South, Halenberg [2615 CB], Dinter 6632 (holotype: M, photo!; isotypes: BM!

2 sheets, K!, NBG!, PRE!, S!, Z, photo!).

Lebeckia decutiens var /3 glabrata E.Mey., Comm. Pl. Afr. Austr. 1: 31. Feb.1836,

synon. nov.—TYPE: "Karroo inter Goedemanskraal et Kaus" [2917 AD], altit.

2000 ped., DrOge s.n. "III, B" (lectotype: P!, designated here). [Note: The

specimen in paris displays the type locality and the characteristic glabrous

pods of Calobota halenbergensis.]

Multi-stemmed shrub up to 1.5 m in height. Branches green; young branches

pilose; older branches pilose or glabrous with light brown bark. Leaves digitately

trifoliolate; petiole 18-40 mm long, longer than leaflets; leaflets narrowly

oblanceolate to linear, alternate, pilose on both surfaces, subsessile, terminal leaflet

5-30 mm long, 0.5-1.5 (-2.0) mm wide, lateral leaflets 4-16 mm long, 0.5-1.0 mm wide, apex acute, base cuneate. Inflorescence 47-155 mm long, with 10 to 20 flowers; pedicel 1-2 mm long; bract 1-3 mm long, linear, pubescent on outer surface; bracteoles 0.5-1.5 mm long, linear, pubescent on outer surface. Flowers 7-

11 mm long, yellow. Calyx (3.5–) 4-6 mm long, pilose to glabrescent on outer surface; tube 2.5-4.0 mm long; lobes 1-2 mm long, subulate. Standard 7.0-11.5 mm long, claw linear, 2-4 mm long, lamina widely ovate to orbicular, 4-7 mm long,

4.5-8.0 mm wide, pilose along dorsal midrib; apex obtuse. Wings 8-11 mm long, claw 2.5-4.0 mm long, lamina oblong, longer than keel, (4.5–) 5.5-7.0 mm long,

(2.2–) 2.5-3.5 mm wide, glabrous, with 6-9 rows of sculpturing. Keel 7.0-9.5 mm long, claw 2.5-4.0 mm long, lamina boat-shaped, 4.0-5.5 mm long, 2.5-4.0 mm wide, glabrous or sometimes pilose on terminal parts. Pistil subsessile to shortly stipitate, glabrous; ovary linear, 4.5-7.5 mm long, 0.8-1.1 mm wide with (8–) 13 to

120 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

18 (-22) ovules; style longer than ovary, 2-4 mm long. Pods linear, semi-terete, glabrous, subsessile to shortly stipitate, 16-32 mm long, 2-4 mm wide, ±3 to 7- seeded, dehiscent. Seeds oblong reniform to reniform, sometimes somewhat angular, 2.0-3.2 mm long, 1.3-2.0 mm wide, mature seeds light brown to light orange, often mottled with brown, surface smooth (Fig. 6.16). Flowering time:

Specimens in flower and fruit have been collected from March to October. The main flowering period appears to be late winter to spring (June to October).

)11 g

\41 k1

v. a h1 h2 14 k2

FIG. 6.16 Morphology of Calobota halenbergensis: (a) leaf in abaxial view; (b) outer surface of the calyx (upper lobes to the left); (c) flower in lateral view; (d) standard petal; (e) wing petal; (f) keel petal; (g) pistil; (h1) bract; (h2—h3) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a, c—f, h — k2) Boatwright et al. 146 (JRAU); (b) Middlemost

2129 (NBG); (g) Salter 5549 (PRE). Scale bars: (a, k1, k2) 1 cm; (b—j3) 1 mm.

121 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—Calobota halenbergensis is similar to Calobota sericea, but differs in the pilose leaves, glabrous keel petals and glabrous, fewer seeded pods (and pistil). In C. sericea the leaves are sericeous, the keel petals pilose and the pods (and pistil) sericeous with more seeds.

Distribution and habitat—Calobota halenbergensis occurs from the Luderitz area in south-western Namibia southwards into South Africa as far as Garies south of Springbok (Fig. 6.17). It can be found on well-drained sandy loam, rocky sandstone, gravelly sand or loose red sand often in dry riverbeds or along disturbed roadsides at altitudes of between ca. 50 m and 1000 m above sea level. According to Mucina et al. (2006) it is an important component of Namaqualand Hardeveld

(SKn 4 Namaqualand Heuweltjieveld) and Namaqualand Sandveld (SKs 7

Namaqualand Strandveld, SKs 8 Namaqualand Coastal Duneveld, SKs 11

Namaqualand Arid Grassland, SKs 12 Namaqualand Spinescent Grassland). Label information indicates that it is grazed by livestock.

Note—The conservation status of Calobota halenbergensis in Namibia has been assessed as lower risk (LR-nt) by Golding (2002).

Additional specimens examined

—2615 (Luderitz): Halenberg (—CB), Dinter 3805 (BM, BOL 2 sheets, K, NBG); 23 km from

Rotkop Station on power line track (—DC), Kolberg and Maggs 184 (WIND).

—2616 (Aus): plains around Garub (—CA), Strohbach 577 (WIND).

—2618 (Keetmanshoop): river at Spitzkop (—AD), Wiss 2059 (WIND).

—2715 (Bogenfels): 9.8 km south of Grillental on main north-south road between Luderitz and Alexander Bay (—AB), Powrie LWP1163 (WIND); Klinghardt Mountains, dark rock mountain east of a patch of shifting dunes in the valley ca. 5 km north-east of Sargdeckel, on the road south to Oranjemund from Luderitz in the Diamond Area 1 (—BC), Bean and

Oliver 2445 (BOL, NBG, PRE); Klinghardt Mountains (—BC), Dinter 3987 (BOL), Milner 830

122 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(PRE, WIND); Namitsas, south of Klinghardt basin (—BD), Mannheimer CM2107 (WIND); 20 km east of Kakaoberg-Escarpment (—DB), Jurgens 28153 (PRE); Boegoeberg (—DD), Dinter

6552 (BM).

—2716 (Witputz): Klinghardt Mountains, koppie south-west face (—AC), Mannheimer

CM1455 (WIND); Diamond area 1, Polly Mountain (—AD), Wendt 20/2 (WIND); Udabib

Mountain (—BB), Willer 810 (K, PRE, WIND); sandy flats at the base of Rooiberg sandstone mountain (—CB), Bean and Oliver 2386 (NBG, PRE); Diamond Area 1, Aurusberg (—CB),

Burke 96225 (WIND), Oliver 10112 (WIND); Sperrgebiet, central Aurus Valley (—CB),

Sirgens 28205 (PRE); Aurus at new camp (—CB), Mannheimer CM2418 (WIND); Farm

Spitzkop (—DC), Giess 14639 (WIND), Wiss 2059 (WIND 3 sheets); Zebrafontein (—DD),

Mei-mailer and Giess 3310, 28861 (K, PRE, WIND), Miller and Horn 1615 (WIND); Rosh

Pinah, west of airfield (—DD), Zietsman 1831 (WIND).

—2816 (Oranjemund) 15 km west of Rosh Pinah on road to Obib (—BA), Giess 13820 (K,

WIND); Diamond Area 1, Valley between Obib Mountain Peak and Obib dunes (—BA), Van

Wyk 9034 (PRE, WIND); 40 km from Brandkaros (—BC), Petersen s.n. (NBG); between

Brandkaros and Grootderm (—BC), Venter 8199 (PRE); Numees (—BD), Sirgens 22149

(PRE), McDonald 739 (NBG, PRE); Annisvlakte (—BD), Jurgens 22322 (PRE); Oranjemund

(—CB), Men(miiller and Giess 2266 (BM, K, PRE, WIND); 70 km north of Oranjemund (—CB),

Williamson 4900 (WIND); 38 mls [61.14 km] north of Port Nolloth, on way to Alexander Bay

(—DA), Werger 519 (K); Goariep-Passage (—DB), JOrgens 22485 (PRE); Goariepvlakte (—

DB), Jurgens 23119 (PRE); near Doringbank, between Lekkersing and Kuboes (—DB), Van

Wyk 2836 (JRAU, 4 sheets); Holgat River (—DB), Wisura 2499 (NBG); Sukkel (—DD),

Jurgens 22739 (PRE); between Port Nolloth and Holgat (—DD), Pillans 5191 (K).

—2817 (Vioolsdrif): bed of Kubus River (—AC), Hardy and Venter 4757 (PRE, WIND); north- west slope of Noemiesberg down to Grasvlakte (—AD), Smook 7937 (PRE); in flats between

Blackhills and Wildepaardenrant (—CA), Thompson and Le Roux 288 (K, NBG);

Eksteensfontein (—CA), Venter 8053 (PRE); 5 mls [8.05 km] west of Stinkfontein (—CC),

Leistner 3400 (K, WIND); Lekkersing (—CC), Thorne s.n. (NBG); 7 mls [11.26 km] beyond

123 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Stinkfontein on way to Kuboos, near Skouerfontein (—CC), Werger 431 (K, PRE, WIND);

Blomhoek, 5 km from Eksteensfontein on road to Kubus (—DD), Germishuizen 4723 (PRE).

2916 (Port Nolloth): 16 km north of Port Nolloth on road to Alexander Bay (—BB),

Germishuizen 5315 (BOL, PRE); 38 mls [61.14 km] east by south of Port Nolloth (—BD),

Acocks 14236 (K, PRE 2 sheets); Port Nolloth (—BD), Bolus 427 (BM, BOL, K, NBG, PRE,

UPS); 2 mls [3.22 km] east of Port Nolloth (—BD), Marloth 12661 (NBG, PRE); between Port

Nolloth road and Stinkfontein (—BD), Middelmost 2129 (NBG 2 sheets, S); 25 mls [40.23 km] after Anenous Pass (—BD), Schlechter sub STE 10890 (NBG 2 sheets, K, PRE).

—2917 (Springbok): 37.5 km from Port Nolloth on road to Steinkopf (—AA), Clarke 640 (K,

PRE); Anenous Flats along road to Eksteenfontein (—AB), Goldblatt and Manning 9284

(NBG); on road to Port Nolloth from Steinkopf (—AC), Boatwright et al. 149, 190 (JRAU);

Anenous Pass (—BA), Germishuizen 4838 (PRE), Goldblatt 6002 (K, PRE, S), Van Wyk

3086 (JRAU 5 sheets), Van Wyk 6189 (PRE); 10 mls [16.09 km] north-west of Steinkopf (-

BC), Salter 5549 (BM, BOL, K, PRE); 5 mls [8.05 km] south of Komaggas (—CA), Compton

22794 (NBG); Sandkop gate at Kleinsee (—CA), Le Roux 5183 (NBG); Komaggas (—CA),

Maguire 400 (NBG 4 sheets); Grootmis, at Springbok turnoff on Port Nolloth-Kleinsee road

(—CA), Rosch and Le Roux 560 (PRE); just north-east of Grootmis (—CA), Strid and Strid

37734 (NBG); between Spektakel and Kleinsee (—CB), Grobbelaar 1962 (PRE); Farm

Langhoogte on Port Nolloth turn-off from Springbok-Kleinsee road just north of waterpipe (-

CB), Le Roux 3959 (NBG); Spektakel (—DA), Compton 11508 (NBG, PRE), Esterhuysen s.n. sub PRE 53706 (PRE); bottom of Wildepaardehoek Pass (—DC), Van Jaarsveld 1392

(PRE).

2918 (Gamoep): Karuchabpoort, Lekkersing (—BB), Oliver, Tolken and Venter 145 (PRE).

3017 (Hondeklipbaai): Farm Koingnaas 475 on road from Hondeklipbaai to Koingnaas (-

AB), Le Roux and Lloyd 353 (NBG); ca. 6 km north of Koingnaas on Springbok road (—AB),

Van Wyk 2848 (JRAU, 2 sheets); 3 km south-west of Soebatsfontein on road to Wallekraal

(—BA), Le Roux 2899 (NBG); Farm Doornfontein 464, Portion 1, Steenkamp Kraal (—BA), Le

Roux 4557, 4630 (NBG); on road from Garies to Hondeklipbaai (—BC), Boatwright et al. 146

124 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(JRAU); Wallekraal, between Garies and Hondeklipbaai (—BC), Compton 5485 (BOL, NBG);

Wallekraal (—BC), Pillans 17970 (BOL); 5 km from Wallekraal to Garies (—BC), Stirton 6053

(K); Farm Sandkraal, 40 km west of Garies (—DA), Hilton-Taylor 1371 (NBG, PRE); 12.9 km from Garies-Hondeklipbaai road on road to Soutfontein, Farm Sandkraal (—DA), Steiner

2172 (NBG, PRE); in red sand near Garies (—DB), Esterhuysen s.n. (K, NBG 2 sheets,

PRE).

Precise locality unknown: Arragaal, Richtersveld, Here s.n. sub STE 11887 (NBG); between Port Nolloth and Augrabies Poort, Bolus 6457 (K); Doornpoort, Compton 20633

(NBG 3 sheets); Kalkfontein, near river, Marloth 12273 (NBG 4 sheets, PRE); kloof by

Doornpoort, Richtersveld, Herre s.n. sub STE 11865, 11866 (NBG); Noagas, DrOge s.n. "Ill,

B"(BM, K, P); sandveld, Richtersveld, Here s.n. sub STE 11868 (NBG 2 sheets, PRE); without locality, DrOge s.n. sub PRE 9367 (PRE).

12 14 16 18 20 22 24 26 28 30 32 o gi - 18RilliiMe■■■ tmruramm a 118 10 ■■■11 Iranlor. 1111 MN KIM 1111111111111 al MIIIN i 1111111111111111 Illal Milt 111111111111■i Solivfilllifil ' 11110111111 ii ug ri milimmori e mom mi wrialtilliallIgne 0111111111111 lil WITIM 111-1111,1117•11111 '1111111111710 ■ER NI 1111111 1/11111111111 111111111.7 11 • - MN Militird111111 misioncsAM PI am mummi28 111111111LN 21 kii MI II" t. 1111 LS NM gag_invi monor \ 111 111, *I 1 1 111,1111111111 '•■ • 4 1111111111111111R ,611 MI Ini II; iiiirealllialaLIMISII imIllilut, 10 12 14 16 18 III-20 22 24 26 28 30 32 34 36

FIG. 6.17. Known geographical distribution of Calobota halenbergensis.

125 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

7. CALOBOTA LINEARIFOLIA (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia linearifolia E.Mey., Comm. Pl. Afr. Austr. 1:

33. Feb. 1836; Benth. in Hook. Lond. J. Bot. 3: 358. 1844; Harv. in Harv. and

Sond., Fl. Cap. 2: 86. 1862; Merxm. and Schreiber, FSWA 60: 67. 1970;

Polhill, Fl. Zambesiaca: 246. 2003—TYPE: SOUTH AFRICA, Northern Cape

Province, on the Gariep, near Verleptpraam [2817 AA], 300 ped., Drege s.n.

"III, B" (lectotype: P!, designated here; isolectotypes: K!, P!, S!, Z photo!).

[Note: The Paris specimen bears the type locality and was annotated by

Meyer himself.]

Lebeckia dinteri Harms, Feddes. Repert. 16: 360. 1920; Merxm. and Schreib.,

FSWA 60: 67. 1970, synon. nov.—TYPE: SOUTH AFRICA, Namibia, Garub

[2616 CA], Dinter 1057 (lectotype: NBG!, designated here). [Note: No other

specimens of the collection Dinter 1057 could be traced in either B or Z. Both

Dinter 1056 and Dinter 1058 were found in Z, but it is likely that all specimens

of Dinter 1057, except the NBG specimen chosen as lectotype, have been

destroyed.]

Erect or virgate, multi-stemmed, unarmed shrub up to 3 m in height.

Branches green; young branches sericeous or silky; older branches sericeous or silky with light brown bark. Leaves simple; petiole absent; leaflets linear to oblanceolate or spathulate, sericeous, sessile, leaflets 11-45 mm long, 1.0-2.5 mm wide; apex acute; base cuneate. Inflorescence 38-170 mm long, with 5 to 9 flowers; pedicel 1-2 (-8) mm long; bract 1.5-2.5 mm long, linear to narrowly ovate, pubescent; bracteoles 1.0-1.5 mm long, linear to narrowly ovate, pubescent.

Flowers 13-17 mm long, bright yellow. Calyx 5.5-7.5 mm long, sericeous; tube 4-6

126 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

mm long; lobes 1.0-2.5 mm long, deltoid. Standard 11.5-15.0 mm long; claw linear,

2.5-3.5 mm long; lamina ovate, 8.5-12.0 mm long, 8.5-10.5 mm wide, dorsal

surface pubescent; apex acute. Wings 8.0-11.5 mm long, claw 3.0-4.5 mm long,

lamina oblong, longer than keel, 5-7 mm long, 2.5-4.0 mm wide, pilose, with 4-8

rows of sculpturing. Keel 12-15 mm long, claw 3.5-6.0 mm long, lamina boat-

shaped, 7.5-10.0 mm long, 4.0-5.5 mm wide, pubescent. Pistil subsessile to shortly

stipitate, pubescent; ovary linear, 9-11 mm long, 0.8-1.5 mm wide with 12 to 18

ovules; style longer than ovary, 4.8-7.8 mm long. Pods linear, sometimes somewhat

clavate, semi-terete, sericeous, subsessile to shortly stipitate, 25-40 mm long, 2-3

mm wide, ±4 to 8-seeded, dehiscent. Seeds oblong-reniform, 2.5-3.5 mm long, 1.5-

2.5 mm wide, mature seeds light brown or grey, surface smooth (Fig. 6.18).

Flowering time: Mainly between September and November, but flowering and

fruiting specimens have been collected in January, April, May and August.

Diagnostic characters—Calobota linearifolia is similar to C. cinerea but

differs in its virgate to erect habit, simple, linear leaves, fewer flowered

inflorescences, sericeous pods that are constricted between the seeds and the light

brown to grey seeds. Calobota cinerea has a low, spreading habit, obovate, elliptic

or oblanceolate and trifoliolate leaves and the inflorescences more densely flowered. The latter species also has tomentose pods that are not constricted

between the seeds and the seeds are light brown to orange and often mottled.

Distribution and habitat—Calobota linearifolia is widely distributed in the

Northern Cape Province of South Africa and southern Namibia, with outlier populations around the Unjab River, Sima Hill and the Sanitatas area in north- western Namibia (Fig. 6.19). Polhill (2003a) mentioned that this species also occurs in Botswana in the Kgalagadi district, but no specimens from this region were

127 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA available for study. Calobota linearifolia occurs in dune sand, red sand or brackish soil often along rivers or in dry watercourses, washes and gulleys. It occurs at altitudes of between ca. 300 m and 1350 m. It is suspected of poisoning livestock

(according to label information).

e ■ b ,•

421Wa=ommmtzal°5E22°' d hl h2 k2

FIG. 6.18 Morphology of Calobota linearifolia: (a1—a2) leaves in abaxial view; (b) flower in lateral view; (c) standard petal; (d) outer surface of the calyx (upper lobes to the left); (e) wing petal; (f) keel petal; (g) pistil; (hl ) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) intermediate carinal anther; 03) short, dorsifixed anther; (kl) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al, b) Acocks 13195 (PRE); (a2) Giess and Van Vuuren 821

(BOL); (c) Giess 13803 (WIND); (d—g, j1 j3) Dean 673 (JRAU); (hl—i) Evrard 9285 (PRE);

(k1—k2) Keet 1668 (WIND). Scale bars: (al —a2, k1—k2) 1 cm; (b—j3) 1 mm.

128 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Additional specimens examined

—1812 (Sanitatas): 20 mls [32.18 km] south of Orupembe (—CD), Giess and Leippert 7435

(WIND).

—1913 (Sesfontein): 2 km north of Sima Hill, Koakoland (—AA), Braine 68 (WIND); near Sima

Hill, north-east of Mowe Bay (—AA), Moss and Jacobsen K316 (PRE, WIND); between

Hoamib and Unjab Rivers (—AC), Tinley 1625 (WIND).

—2013 (Unjab Mouth): 8 mls [12.87 km] east of Torrabay (—AA), Giess et al. 6180 (PRE,

WIND); Unjab River (—AB), Kers 1875 (S).

—2517 (Gibeon): Orion, Varsfontein (—AC), Giess et al. 7190 (K); between Achterfontein and

Gelwater (—BC), Pearson 9233 (K).

—2520 (Mata Mata): Kalahari Gemsbok National Park (—CA), Schlieben 10433 (PRE).

—2615 (Luderitz): Farm Gawachab KEE 114 (—BB), De Winter and Giess 6424 (BOL, K,

WIND).

2616 (Aus): between Neisip and Eureka (—BC), Merxmiiller and Giess 2986 (WIND); 13 mls [20.92 km] west of Aus (—CA), Giess and Van Vuuren 821 (BOL, K, WIND); Kuibis (-

DA), Bayliss BRI.B.365 (K, PRE 2 sheets).

—2617 (Bethanie): Felshang, near Seeheim (—DD), Men(miiller and Giess 3615 (PRE,

WIND); Seeheim (—DD), Dinter 1205 (NBG).

2618 (Keetmanshoop): 7 mls [11.26 km] south by east of Naute Station (—DB), Acocks

15659 (K, PRE); Sandmodder 73 (—DD), Lensing A87/75 (WIND).

—2619 (Aroab): east of Groot Karas Mountains (—CA), Mannheimer et al. CM771 (WIND).

2620 (Twee Rivieren): Ashkam (—DD), Evrard 9285 (PRE).

2715 (Bogenfels): Sperrgebiet South, 38 km east of Kakaoberg escarpment (—DB),

Jurgens 28176 (PRE); Chameis Valley (—DC), Williamson 2647 (BOL).

—2717 (Chamaites): Mara, Letterklip area in Konkiep River bed (—CD), Craven 2634

(WIND); bed of Vis River, near Seeheim (—DA), Keet 1663 (WIND, 2 sheets).

129 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

2719 (Tranental): Warmfontein 280 (—AB), Lensing J14/76 (WIND), Muller 87 (WIND);

Farm Blaufontein WAR 276 (—AC), Giess et al. 7131 (WIND).

2720 (Noenieput): 20 mls [32.18 km] north of Noenieput on way to Kalahari Gemsbok Park

(—AD), Werger 1489 (NBG, PRE); Askam area, 120 km from Van Zylsrus (—BB), Dlamini et al. EvW568 (K).

2723 (Kuruman): 10 mls [16.09 km] south-west of confluence of the Kuruman and Molopo

Rivers (—AA), Leistner 2255 (K).

—2817 (Vioolsdrif): 5 km west of Nuob River (—AA), Giess 13803 (K, PRE, WIND); Garip (-

AA), Wyley s.n. (S); Richtersveld (—BA), Venter 8069 (PRE).

—2818 (Warmbad): a little south of Dabegabis (—BC); Pearson 4390 (BM 2 sheets, K); near

Dabegabis (—BC), Pearson 4896 (K); mountains between Wolftoon and Henkriesfontein (-

CC), Pearson 3105 (BM 2 sheets, BOL, K 2 sheets); Farm Aluriesfontein 308 (—DA), Owen-

Smith 1252 (WIND).

2819 (Ariamsvlei): Nababis, Guigatsis (—BA), Evrard 9220 (PRE).

—2820 (Kakamas): below Riemvastmaak, Upington (—AD), Barnard s.n. (NBG)

—2821 (Upington): Swartpan, north-west of Upington (—AA), Snijman 214 (NBG); 24 mls

[38.62 km] south-east of Groblershoop (—DD), Leistner and Joynt 2861 (K, PRE).

—2822 (Glen Lyon): 9 mls [14.48 km] west south-west of Volop (—CD), Leistner 1742 (K).

2824 (Kimberly): Gordonia (—BA), Webb 16 (K, PRE).

—2918 (Gamoep): Karasberg (—CB), Van Schauroff s.n. (K).

2921 (Kenhardt): 40 mls [64.36 km] north north-east of Kenhardt (—AC), Acocks 18803 (K).

3020 (Brandvlei): sand dunes along Sak River near Brandvlei (—AD), Acocks 13195 (BOL,

K, PRE).

—3022 (Carnarvon): Van Wyksvlei, Beesdam (—AD), Dean 673 (JRAU 2 sheets).

Precise locality unknown: without locality, Bolus 1935 (BOL), Flanagan s.n. sub PRE

53667 (PRE), Range 1935 (BOL, NBG).

130 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

12 14 16 18 20 22 24em 26 28 30 32 34 aamallosalli. rrilift511111ra no 18 ■Normon ■ 1 alwry Immoum 1 111011111111 I 111_11111111 111111111111 11 2 AISIMME 11111111111111 ga MOM 111111111111111 III MEM 1111111■211 ' P 2 MEM IlrillINIMMIN ii munErwmasWM= MN X m an FAIMIZIM INEMIIIM , isIIII siawm riticlin 10 ME MEM In SIM Ili■ V g MI " MN MI • • MP 1011102 nu 4010mmommom "64 lim'II nigm 11!I". wr ommERSOM MO NM 111111111E1 c r!Rnic' C ' IsmaiiiiiiiM Faciamili im.34 36 10 12 14 16 18 20 22 24 26 28 30 32

FIG. 6.19 Known geographical distribution of Calobota linearifolia.

8. CALOBOTA LOTONONOIDES (Schltr.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr.

J. Bot.: submitted. 2008). Lebeckia lotononoides Schltr. in Engler Bot. Jahr.

27: 143. 1900, as "L. lotonoides"; Germishuizen and Meyer, Plants of

Southern Africa: an annotated checklist, Strelitzia 14: 523. 2003, pro parte,

Boatwr. and Van Wyk, in S. Afr. J. Bot. 73 (4): 664-666. 2007, as "L.

lotononoides", non Goldblatt and Manning. 2000.—TYPE: SOUTH AFRICA,

`In regione namaquensi: In sabulosis montium Karree-Bergen, alt. C. 4500

ped.' [3118 AB], Schlechter 8214 (lectotype: K!, designated here;

isolectotypes: BM!, BOL!, PRE!, S!, Z, photo!). [Note: A large number of

131 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Schlechter's African collections are housed at Kew (Stafleu and Cowan 1985)

and we therefore select the K specimen as lectotype.]

Small, decumbent, multi-stemmed, unarmed shrublet up to 0.5 m in height.

Branches green; young branches sericeous, older branches pilose to glabrous with light brown bark, often buried. Leaves digitately trifoliolate or rarely 5-digitate with two additional leaflets; petiole 10-23 mm long, longer than leaflets; leaflets elliptic to narrowly oblanceolate, secund, alternate, sericeous, conduplicate, subsessile, terminal leaflet 10-17 mm long, 1.5-3.0 mm wide, lateral leaflets 8-14 mm long,

1.5-3.0 mm wide, mucronate, base narrowly cuneate. Inflorescence (15—) 24-54 mm long, with (3—) 4 to 14 flowers; pedicel 1-3 mm long; bract 0.7-1.5 mm long, linear, pubescent; bracteoles 0.3-0.6 mm long, linear, pubescent. Flowers 12-15 mm long, yellow. Calyx 5.0-6.5 mm long, pubescent; tube 3.0-3.5 mm long; lobes

2-3 mm long, subulate. Standard 13-16 mm long; claw linear, 4-7 mm long; lamina widely ovate to transversely oblong, 8-9 mm long, 9.0-12.5 mm wide, pilose along dorsal midrib; emarginate. Wings 14-15 mm long; claw 3.0-4.5 mm long; lamina narrowly oblong, subfalcate, longer than keel, 11.5-15.0 mm long, 2-4 mm wide, glabrous, with 7-8 rows of sculpturing. Keel 9-11 mm long; claw 3-4 mm long; lamina boat-shaped, 5.5-7.0 mm long, 3.0-4.5 mm wide, glabrous. Pistil subsessile to shortly stipitate, pubescent, ovary linear, 6.5-7.5 mm long, 0.7-0.9 mm wide with

15 to 19 ovules; style longer than ovary, 3.0-3.5 mm long. Pods narrowly oblong, sometimes somewhat clavate, sericeous, semi-terete, subsessile to shortly stipitate,

18-37 (-40) mm long, 3-4 mm wide, 7 to 12-seeded, dehiscent. Seeds suborbicular, 2.0-2.5 mm long, 1.5-2.0 mm wide, mature seeds brown with beige

132 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA spots, hilum brown, surface rugose (Fig. 6.20). Flowering time: late winter to late spring.

j1 j2 j3

g1 g2 k1

FIG. 6.20 Morphology of Calobota lotononoides: (al—a2) leaves in abaxial view; (b) flower in lateral view; (c) standard petal; (d) wing petal; (e) keel petal; (f) pistil; (g1) bract; (g2) bracteoles; (h) outer surface of the calyx (upper lobes to the left); (i) androecium; (j1) long, basifixed anther; 0,2) intermediate carinal anther; 03) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al—k2) Boatwright et al. 142 (JRAU). Scale bars: (al—a2, k1— k2) 1 cm; (b—j3) 1 mm.

133 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—Calobota lotononoides superficially resembles C. sericea, notably the broad-leaved form on the West Coast and around Clanwilliam.

In this form the leaflets are broad and sericeous much like those of C. lotononoides.

However, C. lotononoides differs in its smaller, decumbent habit with buried stems and especially in the exceptionally long lamina of the wing petals, which is ± twice as long as those of the glabrous keel petals. The inflorescences are unarmed, and the sericeous pods contain spotted, rugose seeds. Calobota sericea can reach heights of up to 2 m and the wing petals are only slightly longer than the pilose keel petals.

Distribution and habitat—Calobota lotononoides occurs on the west coast of South Africa around Hondeklipbaai and Brandsebaai (Fig. 6.21). The type collection was made by Schlechter at "Karree-Bergen", close to Nuwerus in the

Vanrhynsdorp district according to Leistner and Morris (1976). The species has been recorded on well-drained, sandy soils with the main stems becoming covered with sand. The multi-stemmed, decumbent habit appears to be an adaptation to moving sand dunes, as extensive underground branches enable the plant to emerge above the sand after being covered.

Note—According to Raimondo et al. (in press) the status of this species is declining. Although only a 5% habitat loss has been observed due to heavy mineral sand mining, the loss is still ongoing and could later become problematic.

Additional specimens examined

—3017 (Hondeklipbaai): from Sandkraal, north of Bitter River, 25 km west of Garies (—CB),

Helme et al. 4720 (NBG); Farm Roodeheuwel, 9 km west of Nariep (—DC), Boatwright et al.

142 (JRAU), PeroId 1646 (PRE).

134 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

—3117 (Lepelfontein): Vredendal, south-east of Brandsebaai, on Farm Hartebeestekom (-

BD), Helme 2931 (NBG); Van RhynsdorpNredendal, Brandsebaai (—BD), Van Rooyen 2235

(PRE).

—3118 (Vanrhynsdorp): Karree-Bergen (—AB), Schlechter s.n. sub TRV 1059 (PRE); from Skaapvlei, some 10 km north-east of Olifants River mouth (—AC), He/me 4684 (NBG).

FIG. 6.21. Known geographical distribution of Calobota lotononoides.

9. Calobota namibiensis Boatwr. and B.-E.van Wyk, sp. nov., similar to Calobota

linearifolia (E.Mey.) Boatwr. and B.-E.van Wyk, but differs in the short, elliptic

to ovate leaves and short, tomentose pods. It is also similar to Calobota

saharae (Coss. and Dur.) Boatwr. and B.-E.van Wyk but differs in the elliptic

to ovate leaves, densely pubescent petals and the short, tomentose pods (to

be translated into Latin).—TYPE: 25 mls [40.23 km] east of Luderitz on road

135 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

to Aus [2615 CA], De Winter and Hardy 7919 (holotype: WIND!, specimen on

the right, here designated; isotypes: K!, PRE!).

Lebeckia dinteri auct. non Harms: Merxm. and Schreib., FSWA 60: 67. 1970. [Note:

Harms described Lebeckia dinteri from a specimen that is actually Calobota linearifolia. Subsequent authors, however, referred to the species here described as new as "Lebeckia dinteri' which is distinct from Calobota linearifolia, but this taxon has not been formally described.]

Virgate, multi-stemmed shrubs up to 1.2 m tall. Branches green; young branches sericeous; older branches sericeous or glabrous with light brown bark.

Leaves simple; petiole absent; elliptic to ovate, upper and lower surfaces sericeous, alternate, sessile, 6-12 mm long, 1-2 mm wide; apex acute; base cuneate.

Inflorescences 70-110 mm long, with 3 to 8 flowers; pedicel 1-2 mm long; bract

2.5-3.5 mm long, linear, sericeous; bracteoles 1.5-3.0 mm long, linear, sericeous.

Flowers 10-11 mm long, bright yellow. Calyx 6-8 mm long, densely pubescent on the outer surface; tube 3-4 mm long; lobes 3.0-4.5 mm long, subulate. Standard

9.5-12.0 mm long; claw linear, 2-3 mm long; lamina ovate, 7.5-9.0 mm long, 5-6 mm wide, densely pubescent on dorsal surface; apex acute. Wings 8.0-8.5 mm long; claw 2-3 mm long; lamina oblong, shorter than the keel, 5.5-6.5 mm long,

2.5-3.0 mm wide, with 6-8 rows sculpturing, glabrous. Keel 8.0-11.5 mm long; claw

2.5-3.5 mm long; lamina boat-shaped, 5.5-8.0 mm long, 3.5-4.5 mm wide, pubescent. Pistil shortly stipitate, pubescent; ovary linear, 4-5 mm long, 0.9-1.0 mm wide with about 5 ovules; style longer than the ovary, 5.0-9.5 mm long. Pods linear, semi-terete, 15-17 mm long, 2 mm wide, 3 to 4-seeded, tomentose, ?dehiscent.

136

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Seeds not seen (Fig. 6.22). Flowering time: Flowering specimens have been

collected in March, August and October.

b g j1 j2

k1 d

h1 h2

FIG. 6.22 Morphology of Calobota namibiensis: (a) flower in lateral view; (b) standard petal;

(f) keel petal; (g) pistil; (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther;

(j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2)

pod in dorsal view.

Voucher specimens: (a—c, e—f, h1—h2, j1—j2) Owen-Smith 1268 (WIND); (d) Kolberg

and Maggs 212 (WIND); (g, i) Owen-Smith 117 (WIND); (k1—k2) De Winter and Hardy 7919

(WIND). Scale bars: (d, k1—k2) 1 cm; (a—c, e—j3) 1 mm.

Diagnostic characters—A very poorly known and collected species. The lack of sufficient material makes it difficult to study the variation within this species. It 137 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA is similar to Calobota linearifolia and C. saharae, but differs in the elliptic to ovate leaves, densely pubescent petals and the short, tomentose pods. In both C. linearifolia and C. saharae the leaves are linear. The petals are also pubescent in C. linearifolia while in C. saharae only the standard petal is pilose. The pods of C. linearifolia are sericeous, while those of C. saharae are glabrous and in both species they are longer than those of C. namibiensis.

Distribution and habitat—Calobota namibiensis is endemic to Namibia and occurs in the Luderitz and Aus areas (Fig. 6.23). It grows on dunes in red sand or gravelly sandy soil along roadside up to 1800m above sea-level.

Note—The status of this species has been assessed as vulnerable (VU D2) by Golding (2002; included as Lebeckia dinteri) and as least concern (LC) recently by Loots (2005; included as Lebeckia dinteri).

Additional specimens examined

—2615 (Luderitz): dunes south of Koichab pump station (—AD), Seely 2005 (WIND); Koichab

State Land (—BC), Owen-Smith 117, 1268 (WIND); Halenberg (—CB), Merxm011er and Giess

3088 (WIND); Rotkop, Luderitz (—CB), Dinter 1032 (NBG), Merxm011er and Giess 3141

(WIND).

—2616 (Aus): 13 mls [20.92 km] west of Aus (—CB), Giess and Van Vuuren 821 (PRE,

WIND); koppie south-west of Rotkop Station, next to power-line track (—CB), Kolberg and

Maggs 212 (PRE, WIND).

138

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

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FIG. 6.23 Known geographical distribution of Calobota namibiensis.

10. CALOBOTA OBOVATA (Schinz) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia obovata Schinz, Mem. Herb. Boiss. 1: 126.

1900; Merxm. and Schreiber, FSWA 60: 68. 1970.—TYPE: SOUTH AFRICA,

Namibia, Gansberg [grid unknown], Fleck 75 (lectotype: Z-000022914,

photo!, specimen on the left, designated here; isolectotype: Z-000022915,

photo!). [Note: According to Stafleu and Cowan (1985) Hans Schinz's main

collection is in Z and we therefore designate one of the Z specimens as

lectotype.]

139 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Sericeous, multi-stemmed shrub up to 1 m in height. Branches green; young branches short-sericeous, older branches short-sericeous or glabrous with light brown bark. Leaves unifoliolate or rarely trifoliolate; petiole shorter than leaflets, 5-

24 mm long; leaflets widely obovate to orbicular, rarely ovate, alternate, short- sericeous, subsessile, 12-36 mm long, 10-25 mm wide, apex obtuse or rarely acute, base cuneate. Inflorescence 24-85 mm long, with 7 to 16 flowers; pedicel 4-

5 mm long; bract 3-5 mm long, ovate, pubescent; bracteoles 3-5 mm long, ovate to linear, pubescent. Flowers 13-16 mm long, bright yellow. Calyx 8.0-8.5 mm long, densely pubescent; tube 4.5-5.5 mm long; lobes 3-4 mm long, subulate. Standard

14.5-15.5 mm long, claw linear, 3.5-4.5 mm long, lamina widely ovate, 10.0-11.5 mm long, 12.5-14.5 mm wide, dorsal surface pubescent; apex obtuse. Wings 14.5–

15.5 mm long, claw 3.0-3.5 mm long, lamina oblong, shorter than keel, 11-12 mm long, 5.5-6.0 mm wide, pilose, with 4-5 rows of sculpturing. Keel 14.5-16.5 mm long, claw 4-5 mm long, lamina boat-shaped, 10.0-11.5 mm long, 5.5-6.5 mm wide, pubescent. Pistil subsessile to shortly stipitate, pubescent, ovary linear, 11.5-

12.5 mm long, 1.5-2.0 mm wide with ±15 ovules; style shorter than ovary, 6-7 mm long. Pods linear, semi-terete, subsessile to shortly stipitate, 30-40 mm long, 4-5 mm wide, 5 to 7-seeded, short-sericeous, dehiscent. Seeds not seen (Fig. 6.24).

Flowering time: Flowering specimens have been collected in May, June, July,

November and December suggesting that flowering occurs in spring and summer.

Diagnostic characters—Similar to Calobota elongata but differs in the unifoliolate leaves, racemose, few-flowered racemose inflorescences, pedicellate flowers, obovate bracts and bracteoles, widely ovate standard petal, pubescent wing petals that are shorter than the keel petals and the linear, sericeous pods (C. elongata differs in the trifoliolate leaves, dense, spicate inflorescences with smaller,

140 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA sessile flowers, long, narrow bracts and bracteoles, ovate to elliptic standard petal with a pilose midrib, wing petals that are longer than keel and shorter, tomentose pods with less seeds).

d b

g2 k2 ••65===seurzsm.„..

FIG. 6.24 Morphology of Calobota obovata: (al—a2) leaves in abaxial view; (b) flower in lateral view; (c) standard petal; (d) wing petal; (e) keel petal; (f) outer surface of the calyx

(upper lobes to the left); (g1) bract; (g2) bracteoles; (h) androecium; (i) pistil; (jl) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (kl) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al) Schweidfezer 4315 (WIND); (a2, kl—k2) Dinter 1094

(NBG); (b, f, i) Pearson 8036 (BOL); (c—e, h, jl—j3) Kers 152 (WIND). Scale bars: (al—a2, kl—k2) 1 cm; (b—j3) 1 mm.

141 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Distribution and habitat—Calobota obovata is endemic to Namibia and occurs from Windhoek southwards as far as Aus (Fig. 6.25). It grows on rocky outcrops or granite at altitudes of ca. 1600 m to 2230 m.

Note—The status of this species has been assessed as data deficient (DD) by Golding (2002) and Loots (2005).

Additional specimens examined

—2217 (Windhoek): Farm Regenstein (—CA), Giess and Giess 11765 (NBG, PRE, S, WIND),

Menanaller and Giess 28010 (WIND).

2316 (Nauchas): Gamsberg (—AD), Kers 152 (WIND), Kers 154 (S), Merv-nuIler and Giess

957 (BM, K, PRE, WIND), Kolberg and Tholkes NAM2960-HK1418 (K), Meyer 1016

(WIND), Schweidfezer 4315 (WIND); Farm Hopefield (—AD), Merxrni.iller and Giess 893

(WIND); Farm Weipenfels (—AD), Walter and Walter 1686 (WIND).

2318 (Leonardville): Farm Rietfontein (—CA), Leippert 4537 (WIND).

2516 (Helmeringhausen): Farm Aruab 23 on top edge of Great Escarpment (—DA), Miller

Mil1/036 (WIND).

—2615 (Luderitz): Luderitz (—CA), Rogers 2959 (K).

—2616 (Aus): Urus Plateau, Luderitz (—BA), Logan and Seely 301 (WIND); Aus (—CB), Dinter

1094 (NBG); Aus, rocks east of railway station (—CB), Pearson 8036 (BM, BOL 2 sheets, K,

NBG, PRE).

Precise locality unknown: Ruschberg, Rusch 4379 (S).

142 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

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FIG. 6.25. Known geographical distribution of Calobota obovata.

11. CALOBOTA PSILOLOBA (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Stiza psiloloba E.Mey. Comm. Pl. Afr. Austr. 1: 31.

Feb. 1836; Meisner in Hook. Lond J. Bot. 2: 68. 1843. Lebeckia psiloloba

(E.Mey.) Walp., Linnaea 13: 478. 1839; Benth. in Hook. Lond. J. Bot. 3: 356.

1844; Harv. in Harv. and Sond., Fl. Cap. 2: 84. 1862; Bond and Goldblatt, J.

S. Afr. Bot. 13: 290. 1984; Goldblatt and Manning, Cape Pl.: 493. 2000.—

TYPE: SOUTH AFRICA, without locality, DrOge 6470 (lectotype: P!,

designated here; isolectotype: K!, S!). [Note: The Paris specimen bears a

label with the words "Stiza psiloloba mihi' in Meyer's handwriting, and is

therefore chosen as lectotype. The relatively thin, glabrous twigs and small,

143 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

leathery pods (in an envelope on the specimen) are characteristic of this

species.]

Acanthobotrya pungens sensu Eckl. and Zeyh., Enum. 2: 193. Jan 1836, non

Lebeckia pungens Thunb. Type: South Africa, "Zwartkops— et Zondagsrivier",

Ecklon and Zeyher 1340 (lectotype: NBG!, designated here; isolectotypes: P!,

K!, S!). [Note: The specimen in NBG dislays the type locality and the

Enumeratio label.]

Erect, multi-stemmed, spinescent shrub up to 1.2 m tall. Branches green; young branches glabrous to very sparsely pubescent, older branches glabrous with brown bark. Leaves unifoliolate or rarely trifoliolate; petiole short, 0.5-1.0 mm long

(up to 5 mm long when trifoliate); leaflets slightly obovate to spathulate, alternate, upper and lower surfaces thinly pilose, 5-7 (-11) mm long, 1.5-3.0 (-4.0) mm wide; base attenuate; apex obtuse. Inflorescences 40-50 (-65) mm long, with 8 to 10 (-

12) flowers; pedicel relatively long, (3—) 4-5 (-6) mm long; bract 0.3-0.4 (-0.6) mm long, triangular; bracteoles 0.1-0.3 mm long, linear to slightly ovate. Flowers 8-11 mm long; bright yellow. Calyx 2.2-2.6 mm long, slightly pubescent on the outer surface; tube 1.5-2.0 mm long; lobes widely triangular, 0.5-0.8 mm long; apices broadly acute. Standard 7-9 mm long; claw 2.5-3.0 (-4.0) mm long; lamina oblong to slightly ovate, 4-5 (-6) mm long, 3.0-4.5 mm wide, glabrous; apex emarginate.

Wings 7.0-8.5 mm long; claw 2.5-3.0 (-3.5) mm long; lamina oblong, shorter than the keel, 4.0-4.5 (-5.0) mm long, 2-3 mm wide, with 4-5 rows sculpturing, glabrous.

Keel 8.5-11.0 mm long; claw 3.5-4.0 (-4.5) mm long; lamina boat-shaped, 5-6 (-7) mm long, 3-4 mm wide, glabrous. Pistil stipitate, glabrous; ovary linear to elliptic,

5.5-7.5 mm long, 0.8-1.1 mm wide; ovules 6 to 8; style shorter than ovary, 3.5-5.0

144 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA mm long. Pods oblong, laterally compressed, membraneous, 14-17 (-33) mm long,

4-5 mm wide, glabrous; stipe up to 4 mm long, 2 to 6-seeded, indehiscent. Seeds reniform, 2.5-3.0 mm long, 2.0-2.5 mm wide, light brown, surface smooth (Fig.

6.26). Flowering time: The main flowering time of C. psiloloba is between March and

June, but flowering also sometimes occurs in spring and summer (October to

January).

Diagnostic characters—Calobota psiloloba is similar to Calobota cuspidosa, but differs in that this species only reaches heights of up to 1.2 m and has small flowers that are borne on long, thin pedicels (C. cuspidosa reaches heights of up to

4 m and has much larger flowers on relatively short pedicels). The older stems are almost completely glabrous and the standard petal notched, while in C. cuspidosa the older stems are pubescent and the standard petal is obtuse. The pods and seeds are also smaller than those of C. cuspidosa.

Distribution and habitat—Calobota psiloloba is restricted to the Eastern

Cape Province and occurs mainly around Port Elizabeth, Grahamstown and Addo, but has outlier populations around Cradock at altitudes between 30 and 520 m (Fig.

6.27). It is normally found in open grasslands on sandy soils and is eaten by elephants (according to label information).

145

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

d a

JiT j2 j3

f 'AitfiNT* k1

h1 h2 g1 g2

FIG. 6.26 Morphology of Calobota psiloloba: (a) flower in lateral view; (b) standard petal; (c) outer surface of the calyx (upper lobes to the left); (d) wing petal; (e) keel petal; (f) pistil;

(g1—g2) leaves in abaxial view; (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a) Story 3353 (PRE); (b—f, g1, h1—j3) Van Wyk s.n. (JRAU);

(g2) Zeyher 2317 (NBG); (k1—k2) Le Roux et al. 20 (JRAU). Scale bars: (g1—g2, k1—k2) 1 cm; (a—f, h1—j3) 1 mm.

146 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Additional specimens examined

—3125 (Steynsburg): Grootfontein, Middelburg district (—AC), Burtt-Davy 9665 (PRE).

—3225 (Somerset East): Cradock (—BA), Botany Honours Students 320 (PRE, PRU);

Somerset East, farm Kaboega (—DA), Bayliss 6897 (K, S).

—3324 (Steytlerville): on plains near Zondags River (—DB), Bowie s.n. (BM).

—3325 (Port Elizabeth): in the valleys of the Zuurberg Mountains at Paarde Poort (—AC),

Bolus 2667 (BOL, K), Wilson 422 (BM); Perdepoort (—AC), Theron 685 (PRE); Bassons

Kloof, Annes Villa (—BB), Bayliss 3923 (NBG); Addo Elephant Park (—BD), Archibald 3848

(GRA, K, PRE), Brynard 445 (K, PRE), Repton 5715 (PRE); 15th mile [24.1 km] on the road from Port Elizabeth to Addo (—BD), Long 612 (GRA, K); along road from Grahamstown to

Paterson (—BD), Van Wyk s.n. (JRAU); Uitenhage (—CD), Alexander-Prior s.n. (BM, K 3 sheets); Cradock Place, Port Elizabeth (—DC), Galpin 6430 (GRA, PRE); district Uitenhage,

Zwartkops River (—DC), Ecklon and Zeyher 737 (BM, BOL 2 sheets, K 2 sheets, NBG, PRE

4 sheets), Ecklon and Zeyher 1340 (K, NBG, P 3 sheets, PRE, S); New Brighton near Port

Elizabeth (—DC), Ethel West 395 (BOL); Coega (—DC), Long 945 (GRA, K); Redhouse, near cemetary (—DC), Troughton 425 (PRE 2 sheets); Redhouse (—DC), Boatwright et al. 107

(JRAU), Le Roux et al. 20 (JRAU), Wells 3697 (PRE); Zwartkops River (—DC), Zeyher 2317

(PRE 2 sheets, NBG, S).

—3326 (Grahamstown): Table farm near Grahamstown (—BC), Atherstone 24, 50 (K);

Grahamstown (—BC), MacOwen 1102 (K); Bushman's River bridge, main road from

Grahamstown to Port Elizabeth (—CA), Archibald 3958 (PRE), Story 3353 (K, PRE).

Precise locality unknown: without locality, Barber s.n. (K), Bowie s.n. (K), Zeyher 3317

(PRE).

147 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

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'8111 • le tenrill lams via ono ionssI IIMI " a 11111 all EMI 11111111 II 111111 ' - ■ IN ' 111111111111111111 MB- - 11111111143„11/611111111111 All oriemen ari ..ivirt ':....--. Ism c,...... Imatmoduno .... mii,,,, 10 12 14 16 18 20 22 24 26 28 30 32 34 36

FIG. 6.27 Known geographical distribution of Calobota psiloloba.

12. CALOBOTA PUNGENS (Thunb.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia pungens Thunb., Nov. Gen.: 141. 1800;

Prodr. Pl. Cap.: 122. 1800; Willd., Sp. Pl.: 947. 1800; Thunb., Fl. Cap.: 562.

1823; DC., Prodr. 2: 137. 1825; Benth. in Hook. Lond. J. Bot. 3: 356. 1844;

Harv. in Harv. and Sond., Fl. Cap. 2: 83. 1862; Compton, Trans. Roy. Soc. of

S. A. 19: 294. 1931; Bond and Goldblatt, J. S. Afr. Bot. 13: 290. 1984;

Goldblatt and Manning, Cape Pl.: 494. 2000.—TYPE: SOUTH AFRICA,

Eastern Cape Province, near Olifant's River [close to Oudtshoorn] at

Cannaland [now Little Karoo], Thunberg s.n. sub THUNB-UPS 16417

(lectotype: UPS!, designated here). [Note: This is the only specimen in

148 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Thunberg's herbarium and the type locality is written on the back of the

sheets in Thunberg's handwriting. This specimen is therefore chosen as

lectotype.]

Stiza erioloba E.Mey. Comm. Pl. Afr. Austr. 1: 31. Feb. 1836.—TYPE: SOUTH

AFRICA, Eastern Cape Province, near Klaarstroom [close to Prince Albert,

3322BC], near the foot of the "Groote Zwartebergen" [Great Swartberg],

Drege s.n. (lectotype: P!, specimen on the right, designated here;

isolectotypes: K!, P!, S!). [Note: The Drege specimen in P is chosen as

lectotype because it displays the type locality as well as the characteristic

tomentose pods from which Meyer clearly derived the name for this species].

Erect, multi-stemmed woody spinescent shrub up to 2.5 m tall. Branches green; young branches densely sericeous; older branches sericeous with brown bark. Leaves unifoliolate or rarely trifolioilate; petiole short, 0.5-2.0 mm long (up to 5 mm long when leaves are trifoliate); leaflets obovate to broadly obovate, upper and lower surfaces sericeous, alternate, 3-13 (-18) mm long, 2-5 (-8) mm wide; base attenuate; apex retuse. Inflorescences (22–) 25-40 (-55) mm long, with (4–) 6-10 flowers; pedicel 2-4 mm long; bract 0.6-1.1 mm long, triangular; bracteoles 0.4-0.6 mm long, narrowly ovate. Flowers 10-13 (-14) mm long, bright yellow. Calyx 2.5-

3.7 mm long, pubescent on the outer surface; lobes triangular, 0.4-1.0 mm long; tube 2-3 mm long, apices acute. Standard 9.5-12.0 mm long; claw 3-4 mm long; lamina widely ovate, 6-7 (-8) mm long, 4-6 (-9) mm wide, pilose along the dorsal midrib; apex broadly acute. Wings 7-12 mm long; claw (2.0–) 3.5-5.0 mm long; lamina oblong, shorter than the keel, 4-6 (-7) mm long, 2-3 (-4) mm wide with 4-5 rows of sculpturing, pilose. Keel 10-14 mm long; claw 4.0-5.5 mm long; lamina

149 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA oblong, 6.0-8.5 mm long, (3.0–) 4.0-5.5 mm wide, pilose. Pistil subsessile to shortly stipitate, pubescent; ovary linear, 6.0-8.5 mm long, 0.8-1.0 mm wide; ovules 6 to 9; style shorter than ovary, 4.0-5.5 mm long. Pods oblong to slightly falcate, laterally compressed, membraneous, 11-21 (-28) mm long, (2.5–) 3.0-4.0 (-6.0) mm wide, densely tomentose; stipe very short (less than 1 mm long), ±4 to 6-seeded, indehiscent. Seeds reniform, 2.0-2.5 mm long, 1.5-2.0 mm wide, colour, surface smooth (Fig. 6.28). Flowering time: Flowering occurs from spring to summer

(September to December).

Diagnostic characters—Calobota pungens is similar to Calobota cuspidosa and C. psiloloba, but differs from both in the sericeous stems, sericeous obovate leaves, pilose petals, pubescent ovary and tomentose pods (in C. cuspidosa the stems are pubescent and the leaves linear to slightly obovate, while in C. psiloloba the stems are glabrous or slightly pilose and the leaves slightly obovate to spathulate. In both these species the ovary and pods are glabrous).

Distribution and habitat—Calobota pungens occurs from Laingsburg through Ladysmith and Oudtshoorn to Baviaanskloof and Willowmore in the east at altitudes of between 790 m and 1100 m (Fig. 6.29). It is typically a fynbos species and occurs on rocky, loamy or quartzitic, soils.

150

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

e a

j1 j2 j3

g k1

h2 aleenaMftsib h1 k2 dl d2

FIG. 6.28 Morphology of Calobota pungens: (a) flower in lateral view; (b) standard petal; (c) outer surface of the calyx (upper lobes to the left); (d1—d2) leaves in abaxial view; (e) wing petal; (f) keel petal; (g) pistil; (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view;

(k2) pod in dorsal view.

Voucher specimens: (a, b, dl, e—g, h2, i , k1—k2) Boatwright et al. 106 (JRAU); (c)

Taylor 9386 (NBG); (d2) Barker 9948 (NBG); (h1 ) Acocks 15547 (PRE); (j1—j3) Van Wyk

3252 (JRAU). Scale bars: (d1—d2, k1—k2) 1 cm; (a—c, e—j3) 1 mm.

Additional Specimens Examined

—3320 (Montagu): Cadibu (—AB), Bond 12112 (BOL, NBG); Matjiesfontein (—BA), Compton

2714, 3315 (BOL, K), Compton 22233 (NBG 2 sheets), Foley 58 (PRE), Marloth 10764

151 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(NBG 2 sheets, PRE); top of Koppie near Matjiesfontein (—BA), Thoday and DeIf (NBG);

Rooikloof (—BD), Van Wyk 2147 (JRAU).

—3321 (Ladysmith): Farm Withoek (—AC), Van Wyk 2154 (JRAU); road from Seweweeks

Poort (—AD), Levyns 2334 (BOL); Touwsberg, farm Rietfontein (—CA), Germishuizen 6956

(PRE); Touwsberg, farm Wolwefontein (—CA), Van Wyk et al. 3447 (JRAU, PRE).

—3322 (Oudtshoorn): 14 mls [22.5 km] east of Prince Albert (—AA), Sidey 1906 (PRE, S);

Swartberg Pass (—AC), Bond 1537 (NBG, PRE), Markotter 9951 (NBG); Congo Valley close

to Oudtshoorn (—AC), Dreyer 18618 (NBG); northern foothills of Swartberg above farm

Rietfontein (—AC), Vlok 1152 (K, PRE); Venterskloof near Klaarstroom (—AD), Taylor 9386

(K, NBG); 20 mls [32.2 km] west of Klaarstroom (—AD), Acocks 15547 (K, PRE); on road

between Prins Albert and Klaarstroom (—AD), Van Wyk EvW0151 (K); Meiringspoort (—BC),

Boatwright et al. 106 (JRAU), Levyns 6619 (BOL), Stokoe 61389 (NBG); outside

Meiringspoort, past Klaarstroom turn-off along roadsides (—BC), Boatwright et al. 111

(JRAU); foot of Swartberg Mountains, near Klaarstroom (—BC), DrOge s.n. (S, K);

Klaarstroom, Prince Albert (—BC), Barker 9948 (NBG 2 sheets).

—3323 (Willowmore): Hot Springs (—AC), Fourcade 5841 (NBG), Fourcade 6086 (BOL,

NBG); flats between Hot Springs and Toorwater (—AC), Thompson 1398 (PRE); lower

slopes of Zuurberg at Georgida (—AD), Fourcade 4410 (BOL, K); 7 mls [11.26 km] south- west of Willowmore (—AD), Horn s.n. (PRE); 6-8 mls [9.6-12.8 km] south-east of

Willowmore on Baviaanskloof road (—AD), Wells 2606 (PRE); 7 mls [11.2 km] on

Baviaanspoort road from Willowmore (—AD), Wells 2838 (GRA, PRE); 18 ml [28.8 km] from

Uniondale on Willowmore road (—AD), Wells 3699 A and B (GRA, PRE); along road from

Willowmore to Baviaanskloof near Uniondale turnoff (—BC), Snijman 353 (NBG, PRE);

Kammanassie hill sides 7 mls [11.3 km] from Uniondale (—CA), Fourcade 3594 (BOL 2 sheets, K); Vet Vlei, 8 mls [12.8 km] from Uniondale (—CA), Fourcade 6255 (BOL);

Pietselaagte on road to Uniondale (—CA), Van Wyk 3180 (JRAU); dirt road between De Rust and Uniondale (—CA), Van Wyk 3252 (JRAU).

152 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

FIG. 6.29 Known geographical distribution of Calobota pungens.

13. CALOBOTA SAHARAE (Coss. and Dur.) Boatwr. and B.-E.van Wyk, comb. nov. (S.

Afr. J. Bot.: submitted. 2008). Genista saharae Coss. and Dur., B. Soc. Bot.

France 2: 247. 1855. Spartidium saharae (Coss. and Dur.) Pomel, Nouv. Mat.

FL Atl.: 173. 1874.—TYPE: ALGERIA, D'Oran Province, Cosson s.n. (P-

COS?, type not seen).

Erect or virgate, multi-stemmed, unarmed shrub up to 2 m in height.

Branches green; young branches densely pilose or glabrous; older branches pilose or glabrous with light brown bark. Leaves simple; petiole absent; linear, alternate, pilose on both surfaces, sessile, 6-10 x 1-2 mm, apex acute, base cuneate.

Inflorescence 75-150 mm long, with 5 to 12 flowers; pedicel 1-2 mm long; bract

1.5-2.0 mm long, linear, pubescent; bracteoles 0.5-1.0 mm long, linear, pubescent.

Flowers 13-17 mm long, yellow. Calyx 4.5-7.0 mm long, pilose; tube 3-4 mm long;

153 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

lobes 1.5-3.0 mm long, subulate. Standard 13-15 mm long, claw linear, 3.5-4.0 mm

long, lamina ovate, 9.5-11.0 mm long, 7.5-8.0 mm wide, dorsal surface pubescent;

apex obtuse. Wings 11.5-14.0 mm long, claw 4.0-4.5 mm long, lamina oblong,

longer than keel, 7-9 mm long, 3-4 mm wide, glabrous, with 4 rows of sculpturing.

Keel 12.5-13.0 mm long, claw 4-5 mm long, lamina boat-shaped, 8.0-8.5 mm long,

4.5-5.0 mm wide, glabrous. Pistil subsessile to shortly stipitate, glabrous; ovary

linear, 9.0-11.5 mm long, 1.0-1.3 mm wide with 7 to 8 ovules; style shorter than

ovary, 4.5-5.5 mm long. Pods oblong-linear, laterally compressed, membraneous,

subsessile to shortly stipitate, 30-40 mm long, 6-10 mm wide, 4 to 6-seeded,

indehiscent. Seeds reniform to oblong-reniform, 2.5-3.0 mm long, 1.5-2.0 mm wide,

mature seeds light brown, surface smooth (Fig. 6.30). Flowering time: Flowering

occurs between from March to May.

Diagnostic characters—Calobota saharae is similar to Calobota cuspidosa

and C. psiloloba in the laterally compressed, membraneous pods, but differs in the

unarmed habit, linear leaves and pubescent standard petal (in C. cuspidosa and C.

psiloloba the branches are strongly spinescent, the leaves slightly obovate to elliptic

or spathulate and the petals pubescent). Calobota saharae is also similar to C.

namibiensis, but differs in the linear leaves, glabrous, the glabrous keel petals and

the membraneous pods (in C. namibiensis the leaves are elliptic to ovate, the keel

petals densely pubescent and the pods thick-walled and tomentose).

Distribution and habitat—Calobota saharae is endemic to North Africa and occurs in Morocco, Algeria, Libya and Tunisia. It is found on desert hills or dunes at altitudes of ca. 200m. (Fig. 6.31)

154 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

h1 2 44 k2

FIG. 6.30 Morphology of Calobota saharae: (a) flower in lateral view; (b) standard petal; (c) outer surface of the calyx (upper lobes to the left); (d) leaf in abaxial view; (e) wing petal; (f) keel petal; (g) pistil; (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther;

(j2) intermediate carinal anther; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a, e, g) Keith 181 (K); (b, c, f, h1—j3) Hill s.n. (K); (d) Pitard

3276 (K); (k1—k2) Alleizette s.n. sub K 2007/014804 (K). Scale bars: (d, k1—k2) 1 cm; (a—c, e—j3) 1 mm.

Additional specimens examined

Algeria: Ain Sefra, Alleizette s.n. sub K 2007/014804 (K), Cosson s.n. sub K 000227004

(K), Hill s.n. (K), Bourgeau 210 (K 2 sheets), Le Cesve 5429 (BM), Rothschild and Hartert s.n. (BM), Samuelsson 6994 (K); Constantine Province, Oued Souf, Cosson s.n. sub K

000227008 (BM, K); Djebel Milogh, Cosson s.n. sub K 000227003 (K); Oran Province, /kin

Sefifsifa, Cosson s.n. sub K 000227005 (K); Oran Province, Abiod-Sisi-Cheihk, Cosson s.n.

155 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA sub K 000227006 (K); Wadi Rummel, above Sidi Benour, Guichard KG/Lib/289 (K); El

Ardja, Pitard 3275 (K); Oued el Khezoua/EI Oued, Pitard 3276 (K).

Tunisia: Nefta, Cosson s.n. sub K 000227007 (K).

Libya: /kin Zara, near Pripoli, Brown s.n. (K); Jebel Nefoussa, Azizia-Rhnem, Davis 49544

(K); Jebel, north of Jefren, Keith 181 (K).

20 10 10 20 30 40 50 60 11111161111R1111M11111111 1

50 50

40 mempubwmaimirrarwth dint 40

30 M.% 30 AMIN 1111111111111VIM=

20 41111 1111111111111=11111b 20 Ppm yip=

10 11111111e...ig 10 ELM 111711

10 10

20 11111111111111=11111111.1101111 20 111111111111111=11111 30 1111, 30

NB! 1991 1 10 10 20 30 40 50 60

FIG. 6.31 Approximate geographical distribution of Calobota saharae.

156 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

14. CALOBOTA SERICEA (Ait.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J. Bot.:

submitted. 2008). Spartium sericeum Ait. in Hort. Kew. 3: 12. 1789.—TYPE:

SOUTH AFRICA, Cape of Good Hope, Masson s.n. sub BM000794145

(lectotype: BM, photo!, specimen on the left, designated here). [The two

specimens on the left were collected by Masson and are marked with '1' on

the tape on the stems. The specimen on the right was grown in the gardens

at Kew and has the note 'Hort. Kew 1781' associated with it. According to

Stafleu and Cowan (1976), most of the Aiton types are in the Banks collection

in BM).]

Lebeckia angustifolia E.Mey. in Linnaea 7: 155. 1832. Acanthobottya angustifolia

(E.Mey.) Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 194. Jan. 1836.—TYPE:

SOUTH AFRICA, without locality, Ecklon s.n. (lectotype: S!, designated

here). [Note: the type is represented by the two branches mounted at the

bottom right hand side of the sheet – one with several flowers, the other with

a single bud. Meyer (1932) incorrectly cites the type locality as "Uitenhage?"

but expresses doubt by placing a question mark after the locality. The

specimen in S comprises mixed elements but the two branches referred to

above are accompanied by a label in Meyer's handwriting stating "Lebeckia

angustifolia mihr.]

Lebeckia multiflora E.Mey., Comm. Pl. Afr. Austr. 1: 34. Feb. 1836; Benth. in Hook.

Lond. J. Bot. 3: 360. 1844; Walp., Rep. Bot. Syst.: 453. 1845; Harv. in Harv.

and Sond., Fl. Cap. 2: 88. 1862; Merxm. and Schreiber, FSWA 60: 68. 1970;

Bond and Goldblatt, J. S. Afr. 13: 290. 1984; Goldblatt and Manning, Cape

Pl.: 493. 2000.—TYPE: SOUTH AFRICA, between Holrivier and

Mierenkasteel [3118 CB], DrOge 6474 "III, C" (lectotype: P!, designated here;

157 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

isolectotypes: BM!, K!, PRE!, S!). [Note: This specimen bears the type

locality, shows the features of the plant described by Meyer and bears the

characteristic Drege field labels]

Lebeckia decutiens var. a canescens E.Mey., Comm. Pl. Afr. Austr. 1: 34. Feb.

1836.—TYPE: SOUTH AFRICA, near Heerenlogement [3118 DC], DrOge s.n.

"III, E, a"(lectotype: P!, designated here; isolectotype: BM!, K!). [Note: The

specimen displays the diagnostic characters described by Meyer and bears

the type locality.]

Lebeckia parvifolia (Schinz) Harms in Engl. Pflanzenw. Afr. 3(1) (Engl. and Drude,

Veg. der Erde 9): 545 (1915). Lebeckia multiflora var parviflora Schinz in

Abhandl. Bot. Ver. Prov. Brandenb. 30: 157. 1888.—TYPE: not seen.

Erect, multi-stemmed, unarmed or rarely spinescent shrub up to 2 m in height. Branches green; young branches sericeous, older branches sericeous or glabrous with brown bark. Leaves digitately trifoliolate; petiole longer or shorter than leaflets, 8-45 mm long; leaflets linear to oblanceolate, alternate, sericeous, subsessile, terminal leaflet 5-30 mm long, 0.5-5.0 mm wide, lateral leaflets 4-25 mm long, 0.5-4.0 mm wide, apex acute, base cuneate. Inflorescence 20-200 mm long, with 7 to 40 flowers; pedicel 1-2 mm long; bract 1-3 mm long, linear, pubescent; bracteoles 0.5-2.0 mm long, linear, pubescent. Flowers 7-11 mm long, bright yellow. Calyx 4.0-6.5 mm long, pubescent; tube 2.5-4.5 mm long; lobes 1.0-

3.5 mm long, deltoid. Standard 6.5-13.0 mm long; claw linear, 2.0-4.5 mm long; lamina widely ovate, 4-9 mm long, (3.8–) 4.5-10.0 mm wide, pilose along dorsal midrib; apex obtuse. Wings 7-13 (-14) mm long; claw 2.5-4.5 (-5.5) mm long; lamina oblong, as long as but more often longer than keel, 4.5-8.5 mm long, 2.0-4.5

158 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA mm wide, glabrous, with 4-12 rows of sculpturing. Keel 6.5-12.5 mm long; claw

2.5-4.5 (-5.5) mm long; lamina boat-shaped, 4-8 mm long, 2.5-5.0 mm wide, glabrous. Pistil subsessile to shortly stipitate, pubescent; ovary linear, 5.0-9.5 mm long, 0.5-1.1 mm wide with 11 to 20 ovules; style shorter than ovary, 2.5-5.0 mm long. Pods linear, semi-terete, sericeous, subsessile to shortly stipitate, 18-35 mm long, 2-4 mm wide, 5 to 12-seeded, dehiscent. Seeds oblong-reniform, 2.5-3.5 mm long, 1.5-2.0 mm wide, mature seeds light brown or light orange, sometimes mottled with brown, surface smooth (Fig. 6.32). Flowering time: Flowers for most of the year January to November.

Diagnostic characters— Calobota sericea is similar to C. lotononoides but differs in that it is an erect, divaricately branched shrub usually more than 1 m in height with inflorescences that often terminate in a spine; the wing petals are almost as long as or slightly longer than the pilose keel petals and the seeds are pale pink with a smooth surface (C. lotononoides is a small, decumbent shrublet with unarmed inflorescences, extremely long wing petals, glabrous keel petals and the seeds are brown with beige spots and a rugose surface).

It is also similar to C. halenbergensis but differs in the sericeous leaves, pilose keel petals, hairy ovary and sericeous pods with more seeds (in C. halenbergensis the leaves are pilose, the keel petals glabrous and the ovary and pods glabrous with fewer seeds in the latter).

159 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

d b

jl j2

k2 —

FIG. 6.32 Morphology of Calobota sericea: (al)) leaf in abaxial view (typical form); (a2) leaf in abaxial view (Oranjemund form); (a3) leaf in abaxial view (short-leaved form); (a4) leaf in abaxial view (broad-leaved form); (b) flower in lateral view; (c) standard petal; (d) wing petal;

(e) keel petal; (f) pistil; (g) outer surface of the calyx (upper lobes to the left); (h) androecium; (i1) long, basifixed anther; (i2) intermediate carinal anther; (i3) short, dorsifixed anther; (j1) bract; (j2) bracteoles; (kl ) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al, kl—k2) Boatwright et al. 138 (JRAU); (a2) Ward 12435

(WIND); (a3) Giess and Robinson 13177 (WIND); (a4) Stirton and Zantovska 11426 (NBG);

(b, d—f, h—i) Boatwright et al. 139 (JRAU); (c) Van Wyk 2847 (JRAU); (g) Lewis 4705 (NBG);

(j1—j2) Whitehead 80.9.8 (WIND). Scale bars: (al—a4, kl—k2) 1 cm; (a—c, ej3) 1 mm.

160 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Distribution and habitat—Calobota sericea is widely distributed along the west coast of South Africa and Namibia (Fig. 6.33). It forms an important component of Western Strandveld (FS 1 Lambert's Bay Strandveld, FS 5 Langebaan Dune

Strandveld; Rebelo et al. 2006), Richtersveld (SKr 3 Goariep Mountain Succulent

Shrubland and disturbed areas of SKr 13 Southern Richtersveld Scorpionstailveld;

Mucina et al. 2006) and Namaqualand Sandveld (SKs 2 Northern Richtersveld

Yellow Duneveld, SKs 4 Richtersveld Sandy Coastal Scorpionstailveld, SKs 5

Richtersveld Red Duneveld, SKs 6 Oograbies Plains Sandy Grassland and SKs 13

Klawer Sandy Shrubland, the latter recorded as C. halenbergensis instead of C. sericea; Mucina et al. 2006). Rebelo et al. (2006) recorded C. thunbergii as an important component of FS 4 Saldanha Limestone Strandveld, but it is C. sericea that occurs in that area.

Regional variation—Four regional forms can be recognized based on leaf morphology and geographical distribution:

FORM A (Typical form)

Erect shrubs up to 1.8 m in height, often spinescent. Leaves linear to narrowly oblanceolate, sericeous, terminal 10-30 mm long, 0.5-2.0 (-2.5) mm wide, lateral

8-23 mm long, 0.5-2.0 (-2.5) mm wide; petiole 11-35 mm. Inflorescence 27-90 mm; with 10 to 27 flowers; pedicel 1-2 mm; bract 1.5-2.0 mm, bracteole 0.5-1.5 mm. Flowers 9-11 mm. Calyx 4.5-6.0 mm long; tube 3.5-4.5 mm long; lobes 1-3 mm long. Standard 9-13 mm long; claw 2.5-4.5 mm long; lamina 5.5-9.0 mm long,

(-3.5) 4.5-10.0 mm wide. Wings 8.5-13.0 (-14.0) mm long; claw 3.0-4.5 (-5.5) mm long; lamina 5.5-8.5 mm long, 2.0-3.5 mm wide, glabrous, with 5-8 rows of sculpturing. Keel 8.0-12.5 (-13.0) mm long; claw 3.5-4.5 (-5.5) mm long; lamina

161 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

4.5-7.5 mm long, 2.5-4.5 mm wide, glabrous. Pistil ovary 5.5-9.5 mm long, 0.5-1.1

mm wide with 11 to 20 ovules; style 3-5 mm long. Pods sericeous, 18-35 x 2-3

mm.

Diagnostic characters—Populations of this form around Klawer and Van

Rhynsdorp have very long wing petals and small flowers. It differs from the other forms in the long, narrow leaflets and relatively short pods. The inflorescences are

often spine-tipped.

Distribution and habitat—This form occurs around Berg River, Piquetberg,

Clanwilliam, Velddrif, Lambert's Bay, Vredenburg, Hopefield, Langebaan, Saldanha,

Vanrhynsdorp, Heerenlogement, Wallekraal, Hondeklipbaai, Garies and Kleinsee on the West Coast of South Africa and does not extend into Namibia. It grows in well-

drained sand, sandy loam red sand or sand dunes between sea level and 500 m

above.

FORM B (Broad-leaved form)

Erect or spreading shrubs up to 2 m in height, sometimes spinescent. Leaves with

petiole 8-30 mm long; leaflets obovate to widely oblanceolate, sericeous, terminal

5-25 mm long, 2-5 mm wide, lateral 5-25 mm long, 2-4 mm wide. Inflorescence

20-65 mm; with 7 to 18 flowers; pedicel 1-2 mm; bract 2 mm long, bracteole 1.0–

1.5 mm long. Flowers 9-11 mm long. Calyx 5.0-6.5 mm long; tube 3.5-4.0 mm

long; lobes 1.0-2.5 mm long. Standard 10-11 mm long; claw 3.0-3.5 mm long;

lamina 6.5-8.0 mm long, 7.5-10.0 mm wide. Wings 10.5-11.5 mm long; claw 3-4

mm long; lamina 6.5-8.0 mm long, 2.5-4.5 mm wide, glabrous, with 4-7 rows of sculpturing. Keel 10-11 mm long; claw 3.5-4.0 mm long; lamina 6.5-8.0 mm long,

3.5-5.0 mm wide, glabrous. Pistil with ovary 7.0-8.5 mm long, 0.7-1.1 mm wide

162 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA with 11 to 15 ovules; style 4.5-5.0 mm long. Pods sericeous, 25-35 mm long, 3 mm wide.

Diagnostic characters—This form differs from the others in the broad leaflets (always more than 2 mm wide) and relatively short inflorescences.

Distribution and habitat—Occurs from the Saldanha area, Elands Bay,

Clanwilliam, Lambert's Bay, Doring Bay, mouth of Olifants River, Brandsebaai,

Garies, Wallekraal, Port Nolloth and Walvis Bay along the West Coast of South

Africa and Namibia. It grows in deep limestone or calcrete sand or dunes at altitudes of between 4 m and 50 m and is grazed by livestock.

FORM C (Short-leaved form)

Erect or virgate shrubs up to 2 m in height. Leaves with petiole relatively long, 9-45 mm long; leaflets linear to oblanceolate, very densely sericeous, terminal 2-13 mm long, 1-2 mm wide, lateral 2-7 mm long, 0.5-1.0 mm wide. Inflorescence 30-200 mm; with 8 to 40 flowers; pedicel 1 mm; bract 2-3 mm long, bracteole 1.5-2.0 mm long. Flowers 7-10 mm long. Calyx 4-6 mm long; tube 2.5-4.0 mm long; lobes 1.5-

3.5 mm long. Standard 7.5-9.0 mm long; claw 2-3 mm long; lamina 5.5-6.5 mm long, 5.5-7.0 mm wide. Wings 8.5-10.0 mm long; claw 2.5-4.0 mm long; lamina 6-

7 mm long, 3-4 mm wide, glabrous, with 4-10 rows of sculpturing. Keel 7.5-8.5 mm long; claw 2.5-3.5 mm long; semi-circular, 4-5 mm long, 3.0-3.5 mm wide, glabrous. Pistil with ovary 5.5-6.5 mm long, 0.7-1.0 mm wide with 13 to 18 ovules; style 3.0-3.5 mm long. Pods densely sericeous, 20-40 mm long, 3-4 mm wide.

Diagnostic characters—In this form the leaflets are very short with relatively long petioles.

163 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Distribution and habitat—This form of C. sericea is restricted to Namibia

and occurs in the Luderitz Bay area, Boegoeberg, Bogenfels, Pomona, Chamais,

Alexander Bay and Angra Pequena. It grows in well-drained sand or coarse sand,

sometimes on rocky soil or dolomite outcrops at altitudes of between 20 m and 120

m. It is heavily grazed by livestock.

FORM D (Oranjemund form)

Erect shrubs up to 1.5 m (-1.8 m) in height. Leaves with petiole 12-35 mm; leaflets

narrowly oblanceolate, sericeous, terminal 5-14 mm long, 0.5-1.0 mm wide, lateral

5-9 mm long, 0.5-1.0 mm wide. Inflorescence 30-100 mm; with 13 to 15 flowers;

pedicel 1-2 mm long; bract 1-3 mm long, bracteole 1.0-1.5 mm long. Flowers 7-10

mm long. Calyx 4.5-5.0 mm long; tube 3.0-3.5 mm long; lobes 1-2 mm long.

Standard 6.5-8.0 mm long; claw 2-3 mm long; lamina 4.0-5.5 mm long, 5.5-6.5

mm wide. Wings 7.0-8.5 mm long; claw 2.5-3.5 mm long; lamina 4.5-5.5 mm long,

3.0-3.5 mm wide, glabrous, with 8-12 rows of sculpturing. Keel 6.5-8.0 mm long;

claw 2.5-3.5 mm long; lamina 4.0-4.5 mm long, 3.0-3.5 mm wide, glabrous. Pistil

with ovary 5-6 mm long, 0.8-1.0 mm wide with 12 to 19 ovules; style 2.5-3.5 mm

long. Pods sericeous, 20-32 mm long, 2-3 mm wide.

Diagnostic characters—Leaflets of the Oranjemund form are short and very

narrow and the pods also relatively short.

Distribution and habitat—This form is centered in the Oranjemund district. It

occurs in sandy areas, alluvial sand and dunes near the mouth of the Orange River

and in the Oranjemund mining area at altitudes of between 10 m and 100m.

Additional specimens examined

–2214 (Swakopmund): Walvis Bay (–DC), Heywood s.n. (NBG).

164 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

—2514 (Spencer Bay): ca. 2 km east of coast near East Hill between Oyster Cliffs and

Easter point (—BD), Seely and Ward 54 (WIND); Spencer Bay (—DB), Giess and Robinson

13177 (K, S, PRE, WIND), Sydow 10179 (WIND).

—2516 (Helmeringhausen): Halbinsel (—CA), Walter 56 (WIND).

—2615 (Luderitz): Schurmhaudeberg (—AA), Hardy and Venter 4437 (K, PRE, WIND); Angra

Pequena, Marloth 4663 (NBG, PRE), Schenck 9 (PRE), Schinz 811 (BOL, K); 14 mls [22.53 km] east of Luderitz Bay (—CA), Acocks 15654 (K, PRE 2 sheets); Luderitz peninsula (—CA),

Clark and Mailer 337 (WIND), De Winter and Hardy 7894 (K, WIND), Mannheimer CM2020

(WIND); Luderitz Bay (—CA), De Winter and Giess 6233 (BOL, K, PRE, WIND), Dinter 3838

(BM, BOL, K, NBG, PRE, S), Dinter 5996 (BM, BOL, K, NBG 2 sheets, PRE), Giess and

Van Vuuren 678 (PRE, WIND), Lavranos and Pehleman 19641 (WIND 2 sheets),

Leuenberger et al. 3306 (PRE, WIND), Merxm011er and Giess 28275 (K, PRE, WIND), Metz s.n. (WIND 2 sheets), Peter 47177 (K), Range 363 (BOL), Meyer 56 (WIND), Volk 12825

(WIND), Wall s.n. (S); 13 mls [20.92 km] east of Luderitz, road to Aus, Grasplatz (—CA),

Giess and Van Vuuren 738 (PRE, WIND); Luderitz (—CA), Hobart-Hampden 11 (BM),

Lavranos and Barod 15409 (WIND), Moss 11462 (BM), Rogers 29588 (K 2 sheets); just outside Luderitz along main road (—CA), Mannheimer CM2015 (WIND); Luderitz lagoon (-

CA), Merxm011er and Giess 3133 (WIND); Luderitz, just outside township (—CA), Muller and

Jankowitz 275 (WIND); Essy Bay (—CA), Strohbach 151 (WIND); Luderitz Bay, Halbinsel (-

CA), Walter 56 (WIND), Wendt s.n. (WIND 2 sheets); Aus (—CB), Regius 23c (WIND);

Grasplatz (—CB), Merxmiiller and Giess 3052 (PRE, WIND), Venter 9160 (PRE).

—2715 (Bogenfels): Pomona (—AB), Dinter 6336 (BM, BOL, K, NBG 2 sheets, PRE), Kolle

13560b (NBG, PRE), Wall s.n. (S), Whitehead 80.9.8 (WIND); along road to Pomona (—AB),

Mannheimer et al. CM898 (WIND); blue rocky ridge on way to Pomona town (—AB),

Mannheimer CM 924 (WIND); Bogenfels (—AD), De Winter and Giess 6220 (K, PRE,

WIND), Merxm011er and Giess 28326 (WIND); area and surrounds north-east of Bogenfels

Arch (—AD), Powrie LWP1170 (WIND); Diamond Area 1 (—AD), Watmough 867 (PRE);

Chamais (—DC), Merxmilller and Giess 28319 (K, PRE, WIND); Chamais valley in rock (—

165 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

DC), Williamson 2647 (BOL); 3 km east of Chamais Bay (—DC), Williamson 4980 (WIND);

Boegoeberg (—DD), Merxm011er and Giess 28304 (PRE, WIND), Williamson 2575 (BOL).

—2816 (Oranjemund): between Oranjemund and check point (—BC), Gess and Gess

97/98/68 (J, WIND); Oranjemund (—CB), Merxmiiller and Giess 2267 (BM, K, WIND); sandy area near mouth of Orange at Oranjemund (—CB), Metelerkamp 290 (BOL), Schenck 238

(PRE); north of Orange River (—CB), O'Callaghan et al. 5 (NBG, PRE); hills north-east of

Alexander Bay (—CB), O'Callaghan et al. 67 (NBG, PRE); south bank of lagoon at mouth of

Orange River (—CB), Pillans 5636 (BOL); near airport turnoff from main entrance road (-

CB), Ward 11136 (K, PRE, WIND); Orange River, upper estuary (—CB), Ward 12435 (PRE); mining area 1 (—CB), Williamson 4844, 4851 (WIND); k122 mining area Ev5 control plot (-

CB), Williamson 4884 (WIND); Witbank (—DC), Pillans 5207 (BOL, K), 40 km from Port

Nolloth on road to Alexander Bay (—DD), Germishuizen 4770 (PRE, WIND), Germishuizen

4771 (PRE); 32 km from Port Nolloth on road to Alexander Bay (—DD), Germishuizen 4813

(PRE).

—2817 (Vioolsdrif): "ad ostia fluvii Garip" (—AA), DrOge s.n. "Ill, B"(P, K); sides of dry river bed at Kuboos (—AC), Pillans 5429 (BOL).

—2820 (Kakamas): Farm Harperspot, east of Riemvasmaak (—AD), Gubb KMG13032

(WIND).

—2916 (Port Nolloth): Port Nolloth (—BD), Bolus 9503 (K), Galpin and Pearson 7605 (BOL,

K, NBG, PRE), Marloth 4747 (NBG, PRE); 1 km south of McDougalls Bay (—BD), Raitt 302

(NBG); between Port Nolloth and Holgat (—BD), Schlechter s.n. sub STE 10850 (NBG 2 sheets); Port Nolloth, at turn-off to McDougall Bay (—BD), Van Wyk 2847 (JRAU 2 sheets); 9 km east of Port Nolloth (—BD), Zietsman and Zietsman 823 (PRE).

—2917 (Springbok): sand by roadside between Anenous and Chubiessis outspan (—BA),

Pearson 5978 (BOL, K); Oubeep Bay (—BD), Le Roux and Ramsey 144 (NBG); 2 km south of Kleinsee (—CA), Hugo 2849 (PRE 2 sheets); Buffels River mouth (—CA), Le Roux and

Parsons 23 (PRE); Kleinsee (—CA), Theron 3858 (PRE); between Brakwater and Komaggas

(—DC), Compton 22071 (BOL, NBG).

166 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

—3017 (Hondeklipbaai): 6 mls [9.65 km] south-east of Hondeklipbaai (—AD), Hall 899 (BOL,

NBG); Farm Oubees 339, 1.2 km east of Oubees/Wildepaardehoek boundary fence and 4.7 km east of Springbok, Soebatsfontein road (—BA), Le Roux and Lloyd 670 (NBG); between

Brakwater and Komaggas (—BB), Compton 22071 (BOL, NBG); 3 mls [4.83 km] north-west of Wallekraal Post Office (—BC), Acocks 14933 (K, PRE 2 sheets); 5 mls [8.05 km] north- west of Wallekraal (—BC), Hall 895 (NBG 2 sheets); Wallekraal, near Hondeklipbaai (—BC),

Stirton 6036b (K, PRE); 15 mls [24.14 km] east of Wallekraal on road to Garies (—BC),

Thompson 1090 (K, NBG); Groen River Mouth (—DC), Le Roux and Ramsey 276 (K, NBG,

PRE), Le Roux and Ramsey 280 (NBG).

—3018 (Kamiesberg): Ezelkop, Garies (—AC), Scheffler 266 (NBG).

—3117 (Lepelfontein): Brandsebaai (—BD), De Villiers 34 (PRE, WIND), De Villiers 48, 49

(PRE), Van Rooyen 2200 (PRE).

—3118 (Vanrhynsdorp): mouth of Olifants River, Papendorp (—CA), Arnold 916 (PRE);

Vredendal, Olifants River Mouth (—CA), Le Roux and Ramsey 62 (NBG); Olifants River

Mouth (—CA), O'Callaghan 657 (NBG, PRE); Vredendal road (—CB), Steyn 464 (NBG);

Doringbaai (—CC), Boucher 4054 (NBG, PRE), Hugo 2924 (PRE); Holrivier (—CB), DrOge s.n. (S); between Vredendal and Lambert's Bay (—CD), Lewis 4705 (NBG), Schutte 269

(JRAU); 10.7 km from Doringbaai to Donkinbaai (—CD), Stirton 6077 (K, PRE); Zandkraal (-

DA), Acocks 14840 (PRE); 11.7 km south of Redelinghuys (—DA), Acocks 24284 (K, PRE);

Kys, south-east of Vredendal (—DA), Bayer 6396 (PRE); Farm Liebendal, Vredendal (—DA),

Hall 3701 (NBG 2 sheets); Klawer (—DC), Andreae 474 (NBG, PRE); Heerenlogement (-

DC), DrOge s.n. "Ill, E, a" (BM, K, P), Esterhuysen 5547 (BOL); Vanrhynsdorp, foot of

Nardousberge below Witbakenkop on old Clanwilliam, Vanrhynsdorp road (—DC), Hilton-

Taylor 1613 (NBG); western aspect of koppie on Vanrhynsdorp road, Klawer (—DC), Lavis

20236 (BOL, K 2 sheets); 6 mls [9.65 km] north of Klawer (—DC), Maguire 142 (NBG 2 sheets); 6 mls. [9.65 km] south of Klawer (—DC), Salter 5612 (BM, BOL, K, PRE).

—3217 (Vredenberg): Stompneus Point (—DB), Taylor 1515 (NBG); Brittania Bay (—DB),

Taylor 5189 (NBG, K, PRE); Farm Trekossenkraal (—DD), Boucher 7065 (NBG); Cape

167 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Columbine Lighthouse (—DD), Horrocks 182 (NBG 2 sheets); slopes south of Vredenburg (-

DD), Hutchinson 247 (BM, BOL, K, PRE); granite hills near Vredenburg (—DD), Marloth

7945 (PRE).

—3218 (Clanwilliam): Farm Brakke Kuil 9, on road from Laaiplek to Elands Bay (—AB),

Bosenberg and Rutherford 394 (NBG); Lamberts Bay (—AB), Acocks 14182 (K, PRE),

Clarke 633 (K, PRE), Henrici 3309 (PRE), Muir s.n. (NBG), Pole-Evans 26 (PRE), Walsh s.n. (PRE); Groendam (—AB), Stirton 9349 (PRE); 5 km from Lamberts Bay to Elands Bay (-

AB), Stirton and Zantovska 11426 (JRAU, NBG, PRE); Wadrifsoutpan (—AB), Stirton 9340

(PRE); Farm Middelpos (—AB), Stirton 9387 (PRE); Nortier Experimental Station, Lambert's

Bay (—AB), Van Breda 614 (NBG), Van Breda 4435 (PRE); 2 mls [3.22 km] from

Redelinghuys on Aurora road (—AD), Barker 9722 (NBG); Elands Bay (—AD), Barker 2627

(NBG), Britton 22 (NBG, PRE), Pillans 7962 (BOL), Taylor 3912 (NBG), Taylor 3915 (PRE),

Walsh s.n. (PRE); Vredendal road from Graafwater (—BA), Barker 8545 (NBG 3 sheets); between Leipoldtville and Graafwater (—BA), Esterhuysen 3765 (NBG), Leipoldt 3765 (BOL,

NBG); Nieuwoudtville turn-off after Graafwater on road to Lamberts Bay (—BA), Stirton 9197

(PRE); Farm Rieffontein, Graafwater (—BA), Van Blerk 38 (PRE); 5 mls [8.05 km] from

Clanwilliam on Lamberts Bay road (—BB), Gillett 4045 (PRE); Rocher Pan Nature Reserve

(—CB), Hey! 44 (NBG, PRE), Le Roux and Van Rooyen 1 (NBG); on road to Hopefield from

Velddrif (—CC), Boatwright et al. 138 (JRAU); road from Velddrif to Saldanha Bay (—CC),

Boatwright et al. 139 (JRAU); Velddrif (—CC), Compton 15940 (NBG), Goldblatt 6016 (PRE),

Lewis 899 (NBG), Mauve 5356 (K, PRE); 1 km from road between Velddrif and Vredenburg on road to Stompneusbaai (—CC), Grobbelaar 2546 (PRE); Velddrif, next to road before the bridge going over the Berg River (—CC), Le Roux 3 (JRAU 4 sheets); between Hopefield and Paternoster (—CC), Leipoldt 3764 (BOL); 1 ml [1.61 km] from Velddrif, on banks of Berg

River (—CC), Marsch 1265 (K, NBG, PRE); ca. 6 km east of Velddrif, near Ouwerf (—CC),

O'Callaghan 1237 (NBG); Berg River Station (—CD), Barker 4050 (NBG); railway enclosure along road 0.5 mls [0.8 km] from Berg River Station (—CD), Boucher 83 (NBG, PRE); South

168 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA side of Berg River (—CD), Leighton 614 (BOL); Piquetberg (—DA), Van Aarde s.n. (NBG);

Kapteinskloof (—DC), Stirton 6132b (K, PRE).

—3219 (Wuppertal): Doom River Mouth (—DA), Compton 11036 (NBG), Doorn River (—DA),

Schlechter 8059 (BM, BOL, K, PRE 2 sheets).

—3317 (Saldanha): 20 m off road in shooting range camp parallel to dunes (—BB), Blake 91

(NBG); Hoedjies Bay (—BB), Bolus 12652 (PRE); Saldanha Bay (—BB), Grey s.n. (K),

Marloth 10188 (PRE); koppie north of Saldanha Bay (—BB), Van Wyk 2696 (JRAU 5 sheets).

—3318 (Cape Town): Lagoonside, Langebaan (—AA), Axelson 445 (NBG); roadside south of

Langebaan village (—AA), Boucher 2811a (NBG); near Langebaan (—AA), Chaplin s.n.

(NBG), Marloth 13176 (PRE); Geelbek (—AA), Compton 19906 (NBG); Langebaan peninsula above Preekstoel (—AA), O'Callaghan and Steensma 1650 (NBG); Hopefield (-

AB), Bachmann 1889 (BM, K); 10.7 mls [17.22 km] from Hopefield to Vredenberg (—AB),

Marsch 182 (NBG, PRE); 14 km from Hopefield to Langebaanweg (—AB), Stirton 10709 (K);

Schrywers Hoek (—AD), Macnac 1020 (PRE); near Bergrivier and Zwartland (—BD), Ecklon and Zeyher 1346 (S); sandy hill slopes behind the village Langebaan (—DC), Leighton s.n. sub BOL 32422 (BOL).

—3322 (Oudtshoorn): Waenskloof, Cango Valley (—AC), Moffett 494 (PRE).

Precise locality unknown: Olifantsrivier, DrOge s.n. (P); without locality, Ga/pin and

Pearson 7579 (PRE), Leipoldt s.n. (NBG), Range 1941 (NBG); Bokkeveld Mountains,

Leipoldt 730 (PRE); Little Namaqualand, Krapohl 11142 (PRE); Lus district, Merxmilller and

Giess 2349 (WIND); sandy ravine below Doornpoort, Pearson 6017 (BOL, K); Waterkloof at

Doornpoort, Pillans 5377 (BOL); Namaqualand, Scully s.n. (NBG).

169

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

12 14 16 18 20 22 24 26 28 30 32aim 34 AIM ..11_11maciromin ammil8 101111111111111111 111111112110111111 1 111111111111111 • MIT 1111111Si , 111111111111111111 171 mai 111111111111111111 Sp 11/111111h ..16-13 1111,11i ' 11111111511111111111 in 11111111111 r41 1111111113 1111 orammismuma '' IIMIIIIIIIIIII ■II■VANN ■ 111A1111111M111111 11111111111111111 II 1111/1 ■ +61 1111./1111 WWI 11111111111111 • 11111111111011 • 1111 1111116/41111111 1111116 ■ ■ ■ NMI 11J11111111 ' 11111111111111 ■ II ■II . 2 VII IN o a 1111V111 MI REIM ■ 4 1111111111110 , 1111111 1 I ri lo111111•is a r=I 4r. INN'-• MI MI MI• 2111_111M sersiMens 4 II1111111111 mine < mliaiiiii1111 ialAriling 34 36 10 12 14 16 18 20 22 24 26 28 30 32

FIG. 6.33. Known geographical distribution of Calobota sericea.

15. CALOBOTA SPINESCENS (Harv.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia spinescens Harv. in Harv. and Sond., Fl.

Cap. 2: 88. 1862; Wilman, Preliminary checklist of flowering plants and ferns

of Griqualand West (South Africa): 53. 1946; Merxm. and Schreib., FSWA 60:

68. 1970; Goldblatt and Manning, Cape Pl.: 494. 2000.—TYPE: SOUTH

AFRICA, Dwaka River, Burke s.n. (lectotype: K!, designated here;

isolectotypes: BM! 2 sheets). [Note: The Burke specimen in Kew was

determined by Harvey himself as "Lebeckia spinescens Harv." and is here

chosen as lectotype.]

Lebeckia armata auct. non Thunb.: E.Mey., Comm. Pl. Afr. Austr. 1: 35. Feb. 1836.

170 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Lebeckia elongata Hutch. in Ann. Bol. Herb. 3: 2. 1923.—TYPE: NAMIBIA, without

locality, Pearson 9899 (holotype: BOL!, isotype: K!).

Erect to spreading, multi-stemmed, spinescent shrub up to 1.0 m in height.

Branches green; young branches sericeous; older branches sericeous or glabrous with light brown bark. Leaves digitately trifoliolate; petiole longer than leaflets, 6-25 mm long; leaflets linear to oblanceolate or spathulate, alternate, sericeous, subsessile, terminal leaflet 3-12 mm long, 0.5-2.0 mm wide, lateral leaflets 2-11 mm long, 0.5-2.0 mm wide; apex acute; base cuneate. Inflorescence 25-105 mm long, with 3 to 10 flowers; pedicel 1-3 mm long; bract 1-2 mm long, linear, pubescent, caducous; bracteoles 0.5-1.5 mm long, linear, pubescent, caducous.

Flowers 10-16 mm long, bright yellow. Calyx 4.5-7.0 mm long, pubescent; tube 3-5 mm long; lobes 1.0-2.5 mm long, subulate. Standard 10.0-15.5 mm long; claw linear, 2-4 mm long; lamina ovate, 8.5-12.0 mm long, 7.5-11.0 mm wide, dorsal surface sericeous; apex acute. Wings 10.5-14.0 mm long; claw 2.5-4.0 mm long; lamina oblong, as long as or shorter than keel, 7.5-10.0 mm long, 2-4 mm wide, glabrous or rarely pilose, with 4-6 rows of sculpturing. Keel 11-15 mm long; claw

3.5-5.0 mm long; lamina boat-shaped, 7-10 mm long, 3.5-6.0 mm wide, glabrous or rarely pilose. Pistil subsessile to shortly stipitate, pubescent or glabrous with pilose hairs on margins; ovary linear, 7.5-11.5 mm long, 1.0-1.5 mm wide with 10 to

16 ovules; style longer than ovary, 3.5-6.0 mm long. Pods linear, somewhat falcate, laterally compressed, subsessile to shortly stipitate, 20-40 mm long, 3-4 mm wide,

±4 to 6-seeded, either sericeous or glabrous and pilose along upper suture, dehiscent. Seeds suborbicular to reniform, 2.4-2.5 mm long, 2.1-2.2 mm wide (only one specimen available for measurement), mature seeds light brown, surface

171

CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA smooth (Fig. 6.34). Flowering time: Flowering occurs from February to May and

September to October. One specimen collected in December (summer and spring flowering).

■11•1•11

f j1 j2 v j3

g1 g2

FIG. 6.34 Morphology of Calobota spinescens: (al—a2) leaves in abaxial view; (b) flower in lateral view; (c) standard petal; (d) wing petal; (e) keel petal; (f) pistil; (g1) bract; (g2) bracteoles; (h) outer surface of the calyx (upper lobes to the left); (i) androecium; (j1) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (kl) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al, b—e, gl—k2) Boatwright et al. 158 (JRAU); (a2) Koekemoer

248 (JRAU); (f) Zietsman 3489 (PRE). Scale bars: (al —a2, kl—k2) 1 cm; (b—j3) 1 mm.

172 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—This species is similar to Calobota acanthoclada but differs in the green young branches, trifoliolate leaves, larger flowers, short calyx lobes and pods that are constricted between the seeds (in C. acanthoclada the young branches are brown, the leaves simple, the flowers smaller and the calyx lobes longer than in C. spinescens, and the pods not constricted between the seeds).

Distribution and habitat—Calobota spinescens is widely spread throughout the Cape and southern Namibia (Fig. 6.35). According to Mucina et al. (2006) it is an important component of Namaqualand sandveld (SKs 7 Namaqualand Strandveld) and Trans-Escarpment Succlent Karoo (SKt 1 Western Bushmanland Klipveld).

Regional Variation—Two forms can be distinguished based on the pubescens of the ovary and fruit:

FORM A (Typical form)

Leaves with petiole 6-25 mm long; leaflets with terminal 3-12 mm long, 0.5-2.0 mm wide, lateral 2-11 mm long, 0.5-2.0 mm wide. Inflorescence 25-105 mm; with 3 to

10 flowers. Flowers 11-15 mm. Calyx 5.5-6.5 mm long, pubescent or sericeous; tube 3.5-5.0 mm long; lobes 1.5-2.5 mm long. Standard 12.5-15.5 mm long; claw

3-4 mm long; lamina 9.5-12.0 mm long, 7.5-9.5 mm long. Wings 10.5-14.0 mm long; claw 2.5-4.0 mm long; lamina 7.5-10.0 mm long, 2.5-3.5 mm wide, with 4-5 rows of sculpturing, glabrous or rarely pilose. Keel 11-15 mm long; claw 4-5 mm long; lamina 7-10 mm long, 3.5-5.0 mm wide, glabrous or rarely pilose. Pistil pubescent; ovary 7.5-11.5 mm long, 1.0-1.5 mm wide with 10 to 16 ovules; style

3.5-5.5 mm long. Pods sericeous, 20-40 mm long, 3-4 mm wide.

173 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Diagnostic characters—The ovary and pods of this form are densely sericeous.

Distribution and habitat—The typical form of Calobota spinescens is widely distributed in southern Namibia and the Northern Cape Province of South Africa, with some populations in the Western and Eastern Cape. It occurs on sandy soils along roadsides, in riverbeds or on sand dunes. It also grows on sand overlying limestone or granite koppies or calcareous sand at altitudes of between 600 m and

1100 m and is heavily grazed livestock.

FORM B (Prieska form)

Leaves with petiole 6-14 mm long; leaflets with terminal 3-10 mm long, 0.5-1.5 mm wide, lateral 2-10 mm long, 0.5-1.0 mm wide;. Inflorescence 30-70 mm; with 5 to 7 flowers. Flowers 10-16 mm. Calyx 4.5-7.0 mm long, glabrescent or pilose; tube 3-5 mm long; lobes 1.0-2.5 mm long. Standard 10-15 mm long; claw 2-4 mm long; lamina 8.5-11.5 mm long, 7.5-11.0 mm wide. Wings 10.5-13.0 mm long; claw 2.5-

4.0 mm long; lamina 7.5-10.0 mm long, 2-4 mm wide, with 4-6 rows of sculpturing, glabrous. Keel 11-14 mm long; claw 3.5-5.0 mm long; lamina 7.5-10.0 mm long,

3.5-6.0 mm wide, glabrous. Pistil glabrous with pilose hairs along margins; ovary

9.5-11.0 mm long, 1.0-1.5 mm wide with 12-16 ovules; style 4.5-6.0 mm long.

Pods glabrous and pilose along upper suture, 25-35 mm long, 3-4 mm wide.

Diagnostic characters—In the Prieska form the ovary and pods are largely glabrous with only some hairs along the upper suture.

Distribution and habitat—The Prieska form occurs in the central parts of

South Africa (Prieska, Brandvlei, Britstown, Aberdeen and Willowmore) with an outlier population around Klinghardt. It grows on red-brown, well-drained, sandy soil,

174 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA limestone, deep aeolitic sand banks, on calcretic flats or sand on dolerite rock at altitudes of between 250 m and 1100 m. It is heavily grazed by livestock.

Additional specimens examined

—2416 (Maltahahe): Farm Grootplaas MAL 95 (—DD), Giess et al. 5204 (PRE, WIND).

—2418 (Stampriet): between Hofmeyr and Stamprietfontein, 1 ml [1.61 km] from the latter (-

AD), Wilman 383 (BOL, NBG, PRE).

—2516 (Helmeringhausen): Grootfontein MAL 91 (—BB), Volk 12875b (WIND).

—2618 (Keetmanshoop): west of Gellap-Ost 15 km north-west of Keetmanshoop (—CA),

Maggs 111 (WIND).

—2619 (Aroab): 20.5 mls. [32.98 km] south-east of Aroab on Klipdam road (—DC), De Winter

3440 (K, PRE, WIND); 3.6 mls [5.79 km] from Aroab on road to Koes (—DC), De Winter

3458 (K, PRE, WIND).

—2620 (Twee Rivieren): Twee Rivieren in bed of Nossob River (—BC), Leistner 1497 (BOL,

K, PRE); Kalahari-Gemsbok National Park (—BC), Van Rooyen and Bredenkamp 76 (PRE);

40 mls [64.36 km] from Aroab (—CC), Grobbelaar 1862 (K); 50 km south of the Kalahari

Gemsbok National Park (—DA), Koekemoer 248 (JRAU 6 sheets).

—2715 (Bogenfels): Klinghardt Mountain, Diamond Area no. 1 (—BB), Ward and Seely 10223

(WIND).

—2717 (Chamaites): south of Lowen River, at Gawachab (—BB), Pearson 4098 (BM, K);

Wegdraai, Hottentots Plateau (—CB), Helary and Batault 202 (WIND).

—2719 (Tranental): Farm Blinkoog (—CA), Waltu and Waltu 2403 (WIND); Farm Nuwerus (-

CB), Giess et al. 7125 (WIND); Farm Goedemoed (—DD), Strobach et al. 3000 (WIND).

—2722 (Olifantshoek): 4 mls [6.44 km] north of Gamotep Pan (—CC), Leistner 1706 (K).

—2816 (Oranjemund): Sendelingsdrif (—BB), Metelerkamp 59 (BOL).

—2818 (Warmbad): Ramansdrift (—CD), Pearson 4046 (BM).

—2819 (Ariamsvlei): Farm Udabis (—AD), Giess et al. 7107 (WIND); Farm Skroef, near hot spring on bank of Orange River (—DA), Van Hoepen 1942 (BOL, PRE).

175 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

2820 (Kakamas): approximately 70 km from Ariamisvlei on road to Upington (—AB),

Grobbelaar 1926 (K, PRE); Augrabies Waterfall National Park (—CB), Werger 345 (K,

WIND), Zietsman 3489, 3920 (PRE); 31 km west of Kakamas along road to Pofadder and

Onseepkans (—DC), Davidse 6161 (K); 6 mls [9.65 km] south of kakamas (—DC), Leistner and Joynt 2828 (K, PRE); north bank of Orange River near Kakamas (—DC), Lewis 198

(NBG); 20 mls [32.18 km] south-east of Kakamas (—DC), Theron 1943 (BOL, K); between

Kakamas and Kenhardt (—DC), Thorne s.n. (NBG).

2821 (Upington): Karakul Research Station near Upington (—AC), Labuschange s.n.

(NBG).

—2822 (Glen Lyon): 8 mls [12.87 km] west-north-west of west entrance of Padkloof (—DA),

Leistner 1750 (K, PRE).

—2824 (Kimberley): 60 mls [96.54 km] from Upington (—AC), Pole-Evans 2160 (PRE); 21.5 mls [34.59 km] north of Witkop (—BA), Pole-Evans 2132 (PRE).

—2919 (Pofadder): Pofadder (—AB), Conradie 25 (NBG); on N14 highway 25 km from

Pofadder from Springbok (—BB), Boatwright et al. 158 (JRAU).

—2921 (Kenhardt): Kenhardt (—AC), Wabberfall 1114 (K).

—2922 (Prieska): Kwakwas (—AB), Acocks 1759 (PRE); on sandy waste behind

Draghoender hotel (—AC), Acocks and Hafstrom H1343 (K, PRE 2 sheets); Boesmanland,

Farm Mames-Pan 107, 0.5 km north of Kielder farmhouse (—CC), Le Roux and Lloyd 173

(K, PRE); Boesmanland, Farm Doonies-Pan 106, 13.3 km north north-west of Copperton (-

CD), Le Roux and Lloyd 138 (PRE); Prieska (—DA), Bryant J263 (K); North bank of Orange

River (—DA), Bryant 5404 (BOL).

2923 (Douglas): Lanyon-Vale (—AC), Acocks 1982 (BOL, K).

3020 (Brandvlei): Boesmanland, Farm Bastiaans Kolk 331, on stabilized sand dunes in

Groot Vloer (—BA), Le Roux and Lloyd 111 (PRE).

—3023 (Britstown): at Vissershoek turn-off 39 km north of Britstown (—BC), Van Wyk 2849

(JRAU 2 sheets); 36.3 km from Britstown to Vissershoek turn-off, at Farm Fonteintjies 8 km

176 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA from house (—BC), Van Wyk 3081 (JRAU); 10 km south on Deelfontein road, Farm

Witfontein (—DA), Le Roux 236 (PRE).

—3120 (Williston): Great Fish River and Zout River, Beaufort (—AD), Zeyher 397 (BM, K,

NBG); Williston Station (—BD), Smith 2443 (K); Rietpoort (—DB), Shearing 81 (K).

—3122 (Loxton): Pampoenpoort (—BA), Henrici 4750 (PRE).

—3219 (Wuppertal): Grasberg South, Tanqua National Park (—BC), Bayer 6244 (PRE);

Sandkop (—BC), Henrici 3333 (PRE).

—3221 (Merweville): rocks at Driekoppe (—DC), DrOge s.n. "II, f' (P).

—3224 (Graaff-Reinet): 40 mls [64.36 km] from Aberdeen (—AC), Hutchinson 3135 (BM, K 2 sheets); south-west of Aberdeen (—AC), Maguire 742 (NBG).

—3321 (Ladismith): near Grootfontein, 30 mls [48.27 km] north east of Laingsburg (—AB),

Wilman 1048 (BOL).

—3322 (Oudtshoorn): Prince Albert (—AA), Bolus 11774 (K, PRE), Thoday and Doff 66

(NBG).

—3323 (Willowmore): Willowmore, near station Vondeling on banks of Olifantsriver (—AC),

Vlok 1932 (JRAU, PRE).

Precise locality unknown: without locality, Acocks 1701 (PRE); Griqualand West, Burchell

1965 (K); between Dweka River and Zwartbulletjie, Drege s.n. (S); Bushmanland, Pearson

3633 (K).

177 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA 12 14 16 seammErams.18 20 22 24 26 28 30 mum32 34 12-fiva 11111111M11111 'Raw 1 8 11111111111 MOM 11111 NUM 111111111111 lisi ormal 111111111 EMI • MI MN& e 2..1 11111 Miiiiiallin 112 1111111111wil , 111111111111 • wrimpumnimm e ossori• 1 simbi m WM IVEMYMMANN 1 snimminixo no■MONMOMF OMMINWIEW 4 -111 MUM , mosistoriV3 • MI ME WAIMMIN28 111111111116 713 lirg Pall III NM EMI Rim mow " owMIMI Arc Ili rpm !In 1111111101111 ME- PIM EN 11111 IN 4 wannemon ma PM NMI < ' ilIiiirdigill iralailling 111111 ire!: 14 ' 10 12 14 16 18 20 22 24 26 28 30 32 34 36

FIG. 6.35 Known geographical distribution of Calobota spinescens.

16. CALOBOTA THUNBERGII Boatwr. and B.-E.van Wyk, nom. nov. pro Lebeckia

sericea Thunb. non Calobota sericea (Ait.) Boatwr. and B.-E.van Wyk. (S. Afr.

J. Bot.: submitted. 2008). Lebeckia sericea Thunb., Nov. Gen.: 143. 1800;

Prodr. Pl. Cap.: 122. 1800; Willd., Sp. Pl.: 948. 1800; Thunb., Fl. Cap.: 562.

1823; DC., Prodr. 2: 137. 1825; Benth. in Hook. Lond. J. Bot. 3: 360. 1844;

Harv. in Harv. and Sond., Fl. Cap. 2: 88. 1862; Bond and Goldblatt, J. S. Afr.

Bot. 13: 290. 1984; Goldblatt and Manning, Cape Pl.: 494. 2000.—TYPE:

SOUTH AFRICA, 'e Cap. b. Spei' [grid unknown], Thunberg s.n. sub THUNB-

UPS 16423 (lectotype: UPS!, designated here). [Note: The epithet `sericea'

was applied to Lebeckia multiflora E.Mey. (now Calobota sericea) by Aiton

178 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

(1789), the basionym of which is Spartium sericeum Ait., and is thus not

available for the current taxon. We therefore propose a new name in memory

of Thunberg who first described this species]. [Note: The specimen listed is

the only specimen in Thunberg's herbarium and is here chosen as lectotype].

Lebeckia flexuosa E.Mey., Comm. Pl. Afr. Austr. 1:34. Feb. 1836; Walp in Linnaea

13: 478. 1839.—TYPE: SOUTH AFRICA, "In rupestribus prope Kwekrivier"

[grid unknown], altit. 700 ped., Drege s.n. "III, C" (lectotype: P!, designated

here; isolectotype: K!, P!). [Note:The specimen bears the diagnostic features

described by Meyer and also the type locality.]

Lebeckia decipiens E.Mey., Comm. Pl. Afr. Austr. 1:35. Feb. 1836.—TYPE: SOUTH

AFRICA, Karroo prope Mierenkasteel [grid unknown], Drege s.n. "III, C"

(lectotype: P!, designated here; isolectotype: BM!, K!, NBG!, P!, PRE!, S!).

[Note: The P specimen bears the type locality and was annotated by Meyer

himself.]

Sericeous, multi-stemmed unarmed shrub up to 2 m in height. Branches green; young branches sericeous; older branches sericeous or glabrous with brown bark. Leaves digitately trifoliolate; petiole longer or shorter than leaflets, 13-55 mm long; leaflets elliptic to narrowly oblanceolate, alternate, sericeous, subsessile, terminal leaflet (15–) 17-50 mm long, (1–) 2-8 mm wide, lateral leaflets 10-45 mm long, (1–) 2-5 mm wide, apex acute, base angustate. Inflorescence 70-220 mm long, with 10 to 28 flowers; pedicel 2-4 mm long; bract 2-7 mm long, linear, pubescent; bracteoles 1-5 mm long, linear, pubescent. Flowers 13-24 mm long, pale or bright yellow. Calyx 6.0-9.5 mm long, sericeous; tube 4.0-6.5 mm long; lobes 1-4 mm long, subulate. Standard (11.5–) 13.0-19.0 mm long; claw linear, 3-5

179 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

mm long; lamina widely ovate, 8.0-14.5 mm long, 9.5-17.0 mm wide, pilose along

dorsal midrib; apex obtuse. Wings 12.0-17.5 mm long; claw 3.0-5.5 mm long;

lamina oblong, as long as or shorter than keel, 8-12 mm long, 4.0-6.5 mm wide,

glabrous, with (5–) 8-14 rows of sculpturing. Keel 12.5-19.5 mm long, claw 3.5-7.0

mm long, lamina boat-shaped, 8.5-13.0 mm long, 4.5-6.5 mm wide, pilose on

terminal parts. Pistil subsessile to shortly stipitate, pubescent; ovary linear, 8.5-15.0

mm long, 0.8-1.5 mm wide with 19 to 28 ovules; style longer than ovary, 3.5-8.5

mm long. Pods linear, semi-terete, subsessile to shortly stipitate, 30-50 mm long, 3-

4 mm wide, 7 to 14-seeded, pubescent, dehiscent. Seeds oblong-reniform, 2.3-3.5

mm long, 1.7-2.5 mm wide, mature seeds light brown to light pink, surface smooth

(Fig. 6.36). Flowering time: C. thunbergii flowers mainly from May to October. One

specimen collected in March was in flower.

Diagnostic characters—Similar to Calobota cytisoides but differs in the

acute apices of the leaflets, short pedicels and smaller flowers, linear bracts and

bracteoles, sericeous calyx and sericeous pistil and pods (in C. cytisoides the apices of the leaflets are mucronulate, the pedicels longer and the flowers larger. The

bracts and bracteoles are often ovate, the calyx glabrous and the pistil and pods glabrous).

Distribution and habitat—Calobota thunbergii is widely distributed from

Wupperthal south as far as the Oranjemund area (Fig. 6.37). It grows in well-drained sandy loamy soil, granite, rocky sand, sandy dunes, in riverbeds or along disturbed roadsides between 300 m and 1200 m above sea-level. This species, according to

Mucina et al. (2006) and Jiirgens (2006), is an important component of Richtersveld

(SKr 14 Southern Richtersveld Inselberg Shrubland), Namaqualand Hardeveld (SKn

1 Namaqualand Klipkoppe Shrubland, SKn 2 Namaqualand Shale Shrubland, SKn 6

180 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Kamiesberg Mountains Shrubland), Namaqualand Sandveld (SKs 9 Namaqualand

Inland Duneveld) and Southern Namib Desert (Dn 1 Alexander Bay Coastal

Duneveld).

jl j2

AeMERS5E1M kl

bl b k2

FIG. 6.36 Morphology of Calobota thunbergii: (a1 —a2) leaves in abaxial view; (b1) bract; (b2) bracteoles; (c) flower in lateral view; (d) standard petal; (e) wing petal; (f) keel petal; (g) outer surface of the calyx (upper lobes to the left); (h) androecium; (i) pistil; 01) long, basifixed anther; (j2) intermediate carinal anther; (j3) short, dorsifixed anther; (kl) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al , bl—j3) Boatwright et al. 151 (JRAU); (a2, kl—k2) Van Wyk

3119 (JRAU). Scale bars: (al—a2, kl—k2) 1 cm; (b—j3) 1 mm.

181 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Additional specimens examined

2816 (Oranjemund): sand drifts on Witbank (—DC), Pillans 5542 (BOL); sandy coast belt between Port Nolloth and Holgat River (—DD), Pillans 5627 (BOL, K).

2817 (Vioolsdrif): Kodaspiek (—AA), Oliver et al. 451 (K, NBG); Vandersterrberg north-east of Khubus north end south-east of the top of Helskloof and south-east of Paradysberg western slopes to summit (—AC), Oliver et al. 151 (NBG, K, PRE); Vioolsdrif, Ridge above

Koeskop, summit of Vandersterberg, Richterveld National Park (—AC), Williamson and

Williamson 5764 (NBG); Cornellsberg (—CA), Bean 1258 (BOL), Bean 1328 (S), Van

Jaarsveld 11977 (PRE), Viviers 2046 (NBG); Jenkins Hill, 20 km from Eksteenfontein on road to Mount Stewart (—CB), Germishuizen 4679a (PRE 2 sheets); Spektakel Hill (—DA),

Johnson 216 (NBG).

—2916 (Port Nolloth): sandy plain 2 mls [3.22 km] east of Port Nolloth (—BD), Marloth 12660

(NBG, PRE).

—2917 (Springbok): along roadside on road from Port Nolloth to Steinkopf (—AC), Boatwright et al. 151 (JRAU); Klipfontein (—BA), Bolus 429 (BM, BOL, K, NBG, PRE), Bolus 6549 (K),

Hutchinson 904 (BM, BOL, K, PRE); Steinkopf (—BA), Herre s.n. sub STE 11889 (NBG),

Meyer s.n. sub STE9068 (NBG); Anenous Pass (—BA), Germishuizen 4839 (PRE, WIND),

Schutte 591 (JRAU 2 sheets), Van Wyk 6266 (PRE); Spitskop, 6 mls [9.65 km] south of

Steinkopf (—BD), Lewis 5487 (NBG 2 sheets); Klipfontein Koppie (—CA), Compton 5486

(BOL, NBG); Stinkfontein Mountains south-west of Van Zylsrus (—CA), Oliver et al. 625

(NBG, PRE); Spektakel (—DA), Compton 11524 (NBG); Spektakel Pass (—DA), Eliovson 37

(NBG); 25 km from Springbok towards Spektakelberg Pass (—DA), Glen 1463 (BOL, PRE);

Spektakel Pass, 26 km west of Springbok (—DA), Goldblatt 3651 (PRE); top of

Spektakelberg Pass (—DA), Grobbelaar 2584 (PRE); Farm Ezelfontein 214, 2 km north of

Naries farmstead (—DA), Le Roux 3233 (NBG); 21.7 km fro Springbok to Spektakel (—DA),

Stirton 6019 (K); 24 km from Springbok to Kleinsee (—DA), Stirton 10149 (NBG); 9 mls

[14.81 km] south-east by east of Springbok (—DB), Acocks 19237 (K, PRE); 8.5 mls [13.68

182 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

km] west by south of Springbok (—DB), Acocks 19550 (BOL, K, PRE); near O'kiep (—DB),

Barker 6244 (NBG 3 sheets); 1ml [1.61 km] south of Springbok (—DB), Barker 8389 (NBG 2

sheets); 2 km east of Springbok (—DB), Bellamy 12 (PRE); Springbok (—DB), Bester and

Pretorius 8 (JRAU), Crosby 20 (PRE), Salter 895 (BM, K), Small et al. 82 (PRE), Van Blerk

7 (PRE), Van Hille 8 (BOL), Venter 8820 (PRE); Nababeep, road to O'kiep (—DB), Burrows

2950 (GRA); ±8 km north-east of Nababeep on gravel road to junction with tar road (—DB),

Botha 3170 (PRE); 2 mls [3.22 km] north-east of Springbok (—DB), Compton 22012 (NBG);

Concordia, near Springbok (—DB), Clarke 687 (PRE), Thorns s.n. (NBG); just outside

nababeep on road to O'kiep (—DB), Germishuizen 4515 (PRE); Nababeep (—DB), Gess and

Gess 89/90/67 (GRA); Hester Malan Wild Flower Reserve north of Oxaliskoppie (—DB), Le

Roux 918, 1231 (PRE); Nababeep, near golf course (—DB), Le Roux 2674 (BOL); 2 mls

[3.22 km] north of Springbok (—DB), Lewis 3392 (NBG), Schelpe 240 (BM, BOL); 4 mls [6.44

km] west of Springbok (—DB), Macguire 369 (BOL, NBG 2 sheets); Springbok, weather

station hill on west of town (—DB), Manning 2774 (NBG); Okiep (—DB), Dammel s.n. (K),

Marloth 6712 (NBG 3 sheets, PRE), Morris 5623 (BM, BOL, NBG); 15 mls [24.14 km] north

of Springbok (—DB), Schlieben 9059 (BM, K, PRE); Springbok T'Karoep, on road between

Nababeep and Okiep (—DB), Scholtz 20 (NBG, PRE); Hester Malan Nature Reserve (—DB),

Stirton 6004 (PRE), Struck 113 (NBG), Van Wyk 5728 (PRE); 5 mls [8.05 km] north of

O'kiep (—DB), Van der Schijff and Schweickerd 5744 (K, PRE); 2 mls [3.22 km] north of

Springbok (—DB), Van Niekerk 240 (PRE, 2 sheets); 11 km from Springbok to Pofadder (—

DB), Van Wyk 3078 (JRAU 2 sheets); Modderfontein (—DB), Whitehead s.n. (S); 3 km east

of Springbok (—DB), Zietsman and Zietsman 667 (PRE); Messelpad between Springbok and

Wallekraal (—DC) Compton 20664 (NBG 2 sheets); near Komaggas (—DC), Eliovson 69 (J);

north slopes of Rooiberg above Buffels River on eastern approach to the Meselpad (—DC),

Hilton-Taylor 2140 (NBG, PRE); ca. 11 km from Springbok on sand road to Hondeklipbaai

(—DC), Joffe 96 (NBG, PRE); on road to Hondeklipbaai, 18 mls [28.96 km] from turn-off from

national road to Springbok (—DC), Van der Merwe 215 (NBG); 29 mls [46.66 km] from

Springbok on road to Caries (—DD), Grobbelaar 1119 (PRE); near Mesklip (—DD), Leighton

183 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

1156 (BOL); on roadside of N7 highway 13 km south of Springbok, opposite 33.4 marker (-

DD), Powrie 652 (NBG 2 sheets, PRE); 30 mls [48.27 km] south of Springbok (—DD),

Rynolds 5440 (K); 20 mls [32.18 km] south-west of Springbok (—DD), Verdoorn and Dyer

1791 (BOL, K 2 sheets).

—2918 (Gamoep): Zilverfontein (—CC), Drege s.n. (BM, K, P, S).

—3017 (Hondeklipbaai): Diknek (—AD), Van Breda 4073 (PRE); Farm Doornfontein 464,

Portion 1, Steenkamp Kraal (—BA), Le Roux 4555 (NBG); on road to Leliefontein from

Kamieskroon (—BB), Boatwright et al. 219 (JRAU); Grootvlei, hillsides, Kamieskroon (—BB),

Compton 6839 (NBG); Skilpad Wild Flower Reserve, 100m from house (—BB), Cruz 85

(NBG); Studer Pass (—BB), Evrard 8960 (PRE); Kamieskroon (—BB), Hutchinson 841 (BM, K

2 sheets, PRE), Salter 863 (BM), Salter 865 (K), Thorne s.n. (NBG), Van Rooyen 2416

(PRE); portion of Wolwepoort 459, Skilpad Farm, north-west of Kamieskroon (—BB),

Johannesburg Botanical Garden 3877 (PRE); between Kamieskroon and Soebatfontein (-

BB), Meyer 7331 (PRE); ±4 km from Kamieskroon (—BB), Meyer 7349 (PRE); Kamiesberg

Pass (—BB), Schutte 593 (JRAU), Stirton 5993 (PRE), Strid and Strid 37780 (NBG), Van

Wyk 2540, 3094 (JRAU); Farm Olienfontein (—BB), Zietsman and Zietsman 1017 (PRE); 2.3

km from start of Kamiesberg Pass (—BB), Van Wyk 2352 (JRAU 2 sheets); 5 km from

Wallekraal to Garies (—BC), Stirton 6054d (K); Garies (—BD), Compton 17180 (NBG), Kruger

K5 (K, NBG, PRE), Lynes 1944 (BM); Brakdam (—BD), Johnson 200 (NBG 2 sheets); 5 mls

[8.05 km] north of Garies, top of pass (—BD), Lewis 1308 (NBG); between Nuwerus and

Karkams (—BD), Pearson 6500 (K); Brakdam (—BD), Schlechter 11150 (BM, BOL, K, PRE);

20 km from Kamieskroon to Garies (—BD), Stirton 5982 (K); 16 km north of Garies (—BD),

Wisura 2924 (NBG); between Garies and Kamieskroon (—DB), Jordaan 1225 (NBG), Van

der Schijff 6994 (PRE); 4 km north of Garies on road to Springbok (—DB), Le Roux 2630

(BOL); 6 mls [9.65 km] north of Garies (—DB), Leighton 1127 (BOL); near Garies (—DB),

Levyns 6984 (BOL), Markotter s.n. (NBG), Stirton 5978 (K, PRE); 2.1 km from Garies to

Bitterfontein (—DB), Marsch 343 (NBG, PRE); 3 km north of Garies on the main road to

Springbok, Garieshoogte (—DB), McDonald 1329 (NBG, PRE, UPS); 5 km from Kotzerus 184 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

turn-off from Wallekraal to Caries (—DB) Stirton 6055 (K); 6mIs [9.65 km] from Garies to

Kamiesberg (—DB), Thompson 415 (NBC, PRE); 23 km north of Caries beside N7 (—DB),

Van Wyk 2339 (JRAU); 24 km from Caries on main road to Springbok (—DB), Van Wyk 2424

(JRAU); between Caries and Springbok (—DB), Werdermann and Oberdieck 598 (K);

Kotzerus (—DD), Stirton 6058 (K); road between Kotzerus and Bitterfontein (—DD), Van Wyk

3115 (JRAU).

—3018 (Kamiesberg): Farm Pedroskloof, 18 km east of Kamieskroon (—AA), Hilton-Taylor

2122 (NBG); north of Leliefontein (—AB), Liede and Here s.n. (JRAU); near Eselsfontein (-

AC), Van der Schijff and Schweikerd 5780 (PRE); on farm near Caries (—AC), Van Der Walt

159 (NBG); slopes of hills near Caries (—CA), Thorne s.n. (NBC); on N7 towards Caries 30

km from town (—CC), Boatwright et al. 140 (JRAU); 10 km from Caries on N7 (—CC),

Boatwright et al. 141 (JRAU); along N7 to Springbok 20 km from Caries (—CC), Boatwright

et al. 183 (JRAU); 35 mls [56.32 km] south of Garies (—CC), Thompson 1014 (NBC, PRE);

along main road 5 mls [8.05 km] from Caries to Springbok (—CC), Van Breda 1283 (K,

PRE); Kliprand (—DA), Hall 891 (BOL, NBC), Van Breda 1607 (K); 5.6 mls [9.01 km] from

Bitterfontein to Pofadder (—DB), Marsch 444 (NBC).

—3117 (Lepelfontein): Lepelfontein (—BB), Kolbe s.n. sub BOL 14269 (BOL).

—3118 (Vanrhynsdorp): Nuwerus, Elandsfontein (—AB), Barnard 307 (NBC); Bitterfontein (-

AB), Bond 1093 (NBC); Farm Quaggas Kop 125. 6 km west of Nuwerus (—AB), Le Roux

2278 (NBC); slopes of Spitsberg south-east of Nuwerus (—AB), Oliver 5930 (NBC, PRE);

Karree-Bergen (—AB), Schlechter s.ri. sub TRV 1057 (PRE), Schlechter 8182 (BM, BOL, K,

PRE); 3 mls [4.83 km] north of Bitterfontein (—AB), Schelpe 131 (BM, BOL, K); Nuwerus (-

AB), Steyn 472 (NBC); Vanrhynsdorp (—DA), Van der Byl s.n. sub STE 17419 (NBC); top of

Klein Kobee Pass (—DB), Manning s.n. (K, NBG 2 sheets); Klapmuts (—DD), Schlechter 375

(BOL).

—3119 (Calvinia): Bottom of Vanrhyns Pass (—AC), Marsch et al. 312 (NBG, PRE);

Nieuwoudtville (—AC), Pearson 3468 (K); Vanrhyns Pass before Nieuwoudtville (—AC),

Schutte 285 (JRAU), Van Wyk 3119 (JRAU); Vanrhyns Pass between Nieuwoudtville and

185 CHAPTER 6: TAXONOMY OF THE GENUS CALOBOTA

Vanrhynsdorp (—AC), Van der Schijff 7183 (PRE), Van Wyk 2584 (JRAU 2 sheets, NBG,

PRE).

—3218 (Clanwilliam): Nortier Experimental Station near Lamberts Bay (—AB), Van Breda

4435 (PRE); near Nuwerus (—DC), Acocks 14188 (K, PRE); few miles north of Nuwerus (-

DC), Wilman 702 (BOL).

—3219 (Wuppetal): Citrusdal (—CA), Thorns s.n. (NBG); Doorn River (—DA), Herre s.n. sub

STE 11890 (NBG); Doom River bridge (—DA), Pillans 6305 (BOL, K).

Precise locality unknown: Platberg, DrOge s.n. "Ill, A, b" (P); without locality, Leipoldt 739

(NBG), Marloth 11143 (NBG), Zeyher 400 (K 2 sheets, S); north of Darters Grave, Maguire

296 (NBG); Namaqualand, Scully 7 (BM).

FIG. 6.37 Known geographical distribution of Calobota thunbergii.

186 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

7.1 INTRODUCTION

Wiborgiella is here proposed as a new generic concept to accommodate nine species that are endemic to the Cape region of South Africa. The description of the genus followed a phylogenetic study of the tribe Crotalarieae (Chapter 2) which showed that the genus Lebeckia, of which these species formed a part, is polyphyletic

(Boatwright et al. 2008a). These results together with morphological and anatomical data indicated that Lebeckia s.l. should be divided into three genera: Lebeckia s.s.,

Calobota and Wiborgiella. Wiborgiella comprises all the species that were previously included in Lebeckia section Viborgioides, along with the anomalous Lebeckia inflata,

L.- mucronata and Wiborgia humilis. Although the monophyly of Wiborgiella did not receive support in the phylogenetic analyses, a combination of salient morphological and anatomical characters support the recognition of this group at the generic level.

Lebeckia section Viborgioides was studied by Dahlgren (1975) in his revision of

Wiborgia due to the superficial similarity between the former and Wiborgia species. He suggested that the taxa might be con-generic, but from the data in Chapter 2

(Boatwright et al. 2008a) it is clear that Wiborgia is strongly supported as a separate genus. Wiborgiella can be distinguished from Calobota, Lebeckia s.s. and Wiborgia by a unique combination of characters: brown or grey bark on young branches (the short- lived W. inflata and 'W. vlokii' have green stems), laminar trifoliolate leaves, glabrous petals, 4+6 anther arrangement and oblong, wingless, much inflated fruit.

187 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

This chapter is aimed at describing the genus Wiborgiella and presenting a taxonomic revision of its species which includes a key, complete nomenclature, typification, formal descriptions, the known geographical distributions and illustrations of all the species.

FIG. 7.1. Habit of Wiborgiella species. (a, d, e) W. sessilifolia. (b) W. inflata. (c) W. humilis. (f, g, h) W. leipoldtiana.

188 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

7.2 DISCUSSION OF CHARACTERS

7.2.1 Vegetative morphology and anatomy— HABIT AND BRANCHES —The species of Wiborgiella are either long-lived perennials or short-lived fire-weeds. In the former the branches are brown or grey and covered with longitudinally splitting bark whereas in the short-lived species the branches are green and the plants do not reach heights of more than 0.3 m. The perennial species are rigid shrubs or shrublets that are between 0.5 m and 1.5 m in height (Fig. 7.1).

The stem anatomy of Wiborgiella species revealed the presence of several chlorenchyma layers in the species with green stems that are absent in those species with brown, bark-covered stems. The genera Calobota and Lebeckia differ from most species of Wiborgiella in that they also have green stems, i.e. with chlorenchyma, but

Wiborgia species have a similar habit and stem structure to that of Wiborgiella (Fig.

7.2).

Leaves of Wiborgiella species are invariably laminar, trifoliolate and petiolate

(sessile on older branches of W. sessilifolia). The petioles of the shrubby species have a swollen base or tubercle and are persistent even after the leaflets have been shed.

The branches in most perennial species become hard and pungent after flowering and have a thorn-like appearance similar to most Wiborgia species (Dahlgren 1975). The persistent petioles along with the somewhat spinescent branches give these plants a very rigid appearance. Transverse sections through the leaves of all Wiborgiella species revealed that the leaves are dorsiventral (clearly differentiated into palisade parenchyma adaxially and spongy parenchyma abaxially) and that they have mucilage

189 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA cells in the epidermis (Fig. 7.2). This is a very reliable distinguishing character to separate Wiborgiella from Lebeckia and Calobota. Lebeckia has acicular leaves with only palisade parenchyma and Calobota has laminar, but isobilateral leaflets with no differentiation into palisade or spongy parenchyma.

7.2.2 Reproductive morphology and anatomy—INFLORESCENCE—Polhill

(1976) mentions that the inflorescence tends to be an unreliable character in papilionoid legumes. In Wiborgiella, as in most Crotalarieae, the inflorescences are terminal racemes, except sometimes in W. bowieana where the flowers may be solitary. The inflorescences are either short as in W. dahlgrenii and W. leipoldtiana or long as in W. inflata and W. mucronata. The number of flowers per raceme also differs between species.

FLOWERS—The flowers are relatively large (more than 10 mm long) in W. bowieana, W. leipoldtiana, W. mucronata and W. sessilifolia and 10 mm or less in W. dahlgrenii, W. fasciculata, W. humilis, W. inflata and W. vlokii. The pedicel is short and less than 1.5 mm in W. dahlgrenii, W. bowieana and W. fasciculata but may be longer in the other species. The flowers in the genus are yellow, although the label information from the only available flowering specimen of W. dahlgrenii suggests that its flowers may be pale pink. The older petals in W. inflata turn purple and in W. sessilifolia and W. vlokii they turn orange. Gess and Gess (1994) suggest that this change in colour follows pollination and is a very effective pollination strategy as it discourages non- productive flower visits thereby increasing pollination efficiency. The petals are generally glabrous in Wiborgiella (although a few hairs may very rarely be present on the back of the standard petal).

190 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.2 Transverse sections through the stems and leaves of Wiborgiella species. (a) Portion of the stem of W. humilis; (b) Portion of the stem of W. sessilifolia; (c) Portion of the stem of W. inflata; (d) Petiole of W. inflata; (e) Petiole of W. leipoldtiana. (f) Petiole of W. mucronata; (g)

Portion of the leaf of W. inflata; (h) Portion of the leaf of W. sessilifolia; Voucher specimens: (a)

Boatwright et aL 129 (JRAU); (b, h) Van Wyk 2120 (JRAU); (c, d, g) Johns 162 (JRAU); (e)

Boatwright et al. 123 (JRAU); (f) Vlok 1726 (JRAU). Scale bars: (a—c) 0.1 mm; (d—h) 0.2 mm.

191 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

The bracts and bracteoles are linear to slightly lanceolate or ovate. Those of

Wiborgiella bowieana, W. mucronata and W. sessilifolia are relatively long (generally more than 3 mm). The bracts and bracteoles may be either pilose or sericeous on the adaxial surface and are caducous.

The calyces of Wiborgiella species are subequally lobed with the upper sinus often deeper than the lateral or lower sinuses and the carinal lobe invariably narrower than the others. The outer surface may be either pubescent or glabrous, while the inner surface of the lobe tips is always minutely pubescent. This feature is present in many of the genera of the Crotalarieae but known to be absent in Aspalathus. The calyces are relatively large in Wiborgiella bowieana and W. sessilifolia (9.0-10.5 mm long) with very long lobes that are between 3.0 and 5.5 mm long and the calyx tubes that are longer than in the other species (always exceeding 4.5 mm).

The standard petal is ovate to widely ovate in most species, except in

Wiborgiella dahlgrenii where it is obovate to cordate. The apex is shallowly emarginate in the former species, acute in W. humilis and obtuse in all the other species. The standard petal is erect in W. dahlgrenii, W. bowieana, W. inflata, W. mucronata, W. sessilifolia and W. vlokii, while it is reflexed in W. fasciculata, W. humilis and W. leipoldtiana. Reflexed standard petals are also present in Wiborgia (Dahlgren 1975).

The claws are relatively long in W. bowieana, W. fasciculata, W. inflata, W. leipoldtiana and W. vlokii.

The wing petals of Wiborgiella species are oblong in W. dahlgrenii and W. bowieana, slightly down-curved in the former and longer than the keel petals in both species. In W. fasciculata, W. humilis, W. leipoldtiana and W. mucronata they are

192 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA oblong to slightly elliptic and generally shorter than the keel petals. Wiborgiella inflata and W vlokii have wing petals that are oblong and longer than the keel petals, while in

W. sessilifolia they are obovate, longer than the keel petals and broaden towards the apex. All species have sculpturing on the upper basal parts of their wing petals. In W. fasciculata, W. humilis and W leipoldtiana the wing petals are broadest near the middle, while in the other species they are broadest near the apex or base. The claws are relatively long in W. bowieana, W. fasciculata, W. inflata, W. leipoldtiana and W. vlokii.

The keel petals are boat-shaped in some species with the apex slightly upcurved, while they are lunate in the others generally with acute apices. Pockets are present in all species, except W. inflata and W. vlokii. The apices are acute in W.

dahlgrenii, W. bowieana, W. inflata, and W. vlokii, sub - acute in W. fasciculata and W. leipoltiana and obtuse in W. humilis, W. mucronata and W. sessilifolia. The claws are relatively long in W. bowieana, W. fasciculata, W. inflata, W. leipoldtiana and W. vlokii.

Anther arrangement is an important character that distinguishes Wiborgiella from

Lebeckia, Calobota and Wiborgia (Chapter 5; Boatwright et al. submitted). The anthers are dimorphic with alternatingly basifixed and dorsifixed anthers as in practically all other genera. However, the size and shape of the carinal anther is critical. Species of

Wiborgiella have a 4+6 arrangement where the carinal anther resembles the short dorsifixed anthers, as opposed to the 4+1+5 arrangement found in Calobota and

Wiborgia (the carinal anther is intermediate in size and shape between the long basifixed and short dorsifixed anthers) and the 5+5 arrangement of Lebeckia (carinal anther resembles the long basifixed anthers).

193 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

The pistil is subsessile to stipitate (long-stipitate in W. humilis). The ovary may be either linear (W. fasciculata, W. leipoldtiana, W. mucronata and W. sessilifolia), ovate (W. dahlgrenii and W. bowieana) or elliptic (W. humilis, W. inflata and W. vlokii) with from four up to 21 ovules. The style is always glabrous and shorter than the ovary, except in W. dahlgrenii and W. bowieana where the style exceeds the ovary in length.

The pods of species of Wiborgiella are generally inflated or turgid, except in W. mucronata and W. vlokii. In these two species the pods are linear and laterally compressed. The anatomy of the fruit wall showed that the pods of all species have a thin mesocarp with several layers of schlerenchyma adjacent to the endocarp. Mucilage cells are present in the exocarp of some species. The endocarp is composed of one or rarely two rows of periclinally elongated or round cells (Fig. 7.3). In some species of

Calobota and Lebeckia the endocarp may be composed of several layers of loosely arranged cells that have a spongy macroscopic appearance. This feature is never present in Wiborgiella. The pods are few-seeded (one- or two-seeded) in W. dahlgrenii and W. humilis and many-seeded in the other species.

Seeds of most of the species were not available for study. The seeds of the two short-lived fire-weeds W. inflata and W. vlokii are black mottled with white and have a rugose surface. These are similar to those of two Lebeckia species that are also short- lived fire-weeds, namely L. uniflora and L. wrightii. In these two species the seeds are also black mottled with white and have rugose surfaces (Le Roux and Van Wyk 2009).

As these short-lived species occur in recently burnt vegetation, the black colour could serve to camouflage the seeds as there would be no undergrowth to cover them. The seeds of W. leipoldtiana are light pink with a smooth surface, while those of W. humilis

194 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA are light brown or light orange and mottled with brown with a rugose surface. It would be interesting to obtain seeds of all species to ascertain whether a smooth surface is more common than a rugose surface and to assess the colour variation in the genus.

FIG 7.3 Transverse sections through the fruit walls of Wiborgiella species. (a) W. sessilifolia; (b)

W. inflata; (c) W. vlokii; (d) W. humilis (young fruit); (e) W. bowieana. Voucher specimens: (a)

Taylor 4329 (PRE); (b) Johns 162 (JRAU); (c) Vlok 2045 (PRE); (d) Boatwright et al. 212

(JRAU); (e) Streicher s.n. sub Schulte 831 (JRAU). Scale bars: (a—c, e) 0.2 mm; (d) 0.1 mm.

195 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

9.2.3 Phylogenetic relationships—MORPHOLOGICAL DATA—Infrageneric relationships within Wiborgiella were assessed using 13 morphological characters scored for 14 taxa of the "Cape group" of the Crotalarieae (Table 7.1). Calobota was used as outgroup and Wiborgia and Aspalathus included as ingroups in the analyses based on the close relationship between these genera and Wiborgiella indicated by the molecular study in Chapter 3 (see also Boatwright et al. 2008a). All 13 included characters were parsimony informative and the analysis resulted in three equally parsimonious trees with a length of 20 steps, a CI of 0.65 and a RI of 0.82 (Fig. 7.4).

Dahlgren (1975) discussed the close similarity of the shrubby species of Wiborgiella to species of Wiborgia. This and the high levels of convergence within the crotalarioid genera (Dahlgren 1970a) complicated the morphological analysis due to the lack of apomorphic states for genera and the extreme overlap of morphological characters. As a result a combination of characters is needed to define genera.

The monophyly of Wiborgiella is supported by the anther arrangement of 4+6, but the group lacks bootstrap support. Wiborgiella bowieana and W. sessilifolia are supported as sister taxa (61 BP) by the long calyx tube and long wing petals (shared with W. dahlgrenfi) found in both species. The shrubby species of Wiborgiella form a

Glade which is supported by the presence of rigid branches and tuberculate petioles in these species. These characters are shared by Wiborgia tetraptera and other species of Wiborgia. The short-lived species of the genus, Wiborgiella inflata and W. vlokii, are sister based on their life history and long wing petals (70 BP). The position of

Wiborgiella mucronata is unresolved due to the fact that it shares some characters with the shrubby species of the genus (e.g. brown stems), and others with the short-lived

196 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA species (e.g. petioles that are neither tuberculate nor persistent). The close relationship between Wiborgiella fasciculata, W. humilis and W. leipoldtiana is supported (65 BP) by the wing petals that are broadest near the middle and the reflexed standard petals that are, however, also found in Wiborgia. Aspalathus and Wiborgia are sister genera based on the few ovules in the ovary (also in Wiborgiella humilis). The monophyly of

Aspalathus is supported by the sessile leaves and glabrous inner surface of the calyx lobes (84 BP) and that of Wiborgia is supported by the few-seeded samaras and wings on the fruit (70 BP).

MOLECULAR DATA—Wiborgiella is closely related to Wiborgia and Aspalathus based on combined rbcL/ITS/morphological data presented in Chapter 3 (Boatwright et al. 2008a). Although the monophyly of the genus lacks support in these analyses due to the unresolved position of Wiborgiella mucronata and low sequence divergence within the "Cape group" (see Fig. 3.4), it is supported by a combination of morphological and anatomical characters that is unique to this group and not found in the other genera of the Crotalarieae. The close relationship between the shrubby species of Wiborgiella is strongly supported by these data.

197 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

TABLE 7.1 Characters and character polarisations in Aspalathus, Calobota, Wiborgia and

Wiborgiella. Characters and character states are explained at the end of the table.

1 2 3 4 5 6 7 8 9 10 11 12 13 Wiborgiella bowieana 0 1 1 0 1 0 0 1 0 1 0 0 0 W. dahlgrenii 0 1 1 0 0 0 0 1 0 1 0 0 0 W. fasciculata 0 1 1 0 0 0 1 0 1 1 0 0 0 W. humilis 0 1 1 0 0 0 1 0 1 1 1 0 0 W. inflata 1 0 0 0 0 0 0 1 0 1 0 0 0 W. leipoldtiana 0 1 1 0 0 0 1 0 1 1 0 0 0 W. mucronata 0 0 0 0 0 0 0 0 0 1 0 0 0 W. sessilifolia 0 1 1 1 1 0 0 1 0 1 0 0 0 W. vlokii 1 0 0 0 0 0 0 1 0 1 0 0 0 Wiborgia fusca 0 0 0 0 0 0 1 0 0 0 1 1 0 W. tetraptera 0 1 1 0 0 0 1 0 0 0 1 1 1 Aspalathus cordata 0 0 0 1 0 1 0 1 0 0 1 0 1 A. nivea 0 0 0 1 0 1 0 0 0 0 1 0 0 Calobota cytisoides 0 0 0 0 0 0 0 0 0 0 0 0 0

(1) Persistence: perennial = 0, short-lived fireweeds = 1. (2) Branches: not rigid = 0, rigid = 1. (3) Petiole: not tuberculate = 0, tuberculate = 1. (4) Petiole: present = 0, absent = 1. (5) Calyx tube: short = 0, long = 1. (6) Inner surface of calyx lobes: pubescent = 0, glabrous = 1. (7) Standard petal: upright = 0, reflexed = 1. (8) Wing petals: shorter than keel = 0, longer than keel = 1. (9) Wing petals: broadest near base or apex = 0, broadest near middle = 1. (10) Anther arrangement: 4+5+1 = 0, 4+6 = 1. (11) Ovules: many = 0, few = 1. (12) Fruit type: legume = 0, samara = 1. (13) Fruit: without wings = 0, winged = 1.

198 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

61 Wiborgiella bowieana Wiborgiella sessilifolia Wiborgiella leipoldtiana 65 Wiborgiella fasciculata Wiborgiella humilis Wiborgiella dahlgrenii

70 Wiborgiella inflata Wiborgiella vlokii 100 Wiborgiella mucronata

70 Wiborgia fusca Wiborgia tetraptera 8 Aspalathus cordata Aspalathus nivea Calobota cytisoides

FIG. 7.4. Strict consensus of three trees produced by the analysis of 13 morphological characters for Aspalathus, Wiborgia and Wiborgiella. Values above the branches are bootstrap percentages above 50%.

7.3 TAXONOMIC TREATMENT

WIBORGIELLA Boatwr. and B.-E.van Wyk, gen. nov. (S. Afr. J. Bot.: submitted. 2008),

Wiborgiae Thunb. similis sed fructibus oblongis non alatis valde inflatis, anthera

carinali breve (aliis in eodem fore brevibus dorsifixis simile) differt; Calobotae

Eckl. and Zeyh. similis sed petalis ubique glabris, antheris ut supra et foliis

dorsiventralibus differt; Lebeckiae Thunb. sensu stricto similis sed foliolis planis

199 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

non acicularibus valde differt.—TYPE species: Wiborgiella leipoldtiana

(R.Dahlgren ex Schltr.) Boatwr. and B.-E.van Wyk.

Lebeckia section Viborgioides Benth. in Hook. Lond. J. Bot. 3: 361. 1844. —

LECTOTYPE species (designated here): Wiborgiella fasciculata (Benth.) Boatwr.

and B.-E.van Wyk. [Note: As there is no indication as to which species would be

a better choice of lectotype, we here choose the first listed species.]

[Note: the new genus proposed here conforms to Bentham's (1844) concept of

Lebeckia section Viborgioides, a taxon for which no name is available at generic level.

The name Wiborgiella reflects the vegetative similarity and close relation to the genus

Wiborgia.]

Rigid, resprouting, woody shrubs (rarely a lignotuberous shrublet or short-lived fireweed). Branches thick and woody (except in two short-lived species); young branches brown, covered with bark (rarely green in two species), pubescent. Stipules absent. Leaves digitately trifoliolate; petioles shorter than leaflets, pubescent, usually persistent and becoming woody after leaflets are shed; leaflets linear to widely oblanceolate or obovate, pubescent. Inflorescence terminal, multi-flowered racemes or rarely single-flowered. Pedicel pubescent. Bracts linear to lanceolate or elliptic, at least slightly pubescent, caducous. Bracteoles linear to lanceolate, at least slightly pubescent, caducous. Corolla yellow, older petals fading purple or orange, glabrous.

Calyx subequally lobed, upper sinus often deeper than the lateral or lower sinuses, carinal lobe narrower than the others, pubescent or glabrous. Standard ovate to widely

200 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA ovate, obovate or cordate. Wing petals oblong, ovate or rarely obovate, shorter, as long as or longer than the keel petals, glabrous; apex obtuse. Keel petals boat-shaped or lunate, pockets present or absent, glabrous; apex obtuse, acute or sub-acute. Anthers dimorphic, four long, basifixed anthers alternating with five ovate, dorsifixed anthers, carinal anther resembling the ovate, dorsifixed anthers (4+6 configuration). Pistil subsessile to long-stipitate, ovary linear to slightly elliptic, with 4 to many ovules, glabrous, very rarely slightly pubescent on the upper basal parts; style shorter or longer than the ovary, curved upwards, glabrous. Pods inflated or turgid, rarely laterally compressed in W. mucronata and W. vlokii, linear to oblong to oblanceolate or lanceolate or elliptic, few- to many-seeded, glabrous, dehiscent (rarely indehiscent).

Seeds reniform or suborbicular, light pink (W. leipoldtiana), light brown or orange mottled with brown (W. humilis), or black with white or light brown spots (W. inflata and

W vlokii), surface smooth or rugose (seeds of W. bowieana, W. fasciculata, W mucronata and W. sessilifolia not seen).

Diagnostic characters—Bark formation is early and the young twigs are brown and covered with bark as opposed to the green twigs of Calobota (the short-lived

Wiborgiella inflata and W. vlokii have green stems due to their herbaceous habit). The genus is similar to Wiborgia, but differs in the oblong, wingless, much inflated fruit

(ovate to orbicular, markedly winged and laterally compressed in Wiborgia). It also differs in the carinal anther that resembles the short, dorsifixed anthers (carinal anther intermediate in' both Wiborgia and Calobota). It further differs from Calobota in the totally glabrous petals (in Calobota, at least the standard petal has a few apical hairs)

201 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA and in the dorsiventral leaves (isobilateral in Calobota). It differs markedly from

Lebeckia sensu stricto in the flat leaflets (invariably acicular in Lebeckia).

Notes on distribution —The genus is endemic to the Cape region of South

Africa.

KEY TO THE SPECIES OF WIBORGIELLA

1. Branches green, short-lived fireweeds 2

2. Pods inflated, leaves lanceolate W. inflata

Pods laterally compressed, leaves oblanceolate W. vlokii

1 Branches brown, long-lived shrubs or resprouting shrublets 3

Calyx 9.0-10.5 mm long, calyx lobes 3.0-5.5 mm long 4

Inflorescences with 8 to 15 flowers, calyx lobes 4.0-5.5 mm long, claws

of all petals relatively short, ovary linear, style shorter than

ovary W sessilifolia

4. Inflorescences with(1–) 2 to 5 flowers, calyx lobes 3.0-3 5 mm long,

claws of all petals relatively long, ovary elliptic, style longer than

ovary W. bowieana

3. Calyx <9 mm long, calyx lobes mm long 5

5. Wings longer than the keel and down-curved, pods narrowly ovate, flowers pink, plants restricted to Saldanha Bay and Hopefield W. dahlgrenii

202 CHAPTER 7: TAXONOMY OF. THE GENUS WIBORGIELLA

5. Wings shorter than the keel and straight, pods oblanceolate to elliptic, linear or obovate, flowers yellow, plants occurring in the Eastern Cape, Niewoudtville, Gifberg,

Albertinia or from Sutherland north to Vioolsdrif 6

6. Pods laterally compressed, petioles not tuberculate or persistent

W. mucronata

6. Pods inflated, petioles tuberculate and persistent 7

7. Leaves linear to spathulate, pods long-stipitate, pods 4-6 mm

long W. humilis

7. Leaves linear-oblanceolate to ovate, pods short stipitate, pods

more than 14 mm long 8

8. Pods linear, flowers 9-10 mm long, Albertinia

W. fasciculata

8. Pods oblanceolate to elliptic, flowers 10-13 mm long,

from Sutherland area northwards to Vioolsdrif

(Nababeepberg) W. leipoldtiana

1 WIBORGIELLA BOWIEANA (Benth.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia bowieana Benth. in Hook. Lond J. Bot. 3: 362-

363. 1844; Walp., Rep. Bot. Syst.: 454. 1845; Harv. in Harv. and Sond., Fl. Cap.

2: 89. 1862; R.Dahlgren, Op. Bot. 38: 79. 1975; Bond and Goldblatt, J. S. Afr.

Bot. 13: 289. 1984; Goldblatt and Manning, Cape Pl.: 493. 2000.—TYPE:

SOUTH AFRICA, 'Cape' [grid unknown], Bowie s.n. (lectotype: K!, designated by

Dahlgren [1975]; isolectotype: BM!).

203 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Erect, rigid, woody, single-stemmed shrub up to 1.5 m in height. Branches greyish-brown; young branches sericeous; older branches covered in longitudinally splitting brown-grey bark. Leaves digitately trifoliolate; petiole 1-2 mm long, tuberculate, becoming hard and persistent on older branches after leaflets have been shed; leaflets oblanceolate to spathulate, sericeous on both surfaces, subsessile, terminal leaflet 4-6 mm long, 1-2 mm wide, lateral leaflets 3-5 mm long, 1-2 mm wide; apex acute to apiculate or rarely retuse; base cuneate. Inflorescence 18-35 mm long, with flowers solitary or 2 to 5 per raceme; pedicel 1.0-1.5 mm long; bract 4-5 mm long, linear to lanceolate, relatively long, sericeous on adaxial surface; bracteoles 3-5 mm long, linear to lanceolate, relatively long, sericeous on adaxial surface. Flowers 10-15 mm long, yellow with brown markings on abaxial surface of standard (according to label information). Calyx 9.0-10.5 mm long, glabrous or rarely sparsely pilose; tube relatively long, 5.5-7.0 mm long; lobes 3.0-3.5 mm long, deltoid. Standard 16.0-16.5 mm long, glabrous, very rarely sparsely pilose along the dorsal midrib; claw linear, relatively long,

6.0-6.5 mm long; lamina ovate, 10.0-10.5 mm long, 7.5-8.0 mm wide; apex obtuse.

Wings 15-16 mm long, glabrous; claw relatively long, 6-7 mm long; lamina oblong, 9-

10 mm long, 4.0-4.5 mm wide, longer than keel, with 8-11 rows of sculpturing. Keel

14-15 mm long, glabrous; claw relatively long, 6-7 mm long, lamina lunate, 6.5-8.0 mm long, 4.0-4.5 mm wide; apex acute; pocket present. Pistil subsessile, glabrous; ovary elliptic, 5.5-6.5 mm long, 1.3-1.5 mm wide with ±8 ovules; style longer than ovary, 9-10 mm long. Pods ovate to lanceolate, dark brown, rugulose, strongly inflated, subsessile, 15-20 mm long, 5-7 mm wide, up to ?5 seeded, dehiscent. Seeds not seen

(Fig. 7.5). Flowering time: May to July.

204 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

g d

jl j2 j3

1■■

FIG. 7.5 Morphology of Wiborgiella bowieana: (al)) bract; (a2) bracteoles; (b) leaf in abaxial view; (c) flower in lateral view; (d) standard petal; (e) wing petal; (f) keel petal; (g) outer surface of the calyx (upper lobes to the left); (h) androecium; (i) pistil; (j1) long, basifixed anther; 02) carinal anther resembling the short, dorsifixed anthers; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (al—j3) Streicher s.n. (JRAU); (k1—k2) Van Wyk 2106 (JRAU).

Scale bars: 1 mm.

205 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Diagnostic characters—Wiborgiella bowieana is similar to W. sessilifolia, but differs in the petiolate leaves on older branches, narrower leaflets, few-flowered inflorescences, largely glabrous calyx, shorter calyx lobes, long petal claws and the elliptic ovary with the style longer than the ovary (in W. sessilifolia the leaves on the older branches are sessile, the leaflets broader, the inflorescences more densely flowered, the calyx pilose with longer lobes, the petals claws relatively short and the linear ovary is longer than the style). It is also similar to W. dahlgrenii, but differs from this species in the larger leaflets, long calyx tube and calyx lobes, larger flowers, long petal claws and larger pods (in W. dahlgrenii the leaflets are smaller, the calyx tube and lobes shorter, the petal claws shorter and the flowers and pods smaller than in W. bowieana).

Distribution and habitat—Wiborgiella bowieana is a rare and highly restricted species that occurs on rocky soils from Worcester in the west to around Bredasdorp in the east (Fig. 7.6).

Note—Raimondo et al. (in press) assessed the conservation status of this species as critically endangered (CR A2bc; C2a(i); D). They speculate that the habitat of this species has been transformed for wheat farming over the last few decades and as few as 50 individuals may be in existence today. Overgrazing leads to ongoing degradation of its habitat.

Additional specimens examined

3319 (Worcester): Robertson (—DD), Schmidt 30 (PRE).

3321 (Ladismith): Attaquaskloof (—DD), Zeyher 2345 (K, NBG, S).

3420 (Bredasdorp): Uitvlug Farm, north of Bredasdorp (—AB), Streichers.n. (JRAU 2 sheets),

Streichers.n. sub Schutte 831 (JRAU), Van Wyk 2106 (JRAU).

206 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.6 Known geographical distribution of Wiborgiella bowieana.

2. Wiborgiella dahlgrenii Boatwr. and B.-E.van Wyk, sp. nov., similar to W. bowieana

(Benth.) Boatwr. and B.-E.van Wyk but differs in the smaller leaves, shorter

bracts and bracteoles, smaller flowers, relatively short claws, obovate to cordate

standard petal lamina, down-curved wing petals, pilose calyx and shorter, few-

seeded pods. It is also similar to W. leipoldtiana (Schltr. ex R.Dahlgren) Boatwr.

and B.-E.van Wyk, but differs in the smaller flowers, obovate to cordate standard

petal lamina, long, down-curved wing petals, style that is longer than the ovary

and narrowly ovate, few-seeded pods (to be translated into Latin).—TYPE:

SOUTH AFRICA, Langebaan, Abbus Farm [3318 AA], Baker 10407 (holotype:

NBG!).

207 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Rigid, woody, multi-stemmed shrub of unknown height. Branches brown; young branches sericeous; older branches covered in longitudinally splitting light brown bark.

Leaves digitately trifoliolate; petiole 2-8 mm long, somewhat tuberculate, not persistent although base or tubercle persistent on older branches after leaflets have been shed; leaflets obovate to oblanceolate or rarely elliptic, sericeous to sparsely sericeous on both surfaces, subsessile, terminal leaflet 3.5-7.0 mm long, 1.5-4.0 mm wide, lateral leaflets 3.0-5.5 mm long, 1.5-3.0 mm wide; apex retuse to obcordate or rarely acute; base cuneate. Inflorescence 22-40 mm long, with 4 to 8 flowers; pedicel 1 mm long; bract 1.5-2.0 mm long, linear to lanceolate, sericeous on adaxial surface; bracteoles

1.0-1.5 mm long, linear to lanceolate, sericeous on adaxial surface. Flowers 7-9 mm long, light pink (according to label information) or possibly yellow with older petals fading pink. Calyx 4.5-5.0 mm long, pilose; tube 3.0-3.5 mm long; lobes 1.5-2.0 mm long, deltoid. Standard 8.0-9.5 mm long, rarely sparsely pilose along dorsal midrib; claw linear, 3-4 mm long; lamina obovate to cordate, 5.0-5.5 mm long, 4.5-5.0 mm wide; apex shallowly emarginate. Wings 8.5-9.0 mm long, glabrous; claw 2.5-3.0 mm long; lamina oblong and somewhat down-curved, 5.5-6.5 mm long, 2.0-2.5 mm wide, longer than keel, with 4 rows of sculpturing. Keel 6-7 mm long, glabrous; claw 2.0-2.5 mm long; lamina lunate, 3.5-4.0 mm long, 2.0-2.5 mm wide; slight pocket present; apex acute. Pistil subsessile to shortly stipitate; ovary ovate, 3.0-3.5 mm long, 1.0-1.2 mm wide with 4 to 5 ovules; style as long as or longer than ovary, 3.5-4.0 mm long.

Pods narrowly ovate, rugulose, inflated, subsessile to shortly stipitate, dark brown, 10-

12 mm long, 3-5 mm wide, ±2-seeded, dehiscent. Seeds not seen (Fig. 7.7). Flowering time: There were two specimens of this species available for study. The single flowering

208 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA specimen was collected in August, suggesting that the species flowers in early spring.

The fruiting specimen was collected in September.

Diagnostic characters—This species is similar to Wiborgiella bowieana but differs in the smaller leaves, shorter bracts and bracteoles, smaller flowers, the pink flower colour, relatively short claws, obovate to cordate standard petal lamina, down- curved wing petals, pilose calyx and shorter, few-seeded pods. It is also similar to W. leipoldtiana, but differs in the smaller, pink flowers, obovate to cordate standard petal lamina, long, down-curved wing petals, style that is longer than the ovary and narrowly ovate, few-seeded pods. In both W. bowieana and W. leipoldtiana the flowers are yellow and larger than in W. dahlgrenii, the wing petals not down-curved and the standard petals ovate. In W. bowieana the calyx is large and pilose and the pods larger than in W. dahlgrenii. The pods are many seeded in W. leipoldtiana and the style shorter than the ovary.

209 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

d a

e J1 12 j3

b f

k2 c1 h1 h2

FIG. 7.7 Morphology of Wiborgiella dahlgrenii: (a) flower in lateral view; (b) standard petal; (c1, c2) leaves in abaxial view; (d) wing petal; (e) keel petal; (f) pistil; (g) outer surface of the calyx

(upper lobes to the left); (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther;

(j2) carinal anther resembling the short, dorsifixed anthers; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a—c1, d—j3) Barker 10407 (NBG); (c1, k1—k2) Bolus 12915 (BOL).

Scale bars: 1 mm.

210 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Distribution and habitat— This is a very rare species which is poorly collected and known from only two collections (Fig. 7.8). It occurs in Strandveld vegetation around Saldanha Bay and Hopefield and has been recorded on sandy soil at an altitude of 15 m.

Note—This species is in desperate need of conservation attention. Collection attempts have been unsuccessful and it is possible that the plant is either very highly localised or extinct due to severe habitat disturbance. It is known only from the

Saldanha Bay and Langebaan areas where the natural vegetation is severely fragmented.

Additional specimens examined

-3317 (Saldanha): near Hoetje's Bay, Saldanha Bay (-BB), Bolus 12915 (BOL) (fruiting material).

FIG. 7.8. Known geographical distribution of Wiborgiella dahlgrenii.

211 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

3. WIBORGIELLA FASCICULATA (Benth.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia fasciculata Benth. in Hook. Lond. J. Bot. 3: 361.

1844; Walp., Rep. Bot. Syst.: 453. 1845; R.Dahlgren, Op. Bot. 38: 74. 1975;

Bond and Goldblatt, J. S. Bot. 13: 289. 1984; Goldblatt and Manning, Cape Pl.:

493. 2000.—TYPE: SOUTH AFRICA, 'Cape' [grid unknown], Bowie s.n.

(lectotype: K!, designated by Dahlgren [1975]; isolectotype: BM!).

Small, erect, rigid, multi-stemmed shrublet up to ?0.5 m in height. Branches greyish-brown; young branches densely pilose; older branches covered in longitudinally splitting brown-grey bark. Leaves digitately trifoliolate; petiole 0.5-1.0 mm long, tuberculate, becoming hard and persistent on older branches after leaflets have been shed; leaflets linear-oblanceolate to oblanceolate, pilose on both surfaces, subsessile, terminal leaflet 3-5 mm long, 1.0-1.5 mm wide, lateral leaflets 2-3 mm long, 1.0-1.5 mm wide; apex truncate or acute; base cuneate. Inflorescence 28-30 mm long, with 5 to 15 flowers; pedicel 1.0-1.5 mm long; bract 1.0-1.4 mm long, lanceolate to ovate, pilose along margins and adaxial surface; bracteoles 1.0-1.4 mm long, elliptic to lanceolate, pilose along margins and adaxial surface. Flowers 9-10 mm long, yellow.

Calyx 4.0-4.5 mm long, glabrous; tube 2.5-3.0 mm long; lobes 1.0-1.5 mm long, deltoid. Standard 8.5-10.0 mm long, glabrous; claw linear, 3.0-3.5 mm long; lamina ovate, 5.5-6.5 mm long, 4.0-4.5 mm wide; apex obtuse. Wings 8.0-8.5 mm long, glabrous; claw 3.0-3.5 mm long; lamina oblong, 4.5-5.0 mm long, 2.0-2.5 mm, shorter than keel, wide with 6-7 rows of sculpturing. Keel 10.5-11.0 mm long, glabrous; claw

3.5-4.0 mm long; lamina boat-shaped, 6.5-7.5 mm long, 3.0-3.5 mm wide; apex sub-

212 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA acute, pocket present. Pistil subsessile, glabrous; ovary linear, 6.0-6.2 mm long, 0.7 mm wide with 7 to 9 ovules; style shorter than ovary, ±4.3 mm long. Pods linear, rugulose, inflated, subsessile, 14-17 mm long, 3-4 mm wide, number of seeds unknown, dark brown to black, dehiscent. Seeds not seen (Fig. 7.9). Flowering time:

September (spring).

Diagnostic characters—Wiborgiella fasciculata is similar to Wiborgiella leipoldtiana, but differs in the smaller, pilose leaves, smaller flowers and shorter, linear pods. In W. leipoldtiana the leaves and flowers are much larger and the pods oblanceolate to elliptic.

Distribution and habitat—Wiborgiella fasciculata is a very rare and localised species that occurs in the Riversdale area around Albertinia (Fig. 7.10).

Note—Raimondo et al. (in press) assessed the status of Wiborgiella fasciculata as critically endangered (CR B1ab(iii)+2ab(iii); D). The populations of this species are severely fragmented and declining due to the exclusion of fire. A total of about 13 plants are still in existence.

Additional specimens examined

—3421 (Riversdale): Onverwacht, Albertinia, Riversdale division (—BA), Muir 859 (BOL, PRE),

Muir 4775 (NBG).

213 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

d a

b f j1 Qi j2 j3 9

h1 h2 k2

FIG. 7.9 Morphology of Wiborgiella fasciculata: (a) flower in lateral view; (b) standard petal; (c) leaf in abaxial view; (d) wing petal; (e) keel petal; (f) pistil; (g) outer surface of the calyx (upper lobes to the left); (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) carinal anther resembling the short, dorsifixed anthers; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a—j3) Muir 859 (NBG); (k1—k2) Bowie s.n. (BM). Scale bars: 1 mm.

214 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.10 Known geographical distribution of Wiborgiella fasciculata.

4. WIBORGIELLA HUMILIS (Thunb.) Boatwr. and B.-E.van Wyk in S. Afr. J. Bot.: submitted.

2008. Lebeckia humilis Thunb., Nov. Gen.: 142. 1800., Fl. Cap.: 562. 1823;

Harv. in Harv. and Sond., Fl. Cap. 2: 89. 1862. Wiborgia humilis (Thunb.)

R.Dahlgren, Op. Bot. 38: 31. 1975.—TYPE: SOUTH AFRICA, e Cap. B. Spei

[grid unknown], Thunberg s.n. sub THUNB-UPS 16416 (lectotype: UPS!,

designated by Dahlgren [1975]).

Wiborgia apterophora R.Dahlgren, Bot. Notiser 123: 112. 1970.—TYPE: SOUTH

AFRICA, Gifberg [3118 DC], sand at edge of lands, Esterhuysen 22042

(holotype: BOL; isotype: K!).

215 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Small, erect or decumbent, multi-stemmed shrublet up to 0.5 m in height.

Branches brown; young branches densely pilose; older branches covered in longitudinally splitting brown bark. Leaves digitately trifoliolate; petiole 0.5 or less-1.5 mm long, tuberculate, becoming hard and persistent on older branches after leaflets have been shed; leaflets linear-spathulate, pilose on abaxial surface, adaxial surface glabrous, subsessile, terminal leaflet 5-12 mm long, 0.8-1.2 mm wide, lateral leaflets

5-10 mm long, 0.8-1.0 mm wide; apex truncate to retuse or acute, base cuneate.

Inflorescence (32—) 35-70 mm long, with (5—) 8 to ±13 flowers; pedicel 1.0-2.5 mm long; bract 1.0-1.5 mm long, lanceolate to linear, pilose on adaxial surface; bracteoles

0.7-1.0 mm long, lanceolate to linear, pilose on adaxial surface. Flowers 6.5-10.0 mm long, yellow. Calyx 2.5-4.0 mm long, glabrescent; tube 2.0-3.0 mm long; lobes 0.8-1.1 mm long, deltoid. Standard 8.0-8.5 mm long, glabrous; claw linear, 2.0-2.5 mm long; lamina ovate, 5.0-6.0 mm long, 3.5-4.0 mm wide, apex acute. Wings 7.5-8.5 mm long, glabrous; claw 2.5-3.0 mm long; lamina oblong, 4.5-6.0 mm long, 2.5-3.5 mm wide, shorter than or as long as keel, with 6-7 rows of sculpturing. Keel 8.0-9.0 mm long, glabrous; claw 2.5-3.0 mm long; lamina boat-shaped, 6.0-6.5 mm long, 3.0-4.0 mm wide, apex obtuse; pocket present. Pistil long-stipitate (stipe 2.5-3.0 mm long), glabrous, sometimes pilose on upper basal parts; ovary elliptic, 5-6 mm long, 0.8-0.9 mm wide with ±5 ovules; style shorter than ovary, 3-4 mm long. Pods elliptic to obovate, rugulose, inflated, long-stipitate, 4-6 mm long, 1.0-1.5 mm wide, ±2-seeded, dehiscent. Seeds reniform, 2.0-2.7 mm long, 1.5-2.3 mm wide, mature seeds light brown or light orange mottled with brown, surface rugose (Fig. 7.11). Flowering time:

Spring to summer (September to November).

216 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

a d

O i1 (-( i2R i3 a b

J kl

g1 g2 ft 6 k2 C

FIG. 7.11 Morphology of Wiborgiella humilis: (a) flower in lateral view; (b) standard petal; (c) leaf in abaxial view; (d) wing petal; (e) keel petal; (f) outer surface of the calyx (upper lobes to the left); (g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) carinal anther resembling the short, dorsifixed anthers; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a—e, k1—k2) Boatwright et al. 216 (JRAU); (f—i3) Van Wyk 3530 (JRAU).

Scale bars: 1 mm.

217 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Diagnostic characters—This species is similar to W. leipoldtiana, but differs in the linear-spathulate leaves, smaller flowers, shorter calyx tube and calyx lobes, long- stipitate fruit and pistil, small, few-seeded pods and rugose seeds (W. leipoldtiana has oblanceolate to obovate leaves, larger flowers, a longer calyx tube and calyx lobes, short-stipitate fruit and pistil, many-seeded pods with smooth seeds).

Distribution and habitat—Wiborgiella humilis is a highly restricted species that occurs around Gifberg and Nieuwoudtville (Fig. 7.12). It has been recorded in sandy or rocky soils, often along streams, at altitudes of between 480 m and 800 m in fynbos.

Additional specimens examined

3118 (Vanrhynsdorp): Gifberg Plateau (—DC), Acocks 14892 (PRE), Acocks 14898 (K, PRE).

3119 (Calvinia): Keyzerfontein, Sanddraai on Farm Oorlogskloof (—AC), Boatwright et al. 129

(JRAU); Farm Oorlogskloof (—AC), Boatwright et al. 216 (JRAU); on road to Keyzerfontein, south of the Calvinia road, close to Nieuwoudtville (—AC), Goldblatt 7068 (PRE); Farm

Kleinarendskraal, close to edge of Nieuwoudtville plato (—AC), Niemand 8 (JRAU); Oorlogskloof

Nature Reserve, 15 km south-west of Nieuwoudtville, Farm Gipherkop 804, 300 m south of waterfall on burnt area (—AC), Pretorius 392 (NBG); Kranskloof 792 (—AC), Pretorius 646 (K,

NBG, PRE); Nieuwoudtville, Arendskraal, Rooi pad (—AC), Van Wyk 3530 (JRAU).

218 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.12 Known geographical distribution of Wiborgiella humilis.

5. WIBORGIELLA INFLATA (H.Bolus) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia inflate H.Bolus in Hook. !cones Pl. 16: pl. 1576. 1887;

Salter in Adamson and Salter, Fl. Cape Peninsula: 471. 1950; Bond and Goldblatt, J. S.

Afr. Bot. 13: 289. 1984; Goldblatt and Manning, Cape Pl.: 493. 2000.—TYPE: SOUTH

AFRICA, Western Cape Province, eastern slopes of Devil's Peak [3318 CD], Bolus

4826 (holotype: BOL!; isotypes: K!, NBG!).

Small, erect or decumbent, short-lived herb up to 0.3 m in height. Branches green, angular, densely to sparsely pilose. Leaves digitately trifoliolate; petiole 7-20 mm long, not tuberculate or persistent; leaflets lanceolate, subsessile, densely to

219 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA sparsely pilose on both surfaces, terminal leaflet 14-36 mm long, 1.5-3.5 mm wide; lateral leaflets 10-23 mm long, 1-2 mm wide; apex acute; base cuneate. Inflorescence

(27—) 50-135 (-255) mm long, with 4 to 13 flowers; pedicel 2-5 mm long; bract 1-2 mm long, elliptic to lanceolate, pilose on adaxial surface; bracteoles 1.5-2.5 mm long, linear, pilose on adaxial surface. Flowers 8-10 mm long, yellow, older petals fading purple. Calyx 3.5-5.0 mm long, glabrous or very sparsely pilose; tube 2.5-3.0 mm long; lobes 1-3 mm long, deltoid; acute. Standard 7-12 mm long, glabrous; claw linear, 3.0-

5.5 mm long; lamina ovate, 4.0-7.5 mm long, 4.5-8.0 mm wide; apex obtuse. Wing petals 8-10 mm long, glabrous; claw 3.54.5 mm long; lamina oblong, 4.5-6.5 mm long, 2.0-3.0 mm, as long as or longer than the keel, wide with 6 to 8 rows of sculpturing. Keel petals 7.5-8.5 mm long, glabrous; claw 3.0-4.0 mm long; lamina lunate, 4.0-5.0 mm long, 2.0-2.5 mm wide, apex acute; pocket absent. Pistil subsessile, glabrous; ovary elliptic, 5.0-6.5 mm long, 0.8-1.5 mm wide with up to ±21 ovules; style 2-3 mm long, shorter than the ovary. Pods sessile, oblong, smooth, inflated, 28-35 mm long, 5.5-10.0 mm wide, glabrous. Seed suborbicular, 1.5-2.1 mm long, 1.7-2.0 mm wide, rugose, black mottled with white (Fig. 7.13). Flowering time:

Spring to early summer (October to November).

220 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

a d

e i2 6Di3 1J) i1

g1 g2 k2

FIG. 7.13 Morphology of Wiborgiella inflata: (a) flower in lateral view; (b) standard petal; (c) leaf in adaxial and abaxial view; (d) wing petal; (e) keel petal; (f) outer surface of the calyx (upper lobes to the left); (g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) carinal anther resembling the short, dorsifixed anthers; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a, b, d—f) Vlok et al. 2 (JRAU); (c, k1—k2) Johns 162 (JRAU); (g1—j)

Haynes H420 (PRE). Scale bars: 1 mm

221 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Diagnostic characters—Wiborgiella inflata is closely related to W. vlokii, but differs in the inflated, oblong pods and lanceolate leaves. The latter species has oblanceolate leaves and laterally compressed, linear pods.

Distribution and habitat—Wiborgiella inflata occurs from the Cape Peninsula eastwards to the Kogelberg, the Hottentots Holland Mountains and also the Kleinrivier

Mountains at Hermanus. It typically grows in recently burnt vegetation, one to two years after fire (Fig. 7.14). It can be found on Table Mountain Sandstones (TMS), in damp stream banks on clay soils or on rocky soils and east to south-west facing slopes between 40 m to 750 m. It is grazed by livestock.

Additional specimens examined

3318 (Cape Town): Kirstenbosch, Nursery Gorge (—CD), Esterhuysen 6143 (BOL, K, NBG,

PRE 2 sheets); Table Mountain, slopes above south end of Kirstenbosch (—CD), Esterhuysen

22402 (BOL).

3418 (Simonstown): Cape Peninsula, between Constantia Corner and Castle Rock (—AB),

Lewis 1816 (NBG); locally common on upper south slopes of Gerbera Hill, west of the

Muizenberg (—AB), Pillans 3465 (BOL); Nuweberg south facing slopes of Dwarsrivier

Mountains, above jeep track, Kogelberg Nature Reserve (—BD), Le Maitre 578 (NBG, PRE).

3419 (Caledon): Lebanon Forest Reserve, Jakkalsrivier Catchment (—AA), Haynes H420

(NBG, PRE), Haynes 474 (PRE); Glenfrum Onrus Mountain, Hermanus (—AD), Barker 204

(JRAU); Vogelgat Nature Reserve (—AD), Stirton 11170 (K); Vogelgat above Dragon Fly Pool (-

AD), Williams 3693 (NBG 2 sheets, PRE); Kogelberg Nature Reserve (—BD), Johns 162

(JRAU), Vlok et al. 2 (JRAU).

222 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.14 Known geographical distribution of Wiborgiella inflata.

6. WIBORGIELLA LEIPOLDTIANA (Schltr. ex R.Dahlgren) Boatwr. and B.-E.van Wyk, comb.

nov. (S. Afr. J. Bot.: submitted. 2008). Lebeckia leipoldtiana Schltr. ex

R.Dahlgren, Op. Bot. 38: 72. 1975; Bond and Goldblatt, J. S. Afr. Bot. 13: 289.

1984; Goldblatt and Manning, Cape Pl.: 493. 2000.—TYPE: SOUTH AFRICA,

Western Cape Province, Calvinia Div.: 'Inter Grasberg et Nieuwoudtville' [3119

AC], Lewis 5839 (holotype: NBG!; isotype: NBG!).

Crotalaria aspalathoides auct. non Lamarck: E.Mey., Comm. Pl. Afr. Austr. 1: 25. Feb.

1836; Benth. in Hook. Lond. J. Bot. 2: 591. 1843; Harv. in Harv. and Sond., Fl.

Cap. 2: 46. 1862.

223 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Erect, rigid, woody, multi-stemmed shrub up to 1.2 m in height. Branches brown; young branches sericeous; older branches covered in longitudinally splitting brown bark. Leaves digitately trifoliolate; petiole 1-3 mm long, tuberculate, becoming hard and persistent on older branches after leaflets have been shed; leaflets oblanceolate to obovate, sericeous to sparsely sericeous on both surfaces, subsessile, terminal leaflet

4-11 mm long, 1.5-4.0 mm wide, lateral leaflets 4-10 (-11) mm long, 1.5-3.0 mm wide; apex retuse to obcordate, base cuneate. Inflorescence (16—) 20-40 mm long, with 4 to

9 flowers; pedicel 1-3 mm long; bract 1.5-2.2 mm long, linear to lanceolate, sericeous on adaxial surface; bracteoles 1.0-1.5 mm long, linear to lanceolate, sericeous on adaxial surface. Flowers 10-13 mm long, yellow. Calyx 4.5-6.5 mm long, pilose; tube

3.0-4.5 mm long; lobes 1-2 mm long, deltoid. Standard 12.5-16.0 mm long, glabrous or very rarely sparsely pilose along dorsal midrib; claw linear, 4.0-6.5 mm long; lamina ovate, 8.5-9.5 mm long, 5.5-6.0 mm wide; apex obtuse or sub-acute. Wings 10.0-12.5 mm long, glabrous; claw 4.0-5.0 mm long; lamina oblong, 6.0-7.5 mm long, 3-4 mm wide, shorter than keel, with 4-6 rows of sculpturing. Keel 11.5-14.0 mm long, glabrous; claw 4.0-5.5 mm long; lamina boat-shaped, 8.0-9.5 mm long, 4-5 mm wide, apex acute; pocket present. Pistil subsessile to shortly stipitate, rarely pubescent on the upper basal parts; ovary linear, 8.5-10.0 mm long, 1.0-1.3 mm wide with 10 to 12 ovules; style shorter than ovary, 5-6 mm long. Pods oblanceolate to elliptic, inflated, smooth, subsessile to shortly stipitate, dark brown, 16-26 mm long, 3-6 mm wide, ±7- seeded, dehiscent. Seeds reniform, 2.5-3.0 mm long, 2.0-2.6 mm wide, mature seeds light pink, surface smooth (Fig. 7.15). Flowering time: Spring, August to October.

224 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

j1 j2 j3 il

k2 h1 h2 d

FIG. 7.15 Morphology of Wiborgiella leipoldtiana: (a) flower in lateral view; (b) standard petal;

(c) outer surface of the calyx (upper lobes to the left); (d) leaf in abaxial view; (e) wing petal; (f) keel petal; (g) pistil; (h1) bract; (h2) bracteoles; (i) androecium; (j1) long, basifixed anther; (j2) carinal anther resembling the short, dorsifixed anthers; (j3) short, dorsifixed anther; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a, c—f, j1—j3) Boatwright et al. 123 (JRAU); (b, h1—i) Schutte 295 (JRAU);

(g) Hanekom 2125 (PRE); (k1—k2) Van Wyk 3278 (JRAU). Scale bars: 1 mm

225 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Diagnostic characters—This species is similar to Wiborgiella fasciculata, but differs in the longer leaflets, larger flowers and oblanceolate to elliptic, larger pods (the leaflets are short, the flowers small and the pods linear and shorter in W. fasciculata). It differs from W. dahlgrenii in the larger yellow flowers, ovate standard petal, short wing petals, many-seeded oblanceolate to elliptic pods and style that is shorter than the ovary (W. dahlgrenii has smaller pink flowers, an obovate to cordate standard petal lamina, long, down-curved wing petals, the style that is longer than the ovary and narrowly ovate, few-seeded pods). It is also similar to W. humilis, but differs in the oblanceolate to obovate leaves, larger flowers, longer calyx tube and calyx lobes, many-seeded, much larger pods and smooth seeds (in W. humilis the leaves are linear- spathulate, the flowers smaller, the calyx tube and calyx lobes shorter, the pods small and few-seeded and the seeds rugose). Furthermore, it differs from W. mucronata in the sericeous oblanceolate to obovate leaflets, shorter bracts and bracteoles, sub-acute keel petals and inflated, oblong to elliptic pods (W. mucronata has sparsely pilose, linear to oblanceolate leaflets, long bracts and bracteoles, obtuse keel petals and laterally compressed, linear pods).

Distribution and habitat—Wiborgiella leipoldtiana is widely distributed from the

Sutherland area northwards as far as Nababeepberg in the Vioolsdrif area (Fig. 7.16). It occurs in Renosterveld on well-drained sandy loam, red clay or rocky slopes or disturbed roadsides between 600m and 850 m and is grazed by livestock.

Additional specimens examined

-2917 (Springbok): Steinkopf (-BC), Horrocks 14 (NBG); Anenous Mountain (-BC), Taylor

1161 (BOL); Spektakel (-DA), Bolus 9494 (PRE), Bolus 9495 (BOL); Spektakel Hill (-DA),

Maguire 1013 (NBG); Nababeepberg, near top of mountain (-DC), Van Wyk 3278 (JRAU).

226 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

—3118 (Vanrhynsdorp): Kromme River (—BB), Esterhuysen 20532 (BOL).

3119 (Calvinia): Grasberg road, 300 m along turn-off from tar near Nieuwoudtville (—AA),

Taylor 11787 (PRE); on Farm Oorlogskloof at Gletservloer (—AC), Boatwright et al. 123 (JRAU);

Redclay, Klip Koppies, trails northeast of Nieuwoudtville (—AC), Bolus 19558 (BOL 2 sheets, K);

Vanrhyns Pass (—AC), Esterhuysen 5311 (BOL), Bond 1148 (NBG); north-west of

Nieuwoudtville in sandy soil near Grasberg (—AC), Goldblatt 5842 (PRE); Namaqualand,

Remhoogte 416, western slopes of northern end of Grasberg (—AC), Le Roux 4346 (NBG);

Oorlogskloof Nature Reserve, Nieuwoudtville (—AC), Pretorius 104 (NBG, PRE, WIND),

Pretorius 133 (NBG, PRE); between Nieuwoudtville and Grasberg (—AC), Schutte 295 (JRAU 2 sheets), Taylor 11331 (JRAU, NBG, PRE 2 sheets).

—3120 (Williston): Roggeveld Escarpment near Middelpos (—CC), Levyns 9476 (BOL).

3219 (Wuppertal): Citadel Kop (—AA), Compton 24254 (NBG); Biedouw Valley, Mertenhof (-

AA), Hanekom 3179 (NBG), Van Breda 4311 (K); Pakhuis, along the road to Wuppertal (—AA),

Leipoldt 444 (BOL, K); Diamond Drift, Biedouw River between Pakhuis and Wuppertal (—AA),

Leipoldt 3131 (BOL); Brandewynrivier (—AA), Schlechter 10823 (BM, BOL, K); Pakhuis Pass (-

AA), Stokoe s.n. (NBG); Koudeberg Mountains, near Wuppertal (—AC), Bolus 8974 (BOL, K,

NBG); ca. 5 km northwest of Wuppertal (—AC), Strid and Strid 37902 (NBG); Citrusdal (—CA),

Taylor 1219 (BOL).

3220 (Sutherland): Kleinfontein (Agterkop), peak Middelpos (—AA), Hanekom 2125 (PRE);

13.5 mls [21.72 km] south south-west of Komsberg Pass (—DB), Acocks 18449 (K).

227 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.16 Known geographical distribution of Wiborgiella leipoldtiana.

7. WIBORGIELLA MUCRONATA (Benth.) Boatwr. and B.-E.van Wyk, comb. nov. (S. Afr. J.

Bot.: submitted. 2008). Lebeckia mucronata Benth. in Hook. Lond. J. Bot. 3: 359.

1844; Walp., Rep. Bot. Syst.: 453. 1845; Harv. in Harv. and Sond., Fl. Cap. 2:

87. 1862; Bond and Goldblatt, J. S. Afr. Bot. 13: 290. 1984; Goldblatt and

Manning, Cape Pl.: 493. 2000.—TYPE: SOUTH AFRICA, Uitenhage [3325 CD],

Zeyher 344 (lectotype: K!, designated here; isolectotypes: BM!, TCD, photo!).

[Note: The K specimen is chosen as lectotype as most of Bentham's type

material is housed in this herbarium (Stafleu and Cowan 1976).]

Lebeckia leptophylla Benth. in Hook. Lond J. Bot. 3: 359. 1844; Walp., Rep. Bot. Syst.:

453. 1845; Harv. in Harv. and Sond., Fl. Cap. 2: 87. 1862; Bond and Goldblatt, J.

228 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

S. Afr. Bot. 13: 289. 1984; Goldblatt and Manning, Cape Pl.: 493. 2000, synon.

nov.—TYPE: SOUTH AFRICA, 'Grassy subalpine situations near Swellendam'

[3420 AB], Mundt 87 (lectotype: K!, designated here; isolectotype: S!). [Note:

The K specimen is very rich with flowers and fruit and is a better choice of

lectotype than the fairly poor specimen in S.]

Small, erect, multi-stemmed shrublet up to 0.3 m in height. Branches brown; young branches densely pilose; older branches covered in longitudinally splitting brown bark. Leaves digitately trifoliolate, petiole 2-10 (-15) mm long, not tuberculate or persistent; leaflets linear to oblanceolate, sparsely pilose, subsessile, terminal leaflet

(4–) 7-23 mm long, 1-4 mm wide, lateral leaflets 4-12 mm long, 1.0-3.5 mm wide; apex mucronate; base angustate. Inflorescence (-23) 35-75 (-115) mm long, with (5–)

8 to 15 (-21) flowers; pedicel 1-2 mm long; bract 4.0-5.5 mm long, linear, pilose on adaxial surface; bracteoles 1.5-3.5 mm long, linear, pilose on adaxial surface. Flowers

8-13 mm long, yellow with back of standard reddish brown (according to label information). Calyx 5-7 mm long, pubescent; tube 3.0-4.5 mm long; lobes 1.5-3.0 mm long, deltoid. Standard 11.5-13.5 mm long, glabrous; claw linear, 2.5-4.5 mm long; lamina ovate, 7-9 mm long, 7.5-9.0 mm wide; apex obtuse. Wings 11.0-12.5 mm long, glabrous; claw 3.5-4.5 mm long; lamina oblong, 6.5-8.0 mm long, 3.5-5.0 mm wide, shorter than or as long as keel, with 5-7 rows of sculpturing. Keel 11.5-13.0 mm long, glabrous; claw 3.5-5.0 mm long; lamina boat-shaped, 7.5-8.5 mm long, 5-6 mm wide, apex obtuse; pocket present. Pistil subsessile to shortly stipitate, glabrous; ovary linear,

7.5-9.5 mm long, 1.3-1.5 mm wide with 16 to 22 ovules; style shorter than ovary, 4.5–

229 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

5.5 mm long. Pods linear to somewhat clavate, smooth, laterally compressed, subsessile to shortly stipitate, 20-25 mm long, 4-5 mm wide, ±10-seeded, dehiscent.

Seeds not seen (Fig. 7.17). Flowering time: Most of the flowering specimens were collected in spring (September to October) and summer (November) and one in May.

Wiborgiella mucronata appears to flower in response to fire.

Diagnostic characters — Wiborgiella mucronata is similar to Wiborgiella leipoldtiana, but differs in the sparsely pilose, linear to oblanceolate leaflets, long bracts and bracteoles, obtuse keel petals and laterally compressed, linear pods. In W. leipoldtiana the leaflets are sericeous and oblanceolate to obovate, the bracts and bracteoles shorter, the keel petals sub-acute and the pods inflated and oblong to elliptic.

Distribution and habitat— Wiborgiella mucronata occurs mainly in the Eastern

Cape Province around Grahamstown, Port Elizabeth and Oudtshoorn, with some outlier populations around Bredasdorp (Fig. 7.18). It occurs in recently burnt vegetation up to the second or third year after fire. This species is found on south facing slopes on sandy soil or in moist fynbos at altitudes of between 300 m and 750 m.

230 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

i1 i2 i3 b

g1 g2

FIG. 7.17 Morphology of Wiborgiella mucronata: (a) flower in lateral view; (b) standard petal; (c) outer surface of the calyx (upper lobes to the left); (dl, d2) leaves in abaxial view; (e) wing petal; (f) keel petal; (g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) carinal anther resembling the short, dorsifixed anthers; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view; (k2) pod in dorsal view.

231 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Voucher specimens: (a—c, e, f, h, i1—i3, j) Stirton 10880 (JRAU); (d1) Esterhuysen 27306

(BOL); (d2) Vlok 1726 (JRAU), (g1—g2) Stirton and Zantovska 11608 (PRE); (k1—k2)

Esterhuysen 6880 (BOL). Scale bars: 1 mm

Additional specimens examined

—3322 (Oudtshoorn): northern foothills of Outeniqua Mountains, about 3 km south-west of

Herold at Camferskloof (—CD), Vlok 1726 (JRAU, PRE).

3323 (Willowmore): Wagenbooms River (—CD), Fourcade 2340 (BOL, K); Kouga Mountains,

near Smutsburg (—DB), Esterhuysen 10749 (BOL); De Hoek, northern base of Zitzikamma

Mountains near Joubertina (—DC), Esterhuysen 27306 (BOL, K); Joubertina (—DD),

Esterhuysen 6880 (BOL), Esterhuysen 6904 (BOL, PRE).

3324 (Steytlerville): top of Elandsberg on road to Patensie above Erasmuskraal farm (—BD),

Schrire 2011 (GRA); Kareedouw Pass (—CD), Gillett 2280 (BOL, NBG); poort between Patensie

and Cambria (—DA), Thompson 1889 (NBG); Elandsberg Mountains, north of Patensie (—DB),

Stirton 10880 (JRAU 2 sheets).

3325 (Port Elizabeth): Ten Stop Hill, Groendal, Uitenhage (—CA), Olivier 3196 (PRE); Van

Stadens River Mountains (—CC), Bolus 1605 (BOL), Zeyher 2318 (NBG, S); Van Stadensberg

Mountains, Longmore Forest Station far eastern end (—CC), Stirton and Zantovska 11608

(NBG, PRE).

—3326 (Grahamstown): Howieson's Poort (—AD), Britten 5157 (GRA); Faraway, portion 3 of

Coldsprings (—AD), Jacot Guillarmod 8387, 9234 (GRA); Grahamstown (—BC), Britten 5064

(GRA), MacOwen 692 (K); Featherstone's Kloof (—BC), Galpin 253 (GRA, PRE), MacOwen 692

(NBG).

3420 (Bredasdorp): Marloth Nature Reserve (—AB), Taylor 7688 (NBG); hill beyond

Lemmetjiesdorp (—AB), Wurts 470 (NBG).

232 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.18 Known geographical distribution of Wiborgiella mucronata.

8. WIBORGIELLA SESSILIFOLIA (Eckl. and Zeyh.) Boatwr. and B.-E.van Wyk, comb. nov.

(S. Afr. J. Bot.: submitted. 2008). Acanthobotrya sessilifolia Eckl. and Zeyh.,

Enum. Pl. Afric. Austral. 2: 193. Jan. 1836. Wiborgia (Viborgia) grandiflora

E.Mey., Comm. Pl. Afr. Austr. 1: 31. Feb. 1836. Lebeckia sessilifolia (Eckl. and

Zeyh.) Benth. in Hook. Lond. J. Bot. 3: 362. 1844; Walp., Rep. Bot. Syst.: 453.

1845; Harv. in Harv. and Sond., Fl. Cap. 2: 89. 1862; R.Dahlgren, Op. Bot. 9:

261-262. 1963; R.Dahlgren, Op. Bot. 38: 76. 1975; Bond and Goldblatt, J. S.

Bot. 13: 290. 1984; Goldblatt and Manning, Cape Pl.: 494. 2000.—TYPE:

SOUTH AFRICA, 'between Breede River and Duyvelshoek', Ecklon and Zeyher

1344 (lectotype: S!, designated by Dahlgren [1975]; isolectotypes: K!, SAM!).

233 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Erect, rigid, woody, multi-stemmed shrub up to 1.5 m in height. Branches brown- grey; young branches densely sericeous; older branches covered in longitudinally splitting brown-grey bark. Leaves digitately trifoliolate; petiole tuberculate, becoming hard and persistent on older branches after leaflets have been shed, 1.5-3.0 mm long; leaflets widely oblanceolate, sericeous to sparsely pilose on both surfaces, those that appear first are subsessile and subsequently followed by leaves arising from the axil of the petiole that are sessile, terminal leaflet 4-13 mm long, 1.5-4.0 mm wide, lateral leaflets 4-12 mm long, 1-3 mm wide; apex retuse or truncate; base cuneate.

Inflorescence 25-60 (-70) mm long, with 8 to 15 flowers; pedicel 1-3 mm long; bract

3-5 mm long, linear to lanceolate, pilose on adaxial surface; bracteoles 3-5 mm long, linear to lanceolate, pilose on adaxial surface. Flowers (-8) 10-12 mm long, yellow, older petals turning orange. Calyx 8.0-10.5 mm long, densely pilose; tube 4.5-6.0 mm long; lobes 4.0-5.5 mm long, subulate. Standard 11-13 mm long, glabrous; claw linear,

3.5-4.5 mm long; lamina widely ovate, 7.5-8.5 mm long, 9.5-11.0 mm wide; apex obtuse. Wings 12.0-14.5 mm long, glabrous; claw 4-5 mm long; lamina obovate, 8-10 mm long, 4-6 mm wide, longer than keel, with 7-8 rows of sculpturing. Keel 10.5-12.0 mm long, glabrous; claw 4.0-4.5 mm long; lamina boat-shaped, 6.0-7.5 mm long, 4-5 mm wide, apex obtuse; pocket present. Pistil subsessile to shortly stipitate, glabrous; ovary linear broadening towards the base, 7.5-9.0 mm long, 1.5-2.0 mm wide with 8 to

13 ovules; style shorter than ovary, 4.5-5.0 mm long. Pods linear-oblong to lanceolate, smooth, inflated, subsessile to shortly stipitate, 19-21 mm long, 4-5 mm wide, ±5- seeded, dehiscent. Seeds not seen (Fig. 7.19). Flowering time: Spring, mainly in

September and October.

234 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Diagnostic characters — Wiborgiella sessilifolia is similar to W. bowieana, but differs in the sessile leaves on older branches, many-flowered inflorescences, pilose calyx with longer calyx lobes, relatively short petal claws, linear ovary with the style shorter than the ovary and the linear oblong to lanceolate pods (W. bowieana has petiolate leaves on the older branches, few-flowered inflorescences, a largely glabrous calyx, shorter calyx lobes, long petal claws and an elliptic ovary with the style longer than the ovary).

Distribution and habitat— Wiborgiella sessilifolia occurs along the Cape coast from Bredasdorp to Still Bay. It appears to be restricted to limestone hills and grows on south or south-west facing slopes at altitudes of 100 m to 200 m (Fig. 7.20).

Additional specimens examined

–3420 (Bredasdorp): 2.5 mls [4.02 km] south-west of Wydgelegen Post Office (–AD), Acocks

23002 (K, PRE); Potberg Nature Reserve, Windhoek, Fonteinkloof on west side of De Hoopvlei,

De Hoop Nature Reserve (–AD), Burgers 1252 (NBG, PRE); De Hoop Hills, 3.10 mls [4.99 km] from Wydgelegen towards De Hoop (–AD), Taylor 4329 (PRE); Wydgelee, on road to De Hoop

(–AD), Thompson 3262 (K, NBG); on limestone hills after entrance to De Hoop Nature Reserve

(–BC), Boatwright et al. 170, 198 (JRAU); Hamerkop (–BC), Van Wyk 1907 (NBG); along entrance road to De Hoop just after the main gate (–BC), Van Wyk 2120 (JRAU 5 sheets);

Zoetendals Valley, south Bredasdorp neat the old cemetery (–CA), Albertyn 498a (NBG);

Grasrug downs on south side of Karsrivier near Drie Mond (–CA), Albertyn 498b (NBG, PRE);

Die Mond (–CA), Martin 591 (NBG); Agulhas, De Mond Nature Reserve on limestone outcrop

on the west side of the river (–CC), Euston - Brown 201 (NBG); Struisbaai, ridge west of

Hotagterklip on Paapekuilfontein 281 (–CC), Helme 3696 (NBG).

235 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

—3421 (Riversdale): 17 mls (27.4 km) south of Riversdale (—AC), Acocks 24111 (K, NBG, PRE);

Stillbay, ridge east of bridge (—AD), Bohnen 4203 (NBG, PRE); Stillbay rifle range kloof (—AD),

Bohnen 4832 (NBG); Hills at Melkhoutfontein, near Stillbay (—BD), Muir 2444 (PRE).

Precise locality unknown: Caledon, Mundt 22 (K).

a h IMIIMIMI

il i2 i3

kl ••■

g2 k2

FIG. 7.19 Morphology of Wiborgiella sessilifolia: (a) flower in lateral view; (b) standard petal; (c) leaves in abaxial view; (d) wing petal; (e) keel petal; (f) outer surface of the calyx (upper lobes to the left); (g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) carinal anther resembling the short, dorsifixed anthers; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view; (k2) pod in dorsal view.

236 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Voucher specimens: (a—i3) Van Wyk 2120 (JRAU); (j) Albertyn 498b (NBG); (k1—k2) Taylor

4329 (PRE). Scale bars: 1 mm

FIG. 7.20 Known geographical distribution of Wiborgiella sessilifolia.

9. Wiborgiella vlokii Boatwr. and B.-E.van Wyk, sp. nov., similar to Wiborgiella inflata

(H. Bolus) Boatwr. and B.-E.van Wyk but differs in the laterally compressed pods

and shorter, oblanceolate leaflets of up to 13 mm (in W. inflata the pods are

strongly inflated, the leaflets are lanceolate and up to 36 mm long) (To be

translated into Latin).—TYPE: SOUTH AFRICA, Western Cape Province,

Langeberg Mountains, next to forestry track to Helderfontein [3320 DD], Vlok

2045 (holotype: PRE!, designated here; isotype: PRE!).

237 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

Small, erect or decumbent, short-lived herb up to 0.15 m in height. Branches green, angular, densely to sparsely pilose. Leaves digitately trifoliolate; petiole 4-9 mm long, not tuberculate or persistent; leaflets oblanceolate, densely to sparsely pilose on both surfaces, subsessile, terminal leaflet 8-13 mm long, 2.5-3.5 mm wide; lateral leaflets 6-10 mm long, 2-3 mm wide; oblanceolate; apex apiculate or retuse; base cuneate. Inflorescence 43-90 mm long, with 3 to 5 flowers; pedicel 2-3 mm long; bract

±1.5 mm long, ovate to linear, pilose on adaxial surface; bracteoles ±1.1 mm long, ovate to linear, pilose on adaxial surface. Flowers 9-10 mm long, yellow, older petals turning orange (according to label information). Calyx 4.5-5.0 mm long, glabrous on the outer surface; tube 2.5-3.0 mm long; lobes 1.0-1.5 mm long, deltoid. Standard 9.5-

10.0 mm long, glabrous; claw 3.5-4.0 mm long; lamina ovate, 5.5-6.0 mm long, 4.0-

4.5 mm wide; apex obtuse. Wing petals 9.0-10.5 mm long, glabrous; claw 3.5-4.5 mm long; lamina oblong, 4.5-5.5 mm long, 2.0 mm wide, ± as long as the keel or longer, with 6 to 8 rows of sculpturing. Keel petals 8.5-10.0 mm long, glabrous; claw 3.5-4.5 mm long; lamina boat-shaped, 4.5-5.5 mm long, 2.5-3.0 mm wide; apex acute; pocket absent. Pistil sub-sessile, glabrous; ovary elliptic, 7.0-7.5 mm long, 1.0-1.2 mm wide, with 13 to 18 ovules; style shorter than the ovary, 2.5-3.0 mm long. Pods sessile, oblong, smooth, laterally compressed, 23-31 mm long, 4.0-5.5 mm wide, ±10-seeded, glabrous. Seed suborbicular, 2.3-2.4 mm long, 2.2-2.7 mm wide, rugose, black mottled with white (Fig. 7.21). Flowering time: Spring and summer. The only two available specimens with flowers and fruit and were collected in November.

Diagnostic characters — Wiborgiella vlokii is only known from the type collection. It is very similar to W. inflate, but differs in the shorter, oblanceolate leaves

238 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA and linear, laterally compressed pods. Wiborgiella inflate has lanceolate leaves and inflated, oblong pods.

Distribution and habitat—Wiborgiella vlokii is a very rare species and is only known from the Langeberg Mountains where it occurs in deep, peaty soil on southern slopes in recently burnt mountain fynbos at an altitude of ca. 808 m (Fig. 7.22).

Note—This species is in need of conservation attention as it is known from a single collection in the Langeberg Mountains and has not subsequently been recollected.

b i2 i3

g2 kl gl k2

239 CHAPTER 7: TAXONOMY OF THE GENUS WIBORGIELLA

FIG. 7.21 Morphology of Wiborgiella vlokii: (a) flower in lateral view; (b) standard petal; (c) leaf in abaxial view; (d) wing petal; (e) keel petal; (f) outer surface of the calyx (upper lobes to the left); (g1) bract; (g2) bracteoles; (h) androecium; (i1) long, basifixed anther; (i2) carinal anther resembling the short, dorsifixed anthers; (i3) short, dorsifixed anther; (j) pistil; (k1) pod in lateral view; (k2) pod in dorsal view.

Voucher specimens: (a—k2) Vlok 2045 (PRE). Scale bars: 1 mm

FIG. 7.22 Known geographical distribution of Wiborgiella vlokii.

240 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS, REINSTATEMENT OF EUCHLORA,

LEOBORDEA AND LISTIA AND A NEW GENUS OF THE TRIBE CROTALARIEAE (FABACEAE)

8.1 INTRODUCTION

Lotononis is a large and complex genus of the tribe Crotalarieae with approximately 151 species [one recently described from Namibia by Van Wyk and

Kolberg (2008)], 145 of which occur in South Africa. The classification system of the genus has been somewhat volatile since the original concept was established by De

Candolle in 1925, published as Ononis section Lotononis. This section was thereafter raised to generic level by Ecklon and Zeyher (1836) who, along with

Meyer (1836), proposed numerous other genera to accommodate the species of what is today Lotononis. The genus Euchlora was described by Ecklon and Zeyher

(1836) to accommodate an anomalous plant that was placed in the genus Ononis by

Thunberg (1800) and in Crotalaria and later Microtropis by Meyer (1832, 1836).

Bentham (1843) expanded the concept of Lotononis to include the genera described by Ecklon and Zeyher (1836) and Meyer (1836), with the exception of Euchlora, and a sectional classification was proposed which Harvey (1862) also followed in the treatment of the genus in the Flora Capensis. Dahlgren (1964) discussed the taxonomic history and synonymy of Euchlora and mentioned the similarities between it and Lotononis, such as trifoliolate, stipulate leaves (in some forms of

Lotononis hirsuta), a similar hair type and the warty upper suture of the pod.

Euchlora serpens was subsequently transferred to Lotononis (as L. hirsuta) by

Dahlgren (1964). The most recent revision of Lotononis by Van Wyk (1991b), follows Bentham (1843) to some extent but extended the generic concept to include

241 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Buchenroedera Eckl. and Zeyh. Based on cladistic analyses of morphlogical, cytological and chemical data, a detailed infrageneric classification system was proposed, comprising 15 sections (Van Wyk 1991b). This detailed study, published as a series of papers and as a synopsis in 1991, represents the most rigorous treatment of the genus to date, and provided a framework for future studies on

Lotononis.

Lotononis shares similarities with many of the genera in the tribe (viz.

Crotalaria, Lebeckia, Pearsonia, Rothia) and a sister relationship with Crotalaria was suggested by Van Wyk (1991a) based on the presence of macrocyclic pyrrolizidine alkaloids and rugose seeds.

The data presented in Chapter 3 (Boatwright et al. 2008a) showed that

Lotononis is polyphyletic, through the analysis of nrITS and plastid rbcL sequence data which included 52 species of Lotononis representing all the currently

recognized sections of the genus. However, the analysis of morphological characters in combination with the sequence data indicated that Lotononis was in fact weakly supported to be monophyletic if L. hirsuta (L. section Euchlora) was excluded. However, the anomalous Lotononis macrocarpa could not be included in this study seeing that it is an extremely rare species and several attempts to collect this plant have failed. Lotononis macrocarpa possesses a unique combination of characters: paired stipules that are equal in size, a 5+5 anther arrangement, presence of bracteoles and large fruit. Subsequent to the study of Boatwright et al.

(2008a) material of this species was obtained from a herbarium specimen of a recent collection of this plant. This in combination with available data from

Boatwright et al. (2008a) has allowed for a re-evaluation of the generic circumscription of Lotononis.

242 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

This chapter is aimed at presenting a new generic classification system for

Lotononis s.l. based on molecular and morphological evidence. A discussion of critical characters is also presented along with systematic data on the placement of the anomalous Lotononis macrocarpa. A key to the genera of the Crotalarieae is

also presented in which the changes at generic level formalised in this chapter are

incorporated.

8.2 RESULTS

8.2.1 Combined ITS/rbcL data set—The combined molecular data set

consisted of 1,854 included characters, 1,401 of which were constant, 453 variable

and 276 parsimony informative. A total of 560 trees were retrieved using MP with a tree length (TL) of 1,056, consistency index (CI) of 0.55 and a retention index (RI) of

0.82 (Fig. 8.1a). Lotononis was polyphyletic with Lotononis s.s. (L. section Lotononis

and allies) sister to Pearsonia, Robynsiophyton and Rothia, i.e. the Pearsonia Glade

(51 bootstrap percentage [BP]). Lotononis s.s. was strongly supported to be

monophyletic (100 BP). The Leobordea Glade (Lotononis section Leobordea and

allies) was sister to the Listia Glade (Lotononis section Listia excluding L.

macrocarpa) with a bootstrap percentage of 75 and both these groups were strongly supported as monophyletic (99 BP and 100 BP, respectively). Lotononis hirsuta

(section Euchlora) was strongly supported as sister to Bolusia and Crotalaria (100

BP), while L. macrocarpa was placed within the "Cape group", albeit without support.

8.2.2 Combined ITS/rbcL/morphological data set—The analysis of the combined molecular and morphological matrix included 1,885 characters, 1,401 of

243 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS which were constant, 484 variable and 404 parsimony informative. The resulting 370 trees from the MP analysis had a TL of 1,185 steps, CI of 0.52 and a RI of 0.84 (Fig.

8.1b). The topology differed slightly from that obtained by Boatwright et al. (2008a;

Chapter 3), largely within the "Cape group". Aspalathus and Wiborgia were sister to each other and both monophyletic (98 BP and 94 BP, respectively). Successively sister was the genus Wiborgiella, which was also monophyletic, albeit without support. The following Glade comprised Lebeckia and Rafnia that were sister taxa

(without support) and moderately to strongly supported as monophyletic (82 BP and

99 BP, respectively). Calobota formed the next Glade and was strongly supported as

monophyletic (94 BP). The "Cape group" was weakly supported (69 BP) and

Lotononis macrocarpa placed in a position sister to this group, but without bootstrap support. With the exclusion of Lotononis hirsuta and L. macrocarpa, the rest of

Lotononis s.l. was weakly supported as monophyletic, as opposed to being

polyphyletic in the combined molecular analysis. Lotononis s.s, the Leobordea Glade

and the Listia Glade were strongly supported (100 BP; 100 BP; 100 BP, respectively) with the Leobordea and Listia clades strongly supported to be sister (86 BP).

Lotononis hirsuta (section Euchlora) was strongly supported as sister to Bolusia and

Crotalaria (100 BP).

8.2.3 Evolution of morphological characters—The reconstructions of the

12 morphological characters onto the minimal length trees are indicated in Fig. 8.2-

8.13. These reconstructions are represented on one of the 560 or 370 equally most

parsimonious trees from the combined molecular or combined molecular/

morphological analyses, respectively, but if more than one reconstruction was

possible these are also shown. The large tuber (character 1) was reconstructed as an apomorphy for Lotononis hirsuta (Fig. 8.2). Digitate leaves are predominant in

244 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS the basal taxa of the tribe, but in the "Cape group" some variation was found, notably in the reduction to simple or unifoliolate leaves (character 2; Fig. 8.3).

Lotononis hirsuta has predominantly simple leaves, but digitate leaves are found in some forms. Asymmetrical stipules (character 3; Fig. 8.4) are found mostly in

Lotononis s.l. Only one species of Rothia, R. hirsuta has single stipules. In the combined molecular analysis this character is convergent between Lotononis s.s. and the Leobordea and Listia clades, while in the combined molecular and morphological analysis it is reconstructed as an apomorphy for Lotononis s.l. with a

reversal in L. sections Aulacinthus and Polylobium. Loss of bracteoles (character 4;

Fig. 8.5) is convergent between Lotononis hirsuta and the other Lotononis clades, while the Listia Glade is unique in having bracteoles. A zygomorphic calyx (character

5; Fig. 8.6) is reconstructed as a synapomorphy for Lotononis s.s., the Leobordea

and Listia clades and Pearsonia with a reversal to an equal or subequal calyx in

Robynsiophyton and Rothia. Rostrate or coiled keel petals (character 6; Fig. 8.7)

evolved multiple times in Crotalarieae and are found in Crotalaria, Bolusia,

Lotononis section Oxydium, Lebeckia and in some species of Rafnia and

Aspalathus. An anther arrangement (character 7; Fig. 8.8) of 4+1+5 is most frequent

in the "Cape group" with Lebeckia and Wiborgiella distinctly unique with a 5+5 and

4+6 arrangement, respectively. This character is variable within Lotononis s.l. with

only the Listia Glade having an exclusively 4+1+5 arrangement. A verrusoce upper

suture of the pod (character 8; Fig. 8.9) is extremely convergent within the tribe and

present in species of all the Lotononis clades, except the Listia Glade. The presence

of long funicles (character 9; Fig. 8.10) was reconstructed as an apomorphy for

Lotononis s.l. in the combined molecular and morphological analysis, but is also

present in Lotononis macrocarpa. A chromosome number (character 10; Fig. 8.11)

245

CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

of x=9 appears to be the base number in the Crotalarieae with a reduction to x=8 in

Crotalaria, Rafnia and some Aspalathus, while x=7 is found in Lotononis s.s. and

Pearsonia. Cyanogenesis (character 11; Fig. 8.12) is unique to Lotononis s.s. with a

reversal in L. section Cleistogama, while macrocyclic pyrrolizidine alkaloids

(character 12; Fig. 8.13) are present in Crotalaria, Lotononis s.s. and some species

of the Leobordea Glade (Lotononis section Synclistus).

Agastalles Ow. 1 100 A7,13411116a OMNI. 1 055515858 807.88.0 8800810 Aspokt. amen Aspetatnus 5151.853 mom. 5555151185.0ermana AspelaDne maws 7ir.,....------,...... 28881.988 9808019. 09000. 881.88 ----...:-PeP'..7""'''' 2.80•1•1188 5'''''''85588511 '05"".11555 71.901999 087809.8580 208091. .890 atop. 558.5 Aspalailurs rAd* 810.1m gm.* 1958 41.2960 Aspalathus carhas Aab5b51.4 57.9 MU 9B 0- 80990/789 pats 8.09 pit-,751 .87009080 hystna 2-855558188 oentsta 45 n 51151885 y• A.m.. 8.81500 08851515 555184518 15155715 Ms. 1 1888•A'":1= Mon 8 5 1584885 W., 5 • torn 1 80279,815 Om. 508551. 815548 01 .=15,80.881 1505 55511. 1 I 97 Near& 89990810/9 1 Er8r2' Mar& 90890079 1 1188.2.589 muctonala 02 812894070 Damp1 1111,9619 011887 1515 a 4 Hi a larpoidtlane 1 118,18005,81 8.505 1 1515548115 558.5558tor 1.18002888 Moo.* 1 8818000885 kw* 1 00:099688 00.8782. M.O. 8007. 2 1818089080115.088 1 51058419 18008125 8028919,5 2 r="csmosa 2 51 acuminats jarr4- 1.85.1.18 m•yellen• 1 ROBB 80.1958 8080720089280.085 1 87 /24599088 Lebeckda ptarolbr a 2 BOBO Otto. L•00.41 5.08 1 151.1.71195 L... be5.595 8 I555885 8.000a 2 C ,_L Law. mayeran• 1 Gob.. plIonstiana 68 "05820.77curninata."''''''' ''' L51558,18 885,8855. Lizir.= 8188.7090988C 1.80•0815518858 Rah. 555818 Lob.* /9951a 1 Le** woo. = 807589'''' Gebbota pungens 2 100 Ca.. 8,908 2 Ga.. asp.. 2 5' C.O. 0090918 2 00080 8951051 Cat* .091000 " C9,98010 87800.5 CalobolC"rn cyfuoidea"'' 22 75 41.•• 01 as.. *we 6l1952 lk..1012/02 1.51518 Icecoonci. 8518.08208515 2 C... 5& 5M., .• 5.58.8555550...... 2 0 0•95019 spinescens 55 Calobola wend Cal.. chorea C... *a Caktars saber. 1.8855554 51•019.058 880598728 tnactoca.• 89.800 dam 1 ,.. Et-0=c= 1.58.80159 faica 1 Lor,v1Ords enachy8,558 ladenonis spanolon Lamm spersilkoa 2 Lonna. 8,555 58 5015nords 4+18182. 5.08515158 roshate 9,588. Ma 5531511155055. nern1.o* 1.058518 *lab ■ Anlicut, 20.0.1. LOtOlena 89 87c 1.08080 meow.. 92 73= .6999015 5199659 a Lolonanla made Lama. 218198 Lolononis maculala 08 1.555,585 55.1515 1- Lao.. cans loionsrls 551885. ICC 10985080 6570.10. nc orson5 58655,55 2 , [01=1.•0o5s 05815/9. 2 58 100 Lama. 5.5815115.5 1 Law* pangs. 2 Lalmanes pmarts 2 1.9nonla Orouleda BO ..===.14., Loten.la seficophytla 1 1 loknorsa puthera Lownonos woad 1- Law* rne,r1 1 cotron41 m•yon 1.05810,10 dans& sob. 9599.098 009r55.75. 1.0105558 85.858841 IL la 1.545551.8558588118 mama .05584518855crea 58888 pectoncularis Md 51/9/2. 95.188511. tax* 8.58535i i1 Lalanonb5langlata 97 81c 1,..018999.5 70188 Involcom,55 851:88 585,555815 _8--.082ore15555.5 Lottrom 9800009091 Oles,ta P.m* 58552.110 1 Loom. 1051850079 5 Prersonda eristad ;Wawa 510d5568 ImowW toboaa 2 Powspral 54858555 10/8/805..0910 /288. .54. SB /a* acfposs• Rol,. 5 5 .50er ystil Lalononla 05585 580 Won* 91588880 5 tnion08155140588 LoM5 585.• 191958558 Lc... Mad 1 2002/10,112 Man. ,884, Latononia 58,558 0 1.588.18 0355515 Latex. ealkaps Worm 8558558•15 2 Lotman. 0485.8.8 240270. 20117:12nA2 to0555.5 Musk 20/123/111214 Lobo* mais 1 -Lcanona d5155. Loisnona 5555118 “tonenla 1.08800 .0.00.5 Lotonce. 1.0,011.11,001111.1. Laces. OM.= LOAMIS 2aratc 75 luf1141E2 latononi5 1958 8658. 1.0108055 Cm ladonoms 050. 1 5010nerIa Lotman 00 Pearsove NW.* 1 LotOncr. anat. lgo.. MUM. 151 87 pram.* Loran.. statata Pearsans5 mgr.. 2 4959818 8572 /79895 WIMPS* rnadoth2 8•805/88/1 Warm. 0885 1 1.0551.118 8s09 1.800. 1994 1 Caton* Y.. Euchlas Cacalaro 070ans 085.8418:50,8 8.8801815 poposnos 00 52 0,800.81007855,89 Coata-081.955.00 1 6907708 sows. 1 Cn514545 v58.88 1 C.0,881800 9.801918 87.2881.8 hurnlis 1 0.2008.8.8005 Gob., "Kb" 828..19 889880800 Bata *con= 1 DoSesia eanboonta 1 090085 098.80 0882.

FIG. 8.1 (a) Strict consensus of the 560 most parsimonious trees from the combined

molecular analysis; (b) Strict consensus of the 370 equally parsimonious trees from the

246 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

combined molecular/morphological analysis. Numbers above the branches are bootstrap

percentages above 50%.

8.3 DISCUSSION

8.3.1 Evolution of characters in the tribe Crotalarieae—UNDERGROUND

PARTS — The enormous underground tuber found in Lotononis hirsuta is

reconstructed as an apomorphy for this taxon (Fig. 8.2). Lotononis sections

Polylobium and Lipozygis are also somewhat tuberous with annual shoots produced

from the underground tuber, but the extensive underground system of Lotononis

hirsuta is not found in any of these sections. In Lipozygis the tuberous habit could be

an adaptation to recurrent fires in the grassland habitat where these plants occur

(Van Wyk 1991b).

LEAF TYPE—A diverse array of leaf-types are found within the Crotalarieae

ranging from digitate with five, three or single leaflets, and simple or phyllodinous

leaves. Lebeckia is the only example in the tribe where the acicular leaves of some

species are articulated or "jointed" near the middle or completely reduced to petioles

(without an articulation) and the leaves are phyllodinous (Dahlgren, 1970a). The

genera of the "Cape group" of the Crotalarieae show an extreme diversity in leaf- types when compared to those genera outside the Cape region where the leaves

are generally petiolate, trifoliolate and stipulate (Dahlgren, 1970a, Polhill, 1976).

Lotononis section Euchlora has both simple, sessile leaves and digitately trifoliolate

leaves with paired stipules in the latter (Dahlgren 1964; Van Wyk 1991b). Leaf type,

however, does not provide any informative characters to circumscribe groups within

Lotononis s.l. (Fig. 8.3).

247 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

A. Immix A Moans = absent pandula A. peroduta s A. samosa A. samosa pebea A garama = present madanme A mamantha eOternorriana A. madenorsiana . confusa A. contuse perfollata autep. aniLipsii A perfollata subsP. frbiBPW pw. nmaa A. denote p.A. Leonia*. A. yermiatata - Mama raw** A. era...dou ram* LI- : ercoobelacfLtp..i. is A. larictfolia wisp. A he. subsp. bitta A bran, fra= A mow Pt PaohYlLtUa A pachylobe foam WM Not baWma Mb. talreptera . sericea wa stoic. Wdo obconlata 000 monoptara Mb. monoptata Incumata Wt. inamorata 1445. axon:lea LIMO/. muconata 131131. mean Mbl leiptlrftierta Wibt tamells 141bl sassiffogs Wed. telpotrifiarta bowiaana .__LjNitt smart:ea WWI Witt bowierana Mal maraca. u Lab. partaken Lab. samosa acuminate Lab. me.r rian.j..a I- 311raff. mammas Lab. px spicata r obot pauciflora . globose erighti Rat diffuser ab. bremicarps Lob. samosa abua. ambig meyeriana Lab. sepiaria plukenoterna Rat acurninata Lab. brevicame Rat tersonosa Lab. malaria Rat spicata Lob. ambigua Rat &boss gab. wegfsbi Rat. Warn Cat pungens CM P.O.= Cal. cuspidosa Cal psioblo Cat. pOolOba --LICot =As Cal. sedum Cal. sense Cal lotoranoidas CM. lotononoidas Cat cermet Cot drama nalanbargonsis Cal radenbargansis Cal smnescons CM spinescans ICal rename u Cat whams ItLot narmsarys Lot macrame. Lot Warta Lot. framer Lot Lot parvillour 1-7 1lLot brachyantha Lot fruticoklas Lod sparstlora At aparsatora 0.71Mall omptara Lot rostrata subs. nanumuonsis 1lLot. rostrata subsp. mena9mosis Lot lonticula Lot lantkula 1, gat macMata tit& maculate ‘.]Lot. Lot midi LOt lase 13 Lot. Ls. - Lat. mactosapala Lot. &bra Lot. micrantlet Lot. numatepala Mat &bra Lot. nalcaranlha last sable., sl Ltd sabaosa lapforoba rte , ri Lot. loptolobo ot. masimillani Lat. maximiliani &macaw La. Pomona ot wricophytta Lot..fivaricata}.. lotononordes Lot. pulchella a Lao pukhella Lot meyan me yed strfota Lot lotononokMs Lot camosa Lot stride II LOt pungens Lot. samosa Lot aeon subsp. lemodada Lot. dense steam UmbeCtaaa Lot malaria. Lot. earstipulam Lot. %Mille:MS Lot. Involubata Lot. imolucrata Wm. padunculans Lof. Intolumala subsp. peduncularls filiformis Lot. Wheats 13Lot. Wongata Lot. elongate P. Lot platycarps P. omega. La P. pa Lot. 1o/rasa P. caMMtalia Lot mamba 1-1-71Rot o'stelab Lot admessa vandrarysti Lot dacumbons Lot platycarpa Lot acsecarpa Lot. calydna Lot boluse of. folios, Lot. quinata Lot. Menthe M Lot Warms° Lot an of acullcansa Lot. molts Lot dattanbens Lot. bolusil El =Mans Lot. mobs Lot Malta. Lot. pleats L ot rLot. guinea Lot = PetsaPnola Lot soliesenis parrtaphyda subriata or. magnifica Lot. non ❑Lot. bentharniana La. mmloba La P sessaablia La =as a P arislata Lot solitudinis P. gram/Cada Lot. substrata P. cajantrolla Lot. Hail R ob Lot =dog* Rob. yandaryste Lot IMAM Lot MPArta Co dams G. deems Cr. gnmensis Cr anaconda CrCo Camera Cr capensis adrpultaks Cr virgultalls Cr. Mantis Cr Monies Cr lanceolate Cr lonmeloto arnboansls arnboansis C areas b 13 gractis

FIG. 8.2 Parsimony-based reconstructions of the presence or absence of a large underground tuber (character 1) on the 560 and 370 minimal length trees from (a) the combined molecular and (b) the combined molecular/morphological analyses, respectively.

248 • • •

CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

= digitate comae A. comae A gobsto A ammo = unifoliolate A rnacrontho A. robbromiena A. oon. 1-1"11.4.Marlariarana = simple or phyltodinous A. cora wow. pO.o. A. pens. A "Wog. sub.. priapai A vonnicasto A. crenate A Oa* *lap alma A. rtrorFuito A. brick& =bop. lariatote lreta sue.. noto A. fi÷aret:t "t a J''''"'** A Hued A pachylobe

,.....L.Re.:WD. obcordsto haelds Ylebt lopoletana

7 retie

Re mu... Lae. camas. Re mamma LabLacl. Fit ottco Rot Wire Lab montii Lab. camas. Leo. brovIcarpa Lab envenom Lab. pAranatins Lob. "MCP. tae. sopiorio rev,..xmose ab. orobio• tott werghOi Rft' A Ca purgora at. a capable, Gat prob. Col cuspeoso CP. oat.. c.1..= Cal. saris Cot = Cat lotomnokles Cot swim. Col dm. Card. lot000noldos CO beenteepanes Col cinema torentaepartes Cal alAnee.R. cr. sancta* Iasi Lot ovoroct000 CV. aaharo Lot takers Lot maao3vpa to heroes, tot mean ' Me Lot Orarbyeatha Lot spastic/a La lobar. La. coyote," Lol rr Lot menae so., 0000PPAPP.P Lot ten*. ror"*.'strate"'" sobs, aro 1.ot owe. Lot kunkaAn La cue Lot L01. laxe Lot code . moot:epee 1-1--... L"Lot mkrelabe ■ I Le sa psabaose Lot. meant. Lot lope*. La, aaeabw n , - Lot ~taco I -.911r- ea. : Loot serleepayee La Lot butcher Lot mere Lot cameo odes PiVene Lot. Mose oar. rower. Lot antipasto Lot (0.003 E4.... La I. *rate Lot axsabuito Lot lovolucrate boa,. padonoulads c, a Lot leronnis Lot involocrata beramartris Lot. Worms Lot 00017ei p same olla Lot plalyCerm Wee° onaTtloaa. La cabcino ... tor Wass Rot hes. Lot. fate. Rob. vanderysei Lot otetyceopa Lot LOLscarama Lot c✓ythr Lot Wass Lan toner Water Wt tuner Lot **now Lot we,. as Lot Orpontrns o"?.. Lot bolusa ot. moan tor PAro.1.4go Let eart1.117 171. ablate Lot: romonaca Lot bontnomiono tthdotti p Lot epee eobberla 1.?entaata La. subarea or abloaa P cale irk La. WI arndar Lot meactlei Rot. Maeda La. Note* Rob vereryea Cr. deans Lot. Mita C., grew* Cr. Agra Cr upon.* Cr gequeng, CL Kook. Cr Menet Cr. /worm Cr. 0.frouttolis Cr Isrlcoolate Cr. boner amboan Cr. lanceolate 0 grOCPsis 8 amboartals 0 p5000

FIG. 8.3 Parsimony-based reconstructions of leaf type (character 2) on the 560 and 370

minimal length trees from (a) the combined molecular and (b) the combined

molecular/morphological analyses, respectively.

249 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

STIPULE SYMMETRY—The stipules are often dimorhic in Lotononis s.s., the

Leobordea Glade and the Listia Glade, a character which was regarded as an

apomorphy for Lotononis s.l. In the trees from the combined molecular analysis the

presence of asymmetrical or single stipules was reconstructed as a convergence

between Lotononis s.s. and the Leobordea and Listia clades (Fig. 8.4). Lotononis

macrocarpa and L. hirsuta are conspicuously different in having symmetrical stipules

(when present in the latter). In Rothia hirsuta, the stipules are single as opposed to

being paired in R. indica (Chapter 10; Boatwright et al. 2008c).Asymmetrical stipules

were reconstructed as an apomorphy for Lotononis s.l. in the combined

molecular/morphological analysis, with a reversal in species from sections

Aulacinthus and Polylobium.

BRACTEOLE PRESENCE—The loss of bracteoles appears to be a largely

convergent character within the tribe and not unique to Lotononis s.l. Bracteoles are

absent in Lotononis section Euchlora, Leptis and Lotononis, as well as Rothia and

Robynsiophyton (Van Wyk 1991b; Chapter 10; Boatwright et al. 2008c; Chapter 9;

Boatwright and Van Wyk 2009). This character was reconstructed as convergences

between the groups mentioned before in both the combined molecular and combined molecular/morphological analyses (Fig. 8.5).

CALYX TYPE—The tribe Crotalarieae can be distinguished from Genisteae by the trifid lower lip of the calyx found in the latter. However, two calyx types are found within the Crotalarieae; the lebeckioid calyx type (equally or sub-equally lobed) and the Iotononoid calyx type (upper and/or lateral lobes fused higher up than the lower

lobes to varying degrees). Lotononis s.s., the Leobordea Glade, the Listia Glade and

Pearsonia all have calyces of the lotononoid type, while the remaining genera have

250 CHAPTER 8: THE GENERIC CONCEPT OF LOTONON1S the lebeckioid calyx type. The distinction of Lotononis hirsuta and L. macrocarpa

from Lotononis s.l. based on this character is notable (Fig. 8.6).

KEEL SHAPE— The shape of the keel is very important to distinguish Crotalaria

and Bolusia from the other genera in Crotalarieae. An extremely rostrate or beaked

keel is characteristic of Crotalaria, while in Bolusia the keel (and style) is twisted

through several turns and helically coiled as a result. Lotononis section Oxydium

has often been confused with Crotalaria due to the presence of rostrate keels, but

can be distinguished from the latter genus by the single stipules and loss of

bracteoles (Van Wyk 1991b). The Listia and Leobordea clades also differ from

Lotononis s.s. in their obtuse keel petals. Rostrate keel petals are shown to have

evolved several times in different lineages of the Crotalarieae by the reconstructions

on both the combined molecular and molecular/morphological trees (Fig. 8.7) and

this character is convergent between Crotalaria and Lotononis section Oxydium.

251 U) a) -C

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CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

lime. = present A pen.. IA. samosa A geleata gab., = absent A inacranthe _J macrarithe wAlanowiene AZ...arta •• I A.confuse confuse r= A perlotiate wasp. pantlail perf abate sotow. phatassi I IA. cronata crenat. vamoutato , vertnicalato sham 1 gip: weft. • Welts mblb hirta wasp. larta hee slita nine :IA moos bA "Woe pechybbe WM/ob. Wt. Iowa "aivm fusca tottbototo tetraptere Va. somas ark. N s WA monopb. WA bmb0000mdpt.sabm WM incur.te WL iarm obcOrdala WU. =Atone. NW. hurnEts • Witt /Viola loipakttiana WM trines sesedone labotdtiene . bowies. Wml sessdfolta ,r1 Ibis.. inflate .21. babies. bW:01. mucrwats Lob. m.o.. _1.)Lb samba Rat amersinefe • r • Rel. Mocenose &Ameba. spicete ob pluicznore Rel. plobotta t eb AV. Rat o7ffuse Lr.-1 Etb braviceipa Leb samosa eb embigun Lab. maw.. ab sePlada Le& plubenetlens brenterpe rscmnwe Leb. enplane Lab. ambigua gbbosa Lg. wig.° Abuse Cal pungens Cat pungens ACM. ' =•=—L—lljCal cuspid:. Cal = Cal ...be = Ctt Cal s=t satibea senses Cat lotononoides Cat tolononoidas Cat cinema cinema Cal. ludenbergensis halonteoponsia Cal spines.. Ca/ spew= Cat sabers. Veal saber. u law. Inebocarpe sl Lot rrAcrocarpe Lot falcate Lot falcate Lot huticobse Lot pewit/ore 1:t =2:rare' Lot .whys.. Lot buticoidas Lot spats.. Lot spared/orb oxyptera Lot L.' °MM. Lot rostrata subsp. nainequensis Lot rostrata nutop. namequensis Lot lenticule Lot len Lot inecubte Lot maculata Lot curtg Lot CUrti Lot. tare Lot la. Lot. macrosepate Lot glebra Lot intone.. Lot macnesepate000 Lot &bra Lol 01(0000. Lot sebubse Lot &Idiom. Lot beptolobe Lot leptolobe Lot. maximillent ol marimiliani Lot &sensate Lot pungens Lot sericophyda Lot dimricate Lot. lotmonoides Lot sencophyde Lot pub.. Lot pulchelle Lot. = Lot may. Lot lotonormides Lot. samosa Lot striae Lot. M... Lot samosa LPL dense soap. laucoclada Lot dense subsp. lemodada Lot bedbug. Lot e.lipadate Lo. inrolusrate Lot imolucrele Lot. imam:rate wino. pedunculans Lot involusrate sub. Peclunst&ft Lot. Worm. Lot Alberni, Lot elongate Lot elongate Lot. pletycame Lot catycina grandifolia Lot folio. cajanIfolie Lot Menthe Rot Mau. Lot ad... Rob. render.. Lot ...bens Lot Lot actitte,pa Lot calycina Lot bobs° Lot folioed Lot (Abate Lot. enenthe PaYOMbela Lot ed... Lbw PentsPhYps Lot .insane. Lot mobs Lot. decurnbens Lgt, LA bobs. Lot mot. Lot. plicate o mo= Lot. mina. Lot. polycephala solitudinis Lot pants". sububte. . Lot Alattoarna Lot benalarniana I taw mat.. Lot digitate Lot globulose Lot solitudinle t ie Lot subulate Lot. Otto Lot. mart.. . vender.. Lot him. lot. hirsute Cr. distans Cr. dtstens Cr. O•duensis gMbensis _LIG,. ...is _j—L1--JCr. irgui.a Cr. Men. Cr. hm. • Cr. lanceolate • Cr lanceolate p B. =bonnets ■ 113. ID. °recalls

FIG. 8.5 Parsimony-based reconstructions of the presence or absence of bracteoles

(character 4) on the 560 and 370 minimal length trees from (a) the combined molecular and

(b) the combined molecular/morphological analyses, respectively.

253 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

A. lemma A linearis = lateral lobes not fused . pandula A. pendula A. semosa el A. manse A. pabata m A. galoata = lateral lobes fused A. matrontha A. =crenate A. weldenowiana A. cledenovriene A. confuse A. sodas. A. perkitate subsp. phellpsi A pedollata subsp. phelpsi A. somata A comb A. carriage. vannsulata e A. share* 711b3p..t...7 o szzleo=fr. —TA. labor/oda esop. lark/bra tj A bole Weep Oita Aboa m subsp

1 1=1. UA A pennants, 1A. pochybba WM luxes 740, Wee ff optem Meacham bib. senses Be. senses Inge on:argon telb. monopeefe Vale monopfera Wits locum. Neb. innovate . 17410. obsonlata LW& murnenata 1,144,1 numbs nflato p_uliebt Sabo:dem o Mb Months Witel sessibblia ielpoldtiona WPC bowieana sessibelia Mb baba WSW bowieene 7401 snuentorita u Leb pentagon Lab. carnosa -u Rat acumnato Lob. mobs/lane Rat. mamma Leb. plukeneflone Rat sok. Lab. povsibore of plobosa ulna writing Ref &Musa eb. trevisame Lab. samosa bleb. ambigua Lab. rnmenarte Lab. =Om* Lob. phiwtatiana uRat acuminate, Lab. brovkarbe eiRed. facemose Lan sopione spicata Lab ambigua Rat pbbosa I Leb might" Rat Musa Ca/. sampan, =_IJCW. Ammons Cat nopirlose L_IJ Cal. Cal. polo/Goa Munbengi UL Ca. cyasoide. Cm, ty0ende Cat saris. II Cal. canc. lolononoiebs CaL lotononokles Cal cinema Cal. acetate Cal halenbergensis Cal belembergensis • Cal spbestens Cal spews:es Cal Bahama Cal sabaroa Lot mosinsame 1Lot macrosama Lot Lot. Woofs Lot Indicordas Lot b=aa Let. widens La. Lot nochyanna Lot Intestines Lot abominate Lot spersillore Lot. aomtere Lot. oryptora Lot subsp. nameguonslo Lot =Orate subb. namapumaia Lot lentisula Lot Ionians La, mesulata Lot. menden, La mai Lot non Lot. lace Lot. taxa Lot. rnannsepala Lot. &bra La mina he Int macron:mob Lot plabra Lot. nestentia Lot sabulosa 1010111050 Lot.. bprolobe Lot. lopt01000 Lot. maxlmillani Lot. maximilianl Lot

FIG. 8.6 Parsimony-based reconstructions of the calyx type (character 5) on the 560 and

370 minimal length trees from (a) the combined molecular and (b) the combined molecular/morphological analyses, respectively.

254 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

ANTHER ARRANGEMENT—Anther dimorphism and arrangement are important within Crotalarieae and informative at the generic level (Chapter 3; Boatwright et al.,

2008a; Chapter 5; Boatwright et al., submitted). Pearsonia, Rothia and

Robynsiophyton are unique within the tribe in having uniform anthers that are similar in shape and size. The rest of the tribe have dimorphic anthers with alternating dorsifixed and basifixed stamens. The size, shape and attachment of the carinal anther is important and three arrangements are found: 5+5 (carinal anther

resembles the basifixed anthers) present in Bolusia, Crotalaria, Lebeckia and

Lotononis macrocarpa; 4+1+5 (the carinal anther intermediate in between the dorsifixed and basifixed ones) present in Aspalathus, Calobota, Lotononis s.s., the

Listia Glade, Rafnia and Wiborgia; 6+4 (carinal anther resembling the dorsifixed anthers) present in Lotononis hirsuta, the Leobordea Glade, Lotononis s.s. and

Wiborgiella. Anther arrangement is fairly diagnostic for the genera within the "Cape group", but the distinction within Lotononis s.l. is not as clear (Fig. 8.8).

VERRUCOSE UPPER SUTURE OF POD—A verrucose upper suture of the pod was thought to represent a generic apomorphy for Lotononis s.I., but this character was

reconstructed as a convergence between the Leobordea Glade, Lotononis s.s., L.

hirsuta and L. macrocarpa (Fig. 8.9).

FUNICLE LENGTH—Exceptionally long funicles are only found in species of

Lotononis s.l. On the trees from the combined molecular analysis this character was reconstructed as a convergence between several lineages (Fig. 8.10). In the combined molecular/morphological analysis it is reconstructed as an apomorphy for

Lotononis s.l., but it is also present in L. macrocarpa (Fig. 8.10).

CHROMOSOME BASE NUMBER—Goldblatt (1981) suggested a base number of x=9 for the tribe Crotalarieae, which seems to be likely from the reconstructions

255 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS presented here with a reduced number in some genera (Fig. 8.11). Crotalaria,

Rafnia and some Aspalathus species have a number of x=8, while some species of

Lotononis s.s., Pearsonia and Rothia have a base number of x=7.

CYANOGENESIS—Chemical data are of important systematic value in the

Crotalarieae (Van Wyk 2003b). Cyanogenesis was reconstructed as an apomorphy for Lotononis s.s. and is absent from the Leobordea and Listia clades, Lotononis hirsuta and L. macrocarpa (Fig. 8.12). The superficially similar genera Pearsonia,

Rothia and Robynsiophyton are acyanogenic and accumulate esters of hydroxylupanine, a unique character in the Crotalarieae.

MACROCYCLIC PYRROLIZIDINE ALKALOIDS—The presence of pyrrolizidine alkaloids in some sections of Lotononis was thought to suggest a sister relationship with Crotalaria that also produces such alkaloids (Van Wyk, 1991b). The presence of these alkaloids was reconstructed as a convergence between Lotononis s.s. and

Crotalaria in both the combined molecular and molecular/morphological analyses

(Fig. 8.13).

256

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CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

. enamels ❑ = normal length 1 A .arena A carnosa .seats A gale. = exceptionally long madvnpa A. rnacranthe —11-44A: A confusa A. porkilata gimp_ paipsi A polio:iota tatOP. PlolLOsi A. crenate A. drn venniculasa vernecteata ■ A. alseni 'atop. shared _7,-.7 :. "lanoTtgorsa sp.: Lerlorfoi a laaldroas mew. larIcaoaa artoeubso. tuna A. halo sutreo. larta Flaystrts laws A= CA.MO DeCtlabba fusee A pactortba Mb Leta Mo. Moaners sori:os Wit mac. M0 awn:1sta rnonootare rnonoplara Inarvata 1500 oboortlata mwtonata hunatte r Ines. Walt lk➢catiCana r itabl. ewes easstIctia iatelagana WV. batman,' staafectie NW. IMAM Nta boveasne IN& mucronate r Lab. peualtora Ls. comma Rd. Lab. rneyeriane R•fs Leb. plakenetiena Rat spice's Leo. peoceore Le. IMantii L—P— Trf. oefloss O. lesvicama .Lea canyon LM ambigua lab. 0,0yan.na Lab. set** Lab. plokenetan• Rat SOLIMASSO Lab. Movicarpa Rat raceme. Lab. aeolana Rat Ocala Lab. tontipsa Rat Oates. Lab wag,* Rat Whoa Cat pungens Cal Wog?. Cat a Cd. Cat a= Cal. mime CA" WA.] Cd lotorronoiclos Cat lolononoilea Cal. caw. Cd cam. Cat halanborpansis Cal hatenterpenan Cat spinescena Ca/ apinassana Cal seherse Ca/ sonatoo Lot rnsonOarpa Lai macron/pa Lot falcate ■ Lot falcate Lot. tnalcoake di arsolp.=t al Lot pervIttore li Lot arachyantha Lot fnrticoicles Lot sparasfiora Lot Span:Wont Lot oawaren Lot. coypeara Lot rostrata site° natnaquenais ot rostrata Subap. eamiloe.,a Lot lentecula Lot lentlaia Lot. moculata L ot Lot. coal Lot coal Lot. taro Lot. less Les. maaosepata Lot glabra Lot gnientnillatbra Lot. macrosepola Lot. g Lot. MAMA. Lot. madams Lot ...tom Lot leptolooa LOC lepaMbe Lot maxlmillanl Lot nveeinaliara Lot clivaricata Lot Poo f lona Lot. sarkophylfa Lot divaricata Lot. lotononoiles Lot. saricaohylle Lot putonella Let pu/cheaa Lot. awed 2.t.= ta odes Lot carnosa LotLo stn. Lot pawns ot oae Lot dame exesp. Paco.. Lot dense wimp teaoaclada Lot stelOtdra Lot. oaskoulata Lat involoontta Lot. ltmluctota Lot. involuerata Bubo. peclunCelada Lot koolocrata instep partuncutarte Lot. Worn* Lot. eleonnia Lot.P .oso MOM.. Lot Now. P. maaaabsa r Lot PRAYotalaa Lot calycloa Lot. rob's@ I4 cyaatda Lot. ensnare Lot eaprossa Ro0. ao:orwe' Lot. clactsnbons 'Lot ecutiotope i ci Pe,b aaian""a Lot. 0.0 Lot. teams Lt anerale Lot. Poteosphate Lot. perSaMlySs au.'"Came Lot. moats Lo Oecumens Lot. Waal Lot WIltnIens Lot. moms Lot acta. Lot plicate Lot. pun. 11= LapolyseplIala Lot. SOALEY. LotOontalAYIL, Lot sod!. Lotmagolaca Lot Lotbentharniana Lot. manatee Lot &Vete Lct globule= P. erhtab La solludnis Lot mandato Lot real Rol. his. Lot rnallotall xl La. needs I, tot Irks. . tlbme Cr cf.. gfintentsis Cr gnouensta Cr wefts =-1.1 Cr composts viroultaas Cr. virpotteas Cr. holies Cr. human l amed. t , Cr. tan ease artrOoensle x113 arnt LID radii, 0 gracile

FIG. 8.10 Parsimony-based reconstructions of the funicle length (character 9) on the

560 and 370 minimal length trees from (a) the combined molecular and (b) the combined molecular/morphological analyses, respectively.

260 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

= base number 7 Peolt =1.= M' A. cam. =IC .nose A gabote A. gaboto 0 = base number 8 A mecronthe A. macranthe A. tbadenowlena A scamp A. conlose"a'''''" 1 = base number 9 A. pedals. WM. PnbbPtd A. Pastes subs', 000s, aeon. A. crenate A VOX MiCale8 A rennictAbla A. Mang alloop. A staled subso sherd A brio:ben sistop. Parkiblia AA. la''aaArro sattor. RateP *1'1'5' A nine outdo tote A SAA.Ayserb

A pachybbe A. pLprybba INID. baba W. totraptere Isesptara Wt. sada. bib steno. Wel °boor.. WO. monoptere Mb. monopoles Wed. ilcsrofe incurvata Wes. pdmrdad mudone Mg. baste es bild. MM. Sunda sessafofia Wd2/. loipoelorto WU boomer. sesstione W.N. Melo. Wed bow.. Witt ent.onate Lab. pouring. Ras won.. Re. racemes LYE. pbkarMare Rol. std.. rRef.I &Wu ob. nig,. L Rol Muse ob. ...goo SO. replan& meTh.Y..."., LeOplukonetiana awnbaa Led. Led. '""ana at sp1.1. Lab. magus Lab. mighti CO. 1.07ans Cat puno,n3 = t"

Cat seises ca. wooed Cal. btononobles ■ Cal. lotononosies Cal come Col. cinema Cal baktnbenrom CaL ludenbotganes Cot oPiOSsCons spinescens Cot anhemo Cal. sehamo to moaccame Lot 1718a0004. Lot falcate Lot retch Lot h.lcoides Lot. becnyentrta LoL pervinora Lot penacre L. braddarobe tot Ineacados Lot. spandore Lot. spe.figra lat OAYP,.. tht Lot ro.sto aubsp. nooloao...4 Lot to.O. subsp. nomnpuensb Lot. len.. Lot baba. Lol. macubto Lot mocutata Lot maul Lot curb L. b. Lot la. L. macros.. Lot globe Lot micron". Le a. macrosepals Lot (dab. Lot. nermenthe Lot .bubse I. 6.10. Lot Oopro1o0a Lot leptobbe L. med... LOL Lot drama. Lot pungens Lot. dead.. El— ''‘''. I= Lot. sarko.rybo Lot. p✓cnolle LoL butcher:la Lot = Lot awed Lot boononodes Lot camp. L. dncte I.. ',Lawn Lot *arose Lot. dense sub. ioueociado Lot dance sub. Wow/ads Lot ...Opole. ffi.:= Lot Inmexrata Lot bedtimes &bap. parimares Lot. emits:tote sub*. pedurrembfis 1--1 1._—_,.., Lot fildonnis tadorrals Lii Lot. elongate elongate P sesslefona Lot platycame P mtioto Lot ce/ycino P. gronddolle Lot lobo. P. ...bee L. Menthe Rot. hirsute Lot Depress& Rob. vandetystil Lot decorate. Lot Potycoom LoL acuticama lot LoL Dodo. Lot. Mime Lot p.n. Lot. atone.. La. polycephola Lot adontsso Let pomades Lot ecuticarpo Lot mobs t otdectanbens Lot qt. Lot. Doane Lot bentnandena L.. mobs Lot digitate Lot pedalo a. mamAta Lot. pub. la bbrdlse Lot pOrycepattla saeweu Lot pentaprryte Lot WARMS Lot. magnifica Lot Fs. Lot benthondana Lot mortal., Lot diger. sesseboth !Lot glob.ose ,Aanstftit. Lot. soleudids Lot sub.. cab Lot Ada how. Lot mattond Rob. wedorysffi Lot S... tot hirsuro _ ., C.r. Lesbos Cr (Mammas Cr omens., Cr. Legueobs Cr hurne7b .0 kroLvolsto t1Cr. langeolata a emboonsie ID padre OrOCSS'nb'M''

FIG. 8.11 Parsimony-based reconstructions of the chromosome base number

(character 10) on the 560 and 370 minimal length trees from (a) the combined molecular and (b) the combined molecular/morphological analyses, respectively.

261 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

A. Ilnaara = absent 11.=, pond. A. comm. A. A. golooto nA. polo. 1111 = present A. macro.* A. aditdonowiona 00,09. ± 0. porfolato .69*. WA:112a A. coma. AA mango A. von:Luba vamiculoto

aAdkC armAOYYa . Oita wasp. larto A larto su top. hob A hystria A bytes n 0300nruO,.c. A. Poonytobtt 0:87 bag r,=1140 notraptera IA4b. totroptana 1__LINALr. "now W.D. sonar PAO. otroccilata mongstora monomoro INIb. bauveba obc"'rolo"TO muoonoto tomails MO Ababa IONIA loosoldliana 142. lamas JASa. 6eamr0M 6.93L ia,polabono ARM. bovnoono sossettn• WItaL Odom Wprt traofieffnat .,W61 mamma Lob poocelont Rol barLOoto Rot ra.00aa 1- 11'thob rA71 " Rat spostS Rot ansbaaO O Omuta/pa RoLab. rno asa b ambigua Lob. mayenona PA.10.300 • W'd'oeurninato bornorpa racornosa Lob. sepioria op/181a . ambigua 91.0. In VIA*.1 ((800 RION Whoa GA porsoons Cot pungens

get Ca ~Da I A Cal cm= Cot sot. cat lotononottat szcoglil iizo.c."lotonono:.,.:n. „.: Cat ammo Ca/ bolanacovonsis

st =home ,amebae 1 Lot rnacmcorpo LCol rnacmca Lot talc. u‘tot lama Lot fruttooklos Lot parabola Lot parAbora Lat. dada... Lot aubcoitios LOt Mao:Cora LOt var./on .YPtaro Lot rostrata autrap. narnaguensis 18180'6 sub.. namorpronsis Lct lenbant Lot lenbudo LaMal. Lot moculoto Loa Lot cat Lot Ia. LO. moctosepata Lot aroon. Lot1. ,,,,,,...,.91fn":crosopolo • Lot plabra Lot. mierantba Lat. madam Lot sobuloso 100.101:. Lot loptOobo Lot. mazineloo Lot frOOOTIFOSOi Lot dram. MIN ot oadCapN00 Lot divaOcata Lot btonomatas La. soraophylla La pulchella Lot pukheAs Lo. most Lat. Orsto tn: Ictononoilam"rM s Lot samosa 4■1 Lot ark. Lot panacea Lot bansa sub.. taucoclodo LO. dans, sub.. Macao/ado 1.:t. 47:dab

Lat. imolutrota Wm. potanc.ona LOt. a sub. pi...40ra La. 00060r63 Lat.03 baron. Lot *bowl. P. seuilifolla P. anstoto AL c*6''todosa gaiVrel: Lot Rd. laratOo Lot 4.0ressa Rob. varularyall Lot Oa...as tot plabroabo Lot bonkarba Lot C611.18 L01.11OIUSO tat lobos. Lot auto. L. ...ha 1 Lot adores,. UAL' pentaphylll Lai welcome Lot. molls Lot documboos La ado" Lot.=Mana La. malls Lot Olgazda Lat. p.m COIL' poq''.41pbola Lot xerea00. Lot subutoto Lot. manners Lot tali Lot. bantharniana Lot mance* Lot Makes Ej--0, 1;sosoillra is s0 0 1 La. subutota copnbatio Of. /111.111 Lot madothil Rob. vandarystii Lot bus. al Lot tomato Cr. distant ___.., Cr tfttarii Onbuonsts CC WWWISIS b. copensts r .LL .li g capons. vmultall Cr. Monks Cr. humks Cr lanced. ■ .Cr lona.. 8 arnbconsis , 8 amboanso comogla D was&

FIG. 8.12 Parsimony-based reconstructions of the presence or absence of cyanogenic glucosides (character 11) on the 560 and 370 minimal length trees from

(a) the combined molecular and (b) the combined molecular/morphological analyses, respectively.

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II r CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

8.3.2 Lotononis hirsuta— Lotononis section Euchlora is only distantly related to the rest of Lotononis. It is placed within a Glade comprising

Crotalaria and Bolusia (100 BP) and this Glade represents the earliest diverging lineage within the tribe. The placement of Lotononis hirsuta in this

Glade is supported by the strongly inflated pods and trifoliolate leaves (if

present) with paired stipules that are equal in size found in these genera.

Dahlgren (1964) transferred this species to Lotononis where it was treated as

a section by Van Wyk (1991b). This anomalous plant differs markedly from

species of Lotononis s.s., the Leobordea Glade and the Listia Glade by the

large underground tuber (geophytic habit), simple, sessile leaves (in some

forms), large pods and the subequally lobed calyx. The position of Lotononis

hirsuta renders two important generic apomorphies for Lotononis s.l., to be

the result of convergence, namely the loss of bracteoles and the verrucose

upper suture of the pod (Van Wyk, 1991b).

8.3.3 Lotononis s.s.—Although the resolution is low in Lotononis s.s.,

some sectional relationships were observed. Lotononis section Oxydium (BP

98) is sister to a Glade with representatives of L. sections Aulacinthus,

Buchenroedera, Cleistogama, Krebsia, Lotononis, Monocarpa and

Polylobium. Lotononis section Oxydium is a large group comprising 35

species, and is shown to be monophyletic. This section is unique within the

Lotononis s.s. in the chromosome base number of x=9 (the rest have x=7),

the presence of long funicles and also its distribution throughout tropical

Africa. The sections Monocarpa and Cleistogama are sister to each other as

was also shown in the chosen cladogram of Van Wyk (1991b) in Fig. 8.14

where this combined Glade was also shown to be sister to the section

264 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Oxydium. However, Van Wyk (1991b) concluded that the sections Monocarpa and Cleistogama have their real affinities within the larger group of sections around sections Polylobium and Aulacinthus and not Oxydium. The shared presence of long funicles and a chromosome base number of x=9 in L. section Monocarpa is reconstructed as the result of convergence rather than common ancestry in Figs. 8.10 and 8.11. Lotononis section Buchenroedera is embedded within L. section Krebsia, albeit with low support. These two sections have paired stipules, a character shared only by L. sections

Polylobium and the Listia Glade. Lotononis sections Aulacinthus and

Polylobium are weakly supported to be sister groups, the latter differing from section Polylobium in its single stipules and shrubby habit (Van Wyk 1991b).

With the exception of section Oxydium, Lotononis s.s. is endemic to southern

Africa. Two sections are mainly found in the eastern parts of southern Africa, namely sections Krebsia and Buchenroedera, while all the remaining sections are restricted to the Cape and Namaqualand. Lotononis section Monocarpa occurs in the north-western Cape, while sections Lotononis, Aulacinthus and

Polylobium are more or less restricted to the Western Cape Province.

Lotononis section Cleistogama has a more eastern distribution in the Cape region (Van Wyk 1991ba). Lotononis s.s. is chemically distinct from the

Leobordea and Listia clades in that its members are cyanogenic (except for L. section Cleistogama) and accumulate macrocyclic pyrrolizidine alkaloids, the first of which is a synapomorphy for this group.

8.3.4 The Leobordea Glade—The Leobordea Glade comprises L. section Leobordea and its apparent relatives from L. sections Digitata, Leptis,

Lipozygis, and Synclistus. This Glade is well-supported as sister to the Listia

265 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Glade as was also shown by Van Wyk (1991b) in his chosen cladogram (Fig.

8.14). The Leobordea Glade shares with the Listia Glade (and differs from

Lotononis s.s.) the rounded keel petals, acyanogenesis and a chromosome base number of x=9, all pleisiomorphic states. The two groups differ in the non-stoloniferous habit, single stipules that are sometimes similar to the leaflets, pubescent vegetative and reproductive parts, absence of bracteoles

(except in two species) and 4+6 anther arrangement (very rarely 4+1+5) found in the Leobordea Glade. The latter group is also characterised by its wide distribution range throughout southern and tropical Africa. Sections

Leptis and Leobordea extend into the Mediterranean region of Africa, with L. genistoides (section Leptis) extending into Europe and L. platycarpa (section

Leobordea) extending into Pakistan and the Cape Verde Islands. Sections

Synclistus and Digitata both occur in the north-western Cape, while section

Lipozygis occurs in the eastern parts of southern Africa (Van Wyk 1991b).

8.3.5 The Listia Glade—The Listia Glade is a very distinct group with a unique combination of characters: stoloniferous habit, paired stipules, presence of bracteoles and a 4+1+5 anther arrangement (Van Wyk 1991b).

The concept of Meyer's (1836) monotypic genus Listia was broadened by Van

Wyk (1991b) and included as a section of Lotononis. It is interesting to note that the species of Listia have lupinoid (sleeve-like) root nodules (as are also found in the genus Lupinus) and not the conventional types that are present in all other species of Lotononis (and indeed all Crotalarieae) hitherto investigated (Yates et al. 2007). The species of this group are distributed throughout southern and tropical Africa.

266 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

8.3.6 Lotononis macrocarpa— Lotononis macrocarpa was included in

Lotononis section Listia as a distinct subsection (subsection Macrocarpa) by

Van Wyk (1991b) based mainly on the presence of bracteoles and superficial similarities with the other species in Listia that were taken at face value.

However, this anomalous species is unique in the 5+5 anther arrangement, equally lobed calyx, large fruit and systematic placement close to the "Cape group" of the Crotalarieae demonstrated in this study. It is endemic to the south-western Cape and geographically isolated from the rest of the Listia group. Based on the unique combination of characters found in this plant and the fact that it allies with the "Cape group" in the molecular study, it is here recognized as a monotypic genus.

8.3.7 Generic circumscription—The close agreement of the relationships within Lotononis s.s. found in Chapter 3 (Boatwright et al. 2008a) and this study with those proposed by Van Wyk (1991a) is notable (Fig. 8.14).

The only difference is the inclusion of L. section Euchlora in Lotononis and the sister relationship proposed between section Oxydium and sections

Cleistogama and Monocarpa. The cladogram from Van Wyk (1991b), based on vegetative and reproductive morphological, cytological and chemical characters (Fig. 8.14), also shows Lotononis section Listia (the Listia Glade) as sister to L. sections Digitata, Leobordea, Leptis, Lipozygis, and Synclistus, the Leobordea Glade. This Glade is sister to L. sections Aulacinthus,

Buchenroedera, Cleistogama, Euchlora, Krebsia, Lotononis, Monocarpa,

Oxydium and Polylobium, i.e. Lotononis s.s. (excluding section Euchlora).

Dahlgren (1970a) discussed some examples of convergence and parallelisms in the tribe, but the results presented in this study have

267 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS uncovered that the relationships within the Crotalarieae are even more complex (Chapter 3; Boatwright et al. 2008a; Chapter 5; Boatwright et al. submitted) and revealed more examples of convergent evolution of morphological and chemical characters. Characters thought to be unique to certain groups, especially Lotononis s.I., have proven to be shared by distantly related groups. Due to the extreme overlap of character states in the genera of the Crotalarieae, unique generic apomorphies are frequently not available and a combination of characters is necessary for generic circumscription. The original generic concept of Lotononis s.l. is here shown to be polyphyletic, largely due to the positions of L. macrocarpa and L. hirsuta and also the paraphyly of Lotononis s.l. shown by combined molecular

evidence. Despite weak support for the monophyly of Lotononis s.l. in the combined molecular/morphological analysis a narrower concept of Lotononis

can be better circumscribed in light of the lack of generic apomorphies for

Lotononis s.l. The recognition of smaller, monophyletic groups is now more

practical and therefore the reinstatement of Euchlora, Leobordea and Listia is

proposed and Ezoloba described as a new genus to accommodate Lotononis

macrocarpa. Unique combinations of characters are available to circumscribe these genera that are strongly supported as monophyletic in the phylogenetic

analyses presented in this Chapter and Chapter 3 (Boatwright et al. 2008a).

268 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Sect. Listia

Sect. Digitata

Sect. Lipozygis

Sect. Leptis

Sect. Leobordea

Sect. Synclistus

Sect. Oxydium

Sect. Monocarpa

Sect. Cleistogama

Sect. Euchlora

Sect. Polylobium

Sect. Lotononis

Sect. Aulacinthus

Sect. Krebsia

Sect. Buchenroedera

FIG. 8.14 Chosen cladogram (Fig. 17) from Van Wyk (1991b) based on morphological, cytological and chemical data.

269 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

8.4 TAXONOMIC TREATMENT

8.4.1 Key to the genera of the Crotalarieae

1. Stipules absent (if stipules rarely present then leaves acicular and keel

spirally twisted) 2

2. Calyx zygomorphic (upper and lateral lobes on either side fused

higher up in pairs) Lotononis (partly)

2. Calyx subequally lobed 3

3. Style with 1-2 lines of hairs Crotalaria

3. Style glabrous 4

4. Bracteoles absent, leaves (at least the basal ones)

simple, flat and sessile; geophyte with large underground

tuber Euchiora

4. Bracteoles present, leaves (is simple and flat) not

sessile; annuals, suffrutices or shrubs 5

5. Leaves acicular, terete 6

6. Ovary 2 to 4 ovulate, pods 1 to ±2-seeded Aspalath us

Ovary with more than 6 ovules, pods many-seeded..Lebeckia

5. Leaves digitate, unifoliolate or simple (flat, never terete) 7

Upper suture of pod asymmetrically convex 8

Plants glabrous except occasionally on bracts and

bracteoles, usually turning black when dried Rafnia

270 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Plants usually pubescent on all parts, if leaves

glabrous then standard petal hairy and inner surface of

calyx glabrous Aspalathus

7. Upper suture of pod symmetrically convex 9

Petals pubescent, if glabrous then plants strongly

spinescent shrubs; twigs green (bark formation late);

leaves isobilateral Calobota

9. Petals glabrous; twigs brown (bark formation early), if

twigs rarely green then plant an annual fireweed; leaves

dorsiventral 10

Fruits winged, indehiscent; carinal anther

intermediate (anthers 4+1+5) Wiborgia

10. Fruits without wings, dehiscent (if rarely

indehiscent then ovary and fruit distinctly stalked);

carinal anther resembles short anthers (anthers

4+6) Wiborgiella

1 Stipules present 11

Style straight or rarely down-curved, anthers similar in size and

shape 12

Stamens nine (five fertile and four lacking

anthers) Robynsiophyton

12. Stamens 10 (all fertile) 13

Anthers monomorphic, prostrate annuals ...... Rothia

271 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

13. Anthers slightly dimorphic, four basifixed, six attached

slightly higher up, all elongate, perennial herbs or

shrubs Pearsonia

11. Style curved upwards, anthers dimorphic 14

14. Bracteoles absent 15

15. Geophyte with woody tuber, leaves (at least the basal

ones) flat, simple and sessile Euchiora

15. Annuals, suffrutices, shrubs, leaves is flat and simple

then not sessile 16

16. Stipules paired or absent 17

17. Keel obtuse, hairy Leobordea

Keel beaked, glabrous Lotononis

16. Stipules dimorphic or single 18

Keel obtuse, hairy Leobordea

18. Keel beaked, glabrous Lotononis

14. Bracteoles present 19

19. Keel obtuse 20

20. Calyx subequally lobed; fruit more than 20 mm

long; stems without adventitious roots; seeds ± 4

mm long; anthers 5+5 Ezoloba

Calyx zygomorphic (upper and lateral lobes on

either side fused higher up in pairs); fruit less than

20 mm long; stems often with adventitious roots;

seeds ± 1 mm long; anthers 4+1+5 . Listia

Keel beaked or helically coiled 21

272 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

21. Keel and style helically coiled through several

turns Bolusia

21. Keel and style not helically coiled Crotalaria

8.4.2 Taxonomy

1. EZOLOBA B.-E.van Wyk and Boatwr., gen. nov., similar to Listia E.Mey. but

differs in the equally sized stipules, sub-equally lobed calyx, 5+5 anther

arrangement, large pods (more than 20 mm long) and seeds (more

than 4 mm long) and the non-stoloniferous habit (to be translated into

Latin).—TYPE species: Ezoloba macrocarpa (Eckl. and Zeyh.) B.-

E.van Wyk and Boatwr.

The generic concept proposed here is based on a unique combination

of characters, namely the stipules that are equal in size, the sub-equally lobed

calyx, presence of bracteoles, 5+5 anther arrangement, the very large fruit

with a warty upper suture and the large seeds. The molecular and

morphological evidence presented here indicates that this species is more

closely related to the "Cape group" of the Crotalarieae than to Lotononis s.l.

The single species is known from only a few localities in the Western Cape

Province of South Africa. The generic name commemorates Ecklon and

Zeyher who first described this species.

EZOLOBA MACROCARPA (Eckl. and Zeyh.) B.-E.van Wyk and Boatwr., comb.

nov. Lotononis macrocarpa Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 176.

273 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

1836.—TYPE: SOUTH AFRICA, Cape Province, Brackfontein, Clanwilliam,

Ecklon and Zeyher 1271 (lectotype: S!, designated by Van Wyk [1991b];

isolectotypes: C!, S!, SAM!, isolecto.).

2. EUCHLORA Eckl. and Zeyh., Enum Pl. Afr. Austr. 2: 171. 1836. Microtropis

E.Mey., Comm. Pl. Afr. Austr. 1(1): 65. 1836. Lotononis section

Euchlora (Eckl. and Zeyh.) B.-E.van Wyk, Contr. Bol. Herb. 14: 213.

1991b.—TYPE species: Euchlora serpens (E.Mey.) Eckl. and Zeyh.

The genus is easily recognisable by the large underground tuber, simple, sessile leaves (in some forms) and the large, inflated pods. The

results presented here indicate that this genus forms part of the early diverging elements of the Crotalarieae and is closely related to Bolusia and

Crotalaria. The single species occurs in the Northern and Western Cape

Provinces of South Africa.

EUCHLORA HIRSUTA (Thunb.) Druce in Rep. Bot. Exch. Cl. Brit. Isles 1916: 622.

1917. m Ononis hirsuta Thunb., Prodr. Pl. Cap.: 129. 1800. Microtropis hirsuta

(Thunb.) E.Mey., Comm. Pl. Afr. Austr. 1(1): 65. 1836. Lotononis hirsuta

(Thunb.) D. Dietr., Syn. Pl. 4: 960. 1847.—TYPE: SOUTH AFRICA, Cape

Province, `grope Cap juxta Leuwestaart', Thunberg s.n. sub THUNB-UPS

16614 (lectotype: UPS!, designated by Dahlgren [1964]).

Crotalaria serpens E.Mey. in Linnaea 7: 153. 1832. Euchlora serpens

(E.Mey.) Eckl. and Zeyh., Enum Pl. Afr. Austr. 2: 171 .1836. Lotononis serpens (E.Mey.) Dahlgr. in Bot. Not. 117: 373. 1964. nom superfl.—TYPE:

274 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

SOUTH AFRICA, Cape Province, `Sandige Stellen unweit Salzrivier', Ecklon s.n. (lectotype: S!, designated by Dahlgren [1964]).

3. LISTIA E.Mey., Comm. Pl. Afr. Austr. 1(1): 80. 1836. Lotononis section Listia

(E.Mey.) B.-E.van Wyk, Contr. Bol. Herb. 14: 99. 1991b.—TYPE

species: Listia heterophylla E. Mey.

The concept of Listia proposed here conforms to that of Lotononis section Listia subsection Listia as described by Van Wyk (1991b), which includes seven species. The genus can be distinguished from especially

Ezoloba, Lotononis s.s. and Leobordea by a unique combination of characters: stoloniferous habit, paired, dimorphic stipules, presence of bracteoles, 4+5+1 anther arrangement, largely glabrous petals and pods, the latter often folded like a concertina. The root nodules differ from all other members of the Crotalarieae in being lupinoid (sleeve-like). The species mainly occur in the interior of southern Africa, but L. angolensis and L. heterophylla extend into central Africa.

LISTIA ANGOLENSIS (Welw. ex Bak.) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis angolensis Welw. ex. Bak. in Oliv. Fl. Trop. Afr. 2: 6. 1871.—TYPE:

ANGOLA, Huilla District, Welwitsch 1896 (lectotype: BM!, designated by Van

Wyk [1991b]; isolectotypes: BM!, C!, K!); Angola, Pungo Andongo District,

Welwitsch 1895 (BM!).

275 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LISTIA BAINESII (Bak.) B.-E.van Wyk and Boatwr., comb. nov. Lotononis bainesii Bak. in Oliv., Fl. Trop. Afr. 2: 6. 1871.—TYPE: In the interior near the

Tropic of Capricorn, Chapman and Baines s.n. (holotype: K!).

LISTIA HETEROPHYLLA E.Mey., Comm. Pl. Afr. Austr. 1(1): 81. 1836. Lotononis listii Polhill in Bot. Syst. 1: 324. 1976.—TYPE: SOUTH AFRICA, Cape

Province, Gaatje, Drege s.n. a (lectotype: K!, Herb. Benth. specimen, designated by Van Wyk [1991b]; isolectotypes: BM!, K!, Herb. Hook. specimen, S!); Cape Province, `grope Wild. schutshoek', Drege s.n. b (BM!,

K!, S!).

LISTIA MARLOTHII (Engl.) B.-E.van Wyk and Boatwr., comb. nov. Lotononis

marlothii Engl. in Engl., Bot. Jahrb. 10: 26. 1888.—TYPE: SOUTH AFRICA,

Cape Province, Griqualand West, Kimberley, Marloth 765 (lectotype: K!, Herb.

Engl. specimen, designated by Van Wyk [1991b]; isolectotypes: BOL!, K!,

PRE!).

LISTIA MINIMA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov. Lotononis minima B.-E.van Wyk in S. Afr. J. Bot. 54(6): 628. 1988.—TYPE: SOUTH

AFRICA, Cape Province, Kenhardt Div., Jagbult, floor of Uilpan, Acocks

12664 (holotype: PRE!; isotype: K!).

LISTIA SOLITUDINIS (Dummer) B.-E.van Wyk and Boatwr., comb. nov. Lotononis solitudinis Diimmer in Trans. Roy. Soc. S. Afr. 3(2): 297. 1913.—TYPE:

SOUTH AFRICA, Vaal River, Wilms 400 (holotype: BM!).

276 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LISTIA SUBULATA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis subulata B.-E.van Wyk in Bothalia 20(1): 79. 1990. —TYPE:

SOUTH AFRICA, Transvaal, Parys, near bridge over Vaal River on

Potchefstroom Road, B.-E. van Wyk 2884 (holotype: PRE!; isotypes: JRAU!,

K!, MO!, 5!).

4. LEOBORDEA Del. in Laborde, Fragm. fl. Arabie Petree (1830) emend. B.-

E.van Wyk and Boatwr., emend nov. Lotononis section Leobordea

(Del.) Benth. in Hook., Lond. J. Bot. 2: 607 (1843).—LECTOTYPE

species: Leobordea lotoidea Del. [Lotononis platycarpa (Viv.) Pic.–

Serm.].

Leptis E.Mey. ex Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 174 (Jan. 1836).

Lotononis section Leptis (E.Mey. ex Eckl. and Zeyh.) Benth. emend.

B.-E van Wyk in Contr. Bol. Herb. 14: 124. 1991b.—LECTOTYPE

species: Leptis debilis Eckl. and Zeyh. [Lotononis prolifera (E.Mey.) B.-

E.van Wyk].

Lotononis section Digitata B.-E.van Wyk, Contr. Bol. Herb. 14: 107. 1991b.—

TYPE species: Lotononis digitata Harv.

Lipozygis E.Mey., Comm. Pl. Afr. Austr. 1(1): 80 (1836, Feb. or later)

Lotononis section Lipozygis (E.Mey.) Benth. emend. B.-E.van Wyk,

Contr. Bol. Herb. 14: 114. 1991b.—LECTOTYPE species: Lipozygis

corymbosa E.Mey. [now Leobordea corymbosa (E.Mey.) B.-E.van Wyk

and Boatwr.].

277 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

Lotononis section Synclistis B.-E.van Wyk, Contr. Bol. Herb. 14: 157.

1991b.—TYPE species: Lotononis longicephala B.-E.van Wyk

[Leobordea longicephala (B.-E.van Wyk) B.-E.van Wyk and Boatwr.].

[Note: Leobordea is here reinstated and its concept broadened to include

Lotononis sections Digitata, Lipozygis, Leptis, Leobordea and Synclistus. Of the names available for the generic concept proposed here, Amphinomia is the oldest. This genus was described by De Candolle (1925) based on a plate

of Connarus decumbens Thunb. in Roem. Arch. Bot. 1, 1: 1796. The identity

of this species is, however, has until now been unclear. Gillet and Bullock

(1957) mentioned that Connarus decumbens might not even be a legume and that Amphinomia is possibly not a synonym of Lotononis. A reinvestigation of the type specimen of Connarus decumbens in UPS has provided clarity

regarding the identity of this plant. The unusual vestiture of the calyx (hairs in

rows along the veins) and the short, few-seeded pods of the specimen are

diagnostic characters of Lotononis pallens (Eckl. and Zeyh.) Benth. In light of this discovery, Amphinomia in actual fact represents a synonym of Lotononis

s.s. Since Amphinomia has already been rejected in favour of Lotononis and

Leobordea (Rickett 1960; Lanjouw et al. 1966), Leobordea is thus the oldest

available name that can here be considered for reinstatement.]

The genus Leobordea now includes 51 species that occur mostly in the

eastern parts of South Africa and extend into tropical Africa and the

Meditteranean region. They can be distinguished from especially Listia by the

habit that is never stoloniferous, the single stipules that are sometimes similar to the leaflets, pubescent vegetative and reproductive parts, abscence of

278 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

bracteoles (except in two species) and 4+6 anther arrangement (very rarely

4+5+1). After Lotononis s.s. which now comprises 90 species, this is the

largest of the genera proposed here. The species have been classified into distinct sections by Van Wyk (1991b). The number of species sampled for the

molecular studies does not allow for a proper evaluation of the infrageneric classification system. Pending further studies, the species are simply listed

alphabetically below. See Van Wyk (1991b) for complete synonymies as well

as keys to the sections and species. Fortunately, the nomenclature of the 90

species remaining in Lotononis s.s. are not affected (see Van Wyk 1991b).

LEOBORDEA ACUTICARPA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb.

nov. Lotononis acuticarpa B.-E.van Wyk in Bothalia 20: 21. 1990. TYPE:

SOUTH AFRICA, Gauteng, Springs district, 5 km from Devon radar station to

Leandra, Van Wyk 1815 (holotype: PRE!; isotypes: JRAU!, K!, MO!, NBG!).

LEOBORDEA ADPRESSA (N. E. Br.) B.-E.van Wyk and Boatwr., comb. nov.

Two subspecies are recognised:

LEOBORDEA ADPRESSA (N. E. Br.) B.-E.van Wyk and Boatwr. subsp.

ADPRESSA. Lotononis adpressa N. E. Br. subsp. adpressa. Lotononis adpressa

N. E. Br. in Kew. Bull. 1906: 18. 1906.—TYPE: SOUTH AFRICA, Natal, Stony

hill near Charlestown, Wood 5712 (holotype: K!).

LEOBORDEA ADPRESSA (N. E. Br) B.-E.van Wyk and Boatwr. subsp.

LEPTANTHA B.-E.van Wyk. Lotononis adpressa N. E. Br. subsp. leptantha B.-

279 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

E.van Wyk in Contr. Bol. Herb. 14: 141. 1991b.—TYPE: SOUTH AFRICA,

Gauteng, Farm Waterval, 2 miles [3.2 km] WNW of Krugersdorp, Mogg 22844

(holotype: PRE!; isotypes: PRE!, 2 sheets).

LEOBORDEA ANTHYLLOIDES (Harv.) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis anthylloides Harv. in Harv. and Sond., Fl. Cap. 2: 59. 1862.

Lotononis anthyllopsis B.-E.van Wyk in Contr. Bol. Herb. 14: 164. 1991b.—

TYPE: SOUTH AFRICA, Cape Province, Namaqualand, Wyley s.n. (holotype:

TCD; isotypes K!, S!).

LEOBORDEA ARIDA (DCImmer) B.-E. van Wyk and Boatwr., comb. nov.

Lotononis arida Durnmer in Trans R. Soc. S. Afr. 3(2): 324. 1913.—TYPE:

SOUTH AFRICA, Cape Province, mountain tops, Eland's Hoek near Aliwal

North, F. Bolus 31 sub BOL 10559 (lectotype: K!, designated by Van Wyk

[1991b]; isolectotype: BOL!).

LEOBORDEA BENTHAMIANA (Durnmer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis benthamiana DOmmer in Trans. Roy. Soc. S. Afr. 3(2): 294.

1913.—TYPE: SOUTH AFRICA, Western Region, Little Namaqualand: Near

Ookiep, Scully s.n. sub Herb. Norm. Austr. Afr. 1127 (lectotype: K!, designated by Van Wyk [1991b]; isolectotypes: BOL!, K!), Scully 150

(isosyntype: BM!), Morris s.n. sub BOL 5622 (isosyntypes: BOL!, K!);

Steinkopf, Schlecther 39 (isosyntypes: BM!, BOL!, GRA!, MO!).

280 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA BOLUS!' (Diimmer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis bolusii Demmer in Trans. Roy. Soc. S. Afr. 3(2): 306. 1913.—

TYPE: SOUTH AFRICA, Cape Province, near Piquetberg, Bolus 8431

(lectotype: K!, designated by Van Wyk [1991b]; isolectotype: Z).

LEOBORDEA BRACTEOSA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis bracteosa B.-E.van Wyk in Bothalia 20: 73. 1990.—TYPE:

NAMIBIA, Outjo district, mountains 14 miles [22.4 km] east of Torra Bay,

Giess, Vlok and Bleissner 6198 (holotype: PRE!; isotypes: M!, PRE!, WIND!).

LEOBORDEA BULLONII (Emberger and Maire) B.-E.van Wyk and Boatwr., comb. nov. Lotononis bullonii Emberger and Maire, Pl. Marocc. Nov. (Arch. Sc.

Maroc.) Fasc. 1: 1. 1929.—TYPE: not seen.

LEOBORDEA CARINATA (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov.

Lipozygis carinata E.Mey., Comm. Pl. Afr. Austr. 1(1): 80. 1836. Lotononis carinata (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 609. 1843.—TYPE: SOUTH

AFRICA: North-eastern Transkei, between `Omsamculo' and `Umcomas',

Drege s.n. (lectotype: K!, designated by Van Wyk [1991b]; isolectotypes: MO!,

S!).

LEOBORDEA CORYMBOSA (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov.

Lipozygis corymbosa E.Mey., Comm. Pl. Afr. Austr. 1(1): 80. 1836. Lotononis corymbosa (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 605. 1843 – Lectotype:

281 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

South Africa, Transkei, `grope Omtata', DrOge s.n. (lectotype: S!, designated by Van Wyk [1991b]; isolectotypes: BM!, K!, 2 sheets, M!, 5!).

LEOBORDEA DECUMBENS (Thunb.) B.-E.van Wyk and Boatwr., comb. nov.

Two subspecies are recognised:

LEOBORDEA DECUMBENS (Thunb.) B.-E.van Wyk and Boatwr. subsp.

DECUMBENS. Ononis decumbensThunb., Prodr. Pl. Cap.: 129. 1800.

Lotononis decumbens (Thunb.) B.-E.van Wyk subsp. decumbens. —TYPE:

SOUTH AFRICA, `Roggeveld', Thunberg s.n. sub THUNB-UPS 16604

(lectotype: UPS!, designated by Van Wyk [1991b]).

LEOBORDEA DECUMBENS (Thunb.) B.-E.van Wyk and Boatwr. subsp.

REHMANNII (Dummer) B.-E.van Wyk, Contr. Bol. Herb. 14: 139. 1991b.

Lotononis rehmannii DOmmer in Trans. R. Soc. S. Afr. 3(2): 326. 1913.

Lotononis decumbens (Thunb.) B.-E.van Wyk subsp. rehmannii (Dummer) B.-

E.van Wyk, Contr. Bol. Herb. 14: 139. 1991b.—TYPE: SOUTH AFRICA,

Gauteng, `Rogge Veld, Perekopberg', Rehmann 6831 (lectotype: K!, designated by Van Wyk [1991b]; isolectotypes: BM!, Z).

LEOBORDEA DIFFORMIS (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis difformis B.-E.van Wyk in S. Afr. J. Bot. 55(5): 529. 1989.—TYPE:

SOUTH AFRICA, Gauteng, Piet Retief District, Iswepe, Sidey 1609 (holotype:

PRE!; isotype: S).

282 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA DIGITATA (Harv.) B.-E.van Wyk and Boatwr., comb. nov. Lotononis digitata Harv. in Harv. and Sond., Fl. Cap. 2: 52. 1862.—TYPE: SOUTH

AFRICA, in some part of the eastern provinces, Capt. Carmichael s.n.

(holotype: TCD!).

LEOBORDEA DIVARICATA Eckl. and Zeyh., Enum. Pl. Afr. Austr.: 175. 1836.—

TYPE: SOUTH AFRICA, Cape Province, side of Bothasberg, not far from

Vischrivier', Ecklon and Zeyher 1266 (lectotype: S!, designated by Van Wyk

[1991b]; isolectotype: C!; M!; S!, SAM!).

Lipozygis calycina E.Mey., Comm. Pl. Afr. Austr. 1(1): 78. 1836.

Lotononis calycina (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 611. 1843.—

TYPE: SOUTH AFRICA: Cape Province, Katberg, Drege s.n. a (lectotype: K!,

Herb. Benth., designated by Van Wyk [1991b]; isolectotype: BM!, K!, Herb.

Hook.); Klipplaatrivier, Drege s.n. c (Sp; Bothasberg, Drege s.n. d (not seen).

LEOBORDEA ERIANTHA (Benth.) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis eriantha Benth. in Hook. Lond. J. Bot. 2: 605. 1843.—TYPE:

SOUTH AFRICA, `Macalisberg', Burke 383 (holotype: K!, Herb. Benth.

specimen; isotypes: K!, Herb. Hook. specimen, PRE!).

LEOBORDEA ESTERHUYSEANA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb.

nov. Lotononis esterhuyseana B.-E.van Wyk in Bothalia 20: 70 (1990).—

TYPE: SOUTH AFRICA, Cape Province, Ceres District, Stompiesvlei,

Swartruggens, Esterhuysen 29341 (holotype: B011; isotypes: C!, K!, MO!).

283 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA FOLIOSA (H. Bolus) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis foliosa H. Bolus in J. Linn. Soc. (Bot.) 24: 173. 1887.—TYPE:

SOUTH AFRICA, Gauteng, `prope Pretoria', Maclea s.n. sub BOL 5620

(holotype: BOL!; isotype: K!).

LEOBORDEA FURCATA (MerXM011er and Schreiber) B.-E.van Wyk and Boatwr., comb. nov. Lotononis furcata (MerxmCiller and Schreiber) Schreiber in Mitt.

Bot. Staatssamm. Muenchen 3: 613. 1960.—TYPE: NAMIBIA, Rehoboth district, Buellsport, Strey 2614 (holotype: M!; isotype: K!, M!, PRE!, SAM!)

LEOBORDEA GENISTOIDES Fenzl, Pug. Pl. Nov. Syr.: 6. 1842. Lotononis genistoides (Fenzl) Benth. in Hook. Lond. J. Bot. 2: 607. 1843.—TYPE:

TURKEY, 'in monte Tauro prope Gulek', Kotschy 159 (K!, 2 sheets).

LEOBORDEA GLOBULOSA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis globulosa B.-E.van Wyk in Bothalia 20: 2. 1990.—TYPE: SOUTH

AFRICA, Cape Province, 29.5 km from Touws River to Laingsburg, near

Tweedside, B.-E. van Wyk 2210 (holotype: PRE!).

LEOBORDEA GRANDIS (DUmmer and Jennings) B.-E. van Wyk and Boatwr., comb. nov. Lotononis grandis Dummer and Jennings in Trans. Roy. Soc. S.

Afr. 3(2): 310. 1913.—TYPE: SOUTH AFRICA, Natal, at the Umzinyati River,

Wylie s.n. sub Wood 11525 (holotype: K!).

284 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA LANCEOLATA (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov.

Aspalathus lanceolatus E.Mey., Comm. Pl. Afr. Austr. 1(1): 37. 1836.

Lotononis lanceolate (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 606. 1843.—

TYPE: SOUTH AFRICA, Cape Province, Witbergen ... prope Leeuwenspruit',

Drege s.n. (lectotype: K!, designated by Van Wyk [1991b]; isolectotype: BM!,

K!, PRE!, MO!, S!, SAM!).

LEOBORDEA LATICEPS (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis laticeps B.-E.van Wyk in Bothalia 20: 3. 1990.—TYPE: SOUTH

AFRICA, Cape Province, Ceres District, Stompiesvlei, Swartruggens,

Esterhuysen 29334 (holotype: BOL!; isotypes: C!, K!, M!, MO!, 5!).

LEOBORDEA LONGICEPHALA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov. Lotononis longicephala B.-E.van Wyk in Bothalia 20: 5. 1990.—TYPE:

SOUTH AFRICA, Flats east of Prince Alfred's Hamlet, Oliver 5063 (holotype:

PRE!; isotypes: K!, MO!, STE!).

LEOBORDEA LONGIFLORA (H. Bolus) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis longiflora H. Bolus in J. Linn. Soc. 25: 159. 1889.—TYPE: SOUTH

AFRICA, Cape Province, Namaqualand, Dowdle s.n. sub BOL 6568

(holotype: BOL!; isotype: K!).

LEOBORDEA LUPINIFOLIA Boiss. in Bibl. Univ. Geneve, Elench. Pl. Nov.: 36.

1838. Lotononis lupinifolia (Boiss.) Benth. in Hook. Lond. J. Bot. 2: 607.

285 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

1843.—TYPE: SPAIN, Malaga Province, 'in arenis torrentium Malaga, Motril',

Boissier El. 61 (G; K! 2 sheets, M! 2 sheets).

LEOBORDEA MAGNIFICA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis magnifica B.-E.van Wyk in S. Afr. J. Bot. 55(6): 647. 1989.—TYPE:

SOUTH AFRICA, Cape Province, Summit of Khamiesberg, 3 km south of radio tower, B.-E. van Wyk 2421 (holotype: PRE!; isotypes: K!, MO!, NBG!).

LEOBORDEA MAROCCANA (Ball) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis maroccana Ball in J. Bot. 11: 302. 1873.—TYPE: `Ourika', Ball s.n.

(lectotype: K!, upper specimen on sheet with illustration, designated by Van

Wyk [1991b]; isolectotypes: BM!, K!, bottom of second sheet), Hooker s.n.

(isosyntype: K!); Marocco, 'in regione inferiori Atlantis Majoris: prope

Tasseremout' Ball s.n. (isosyntype: K!); 'Aft Mesan', Ball s.n. (isosyntypes: K!

2 sheets); `Amsmiz', Ball s.n. (isosyntypes: K!, 2 sheets), Hooker s.n.

(isosyntype: K!).

LEOBORDEA MIRABILIS (Dinter) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis mirabilis Dinter in Feddes, Repert. 30: 200. 1932.—TYPE:

NAMIBIA, `Granitflachberge von Aus and Gubub', Dinter 3597 (lectotype:

SAM, designated by Van Wyk [1991b]; isolectotypes: BOL!, K!, PRE!);

`Granitberge von Zwartaus (6 km nordl. Aus)', Dinter 6098 (BM!, BOL!, K!, M!,

PRE!, S!, SAM!).

286 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA MOWS (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov. Lipozygis moffis E.Mey., Comm. Pl. Afr. Austr. 1(1): 79. 1836. Lotononis moffis (E.Mey)

Benth. in Hook. Lond. J. Bot. 2: 609. 1843.—TYPE: SOUTH AFRICA, Cape

Province, Leliefontein, Drege s.n. (lectotype: K!, designated by Van Wyk

[1991b]; isolectotypes: MO!, 5!).

LEOBORDEA MUCRONATA (Conrath) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis mucronata Conrath in Kew Bull. 1908: 222. 1908.—TYPE: SOUTH

AFRICA, Gauteng, Modderfontein, Conrath 124 (holotype: K!).

LEOBORDEA NEWTONII (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis newtonii Dummer in Trans. Roy. Soc. S. Afr. 3(2): 303. 1913.—

TYPE: ANGOLA, Mossamedes, Moulino, Newton 95 (lectotype: K!, designated by Van Wyk [1991b]; isolectotype: Z).

LEOBORDEA OLIGOCEPHALA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov. Lotononis oligocephala B.-E.van Wyk in Bothalia 20: 1. 1990.—TYPE:

SOUTH AFRICA, Cape Province, Areb, ± 27 miles [43.2 km] NE of Springbok,

Van der Westhuizen 276 (holotype: PRE!; isotypes: K!, MO!).

LEOBORDEA PARIFLORA (N. E. Br.) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis pariflora N. E. Br. in Burtt Davy, Man. Pl. Transvaal: 388. 1932.—

TYPE: SOUTH AFRICA, Gauteng, Pietersburg District, The Downs, Rogers

21994 (lectotype: K!, designated by Van Wyk [1991b]; isolectotype: PRE!);

Rogers 22017 (PRE!).

287 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA PENTAPHYLLA (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov.

Lipozygis pentaphylla E.Mey., Comm. Pl. Afr. Austr. 1(1): 79. 1836. Lotononis pentaphylla (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 605. 1843.—TYPE:

SOUTH AFRICA, Cape Province, `Karakuis', Drege s.n. (lectotype: K!,

designated by Van Wyk [1991b]; isolectotype: MO!, S!).

LEOBORDEA PLATYCARPA (ViV.) B.-E.van Wyk and Boatwr., comb. nov. Lotus

platycarpos Viv., Pl. Aegypt., Dec IV: 14, t. 2, fig. 9. 1830. Lotononis

platycarpa (Viv.) Pic-Serm. in Webbia 7: 331. 1950.—TYPE: 'in desertis prope

Kahirum', Figari s.n. (holotype: G).

LEOBORDEA PLICATA (B.-E.van Wyk) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis plicata B.-E.van Wyk in S. Afr. J. Bot. 55(6): 649. 1989.—TYPE:

SOUTH AFRICA, Cape Province, near Bitterfontein, between Vanrhynsdorp

and Bitterfontein, Salter 1601 (holotype: K!; isotype: BOL!).

LEOBORDEA POLYCEPHALA (E.Mey.) B.-E.van Wyk and Boatwr., comb. nov.

Lipozygis polycephala E.Mey., Comm. Pl. Afr. Austr. 1(1): 79. 1836. Lotononis

polycephala (E.Mey.) Benth. in Hook. Lond. J. Bot. 2: 605. 1843.—TYPE:

SOUTH AFRICA, Cape Province, `Khamiesbergen', Drege s.n. (lectotype: K!,

designated by Van Wyk [1991b]; isolectotype: BM!, MO!, S!).

LEOBORDEA PROCUMBENS (H. Bolus) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis procumbens H. Bolus in J. Bot. 1896: 18. 1896.—TYPE: SOUTH

288 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

AFRICA, Free State, Harrismith district, plateau and mountains at Bester's

Vlei, Bolus 8139 (holotype: BOL!; isotype: NBG!).

LEOBORDEA PROLIFERA (E.Mey.) Eckl. and Zeyh., Enum. Pl. Afr. Austr.: 175.

1836. Crotalaria prolifera E.Mey. in Linnaea 7: 152. 1832. Lotononis prolifera

(E. Mey.) B.-E.van Wyk in Contr. Bol. Herb. 14: 135. 1991b.—TYPE: SOUTH

AFRICA, Cape Province, 'Nieuwe feld', Distr. Beaufort, Drege s.n. (lectotype:

S!, designated by Van Wyk [1991b]).

LEOBORDEA PULCHRA (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis pulchra Dummer in Trans. Roy. Soc. S. Afr. 3(2): 308. 1913.—

TYPE: SOUTH AFRICA, Gauteng, Lydenburg, Wilms 280 (lectotype: K!, designated by Van Wyk [1991b]; isolectotype: BM!), Wilms 279a p.p. (BM!),

Atherstone s.n. (not seen).

LEOBORDEA PUSILLA (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis pusilla Dummer in Trans. Roy. Soc. S. Afr. 3(2): 324. 1913.—

TYPE: SOUTH AFRICA, Cape Province, slopes of mountains, Eland's hoek near Aliwal North, F. Bolus 122 sub BOL 10535 (lectotype: K!, designated by

Van Wyk [1991b]); F. Bolus 122 sub BOL 8141 (isolectotype: BOL!), F. Bolus

122 (isolectotype: PRE!).

LEOBORDEA QUINATA (Thunb.) B.-E.van Wyk and Boatwr., comb. nov. Ononis quinata Thunb., Prodr. Pl. Cap.: 130. 1800. Lotononis quinata (Thunb.) Benth.

289 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS in Hook., Lond. J. Bot. 2: 608. 1843.—TYPE: 'e Cap. B. Spei', Thunberg s.n. sub THUNB-UPS 16636 (lectotype: UPS!, designated by Van Wyk [1991b]).

LEOBORDEA ROSEA (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis rosea Dummer in Trans. Roy. Soc. S.Afr. 3(2): 305. 1913.—TYPE:

SOUTH AFRICA, Cape Province, Clanwilliam, Mader 207 (lectotype: K!, designated by Van Wyk [1991b]; isolectotype: GRA!).

LEOBORDEA SCHOENFELDERI (Dinter ex MerxmUller and Schreiber) B.-E.van

Wyk and Boatwr., comb. nov. Amphinomia schoenfelderi Dinter ex Merxmuller

and Schreiber in Bull. Jard. Bot. Brux. 27: 273. 1957. Lotononis schoenfelderi

(Dinter ex Merxmaller and Schreiber) Schreiber in Mitt. Bot. Staatssamm.

Muenchen 3: 613. 1960.—TYPE: NAMIBIA, Grootfontein district, 'Gross Huis',

Dinter 7383 (holotype: M!; isotypes: BM!, BOL!, K!, M!, PRE!, S!, WIND!).

LEOBORDEA SPICATA (Compton) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis spicata Compton in J. S. Afr. Bot. 41(1): 48. 1975.—TYPE:

SWAZILAND, Mpaleni, Compton 32111 (holotype: NBG!; isotypes: K!, 2

sheets, PRE!).

LEOBORDEA STIPULOSA (Bak. f.) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis stipulosa Bak. f., Leg. Trop. Afr. 1: 18. 1926.—TYPE: RHODESIA

[ZIMBABWE], Macheke, Eyles 2020 (holotype: K!; isotypes: K!, SAM!)

290 CHAPTER 8: THE GENERIC CONCEPT OF LOTONONIS

LEOBORDEA STOLZII (Harms) B.-E.van Wyk and Boatwr., comb. nov. Lotononis stolzii Harms in Engl., Bot. Jahrb. 54: 379. 1917.—TYPE: TANZANIA, (North of Lake Nyasa, Kyimbila District), Western Njombe District, Madehani, Stolz

2602 (Bt; lectotype: K!, designated by Van Wyk [1991b]; isolectotypes: BM!,

BOL!, C!, K!, MO!, PRE!).

LEOBORDEA SUTHERLAND!! (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis sutherlandii Diimmer in Trans. Roy. Soc. S. Afr. 3(2): 307. 1913.—

TYPE: SOUTH AFRICA, Natal, without precise locality, Sutherland s.n.

(holotype: K!).

LEOBORDEA TAPETIFORMIS (Emberger and Maire) B.-E.van Wyk and Boatwr., comb. nov. Lotononis tapetiformis Emberger and Maire in Bull. Soc. Hist. Nat.

Afrique Nord 28(6): 349. 1937.—TYPE: MOROCCO, ... 'Atlantis Majoris orientalis ad radices australes montis Masker, inter Tagoudimt et Anemzi, ad alt. 2200-2400 m', Emberger and Maire s.n. 1936 (not seen).

LEOBORDEA WILMS!! (Dummer) B.-E.van Wyk and Boatwr., comb. nov.

Lotononis wilmsii Dummer in Trans. Roy. Soc. S. Afr. 3(2): 307. 1913.—

TYPE: SOUTH AFRICA, Gauteng, between Middelburg and Crocodile River,

Wilms 277 (holotype: K!).

291 TABLE 8. 1 Characters states for 12 morp hological, chem ical and cytological characters scored for the accessions includedin the combined molecular and morphological analyses. Characters and character states are explained at the end of the table. 1— .1— 1— N 0)000000 7-- 0 N Cr)000000 •:1' CO N. CO 1— LC)000000 0 0 C \I CD 1— 000000 0000000000 OON Aspalathus carnosa 000000 0 0 N 0 Aspalathus confusa 1— 1— 1— 0 0 CM c \I Aspalathus crenata 0 0 0 .1 000 Aspalathus galeata 0 0 cv 0 Aspalathus hirta subsp. hirta 1— 0 0 cm 0 0 Aspalathus hystrix 1— 0 0 0 cm T— 00 01— OCD 00CD Aspalathus laricifolia subsp. laricifolia 00 0 0 C \I — 1— 1— CDO Aspalathus linearis 0 0 0 0 0 0 1— (1) 00 00 Aspalathus macrantha 0001—N00000 0 0 0 0 0 0 cN 0 1— 0 Aspalathus nivea 0 0 NNNN 0 1— 00 00 0 040 Aspalathus pachyloba 00 0 0 CD .1— 0 Aspalathus pendula 0 0 (0 000 0 0 1— 0 00 CZ) Aspalathus perfoliata subsp. perfoliata 0 0 0 0 0 T— 00000 00 CDCD Aspalathus shawii subsp. shawii 00 Cs— 0 0 1— Aspalathus vermiculata 0 0 0 000 1— CD T 000 0 0 0 . Aspalathus willdenowiana 1— 0 N . 0 00 00 QC) Bolusia amboensis T T— 0 0 1— T 0 . Crotalaria capensis 1— 0 1— ILD 0000 C) r r r 0000 Q000 C,OCD I Crotalaria distans 0000 1— 0 1— 00 0 I Crotalaria griquensis 000 1— 0 0 — 1— 1— 0 1...

I Crotalaria humilis 1— 0 *— 0 Crotalaria lanceolata 0 1- 00000000000000000000001- 1- 0

0000000000000000000000001- 0

I-NNNNNNNO-NNNNNNNNC\INNNNNNNN

000000000000000000001- 1- 1- 1- T- 'I

O O O Q Q Q O Q O O Q C Q Q Q O O O Q Q Q Q Q O

0 T T T T 1. T- /-. T T ., T a 0 0 0 0 0 a 0 NN Ni-NN

1- 000000000001-1-1-1-1-1-1- 1- OCD 001- 0

000000000000000000001- 1- 1- 1- 1-1-

000000000000000000001- 1- 1- 1- 1- 1-

C) 000000000000000000001- T-1-1-1-1-

1-01-0001- 1-NOOONNNNNNNNO0 0 CDOO

00000000000000000000000000

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h 0cowa) hilus 0. t fn fn fn fn fn by )94 IL) ca_ Wiborgie Wibo Wiborg C7) Wiborg Wibo Wiborg Wiborg Wiborg Wibo Wiborg Wibo Wibo Dic Ro Ro Ra Ra Ra Ra Ra CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

9.1. INTRODUCTION

Robynsiophyton is a poorly known, monotypic genus that occurs in south-central tropical Africa. It is morphologically similar to Pearsonia and Rothia, suggesting a close relationship with these genera (Polhill 1976, 1981; Van Wyk 1991a). Pearsonia occurs in central and southern tropical Africa, while Rothia is widely distributed throughout tropical Africa, Asia and Australia (Polhill 1974; Chapter 10; Boatwright et al. 2008c).

These three genera differ from the rest of the tribe in their monomorphic anthers and straight styles (Polhill 1976; Van Wyk 1991a; Van Wyk and Schutte 1995; Chapter 10;

Boatwright et al. 2008c). Chemically they are also unique within Crotalarieae in accumulating angelate esters of hydroxylupanine-type alkaloids (Van Wyk and

Verdoorn 1991b). In Chapter 3 (Boatwright et al. 2008a) the study of molecular (ITS and rbcL) and morphological data, demonstrated that Pearsonia, Robynsiophyton and

Rothia form a strongly supported Glade and that the latter two are strongly supported as sister genera (Fig. 9.1). Doubt regarding the generic status of Robynsiophyton has been expressed by previous authors (Polhill 1976; Van Wyk 1991a), who suggested that it could merely be a local derivative of Pearsonia. However, as shown in Chapter 3

(Boatwright et al. 2008a) the genus is not embedded within Pearsonia, but sister to

Rothia with both of these subsequently sister to Pearsonia. These data in combination with the annual life history and the unusual androecium (nine stamens with only five fertile anthers) support the generic concept of Robynsiophyton.

297 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

The aim of this chapter is to present a revision of Robynsiophyton with illustrations, a discussion on diagnostic characters and a distribution map.

100 E Pearsonia sessilifolia

99 Pearsonia aristata Pearsonia grandifolia 100 Pearsonia cajanifolia 9: 7 Rothia hirsuta Robynsiophyton vanderystii

FIG. 9.1 Phylogenetic position of Robynsiophyton vanderystii based on gene sequences (ITS and rbcL) and morphological data (strict consensus of 370 trees Chapter 2 — see Boatwright et al. 2008a; tree length = 1166; consistency index = 0.53; retention index = 0.84). Numbers above the branches are bootstrap percentages above 50%.

9.2 DISCUSSION OF CHARACTERS

A summary of the diagnostic characters for Robynsiophyton, Rothia and

Pearsonia is presented in Table 9.1. Robynsiophyton vanderystii Wilczek is a small annual (or rarely a short-lived perennial) with hairy, reddish-brown branches. It shares with Rothia the annual life history, a trait which is not found in Pearsonia, a genus comprising perennial herbs or small shrubs. The leaves are digitately trifoliolate and sparsely pubescent adaxially but densely pubescent on the abaxial surface. The narrow stipules are paired at the base of the petiole. Rothia hirsuta has single stipules at each leaf; Rothia indica has paired stipules; and Pearsonia has the stipules paired or less often absent (Polhill 1974; Chapter 10; Boatwright et al. 2008c).

298 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

The flowers of Pearsonia, Robynsiophyton and Rothia are all relatively unspecialised, with straight or even down-curved styles and monomorphic anthers

(Polhill 1976). In Rothia the anthers are all small and rounded as opposed to the large, elongate anthers of Pearsonia, six of which are attached slightly higher up to the filament (Polhill 1974, 1976). Robynsiophyton has a reduced number of stamens (from

10 to nine) and only five fertile stamens (the other four stamens lack anthers and are sterile). This is the most notable feature of the genus. In the Crotalarieae, Genisteae and Podalyrieae there is great variation in staminal arrangement from completely free stamens (Podalyrieae) to those joined in either an open sheath (Crotalarieae) or closed tube (most Genisteae). The anthers may be either dimorphic with alternating basifixed and dorsifixed anthers, or monomorphic as found in Pearsonia, Robynsiophyton and

Rothia (a feature that is unique to this Glade within the Crotalarieae). Anther characters are very reliable in legumes and usually consistent with other characters, thus providing important insight especially at generic and tribal levels (Bentham 1843; Polhill 1976;

Chapter 3; Chapter 10; Boatwright et al. 2008a, 2008c).

The calyx of Robynsiophyton is equally lobed, while in Pearsonia and Rothia the upper lobes are larger than the three lower lobes. In Pearsonia, the upper and lateral lobes on either side are often fused higher up (the so-called "lotononoid" calyx type).

Bracts are present in all three genera, but bracteoles are generally lacking in Pearsonia

(if present very small) and completely absent in Robynsiophyton and Rothia (Polhill

1974, 1976).

The pods of Robynsiophyton are relatively short and few-seeded when compared to those of Rothia. The mature seeds are brown, smooth and similar in size

299 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON to those of Rothia indica. Robynsiophyton vanderystii has larger seeds than Rothia hirsuta that are even coloured with a smooth surface, whereas those of Rothia hirsuta are mottled with a somewhat rugose surface (Chapter 10; Boatwright et al. 2008c).

TABLE 9.1 Summary of diagnostic characters for Robynsiophyton, Rothia and Pearsonia

Character Robynsiophyton Rothia Pearsonia Life history annual or short- annual perennial lived perennial Stipules paired paired or single paired or absent Bracteoles absent absent small or absent Calyx equally lobed sub-equally lobed zygomorphic Androecium 9 stamens, 5 10 stamens, 10 10 stamens, 10 rounded anthers, rounded anthers, elongated anthers, 4 staminodes 0 staminodes 0 staminodes Fruit oblong to ovate linear to ovate or ellipsoid to linear- falcate oblong Seeds brown, smooth brown or mottled, light or dark smooth or rugose brown, sometimes mottled, smooth

9.3 TAXONOMIC TREATMENT

ROBYNSIOPHYTON Wilczek in Bull. Jard. Bot. Etat. 23: 128. 1953.; in F. C. B. 4: 286.

1953; Hutchinson, Genera of Flowering Plants: 362. 1964; Polhill in Bot. Syst. 1:

326. 1976; Van Wyk and Schutte, Advances in Legume Systematics 7: 306.

1995; Polhill, Fl. Zam. 3 (7): 66. 2003; Leistner, Seed Plants of southern Tropical

Africa: Families and Genera: 202. 2005; Van Wyk in Legumes of the World,

Lewis et al. (Eds): 281. 2005.—TYPE species: Robynsiophyton vanderystii

Wilczek.

300 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

The genus is monotypic and distinguished by the reduced number of stamens

(nine instead of 10) and presence of four staminodes (Table 9.1). It occurs in central tropical Africa extending from Angola to Zambia in the east.

1 ROBYNSIOPHYTON VANDERYSTII Wilczek in Bull. Jard. Bot. Etat. 23: 128. 1953; in F. C.

B. 4: 286, t. 17. 1953; White, F. F. N. R.: 164. 1962; Torre in C. F. A. 3: 6. 1962;

Lock, Leg. Afr. Check-list: 232. 1989; Polhill, Fl. Zam. 3 (7): 66. 2003.—TYPE:

Democratic Republic of Congo, Lazaret du Sacre-Coeur [1025 BA], Vanderyst

s.n. (holotype: BR, photo!).

Small, prostrate or ascending annual or short-lived perennial up to ±0.3 m in height. Branches reddish-brown, pubescent. Stipules 5-8 mm long, linear to lanceolate, invariably paired, pubescent. Leaves digitately trifoliolate; petiole shorter than leaflets,

4-7 mm long; leaflets elliptic to oblanceolate, subsessile, sparsely pubescent adaxially and densely so abaxially, terminal leaflet 14-22 mm long, 4-9 mm wide, lateral leaflets

7-16 mm long, 2.5-6.0 mm wide, obtuse, base cuneate. Inflorescence axillary or rarely terminal congested racemes, with (2–) 5 to 9 (-10) flowers; pedicel less than 1 mm long; bract linear, 1.5-3.0 mm long, pubescent, caducous; bracteoles absent. Flowers pale yellow, 3-6 mm long. Calyx equally lobed, pubescent, 4-5 mm long; tube 1.5-2.0 mm long; lobes subulate, 1.5-3.0 mm long, tips minutely pubescent on inner surface.

Standard 3-5 mm long; claw 1.0-1.5 mm long; lamina elliptic to ovate, 2-4 mm long,

1.0-1.5 mm wide, obtuse to very slightly emarginate, pilose along dorsal midrib. Wings

2.5-4.5 mm long; claw 1.5-1.8 mm long; lamina oblong to obovate, as long as or

301 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON slightly longer than keel, 1.0-2.5 mm long, 0.5-1.0 mm wide, obtuse, glabrous, with 1-

2 rows of sculpturing. Keel 3-4 mm long; claw 1-2 mm long; lamina boat-shaped, 1.5-

3.0 mm long, 0.5-0.8 mm wide, obtuse, glabrous, sometimes with a very slight pocket.

Stamens 9, anthers monomorphic with 5, sub-basifixed anthers alternating with 4 staminodes. Pistil subsessile, pubescent, ovary elliptic, 1.5-2.0 mm long, 0.5-0.8 mm wide with ±5 to 8 ovules; style straight, 1.5-1.8 mm long, glabrous. Pods oblong to ovate, laterally compressed, subsessile, 7-10 mm long, 3-4 mm wide, ±2 to 8 seeded, dehiscent. Seeds oblique-cordiform, 1.0-1.5 mm long, 1.0-1.2 mm wide, brown, smooth (Fig. 9.2). Flowering time: April. Plants appear to be inconspicuous and are poorly collected so that the flowering time needs confirmation.

Distribution and habitat— Robynsiophyton vanderystii occurs in moist, sandy soils and is especially common along roadsides. Very few specimens are available for study and these include collections from Angola, the Democratic Republic of Congo and Zambia, but it is likely that the distribution range is more extensive (Fig. 9.3).

302 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

FIG. 9.2. Morphology of Robynsiophyton vandetystii: (a) leaf in abaxial view; (b1—b2) stipules;

(c) flower in lateral view; (d) bract; (e) outer surface of the calyx (upper lobes to the left); (f) standard petal; (g) wing petal; (h) keel petal; (i) androecium showing the five uniform anthers and four staminodes; (j) anthers (carinal anther on the left); (k) pistil; (I) pod in lateral view; (m) seed in lateral view.

Voucher specimens: (c, d, f, g, h, i, j, k) McCallum Weston 717 (K); (e) Lisowski 20326

(K); (I) Richards 9321 (K); (a, b1 —b2) Richards 18144 (K); (m) Exell and Mendonga 657(K).

Scale bars: (a—m) 1 mm.

303 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

Additional specimens examined

Angola:

0614: Kimbambu, Madimba (—AA), Pauwels 6470 (PRE).

0720: Lunda, Chicapa (—BC), Exell and Mendonca 657 (BM, K, LISC 3 sheets).

Democratic Republic of Congo:

0416: Bakama (—AB), Flamigni s.n. (K).

0515: Bas-Congo, Kisantu (—AA), Vanderyst s.n. (BM, K); Bas-Congo, Kimpako

(—AB), Vanderyst 42290 (K).

1025: Haut-Shaba, Kolwezi (—DA), Lisowski 20326, 20333 (K).

Zambia:

0831: Chilongowelo, Abercorn (—CA), McCallum Weston 717 (K); Mpulungu

Abercorn road, close to Chilongowelo turning (—CA), Richards 5309, 5317, 18144 (K),

Richards 11097 (K, PRE).

0929: Kawambwa, Timnatushi Falls (—CC), Richards 9319a, 9321 (K).

1124: Mwinilunga (—CD), Mutimushi 3416 (K).

304 CHAPTER 9: TAXONOMY OF THE GENUS ROBYNSIOPHYTON

0 Gabon ,:. ------f... Kenya ... • **--.. .; Conc. o : Democratic Republic of Congc •. , ' • i Tanzania • • ''•.• • ...... 10 10 ...... • .. •••..•• ..., Angola "..... i : Mozambiq Zambia . • •....

" Madagascar

:::::". .. • ..1-"' ••••• • Zimbribwe f 20 20 *:...

.1 Botswana : • Namibia

. • 1

South Af ca :.*** 30 30

10 20 30 40 50

FIG. 9.3. Known geographical distribution of Robynsiophyton vanderystii.

305 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

10.1 INTRODUCTION

The genus Rothia was described in 1806 and comprises two species, R. hirsuta and R. indica. The species have been treated in several floristic studies [e.g. The Flora of Tropical Africa (Baker 1926), Flora of West Tropical Africa (Hepper 1958), Flora of

Tropical East Africa (Milne-Redhead 1971), Flora Zambesiaca (Polhill 2003b), A revised Handbook of the Flora of Ceylon (Rudd 1991)] but no comprehensive revision of the genus exists. An Australian-endemic subspecies of R. indica namely R. indica subsp. australis was described by Holland (1997). It is morphologically distinct from the typical Asian subspecies.

The genus is a member of the Papilionoideae and is placed within the tribe

Crotalarieae. The Crotalarieae form part of a monophyletic assemblage of tribes, the

"core" genistoid legumes (Crisp et al. 2000; Boatwright et al. 2008b), which comprises the Crotalarieae, Euchresteae, Genisteae, Podalyrieae, Sophoreae (in part) and

Thermopsideae. The Crotalarieae are sub-endemic to Africa with a few species of

Crotalaria, Lotononis and Rothia extending to other continents. The study of the tribe based on gene sequences (ITS and rbcL) and morphological data (Chapter 3;

Boatwright et al. 2008ab), showed Rothia to be in a well-supported Glade along with

Pearsonia and Robynsiophyton, i.e. the Pearsonia Glade. A sister relationship between

Robynsiophyton and Rothia was strongly supported (Fig. 10.1). This confirmed hypotheses by Polhill (1976), Van Wyk (1991a) and Van Wyk and Schutte (1995) who

306 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA suggested an affinity between these genera. The close relationship between Pearsonia,

Robynsiophyton and Rothia is supported by three synapomorphies, i.e. uniform anthers, straight or down-curved styles and the presence of angelate esters of lupanine-type alkaloids. Rothia can be distinguished from Pearsonia and

Robynsiophyton by its 10 small, rounded anthers and sub-equally lobed calyx.

In this chapter a taxonomic account of Rothia is presented with illustrations, descriptions and distribution maps of the species. The results of anatomical studies of the leaves and fruits of Rothia and Robynsiophyton are also presented.

100 E Pearsonia sessilifolia H__ 99 Pearsonia aristata Pearsonia grandifolia 100 Pearsonia cajanifolia 9997 Rothia hirsuta Robynsiophyton vanderystii

FIG. 10.1 The Pearsonia Glade taken from a strict consensus of 370 trees showing the relationship between Pearsonia, Robynsiophyton and Rothia based on the combined analysis of molecular (ITS and rbcL) and morphological data (Chapter 3; see also Boatwright et al.

2008a; tree length = 1166; consistency index = 0.53; retention index = 0.84). Numbers above the branches are bootstrap percentages above 50%.

307 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

10.2 DISCUSSION OF CHARACTERS

10.2.1 Generic relationships — In Chapter 3 (Boatwright et al. 2008a), the

Pearsonia clade is well-separated from the "Cape group" of the Crotalarieae

(Aspalathus, Lebeckia, Rafnia, Spartidium and Wiborgia) and placed closer to

Lotononis, a genus with which they share the lotononoid or zygomorphic calyx.

However, the three genera of the Pearsonia Glade possess a 17 base-pair deletion at positions 179-196 in the aligned ITS matrix and unique floral characters such as straight styles and uniform anthers. The Crotalaria Glade [Bolusia, Crotalaria and

Lotononis section Euchlora] is early diverging within the Crotalarieae and all have strongly inflated pods (Chapter 3; Boatwright et al. 2008a).

Van Wyk (2003b) discussed the importance of chemical characters in genistoid legumes and mentions that their distribution is not random, but in fact provides reliable information supporting phylogenetic relationships retrieved by studies of DNA sequence data. The Pearsonia Glade is chemically distinct within the tribe Crotalarieae in having unique alkaloids (Van Wyk and Verdoorn 1990; Van Wyk 1991a; Van Wyk and Schutte

1995; Van Wyk 2005). Pearsonia, Robynsiophyton and Rothia produce tetracyclic quinolizidine alkaloids such as lupanine together with angelate esters such as lupanine-

13a-angelate, cajanifoline, sessilifoline and pearsonine (Hussain et al. 1988; Van Wyk and Verdoorn 1989d, 1991a). These chemical characters are especially useful in distinguishing this Glade from the morphologically similar genus Lotononis where pyrrolizidine and quinolizidine alkaloids are present but no esters of lupanine-type

308 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA alkaloids (Van Wyk 1991b). Also, some sections of Lotononis are cyanogenic, a trait which is unique within Crotalarieae (Van Wyk 1991b).

10.2.2 Vegetative morphology and anatomy —The species of Rothia are prostrate to procumbent or erect annuals. They share this life history with the monotypic African genus Robynsiophyton. This character lends support to the sister relationship found between these genera in Chapter 3 (Boatwright et al. 2008b) and distinguishes them from the closely related genus Pearsonia which consists of perennial herbs or shrubs.

The leaves of Rothia are digitately trifoliolate as in many of the Crotalarieae, and pubescent both adaxially and abaxially. Rothia indica subsp. australis has densely pubescent leaves with long whitish hairs that are more or less spreading. In contrast, the leaves of R. indica subsp. indica are moderately hairy with shorter, more or less appressed hairs. The stipules of Rothia hirsuta are linear to slightly falcate and single at each node, whereas those of Rothia indica are elliptic to lanceolate or ovate and invariably paired.

Anatomical studies of the lamina (Fig. 10.2 a, b) and petiole (Fig. 10.2 c, d) revealed no informative differences between Rothia hirsuta and R. indica, or even between these and Robynsiophyton vanderystii. The leaves are dorsiventral with mucilage cells present in the epidermis in both Rothia and Robynsiophyton (Fig. 10.2 a, b). These cells are thought to contribute to the retention of water and reduction of transpiration but their function has not been accurately ascertained (Gregory and Baas

1989).

309 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

c d

FIG. 2 Transverse sections through the leaves and fruit of Rothia and Robynsiophyton.

(a) Portion of leaf of Robynsiophyton vanderystii (scale bar = 0.1 mm). (b) Portion of leaf of

Rothia indica (scale bar = 0.1 mm). (c) Petiole of Robynsiophyton vanderystii (scale bar = 0.2 mm). (d) Petiole of Rothia indica (scale bar = 0.2 mm). (e) Portion of fruit wall of

Robynsiophyton vanderystii (scale bar = 0.07 mm). (f) Portion of fruit wall of Rothia indica

(scale bar = 0.07 mm). Voucher specimens. (a, e) Lisowski 20326 (K). (b, d, f) Latz 16126

(MEL). (c) Exell and Mendonga 657 (K).

10.2.3 Reproductive morphology and anatomy—Both species of Rothia have axillary racemes. Rothia hirsuta has up to seven flowers per raceme as opposed to R. indica where up to four flowers per raceme are found, although the flowers are often solitary or in pairs. Terminal racemes are found in most genera of the Crotalarieae

(Polhill 1976), but are either terminal or axillary in Pearsonia and strictly axillary in

Rothia and Robynsiophyton. Bracts are present in Rothia, but bracteoles are lacking.

310 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

Bracteoles are also absent in Robynsiophyton and most species of the genus

Lotononis, but present in almost all Pearsonia species.

The flowers of Rothia and its relatives are relatively unspecialised compared to the rest of the Crotalarieae. Rothia, Robynsiophyton and Pearsonia have a "gullet" type flower where the style is straight or down-curved and the anthers uniform and/or further reduced, as opposed to the more specialised and reflexed floral parts found in the other crotalarioid genera (Polhill 1976). In Rothia the two upper calyx lobes are larger than the other three lobes and distinctly falcate. The calyces of Pearsonia species have the upper sinus often shallower than the others or in some species the lateral sinus is the shallowest with the carinal lobe always narrower than the other lobes (Polhill 1974).

Robynsiophyton differs from both these genera in having an equally lobed calyx

(Chapter 9; Boatwright and Van Wyk 2009).

The most useful generic character to distinguish between Pearsonia,

Robynsiophyton and Rothia is the androecium. These genera are unique within the

Crotalarieae in having uniform anthers as opposed to the dimorphic anthers found in other genera of the tribe with alternating basifixed and dorsifixed anthers (Polhill 1976).

The anthers of Pearsonia are elongate and large, whereas in Rothia and

Robynsiophyton they are small and rounded (Fig. 10.3). Robynsiophyton has a reduced number of stamens (nine rather than 10) and four anthers are infertile (Fig. 10.3).

Although the generic concepts of Rothia and Robynsiophyton have been questioned by previous authors, the informative value of androecial characters (the main distinction between these three genera) at both tribal and generic level is usually closely correlated with relationships suggested by both molecular and morphological data

311 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

(Polhill 1976; Chapter 3; Boatwright et al. 2008b) and should be taken at face value as strong, sound generic apomorphies.

FIG. 10.3 Androecia and anthers of Pearsonia (a, el), Robynsiophyton (b, e2) and

Rothia (c, e3). Voucher specimens: (a, el) Swanepoel 1 (JRAU). (b, e2) McCallum Weston 717

(K). (c, e3) Greenway and Kanuri 14269 (K). All scale bars represent 1 mm.

The pods of Rothia hirsuta are ovate to falcate and much shorter and fewer seeded (up to 18 mm long with ±25 seeds) than the linear pods of R. indica that are up to 55 mm long with ±35 seeds. The pods of Rothia and Robynsiophyton are thin-walled

312 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA and sclerified on the inner surface with no apparent anatomical differences (Fig. 10.2 e, f).

Rothia indica and Robynsiophyton have similar sized seeds that are larger than those of Rothia hirsuta (Chapter 9; Boatwright and Van Wyk 2009). The seeds of the latter are light brown with dark mottling and have a rugose surface in contrast to the brown, smooth seeds of R. indica and Robynsiophyton vandetystii.

10.3 TAXONOMIC TREATMENT

Key to the genus Rothia

1. Anthers dimorphic, styles upcurved other Crotalarieae

1. Anthers uniform, styles straight or down-curved 2

2. Perennial herbs or small shrubs, anthers large and elongate Pearsonia

2. Annual herbs, anthers small and rounded 3

3. Upper two calyx lobes larger than others (calyx sub-equally lobed),

stamens 10, all fertile Rothia

3. Upper two calyx lobes similar to others (calyx equally lobed), stamens

nine, five fertile, four reduced and infertile Robynsiophyton

ROTHIA Pers., Syn. Pl. 2: 302, 638. 1807. nom. conserv.; Benth. in Hook., Lond. J. Bot.

3: 338. 1844; Benth. and Muller, Fl. Austral. 2: 185. 1864; Baker, Fl. Trop. Africa:

7. 1871; Trimen, Handb. FL Ceylon: 7. 1894; Thonner, Flowering Plants of

Africa: 274. 1915; Baker, Leg. Trop. Africa: 21. 1926; Hutchinson and Dalziel, Fl.

313 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

West Trop. Africa: 543. 1954; Hutchinson, The Genera of Flowering Plants: 361.

1964; D'Orey and Liberato, Fl. Guine Portuguesa: 71. 1971; Polhill in Bot. Syst.

1: 326. 1976; Thulin in Op. Bot. 68: 153. 1983; Hedberg and Edwards, Fl.

Ethiopia: 195. 1989; Rudd, A revised handbook to the flora of Ceylon: 184. 1991;

Van Wyk and Schutte, Advances in Legume Systematics 7, M. D. Crisp and J. J.

Doyle (Eds.): 306. 1995; Leistner, Seed Plants of southern Africa: Families and

Genera: 295. 2000; Du Puy et al., The Leguminosae of Madagascar: 671. 2002;

Polhill, Fl. Zambesiaca: 64. 2003; Leistner, Seed Plants of southern Tropical

Africa: Families and Genera: 202. 2005; Van Wyk in Legumes of the World,

Lewis et al. (Eds.), 281. 2005.—TYPE species: R. trifoliata (Roth) Pers.

(=Dillwynia trifoliata Roth)

Dillwynia Roth, Catal. Bot. 3: 71. 1806 non Dillwynia Sm. in Koenig and Sims, Ann. Bot.

1: 510 (1805).—TYPE species: D. trifoliata Roth

Westonia Spreng., Syst. Veg. 3: 152, 230. 1826 nom. illeg. —TYPE species: W.

humifusa (Willd.) Spreng.

Goetzea Reichb., Consp. Regn. Veg.: 150. 1828 nom. rej.

Xerocarpus Guill. and Perr., Fl. Seneg. Tent.: 169. 1832.—TYPE species: X. hirsutus

Guill. and Perr.

Harpelema J.Jacq., Ecl. Pl. Rar. 2: t. 129 (1844).—TYPE species: H. dillwynia J.Jacq.

Prostrate, annual herbs. Branches up to 40 cm long, densely to sparsely pubescent. Leaves digitately trifoliolate, elliptic to oblanceolate or obovate, pubescent; leaflets sub-sessile, apex apiculate, obtuse or slightly acute, base cuneate; petioles

314 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA shorter than leaflets, pubescent. Stipules linear to slightly falcate or elliptic to lanceolate or ovate, single in R. hirsuta, paired in R. indica, pubescent. Inflorescence axillary, racemose, flowers solitary or up to 7 per raceme; pedicel up to 2 mm long; bract linear, pubescent, caducous; bracteoles absent. Flowers pale yellow to white. Calyx pubescent, sub-equally lobed; lobes triangular to lanceolate, upper lobes wider than others, falcate, carinal lobe slightly narrower than others, tips minutely pubescent inside. Standard elliptic to ovate, pilose along dorsal midrib; claw linear; apex obtuse.

Wing petals oblong to obovate, glabrous, with 2 to 3 rows of sculpturing, as long as keel; apex obtuse. Keel boat-shaped, glabrous, pocket sometimes present; apex obtuse. Stamens 10, anthers uniform, sub-basifixed. Pistil sub-sessile, pubescent; ovary linear to narrowly ovate; style straight, glabrous. Pods linear to ovate or falcate, laterally compressed, sub-sessile, many-seeded, pubescent, dehiscent. Seeds oblique- cordiform, brown or brown mottled with dark brown, surface smooth or rugose.

Etymology—The genus is named in honour of the German physician and botanist Albrecht Wilhelm Roth (1757-1834).

Diagnostic characters — Rothia can be distinguished from Robynsiophyton in that the latter has an equally lobed calyx (in Rothia the upper two lobes are larger than the others) and the 10 stamens with 10 fertile anthers found in Rothia (in

Robynsiophyton the stamens are reduced to nine and only five anthers are fertile).

Furthermore it differs from Pearsonia by its sub-equally lobed calyx and small, rounded anthers in contrast to the lotononoid' calyx (zygomorphic due to the fusion of the upper and lateral lobes on either side) and large, elongate anthers of Pearsonia.

315 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

Chromosome number—A count of n = 7 has been reported for Rothia indica

(Goldblatt 1981; Bairiganjan and Patnaik 1989).

Distribution —The genus occurs in tropical Africa, Asia and Australia (Fig. 10.5 and Fig. 10.7). It appears to have been well-collected in areas for which regional floras have been compiled, e.g. the Flora of Tropical East Africa and Flora of West Tropical

Africa.

Key to the species of Rothia

1. Stipules linear and single at each node, up to seven flowers per inflorescence, pods

ovate to falcate and up to 18 mm long, seeds light brown with dark mottling and

a rugose surface, Africa and Madagascar 1. R. hirsuta

1. Stipules elliptic to lanceolate or ovate and paired at each node, up to four flowers per

inflorescence, pods linear or slightly falcate and up to 55 mm long, seeds brown with

a smooth surface, Asia and Australia 2. R. indica

1 RoTHIA HIRSUTA (Guill. and Perr.) Baker in Fl. Trop. Africa 2: 7. 1871; Eyles in Trans.

Roy. Soc. South Africa 5: 369. 1916; E.G.Baker, Leg. Trop. Africa: 21. 1926;

Broun and Massey, Fl. Sudan: 176. 1929; Andrews, The Flowering Plants of the

Anglo-Egyptian Sudan: 230. 1952; Hepper in Fl. West Trop. Africa, ed. 2, 1: 543.

1958; Torre, Consp. Fl. Angolensis 3: 5. 1962; White, Forest Fl. Northern

Rhodesia: 164. 1962; Schreiber in Merxmaller, Prodr. Fl. SW. Africa, fam. 60:

107. 1970; Milne-Redhead, Fl. Trop. East Africa, Leguminosae, Pap.: 811. 1971;

Drummond in Kirkia 8: 226. 1972; Thulin in Op. Bot. 68: 153. 1983; Hedberg and

316 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

Edwards, Fl. Ethiopia: 195. 1989; Du Puy et al., The Leguminosae of

Madagascar: 671. 2002. Xerocarpus hirsutus Guill. and Perr. in Guillemin,

Perrottet and Richard, Fl. Seneg. Tent.: 44 (1832).—TYPE: SENEGAL, Leprieur

s.n. (lectotype: P photo!, designated here). [Note: According to Stafleu and

Cowan (1983) the type material of Perrottet is housed in P or G. No specimens

were traced in G and the Paris specimen is therefore designated as lectotype.]

Amphinomia desertorum (Dummer) Schreiber in Mitt. Bot. Munchen 2: 286. 1957;

Lotononis desertorum Dummer in Trans. Roy. Soc. S. Africa 3: 316. 1913.—

TYPE: Great Barmen, Buschsteppe, 23.iv.1907, Dinter 518 (lectotype: Z photo!,

designated here; isolectotypes: BM!, BOL!, NBG!, K!). [Note: Dummer was

based at K during his revision of Lotononis. However, the K isotype is a small

piece taken from the specimen in Z (as annotated by Dummer himself). The Z

specimen is therefore chosen as lectotype.]

Small, pubescent, prostrate or semi-erect to procumbent herb. Branches ±10-40 cm long, brown, sparsely hairy (pilose). Leaves with leaflets elliptic to oblanceolate, terminal leaflet 10-30 mm long, 3.5-12.0 mm wide, lateral leaflets 6-20 mm long, 2-7 mm wide; apex apiculate; base cuneate; petiole 3-13 mm long. Stipules linear to slightly falcate, 2-8 mm long, single. Inflorescence with 2 to 7 flowers; pedicel up to 1 mm long; bract 1.0-2.5 mm long. Flowers 4-8 mm long. Calyx 5-7 mm long; tube 1.5-

2.0 mm long; lobes 3-4 mm long. Standard 4.0-5.5 mm long; claw 1.0-1.5 mm long; lamina 3.0-4.5 mm long, 1-2 mm wide. Wings 4-5 mm long; claw 1.5-2.0 mm long; lamina 2.5-4.0 mm long, 0.5-0.7 mm wide. Keel 2.5-4.0 mm long; claw 1.5-2.0 mm

317 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA long; lamina 1-3 mm long, 0.5-1.0 mm wide. Pistil ovary narrowly ovate, 2-3 mm long,

0.5-0.7 mm wide with ±7 to 13 ovules; style 1.3-1.6 mm long. Pods ovate, falcate, tapering towards apex, 10-18 mm long, 2-3 mm wide with up to ±25 seeds per pod.

Seeds 1.1-1.5 mm long, 0.8-1.1 mm wide, mature seeds light brown with dark mottling, surface rugose (Fig. 10.4). Flowering time: Rothia hirsuta flowers mainly from

February to November, but flowering and fruiting specimens have been recorded throughout the year.

Distribution and ecology—Rothia hirsuta is widespread in the dry parts of tropical Africa from South Africa northwards to Eritrea and westwards up to Guinea

(Fig. 10.5). Plants occur on sandy loam or clay soils, often in grassland or forest openings. It is common on sandy soils along watercourses or disturbed roadsides.

Diagnostic characters—The linear stipules are invariably single at each node in Rothia hirsuta, whereas they are paired and elliptic to ovate in Rothia indica. The inflorescences are more densely flowered (up to 7 flowers) in R. hirsuta as opposed to

R. indica where the flowers are often solitary or up to 4 per inflorescence. The linear pods of R. indica are three times longer (up to 55 mm) than the ovate pods of R. hirsuta that are only up to 18 mm long. In R. hirsuta the seeds are brown with dark mottling and have a rugose surface, while those of R. indica are brown (unmottled) with a smooth surface.

318 CHAPTER 10: TAXONOMY OF THE GENUS ROTH/A

FIG. 10.4 Morphology of Rothia hirsuta. (a) Standard petal. (b) Seed in lateral view. (c) Outer surface of the calyx (upper lobes to the left). (d) Wing petal. (e) Keel petal. (f) Pistil. (g) Flower in lateral view. (h) Pod in lateral view. (i) Bract. (j) Anther. (k) Androecium. (I) Leaf in abaxial view. (m) Stipule. Voucher specimens. (a, c) Davey 173 (K). (b) Milne-Redhead and Taylor

11245 (K). (d, g, h, j, k) Greenway and Kanuri 14269 (K). (e) Jackson and Apejoye 413973 (K).

(f) Hepper 1021 (K). (i) Polhill and Paulo 2089 (K). (I, m) Siame 162 (K). All scale bars represent 1 mm.

319 CHAPTER 10: TAXONOMY OF THE GENUS ROTH1A

Selected specimens

ERITREA: Bocos, Cheren, Pappi 2527, 10.xi.1902 (BM 2 sheets, K). SUDAN: Kordofan, Kotsch

420, without date (K). CHAD: Dioura, Macina, Davey 173, Oct. 1954 (K). NIGER: Niamey,

Olufsen 476, 8.x.1947 (BM, S). GUINEA: Guine-Bissau, Pussuli, Santos 1438, 17.xii.1942 (K).

BURKINA FASO: Haute Volta, Lergo, Ake Assi 13587, 13.xi.1976 (K). GHANA: Tefle, on Volta

River, Akpabla 1982, 13.8.1959 (K). NIGERIA: Naraguta, Jos Plateau, rough ground near

Forestry Department, amongst grass, Hepper 1021, 14.x.1957 (K). CAMEROON: Ganglani village, 6 mls [9.65 km] from confluence of Kam and Kimiri Rivers, Vogel Peak area, Hepper

1441, 23.xi.1957 (K). DEMOCRATIC REPUBLIC OF CONGO: 50 km north-east of Mokolo, on route to Mora, Bounougou 173, 18.ix.1964 (K, S). UGANDA: Matuga, Robertson 3312,

29.vii.1982 (K). KENYA: Thika-Garissa road, 2.6 km towards Garissa after junction with Kitui road, near Kongonde, Faden and Faden 74/737, 7.vi.1974 (K, PRE). TANZANIA: !magi Hill, 1 ml. [1.61 km] south of Dodoma, Polhill and Paulo 2089, 20.iv.1962 (K, PRE). ZAMBIA:

Livingstone, Chief Mukuni area, 2.1 km along road to Chief Mukuni Palace from junction with

Victoria Falls road in Baikiae forest, Zimba et al. 913, 19.ii.1997 (K). MALAWI: Lilongwe, Banda

250, 14.ix.1956 (BM). MOZAMBIQUE: Niassa, Mossuril para Lumbo, 4.5 km na estrada de

Nampula, Pedro and Pedrogao 3131, 5.v.1948 (K, PRE). ANGOLA: Huilla, Welwitsch 1904,

Apr. 1860 (K, BM). NAMIBIA: Grey sandy flats below Musese Camp west of Lupala Mission

Station; De Winter and Marais 4978, 4.iii.1969 (K). ZIMBABWE: Gwanda, Sezane Reservoirs,

Davies 1296, May 1955 (K, PRE). SOUTH AFRICA: Kruger National Park, Punda Milia, Van

Rooyen 494, 19.iii.1976 (PRE). MADAGASCAR: Ankazoabo, Bosser 17241, Feb. 1963 (K).

(150 specimens examined.)

320 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA 20 mitimpwainimato 10 20 30 40 50 60

50 111 11•111111111 50 I 1101114 ANIIIIIII• 6

40 Mt ONIMMIlliti 40 illffille.. "4111M1111110111111 30 11111111111111111 30 20 1111.11111111.11EME 20 lmouppm Imokuirm

10 —mElleru islin" km 10 1111110 1114 II 10 20 minumAraryipqsaw 20

30 Iin rep•mila 30

N131 991 10 0 10 20 30 40 50 60

FIG. 10.5 Known geographical distribution of Rothia hirsuta.

321 CHAPTER 10: TAXONOMY OF THE GENUS ROTH/A

2. ROTHIA INDICA (L.) Druce in Bot. Exch. Club Soc. Brit. Isles 3: 423. 1914; Rudd, A

revised handbook to the flora of Ceylon: 184. 1991; Holland, Austrobaileya 5: 93

(1997); Trigonella indica L., Sp. Pl. 2: 778. 1753.—TYPE: Herb. Hermann 3: 24

no. 285 BM-000621888 (lectotype: BM photo!, designated by Rudd [1991])

Lotus indicus Desr. in Lam., Encyc. 3: 606. 1759.—TYPE: as for T. indica

Hosackia indica Graham in Wallich, Numer. List n. 5940. 1831-1832. nom. nud.

Glycine leptocarpa Graham in Wallich, Numer. List n. 5515. 1831-1832. nom. nud.

Rothia trifoliata (Roth) Pers., Syn. Pl. 2: 638 and 659. 1807; Benth. in Hook., Lond. J.

Bot. 3: 339. 1844; Bentham and Muller, Fl. Austr. 2: 185. 1864; Trimen, Handb.

Fl. Ceylon: 7. 1894; Dillwynia trifoliata Roth, Catal. Bot. 3: 71. 1806.—TYPE:

`horto medico Amsteledamensi' (Bt?)

Small, pubescent, prostrate, herb. Branches up to 40 cm long, brown, either covered with dense, woolly hairs or sparsely hairy (pilose). Leaves with leaflets elliptic to obovate, terminal leaflet (7–) 9-26 mm long, (2–) 3-5 mm wide, lateral leaflets (5–) 8-

20 mm long, (1.5–) 2.5-6.0 mm wide; apex obtuse or slightly acute; base cuneate; petiole 4-13 mm long. Stipules elliptic to lanceolate or ovate, 2-8 (-10) mm long, paired. Inflorescence with flowers solitary or up to 4 per raceme; pedicel up to 2 mm long; bract 1-3 mm long. Flowers 5-8 mm long. Calyx 4-7 mm long; tube 1.5-4.0 mm long; lobes 1-4 mm long. Standard 4.0-6.5 mm long; claw 1.5-2.5 mm long; lamina

2.5-4.0 mm long, 2.0-2.5 mm wide. Wings 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Keel 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Pistil ovary linear, 3.0-4.5 mm long,

322 CHAPTER 10: TAXONOMY OF THE GENUS ROTH1A

0.5-1.0 mm wide with ±13 to 18 ovules; style 1.4-1.7 mm long. Pods linear or slightly falcate, (30—) 35-55 mm long, 1.5-3.0 mm wide with up to ±35 seeds per pod. Seeds

1.3-1.8 mm long, 1.0-1.3 mm wide, mature seeds brown, surface smooth (Fig. 10.6).

Flowering time: Flowering and fruiting all year round from January to December.

Distribution and ecology —The species occurs in India, Sri Lanka, China,

Vietnam, Malaysia and Australia (Fig. 7).

Diagnostic characters —The paired, ovate stipules, sparse inflorescences, long, linear pods and smooth, brown seeds distinguish Rothia indica from R. hirsuta.

Notes —Two subspecies can be recognised based predominantly on their geographical distribution, size and pubescence of vegetative and reproductive parts.

1. Vegetative parts sparsely hairy, terminal leaflets (7—) 10-26 x (2—) 4-8 mm, lateral

leaflets (5—) 8-20 x (1.5—) 3.0-6.0 mm, petiole 5-13 mm long, stipules elliptic to

lanceolate 2-8 (-10) mm long, pods (-33) 35-55 x (1.5—) 2.0-3.0 mm, seeds 1.3—

1.6 x 1.0-1.3 mm, India, Sri Lanka, China, Vietnam and

Malaysia 2a subsp. indica

1. Vegetative parts covered with dense, woolly hairs, terminal leaflets 9-15 x 3-5 mm,

lateral leaflets 9-12 x 2.5-4.0 mm, petiole 4-8 mm long, stipules elliptic to ovate 2-5

mm long, pods (30—) 35-45 x 1.5-2.0 mm, seeds 1.5-1.8 x 1.2-1.3 mm, restricted

to Australia 2b. subsp. australis

323 CHAPTER 10: TAXONOMY OF THE GENUS ROTH/A

FIG. 10.6 Morphology of Rothia indica. (a) Standard petal. (b) Pod in lateral view. (c) Leaf of R. indica subsp. australis in abaxial view. (dl, d2) Stipules of R. indica subsp. australis. (e) Leaf of

R. indica subsp. indica in abaxial view. (f1, f2) Stipules of R. indica subsp. indica. (g) Pistil. (h)

Wing petal. (i) Keel petal. (j) Androecium. (k) Flower in lateral view. (I) Anther. (m) Bract. (n)

Seed in lateral view. (o) Outer surface of the calyx (upper lobes to the left). Voucher specimens.

(a, g, h, i, k) Cooray 70020117R (K). (b, n) Gamble 13771 (K). (c, dl, d2) Latz 16126 (MEL). (e, fl, f2) Barber 55 (K). (j, /, o) Wight 828. (K). (m) Wight 571 (K). All scale bars represent 1 mm.

324 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

40 0 700 1400 km 60 70 80 90 100 110 120 130 140 150 160

FIG. 10.7. Known geographical distribution of Rothia indica subsp. indica (triangles) and

R. indica subsp. australis (dots). Specimens cited in Chen and Li (1997) from China are represented by stars.

2a. Rothia indica (L.) Druce subsp. indica

Leaves with terminal leaflet (7—) 10-26 long, (2—) 4-8 mm wide, lateral leaflets (5—) 8-

20 mm long, (1.5—) 3.0-6.0 mm wide; petiole 5-13 mm long. Stipules elliptic to lanceolate, 2-8 (-10) mm long. Inflorescence with flowers solitary or 2 to 4 per raceme.

Calyx 5.0-6.5 mm long; tube 2-4 mm long; lobes 1-3 mm long. Standard 4.0-6.5 mm

325 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA long; claw 1.5-2.0 mm long; lamina 2.5-4.0 mm long, 2.0-2.5 mm wide. Wings 4-6 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.0 mm long, 1.0-1.5 mm wide. Keel 4-6 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.0 mm long, 1.0-1.5 mm wide. Pistil with ovary 3-4 mm long, 0.5-0.7 mm wide with ±14 to 18 ovules; style 1.4-1.6 mm long.

Pods (-33) 35-55 mm long, (1.5—) 2.0-3.0 mm wide with up to ±35 seeds per pod.

Seeds 1.3-1.6 mm long, 1.0-1.3 mm wide.

Distribution and ecology—Collections from India, Sri Lanka, Vietnam and

Malaysia were studied (Fig. 10.7), but this subspecies has also been recorded from

China (Chen and Li 1997) and the specimens cited by these authors were included on the map. The subspecies favours sandy soils, grassy slopes or moist open fields. It is common in disturbed areas such as roadsides.

Selected specimens

INDIA: Chinglepul, Barber 55, 10.i.1899 (K); Bombay (Mumbai), Dalzell s.n., Apr. 1878 (K);

Ganjam district, Chennai, Gamble 13771, Feb. 1884 (K); Bengaluru, Gough 162, Dec 1937 (K);

Tirunelveli, Tamil Nadu, Matthew 16388, 26.i.1979 (BM); Denkanikotta, Dharmapuri,

Denkanikotta Rest House, Matthew RHT 24647, 15.xi.1979 (K); Kodaikanal, Dindigul, Law's

Ghat road, Matthew RHT 51949, 4.xii.1987 (K); Mysore, Mimso 607, Oct. 1837 (K); Hassan,

Mysore, Belvathally, Ramamoorthy 1956, 13.xii.1971 (K); 70 km north of Munnar, Palani hills,

Tamilnadu, Van der Maesen 3453, 20.i.1979 (K). VIETNAM: Tourane (now Da Nang), Clemens and Clemens 4294, May—Jul 1927 (BM). SRI LANKA: Ruhuna National Park, block 1 near

Buttawa Bungalow, Cooray and Balakrishnan 69010909R, 9.vi.1969 (BM); Trincomalee, near

China Bay Airport, Rudd and Balakrishnan 3134, 24.ii.1970 (K); Hambantota, Yala, Fagerlind

1695, Jan. 1974 (S). MALAYSIA: Pulau Pinang, Haihtmu 2996, 21.xii.1917 (K). (33 specimens examined.)

326 CHAPTER 10: TAXONOMY OF THE GENUS ROTHIA

2b. Rothia indica subsp. australis A.E.Holland, Austrobaileya 5: 94 (1997)

Type: Queensland, Gregory South District, Site 195, Warlus 1, 7 Aug 1971, Boyland

4016; holo BRI (photo seen)

Leaves with terminal leaflet 9-15 mm long, 3-5 mm wide, lateral leaflets 9-12 mm long, 2.5-4.0 mm wide; petiole 4-8 mm long. Stipules elliptic to ovate 2-5 mm long.

Inflorescence with flowers solitary or 2 per raceme. Calyx 4-7 mm long; tube 1.5-3.0 mm long; lobes 3.5-4.0 mm long. Standard 4.0-6.5 mm long; claw 1.5-2.0 mm long; lamina 2.5-4.0 mm long, 2.0-2.5 mm wide. Wings 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Keel 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Pistil with ovary 4.0-4.5 mm long,

0.6-1.0 mm wide with ±13 to 17 ovules; style 1.5-1.7 mm long. Pods (30—) 35-45 mm long, 1.5-2.0 mm wide with up to ±30 seeds per pod. Seeds 1.5-1.8 mm long, 1.2-1.3 mm wide.

Distribution and ecology—The subspecies is restricted to Australia where it occurs on sandy hills and flats (Fig. 10.7).

Specimens examined

AUSTRALIA: 100 mls [160.9 km] west of Windorah, Everist 3898, 9.vi.1949 (K); Boomerang

Waterhole, Lander River, 20°38'00"S 132°11'00"E, Latz 16126, 10.v.2000 (MEL); Upper

Victoria River, von Muller s.n., without date (K, MEL 2 sheets); near the Ord River, O'Donnell s.n., without date (MEL 2 sheets).

327 CHAPTER 11: GENERAL CONCLUSIONS

The results presented in this study have provided clarity on relationships within the Crotalarieae and also generic circumscriptions within the tribe. The new insights gained have resulted in new generic classifications for both Lebeckia s.l. and Lotononis s.l. As a result the number of genera included in the tribe has increased from 11 to 16.

The new hypothesis of phylogenetic relationships (Fig. 11.1) provides the opportunity to

re-interpret the patterns of evolution of various characters within the tribe and to ascertain the apomorphic states for the various genera. Polhill (1976) and Van Wyk and

Schutte (1995) discussed the distribution of characters within the Crotalarieae and related tribes, but given the high incidence of convergence, some doubt has remained about the circumscription of genera, particularly Lebeckia s.l. The new insights into generic delimitations and relationships allow for a re-evaluation of diagnostic characters and apomorphic states. The characters that are most informative for generic circumscription were therefore reconstructed onto the minimal length trees from the combined molecular/morphological analysis (some already presented in Chapter 8).

The characters included are listed in Table 8.1 (Chapter 8) and Appendix A5. Where more than one reconstruction is possible two or more trees are shown. The reconstructions were done on one of the 370 most parsimonious trees from the combined molecular/morphological analysis presented in Chapter 8 in order to examine the evolution of the specific characters (Appendix A5) on that particular hypothesis of relationships.

328 CHAPTER 11: GENERAL CONCLUSIONS

HABIT AND PERSISTENCE—MOSt of the basally diverging lineages of the Crotalarieae have a herbaceous or suffrutescent habit and woodiness or a shrubby habit appears to have become pronounced in especially the "Cape group". Most of the species are, however, perennial and the development of a short-lived or annual life history appears to be a secondary and convergent character, possibly to survive recurrent fires, periods of aridity or unfavourable weather conditions (Figs. 11.2 and 11.3). Van Wyk and

Schutte (1995) discussed the taxonomic value of the secondary xylem within the

Podalyrieae and identified two types of arrangement, one where the vessels are arranged in small isolated radial or tangential groups and another where the vessels are arranged in large confluent or dendritic groups. The latter has been suggested to be associated with strongly seasonal habitats or Mediterranean climates. The great variety of habitats in which the Crotalarieae occur indicate that a comparative study of wood anatomy would be of considerable interest. It will allow a comparison of the patterns of vessel arrangement with those in the closely related Podalyrieae and Genisteae.

LEAVES—The evolution of leaf types within the Crotalarieae was explored in

Chapter 8 (Fig. 8.3) and indicated that within the Crotalaria and Lotononis clades the leaves are mostly digitate with little variation. The diversity of leaf types increases in the

"Cape group" where unifoliolate and simple leaves become pronounced in groups such as Rafnia and Lebeckia, with the latter having a further development to phyllodinous leaves. In the study of the lebeckioid genera Calobota, Lebeckia and Wiborgiella, lamina anatomy provided important characters that were informative at the generic level (Chapter 5; Boatwright et al. submitted) especially with regard to the arrangement of the mesophyll (isobilateral or dorsiventral) and presence or absence of mucilage

329 CHAPTER 11: GENERAL CONCLUSIONS cells. These characters may potentially be informative throughout the Crotalarieae and should in future be explored with a larger sampling of genera within the tribe to ascertain their taxonomic value.

98 Aspalathus 94 Wiborgia Wiborgiella 82 "Cape group" Lebeckia 69 Rafnia 94 Calobota Ezoloba

98 100 Lotononis 72 100 86 Leobordea Lotononis 51 100 Listia Glade

98 Pearsonia 100 100 Rothia Robynsiophyton

Euchlora 100 100 Crotalaria 90 Crotalaria Glade Bolusia

Genisteae

FIG. 11.1 Hypothesis of phylogenetic relationships in the tribe Crotalarieae. The cladogram is a simplified strict consensus tree of the 370 trees from the combined molecular and morphological analysis presented in Chapter 8.

330 CHAPTER 11: GENERAL CONCLUSIONS

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Lift0L:= tot insolucrots wasp. padunctiMs t I Laflaltstals p.„ 0 j--' =—..." j7t. 7.= ebng''''"ifolLtocin' Lot Month* Lot Mmtm LOL 00.11700,12 Lot sevramps Lot boltati tat Mrs.

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LiOrrLS" a n a Lag digAHLB

al Lot salt/d ais subutato

S sasseritda R meats RstranctSM vooneolla , 11 Rot Ms. .. 1.1Rab. veaMyad L.L. PM. Mims Cr. Mtgranals Cr. commis Cr. si:Asti, Cr nunets Fl .f Cr Sarnia= 13. onr.rtstS o Leptifis

FIG. 11.2 Parsimony-based reconstruction of habit on the 370 minimal length trees from the combined molecular/morphological analysis presented in Chapter 8.

331

CHAPTER 11: GENERAL CONCLUSIONS

= perennial A cannon = short-lived t A. cannon et A. plena or annual A. /1700027VIS A. walonortiona A. contuse A. partaken wasp. ploktedi A. crenate I A. vennieulate , L A. &wog stew. Mo. A. latidkela subcp. lancings A lartesubenana hysofx n A. twee 11 A- Peclglobe WM. final We. tonna. We mime Slab. annoptera l-i1— WID.117011.8 I r II Ile obcontute r_r_r_L,f--D, AVAhl. Image Ina leinadnene NW. oesefolb WM Maim WIt ingots WIPI noconato L.I.:.....p.uci.aocontaos..... Lab. moyedatto --1-1-r-' Lea. t•PODD LW ambigua ,,,,,Log,,0, r...... sepinia

_,J PR:el. globOwWk''„ R . Musa

I--C"- C:= E,.1= Ca sencee Cw0.Ca. lo ono o , la . Ca creme - ._r-H-r- Cal sposseatte Cal senates l , Lot rnacrocame Lot Moan ig),.:=..... Lot spetruecw La. onmete Lot roWeat suing. nemequensts La. lenaada La famine La. mail La lam Lot abbr.. .1 Lot rtranosepale La mioanthe Lot 'abeam tfoRLot. leptolobe lLot. mexinaliani Lot paws __J Lot divaricata 1------L. Lot. solcoptatte Lot. puking, Lot. awed --1—r-' Lot lotononoides 1 Lot. sante La. till70.11 Lot. Most wimp. tootoctoda Lot. malpulata nFL:r- ... Lilt iit0tze..1crata .0 . 00,05. peduntsulate .1 La. filiformie Lot elongate , .,La platycoupe La 1--1 Lot lobo. r-i Lot onontno Lot Mpassa

' ea' acuad"Taneb''

LotLc,u1lEpotycoOPt17:b Ip. :,P to,,t, Wa...n. Lat. La ll Lot. windage 1_,—,- La. sot:Meta Lot OM La narkaln 1-1-0 ...2 ensfata"'46*0 P. weaned, P. attenikela Pot !Omuta Rob. vonctrystii Lot. hewn Cr. drams M Cr tptoonsis --Li Cr. womb CC amen

Cr lanceolate L Li annotate:Is " D. groan

FIG. 11.3 Parsimony-based reconstruction of life history on the 370 minimal length trees from

the combined molecular/morphological analysis presented in Chapter 8.

332 CHAPTER 11: GENERAL CONCLUSIONS

STIPULES—The loss of stipules is reconstructed as an apomorphy for the "Cape group" and stipules are only present in Lebeckia wrightii and L. uniflora. A loss of stipules is autapomorphic for some species of the Leobordea Glade, Pearsonia and

Crotalaria where stipules are largely present (Fig. 11.4). Asymmetrical stipules are unique to the lotononoid groups, i.e. Lotononis s.s., together with the Leobordea and

Listia clades (Fig. 8.4) and Rothia hirsuta, while Euchlora and Ezoloba are conspicuously different in having symmetrical stipules (when present in the former).

BRACTEOLES—Bracteoles are present in all genera of the "Cape group", Ezoloba,

Listia, Pearsonia, Bolusia and Crotalaria. A loss of bracteoles has been regarded as an apomorphy for Lotononis s.l., but this character is reconstructed as a convergence between Lotononis s.s., Leobordea, Rothia, Robynsiophyton and Euchlora (Fig. 8.5).

Prenner (2004) mentions that "initiated but suppressed" bracteoles represents a new character state in Papilionoideae and is a refinement of the traditional presence or absence of bracteoles as presented here. However, ontogenetic studies (investigation of the floral primordium) are necessary to evaluate whether bracteoles are indeed initiated and later suppressed in Lotononis s.s., Leobordea, Rothia, Robynsiophyton and Euchlora or whether a total absence is found in Crotalarieae.

333

CHAPTER 11: GENERAL CONCLUSIONS

= present = 1.. = A..tnose A patents = absent A massentha A padienowisno A. confuse A pelf... Wasp. phAnpag A. crenate A. vertnissAste A. shwa subsp. stu. A. brisk:es wimp. IsfictroSs A Siete subsp. ners IMI A bra. A nate, IA pechylobe IMI Mb. how 114b. tanagers PAO semen Mb. monoptera Witt income. Was obakdata Sinttl hungtO WI'S letpattliene 80. sessedofis NW NW Wk. MOO rrtoponale I.b. samosa Lett mar... Lab. plutons.. Lob. mikes 1 I Lob. orksik bey:saw le& =btu@ L.. sapiens Rat esuntineas Ref. ...sae Rat spice. Rat. globose Rel. diffuse C. pavans Cat C ad. . e Ca p C.huts.v= Cat senses Cat totonanokles Cot cinema Cal. hatenbergent. Cal. spi.scents cot. sonar. T 1 Lot nascroa nge _L Lott fal sete Lo =the Lot frutiosicies Lot .anstnore JL L. oxygen, it Lot. m.ofa subep. nernaquensb Lot knexas , LI Lot. messists IJ Lot. tuna 1— , IJ Lot. la. EcIfnectb'osepela Lot min/ann. 111.oL Sabulase r.--.1 ri Lot. Rotolo. Lot. mannatiani Lot pungens r, Lot divaricata L.. LI Lot. sencopnytte Lot outshone Lot moyeri —1—j--' tot lotononddes Lot. Mk. Lot samosa Lot knee stens. lauc.lade Lot : ox.,= Lot inyolucrata eubso pactunatterts r 1"--L, Lot. reenters i_JI., LkLot elongate n plat0C.P. tol = Lot. &torahs LI ecIptessa .11.1 ea/0.w Lot Wean Lot puinsta I.ol polwephats L pontephyne -= 1—. ,—...1.1 lift mans 1 L. =In' Sew Lot SOO. st —1 ILI= i . Lot sontudinie r.....r--- Lot sax.. L ,. 1_7. 1.21., Its. P sessinfona P. aristate P pentad°. P. canuafolie , I I Rob &____U Rob. vendesystil p let tam. Cr. distane Cs griquensts Cr cepa..

' 7*°""14, Linl Cr.r lanceolate Ol 8. ornhoensis il O. prangs

FIG. 11.4 Parsimony-based reconstruction of the presence or absence of stipules on the 370

minimal length trees from the combined molecular/morphological analysis presented in Chapter

334 CHAPTER 11: GENERAL CONCLUSIONS

FLOWERS—Flowers often provide important characters in legumes (Polhill 1976).

In the Crotalarieae two floral types are found. Most genera have specialised flowers with petals that are reflexed to various degrees. Pearsonia, Robynsiophyton and Rothia have gullet type flowers where the floral parts are rather unspecialised with a straight or down-curved style and uniform anthers. Overall the flowers are largely yellow, but some species of Aspalathus, Lotononis s.s., Pearsonia and Crotalaria have blue, purple, white or pink flowers. It may be interesting to investigate anthocyanins in these groups to identify the compounds responsible for colour variation and their homology.

CALYx—The Crotalarieae differ from Genisteae in the absence of a two-lipped calyx. The lotononoid calyx found in some Crotalarieae was reconstructed as an apomorphy for the Lotononis clades and Pearsonia, whereas a reversal to the more lebeckioid calyx type is found in Rothia and Robynsiophyton (Fig. 8.6). The carinal lobe of the calyx is narrower than the other lobes in most genera of the tribe and very pronounced in some Rafnia species where the carinal lobe is extremely thin and almost hair-like (Campbell and Van Wyk 2001). Lebeckia, however, has the carinal lobe equal to the other lobes, i.e. an equally lobe calyx (Le Roux 2006).

KEEL—Rostrate keel petals appear to have developed several times in the

Crotalarieae (Fig. 8.7). A beaked keel is often associated with a more pronounced differentiation between the stamens (Polhill 1976) where species with beaked keel petals tend to have the carinal stamen either similar to the long, basifixed anthers or somewhat intermediate between the long and short stamens.

ANTHER ARRANGEMENT—Three basic arrangements were identified within the

Crotalarieae (Fig. 8.8): 5+5 (carinal anther resembles the basifixed anthers) present in

335 CHAPTER 11: GENERAL CONCLUSIONS

Bolusia, Crotalaria, Ezoloba and Lebeckia; 4+1+5 (the carinal anther intermediate in between the dorsifixed and basifixed ones) present in Aspalathus, Calobota, Lotononis,

Listia, Rafnia and Wiborgia; 4+6 (carinal anther resembling the dorsifixed anthers) present in Euchlora, Leptis, Lotononis and Wiborgiella. Anther characters in combination with other data are very useful at the generic level and were very important in segregating Calobota and Wiborgiella from Lebeckia s.s. Most genera of the "Cape group" as well as Lotononis macrocarpa have a 4+1+5 anther arrangement with the exception of Lebeckia (5+5 configuration) and Wiborgiella (4+6 configuration). In

Lotononis s.s. and the Leobordea Glade both a 4+1+5 and 4+6 is found, while the Listia group has an exclusively 4+1+5 arrangement. Crotalaria and Bolusia share a 5+5 anrrangement, while Lotononis hirsuta differs from these in its 4+6 arrangement.

GYNOECIUM—Straight styles are a unique feature found in Pearsonia, Rothia and

Robynsiophyton. The other genera in the tribe Crotalarieae have upcurved styles.

Crotalaria is unique within the tribe in having hairs on the style as opposed to the glabrous styles of the other genera. This style-brush facilitates the "pollen pump" mechanism which is found in flowers of Crotalaria where the pollen is shed into the keel before the flowers are fully open and then pushed out by the hairs of the style when the keel is depressed by a pollinator (Polhill 1982).

FRUIT—Fruit types within the Crotalarieae are variously modified, but some general trends can be observed. A verrucose upper suture of the pod was thought to be unique to Lotononis s.l., but in light of the new generic classification for the genus this character is convergent between the Leobordea Glade, Lotononis s.s., L. hirsuta

(section Euchlora) and L. macrocarpa (Fig. 8.9).

336 CHAPTER 11: GENERAL CONCLUSIONS

Aspalathus and Wiborgia both have few-seeded (1 to 2-seeded) fruit (Fig. 11.5) that are sporadically found in Calobota and some species of the Leobordea Glade

(Lotononis section Synclistus).

The shape of the pods in transverse section (inflated, terete or laterally compressed) is informative at especially the generic level, with Bolusia, Crotalaria and

Euchlora, i.e. the Crotalaria Glade, all having inflated pods, as well as most species of

Wiborgiella (Fig. 11.6). The Lotononis clades have predominantly semi-terete pods and the Pearsonia Glade differ in their laterally compressed pods. In most of the "Cape group" the pods are laterally compressed. The samaras found in Wiborgia represent a unique fruit-type in the tribe and also a generic apomorphy for the genus. Lebeckia meyeriana, L. plukenetiana and some Rafnia species are the only other examples of winged fruit in the Crotalarieae and their pods are oblong, in contrast to the ovoid pods of Wiborgia species.

The anatomy of the fruit wall is an interesting character that deserves further investigation within the Crotalarieae. Most of the genera have thin-walled pods, but the fruit wall structure is not necessarily similar. The presence or absence of sclerenchyma and the amount of sclerification when present differ between genera. Thick-walled pods with a spongy inner surface are found in Lebeckia and Calobota but may also be present in species of other genera such as Lotononis s.s. (e.g. L. pallens).

SEEDS—The seeds of members of Genisteae are usually transversely oblong in contrast to the oblong seeds found in Crotalarieae. Seed characters are not very informative between genera but may be valuable at the species level. A rugose seed surface, for example, is an important generic character for Lebeckia, but occurs

337 CHAPTER 11: GENERAL CONCLUSIONS sporadically in other genera such as Calobota, Lotononis s.I., Rafnia, Rothia and

Wiborgiella.

CHROMOSOME BASE NUMBER—The reconstruction of chromosome base numbers is shown in Fig. 8.11. A base number of x=9 seems probably for the tribe with x=8 in

Crotalaria, Rafnia and some species of Aspalathus and x=7 in Pearsonia, Rothia and

Lotononis s.s. It is possible that this pattern may become more complex as more counts become available if for example material of L. macrocarpa (Ezoloba) can be found.

CHEMISTRY—The chemical data available for the Crotalarieae are very informative and provide additional insight into generic relationships. Quinolizidine alkaloids are present in most genera of the tribe, but absent from Lotononis s.s. and

Crotalaria (Fig. 11.7). This is due to the presence of macrocyclic pyrrolizidine alkaloids in these genera (see Fig. 8.13). In Lotononis s.s. and Crotalaria the quinolizidine alkaloid biogenetic pathway appears to have been replaced by a macrocyclic pyrrolizidine pathway. Species of Lotononis s.s. furthermore produce cyanogenic glucosides (Fig. 8.12), a trait not found in any other genus of the tribe. The accumulation of esters of alkaloids in Pearsonia, Robynsiophyton and Rothia lends support to the close relationship shown between these genera. Seed alkaloids and seed flavonoids are worth while exploring in future due to the conservative nature of these compounds in the seeds versus the leaves. This may erradicate the problem of overly complex chemical patterns that are not chemotaxonomically useful. The flavonoids of the Crotalarieae are especially poorly studied and in light of the utilisation of some species as teas it would be interesting to ascertain their phenolic chemistry.

338 CHAPTER 11: GENERAL CONCLUSIONS

The diversity and taxonomic complexity of the Crotalarieae mentioned by Polhill

(1976), Dahlgren (1970a) and Van Wyk (1991a) have been highlighted by the results of this study. Molecular characters in combination with the vast volume of morphological and chemical data for the Crotalarieae were extremely useful in unravelling the phylogenetic structure of the tribe and has led to a new generic classification that is both practical and highly predictable. In light of the overlap of morphological characters and lack of generic apomorphies informative combinations of characters are available to distinguish between the crotalarioid genera.

339 •

CHAPTER 11: GENERAL CONCLUSIONS

= many-seeded t=e CAS A samosa .1mM A Gamow = few-seeded ;Meets A. 500000 A. nuonarthe A roncrenthe A. readanandana A mettlenociene A obtuse INN A. =Was A. pertelbs, sump. pl=pell A pedalo:a onnsp. A. crenate A. ',coals A. awricarate A. vormicubta A staaal stag Mewl' A. Waal euteg sheaf A larraloPe wimp. lencdoele A. Oancelake .00w. Waal, A. Moe slaw. Mte A. WM sub*, lata ia Walla WWII A now A.Mmaybom be A Pectlybbe Woe 1Wb. Stnutea e WM. tetraptere 4 Mb. settee WM. mime . ..oPer a WM.rnonopteras WtMW®. Wor e W50.int:uncle WC obcordets obconlate WIN. Menge WW. WU besoldtlene 11/101.Woo:Wane WM awe-WM WW. sessIttofia WU1405 barbene Wbt bombs. n .. 'Meta , 1 1.44bl. abets Illibl mumurate Mat mumwele s camas J Lab. samosa Isb. mummy Lab. oblemationa Lob. plukenotlene LAA laualCore , I Leb. pawns" encl.* 1_1_1 tell bramerpe Lab. bre* "tts pate Lea. ambigua eb. septa.. ,Ref wombat', RM. ocumbete Ratwows. P.O. mama. Rat Waste Ret. globose A. SPbt0000 Re! Wane Rat &Rosa ,—,...s 1,1 Cal pungen I =_CCal pungens ..77,. __Cel. cusoldose Cat OLIO. .1. Gl =1 Cal. Cal. sottea =se icsa ' Cat Oafananaides Ca tolonanoider, Cal ammo anew = --H—r— Cal hnlenhergensa Col helenbargensb Ca, winescons Cal spineacens Col sohereo CM whew, Lot memouspe I. Lot nuaroceme Lot lekata L.Lotw:. 1:::....h r—F-4 1-1=='"' n od. Lot. panbore LOt wawa. Lot spaadlom Lot wawa LOI. Lot manta subsp. newouensis Lot =maw. nartuesuenbs Lot *Ma& Lot 'engage Lot. meculate 0000140 Lot awl 0. court.il Lot. Ora r--.1 Li Lotlem Ld. Lot. Lot memwepais Lot. memosepale Lot mamba LU'ret. sabab"'n'Ir II Lot satoulosa loptokaa Lac bobsled, Lot noWnIllonl Lothti mealmabn‘.,... 1 Lot pungens LotWakens ,__Lj Lot dense Wrap. butoolada Lot seticephyea La.= Lot pulchate Lac mewl kmolumea wasp. pedunculana Lot lotononobee Lot. 'Womb Lot strate Lot. along,. 1-1.1 La samosa . Lot. &web= ,_1.1 Lot. dense sulaP. lensocisde Lot serkapnylle EI—IJ Lot. erstlaWate Lot pieta& 11 Lot maim:rate Lot owed Lot involwate rant, pedunoulerts Lot lotonowedes Lot Mamie -Jer Lot. WWII Lct elongate Lot. samosa LOt plotybarpe -j n Lot pletycabe = 1J Lotolts'. celyelne Lot cWow Ld. +0000 Lot. 'Menthe Lot otct Lot. nomenbona can 'I Lot scutterce Lot earticame : bolusil toy = *nob Lot coyote's/ale L polYwPW" Lot. penteptctle Lot pontephylle ... Lot. nada Lot. mobs Lot scare r—r7:Ii tot.

lot dgdab Lot regratce r Lm tot. glonutoen Lot gionulose Loa wbuttints 'lot sokfudinis Lot.Lot 1010 WWI. Lotsubulets Lot £0141 Lot ~WM medal* P. sesseabla P. sewisbrie erlafete X11 P. =at cAig""dlenifolit" Rot raw. Rot Wawa Rot. vondarysfil Rob. vandaryth/ , Lot hirwu Lot hirsute Cf. datana Cr. &stem , Cr gamonels Jr 91_G OWW.W. mews. -•• Cr. vbcurtails ., Cr humills Cr. Walla Cr Lanceolate Ca lanceolate emboormle _ emboansis 0. great. .1 O grease

FIG. 11.5 Parsimony-based reconstruction of few- versus many seeded fruit on the 370 minimal

length trees from the combined molecular/morphological analysis presented in chapter 8.

340 •

CHAPTER 11: GENERAL CONCLUSIONS

:1 = laterally compressed 1 A bowls = semi-terete A =toss 0 A. esteem = tents .;l1be v rre = inflated r=, r—L PlAreloff •ia= h'" ie=tels. febiesi A. verecceles roorsaLlalana3 A.10009 subsp. ewes A. shooll mese. Omni A. ebb" sm. lartatila torrobea BRap. Ware's . IOWA sub.. hela A /weft ,11 i AlsA..r- ebetshr:o;:. teto A Num I A. pacheota wmWitt. twoomlbsca WO. Setweett lAbb. seems Mb maned monoptora Wit. mance= Web. locums. WasWooten Witx aboorties Mb. Weeds. Wit Moan Pesseles WLY labsaddllorm letperkent WA amoeba!' bosierans ima Irtb3117 seal mtmanah Lob. comma Lat. n7d1,2411.1a PUNINICSTIO peuetbro Vb. peoclbre L-1 LO. boMemPa ark.'"amoros'Wgh:I'W' Rat menials Rat mammas II 1—=', raeonsa R Tetlebr — 1 Red dates, RRWEZI RCM. Cal pongees =_.1CoLeonora. 1--m —, 1 Col= Met fl= Pat'. 14t= 13 Cal. *atm Cat Cal lolonanaldes lototanoides Cal ammo Cal bakmbamaneto Cal befortbeeponala C. spettcoms C. lam. Catmb ar. Cal soh.. LlmanLorrea II Let mormeellta balbeleal, 1---1-1-1 Lot. bpr=he 1-1 2 01. Ows, Lolta lrefisAlos La a --MGM apersakto L•gaponelors 174-1-11,...Th i to:i Maids. E,subsp. BaBlarplefld.B LLZ. rem sctsp. nentsawses Lot 0000macesta tot meat. Lot arbt usLat:1am 0. LoL Aos r-- Les. mesosepots Lot mosesspala totmiorretts 1.6. LoL blabra Lot SII,XIOda osbuttms La laplekSta " .. 13 Lot *Maw Lot. modmillsol Lot maximlbooi purports Lot memo. Lo oiwerketa ot ia Lot cerleatebta Lot etleltalia Lot polchabis mspto _at latornmokleam"W Lot loteommilso —El Lel. strIcta Lot. OKRA Lot aroma Lot corm. Lot. dams weep. leamadola Lot amca sotcp. Samoa*. Lot. La. ee(CVlob 0000 e.wluaab~ (atop. too nocrorA Lot braSoorrls sabste Lot tOsors. m etartgoto Lot pta, yeama L S. Wpm(( L. Wpm"?' Lot docombans Lot aralboeses Lot Wu. Lot me/WA polmonola G mmtabbytt

Lot bestbantiona Lot CNVIO Lct IrrarifICO Lot ;Must., ' msbuttallo Lot auladola Lot marlatlis. sambablia P. grandee& eataaltato Rot. Ro reedryalb t. UM. Cr. &dans Cr "sorrels Cr emends Cr. Wombat Cr Meath 00 lanceolsba D. owes

FIG. 11.6 Parsimony-based reconstruction of fruit shape in transverse section on the 370

minimal length trees from the combined molecular/morphological analysis presented in chapter

341

CHAPTER 11: GENERAL CONCLUSIONS

g---r AP.Pdc.'nm"! LJ = present I 'll:Zr 1.1 = absent _i_r_i_c__, ate"1,' I 4sirp. — A. pagoolMt ode?. ph.Cloa , '—' A. clones ii--1 `JJ. 4..1 ...,sortobtletozz 1_--1—; , ;:rd''' I= Iiir ft.rx: .fr 1 t lotoototro 1---1 E_, at Mt aortae ttib. tootnato LIRJ otvorstra , t---- WS.t. brorCa _1 -, — A.L. sto.ltortet Sliti...... 11. hxlcosto WX Mora I 11.10,...groextro L LeS:. to Lott p = 7: ' t ' Le'1.46. Wvt'Putooltdr.k. -ob. troptaeht I_____I ■ ...._,i,_,,'' c Ro....7. ostOORtoo'''''''... .- t—. ttle. &boo. Re. a.... _Li EX MAWR

I L"-- CV(X" MI'ottrzett7

LI —7-7-7' .--LL Catt. totorrsCdes: Cat ass. Li 1-7 : ri 0 CAO spRottosea Ca' .htusa LS. mayo.. ..ifi 11 toe. Citz ... n.

Lot re. L.... tot. trer. Lot f•o MIMS, notragoonsis Lot lortOrtifx La menads Lat. cprli 11.1. it Orgy* L.L messonosto LAt stkreolha 11 L. =Mt= ktoteriX&t i la otodootom 1.0. Amon, Lot Lot tortetrOoat pokboa Lot. rrtoyen tot. banonottoo Lot. &trio La carom La. rim. whop. lostodszte Lot croftput. tot ftwtoot

Lot Lce.

I F-1. flamers/a t, Lot otottorpo Llott oo., _f Li= Let. -mdetw. T t Lot ottotootooto — tce.f ft moth at. borilhafRooe / crt otOrtoo r iot = tot axfa .. boo sataoto Utt. No* Lot, mend. !R ...Clots ' P oICdL P 6 P Otlank.A. Rot. hirouto Rob. tand000dt Lot &tato Co atatsa Cr. Lsty.... at mortars Cf. Royettfis MsfLa Cr. tinumktIo ort.tors. U omen,

FIG. 11.7 Parsimony-based reconstruction of the presence or absence of quinolizidine alkaloids

on the 370 minimal length trees from the combined molecular/morphological analysis presented

in chapter 8.

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364 APPENDIX Al. Voucher specimens from which floral dissections were made in order to study and illustrate reproductive morphology.

Calobota acanthoclada: Dinter 6269 (BOL); Mannheimer CM2012a (WIND);

Williams 2594a (BOL); Williams 3376 (BOL). C. cinerea: Boatwright et al. 150

(JRAU); Dinter 3998 (BOL); Jurgens 2234 (PRE); Le Roux 2066 (NBG);

Mannheimer et al. CM 899 (WIND); Merxmilller and Giess 3089 (WIND). C. cuspidosa: Acocks 2455 (BOL); Ethel Anderson 591 (BOL); Gubb 12584 (PRE);

Van Wyk BSA1241 (PRE); Van Wyk 3055 (JRAU) 3 dissections. C. cytisoides:

Andrag 272 (PRE); Barker 6426 (NBG); Barker 9694 (NBG); Boatwright et al. 114

(JRAU); Mauve and Oliver 136 (PRE); McDonald 1568 (PRE); Pillans 7701 (BOL);

Pretorius 105 (NBG); Schutte 262 (JRAU); Schutte 286 (JRAU); Stirton 6167 (PRE);

Van Breda SKF574 (PRE); Van Wyk 2568 (JRAU). C. elongata: Hall 177 (NBG);

Van Breda 4486 (PRE); Van Wyk 2229 (JRAU). C. halenbergensis: Bean and

Oliver 2445 (BOL); Boatwright et al. 146 (JRAU); Esterhuysen s.n. (NBG); Giess

13820 (WIND); Johnson 22071 (NBG); Jurgens 22739 (PRE); Mannheimer CM2418

(WIND); McDonald 739 (NBG); Middlemost 2129 (NBG); Muller 810 (PRE) 2 dissections; Muller 830 (WIND); Van Wyk 3086 (JRAU); Werger 431 (PRE); Salter

5549 (PRE). C. linearifolia: Acocks 13195 (PRE); Braine 98 (WIND); Dean 673

(JRAU); Evrard 9285 (PRE); Giess 13803 (WIND); Keet 1663 (WIND); Merxm011er and Giess 3615 (PRE); Snijman 214 (NBG). C. lotononoides: Boatwright et al. 142

(JRAU); Perold 1646 (PRE); Schlechter s.n. sub TRV 1059 (PRE); Schlechter 8214

(PRE). C. namibiensis: Kolberg and Maggs 12 (WIND); Owen-Smith 117 (WIND);

Owen-Smith 1268 (WIND). C. obovata: Kers 152 (WIND); Pearson 8036 (BOL). C. psiloloba: Archibald 3848 (GRA); Story 3353 (PRE); Troughton 425 (PRE); Van

365 APPENDIX Al

Wyk s.n. (JRAU) 3 dissections; Zeyher 31438 (NBG). C. pungens: Boatwright et al.

106 (JRAU); Bond 12112 (NBG); Markotter 9951 (NBG); Sidey 1906 (PRE);

Snijman 353 (NBG); Taylor 19386 (NBG); Thompson 9948 (NBG); Van Wyk 2147

(JRAU); Van Wyk 3252 (JRAU). C. saharae: Hill 1910 (K); Keith 181 (K) 2 dissections. C. sericea: TYPICAL FORM—Boatwright et al. 139 (JRAU); Compton

19906 (NBG); Hall 899 (NBG); Hall 3701 (NBG); Hilton Taylor 1366 (NBG, PRE);

Leipoldt 3765 (NBG); Lewis 4705 (NBG); O'Callaghan and Steensma 1650 (NBG);

Salter 5612 (BOL). BROAD-LEAVED FORM—Arnold 916 (PRE); Clark 633 (PRE);

Heywood s.n. (NBG); Van Wyk 2847 (JRAU). SHORT-LEAVED FORM—Clark and

Mailer 337 (WIND); Giess and Van Vuuren 678 (WIND); Giess and Robinson 13177

(WIND); Mannheimer CM2020 (WIND); Watmough 867 (WIND); Whitehead 80.9.8

(WIND); Williamson 2575 (BOL).ORANJEMUND FORM—Gess and Gess 97/98/68

(WIND); Ward 11136 (WIND); Williamson 4884 (WIND). C. spinescens: TYPICAL

FORM—Boatwright et al. 158 (JRAU); Koekemoer 248 (JRAU); Leistner 1897 (BOL);

Wilman 838 (PRE); Zietsman 3489 (PRE). PRIESKA FORM—Bryant 5404 (BOL);

Maguire 742 (NBG); Van Wyk 3081 (JRAU); Vlok 1932 (JRAU); Williams 2594a

(BOL). C. thunbergii: Boatwright et al. 141 (JRAU); Boatwright et al. 151 (JRAU);

Germishuizen 4839 (PRE); Hall 891 (NBG); Johnson 200 (NBG); Manning s.n.

(NBG); Schutte 591 (JRAU); Schutte 593 (JRAU); Thorns s.n. (NBG); Van Jaarsveld

11977 (PRE); Van Wyk 2584 (JRAU). Pearsonia cajanifolia subsp. cryptantha:

Posthumus 1b (JRAU). Pearsonia sessilifolia: Swanepoel 1 (JRAU).

Robynsiophyton vanderystii: Lisowski 20326 (K); McCallum Weston 717 (K);

Richards 5309 (K); Richards 9321 (K). Rothia hirsuta: Akpabla 1982 (K); Bidgood et al. 1194 (K); Davies 1296 (K); Davey 173 (K); Greenway & Kanuri 14269 (K);

Hepper 1021 (K); Horndy 20206 (K); Jackson and Apejoye 413973 (K); Polhill and

366 APPENDIX Al

Paulo 2089 (K); Robinson 151 (K); Saadou 1798 (K); Saime 162 (K). R. indica:

Cooray 70020117R (K); Latz 16126 (MEL); Wight 571 (MEL); Wight 828 (K); Wight

5821 (K); Von Muller s.n. (MEL). Wiborgiella aspalathoides: Barker 10407 (NBG)

3 dissections. W. bowieana: Streicher s.n. (JRAU) 2 dissections. W. fasciculata:

Muir 4775 (NBG) 2 dissections. W. humilis: Boatwright et al. 216 (JRAU); Niemand

8 (JRAU); Van Wyk 3530 (JRAU). W. inflata: Belle Barker 204 (JRAU); Haynes

H420 (PRE) 2 dissections; Lewis 1816 (NBG); Vlok et al. 2 (JRAU). W. leipoldtiana: Boatwright et al. 123 (JRAU); Bolus 8974 (BOL); Hanekom 2125

(PRE); Schutte 295 (JRAU). W. mucronata: Guillarmod 8387 (GRA); Stirton 10880

(JRAU); Stirton and Zantovska 11608 (PRE). W. sessilifolia: Albertyn 498b (NBG);

Muir 2144 (PRE); Van Wyk 2120 (JRAU). W. vlokii: Vlok 2045 (PRE) 2 dissections.

367 APPENDIX A2. Voucher information for species of Calobota, Lebeckia,

Robynsiophyton, Rothia, Wiborgia and Wiborgiella used to study stem, petiole, leaf and fruit wall anatomy.

Calobota cinerea: Stem—Boatwright et al. 150 (JRAU); Petiole—Boatwright et al. 150

(JRAU); Leaf— Boatwright et al. 150 (JRAU); Fruit wall—Boatwright et al. 150 (JRAU).

C. cuspidosa: Stem — Boatwright et al. 92 (JRAU); Petiole— Boatwright et al. 92

(JRAU); Leaf— Boatwright et al. 92 (JRAU); Fruit wall—Boatwright et al. 92 (JRAU). C. cytisoides: Stem—Boatwright et al. 114 (JRAU); Petiole—Boatwright et al. 114

(JRAU); Leaf—Boatwright et al. 114 (JRAU); Fruit wall—Boatwright et al. 114 (JRAU).

C. elongata: Petiole—Van Wyk 2562b (JRAU); Leaf—Van Wyk 2562b (JRAU); Fruit

wall — Van Wyk 2562b (JRAU). C. halenbergensis: Stem — Boatwright et al. 149

(JRAU); Petiole—Boatwright et al. 149 (JRAU); Leaf— Boatwright et al. 149 (JRAU);

Fruit wall—Boatwright et al. 149 (JRAU). C. linearifolia: Leaf— Dean 673 (JRAU); Fruit wall—Giess et al. 6180 (WIND). C. lotononoides: Stem—Boatwright et al. 142

(JRAU); Petiole—Boatwright et al. 142 (JRAU); Leaf— Boatwright et al. 142 (JRAU);

Fruit wall—Boatwright et al. 142 (JRAU). C. namibiensis: Leaf—De Winter and Hardy

7919 (WIND); Fruit wall—De Winter and Hardy 7919 (WIND). C. obovata: Petiole-

Kers 152 (WIND); Leaf—Kers 152 (WIND). C. pungens: Stem—Boatwright et al. 106

(JRAU); Petiole—Boatwright et al. 106 (JRAU); Leaf— Boatwright et al. 106 (JRAU);

Fruit wall—Boatwright et al. 106 (JRAU). C. psiloloba: Stem—Le Roux et al. 20

(JRAU); Petiole—Le Roux et al. 20 (JRAU); Leaf—Van Wyk s.n. (JRAU); Fruit wall—Le

Roux et al. 20 (JRAU). C. saharae: Stem — Keith 181 (K), Alleizette s.n. sub K

368 APPENDIX A2

2007/014804 (K); Leaf—Pitard 3276 (K), Alleizette s.n. sub K 2007/014804 (K),

Bourgeau 210 (K); Fruit wall — Alleizette s.n. sub K 2007/014804 (K), Davis 49544 (K).

C. sericea: Stem—Boatwright et al. 138 (JRAU); Petiole—Boatwright et al. 138

(JRAU); Leaf—Boatwright et al. 138 (JRAU); Fruit wall—Boatwright et al. 138 (JRAU).

C. spinescens: Stem—Boatwright et al. 158 (JRAU); Petiole—Boatwright et al. 158

(JRAU); Leaf—Boatwright et al. 158 (JRAU); Fruit wall—Boatwright et al. 158 (JRAU).

C. thunbergii: Stem—Boatwright et al. 151 (JRAU); Petiole—Boatwright et al. 141

(JRAU), Boatwright et al. 151 (JRAU); Leaf—Boatwright et al. 151 (JRAU); Fruit wall—

Boatwright et al. 151 (JRAU). Crotalaria sp.: Fruit wall—De Villiers 106 (JRAU).

Lebeckia ambigua: Stem—Boatwright et al. 132 (JRAU); Fruit wall—Boatwright et al.

131 (JRAU). L. brevicarpa: Fruit wall—Le Roux et al. 4 (JRAU). L. contaminata: Fruit wall—Le Roux et al. 16 (JRAU). L. pauciflora: Fruit wall—Le Roux et al. 12 (JRAU). L. wrightii: Fruit wall—Johns 163 (JRAU). Melolobium alpinum: Fruit wall—Schulte 160

(JRAU). Rafnia amplexicaulis: Fruit wall—Campbell and Van Wyk 40 (JRAU). Rothia hirsuta: Stem—Davies 1296 (K); Petiole—Davies 1296 (K), Greenway and Kanuri

14269 (K), Bogdon 2205 (K); Leaf—Davies 1296 (K), Greenway and Kanuri 14269 (K),

Bogdon 2205 (K); Fruit wall — Bogdon 2205 (K), Davies 1296 (K), Greenway and Kanuri

14269 (K). R. indica: Stem — — ; Petiole — Ramamoorthy 1956 (K), Gamble 13771 (K),

Latz 16126 (MEL); Leaf—Ramamoorthy 1956 (K), Gamble 13771 (K), Latz 16126

(MEL); Fruit wall—Ramamoorthy 1956 (K), Gamble 13771 (K), Latz 16126 (MEL).

Robynsiophyton vanderystii: Stem—Lisowski 20326 (K); Petiole—Exell and

Mendonga 657 (K), Richards 18144 (K), Lisowski 20326 (K); Leaf— Exell and

Mendonga 657 (K), Lisowski 20326 (K), Richards 18144 (K); Fruit wall — Exell and

369 APPENDIX A2

Mendonga 657 (K), Lisowski 20326 (K), Richards 18144 (K). Wiborgia monoptera:

Fruit wall—Boatwright et al. 152 (JRAU). W. sericea: Stem—Boatwright et al. 124

(JRAU); Petiole—Boatwright et al. 124 (JRAU); Leaf—Boatwright et al. 124 (JRAU);

Fruit wall—Boatwright et al. 124 (JRAU). Wiborgiella bowieana: Petiole—Streicher s.n. sub Schutte 831 (JRAU); Leaf—Streicher s.n. sub Schutte 831 (JRAU); Fruit wall-

Streicher s.n. sub Schutte 831 (JRAU). W. humilis: Stem—Boatwright et al. 129

(JRAU); Petiole—Boatwright et al. 129 (JRAU); Leaf— Boatwright et al. 129 (JRAU);

Fruit wall—Boatwright et al. 212 (JRAU). W. inflata: Stem—Johns 162 (JRAU);

Petiole—Johns 162 (JRAU); Leaf—Johns 162 (JRAU); Fruit wall—Johns 162 (JRAU).

W. leipoldtiana: Stem—Boatwright et al. 123 (JRAU); Petiole—Boatwright et al. 123

(JRAU); Leaf—Boatwright et al. 123 (JRAU); Fruit wall—Boatwright et al. 123 (JRAU).

W. mucronata: Petiole—Vlok 1726 (JRAU); Leaf—V/ok 1726 (JRAU); Fruit wall-

Esterhuysen 6880 (BOL). W. sessilifolia: Stem—Van Wyk 2120 (JRAU); Leaf—Van

Wyk 2120 (JRAU); Fruit wall—Taylor 4329 (PRE). W. vlokii: Petiole—V/ok 2045 (PRE);

Leaf—V/ok 2045 (PRE); Fruit wall—V/ok 2045 (PRE).

370 APPENDIX A3. Voucher information and GenBank accession numbers of the taxa sampled in this study. Voucher specimens are deposited in the following herbaria:

Australian National Herbarium (CANB), University of Johannesburg Herbarium (JRAU),

Kew Herbarium (K), National Herbarium, Pretoria (PRE), Southern Cape Herbarium

(SCHG). The information is listed as follows: taxon— GenBank accessions: rbcL, ITS;

Voucher specimen. Those regions not sampled for a taxon are represented by a dash

['Boatwright et al. (2008b); 2Crisp et al. (2000); 3Kass and Wink (1997); 4Doyle et al.

(1997); 5Pardo et al. (2004); 6Motsi (2004); 'Van der Bank et al. (2002); 5Le Roux

(2006); *Sequenced by J.S. Boatwright; **Sequenced at University of Heidelberg].

Amphithalea Eckl. & Zeyh.: A. alba Granby— AM180171 1 , AM261217 1 ; Van Wyk

2125 (SCHG). A. micrantha (E.Mey.) Walp.— AM180182 1 , AM261227 1 ; Schutte 751

(SCHG). A. williamsonii Harv.— AM177372 1 , AM261436 1 ; Euston - Brown s.n. (SCHG).

Argyrolobium Eckl. & Zeyh.: A. harmsianum Schltr. ex Harms— —, AF287685 2; Crisp

9042 (CANB). A. lunare (L.) Druce 1— —, AF287686 2; Crisp 9039 (CANB). A. lunare

(L.) Druce subsp. sericeum (Thunb.) T.J.Edwards 2— EU348013**, —; Van Wyk 2080

(JRAU). A. marginatum Bolus— EU348014**, —; Van Wyk 3038 (JRAU). A.

marginatum Bolus — Z955473, — ; Edwards 471. A. megarrhizum Bolus— EU348015**,

—; Germishuizen s.n. (JRAU). Argyrolobium sp. — EU348016**, — ; Van Wyk 2173

(JRAU). Aspalathus L.: A. acutiflora R.Dahlgren — — , EU347735*; Boatwright &

Magee 22 (JRAU). A. carnosa Berg. — EU348022*, EU347723*; Boatwright & Magee

61 (JRAU). A. confusa R.Dahlgren— EU348031*, EU347736*; Boatwright et al. 97

(JRAU). A. crenata (L.) R.Dahlgren— EU348028*, EU347729*; Boatwright & Magee 70

371 APPENDIX A3

(JRAU). A. galeata E.Mey.— EU348032*, EU347726*; Boatwright et al. 100 (JRAU). A. hirta E.Mey. subsp. hirta— EU348025*, EU347732*; Boatwright & Magee 43 (JRAU).

A. hystrix L.f.— EU348023*, EU347734*; Boatwright & Magee 30 (JRAU). A. laricifolia

Berg. subsp. laricifolia— EU348026*, EU347731*; Boatwright & Magee 46 (JRAU). A. linearis (Burm.f.) R.Dahlgren 1— EU348020*, EU347722*; Van Wyk et al. 4192

(JRAU). A. linearis (Burm.f.) R.Dahlgren 2— EU348019**, EU347739**; Van Wyk s.n.

(JRAU). A. macrantha Harv.— EU348029*, EU347728*; Boatwright & Magee 71

(JRAU). A. nivea Thunb.— EU348018**, EU347737**; Van Wyk 2938 (JRAU). A. pachyloba Benth.— EU348024*, EU347733*; Boatwright & Magee 42 (JRAU). A. pendula R.Dahlgren— EU348017**, EU347738**; Van Wyk s.n. (JRAU). A. perfoliata

(Lam.) R.Dahlgren subsp. phillipsii R.Dahlgren— EU348030*, EU347727*; Boatwright

& Magee 82 (JRAU). A. shawii Bol. subsp. shawii— EU348033*, EU347725*;

Boatwright & Magee 27 (JRAU). A. vermiculata Lam.— EU348021*, EU34772*;

Boatwright & Magee 15 (JRAU). A. willdenowiana Benth.— EU348027*, EU347730*;

Boatwright & Magee 69 (JRAU). Bolusia Benth.: B. acuminata (DC.) Polhill-

EU347944**, —; Morris 1193 (PRE). B. amboensis (Schinz) Harms 1— EU347943**,

EU347891**; Robinson 1558 (K). B. amboensis (Schinz) Harms 2— EU347943**, —;

Giess 10091 (K). Cadia Forssk.: C. purpurea (Ait.) Forssk.— AM260751 1 , AM261740

1 ; Beckett 1702 (K). Calpurnia E.Mey.: C. sericea Harv.— AM177374 1 , AM268374' &

AM268375 1 ; Boatwright 86 (JRAU). Crotalaria L.: C. capensis Jacq. 1— EU348034**,

EU347884**; Van Wyk 2933a (JRAU). C. capensis Jacq. 2— EU348045*, EU347888*;

Van der Bank 14 (JRAU). C. distans Benth.— EU348038**, EU347882**; de Castro

137 (JRAU). C. griquensis Bolus— EU348043*, EU347887*; Van Wyk et al. 4173

372 APPENDIX A3

(JRAU). C. humilis Eckl. & Zeyh. 1— EU348041*, EU347889*; Boatwright et al. 143

(JRAU). C. humilis Eckl. & Zeyh. 2— EU348042*, EU347890*; Boatwright et al. 156

(JRAU). C. lanceolata E.Mey.— EU348040**, EU347885**; Van Wyk 1985 (JRAU). C. lebeckioides Bond— EU348036**, —; Van Wyk 3315e (JRAU). C. lotoides Benth.—

EU348035**, —; Van Wyk s.n. (JRAU). Crotalaria sp.— EU348039**, EU347883**; Van

Wyk 3367 (JRAU). C. virgultalis Burch. ex DC. 1— EU348044*, EU347886*; Van Wyk et al. 4175 (JRAU). C. virgultalis Burch. ex DC. 2— EU348037**, —; Van Wyk 3060

(JRAU). Cyclopia Vent.: C. genistoides (L.) R.Br.— AM261716 1 , AM050819 1 ;

Boatwright & Magee 53 (JRAU). C. subternata Vogel— AM261725 1 , AM050821 1 ;

Boatwright & Magee 35 (JRAU). Dichilus DC.: D. gracilis Eckl. & Zeyh.— EU347962**,

EU347893**; Schutte 247 (JRAU). D. lebeckioides DC.— EU347963**, EU347894**;

Schutte 151 (JRAU). D. pilosus Conrath ex Shinz— EU347959**, EU347892**; Schutte

363 (JRAU). D. reflexus (N.E.Br.) A.L.Schutte— EU347961**, —; Schutte 177 (JRAU).

D. strictus E.Mey.— —, AJ2876842; Crisp 9073 (CANB). D. strictus E.Mey.—

EU347960**, —; Schutte 150 (JRAU). Genista L.: G. teretifolia Willk.— —, AY263668 5;

MAF 162924. Lebeckia Thunb.: L. ambigua E.Mey. 1— EU347917**, EU347852**;

Van Wyk 2900 (JRAU). L. ambigua E.Mey. 2— EU3479348, EU347851 8; Le Roux et al.

6 (JRAU). L. bowieana Benth. [Wiborgiella bowieana (Benth.) Boatwr. & B.-E.van Wyk]

1— EU347909*, EU347868*; Streicher s.n. (JRAU). L. bowieana Benth. 2—

EU347910**, EU347869**; Van Wyk 2106 (JRAU). L. brevicarpa M.M.le Roux & B.-

E.van Wyk— EU347933 8, EU3478508; Le Roux et al. 4 (JRAU). L. carnosa (E.Mey.)

Druce 1— EU347900**, EU347845**; Vlok et al. s.n. (JRAU). L. carnosa (E.Mey.)

Druce 2— EU347937 8, EU3478468; Le Roux et al. 15 (JRAU). L. carnosa (E.Mey.)

373 APPENDIX A3

Druce 3— —, EU3478478; Le Roux et al. 16 (JRAU). L. cinerea E.Mey. [Calobota cinerea (E.Mey.) Boatwr. & B.-E.van Wyk]— EU347930*, EU347840*; Boatwright et al.

150 (JRAU). L. cytisoides Thunb. [Calobota cytisoides (Berg.) Eckl. & Zeyh.] 1—

EU347903**, EU347837**; Van Wyk 2313 (JRAU). L. cytisoides Thunb. 2—

EU3479258, EU3478388; Le Roux et al. 2 (JRAU). L. gracilis Eckl. & Zeyh.—

EU3478558; Le Roux et al. 17 (JRAU). L. halenbergensis Merxm. & A.Schreib.

[Calobota halenbergensis (Merxm. & A.Schreib.) Boatwr. & B.-E.van Wyk] 1—

EU347908**, EU347836**; Van Wyk 3086 (JRAU). L. halenbergensis Merxm. &

A.Schreib. 2— EU347927*, EU347842*; Boatwright et al. 146 (JRAU). L. inflata H.

Bolus [Wiborgiella inflata (H. Bolus) Boatwr. & B.-E.van Wyk] 1— EU347901**,

EU347863**; Vlok et al. 2 (JRAU). L. inflata H. Bolus 2— EU347940*, EU347864*;

Johns 162 (JRAU). L. leipoldtiana Schltr. ex R.Dahlgren [Wiborgiella leipoldtiana

(Schltr. ex R.Dahlgren) Boatwr. & B.-E.van Wyk] 1— EU347939*, EU347866*;

Boatwright et al. 123 (JRAU). L. leipoldtiana Schltr. ex R.Dahlgren 2— EU347914**,

EU347865**; Van Wyk 3278 (JRAU). L. lotononoides Schltr. [Calobota lotononoides

(Schltr.) Boatwr. & B.-E.van Wyk]— EU347928*, EU347843*; Boatwright et al. 142

(JRAU). L. macrantha Harv. [Calobota cuspidosa (Burch.) Boatwr. & B.-E.van Wyk] 1—

EU347916, EU347828; Van Wyk s.n. (JRAU). L. macrantha Harv. 2— EU347922*,

EU347831*; Boatwright et al. 92 (JRAU). L. melilotoides R.Dahlgren [Calobota elongata

(Thunb.) Boatwr. & B.-E.van Wyk] 1— EU347919**, —; Van Wyk 2562b (JRAU). L. melilotoides R.Dahlgren 2— —, EU347841*; Van Wyk 2229 (JRAU). L. meyeriana

Eckl. & Zeyh. ex Harv. 1— EU347904**, EU347856**; Van Wyk 3351 (JRAU). L. meyeriana Eckl. & Zeyh. ex Harv. 2— EU347905**, —; Van Wyk 3009 (JRAU). L.

374 APPENDIX A3 meyeriana Eckl. & Zeyh. ex Harv. 3— —, EU3478578 ; Van Wyk 4043 (JRAU). L. mucronata Benth. [Wiborgiella mucronata (Benth.) Boatwr. & B.-E.van Wyk]—

EU347941*, EU347870*; Vlok 1726 (JRAU). L. multiflora E.Mey. [Calobota sericea

(Ait.) Boatwr. & B.-E.van Wyk]— EU347926*, EU347833*; Boatwright et al. 138

(JRAU). L. pauciflora Eckl. & Zeyh. 1— EU347902**, EU347861**; Van Wyk 3024

(JRAU). L. pauciflora Eckl. & Zeyh. 2— EU3479358 , EU3478628; Le Roux et al. 7

(JRAU). L. plukenetiana E.Mey. 1— EU347932*, EU347849*; Le Roux et al. 24

(JRAU). L. plukenetiana E.Mey. 2— EU347906**, EU347848**; Van Wyk 4043 (JRAU).

L. psiloloba (E.Mey.) Walp. [Calobota psiloloba (E.Mey.) Boatwr. & B.-E.van Wyk]—

EU347923*, EU347830*; Le Roux et al. 20 (JRAU). L. pungens Thunb. [Calobota pungens (Thunb.) Boatwr. & B.-E.van Wyk] 1— EU347918**, EU347827**; Van Wyk s.n. (JRAU). L. pungens Thunb. 2— EU347921*, EU347829*; Boatwright et al. 106

(JRAU). L. sepiaria (L.) Thunb. 1— EU3479368 , EU3478538 ; Le Roux et al. 10 (JRAU).

L. sepiaria (L.) Thunb. 2— EU347920**, EU347854**; Van Wyk 2979 (JRAU). L. sericea Thunb. [Calobota thunbergii Boatwr. & B.-E.van Wyk] 1— EU347907**,

EU347832**; Van Wyk 3115 (JRAU). L. sericea Thunb. 2— EU347924*, EU347834*;

Boatwright et al. 151 (JRAU). L. sericea Thunb. 3— EU347915**, EU347835**; Van

Wyk 3119 (JRAU). L. sessililfolia (Eckl. & Zeyh.) Benth. [Wiborgiella sessilifolia (Eckl. &

Zeyh.) Boatwr. & B.-E.van Wyk]— EU347938*, EU347867*; Boatwright et al. 170

(JRAU). L. spinescens Harv. [Calobota spinescens (Harv.) Boatwr. & B.-E.van Wyk]—

EU347929*, EU347839*; Boatwright et al. 158 (JRAU). L. wrightii (Harv.) Bolus 1—

EU347912**, EU347858**; Vlok et al. s.n. (JRAU). L. wrightii (Harv.) Bolus 2—

EU347913**, EU347860**; Vlok et al. s.n. (JRAU). L. wrightii (Harv.) Bolus 3—

375 APPENDIX A3

EU347911 8, EU3478598 ; Van Wyk 3354 (JRAU). Liparia L.: L. angustifolia (Eckl. &

Zeyh.) A.L.Schutte— AM177376 1 , AM261478 1 ; Boatwright & Magee 66 (JRAU). L. hirsuta Thunb.— AM259357 1 , AM261486 1 ; Boatwright & Magee 33 (JRAU).

Lotononis (DC.) Eckl. & Zeyh.: L. acuticarpa B.-E.van Wyk— EU348047**,

EU347770**; Van Wyk 2625 (JRAU). L. adpressa N.E.Br. subsp. adpressa-

EU348100**, EU347780**; Van Wyk 1899 (JRAU). L. alpina (Eckl. & Zeyh.) B.-E.van

Wyk— —, AM262446 6; Van Wyk & Van Wyk 1478 (JRAU). L. benthamiana Dummer-

EU348051**, EU347771**; Van Wyk 2528 (JRAU). L. bolusii Dummer— EU348046**,

EU347761**; Van Wyk 2443 (JRAU). L. brachyantha Harms— EU347998**,

EU347816**; Boatwright et al. 117 (JRAU). L. brevicaulis B.-E.van Wyk— EU348080**,

—; Schutte 447 (JRAU). L. calycina (E.Mey.) Benth.— EU348052**, EU347762**; Van

Wyk 1433 (JRAU). L. carnosa (Eckl. & Zeyh.) Benth.— EU348083**, EU347800**; Van

Wyk 1663 (JRAU). L. curtii Harms— EU348006**, EU347815**; Van Wyk et al. 4172

(JRAU). L. crumanina Burch. ex Benth.— EU348062**, —; Van Wyk 3057 (JRAU). L. decumbens (Thunb.) B.-E.van Wyk subsp. decumbens— EU348058**, EU347778**;

Van Wyk 1701 (JRAU). L. densa (Thunb.) Harv. subsp. leucoclada (Schltr.) B.-E.van

Wyk— EU348048**, EU347792**; Van Wyk 2430 (JRAU). L. digitata Harv. 1—

EU348071**, EU347772**; Van Wyk 2350 (JRAU). L. digitata Harv. 2— EU348057**,

EU347773**; Van Wyk 2342 (JRAU). L. divaricata (Eckl. & Zeyh.) Benth.—

EU348092**, EU347788**; Van Wyk 2484 (JRAU). L. elongata (Thunb.) D.Dietr.—

EU348087**, EU347821**; Van Wyk 2573 (JRAU). L. eriantha Benth.— EU348086**,

EU347784**; Schutte 383 (JRAU). L. eriocarpa (E.Mey.) B.-E.van Wyk— EU348084**,

—; Van Wyk 1952 (JRAU). L. exstipulata Bolus— EU348055**, EU347796**; Van Wyk

376 APPENDIX A3

2271 (JRAU). L. falcata (E.Mey.) Benth. 1— EU348103*, EU347756*; Boatwright et al.

184 (JRAU). L. falcata (E.Mey.) Benth. 2— EU347999**, EU347817**; Boatwright et al.

120 (JRAU). L. filiformis B.-E.van Wyk— EU348074**, EU347794**; Vlok 2030 (JRAU).

L. foliosa Bolus 1— EU348073**, EU347776**; Van Wyk 2481 (JRAU). L. foliosa Bolus

2— EU347993**, EU347819**; Boatwright & Magee 88 (JRAU). L. fruticoides B.-E.van

Wyk— EU348085**, EU347801**; Van Wyk 2021 (JRAU). L. galpinii Diimmer-

Z95538 3 , -; T. Edwards 480. L. glabra (Thunb.) D.Dietr.— EU348005**, EU347814**;

Van Wyk 2014 (JRAU). L. globulosa B.-E.van Wyk— EU348075**, EU347777**; Van

Wyk 2210 (JRAU). L. hirsuta (Thunb.) D.Dietr. 1— EU348069**, EU347879**; Van Wyk

1338 (JRAU). L. hirsuta (Thunb.) D.Dietr. 2— EU348070**, EU347880**; Van Wyk s.n.

(JRAU). L. hirsuta (Thunb.) D.Dietr. 3— EU348102**, EU347881**; Schutte 290

(JRAU). L. involucrata (Berg.) Benth.— EU947997**, EU347809**; Boatwright et al.

116 (JRAU). L. involucrata (Berg.) Benth. subsp. digitata B.-E.van Wyk — EU348076**,

EU347805**; Van Wyk 2873 (JRAU). L. involucrata (Berg.) Benth. subsp. peduncularis

(E.Mey.) B.-E.van Wyk— EU348007**, EU347806**; Van Wyk et al. 4195 (JRAU). L. laxa Eckl. & Zeyh. 1— EU348059**, EU347795**; Van Wyk 2608 (JRAU). L. laxa Eckl.

& Zeyh. 2— EU348060**, —; Van Wyk 1726 (JRAU). L. laxa Eckl. & Zeyh. 3— —,

AF287677 2 ; Crisp 9075 (CANB). L. lenticula (E.Mey.) Benth.— EU348088**,

EU347802**; Van Wyk 2017 (JRAU). L. leptoloba Bolus 1— EU348104*, EU347757*;

Boatwright et al. 185 (JRAU). L. leptoloba Bolus 2— EU348077**, EU347804**;

Schutte 276 (JRAU). L. listii Polhill— EU348012**, EU347826**; Van Wyk et al. 4207

(JRAU). L. lotononoides (Scott Elliot) B.-E.van Wyk— EU348089**, EU347822**; Van

Wyk 1962 (JRAU). L. macrocarpa Eckl. & Zeyh.— FM875934*, FM875935* &

377 APPENDIX A3

FM875936*; Helme 2076 (NBG). L. macrosepala Conrath— EU348056**, EU347810**;

Van Wyk 2622 (JRAU). L. maculata Diimmer— EU348004**, EU347813**; Van Wyk et al. 4169 (JRAU). L. magnifica B.-E.van Wyk— EU348079**, EU347779**; Van Wyk

2421 (JRAU). L. marlothii Engl.— EU348010**, EU347825**; Van Wyk et al. 4203

(JRAU). L. maximiliani Schltr. ex De Wildeman 1— EU348078**, EU347799**; Schutte

282 (JRAU). L. maximiliani Schltr. ex De Wildeman 2— EU348098**, —; Schutte 271

(JRAU). L. meyeri (Pres!) B.-E.van Wyk 1— EU348095**, EU347811**; Van Wyk 1766

(JRAU). L. meyeri (Presl) B.-E.van Wyk 2— EU348094**, —; Van Wyk 1765 (JRAU). L. micrantha Eckl. & Zeyh.— EU348049**, EU347798**; Van Wyk 2481 (JRAU). L. mollis

(E.Mey.) Benth. 1— EU348068**, EU347763**; Koekemoer 524 (PRE). L. mollis

(E.Mey.) Benth. 2— EU348067**, EU347764**; Van Wyk 3113 (JRAU). L. oxyptera

(E.Mey.) Benth.— EU347992**, EU347787**; Boatwright s.n. (JRAU). L. parviflora

(Berg.) D.Dietr.— EU348050**, EU347797**; Van Wyk 2442 (JRAU). L. pentaphylla

(E.Mey.) Benth.— EU348001**, EU347783**; Boatwright et al. 148 (JRAU). L. platycarpa (Viv.) Pic.-Serm. 1— EU348105*, EU347759*; Boatwright et al. 192 (JRAU).

L. platycarpa (Viv.) Pic.-Serm. 2— EU348063**, EU347766**; Van Wyk 3066 (JRAU).

L. platycarpa (Viv.) Pic.-Serm. 3— EU348064**, EU347767**; Koekemoer 942 (JRAU).

L. platycarpa (Viv.) Pic.-Serm. 4— EU348011**, EU347768**; Boatwright et al. 159

(JRAU). L. platycarpa (Viv.) Pic.-Serm. 5— EU348003**, EU347769**; Van Wyk et al.

4204 (JRAU). L. platycarpa (Viv.) Pic.-Serm. 6— EU348065*, EU347765*; Friis et al.

9080 (K). L. plicata B.-E.van Wyk— EU348072**, EU347775**; Van Wyk 3382 (JRAU).

L. polycephala (E.Mey.) Benth. 1— EU348082**, EU347781**; Van Wyk 2408 (JRAU).

L. polycephala (E.Mey.) Benth. 2— EU347994**, EU347782**; Boatwright et al. 94

378 APPENDIX A3

(JRAU). L. prostrata (L.) Benth. 1— EU348054**, —; Van Wyk 3229 (JRAU). L. prostrata (L.) Benth. 2— —, EU347808**; Boatwright et al. 115 (JRAU). L. pulchella

(E.Mey.) B.-E.van Wyk 1— EU348090**, EU347785**; Van Wyk 1659 (JRAU). L. pulchella (E.Mey.) B.-E.van Wyk 2— EU348066**, —; Van Wyk 1659 (JRAU). L. pungens Eckl. & Zeyh. 1— EU348096**, EU347790**; Van Wyk 1725 (JRAU). L. pungens Eckl. & Zeyh. 2— EU347996**, EU347807**; Boatwright et al. 105 (JRAU). L. quinata (Thunb.) Benth.— EU347995**, EU347774**; Boatwright et al. 95 (JRAU). L. rabenaviana Dinter & Harms— —, EU347758*; Boatwright et al. 191 (JRAU). L. rigida

(E.Mey.) Benth. 1— EU348101**, —; Van Wyk 2700 (JRAU). L. rigida (E.Mey.) Benth.

2— —, EU347791**; Van Wyk s.n. (JRAU). L. rostrata Benth. subsp. namaquensis

(Bolus) B.-E.van Wyk — EU348002**, EU347812**; Boatwright et al. 155 (JRAU). L. rostrata Benth.— EU348000**, EU347818**; Boatwright et al. 147 (JRAU). L. sabulosa

Salter— EU348081**, EU347820**; Van Wyk 2325 (JRAU). L. sericophylla Benth. 1—

EU348093**, EU347789**; Van Wyk 1956 (JRAU). L. sericophylla Benth. 2—

EU348053**, EU347803**; Van Wyk 1917 (JRAU). L. sericophylla Benth. 3—

EU348097**, —; Van Wyk 1958 (JRAU). L. solitudinis Dummer— EU348008**,

EU347823**; Van Wyk et al. 4198 (JRAU). L. sparsiflora (E.Mey.) B.-E.van Wyk-

EU348099**, EU347786**; Van Wyk 2057 (JRAU). L. stricta (Eckl.& Zeyh.) B.-E.van

Wyk— EU348091**, EU347793**; Van Wyk 1718 (JRAU). L. subulata B.-E.van Wyk-

EU348009**, EU347824**; Van Wyk et al. 4202 (JRAU). L. umbellata (L.) Benth. 1—

EU348061**, —; Van Wyk 3234 (JRAU). L. umbellata (L.) Benth. 2— —, EU347760*;

Boatwright s.n. (JRAU). Lupinus L.: L. polyphyllus Lindley— Z70052 3 , -; Planchuelo

95. Melolobium Eckl. & Zeyh.: M. adenodes Eckl. & Zeyh. 1— —, AM050832 6 ; Van

379 APPENDIX A3

Wyk 4036 (JRAU). M. adenodes Eckl. & Zeyh. 2— EU347976**, —; Van Wyk 2159

(JRAU). M. adenodes Eckl. & Zeyh. 3— EU347986**, —; Van Wyk 4036 (JRAU). M. aethiopicum (L.) Druce 1— EU347977**, —; Strydom 5 (PRE). M. aethiopicum (L.)

Druce 2— EU347987**, —; Van Wyk 4040 (JRAU). M. alpinum Eckl. & Zeyh.—

EU347978**, —; Schutte 158 (JRAU). M. candicans (E.Mey.) Eckl. & Zeyh. 1— —,

AM050833 6 , Van Wyk 4016 (JRAU). M. candicans (E.Mey.) Eckl. & Zeyh. 2—

EU347980**, —; Van Wyk 3072 (JRAU). M. candicans (E.Mey.) Eckl. & Zeyh. 3—

EU347991*, —; Boatwright et al. 187 (JRAU). M. canescens (E.Mey.) Benth.—

EU347981**, —; Schutte 303 (JRAU). M. exudans Harv.— EU347990**, —; Van Wyk

2234 (JRAU). M. humile Eckl. & Zeyh.— EU347982**, —; Van Wyk 2543 (JRAU). M. macrocalyx Dummer— EU347989**, —; Van Wyk 3061b. M. microphyllum (L.f.) Eckl. &

Zeyh. 1— EU347983**, —; Van Wyk 4006 (JRAU). M. microphyllum (L.f.) Eckl. & Zeyh.

2— EU347979**, —; Moteetee & Van Wyk 3 (JRAU). M. microphyllum (L.f.) Eckl. &

Zeyh. 3— Z955393, —; Edwards 470. M. obcordatum Harv.— Z955403, —; Edwards

470. M. stipulatum (Thunb.) Harv.— EU347988**, —; Van Wyk 4037 (JRAU). M. subspicatum Conrath— EU347984**, —; Van Wyk 1779 (JRAU). M. wi/msii Harms-

EU347985**, —; Van Wyk 2724 (JRAU). Pearsonia Diimmer P. aristata (Shinz)

Mummer— EU347945**, EU347873**; De Castro 346 (JRAU). P. cajanifolia (Harv.)

Polhill 1— EU347948**, EU347875**; Posthumus 7 (JRAU). P. cajanifolia (Harv.)

Polhill 2— EU347950*, EU347876*; Boatwright & Magee 90 (JRAU). P. grandifolia

(Bolus) Polhill— EU347946**, EU347874**; Van Wyk 3047 (JRAU). P. sessilifolia

(Harv.) Dimmer 1— EU347947**, EU347871**; Schutte 463 (JRAU). P. sessilifolia

(Harv.) Diimmer 2— —, EU347872**; Schutte 589b (JRAU). P. sessilifolia (Harv.)

380 APPENDIX A3

Dlimmer 3— —, AJ287675 2; Crisp 9078 (CANB). P. uniflora (Kensil) Polhill-

EU347949**, —; Van Wyk 3033 (JRAU). Podalyria Willd.: P. buxifolia (Retz.) Lam.—

AM261693 1 , AM261496 1 ; Boatwright & Magee 34 (JRAU). P. oleaefolia Salisb.—

AM261708 1 , AM261667 1 ; Boatwright & Magee 79 (JRAU). Polhillia C.H.Stirton: P. brevicalyx (C.H.Stirton) B.-E.van Wyk & A.L.Schutte— EU347954**, —; Van Wyk 2100

(JRAU). P. canescens C.H.Stirton— EU347957**, —; Van Wyk 2092 (JRAU). P. involucratum (Thunb.) B.-E.van Wyk & A.L.Schutte— EU347956**, —; Schutte 339

(JRAU). P. obsoleta (Thunb.) B.-E.van Wyk— EU347955**, —; Van Wyk 2701 (JRAU).

P. pallens C.H.Stirton— EU347958**, —; Van Wyk 2095 (JRAU). Rafnia Thunb.: R. acuminata (E.Mey.) G.J.Campbell & B.-E.van Wyk— AM931036**, AM931034** &

AM931035**; Schutte 437 (JRAU). R. angulata Thunb.— EU347896*, —; Boatwright et al. 103 (JRAU). R. capensis (L.) Schinz— —, EU347742*; Boatwright & Magee 26

(JRAU). R. diffusa Thunb.— EU347895*, AJ744944 6 ; Campbell & Van Wyk 44 (JRAU).

R. globosa G.J.Campbell & B.-E.van Wyk — EU347899*, EU347743*; Boatwright et al.

180 (JRAU). R. racemosa Eckl. & Zeyh.— EU347895*, EU347741*; Boatwright &

Magee 29 (JRAU). R. spicata Thunb.— EU347897*, EU347740*; Boatwright et al. 102

(JRAU). Robynsiophyton R. Wilczek: R. vanderystii R. Wilczek— EU347952*,

EU347878*; Lisowski 20326 (K). Rothia Pers.: R. hirsuta (Guill. & Perr.) Baker-

EU347953*, EU347877*; Saadou 1798 (K). Spartidium Pomel: S. saharae (Coss. &

Dur.) Pomel [Calobota saharae (Coss. & Dur.) Boatwr. & B.-E.van Wyk]— EU347931*,

EU347844*; Davis 49544 (K). Stirtonanthus B.-E.van Wyk & A.L.Schutte: S. chrysanthus (Adamson) B.-E.van Wyk & A.L.Schutte— AM259367 1 , AM268386 1 &

AM268387'; Van Wyk & Schutte 3297 (JRAU). Ulex L.: U. europaeus L.— Z70111 3 ,

381 APPENDIX A3

—; Botanical Garden Heidelberg, Germany. Virgilia Poir.: V. divaricata Adamson-

AM260737 1 , AJ409910 7; Van Wyk 879-888 (JRAU). Wiborgia Thunb.: W. fusca

Thunb. 1— EU347966**, EU347744**; Schutte 736 (JRAU). W. fusca Thunb. 2—

EU347967*, EU347754*; Van Wyk et al. 4186 (JRAU). W. humilis (Thunb.) R.Dahlgren

[Wiborgiella humilis (Thunb.) Boatwr. & B.-E.van Wyk] 1— EU347969*, EU347752*;

Boatwright et al. 129 (JRAU). W. humilis (Thunb.) R.Dahlgren 2— EU347970*,

EU347753*; Van Wyk 3530 (JRAU). W. incurvata Thunb.— EU347975*, EU347751*;

Boatwright et al. 188 (JRAU). W. monoptera E.Mey. 1— EU347971*, EU347749*;

Boatwright et al. 152 (JRAU). W. monoptera E.Mey. 2— EU347974*, EU347750*;

Boatwright et al. 153 (JRAU). W. obcordata (Berg.) Thunb. 1— EU347965**,

EU347746**; Van Wyk 2686 (JRAU). W. obcordata (Berg.) Thunb. 2— EU347972*,

EU347748*; Boatwright et al. 98 (JRAU). W. sericea Thunb.— EU347968*,

EU347755*; Boatwright et al. 124 (JRAU). W. tetraptera E.Mey. 1— EU347964**,

EU347745**; Schutte 737 (JRAU). W. tetraptera E.Mey. 2— EU347973*, EU347747*;

Boatwright et al. 104 (JRAU). Xiphotheca Eckl. & Zeyh.: X. guthriei (Bolus)

A.L.Schutte & B.-E.van Wyk— AM260744 1 , AM261741 1 ; Vlok & Schutte 4 (SCHG). X. reflexa (Thunb.) A.L.Schutte & B.-E.van Wyk— AM260747 1 AM261744 1 ; Schutte 760

(JRAU).

382 APPENDIX A4. List of morphological characters and character states used in the morphological analysis of the Crotalarieae in Chapter 3.

1. Habit: herbs or suffrutices = 0; shrubs = 1. 2. Persistence: perennial = 0; annual =

1. 3. Young twigs: without bark formation (bark formation late or absent) = 0; with bark formation (bark formation early) = 1. 4. Leaves: digitate = 0; unifoliolate = 1; simple or phyllodinous = 2. 5. Lamina: flat = 0; terete (acicular) = 1. 6. Petiole presence: present = 0; absent (leaves sessile) = 1. 7. Petiole base: normal or leaves sessile = 0; tuberculate or with persistent spur = 1. 8. Stipules presence: present = 0; vestigial or absent = 1. 9. Stipules symmetry: symmetrical or absent = 0; asymmetrical (dissimilar in size or single) = 1. 10. Bracteole presence: present = 0; vestigial or absent = 1. 11.

Standard petal vestiture: hairy at least along the dorsal midrib = 0; totally glabrous =

1. 12. Keel shape: not rostrate = 0; markedly rostrate or helically coiled = 1. 13. Calyx lower lobes: with trifid lower lip = 0; without trifid lower lip = 1. 14. Calyx lateral lobes: not fused higher up = 0; fused higher up = 1. 15. Anthers: dimorphic = 0; uniform = 1.

16. Carinal anther: resembling basifixed anthers = 0; intermediate = 1; resembling dorsifixed anthers = 2. 17. Gynoecium base: sessile or subsessile = 0; stipitate = 1.

18. Style: upcurved = 0; straight = 1; helically coiled = 2. 19. Style vestiture: glabrous

= 0; hairy = 1. 20. Fruit type: not a samara = 0; fruit an ovoid, winged samara = 1. 21

Fruit upper suture: upper suture straight or symmetrically convex = 0; upper suture asymmetrically convex = 1. 22. Fruit: many-seeded = 0; few-seeded (1-2) = 1. 23.

Funicle length: normal length = 0; exceptionally long = 1. 24. Seed shape: transversely oblong = 0; oblong = 1. 25. Seed surface: smooth = 0; rugose = 1. 26.

383 APPENDIX A4

Chromosome base number: 7 = 0; 8 = 1; 9 = 2. 27. Cyanogenesis: absent = 0; present = 1. 28. Quinolizidine and piperidyl alkaloids (Lysine pathway): present =

0; absent = 1. 29. Sparteine: present = 0; absent = 1. 30. Lupanine type esters:

±absent = 0; present as major alkaloid = 1. 31. Macrocyclic pyrrolizidine alkaloids: absent = 0; present = 1.

384 APPENDIX A5. Character states for 6 morphological and chemical characters scored for the taxa included in the combined molecular and morphological analysis presented in Chapter 8. Characters and character states are explained at the end of the table.

1 2 3 4 5 6 Aspalathus carnosa 1 0 1 0 1 0 Aspalathus confusa 1 0 1 0 1 0 Aspalathus crenata 1 0 1 0 1 0 Aspalathus galeata 1 0 1 0 1 0 Aspalathus hirta subsp. hirta 1 0 1 0 1 0 Aspalathus hystrix 1 0 1 0 1 0 Aspalathus laricifolia subsp. laricifolia 1 0 1 0 1 0 Aspalathus linearis 1 0 1 0 1 0 Aspalathus macrantha 1 0 1 0 1 0 Aspalathus nivea 1 0 1 0 1 0 Aspalathus pachyloba 1 0 1 0 1 0 Aspalathus pendula 1 0 1 0 1 0 Aspalathus perfoliata subsp. phillipsii 1 0 1 0 1 0 Aspalathus shawii subsp. shawii 1 0 1 0 1 0 Aspalathus vermiculata 1 0 1 0 1 0 Aspalathus willdenowiana 1 0 1 0 1 0 Bolusia amboensis 0 0 0 3 0 1 Crotalaria capensis 1 0 0 3 0 1 Crotalaria distans 0 1 0 3 0 1 Crotalaria griquensis 1 0 0 3 0 1 Crotalaria humilis 0 1 0 3 0 1 Crotalaria lanceolata 0 1 1 3 0 1 Crotalaria virgultalis 1 0 1 3 0 1 Calobota cinerea 1 0 1 1 0 0 Calobota cuspidosa 1 0 1 0 0 0 Calobota cytisoides 1 0 1 1 0 0 Calobota halenbergensis 1 0 1 1 0 0 Calobota lotononoides 1 0 1 1 0 0 Calobota pungens 1 0 1 0 0 0 Calobota psiloloba 1 0 1 0 0 0 Calobota saharae 1 0 1 0 0 0 Calobota sericea 1 0 1 1 0 0 Calobota spinescens 1 0 1 0 0 0 Calobota thunbergii 1 0 1 1 0 0 Lebeckia ambigua 0 0 1 1 0 0

385 APPENDIX A5

Lebeckia brevicarpa 0 0 1 2 0 0 Lebeckia carnosa 0 0 1 0 0 0 Lebeckia meyeriana 0 0 1 0 0 0 Lebeckia pauciflora 0 0 1 1 0 0 Lebeckia plukenetiana 0 0 1 0 0 0 Lebeckia sepiaria 0 0 1 2 0 0 Lebeckia wrightii 0 1 0 0 0 0 Lotononis adpressa 0 0 0 1 0 0 Lotononis acuticarpa 0 1 0 1 0 0 Lotononis benthamiana 0 0 0 1 0 0 Lotononis bolusii 0 1 0 1 1 0 Lotononis brachyantha 0 1 0 1 0 1 Lotononis calycina 0 0 0 1 0 0 Lotononis carnosa 1 0 0 1 0 1 Lotononis curtii 0 1 0 1 0 1 Lotononis decumbens 0 0 0 1 0 0 Lotononis densa subsp. leucoclada 1 0 1 3 0 1 Lotononis digitata 0 0 0 1 0 0 Lotononis divaricata 1 0 0 1 0 1 Lotononis elongata 0 0 0 1 0 1 Lotononis eriantha 0 0 0 1 0 0 Lotononis exstipulata 0 0 1 3 0 1 Lotononis falcata 0 1 0 1 0 1 Lotononis filiformis 0 0 0 1 0 1 Lotononis foliosa 0 0 0 1 0 0 Lotononis fruticoides 0 1 0 1 0 1 Lotononis glabra 0 1 0 1 0 1 Lotononis globulosa 0 1 0 1 1 0 Lotononis hirsuta 0 0 0/1 3 0 1 Lotononis involucrata 0 0 0 3 0 1 Lotononis involucrata subsp. peduncularis 0 0 0 3 0 1 Lotononis laxa 0 0 0 1 0 1 Lotononis lenticula 0 1 0 1 0 1 Lotononis leptoloba 0 1 0 1 0 1 Lotononis fistii 0 0 0 1 0 0 Lotononis lotononoides 1 0 0 1 0 1 Lotononis macrocarpa 0 0 0 1 0 ? Lotononis macrosepala 0 0 0 1 0 1 Lotononis maculata 0 1 0 1 0 1 Lotononis magnifica 0 0 0 1 0 0 Lotononis marlothii 0 0 0 1 0 0 Lotononis maximiliani 0 1 0 1 0 1 Lotononis meyeri 1 0 0 1 0 1 Lotononis micrantha 0 1 0 1 0 1

386 APPENDIX A5

Lotononis mollis 0 0 0 1 0 0 Lotononis oxyptera 0 1 0 1 0 1 Lotononis parviflora 0 1 0 1 0 1 Lotononis platycarpa 0 1 0 1 0 0 Lotononis plicata 0 0 0 1 0 0 Lotononis pentaphylla 0 1 0 1 1 0 Lotononis polycephala 0 0 0 1 1 0 Lotononis pulchella 1 0 0 1 0 1 Lotononis pungens 0 0 0 3 0 1 Lotononis quinata 0 0 0 1 0 0 Lotononis rostrata subsp. namaquensis 0 1 0 1 0 1 Lotononis sabulosa 0 1 0 1 0 1 Lotononis sericophylla 1 0 0 1 0 1 Lotononis solitudinis 0 0 0 1 0 0 Lotononis sparsiflora 0 1 0 1 0 1 Lotononis stricta 1 0 0 1 0 1 Lotononis subulata 0 0 0 1 0 0 Pearsonia aristata 1 0 1 0 0 0 Pearsonia cajanifolia 1 0 0 0 0 0 Pearsonia grandifolia 1 0 0 0 0 0 Pearsonia sessilifolia 1 0 1 0 0 0 Rafnia acuminata 1 0 1 0 0 0 Rafnia diffusa 1 0 1 0 0 0 Rafnia globosa 1 0 1 0 0 0 Rafnia racemosa 1 0 1 0 0 0 Rafnia spicata 1 0 1 0 0 0 Robynsiophyton vanderystfi 0 1 0 0 0 0 Rothia hirsuta 0 1 0 0 0 0 Wiborgia fusca 1 0 1 0 1 0 Wiborgia incurvata 1 0 1 0 1 0 Wiborgia monoptera 1 0 1 0 1 0 Wiborgia obcordata 1 0 1 0 1 0 Wiborgia sericea 1 0 1 0 1 0 Wiborgia tetraptera 1 0 1 0 1 0 Wiborgiella bowieana 1 0 1 3 0 0 Wiborgiella inflata 0 1 1 3 0 ? Wiborgiella humilis 1 0 1 3 1 0 Wiborgiella leipoldtiana 1 0 1 3 0 0 Wiborgiella mucronata 1 0 1 0 0 ? Wiborgiella sessililfolia 1 0 1 3 0 0 Dichilus gracilis 0 0 0 0 0 0 (1) Habit: herbs or suffrutices = 0, shrubs = 1. (2) Persistence: perennial = 0, short-lived or annual = 1. (3) Stipules: present = 0, absent = 1. (4) Fruit: laterally compressed = 0, semi-terete = 1, terete = 2, inflated = 3. (5) Fruit: many-seeded = 0, few-seeded = 1. (6) Quinolizidine alkaloids: present =0, absent = 1.

387 APPENDIX B. Publications to date resulting from this PhD study:

Appendix B1. Boatwright, J. S. and B.-E. van Wyk. 2007. A new species of

Rafnia (Crotalarieae, Fabaceae). South African Journal of Botany 73 (3): 471-

473.

Appendix B2. Boatwright, J. S. and B.-E. van Wyk. 2007. The identity of Lebeckia lotononoides (Crotalarieae, Fabaceae). South African Journal of Botany 73 (4):

664-666.

Appendix B3. Boatwright, J. S., M. M. le Roux, M. Wink, T. Morozova and B.-E. van Wyk. 2008. Phylogenetic relationships of the tribe Crotalarieae (Fabaceae) inferred from DNA sequences and morphology. Systematic Botany 33 (4): 752-

761.

Appendix B4. Boatwright, J. S., P. M. Tilney and B.-E. van Wyk. 2008. A taxonomic revision of the genus Rothia (Crotalarieae, Fabaceae). Australian

Systematic Botany 21 (6): 422 -430.

Appendix B5. Boatwright, J. S. and B. - E. van Wyk. 2009. A revision of the African genus Robynsiophyton (Crotalarieae, Fabaceae). South African Journal of

Botany 75 (2): 367-370.

388 Appendix B6. Boatwright, J. S., P. M. Tilney and B.-E. van Wyk. The generic concept of Lebeckia, reinstatement of the genus Calobota and a new genus of the tribe Crotalarieae (Fabaceae). South African Journal of Botany, submitted for publication.

389 Appendix B1. Boatwright, J. S. and B. - E. van Wyk. 2007. A new species of

Rafnia (Crotalarieae, Fabaceae). South African Journal of Botany 73 (3):

471-473.

Available online at www.sciencedirect.com

SOUTH AFRICAN °,07° ScienceDirect JOURNAL OF BOTANY

ELSEVIER South African Journal of Botany 73 (2007) 471-473 www.elsevier.com/locate/sajb

Short communication A new species of Rafnia (Crotalarieae, Fabaceae)

J.S. Boatwright *, B.-E. Van Wyk

Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland Park 2006, South Africa Received 6 January 2007; received in revised form 19 February 2007; accepted 20 February 2007

Abstract

The new species Rafnia lebeckioides J.S. Boatwright and B.-E. Van Wyk is described. It is known only from a few collections in the mountains around Worcester in the Western Cape Province of South Africa. The species differs from all others in the genus in its linear fruits, which are similar to those found in species of Lebeckia Thunb. © 2007 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Crotalarieae; Fabaceae; Rafnia; new species

1. Introduction It is unfortunate that the only known collections of this seemingly rare species were filed among the unidentified The genus Rafnia Thunb. comprises 19 species endemic to specimens of Lebeckia in both the Bolus herbarium and the the fynbos regions of the Western and Eastern Cape Provinces herbarium of the Royal Botanic Gardens, Kew, and were thus (Campbell and Van Wyk, 2001). Rafnia differs from all other overlooked in the recent revision of the genus Rafnia (Campbell Cape legumes of the tribe Crotalarieae in the simple, sessile and Van Wyk, 2001). During ongoing studies in the genus Le- leaves combined with an almost complete absence of hairs, beckia it became clear that the material was misplaced in Le- except at the tips of the calyx lobes and sometimes on the bracts beckia and that it represented an undescribed species of Rafnia. and bracteoles (Van Wyk and Schutte, 1995; Campbell and Van Wyk, 2001). All the species are resprouting shrubs that 2. Species treatment regenerate from an underground lignotuber after fire. An unusual and rare new species that is superficially similar Rafnia lebeckioides J.S. Boatwright and B.-E. Van Wyk, to species of Lebeckia is described here as Rafnia lebeckioides. sp. nov., a speciebus omnibus aliis leguminis linearibus The linear leaves of this taxon resemble the acicular leaves multiseminibus, et seminibus valde angularibus differt. Non- found in L. sect. Lebeckia but differ in being flat and not terete. nihil R. spicatae Thunb., R. angulatae Thunb. et R. capensi (L.) The fruits are especially unusual for Rafnia in being linear and Schinz similis habitu parvo et foliis parvis, sed inflorescentiis flat, almost exactly like the fruits of some Lebeckia species, racemosis non pseudo racemosis ex inflorescentiis 1-6-floris including L. mucronata Benth. and L. gracilis Eckl. and Zeyh. factis differt. However, the sessile, simple, flat and glabrous leaves are typical TYPE — South Africa, Western Cape Province, slopes of for Rafnia and are not found in any species of Lebeckia. Fonteintjiesberg, south aspect [3319 CB], Esterhuysen 30582 R. lebeckioides is superficially similar to those species of Le- (BOL!, holo.; K!, iso.). beckia with a 5+5 or 6+4 arrangement of the anthers, but Small, glabrous, multi-stemmed perennial up to ?0.3 m. differs in its 5+4+1 anther configuration, which is found in all Branches densely leafy. Leaves simple, markedly secund, linear other species of Rafnia. to subspathulate, (15–) 17-23 (-27) x 1.5-3.0 mm, sessile, acute, base narrowly cuneate, glabrous. Stipules absent. Inflor- escence terminal, racemose, on slender peduncle 40-70 mm * Corresponding author. long; racemes 20-40 mm long, with 9-13 flowers; pedicels E-mail address: [email protected] (J.S. Boatwright). 1.0-1.5 mm long; bract narrowly lanceolate, leaf-like, 3.0–

0254-6299/$ - see front matter CO 2007 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2007.02.190 472 J.S. Boatwright, B.-E. Van Wyk / South African Journal of Botany 73 (2007) 471-473

3.5 mm long, glabrous; bracteoles 0.9-1.5 mm long, glabrous. surface rugose (Fig. 1). Flowering occurs in summer, mainly Flowers 8-9 mm long, yellow, standard with brown streaking in December. and wing petals turning red-brown (according to label information). Calyx 3.7-4.0 mm long, glabrous on outer 3. Diagnostic characters and relationships surface, subequally lobed but upper sinus slightly deeper than lateral and lower sinuses, tube 1.4-1.6 mm long, lobes 1.6- R. lebeckioides differs from all other species in the linear, 2.3 mm long, subulate, with carinal lobe narrower than the many-seeded (five- to 10-seeded) pods and the markedly others, tips minutely pubescent on inner surface. Standard angular seeds. In all other Rafnia species the pods are obliquely suborbicular, 5.8-6.0 x 5.5-5.8 mm, slightly emarginate, glab- lanceolate to oblanceolate or oblong and one- to five-seeded. rous, claw 1.0-1.2 mm long. Wings oblong, shorter than Furthermore, the seeds are oblong-reniform to oblong-cordi- keel, 6.0-6.3 x 2.5-2.8 mm, obtuse, glabrous, with 7-8 rows form (Campbell and Van Wyk, 2001). of sculpturing, claw 1.8-2.2 mm long. Keel slightly rostrate, R. lebeckioides has the subequally lobed calyx, sessile and 6.7-6.9 x 2.8-3.1 mm, obtuse, glabrous, pocket absent, claw simple leaves, and glabrous vegetative and reproductive parts 2.0-2.2 mm long. Anthers dimorphic, four long, basifixed typical of all other Rafnia species. Within the genus, it shares anthers alternating with five ovate, dorsifixed anthers, carinal some apomorphies with species of section Rafnia, notably the anther intermediate in size and shape. Pistil subsessile, calyx without a trifid upper lip, sculpted wing petals and rostrate glabrous, ovary linear, 5.0-5.3 x 0.7-0.9 mm with 9-10 ovules; keel petals without pockets (Campbell and Van Wyk, 2001). It style 3.7-4.5 mm long, curved upwards, glabrous. Pods is one of only two species with a multi-flowered raceme — the laterally compressed, subsessile, linear, 22-28 x 2.5-4.0 mm, other is R. racemosa Eckl. and Zeyh. (a large woody shrub). 5-10 seeded, indehiscent. Seeds markedly angular, 2.5- R. lebeckioides is small in stature and appears to only reach 2.6x2.2-2.3 mm, predominantly black with white spots, heights of up to 0.3 m (exact height unknown). As it is unlike

Fig. I . Morphology of Rafizia lebeckioides (drawings by JSB): (a) lateral view of flower; (b) wing petal; (c) keel petal; (d) standard petal; (e) outer surface of the calyx (upper lobes to the left); (0 short, dorsifixed anther, intermediate carinal anther and long, basifixed anther; (g) androecium; (h) lateral view of pod; (i) pistil; (j) bract and bracteoles; (k) lateral view of seed; (I) abaxial surface of leaf; (m) flowering branch, showing the densely leafy branches and racemose inflorescence structure. Voucher specimens: (a, c, d, f, g, i , j bract only, I, m) Esterhuysen 8197 (BOL); (b, e, j bracteoles only) Esterhuysen 30581 (BOL); (h, k) Esterhuysen 28900 (BOL). Scale bars: (a—I) I mm; (m) I0 mm. J.S. Boatwright, B.-E. Van Wyk / South African Journal of Botany 73 (2007) 471-473 473

-OW OMMEMISSIMPOWNPORWROMInISEMMIlmemerr/ ftie,Mmusim-NRAmaimmogm L-Pwim 41 NTMOMPEivow,el r ft4ftEMPITATI SMOVAMMA*MANKEMeaaggameamgraile• All vailOWANIUMWMPNOMMEMOMIRENOMNIOMMOINOM kamOMMOOMMERWOMMEMMOMM Ljorpiarmonmmummi'MEM MIOMEMEMMUMZ Mammilammimmarim=v

Fig. 2. The known geographical distribution of Rafnia lebeckioides. any of the species of Rafnia it is not clear to which members of Esterhuysen 28900 (BOL); Du Toit's Peak, summit ridge the genus it is related. Obtaining fresh material for molecular (—CC), Esterhuysen 29038 (BOL). systematic studies is imperative to assess the phylogenetic position of this species. Acknowledgements

4. Distribution and habitat The National Research Foundation (NRF) and Univer- sity of Johannesburg are gratefully acknowledged for R. lebeckioides occurs on the Hex River and Du Toit's Moun- funding. We thank Dr. H.F. Glen for kindly translating tains around Worcester in the South Western Cape Province at the diagnosis. altitudes between 1500 and 1800 m (Fig. 2). It has mainly been collected on southern to south-eastern slopes in recently burnt References vegetation, where it has been recorded on stony, slightly marshy soil. Campbell, G.J., Van Wyk, B.-E., 2001. A taxonomic revision of Rafilia (Fabaceae, Crotalarieae). South African Journal of Botany 67,90-149. Van Wyk, B.-E., Schutte, A.L., 1995. Phylogenetic relationships in the tribes Additional specimens examined Podalyrieae, Liparieae and Crotalarieae. In: Crisp, M., Doyle, J.J. (Eds.), Advances in Legume Systematics 7, Phylogeny. Royal Botanic Gardens, —3319 (Worcester): Waaihoek Mountain (—AD), Esterhuy- Kew, pp. 283-308. sen 8297 (BOL); Waaihoek Peak, south-eastern slopes (—AD),

Edited by JC Manning Appendix B2. Boatwright, J. S. and B.-E. van Wyk. 2007. The identity of

Lebeckia lotononoides (Crotalarieae, Fabaceae). South African Journal of

Botany 73 (4): 664-666.

Available online at www.sciencedirect.com e7 62 SOUTH AFRICAN c),' ScienceDirect IOURNAL OF BOTANY

ELSEVIER South African Journal of Botany 73 (2007) 664-666 www.elseviercom/locate/sajb

Research note The identity of Lebeckia lotononoides (Crotalarieae, Fabaceae)

J.S. Boatwright * , B.-E. Van Wyk

Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland Park 2006, South Africa Received 28 February 2007; received in revised form 12 April 2007; accepted 19 April 2007

Abstract

Lebeckia lotononoides Schltr. is a rare species of the Western Cape Province of South Africa. It is anomalous in the genus in its decumbent habit with buried stems and unique wing- to keel lamina ratio. Poorly collected, the species has been confused with Lebeckia multiflora E. Mey. Recent collections have increased our understanding of the taxon and provided insight into its sectional placement. The trifoliolate leaves with long petioles, pubescent standard petal, 5+4+1 anther arrangement, and semi-terete, thick-walled pods place L. lotononoides in section Calobota (Eckl. & Zeyh.) Benth. L. lotononoides differs from L. multtflora in its decumbent habit, wing- to keel lamina ratio of ±2:1, and rugose, spotted seeds. © 2007 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Crotalarieae; Fabaceae; Lebeckia lotononoides; Lebeckia multiflora

Introduction Southern Africa: an annotated checklist, Strelitzia 14: 523 (2003), pro parte, non Goldblatt and Manning (2000). Type: Lebeckia Thunb. is a genus of legumes, comprised of ±36 South Africa, [Western Cape Province], "In regione nama- species, that occurs in the southern and western parts of South quensi: In sabulosis montium Karree-Bergen, alt. c. 4500 ped.", Africa and Namibia. The last comprehensive revision of the Schlechter 8214 (BM!, BOL!, K!, PRE!, S!, syn.). genus was that of Harvey (1862), who followed the sectional [Note: In accordance with Art. 60 and Art. 61 of the Vienna classification proposed by Bentham (1844). Since then several Code (www.ibot.say.sk), it is here formally proposed that the new species have been described with some doubt as to their name Lebeckia lotonoides be regarded as a correctable, correct systematic placement within the genus. One such orthographic error and that L. lotononoides be accepted as the species is Lebeckia lotononoides, described by Schlechter correct form of the name. Schlechter clearly intended to name (1900). Until recently this very rare species was known only the species after the genus Lotononis (DC.) Eckl. & Zeyh., with from the type material. Recent collections as well as studies of which it has a superficial similarity, as is evidenced by his plants in the wild have increased our understanding of the handwritten labels on all the type specimens cited above. The species, which has been confused with other species of Le- inadvertent elision of a few letters has reduced Schlecther's beckia such as Lebeckia multiflora. In this paper we give a meaningful epithet to a jumble of nonsense syllables. It is easy complete description of the species, correct the spelling of its to see that this tongue-twisting epithet could have been misspelt name, and provide notes on its distribution, habit and diagnostic by a non-specialist copy-editor or typesetter. The Code characters. recommends that "The liberty of correcting a name is to be used with reserve, especially if the change affects the first Taxonomy syllable and, above all, the first letter of the name" (Art. 60.3). In this case the change is deep within the word and has a Lebeckia lotononoides Schltr. in Engl. Bot. Jahrb. 27: 143 minimal effect on indexing.] (1900), as "L. lotonoides"; Germishuizen and Meyer, Plants of Small, sericeous, multi-stemmed, decumbent perennial up to 0.5 m in height. Branches woody at base, often buried. Leaves * Corresponding author. digitately trifoliolate (rarely 5-digitate with two additional E-mail address: [email protected] (J.S. Boatwright). leaflets), secund, sericeous; leaflets elliptic to narrowly

0254-6299/S - see front matter © 2007 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2007.04.064 J.S. Boatwright, B.-E. Van Wyk / South African Journal of Botany 73 (2007) 664-666 665

oblanceolate, conduplicate, subsessile, terminal leaflet 10 3 mm long, subulate, carinal lobe narrower than others, tips 17 x 1.5-3.0 mm, lateral leaflets 8-14 x 1.5-3.0 mm, mucro- minutely pubescent on inner surface. Standard 13-16 mm long, nate, base narrowly cuneate; petiole longer than leaflets, 10- claw linear, 4-7 mm long, lamina widely ovate to transversely 23 mm long. Stipules absent. Inflorescence terminal, racemose, oblong, 8-9 x 9.0-12.5 mm, emarginate, pilose along dorsal (15—) 24-54 mm long, with (3—) 4-14 flowers; pedicel 1-3 mm midrib. Wings 14-15 mm long, claw 3.0-4.5 mm long, lamina long; bract linear, 0.7-1.5 mm long, pubescent, caducous; narrowly oblong, subfalcate, longer than keel, 11.5-15.0 x 2- bracteoles linear, 0.3-0.6 mm long, pubescent, caducous. Flo- 4 mm, obtuse, glabrous, with 7-8 rows of sculpturing. Keel 9- wers 12-15 mm long, yellow. Calyx 5.0-6.5 mm long, 11 mm long, claw 3-4 mm long, lamina semi-circular, 5.5- pubescent, subequally lobed but upper sinus slightly deeper 7.0 x 3.0-4.5 mm, obtuse, glabrous, pocket present, 1.5-3.0 mm than lateral and lower sinuses, tube 3.0-3.5 mm long, lobes 2— long. Anthers dimorphic, four long, basifixed anthers alternating

a/P b2

Fig. I. Morphology, habit and distribution of Lebeckia lotononoides: (al —a3) leaves in abaxial view: (al and a2) trifoliolate leaves, (a3) 5-digitate leaf; (b I —b3) pods: (b l and b2) lateral view (note the slightly clavate shape), (b3) dorsal view; (c I —c3) seeds in lateral view; (dl —d3) petals: (d1) standard petal (note the long, linear claw), (d2) wing petal (note the exceptionally long lamina length), (d3) keel petal; (e) flower in lateral view showing the long wing petals; (0 flowers of L. lotononoides; (g) decumbent habit of L. lotononoides; (h) known geographical distribution. Photographs taken by J.S. Boatwright at Roodeheuwel Farm. Voucher specimen: (a—d) Boatwright et al. 142 (JRAU). Scale bars: (a, b) 10 mm; (c) 1 mm; (d) 4 mm. 666 J.S. Boatwright, B.-E. Van 1+54c/ South African Journal of Botany 73 (2007) 664-666 with five ovate, dorsifixed anthers, carinal anther intermediate was made by Schlechter at "Karree-Bergen", close to Nuwerus in size and shape. Pistil subsessile to shortly stipitate, pu- in the Vanrhynsdorp district according to Leistner and Morris bescent, ovary linear, 6.5-7.5 x 0.7-0.9 mm with 15-19 ovules; (1976) (Fig. 1h). The species has been recorded on well- style 3.0-3.5 mm long, curved upwards, glabrous. Pods drained, sandy soils with the main stems becoming covered • narrowly oblong, sometimes somewhat clavate, semi-terete, with sand. The multi-stemmed, decumbent habit (Fig. 1g) subsessile to shortly stipitate, 18-37 (-40) x 3-4 mm, 7-12- appears to be an adaptation to moving sand dunes, as extensive seeded, dehiscent. Seeds suborbicular, 2.0-2.5 x 1.5-2.0 mm, underground branches enable the plant to emerge above the mature seeds brown with beige spots, hilum brown, surface sand after being covered. rugose (Fig. I). Flowering time: late winter to late spring. 4.1. Additional specimens examined Diagnostic characters 3017 (Hondeklipbaai): Farm Roodeheuwel, 9 km west of L. lotononoides superficially resembles broad-leaved forms Nariep (—DC), Boatwright et al. 142 (JRAU), Perold 1646 of L. multiflora, notably some northern populations on the West (PRE). Coast and around Clanwilliam. In these forms the leaflets are 3117 (Lepelfontein): Vredendal, south-east of Brandsebaai, broad and sericeous much like those of L. lotononoides. on Farm Hartebeestekom (—BD), Helme 2931 (NBG); Van However, L. lotononoides differs in its smaller, decumbent RhynsdorpNredendal, Brandsebaai (—BD), Van Rooyen 2235 habit with buried stems and especially in the exceptionally long (PRE). lamina of the wing petals, which is ± twice as long as those of —3118 (Vanrhynsdorp): Karree-Bergen (—AB), Schlechter the glabrous keel petals. The inflorescences are unarmed, and s.n. sub TRV 1059 (PRE). the sericeous pods contain spotted, rugose seeds. Lebeckia multiflora, in contrast, is an erect, divaricately branched shrub Acknowledgments usually more than 1 m in height with inflorescences that often terminate in a spine; the wing petals are almost as long as or Funding from the National Research Foundation (NRF) and slightly longer than the pilose keel petals; the pods are University of Johannesburg are gratefully acknowledged. The pubescent; and the seeds are pale pink with a smooth surface. authors would like to thank Mrs. B. van Zyl for kindly allowing The trifoliolate leaves with long petioles, pubescent standard us to collect specimens on the farm and for sending seed petal, 5+4 +1 anther arrangement, and semi-terete, thick-walled material for this study. Dr H.F. Glen is thanked for a useful pods place L. lotononoides in section Calobota (Eckl. & Zeyh.) discussion on the nomenclature of the species. Benth. The shape and relative size of the petals are most unusual in References this species (Fig. 1d). Not only that the lamina of the wing petals is exceptionally long but also that the standard petal is strongly Bentham, G., 1844. Enumeration of Leguminosae indigenous to southern Asia differentiated into a long, linear claw and a transversely oblong and central and southern Africa. Hooker's London Journal of Botany 3, 355-363. lamina. In other species of Lebeckia section Calobota, the wing Harvey, W.H., 1862. Leguminosae. In: Harvey, W.H., Sonder, O.W. (Eds.), petal laminas are never more than 1.3 times as long as the Flora Capensis, vol. 2. Hodges, Smith, and Co., Dublin, pp. 82-89. lamina of the keel petals, whereas in L. lotononoides these are Leistner, 0.A., Morris, J.M., 1976. South African place names. Annals of the almost twice the length of the keel lamina. Cape Provincial Museums 12, 199. Schlechter, F.R., 1900. Plantae Schlechterianae novae vel minus cognitae describunter vol. 2. Engler Botanische Jahrbucher 27, 143. Distribution and habitat

L. lotononoides occurs on the west coast of South Africa around Hondeklipbaai and Brand se baai. The type collection

Edited by JC Manning Appendix B3. Boatwright, J. S., M. M. le Roux, M. Wink, T. Morozova and B.-

E. van Wyk. 2008. Phylogenetic relationships of the tribe Crotalarieae

(Fabaceae) inferred from DNA sequences and morphology. Systematic

Botany 33 (4): 752-761. Systematic Botany (2008), 33(4): pp. 752-761 © Copyright 2008 by the American Society of Plant Taxonomists Phylogenetic Relationships of Tribe Crotalarieae (Fabaceae) Inferred from DNA Sequences and Morphology

James S. Boatwright, 1-3 Marianne M. le Roux; Michael Wink,2 Tatjana Morozova,2 and Ben-Erik van Wyk l 'Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland Park 2006, Johannesburg, South Africa 2lnstitute of Pharmacy and Molecular Biotechnology, University of Heidelberg, Im Neuenheimer Feld 364, 69120 Heidelberg, Germany 3Author for correspondence ([email protected] )

Communicating Editor: Gregory M. Plunkett

Abstract—Tribe Crotalarieae is a large and diverse group of papilionoid legumes that largely occur in Africa. A systematic study of generic relationships within the tribe was undertaken using nucleotide sequences from the internal transcribed spacer (ITS) of nuclear ribosomal DNA, the plastid gene rbcL, and morphological data. The Crotalarieae are supported strongly as monophyletic and sister to the tribe Genisteae. Lebeckia, Lotononis, and Wiborgia are all paraphyletic in the molecular analyses and morphological data support the division of Lebeckia into three more natural genera (one of which includes the monotypic North African Spartidium). Four major lineages were identified within the tribe based on sequence data: the "Cape" group, comprising Aspalathus, Lebeckia, Rafnia, Spartidium, and Wiborgia; the Lotononis group, comprising Lotononis pro parte, Pearsonia, Robynsiophyton, and Rothia; a group comprising Lotononis section Leptis, L. section Listia, and allies; and the Crotalaria group, comprising Bolusia, Crotalaria, and Lotononis hirsuta (Lotononis section Euchlora). Morphological analysis yields a similar topology, except that Lotononis is monophyletic if L. hirsuta were excluded. When the molecular and morphological data sets are combined, the same major clades are retrieved as in the molecular analysis, with the notable exception that Lotononis and Lebeckia senso stricto are supported as monophyletic. The results from this study have important implications for the classification of the tribe Crotalarieae and present an important step towards a natural and phylogenetic generic classification for the tribe.

Keywords—Crotalarieae, Fabaceae, genistoid legumes, ITS, morphology, phylogeny, rbcL.

Crotalarieae (Benth.) Hutch. is a tribe of legumes that cur- have shown that the genus is unlikely to be monophyletic (Le rently comprises 11 genera and ca. 1204 species (van Wyk Roux 2006; Le Roux and van Wyk 2007; Boatwright, in prep.). 2005). It represents the largest tribe of papilionoid legumes in Several molecular systematic studies of Fabaceae and spe- Africa and also within the genistoid alliance, comprising cifically the genistoid legumes (sensu Polhill 1976, 1981) have about 51% of genistoid legumes (species totals in Lewis et al. been conducted in recent years (Kass and Wink 1995, 1996, 2005 used for calculation), largely due to the fact that the 1997; Crisp et al. 2000; Doyle et al. 2000; Kajita et al. 2001; genus Crotalaria contains ca. 600 species (Polhill 1982). Some Wink and Mohamed 2003; Wojciechowski et al. 2004; Boat- of the species in the tribe are important commercially such as wright et al. 2008). These studies place Crotalarieae within the "core" genistoid Glade together with, among others, the Aspalathus linearis, which is used for the production of Rooi- South. African tribes Podalyrieae Benth. and Genisteae bos tea (van Wyk et al. 1997), and Lotononis bainesii Bak., an (Bronn) Dumort., and confirm the exclusion of the Argyrolo- important fodder plant (Bryan 1961). Some Crotalaria and bium group (Argyrolobium, Dichilus, Melolobium, Polhillia) and Lotononis species have been reported to have medicinal prop- the rest of the Genisteae from Crotalarieae (van Wyk and erties (van Wyk 2005) and a few are used as what is called Schutte 1995). A sister relationship between Crotalarieae and 'Musa -pelo in Lesotho traditional medicine to cure or ease a Genisteae, with Podalyrieae successively sister to these, is broken heart (Moteetee and van Wyk 2007), while others shown by the molecular studies cited above, but sampling from the same genera are poisonous (van Wyk et al. 2002). limitations did not allow detailed evaluations at the generic The Crotalarieae are subendemic to Africa, with only a few level. species of Crotalaria, Lotononis, and Rothia occurring on other The present study is aimed at exploring generic circum- continents. Aspalathus, Rafnia, and Wiborgia are endemic to scriptions and relationships within Crotalarieae, using the Cape Floristic Region, while Lebeckia (as currently circum- nrDNA ITS sequences, plastid rbcL sequences, and morpho- scribed) is distributed throughout the Cape and extends into logical data, based on a sample of 135 species representing all the south-western and central parts of Namibia. Generic cir- of the 11 genera currently recognized, as well as major infra- cumscriptions and relationships within the tribe are known generic groups within some of the genera. The aim was to to be complicated, with evidence of reticulation and conver- assess the monophyly and relationships of the individual gence (Dahlgren 1970a; Polhill 1976, 1981; van Wyk 1991a). genera. The most recent evaluations of generic relationships in the tribe are those of Polhill (1976, 1981), van Wyk (1991a), and van Wyk and Schutte (1989, 1995). According to van Wyk MATERIALS AND METHODS (1991a), two main groups can be identified within the tribe, Plant Accessions and Choice of Outgroups—A total of 175 sequences of ITS (88% of the total ITS matrix) and 207 sequences for rbcL (91% of the namely the "Cape" group comprising Aspalathus, Lebeckia, total rbcL matrix) were produced from taxa of the Crotalarieae and com- Rafnia, and Wiborgia, and the Lotononis group comprising bined with previously published sequences from the tribes Genisteae and Lotononis, Pearsonia, Robynsiophyton, and Rothia. The place- Podalyrieae as outgroups (van Wyk and Schutte 1995; Crisp et al. 2000; ment of Bolusia, Crotalaria, and Spartidium within these two Boatwright et al. 2008). Sequences of both the ITS and rbcL regions were available for 161 taxa of the Crotalarieae and these were used for the groups is not clear, but a close relationship between the combined molecular analysis. A morphological matrix was compiled for former two genera has been suggested. Recent revisionary those species included in the combined molecular analysis, based on 31 studies of Lebeckia (the first since Harvey's treatment of 1862) characters (including a few chemical and cytological characters) that were

752 2008] BOATWRIGHT ET AL.: CROTALARIEAE PHYLOGENY 753 scored from examining specimens from BM, BOL, GRA, JRAU, K, NBG quences were not available for some of the Genisteae included as out- (including SAM and STE), P, PRE, S, UPS, and WIND for Lebeckia, Robyn- groups. siophyton, Rothia, and Spartidium and from literature for the other genera Morphological Analysis—Much debate currently surrounds the inclu- (Polhill 1974, 1976, 1982; Dahlgren 1975, 1988; Schutte and van Wyk 1988; sion of morphological characters in phylogenetic analyses (e.g. Scotland van Wyk 1991b; van Wyk and Schutte 1995; Campbell and van Wyk 2001; et al. 2003). However, Wiens (2004) discusses the importance of morpho- van Wyk 2003). These data, along with sequences of the same species, logical characters in phylogenetics, suggesting that their implementation were used to perform a combined molecular/morphological analysis. in separate and combined analyses may provide more rigorous phylog- DNA Extraction, Amplification, and Sequencing—Sequencing of the enies, while improved resolution has been shown by Wortley and Scot- gene regions was carried out in two independent laboratories, therefore land (2006) when morphology is combined with molecular data. Charac- the strategies differed slightly. The gene sequencing methods of Kass and ters and character states used for the cladistic analysis are given in Ap- pendix 2. The character states were polarised using the method of Wink (1997) were followed at University of Heidelberg, whereas se- outgroup comparison. Phylogenetic analyses were performed in PAUP* quences generated at the University of Johannesburg were gathered ac- (Swofford 2002) with the characters treated as unordered and equally cording to the methods described below. weighted (Fitch 1971). A heuristic search with 1,000 random addition DNA was extracted from silica-dried or herbarium leaf material using sequences, TBR, and the MULTREES option off was performed with a the 2x hexadecyltrimethylammonium bromide (CTAB) method of Doyle limit of 10 trees held per replicate. Internal support was assessed using and Doyle (1987) and purified through QlAquick silica columns (Qiagen 1,000 bootstrap replicates (Felsenstein 1985) as described above. Dichilus Inc., Hilden, Germany). Sources of material used in the study are listed in gracilis, a suffrutex, was selected as the outgroup for the analysis. A Appendix 1. The internal transcribed spacers (ITS) of nuclear rDNA were second analysis was performed where Genista tinctoria L., a shrub, was amplified using the primers of White et al. (1990) and Sun et al. (1994), used as outgroup to test whether polarising the shrubby habit as ancestral while for rbcL those of Fay et al. (1997) were used. Amplification reactions had an effect on the topology, but no difference was found in this analysis were carried out using polymerase chain reactions (PCR), in 25 u.1 reac- (tree not shown). tions containing: 22.5 ul ABgene 1.1x PCR Mastermix (ABgene House, Blenheim Road, Epsom, Surrey, KT19 9AP, U.K.) consisting of 1.25 units Thermoprime Plus DNA Polymerase, 75 mM Tris-HC1 (pH 8.8), 20 mM RESULTS (NH4),SO4, 1.5 mM MgC12, 0.01% (v/v) Tween® 20, 0.2 mM each of deoxyribonucleotide triphosphates (dNTPs); 0.3 ul of both forward and rbcL Data Set—The rbcL matrix consisted of 1,296 aligned reverse primers (0.1 ng/u1); 0.8 ul 0.004% bovine serum albumin (BSA); positions with 222 variable and 167 parsimony informative 1.2% dimethyl sulfoxide (DMSO; for ITS only); 20-50 ng DNA template; characters. Tree searches produced 2,650 equally parsimoni- and sterile distilled water to make up a final volume of 25 p.I. The DMSO ous trees of 527 steps, a consistency index (CI) of 0.50, and a was added to the amplification reactions as this may improve sequencing of ITS due to relaxation of the template secondary structure during am- retention index (RI) of 0.92. Overall the strict consensus tree plification (Alvarez and Wendel 2003). The PCR cycles and cycle sequenc- (not shown) is poorly resolved with a few supported clades. ing reactions followed Boatwright et al. (2008). A Glade consisting of Lebeckia section Calobota (Eckl. & Zeyh.) Sequence Alignment and Phylogenetic Analyses—Complementary Benth., L. section Stiza (E.Mey) Benth., and Spartidium saharae strands of the sequenced genes were edited in Sequencher v. 3.1.2 (Gene Lotononis s.s. (L. section Lotono- Codes Corporation), and aligned manually in PAUP* version 4.0b10 is weakly supported (60 BP). (Swofford 2002). Alignments used in this study are available from Tree- nis and allies) is moderately supported (74 BP), while the rest BASE (study number S2070). Insertions and deletions (indels) of nucleo- of the genus is unresolved except for L. section Listia (E.Mey.) tides were scored as missing data and thus did not contribute to the B.-E.van Wyk, which has high support (89 BP). The Crotalaria combined analysis, but an additional search with gaps from the ITS data Glade is strongly supported (88 BP) and the sister relationship set coded as binary characters was performed (no gaps were present in is also strongly supported the rbcL dataset). Gaps were coded in SeqState version 1.32 (Muller 2005) between Bolusia and Crotalaria using simple indel coding as described by Simmons and Ochoterena with a BP of 90. Both these genera are well-supported to be (2000). Maximum parsimony analyses (MP; Fitch 1971) and Bayesian monophyletic (Bolusia 98 BP; Crotalaria 90 BP) and they are MCMC analysis (BI; Yang and Rannala 1997; Huelsenbeck and Ronquist successively sister to Lotononis hirsuta [L. section Euchlora 2001; Ronquist and Huelsenbeck 2003; MRBAYES version 3.1.2) were (Eckl. & Zeyh.) B.-E.van Wyk]. Tribe Crotalarieae as a whole performed as described in Boatwright et al. (2008). Internal support for MP was estimated with 1,000 bootstrap replicates (Felsenstein 1985) using is strongly supported as monophyletic (94 BP) and sister to tree bisection-reconnection (TBR) and holding 10 trees per replicate. The the Genisteae (100 BP). Genisteae and Podalyrieae are both following scale for bootstrap support percentages (BP) was used: 50-74%, supported as monophyletic (89 BP and 78 BP, respectively). low; 75-84%, moderate; 85-100%, strong. Congruence of the separate ITS Data Set—The analysis of the ITS data set included 560 datasets was assessed by examining the individual bootstrap consensus trees in order to compile a combined ITS and rbcL matrix for the 161 taxa characters, of which 353 were variable and 256 parsimony where both sequenced regions were available. The bootstrap trees were informative. Analysis resulted in 5,970 equally parsimonious considered incongruent only if they displayed 'hard' (i.e. strong boot- trees with a tree length (TL) of 1,031 steps, a CI of 0.51, and strap support) rather than 'soft' (i.e. low bootstrap support) incongruence a RI of 0.85. The analysis in which gaps were coded resulted (Seelanan et al. 1997; Wiens 1998). In addition, incongruence length dif- in identical Glade resolution as the analysis without coded ference tests (ILD, Farris et al. 1995) were performed using the partition- homogeneity test of PAUP*. The test was implemented with 1,000 repli- gaps, although some clades had stronger or weaker support cate analyses, using the heuristic search option with simple addition of (not shown). The analysis with coded gaps resulted in 2,540 taxa, TBR, and the MULTREES option in effect. trees (TL = 1,340; CI = 0.51; RI = 0.84). The strict consensus The GTR + I + G model [selected by MODELTEST v. 3.06 (Posada and tree based on ITS data (not shown) is better resolved than Crandall 1998) using the corrected AKAIKE information criterion (AICc)] was implemented for the BI analysis and a total of three million genera- that of rbcL, with several well-supported clades. Aspalathus tions were performed with a sampling frequency of 10. A majority rule was weakly supported as monophyletic (64 BP), although consensus tree was produced to determine the posterior probabilities this support percentage is higher when the gap characters are (PP) of all observed bipartitions (only PPs above 0.5 are reported on the included (78 BP). Lebeckia and Wiborgia are both paraphyletic. tree). The following scale was used to evaluate the PPs: 0.50-0.84, low; section Viborgioides Benth. including Wiborgia humilis 0.85-0.94, moderate; 0.95-1.0, strong. Lebeckia Despite repeated attempts, we were only able to amplify ITS1 for some is moderately supported (80 BP), while the positions of L. samples of Lotononis marlothii, L. pentaphylla, L. platycarpa, and L. rostrata. inflata and L. mucronata are unresolved. The remaining spe- We could also amplify only the first half of rbcL for some samples of cies of Wiborgia form a weakly supported Glade (61 BP; 79 BP shawii, Lotononis Aspalathus laricifolia subsp. laricifolia, A. shawii subsp. with gap coding). Rafnia is weakly supported as monophy- falcata, L. platycarpa, Robynsiophyton vanderystii, Rothia hirsuta, Spartidium saharae, and Wiborgia humilis. Missing data represented 4.3% of the entire letic (75 BP) and Lebeckia section Calobota along with L. section combined molecular matrix due to the fact that both ITS and rbcL se- Stiza and Spartidium saharae also has weak support (59 BP).

754 SYSTEMATIC BOTANY [Volume 33

LO10,10,01:-.11019 1 100 E Aspatathus kneads 1 LOWerttalis panarcaa Aspalathus ponduta Locanonts tuticaklas 1.0 Aspalathus (Means 2 0 Lorartanis 0,000yanttsa Aspatathus cameo) Laformelbc 0LI:cue 2 Aspatathus galeata LotanasCia cLva--Soca 73 Lataranis aryjcsbra Aspalathus macrantha 99 la Latacsaras =Data subcp. namaquensis Aspatafflus wi,b7 lonoriana . is rostra/a Aspatathus oonfusa n 94 a Lotononis taccucata Aspalathus fro:foliate subsp, phfflipsii 1.0 LO.Ora0.4 boa Aspalathus cronata 9 , 91, LaDanaris tatrca0ho Aspatathus vermiculate la Lacanaces rnararasepa. Aspatathus shang subsp. shawl; Lot990,83 trafra 99 Lae:nods m=aaccca Aspalathus laricifetia subsp. lark:doh° tmencos euMi Aspatathus hitta subsp. trine LatcaracVs •^71+,,,, Aspalathus hystrix go Aspalathus nmea 00 LC:I:Cc:cc:L:2,T 2). I Aspatathus pachytabo 00 to:mans i'F.V.atob .7 2 1 C Lotontees pungens I Wibargia fusee 1 1 6.1171■Vittorgla tetragon) 1 0.81 1-- alturani r8912464 1 5,55 Wibmpla totraptera 2 LotonanW*rWtt scrfc001,d6'Wean'yfb 1 cs 0.99 0 521 Witorgia sencea tononabis SalicOphy173 2 65 ro.... Wibergia obcoldata 1 730 c totonants putheea 1 1 totorsonIS meyori 1 0.99 L— Wtborgia oboe/data 2 1 96 1 Weborgta monoptera 1 100 to:wards demo subsp. eausoctscla Wibergla menoptora 2 9 54 4 tokmanis 0,49putato Lc....ronon,2 f',,,- ;.,-..89,1 :.- 98:9 6..f.1.!..9a , 1 .0 I— Witco& Incunatta 10 094 Lc.nrracra in:re:1Mb C.Utt. "".1....0704. :7 lAlborgia fusee 2 L8.4-,89,a ,Arritt=!..1 I-- Lefacckla inflata 1 Lebeck/a fn ats 95 Lotononis E. 2,948 ,— Leiabe...ea Jnkata 2 1.0 Worry= eangXo Lebectca 11111Cranata Lebockfa mucronata 18828ents ct&ta = Lotcoonts danosa 60 Lebeckia purmans 1 Loton8,94 lolonancestas Lebecka pavans 2 /8=8,8,1 sas2/iaa 1 Lebeckia macrantha 1 Poafaarnb tutztrta Lebt•cke POroSaribgralletA:da fitccranztra 2 Lcbeckia p.s4Woba Pearaarea c*rclea 1 Paacsartfaon. W..!anif 2 Lebeckia sericco 1 Ra110., hircrais' Lcbeckia sencaa 2 Rotynsialchyron vanderysto tobeck42 sencea 3 -..8.2 larieae larieae p!.-..15= Leber-km cyttsoides 1 /atm,: t pt.8.c.:9•99 2 Lebec-PM cytisattes 2 ta tWancr..'s 8'9783,93 3 Lcbcckia =Mk= La:0mm* p'Zirtopa 4 , LetartOniS r...'3:y:z.r.1-7 5 Lebeckia lotonorrecfes Cro Leheckia tra!enberpensis La.an0nis Woad altloor.a Lebeckia haleabcrgonsis 2 1.81Cutzr,'s to:817J 1 LebeckJa spfnescens 1.01'CrOr1,11112ma 2 I 0 85 Lebeckfa cfnea.13 0 0 Latanan 00,0+100) Spartidiuni =Mama acon.z,sa Rents acumtneta Lbt1r1+11:.,.5 Cm:ft.:pa Losolem:9 9c-ccimtsana Rafnia tacemosa 941 Rofnia spicata toSononis mega I iD 18 1 Lotenents no/:s 2 1 Rafnia globose LCAntm:',, 0 j-• Rabbet diffuse Lotorpenis bonftpwriCarta VViborgia humbis 1 Lotonotral dvlota 1 Wtbonya humdis 2 1.013:705 Mara 2 1 i Lebeckia letpoldtiana 1 77 1.0.onsars goners Lebeckia bowtesna 1 10 Leonters Lebockla bowfoana 2 1.0X910111S Obt.s10S., Laarzo,Us rplayefico 1.0 Lebeckia lelpoldliona 2 0.98 CI— Lebeckia sesstlifolia LCuX/120.L"na",5 = 2 Lebeckia cantasa 1 Lolanoiris pent..-98yZa Lebeckia cantos° 2 L0.48,989 SO:Wand Lebeckia moyertana 1 M° 91 Leone:lb sub :i4, 1 .0 , LW:mortis arta Lebeckia pkikenetiana 1 lc Laconor ear rrtancet2 ------100 Lebcckia paucifictra 1 100 E Lolonores /88:91a I Lebeckia paucitiora 2 Latanorsts Airs1.48 2 Lebeckia wrfetatil 1 1.0 53 100 164= Lobockia voighttif 2 Lebeckia wrighttit 3 1.0 99 1990 10099 Lebeckia plukanctiana 2 1.0 CrWar°Lciamniscavar.CnatCmlaatat,4:4' d4tw91417c.f.i"3 3. C1,—. tebeckia ambigua 1 00 98 18 Cnvafarta capeasia2 Lebeckia brevicarpa Cratatarta L004.etaf 1 Lebockta ambigua 2 92 Lebeckia solaria 1 n Cm4acraq'atartad'hbuireEisumls Lebeckia septada 2 8,0188814 lanceobta &Anis amboensis P0010,1;)th"a 010Par6:0050, FIG. 1. Strict consensus tree of 560 shortest len gth trees from the com- 1X0R2a3 *bauS 04ctaBus kb:skit:Arcs bined analysis of the ITS and rbcL data sets (TL = 1,473 ; CI = 0.50; RI = 85 acklus graCclis Metobblum candicaas 1 0.86). Numbers above the branches are bootstrap percenta ges above 50% 1.0 Mobbbilim actonodas 1 C itrgyroiatxum funoro from the maximum parsimon y analysis and numbers below the branches Argyrott9Nro llarmstaort Argrolobaum tartocolatum are posterior probabilities above 0.5 from the Ba yesian analysis. The grey Avoceatmen rnagarrnizurn 6 Arr43111111skrJ names or lines indicate where the topolo gy differed in the Ba yesian AcricicallaW nth:bran:ft Ampirahals0 weiancsacil analysis. 0 97 Xioaftothaco culbra a) Xephottroca purteroi to Y. 138 Up311,3 ongurak193 9 40=1 /8=0 54 ,8 Podatyria Ocadalca To 93 a P00a0rfa Lebeckia section Lebeckia is unresolved. Lotononis is left unre- 0 StrIcoargRos chey9989,98 100 C)T19819 Var,8114 solved and L. hirsuta is well separated from the rest of the genus. Three main clades can be distinguished. The first con- rambCafffa 044.31:01003erl."1a sists of Lotononis s.s. (also supported in the rbcL tree) with FIG. 1. (Continued) moderate support (84 BP). The second Glade, comprising taxa from Lotononis section Leptis (E.Mey. ex Eckl. & Zeyh.) Benth., L. section Listia, and their allies also has low support (64 BP). Combined ITS/rbcL Data Set—Visual inspection of the The third Glade of Lotononis groups with Bolusia and Crota- bootstrap consensus trees resulting from separate analyses of laria with strong support (87 BP) and comprises one anoma- ITS and rbcL sequences presented no strongly supported in- lous species, L. hirsuta, which makes up the monotypic sec- congruent patterns. The ILD test indicated significant differ- tion Euchlora. Crotalaria and Bolusia are weakly supported as ence between the two datasets (p=0.001). Following the sug- sister taxa (64 BP) and Crotalaria is strongly supported to be gestions of Seelanan et al. (1997) and Wiens (1998), together monophyletic (99 BP). Pearsonia, Robynsiophyton, and Rothia with suggestions that the ILD test may be unreliable (Reeves form a strongly supported Glade (98 BP) and the latter two et al. 2001; Yoder et al. 2001) these datasets were combined are strongly supported to be sister genera (97 BP). The taxa of directly. The combined ITS and rbcL matrix consisted of 1,854 this Glade all possess a 17 base-pair deletion at positions 179- included positions, of which 540 were variable and 404 par- 196 in the aligned ITS matrix. The Crotalarieae are strongly simony informative. The MP analysis produced 560 equally supported to be monophyletic (81 BP) and sister to Genisteae parsimonious trees (Fig. 1 ; TL = 1,473; CI = 0.50; RI = 0.86). (100 BP). In both the MP and BI analyses, the same major clades 2008] BOATWRIGHT ET AL.: CROTALARIEAE PHYLOGENY 755 could be observed: the "Cape" group consisting of Aspala- within this group. Lotononis hirsuta is not included in Lotono- thus, Lebeckia, Rafnia, Spartidium, and Wiborgia (53 BP; PP 1.0); nis, but groups with Crotalaria (63 BP) and Bolusia as in the the Lotononis group consisting of Lotononis s.s., Pearsonia, molecular analysis. The rest of Lotononis is monophyletic (56 Robynsiophyton, and Rothia (PP 0.94); a second Lotononis Glade BP) in the morphological analysis as opposed to paraphyletic consisting of Lotononis section Leptis, L. section Listia, and in the combined molecular analysis, but relationships within allies (77 BP; PP 1.0); and the Crotalaria group consisting of the genus are largely unresolved. Pearsonia, Rothia, and Bolusia, Crotalaria, and Lotononis hirsuta/L. section Euchlora (99 Robynsiophyton form a strongly supported Glade (87 BP) and BP; PP 1.0). Lebeckia, Lotononis, and Wiborgia are all paraphy- a sister relationship between Robynsiophyton and Rothia is letic and Lebeckia section Lebeckia is unresolved in both the strongly supported (88 BP). analyses. Combined ITS/rbcL/Morphological Data Set—The boot- Generic relationships are unresolved within the "Cape" strap consensus trees from the combined analysis of ITS and group and represented as a polytomy at the base of this Glade rbcL and the morphological analysis showed no strongly sup- in the bootstrap consensus tree of the parsimony analysis ported incongruent patterns, although an ILD test indicated (not shown). In the BI tree, Aspalathus (88 BP; PP 1.0) and significant difference between the data sets (p=0.002). The Wiborgia (70 BP; PP 1.0) are sister to each other, with L. mu- suggestions of Seelanan et al. (1997) and Wiens (1998) were cronata (PP 0.96) and Lebeckia inflata included in this Glade. followed and the data sets combined directly. The combined These positions are unresolved in the MP analysis. Lebeckia matrix consisted of 1,885 characters, 1,405 of which were con- section Calobota (including L. section Stiza and Spartidium sa- stant, 480 variable, and 303 parsimony informative. The MP harae; 93 BP; PP 1.0) is sister to Aspalathus and Wiborgia, fol- analysis resulted in 370 trees (TL = 1,166, CI = 0.53, RI = 0.84; lowed by L. section Viborgioides (including Wiborgia humilis; Fig. 2 ). 64 BP; PP 1.0) and Rafnia (94 BP; PP 1.0). Lebeckia section The trees resulting from this analysis are similar to those Lebeckia is unresolved in both the MP and BI trees. from the combined molecular analysis, except that Lotononis Lotononis is paraphyletic in both the MP and 131 trees. The and Lebeckia section Lebeckia are monophyletic. The "Cape" Lotononis group is moderately supported in the BI tree (PP group (73 BP), Lotononis group (including Pearsonia, Robyn- 0.94) and consists of those species of Lotononis from the sec- sionphyton, and Rothia), and Crotalaria group (100 BP) found tions Aulacinthus (E.Mey.) Benth., Buchenroedera (Eckl. & in the molecular analysis were also retrieved in the combined Zeyh.) B.-E.van Wyk, Cleistogama B.-E.van Wyk, Krebsia (Eckl. data set of molecular plus morphological characters. Within & Zeyh.) Benth., Lotononis, Monocarpa B.-E.van Wyk, Oxy- the "Cape" group, Aspalathus, Lebeckia section Calobota (in- dium Benth., and Polylobium (Eckl. & Zeyh.) Benth. (100 BP; cluding L. section Stiza and Spartidium saharae), L. section PP 1.0). These are sister to a strongly supported Glade (BP 100; Lebeckia, L. section Viborgioides (including Wiborgia humilis), PP 1.0) consisting of Pearsonia (98 BP; PP 1.0), Robynsiophyton, Rafnia, and Wiborgia (excluding Wiborgia humilis) received and Rothia. The sister relationship of the latter two genera has moderate to strong support as monophyletic (99 BP; 95 BP; 77 high bootstrap support (97%) and PP (1.0). BP; 87 BP; 98 BP; 94 BP, respectively). Lotononis is monophy- The next Glade consists of species of Lotononis section Listia letic (75 BP; excluding L. hirsuta) and the groups retrieved (100 BP; PP 1.0) and sections Digitata B.-E.van Wyk, Leobordea within the genus are identical to the separate clades found in (Del.) Benth., Leptis, Lipozygis (E.Mey.) Benth., and Synclistus the molecular analysis, namely Lotononis s.s. (100 BP), Lotono- B.-E.van Wyk (99 BP; PP 1.0), and these are moderately to nis section Leptis and allies (100 BP), and Lotononis section strongly supported as sister (77 BP; 1.0 PP). Listia (100 BP). The sister relationship between the latter two The Crotalaria Glade consists of Bolusia and Crotalaria (100 is moderately supported with a BP of 88. Pearsonia is strongly 13P; PP 1.0), strongly supported as sister groups (92 BP; PP supported as monophyletic (99 BP) and sister to Robynsiophy- 1.0), and Lotononis hirsuta (corresponding to Lotononis section ton and Rothia (100 BP). The sister relationship between Euchlora). This Glade receives high support in both the MP Robynsiophyton and Rothia is strongly supported (99 BP). The and BI analyses (99 BP; PP 1.0). Crotalaria group is strongly supported (100 BP) and consists The Crotalarieae are strongly supported as being mono- of Bolusia, Crotalaria and Lotononis hirsuta. Crotalaria is phyletic (97 BP; PP 1.0) and sister to Genisteae (100 BP; PP strongly supported to be monophyletic (100 BP) and is sister 1.0). The monophyly of the latter tribe receives low to strong to Bolusia (94 BP). support (65 BP; PP 1.0). The Podalyrieae s.s. are strongly supported as monophyletic (93 BP; PP 1.0) and the relationships within the tribe conform to those of previous studies DISCUSSION (Boatwright et al. 2008). Phylogenetic Relationships—The phylogenetic hypoth- Morphological Analysis—The morphological matrix in- eses presented in this study are based on a complete sample cluded 31 polarised characters, 29 of which were parsimony of Crotalarieae at the generic level and a representative informative. Parsimony analysis yielded 261 trees (not sample of most of the taxonomic and morphological varia- shown) of 101 steps (CI = 0.36, RI = 0.93). In the strict con- tion within these genera. The data were based not only on sensus tree (not shown), Aspalathus and Rafnia group to- DNA sequences but also on salient morphological characters gether, albeit without support. Aspalathus (excluding A. cre- that were carefully polarised and proved to be informative at nata and A. perfoliata) and Rafnia are weakly supported as the generic level. The low resolution within the Crotalarieae monophyletic (55 BP and 71 BP, respectively). Lebeckia section based on morphology and some chemical and cytological Viborgioides, L. mucronata, and Wiborgia s.s. group together, characters reflects the somewhat reticulate relationships while L. inflata groups with Lebeckia section Lebeckia, but with- within the tribe (also mentioned by van Wyk 1991a; van Wyk out support. Only Wiborgia is weakly supported as mono- 2005) with remarkable examples of parallelism, convergence, phyletic (78 BP). Lebeckia section Lebeckia is moderately sup- and analogy with regard to vegetative and reproductive mor- ported as monophyletic (84 BP), but there is no resolution phology as outlined by Dahlgren (1970a). The presence of

756 SYSTEMATIC BOTANY [Volume 33

Loasnotas &taut 1 100 Aspefathus 'Meads LeareuaLa parr..14ara C Aspalathus pondula -- Loa:awes Ote.tot4-5 Aspatathus camosa —1..otonams brxtl ysrora Aspatathus veradadals toxer57..s sp.ma1a42 0007Oryp;y.1 67 Aspatathus shawii subsp. !i8 L0 Utonart. M4600 74194p. frartiVurrtS13 63 Aspatethus latfcilem subsp. lartailette a Lceonc4r:4 krt10.44 Aspalathus NMI subsp. Mao 1.45.015,45 bra Aspatathus hystrix 93 73E 1.13:00an:3 512croscp84a 52 Aspalathus gataate 87 14:1072/r:517....5,0 62 Aspalathus mar:fondle Lamm:3 moccesto 94r Aspalathus witkienowiarta L— Ladortoes carte Aspalathus confine LakonoM wine= 96 1-Aspatathus perfoliate subsp. 99 1.5•:4555fs epatabo 2 L.4554445.4 maJorre....;5.5, 1 Aspatathus 0(00079 tO0 La•anaras mavens 2 Aspatathus pachyloba Lamat dam:r..- ASp010111u5 alma I Lame:was sencophyCa 1 Wiborgia testa 1 ■ _FE 74 (0700Ortit fr.eCte.. Wibotgia fetrepteto 1 LoZonenis maven 1 Martaaatfas Kaborgla senae8 97 Wiborgia obcordata 1 fie r— Latonortisatrkaa (emmas maim bogie menoptera 1 190 IMancets dam Wm. t-e.,,e,e*.eta C Naborgta incumato 70 Laamm cleVotesia 93 Whergta humills 1 LoMom ierMacraal 0) 887 10700000 in...Marata Mm. jaMmegarts 87 Le:made kapoldtiana 1 01) LC40404:5 P040,44.$ Lebeckia sessilitoffa co (6 75 C Lcconem caarqata Lebeckia bowieena 1 (6 o! Lononmis Xaryearpo 5 Lebeckia inflate 2 Comma catyrina Lebeckia mucronata 0 : G25045,44 7e-oso 2 Lebeckia pungens 2 Lamortia cosaMa 100 Mmrx Wipe= Lebeckia amaranths 2 59 La. Lobockto psdototta Lammas dammens 73 L.Mrawas actaloam LabOCkfil Sericea 2 Lammas boaera7 100 Lebeckia cylisolies 2 95 Lebeckia muttiffora —E to 4ta I Lebeckia totoneneides Lotortcaaa 9441710 81 Lebeckia halenbergensis 2 Lotanolimaeycapaab 2 Lotoncraa amMaryff. a Lebeckia spinescons Latoemai benth2mis.4 Lebeckia cinema Lotonones cavaata 1 Sparticawn saheme Le:gamma razgralta Lebeckia camosa 2 Latonor4s obbixasa Lotarmes ste;Zurtnet Lebockio moyerfarta 1 100 99 LotomnIsepubt&ta Lebeckia phakenotians 1 100 otarton:s GsGi 7 Lebockia pauciffera 2 Lotanom wow,* Lebockta ntrightii 1 100 PezmW.5 se..22&.4 1 LebeckM broyicarpa 99 Poarso,42 aeivato 97 Partraam warataM Lebockia ambigua 1 100 PavraM a*Mala 2 Lebeckia septarfa 1 49 7752041621451 9 Rafnia OCuminata Raba:tame:Ma vandayalta lh Rata, recomese La:amnia nested 1 I Rotate spicate 103 r— CrataMa OEMs CrotaMa greavansil 71 Raffia globose 100 52 97 Ceataaaa caper= 1 Reale diffuse 100 00 Ceotarea viepaaLs 94 0MM:a 7um2:s 1 FIG. 2. Strict consensus tree from the combined molecular (ITS and Croteranio bnax0a14 80000 ainDoens:s 1 rbcL) and morphological analysis. Numbers above the branches are boot- 0:04.,3970777:6 Genisteae strap percentages above 50% (no. of trees = 370; TL = 1,166; CI = 0.53; RI = 0.84). FIG. 2. (Continued) seemingly identical apomorphic states in unrelated genera suggested by Polhill (1976) and van Wyk (1991a), while Spar- (e.g. hairy petals in Lebeckia section Calobota and Lotononis tidium was thought to either be closely allied to Genisteae or section Leptis) complicates the cladistic analysis. In the ab- to Lebeckia, but the affinities remained unclear (Polhill 1976). sence of single apomorphies with diagnostic value, natural The results of the current study indicate four major lineages groups can only be delimited using combinations of charac- within the Crotalarieae comprising the "Cape" group (As- ters. Our results (both molecular and morphological) support palathus, Lebeckia, Rafnia, Wiborgia, including Spartidium sah- some of the current generic concepts within the tribe, but arae), two lineages comprising genera with zygomorphic ca- provide novel insights into as yet phylogenetically unstudied lyces (Lotononis, Pearsonia, Robynsiophyton and Rothia), plus groups such as Lebeckia and the smaller genera of the Crota- Bolusia, Crotalaria, and Lotononis section Euchlora. larieae, namely Robynsiophyton, Rothia, and Spartidium. The "Cape" Group— The lack of resolution between genera Previous studies on relationships within the Crotalarieae in this group is caused by the low sequence divergence have all suggested a close relationship between the "Cape" among these taxa. Aspalathus and Rafnia are both strongly genera (viz., Aspalathus, Lebeckia, Rafnia, and Wiborgia), as supported as monophyletic and indicated as sister taxa by well as the genera that have a zygomorphic calyx (viz., the morphological data because they share sessile leaves and Lotononis, Pearsonia, Robynsiophyton, and Rothia; Dahlgren an asymmetrical upper suture of the pod (Campbell and van 1963, 1967, 1970a; Polhill 1981; van Wyk 1991a). The place- Wyk 2001). Aspalathus is placed closer to Wiborgia by both the ment of Bolusia, Crotalaria, and Spartidium has been uncertain. MP and BI analyses. Although the sample of Aspalathus is not A sister relationship between the former two genera was at all comprehensive, some clades of interest can be noted. 2008] BOATWRIGHT ET AL.: CROTALARIEAE PHYLOGENY 757

Aspalathus linearis and A. pendula both have simple, terete The positions of both L. inflata and L. mucronata are unre- leaves and this Glade corresponds to the Lebeckiiformes solved in the combined ITS and rbcL analysis. Lebeckia mu- group (Dahlgren 1988), named for its similarity to Lebeckia cronata is currently placed within L. section Calobota. Lebeckia section Lebeckia. Aspalathus laricifolia, A. hirta, A. hystrix, and inflata was described by Bolus (1887) without any reference to A. shawii all have spine-tipped leaves (corresponding to the its position in the infrageneric systems of Bentham (1844) and Laterales group of Dahlgren 1988) and form a Glade, which is Harvey (1862). Both of these species, however, share several well-supported in the BI tree. The Borboniae group (Dahl- anatomical and morphological characters with species of L. gren 1988) represented by A. perfoliata and A. crenata form a section Viborgioides, such as a 6 + 4 anther arrangement, gla- weak to strongly supported Glade. Within Rafnia, two clades brous petals, and the dorsiventral leaves with mucilage cells. can be identified that correspond to the two sections of the When morphological characters were combined with the mo- genus described by Campbell and van Wyk (2001): the R. lecular ITS and rbcL data, L. inflata was included in a Glade acuminata Glade representing R. section Rafnia, and the R. along with L. section Viborgioides and Wiborgia humilis, while spicata Glade representing R. section Colobotropis E.Mey. These the position of L. mucronata remained unresolved. two sections differ mainly in the rostrate keel petals and The Lotononis Clades—Based on the molecular data, unequal calyx lobes of R. section Colobotropis. Lotononis is paraphyletic and consists of three clades. Lotono- It is clear from both the MP and BI analyses that Lebeckia nis s.s. groups with the Pearsonia Glade (albeit without sup- and Wiborgia are not monophyletic. Lebeckia is represented by port in the MP analysis) and is strongly supported as mono- several smaller clades within the "Cape" group, while Wibor- phyletic. Two groups were noted within this Glade. The first gia humilis is nested within the Lebeckia leipoldtiana Glade and consists of representatives of L. section Oxydium and the sec- well-separated from the main Wiborgia Glade. ond of representatives of L. sections Aulacinthus, Buchenroed- The species of Lebeckia section Calobota form a well- era , Cleistogama, Krebsia, Lotononis, Monocarpa, and Polylobium. supported Glade with those of L. section Stiza and the mono- Lotononis section Oxydium is strongly supported to be monotypic genus Spartidium. These species are all shrubs with tri- phyletic in the molecular analyses and currently consists of foliolate or unifoliolate leaves and green mature stems. The 35 species (accommodated in 14 subsections as described by petals are usually pubescent and they possess a 5 + 4 + 1 van Wyk 1991b) distinguished from other sections by several anther configuration (five short dorsifixed anthers, four long characters, including single stipules at each leaf base and basifixed anthers, and an intermediate carinal anther). Some glabrous wing and keel petals. In the second group, L. section of these character states also occur in Lotononis species and Cleistogama is sister to L. section Monocarpa, while L. sections are thus not informative in the morphological analysis. The Aulacinthus, Lotononis, and Polylobium form a Glade without species of L. section Stiza and L. section Calobota occur in the support. The resolution in the rest of the group is low and Cape and those of the latter also extend northwards into relationships not clear. In the molecular analysis, Lotononis Namibia. Spartidium saharae is a North African plant that has s.s. and the Pearsonia Glade clearly share similarities in their had uncertain affinities within the tribe. The close relation- sequences but morphologically Lotononis s.s. allies with the ship of this species to L. section Calobota has also been dem- rest of the genus to form a monophyletic assemblage. From onstrated independently by Edwards and Hawkins (2007) these results it is clear that some generic apomorphies sup- based on ITS data. port the current concept of the genus. When the molecular Wiborgia and species of Lebeckia section Viborgioides are su- data were combined with the morphology, Lotononis re- perficially very similar in their rigid, woody habit, glabrous mained monophyletic, although with moderate support. It is petals and highly reflexed standard petals in most species. interesting to note the close agreement between the sectional However, they differ markedly in the winged, few-seeded classification proposed by van Wyk (1991b) and the results of samaras that are typical of Wiborgia species (Dahlgren 1975). this study. Pearsonia, Robynsiophyton, and Rothia share several However, Wiborgia humilis has thin-walled, inflated pods that morphological characters that are unique among the genera lack the dorsal wing characteristic of Wiborgia (Dahlgren of Crotalarieae, such as the uniform anthers, straight styles, 1970b). The fruits of Lebeckia section Viborgioides are typically and presence of angelate esters of hydroxylupanine. The ge- turgid, thin-walled, many-seeded, and wingless. Wiborgia hu- neric concepts of Robynsiophyton and Rothia, based mainly on milis is the only species of Wiborgia with a 6 + 4 anther ar- anther characters, have been doubtful and Robynsiophyton rangement (6 short dorsifixed and 4 long basifixed anthers) was thought to be a local derivative of Pearsonia (van Wyk instead of the 5 + 4 + 1 arrangement found in all other spe- 1991a). Taxonomic studies of the former two genera and a cies. These characters therefore support the placement of W. thorough examination of their morphology and anatomy re- humilis in the L. section Viborgioides Glade. vealed that these generic concepts are indeed sound and em- Lebeckia section Lebeckia is unresolved in the BI and MP phasizes the value of androecial characters in the Crotalar- analyses. This section is morphologically very distinct from ieae (Boatwright and van Wyk, unpubl.; Boatwright et al., the other sections in the genus and its monophyly is sup- unpubl.). ported by four apomorphies, namely the suffrutescent habit, The second Glade comprises those species from Lotononis acicular leaves, a 5 + 5 anther arrangement, and rugose seeds section Leptis, Lotononis section Listia, and allies. This Glade is (Le Roux 2006; Le Roux and van Wyk 2007). This is clear from moderately to strongly supported in the molecular analysis the moderate support this group received in the morphologi- but lacks resolution in the morphological analysis. The con- cal analysis as well as the combined morphological and mo- cept of the hitherto monotypic Listia was broadened by van lecular analysis. Lebeckia lotononoides and L. inflata are the Wyk (1991b) and treated as a section of Lotononis with several only other species of Lebeckia with rugose seeds, but these distinct characters. This broadened concept is supported in species share characters with L. section Calobota and L. sec- the analyses presented in this study. The absence of suitable tion Viborgioides, respectively (Boatwright and van Wyk 2007; material of the rare Lotononis macrocarpa Eckl. & Zeyh. (an Boatwright, pers. obs.). anomalous species within this group) prevented its inclusion 758 SYSTEMATIC BOTANY [Volume 33 in this study. Special efforts should be made to obtain DNA Bryan, W. W. 1961. Lotononis bainesii Baker - a legume for subtropical from this species in order to evaluate its placement within L. pastures. Australian Journal of Experimental Agriculture and Animal Husbandry 1: 4-10. section van Wyk (1991b) mentions that no apomor- Listia. Campbell, G. J. and B.-E. van Wyk. 2001. A taxonomic revision of Rafnia phies exist for L. section Leptis, which is clear from the mor- (Fabaceae, Crotalarieae). South African Journal of Botany 67: 90-149. phological analysis where this section is left unresolved. The Crisp, M. D., S. Gilmore, and B.-E. van Wyk. 2000. Molecular phyloge- molecular results, however, indicate a close relationship be- netics of the genistoid tribes of Papilionoid Legumes. Pp. 249-276 in tween this section and L. sections Digitata, Leobordea, Lipo- Advances in Legume Systematics 9, eds. P. S. Herendeen and A. Bru- neau. Kew: Royal Botanic Gardens. zygis, and Synclistus, all of which are successively sister to L. Dahlgren, R. 1963. Studies on Aspalathus and some related genera in section Listia. The third Glade comprises L. hirsuta, which South Africa. Opera Botanica 9: 1-301. groups with the Crotalaria Glade. Dahlgren, R. 1967. Chromosome numbers in some South African genera of the tribe Genisteae s. lat. (Leguminosae). Botaniska Notiser 120: The Crotalaria Clade -The close relationship between Cro- talaria and Bolusia has been mentioned by both Polhill (1982) 149-160. Dahlgren, R. 1970a. Current topics - parallelism, convergence, and anal- and van Wyk (1991a) and these authors suggested that Bolu- ogy in some South African genera of Leguminosae. Botaniska Notiser sia could merely be a local derivative of Crotalaria. Data from 123: 551-568. the present study indicates that Bolusia is sister to and not Dahlgren, R. 1970b. Wiborgia apterophora R. Dahlgr., a new species of embedded within Crotalaria (rbcL analysis with multiple rep- Leguminosae from the Cape Province. Botaniska Notiser 123: 112-114. Wiborgia Thunb. and related species of resentatives), so that its generic status seems justified. Bolusia Dahlgren, R. 1975. Studies on Lebeckia Thunb. (Fabaceae). Opera Botanica 38: 6-83. differs from Crotalaria mainly in its glabrous style and Dahlgren, R. 1988. Crotalarieae. Pp. 1-430 in Flora of southern Africa, vol. strongly coiled keel petals. Also included in this Glade is 16, part 3, fascicle 6, ed. 0. A. Leistner. Pretoria: CTP Book Printers. Lotononis hirsuta, representing the monotypic section Eu- Doyle, J. J. and J. L. Doyle. 1987. A rapid DNA isolation procedure for chlora. This species differs from all other Lotononis species in small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11-15. Doyle, J. J., J. A. Chappill, C. D. Bailey, and T. Kajita. 2000. Towards a its peculiar tuberous habit and from most others in the very comprehensive phylogeny of legumes: evidence from rbcL sequences large, inflated pods, equally lobed calyx, and very large seeds and non-molecular data. Pp. 1-20 in Advances in Legume Systematics with smooth surfaces. It shares with Crotalaria and Bolusia the 9, eds. P. S. Herendeen and A. Bruneau. Kew: Royal Botanic Gar- strongly inflated pods. dens. Doyle, J. J., J. L. Doyle, J. A. Ballenger, E. E. Dickson, T. Kajita, and H. Implications for Generic Classification -The results gen- Ohashi. 1997. A phylogeny of the chloroplast gene rbcL in the Legu- erated in this study have important implications for the ge- minosae: Taxonomic correlations and insights into the evolution of neric classification system of the Crotalarieae. Detailed stud- nodulation. American Journal of Botany 84: 541-554. ies of character variation in the tribe over a period of more Edwards, D. and J. A. Hawkins. 2007. Are Cape Floral Clades the same than 23 yr are now nearing completion, allowing us to make age? Contemporaneous origins of two lineages in the genistoids s./. an informed evaluation of the congruence between morpho- (Fabaceae). Molecular Phylogenetics and Evolution 45: 952-970. Farris, J. S., M. Kallersjo, A. G. Kluge, and C. Bult. 1995. Constructing a logical and molecular patterns. The results clearly show that significance test for incongruence. Systematic Biology 44: 570-572. the genera Calobota Eckl. & Zeyh. and Euchlora Eckl. & Zeyh. Fay, M. F., S. M. Swensen, and M. W. Chase. 1997. Taxonomic affinities of should be reinstated. A new genus should be described that Medusagyne oppositifolia (Medusagynaceae). Kew Bulletin 52: 111-120. will include Lebeckia sect. Viborgioides, as well as L. inflata and Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using L. mucronata. Spartidium will be transferred to Calobota and the bootstrap. Evolution 39: 783-791. Fitch, W. M. 1971. Towards defining the course of evolution: minimum Wiborgia humilis to the new genus (Boatwright et al., un- change for a specified tree topology. Systematic Zoology 20: 406-416. publ.). These changes, about to be formalized, will clearly Harvey, W. H. 1862. Leguminosae. Pp. 82-89 Flora Capensis, vol. 2, eds. result in a more practical and predictable generic classifica- W. H. Harvey and 0. W. Sonder. Dublin: Hodges, Smith, and Co. tion system for the tribe Crotalarieae. Huelsenbeck, J. P. and F. Ronquist. 2001. MRBAYES: Bayesian inference of phylogeny. Bioinformatics (Oxford, England) 17: 754-755. Kajita, T., H. Ohashi, Y. Tateishi, C. D. Bailey, and J. J. Doyle. 2001. rbcL ACKNOWLEDGMENTS. The authors thank the National Research Foun- and legume phylogeny, with particular reference to Phaseoleae, Mil- dation and University of Johannesburg for funding. We thank Kew Her- lettieae, and allies. Systematic Botany 26: 515-536. barium and Amida Johns for plant samples and the Jodrell Laboratory for Kass, E. and M. Wink. 1995. Molecular phylogeny of the Papilionoideae extraction of the Kew material. The Molecular Systematics Laboratory at (Family Leguminosae): rbcL gene sequences versus chemical tax- the University of Johannesburg is greatfully acknowledged for the use of onomy. Botanica Acta 108: 149-162. their facilities. The curators and staff of the listed herbaria are thanked for Kass, E. and M. Wink. 1996. Molecular evolution of the Leguminosae: making specimens available for study. Some of the sequences generated Phylogeny of the three subfamilies based on rbcL-sequences. Bio- at the University of Heidelberg came from Ernst Kass and Gamal I. A. chemical Systematics and Ecology 24: 365-378. Mohamed. Kass, E. and M. Wink. 1997. Phylogenetic relationships in the Papilion- oideae (Family Leguminosae) based on nucleotide sequences of Molecular Phylogenetics and LITERATURE CITED cpDNA (rbcL) and ncDNA (ITS 1 and 2). Evolution 8: 65-88. Alvarez, I. and J. F. Wendel. 2003. Ribosomal ITS sequences and plant Le Roux, M. M. 2006. A taxonomic study of the type section of the genus phylogenetic inference. Molecular Phylogenetics and Evolution 29: 417- Lebeckia Thunb. (Fabaceae, Crotalarieae). M.S. thesis. Johannesburg: 434. University of Johannesburg. Bentham, G. 1844. Enumeration of Leguminosae indigenous to southern Le Roux, M. M. and B.-E. van Wyk. 2007. A revision of Lebeckia sect. Asia and central and southern Africa. Hooker's London Journal of Lebeckia: The L. sepiaria group. South African Journal of Botany 73: Botany 3: 355-363. 118-130. Boatwright, J. S. and B.-E. van Wyk. 2007. The identity of Lebeckia lotono- Lewis, G., B. Schrire, B. Mackinder, and M. Lock. [eds.]. 2005. Legumes of noides (Crotalarieae, Fabaceae). South African Journal of Botany 73: the World. Kew: Royal Botanic Gardens. 664-666. Moteetee, A. N. and B.-E. van Wyk. 2007. The concept of 'Musa-pelo and Boatwright, J. S., V. Savolainen, B.-E. van Wyk, A. L. Schutte, F. Forest, the medicinal use of shrubby legumes (Fabaceae) in Lesotho. Bothalia and M. van der Bank. 2008. Systematic position of the anomalous 37: 75-77. genus Cadia and the phylogeny of the tribe Podalyrieae (Fabaceae). Motsi, M. C. 2004. Molecular phylogenetics of the genus Rafnia Thunb. (Fa- Systematic Botany 33: 133-147. baceae, Crotalarieae). M.S. thesis. Johannesburg: Rand Afrikaans Uni- Bolus, H. 1887. Lebeckia inflata. Hooker's Icones plantarum 16, plate 1576. versity. 2008] BOATWRIGHT ET AL.: CROTALARIEAE PHYLOGENY 759

Miiller, K. 2005. SeqState — primer design and sequence statistics for Wojciechowski, M., M. Lavin, and M. J. Sanderson. 2004. A phylogeny of phylogenetic data sets. Applied Bioinformatics 4: 65-69. legumes (Leguminosae) based on analysis of the plastid matK gene Pardo, C., P. Cubas, and H. Tahiri. 2004. Molecular phylogeny and sys- resolves many well-supported subclades within the family. American tematics of Genista (Leguminosae) and related genera based on Journal of Botany 91: 1846-1862. nucleotide sequences of nrDNA (ITS region) and cpDNA (trnL-trnF Wortley, A. H. and R. W. Scotland. 2006. The effect of combining mo- intergenic spacer). Plant Systematics and Evolution 244: 93-119. lecular and morphological data in published phylogenetic analyses. Polhill, R. M. 1974. A revision of Pearsonia (Leguminosae-Papilionoideae). Systematic Biology 55: 677-685. Kew Bulletin 29: 383-410. Yang, Z. and B. Rannala. 1997. Bayesian phylogenetic inference using Polhill, R. M. 1976. Genisteae (Adans.) Benth. and related tribes (Legu- DNA sequences: a Markov Chain Monte Carlo method. Molecular minosae). Botanical Systematics 1: 143-368 Biology and Evolution 14: 717-724. Polhill, R. M. 1981. Tribe 29. Crotalarieae (Benth.) Hutch. Pp. 399-402 in Yoder, A. D., J. A. Irwin, and B. A. Payseur. 2001. Failure of the ILD to Advances in Legume Systematics 1, eds. R. M. Polhill and P. H. Raven. determine data combinability for slow loris phylogeny. Systematic Kew: Royal Botanic Gardens. Biology 50: 408-424. Polhill, R. M. 1982. Crotalaria in Africa and Madagascar.Netherlands: A. A. Balkema Publishers, Rotterdam. Posada, D. and K. A. Crandall. 1998. Modeltest: testing the model of DNA APPENDIX 1. Voucher information and GenBank accession numbers of substitution. Bioinformatics (Oxford, England) 14: 817-818. the taxa sampled in this study. Voucher specimens are deposited in the Reeves, G., M. W. Chase, P. Goldblatt, M. F. Fay, A. V. Cox, B. Lejeune, following herbaria: Australian National Herbarium (CANB), University and T. Souza-Chies. 2001. Molecular systematics of Iridaceae: evi- of Johannesburg Herbarium (JRAU), Kew Herbarium (K), National Her- dence from four plastid DNA regions. American Journal of Botany 88: barium, Pretoria (PRE), Southern Cape Herbarium (SCHG). The informa- 2074-2087. tion is listed as follows: taxon—GenBank accessions: rbcL, ITS; Voucher Ronquist, F. and J. P. Huelsenbeck. 2003. MRBAYES 3: Bayesian phylo- specimen. Those regions not sampled for a taxon are represented by a genetic inference under mixed models. Bioinformatics (Oxford, En- dash ['Boatwright et al. (2008); 'Crisp et al. (2000); 'Kass and Wink (1997); gland) 19: 1572-1574. 'Doyle et al. (1997); 'Pardo et al. (2004); 6Motsi (2004); 'Van der Bank et Schutte, A. L. and B.-E. van Wyk. 1988. A synopsis of the genus Dichilus al. (2002)]. (Fabaceae-Crotalarieae). South African Journal of Botany 54: 182-184. Amphithalea A. alba Granby— AM180171 1, AM261217 1; Scotland, R. W., R. G. Olmstead, and J. R. Bennett. 2003. Phylogeny re- Eckl. & Zeyh.: 2125 (SCHG). (E.Mey.) Walp.— AM180182 1 , construction: the role of morphology. Systematic Biology 52: 539-548. Van Wyk A. micrantha AM261227 1; Schutte 751 (SCHG). A. williamsonii Harv.— AM177372 1 , Seelanan, T., A. Schnabel, and J. F. Wendel. 1997. Congruence and con- AM261436 1 ; Euston-Brown s.n. (SCHG). Argyrolobium Eckl. & Zeyh.: A. sensus in the cotton tribe (Malvaceae). Systematic Botany 22: 259-290. harmsianum Schltr. ex Harms— —, AF287685 2; Crisp 9042 (CANB). A. Simmons, M. P. and H. Ochoterena. 2000. Gaps as characters in sequence lunare (L.) Druce 1— —, AF287686 2; Crisp 9039 (CANB). A. lunare (L.) based phylogenetic analyses. Systematic Biology 49: 369-381. Druce subsp. sericeum (Thunb.) T.J.Edwards 2— EU348013, —; Van Wyk Sun, Y., D. Z. Skinner, G. H. Liang, and S. H. Hulbert. 1994. Phylogenetic 2080 (JRAU). A. marginatum Bolus— EU348014, —; Van Wyk 3038 (JRAU). analysis of sorghum and related taxa using internal transcribed spac- A. marginatum Bolus— Z955473, —; Edwards 471. A. megarrhizum Bolus- ers of nuclear ribosomal DNA. Theoretical and Applied Genetics 89: EU348015, —; Germishuizen s.n. (JRAU). Argyrolobium sp.— EU348016, —; 26-32. Van Wyk 2173 (JRAU). Asp alathus L.: A. acutiflora Dahlgren— —, Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony EU347735; Boatwright & Magee 22 (JRAU). A. carnosa Berg. — EU348022, (*and other methods). Sunderland: Sinauer Associates. EU347723; Boatwright & Magee 61 (JRAU). A. confusa Dahlgren- van der Bank, M., M. W. Chase, B.-E. van Wyk, M. F. Fay, F. H. van der EU348031, EU347736; Boatwright et al. 97 (JRAU). A. crenata (L.) Dahl- Bank, G. Reeves, and A. Hulme. 2002. Systematics of the tribe Po- gren— EU348028, EU347729; Boatwright & Magee 70 (JRAU). A. galeata dalyrieae (Fabaceae) based on DNA, morphological and chemical E.Mey.— EU348032, EU347726; Boatwright et al. 100 (JRAU). A. hirta data. Botanical Journal of the Linnean Society 139: 159-170. E.Mey. subsp. hirta— EU348025, EU347732; Boatwright & Magee 43 van Wyk, B.-E. 1991a. A review of the tribe Crotalarieae (Fabaceae). Con- (JRAU). A. hystrix L.f.— EU348023, EU347734; Boatwright & Magee 30 tributions from the Bolus Herbarium 13: 265-288. ORAU). A. laricifolia Berg. subsp. laricifolia— EU348026, EU347731; Boat- van Wyk, B.-E. 1991b. A synopsis of the genus Lotononis (Fabaceae: Cro- wright & Magee 46 (JRAU). A. linearis (Burm.f.) Dahlgren 1— EU348020, talarieae). Contributions from the Bolus Herbarium 14: 1-292. EU347722; Van Wyk et al. 4192 GRAU). A. linearis (Burm.f.) Dahlgren 2— van Wyk, B.-E. 2003. Bolusia. Pp. 228-233 in Flora Zambesiaca vol. 3 (7), eds. EU348019, EU347739; Van Wyk s.n. (JRAU). A. macrantha Harv.— G. V. Pope, R. M. Polhill, and E. S. Martins. Kew: Royal Botanic EU348029, EU347728; Boatwright & Magee 71 (JRAU). A. nivea Thunb.— Gardens. EU348018, EU347737; Van Wyk 2938 (JRAU). A. pachyloba Benth.— van Wyk, B.-E. 2005. Crotalarieae. Pp. 273-281 in Legumes of the World, EU348024, EU347733; Boatwright & Magee 42 (JRAU). A. pendula Dahl- eds. G. Lewis, B. Schrire, B. Mackinder, and M. Lock. Kew: Royal gren— EU348017, EU347738; Van Wyk s.n. (JRAU). A. perfoliata (Lam.) Botanic Gardens. Dahlgren subsp. phillipsii Dahlgren— EU348030, EU347727; Boatwright & van Wyk, B.-E. and A. L. Schutte. 1989. Taxonomic relationships amongst Magee 82 (JRAU). A. shawii Bol. subsp. shawii— EU348033, EU347725; some genera of Leguminosae tribe Crotalarieae and Agyrolobium Boatwright & Magee 27 (JRAU). A. vermiculata Lam.— EU348021, EU34772; (Genisteae). Kew Bulletin 44: 397-423. Boatwright & Magee 15 (JRAU). A. willdenowiana Benth.— EU348027, van Wyk, B.-E. and A. L. Schutte. 1995. Phylogenetic relationships in the EU347730; Boatwright & Magee 69 (JRAU). Bolusia Benth.: B. acuminata tribes Podalyrieae, Liparieae and Crotalarieae. Pp. 283-308 in Ad- (DC.) Polhill— EU347944, —; Morris 1193 (PRE). B. amboensis (Schinz) vances in Legume Systematics 7, eds. M. Crisp and J. J. Doyle. Kew: Harms 1— EU347943, EU347891; Robinson 1558 (K). B. amboensis (Schinz) Royal Botanic Gardens. Harms 2— EU347943, —; Giess 10091 (K). Cadia Forssk.: C. purpurea (Ait.) van Wyk, B.-E., B. van Oudtshoorn, and N. Gericke. 1997. Medicinal Plants Forssk.— AM260751 1 , AM261740 1; Beckett 1702 (K). Calpurnia E.Mey.: of South Africa. Pretoria: Briza Publications. C. sericea Harv.— AM177374 1 , AM268374 1 & AM268375 1; Boatwright 86 van Wyk, B.-E., F. van Heerden, and B. van Oudtshoorn. 2002. Poisonous (JRAU). Crotalaria L.: C. capensis Jacq. 1— EU348034, EU347884; Van Wyk Plants of South Africa. Pretoria: Briza Publications. 2933a (JRAU). C. capensis Jacq. 2— EU348045, EU347888; Van der Bank 14 White, T. J., T. Bruns, S. Lee, and J. Taylor. 1990. Amplification and direct (JRAU). C. distans Benth.— EU348038, EU347882; de Castro 137 (JRAU). C. sequencing of fungal ribosomal RNA genes for phylogenetics. Pp. griquensis Bolus— EU348043, EU347887; Van Wyk et al. 4173 (JRAU). C. 315-322 in PCR Protocols, eds. M. A. Innas, M. A. Gelfand, J. J. Snin- humilis Eckl. & Zeyh. 1— EU348041, EU347889; Boatwright et al. 143 sky, and T. J. White. New York: Academic Press. (JRAU). C. humilis Eckl. Sr Zeyh. 2— EU348042, EU347890; Boatwright et al. Wiens, J. J. 1998. Combining data sets with different phylogenetic histo- 156 °RAU). C. lanceolata E.Mey.— EU348040, EU347885; Van Wyk 1985 ries. Systematic Biology 47: 568-581. (JRAU). C. lebeckioides Bond— EU348036, —; Van Wyk 3315e (JRAU). C. Wiens, J. J. 2004. The role of morphological data in phylogeny reconstruc- lotoides Benth.— EU348035, —; Van Wyk 5.11. (JRAU). Crotalaria sp.— tion. Systematic Biology 53: 653-661. EU348039, EU347883; Van Wyk 3367 (JRAU). C. virgultalis Burch. ex DC. Wink, M. and G. I. A. Mohamed. 2003. Evolution of chemical defense 1— EU348044, EU347886; Van Wyk et al. 4175 (JRAU). C. virgultalis Burch. traits in the Leguminosae: mapping of distribution patterns of sec- ex DC. 2— EU348037, —; Van Wyk 3060 (JRAU). Cyclopia Vent.: C. genis- ondary metabolites on a molecular phylogeny from nucleotide se- toides (L.) R.Br.— AM261716 AM050819 1 ; Boatwright & Magee 53 quences of the rbcL gene. Biochemical Systematics and Ecology 31: 897- (JRAU). C. subternata Vogel— AM261725 1 , AM050821 1 ; Boatwright & 917. Magee 35 (JRAU). Dichilus DC.: D. gracilis Eckl. & Zeyh.— EU347962, 760 SYSTEMATIC BOTANY [Volume 33

EU347893; Schutte 247 (JRAU). D. lebeckioides DC.- EU347963, EU347894; (E.Mey.) Benth. 1- EU348103, EU347756; Boatwright et al. 184 (JRAU). L. Schutte 151 GRAU). D. pilosus Conrath ex Shinz- EU347959, EU347892; falcata (E.Mey.) Benth. 2- EU347999, EU347817; Boatwright et al. 120 Schutte 363 (JRAU). D. reflexus (N.E.Br.) A.L.Schutte- EU347961, -; Sch- °RAU). L. filiformis B.-E.van Wyk- EU348074, EU347794; Vlok 2030 utte 177 GRAU). D. strictus E.Mey.- AJ2876842; Crisp 9073 (CANB). D. (JRAU). L. foliosa Bolus 1- EU348073, EU347776; Van Wyk 2481 GRAU). strictus E.Mey.- EU347960, -; Schutte 150 (JRAU). Genista L.: G. tereti- L. foliosa Bolus 2- EU347993, EU347819; Boatwright & Magee 88 (JRAU). folia Willk.- AY2636685; MAF 162924. Lebeckia Thunb.: L. ambigua L. fruticoides B.-E.van Wyk- EU348085, EU347801; Van Wyk 2021 (JRAU).

E.Mey. 1- EU347917, EU347852; Van Wyk 2900 (JRAU). L. ambigua L. galpinii Diimmer- Z95538 3, -; T. Edwards 480. L. glabra (Thunb.) E.Mey. 2- EU347934, EU347851; Le Roux et al. 6 (JRAU). L. bowieana D.Dietr.- EU348005, EU347814; Van Wyk 2014 (JRAU). L. globulosa B.- Benth. 1- EU347909, EU347868; Streicher s.n. (JRAU). L. bowieana Benth. E.van Wyk- EU348075, EU347777; Van Wyk 2210 GRAU). L. hirsuta 2- EU347910, EU347869; Van Wyk 2106 (JRAU). L. brevicarpa M.M.Le (Thunb.) D.Dietr. 1- EU348069, EU347879; Van Wyk 1338 (JRAU). L. Roux & B.-E.van Wyk- EU347933, EU347850; Le Roux et al. 4 (JRAU). L. hirsuta (Thunb.) D.Dietr. 2- EU348070, EU347880; Van Wyk s.n. GRAU). contaminata (L.) Thunb. 1- EU347900, EU347845; Vlok et al. s.n. (JRAU). L. L. hirsuta (Thunb.) D.Dietr. 3- EU348102, EU347881; Schutte 290 (JRAU). contaminata (L.) Thunb. 2- EU347937, EU347846; Le Roux et al. 15 GRAU). L. involucrata (Berg.) Benth.- EU947997, EU347809; Boatwright et al. 116 L. contaminata (L.) Thunb. 3- -, EU347847; Le Roux et al. 16 (FRAU). L. (JRAU). L. involucrata (Berg.) Benth. subsp. digitata B.-E.van Wyk - cinerea E.Mey.- EU347930, EU347840; Boatwright et al. 150 (JRAU). L. EU348076, EU347805; Van Wyk 2873 °RAU). L. involucrata (Berg.) Benth. cytisoides Thunb. 1- EU347903, EU347837; Van Wyk 2313 (JRAU). L. cy- subsp. peduncularis (E.Mey.) B.-E.van Wyk- EU348007, EU347806; Van tisoides Thunb. 2- EU347925, EU347838; Le Roux et al. 2 (JRAU). L. gracilis Wyk et al. 4195 (JRAU). L. taxa Eckl. & Zeyh. 1- EU348059, EU347795; Van Eckl. & Zeyh.- EU347855; Le Roux et al. 17 (JRAU). L. halenbergensis Wyk 2608 (JRAU). L. laxa Eckl. & Zeyh. 2- EU348060, -; Van Wyk 1726 Merxm. & A.Schreib. 1- EU347908, EU347836; Van Wyk 3086 GRAU). L. (JRAU). L. laxa Eckl. & Zeyh. 3- -, AF287677 2; Crisp 9075 (CANB). L. halenbergensis Merxm. & A.Schreib. 2- EU347927, EU347842; Boatwright lenticula (E.Mey.) Benth.- EU348088, EU347802; Van Wyk 2017 (JRAU). L. et al. 146 (JRAU). L. inflata Bolus 1- EU347901, EU347863; Vlok et al. 2 leptoloba Bolus 1- EU348104, EU347757; Boatwright et al. 185 (JRAU). L. (JRAU). L. inflata Bolus 2- EU347940, EU347864; Johns 162 (JRAU). L. leptoloba Bolus 2- EU348077, EU347804; Schutte 276 (JRAU). L. listii Pol- leipoldtiana Schltr. ex Dahlgren 1- EU347939, EU347866; Boatwright et al. hill- EU348012, EU347826; Van Wyk et al. 4207 (JRAU). L. lotononoides 123 (JRAU). L. leipoldtiana Schltr. ex Dahlgren 2- EU347914, EU347865; (Scott Elliot) B.-E.van Wyk- EU348089, EU347822; Van Wyk 1962 (JRAU). Van Wyk 3278 GRAU). L. lotononoides Schltr.- EU347928, EU347843; Boat- L. macrosepala Conrath- EU348056, EU347810; Van Wyk 2622 (JRAU). L. wright et al. 142 (JRAU). L. macrantha Harv. 1- EU347916, EU347828; Van maculata Diimmer- EU348004, EU347813; Van Wyk et al. 4169 (JRAU). L. Wyk s.n. °RAU). L. macrantha Harv. 2- EU347922, EU347831; Boatwright magnifica B.-E.van Wyk- EU348079, EU347779; Van Wyk 2421 GRAU). L. et al. 92 (JRAU). L. melilotoides Dahlgren 1- EU347919, -; Van Wyk 2562b marlothii Engl.- EU348010, EU347825; Van Wyk et al. 4203 (JRAU). L. (JRAU). L. melilotoides Dahlgren 2- -, EU347841; Van Wyk 2229 (JRAU). maximiliani Schltr. ex De Wildeman 1- EU348078, EU347799; Schutte 282 L. meyeriana Eckl. & Zeyh. ex Harv. 1- EU347904, EU347856; Van Wyk °RAU). L. maximiliani Schltr. ex De Wildeman 2- EU348098, -; Schutte 3351 (JRAU). L. meyeriana Eckl. & Zeyh. ex Harv. 2- EU347905, -; Van 271 (JRAU). L. meyeri (Presl) B.-E.van Wyk 1- EU348095, EU347811; Van Wyk 3009 (JRAU). L. meyeriana Eckl. & Zeyh. ex Harv. 3- -, EU347857; Wyk 1766 (JRAU). L. meyeri (Pres') B.-E.van Wyk 2- EU348094, -; Van Van Wyk 4043 (JRAU). L. mucronata Benth.- EU347941, EU347870; Vlok Wyk 1765 (JRAU). L. micrantha Eckl. & Zeyh.- EU348049, EU347798; Van 1726 GRAU). L. multiflora E.Mey.- EU347926, EU347833; Boatwright et al. Wyk 2481 (JRAU). L. mo/lis (E.Mey.) Benth. 1- EU348068, EU347763; 138 (JRAU). L. pauciflora Eckl. & Zeyh. 1- EU347902, EU347861; Van Wyk Koekemoer 524 (PRE). L. mo//is (E.Mey.) Benth. 2- EU348067, EU347764; 3024 (JRAU). L. pauciflora Eckl. & Zeyh. 2- EU347935, EU347862; Le Roux Van Wyk 3113 (JRAU). L. oxyptera (E.Mey.) Benth.- EU347992, EU347787; et al. 7 (JRAU). L. plukenetiana E.Mey. 1- EU347932, EU347849; Le Roux et Boatwright s.n. (JRAU). L. parviflora (Berg.) D.Dietr.- EU348050, al. 24 (JRAU). L. plukenetiana E.Mey. 2- EU347906, EU347848; Van Wyk EU347797; Van Wyk 2442 (JRAU). L. pentaphylla (E.Mey.) Benth.- 4043 (JRAU). L. psiloloba Walp.- EU347923, EU347830; Le Roux et al. 20 EU348001, EU347783; Boatwright et al. 148 (JRAU). L. platycarpa (Viv.) (JRAU). L. pungens Thunb. 1- EU347918, EU347827; Van Wyk s.n. Pic.-Serm. 1- EU348105, EU347759; Boatwright et al. 192 (JRAU). L. platy- (JRAU). L. pungens Thunb. 2- EU347921, EU347829; Boatwright et al. 106 carpa (Viv.) Pic.-Serm. 2- EU348063, EU347766; Van Wyk 3066 (JRAU). L. (JRAU). L. sepiaria (L.) Thunb. 1- EU347936, EU347853; Le Roux et al. 10 platycarpa (Viv.) Pic.-Serm. 3- EU348064, EU347767; Koekemoer 942 (JRAU). L. sepiaria (L.) Thunb. 2- EU347920, EU347854; Van Wyk 2979 (JRAU). L. platycarpa (Viv.) Pic.-Serm. 4- EU348011, EU347768; Boat- (JRAU). L. sericea Thunb. 1- EU347907, EU347832; Van Wyk 3115 (JRAU). wright et al. 159 (JRAU). L. platycarpa (Viv.) Pic.-Serra. 5- EU348003, L. sericea Thunb. 2- EU347924, EU347834; Boatwright et al. 151 (JRAU). L. EU347769; Van Wyk et al. 4204 (JRAU). L. platycarpa (Viv.) Pic.-Serm. 6- sericea Thunb. 3- EU347915, EU347835; Van Wyk 3119 (JRAU). L. sessil- EU348065, EU347765; Friis et al. 9080 (K). L. plicata B.-E.van Wyk- ilfolia (Eckl. & Zeyh.) Benth.- EU347938, EU347867; Boatwright et al. 170 EU348072, EU347775; Van Wyk 3382 (JRAU). L. polycephala (E.Mey.) (JRAU). L. spinescens Harv.- EU347929, EU347839; Boatwright et al. 158 Benth. 1- EU348082, EU347781; Van Wyk 2408 (JRAU). L. polycephala (JRAU). L. wrightii (Harv.) Bolus 1- EU347912, EU347858; Vlok et al. s.n. (E.Mey.) Benth. 2- EU347994, EU347782; Boatwright et al. 94 (JRAU). L. (JRAU). L. wrightii (Harv.) Bolus 2- EU347913, EU347860; Vlok et al. s.n. prostrata (L.) Benth. 1- EU348054, -; Van Wyk 3229 (JRAU). L. prostrata (JRAU). L. wrightii (Harv.) Bolus 3- EU347911, EU347859; Van Wyk 3354 (L.) Benth. 2- -, EU347808; Boatwright et al. 115 (JRAU). L. pulchella (JRAU). Liparia L.: L. angustifolia (Eckl. & Zeyh.) A.L.Schutte- (E.Mey.) B.-E.van Wyk 1- EU348090, EU347785; Van Wyk 1659 (JRAU). AM177376 I, AM261478 1 ; Boatwright & Magee 66 (JRAU). L. hirsuta L. pulchella (E.Mey.) B.-E.van Wyk 2- EU348066, -; Van Wyk 1659 Thunb.- AM259357 AM261486 I ; Boatwright & Magee 33 (JRAU). (JRAU). L. pungens Eckl. & Zeyh. 1- EU348096, EU347790; Van Wyk 1725 Lotononis (DC.) Eckl. & Zeyh.: L. acuticarpa B.-E.van Wyk- EU348047, (JRAU). L. pungens Eckl. & Zeyh. 2- EU347996, EU347807; Boatwright et EU347770; Van Wyk 2625 (JRAU). L. adpressa N.E.Br. subsp. adpressa- al. 105 (JRAU). L. quinata (Thunb.) Benth.- EU347995, EU347774; Boat- EU348100, EU347780; Van Wyk 1899 (JRAU). L. alpina (Eckl. & Zeyh.) wright et al. 95 (JRAU). L. rabenaviana Dinter & Harms- -, EU347758; B.-E.van Wyk- -, AM262446 6; Van Wyk & Van Wyk 1478 GRAU). L. Boatwright et al. 191 GRAU). L. rigida (E.Mey.) Benth. 1- EU348101, -; benthamiana Diimmer- EU348051, EU347771; Van Wyk 2528 (JRAU). L. Van Wyk 2700 (JRAU). L. rigida (E.Mey.) Benth. 2- -, EU347791; Van bolusii Diimmer- EU348046, EU347761; Van Wyk 2443 GRAU). L. brachy- Wyk s.n. (JRAU). L. rostrata Benth. subsp. namaquensis (Bolus) B.-E.van antha Harms- EU347998, EU347816; Boatwright et al. 117 (JRAU). L. brevi- Wyk - EU348002, EU347812; Boatwright et al. 155 GRAU). L. rostrata caulis B.-E.van Wyk- EU348080, -; Schutte 447 (JRAU). L. calycina Benth.- EU348000, EU347818; Boatwright et al. 147 GRAU). L. sabulosa (E.Mey.) Benth.- EU348052, EU347762; Van Wyk 1433 GRAU). L. carnosa Salter- EU348081, EU347820; Van Wyk 2325 (JRAU). L. sericophylla Benth. (Eckl. & Zeyh.) Benth.- EU348083, EU347800; Van Wyk 1663 °RAU). L. 1- EU348093, EU347789; Van Wyk 1956 (JRAU). L. sericophylla Benth. 2- curtii Harms- EU348006, EU347815; Van Wyk et al. 4172 (JRAU). L. cru- EU348053, EU347803; Van Wyk 1917 (JRAU). L. sericophylla Benth. 3- manina Burch. ex Benth.- EU348062, -; Van Wyk 3057 (JRAU). L. decum- EU348097, -; Van Wyk 1958 GRAU). L. solitudinis Diimmer- EU348008, bens (Thunb.) B.-E.van Wyk subsp. decumbens- EU348058, EU347778; EU347823; Van Wyk et al. 4198 (JRAU). L. sparsiflora (E.Mey.) B.-E.van Van Wyk 1701 (JRAU). L. densa (Thunb.) Harv. subsp. leucoclada (Schltr.) Wyk- EU348099, EU347786; Van Wyk 2057 (JRAU). L. stricta (Eckl.& B.-E.van Wyk- EU348048, EU347792; Van Wyk 2430 (JRAU). L. digitata Zeyh.) B.-E.van Wyk- EU348091, EU347793; Van Wyk 1718 (JRAU). L. Harv. 1- EU348071, EU347772; Van Wyk 2350 GRAU). L. digitata Harv. subulata B.-E.van Wyk- EU348009, EU347824; Van Wyk et al. 4202 2- EU348057, EU347773; Van Wyk 2342 (JRAU). L. divaricata (Eckl. & (JRAU). L. umbellata (L.) Benth. 1- EU348061, -; Van Wyk 3234 GRAU). Zeyh.) Benth.- EU348092, EU347788; Van Wyk 2484 °RAU). L. elongata L. umbellata (L.) Benth. 2- -, EU347760; Boatwright s.n. GRAU). Lupinus

(Thunb.) D.Dietr.- EU348087, EU347821; Van Wyk 2573 (JRAU). L. eri- L.: L. polyphyllus Lindley- Z70052 3, -; Planchuelo 95. Melolobium Eckl. antha Benth.- EU348086, EU347784; Schutte 383 (JRAU). L. eriocarpa & Zeyh.: M. adenodes Eckl. & Zeyh. 1- -, AM050832 6; Van Wyk 4036 (E.Mey.) B.-E.van Wyk- EU348084, -; Van Wyk 1952 (JRAU). L. exstipu- (JRAU). M. adenodes Eckl. & Zeyh. 2- EU347976, -; Van Wyk 2159 lata Bolus- EU348055, EU347796; Van Wyk 2271 (JRAU). L. falcata PAU). M. adenodes Eckl. & Zeyh. 3- EU347986, -; Van Wyk 4036 2008] BOATWRIGHT ET AL.: CROTALARIEAE PHYLOGENY 761

(JRAU). M. aethiopicum (L.) Druce 1— EU347977, —; Strydom 5 (PRE). M. fusca Thunb. 2— EU347967, EU347754; Van Wyk et al. 4186 (JRAU). W. aethiopicum (L.) Druce 2— EU347987, —; Van Wyk 4040 (JRAU). M. alpi- humilis (Thunb.) Dahlgren 1— EU347969, EU347752; Boatwright et al. 129 num Eckl. & Zeyh.— EU347978, —; Schutte 158 URAU). M. candicans URAU). W. humilis (Thunb.) Dahlgren 2— EU347970, EU347753; Van Wyk (E.Mey.) Eckl. & Zeyh. 1— —, AM050833 6, Van Wyk 4016 (JRAU). M. 3530 (JRAU). W. incurvata Thunb.— EU347975, EU347751; Boatwright et al. candicans (E.Mey.) Eckl. & Zeyh. 2— EU347980, —; Van Wyk 3072 (JRAU). 188 (JRAU). W. monoptera E.Mey. 1— EU347971, EU347749; Boatwright et M. candicans (E.Mey.) Eckl. & Zeyh. 3— EU347991, —; Boatwright et al. 187 al. 152 (JRAU). W. monoptera E.Mey. 2— EU347974, EU347750; Boatwright (JRAU). M. canescens (E.Mey.) Benth.— EU347981, —; Schutte 303 URAU). et al. 153 (JRAU). W. obcordata (Berg.) Thunb. 1— EU347965, EU347746; M. exudans Harv.— EU347990, —; Van Wyk 2234 URAU). M. humile Eckl. Van Wyk 2686 (JRAU). W. obcordata (Berg.) Thunb. 2— EU347972, & Zeyh.— EU347982, —; (JRAU). M. Diimmer- Van Wyk 2543 macrocalyx EU347748; Boatwright et al. 98 URAU). W. sericea Thunb.— EU347968, EU347989, —; Van Wyk 3061b. M. microphyllum (L.f.) Eckl. & Zeyh. 1— EU347755; Boatwright et al. 124 (JRAU). W. tetraptera E.Mey. 1— EU347983, —; Van Wyk 4006 (JRAU). M. microphyllum (L.f.) Eckl. & Zeyh. EU347964, EU347745; Schutte 737 (JRAU). W. tetraptera E.Mey. 2— EU347979, —; Moteetee & Van Wyk 3 (JRAU). M. microphyllum (L.f.) EU347973, EU347747; Boatwright et al. 104 URAU). Xiphotheca Eckl. & Eckl. & Zeyh. 3— Z955393, —; Edwards 470. M. obcordatum Harv.- Z955403, —; Edwards 470. M. stipulatum (Thunb.) Harv.— EU347988, —; Zeyh.: X. guthriei (Bolus) A.L.Schutte & B.-E.van Wyk— AM260744 Van Wyk 4037 (JRAU). M. subspicatum Conrath— EU347984, —; Van Wyk AM261741 Vlok & Schutte 4 (SCHG). X. reflexa (Thunb.) A.L.Schutte & 1779 (JRAU). M. wilmsii Harms— EU347985, —; Van Wyk 2724 (JRAU). B.-E.van Wyk— AM260747 1, AM261744 Schutte 760 (JRAU). Pearsonia Diimmer: P. aristata (Shin) Dlimmer— EU347945, EU347873; De Castro 346 (JRAU). P. cajanifolia (Harv.) Polhill 1— EU347948, APPENDIX 2. List of morphological characters and character states used EU347875; (JRAU). (Harv.) Polhill 2— EU347950, Posthumus 7 P. cajanifolia in the morphological analysis. EU347876; Boatwright & Magee 90 (JRAU). P. grandifolia (Bolus) Polhill- EU347946, EU347874; Van Wyk 3047 (JRAU). P. sessilifolia (Harv.) Dilm- 1. Habit— herbs or suffrutices = 0; shrubs = 1. 2. Persistence— peren- mer 1— EU347947, EU347871; Schutte 463 (JRAU). P. sessilifolia (Harv.) nial = 0; annual = 1. 3. Young twigs— without bark formation (bark Diimmer 2— —, EU347872; Schutte 589b. P. sessilifolia (Harv.) Diimmer formation late or absent) = 0; with bark formation (bark formation —, AJ2876752; Crisp 9078 (CANB). P. uniflora (Kensil) Polhill- early) = 1. 4. Leaves— digitate = 0; unifoliolate = 1; simple or phyllodi- EU347949, —; Van Wyk 3033 (JRAU). Podalyria Willd.: P. buxifolia (Retz.) nous = 2. 5. Lamina— flat = 0; terete (acicular) = 1. 6. Petiole presence— Lam.— AM261693 1, AM261496 1; Boatwright & Magee 34 (JRAU). P. ole- present = 0; absent (leaves sessile) = 1. 7. Petiole base— normal or leaves aefolia Salisb.— AM261708 AM261667 1; Boatwright & Magee 79 URAU). sessile = 0; tuberculate or with persistent spur = 1. 8. Stipules presence— Polhillia C.H.Stirton: P. brevicalyx (C.H.Stirton) B.-E.van Wyk & A.L.Sch- present = 0; vestigial or absent = 1. 9. Stipules symmetry— symmetrical utte— EU347954, —; Van Wyk 2100 (JRAU). P. canescens C.H.Stirton- or absent = 0; asymmetrical (dissimilar in size or single) = 1. 10. Bracteole EU347957, —; Van Wyk 2092 URAU). P. involucratum (Thunb.) B.-E.van presence— present = 0; vestigial or absent = 1. 11. Standard petal vesti- Wyk & A.L.Schutte— EU347956, —; Schutte 339 URAU). P. obsoleta ture— hairy at least along the dorsal midrib = 0; totally glabrous = 1. 12. (Thunb.) B.-E.van Wyk— EU347955, —; Van Wyk 2701 URAU). P. pallens Keel shape— not rostrate = 0; markedly rostrate or helically coiled = 1. C.H.Stirton— EU347958, —; Van Wyk 2095 URAU). Rafnia Thunb.: R. 13. Calyx lower lobes— with trifid lower lip = 0; without trifid lower acuminata (E.Mey.) G.J.Campbell & B.-E.van Wyk— AM931036, ip = 1. 14. Calyx lateral lobes— not fused higher up = 0; fused higher AM931034 & AM931035; Schutte 437 (JRAU). R. angulata Thunb.— up = 1. 15. Anthers— dimorphic = 0; uniform = 1. 16. Carinal anther— EU347896, —; Boatwright et al. 103 URAU). R. capensis (L.) Schinz- EU347742; Boatwright & Magee 26 (JRAU). R. diffusa Thunb.— EU347895, resembling basifixed anthers = 0; intermediate = 1; resembling dorsifixed AJ744944 6; Campbell & Van Wyk 44 URAU). R. globosa G.J.Campbell & anthers = 2. 17. Gynoecium base— sessile or subsessile = 0; stipitate = 1. B.-E.van Wyk — EU347899, EU347743; Boatwright et al. 180 (JRAU). R. 18. Style— upcurved = 0; straight = 1; helically coiled = 2. 19. Style racemosa Eckl. & Zeyh.— EU347895, EU347741; Boatwright & Magee 29 vestiture— glabrous = 0; hairy = 1. 20. Fruit type— not a samara = 0; fruit URAU). R. spicata Thunb.— EU347897, EU347740; Boatwright et al. 102 an ovoid, winged samara = 1. 21 Fruit upper suture— upper suture (JRAU). Robynsiophyton R. Wilczek: R. vanderystii R. Wilczek- straight or symmetrically convex = 0; upper suture asymmetrically con- EU347952, EU347878; Lisowski 20326 (K). Rothia Pers.: R. hirsuta (Guill. & vex = 1. 22. Fruit— many-seeded = 0; few-seeded (1-2) = 1. 23. Funicle Perr.) Baker— EU347953, EU347877; Saadou 1798 (K). Spartidium Pomel: length— normal length = 0; exceptionally long = 1. 24. Seed shape— S. saharae (Coss. & Dur.) Pomel— EU347931, EU347844; Davis 49544 (K). transversely oblong = 0; oblong = 1. 25. Seed surface— smooth = 0; Stirtonanthus B.-E.van Wyk & A.L.Schutte: S. chrysanthus (Adamson) rugose = 1. 26. Chromosome base number-7 = 0; 8 = 1; 9 = 2. 27. B.-E.van Wyk & A.L.Schutte— AM259367 AM268386 1 & AM268387 Cyanogenesis— absent = 0; present = 1. 28. Quinolizidine and piperidyl Van Wyk & Schutte 3297 (JRAU). Ulex L.: U. europaeus L.— Z70111 3, —; alkaloids (Lysine pathway)— present = 0; absent = 1. 29. Sparteine- Botanical Garden Heidelberg, Germany. Virgilia Poir.: V. divaricata Adam- present = 0; absent = 1. 30. Lupanine type esters— ±absent = 0; present son— AM260737 AJ409910 7; Van Wyk 879-888 (JRAU). Wiborgia as major alkaloid = 1. 31. Macrocyclic pyrrolizidine alkaloids— absent = Thunb.: W. fusca Thunb. 1— EU347966, EU347744; Schutte 736 URAU). W. 0; present = 1. Appendix B4. Boatwright, J. S., P. M. Tilney and B. - E. van Wyk. 2008. A taxonomic revision of the genus Rothia (Crotalarieae, Fabaceae). Australian

Systematic Botany 21(6): 422-430. CSIRO PUBLISHING www.publish.csiro.aujournals/asb Australian Systematic Botany, 21, 422-430

A taxonomic revision of the genus Rothia (Crotalarieae, Fabaceae)

J. S. Boatwright", P. M. Tilney A and B.-E. van WykA

A Department of Botany and Plant Biotechnology, University of Johannesburg, PO Box 524, Auckland Park 2006, Johannesburg, South Africa. B Corresponding author. Email: [email protected]

Abstract. Rothia Pers. is a genus of papilionoid legumes that consists of two species, R. indica (L.) Druce and R. hirsuta (Guill. & Pen.) Baker. The genus is a member of the tribe Crotalarieae and is widely distributed in Africa, Asia and Australia. Recent molecular systematic studies have shown the genus to be closely related to Robynsiophyton Wilczek and Pearsonia Diimmer; however, it is easily distinguished from these by its 10 small, rounded anthers and subequally lobed calyx (as opposed to the large, elongate anthers of Pearsonia or the 9 stamens of Robynsiophyton). These three genera share characters such as uniform anthers, straight styles and the presence of angelate esters oflupanine-type alkaloids. Leaf and fruit anatomy of Rothia and Robynsiophyton were also studied but revealed no informative differences. Both genera have dorsiventral leaves with mucilage cells in the epidermis and thin-walled fruits. A taxonomic revision of Rothia is presented, including a key to the species, correct nomenclature, descriptions, illustrations and distribution maps.

Introduction supported by three synapomorphies, i.e. uniform anthers, straight or down-curved styles and the presence of angelate esters of The genus Rothia Pers. was described in 1806 and comprises lupanine-type alkaloids. Rothia can be distinguished from two species, R. hirsuta (Guill. & Pen.) Baker and R. indica (L.) and by its 10 small, rounded Druce. The species have been treated in several floristic studies Pearsonia Robynsiophyton (e.g. The Flora of Tropical Africa (Baker 1926), Flora of West anthers and subequally lobed calyx. In this paper we present a taxonomic account of Rothia, with Tropical Africa (Hepper 1958), Flora of Tropical East Africa illustrations, descriptions and distribution maps of the species. (Milne-Redhead 1971), Flora • Zambesiaca (Polhill 2003), The results of anatomical studies of the leaves and fruits ofRothia A revised Handbook of the Flora of Ceylon (Rudd 1991)); and are also presented. however, no comprehensive revision of the genus exists. An Robynsiophyton Australian-endemic subspecies ofR. indica, namely R. indica (L.) Druce subsp. australis A.E.Holland, was described by Holland Materials and methods (1997). It is morphologically distinct from the typical Asian Morphology subspecies. Morphological characters were assessed through the study of The genus is a member of the Papilionoideae and is placed material from the following herbaria: BM, BOL, K, MEL, NBG within the tribe Crotalarieae. The Crotalarieae form part of a (including SAM), PRE, S, UPS (abbreviations according to monophyletic assemblage of tribes, the 'core' genistoid legumes Holmgren et al. 1990). Drawings of reproductive features (all (Crisp et al. 2000; Boatwright et al. 2008a), which comprises by JSB) were done by using a stereoscope (WILD M3Z) with a the Crotalarieae, Euchresteae, Genisteae, Podalyrieae, camera lucida attachment. Sophoreae (in part) and Thermopsideae. The Crotalarieae are subendemic to Africa, with a few species of Crotalaria L., Lotononis (DC.) Eckl. & Zeyh. and Rothia extending to other Anatomy continents. A recent study of the tribe by Boatwright et al. Two or three samples per species of the leaves and fruits ofRothia (2008b), based on gene sequences (ITS and rbcL) and and Robynsiophyton were studied. Dried material was first morphological data, showed Rothia to be in a well supported rehydrated and then fixed in formaldehyde : acetic acid : 96% Glade along with Pearsonia Diimmer and Robynsiophyton alcohol : water (10 : 5 : 50 : 35; FAA) for 24 h. The method of Wilczek, i.e. the Pearsonia Glade. A sister relationship Feder and O'Brien (1968) for embedding in glycol methacrylate between Robynsiophyton and Rothia was strongly supported (GMA) was used except that the final infiltration in GMA was (Fig. 1). This confirmed hypotheses by Polhill (1976), Van done for a minimum of 5 days. Sections were stained according to Wyk (1991a) and Van Wyk and Schutte (1995) who the periodic acid Schiff/toluidine blue (PAS/TB) staining method suggested an affinity between these genera. The close and mounted. Photographs were taken with a JVC KY-F1030 relationship between Pearsonia, Robynsiophyton and Rothia is digital camera. CSIRO 23 December 2008 10.1071/SB08033 1030-1887/08/060422 Revision of the genus Rothia Australian Systematic Botany 423

100 Pearsonia sessilifolia R. indica subsp. australis has densely pubescent leaves with long whitish hairs that are more or less spreading. In contrast, the leaves 99 Pearsonia aristata of R. indica subsp. indica are moderately hairy with shorter, Pearsonia grandifolia 100 more or less appressed hairs. The stipules of R. hirsuta are linear Pearsonia cajanifolia to slightly falcate and single at each node, whereas those of 9997 Rothia hirsuta R. indica are elliptic to lanceolate or ovate and invariably paired. Robynsiophyton vanderystii Anatomical studies of the lamina (Fig. 2a, b) and petiole (Fig. 2c, d) revealed no informative differences between Fig. 1. The Pearsonia Glade taken from a strict consensus of 370 trees, R. hirsuta and R. indica, or even between these and showing the relationship between Pearsonia, Robynsiophyton and Rothia, Robynsiophyton vanderystii Wilczek. The leaves are based on the combined analysis of molecular (ITS and rbcL) and dorsiventral with mucilage cells present in the epidermis in morphological data (from Boatwright et al. 2008b; tree length = 1166; both Rothia and Robynsiophyton (Fig. 2a, b). These cells are consistency index = 0.53; retention index = 0.84). Numbers above the branches are bootstrap percentages above 50%. thought to contribute to the retention of water and reduction of transpiration, although their function has not been accurately ascertained (Gregory and Baas 1989). Results and discussion Generic relationships Reproductive morphology and anatomy In the study of Boatwright et al. (2008b), the Pearsonia Glade is well separated from the 'Cape' group of the Crotalarieae Both species of Rothia have axillary racemes. R. hirsuta has up to (Aspalathus L., Lebeckia Thunb., Rafnia Thunb., Spartidium seven flowers per raceme as opposed to R. indica where up to four Pomel and Wiborgia Thunb.) and placed closer to Lotononis, a flowers per raceme are found, although the flowers are often genus with which they share the lotononoid or zygomorphic solitary or in pairs. Terminal racemes are found in most genera of calyx. However, the three genera of the Pearsonia Glade possess the Crotalarieae (Polhill 1976) and they are either terminal or a 17 base-pair deletion at positions 179-196 in the aligned axillary in Pearsonia and strictly axillary in Rothia and ITS matrix and unique floral characters such as straight styles Robynsiophyton. Bracts are present in Rothia, but bracteoles and uniform anthers. The Crotalaria Glade (Bolusia Benth., are lacking. Bracteoles are also absent in Robynsiophyton and Crotalaria and Lotononis section Euchlora (Eckl. & Zeyh.) most species of the genus Lotononis, but present in almost all B.-E.van Wyk) is early diverging within the Crotalarieae and Pearsonia species. all members of this Glade have strongly inflated pods (Boatwright The flowers of Rothia and its relatives are relatively et al. 2008b). unspecialised compared with the rest of the Crotalarieae. Van Wyk (2003) discussed the importance of chemical Rothia, Robynsiophyton and Pearsonia have a 'gullet' -type characters in genistoid legumes and mentioned that their flower where the style is straight or down-curved and the distribution is not random, but in fact provides reliable anthers uniform and/or further reduced, as opposed to the information supporting phylogenetic relationships retrieved by more specialised and reflexed floral parts found in the other studies ofDNA sequence data. The Pearsonia Glade is chemically crotalarioid genera (Polhill 1976). In Rothia the two upper distinct within the tribe Crotalarieae in having unique alkaloids calyx lobes are larger than the other three lobes and distinctly (Van Wyk and Verdoorn 1990; Van Wyk 1991a, 2005; Van Wyk falcate. The calyces of Pearsonia species have the upper sinus and Schutte 1995). Pearsonia, Robynsiophyton and Rothia often shallower than the others or in some species the lateral sinus produce tetracyclic quinolizidine alkaloids such as lupanine, is the shallowest with the carinal lobe always narrower than the together with angelate esters such as lupanine-13a-angelate, other lobes (Polhill 1974). Robynsiophyton differs from both cajanifoline, sessilifoline and pearsonine (Hussain et a/. 1988; these genera in having an equally lobed calyx (J. S. Boatwright Van Wyk and Verdoorn 1989, 1991). These chemical characters and B.-E. van Wyk, unpubl. data). are especially useful in distinguishing this Glade from the The most useful generic character to distinguish among morphologically similar genus Lotononis where pyrrolizidine Pearsonia, Robynsiophyton and Rothia is the androecium. and quinolizidine alkaloids are present but where no esters of These genera are unique within the Crotalarieae in having lupanine-type alkaloids are found (Van Wyk 1991b). Also, some uniform anthers, as opposed to the dimorphic anthers found in sections of Lotononis are cyanogenic, a trait which is unique other genera of the tribe with alternating basifixed and dorsifixed within Crotalarieae (Van Wyk 1991b). anthers (Polhill 1976). The anthers of Pearsonia are elongate and large, whereas in Rothia and Robynsiophyton they are small and rounded (Fig. 3). Robynsiophyton has a reduced number of Vegetative morphology and anatomy stamens (9 rather than 10) and four anthers are infertile The species of Rothia are prostrate to procumbent or erect (Fig. 3). Although the generic concepts of Rothia and annuals. They share this life history with the monotypic Robynsiophyton have been questioned by previous authors, the African Robynsiophyton. This character lends support to the informative value of androecial characters (the main distinction sister relationship found between these genera by Boatwright among these three genera) at both tribal and generic level is et al. (2008b) and distinguishes them from the closely related usually closely correlated with relationships suggested by genus Pearsonia which consists of perennial herbs or shrubs. both molecular and morphological data (Polhill 1976; The leaves of Rothia are digitately trifoliolate as in many of Boatwright et a/. 2008b) and should be taken at face value as the Crotalarieae, and pubescent both adaxially and abaxially. strong, sound generic apomorphies. 424 Australian Systematic Botany J. S. Boatwright et al.

(d)

Fig. 2. Transverse sections through the leaves and fruit ofRothia and Robynsiophyton. (a) Portion of leaf of Robynsiophyton vanderystii (scale bar = 0.1 mm). (b) Portion of leaf of Rothia indica (scale bar =0.1 mm). (c) Petiole of Robynsiophyton vanderystii (scale bar= 0.2 mm). (a') Petiole of Rothia indica (scale bar =0.2 mm). (e) Portion of fruit wall of Robynsiophyton vanderystii (scale bar= 0.07 mm). (1) Portion of fruit wall of Rothia indica (scale bar =0.07mm). Voucher specimens. (a, e) Lisowski 20326 (K). (b, d, f) Latz 16126 (MEL). (c) Exell & Mendonca 657 (K).

The pods of Rothia hirsuta are ovate to falcate and much Rothia Pers., Syn. Pl. 2: 302, 638 (1807). nom. conserv.; shorter and fewer-seeded (up to 18 mm long with ±25 seeds) than Benth. in Hook., Lond. J. Bot. 3: 338 (1844); Benth. & the linear pods of R. indica that are up to 55 mm long with ±35 Fl. Austral. 2: 185 (1864); Baker, Fl. Trop. Africa: 7 (1871); seeds. The pods of Rothia and Robynsiophyton are thin-walled Trimen, Handb. Fl. Ceylon: 7 (1894); Thonner, Flowering and sclerified on the inner surface, with no apparent anatomical Plants of Africa: 274 (1915); Baker, Leg. Trop. Africa: differences (Fig. 2e, f). 21 (1926); Hutchinson & Dalziel, Fl. West Trop. Africa: Rothia indica and Robynsiophyton have similar-sized seeds 543 (1954); Hutchinson, The Genera of Flowering Plants: that are larger than those of Rothia hirsuta (J. S. Boatwright and 361 (1964); D 'Orey & Liberato, Fl. Guine Portuguesa: B.-E. van Wyk, unpubl. data). The seeds of the latter are light 71 (1971); Polhill in Bot. Syst. /: 326 (1976); Thulin in Op. brown with dark mottling and have a rugose surface, in contrast to Bot. 68: 153 (1983); Hedberg & Edwards, Fl. Ethiopia: the brown, smooth seeds of Rothia indica and Robynsiophyton 195 (1989); Rudd, A Revised Handbook to the Flora vanderystii. of Ceylon: 184 (1991); Van Wyk & Schutte, Advances in Legume Systematics 7, M. D. Crisp & J. J. Doyle (Eds): 306 (1995); Leistner, Seed Plants of Southern Africa: Families and Taxonomic treatment Genera: 295 (2000); Du Puy et al., The Leguminosae of Key to the genus Rothia Madagascar: 671 (2002); Polhill, Fl. Zambesiaca: 64 (2003); Leistner, Seed Plants of Southern Tropical Africa: I. Anthers dimorphic, styles upcurved other Crotalarieae Families and Genera: 202 (2005); Van Wyk in Legumes Anthers uniform, styles straight or down-curved 2 of the World, Lewis et al. (Eds), 281 (2005) Perennial herbs or small shrubs, anthers large and elongate ....Pearsonia Annual herbs, anthers small and rounded 3 Type species: R. trifoliata (Roth) Pers. (= Dillwynia trifoliata Upper two calyx lobes larger than others (calyx subequally lobed), Roth). stamens 10, all fertile Rothia Dillwynia Roth, Catal. Bot. 3: 71 (1806) non Dillwynia Sm. Upper two calyx lobes similar to others (calyx equally lobed), stamens in Koenig & Sims, Ann. Bot. 1: 510 (1805). Type species: nine, five fertile, four reduced and infertile Robynsiophyton D. trifoliata Roth. Revision of the genus Rothia Australian Systematic Botany 425

Diagnostic characters: Rothia can be distinguished from Robynsiophyton in that the latter has an equally lobed calyx (in Rothia the upper two lobes are larger than the others) and in that there are 10 stamens with 10 fertile anthers in Rothia (in Robynsiophyton the stamens are reduced to nine and only five e3 go anthers are fertile). Furthermore, it differs from Pearsonia by its subequally lobed calyx and small, rounded anthers in contrast to the lotononoid' calyx (zygomorphic owing to the fusion of the upper and lateral lobes on either side) and large, elongate anthers of Pearsonia. Chromosome number: a count of n= 7 has been reported for Rothia indica (Goldblatt 1981; Bairiganjan and Patnaik 1989). Distribution: the genus occurs in tropical Africa, Asia and Australia (Figs 5 and 7). It appears to have been well collected in areas for which regional floras have been compiled, e.g. the Flora of Tropical East Africa and Flora of West Tropical Africa. Fig. 3. (a, el) Androecia and anthers of Pearsonia, (b, e2) Robynsiophyton Voucher specimens: (a, el) Swanepoel 1 (JRAU), (b, e2) and (c, e3) Rothia. Key to the species of Rothia McCallum Weston 717 (K), (c, e3) Greenway & Kanuri 14269 (K). All scale bars represent 1 mm. I. Stipules linear and single at each node, up to 7 flowers per inflorescence, pods ovate to falcate and up to 18 mm long, seeds light brown with dark mottling and a rugose surface, Africa and Madagascar.... I. R. hirsuta Westonia Spreng., Syst. Veg. 3: 152, 230 (1826) nom. illeg. Type Stipules elliptic to lanceolate or ovate and paired at each node, up to species: W. humifusa (Willd.) Spreng. 4 flowers per inflorescence, pods linear or slightly falcate and up to 55 mm long, seeds brown with a smooth surface, Asia and Goetzea Reichb., Consp. Regn. Veg.: 150 (1828) nom. rej. Australia 2. R. indica Xerocarpus Guill. & Perr., Fl. Seneg. Tent.: 169 (1832). Type species: X hirsutus Guilt. & Pen. Fl. Trop. Africa 2: Harpelema J.Jacq., Ecl. Pl. Rar. 2: t. 129 (1844). Type species: 1. Rothia hirsuta (Guilt. & Perr.) Baker in H. dillwynia J.Jacq. 7 (1871); Eyles in Trans. Roy. Soc. South Africa 5: 369 (1916); E.G.Baker, Leg. Trop. Africa: 21 (1926); Broun & Massey, Prostrate, annual herbs. Branches up to 40 cm long, densely to Fl. Sudan: 176 (1929); Andrews, The Flowering Plants of the sparsely pubescent. Leaves digitately trifoliolate, elliptic to Anglo-Egyptian Sudan: 230 (1952); Hepper in Fl. West Trop. oblanceolate or obovate, pubescent; leaflets subsessile, apex Africa, ed. 2, 1: 543 (1958); Torre, Consp. Fl. Angolensis 3: apiculate, obtuse or slightly acute, base cuneate; petioles 5 (1962); White, Forest Fl. Northern Rhodesia: 164 (1962); shorter than leaflets, pubescent. Stipules linear to slightly Schreiber in Merxmiiller, Prodr. Fl. SW. Africa, fam. 60: 107 falcate or elliptic to lanceolate or ovate, single in R. hirsuta, (1970); Milne-Redhead, Fl. Trop. East Africa, Leguminosae, paired in R. indica, pubescent. Inflorescence axillary, racemose, Pap.: 811 (1971); Drummond in Kirkia 8: 226 (1972); Thulin flowers solitary or up to 7 per raceme; pedicel up to 2 mm long; in Op. Bot. 68: 153 (1983); Hedberg & Edwards, Fl. Ethiopia: bract linear, pubescent, caducous; bracteoles absent. Flowers pale 195 (1989); Du Puy et al., The Leguminosae of Madagascar: yellow to white. Calyx pubescent, subequally lobed; lobes 671 (2002) triangular to lanceolate, upper lobes wider than others, falcate, carinal lobe slightly narrower than others, tips minutely pubescent Xerocarpus hirsutus Guill. & Perr. in Guillemin, Perrottet & Richard, 44 (1832). Type: Senegal, Leprieur s.n. lecto (here inside. Standard elliptic to ovate, pilose along dorsal midrib; claw Fl. Seneg. Tent.: designated) P (photo seen). linear; apex obtuse. Wing petals oblong to obovate, glabrous, with 2 or 3 rows of sculpturing, as long as keel; apex obtuse. Keel boat- [Note: according to Stafleu and Cowan (1983) the type material of shaped, glabrous, pocket sometimes present; apex obtuse. Perrottet is housed in P or G. No specimens were traced in G and we therefore designate the Paris specimen as lectotype.] Stamens 10, anthers uniform, sub-basifixed. Pistil subsessile, pubescent; ovary linear to narrowly ovate; style straight, Amphinomia desertorum (Diimmer) Schreiber in Mitt. Bot. Munchen glabrous. Pods linear to ovate or falcate, laterally compressed, 2: 286 (1957); Lotononis desertorum Diimmer in Trans. Roy. Soc. subsessile, many-seeded, pubescent, dehiscent. Seeds oblique- S. Africa 3: 316 (1913). Type: Great Barmen, Buschsteppe, 23. cordiform, brown or brown mottled with dark brown, surface iv.1907, Dinter 518; lecto (here designated) Z (photo seen), isolecto BM!, BOL!, NBG!, K! smooth or rugose. [Note: Dimmer was based at K during his revision of Lotononis. Etymology: the genus is named in honour of the However, the K isotype is a small piece taken from the specimen in Z German physician and botanist Albrecht Wilhelm Roth (as annotated by Diimmer himself). The Z specimen is therefore chosen (1757-1834). as lectotype.] 426 Australian Systematic Botany J. S. Boatwright et al.

(i)

(c)

Fig. 4. Morphology ofRothia hirsuta. (a) Standard petal. (b) Seed in lateral view. (c) Outer surface of the calyx (upper lobes to the left). (d) Wing petal. (e) Keel petal. (f )Pistil. (g) Flower in lateral view. (h) Pod in lateral view. (i) Bract. (j) Anther. (k) Androecium. (I) Leaf in abaxial view. (m) Stipule. Voucher specimens. (a, c) Davey 173 (K). (b) Milne-Redhead & Taylor 11245 (K). (d,g, h,j, k) Greenway & Kanuri 14269 (K). (e)Jackson & Apejoye 413973 (K). (f )Hepper 1021 (K). (i) Polhill & Paulo 2089 (K). (I, m) Siame 162 (K). All scale bars represent 1 mm.

Small, pubescent, prostrate or semi-erect to procumbent herb. Distribution and ecology: widespread in the dry parts of Branches ±10-40 cm long, brown, sparsely hairy (pilose). tropical Africa from South Africa north to Eritrea and west up Leaves with leaflets elliptic to oblanceolate, terminal leaflet to Guinea (Fig. 5). Occurs on sandy loam or clay soils, often in 10-30 mm long, 3.5-12.0 mm wide, lateral leaflets 6-20 mm grassland or forest openings. Common on sandy soils along long, 2-7 mm wide; apex apiculate; base cuneate; petiole watercourses or disturbed roadsides. 3-13 mm long. Stipules linear to slightly falcate, 2-8 mm long, the linear stipules are invariably single single. Inflorescence with 2-7 flowers; pedicel up to 1 mm long; Diagnostic characters: whereas they are paired and elliptic bract 1.0-2.5 mm long. Flowers 4-8 mm long. Calyx 5-7 mm at each node in Rothia hirsuta, The inflorescences are more densely long; tube 1.5-2.0 mm long; lobes 3-4 mm long. Standard to ovate in R. indica. R. hirsuta than in R. indica 4.0-5.5 nun long; claw 1.0-1.5 mm long; lamina 3.0-4.5 mm flowered (up to 7 flowers) in long, 1-2 mm wide. Wings 4-5 nun long; claw 1.5-2.0 nun where the flowers are often solitary or up to 4 per inflorescence. The linear pods of R. indica are three times long; lamina 2.5-4.0 mm long, 0.5-0.7 nun wide. Keel R. hirsuta that are 2.5-4.0 mm long; claw 1.5-2.0 mm long; lamina 1-3 mm long, longer (up to 55 mm) than the ovate pods of the seeds are brown with dark 0.5-1.0 mm wide. Pistil with ovary narrowly ovate, 2-3 mm long, only up to 18 mm long. In R. hirsuta indica are 0.5-0.7 ram wide with ±7-13 ovules; style 1.3-1.6 mm mottling and have a rugose surface, whereas those ofR. long. Pods ovate, falcate, tapering towards apex, 10-18 mm brown (unmottled) with a smooth surface. long, 2-3 mm wide with up to ±25 seeds per pod. Seeds Flowering period: R. hirsuta flowers mainly from February to 1.1-1.5 mm long, 0.8-1.1 mm wide, mature seeds light brown November, but flowering and fruiting specimens have been with dark mottling, surface rugose (Fig. 4). recorded throughout the year.

Revision of the genus Rothia Australian Systematic Botany 427

Bot. Exch. Club Soc. Brit. Isles 3: 20 10 20 60 2. Rothia indica (L.) Druce in 423 (1914); Rudd, A Revised Handbook to the Flora of Ceylon: 184 (1991); Holland, Austrobaileya 5: 93 11119.1111111111111111 (1997); Trigonella indica L., Sp. Pl. 2: 778 (1753) Type: Herb. Hermann 3: 24 no. 285 BM-000621888, lecto 1111M111111111119111111L (designated by Rudd 1991) BM (photo seen). Lotus indicus Desr. in Lam., Encyc. 3: 606 (1759). Type: as for 40 111111121:4111111111111117 4 sisomi1100111111111111 T. indica. Hosackia indica Graham in Wallich, Numer. List n. 5940 30 sramiramentiffen 30 (1831-1832). nom. nud. 111111111111MMIBM Glycine leptocarpa Graham in Wallich, Numer. List n. 5515 (1831-1832). nom. nud. 0 11111111MENIMIllb 20 MI ■ Rothia trifoliata (Roth) Pers., Syn. Pl. 2: 638 & 659 (1807); Benth. in niumorniorr Hook., Lond. J. Bot. 3: 339 (1844); Bentham & Muller, Fl. Austr. 2: 185 111111111M11111111MINEM (1864); Trimen, Handb. Fl. Ceylon: 7 (1894); Dillwynia trifoliata Roth, Catal. Bot. 3: 71 (1806). Type: 'horto medico Amsteledamensi' (Bt?). 11111111111/1111111.11M1 Small, pubescent, prostrate, herb. Branches up to 40 cm long, MIONNEWIIME brown, either covered with dense, woolly hairs or sparsely hairy (pilose). Leaves with leaflets elliptic to obovate, terminal 111111111MMOMMIN leaflet (7-)9-26 mm long, (2-)3-5 mm wide, lateral leaflets 1111111•1111111211111EN (5-)8-20 mm long, (1.5-)2.5-6.0 mm wide; apex obtuse or base cuneate; petiole 4-13 mm long. Stipules 20 INLEIL0110110111- 0 slightly acute; elliptic to lanceolate or ovate, 2-8(-10) mm long, paired. 1111111111LIMEMEM Inflorescence with flowers solitary or up to 4 per raceme; 30 pedicel up to 2 mm long; bract 1-3 mm long. Flowers 5-8 mm IP! long. Calyx 4-7 mm long; tube 1.5-4.0 mm long; lobes 1-4 mm long. Standard 4.0-6.5 mm long; claw 1.5-2.5 mm long; lamina N81 1991 10 20 30 4 50 6 2.5-4.0 mm long, 2.0-2.5 mm wide. Wings 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 min Fig. 5. Known geographical distribution of Rothia hirsuta. wide. Keel 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Pistil with ovary linear,

Selected specimens (150 specimens examined) 3.0-4.5 mm long, 0.5-1.0 mm wide with ±13 - 18 ovules; style ERITREA: Bocos, Cheren, Pappi 2527, 10.xi.1902 (BM 2 sheets, K). 1.4-1.7 mm long. Pods linear or slightly falcate, (30 —)35 - 55 mm SUDAN: Kordofan, Kotsch 420, without date (K). CHAD: Dioura, long, 1.5-3.0 mm wide with up to ±35 seeds per pod. Seeds Macina, Davey 173, Oct. 1954 (K). NIGER: Niamey, Olufsen 476, 1.3-1.8 mm long, 1.0-1.3 mm wide, mature seeds brown, surface 8.x.1947 (BM, S). GUINEA: Guine-Bissau, Pussuli, Santos 1438, smooth (Fig. 6). 17.xii.1942 (K). BURKINA FASO: Haute Volta, Lergo, Ake Assi 13587, 13.xi.1976 (K). GHANA: Tefle, on Volta River, Akpabla Distribution and ecology: occurs in India, Sri Lanka, China, 1982, 13.8.1959 (K). NIGERIA: Naraguta, Jos Plateau, rough ground Vietnam, Malaysia and Australia (Fig. 7). near Forestry Department, among grass, Hepper 1021, 14.x.1957 (K). Diagnostic characters: the paired, ovate stipules, sparse CAMEROON: Ganglani village, 6 miles [9.65 km] from confluence of inflorescences, long, linear pods and smooth, brown seeds Kam and Kimiri Rivers, Vogel Peak area, Hepper 1441, 23.xi.1957 (K). distinguish Rothia indica from R. hirsuta. DEMOCRATIC REPUBLIC OF CONGO: 50 km NE of Mokolo, on route to Mora, Bounougou 173, 18.ix.1964 (K, S). UGANDA: Matuga, Flowering period: flowering and fruiting all year round from Robertson 3312, 29.vii.1982 (K). KENYA: Thika-Garissa road, 2.6 km January to December. towards Garissa after junction with Kitui Rd, near Kongonde, Faden & two subspecies can be recognised based predominantly Faden 74/737, 7.vi.1974 (K, PRE). TANZANIA: Imagi Hill, 1 mile Notes: [1.61 km] S of Dodoma, Polhill & Paulo 2089, 20.iv.1962 (K, PRE). on their geographical distribution, size and pubescence of ZAMBIA: Livingstone, Chief Mukuni area, 2.1 km along road to Chief vegetative and reproductive parts. Mukuni Palace from junction with Victoria Falls Rd in Baikiae forest, Zimba et al. 913, 19.ii.1997 (K). MALAWI: Lilongwe, Banda 250, 1. Vegetative parts sparsely hairy, terminal leaflets (7-)10-26 x (2-) 14.ix.1956 (BM). MOZAMBIQUE: Niassa, Mossuril para Lumbo, 4-8 mm, lateral leaflets (5-)8-20 x (1.5-)3.0-6.0 mm, petiole 4.5 km na estrada de Nampula, Pedro & Pedrogao 3131, 5.v.1948 5-13mm long, stipules elliptic to lanceolate 2-8(-I 0) mm long, pods (K, PRE). ANGOLA: Huilla, Welwitsch 1904, Apr. 1860 (K, BM). (-33)35-55 x (1.5-)2.0-3.0 mm, seeds 1.3-1.6 x 1.0-1.3 mm, India, NAMIBIA: grey sandy flats below Musese Camp, W of Lupala Mission Sri Lanka, China, Vietnam and Malaysia 2a. subsp. indica Station, De Winter & Marais 4978, 4.iii.1969 (K). ZIMBABWE: Vegetative parts covered with dense, woolly hairs, terminal leaflets Gwanda, Sezane Reservoirs, Davies 1296, May 1955 (K, PRE). 9-15 x 3-5 mm, lateral leaflets 9-12 x 2.5-4.0 mm, petiole 4-8 mm SOUTH AFRICA: Kruger NP, Punda Milia, Van Rooven 494, long, stipules elliptic to ovate 2-5 mm long, pods (30-)35-45 x 19.iii.1976 (PRE). MADAGASCAR: Ankazoabo, Bosser 17241, 1.5-2.0 mm, seeds 1.5-1.8 x 1.2-1.3 mm, restricted to Australia Feb. 1963 (K). 2b. subsp. australis 428 Australian Systematic Botany J. S. Boatwright et al.

Fig. 6. Morphology of Rothia indica. (a) Standard petal. (b) Pod in lateral view. (c) Leaf of R. indica subsp. australis in abaxial view. (d 1 , d2) Stipules of R. indica subsp. australis. (e) Leaf ofR. indica subsp. indica in abaxial view. (fl ,f2) Stipules ofR. indica subsp. indica. (g) Pistil. (h) Wing petal. (i) Keel petal. (j) Androecium. (k) Flower in lateral view. (1) Anther. (m) Bract. (n) Seed in lateral view. (o) Outer surface of the calyx (upper lobes to the left). Voucher

specimens. (a, g, h, i, k) Cooray 70020117R (K). (b, n) Gamble 13771 (K). (c, d 1 , d2) Latz 16126 (MEL). (e, fl, f2) Barber 55 (K). ( j, 1, o) Wight 828. (K). (m) Wight 571 (K). All scale bars represent I mm.

2a. Rothia indica (L.) Druce subsp. indica also been recorded from China (Chen and Li 1997) and the Leaves with terminal leaflet (7—)10-26 long, (2—)4-8 mm wide, specimens cited by these authors were included on the map. It lateral leaflets (5—)8-20 mm long, (1.5—)3.0-6.0 mm wide; favours sandy soils, grassy slopes or moist open fields. Common petiole 5-13 mm long. Stipules elliptic to lanceolate, in disturbed areas such as roadsides. 2-8(-10) mm long. Inflorescence with flowers solitary or 2-4 per raceme. Calyx 5.0-6.5 mm long; tube 2-4 mm long; Selected specimens (33 specimens examined) lobes 1-3 mm long. Standard 4.0-6.5 mm long; claw 1.5-2.0 mm INDIA: Chinglepul, Barber 55, 10.i.1899 (K); Bombay (Mumbai), long; lamina 2.5-4.0 mm long, 2.0-2.5 mm wide. Wings 4-6 mm Dalzell s.n., iv.1878 (K); Ganjam district, Chennai, Gamble 13771, long; claw 2.0-2.5 mm long; lamina 2.5-4.0 mm long, ii.1884 (K); Bengaluru, Gough 162, Dec 1937 (K); Tirunelveli, 1.0-1.5 mm wide. Keel 4-6 mm long; claw 2.0-2.5 mm long; Tamil Nadu, Matthew 16388, 26.i.1979 (BM); Denkanikotta, lamina 2.5-4.0 mm long, 1.0-1.5 mm wide. Pistil with ovary Dhannapuri, Denkanikotta Rest House, Matthew RHT 24647, 15.xi.1979 3-4 mm long, 0.5-0.7 mm wide with ±14-18 ovules; style (K); Kodaikanal, Dindigul, Law's Ghat Rd, Matthew RHT 51949, 1.4-1.6 mm long. Pods (-33)35-55 mm long, (1.5—)2.0-3.0 mm 4.xii.1987 (K); Mysore, Mimso 607, x.1837 (K); Hassan, Mysore, wide with up to ±35 seeds per pod. Seeds 1.3-1.6 mm long, Belvathally, Ramamoorthy 1956, 13.xii.1971 (K); 70 km N of 1.0-1.3 mm wide. Munnar, Palani hills, Tamilnadu, Van der Maesen 3453, 20.i.1979 (K). VIETNAM: Tourane (now Da Nang), Clemens & Clemens 4294, Distribution and ecology: collections from India, Sri Lanka, v—vii.1927 (BM). SRI LANKA: Ruhuna NP, Block 1 near Buttawa Vietnam and Malaysia were studied (Fig. 7); this subspecies has Bungalow, Cooray & Balakrishnan 69010909R, 9.vi.1969 (BM); Revision of the genus Rothia Australian Systematic Botany 429

thanked for making specimens available for loan or study. Lorna Ngugi 40 (Queensland Herbarium, Australia) is thanked for supplying digital images of the type specimen of Rothia indica subsp. australis and Dr J.-N. Labat 30 (Museum National d'Histoire Naturelle) for images of the type specimen of R. hirsuta. Hester Steyn (National Herbarium, Pretoria) kindly provided the

20 map of Asia and Australia.

10 References Bairiganjan GC, Patnaik SN (1989) Chromosomal evolution in Fabaceae. Cytologia 54, 54-64. 10 Baker EG (1926) 'The Leguminosae of tropical Africa. 1' (Erasmus Press: Ghent, Belgium) 20 Boatwright JS, Savolainen V, Van Wyk B-E, Schutte-Vlok AL, Forest F, Van der Bank M (2008a) Systematic position of the anomalous 30 genus Cadia and the phylogeny of the tribe Podalyrieae (Fabaceae). Systematic Botany 33, 133-147.

40 0 700 1400 km doi: 10.1600/036364408783887500 Boatwright JS, Le Roux MM, Wink M, Morozova T, Van Wyk B-E (2008b) 60 70 80 100 110 120 130 140 150 160 Phylogenetic relationships of the tribe Crotalarieae (Fabaceae) inferred from DNA sequences and morphology. Systematic Botany 33, Fig. 7. Known geographical distribution of Rothia indica subsp. indica 752-761. doi: 10.1600/036364408786500271 (triangles) and R. indica subsp. australis (dots). Specimens cited in Chen and Chen T, Li Z (1997) Rothia, a newly recorded genus of Leguminosae from Li (1997) from China are represented by stars. China. Journal of Tropical and Subtropical Botany 5, 11-16. Crisp MD, Gilmore S, Van Wyk B-E (2000). Molecular phylogeny of the Trincomalee, near China Bay Airport, Rudd & Balakrishnan 3134, 24. genistoid tribes of papilionoid legumes. In 'Advances in legume ii.1970 (K); Hambantota, Yala, Fagerlind 1695, i.1974 (S). systematics 9'. (Eds PS Herendeen, A Bruneau) pp. 249-276. (Royal MALAYSIA: Pulau Pinang, Haihtmu 2996, 21.xii.1917 (K). Botanic Gardens, Kew: London) Feder N, O'Brien TP (1968) Plant microtechnique: some principles and new 2b. Rothia indica subsp. australis A. E. Ho I land, Austrobaileya methods. American Journal of Botany 55, 123-142. 5:94 (1997) doi: 10.2307/2440500 Type: Queensland, Gregory South District, Site 195, Warlus 1, 7 Goldblatt P (1981) Cytology and the phylogeny of Leguminosae. Aug. 1971, Boyland 4016; holo BRI (photo seen). In 'Advances in legume systematics 1'. (Eds RM Polhill, PH Raven) pp. 427-463. (Royal Botanic Gardens, Kew: London) Leaves with terminal leaflet 9-15 mm long, 3-5 mm wide, Gregory M, Baas P (1989) A survey of mucilage cells in vegetative organs lateral leaflets 9-12 mm long, 2.5-4.0 mm wide; petiole 4-8 mm of the dicotyledons. Israel Journal of Botany 38, 125-174. long. Stipules elliptic to ovate, 2-5 mm long. Inflorescence with Hepper FN (1958) Papilionoideae. In 'Flora of west tropical Africa'. flowers solitary or 2 per raceme. Calyx 4-7 mm long; tube (Ed. RWJ Keay) pp. 543-544. (Crown Agents: London) 1.5-3.0 mm long; lobes 3.5-4.0 mm long. Standard Holland AE (1997) Rothia indica subsp. australis A.E.Holland (Fabaceae: 4.0-6.5 mm long; claw 1.5-2.0 mm long; lamina 2.5-4.0 mm Crotalarieae), a new subspecies occurring in Australia. Austrobaileya 5, 93-96. long, 2.0-2.5 mm wide. Wings 4.0-6.5 mm long; claw Holmgren PK, Holmgren NH, Barnett LC (Eds) (1990) 'Index Herbariorum. 2.0-2.5 mm long; lamina 2.5-4.5 mm long, 1.0-1.5 mm wide. Part I: the herbaria of the world.' 8th edn. Regnum Vegetabile Vol. 120. Keel 4.0-6.5 mm long; claw 2.0-2.5 mm long; lamina 704 pp. (Botanical Garden Press: New York) 2.5-4.5 mm long, 1.0-1.5 mm wide. Pistil with ovary Hussain RA, Kinghom AD, Molyneux RJ (1988) Alkaloids of Rothia 4.0-4.5 mm long, 0.6-1.0 mm wide with ±13-17 ovules; style trifoliata and Rothia hirsuta. Journal of Natural Products 51, 809-811. 1.5-1.7 mm long. Pods (30—)35-45 mm long, 1.5-2.0 mm wide doi: 10.1021/np50058a033 with up to ±30 seeds per pod. Seeds 1.5-1.8 mm long, Milne-Redhead E (1971) Rothia. In 'Flora of tropical east Africa'. 1.2-1.3 mm wide. (Eds JB Gillett, RM Polhill, B Verdcourt) pp. 811. (Crown Agents: London) Distribution and ecology: restricted to Australia where it Polhill RM (1974) A revision of Pearsonia (Leguminosae-Papilionoideae). occurs on sandy hills and flats (Fig. 7). Kew Bulletin 29, 383-410. doi: 10.2307/4108548 Polhill RM (1976) Genisteae (Adans.) Benth. and related tribes Specimens examined (Leguminosae). Botanical Systematics 1, 143-368. AUSTRALIA: 100 miles [160.9 km] W of Windorah, Everist 3898, Polhill RM (2003) Rothia. In 'Flora Zambesiaca 3'. (Eds GV Pope, 9.vi.1949 (K); Boomerang Waterhole, Lander River, 20°38'00"S RM Polhill, ES Martins) p. 64. (Royal Botanic Gardens, Kew: 132°1 1 '00"E, Latz 16126, 10.v.2000 (MEL); Upper Victoria River, London) von Muller s.n., without date (K, MEL 2 sheets); near the Ord River, Rudd VE (1991) Fabaceae. In 'A revised handbook of the flora of Ceylon, 7'. O'Donnell s.n., without date (MEL 2 sheets). (Eds MD Dassanayake, FR Fosberg) pp. 184-185. (Amerind Publishing: New Delhi) Stafleu FA, Cowan RS (1983) 'Taxonomic literature vol 4: P-Sak.' 2nd edn. Acknowledgements (Bohn, Scheltema & Holkema: Utrecht, The Netherlands) The National Research Foundation (NRF) and University ofJohannesburg are Van Wyk B-E (1991a) A review of the tribe Crotalarieae (Fabaceae). thanked for financial support. The curators and staff of the listed herbaria are Contributions from the Bolus Herbarium 13, 265-288. 430 Australian Systematic Botany J. S. Boatwright et al.

Van Wyk B-E (1991b) A synopsis of the genus Lotononis (Fabaceae: Van Wyk B-E, Verdoom GH (1989) Chemotaxonomic significance of Crotalarieae). Contributions from the Bolus Herbarium 14, 1-292. alkaloids in the genus Pearsonia. Biochemical Systematics and Van Wyk B-E (2003) The value of chemosystematics in clarifying Ecology 17, 391-394. doi: 10.1016/0305-1978(89)90054-9 relationships in the genistoid tribes of papilionoid legumes. Van Wyk B-E, Verdoorn GH (1990) Alkaloids as taxonomic characters in Biochemical Systematics and Ecology 31, 875-884. the tribe Crotalarieae (Fabaceae). Biochemical Systematics and Ecology doi: 10.1016/S0305-1978(03)00083-8 18, 503-515. doi: 10.1016/0305-1978(90)90122-V Van Wyk B-E (2005) Crotalarieae. In 'Legumes of the World'. (Eds G Lewis, Van Wyk B-E, Verdoom GH (1991) Chemotaxonomic significance of B Schrire, B Mackinder, M Lock) pp. 273-281. (Royal Botanic Gardens, alkaloids in the genus Robynsiophyton. Biochemical Systematics and Kew: London) Ecology 19, 681-683. doi: 10.1016/0305-1978(91)90086-F Van Wyk B-E, Schutte AL (1995) Phylogenetic relationships in the tribes Podalyrieae, Liparieae and Crotalarieae. In 'Advances in legume systematics 7: phylogeny'. (Eds M Crisp, JJ Doyle) pp. 283-308. (Royal Botanic Gardens, Kew: London) Manuscript received 24 July 2008, accepted 27 October 2008

http://www.publish.csiro.au/joumals/asb Appendix B5. Boatwright, J. S. and B.-E. van Wyk. 2009. A revision of the

African genus Robynsiophyton (Crotalarieae, Fabaceae). South African

Journal of Botany 75: 367 - 370.

Available online at www.sciencedirect.com -,0 SOUTH AFRICAN °°,7' ScienceDirect JOURNAL OF BOTANY

ELSEVIER South African Journal of Botany 75 (2009) 367-370 www.elsevier.com/locate/sajb

Short communication A revision of the African genus Robynsiophyton (Crotalarieae, Fabaceae)

J.S. Boatwright*, B.-E. Van Wyk

Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland Park 2006, South Africa

Received 23 July 2008; received in revised form 24 November 2008; accepted 27 November 2008

Abstract

The monotypic genus Robynsiophyton is revised. Robynsiophyton vanderystii occurs in central and southern tropical Africa and is unique in the reduction of the androecium to nine stamens and five fertile anthers. It is morphologically similar to Pearsonia and Rothia and sister to the latter based on morphological and DNA sequence data. A revision of the genus is presented, including illustrations of vegetative and reproductive features and distributional information. 2008 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Crotalarieae; Fabaceae; Robynsiophyton vanderystii; Taxonomic revision

1. Introduction within Pearsonia, but sister to Rothia with both of these subsequently sister to Pearsonia. These data in combination Robynsiophyton Wilczek is a poorly known, monotypic with the annual life history and the unusual androecium (nine genus that occurs in south-central tropical Africa. It is stamens with only five fertile anthers) support the generic morphologically similar to Pearsonia Diimmer and Rothia concept of Robynsiophyton. Pers., suggesting a close relationship with these genera (Polhill, The aim of this paper is to present a revision of Robynsio- 1976, 1981; Van Wyk, 1991). Pearsonia occurs in central and phyton with illustrations, a discussion on diagnostic characters southern tropical Africa, while Rothia is widely distributed and a distribution map. throughout tropical Africa, Asia and Australia (Polhill, 1974; Boatwright et al., in press). These three genera differ from the Materials and methods rest of the tribe in their monomorphic anthers and straight styles (Polhill, 1976; Van Wyk, 1991; Van Wyk and Schutte, 1995; 2.1. Morphology Boatwright et al., in press). Chemically they are also unique within Crotalarieae in accumulating angelate esters of hydro- Morphological data were obtained through the study of xylupanine-type alkaloids (Van Wyk and Verdoorn, 1991). herbarium material from BM, K and PRE (abbreviations Recently, Boatwright et al. (2008), through the study of according to Holmgren et al., 1990). Digital images of critical molecular (ITS and rbcL) and morphological data, demon- specimens were obtained from BR and LISC. Illustrations were strated that Pearsonia, Robynsiophyton and Rothia form a prepared using a stereoscope (WILD M3Z) with a camera lucida strongly supported Glade and that the latter two are strongly attachment. supported as sister genera (Fig. 1). Doubt regarding the generic status of Robynsiophyton has been expressed by previous Results and discussion authors (Polhill, 1976; Van Wyk, 1991), who suggested that it could merely be a local derivative of Pearsonia. However, A summary of the diagnostic characters for Robynsiophyton, according to Boatwright et al. (2008) the genus is not embedded Rothia and Pearsonia is presented in Table 1. Robynsiophyton vanderystii Wilczek is a small annual (or rarely a short-lived • Corresponding author. perennial) with hairy, reddish-brown branches. It shares with E-mail address: [email protected] (J.S. Boatwright). Rothia the annual life history, a trait which is not found in

0254-6299/$ - see front matter © 2008 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2008.11.005 368 J.S. Boatwright, B.-E. Van 1114 / South African Journal of Botany 75 (2009) 367-370

100 Pearsonia sessilifolia 4. Taxonomy 99 Pearsonia atistata 97 Pearsonia grandifolia 100 4.1. Robynsiophyton Pearsonia cajanifolia 99 E Rothia hirsuta Wilczek in Bull. Jard. Bot. Etat. 23: 128 (1953); in F. C. B. 4: Robynsiophyton vanderystii 286 (1953); Hutchinson, Genera of Flowering Plants: 362 (1964); Polhill in Bot. Syst. 1: 326 (1976); Van Wyk and Fig. I. Phylogenetic position of Robynsiophyton vanderystii based on gene Schutte, Advances in Legume Systematics 7: 306 (1995); sequences (ITS and rbcL) and morphological data (strict consensus of 370 trees Polhill, Fl. Zam. 3 (7): 66 (2003); Leistner, Seed Plants of from Boatwright et al., 2008; tree length = 1166; consistency index = 0.53; southern Tropical Africa: Families and Genera: 202 (2005); Van retention index = 0.84). Numbers above the branches are bootstrap percentages above 50%. Wyk in Legumes of the World, Lewis et al. (Eds): 281 (2005). Type species: R. vanderystii Wilczek. The genus is monotypic and distinguished by the reduced Pearsonia, a genus comprising perennial herbs or small shrubs. number of stamens (nine instead of 10) and presence of four The leaves are digitately trifoliolate and sparsely pubescent staminodes (Table 1). It occurs in central tropical Africa adaxially but densely pubescent on the abaxial surface. The extending from Angola to Zambia in the east. narrow stipules are paired at the base of the petiole. Rothia hirsuta (Guill. & Perr.) Bak. has single stipules at each leaf; 4.2. R. vanderystii Rothia indica (L.) Druce has paired stipules; and Pearsonia has the stipules paired or less often absent (Polhill, 1974; Wilczek in Bull. Jard. Bot. Etat. 23: 128 (1953); in F. C. B. 4: Boatwright et al., in press). 286, t. 17 (1953); White, F. F. N. R.: 164 (1962); Tone in C. F. A. The flowers of Pearsonia, Robynsiophyton and Rothia are 3: 6 (1962); Lock, Leg. Afr. Check-list: 232 (1989); Polhill, Fl. all relatively unspecialised, with straight or even down-curved Zam. 3 (7): 66 (2003). styles and monomorphic anthers (Polhill, 1976). In Rothia the Type: Democratic Republic of Congo, Lazaret du Sacre- anthers are all small and rounded as opposed to the large, Coeur [1025 BA], Vanderyst s.n. (BR, holo., photo!). elongate anthers of Pearsonia, six of which are attached slightly Small, prostrate or ascending annual or short-lived perennial higher up to the filament (Polhill, 1974, 1976). Robynsiophyton up to ±0.3 m in height. Branches reddish-brown, pubescent. has a reduced number of stamens (from 10 to nine) and only five Stipules 5-8 mm long, linear to lanceolate, invariably paired, fertile stamens (the other four stamens lack anthers and are pubescent. Leaves digitately trifoliolate; petiole shorter than sterile). This is the most notable feature of the genus. In the leaflets, 4-7 mm long; leaflets elliptic to oblanceolate, Crotalarieae, Genisteae and Podalyrieae there is great variation subsessile, sparsely pubescent adaxially and densely so in stamina] arrangement from completely free stamens abaxially, terminal leaflet 14-22 x 4-9 mm, lateral leaflets 7- (Podalyrieae) to those joined in either an open sheath 16 x 2.5-6.0 mm, obtuse, base cuneate. Inflorescence axillary (Crotalarieae) or closed tube (most Genisteae). The anthers or rarely terminal congested racemes, with (2—) 5 to 9 (-10) may be either dimorphic with alternating basifixed and flowers; pedicel less than 1 mm long; bract linear, 1.5-3.0 mm dorsifixed anthers, or monomorphic as found in Pearsonia, long, pubescent, caducous; bracteoles absent. Flowers pale Robynsiophyton and Rothia (a feature that is unique to this yellow, 3-6 mm long. Calyx equally lobed, pubescent, 4-5 mm Glade within the Crotalarieae). Anther characters are very long; tube 1.5-2.0 mm long; lobes subulate, 1.5-3.0 mm long, reliable in legumes and usually consistent with other characters, tips minutely pubescent on inner surface. Standard 3-5 mm thus providing important insight especially at generic and tribal long; claw 1.0-1.5 mm long; lamina elliptic to ovate, 2- levels (Bentham, 1843; Polhill, 1976; Boatwright et al., 2008, in 4 x 1.0-1.5 mm, obtuse to very slightly emarginate, pilose along press). dorsal midrib. Wings 2.5-4.5 mm long; claw 1.5-1.8 mm long; The calyx of Robynsiophyton is equally lobed, while in Pearsonia and Rothia the upper lobes are larger than the three Table I lower lobes. In Pearsonia, the upper and lateral lobes on either Summary of diagnostic characters for Robynsiophyton, Rothia and Pearsonia. side are often fused higher up (the so-called "lotononoid" calyx Character Robynsiophyton Rothia Pearsonia type). Bracts are present in all three genera, but bracteoles are Life history Annual or short- Annual Perennial generally lacking in Pearsonia (if present very small) and lived perennial completely absent in Robynsiophyton and Rothia (Polhill, Stipules Paired Paired or single Paired or absent 1974, 1976). Bracteoles Absent Absent Small or absent Calyx Equally lobed Sub-equally lobed Zygomorphic The pods of Robynsiophyton are relatively short and few- seeded when compared to those of Rothia. The mature seeds Androecium 9 Stamens, 10 Stamens, 10 Stamens, 5 rounded anthers, 10 rounded anthers, 10 elongated anthers, are brown, smooth and similar in size to those of R. indica. 4 staminodes 0 staminodes 0 staminodes Robynsiophyton vanderystii has larger seeds than Rothia hirsuta Fruit Oblong to ovate Linear to ovate or Ellipsoid to linear-oblong that are even coloured with a smooth surface, whereas those of falcate R. hirsuta are mottled with a somewhat rugose surface Seeds Brown, smooth Brown or mottled, Light or dark brown, (Boatwright et al., in press). smooth or rugose sometimes mottled, smooth J.S. Boatwright, B.-E. Van 1154c / South African Journal of Botany 75 (2009) 367-370 369 lamina oblong to obovate, as long as or slightly longer than 5. Distribution and habitat keel, 1.0-2.5 x 0.5-1.0 mm, obtuse, glabrous, with 1-2 rows of sculpturing. Keel 3-4 mm long; claw 1-2 mm long; lamina R. vanderystii occurs in moist, sandy soils and is especially boat-shaped, 1.5-3.0 x 0.5-0.8 mm, obtuse, glabrous, some- common along roadsides. Very few specimens are available for times with a very slight pocket. Stamens 9, anthers mono- study and these include collections from Angola, the Demo- morphic with 5, sub-basifixed anthers alternating with 4 cratic Republic of Congo and Zambia, but it is likely that the staminodes. Pistil subsessile, pubescent, ovary elliptic, 1.5- distribution range is more extensive (Fig. 3). 2.0 x 0.5-0.8 mm with ±5 to 8 ovules; style straight, 1.5- 1.8 mm long, glabrous. Pods oblong to ovate, laterally 5.1. Additional specimens examined compressed, subsessile, 7-10 x 3-4 mm, ±2 to 8 seeded, dehiscent. Seeds oblique-cordiform, 1.0-1.5 x 1.0-1.2 mm, Angola: brown, smooth (Fig. 2). Flowering time: April. Plants appear 0614: Kimbambu, Madimba (—AA), Pauwels 6470 (PRE). to be inconspicuous and are poorly collected so that the 0720: Lunda, Chicapa (—BC), Exell and Mendonca 657 flowering time needs confirmation. (BM, K, LISC 3 sheets).

Fig. 2. Morphology of Rokynsiophyton vanderystii (drawings by JSB): (a) leaf in abaxial view; (bl—b2) stipules; (c) flower in lateral view; (d) bract; (e) outer surface of the calyx (upper lobes to the left); (f) standard petal; (g) wing petal; (h) keel petal; (i) androecium showing the five uniform anthers and four staminodes; (j) anthers

(carinal anther on the left); (k) pistil; (I) pod in lateral view; (m) seed in lateral view. Voucher specimens: (c, d, f, g, h, i , j, k) McCallum Weston 717 (K); (e) Lisowski 20326 (K); (1) Richards 9321 (K); (a, bl —b2) Richards 18/44 (K); (m) Krell and Mendonca 657 (K). Scale bars: (a—m) I mm. 370 J.S. Boatwright, B.-E. Van IlYk . / South African Journal of Botany 75 (2009) 367-370

0 Gabon ' .- ...... - . Kenya ..:•....:' I ..; Conc. ° : Democratic Re 3ublic • of Congc : .1 ...... ',. Tanz. nia • • • ..• •

. 10 10 % J 1...... Angola : • ' • : '... " ...,. • • Mozambiq - . Zambia • . ••• .• .... . .. ..•••.. .• ,..... dagascar 1::::: .... fir ';. Zimb bwe 20 20

•••• Botsw na ,- Namibia ..... '..

*. :•"4 "...- •• : :

....,.- South Af Ica :".. : 30 30

10 20 30 40 50

Fig. 3. Known geographical distribution of Robynsiophyton vandetystii.

Democratic Republic of Congo: Boatwright, J.S., Tilney, P.M., Van Wyk, B.-E., in press. A taxonomic revision of the 0416: Bakama (—AB), Flamigni s.n. (K). genus Rothia (Crotalarieae, Fabaceae). Australian Systematic Botany 21 (6). Boatwright, J.S., Le Roux, M.M., Wink, M., Morozova, T., Van Wyk, B.-E., 0515: Bas-Congo, Kisantu (—AA), Vanderyst s.n. (BM, K); 2008. Phylogenetic relationships of tribe Crotalarieae (Fabaceae) inferred Bas-Congo, Kimpako (—AB), Vanderyst 42290 (K). from DNA sequences and morphology. Systematic Botany 33, 752-761. 1025: Haut-Shaba, Kolwezi (—DA), Lisowski 20326, Holmgien, P.K., Holmgren, N.H., Barnett, L.C., 1990. Index Herbariorum Part 1: 20333 (K). The Herbaria of the World, 8th ed. Regnum Vegetabile, vol. 120. Zambia: Polhill, R.M., 1974. A revision of Pearsonia (Leguminosae-Papilionoideae). Kew Bulletin 29, 383-410. 0831: Chilongowelo, Abercorn (—CA), McCallum Polhill, R.M., 1976. Genisteae (Adans.) Benth. and related tribes (Leguminosae). Weston 717 (K); Mpulungu Abercom road, close to Chilongo- Botanical Systematics I, 143-368. welo turning (—CA), Richards 5309, 5317, 18144 (K), Polhill, R.M., 1981. Tribe 29. Crotalarieae (Benth.) Hutch. In: Polhill, R.M., Richards 11097 (K, PRE). Raven, P.H. (Eds.), Advances in Legume Systematics I. Royal Botanic 0929: Kawambwa, Timnatushi Falls (—CC), Richards Gardens, Kew, pp. 399-402. Van Wyk, B.-E., 1991. A review of the tribe Crotalarieae (Fabaceae). 9319a, 9321 (K). Contributions from the Bolus Herbarium 13, 265-288. 1124: Mwinilunga (—CD), Mutimushi 3416 (K). Van Wyk, B.-E., Verdoom, G.H., 1991. Chemotaxonomic significance of alkaloids in the genus Robynsiophyton. Biochemical Systematics and Acknowledgements Ecology 19, 681-683. Van Wyk, B.-E., Schulte, A.L., 1995. Phylogenetic relationships in the tribes Podalyrieae, Liparieae and Crotalarieae. In: Crisp, M., Doyle, J.J. (Eds.), Funding from the National Research Foundation (NRF) and Advances in Legume Systematics 7, Phylogeny. Royal Botanic Gardens, University of Johannesburg are gratefully acknowledged. The Kew, pp. 283-308. authors would like to thank the curators and staff of the listed herbaria for making specimens available for study or on loan.

References

Bentham, G., 1843. Enumeration of Leguminosae, indigenous to southern Asia, and central and southern Africa. Hooker's London Journal of Botany 2.

Edited by JC Manning Appendix B6. Boatwright, J. S., P. M. Tilney and B.-E. van Wyk. The generic concept of Lebeckia, reinstatement of the genus Calobota and a new genus of the tribe Crotalarieae (Fabaceae). South African Journal of Botany, submitted for publication. The generic concept of Lebeckia, reinstatement of the genus Calobota

and a new genus of the tribe Crotalarieae (Fabaceae)

J.S. Boatwright*, P.M. Tilney, B.-E. van Wyk

Department of Botany and Plant Biotechnology, University of Johannesburg,

P.O. Box 524, Auckland Park 2006, South Africa

*Corresponding author.

E-mail address: [email protected] (J.S. Boatwright).

Abstract

The reinstatement of the genus Calobota Eckl. and Zeyh. is proposed

and Wiborgiella Boatwr. and B.-E.van Wyk described as new. Evidence from

the internal transcribed spacer (ITS) of nuclear ribosomal DNA and plastid

rbcL data revealed that the genus Lebeckia Thunb. is not monophyletic.

These analyses, along with morphological and anatomical data, showed that the genus should be divided into three genera: (1) Lebeckia senso stricto (L.

section Lebeckia); (2) Calobota [L. section Calobota (Eckl. and Zeyh.) Benth. and L. section Stiza (E.Mey.) Benth.] including the monotypic, North African genus Spartidium Pomel; (3) a new genus, Wiborgiella [L. section

Viborgioides Benth., together with L. inflata Bolus, L. mucronata Benth. and

Wiborgia humilis (Thunb.) Dahlgr.]. Brief synopses of the genera Calobota

1 and Wiborgiella, including nomenclature, synonymy, descriptions and diagnostic characters are presented.

Keywords: Calobota, Crotalarieae, Fabaceae, Lebeckia, new genus,

Wiborgiella

2 1. Introduction

The current broad generic concept of Lebeckia Thunb. dates back to

Bentham (1844) and Harvey (1862) and refers to a group of ca. 36 species of

papilionoid legumes that occur mainly in the southern and western parts of

South Africa, with some extending into Namibia. The group is particularly

common in the Cape Floristic Region (CFR). Bentham (1844) reduced several

genera described by Meyer (1836) and Ecklon and Zeyher (1836) to the

synonymy of an expanded Lebeckia. This broadened concept included

Meyer's Stiza and Sarcophyllum, together with Ecklon and Zeyher's

Acanthobotrya and Calobota. A new sectional classification was proposed,

based mainly on the shape of the keel and the morphology of the fruit. This

comprised five sections, viz. section Calobota (Eckl. and Zeyh.) Benth.,

section Eulebeckia Benth., section Phyllodiastrum Benth., section Stiza

(E.Mey.) Benth., and section Viborgioides Benth. Harvey (1862) followed this

sectional classification of Lebeckia for his treatment in the Flora Capensis. For

nearly 150 years, the generic concept and relationships between the

morphologically rather diverse species were never studied in depth. A revision

of Lebeckia sensu stricto (sections Eulebeckia and Phyllodiastrum) was

recently completed by Le Roux and Van Wyk (2007, 2008, in press). Their

results show that the 14 species of Lebeckia s.s. can easily be distinguished

by their acicular leaves and 5+5 anther arrangement, not only from all other species of the so-called "Cape" group of the tribe Crotalarieae (i.e. Aspalathus

L., Rafnia Thunb. and Wiborgia Thunb.) but also from all other species hitherto included in Lebeckia.

3 Spartidium Pomel is a monotypic genus that occurs in North Africa.

The affinities of the genus within the genistoid legumes have been unclear for some time as a possible relationship with both Retama Raf. and Lebeckia have been suggested (Polhill, 1976). The genus is currently placed within the

Crotalarieae based on the open androecial sheath, but an evaluation of its systematic position is clearly desirable. Polhill (1981) considered Spartidium to be "virtually indistinguishable from Lebeckia". He used the orientation of the seeds in the fruit and the North African distribution as the only key characters to distinguish Spartidium.

In a broad study of molecular (ITS, rbcL) and morphological data of

117 species representing all the genera of the Crotalarieae, Boatwright et al.

(in press) discovered that Lebeckia is paraphyletic (Fig. 1 a and b). This study revealed important new relationships within the Crotalarieae as well as

Lebeckia and showed the need for new generic circumscriptions. The species can be readily accommodated in three easily recognizable morphological groups (genera): (1) Lebeckia s.s. (L. section Lebeckia, including sections

Phyllodiastrum and Eulebeckia); (2) Calobota Eckl. and Zeyh. [L. section

Calobota (Eckl. and Zeyh.) Benth. and L. section Stiza (E.Mey.) Benth., together with the monotypic North African Spartidium Pomel] and (3)

"Wiborgiella" [L. section Viborgioides Benth., together with L. inflata Bolus, L. mucronata Benth. and Wiborgia humilis (Thunb.) Dahlgr.].

In this paper, new evidence is presented to show that Bentham's

(1844) broad concept of Lebeckia is paraphyletic. We propose a new generic classification system to reflect new insights into morphological discontinuities

4 within the tribe which are also supported by molecular systematic evidence

(Boatwright et al., in press).

2. Materials and methods

2.1. Morphology

Morphological data were obtained from herbarium specimens as well

as fresh material collected while studying various taxa of the Crotalarieae in

the field. Specimens of Lebeckia from the following herbaria were studied:

BM, BOL, GRA, J, JRAU, K, NBG (including SAM and STE), P, PRE, S, SBT,

UPS and WIND (abbreviations according to Holmgren et al., 1990). Online

photographs of the collections of B, M and Z were studied. Drawings of

reproductive structures (all by JSB) were done using a stereoscope (WILD

M3Z) with a camera lucida attachment.

2.2. Anatomy

For anatomical studies, material was fixed in FAA (formaldehyde:

acetic acid: 96% alcohol: water; 10:5:50:35 ) or dried material was rehydrated

and then fixed in FAA for 24 hours. A modification of the method of Feder and

O'Brien (1968) was used for embedding in glycol methacrylate (GMA). A

minimum of five days was used for the third infiltration in GMA. Sections were stained according to the periodic acid Schiff/toluidine blue (PAS/TB) staining method.

5 3. Morphological and anatomical evidence for the paraphyly of Lebeckia

3.1. Habit and branches

Most genera of the Crotalarieae have a shrubby habit (Polhill, 1976), but variation in habit is quite pronounced within Lebeckia. Species of Lebeckia section Lebeckia are predominantly suffrutescent plants that branch mainly at ground level, whereas the species of the remaining sections are almost invariably woody shrubs. A strongly spinescent habit is characteristic of

Lebeckia section Stiza but also of two species of section Calobota, viz. L. acanthoclada Dinter and L. spinescens Harv. Both sections Calobota and

Stiza have green young branches (except L. acanthoclada), as opposed to the woody, rigid, ramified brown to greyish branches of section Viborgioides

(similar to the genus Wiborgia Thunb.). In species with green stems [sections

Calobota, Lebeckia and Stiza, and Spartidium saharae (Coss. and Dur.)

Pomel] a large part of the cortex is composed of chlorenchyma which serves a photosynthetic function (Metcalfe and Chalk, 1950). This layer is absent in species of section Viborgioides and also L. mucronata.

3.2. Leaves

Leaves are extremely variable, ranging from trifoliolate and petiolate

(sections Calobota and Viborgioides), to unifoliolate (section Stiza) or simple

(section Calobota and Spartidium) and completely phyllodinous (and acicular)

6 in section Lebeckia. Cultivated plants of the section Stiza showed that the juvenile leaves are petiolate and trifoliolate, but a loss of the lateral leaflets and shortening of the petioles result in unifoliolate leaves on the older branches, as was also mentioned by Polhill (1976) and Dahlgren (1970). This transition is also seen in L. obovata Schinz. Stipules are absent in all but two species of Lebeckia s.I.: L. wrightii (Harv.) Bolus and L. uniflora M.M.le Roux and B.-E.van Wyk (Le Roux and Van Wyk, in press).

Transverse sections through the leaves of Lebeckia species (Fig. 2) revealed a remarkable difference between the species of

Calobota/Stiza/Spartidium on the one hand (hereafter called the Calobota group) and Viborgioides/L. inflata/L. mucronata/Wiborgia humilis on the other hand (hereafter called the Viborgioides group), namely that the former group has isobilateral leaves, while the latter has dorsiventral leaves. This difference was unexpected, as anatomical characters are generally regarded as conservative. Even more remarkable was the exact congruence between the anther arrangements of the two groups (respectively 4+1+5 and 4+6 — see later). Leaves of Lebeckia s.s. are completely different from the species mentioned above, as they are phyllodinous, acicular and terete in transverse section.

Another interesting difference between the Calobota group and the

Viborgioides group is the presence of mucilage cells in the latter but not the former (Fig. 2). In this character, the Viborgioides group agrees with other

Cape genera (Aspalathus, Rafnia and Wiborgia, as well as the predominantly

Cape Lebeckia s.s.). In contrast, the Calobota group, which extends into arid regions (Northern Cape and Namibia), does not have mucilage cells in mature

7 leaves. The North African Spartidium, however, has mucilage cells in the epidermis, as do immature leaves of L. pungens, suggesting that there is a loss of these cells as the leaves mature (Boatwright, pers. obs.). Mucilage cells are widely distributed throughout the various plant families and several authors have speculated about their function (Gregory and Baas, 1989).

Although experimental evidence is lacking, mucilage cells are thought to aid in water storage and reduction of transpiration, protection against intensive radiation and also against herbivory (Gregory and Baas, 1989). Mucilage cells are often associated with plants that occur in Mediterranean climates (Van der

Merwe et al., 1994). Bredenkamp and Van Wyk (1999) speculated that in

Passerina L. (Thymelaeaceae) the mucilage serves as a regulator of hydration in the leaves, protecting them against water loss, but also helping to accumulate reserve water in the leaves. Lebeckia s.s. and the Viborgioides group are restricted to the CFR and therefore diversifying in a Mediterranean climate could explain the presence of the mucilage cells in these taxa. It is interesting to note the presence of mucilage cells in Spartidium saharae

(Coss. and Dur.) Pomel, which also occurs in a Mediterranean climate. The

Calobota group (excluding Spartidium) extend out of the CFR into summer rainfall, more arid regions and show a different adaptation to drought, viz. isobilateral leaves (Fig. 2 a, b; also in Spartidium). Van der Merwe et al.

(1994) mention that the presence of more layers of palisade parenchyma improves the transport of water through the leaves and also offers increased protection to the chloroplasts. In contrast, the leaves of species within the

Viborgioides group are dorsiventral (Fig. 2 c, e, f). In Lebeckia s.s., the

8 acicular leaves have a complete circular zone of palisade cells, with no spongy parenchyma (Fig. 2 d).

The petioles are always shorter than the leaflets in the Viborgioides group. In the Viborgioides group the persistent petioles become hard and woody when the leaflets are shed, contributing to the rigid and somewhat thorny appearance of the branches. These characters are also found in some species of Wiborgia s.s. (Dahlgren, 1975). In the Calobota group, the petioles are either longer or shorter than the leaflets and sometimes persistent, but never rigid and spinescent. The acicular leaves of some species of Lebeckia s.s. are articulated or "jointed" near the middle or reduced to petioles in species with unarticulated leaves, i.e. the leaves are phyllodinous (Dahlgren,

1970), which serves as a distinct synapomorphy for this group.

3.3. Inflorescences

As in most Crotalarieae, the inflorescences in Lebeckia are all terminal, multi-flowered racemes, varying in length and number of flowers. In Lebeckia s.s., the inflorescences may be relatively long and are often densely flowered with up to 93 flowers per raceme, for example in L. brevicarpa M.M.le Roux and B.-E.van Wyk (Le Roux and Van Wyk, 2007). Species of section

Calobota generally have fewer flowers per inflorescence, except in L. melilotoides Dahlgr., where more than 100 flowers are found on the elongated racemes (Dahlgren, 1967). The racemes of the three species of section Stiza are characteristically spine-tipped. Very short and few-flowered inflorescences

9 are found in L. section Viborgioides, with the flowers often solitary in L. bowieana Benth.

3.4. Flowers

The calyx structure is often an important generic character in the

Crotalarieae. Lebeckia species normally have equally lobed or "lebeckioid" calyces (Polhill, 1976), as opposed to the zygomorphic calyx ("lotononoid") in

Lotononis (DC.) Eckl. and Zeyh. and Pearsonia Durnmer or the bilabiate calyx typical of all members of the tribe Genisteae (Van Wyk, 1991a, Van Wyk and

Schutte, 1995). In the Calobota and Viborgioides groups, the carinal lobe is slightly narrower than the upper and lateral lobes, whereas in Lebeckia s.s. it is equal to the other lobes (Fig. 3). In the former two groups, the calyces are pubescent or at least glabrescent as opposed to the usually glabrous calyces found in section Lebeckia (L. wrightii is an exception in having a sparsely pubescent calyx).

The pubescence of the petals, the shape of the keel petal and the arrangement of the anthers closely follow the three major groups within

Lebeckia s.l. Glabrous petals are found in Lebeckia s.s. and the Viborgioides group, whereas species of the Calobota group generally have pubescent petals (or at least pilose along the dorsal midrib of the standard petal).

Lebeckia macrantha Harv. and L. psiloloba (E.Mey.) Walp. are the only exceptions and have totally glabrous petals. The keel petals in Lebeckia s.s. are characteristically rostrate as opposed to the obtuse keel petals found in the other groups (Fig. 3).

10 3.5. Anthers

Surprisingly, it was found that the size and shape of the carinal anther are diagnostic for each of the three groups in Lebeckia s.l. In Lebeckia s.s., the carinal anther resembles the long, basifixed anthers (Fig. 3 h3) resulting in a 5+5 arrangement. In the Calobota group (including Spartidium saharae), the carinal anther is intermediate between the dorsifixed and basifixed anthers

(Fig. 3 e3, f3, g3), resulting in a 4+1+5 anther configuration (i.e., four long basifixed anthers, an intermediate carinal anther and five short dorsifixed anthers). The carinal anther is usually attached a little higher up. In the

Viborgioides group, the carinal anther resembles the short, dorsifixed anthers

(Fig. 3 a3, b3, c3, d3), resulting in a 4+6 anther arrangement (i.e., four long anthers and six short anthers). The anther arrangement is therefore in perfect agreement with the leaf anatomy of the three groups.

3.6. Fruit and seeds

Pods in the Crotalarieae are an important source of systematic information, as specialisations for seed protection and dispersal may result in structural differences. Lebeckia s.l. displays a great diversity of fruit structure, including dehiscent and indehiscent fruits that are either inflated or laterally compressed, and with or without wings (Polhill, 1976). Fruits of Lebeckia s.s. are terete to semi-terete and thick- or thin-walled, with wings on the upper suture in some species, e.g. L. meyeriana Eckl. and Zeyh. ex Harv. Species of the Calobota group generally have terete or semi-terete pods that are thick-

11 walled and spongy or the fruits are thin-walled (membranous), laterally

compressed and pubescent or glabrous, as is also found in Spartidium

saharae. In contrast, the fruits of section Viborgioides (and L. inflata) are

inflated and always glabrous, with highly sclerified, thin walls. The placement

of Wiborgia humilis within the Viborgioides group is supported by fruit

structure. Wiborgia humilis has inflated pods that lack wings on the upper

suture and do not have highly sclerified fruit walls. In contrast, the winged

samaras of the rest of Wiborgia are laterally compressed and have highly

sclerified fruit walls in most species (Dahlgren, 1975).

Polhill (1976, 1981) used the orientation of the seed in Spartidium as

the only diagnostic character to separate this genus from Lebeckia s.l. A study

of most of the species showed that at least three species of the Calobota

group (L. macrantha, L. psiloloba and L. pungens Thunb.) have the seed

oriented at right angles to the placenta, exactly as in Spartidum saharae.

The seed surface of species in Lebeckia s.s. is invariably rugose, while

the seeds of only one species in the Calobota group (L. lotononoides Schltr.)

and one species in the Viborgioides group (L. inflata) have rugose seeds (Le

Roux and Van Wyk, 2007, 2008, in press; Boatwright and Van Wyk, 2007).

4. Molecular evidence for the paraphyly of Lebeckia

4.1. Combined molecular analyses

In the combined molecular analyses (Parsimony and Bayesian analyses; Fig. 1 a) of Boatwright et al. (in press), the Calobota group is clearly

monophyletic with very strong support. In the Bayesian analysis, Spartidium

12 groups with the Lebeckia multiflora group with strong support (Fig. 1 a). The

Viborgioides group is partly monophyletic with weak to strong support, but the positions of L. inflata and L. mucronata are unresolved. It is interesting to note that molecular evidence strongly supports the exclusion of Wiborgia humilis from Wiborgia and the transfer of this species to the Viborgioides group.

Surprisingly, Lebeckia s.s. is unresolved in the molecular analyses.

Morphologically this group is very distinct from the other groups of Lebeckia s.I. and all other Cape Crotalarieae.

4.2. Combined molecular and morphological analysis

When the molecular data were combined with morphological data (Fig.

1 b), the resolution within the "Cape" group and among the three groups of

Lebeckia s.I. was much improved. The Calobota group is again strongly supported as monophyletic. The Viborgioides group is partly monophyletic with strong support and L. inflata is sister to this group, albeit without support.

The position of L. mucronata, however, remains unresolved in this analysis, although abundant morphological and anatomical evidence suggests its placement within this group. The inclusion of W. humilis in the Viborgioides group again receives strong support. With the addition of morphological data

Lebeckia s.s. is moderately supported as monophyletic as opposed to being unresolved in the molecular analyses (Boatwright et al., 2008).

5. Taxonomic treatment

13 Major continuities and discontinuities in morphological characters

amongst the three groups discussed above closely agree with the three main

clades revealed by genetic analysis (Fig. 1 a and b). The new system

proposed here is based on a wider consideration of the intricate relationships

amongst all genera of the tribe Crotalarieae, all of which have been revised in

recent years (Pearsonia — Polhill, 1974; Wiborgia — Dahlgren, 1975; Crotalaria

— Polhill 1982; Aspalathus — Dahlgren 1988; Lotononis — Van Wyk, 1991b;

Rafnia — Campbell and Van Wyk, 2001; Bolusia — Van Wyk, 2003, Van Wyk

et al., submitted; Lebeckia s.s. — Le Roux and Van Wyk 2007, 2008, in press;

Robynsiophyton — Boatwright and Van Wyk, submitted; Rothia — Boatwright et

al., submitted). Dahlgren (1970) and Polhill (1976) both discussed the high

incidence of convergence in the tribe Crotalarieae and the intricate

relationships between the genera. The difficulty in determining generic limits

within the Crotalarieae is discussed by Polhill (1976), who emphasized that it

would be unwise to propose modifications to the system without a clear

understanding of the patterns of character state distributions. Furthermore,

alterations to the system should result in a more predictive and useful system without running the risk of instability of circumscriptions and nomenclature. In existing keys to the genera of the Crotalarieae (Polhill, 1981; Van Wyk and

Schutte, 1995), the lack of uniformity and clearcut diagnostic characters for

Lebeckia s.l. is clearly reflected in the fact that the genus keys out no less than three times (in both keys). Unique combinations of morphological characters have now been identified for the three main groups that we are convinced should be given generic status. The improvement in generic delimitations is also reflected in the following key.

14 5.1. Key to the genera of the Crotalarieae

la. Leaves acicular, terete:

Ovary 2 to 4 ovulate, pods 1 to 8-seeded Aspalathus

Ovary with more than 6 ovules, pods many-seeded Lebeckia s.s.

1 b. Leaves digitate, unifoliolate or simple (flat, never terete):

3a. Stipules present:

4a. Style curved upwards; anthers dimorphic:

Stipules asymmetrical or single; style glabrous and not helically coiled;

keel obtuse or rostrate; anther arrangement 4+6, 4+1+5 or very rarely

5+5 Lotononis

Stipules symmetrical when present; style with 1 or 2 lines of hairs or glabrous and helically coiled; keel strongly rostrate (often at right angles) or

helically coiled; anther arrangement 5+5:

Stipules dentate; beak of keel and style helically coiled, style glabrous Bolusia

Stipules entire; beak of keel and style not helically coiled, style with 1 or 2 lines of hairs Crotalaria

4b. Style straight or rarely down-curved; anthers similar in size and shape:

Stamens 9 (5 fertile and 4 lacking anthers) Robynsiophyton

Stamens 10 (all fertile):

Anthers all rounded; prostrate annuals Rothia

4 anthers basifixed, 6 attached slightly higher up, all elongate;

perennial herbs or shrubs Pearsonia

15 3b. Stipules absent:

Calyx with upper and lateral lobes fused on either side Lotononis

Calyx sub-equal (upper and lateral lobes not fused):

Leaves sessile; upper suture of pod asymmetrically convex:

11 a. Plants glabrous except occasionally on bracts and bracteoles; usually

turning black when dried Rafnia

11b. Plants usually pubescent on all parts, if leaves glabrous then standard

petal hairy and inner surface of calyx glabrous; not turning black when

dried Aspalathus

Leaves usually petiolate, if leaves sessile then plants with many-

seeded pods and at least some hairs on leaves or stems; upper suture

of pod symmetrically convex:

Petals pubescent; at least on the dorsal midrib of the standard petal (if

glabrous then plants strongly spinescent, woody, practically leafless

shrubs); twigs green (bark formation late. chlorenchyma present);

leaves isobilateral Calobota

Petals glabrous; twigs brown (bark formation early, chlorenchyma

absent; if twigs rarely green then plant a short-lived fireweed); leaves

dorsiventral:

Fruits winged, indehiscent; carinal anther intermediate (anthers 4+1+5)

Wiborgia

Fruits without wings, dehiscent (if rarely indehiscent then ovary and

fruit distinctly stalked); carinal anther resembles short anthers (anthers

4+6) Wiborgiella

16 5.2. Calobota

Calobota Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 192 (Jan. 1836) emend. Boatwr. and B.-E.van Wyk, emend. nov., Lebeckia section Calobota

(Eckl. and Zeyh.) Benth. in Hook. Lond. J. Bot. 3: 358-361 (1844) pro parte majore. Lectotype species (here designated): Calobota cytisoides (Berg.)

Boatwr. and B.-E.van Wyk [Note: This species is chosen as lectotype as it is the only species included in Ecklon and Zeyher's original concept of

Calobota].

Acanthobotrya Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2: 192 (Jan 1836) pro parte. Lectotype species (here designated): A. pungens sensu Eckl. and

Zeyh. [now Calobota psiloloba (E.Mey.) Boatwr. and B.-E.van Wyk]. [Note: As mentioned by Bentham (1844), Acanthobotrya is a mixed concept representing at least four different genera. However, the diagnosis agrees with the concept of Stiza E.Mey. (e.g. linear-oblong, compressed fruits); furthermore, C. psiloloba (at the time still part of the concept of A. pungens), which has all the diagnostic characters mentioned in the diagnosis, is listed first].

Stiza E.Mey., Comm. Pl. Afr. Austr. 1: 31 (Feb. 1836), synon. nov.

Lebeckia section Stiza (E.Mey.) Benth. in Hook. Lond. J. Bot. 3: 355-356

(1844). Lectotype species (here designated): Stiza erioloba E.Mey. [now

Calobota pungens (Thunb.) Boatwr. and B.-E.van Wyk]. [Note: Meyer (1836) included only Stiza erioloba and S. psiloloba E.Mey. (now Calobota psiloloba) in Stiza. The original description of Stiza is too general to allow for a

17 considered choice of lectotype, so we here choose C. pungens simply as it is the first-mentioned species.].

Spartidium Pomel, Nouv. Mat. Fl. Atl.: 173 (1874), synon. nov. Type species: Spartidium saharae (Coss. and Dur.) Pomel. [Note: Spartidium is monotypic].

[Note: The concept of Calobota is here expanded to include the genus Stiza and also the monotypic North African Spartidium (but excluding Lebeckia mucronata)].

Woody, sometimes spinescent shrubs or shrublets. Branches thick and woody; young branches green, lacking bark (except in C. acanthoclada), pubescent, often sericeous. Leaves unifoliolate, digitately trifoliolate (rarely 4- or 5-foliolate in some species) or simple; leaflets flat, oblanceolate to elliptic or obovate to suborbicular, less often spathulate, pubescent; petioles shorter or longer than leaflets. Stipules absent. Inflorescence terminal, few to multi- flowered racemes. Bracts linear to obovate, pubescent. Bracteoles linear, pubescent. Corolla yellow, usually pubescent or at least pilose along the median section of the standard petal (except in C. cuspidosa (Burch.) Boatwr. and B.-E.van Wyk and C. psiloloba). Calyx subequally lobed, upper sinus often deeper than the lateral or lower sinuses, carinal lobe narrower than the others, pubescent or at least glabrescent. Standard linear to widely ovate, with basal callosities in C. cytisoides. Wing petals narrowly oblong to oblong or slightly ovate, longer or shorter than the keel, terminal parts sometimes pubescent; apex obtuse. Keel petals oblong, pockets invariably present,

18 terminal parts sometimes pubescent; apex obtuse. Anthers dimorphic, four

long, basifixed anthers alternating with five ovate, dorsifixed anthers, carinal

anther intermediate in size and shape between the basifixed and dorsifixed

anthers (4+1+5 configuration). Pistil sub-sessile to very shortly stipitate; ovary

linear to slightly elliptic, with four to many ovules, pubescent or glabrous; style

shorter than the ovary, curved upwards, glabrous. Pods laterally compressed,

semi-terete or terete, linear to oblong, few- to many-seeded, glabrous or

pubescent, dehiscent or indehiscent. Seeds reniform to oblong-reniform, or

less often suborbicular; colour variable, light pink to pink, sometimes mottled with brown; hilum round, brown or black; surface smooth [rugose only in C. lotononoides (Schltr.) Boatwr. and B.-E.van Wyk; seeds of C. dinteri (Harms)

Boatwr. and B.-E.van Wyk, C. obovata (Schinz) Boatwr. and B.-E.van Wyk and C. saharae (Coss. and Durr.) Boatwr. and B.-E.van Wyk not seen].

5.2.1. Diagnostic characters

Bark formation is late, so that the twigs remain green and photosynthesizing, whereas it is early in most species of Wiborgiella, so that even the young twigs are not green but covered in brown bark (as in Wiborgia species). The green twigs are a useful diagnostic character, visible even in sterile herbarium specimens. Calobota also differs from Wiborgiella in the hairy petals (C. cupidosa and C. psiloloba are exceptions); they are glabrous in Wiborgiella. The most reliable diagnostic character to distinguish Calobota from Wiborgiella is the anther configuration of 4+1+5 (4+6 in the latter). The

19 pods are never inflated in Calobota and usually pubescent (pods inflated in most species of Wiborgiella and always glabrous).

5.2.2. Notes on distribution

The species of Calobota occur in the Eastern and Western Cape

Provinces, with some restricted to Namibia. One species, Calobota saharae, is endemic to North Africa where it occurs on sand dunes from Libya to

Algeria and Morocco (Polhill, 1976).

5.3. The species of Calobota

The following 16 species are recognised:

5.3.1. Calobota acanthoclada (Dinter) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia acanthoclada Dinter, Feddes Repert. 30: 196 (1932). Type—South

Africa, Namibia, Kleinfonteiner Flache, Dinter 6269 (B, photo!); Kovisberge,

Dinter 6293 (B, photo!, BM!); Buchuberge, Dinter 6574 (B, photo!, BM!);

Granietberge, Dinter 6694 (B, photo!).

Lebeckia spathulifolia Dinter, Feddes Repert. 30: 197 (1932) . Type—

South Africa, Namibia, Kleinfonteiner Flache, Dinter 3735 (B, photo!, SAM!, Z, photo! 2 sheets).

Lebeckia candicans Dinter, Feddes Repert. 30: 198 (1932), synon. nov.. Type—South Africa, Namibia, Kleinfonteiner Flache, Dinter 3737 (B, photo!, BOL!, NBG!, PRE!, S!, Z, photo!).

20 5.3.2. Calobota cinerea (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia cinerea E.Mey., Comm. Pl. Afr. Austr. 1: 35 (Feb. 1836). Type—

South Africa, hills near Noagas and near Aris on the Gariep, Drege s.n. (P!).

5.3.3. Calobota cuspidosa (Burch.) Boatwr. and B.-E.van Wyk, comb. nov.

Spartium cuspidosum Burch., Tray. S. Africa 1: 348 (1822). Type—South

Africa, Between `Gatikamma' and `Klaarwater [now Griquatown], Burchell

1697 (K!).

Lebeckia macrantha Harv. in Harv. and Sond., Fl. Cap. 2: 83-84

(1862), synon. nov. Type—South Africa, 'Zooloo country', Miss Owen s.n.

(TCD, photo!).

5.3.4. Calobota cytisoides (Berg.) Boatwr. and B.-E.van Wyk, comb. nov.

Spartium cytisoides Berg., Descr. Pl.: 199. 1767; L.f., Suppl. Pl. 320 (1781).

Type—South Africa, 'e Cap. b. Spei', Grubb s.n. sub Bergius 236.57 (SBT!).

Ebenus capensis L., Mant II: 264 (1771) nom. illegit.

Lebeckia cytisoides Thunb., Nov. Gen.: 143 (1800). [Note: Thunberg incorrectly cited Spartium cytisoides L.f.]

5.3.5. Calobota elongata (Thunb.) Boatwr. and B.-E.van Wyk, comb. nov.

Crotalaria elongata Thunb., Fl. Cap.: 571 (1823). Type—South Africa, `Carro prope Bockeveld', Thunberg s.n. sub THUNB-UPS 16544 (UPS!).

21 Lebeckia melilotoides Dahlgr., Bot. Notiser 120: 268-271 (1967).

Type—South Africa, Platfontein, E of Hottentots Holland Kloof, H. Hall 177

(NBG! , 2 sheets, SO.

5.3.6. Calobota halenbergensis (Merxm. and Schreib.) Boatwr. and B.-E.van

Wyk, comb. nov. Lebeckia halenbergensis Merxm. and Schreib., Bull. Jard.

Bot. Bruxelles 27: 276 (1957). Type—South Africa, Namibia, Luderitz South,

Halenberg, Dinter 6632 (M, photo!; K!, NBG!, PRE!, S!, Z, photo!).

5.3.7. Calobota linearifolia (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia linearifolia E.Mey., Comm. Pl. Afr. Austr. 1: 33 (Feb. 1836). Type—

South Africa, Northern Cape Province, on the Gariep, near Verleptpraam,

Drege s.n. (P!, S!).

Lebeckia dinteri Harms, Feddes. Repert. 16: 360 (1920), synon. nov.

Type—South Africa, Namibia, Garub, Dinter 1057 (NBG!).

5.3.8. Calobota lotononoides (Schltr.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia lotononoides Schltr. in Engler Bot. Jahr. 27: 143 (1900). Type—

South Africa, 'In regione namaquensi: In sabulosis montium Karree-Bergen, alt. C. 4500 ped.', Schlechter 8214 (BM!, BOL!, K!, PRE!, S!, Z, photo!).

5.3.9. Calobota obovata (Schinz) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia obovata Schinz, Mem. Herb. Boiss. 1: 126 (1900). Type—South

Africa, Namibia, Gansberg, Fleck 75 (Z, 2 sheets, photo!).

22 5.3.10. Calobota psiloloba (E.Mey.) Boatwr. and B.-E.van Wyk, comb. nov.

Stiza psiloloba E.Mey. Comm. Pl. Afr. Austr. 1: 31 (Feb. 1836). Lebeckia

psiloloba (E.Mey.) Walp., Linnaea 13: 478 (1839). Type—South Africa,

without locality, Drege 6470 (P!, K!, S!).

5.3.11. Calobota pungens (Thunb.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia pungens Thunb., Nov. Gen.: 141 (1800). Type—South Africa,

Eastern Cape Province, near Olifant's River [close to Oudtshoorn] at

Cannaland [now Little Karoo], Thunberg s.n. sub THUNB-UPS 16417 (UPS!).

5.3.12. Calobota saharae (Coss. and Dur.) Boatwr. and B.-E.van Wyk, comb.

nov. Genista saharae Coss. and Dur., B. Soc. Bot. France 2: 247 (1855).

Spartidium saharae (Coss. and Dur.) Pomel, Nouv. Mat. Fl. Atl.: 173 (1874).

Type—Algeria, D'Oran Province, Cosson s.n. (K!, P).

5.3.13. Calobota sericea (Ait.) Boatwr. and B.-E.van Wyk, comb. nov.

Spartium sericeum Ait. in Hort. Kew. 3: 12 (1789). Type—South Africa, Cape of Good Hope, Masson s.n. sub BM000794145 (BM, photo!). [The two specimens on the left were collected by Masson and are marked with '1' on the tape on the stems. The specimen on the right was grown in the gardens at

Kew and has the note 'Hort. Kew 1781' associated with it. According to

Stafleu and Cowan (1976), most of the Aiton types are in the Banks collection in BM)].

Lebeckia angustifolia E.Mey. in Linnaea 7: 155 (1832). Type—South

Africa, without locality, Ecklon s.n. (S!) [Note: the type is represented by the

23 two branches mounted at the bottom right hand side of the sheet – one with several flowers, the other with a single bud. Meyer (1932) incorrectly cites the type locality as "Uitenhage?" but expresses doubt by placing a question mark after the locality. The specimen in S comprises mixed elements but the two branches referred to above are accompanied by a label in Meyer's handwriting stating "Lebeckia angustifolia miff].

Lebeckia multiflora E.Mey., Comm. Pl. Afr. Austr. 1: 34 (Feb. 1836).

Type—South Africa, near Heerenlogement, Drege s.n. (P!); Olifantsrivier,

Drege s.n. (P!); between Holrivier and Mierenkasteel, Drege 6474 (P!, S!).

5.3.14. Calobota spinescens (Harv.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia spinescens Harv. in Harv. and Sond., Fl. Cap. 2: 88 (1862). Type—

South Africa, Rocks of Driekoppe, Drege s.n. (K, P!), Dwaka River, Burke s.n.

(K!), Great Fish River and Zout River, Beaufort, Zeyher 397 (K).

5.3.15. Calobota thunbergii Boatwr. and B.-E.van Wyk, nom. nov. pro

Lebeckia sericea Thunb. non Calobota sericea (Ait.) Boatwr. and B.-E.van

Wyk. Lebeckia sericea Thunb., Nov. Gen.: 143 (1800); Prodr. Pl. Cap.: 122

(1800). Type—South Africa, 'e Cap. b. Spei', Thunberg s.n. sub THUNB-UPS

16423 (UPS!) [Note: The epiphet `sericea' was applied to Lebeckia multiflora

E.Mey. (now Calobota sericea) by Aiton (1789), the basionym of which is

Spartium sericeum Ait., and is thus not available for the current taxon. We therefore propose a new name in memory of Thunberg who first described this species].

24 5.3.16. Calobota sp. 1. De Winter and Hardy 7919 (WIND!, K!, PRE!).

5.4. Wiborgiella

Wiborgiella Boatwr. and B.-E.van Wyk, gen. nov., Wiborgiae Thunb. similis sed fructibus oblongis non alatis valde inflatis, anthera carinali breve

(aliis in eodem fore brevibus dorsifixis simile) differt; Calobotae Eckl. and

Zeyh. similis sed petalis ubique glabris, antheris ut supra et foliis dorsiventralibus differt; Lebeckiae Thunb. sensu stricto similis sed foliolis planis non acicularibus valde differt. Type species: Wiborgiella leipoldtiana

(Dahlgr. ex Schltr.) Boatwr. and B.-E.van Wyk.

Lebeckia section Viborgioides Benth. in Hook. Lond. J. Bot. 3: 361

(1844). Lectotype species (here designated): Wiborgiella fasciculata (Benth.)

Boatwr. and B.-E.van Wyk [Note: As there is no indication as to which species would be a better choice of lectotype, we here choose the first listed species].

[Note: the new genus proposed here conforms to Bentham's (1844) concept of Lebeckia section Viborgioides, a taxon for which no name is available at generic level. The name Wiborgiella reflects the vegetative similarity and close relation to the genus Wiborgia].

Rigid, resprouting, woody shrubs (rarely lignotuberous shrublets or short-lived fireweeds). Branches thick and woody (except in two short-lived species); young branches brown, covered with bark (green in only two species), pubescent. Leaves digitately trifoliolate; leaflets flat, linear to widely oblanceolate or obovate, pubescent; petioles shorter than leaflets, usually

25 persistent and becoming woody after leaflets are shed, with prominent tubercles in some species. Stipules absent. Inflorescence terminal, multi- flowered racemes or rarely single-flowered. Bracts linear to lanceolate or elliptic, at least slightly pubescent. Bracteoles linear to lanceolate, at least slightly pubescent. Corolla yellow, glabrous, very rarely a few hairs along the dorsal midrib. Calyx subequally lobed, upper sinus often deeper than the lateral or lower sinuses, carinal lobe narrower than the others, pubescent or glabrous. Standard ovate to widely ovate to oblong. Wing petals oblong, ovate or rarely obovate, shorter, as long as or longer than the keel petals, glabrous; apex obtuse. Keel petals oblong or lunate, pockets present or absent, glabrous; apex obtuse. Anthers dimorphic, four long, basifixed anthers alternating with five ovate, dorsifixed anthers, carinal anther resembling the ovate, dorsifixed anthers (4+6 configuration). Pistil subsessile to stipitate; ovary linear to slightly elliptic, with 4 to many ovules, glabrous, very rarely slightly pubescent on the upper basal parts; style shorter or rarely longer than the ovary, curved upwards, glabrous. Pods terete or semi-terete, without wings, inflated or turgid (laterally compressed only in W. mucronata (Benth.)

Boatwr. and B.-E.van Wyk and Wiborgiella sp. 1), oblong to oblanceolate or lanceolate or elliptic, few- to many-seeded, glabrous, dehiscent (rarely indehiscent). Seeds reniform, light pink (W. leipoldtiana) or black with white or light brown spots (W. inflata and Wiborgiella sp. 1), surface smooth or rugose

[seeds of W. bowieana, W. fasciculata, W. humilis (Thunb.) Boatwr. and B.-

E.van Wyk, W. mucronata and W. sessilifolia (Eckl. and Zeyh.) Boatwr. and

B.-E.van Wyk not seen].

26 5.4.1 Diagnostic characters

The genus is similar to Wiborgia, but differs in the oblong, wingless, much inflated fruits (ovate to orbicular, markedly winged, samara-like and laterally compressed in Wiborgia). It also differs in that the carinal anther resembles the short, dorsifixed anthers (carinal anther intermediate in both

Wiborgia and Calobota). It further differs from Calobota in the glabrous petals

(in Calobota, at least the standard petal has a few apical hairs) and in the dorsiventral leaves (isobilateral in Calobota). It differs markedly from Lebeckia s.s. in the flat leaflets (invariably acicular in Lebeckia). Bark formation is early and the young twigs are brown and covered with bark as opposed to the green twigs of Calobota (the short-lived W. inflata and Wiborgiella sp. 1 have green stems).

5.4.2. Notes on distribution

The genus is endemic to the Greater Cape Floristic Region.

5.5. The species of Wiborgiella

The following eight species are recognised:

5.5.1. Wiborgiella bowieana (Benth.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia bowieana Benth. in Hook. Lond. J. Bot. 3: 362-363 (1844). Type—

South Africa, 'Cape', Bowie s.n. [K!, lectotype, designated by Dahlgren

(1975)].

27 5.5.2. Wiborgiella fasciculata (Benth.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia fasciculata Benth. in Hook. Lond. J. Bot. 3: 361 (1844). Type—

South Africa, 'Cape', Bowie s.n. [K!, lectotype, designated by Dahlgren

(1975)].

5.5.3. Wiborgiella humilis (Thunb.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia humilis Thunb., Nov. Gen.: 143 (1800). Wiborgia humilis (Thunb.)

Dahlgr., Op. Bot. 38: 31 (1975). Type—South Africa, `e Cap. b. Spei',

Thunberg s.n. sub THUNB-UPS 16416 [UPS!, lectotype, designated by

Dahlgren (1975)].

Wiborgia apterophora Dahlgr., Bot. Notiser 123: 112 (1970). Type—

South Africa, Gifberg, sand at edge of lands, Esterhuysen 22042 (BOL).

5.5.4. Wiborgiella inflata (Bolus) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia inflata Bolus in Hook. (cones Pl. 16: pl. 1576 (1887). Type—South

Africa, Western Cape Province, eastern slopes of Devil's Peak, Bolus 4826

(BOL!, K!, NBG!).

5.5.5. Wiborgiella leipoldtiana (Schltr. ex Dahlgr.) Boatwr. and B.-E.van Wyk,

comb. nov. Lebeckia leipoldtiana Schltr. ex Dahlgr., Op. Bot. 38: 72 (1975).

Type—South Africa, Western Cape Province, Calvinia Div.: 'Inter Grasberg et

Nieuwoudtville', Lewis 5839 (NBG!).

28 5.5.6. Wiborgiella mucronata (Benth.) Boatwr. and B.-E.van Wyk, comb. nov.

Lebeckia mucronata Benth. in Hook. Lond. J. Bot. 3: 359 (1844). Type—

South Africa, Uitenhage, Zeyher 344 (K!, TCD, photo!); Van Stadensberg,

Zeyher 2318 (S!, SAM!).

Lebeckia leptophylla Benth. in Hook. Lond. J. Bot. 3: 359 (1844), synon. nov. Type—South Africa, 'Grassy subalpine situations near

Swellendam', Mundt 87 (K!, 5!).

5.5.7. Wiborgiella sessilifolia (Eckl. and Zeyh.) Boatwr. and B.-E.van Wyk, comb. nov. Acanthobottya sessilifolia Eckl. and Zeyh., Enum. Pl. Afr. Austr. 2:

193 (Jan. 1836). Lebeckia sessilifolia (Eckl. and Zeyh.) Benth. in Hook. Lond.

J. Bot. 3: 362 (1844). Type—South Africa, 'between Breede River and

Duyvelshoek', Ecklon and Zeyher 1344 [S!, lecto.; K!, SAM!, isolecto., designated by Dahlgren (1975)].

5.5.8. Wiborgiella sp. 1. Vlok 2045 (PRE!, 2 sheets).

5.5.9. Wiborgiella sp. 2. Baker 10407 (NBG!).

Acknowledgements

The authors thank the National Research Foundation (NRF) and the

University of Johannesburg for funding. Curators and staff of the herbaria are thanked for their kind hospitality during study visits, mainly in 2006 and 2007

(BM, BOL, GRA, J, K, NBG, P, PRE, S, SBT, UPS and WIND) or for making

29 specimens available on loan (BOL, GRA, NBG, P, PRE, S and WIND). We also thank Dr John Manning (Compton Herbarium) for the suggestion of the name for the new genus. Dr Hugh Glen (National Biodiversity Institute,

Durban) is thanked for translating the Latin diagnosis. We also wish to acknowledge the kind help of Victoria Papworth (Curator of the General

Herbarium, Department of Botany, Natural History Museum, London) in sending us high resolution images of specimens in the Banks collection.

References

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33 Figure captions:

Fig. 1. The "Cape group" taken from (a) a strict consensus tree of the combined analysis of ITS and rbcL data(no. trees = 560; tree length = 1473 steps; consistency index = 0.50; retention index = 0.86); and (b) a strict consensus tree of the combined analysis of molecular (ITS and rbcL) and morphological data for the tribe Crotalarieae from Boatwright et al. (in press; no. trees = 370; tree length = 1166 steps; consistency index = 0.53; retention index = 0.84). Bootstrap percentages are given above the branches and

Bayesian posterior probabilities below the branches. Grey dotted lines indicate alternative topologies in the Bayesian analysis.

Fig. 2. Transverse sections through the leaves of Lebeckia species, showing isobilateral (a, b, Calobota group) and dorsiventral (c, e, f, Viborgioides group) leaflet laminas and an acicular, terete, phyllode with a circular arrangement of palisade cells (d, Lebeckia s.s.). Note the presence of mucilage cells in the epidermis. (a) L. cytisoides; (b) L. pungens; (c) L. sessilifolia; (d) L. sepiaria;

(e) L. inflata; (f) L. mucronata.

Voucher specimens: (a) Boatwright et al. 107 (JRAU); (b) Boatwright et al. 106 (JRAU); (c) Van Wyk 2120 (JRAU); (d) Le Roux et al. 10 (JRAU); (e)

Johns 162 (JRAU); (f) V/ok 1726 (JRAU). Scale bar = 0.2 mm.

Fig. 3. Drawings of the calyces (vestiture not shown), keel petals and anthers of selected species of Lebeckia s.I., Spartidium saharae and Wiborgia humi/is.

Note the relative size of the carinal lobe of the calyx, the apex and vestiture of

34 the keel petal and the relative size of the carinal anther. (al—a3) W. humilis;

(bl—b3) L. mucronata; (cl—c3) L. inflata; (dl—d3) L. leipoldtiana, (el—e3) L. cytisoides; (fl—f3) L. pungens; (gl—g3) S. saharae; (hl—h3) L. sepiaria.

Voucher specimens: (al, a3) Van Wyk 3530 (JRAU); (a2) Boatwright et al., 216 (JRAU); (bl—b3) Stirton 10880 (JRAU); (cl—c3) Vlok et al. 2 (JRAU);

(dl—d3) Boatwright et al. 123 (JRAU); (el—e3) Boatwright et al. 107 (JRAU);

(fl) Taylor 9386 (NBG); (f2) Boatwright et al. 106 (JRAU); (f3) Van Wyk 3252

(JRAU); (gl—g3) Hill 1910 (K); (hl, h3) Barker 6515 (NBG); (h2) Le Roux et al. 24 (JRAU). Scale bars (for all calyces, all keel petals and all anthers, respectively) = 1.0 mm.

35 im Aspeffiwa Mewl, 1

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Fig. 1

36 Fig. 2

37 b 1 dl 1

a2 b2 c2

03 lAG43 b3 k_)(d) ççip d3

Fig. 3

ca Asparagus greseis 1 99 Aspelathua tow& 1 I Alothusma penguin 1.0 E Asparaihus finearts 2 Amersours muck& A66626666 camese Asparagus camose kapabthus gokrata AtVelattna remota& Aspalatims nuterantha Aspahrthus wiDdenewiana 6 —Asperses. bum* *rem shake Aspakethus confuse Amesstnu s &sato& =tsp. iandlotla A6PW6616, perfoiata subsp. phOpsli Aspalabrus crenate —Awake& a NM odes, late Aspatothua vermiculata AwsMinia 9ys01), Asparagus shawl subsp. cher& Aspabhus briploka subs?. lad& olis 52 Asperse,ea gamete Asparaffora he& subsp. he& 42ksPerarnus mambo I Aspidathus brads s. kmaterhus wedenewiana 66p6,6thus twee Asparagus, pasis1oba Asparagus confuse kk,basOie &sea 1 —AVekstnus pada. ...tap. WOO Mob* tempters 1 r 7-41110 WtsaIIM Mbarstere 2 `—Ametarbus aurora M.O. Self0011 Aspaleva parhybhe 65 I— W10001k robcordata 1 AspeWhin /oho 70 009 .-- /16666;6, obconlafa 2 fusca 1 EIMM"q* borgramorrop"'naPrra 21 .0.4. telnplers 1 1. 06 Wermata &curve& 140 Moms soiree 00 r—lebeckiainflata 2 1 &Mock& Mate 400.1:6• oacantata 1 1.0 ,— Lebec& a Mlle& 2 Lebeckia =crenae Labeekta manna& 03 I— Mb9,99moapoa I 60 Lebeckie pungens 1 Wiborpla ilcunrsta Lebeska pungens 2 • e 93 (0700 ie hurnilia 1 Lebec's& rnacrentha 1 a Lebeclaa IllaCrantne 2 137 L seed* Apipokillens 1 Lobed& psiloloba Lebecido sessollofie Labeclen ,mono 1 termer& SenCea 2 &beetle Weem' 1 Lebaskia sad.. 3 Lobos& Saba 2 &Meek&cyf &ord. 1 LebecIde moon. Lebeckia &hackles 2 Lebeskia muluflora 2 Lebodie pungerta 2 Lebeckia lotononeides Lebackle 0000000 2 Lebeskie halenbergensis 1 Lebeckia holenbergensis 2 &beck& mask. Lebockia spinaseens L•b•cocie sweep 2 Lebeckie cinema Spored.' saharae Lebeckle &bookies 2 sa Rob& acuminate 95 go Week's MM. Rafts memos,' Lebeckie Mortenciles Relnta spice& 194.o n Refs& *boss Lerma& nalenberpeners 2 Rafrria criffusa Lebadds 63 86 LikborgialMbargi',...b.d.khk.urn7p.w.iri 0 21. , Lebocki. c.eme 52 Spar idlun sellers. Lerwick. borneana 1 Lebaskie &mesas& 2 Lobed& came= 2 ,— &Mackie leipoldtiona 2 Lastlia rneyerlans 0.99 1-- Lebedda sossillfolie iq IE Lebockla pktkeneflans 1 Lebeckle amass 1 —.."—dLebeekia samosa Lebo:kit peudnota 2 beie Lebec. iwightli 1 abeckia=e 'thtlal 1 Lebeckre breakers* 103. Lebodri6 pauciffora 1 Labeckia peueillera 2 Labeckbr emblgue 1 1,162= Lebeckla *21%.2 '1 100 E Wed* sepa/As , 1.0 Lebodda ample& 3 Rafts acuminate Labackia plukenetlana 2 69I— C Rah. mamma Lo ambigua 1 ee Le &sarcoma HLE RIJN. spkate Le ambigua 2 Rafts globose Le &mho& 1 Le &ORM 2 Rah. diffuse

Fig. 1

36 Fig. 2

37 al

a3 d3

93 h3 f3 AD 5R

Fig. 3

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