ISSN 0006 8241 = Botha! ia

Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 18,1 May/Mei 1988

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BOTHALIA

Bothalia is named in honour of General Louis Botha, first Premier

and Minister of Agriculture of the Union of South Africa. This

fmuse journal of the Botanical Research Institute is devoted to

the furtherance of botanical science. The main fields covered

are taxonomy, ecology, anatomy and cytology. Two parts of the

journal and an index to contents, authors and subjects are publish-

ed annually.

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0001, Republiek van Suid-Afrika. ’n Geldige lys van alle beskik-

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Bothalia is vernoem ter ere van Generaal Louis Botha, eerste Eerste

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sitologie. Twee dele van die tydskrif en ’n indeks van die inhoud,

outeurs en onderwerpe verskyn jaarliks.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

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botany. onderwerpe.

THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA

This serial presents colour plates of African plants with ac-

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FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA

A taxonomic treatise on the flora of the Republic of South Africa,

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West Africa/Namibia, Botswana and Venda. The FSA contains

descriptions of families, genera species, infraspecific taxa, keys

to genera and species, synonymy, literature and limited specimen

citations, as well as taxonomic and ecological notes. Also available

in the FSA series are the following:

’n Taksonomiese verhandeling oor die flora van die Republiek

van Suid-Afrika Ciskei, Transkei, Lesotho, Swaziland, Bophu-

thatswana, SWA/Namibie, Botswana en Venda. Die FSA bevat

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aantekeninge. Ook beskikbaar in die FSA-reeks is die volgende:

The genera of southern African flowering plants by/deur R.A. Dyer, Vol. 1 Dicotyledons (1975); Vol. 2

Monocotyledons (1976).

Keys to families and index to the genera of southern African flowering plants by/deur R.A. Dyer (1977).

Plant exploration of southern Africa by Mary Gunn & L.E. Codd. Obtainable from/Beskikbaar van:

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form of tables and photographic plates depicting fossil popula-

tions. Now available:

’n Palaeoflora met ’n uitleg vergelykbaar met die van die Flora

van suidelike Afrika. Baie van die inligting word aangebied in die

vorm van tabelle en fotografiese plate waarop fossiele populasies

afgebeeld word. Reeds beskikbaar:

Molteno Formation (Triassic) Vol. 1 Introduction. Dicroidium by/deur J.M. & H.M. Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous by/deur J.M. & H.M. Anderson.

Obtainable from/Beskikbaar van: A. A. Balkema Marketing, Box/Posbus 317, Claremont 7735, RSA.

Republic of

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Republiek van

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BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 18,1

Editor/Redakteur: O. A. Leistner

Assisted by B. A. Momberg

Editorial Board/Redaksieraad

D. F. Cutler

B. de Winter

P. H. Raven

J. P. Rourke

M. J. Werger

Royal Botanic Gardens, Kew, UK

Botanical Research Institute, Pretoria, RSA

Missouri Botanical Garden, St Louis, USA

National Botanic Gardens, Kirstenbosch, RSA

University of Utrecht, Utrecht, Netherlands

Editorial Committee/Redaksiekomitee

B. de Winter

D. J. B. Killick

O. A. Leistner

B. A. Momberg

M, C. Rutherford

J.C. Scheepers

ISSN 0006 8241

Issued by the Botanical Research Institute, Department of Agriculture and Water Supply. Private Bag X101 , Pretoria 0001

South Africa

IJiteeeee deur die Navorsingsinstituut vir Plantkunde, Departement van Landbou en Watervoorsiening, Privaatsak X101

Pretoria 000 1 , Suid-Afrika

1988

'

This volume is dedicated to

ROBERT ALLEN DYER

1900-1987

CONTENTS — INHOUD

Volume 18,1

1. Studies in die Ericoideae (Ericaceae). VI. The generic relationship between Erica and Philippia in

southern Africa. E. G. H. OLIVER 1

2. A synopsis of the tribe Desmodieae (Fabaceae) in southern Africa. B. D. SCHRIRE 11

3. The naturalized species of Lupinus (Fabaceae) in southern Africa. C. H. STIRTON 25

4. Studies in the genus Lotononis (Crotalarieae, Fabaceae). 1 . Three new species of the section Aulacinthus

from the Cape Province. B-E. VAN WYK 31

5. Studies in the genus Riccia (Marchantiales) from southern Africa. 8. R. campbelliana (subgenus Riccia),

newly recorded for the region. S. M. PEROLD and O. H. VOLK 37

6. Studies in the genus Riccia (Marchantiales) from southern Africa. 9. R. nigrella and the status of R.

capensis. S. M. PEROLD and O. H. VOLK 43

7. Two closely related species of Caloplaca (Teloschistaceae, Lichenes) from the Namib Desert. I. KARNE-

FELT 51

8. The Cheilanthes hirta complex and allied species (Adiantaceae/Pteridaceae) in southern Africa. W. B. G.

JACOBSEN and N.H.G. JACOBSEN 57

9. Solarium (Solanaceae) in Ghana. Z. R. BUKENYA and J. B. HALL 79

10. Notes on African plants:

Acarosporaceae. Lithoglypha, a new lichen genus from Clarens Sandstone. F. BRUSSE 89

Adiantaceae/Pteridaceae. Doryopteris pilosa, a new record from South Africa. W. B. G. JACOB-

SEN and N. H. G. JACOBSEN 90

Lamiaceae. Resuscitation of Syncolostemon ramulosus E. Mey. ex Benth. L. E. CODD 92

Porpidiaceae. A new species of Porpidia from the Drakensberg. F. BRUSSE 93

Porpidiaceae? Schizodiscus, a new porpidioid lichen genus from the Drakensberg. F. BRUSSE 94

11. The cigarette beetle Lasioderma serricorne (F.) (Coleoptera: Anobiidae): a serious herbarium pest.

E. RETIEF and A. NICHOLAS 97

12. Leaf anatomy of the South African Danthonieae (Poaceae). XVI. The genus Urochlaena. R. P. ELLIS .... 101

13. Ecology and population biology of Euphorbia perangusta (Euphorbiaceae) in the Transvaal, South Africa.

P. A. RAAL 105

14. Miscellaneous notes:

Chromosome studies on African plants. 6. J. J. SPIES and H. DU PLESSIS 1 1 1

Chromosome studies on African plants. 7. J. J. SPIES and S. P. VOGES 114

Chromosome studies on African plants. 8. H. DU PLESSIS and J. J. SPIES 119

New programs for preliminary sequencing of releves and species in phytosociological data sets. R. H.

WESTFALL and B. C. DE WET 122

15. The identification of ‘Isicakathi’ and its medicinal use in Transkei. R. N. BOLOFO and C. T. JOHNSON 125

16. Obituary: Robert Allen Dyer (1900 -1987). L. E. CODD 131

17. Book reviews 135

1 8 . Guide for authors to Bothal ia 137

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume1818unse

Bothalia 18,1: 1-10(1988)

Studies in the Ericoideae (Ericaceae). VI. The generic relationship

between Erica and Philippia in southern Africa

E. G. H. OLIVER*

Keywords: Africa, bract, bracteoles. Erica, Ericaceae, Ericoideae, Philippia, poly phy lesis , recaulescence, taxonomy

ABSTRACT

Problems in placing certain species satisfactorily in Philippia or Erica have led to an investigation of characters

delimiting the genera mainly in southern African species. The only character used in placing problem species was partial

versus total recaulescence of the bract. This separation breaks down completely in a few species in which ericoid and

philippioid flowers occur within the same inflorescence. Not all species of Philippia are closely related, some being more

closely related to various sections within Erica. It is evident that Philippia is an unnatural polyphy letic group It is

concluded that Philippia should be placed in synonymy under Erica.

UITTREKSEL

Probleme met die bevredigende plasing van sekere spesies in Philippia of Erica het gelei tot 'n ondersoek na die

kenmerke wat die genusse hoofsaaklik by spesies in suidelike Afrika omgrens. Die enigste kenmerk wat gebruik is om

probleemspesies te plaas, is gedeeltelike teenoor totale rekaulesens van die skutblaar. By ’n paar spesies met erikoiede en

philippioiede blomme in dieselfde bloeiwyse, verval hierdie onderskeid egter heeltemal. Nie alle spesies van Philippia is

naverwant nie; sommiges is nader verwant aan verskeie seksies van Erica. Philippia word beskou as ’n onnatuurlike

polifiletiese groep. Daar word voorgestel dat Philippia onder Erica in sinonimie geplaas word.

CONTENTS

Introduction 1

Morphology 2

Analysis 6

Philippia 6

Erica 6

Species outside southern Africa 8

Phylogeny of the genus Philippia 8

Conclusions 9

Acknowledgements 9

References 9

INTRODUCTION

Philippia is a widespread genus of ± 65 species native

to Africa and the Madagascan region. Erica occurs from

the Cape to northern Europe and consists of ± 660 spe-

cies, 98 % of which occur in the Cape. Work in progress

on the genera of the subfamily Ericoideae for the Flora

of southern Africa has revealed problems of generic deli-

mitation within this highly diversified group of approxi-

mately 800 species. One such problem area lies between

the extremely large genus Erica and the small genus

Philippia , with 636 and 15 species respectively within

the southern African region.

The genus Philippia was described by Klotzsch (1834)

who distinguished it from Erica on the possession of an

unequal calyx. Ever since then the genus has been uni-

versally upheld. Philippia is a pan- African genus of

* Botanical Research Unit, P.O. Box 471, Stellenbosch 7600.

MS. received; 1987.08.17

some 65 species with the majority (± 40 species) occur-

ring on Madagascar (Figure 1 ).

Erica is distinguished from Philippia by having a

single bract, two bracteoles and a calyx consisting of

four more or less equal segments or lobes. This applies

to the vast majority of the ± 660 species in the genus.

Philippia, on the other hand, is generally described as

having ‘no bract nor bracteoles and a zygomorphic 4-

partite or 4-lobed calyx’, a condition found in most of

the species in the genus.

The first indication of any problems regarding the dis-

tinction between Philippia and Erica was given by Aim

& Fries (1927a: 8) in the only complete revision of the

genus Philippia. They noted ‘In der Tat finden wir bei

der Gattung Erica noch derartige Ubergangsformen',

and further on (p. 36) ‘Diese beiden Arten [i.e. P. ken-

iensis S. Moore and P. jaegeri Engl.] stellen darum fast

Ubergangstypen zwischen den Gattungen Philippia und

Erica dar.’. Nevertheless, in this revision and the revi-

sion of the genus Erica in tropical Africa (Aim & Fries

1927b) they retained both genera. No problems were

recorded in distinguishing Philippia from Erica in

southern Africa until the present revisionary work during

which species were carefully examined and evolutionary

relationships were sought within Erica. Investigations

have revealed a complete series of intermediates within a

few species.

In the African Ericoideae only Erica, Philippia, Blae-

ria L. and Ericinella Klotzsch possess a multiseeded

dehiscent capsule. The genus Philippia seems likely to

have evolved from ancestral stock related to Erica sec-

tion Arsace (Aim & Fries 1927a; Pichi-Sermolli & Hei-

niger 1953), and the majority of species in southern

Africa are related to this section. However, two species,

P. stokoei and P. pallida, are clearly allied to Erica

species in other sections.

2

Bothalia 18,1 (1988)

FIGURE 1 . — Maps showing the approximate distribution of the genera Erica , left, and Philippia, right.

MORPHOLOGY

Aim & Fries (1927a) pointed out that in Erica and

Philippia the bract is recaulescent on the pedicel (Figure

2). This is a well known occurrence in the Angiospermae

(Weberling 1981: 228). However, Aim & Fries postu-

lated that the bract fused with the abaxial sepal — ‘Wir

halten es fur sehr wahrscheinlich . . . [dass] die unpaa-

rige untere Bractee . . . mit dem vorderen Kelchblatte

verschmolzen fist]’ (their floral diagram, 3.c, clearly

shows this feature). They continued ‘Denkbar, aber sehr

unwahrscheinlich ist, dass das vordere Kelchblatt

vollstandig reduziert worden ist.’. Pichi-Sermolli & Hei-

niger (1953: 35), on the other hand, referred to the recau-

lescent bract as the fourth calyx segment, which in Phi-

lippia is replaced by the bract which has undergone a

‘phyletic slide’ upwards and has become part of the ca-

lyx. Ross (1957: 735) found clear evidence that in Phi-

lippia at least, the zygomorphic calyx has arisen by the

migration of bracts up the pedicel, followed by their

fusion with the calyx, but he did not give any evidence to

support his view of fusion. Friedmann (1981: 6) held the

same opinion as Pichi-Sermolli & Heiniger in his revi-

sion of the Ericaceae in the Flore des Mascareignes hav-

ing found that some abnormal flowers in P. abietina

Klotzsch show transitions indicating that the major sepal

is in fact the modified floral bract and later on (p. 8) Me

sepale destine a etre remplace par la bractee a deja re-

gresse totalement’.

I would support the opinion of Pichi-Sermolli & Hei-

niger and of Friedmann that the totally recaulescent bract

does suppress the abaxial sepal and takes over its posi-

tion and ‘function’. No anatomical investigations have

been carried out to support this view but I have observed

morphological variation which suggests it, particularly

in E. peltata Andr. in which there may sometimes be a

thin, hair-like abaxial sepal on the inside of the recaules-

cent bract (Figure 5.4). A similar situation was noted by

Pichi-Sermolli & Heiniger in their P. abyssinica. In

another genus, Coilostigma (C. glabrum Benth.), va-

rious stages in reduction of the abaxial sepal are often

ericoid

ericoid/philippioid

intermediate

rB4, Obr

philippioid

FIGURE 2. — Series of diagrams

showing stages in the recaules-

cence of the bract up the pedicel

in the Ericoideae from axial on

the left to totally recaulescent on

the right (formulae represent

each stage and are explained in

the text).

aB , 2 br

rB2 , 2 br 3

rB4, 2 br5

RB , 0 br

Bothalia 18,1 (1988)

3

FIGURE 3. — Diagrams showing the

arrangement within an inflores-

cence of, left: typical ericoid

flowers; right: typical philip-

pioid flowers and, centre: the

intermediate form occurring fre-

quently in E. ebracteata and E.

peltata (formulae represent the

bract and bracteole arrangement

in each whorl as explained in the

text).

ericoid

philippioid

clearly visible. In its calyx, this is the most variable

species in the subfamily as it often has the adaxial sepal

reduced as well (Oliver 1987a).

Additional evidence to support this opinion was noted

by Pichi-Sermolli & Heiniger (1953: 35) in their P. abys-

sinica, which they stated exhibits a very characteristic

dimorphism of the calyx. The peripheral flowers of an

inflorescence have a regular calyx and a single foliar

bract lower down on the pedicel whereas the central or

upper flowers have a zygomorphic calyx with a totally

recaulescent bract. There is thus a complete gradation

series in the recaulescence of the bract within a single

inflorescence. I have found this gradation occurring in a

number of inflorescences of E. ebracteata H. Bol. from

the Natal Drakensberg, of E. peltata from the southern

Cape Province (Figure 3) and frequently in Coilostigma

glabrum (Oliver 1987a).

It has always been stated that, when the bract is totally

recaulescent, the two bracteoles (prophylls) are missing

or absent. Nobody has queried this statement or dis-

cussed the fate of these two organs. In my investigations

of E. peltata I have come to the conclusion that the

bracteoles do not disappear but also become part of the

calyx. A paper on the fate of the bracteoles is in prepara-

tion (Oliver in prep. b).

In the typical philippioid condition the bract is fully

recaulescent up the pedicel, totally suppressing the ab-

axial sepal and fusing with the calyx, in some cases to

such an extent that it is difficult to distinguish it as not

being a sepal, as in P. simii H. Bol., P. irrorata E. G.

H. Oliver and P. procaviana E. G. H. Oliver

(Figure 4. 1-4.3). Ross (1957: 738), however, stated

that in the east African species with flowers of the philip-

pioid type, the bract-like calyx lobe is always free to the

base and never fused with the others. He added that it is

never inserted at the same level as the other segments but

is always at least slightly lower. This occurs in the sou-

thern African P. stokoei L. Guthrie, P. drakensbergensis

E. G. H. Oliver, P. pallida L. Guthrie and P. alticola E.

G. H. Oliver (Figure 4. 4-4. 7). On the other hand, the

bract is often slightly to considerably larger than the

other lobes of the calyx resulting in an unequal or zygo-

morphic calyx (Figure 5.1— 5.3, P. evansii N.E. Br.).

This condition is found in a number of genera in the

Ericoideae other than Philippia : Ericinella , the Salaxis

Salisb .-Coccosperma K\otzsch-Scyphogyne Brongn.

complex, Coilostigma Klotzsch and the unique monoty-

pic Nagelocarpus Bullock. Except for the single species

Ericinella microdonta (C.H. Wr.) Aim & Fries, these

genera are all Cape endemics.

It is therefore clear that the abaxial ‘sepal’, which is

often enlarged and separate from the rest of the calyx in

the species of Philippia and also of Ericinella, Coilo-

stigma, Salaxis, Coccosperma, Scyphogyne and Nagelo-

carpus, is homologous with the bract in Erica. The bract

thus exhibits three character states: nonrecaulescent

(axial), partially recaulescent and totally recaulescent.

In the distinction between Philippia and Erica, Aim &

Fries (1927a: 36) made use only of the actinomorphic

versus zygomorphic calyx and glossed over those species

that have some flowers with the intermediate condition.

Phillips (1944) re-examined the Ericaceae for the revi-

sion of his The genera of South African flowering plants

(1951) and came to the conclusion that not more than

eight genera could be justified. He sank numerous ge-

nera into what I regard as heterogeneous assemblages.

Philippia was combined with Blaeria and Ericinella, a

very valid step, but was kept separate from Erica,

whereas Thamnus and Coccosperma were included

solely on the basis of ovary and stamen complement.

The last two are, however, very distinct unrelated genera

with indehiscent fruits. Phillips did not consider there to

be any close relationship between Erica and Philippia.

His work was never seriously considered by subsequent

workers and was not taken up in herbaria.

Ross (1957), in preparing the account for the Flora of

tropical east Africa, published some detailed notes on

the genus Philippia in which he not only dealt with the

taxonomic problems of species restricted to that area, but

he also considered generic limits within the whole tribe.

He pointed out several problems, but under the section

dealing with the concept of the genus he did not discuss

the relationship of Philippia and Erica. He retained them

as distinct genera and in his key gave the following dif-

ferentiating characters:

4

Bothalia 18,1 (1988)

FIGURE 4. — Flowers of Philippia showing the bract (on left hand side of each flower) totally recaulescent and fused to lateral sepals in 1,

P. simir, 2, P. irrorata; 3, P. procaviana\ showing the bract (on left hand side of each flower) almost totally recaulescent and free from

lateral sepals in 4, P. stokoei\ 5, P. drakensbergensis ; 6, P. pallida ; 7, P. alticola, all drawn x 20 from type material (BOL, STE).

‘Pedicels with one or more bracts and calyx actinomorphic in

all flowers Erica

Pedicel without bracts and calyx with one sepal larger than the

others in many or all flowers Philippia ’

However, he did mention the problem of the placing

of certain specimens of P. trimera Engl, and P. keniensis

S. Moore which exhibit variation in the bract/calyx

relationship (see later in this article).

For the species occurring in the area covered by the

Flora Zambesiaca there appear to be no problems with

generic distinctions, as Ross (1983: 157) mentioned

none under the generic or specific treatments. However,

in the key he noted for Erica, ‘bract, and often brac-

teoles, present on pedicel’, which indicates that he re-

garded material with only the partially recaulescent bract

present as belonging to Erica, but in his earlier work

(1957: 38) he regarded material having equal proportions

of one bract and no bract as belonging to Philippia. He

noted that all the populations having these proportions

also agree with all other species of Philippia in having a

disciform stigma and anthers without appendages, and

furthermore, the anthers are united before dehiscence

and united anthers occur in a number of species of Phi-

lippia. He thus implied that these were characters he

used to define the genus.

Pichi-Sermolli & Heiniger, in upholding Philippia for

their highly variable species P. abyssinica, stated (1953:

35) that, even if typical actinomorphic flowers are found

in the Abyssinian plant, the presence of zygomorphic

flowers and, at the same time, the shape of the terminal

part of the style head ‘lead us confidently to refer this

species to the genus Philippia' .

The use of the above additional characters to distin-

guish Philippia from Erica is rejected on the grounds

that they also occur within the genus Erica. The enlarged

terminal portion of the style is an adaptation to wind

pollination (Rebelo et at. 1985) and occurs in 5% of the

Bothalia 18,1 (1988)

5

FIGURE 5. — Rowers of Philippia (with bract on left hand side of each flower): 1 , 2 & 3 showing variation in size of the totally recaulescent

bract in P. evansii from (1) Killick 1527 , (2) Edwards 2163 , (3) an isotype, Evans 62 (all PRE); 4, showing a partially recaulescent bract

in position rB? (hatched) removed to reveal the reduced and hair-like abaxial sepal in E. peltaia from Oliver 8815 (STE), (compare with

Figure 4.6); 5, showing a partially recaulescent bract in the position rB:, and with the abaxial sepal suppressed, in P. irrorata from the

type (STE); 6, showing recaulescence of the bract up the pedicel in an unusual flower of P. drakensbergensis from Oliver 8394 (STE); all

drawn X 20.

426 species of Erica occurring in the south-western Cape

Province. This pollination syndrome of small flowers, a

large cyathiform to peltate stigma and the absence of

conspicuous nectaries, although not restricted to the sec-

tion Arsace , is typical of that section. It is also found in

all species of the genera Salaxis , Coccosperma , Scypho-

gyne and Nagelocarpus.

In all species of Ericoideae the anthers are united be-

fore anthesis and in the omithophilous and entomophi-

lous species often remain adhering until disturbed by the

pollinator when an explosive separation causes the pol-

len to be dispersed (Rebelo et al. 1985). There are a few

species of Erica , notably E. embothriifolia Salisb. (with

long tubular flowers), E. coronanthera Compton and

E. syngenesia Compton, in which the anthers remain

fused together. Within the genus Philippia , Aim & Fries

(1927) list 17 of the 40 species as having free anthers. In

southern Africa 12 of the 15 species have free anthers

although some have filaments slightly joined at their

bases.

The above additional characters, used to justify the

separation of variable intermediate specimens, are there-

fore of no value in generic delimitation between Erica

and Philippia.

A detailed examination of bract recaulescence in the

southern African species was undertaken to ascertain the

situation prevailing in the genus there and to assess the

validity of the generic boundaries.

A formula has been developed to express in short

simple terms the number and arrangement of the bract,

bracteoles and calyx within the genera and species of

Ericoideae: B represents the bract; br the bracteoles; a

6

Bothalia 18,1 (1988)

indicates that the bract is axial (situated on the main

axis); r, that the organs are partially recaulescent with

the indices15 indicating the position up the pedicel (1 at

the base, 5 at the top); and R, fully recaulescent; K

represents the calyx. Thus the structure of a typical Erica

(ericoid) flower can be indicated by the formula

aB-rB' \2br \4K and that of a typical Philippia (phi-

lippioid) flower by the formula RB,0br,3K or just

RB,3K (Figures 2 & 3). This is an elaboration of the

‘formula’ briefly mentioned by Ross (1957: 738).

The question arises whether any deviations from these

basic formulae exist. Indeed, within Philippia some

flowers occur in which the bract may suppress the ab-

axial sepal but is free from the calyx (Figure 4.6, P.

pallida) and even as low down the pedicel as rB\3K

(Figure 5.4, P. irrorata)', some flowers may even be

rB3_5,4K which also occurs in some species of Erica (see

later). This is clearly an intermediate Erica! Philippia

condition. A detailed analysis of these bract/bracteole-

/calyx conditions in Erica and Philippia was made and

the results are discussed below.

ANALYSIS

Philippia

The majority of species occurring within southern

Africa are typical of the genus in having a fully recaules-

cent bract forming a zygomorphic calyx represented by

RB,3K; these are P. leeana Klotzsch, P. absinthoides

(Thunb.) E. G. H. Oliver, P. evansii, P. notholeeana E.

G. H. Oliver, P. elsieana E. G. H. Oliver, P. irrorata ,

P . procaviana, P. alticola, P. petrophila E. G. H.

Oliver, P. esterhuyseniae E. G. H. Oliver and P.

drakensbergensis E. G. H. Oliver. This formula does

hold in all except a few flowers. The remaining three

species, P. pallida, P. stokoei and P . tristis, showed

highly significant variations (Table 1).

P. pallida L. Guthrie

Until recently this species was known only from the

type collection from near Riversdale ( Muir 2542 in

BOL*) and this is unfortunately in the bud stage. The

taxon is not closely related to any species within Philip-

pia. During an examination of the type it was recognized

as being identical to Erica peltata Andr. which occurs

along the foothills of the mountain ranges from Rivier-

sonderend to George. An analysis of 269 flowers from a

selection of 15 collections of this species in STE pro-

vided a ratio of 22% ericoid: 38% ericoid/philip-

pioid: 40% philippioid flowers. An analysis was then

made of 900 flowers from three well separated popula-

tions (Oliver 8736, 8737 & 8815 in STE). These showed

considerable variation with the above ratios being

16:40:40, 3:30:67 and 0:38:68 respectively. In the last

population one would probably not hesitate to label the

plants as belonging to P. pallida. The various flower

types were found to be randomly scattered in inflores-

cences, occasionally with all three represented in one

inflorescence.

P. stokoei L. Guthrie

This species is very localized on the low mountains

and hills of the western part of the Bredasdorp District. It

* Abbreviations follow those listed in Holmgren, Keuken & Scho-

field, Index Herbariorum, Regnum Vegetabile 106 (1981).

has no close relatives within Philippia but was found to

have been described as E. accommodata Klotzsch var.

ebracteata H. Bol. (Guthrie & Bolus 1905). Material of

the species was thus filed under two separate genera in

herbaria. The taxon bears some resemblance to E. ac-

commodata Klotzsch, which inhabits the high mountains

of the Riviersonderend Range. It is, however, remark-

ably similar to E. lasciva Salisb. of the Cape Town and

Bredasdorp Districts, which species shows no signs of

any bract/bracteole irregularities. An analysis of several

collections of P. stokoei (250 flowers) in STE provided a

ratio of 0:50:50 with not a single flower having the eri-

coid condition. The only difference between P. stokoei

and E. lasciva is in the absence of bracteoles versus two

bracteoles, otherwise the two taxa are identical in shape,

size, indumentum and texture of the flowers and vegeta-

tive parts.

P. tristis H. Bol.

This species was known only from the type collection

made by Bolus in 1877 in the mountains near Graaff-

Reinet in the central Karoo ( Bolus 2594 in BOL, K). The

specimen in BOL has flowers that are in fruit, but it can

be seen that they are mostly of the RB,3K type with a

few rB,0br,4K. A capsule included on the type sheet in

BOL containing a very small specimen collected farther

east by Weale, appeared to be identical but had flowers

of the rB,0br,4K type which would indicate its being an

Erica/Philippia intermediate following the definition as

given above.

I was sent for comment, several new taxa of Erica

from the north-eastern Cape and Natal by Dr O. M.

Hilliard and Mr B. L. Burtt who intended describing

them for their work on the flora of the southern Drakens-

berg. The material of P. tristis was recognized as being

very similar to one of their species, E. caespitosa Hil-

liard & Burtt. This problem will be discussed later under

Erica.

Erica

E. hispidula L./E. inops H. Bol.

E. hispidula is probably the commonest species of

Erica in the Cape Floral Region occurring from the Cape

Peninsula eastwards to the George District and forming

dense stands on mountain slopes. A survey of 1000 flow-

ers from numerous collections (BOL, NBG, STE) cover-

ing the distribution range east of the Cape Peninsula

provided convincing evidence that it belonged to Erica:

it has a ratio of 100:0:0, i.e. all flowers examined were

of the rB'-^br^K type.

There is, however, a problem on the Cape Peninsula

where the species E. inops occurs. This species was

separated from E. hispidula (Guthrie & Bolus 1905;

Salter 1950) solely because of its possession of a bract

but no bracteoles, i.e. it had the formula rB,0br,4K, the

ericoid/philippioid condition. The problem populations

exist on Constantiaberg. Here an examination of E. his-

pidula ( Baker 992 in BOL) showed that the flowers had a

formula rB1-3 and 2br3“5 or 1 br4^5 or even Obr, all with

4K; 1 10 flowers from all the other specimens had a ratio

of 55:45:0. A collection of E. inops ( Baker 791 in BOL)

had the ratio 0:23:77 and 291 flowers from other collec-

tions 87:13:0, suggesting that some of them were E.

7

Bothalia 18,1 (1988)

hispidula. An examination of 1370 flowers of my own

collection on Constantiaberg (Oliver 8741 in STE) pro-

duced the ratio 3:96:1. It has been concluded from these

findings that the two species should be merged.

E. ebracteata H. Bol.

The very name of this species points to the generic

problem. It is a species confined to the Natal Drakens-

berg. A survey of 238 flowers from 15 specimens (PRE)

gave a ratio of 1 1 :46:43 similar to that found in E. pel-

tata and indicates that 43% of the flowers on the collec-

tions were in fact philippioid flowers and 46% interme-

diate and only 1 1% ericoid. In this species the variation

from partially to totally recaulescent bracts often occurs

within a single inflorescence (Figure 3).

Erica taxa described by Hilliard & Burtt (1985)

In personal communications with the authors, prior to

the publication of their paper, I pointed out the problem

existing between Erica and Philippia, which was high-

lighted by several of their new taxa (see also Hilliard &

Burtt 1985). I have been able to examine their cited

material and some additional collections.

E. caespitosa Hilliard & Burtt

This species is widespread in the eastern parts of the

country from Somerset East in the Karoo region to the

Drakensberg of the north-eastern Orange Free State. A

survey of 256 flowers from 10 collections (BOL, NU,

STE) gave a ratio of 3:83:14. The philippioid condition

was confined to a few scattered collections, one of

which, Ruddock 55 in NU, had only RB,3K flowers.

This latter collection possessed flowers identical to those

of P. tristis mentioned earlier.

E. anomala Hilliard & Burtt

The specific epithet was chosen to indicate that the

flowers were not typical of Erica. A survey of 100 flow-

ers from 5 collections in NU gave a ratio of 0:40:60

indicating that the majority were of the philippioid type.

The species is in fact no more anomalous than the others

referred to in the present paper. It is also peculiar in

having coherent anthers which are exserted in a ring

around the style in the fruiting stage.

E. dissimulans Hilliard & Burtt

Like E. anomala this species is confined to the Natal

Drakensberg. A survey of 160 flowers from 7 collections

in BOL and NU gave a ratio of 2:98:0.

Flora capensis

A survey of the descriptions of the species of Erica

listed in Flora capensis (Guthrie & Bolus 1905) was

made to see if there were any other species with similar

irregularities. (Note that in that work no distinction is

made between the bract and the bracteoles, all being

referred to as bracts).

E. alticola Guth. & Bol.

This very rare species, probably confined to the east-

ern Transvaal near Barberton, is described as having

‘bracts variable, mostly entirely absent, occasionally 2,

sometimes one only’. An examination of the type collec-

tion ( Bolus 7678 in BOL) showed that the flowers are

very variable, most of the visible ones having the for-

mula aB,0br,3K. Two flowers have aB,2br\4K and a

few have rB2~\0br,4K or rB\0br,4K-RB,3K. Too few

flowers were available for this species to be included in

Table 1 .

TABLE 1. — Distribution of bract/bracteole/calyx characters

in nine species of Erica (Ericoideae) in southern Africa*

* Figures express the percentage of flowers in the samples re-

ferred to in the text and Table.

** E. hispidula , sensu stricto, refers to the non-varying popula-

tions outside of the Cape Peninsula; Constantiaberg refers to

the collections from that mountain.

t E. inops : collections Oliver 8741, Baker 791 and all other

collections.

$ E. peltata (= P. pallida)-, collections from Ashton (Oliver

8815), Swellendam (Oliver 8737) and Niekerkshek (Oliver

8736) and 15 other collections from the complete range of

the species.

E. woodii H. Bol.

This widespread species in the eastern parts of the

country is described as having ‘flowers mostly axillary,

solitary, bracts remote, small, occasionally one want-

ing’. There is in fact always one bract and two brac-

teoles, but the bract is large and axial in position appear-

ing like a normal leaf, thus the formula is aB,2br^,4K.

E. leucopelta Tausch

This similarly widespread species is described with

‘bracts remote, mostly 3, one rather large and 2 usually

small or very minute, or perhaps sometimes wanting’.

The bracteoles are always present and the bract is either

aB or rB1.

E. subverticillaris Diels ex Guth. & Bol.

This species is very rare and known from only two

collections in the Drakensberg of the eastern Transvaal.

It is described as having ‘bracts, one foliaceous, two

minute or often wanting’. Davidson (1985) in separating

off var. revoluta as a distinct species, E. revoluta (H.

Bol.) Davidson, noted that E. subverticillaris has ‘brae-

8

teoles 2, or absent, basal to median, hairlike’. It there-

fore possesses flowers with the intermediate ericoid/phi-

lippioid condition in addition to the normal ericoid ones.

E. sparsa Dulfer (=E. floribunda Lodd.).

This species is common on the coastal flats of the

southern Cape. No indication of any irregularities is

given, but during routine examination of herbarium ma-

terial it was found that variation does occur. Most speci-

mens appear to have rB:"\2br \ An examination of 63

flowers of Fourcade 5010 (STE) revealed that the collec-

tion possessed 56 flowers with the rB: \0br,4K condi-

tion, 2 with rB\2br4 and 5 with RB, whereas the 63

flowers of Keet 703 (STE) all had the rB4 \0br condi-

tion. This is very similar to the situation found in

E. peltata which is placed in the same section.

SPECIES OUTSIDE SOUTHERN AFRICA

A brief reference to material of Philippic i (BOL, MO

& PRE) and to literature on the Ericoideae in tropical

Africa (Aim & Fries 1927; Ross 1957; Pichi-Sermolli &

Heiniger 1953), in Madagascar (Perrier de la Bathie

1927) and on the Mascarene Islands (Friedmann 1981)

revealed a similar variation in the bract/bracteole/calyx

arrangement as outlined above for southern Africa, but

perhaps only to the same degree in the east African P.

keniensis S. Moore and E. kingaensis Engl.

Ross (1957; 739) gave a table analysing some 49 col-

lections of the former species and from this table the

ratio 0:34:66, of different flower types can be deduced,

the 34% having the formula rB?,0br,3K (position of

bract not given by Ross). The problem is also reflected in

the history of one species, first described as P. elgonen-

sis Mildbraed (Mildbraed 1922), transferred to Erica by

Aim & Fries (1927a) and back to Philippia (Ross 1957)

asP. keniensis. subsp. elgonensis (Mildbraed) R. Ross.

Ross also mentioned (1957: 738) an intriguing variant

of P. trirnera. He examined 39 gatherings of that species

from Ruwenzori and found no ericoid flowers. However,

he noted that in some collections, notably Fredericq

9325 , the ‘bract-like segment of the calyx’ was inserted

on the pedicel as much as 1 ,0 mm below the calyx which

was always of the 3K type. Sometimes he found ‘as well

as the normal ones (sepals) two narrower ones flanking

the gap on either side’. These in my opinion are the two

bracteoles giving the formula rB?,2br\3K (position of

bract not given by Ross) which is virtually the formula

for many species of Erica. In my investigations of E.

peltata I have found a similar situation. A paper setting

out my views on the fate of the bracteoles in flowers with

a totally recaulescent bract, derived from investigations

of this species, is in preparation (Oliver in prep.b).

In writing about the very widespread E. arborea and

its allies in Africa, Ross (1956) found that many of the

collections in east Africa, which had previously been

allocated to separate species, belonged to one highly

variable species, E. kingaensis. One of the variable char-

acters was the presence or absence of the two bracteoles.

He did not mention the problem of the relationship with

Philippia and retained the following under this species:

subsp. kingaensis — upper bracts often absent (rB,0-

2br);

Bothalia 18,1 (1988)

subsp. rugegensis — upper bracts always present (rB,

2br);

subsp. bequaertii — upper bracts present or absent (rB,

0-2br).

In the collections by Osmaston from Ruwenzori, Ross

noted that all the flowers examined lacked upper bracts.

Therefore some of the material of this species has flow-

ers with the same formula as those in P. keniensis , men-

tioned above.

PHYLOGENY OF THE GENUS PHILIPPIA

In the subfamily Ericoideae only Erica, Philippia,

Blaeria, Ericinella and Bruckenthalia Salisb. possess a

multiseeded dehiscent capsule. Erica is widespread in

Europe and Africa whereas Bruckenthalia is a monotypic

genus confined to the south-eastern highlands of Europe.

The other three genera are endemic to the African conti-

nent (including Madagascar for Philippia) where they

are widespread, more so than is Erica. The remaining 20

genera of the subfamily are confined to the southern tip

of Africa and are characterized by a variety of types of

indehiscent fruit coupled with a reduction in the comple-

ment of floral parts.

The relationships within this group of capsular genera

are very close, with generic distinctions being based on

the flimsiest of morphological characters. They indicate

that these genera have arisen from some ancestral ericoid

stock: Philippia by the total recaulescence of the bract;

Blaeria by the reduction in the number of stamens and

Ericinella by a combination of both processes.

In the southern African representatives of Philippia,

relationships for the majority of species occur within the

section Arsace of Erica. This section is found throughout

Africa and Europe and includes the most widespread

species of Erica, E. arborea L. , which is regarded (Han-

sen 1950) as the most primitive of the European species

based on architecture and inflorescence structure. The

section is characterized by the small coralline urceolate

to cyathiform flowers, mostly dull white or greenish in

colour, and a relatively large expanded and exserted

stigma which is developed for wind pollination. The

flowers being coralline means that the calyx is small,

relative to the corolla and is composed of foliar sepals.

These characters place the section in the subgenus Eu-

erica.

Two species of Philippia, P. stokoei and P. pallida,

have been found to be conspecific with two species of

Erica : P. stokoei with E. lasciva and P. pallida with E.

peltata. In these two taxa the calyces are different from

those in sect. Arsace: they are not small and foliar in

character. The two species are placed in different subge-

nera because of the shape of the corolla and form of the

calyx. E. lasciva is placed in the subgenus Chlamy-

danthe section Elytrostegia because of its calycine flow-

ers with the bracts and sepals relatively large and cartila-

ginous and the stamens exserted. E. peltata, on the other

hand, is placed in the subgenus Platy stoma section Poly-

codon which is characterized by broadly cyathiform co-

rollas and broader, more prominent concolorous sepals.

Both sections contain species adapted to wind pollina-

tion, i.e. with an expanded stigma (Rebelo et al. 1985). I

Bothalia 18,1 (1988)

9

consider the expanded stigma in these two species as the

result of convergent evolution in the pollination syn-

dromes.

Aim & Fries (1927b: 8) regarded section Arsace in

Erica as containing some species that can be described as

‘Ur-Philippien’ (proto-philippias). They stated that there

can be no doubt about the origin of Philippic/. However,

later on (1927: 1 1), in discussing the relationships within

the section Euphilippia , on the grounds that one cannot

see close relations between the species in the southern

and eastern regions of Africa and those of Madagascar

and the Mascarenes, they stated ‘kann es nicht ohne

weiteres angenommen werden, dass die Section einen

einheitlichen Ursprung habe’. They believed that the

Madagascan/Mauritian species and the African species

had a polyphyletic origin whilst the African species were

monophyletic, but earlier on the same page they stated

clearly that they could not find any close relationship

between the Madagascan/Mauritian species and the con-

tinental species nor between the tropical African and the

Cape species.

Pichi-Sermolli & Heiniger (1953: 36) supported the

hypothesis of Aim & Fries that Philippic/ ‘branched off’

from the section Arsace. They regarded their aberrant

ericoid P. abyssir/ica as the species most closely related

to Erica and, being at the northern limit of the genus

Philippic /, ‘the differentiation of Philippic/ .... took place

in the north .... rather than in the South African ones’ .

CONCLUSIONS

It is evident from the above investigation of southern

African species that the sole character for distinguishing

between the genera Erica and Philippic/ , the degree of

recaulescence of the bract, breaks down in a number of

species, more frequently, relatively speaking, in Philip-

pic/ than in the very large genus Erica (Table 1 ).

A ratio of 100:0:0 representing the aB-rB1 \2br' \4K

condition is characteristic of about 99% of the species of

Erica whereas a ratio of 0:0:100 or RB,3K (except for

the occasional aberrant flower) characterizes 12 of the 15

species of Philippic/ occurring in southern Africa. The

possession of flowers falling within the middle portion

(X) of the ratio 0:X:0 would place a specimen interme-

diate between the two genera with the formula

rB2~\0br,3-4K. From Table 1 it can be seen that only

two of the species discussed, E. dissimulans and E. his-

piclula sens, strict., tend to the ericoid side and one spe-

cies, E. anomala , tends towards to the philippioid side.

Six species, E. cc/espitosa (P. tristis ), E. hispiclulal

inops, E. ebractec/ta, E. lasciva (P. stokoei ), E. peltc/ta

(P. pallida) and E. sparsa, have ratios that make it im-

possible to place them satisfactorily in either genus.

It could be argued that a species should be placed in

Erica if it had more flowers that were typically ericoid

than typically philippioid, and vice versa. Thus from

Table 1 and previous discussion E. dissimulans and

E. subverticillaris could be retained in Erica whereas E.

cc/espitosa, E. anomala and E. ebractec/ta should be

transferred to Philippic/. This would require a thorough

analysis of all the flowers on any specimen before it

could be correctly identified. What then would be the

fate of E. alticola, E. hispidulal inops, E. lasciva (P.

stokoei), E. peltc/ta (P . pallida) and E. sparsa ? These are

five totally unrelated species each exhibiting a complete

range of variation from the typical ericoid flower to the

typical philippioid flower. They are all natural species

whose remarkable variability cannot be ascribed to hy-

bridization between species of Erica and Philippic/.

The problem is compounded by geographical con-

siderations because it is not confined to one small area.

This problem as discussed above occurs on the Cape

Peninsula, the southern Cape Province and on the Natal

Drakensberg in southern Africa as well as on the Masca-

rene Islands, the East African highlands and the island of

Sao Tome off West Africa. A polyphyletic origin for the

genus as presently construed is strongly indicated by this

evidence. The genus Philippia can therefore not be

upheld as a natural grouping of species.

I therefore propose to reduce the genus Philippia to

synonymy under Erica. This action will have far-reach-

ing nomenclatural repercussions in the rest of Africa and

Madagascar as pointed out by Hilliard & Burtt (1985:

240). The above decision is therefore not made without

careful consideration of and due regard for these reper-

cussions. A paper formalizing the above proposal and

providing the new combinations for the southern African

material is being published separately (Oliver 1987b).

ACKNOWLEDGEMENTS

I am grateful to the directors of BOL, MO, NBG, NU

and PRE for the loan of material, to Dr Olive M. Hilliard

(formerly of Pietermaritzburg, now Edinburgh) and Mr

B. L. Burtt (Edinburgh) for stimulating correspondence,

to Dr P. F. Stevens (Harvard) and Dr H, P. Linder (Cape

Town) for comments on drafts of this manuscript, and to

Dr L. J. Dorr (Missouri) for a week of fruitful dis-

cussions in Stellenbosch. This work forms part of doc-

toral studies being undertaken through the Bolus Herbar-

ium, University of Cape Town.

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SALTER, T. M. 1950. Ericaceae. In R. S. Adamson & T. M. Salter,

Flora of the Cape Peninsula. Juta, Cape Town.

WEBERLING, F. 1981. Morphologie der Bliiten und der Blii-

tenstdnde. pp. 391. Ulmer, Stuttgart.

Bothalia 18,1: 11-24(1988)

A synopsis of the tribe Desmodieae (Fabaceae) in southern Africa

B. D. SCHRIRE*

Keywords: Alysicarpus, Desmodieae, Desmodium, Fabaceae, key, Lespedeza, Papilionoideae, Pseudarthria, southern Africa, synopsis

ABSTRACT

The tribe Desmodieae has a pantropical distribution and is one of the most advanced tribes in the subfamily Papilionoi-

deae. Its greatest centres of development are in tropical Asia and America. Africa is relatively poorly endowed and only

four genera comprising 16 species occur in the flora of southern Africa. Many of these species are widespread in the Old

World tropics and the few African endemics appear to be closely related to them. A synopsis of the genera Desmodium,

Pseudarthria, Alysicarpus and Lespedeza is given for southern Africa.

UITTREKSEL

Die tribus Desmodieae het ’n pantropiese verspreiding en is een van die mees gevorderde tribusse in die subfamilie

Papilionoideae. Die grootste sentrums van ontwikkeling is in tropiese Asie en Amerika. Die groep is swak verteenwoordig

in Afrika en slegs vier genusse wat 16 spesies insluit, kom in die flora van suidelike Afrika voor. Baie van hierdie spesies is

wydverspreid in die Ou Wereldse trope en dit lyk asof die paar wat endemies in Afrika is, naverwant aan hulle is. ’n

Samevatting van die genusse Desmodium, Pseudarthria, Alysicarpus en Lespedeza word vir suidelike Afrika gegee.

INTRODUCTION

The tribe Desmodieae comprises 27 genera with a

mainly tropical distribution and is distinguished from the

other tribes with jointed indehiscent fruits by the pre-

sence of (1-) 3-foliolate leaves with stipels (Ohashi

1971a).

Three subtribes have been recognized in the latest

treatment of Ohashi, Polhill & Schubert (1981). The

largest and most polymorphic of these is the subtribe

Desmodiinae which is represented in southern Africa by

three genera, Desmodium, Pseudarthria and Alysicar-

pus. The much smaller subtribe Lespedezinae is only

represented in the flora by the naturalized bush-clover

genus, Lespedeza. The type genus, Desmodium, is the

most widespread genus in the tribe and with about 300

species (Ohashi etal. 1981) is also the most numerous. It

occurs widely in the tropical and subtropical regions of

America, Asia, Australia and Africa with two major

centres of diversification: firstly, in Mexico and Brazil,

where there is perhaps the largest number of species with

the most diverse development of plant parts (Schubert

1963); and then in Asia (the Sino-Indian region down

through Malesia to Australia) where there are large

numbers of species in many subgeneric groups. In addi-

tion the least specialized subgenera and the most archaic

genera in the tribe are found in the Asian region, indicat-

ing a possible centre of origin for the genus (Ohashi et

al. 1981).

Of the ten species of Desmodium found in southern

Africa most have an Asian origin, although there are a

few that are endemic in Africa. Some too, are natural-

ized introductions from the New World. Pseudarthria

with 4 to 6 species and Alysicarpus with 25 to 30 species

are in the main, tropical Old World genera. These are

* Botanical Research Institute, Private Bag X 10 1 , Pretoria 0001. Pre-

sently: South African Liaison Officer, Royal Botanic Gardens, Kew,

Richmond, Surrey, TW9 3AE, England.

MS. received: 1985.09.01.

represented in the flora of southern Africa by one and

four species respectively. Southern Africa contains the

tail-end of the tribe’s tropical African distribution as well

as a number of naturalized introductions. All specimens

examined are listed alphabetically according to the name

of the collector after the references at the end of the

article.

The Desmodieae is a tribe of annual herbs or, more

commonly, perennial suffrutices occupying, in southern

Africa, the southerly extension of the large Sudano-Zam-

bezian region of savanna and woodland areas of tropical

Africa, and to a lesser extent the Afromontane and

Indian Ocean coastal belt vegetation types of Werger

(1978). In general the tribe occurs in the warmer and

wetter parts of the Flora area.

TAXONOMIC BACKGROUND

In 1753 Hedysarum L. was published in Species plan-

tarum along with two species occurring in the Flora area,

H. gangeticum and H. vaginale. In 1759 H. barbatum

was published by Linnaeus in Systema naturae. From

then until the early 1800’s various authors described

other species in Hedysarum: Aublet (1775), H. racemo-

sum {—H. incanum)', Swartz (1788), H. adscendens and

H. tortuosum ; Vahl (1791), H. repandum and H. gluma-

ceum ; Willdenow (1802), H. velutinum and H. rugosum

and Poiret (1805), H. salicifolium.

It was recognized that several different genera existed

in Hedysarum and in 1803 Michaux described the genus

Lespedeza. Don transferred Anthyllus cuneata Du Mont,

to this genus in 1832. In 1813 Desvaux described Des-

modium and Alysicarpus and in 1825, A. P. de Candolle

placed all the above-mentioned species, with the excep-

tion of H. barbatum, into one or other of these genera.

A. P. de Candolle also described the genus Nicolsonia

into which Meyer (1836) placed two new species occur-

ring in the Flora area, N. caffra and N. setigera. The

genus Pseudarthria was published at the same time as

12

Bothalia 18,1 (1988)

KEY TO SUBTRIBES

Plants with characteristic uncinate hairs; tertiary venation ± scalariform; stipels present; flowers chasmogamous,

with an explosively dehiscent pollination mechanism; standard with hardly or slightly indexed auricles;

ovary( 1) to many-ovulate subtribe Desmodiinae

Plants without uncinate hairs; tertiary venation reticulate; stipels absent; flowers chasmogamous, not explosively

dehiscent, often mixed with cleistogamous flowers; standard with indexed auricles more persistent than most

Desmodiinae; ovary 1-ovulate subtribe Lespedezinae

the species P. hookeri by Wight & Amott in 1834. Ben-

tham, in his review of the Leguminosae in 1852, placed

Hedysarum barbatum and Nicolsonia caffra under Des-

modium, calling the latter Desmodium dregeanum. Har-

vey (1862), in revising the South African representatives

of the tribe for Flora capensis, placed Nicolsonia seti-

gera under Desmodium and described a new species,

Alysicarpus zeyheri. The genus Nicolsonia was later put

into synonomy with Desmodium.

Other major contributors to the literature on the tribe

include Baker (1871), Schindler (1928), Baker, E. G.

(1929), Schubert & Verdcourt (1971), Ohashi (1971b,

1973), Verdcourt (1974), Ohashi, Polhill & Schubert

(1981) and Schrire (1984).

The subgeneric treatment of Desmodium is as yet

unsatisfactory but for the purposes of this synopsis the

treatment of Ohashi (1973) is followed, i.e. D. repan-

dum is placed in subgenus Podocarpium ; D. setigerum,

D. adscendens, D. barbatum and D. dregeanum in sub-

genus Nicolsonia and the rest in subgenus Heteroloma.

TRIBE DESMODIEAE

Desmodieae (Benth.) Hutch., The genera of flow-

ering plants 1 : 477 (1964), pro parte; Schubert & Verde.:

450 (1971); Verde.: 505 (1974); Ohashi et al.,: 292

(1981). Type genus: Desmodium Desv.

Coronilleae Adans.: 327 (1763), pro parte.

Hedysareae DC.: 307 (1825), pro parte.

Hedysareae subtribe Desmodiinae (as Desmodieae)

Benth.: 449 (1865), pro parte; Taub.: 325 (1894), pro

parte.

Coronilleae subtribe Desmodiinae (Benth.) Schulz-

Menz: 237 (1964), pro parte; Ohashi: 52 (1971a).

Herbs or suffrutices, sometimes shrubs or rarely trees.

Roots woody, often dichotomously branched, cord-like

and producing rhizomes, or more rarely taproots. Leaves

pinnately 3— (9)- or 1-foliolate, petioles sulcate, foliar

and foliolar pulvini present, stipellate (except Lespe-

deza)\ stipules mostly striate. Pseudoracemes variously

contracted, terminal, axillary or occasionally leaf-

opposed, sometimes paniculately branched. Flowers

pedicellate, fasciculate, mostly paired, rarely solitary on

the rhachis, usually subtended by 2 series of bracts and

occasionally bracteoles. Calyx 2-lobed, the upper with 2

connate sepals often slightly bifid at the apex, the lower

3-toothed with the central tooth longer than the laterals,

or the calyx almost equally 5-lobed. Corolla papilionoid.

Stamens mostly diadelphous with the vexillary stamen

free, rarely pseudo-monadelphous; filaments equal or

alternating in length; anthers uniform. Pollen tricol-

porate with a generally thickened endexine and thin or no

foot layer. Fruits transversely jointed into a number of

articles or reduced to one article, or less often 2-valved,

opening along one suture (valves not twisting). Seeds

with a well developed radicular lobe, longer than the

cotyledonary lobe, mostly with a rim aril. Seedlings

generally epigeous with the first two eophylls opposite

and unifoliolate.

Subtribe Desmodiinae

Desmodiinae sensu Ohashi et al. in Advances in

legume systematics 1 : 296 (1981).

Pseudarthrieae Hutch.: 398 (1964), pro parte.

DESMODIUM

Desmodium Desv., Journal de botanique 1: 122,

t.5 (1813) nom. conserv.; DC.: 325 (1825); Benth.: 220

(1852); Harv.: 227 (1862); Benth.: 519 (1865); Harv.:

82 (1868); Bak.: 159 (1871); Bak.: 161 (1876); Taub.:

327 (1894); Schindl.: 263 (1928); Hutch. & Dalz.: 417

(1928); Bak.f.: 324 (1929); Phill.: 332 (1951); Schubert:

287 (1952); Schubert: 180 (1954); Hepper: 582 (1958);

Knaap-van Meeuwen: 240 (1962); Hutch.: 481 (1964);

Torre: 217 (1966); Schreiber: 30 (1970); Schubert: 451

(1971); Ohashi: 87 (1973); Verde.: 506 (1974); Dyer:

263 (1975); Compton: 272 (1976); Schubert: 622 (1980).

Type species: D. scorpiurus (Swartz) Desv. ( Hedysarum

scorpiurus Swartz).

KEY TO GENERA

la Calyx often membranous, never glumaceous or striate, shallowly divided, usually inconspicuous in fruit; fruit

mostly held away from erect; leaves ( I)— 3- (5)- foliolate:

2a Stipules stramineous, striate, erect or spreading but never sharply recurved and velutinous; fruit distinctly

jointed, breaking up into individual articles Desmodium

2b Stipules mostly sharply recurved, dark brown adaxially, silver-velutinous abaxially; fruit splitting into 2

valves Pseudarthria

lb Calyx glumaceous, striate, deeply divided, persistent and often partially enclosing the fruit; fruit held erect,

usually consisting of a number of 'stacked' terete or subterete articles; leaves 1-foliolate Alysicarpus

Bothalia 18,1 (1988)

13

Hedysarum L.: 745 (1753), pro parte (name still cor-

rect for other plants).

Meibomia Heist, ex Fabricius: 168 (1759), nom.

rejic.; Adans.: 509(1763); Kuntze: 195 (1891).

Nicolsonia DC.: 325 (1825).

The name Desmodium was conserved over the name

Meibomia by the International Botanical Congress of

Vienna in 1905 (Schubert 1950). Desmodium is derived

from the Greek ‘desmos’ meaning a band or chain, and

‘hode’ meaning like, in reference to the jointed pod’s

resemblance to the links of a chain.

SYNOPSIS OF SPECIES

1. Desmodium repandum (Vahl) DC., Prodromus

systematis naturalis regni vegetabilis 2: 334 (1825);

Schindl.: 295 (1928); Schubert: 293 (1952); Schubert:

193, 1. 14 (1954); Hepper: 584 (1958); White: 150

(1962); Laundon: 221 (1966); Schubert: 465, t. 65/11

(1971); Verde.: 517 (1974); Compton: 272 (1976).

Type: Yemen, Forsskal (C, holo., fide Schindl.— GH,

photo.!).

Hedysarum repandum Vahl: 32 (1791); Poir.: 408 (1805).

Desmodium scalpe DC.: 334 (1825); Bak.: 164 (1871); Hutch. &

Dalz.: 418 (1928); Bak. f.: 328 (1929); Robyns: 327, t.30 (1948);

Brenan: 420 (1949). Type: Mauritius (Isle de Bourbon), Commerson

(P-LA, holo.; GH, photo.!).

Desmodium strangulatum Wight & Am.: 228 (1834); Harv.: 228

(1862). Type: E Peninsular India, Walker in Wight cat. no. 774 (K,

holo.!;GH).

Desmodium caffrum Eckl. & Zeyh.: 251 (1836) non (E.Mey.)

Druce: 619 (1917), which is a later illegitimate name forZ). dregeanum

Benth. Type: E Cape, Mankazana R., Eckl. & Zeyh. in Enum. 1662

(K, iso.!).

KEY TO SPECIES

la Inflorescences of variously lax to dense, elongated, axillary and terminal pseudoracemes, the terminal ones often

paniculately branched; calyx teeth short, triangular or lanceolate, the adaxial two usually markedly connate

forming a lip:

2a Flowers 8,0—14,0 mm long, brick red; herbs or subshrubs, climbing or scrambling and forming a dense

undergrowth in forest; leaflets rhomboid, deep green venation craspedodromous; inflorescences very lax;

stamens partially monadelphous \.D. repandum

2b Flowers 4,5-8,0 mm long, variously pink, mauve or purple but never red; plants not as above, stamens

always diadelphous:

3a Plants becoming erect to semi-erect suffrutices or subshrubs if not burned; leaflets never obovate, lateral

veins not obviously looping in the distal half of the leaves; floral bracts present; inflorescences variously

contracted:

4a Flowers uniformly deep mauve; pseudoracemes terminal only, and never paniculately branched; leaflets

glossy dark green with characteristic light discolouration along the midrib above; widespread weeds of

disturbed areas in coastal Zululand, Natal, Transkei and eastern Cape to 1 000 m 2.D. incanum

4b Flowers light pink or purple, dark purple only at the tip of the keel; pseudoracemes terminal and often

axillary, the terminal ones often paniculately branched; leaflets various, not as above:

5a Leaves trifoliolate:

6a Venation mixed craspedodromous with principal lateral veins prominent and parallel; leaflets dis-

tinctly saliciform; stipules not as below; articles of the fruit rectangular or oblong, not or slightly

constricted at the isthmi; shrub growing in or near water 4. D. salicifolium

6b Venation eucamptodromous, lateral veins neither prominent nor regular; stipules markedly auri-

culate at the base on the leaf-opposed margin, fused into a collar around the stem when young;

articles of the fruit orbicular to elliptic, strongly constricted at the isthmi; introduced weed with a

restricted distribution 3. D. tortuosum

5b Leaves unifoliolate:

7a Leaflets chartaceous, ovate or cordiform with entire margins, venation eucamptodromous; stipules

gradually narrowing from the base to a slender apex 6. D. gangeticum

7b Leaflets crassate, velutinous, suborbicular, often with sinuate margins; venation craspedodromous;

stipules abruptly narrowing from an auriculate base to a long slender apex 5. D. velutinum

3b Plants procumbent, or stoloniferous creepers; leaflets obovate, lateral veins obviously looping in the distal

half of the leaves; floral bracts usually absent; inflorescences lax:

8a Procumbent herbs rooting from the lower nodes, in open grassland to forest fringes, often drying brown or

grey-green in herbarium specimens; articles of the fruit U-shaped, 1 ,5-2,0 mm wide 7. D. setigerum

8b Stoloniferous herbs of moist forest and forest margin undergrowth, often drying deep green in herbarium

specimens; articles approximately obtriangular, 2,5-3, 0 mm wide 8. D. adscendens

lb Inflorescences congested into dense spicate or capitate pseudoracemes, never paniculately branched but often

with the upper axillary and terminal inflorescences crowded together; calyx teeth long-acuminate, the adaxial

two not markedly connate:

9a Leaves variously 3 or 1-foliolate; inflorescences spicate; pedicels often dark-coloured, visibly deflexed dis-

tally; bracteoles absent; woody herbs in mixed open woodland; northern and eastern

Transvaal 10. D. barbatum

9b Leaves 3-foliolate, conduplicate; inflorescences densely capitate; pedicels not obvious; bracteoles 2,

clasping the calyx; woody herbs mostly of moist grassland 9. D. dregeanum

14

Bothalia 18,1 (1988)

Distribution: 1, Transvaal— a, in Mixed and Sourish

Mixed Bushveld, Acocks Veld Types 18 & 19, from

Zeerust in the west to near Witbank in the east; b, in

similar bushveld or in other types of Sourveld and Sour

Bushveld, Acocks Veld Types 8 & 9, from Louis Tri-

chardt, Pietersburg and Tzaneen in the north, between

Lydenburg and Graskop, Belfast and Barberton, to west-

ern Swaziland in the south. It does not occur in the Low-

veld. 2, Natal— where suitable forest patches occur from

the Drakensberg to almost sea level except in coastal

Dune forest. 3, Transkei and eastern Cape to the coastal

southern Cape at Knysna.

This species is widespread in tropical and subtropical

Africa, Madagascar, the Mascarene Islands, India and

south-east Asia to Malesia.

Vouchers: Compton 27287 (PRE, SAM); Flanagan 515 (BOL,

GRA, NBG, NU, PRE, SAM); B. & C Howlett 16 (NH, NU, PRE);

Kluge 467 (PRE, PRU); Wells 2880 (GRA, NH, PRE).

Notes

(1) Desmodium repandum is found in shaded areas in

moist to dry evergreen mountain, riverine or other gal-

lery forest often near paths or along streams, from ±

3 000 m to near sea level. It occurs as a forest floor herb

or scrambling subshrub. Flowering is from August to

June although most commonly from January to May.

(2) In the latest Asian revision of Desmodium by Oha-

shi (1973), D. repandum is placed in the most advanced

subgenus Podocarpium, because it is characterized by a

monadelphous androecium and long-stipitate pods con-

sisting of indehiscent, approximately obtriangular

articles. However, the androecium cannot be considered

to be truly monadelphous, the vexillary stamen being

coherent (not fused) for part of its length only. The pol-

len grains are also quite distinct from those of all other

species of the subgenus Podocarpium, especially in the

shape of the grains, the colpi and thickness of the exine

(Ohashi 1973). D. repandum is also noted (Ohashi 1973)

for having several characters in common with some

members of the less derived subgenus Dollinera.

From observations on the degree of contraction of the

pseudoraceme in D. repandum it appears that this also

shows a less derived situation. Occasionally one pedicel

may be seen to branch off another, above the fascicle,

indicating incomplete reduction of the botrys. A number

of pedicels also occur in the fascicle, three or four of

which may flower with one or two reduced buds in

between. The fascicle in all the other species in the Flora

area has been reduced to two pedicels with or without

one enclosed median bud.

2. Desmodium incanum DC., Prodromus

systematis naturalis regni vegetabilis 2: 332 (1825);

Bak.: 163 (1871); Hutch. & Dalz.: 418 (1928); Nicol-

son: 365 (1978). Type illustration: Plumier, in J. Bur-

man, PI. Amer.: 140, t. 149, fig. 1 (1757) — Hedysarum

fold’s ternatis, ovatis, floribus spicatis{\).

Hedysarum racemosum Aubl.: 11 A (1775). Type: as for species.

This is the first legitimate name but the epithet is unavailable in Desmo-

dium. Hedysarum incanum Swartz: 107 (1788) reprint 1962; Swartz:

1264 (1806), non Thunb. (1784). Illegitimate renaming of H. racemo-

sum Aubl. Hedysarum canum J. F. Gmel.: 1124 (1791), non Lunan

(1814). Superfluous renaming of H. racemosum Aubl. Desmodium

canum (J. F. Gmel.) Schinz & Thell : 371 (1913); Schinz & Thell.:

428 (1913); Schubert: 184 (1954); Hepper: 584 (1958); Schubert: 456

(1971). Treated as a new name with priority from publication because

of the illegitimate basionym, it is a superfluous renaming of D. inca-

num EXT.

Hedysarum frutescens auct. non L.: Jacq.: 47, t. 89 (1776).

Desmodium frutescens sensu Schindl.: 9 (1928); Bak. f.: 328

(1929). Intended as a new combination of H . frustescens sensu Jacq.

Distribution: Desmodium incanum occurs along the

coast and hinterland up to 1 000 m in Natal, and in

coastal Transkei just into the eastern Cape.

Vouchers: Fisher 590 (NH, NU); Gillen 1203 (BOL, NH); Grobbe-

laar 284 (PRE, PRU); Ward 584 (NU, PRE); Wood 644 (BOL, SAM).

Notes

(1) D. incanum is a spreading herb to woody subshrub

up to 1 m (if protected from fire); the leaf shape may

vary; leaves in deep shade may superficially resemble

those of D. adscendens. Confusion between the two may

occur on forest margins but the partially connate sti-

pules, deep mauve flowers and characteristic fruit of D.

incanum will help to distinguish them. D. incanum is

otherwise readily recognized by its shiny dark green

leaves with a light discolouration along the midrib.

Flowering occurs from September to June.

(2) The plant is a weed in disturbed grassland, wood-

land and forest edges, along streams, paths and roadsides

and often occurs in lawns. It is also invasive in undis-

turbed grassland.

(3) D. incanum is tropical American in origin but is

widely scattered throughout tropical Africa where it is

introduced. It is most likely to have entered the Flora

area along the Natal coast. The Zulu name ‘isinama’

describes the fruit which readily catches and sticks to

clothing. Allen & Allen (1981) refer to the plant as

Kaimi clover or creeping beggarweed and it is favoured

as a forage legume in Hawaii and Florida for its tolerance

to acid soils and wet warm climate.

3. Desmodium tortuosum (Swartz) DC., Prodro-

mus systematis naturalis regni vegetabilis 2: 332 (1825);

Schubert: 202 (1954); Hepper: 585 (1958); Lind & Tal-

lantire: 82, t. 32 (1962); Laundon; 223 (1966); Schubert:

474 (1971); Verde.: 526 (1974). Type: Jamaica, Swartz

(S, lecto.-GH, photo.!).

Hedysarum tortuosum Swartz: 107 (1788) reprint 1962; Swartz:

1271 (1806).

Desmodium spirale auct. non (Swartz) DC.: Bak.: 160 ( 1871 ); Bak.

f.: 331(1929).

Distribution: D. tortuosum is an escape from cultiva-

tion which has now become naturalized in disturbed

grassy areas or along roadsides, in Nelspruit in the

Transvaal, and in the Durban and Port Shepstone Dis-

tricts of Natal.

Vouchers: Coleman 1103 (NH); Mogg, Govt. Herb. 10071 (PRE);

Nichols 433 (NH); Rabie 273 (PRE).

Notes

( 1 ) The Swartz specimen has been chosen as the type

(Schubert pers. comm. 1983) in preference to the Sloan

illustration: 1 16, t. 9 (1696), following the precept in the

Guide for the Determination of Types, ICBN (Stafleu

1983), T. 4b: ‘A specimen is to be given preference over

pre-Linnaean or other cited descriptions or illustrations

when lectotypes of names of species or infra-specific

taxa are designated (see Art. 9.3)’.

Bothalia 18,1 (1988)

(2) D. tortuosum is an herbaceous suffrutex or small

shrub to I m, flowering from December to May. It is

known as Florida beggarweed in the south eastern United

States (Schubert 1971) and is cultivated as a green

manure. It is now naturalized throughout the Old World

tropics coming originally from tropical and subtropical

America. The populations in Durban and Port Shepstone

are large and well established.

4. Desmodium salicifolium (Poir.) DC., Prodro-

mus systematis naturalis regni vegetabilis 2: 337 (1825).

Type: Herb, de I’lnde de M. Poivre (P-Ju 15552,

holo.-GH, photo.!).

Hedysarum salicifolium Poir.: 422 (1805).

Desmodium paleaceum Guill. & Perr.: 209 (1832). Type: Senegal,

Perrottet 246 (P, holo.; W).

Desmodium grande E. Mey.: 124 (1836). Type: Durban (Port

Natal), Drege (B, holo.; W, K!).

var. salicifolium

Schubert in Bulletin du Jardin botanique de l’Etat, a

Bruxelles 22: 294 (1952); Schubert: 470, t. 65/1 (1971);

Verde.: 523 (1974).

Hedysarum salicifolium Poir.: 422 (1805). Desmodium salicifolium

(Poir.) DC.: 337 (1825); Bak.f.: 330 (1929); Schubert: 294 (1952);

Schubert: 198, t. XV (1954); Hepper: 584 (1958); White: 150 (1962);

Laundon: 224, t. 20A (1966); Schubert: 469, t. 65/1 (1971); Verde :

522(1974).

Desmodium paleaceum Guill. & Perr.: 209 (1832); Bak.: 166

(1871); Hutch. & Dalz.: 418 ( 1928).

Desmodium grande E. Mey.: 124 ( 1836); Harv.: 228(1862).

Distribution: D. salicifolium var. salicifolium occurs

in northern South West Africa/Nambia, Caprivi and nor-

thern Botswana, and in Mozambique coming into Koma-

tipoort and Nelspruit in the Transvaal, eastern Swaziland

and southward along coastal Zululand and Natal, from

0- 1 000 m. It occurs widely in tropical Africa, Mada-

gascar and the Mascarene Islands.

Vouchers: Buitendag 1155 (NBG, PRE); Moll & Nel 5610 (NH,

PRE); Venter 5489 (PRE, ZULU); Ward 5560 (NH, NU); Wylie sub

Wood 10335 (GRA, PRE, SAM, STE).

Note

D. salicifolium is a suffrutex or shrub of wet places,

principally along river margins or in swamp forest. It

often occurs in wet clayey soil and flowers from August

to May.

5. Desmodium velutinum (Willd.) DC., Prodro-

mus systematis naturalis regni vegetabilis 2: 328 (1825);

Schubert: 294 (1952); Schubert: 194 (1954); Hepper:

584 (1958); White: 150 (1962); Schubert: 292 (1963);

Laundon: 219 (1966); Schubert: 466, t. 65/7 (1971);

Verde.: 518 (1974). Type: Herb. Willdenow 13763 (B,

holo.-GH, photo.!).

Hedysarum velutinum Willd.: 1 174 ( 1802).

Hedysarum lasiocarpum Beauv.: 32, t. 18 (1805). Desmodium

lasiocarpum (Beauv) DC.: 328 (1825); Bak.: 162 (1871); Hutch. &

Dalz.: 418 ( 1928); Bak.f.: 326 ( 1929). Type: Nigeria, Palisotde Beau-

vois (G, holo. -A, photo.!).

Distribution: D. velutinum is only known from the

north-eastern Lowveld of the Transvaal, up to 750 m,

from near Leydsdorp to the southern boundary of the

15

Kruger National Park. It is widespread throughout the

Old World tropics from Africa and Madagascar to India,

south-east Asia and Malesia.

Vouchers: Junod sub TRV 5279 (PRE); Schrire 655 (NH); Van der

Schijff2659 (PRE).

Note

The plant occurs as a small shrub or suffrutex in

wooded and open grassland, woodland and on forest

margins.

6. Desmodium gangeticum (L.) DC., Prodromus

systematis naturalis regni vegetabilis 2: 327 (1825);

Wight & Am.: 225-226 (1834); Bak.: 161 (1871);

Hutch. & Dalz.: 418 (1928); Bak.f.: 327 (1929); Brenan:

420 (1949); Schubert: 196 (1954); Hepper: 584 (1958);

White: 150 (1962); Laundon: 219 (1966); Schubert: 465,

t. 65/10 (1971); Verde.: 520 (1974). Type: Herb. Lin-

naeus 921 .13 (LINN, holo. — A, photo.!).

Hedysarum gangeticum L.: 746 ( 1753).

Hedysarum maculatum L.: 746 (1753). Desmodium gangeticum (L.)

DC. var. maculatum (L.) Bak.: 168 (1876); Hepper: 584 (1958); Laun-

don: 220 (1966). Type: Herb. Linnaeus 921.14 (LINN, holo. -A,

photo.!).

Desmodium tuitalitium Sond.: 32 (1850); Harv.: 229 (1862);

Schindl.: 285 (1928). Type: South Africa, Port Natal, Gueinzius 203

(Herb. Sond.).

Distribution: D. gangeticum occurs in the Transvaal

from the Soutpansberg in the north, through the Lowveld

between Barberton and Nelspruit to Swaziland in the

south. In Natal this distribution continues through nor-

thern Natal to Zululand and then along the coast south-

wards to the Transkei, from 0— 1 200 m.

Vouchers: McDonald 118 (NU, PRE); Schlechter 3129 (BOL,

GRA, PRE); Van der Schijff965 (PRE, PUC); Ward 2406 (NH, PRE);

Wood 4936 (BOL, PRE, SAM).

Notes

(1) The epithets gangeticum and maculatum were

published on the same date and they were combined

under the name Desmodium gangeticum by Wight &

Arn.: 225-226 (1834).

(2) The plant is a spreading suffrutex to erect subshrub

occurring in wooded grassland, bushveld or thicket or in

open grassland. It readily becomes a weed in semi-dis-

turbed areas i.e. old lands or roadsides, and flowers from

August to March.

(3) This species is widespread throughout the Old

World tropics and has been introduced into America.

Allen & Allen (1981) note that D. gangeticum is used as

a pioneer plant to control erosion of denuded areas. The

roots of D. gangeticum yield a lectone and seven alka-

loids. One of these, hordenine, has been used in experi-

mental medicine as a sympathomimetic and another,

bufotenine, is an hallucinogen.

7. Desmodium setigerum (E. Mey.) Benth. ex

Harv. in Flora capensis 2: 229 (1862); Milne-Redh.: 417

(1937); Schubert: 187, t. HE (1954); Hepper: 585

(1958); Laundon: 222 (1966); Schubert: 460, t. 65/5

(1971); Verde.: 513 (1974). Type: Transkei, Umzim-

vubu River, Drege 446 (B, holo.; W,K!).

Nicolsonia setigera E. Mey.: 124(1836).

16

Bothalia 18,1 (1988)

Desmodium hirtum auct. non Guill. & Perr.: Bak.: 163 (1871), pro

parte; Wood: 14, t. 212 (1902); Hutch. & Dalz.: 418 (1928); Schindl.:

279(1928): Bak.f.: 329(1929), pro parte; Compton: 272(1976).

Distribution: D. setigerum occurs in the Transvaal

from the Soutpansberg and Tzaneen in the north, through

the area between Lydenburg and Sabie to western Swazi-

land in the south. In Natal it is found from sea level to

2 000 m, except in the lowveld of Zululand, and it

extends to coastal Transkei. It is widespread throughout

tropical Africa.

Vouchers: Bos 1253 (PRE, STE); Compton 30608 (NBG, PRE);

Ross 1679 (NH, NU, PRE); Scheepers 581 (PRE, PRU); Strey 2996

(NH, PRE).

Notes

(1) Misinterpretation of types (Schubert 1971) has led

to confusion between two separate species, Desmodium

setigerum (E. Mey.) Benth. ex Harv. and D. hirtum

Guill. & Perr. This began with D. hirtum sensu Bak.:

163(1871).

(2) The plant is a prostrate or low semi-erect suffrutex

of open grassland in damp sites, occurring occasionally

on streambanks, forest margins or on drier rocky hill-

sides. It is also found in semi-disturbed vegetation on

roadsides and flowers from December to May, i.e. it is a

later summer- flowering species.

8. Desmodium adscendens (Swartz) DC., Prodro-

mus systematis naturalis regni vegetabilis 2: 332 (1825);

Bak.: 162 (1871); Hutch. & Dalz.: 418 (1928); Bak.f.:

330 (1929); Brenan: 419 (1949); Laundon: 222, t. 20A1

(1966). Type: West Indies, Swartz (S, holo.— GH,

photo.!).

Hedysarum adscendens Swartz: 106 (1788) reprint 1962; Swartz:

1263(1806).

var. robustum Schubert in Bulletin du Jardin bota-

nique de l’Etat, a Bruxelles 22: 290 (1952); Schubert:

190 (1954); Hepper: 585 (1958); Schubert: 461, t. 65/13

(1971); Verde.: 514 (1974). Type: Congo, Kivu,

Mokoto Lakes, Ghesquiere 4994 (BR, holo.).

Distribution: D. adscendens var. robustum is found in

the Soutpansberg and Woodbush areas of the northern

Transvaal and entering from Mozambique along the

Natal coast to the Transkei border. It is widespread in

tropical Africa but the species is pantropical, from

America, Africa, Asia and Malesia.

Vouchers: Gillen 3156 (BOL, PRE, STE); Moll 2720 (NU, PRE);

Moll & Strey 38 96 (NH, PRE); Strey 7151 (NH, NU, PRE); Venter

5291 (PRE, ZULU).

Note

The plant is a straggling herb or undershrub of shaded

mesic forest floors, and flowers from November to

March. On forest margins along the coast, this species

may be confused with the shade form of D. incanum (see

under that species). Schubert (1971) records that D.

adscendens is an excellent food for stock.

9. Desmodium dregeanum Benth. in Plantae Jung-

huhnianae 2: 222 in adnot. (1852); Harv.: 228 (1862);

Bak.: 165 (1871); Schubert: 476, t. 65/3 (1971); Verde.:

528 (1974). Type: Transkei, Umsimkaba River, Drege

(B, holo.; GH, K!). Based on Nicolsonia caffra E. Mey.

Nicolsonia caffra E. Mey.: 123 (1836).

Desmodium caffrum (E. Mey.) Druce: 619 (1917); Schindl.: 360

(1927); Bak.f.: 331 (1929); Brenan: 419 (1949); Laundon: 225(1966),

non Eckl. & Zeyh. (1836), nom. illegit. D. caffrum Eckl. & Zeyh. is a

later homonym of D. repandum (Vahl) DC.

Distribution: D. dregeanum is predominantly a low

altitude plant from coastal Mozambique to the coast of

northern Transkei. It does however occur in the Lyden-

burg, Barberton and Volksrust Districts of the Transvaal

and the hinterland of Natal from 0-1 800 m. It is an

endemic African species occurring widely in eastern and

southern Africa.

Vouchers: Grobbelaar 1104 (PRE, PRU); Rogers 28284 (GRA,

STE); Ross & Moll 1830 (NH, PRE); Strey 4971 (NH, NU, PRE);

Wood 817 (BOL, SAM).

Note

The plant is an erect or straggling herb in open grass-

land most often in moist or marshy areas in sandy soils.

It can become a woody subshrub to 1 m high if protected

from fire and it is common in semi-disturbed grasslands,

roadsides or forest margins. Flowering may occur

throughout the year.

10. Desmodium barbatum (L.) Benth. in Plantae

Junghuhnianae 2: 224 (1852); Schubert: 477 (1971);

Verde.: 530 (1974). Type: Jamaica?, Herb Linnaeus

921.48 (LINN, holo.).

Hedysarum barbatum L.: 1170 (1759).

10a. var. dimorphum (Welw. ex Bak.) Schubert in

Bulletin du Jardin botanique de l’Etat, a Bruxelles 22:

298 (1952); Schubert: 205 (1954); Hepper: 564 (1958);

Schubert: 478 (1971); Verde.: 531 (1974). Type:

Angola, Cuanza Norte, Golungo Alto, Welwitsch 2165

(LISU, lecto.; K!, BM!); Malawi, Manganya, Kirk (K,

syn.!).

Desmodium dimorphum Welw. ex Bak.: 161 (1871); Hutch. &

Dalz.: 417 (1928); Bak.f.: 332 (1929). Nicolsonia barbata (L.) DC.

var. dimorpha (Welw. ex Bak.) Schindl.: 359 (1927). D. barbatum

(L.) Benth. subsp. dimorphum (Welw. ex Bak.) Laundon: 225, t.

20/ A2 (1966).

Distribution: D. barbatum var. dimorphum is found in

the Transvaal from the Letaba District near Tzaneen

south to Lydenburg, Barberton and Nelspruit, from

100-1 000 m.

Vouchers: Kluge 1222 (PRE); Krige 139 (J); Liebenberg 2897

(PRE); Scheepers 241 (PRE, PRU); Schrire 661 (NH).

Note

The plant is a spreading suffrutex to erect subshrub up

to 1 m high, in the grassland component of open wood-

land, and it flowers from February to June.

10b. var. argyreum (Welw. ex Bak.) Schubert in

Bulletin du Jardin botanique de l’Etat, a Bruxelles 22:

298 ( 1 952); Schubert: 205 ( 1 954); Schubert: 479 ( 1 97 1 );

Verde.: 531 (1974). Type: Angola, Huila, Welwitsch

2162 (LISU, holo.; BM!).

Bothalia 18,1 (1988)

Desmodium dimorphum Welw. ex Bak. var. argyreum Welw. ex

Bak.: 161 (1871); Bak.f.: 332 (1929). Nicolsonia barbata (L.) DC.

var. argyrae (Welw. ex Bak.) Schindl.: 359 (1927). D. barbaium (L.)

Benth. subsp. dimorphum sensu Laundon: 225 (1966), pro parte.

Distribution: D. barbatum var. argyreum has been

recorded from South West Africa/Namibia.

Voucher: Merxmuller <6 Giess 1862 (WIND).

Notes

(1) The plant is erect with silvery-white or fulvous

stems which are appressed silky-pilose. The leaves are

almost all 1-foliolate with the leaflets densely silvery-

silky beneath. It occurs in woodland or grassland from

1 000- 2 000 m.

(2) Schubert (1971) notes that, although this variety is

quite distinguishable from var. dimorphum in the ex-

treme, there are many plants which form a transition

between the two.

Desmodium uncinatum and D. intortum or Silverleaf

and Greenleaf Desmodium, have been deliberately intro-

duced from central America and cultivated as pasture

legumes in the eastern Cape. This is largely out of the

natural range of the genus in southern Africa, and there

is no evidence that they have become naturalized. Des-

modium asperum was once collected on the south coast

of Natal but it has not been seen again. A large number

of other species of Desmodium are being tested in agri-

cultural stations around the country for their pasture

potential.

PSEUDARTHRIA

Pseudarthria Wight & Arn., Prodromus florae

peninsulae Indiae orientalis 1: 209 (1834); Benth.: 521

(1865); Bak.: 167 (1871); Bak.: 153 (1876); Taub.: 329

(1894); Schindl.: 11 (1914); Hutch. & Dalz.: 386

(1928); Bak.f.: 338 (1929); Burtt Davy: 381 (1932);

Phill.: 419 (1951); J. Leonard: 234 (1954b); Hepper: 585

(1958); White: 161 (1962); Hutch.: 399 (1964); Torre:

231 (1966); Verde.: 64 (1970); Verde.: 483 (1971);

Verde.: 534 (1974); Dyer: 264 (1975). Type species: P.

viscida (L.) Wight & Am. ( Hedysarum viscidum L.).

Hedysarum L.: 745 (1753), pro parte.

Desmodium Desv. in DC.: 325 (1825), pro parte.

Anarthrosyne E. Mey.: 124 (1836); Harv.: 229

(1862), pro parte; Harv.: 82 (1868).

The name Pseudarthria is derived from the Greek

‘pseudo’, meaning false, and ‘arthron’ or joint; the con-

strictions between the seeds give the impression of the

pod being falsely jointed. Only one species occurs in

southern Africa.

Pseudarthria hookeri Wight & Arn., Prodromus

florae peninsulae Indiae orientalis 1: 209 (1834); Bak.:

168 (1871); Schindl.: 11 (1914); Hutch. & Dalz.: 386

(1928); Bak.f.: 339 (1929); Robyns: 313 (1948); Brenan:

436 (1949); J. Leonard: 235, t. 15/A-B (1954b); Hep-

per: 586 (1958); White: 161 (1962); Torre: 232 (1966);

Verde.: 65 (1970); Verde.: 484, t. 69/1-9 (1971);

Verde.: 534 (1974). Type: Mauritius, cultivated from

Zanzibar, Telfair (K, ?holo.!).

17

Anarthrosyne robusta E. Mey.: 125 (1836); Harv.: 229 (1862).

Type: Transkei, Umgazana, Drege 452 (K!).

var. hookeri

Verde, in Kew Bulletin 24: 65 (1970); Verde.: 484, t.

69/1-9 (1971); Verde.: 535 (1974).

Distribution: Pseudarthria hookeri var. hookeri

occurs in the Transvaal from Louis Trichardt and Tza-

neen in the north, between Lydenburg and Skukuza, and

between Middelburg and Komatipoort, to Swaziland and

Piet Retief in the south. The species also occurs from

Petersburg through Nylstroom to Thabazimbi in the

Waterberg Mountains. In Natal the range extends south-

wards through northern Natal and Zululand to the coast

and midlands up to Pietermaritzburg, Richmond, Ixopo

and Harding. Further south the range becomes restricted

to coastal Transkei to the eastern Cape border. This

species occurs widely in tropical and southern Africa,

Madagascar, Mauritius and Reunion.

Vouchers: Bos 1300 (PRE, STE); Galpin 776 (GRA, NH, SAM);

Hemm 555 (J, PRE, VENDA); Moll 2744 (NH, NU, PRE); Schlechter

4551 (BOL, GRA).

Note

This species is found most frequently as a suffrutex or

subshrub to 3 m high, in mixed woodland or bushveld,

along forest margins, in open grassland or as a ruderal

along roadsides or in other disturbed areas, from

0— 2 000 m. It is also occasionally associated with

streambanks, vleis or swampy areas. Flowering occurs

between November and May.

ALYSICARPUS

Alysicarpus Desv., Journal de botanique 1: 120,

t.4.f.8. (1813), nom. conserv.; DC.: 352 (1825); Harv.:

230 (1862); Benth.: 522 (1865); Harv.: 82 (1868); Bak.:

169 (1871); Bak.: 157 (1876); Hutch. & Dalz.: 418

(1928); Bak. f.: 341 (1929); Burtt Davy: 427 (1932);

Phill.: 419 (1951); J. Leonard: 84 (1954a); J. Leonard:

223 (1954b); Hepper: 586 (1958); Hutch.: 482 (1964);

Torre: 234 (1966); Schreiber: 13 (1970); Verde.: 491

(1971); Verde. 544 (1974); Dyer: 264 (1975); Compton:

273 (1976); Dillon: 548 (1980). Type species: A. bupleu-

rifolius (L.) DC. (Hedysarum bupleurifolium L.).

Hedysarum L.: 745 (1753), pro parte.

Fabricia Scop.: 307 (1777); Kuntze: 181 (1891);

Taub.: 329 (1894).

Alysicarpus gets its name from the Greek, ‘halysis’,

meaning chain and ‘karpos’ meaning fruit; the pods are

moniliform with the joints in chain formation.

Note

Alysicarpus Neck.: 15 (1790) is not to be treated as a

generic name, according to ICBN Art. 20, unless it has

been published, as such, by a subsequent author. Noted

in Stafleu (1976) is the following: ‘Since Necker desig-

nated this category as ‘species’, the monomial names

(which are in many cases the generic names of previous

authors) are to be regarded as unitary designations of

species and hence to be regarded as not validly pub-

lished’.

18

KEY TO SPECIES

Bothalia 18,1 (1988)

la Fruit with straight margins, not constricted between the articles; calyx lobes narrow, sharply acuminate, not at all

overlapping at the base 1 . A. vaginalis

lb Fruit moniliform, strongly constricted between the articles; calyx lobes ovate-lanceolate, slightly to conspi-

cuously overlapping at the base:

2a Articles smooth; leaves coriaceous with very prominent reticulate venation 2. A. zeyheri

2b Articles with obvious transverse ridge sculpturing; leaves subcoriaceous with reticulate venation only slightly

prominent:

3a Calyx lobes 0,8- 1,5 mm wide, slightly overlapping and imbricate initially, not so in fruit; inflorescences

elongate and spike-like; fruit well exserted from the calyx 4. A. glumaceus

3b Calyx lobes 1 ,5—2,5 mm wide, conspicuously overlapping and imbricate, and rounded or subcordate at the

base; inflorescences often short and compact; fruit only partially exserted from the calyx 3. A. rugosus

SYNOPSIS OF SPECIES

1. Aiysicarpus vaginalis (L.) DC., Prodromus

systematis naturalis regni vegetabilis 2; 353 (1825);

Bak.: 170 (1871), pro parte; Hutch. & Dalz.: 419

(1928), pro parte; Bak.f.: 342 (1929), pro parte; Robyns:

331 (1948); J. Leonard: 84 (1954a); J. Leonard: 224, t.

13/A (1954b); Hepper: 587 (1958); Torre: 234 (1966);

Verde.: 493, t. 71 /A (1971); Verde.: 546 (1974); Comp-

ton: 273 (1976). Syntypes: Ceylon, Hermann (BM-

HERM, 1:27,59, syn.).

Hedysarum vaginale L.: 746 ( 1753).

var. vaginalis

Verde, in Kew Bulletin 24: 67 (1970); Verde.: 493, t.

7 1 / A ( 1 97 1 ); Verde . : 443 ( 1 972); Verde . : 546 ( 1 974) .

Distribution: A. vaginalis occurs in the Transvaal

Lowveld through Swaziland and Tongaland to the Zulu-

land coast. It is widespread throughout the Old World

tropics.

Vouchers: Buitendag 302 (NBG, PRE); Clarke 262 (PRE, PRU);

Rogers 25092 (J); Venter 4593 (BLFU, PRE); Ward 1796 (NH, NU,

PRE).

Notes

( 1 ) The plant occurs as a semi-prostrate or decumbent

trailing herb in the herbaceous vegetation of open areas

in bush veld or in grassland up to 800 m. More com-

monly it becomes a weed of disturbed sandy or rocky

soils in these areas, i.e. in alluvia, old lands or along

roadsides. It flowers nearly all year round from July to

May.

(2) A. vaginalis has been introduced into America

where it is reported to have no economic importance

except as a noxious weed (Schubert 1980). In Africa,

however, Schubert (1980) notes that various species in-

cluding A. vaginalis are used locally as, ‘fodder, for all

kinds of domestic stock, used fresh, but preferably cut

after fruiting and stored as hay’. It is also regarded as an

excellent fodder for horses, but it is said to cause mucous

diarrhoea if given in excess in the young and immature

state during the rains. It is also known as Alyce clover,

and Allen & Allen (1981) state that it is considered as a

good cover crop in Malaysia to prevent erosion on clay

soils of rubber plantations.

2. Aiysicarpus zeyheri Harv. in Flora capensis 2:

230 (1862); Bak.: 170 (1871); Hutch. & Dalz.: 419

(1928); Bak.f.: 343 (1929); Robyns: 332(1948); J. Leo-

nard: 228, t. 13/C (1954b); Hepper: 587 (1958); Torre:

235 (1966); Verde.: 494, t. 71/C (1971); Verde.: 549

(1974). Type: Transvaal, Aapies River, Burke & Zevher

(K, syn.!; PRE!).

Although quoted as Burke & Zevher in Flora

capensis, these are two simultaneous but separate collec-

tions and are therefore syntypes.

Distribution: A. zeyheri occurs in the Transvaal and

northern Natal at altitudes between 900-2 400 m. It is

found from Pietersburg southwards to Pretoria, and from

Pretoria westwards to Zeerust in Sourish Mixed Bush-

veld. East of Pretoria it occurs in the North-eastern

Mountain Sourveld and Lowveld Sour Bushveld from

Lydenburg, Nelspruit and Barberton down through Swa-

ziland. It also occurs in the North-eastern Sandy High-

veld, Piet Retief Sourveld and related veld types from

near Middelburg through Ermelo to Louwsburg and

Mahlabatini in the south.

Vouchers: Du Plessis 1136 (PRE. PRU); Galpin 1158 (GRA, NH,

PRE, SAM); Leendert: 399 (BOL, J, PRE); Rudatis 131 (STE); Stern

995 (NBG).

Notes

(1) The plant is an erect to spreading herb of open

areas in bushveld or in grassland in sandy, rocky or

gravel soils, and often occurs in grazed, seasonally

burned grassland, along roadsides or railway lines and in

old lands. It flowers from October through to April.

(2) Watt & Breyer-Brandwijk (1962) report that A.

zeyheri is used as a remedy in South Africa for impo-

tence, while Allen & Allen (1981) quote a reference in

which the species has been used as a snake-bite remedy

in some African communities.

3. Aiysicarpus rugosus (Willd.) DC., Prodromus

systematis naturalis regni vegetabilis 2: 353 (1825);

Bak.: 171 (1871), pro parte; J. Leonard: 92, t. 12

(1954a); J. Leonard: 229 (1954b); Hepper: 587 (1958);

Lind & Tallantire: 82 (1962); Torre: 236 (1966);

Schreiber: 14 (1970); Verde.: 495 (1971); Verde.: 550

(1974); Compton: 273 (1976). Type: Guinea, Isert in

Herb. Willdenow (B, holo.).

Hedysarum rugosum Willd.: 1172 (1802).

Aiysicarpus violaceus (Forssk.) Schindl . : 13 (1928), pro parte;

Hutch. & Dalz.: 419 ( 1928), pro parte, non Hedysarum violaceum L.:

749 (1753); Forssk.: 136(1775).

KEY TO SUBSPECIES

Erect annual; calyx lobes with white cilia; SWA/Namibia

only subsp. rugosus

Decumbent spreading perennial with many flexuous stems;

calyx lobes with brown to orange cilia subsp. perennirufus

Bothalia 18,1 (1988)

3a. subsp. rugosus

J. Leonard in Bulletin du Jardin botanique de I’Etat, a

Bruxelles 24: 92, t. 12 ( 1954a); J. Leonard: 229 (1954b).

Distribution: the typical subspecies is recorded in the

Flora area only from the Grootfontein District in South

West Africa/Namibia. A specimen, Liebenberg 2789

(PRE), from Barberton in the Transvaal may also be

subsp. rugosus, but it is too underdeveloped to be identi-

fied with certainty. Subsp. rugosus is an erect annual in

open areas in thornveld savanna or in swamps.

Vouchers: Dimer 7628 (BOL, PRE, WIND); Merxmiiller & Giess

30098 (WIND).

3b. subsp. perennirufus J. Leonard in Bulletin du

Jardin botanique de l’Etat, a Bruxelles 24: 95 (1954a); J.

Leonard: 230, t. 13/D (1954b); Torre: 236 (1966);

Verde.: 496, t. 71/D (1971); Verde.: 552 (1974). Type:

Congo, Kivu, Rutshuru, Lebrun 9021 (BR, holo.; K!).

Alysicarpus wallichi Wight & Am.: 234 (1834); Harv.: 230 (1862).

Types: Wallich cat. 5763 a, b (K-WALLICH!).

Alysicarpus glaber E. Mey.: 125 (1836). Type: Transkei, between

Umzimvubu and Umsikaba Rivers, Drege (K!).

Alysicarpus violaceus (Forssk.) Schindl.: 13 (1928), pro parte;

Hutch. & Dalz.: 419 (1928), pro parte, non Hedysarum violaceum L.:

749 (1753); Forssk.: 136(1775).

Distribution: this subspecies is widespread in the east-

ern half of the Flora area, from the Transvaal, Swazi-

land, Natal to the eastern Cape. In the Transvaal it

occurs from Louis Trichardt and Tzaneen in the north

through Pietersburg and Potgietersrust to Lydenburg and

Komatipoort and to Swaziland in the south. Westwards it

occurs from Belfast and Ermelo through Pretoria to Rus-

tenburg with a slight extension into the higher altitude

Bankenveld north of Johannesburg and Potchefstroom.

The distribution in Natal is chiefly northern Natal from

Newcastle to Pongola (an extension of the range in the

south-east Transvaal) through the midlands to the coast.

It does not occur further west of a line extending from

Bergville, Estcourt and Richmond, i.e. into the uplands,

nor does it occur further north than Hluhluwe in Zulu-

land. Along the coast and adjacent interior, particularly

where there are projections inland of Valley Bushveld,

the distribution continues down to East London and

inland to the Harding, Umtata, Engcobo and Fort Beau-

fort Districts.

Vouchers: Acocks 9872 (NH, PRE); Flanagan 416 (GRA, PRE,

SAM); Galpin 12975 (BOL, PRE); Moll 1505 (NH, NU, PRE); Tyson

2002 (BOL, NBG, PRE).

Note

Verdcourt (1971) mentions that intermediates occur

between the two subspecies in the Flora of tropical east

Africa. He notes that a few examples have been found in

the Transvaal.

4. Alysicarpus glumaceus (Vahl) DC., Prodromus

systematis naturalis regni vegetabilis 2: 353 (1825);

Robyns: 332 (1948), pro parte; J. Leonard: 98, t. 13

(1954a); J. Leonard: 231, t. 13/E (1954b); Hepper: 587

(1958); Lind & Tallantire: 82 (1962); Torre: 236 (1966);

Verde.: 497, t. 71/E (1971); Verde.: 553 (1974). Type:

Yemen, Surdud, Forsskdl (C, holo.).

Hedysarum violaceum Forssk.: 136 (1775) non L., nom. illegit.

Type as above.

Hedysarum glumaceum Vahl: 106 ( 1 79 1 ).

19

Alysicarpus violaceus (Forssk.) Schindl.: 13 (1928), pro parte;

Bak.f.: 342 (1929), pro parte.

subsp. glumaceus var. glumaceus

Verde, in Kew Bulletin 24: 68 (1970); Verde.: 497, t.

71/E (1971); Verde.: 553 (1974).

Distribution: Alysicarpus glumaceus is found in the

hot lowlands of northern Botswana, Transvaal, Swazi-

land and Natal. In South Africa it occurs from Phala-

borwa in the north through Setara and between Louw’s

Creek and Komatipoort to eastern Swaziland; and in

Tongaland and coastal Zululand, south to Eshowe.

Vouchers: Bredenkamp 1847 (PRE); Cooley 726 (NH, NU); Schrire

909 (NH); Venter 3765 (ZULU); Ward 1891 (NH, NU).

Notes

(1) Only the typical taxon is found in the Flora area.

(2) This species occurs as an erect annual herb of open

places in bushveld or in grassland, and is often asso-

ciated with moisture, i.e, floodplains, depressions,

saline marshes or near streams. It is also a weed of old

lands or along roadsides. Flowering occurs from

November to June.

(3) Allen & Allen (1981) report that this is also known

as Alyce clover and, like A. vaginalis, is said to be

cultivated as a summer graze for domestic animals. Watt

& Breyer-Brandwijk (1962) state that A. glumaceus is

used in east Africa as a remedy for thrush and veld sores.

In Arabia and India the herb is used as an external appli-

cation to swollen feet.

Subtribe Lespedezinae

Lespedezinae {Hutch.) Schubert in Advances in

legume sy stematics 1 : 300 (1981).

Lespedezeae Hutch.: 486 (1964).

LESPEDEZA

Lespedeza Michx., Flora boreali-americana 2: 70

(1803); Hutch.: 487 (1964); Clewel: 365 (1966). Type

species: L. procumbens Michx.

Hedysarum L.: 745 (1753), pro parte.

The name Lespedeza is derived from Vincento Manuel

de Cespedes, governor of the Florida Colony from

1784-1790 and patron of the botanist Michaux. The

current spelling of the genus name presumably resulted

from illegibility or a printer’s error (Allen & Allen

1981).

Lespedeza cuneata {Du Mont.) G. Don, A general

system of gardening and botany 2: 307 (1832); Wilbur:

185 (1963). Type: Japan, Nagasaki, Oldham 328 (K!).

Hedysarum sericeum Thunb.: 287 (1784), nom. illegit.

Anthyllis cuneata Dumont de Courset: 100 ( 1 8 1 1 ).

Lespedeza sericea auct. non Benth.: 227 (1852): (Thunb.) Miq.: 49

(1867).

Distribution: L. cuneata is an escape from cultivation,

now naturalized in the Natal midlands in the Kokstad.

Ixopo and Pietermaritzburg Districts.

Vouchers: Edwards 12 (NH); Hilliard 3888 (NU); Schrire 1368

(NH); Tainton s.n. (NU).

20

Notes

(1) The plant occurs in old lands, along roadsides or

old railway lines. It is an herbaceous suffrutex to 0,8 m

high, flowering from February to April.

(2) L. cuneata is native to eastern Asia but is widely

cultivated in the United States. Known as Japanese bush-

clover, it is the most widespread and familiar of all the

species and is used extensively to control erosion on

roadside banks and similar areas (Wilbur 1963). It is also

held in high esteem as a forage, hay and protein supple-

ment to stockfeed (Allen & Allen 1981).

ACKNOWLEDGEMENTS

My sincere thanks to the following for much valuable

assistance and constructive criticism: Prof. E. Hennessy,

Dr B. Schubert, Dr H. P. Linder, Mrs E. Potgieter, Mr

G. Nichols, MrC. Buthelezi and Mr A. Ngwenya.

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SPECIMENS EXAMINED

Specimens are listed alphabetically according to the

name of the collector for each genus. Figures in brackets

after the collector’s number refer to the number of the

taxon in the text.

Desmodium

Acocks 9648( 1) PRE; 10118(1) NH; 10888(2) PRE; 10899(9)

PRE; 13332(1) PRE. Ahrens 23(1) NU. Allsopp 969(9) NU;

999(6) NH, NU, PRE; 101 7(9) NH; 1024(1) NH, NU.

Baijnath 185(1) NU, PRE. Balsinhas 2804(2) PRE; 3223a(4)

PRE. Barker 5155(1) NBG; 6138(2) NBG; 10012(9) NBG.

Baur 114(\) BOL, GRA. Bayliss 2757(1) NBG, PRE; 4667(1)

NBG. Blenkinson J 14446(1) J. Bokelmann, NBG 67393(1)

NBG. Bolus 7731(6) BOL; 8882(9) BOL; 8883(1) BOL;

8884(1) BOL; 11827(1) BOL, PRE; STE 24837(1) STE. Bos

1230(1) PRE, STE; 1253(1) PRE, STE. Botha 2583(1) PUC.

Bourquin 122(1) NU; 253(2) NU; 357(9) NU. Bourquin &

Lawson 1168(1) NH, PRE; 1171(9) NH, PRE. Breyer sub TRV

17848(10) PRE; sub TRV 79552(1) PRE. Buitendag 209(6)

NBG, PRE; 277(6) PRE, 7755(4) NBG, PRE.

Clarke 269(6) PRU; 371(1) PRE, PRU; 373(1) PRE, PRU;

464(2) PRE, PRU. Codd 1076(1) PRE; 6412(1) PRE; 9312(9)

PRE; 9339(1) GRA, PRE; 9362(2) NH, PRE. Codd & De

Winter 5144(10) PRE. Codd & Muller 325(1) PRE. Coleman

13(9) NH, PRE; 65(7) NH, PRE; 1103(3) NH. Collins sub TRV

9895(1) PRE. Comins 1956(9) GRA, PRE. Compton 25075(1)

NBG; 25675(1) PRE, SAM; 27257(1) NBG, PRE; 25692(1)

NBG, PRE; 30608(1) NBG, PRE. Cooper 204(1) PRE. Craw-

ford 368(2) PRE. Culverwell 1235(4) PRE.

Denley in J 30699(9) J. Devenish 1647(1) NU; 579(1) NH,

PRE. De Winter & Killick 8922(1) PRE. Dlamini in NBG 491 74

(1) NBG; in NBG 49172(1) NBG; in PRE 55645(1) PRE. Dohse

& Lindahl 94(1) NH, PRE; 95(1) NH, PRE. Dutton 47(9) NH.

Dyer 3153(1) PRE.

22

Bothalia 18,1 (1988)

Edwards D. 1312(1) NU, PRE; 3250(9) NU, PRE. Eicker s.n.( 9)

ZULU. Elan-Puttick 16(1) PRE. Eliovson J 26989(1) J. Ellis

2616a(2) PRE. Esterhuysen 12875(1) BOL, PRE. Evans 112(1)

NH; 181(9) NH.

Fairall 13(6) NBG. Fisher B.S. 590(2) NH, NU; 752(6) NH.

Fisher D.K. SAM 66046(1) SAM. Fitzsimons & Van Dam sub

TRV 26257(1) PRE. Flanagan 575(1) BOL, GRA, NBG, NU,

PRE, SAM; 2574(9) NH, PRE, SAM; 2608(1) PRE, SAM; PRE

55637(1) PRE. Forbes 35(2) NH, PRE; 605(2) NH ; STE 13501

(2) STE. Forrester & Gooyen 187(1) PRE. Fourcade 233(1)

BOL, GRA. Frith 121(7) J.

Galpin 1287(1) PRE; 3479(1) BOL; 6303(1) PRE; 9743(2)

PRE; 70090(1) PRE; 11005(9) PRE; 11019(2) PRE; 11024

(7) PRE; 11444(1) PRE; 14457(1) BOL, PRE; 77555(1) BOL,

PRE; 11933(1) PRE. Gemmell BLFU 5349(1) BLFU; BLFU

6050(9) BLFU. Gerstner 626(1) PRE; 2936(6) NH; 3845(1)

NH; 6005(1) PRE; NH 28779(1) NH. Getliffe 113(2) NU;

NU 27810(9) NU. Giffen 837(1) PRE. Gill NH 22592(1) NH.

Gillett Mrs. 1203(2) BOL, NH. Gillett J.B. 3156(8) BOL, PRE,

STE. Gordon-Gray 94(2) NU; 97(2) NU; 1047(6) NU; 1308

(7) NU. Grobbelaar 59(9) PRU; 66(7) PRE, PRU; 75(7) PRE,

PRU; 727(2) PRE, PRU; 279(9) PRE, PRU; 284(2) PRE, PRU;

649(9) PRE, PRU; 999(9) PRE, PRU; 1104(9) PRE, PRU;

1337(2) PRU; 1807(2) PRU; 7527(9) PRE, PRU; 2312(1)

PRU; 2324(1) PRU; 2329(9) PRU. Guy 34(1) NU, PRE.

Hanekom 2299(1) PRE. Harrison 244(9) NH, PRE. Hay garth

sub Wood 7067(4) BOL, PRE. Heygarth W. STE 9446(1) STE.

Henderson 55(1) BLFU. Hilliard 1095(9) NU; 1134(9) NU;

7227(7) NU; 1336(9) NU; 1344(1) NU; 7565(1) NH, NU;

1951(1) NU; 2639(1) NU. Hilliard & Burtt 3310(1) NU; 6852

(4) NU. Hitchins 391(6) PRE. Hofmeyer PRE 55621(1) PRE.

Holt 207(10) PRE. Howlett 76(1) NH, NU, PRE; 26(1) PRE;

706(1) NH, PRE. Huntley 195(1) NH, PRE; 656(2) NU.

Hutchinson 1759(2) BOL; 2245(8) BOL, PRE.

Jacobs 1757(1) PRE. Jacobsen 2671(1) PRE. Jarman & Guy 71

(2) NU. Jenkins 7068(1) PRE; sub TRV 7072(9) PRE. Johnson

369(2) NBG; 391(9) NBG. Junod 46(1) PRE; 752(1) PRE;

4357(1) PRE; 4382(1) PRE; 5265(7) PRE; 5267(10) PRE;

5270(6) PRE; sub TRV 5279(5) PRE; sub TRV 5284(10) PRE.

Kalf 35(1) NU; 56(9) NU; 55(2) NU. Keit NH 11142(9) NH.

Kemp 1153(1) PRE. Killick 144(1) NU, PRE; 1667(1) PRE.

Kluge 467(1) PRE, PRU; 455(7) PRE, PRU; 7222(10) PRE.

Kotze 57(4) WIND. Krige 139(10) J.

Lang sub TRV 32297(6) PRE. L’Ange 86(1) NU. Lawn 3(9)

NH; 52(2) NH; 72(1) NH; 769(7) NH; 27 7(7) NH; 752(2)

NH; 1807(9) NH; 2057(9) NH. Lawson 307(2) NU; 479(9)

NH; 704(9) NH. Leighton 2993(9) BOL; 5055(2) BOL, PRE.

Leistner 648(1) PRE. Letty 248(1) PRE; 460(1) PRE. Lewis

SAM 61374(1) SAM; SAM 61553(1) SAM; 61554(2) SAM;

SAM 68719(1) SAM; SAM 69720(1) SAM. Liebenberg 2374

(10) PRE; 2557(7) PRE; 2597(10) PRE; 2947(1) PRE; 5045

(9) PRE. Louw 2165(1) PUC, STE; 2730(1) PUC, STE.

Macgregor 64(9) NU. Martin A. J. 061(8) NU. Martin B. 191(2)

NBG. Mauve 4099(9) PRE. McClean 183(1) PRE; 439(9) NH;

496(9) NH. McDonald 118(6) E, K, NU, PRE. Medley Wood 93

(9) BOL, SAM; NH 163(9) NH; 209(9) BOL, SAM; 505(1)

BOL, SAM; 644(2) BOL, SAM; 790(1) BOL; 57 7(9) BOL,

SAM; 5070(4) NH, SAM; 5705(1) NH; 5729(9)NH; 3134(2)

NH; 4936(6) BOL, PRE, SAM; 4936(6) GRA; 5550(1) PRE;

5597(9) PRE; NH 6416(1) NH; 9755(7) NBG; 77545(9) PRE.

Merxmiiller & Giess 1862(10) WIND. Miller PRE 55638(2) PRE.

Miller O.B. 4279(1) PRE. Mogg 4399(1) PRE; 4597(7) PRE;

77927(9) PRE; 72700(9) PRE; 75072(9) PRE; 75455(9) PRE;

55954(1) J; 556277(1)J. Mohle 268(1) PRE. Moll 2471(6)

PRE; 2720(8) NU, PRE; 5459(1) NU, NH; 4767(9) NH, PRE;

5406(1) NH; 5470(2) NH; 5427(1) NH. Moll & Nel 5610(4)

NH, PRE. Moll & Strey 3896(8) NH, PRE. Morgan 70(1) PRE.

Morris 175(6) NU; 790(2) NU; 795(9) NU. Moss 3298(1) J;

3298(2) J, PRE; J 3299(9) J; 5500(1) PRE; J 14213(1) J;

J 15784(1) J; J 75075(1) J. Mtombeni 26(2) ZULU; 40(7)

ZULU. Muller 2352(1) PRE. Muller & Giess 544(4) PRE, WIND.

Netshiungani (1) VEND A. Nichols 423(8) NH; 455(3) NH;

477(1) NH. Nicholson 105(9) NH; 7200(7) PRE.

Oatley 26(9) PRE. Obermeyer sub TRV 29363(6) PRE; sub

TRV 30325(1) PRE; sub TRV 30371(1) PRE. Otto 124(1)

PRE.

Parkhouse NBG 16886(9) NBG. Phillips E.P. PRE 55667(1)

PRE. Phillips J. J 35347(1) J. Pole- Evans 970(1) PRE; 4549(1)

PRE. Pott 5334(6) PRE; 5555(1) PRE. Prosser 1840(1) PRE.

Randles 10(1) NU. Rehmann 8699(1) BOL. 7 lepton 5862(1)

PRE. Rogers 1259(9) PRE; 77606(9)J; 15059(9) J; 15060(2)

SAM; 75067(2) J; 75470(9) PRE; sub TRV 20707(6) PRE;

23564(6) NH; 26050(10) PRE; 28284(9) GRA, STE. 7 loss 78

(2) NU; 208(1) NU; 723(1) BLFU, NH, NU; 775(7) NU; 1679

(1) NH, NU, PRE; 7522(9) NH, PRE; 7994(1) NH, PRE; NU

25765(1) NU; NU 25769(1) NU. Ross & Moll 1830(9) NH,

PRE; 7547(8) NH, PRE; 226 7(2) NH, PRE. Rudatis 68(9) STE;

775(1) STE; 255(9) PRE; 554(1) PRE; STE 2339(1) STE.

Rump NH 21063(9) NH; 14-7-1929(9) NY; Feb. 1931(9) NU.

Rycroft 2420(4) NBG; 2601(2) NBG.

Sanderson 570(1) NH. S.A.R. PRE 55639(2) PRE. Scheepers

247(10) PRE, PRU; 245(1) PRE, PRU; 557(7) PRE, PRU;

Scheepers & Haasbroek Alkaloid survey SKF no. 1140(1) PRE.

Schelpe 51(9) BOL. Schlechter 3129(6) BOL, GRA, PRE; 5554

(7) PRE; 4742(1) PRE; 6665(7) BOL; 72705(8) SAM; 12275

(10) BOL, SAM. Schonken 75(1) STE. Schonland 845(1) GRA,

PRE. Schrire 153(6) NH; 262(2) NH; 263(2) NH; 264(2) NH;

265(2) NH; 266(2) NH; 267(7) NH; 265(8) NH; 272(2) NH;

273(1) NH; 274(1) NH; 275(2) NH; 276(2) NH; 277(9) NH;

278(2) NH; 255(7) NH; 254(9) NH; 285(9) NH; 256(9) NH;

257(9) NH; 255(2) NH; 259(9) NH; 290(8) NH; 297(2) NH;

292(8) NH; 293(2) NH; 294(9) NH; 296(9) NH; 297(8) NH;

552(2) NH; 555(1) NH; 540(2) NH; 550(2) NH; 556(9) NH;

552(9) NH; 555(4) NH; 386(8) NH; 392(2) NH; 596(7) NH;

599(1) NH; 655(5) NH; 667(10) NH; 6 77(2)NH; 776(9) NH;

77 7(2) NH; 555(9) NH; 557(9) NH; 552(7) NH; 555(6) NH.

Shuter 59(9) NU. Sidey 3876(2) PRE; 3976(9) NH, PRE; 4186

(9) PRE. Sim 4057(1) GRA, PRE. Smith 693(4) PRE; 2557(4)

PRE. Stanton 70(1) NU. Stephen, Van Groan & Schwabe 1197

(9) PRE. Stirton 39(1) NU; 462(9) PRE; 505(2) PRE; 556(9)

PRE; 7 755a(l) PRE; 77556(1) PRE; 5562(2) PRE; 5655(10)

NH. Strey 2996(1) NH, PRE; 5725(1) PRE; 3873(9) NH, PRE;

4705(1) PRE; 4559(2) NH, PRE; 4887(9) NH, PRE; 4971(9)

NH, NU, PRE; 5486(1) NH, PRE; 6554(2) NH, PRE; 7151(8)

NH, NU, PRE; 7394(2) NH, PRE; 7545(1) NH; 5550(1) NH,

PRE; 9618(1) NH, NU, PRE; 10752(1) NH, NU, PRE; 10937

(7) NH, PRE. Swierstra PRE 55605(1) PRE.

Taylor R.H. 85(9) NU. Theron 1600(1) PRE; 5672(1) ERE,

PRU. Thode A31 7(1) NH, PRE; STE 3802(9) STE; STE 3803

(1) STE; STE 3805(9) STE; STE 6443(1) STE. Thomcroft

893(1) NH; Gov. Herb. 11361(1) PRE. Thorpe NH 29880(1)

NH. Tiriley 343(8) PRE; 7457(4) WIND. Tomlinson 2/ 115(9)

NH. Trauseld 575(1) PRE. Tyson 1165(1) BOL; 7455(7)

SAM, STE; 7459(1) SAM, STE; 7525(1) PRE, SAM; 2744(1)

BOL; 2752(7) PRE, SAM; 2576(1) SAM; 5757(1) PRE.

Vahrmeijer 541(8) PRE; 667(4) NH, PRE. Vahrmeijer & Hardy

1627(9) PRE. Vahrmeijer & Tolken 559(4) PRE; 5252(9) PRE.

Van der Merwe D.D. PRE 58466(6) PRE. Van der Merwe P.

312(1) PRE. Van der Schijff 965(6) PRE, PUC; 1132(6) PRE;

7525(6) PRE; 2025(9) PRE; 2479(10) PRE; 2659(5) PRE;

Bothalia 18,1 (1988)

23

4332(1) PRE, PRU; 4941(1) PRE, PRU; 5066(1) PRE, PRU;

5976(1) PRE, PRU; 6437(1) PRE, PRU. Van Jaarsveld 12(4)

NBG; 675(4) NBG, PRE; 5500(1) NBG. Van Son sub TRV

30688(1) PRE. Van Vuuren 55(1) PRE. Van Wyk 915(1) PRE,

PUC. Venter F. 697(1) PRE. Venter H.J.T. 7(2) ZULU; 300

(2) BLFU; 570(9) NH; 654(9) NH; 729(9) ZULU; 552(7)

ZULU; 2275(2) ZULU; 2259(7) BLFU; 2424(1) PRE, ZULU;

5505(9) ZULU; 3304(1) ZULU; 5455(1) ZULU; 4547(9)

BLFU; 4993(4) PRE, ZULU; 4996(7) ZULU; 5255(8) BLFU;

5291 ( 8) PRE, ZULU; 5489(4) PRE, ZULU. Verdoom PRE

58467(1) PRE. Vogt PRE 55600(1) PRE. Vorster L.L. 15(1)

PRU. Vorster P. 2623 ( 8) PRE.

Wager H.A. sub TRV 22462(6) PRE; sub TRV 22988(1) PRE;

sub TRV 24101(2) PRE; PRE 55633(1) PRE. Wager V. C45(l)

PRE. Ward 222(9) NU; 584(2) NU, PRE; 755(7) NU, PRE;

1874(9) NH, NU; 1875(9) NH, PRE; 2755(1) NU; 2406(6)

NH, PRE; 4361(9) NH, PRE; 5381(1) NH, PRE; 5389(9) NH,

NU, PRE; 5495(1) NU; 5560(4) NH, NU; 6265(2) NU; 6575

(9) NH, NU, PRE; 6509(9) NH, NU; 7601(1) NU; 7696(9)

NH, NU. Watmough 486(2) PRE. Weeks 82(9) J. Wells 1429

(2) NU; 2550(1) GRA, NH, PRE; 5575(2) GRA, PRE; 4263

(2) NH; 4266(2) PRE. West 1089(1) PRE. Williams 8(1) NU.

Williams Miss sub TRV 7622(1) PRE. Wilson 55(9) NU. Wylie

sub Wood 8990(4) NH; 70555(4) GRA, PRE, SAM, STE.

Young PRE 55596(1) PRE; PRE 55607(1) PRE.

Pseudarthria hookeri

Acocks 1217 PRE; 232136 PRE. Adlam SAM 15562 SAM.

Ahrens 13 NBG. Allsop 800 NU.

Barrett 532 PRE. Baylis 1145 PRE. Biggs 197 PRE. Bolus 8885

BOL; 10949 BOL, PRE. Bos 1300 PRE, STE. Botha 1335

PRE, PUC. Boulle 8 NU. Bourquin 127 NU. Breyer sub TRV

17844 PRE; sub TRV 18387 PRE; PRE 55685 PRE. Britten

4753 GRA, PRE. Brown & Shapiro 464 PRE. Buitendag 498

NBG, PRE. Burtt Davy 1451 PRE; 1546a PRE; 2925 PRE.

Cholmondely PRE 55672 PRE. Clarke 47 PRE, PRU. Codd

5918 PRE; 6015 PRE. Codd <£ Muller 329 PRE. Coetzer211

BLFU, PRU. Coleman 66 NH. Collins sub TRV 9892 PRE.

Comins 88 NH, NU; 1948 GRA, PRE. Compton 24876 NBG;

26606 NBG, PRE; 27393 NBG, PRE; 31263 NBG, PRE. Craw-

ford 434 PRE. Cross 86 NBG.

Davison 91 PRE. Devenish 629 PRE. Dimock Brown 356 PRE;

452 PRE. Dohse & Lindahl 104 NH, PRE. Dyer 3152 PRE.

Eliov sen J 27186 J. Edwards 6 NU. Elan-Puttick 255 PRE.

Flanagan 2534 PRE, SAM; 2571 PRE. Frith 141 J.

Galpin 776 GRA, NH, SAM; 775 PRE; 75727 PRE; 14386

PRE; BOL 32274 BOL; BOL 322 75 BOL. Gerstner 6216 BOL.

Gordon-Gray 1372 NU. Grobbelaar 448 PRE, PRU; 2310 PRU.

Harding sub TRV 5181 PRE. Harrison 450 NH, PRE. Haygarth

STE 214 STE. Hemm s.n. VENDA, J. PRE. Hilliard 1325 NH, NU.

Hitchins 737 NH, PRE. Holt 42 PRE; 42 NH; 792 NH, PRE;

207 NH. Huntley 209 NH, NU, PRE. Hutton 50 GRA.

Indian collector NH 1 7731 NH; sub TR V 34012 PRE.

Jacobsen 2255 PRE. Jenkins sub TRV 7077 PRE; PRE 55702

PRE; PRE 55703 PRE. Johnson 368 NBG. Junod 2369 PRE;

4381 PRE.

KUlick 74 PRE. Kluge 410 PRE, PRU; 7755 PRE.

Lawn 305 NH. Leach 12109 PRE. Letty 441 PRE. Liebenberg

2379 PRE. Louw 2134 PUC, STE. Lovell PRE 55698 PRE.

Macgregor 55 NU. Maguire 689 J. McCallum PRE 55668 PRE.

McClean 129 PRE; 747PRE. Medley Wood 802 SAM, BOL, PRE;

1036 BOL, SAM ; NH 1491 NH; 7402 PRE; 9992 NH. Meeuse

9884 PRE. Moll 7527 NH, NU, PRE; 2744 NH, NU, PRE; 5265

NH, PRE. Morgan 6 7 PRE. Morris 786 NH, NU, PRE. Moss 18448

J . Mudd PRE 556 78 PRE. Muller 2475 PRE. Murray 690 PRE.

Nicholson 236 NH.

Obermeyer 319 PRE; 454 PRE.

Pegler 697 BOL, PRE. Pierce 6 PRE. Pole-Evans H. 1 7030 PRE.

Pooley & Joubert 2013 E, K, NU, MO.

Rehmann 6281 BOL. Repton 3482 PRE. Robbertse 102 PRU.

Rogers STE 13602 STE; 24054 PRE. Ross 85 NU; 277 NU;

777 BLFU; 750 NU. Rudatis 392 STE. Rump NH 10403 NH.

Scheepers 106 PRE, PRU; 520 PRE, PRU. Schlechter4551 BOL,

GRA; sub TRV 2220 PRE; sub TRV 13151 PRE. Schlieben

9478 PRE; 70755 PRE. Schrire 295 NH; 597 NH; 400 NH;

454 NH; 665 NH; 557 NH; 559 NH; 572 NH. Smith PRE

55688 PRE. Smuts 255 PRE; 507 PRE. Stephan 298 PRE;

557 PRE. Stirton 1751b PRE. Strey 3322 PRE; 4656 NH,

PRE; 9721 NH, PRE.

Theron 2351 PUC. Thode STE 3456 STE; STE 6367 STE.

Thomcroft 402 PRE; sub TRV 2849 PRE. Tyson 1440 PRE,

SAM, STE; 2046 SAM.

Venter 744 ZULU; 570 BLFU; 2500 ZULU; 5272 PRU,

ZULU. Verdoom 2442 PRE. Vahrmeijer 2403 PRE. Van Dam

PRE 55684 PRE. Van derMerwe 106 PRE. Van der Schijff 722

PRE; 4577 PRE, PRU. Van Elden 1 PRE. Van Rooyen 6 NU.

Van Son sub TRV 30436 PRE. Van WykA.E. 2568 PRU; 4044

PRE.

Wade NU 52896 NU. Wager C.30 PRE. Ward 2069 NU, PRE;

2510 NH, PRE; 7591 PRE. Watson 3 PRE. Wells 1111 NU,

PRE; 4270 NH; NU 34142 NU. Werdermann & Oberdieck

2776 PRE. Williamson 269 PRE. Wilson 20 NU. Wood 156 NU.

Alysicarpus

Acocks 9872(3) NH, PRE; 13160(2) PRE.

Barker 2742(3) BOL, NBG; 4252(3) NBG. Barrett 348(4) PRE.

Baur 572(3) SAM. Bolus 11074(2) BOL; 11829(2) BOL, PRE.

Bourquin 103(3) NU; 577(1) NU; 650(1) PRE. Bredenkamp

1737(4) PRE; 1739(4) PRE; 1847(4) PRE; 7565(1) PRE.

Bredenkamp <6 Van Vuuren 139(2) PRE; 295(3) PRE. Breyer

sub TRV 17056(1) PRE; sub TRV 17857(1) PRE; sub TRV

20903(3) PRE; PRE 55731(2) PRE; PRE 55733(2) PRE.

Buitendag 502(1) NBG, PRE. Burke PRE 9323(2) PRE. Burtt

Davy 804(2) PRE; 577(3) PRE. Buthelezi 774(3) NH; 142(3)

NH; 150(2) NH; 765(3) NH; 275(3) NH; 252(3) NH.

Clarke 131(2) PRU; 262(1) PRE, PRU; 265(3) PRE, PRU; 470

(3) PRE, PRU; 525(1) PRU. Codd & De Winter 4884(1) PRE;

4964(1) PRE. Coetzee B.J. 6065(4) PRE. Coetzee NH 35014

(3) NH. Coetzer 103(2) PRE; 755(1)PRE. Compton 28332(3)

PRE; 25545(1) NBG, PRE. Comins 436(3) NU; 7779(3)GRA,

PRE. Culverwell 1183(2) PRE; 1196(3) PRE; 7525(1) PRE.

Cuthbert NU 52685(3) NU.

24

Bothalia 18,1 (1988)

Devenish 929(3) PRE. De Winter 7692(3 ) PRE. Dinter 7628(3)

BOL, PRE, WIND. Dixon 21(3) NU. Du Plessis 1136(2) PRE,

PRU.

Elan-Puttick 246(2) PRE. Ellis 2616b(3) PRE.

Fisher 743(3) NH, NU. Flanagan 416(3) GRA, PRE, SAM;

2802(3) NH, PRE. Forbes 738(3) NH. Frankish 66(3) NU.

Galpin 1158(2) GRA, NH, PRE, SAM; 3408(3) BOL, PRE;

5779(3) PRE; 9729(3) PRE; 12975(3) BOL, PRE. Germishui-

zen 388(2) PRE; 557(2) PRE; 654(2) PRE. Gerstner 4209(3)

NH; 6844(4) PRE; NH 33781(2) NH. Gertenbach 7021(4)

PRE. Giffen 1070(3) PRE; 2096(3) PRE. Gordon-Gray 1128

(3) NU. Gordon Truscott 7a( 3) PRE. Grobbelaar 2(3) PRE,

PRU; 5(3) PRU; 323(3) PRE, PRU; 2520(3) PRU.

Hall & Sons PRE 55715(1) PRE. Hanekom 1913(3) PRE. Hil-

liard 1259(3) NU; 1848(3) NU; 1943(3) NU. Hitchins 566(3)

PRE. Huntley 1502(2) PRE. Hutton 1031(3) GRA, NBG.

Jacobsen 1588(3) PRE. Jacobsz 768(3) PRE; 505(3) PRE.

King 57(3) PRE; 64(3) PRE.

Lawn 1120(3) NH; 1908(3) NH. Leeman 23(3) PRE. Leendertz

399(2) BOL, J, PRE; 407(3) PRE, SAM; sub TRV 5985(2)

PRE; sub TR V 8754(2) PRE; sub TRV 11298(2) PRE. Lennox

NU 28057(3) NU. Liebenberg 2788(2) PRE; 5295(1) PRE;

5295(3) PRE; 5505(3) NH; 5572(2)NH; 5452(2) PRE; 8721

(3) PRE.

Mauve 5298(3) NH. Medley Wood 274(3) NH; 6091(3) BOL.

Merxmuller & Giess 1862(3) WIND; 30098(3) WIND. Mogg

5960(3) NH, PRE; 6143(3) PRE; 6605(3) PRE; 15064(3)

PRE. Moll 1057(3) NU, PRE; 1505(3) NH, NU, PRE; 7656(3)

NU, PRE; 2595(3) NU, PRE. Moss 383(3) J ; 8490(2) J ; 75274

(2) J. Moss <£ Rogers 51 7(1) J ; 602(1) J; 995(3) J.

Obermeyer 148(3) PRE; 7244(3)PRE. Obermeyer & VanNou-

huys sub TR V 27805(3) PRE.

Pegler 163(3) PRE; 7459(3) PRE. Pelletier J 30510(3) 1. Penny

Col. Herb. 2388(3) PRE. Pooley 726(4) E, K, NH, NU. Pott

5075(3) PRE; sub TRV 15081(3) PRE; 5545(2) PRE. Preller

139(3) PRE. Puree PRE 55727(1) PRE.

Ranger 302(3) PRE. Rattray 616(3) GRA. Reachmoor PRE

11854(1) PRE. Rehmann 461(3) BOL. Repton 87(3) PRE;

555(3) PRE; 2767(2) PRE; 2755(3) PRE. Rogers 301(3) GRA;

sub TRV 5165(3) PRE; 10931(3) PRE; 14394(3) J, PRE;

18294(3) J; 21921(2) PRE; 24860(3) PRE; 25092(1) J.

Rudatis 131(2) STE; 7565(3) STE; 1729(3) PRE. Rump NH

20290(3) NH; NH 20328(3) NH.

Scheepers 176(3) PRE, PRU. Schlechter 3121(3) PRE; 5227(3)

BOL, PRE. Schrire 269(3) NH; 270(3) NH; 277(3) NH; 405(3)

NH; 667(3) NH; 574(3) NH; 909(4) NH; 7720(2)NH; 1320

(3) NH. Shirley 304(3) NU; NU 32878(3) NU. Sim 19436(3)

PRE; 20175(3) PRE, NU. Smith 1451(3) PRE; 2445(4) PRE.

Smuts & Gillett 2214(3) PRE. Steyn 995(2) NBG. Stirton 5076

(3) PRE; 5190(3) PRE; 5757(3) PRE; 10516(3) NH. Strey

4505(3) NH; 9775(2) NH, PRE.

Taylor 55(4) NH. Theiler PRE 55709(3) PRE. Thode STE 6662

(3) STE. Tinley 751(3) NU. Tyson 23(3) GRA, NH, PRE; 7447

(3) SAM, STE; 1196(3) PRE, SAM; 2002(3) BOL, NBG, PRE.

Vahrmeijer & Tolken 905(1) NH, PRE. Van Dam 21137(4) PRE.

Van der Schijff 437(1) PRE; 7596(1) PRE; 5290(3) PRE, PRU;

5291(2) PRE, PRU; 5297(3) PRE. Van Rensburg 2(3) NU.

Van Vuuren 531(2) NH. Van Wyk 4712(1) PRE. Venter 774(1)

ZULU; 7699(3) NH, PRE; 1969(3) ZULU; 3436(3) ZULU;

5765(4) ZULU; 4595(1) BLFU, PRE. Verdoom 135(3) PRE.

Wahl sub TRV 15525(3) PRE; PRE 55723(3) PRE. Ward 1512

(3) NH, PRE; 1796(1) NH, NU, PRE; 1891(3) NH, NU; 2426

(1) NH, NU, PRE; 5505(1) PRE. Wilms 5873(3) PRE. Wylie

NH 28026(3) NH.

Young A331(3) PRE.

Zey her SAM 32864(2) SAM.

SAGP/SAAB 1/113(3) PRE, PRU.

Lespedeza cuneata

Edwards 12 NH.

Hilliard 3888 NH, PRE.

Schrire 1368 NH.

Tainton s.n. NU.

Bothalia 18,1:25-29(1988)

The naturalized species of Lupinus (Fabaceae) in southern Africa

C. H. STIRTON*

Keywords: Fabaceae, Lupinus, naturalization, southern Africa

ABSTRACT

Four species of Lupinus have become naturalized in southern Africa: L. consentinii Guss., L. angustifolius L., L.

pilosus Murray and L. luteus L. Their morphological variation and geographical distribution is discussed and a key is

provided.

UITTREKSEL

Vier Lupinus-spesies het in suidelike Afrika genaturaliseerd geraak: L. consentinii Guss., L. angustifolius L., L.

pilosus Murray en L. luteus L. Hulle morfologiese variasie en geografiese verspreiding word bespreek en ’n sleutel word

voorsien.

INTRODUCTION

The genus Lupinus comprises some 200 species of

annual and perennial herbs, or sometimes subshrubs

(Polhill 1976; Bisby 1981). It is best represented in the

western parts of North and South America, with a second

centre in Mediterranean Europe with some extensions

into the highlands of eastern Africa (Kay 1979). Twelve

species of European and African origin are recorded

from Africa (Gladstones 1974; Plitmann 1981; Williams,

Demissie & Harbome 1983).

Lupins have been used by man for thousands of years,

principally for fodder, green manure, ornament and as

grain legumes. A few species have become naturalized

weeds in Mediterranean Europe, the south-east United

States of America, the Andes, Australia, New Zealand

and South Africa (Kay 1979).

The taxonomy of the genus has been confused for a

long time. This was exacerbated to some extent by the

development and growing importance of large-seeded

lupins as a source of edible protein (Gladstones 1974).

The most important grain legumes are Lupinus albus L.,

L. angustifolius L., L. luteus L. and L. mutabilis Sweet.

Recent studies on the flavonoid chemistry of the Old

World species of Lupinus provide useful taxonomic

markers (Williams et al. 1983) to separate the species

cultivated or naturalized in southern Africa. L. albus has

flavonols only, L. angustifolius and L. luteus have fla-

vones and flavonols, whereas L. consentinii and L. pilo-

sus are characterized by the presence of flavone C-glyco-

sides, luteolin and novel 2'hydroxyflavone. L. angusti-

folius is chemically distinct from all the other Old World

species as it accumulates diosmetin derivatives as major

leaf constituents (Williams et al. 1983). It is chemically

the most variable species and is divided into a number of

varieties (Plitmann 1966).

* B. A. Krukoff Botanist, The Herbarium, Royal Botanic Gardens,

Kew, Richmond, Surrey, TW9 3AE, UK. Current address: CSIR Unit

for Plant Growth and Development, Department of Botany, University

of Natal, P.O. Box 375, Pietermaritzburg 3200.

MS. received: 1986.09.05.

The commercial exploitation of lupins in South Africa

began early this century but it was C.C.P. Wagener who

first drew attention to the importance of the crop (Van

Vuuren 1962). He used bitter lupins as fodder for sheep.

Prior to that the crop had been used as a green-manuring

crop in vineyards and orchards. By 1947 lupins had be-

come part of crop rotations in the wheatbelt of the Cape

(Preller 1949; Henning 1949). The introduction of sweet

lupins at about that time did much to increase wheat

yields and resulted in a substantial increase in the

number of livestock animals that could be maintained in

the region.

Fourteen cultivars were available in 1962 (Van Vuuren

1962): 1, L. albus: albus (German origin), carstens,

pflugs gela, pflugs ultra, and S.S.K. (white sweet); 2, L.

angustifolius: borre, jackalsfontein, ligvoet, S. E. blue

No. 1 , B. resistant, blue sweet (German origin) and 7002

(white); 3, L. luteus: S. E. yellow No. 1 and weiko III.

The weiko III cultivar is an alkaloid-free, white-seeded

and early maturing strain and is the dominant lupin culti-

vated in a number of European countries. A similar

strain named Stellenbosch-Elsenburg Geel-1 is popular

in South Africa. Lupins are still popular in the western

Cape and have begun to be used in other parts of South

Africa, for example, in the south-eastern Transvaal

Highveld (VanZyl 1973).

It was perhaps inevitable that as lupins became

increasingly popular some species would become natura-

lized. This study, based on herbarium material as well as

fresh specimens collected during three field trips to the

region in 1975, 1976 and 1984, is a first attempt to

monitor the spread of naturalized lupins in the Cape. It

should be stressed, however, that the distribution maps

(Figures 1—4) are only an indication of the extent of

naturalization. Botanists, farmers and extension-officers

are urged to collect herbarium material from areas which

are not recorded in this survey. The author will provide

identifications of any well pressed, vouchered material

that is sent to him.

The descriptions and keys are based on fresh material

and cover naturalized species only.

26

Bothalia 18,1 (1988)

FIGURE 1. — Naturalized distribution of L. angustifolius in South

Africa.

FIGURE 2. — Naturalized distribution of L. consentinii in South Af-

rica.

FIGURE 3. — Naturalized distribution of L. pilosus in South Africa.

KEY TO SPECIES

la Flowers blue, rarely white:

2a Leaflets 8—11, 40-80 X 11-21 mm, flattened out, oblanceolate, upper surface finely sericeous; stipules

10— 1 1 mm long; flowers 18— 19 mm long, keel obtuse to acute; bracteoles extending past the sculpturing

pattern on the wing petals I . consentinii

2b Leaflets 7-9, 25-27 x 4, 5-5,0 mm, folded, linear-spathulate, upper surface glabrous; stipules 7-8 mm

long; flowers 15 mm long, keel rostrate; bracteoles shorter than the sculpturing pattern on the wing petals

2. angustifolius

Bothalia 18,1 (1988)

27

1 b Flowers pink or yellow:

3a Flowers pink; plants densely hirsute, hairs up to 4,5 mm long; leaflets 14—20 mm wide, obovate; bracteoles

pinkish; tip of keel brown; seeds rough 3. pilosus

3b Flowers yellow; plants sparsely hirsute; leaflets 5-12 mm wide, narrowly obovate; bracteoles black; tip of

keel black; seeds smooth 4. luteus

KEY TO DISTINCTIVE FEATURES

A. Apex of petiole, where the leaflets emerge, distinctly

domed consentinii

Apex of petiole, where the leaflets emerge, flattened

angustifolius, luleus, pilosus

B. Stipules recurved angustifolius, luleus

Stipules straight consentinii, pilosus

C. Stipules shaggy pilosus

Stipules pubescent angustifolius, consentinii, luteus

D. Upper surface of leaflets glabrous angustifolius

Upper surface of leaflets pubescent consentinii, luteus, pilosus

E. Flowers blue angustifolius, consentinii

Flowers pink pilosus

Flowers yellow luteus

Flowers white angustifolius

1. Lupinus consentinii Guss ., Florae siculae pro-

dromus 2: 398 ( 1 828). For a detailed synonym listing see

Gladstones: 29 (1972) and Gladstones: 21 (1974).

Robust annual up to 1 m tall, erect with vigorous

lateral branching, covered in shortly villous 1 mm long

patent hairs. Stipules 10—11 mm long, linear-subulate,

straight, arching back towards branch, pubescent. Leaf-

lets 8— 11, 40— 80 x 14—21 mm, oblanceolate, midrib

prominent, apex blunt with a point, base cuneate, finely

sericeous above and below; petiole 85—135 mm long,

terete for most of its length but flattened adaxially to-

wards the base. Inflorescences axillary, verticillate, low-

est verticil quintuplicate sometimes subaltemate,

25— (40)-flowered, 60—180 mm long. Flowers 18—19

mm long, on stout 5 mm long pedicels; bract 6-7 mm

long, lanceolate, early caducous; bracteoles linear. Ca-

lyx 10—12 mm long; upper lobes deeply divided; lower

lobes much longer, fused but shallowly 3-toothed, vil-

lous. Corolla 15-16 x 14-18 mm, dull' violet-blue

darkening with age, nectar patch white turning lilac after

anthesis; scentless. Standard 17—18 mm long and wide,

very broadly ovate, scarcely clawed or auriculate, apex

rounded. Wing petals 15 mm long, barely longer than

keel petals, dull violet-blue; fused at apex; claw 2 mm

long; sculpturing on upper basal parts comprising 10

rows of 14-20 intercostal lunae. Keel petals 14 mm

long; fused halfway along the base to the acute apex, tip

pigmented. Androecium 15 mm long; anthers dimorphic,

comprising 5 basifixed, linear anthers 1 ,2 mm long, and

5 medifixed, reniform anthers 0,5 mm long. Pistil 13

mm long; ovary 8 mm long, densely shaggy, height of

curvature 5 mm. Fruits 40—50 x 13—15 mm, densely

villous, apple-green, rostrate, segmented. Seeds 4,

rough. Figure 5B.

Lupinus consentinii is most commonly confused with

L. angustifolius but it is easily separated trom that spe-

cies by its much longer, wider, open, oblanceolate leaf-

lets and obtuse to acute keel petals. Flowering takes

place in August and September.

2. L. angustifolius L., Species plantarum: 721

(1753); Gladstones: 28 (1972). Holotype: Linnaean Col-

lection 898.7 (LINN).

Erect annual up to 150 mm high, compact, with pro-

fuse lateral branching; finely appressed puberulent. Stip-

ules 7-8 mm long, linear-subulate, recurved, pubes-

cent, glabrous above. Leaflets 7—9, 25-27 x 4,5— 5,0

mm, conduplicate, linear to linear-spathulate, upper sur-

face glabrous, lower surface sparsely sericeous; petiole

35—45 mm long, adaxial surface flattened, somewhat

canaliculate. Inflorescences axillary, on short shoots,

with flowers generally alternate towards base but subver-

ticillate towards apex, 5 — 30-flowered, 50—170 mm

long, subsessile but elongating in fruit. Flowers 12—15

mm long, on 2—4 mm long pedicels; bract 7 mm long,

oblanceolate to ovate, caducous; bracteoles 2-3 mm

long, oblong. Calyx 9- 10 mm long; upper lobes deeply

divided; lower lobes united into a longer lip, entire or

irregularly 3-toothed, bluish, appressed pubescent. Co-

rolla 12-15 mm long, 9 mm wide, lavender to bluish,

rarely pure white, upper margin of wing petals turns pink

after anthesis. Standard 12—13 x 9-10 mm, elliptic,

shortly clawed, auricles absent, apex truncate. Wing pe-

tals 12 mm long, 5 mm wide, longer than keel petals,

lavender, fused at apex; claw 2 mm long; sculpturing on

upper basal parts comprising 5 rows, each with 20—25

intercostal lunae. Keel petals 1 1 mm long, 3,5 mm wide,

rostrate, sharply upcurved, tip deep purple. Androecium

12 mm long; anthers dimorphic, comprising 5 basifixed,

elongated, 0,9 mm long anthers and 5 short, medifixed,

0,3 mm long anthers; pollen orange. Pistil 12 mm long;

ovary 4 mm long, densely shaggy, height of curvature 3

mm. Fruit 40—50 x 7—8 mm, villous, pale apple-

green. Seeds 5—7, smooth. Figure 5A.

The differences between this species and the only

other blue-flowered species, L. consentinii , are given

under the former. White-flowered morphs are known to

occur in L. angustifolius. It is distinguished from the

other three species by the glabrousness of the upper sur-

face of the leaflets.

Flowering occurs between August and October. This

is the most weedy of the naturalized lupins in South

Africa. It can eventually form large stands where it has

become established.

3. L. pilosus Murray in L., Systema vegetabilium,

edn 13: 545 (1774). The complicated synonymy and

typification of this species is outlined in Gladstones

(1974).

Erect, 350—700 mm tall, sparsely branched annual.

Stems, petioles and peduncles all clothed in 4,5 mm

long, soft white hairs. Stipules 10-14 mm long, linear-

subulate, straight, more or less parallel to the subtending

branch, shaggy. Leaflets 9, 40— 60 x 14—20 mm, obo-

vate, finely appressed villous on both sides, partly con-

duplicate; petioles 50-60 mm long, adaxially flattened

28

Bothalia 18,1 (1988)

angustifolius

FIGURE 5. — Naturalized species of Lupinus in South Africa: A, L. angustifolius', B, L. consentinii; C, L. luteus, D, L. pilosus. 1, flower, X

2; 2, bracteole, x 4; 3, standard, X 2; 4, wing petal showing lunate sculpturing, X 2; 5, keel petal, X 2; 6, junction of the petiole with the

stem, showing shape and orientation of the stipule, x 2; 7, upper surface of middle leaflet, x 2; 8, basic shape of leaf; 9, fruit, x 1.

Bothalia 18,1 (1988)

towards the base. Inflorescences axillary, lax, 12—16

mm long; peduncle stout; 20 -30- flowered, subverticil-

late proximally, verticillate distally. Flowers large, 20

mm long, 25 mm high; pedicels 6-7 mm long; bract

10—13 x 5 mm, boat-shaped, caducous; bracteoles 5

mm long, lanceolate. Calyx 13 mm long; upper lobes

deeply cleft, widely spaced; lower lobes united, entire,

longer than upper ones; tinged with pink. Corolla pink.

Standard 22-24 mm long and wide, ovate, pink with

large white nectar guide, turning rose after anthesis,

grooved on inner face; claw 4-5 mm long, broad. Wing

petals 18-19x9-10 mm, pale pink but whiter towards

the claw; claw 4 mm long, twisted. Keel petals 16x5

mm, rostrate; apex brown-tipped. Androecium 18 mm

long; anthers dimorphic, comprising 5 basifixed, elon-

gated anthers 2 mm long, alternating with 5 shorter, 1

mm long, medifixed anthers; pollen orange. Pistil 14

mm long; ovary 10 mm long, shaggy; stigma penicillate,

forward sloping; height of curvature 6 mm. Fruits

50—80 x 20—25 mm, hirsute, yellow-green. Seeds

3-4. Figure 5D.

Lupinus pilosus is distinguished by its shaggy stipules,

densely hairy fruits and pink flowers. Flowering occurs

from late July to October.

4. L. luteus L., Species plantarum: 721 (1753);

Gladstones: 28 (1972) for full synonymy. Holotype: Lin-

naean Collection 898.8 (LINN).

Herbaceous annual up to 700 mm tall; rosetted in-

itially but later becoming erect with vigorous basal

branching; lightly hirsute. Stipules of stem leaves 12-20

x 2—5 mm, smaller on rosette leaves, subulate, re-

curved, pubescent but glabrous on lower half of inner

face. Leaflets 7-9(10), 33-55 x 5-12 mm, narrowly

obovate, mucronate, villous above and below, partly

conduplicate; petioles 70-90 mm long, adaxially flat-

tish, shallowly channelled. Inflorescences axillary,

13— 17 mm long, elongating in fruit, verticillate,

25— 45-flowered; peduncle 5-7 mm long. Flowers

14— 15 mm long, yellow; pedicels 2—3 mm long, weak;

bract 4 mm long, obovate, scoop-like, early caducous,

black; bracteoles 4-5 mm long, linear, black. Calyx

7-8 mm long; lobes equal, pubescent; upper lobes

deeply divided; lower lobes united, shallowly 3-toothed,

tips of lobes tinged with black. Corolla chrome-yellow,

sweetly scented. Standard 15—17 x 10—11 mm, nar-

rowly ovate; claw 3 mm long; apex emarginate, front of

standard channelled; auricles weakly developed. Wing

petals 15-16 x 3-4 mm; claw 3-4 mm long; tips

fused; sculpturing in upper basal parts with 6—7 rows of

15— 20 intercostal lunae. Keel petals 13-14 x 3-4

mm, rostrate, sharply upcurved, tip black. Androecium

14 mm long; anthers dimorphic, comprising 5 basifixed,

elongated, 1,5 mm long anthers alternating with 5 medi-

fixed, ovate, 0,3 mm long anthers; pollen yellow. Pistil

12 mm long; ovary 6 mm long, densely appressed

shaggy; height of curvature 6 mm. Fruits 45-50 x

11 — 13 mm, reticulate, densely villous, olive-green.

Seeds 2-4, smooth. Figure 5C.

29

Lupinus luteus is distinguished by its bright yellow

flowers and distinctly reticulate, densely villous fruits.

Flowering occurs in August and September.

ACKNOWLEDGEMENTS

I would like to thank Miss D. Snijman and Miss P.

Perry (Kirstenbosch) for their generous help during this

study. The guest house facilities at the National Botanic

Gardens, Kirstenbosch have been host to many botanists;

their utility to visiting scientists cannot be overestimated.

I thank the Curators and staff of BM, BOL, GRA, K,

MO, NH, NU, PRE and STE for their help in locating

material and its loan where available. Miss Angela Beau-

mont kindly drew the composite plate, published with

permission of R. B. G. Kew. Finally, I wish to acknowl-

edge the support of the Royal Botanic Gardens, Kew,

and the Missouri Botanical Garden in sustaining my Re-

search Fellowship.

REFERENCES

BISBY, F.A. 1981. Genisteae (Adans.) Benth. In R. M. Polhill &

P. H. Raven, Advances in legume systematics: 409—425. Royal

Botanic Gardens, Kew.

GLADSTONES, J. S. 1972. Lupins in Western Australia. Department

of Agriculture, Western Australia, Bulletin No. 3834.

GLADSTONES, J. S. 1974. Lupins of the Mediterranean region and

Africa. Technical Bulletin No. 26, Western Australian Depart-

ment of Agriculture.

GUSSONE, G. 1828. Florae siculae prodromus 2: 398. Naples.

HENNING, P. O. 1949. Lupines in the winter rainfall area. Farming

in South Africa 24: 227-236.

KAY, D.E. 1979. Food legumes. Tropical Products Institute, London.

LINNAEUS, C. 1753. Species plantarum: 721. Stockholm.

LINNAEUS, C. 1774. Systema vegetabilium, edn 13: 545. Gottingen.

PLITMANN, U. 1966. Studies in the taxonomy of the Leguminosae of

Israel. Israel Journal of Botany 15: 25-30.

PLITMANN, U. 1981. Evolutionary history of the Old World Lu-

pines. Taxon 30: 430-437.

POLHILL, R. M. 1976. Genisteae (Adans.) Benth. and related tribes

(Leguminosae). Botanical Systematics I: 143—368.

PRELLER, J. H. 1949. Lupines. Farming in South Africa 24: 25-29.

VAN VUUREN, P. J. 1962. How to identify lupins. Farming in South

Africa 38,9: 61 - 64.

VAN ZYL, L. G. 1973. Lupins in the south-eastern Transvaal High-

veld. Farming in South Africa 48: 4-7, 10.

WILLIAMS, C. A., DEMISSIE, A. & HARBORNE, J. B. 1983.

Flavonoids as taxonomic markers in Old World Lupinus species.

Biochemical Systematics and Ecology 1 1 : 22 1 -23 1 .

SPECIMENS EXAMINED

Bos 28 (2) STE. Cloete s.n. (4) STE. Markotter s.n. (2) STE 19310.

Snijman 882 (2) K, NBG, PRE; 883 (4) K, NBG, PRE; 884 (2) K,

NBG, PRE; 886 (2) K, NBG, PRE; 903a (2) K, NBG, PRE; 908 (4) K,

NBG, PRE. Stirton 5021 ( 1) PRE; 5036 (4) K, PRE; 5037 (2) K, PRE;

5882 (2) K, PRE; 5940 (2) K, PRE; 6129 (2) K, PRE; 10751 ( 1 ) NBG;

10752 (3) NBG; 10753 (4) NBG; 10762 (2) NBG; 10765 (4) NBG.

Vahrmeijer 2321 (2) PRE. Cultivated specimens: Codd s.n. (3) K.

Henning s.n. ( 1) K.

Bothalia 18,1: 31-36(1988)

Studies in the genus Lotononis (Crotalarieae, Fabaceae). 1. Three new

species of the section A ulacinthus from the Cape Province

B-E. VAN WYK*

Keywords: Cape Province, Lotononis section Aulacinthus, Fabaceae, new taxa

ABSTRACT

Three new species of the section Aulacinthus (E. Mey.) Benth. of Lotononis (DC.) Eckl. & Zeyh. are described: L.

comptonii B-E. van Wyk, L. dahlgrenii B-E. van Wyk and L. dissitinodis B-E. van Wyk. These species are related to L.

gracilis (E. Mey.) Benth. and L. rigida (E. Mey.) Benth. and are all endemic to marginal fynbos areas of the south-western

Cape. The section now comprises seven species.

UnTREKSEL

Drie nuwe soorte van die seksie Aulacinthus (E. Mey.) Benth. van Lotononis (DC.) Eckl. & Zeyh. word beskryf: L.

comptonii B-E. van Wyk, L. dahlgrenii B-E. van Wyk en L. dissitinodis B-E. van Wyk. Hierdie soorte is verwant aan L.

gracilis (E. Mey.) Benth. en L. rigida (E. Mey.) Benth. en is almal endemies in marginale fynbosgebiede van die Suidwes-

Kaap. Die seksie behels nou sewe soorte.

INTRODUCTION

The section Aulacinthus (E. Mey.) Benth. of Lotono-

nis (DC.) Eckl. & Zeyh. comprises a small group of

poorly known fynbos shrubs. Species of this section are

remarkably similar in their leaves and habit to some

species of Lebeckia Thunb., Wiborgia Thunb. and Aspa-

lathus L.

When Bentham (1843) changed the status of Aulacin-

thus E. Mey. to a section of Lotononis, he remarked on

its similarity to other genera of the Crotalarieae but also

noted its distinguishing floral and fruit characters. The

upper calyx lobes are usually fused in pairs on either side

and the fruit is relatively short, laterally inflated and has

the lower suture sunken to various degrees. In terms of

fruit characters, the verrucose upper suture is perhaps the

most significant and useful diagnostic feature. The seeds

of all the species are distinctly tuberculate and not

smooth as in other genera of the Crotalarieae. Bentham

(1843) and Harvey (1862) listed for section Aulacinthus,

three species [L. gracilis (E. Mey.) Benth., L. rigida (E.

Mey.) Benth. and L. viborgioides Benth.] to which

Diimmer (1913) added one [L. leucoclada (Schltr.)

Diimmer, transferred from Lebeckia],

As part of a continuing taxonomic study of the genus

Lotononis, I investigated the section Aulacinthus and

found three undescribed species that clearly belong to

this group. These are described below.

The circumscription of Aulacinthus and its status as a

section need to be reconsidered. As a group it gradually

merges (through yet another group of undescribed spe-

cies) with the section Polylobium (Eckl. & Zeyh.)

Benth. This problem, however, falls beyond the scope of

the present paper.

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

Johannesburg 2000.

MS. received: 1987.08.17.

Lotononis comptonii B-E. van Wyk, sp. nov. L.

gracili (E. Mey.) Benth. valde affinis, sed habitu laxiore

sparsiore, foliis ramunculisque sparse strigillosis (non

dense albo-pubescentibus), stipulo unico in quoque nodo

(in L. gracili stipulis omnino destitutis) et vexillo auran-

tiaco-rubro (in posteriore flavo) differt.

TYPE. — Cape, 3320 (Montagu): Laingsburg to

Seweweekspoort Road, 9,5 km after entrance to Rietvlei

Farm (-AD), 12.10.86, B-E. van Wyk 2160 (PRE,

holo.; BOL, C, JRAU, K, M, MO, NBG, SAAS, STE,

iso.).

Lax, sparsely branched woody shrub, up to 0,4 m tall.

Branches slightly corky, yellow-brown; young twigs

sparsely strigillose. Leaves digitately trifoliolate (very

rarely 4 or 5-foliolate); petiole (7—) 18-25 (-42) mm

long, persistent for some time after the leaflets have been

individually shed; leaflets obovate, narrowly oblong, or

narrowly elliptical, (4—) 8—15 ( — 21) mm long, (1— )

2—3 (-5) mm wide, abaxially sparsely strigillose,

adaxially glabrous. Stipules consistently present, single

at each node, elliptic-oblong to narrowly lanceolate,

(2 — ) 5 — 7 (—9) mm long. Inflorescences terminal,

rarely subterminal, long-pedunculate, 60-150 mm long,

sparsely (3-) 4—6 (—8) -flowered; bracts 2-3 (-5)

mm long; bracteoles absent. Flowers 10-12 mm long,

orange-red and yellow; pedicel 2-5 mm long. Calyx

6-8 mm long, with the lateral lobes on either side fused

much higher up in pairs, minutely strigillose. Standard

broadly ovate to orbicular, 8—12 mm long, longi-

tudinally striate, orange to bright orange-red. Wing

petals broadly oblong, slightly shorter than the keel;

apex obtuse; sculpturing upper basal and upper central,

in 4-5 rows of intercostal lunae. Keel petals broadly

elliptical, auriculate and pocketed near the base. Anthers

dimorphic. Pistil 10—12 mm long; ovary short, oblong-

ovate, 4-6 mm long. Fruit 12-15 mm long, 4-5 mm

wide, laterally much inflated, lower suture sunken,

upper suture verrucose, glabrous. Seed suborbicular in

side view, up to 2 mm long, distinctly tuberculate, red-

dish brown (Figure 1).

32

Bothalia 18,1 (1988)

FIGURE I . -Lolononis comptonii. Al, A2, flowering branches; Bl , B2, leaves, showing the single stipule and vestiture; Cl , C2, calyx

opened out, with the upper lobes to the right (Cl) and to the left (C2), vestiture not shown; Dl, D2, standard petals; El, E2, wing

petals; FI , F2, keel petals; G1 , G2. pistils; HI , H2, fruit, in side and top view; SI. S2, seeds, showing the tuberculate surface; Tl,

T2, bracts. (A I, B2, Cl, C2, D2, E2, F2,G1 &TI from BVW2I60 ; A2 & T2 from BVW 2161 ; Dl, El, FI & G2 from BVW 2185;

B I , H 1 & H2 from Compton 12166; S 1 & S2 from BVW 2190). Scale in mm.

Bothalia 18,1 (1988)

33

This species is closely related to L. gracilis (E. Mey.)

Benth., but differs in the more lax and sparse habit, the

sparsely strigillose leaves and twigs (not densely white-

pubescent), the presence of a single stipule to each node

(stipules totally absent in L. gracilis) and in the orange-

red vexillum (yellow in L. gracilis ).

L. comptonii is known only from two small popula-

tions on the northern slopes of the Witteberg and the

Klein Swartberg (Figure 2), where it was found in dry

fynbos vegetation on shallow, rocky soil. The specific

epithet commemorates Prof. R. H. Compton ( 1 886 —

1979), director of Kirstenbosch Botanical Garden for 34

years. He was responsible for the only collection known

prior to my rediscovery of the species in 1986.

CAPE. — 3320 (Montagu): Bantamskop, Witteberg (-BC),

27.10.1941, Compton 12166 (NBG, PRE); Bantamskop, 3 km from

the top(-BC), 13. 10. 1986, B-E. van Wyk 2185 (JRAU, PRE, NBG),

2186 (BOL, JRAU, PRE, MO, NBG), 2187 (JRAU, PRE), 2188,

2189 (JRAU), 2190 (JRAU, PRE). 3321 (Ladismith): Laingsburg to

Seweweekspoort Road, 9,5 km from entrance of Rietvlei Farm

(-AD), 12.10.1986, B-E. van Wyk 2160 (PRE, holo.; BOL, C,

JRAU, K, M, MO, NBG, SAAS, STE, iso.), 2161 (BOL, JRAU, K,

MO, NBG, STE), 2162 (BOL, GRA, JRAU, KMG, NH, PRE), 2163

(JRAU, PRE).

FIGURE 2. — The known geographical distribution of Lotononis comp-

tonii, ■ ; L. dahlgrenii, #; and L. dissitinoclis , •.

Lotononis dahlgrenii B-E. van Wyk , sp. nov. L.

rigidae (E. Mey.) Benth. similis, sed habitu etiam rigi-

diore lignosioreque, ramis lateralibus brevibus rigidis

aliquantum spinescentibus, racemis longioribus latiore

dispersis unifloratis (2 vel 3-floratis in L. rigida), calyce

sparse puberulo (in L. rigida dense patente pubescens),

vexillo carinaque maioribus, ovario fructuque longiori-

bus, differt. Etiam L. dissitinodi mei similis, sed foliolis

valde brevioribus, minus conduplicatis, stipulis omnino

destitutis, lobis calycis lateralibus connatis et vexillo

valde minore differt.

TYPE. — Cape, 3321 (Ladismith): Ladismith Div.,

near top of Roodeberg Pass (-DA), 19.7.1954, Lewis

4709 (NBG, holo.).

Rigid woody shrub. Branches sparsely leafy; some of

the lateral branches short, stiff and pungent; young twigs

silky-sericeous. Leaves digitately trifoliolate, densely

silky-sericeous; petiole very short, 1-3 mm long; leaf-

lets small, narrowly oblanceolate to linear, 2-5 (-8)

mm long, slightly conduplicate. Stipules absent. Inflo-

rescences subterminal on short lateral branches, 1-

flowered, rarely 2-fiowered; peduncle long, (6-) 10-15

mm in length; bracts small, oblong, up to 1 mm long;

bracteoles absent. Flowers 12—13 mm long, yellow;

pedicel 2—4 mm long. Calyx 6—8 mm long, lobes long,

acuminate, with the lateral ones on either side fused

much higher up in pairs, sparsely puberulous. Standard

ovate and pointed, 10-12 mm long, with a line of hairs

dorsally along the middle. Wing petals shorter than the

keel, oblong; sculpturing upper basal and upper central,

in 7-8 rows of intercostal lunae. Keel petals semicircu-

lar, obtuse, auriculate and pocketed near base. Anthers

dimorphic. Pistil long; ovary linear, 7-12 mm long.

Fruit linear, 12-15 mm long, inflated laterally; lower

suture distinctly sunken, upper suture verrucose. Seed

(immature) densely tuberculate (Figure 3).

This species is similar to L. rigida (E. Mey.) Benth.

but differs in the habit which is even more rigid and

woody, the short stiff and somewhat thorny lateral

branches, the longer, more widely spaced and single-

flowered racemes (2 or 3-flowered in L. rigida), the

sparsely puberulous calyx (densely and patently pubes-

cent in L. rigida), the larger standard and keel and the

longer ovary and fruit. It is also similar to L. dissitinodis

B-E. van Wyk, but differs in the much shorter and less

conduplicate leaflets, the total absence of stipules, the

fused lateral calyx lobes and the much smaller standard.

L. dahlgrenii may easily be confused with species of

Wiborgia and Aspalathus when not in flower. It appears

to be a rare species and is only known from three speci-

mens collected on the Rooiberg near Ladismith. The

species is named after the late Prof. Rolf Dahlgren in

recognition of his valuable contributions to the literature

on Cape legumes.

CAPE. — 3321 (Ladismith): Roodeberg summit ( — CB), 24.3.1940,

Bond 263 (NBG); 3. 1940, Lewis 1211 (SAM); near top of Roodeberg

Pass (-DA), 19.7. 1954, Lewis 4709 (SAM).

Lotononis dissitinodis B-E. van Wyk, sp. nov.,

strictim L. rigidae (E. Mey.) Benth. et L. dahlgrenii mei

similis, sed intemodiis valde longioribus, foliolis

longioribus anguste linearibus conduplicatis, stipulis ad

pedunculorum insertionem semper praesentibus, inflo-

rescentibus foliis oppositis (non subterminalibus), caly-

cis lobis subaequalibus, vexillo maximo, carina minore

acutioreque, fructu minus turgido differt.

TYPE.— Cape, 3320 (Montagu): 14 miles [22,4 km]

SSE of Laingsburg (-BD), 25.7.1959, Acocks 20502

(PRE, holo.; BOL, M, NBG, iso.).

Woody, sparsely branched shrub, up to 0,4 m tall.

Branches with very long intemodes, young twigs

densely sericeous. Leaves widely spaced on the twigs,

digitately trifoliolate (very rarely 4 or 5-foliolate),

densely sericeous; petiole (2-) 5-8 (-10) mm long,

adaxially grooved; leaflets narrowly linear, (5-) 7-12

(-15) mm long, conduplicate, apices recurved at matu-

rity. Stipules rarely present, at the insertion of the pedun-

cles only, single or paired, linear, variable in length,

1-8 mm long. Inflorescences leaf-opposed. 1 -flowered

(rarely 2-flowered); peduncles variable in length, 5-20

mm long; bracts small, up to 1 mm long; bracteoles

absent. Flowers large, 15-20mm long, yellow; pedicel

34

Bothalia 18,1 (1988)

FIGURE 3. — Lotononis dahlgrenii. A, flowering branch; Bl , B3, leaves in adaxial view; B2, B4, leaves in abaxial view; Cl , C2,

calyx opened out with upper lobes to the left, showing lusion of the lateral lobes; Dl, D2, D3, standard petals, showing

variation in shape; El, E2, wing petals; FI , F2, keel petals; G I , G2, pistils; H, Iruit; S, immature seed, showing tuberculate

surface; Tl, T2, T3, bracts. (A, B I , B2, Cl , Dl . E2, F2, G I & T3 from Lewis 4709', B3, B4, D2, FI , G2, FI & T2 from Bund

263, C 2, D3, El, S &TI from Lewis 1211). Scale in mm.

Bothalia 18,1 (1988)

F|CURE 4. Lotonoms dissttinodis A flowering branch, showing the long intemodes and leaf-opposed inflorescences' B1 B7 B3

. 5J’ a.dax,a v,ew’ leaflets opened out, showing paired stipules, B2, abaxial view, with leaflets opened out stipules

fb^! B2 ' fdftXmh V'eW- th0W'ng ‘he n0rma' conduPlicate shape of the leaflets and a single stipule; Cl , C2, calyx^pened out Ljer

lobes to the left show, ng the very shght fusion of the lateral lobes; D1 , D2, standard petals (note the size); E, wing

obovate shape, F, keel petal, showing the small size and pointed apex; V, androecium; G, pistil; H, fruit, showinrverrucose upper

SE r,o“r: surface; Tl ' T2' b"'s- IA" “«P' V * T.'fan W„r,s ,5

36

Bothalia 18,1 (1988)

1—3 mm long. Calyx 8-12 mm long; lobes long, sub-

equal, minutely silky-puberulous. Standard large,

broadly ovate, longer than the keel, 12—18 mm long,

longitudinally striate. Wing petals obovate, longer than

the keel; sculpturing upper basal, in ± 4 rows of inter-

costal lunae and irregular transcostal lamellae. Keel

petals short, semicircular, 8-10 mm long, acute. An-

thers dimorphic. Pistil 9-10 mm long; ovary linear,

6—8 mm long. Fruit oblong-linear, 16—20 mm long, 5

mm wide, not much inflated; lower suture not or only

slightly sunken; upper suture verrucose. Seed (immature)

densely tuberculate (Figure 4).

Superficially the species is similar to L. rigida and L.

dahlgrenii, but it differs in the much longer intemodes,

the longer, narrowly linear, conduplicate leaflets, the

consistent presence of stipules at the insertion of the

peduncles, the leaf-opposed (not subterminal) inflores-

cences, the subequally lobed calyx, the very large stan-

dard, the smaller, more acute keel petals and the less

turgid fruit.

L. dissitinodis appears to be restricted to the Klein

Swartberg (Figure 2) and is only known from four col-

lections. The very long intemodes are a diagnostic

character useful in distinguishing it from other woody

species, hence the specific epithet. Although it is closely

related to other species of the section Aulacinthus, the

very large standard, obovate wing petals and relatively

small, pointed keel approaches the flower structure typi-

cal of the section Telina (E. Mey.) Benth. [L. prostrata

(L.) Benth. and its allies]. This similarity is of interest

since it may help to explain the origin of the Telina-type

flower and inflorescence.

CAPE. — 3320 (Montagu): Buffets Poort Berg, lower slopes (-BD),

5.7.1941, Levyns 7414 (BOL); Klein Swartberg, western end (-BD),

7.8.1957, Wurts 1520 (NBG, 2 sheets); 14 miles SSE of Laingsburg

(-BD), 25.7.1959, Acocks 20502 (BOL, M, PRE); Seweweekspoort,

N foothills of Klein Swartberg, Santo (-AD), 3.9.1982, Viviers &

Vlok 70 (STE).

ACKNOWLEDGEMENTS

I wish to thank Dr H. F. Glen (Botanical Research

Institute, Pretoria) for the Latin translations and the

Directors and staff of the cited herbaria for the loan of

specimens. The taxonomic study of Lotononis is a regis-

tered Ph. D. project at the University of Cape Town.

REFERENCES

BENTHAM, G. 1843. Enumeration of Leguminosae, indigenous to

southern Asia, and central and southern Africa. The London Jour-

nal of Botany 2: 504—613.

DUMMER, R. A. 1913. A synopsis of the species of Lotononis, Eckl.

& Zeyh. , and Pleiospora Harv. Transactions of the Royal Society

of South Africa 3, 2: 275—335.

HARVEY, W. H. 1862. Leguminosae. In W. H. Harvey & O. W.

Sonder, Flora capensis 2: 47—66. Hodges & Smith, Dublin.

Bothalia 18,1:37-42(1988)

Studies in the genus Riccia (Marchantiales) from southern Africa. 8.

R. campbelliana (subgenus Riccia ), newly recorded for the region

S. M. PEROLD* and O. H. VOLK**

Keywords: anatomy, ‘idioblasts’, Marchantiales, Riccia campbelliana, southern Africa, spores

ABSTRACT

R. campbelliana Howe (1899), a rare species originally known from California and later from Georgia, Arkansas

(Jacobs 1951; Wittlake 1954), Kansas and Nebraska (S. Jovet-Ast pers. comm.), as well as from Kazakhstan

(Ladyzhenskaja 1967), has now also been found at a few localities in southern Africa. It is characterized by the distinctive

yellow-brown or rusty colouration of the dorsal surface along the margins and over the proximal parts; enlarged cells,

'idioblasts', which differ in shape and contents from the adjacent cells, are generally found in all parts of the thallus.

UITTREKSEL

R. campbelliana Howe (1899), 'n seldsame spesie oorspronklik bekend van Kalifomie, en later van Georgia, Arkansas

(Jacobs 1951; Wittlake 1954), Kansas en Nebraska (S. Jovet-Ast pers. kom.), sowel as Kazakhstan (Ladyzhenskaja 1967),

kom ook by 'n paar lokaliteite in suidelike Afrika voor. Dit word gekenmerk deur die geel-bruin kleur van die dorsale

oppervlak langs die rande van die tallus en oor die proksimale gedeeltes; vergrote selle, 'idioblaste', wat verskil in vorm en

inhoud van die aangrensende selle, kom ook algemeen in alle dele van die tallus voor.

Riccia campbelliana Howe in Memoirs of Torrey

Botanical Club 7: 26 (1899); Frye & Clark: 20 (1937);

Hassel De Menendez: 267 (1962); Ladyzhenskaja: 316

(1967).

TYPE. — California, near Stanford University, on hills

above Mission Dolores, D. H. Campbell s.n. May 1,

1896 [NY, lecto. !, fide Grolle: 225 (1976)].

Thallus monoecious, perennial, in crowded gregarious

patches or incomplete rosettes or scattered; simple or

once to twice furcate, branches divergent or mostly pa-

rallel, oblong-linear, medium-sized, up to 8 mm long,

1 ,0— 1 ,5( — 2,0) mm broad, once to twice broader than

thick (Figure 1A), segments up to 3,5 mm long; apex

rounded and obtuse, emarginate, dorsal furrow deep, its

sides raised and convex, becoming flatter in older parts

(Figures IB 1-6; 2A, 2B); dorsally pale green, yellow or

rusty brown along margins and more proximally over

entire dorsal surface; margins acute, shortly winged,

slightly undulating, flanks sloping upward and outward,

bronze-brown; ventral surface rounded, green, occasio-

nally flecked with red and brown; when dry, margins

indexed, forming brown lips, flanks with brown scales.

Anatomy of thallus: dorsal epithelium unistratose,

hyaline, cells variously shaped, in younger parts and

towards centre subglobose, 30—45 x 35—50 /zm, late-

rally becoming somewhat flatter and up to 70 /zm wide

(Figures 1C, 2C), brown and collapsed at margins and in

older parts, underlying layer of cells soon losing their

chloroplasts (Figure ID); air pores triangular, ± 7 /zm

wide, often rectangular and larger, ± 12 /zm wide

(Figure IE); in section, assimilation tissue (chloren-

* Botanical Research Institute, Department of Agriculture and Water

Supply , Private Bag X 1 0 1 , Pretoria 000 1 .

** Botanische Anstalten d. Univ. Wurzburg D8700, Germany B.R.D.

MS. received: 1987.04.13.

chyma) occupying about / the thickness of thallus and

consisting of vertical columns of 6—10 cells, 50 — 60 x

35-50 /zm (Figure 1C), frequently some cells, referred

to as ‘idioblasts’ (see p.39), differ from surrounding

ones in the larger size, in the somewhat distended cell

walls and in the contents (Figures ID, 2F), air canals

between assimilation cell columns up to 20 /zm wide;

storage tissue occupying about 'A the thickness of thal-

lus, cells rounded or angular, tightly packed, up to 55

/zm wide, containing chloroplasts and starch granules,

usually with some scattered ‘idioblasts’ (Figure 2E).

Rhizoids hyaline, some smooth, others tuberculate,

15-20 /zm wide, arising from ventral epidermis of thal-

lus and base of scales. Scales imbricate, fragile,

inconspicuous, not projecting above thallus margin (Fig-

ure 2D), up to 750 x 450 /zm; margin hyaline, rest of

scale with groups of brown and different shades of violet

cells, interspersed with single, or with groups of hyaline,

4-6-sided cells, walls straight or sometimes sigmoidal,

cells elongated to nearly isodiametric, up to 65 x 40

/zm, some ‘idioblasts’ may be present, marginal cells

smaller, ± 40 x 40 /zm (Figure IF). Antheridia with

short hyaline or white necks along dorsal groove. Arche-

gonia with purple necks, also along groove. Sporangia

infrequent, single or crowded, large, ± 500 /zm wide,

with about 200 spores each, in median part of thallus,

causing bulging of overlying tissue which turns brown.

Spores (85 — )90 — 1 10( — 1 20) /im in diameter, pale

straw-coloured to brown, semitransparent, triangular-

globular, polar, with wing up to 5 /zm wide (Figure 3B),

pores at angles ±7,5 /zm or more wide, margin ±

smooth, ornamentation on both faces vermicular to irre-

gularly and incompletely reticulate; distal face with

10—15 incomplete areolae across diameter, ± 5 /zm

wide (Figure 3C, F), ridges often sinuous and raised into

blunt papillae at the nodes (Figure 3D), in transmitted

light ridges often appear to have a midline; proximal face

with triradiate mark generally clearly defined (Figure

3E) and with the ridges somewhat lower than those on

38

Bothalia 18,1 (1988)

FIGURE 1 . Riccia campbelliana . Structure of thallus, scales and chromosomes. A, fresh thalli; B1 6, transverse sections of thallus branch

at different distances from apex to older parts; C, transverse section through dorsal epithelium, chlorenchyma and storage tissue; D,

transverse section through collapsed dorsal cells and chlorenchyma, showing ‘idioblasts’ and air canals; E, epithelial cells and narrow air

pores from above, dotted lines indicate contact with chlorenchyma cells below; F, scale; G, chromosomes, the letters A-E identity the

chromosomes (see text): A-D, F, S. M. Perold 888 ; E, Van Rooy & Perold 634 ; G, Van Rooy & Perold 637 \ A-F, by Volk; G, by

Bomefeld. Scale bar on A, B = 2 mm C-F = 100 /am; G = 1 /am.

Bothalia 18,1 (1988)

39

FIGURE 2. — Riccia campbelliana .

Structure of thallus and cells.

A, dorsal surface view of thal-

lus; B, apex and groove; C,

dorsal cells; D, scales at mar-

gin; E, transverse section

through thallus branch, show-

ing ‘idioblasts’; F, dorsal epi-

thelial cells and ‘idioblasts’ in

chlorenchyma. A-F, S. M. Pe-

rold 888. A-D, SEM micro-

graphs by Perold; E & F, LM

micrographs by Volk. Scale bar

= 50 pm.

distal face (Figure 3A). Chromosome number n = 8

[Figure 1G: the letters A— E identify the chromosomes,

T. Bomefeld pers. comm.; Siler (1934)].

In adapting Tatuno’s (1941) method, which in prin-

ciple is also that used by Jovet-Ast (1969; 1986), this

species with n = 8 chromosomes, has the following

karyotype, according to Bornefeld (1984): A, BB, CC,

DD, E. Capital letters of the alphabet are assigned to the

different chromosome types, whereby the largest are

denoted by the letter A and the smallest by the letter E. It

presents the basic set of chromosomes in the genus Ric-

cia, from which all other sets can be derived (Bornefeld

1984).

The ‘idioblasts’ in R. campbelliana are variable in

size, colour and contents; in the assimilation tissue they

are up to 100 gm long x 45-60 gim wide, single or

arranged in groups and are free of chloroplasts (fluores-

cence assay by Volk); those occurring in the epithelium,

the storage tissue and in the scales are also larger than the

surrounding cells. In fresh or dry material, the ‘idio-

blasts’ can be brown, grey or hyaline, and appear to be

clear and homogeneous or granular. They are readily

stained with Heidenhain’s iron haematoxylin (Howe

1899) or with diluted aqueous solutions of Thiacine

stains which are rapidly absorbed and produce metachro-

matic effects. Bienfait & Waterkeyn (1976) reported that

metachromatic effects demonstrate the presence of acid

muco-polysaccharides when Toluidine blue is used as a

stain in histochemical assays. The ‘idioblasts’ in R.

campbelliana are stained blue with Toluidine blue,

whereas other cell walls are violet. It may therefore be

assumed that pectines (derivatives of uronic acid) in the

cell walls, are responsible. With iodine-potassium

iodide, the cells turn brownish. In his original descrip-

tion of R. campbelliana Howe (1899) and R. dictyospora

Howe ( 1901 ), both from North America, Howe regarded

these ‘idioblasts’ as ‘oil cells’. Ladyzhenskaja (1967)

confirmed the presence of ‘oil cells’ in these two species

and also reported their occurrence in the European spe-

cies, R. breidleri Steph. and R. oelandica Jensen. Exam-

ination of four specimens of R. breidleri from Austria

(B ! ; W !) and four from Switzerland (G!) failed however,

to show the presence of ‘idioblasts’, but instead demon-

strated numerous droplets of ‘fatty oil’ in the storage

tissue. ‘Idioblasts’ were not found in R. oelandica (G!)

either, and only rarely could globules of fatty oils be

demonstrated, which also occur in R. campbelliana, but

only after boiling, when they can be stained with Tinc-

ture of Alkanna, Sudan Ill-glycerine or with other stains

for fats. ‘Idioblasts’ similar to those in R. campbelliana.

40

Bothalia 18,1 (1988)

FIGURE 3. — Riccia campbelliana.

Spores. A, proximal face; B,

viewed from side; C, D, F, dis-

tal face; E, centre of triradiate

mark. A-F, S. M. Perold 888.

A-E, SEM micrographs by

Perold; F, LM micrograph by

Volk. Scale bar = 50 gm;

diameter of spore on F, ± 100

gim.

were found in R. dictyospora (G!; PRE!) and in R.

macrocarpa Levier (B ! ; W!) — described in the latter

by Jovet-Ast (1986) as ‘cellules a contenu orange

eparses’, and in specimens of R. nigrella from many

parts of the world. In southern African Ricciae, the pre-

sence of ‘idioblasts’ is a unique feature and only known

in these two species, R. campbelliana and R. nigrella [/?.

capensis auct. non Steph.: S. Amell (1952)], see Perold

& Volk (1988).

According to Howe (1899) the epithelial cells of

R. campbelliana are usually longer than wide, but the

South African specimens have ± globose, to depressed

globose cells. He also reported the antheridial necks to

be 100-170 /u.m long, i.e. more prominent than in our

plants. The SEM micrographs of the spores published by

Steinkamp & Doyle (1979), bear a close resemblance to

those in the present study.

R. campbelliana is new to southern Africa. It is

known from only three localities in central Transvaal,

ranging from Derby, eastwards to Middelburg, and from

a single locality in the eastern Orange Free State, near

Fouriesburg (Figure 4). It grows in summer rainfall areas

with an annual precipitation of 600 - 800 mm and at

altitudes of I 500-1 800 m above sea level. It prefers

shallow, acid soils overlying granitic, quartzitic or

sandstone rock outcrops (pH ± 4,6), fully exposed to the

sun, and is often associated with other Riccia species,

such as R. nigrella DC. and R. okahandjana S. Amell,

with small moss species, e.g. Pleuridium nervosum

(Hook.) Mitt, and with crustaceous lichens.

FIGURE 4. — Map showing distribution of R. campbelliana in southern

Africa.

Bothalia 18,1 (1988)

41

Localities of R. campbelliana outside southern Africa

are the following: USA, California, where it was found

by Campbell, in whose honour Howe named it; Georgia

(Jacobs 1951), Arkansas (Jacobs 1951; Wittlake 1954),

Kansas and Nebraska (S. Jovet-Ast pers. comm); USSR,

Kazakhstan, Western Siberia (Ladyzhenskaja 1967) and

Argentina, Tucuman, [Sleumer 1755 (LIL 19853'.)]. The

latter specimen was referred to R. campbelliana by

Hassel de Menendez (1963) but it is doubtful whether it

has been correctly identified as it lacks ‘idioblasts’ and

the scales are deep red and large, definitely not inconspi-

cuous; only the ornamentation of the spores shows some

resemblance.

R. campbelliana is a very rare species with a strikingly

disjunct distribution, viz. North America, (?) South

America, South Africa and central Asia. These ‘relictual

areas’ (Wulff 1950), may have resulted from interme-

diate localities becoming extinct, as the transport of

spores over such immense distances in modern times by

for example, migratory birds, can hardly be accepted.

Consequently, R. campbelliana is regarded as a very

ancient species, similar to the almost equally rare R.

crustata Trabut with remnant populations likewise

spread over several continents, namely Central Asia

(Kazakhstan), Europe, North Africa (Jovet-Ast 1973)

and, if it were agreed that R. crustata and R. albida

Sullivant ex Austin constitute a single species (Jovet-Ast

1986; Scott & Bradshaw 1986), also occurring in North

America and Australia.

R. campbelliana can be recognized by the yellow-

brown colour along the thallus margins, by the inconspi-

cuous scales, partly hyaline and partly flecked with

brown and violet, and by the sinuous vermicular ridges

on the proximal and distal spore faces. Although small

forms of R. campbelliana bear some resemblance to R.

nigrella, they can be distinguished by the usually wider

and longer branches, by the thinner, slightly attenuate

margins and by the epithelial cells that are not persistent.

R. nigrella is generally a smaller plant and it often grows

in rosettes or partial rosettes; the thallus is dorsally deep

brown except toward the apex, the scales are closely

appressed and shiny black; when it sporulates, it is seen

with overlying masses of dark brown spores; the spores

are smaller and have vermicular ridges on the distal face

and generally on the proximal face too.

SPECIMENS EXAMINED

Riccia campbelliana

Besides the lectotype specimen (NY), the following

specimens were examined:

South Africa

TRANSVAAL. — 2527 (Rustenburg): 2 km to Derby on road from

Rustenburg ( — CC ) S. M. Perold 888 (PRE). 2528 (Pretoria): 19 km N

of Bronkhorstspruit, overlying granite ( — DC). Volk 811024 (M. PRE).

2529 (Witbank): N of Middelburg, on road R35, at plateau near Klein

Olifants River (— CB), S. M. Perold 69, 80, 81 (PRE); Van Rooy &

S. M. Perold 634, 637, 642 (PRE); Volk81/010 (M, PRE).

O.F.S. — 2828 (Bethlehem): 11 km E of Fouriesburg on road from

Clarens, around weathered sandstone outcrops on slope ( — CB ), S. M.

Perold & Germishuizen 1307 (PRE).

North America

NEBRASKA. — Lancaster Co., Lincoln, Waller Kiener 9961 (US).

KANSAS. — Woodson Co., Baker’s Bluff, R. R. Ireland Jr. 83

(US).

CALIFORNIA. — Balboa Park, San Diego, C. C. Haynes 2823

(US).

South America

ARGENTINA. — Tucuman, Sleumer 1755 ( LIL 19853) (doubtfully

referred to R. campbelliana).

Other species examined

R. breidleri from Austria (B; W), Switzerland (G); R. dictyospora

from North America (G; PRE) and R. macrocar pa from Turkey, Italy

and France (B; W).

ACKNOWLEDGEMENTS

The authors wish to thank Mme S. Jovet-Ast, hono-

rary professor, Laboratoire de Cryptogamie, Paris, for

drawing our attention to additional North American loca-

lities of R. campbelliana. Sincere thanks are also due to

Dr habil. T. Bomefeld, Am Reelein 1, D-8706,

Hochberg, Germany, for the chromosome counts and

figures. We are grateful to the curators of B; G; NY; US;

W and Fundacion Miguel Lillo, Tucuman, for the loan

of specimens.

REFERENCES

ARNELL, S. 1952. Hepaticae collected in South and West Africa,

1951. Botaniska Notiser 105: 307-315.

BIENFAIT, A. & WATERKEYN, L. 1976. Sur la nature des parois

sporocytaires chez les mousses et chez quelques Pteridophytes.

Etude comparative. Compte rendu hebdomadaire des seances de

I’ Academie des sciences. Paris, ser. D, 282: 2079-2081.

BORNEFELD, T. 1984. Chromosomenanalyse der Gattung Riccia von

Slid- und SW-Afrika und allgemeine Bemerkungen zur Zytoge-

netik der Lebermoose. Nova Hedwigia 40: 313 — 328.

FRYE, T. C. & CLARK, L. 1937. Hepaticae of North America. Uni-

versity of Washington Publications in Biology 6: 1 — 162.

GROLLE, R. 1976. Verzeichnis der Lebermoose Europas und benach-

barter Gebiete. Feddes Repertorium 87: 171—279.

GROLLE, R. 1983. Hepatics of Europe including the Azores: an anno-

tated list of species, with synonyms from recent literature. Jour-

nal of Bryology 12: 403—459.

HASSEL DE MENENDEZ, G. G. 1962. Estudio de las Anthocero-

tales y Marchantiales de la Argentina. Opera Lilloana 7: 1 -297.

HOWE, M. A. 1899. The Hepaticae and Anthocerotes of California.

Memoirs of the Torrey Botanical Clubl: 13—33.

HOWE, M. A. 1901. Riccia beyrichiana and Riccia dictyospora. Bul-

letin of the Torrey Botanical Club 28: 161-165.

JACOBS, D. L. 1951. Ricciaceae of Arkansas. The Brvologist 54:

274-277.

JOVET-AST, S. 1969. Le caryotype des Ricciaceae. Revue Bryologi-

que et Lichenologique 36: 673—689.

JOVET-AST, S. 1973. Complement a l’etude du Riccia crustata Tra-

but. Presence en Australie. Spores et paroi sporale. Revue Bryo-

logique et Lichenologique 39: 167—174.

JOVET-AST, S. 1986. Les Riccia de la Region Mediterraneenne.

Cryptogamie, Bryologique et Lichenologique 7: 287 — 431 .

LADYZHENSKAJA, K. I. 1967. Riccia campbelliana Howe (Hepati-

cae) in USSR. Novitates systematicae plantarum non vascular-

ium 1967: 316—321.

MULLER, K. 1952. Die Lebermoose Europas. In L. Rabenhorst,

Kryptogamen Flora, edn 3, Band 6,1: 1 —756. Leipzig.

42

PEROLD, S. M. & VOLK, O. H. 1988. Studies in the genus Riccia

(Marchantiales) from southern Africa. 9. R. nigrella DC. and the

status of R. capensis. Bothalia 18,1: 43—49.

SCOTT, G. A. M. & BRADSHAW, J. A. 1986. Australian liverworts

(Hepaticae): annotated list of binomials and checklist of pub-

lished species with bibliography. Brunonia 8: 1 — 171.

SILER, M. B. 1934. Chromosome numbers in certain Ricciaceae.

Proceedings of the National Academy of Science, Washington 20:

603-607.

STEINKAMP, M. P. & DOYLE, W. T. 1979. Spore wall ultrastruc-

ture in four species of the liverwort Riccia. American Journal of

Botany 66: 546—556.

Bothalia 18,1 (1988)

STEPHANI, F. 1882. Riccia ciliifera Link und Riccia breidleri

Juratzka. Hedwigia 21: 76.

STEPHANI, F- 1885. Riccia breidleri Juratzka. Hedwigia 24: 6, 7.

TATUNO, S. 1941. Zytologische Untersuchungen fiber die Leber-

moose von Japan. Journal of Science Hiroshima University 4:

73-187.

WITTLAKE, E. B. 1954. Hepaticae of Arkansas. The Bryologist 57:

7-18.

WULFF, E. V. 1950. Historical plant geography, pp. 223. Chronica

Botanica.

Bothalia 18,1:43-49(1988)

Studies in the genus Riccia (Marchantiales) from southern Africa. 9.

R. nigrella and the status of R. capensis

S. M. PEROLD* and O. H. VOLK**

Keywords: distribution, Marchantiales, Riccia capensis, R. nigrella, southern Africa, spores

ABSTRACT

R. nigrella DC. (Lamarck & De Candolle 1815), with which R. capensis auct. non Steph.: S. Amell (1952, 1963), is

conspecific, is now also recognized from southern Africa. The thallus is brown marginally and dorsally and has shiny black

scales with smooth margins at which coloured and hyaline cells alternate irregularly. Amell's ( 1961 ) records of R. capensis

Steph. from the Canary Islands are to be regarded as erroneous. R. capensis Steph. (Stephani in Brunnthaler 1913) is now

placed in synonymy under/?, limbata Bisch.

UITTREKSEL

R. nigrella DC. (Lamarck & De Candolle 1815), waarmee R. capensis auct. non Steph.: S. Amell (1952, 1963),

konspesifiek is, word nou ook uit suidelike Afrika aangemeld. Die tallus is bruin langs die rande en dorsaal en besit blink,

swart, gaafrandige skubbe met onreelmatig afwisselende, gekleurde en hialine selle. Amell se gegewens oor die versprei-

ding van R. capensis Steph. in die Kanariese eilande moet as foutief beskou word. R. capensis Steph. (Stephani in Brunn-

thaler 1913) is ’n sinoniem van R. limbata Bisch.

Riccia nigrella DC. in Lamarck & De Candolle,

Flore Frangaise 6: 193 (1815); Gottsche et al.\ 605

(1844); Camus: 212 (1892); Stephani: 334 (1898);

Howe: 28 (1899); Boulay: 210 (1904); Casares-Gil: 220

(1919); Macvicar: 24 (1926); Frye & Clark: 21 (1937);

Muller: 465 (1952); Na-Thalang: 93 (1980); Jovet-Ast:

323(1986),

TYPE. — France, Dept, de l’Heraut, in sylvula Gram-

mont prope Monspessulanum, Bouchet 1807 [PC, holo.,

fide Jovet-Ast: 323 (1986); G (G23307), iso.!].

R. aggregata Underw.: 275 (1894) after Howe (1899).

R.pearsoni(i) Steph.: 27 (1898); Steph.: 335, 336(1898).

R.porosa TayL: 416 (1846); pp. fide Na-Thalang: 93 (1980).

R. capensis auct. non Steph.: S. Amell:312 (1952); S. Amell: 28

(1963).

Thallus monoecious, perennial, in complete or incom-

plete rosettes, 8-15 mm across (Figure 2A), or scat-

tered; generally rather small; simple or once to twice

furcate, branches narrowly to moderately divergent, ob-

long to linear-ovate (Figures 1A1-4, 2B), up to 5 mm,

occasionally up to 8 mm long, 0,5- 1 ,0 mm broad, once

to twice broader than thick; margins acute, slightly

recurved, becoming somewhat obtuse toward base; apex

rounded or subacute, emarginate; sulcus narrow and

deep, visible along whole length of thallus (Figures

1B1-1B4, 2C), except at sporangia, sides convex;

dorsally glaucous green to green, shiny, soon turning

brown along margins, over older parts and sporangia;

flanks steep, dark-coloured, covered by closely

appressed, shiny black scales (Figure IE); ventral sur-

face rounded, green, often flecked with brown or purple;

* Botanical Research Institute, Department of Agriculture and Water

Supply, Private Bag X 101 , Pretoria 0001 .

** Botanische Anstalten d. Univ. Wurzburg D8700, Germany. B.R.D.

MS. received: 1987.04.13.

when dry, scales cover most of dorsal surface, sides

tightly indexed distally, clasped together, except over

sporangia and over proximal part of branches; margins

raised and erect toward base. Anatomy of thallus: dorsal

epithelium unistratose, cells 25-35 x 25-32 gm, short

rectangular or subquadrate, dorsally rounded to nearly

flat, persistent, (Figure 1C), often hyaline, but mostly

with metachromatic contents, air pores small, usually

triangular, ± 7,5 gun wide (Figures ID, 2D); assimila-

tion tissue occupying about / the thickness of thallus,

consisting of 6-10 cells in vertical rows (Figure 1C),

cells 25-30 x 20-25 gun, rarely a few cells enlarged

and somewhat distended, up to 62 x 32 gun with hyaline

or brownish contents, referred to as ‘idioblasts’ (Figure

2F); storage tissue with cells closely packed, angular, ±

30 gun wide, many containing chloroplasts; ventral epi-

dermal cells often coloured brown or violet (Figure 2E).

Rhizoids, some smooth, others tuberculate, ± 20 gun

wide. Scales semilunar, imbricate, not extending above

thallus margin (Figure 2C), size up to 550 x 450 gun,

shiny violet-black, distal margin hyaline, sometimes

single cells in interior reddish or hyaline, others deep

purple to black, occasionally coloured and hyaline cells

alternate irregularly at ± smooth margin, resembling a

'battlement’ or dentate outline; cells rectangular or short

hexagonal, in body of scale up to ± 52 x 25 gun, smal-

ler at margin (Figure IF), cell walls straight. Antheridia

with hyaline necks. Archegonia with purple necks, in

rows. Sporangia protruding dorsally, 300-400 gun in

diameter, occupying nearly the whole width of thallus,

in a row or crowded together, containing about 160-230

spores each, overlying tissue dark brown, breaking down

over ripe capsules and leaving a large hollow filled with

many spores. Spores (62,5 — )70, 0-85, 0(— 87,5) gun in

diameter, triangular-globular, polar, light to dark brown,

with wing narrow (Figure 3E), 2, 5-5,0 gun wide,

notched or with a pore at angles, margin finely crenulate;

distal face incompletely reticulate with about 10-12 ir-

regularly shaped areolae across, or vermicular with

thickened, short, sinuous ridges, ± 2,5 gun wide, which

44

Bothalia 18,1 (1988)

FIGURE 1. — Riccia nigrella. Structure of thallus, scales and chromosomes. Al-4, thalli; Bl-4, transverse sections of thallus branch at

intervals along its length; C, transverse section showing dorsal cells, chlorenchyma and storage tissue; D, dorsal cells and air pores from

above; E, flank with scales and rhizoids; F, scales (dotted = dark purple; hatched = reddish); G, chromosomes. (Al, A3, C-G, Volk

8l-2l4c\ A2, Volk 81-289\ A4, Bl-4, 5. M. Perold520). (A-F, by Volk;G, by Bomefeld). Scale baron Al-4, E = 1 mm; Bl^f,C

= 500 fim; D, F, - 100 /am ; G = I /am.

Bothaiia 18,1 (1988)

45

FIGURE 2.—R. nigrella. Structure of thallus and cells. A, rosettes; B, dorsal surface view of thallus; C, apex, groove and

scales; D, dorsal epithelial cells and air pores; E, transverse section through branch, unstained; F, more enlarged

transverse section through thallus branch, showing dorsal epithelial cells with finely granular contents, with unusually

many ‘idioblasts' in bottom left comer). [A, F, Volk 81 -214c ; B-D, Volk 85-802 ; E, S. M. Perold 1147). [A.

photograph by Volk; B-D, SEM micrographs; E & F, LM (light microscope) micrographs by Perold). Scale baron A

= 1 mm; on B-F = 50 /am.

are often roughened with fine granules or papillae (Fig-

ure 3C, D, F); proximal face with triradiate mark dis-

tinct, its arms often wider at juncture with wing, about

30 small, rounded areolae on each of 3 facets, 2, 5-5,0

/i.m wide, ridges thick, slightly raised at nodes (Figure

3A, B). Chromosome number n = 8 (Figure 1G) (Borne-

feld 1984 on Volk 81 -214c (as cf. R. pottsiana)', Na-

Thalang (1980, Australia); Jovet-Ast (1986, France).

In R. nigrella, the dorsal and ventral epithelial cells

frequently contain hyaline or brownish metachromatic

substances which stain blue with dilute aqueous solu-

tions of Toluidine blue, as do the ‘idioblasts’ of R. camp-

belliana (Perold & Volk 1988). Rarely, enlarged cells

with a similar substance are also found in the assimila-

tion and storage tissues of the thallus. In old specimens

the contents of the dorsal cells may have shrunken away

from the cell walls and are bright orange. European

authors excepted, several others have commented on the

striking character and the affinity the persistent dorsal

epithelial cells have for certain stains (Howe 1899; Frye

& Clark 1937; Na-Thalang 1980).

In southern Africa R. nigrella occurs quite rarely in

the summer rainfall areas (with a moderately high annual

precipitation up to 700 mm), of central and southern

Transvaal, Natal, eastern and southern Orange Free

State. It is more common in south-western and north-

46

Bothalia 18,1 (1988)

FIGURE 3. — R. nigrella. Spores. A, proximal face; B, viewed from side; C, distal face; D, distal side; E, more enlarged

view of ridges on distal face; F, distal face. (A, B. Duthie 5019a; C-E, Laxly 5023; F, Arnell 792). (A-E, SEM

micrographs and F, LM micrograph by Perold). Scale bar = 50 /u.m; diameter of spore on F, ± 80 /xm.

western Cape Province, extending to the southernmost

part of South West Africa/Namibia, which are winter

rainfall areas (Figure 4), with low annual precipitation

ranging from less than 200 to 500 mm. It grows at alti-

tudes between 200 and 1 800 m above sea level and

prefers very shallow, sandy to dusty, acid soils overlying

rock outcrops. R. nigrella is able to survive and propa-

gate vegetatively by forming small, round, perennating

turions. It is often associated with other xerophilous Ric-

cia species, e.g. R. okahcmdjana S. Arnell and/or R.

atropurpurea Sim, and occasionally with R. campbel-

liana Howe. It can be distinguished from the other sou-

thern African species with shiny black scales by its gen-

erally smaller size, by scales that are closely appressed

and do not extend above the thallus margins and by the

dark brown dorsal colour. It bears some resemblance to a

small R. campbelliana, which is also brown dorsally,

and occasionally a few ‘idioblasts’ are also present in the

inner tissues of R. nigrella. R. nigrella grows diffusely

or in partial or complete rosettes. When sporulating, it is

seen with overlying clumps of large numbers of dark

brown spores.

This nearly cosmopolitan species (Jovet-Ast 1986) is

found in all countries bordering the Mediterranean, and

in Wales, Cornwall, the Channel Islands, the Macarone-

sian Islands (Madeira, Azores and the Canaries), North

America, Australia, and now also in southern Africa.

Bothalia 18,1 (1988)

47

FIGURE 4. — Map showing distribution of R. nigrella in southern Africa.

THE STATUS OF R. CAPENSIS AUCT. NON STEPH.: S. ARNELL

R. capensis auct. non Steph .: S. Arnell is conspecific

with R. nigrella DC.

While visiting South Africa in the early nineteen fif-

ties, Arnell collected a number of specimens of a small,

black-scaled Riccia species (held at BOL, PRE, S and

UPS). He experienced problems in correctly identifying

his collections and originally named them R. brunomar-

ginata. Later he changed his mind and thought he was

dealing with R. capensis Steph., but when he (Arnell

1952) examined the type specimen of R. capensis

Steph., leg. Brunnt haler, Gnadenthal, from W, the

packet contained only sand and no Riccia could be

found. He obviously did not know of Brunnthaler’s col-

lection [(G 1 3334!) held at G], on which Stephani had

based his diagnosis of R. capensis. His only recourse

therefore, was to rely on Stephani’s drawings in the lat-

ter’s unpublished leones hepaticarum (G; M). As the

transverse sections of the thalli of his collections seemed

to him to be similar to Stephani’s figures of R. capensis,

he chose his specimens Arnell 59 (S) & 162 (BOL) as

types for R. capensis sensu Arnell (Arnell 1952, 1963).

The authors have now found that these and all the other

specimens named R. capensis by Arnell, are in fact R.

nigrella. He thus misapplied the name R. capensis

Steph. when he referred his own and other collections of

R. nigrella to R. capensis Steph. and in so doing, failed

to recognize R. nigrella in the south-western Cape Pro-

vince. R. nigrella DC. is accordingly newly reported

from southern Africa and Arnell’ s collections of R. ca-

pensis have to be renamed as such.

Arnell (1952) noted that his specimens with their

black scales looked like R. limbata Bisch., but were

much smaller and had quite differently ornamented

spores (see below under The status of R. capensis

Steph.)\ he seems to have ignored Stephani’s thallus di-

mensions of 2,5 mm wide, which are definitely those of

a wider plant. Furthermore, he incorrectly (and mislea-

dingly) described the scale margins of his plants as ‘qua-

drangularly serrate’ (1952) and as ‘irregularly dentate’

(1963), which indicates that he did not notice the hyaline

cells in between the dark cells at the ± smooth margins.

Serrate or dentate scale margins rarely occur in the genus

Riccia. The striking colouration of the scales and their

margins in R. nigrella has not been commented on by

previous authors and is only suggested by Macvicar’s

figure (1926: 25). Arnell’s description of the margins

and dorsal surface of the thallus turning yellowish

brown, the deep and sharp dorsal groove with convex

sides and, on transverse section, the rectangular to qua-

drate epithelial cells, indicate that he was referring to R.

nigrella. He reported the spores to be 50-60 p m in

diameter, the ‘ridges often with a brighter midline or two

bright lines and a dark midline’, an effect which is often

seen with thickly ridged aerolar walls.

Besides several places in the Cape Province, Arnell

(1961) also reported his R. capensis from two localities

in the Canary Islands. Despite repeated attempts, these

specimens have not been traced. At UPS only a single

microscope slide labelled ‘ R . capensis Steph. Gran

Canaria, Montana de Tafira', in Arnell’s writing, is

held. On the slide holder he has altered the label to read

R. capensis = macrocarpa. Arnell’s distribution records

for R. capensis in Macaronesia were, however, accepted

and repeated by Eggers (1982), Grolle (1983), Dull

(1984), Sergio (1984). His collections, S. Arnell

281211959 (S; UPS); S. Arnell 4/3/1959 (UPS) and S.

Arnell 5/3/1959 (S; UPS), from La Calzada and Montana

de Tafira, as well as J. Lid 14/2/1954 (UPS), from

Gomera, were seen and the determinations confirmed as

R. nigrella. Two other specimens, S. Arnell 4/3/1959 (S)

and 5. Arnell 7/3/1959 (S), from Montana de Tafira, as

well as one from Las Palmas, S. Arnell 28/12/51 (UPS

20638), that he had referred to R. nigrella, were incor-

rectly determined and are R. trabutiana Steph., which

was not listed by Arnell (1961) as occurring in the

Canary Islands. Some collections of R. atromarginata

and Targionia sp. were also misidentified as R. nigrella.

Arnell furthermore remarked on some resemblance

between R. nigrella and R. atropurpurea Sim (1963: 28)

in the appearance of the thalli, and although they are

both dorsally deeply grooved, somewhat glaucous gray

and shiny when wet, they are quite different in their

margins and scales. The following note which Arnell

wrote to Talbot, was found with a specimen collected by

Pearson 8956 (PRE): T made a mistake in calling this

plant R. nigrella var. austro-africana, as Sim had

already called it R. atropurpurea' , which, however, it is

not either! Arnell identified specimens of R. atropurpu-

rea, Sim July 1920 (PRE; UPS) from the Matopos, Zim-

babwe [Southern Rhodesia], as well as Sim Jan. 1918

(Vereeniging) (PRE) and Volk 2766 Farm Donkerhoek,

SWA (PRE) as R. nigrella var. austro-africana.

THE STATUS OF R. CAPENSIS STEPH.

R. capensis Steph. is now placed in synonymy under R.

limbata Bisch.

R. limbata Bisch. MS. ex Gottsche, Lindenberg et

Nees ab Esenbeck in Synopsis Hepaticarum: 606 (1844);

Stephani: 326 (1898); Sim: 12 (1926); Arnell: 23 (1963).

Type: Cap. Bonae Spei, in locis humidis ad latera Mon-

tium Tafelberg, Duyvelsberg et Leuvenberg, Krauss

Julio 1838 G 1 3 165 (G ,? holo.!); CBS (= Cap. Bonae

Spei), Gnadenthal, Krauss Dec. 1838 (BM! ex Herb.

Hampe).

48

R. capensis Steph. in Denkschriften der Mathematisch-naturwis-

senschaftlichen Klasse der kaiserlichen Akademie der Wissenschaften

88: 724 (1913). Type: Cape Colony, Gnadenthal by Caledon, Brunn-

thaler Oct. 1909 G 13334 (G!) [G, holo. fide Grolle: 226 (1976), fide

Jovet-Ast in lit.]; W [fide Amell: 312 (1952)] (only sand in specimen

packet).

Stephani was evidently unaware that a paratype speci-

men of R. limbata had also previously been gathered at

Genadendal (by Krauss), when he described R. capensis

as a new species (Stephani 1913) from a specimen col-

lected in October 1909 at Genadendal near Caledon, by

Brunnthaler (G13334). His descriptions of R. capensis

(1913) and R. limbata (1898) differ in that he described

the latter plants as large and four times wider than thick

in cross section, with obliquely sloping flanks, whereas

the former are small, twice as wide as thick and have

erect flanks, indicating that they were not yet fully

grown. In both descriptions he however, referred to the

imbricate, purple scales and the thick thalli.

Examination of Brunnthaler’s collection and compari-

son with the type specimens of R. limbata Bisch., leg.

Krauss July 1838 (G) G 1 3 1 65 and Krauss December

1838 (BM) by the authors, clearly show it to be a speci-

men of R. limbata, generally a medium-sized to large

species, possessing shiny black scales that, however, do

extend beyond the thallus margins; the thalli lack ‘idio-

blasts’ and the spores (recovered from the soil in Brunn-

thaler’s apparently sterile specimen) are large, 100-120

gm in diameter, with the ornamentation on the distal

face typically consisting of parallel rows of papillae spi-

ralling outwards to the wing and the proximal face

almost smooth with scattered pores.

Jovet-Ast (1969) reported the chromosome number for

R. capensis Steph. on a specimen collected by Een,

14/3/1967. This gathering from Tanzania, Serengeti

Park, Senora Lodge, contains only R. congoana Steph.

and R. atropurpurea Sim.

ACKNOWLEDGEMENTS

The authors wish to thank Dr P. Geissler (G), Dr R.

Grolle (JE) and Dr S. Jovet-Ast (PC) for additional infor-

mation, also Dr T. Bomefeld (Wurzburg) for chromo-

some counts and figures. We are grateful to the curators

of Herbaria B, BCB, BM, BOL, C, Een (Stockholm),

G. Hebrard (Marseille), LISU, MJG, NAM, Ohio State

University, PC, PRC, S, SYD, TFC, UNSW, UPS, US,

W and Z for the loan of specimens.

REFERENCES

ARNELL, S. 1952. Hepaticae collected in South and West Africa,

1951. Botaniska Notiser 105: 307 — 315.

ARNELL, S. 1961. List of the Hepaticae of the Canary Islands. Svensk

botanisk Tidskrift 55: 379-393.

ARNELL, S. 1963. Hepaticae of South Africa, pp. 411 Swedish

National Scientific Research Council, Stockholm.

BOULAY, M. 1904. Muscinees de la France. Hepatiques. pp. 224.

Paris.

BRUNNTHALER, J. 1913. Ergebnisse einer botanischen Forschungs-

reise nach Deutsch-Ostafrika und Sud-Afrika (Kapland, Natal und

Rhodesien). Teil I, p. 14. (Hepaticae bearbeitet von F. Stephani)

in Denkschriften der Mathematisch-naturwissenschaftlichen

Klasse der kaiserlichen Akademie der Wissenschaften 88: 724.

Bothalia 18,1 (1988)

CAMUS, F. 1892. Sur le Riccia nigrella DC. Bulletin de la Societe

Botanique de France 39: 212—230.

CASARES-GIL, A. 1919. Flora Iberica — Briofitas, part 1, pp. 775.

Madrid.

DULL, R. 1984. Bryogeographische Analyse der Makaronesischen

Inseln mit besonderer Beriicksichtigung der Kanarischen Inseln

Tenerife und La Palma. Proceedings of the Third Meeting of the

bryologists from central and east Europe, Praha, 14 — 18 June,

1982: 177- 190. Univerzita Karlova, Praha.

EGGERS, J. 1982. Artenliste der Moose Makaronesiens. Cryptoga-

mie, Bryologie et Lichenologie 3,4: 283—335.

FRITSCH, R. 1982. Index to plant chromosome numbers — Bryo-

phyta. Regnum Vegetabile: 108. pp. 268.

FRYE, T. C. & CLARK, L. 1937. Hepaticae of North America. Uni-

versity of Washington Publications in Biology 6: 1 — 162.

GEISSLER, P. 1985. Notulae bryofloristicae helveticae. II. Candollea

40: 193 - 200.

GEISSLER, P. 1987. F. Stephani, leones hepaticarum. IDC Micro-

form Publishers, Zug, Switzerland.

GOTTSCHE, C. M., LINDENBERG, J. B. G. & NEES ab ESEN-

BECK, C. G. 1844. Synopsis hepaticarum. pp. 835. Hamburg.

Reprint 1967. Cramer, Lehre.

GROLLE, R. 1976. Verzeichnis der Lebermoose Europas und benach-

barter Gebiete . Feddes Repertorium 87: 171—279.

GROLLE, R. 1983. Hepatics of Europe including the Azores: an anno-

tated list of species, with synonyms from recent literature. Journal

of Bryology 12: 403-459.

GUNN, M. & CODD, L. 1981. Botanical exploration of southern

Africa, pp. 400. Balkema.

HOWE, M. A. 1899. The Hepaticae and Anthocerotes of California.

Memoirs of the Torrey Botanical Club 7: 13—33.

JOVET-AST, S. 1969. Le caryotype des Ricciaceae. Revue Bryo-

logique et Lichenologique 36: 673-689.

JOVET-AST, S. 1986. Les Riccia de la Region Mediterraneenne.

Cryptogamie, Bryologie et Lichenologie 7: 287-431 .

JOVET-AST, S. & BISCHLER, H. 1972. Les Hepatiques de Tunisie.

Enumeration, notes ecologiques et biogeographiques. Revue

Bryologique et Lichenologique 38: 1-125.

LAMARCK, J. B. DE & DE CANDOLLE, A.-P. 1815. Flore Fran-

gaise, edn 5,6: 193, 194.

MACVICAR, S. M. 1926. The student' s handbook of British hepatics.

pp. 464. Eastbourne. Reprinted 1964.

MULLER, K. 1952. Die Lebermoose Europas. In L. Rabenhorst,

Kryptogamen-Flora, 6,1: 1-756, 3rd edn. Leipzig.

NA-THALANG, O. 1980. A revision of the genus Riccia (Hepaticae)

in Australia. Brunonia 3: 61 - 140.

NEES ab ESENBECK, C. G. 1838. Naturgeschichte der europaischen

Lebermoose A: 1—540.

PATON, J. A. 1976. Distribution maps of bryophytes in Britain and

Ireland. Journal of Bryology 9: 107-123.

PEROLD, S. M. & VOLK, O.H. 1988. Studies in the genus Riccia

(Marchantiales) from southern Africa. 8. R. campbelliana, (sub-

genus Riccia), newly recorded for the region. Bothalia 18:

37-42.

SCOTT, G. A. M. & BRADSHAW, J. A. 1986. Australian liverworts

(Hepaticae): annotated list of binomials and checklist of published

species with bibliography. Brunonia 8: 1-171.

SERGIO, C. 1984. The distribution and origin of Macaronesian bryo-

phyte flora. Journal of the Hattori Botanical Laboratory 56:

7-13.

SIM, T. R. 1926. The bryophytes of South Africa. Transactions of the

Royal Society of South Africa 15: 1 -475. Cape Town.

STEPHANI, F. 1876-1907. leones hepaticarum. P. Geissler (ed. ).

IDC Microform Publishers, Zug, Switzerland.

STEPHANI, F. 1898. Species hepaticarum. Bulletin de I'Herbier Bois-

sier 6:309-411.

STEPHANI, F. 1913. See under Brunnthaler.

TAYLOR, T. 1846. New hepaticae. The London Journal of Botany 5:

365.

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Riccia. Botanical Gazette ( Crawfordsville ) 19: 273-278.

Bothalia 18,1 (1988)

49

SPECIMENS EXAMINED

R. nigrella

Southern Africa

SWA/NAMIBIA.— 2718 (Grunau): Noachabeb (-BC), Ortendahl

14. vi. 1931 (UPS).

TRANSVAAL. — 2527 (Rustenburg): 2 km to Derby on road from

Rustenburg (— CC), S. M. Perold 888a (PRE). 2528 (Pretoria): 19 km

N of Bronkhorstspruit (-DC), S. M. Perold 150, 320 (PRE); Volk

811023 (M, PRE). 2530 (Lydenburg): 29 km to Dullstroom on R540

( — AB), S. M. Perold 425 (PRE). 2630 (Carolina): Chrissiemeer, Farm

Knock Dhu (-AD), Smook4892a (PRE). 2729 (Volksrust): 9 km from

Amersfoort on road to Perdekop (— BB), S. M. Perold 1088a (PRE).

O.F.S. — 2729 (Volksrust): 31 km NE of Verkykerskop, on road

from Memel (-CD), S. M. Perold 1274 (PRE); 2827 (Senekal): Alle-

manskraal Dam, on plateau near caravan park (-AC), Volk 811214c

(M, PRE); Senekal, near reservoir on koppie (-BC), S. M. Perold

1336, 1340 (PRE); 6 km N of Clocolan on road to Marquard (-DC),

S. M. Perold 1322 (PRE). 2926 (Bloemfontein): Bloemfontein

(-AA), Duthie 5457a, 5494a (BOL); Bloemfontein, next to Bot. Gar-

den (-AA), Volk 81-289 p.p. (M); 30 km S of Bloemfontein on N1

(-AC), S. M. Perold 956 (PRE). 2927 (Maseru): 10 km from Lady-

brand on R26 ( — AC), J. M. Perold 35b (PRE). 3026 (Aliwal North):

between Zastron and Wesselsdale, Farm Olievenrand (-BB), Van

Rooy 2414, 2417 ( PRE).

NATAL. — 2930 (Pietermaritzburg): Edendale (-CB), Sim CH 1025

(with R. atropurpurea) (PRE).

CAPE. — 2917 (Springbok): Komaggas, top of T'nouroegas — Van

Reenen se water plateau (-DA), Glen 1472, 1585 (PRE). 2924 (Hope-

town): 54 km from Kimberley on road to Hayfields (-BA), Smook

3544a (PRE). 3024 (De Aar): 25 km from De Aaron road to Phillips-

town. Farm Welgegund ( — AC), Smook 3418a (PRE). 3027 (Lady

Grey): 23 km S of Lady Grey (— CC), Van Rooy 2598 (PRE). 3218

(Clanwilliam): 20 km N of Citrusdal, near Farm Hexrivier ( — BD ),

S. M. Perold 533 (PRE); Weltevrede, 7 km W of Piketberg (-DC),

S. M. Perold 501 (PRE); Middelpos (-DC)S. M. Perold 506 (PRE).

3219 (Wuppertal): 18 km S of Citrusdal (-CA), S. M. Perold 520

(PRE). 3225 (Somerset East): Cradock Mtn Zebra Park ( — AD), Smook

6028 (PRE). 3318 (Cape Town): Lion's Head (-CD), Arnell 26, 36

(S): Arnell 49, 59 (S, UPS); Arnell 50 (BOL); Newlands, Pillans’s

garden (-CD), Garside 6650 (BOL); Newlands, Schelpe’s garden

(-CD), Lambert 1 (BOL, PRE); near University of Cape Town

(-CD), Loxly 5023 (BOL); Tamboerskloof (-CD), P.K. 5368 (BOL);

Rosebank (-CD), leg. unknown 5023 (BOL); Stellenbosch ( — DD),

leg. unknown 5019a (BOL); Glen Bawn ( — DD), Duthie 5340 (BOL);

13 km from Franschhoek on road to Paarl (-DD), S. M. Perold 1147

(PRE). 3319 (Worcester): Tulbagh ( — AC), Vogel C682 (MJG, PRE).

3320 (Montagu): ‘Witteberge' turnoff on Nl. before Laingsburg

(-AB), C. M. van Wvk 2523, 2525 (PRE); Cogman’s Kloof (-CC),

Arnell 792 (BOL, PRE, S, UPS); Arnell 802 (BOL, PRE, S). 3418

(Simonstown): Wynberg Park Hotel garden ( — AB), Arnell 150 (BOL,

PRE, S); Arnell 162 (PRE); 186 (S); 189, 302 (BOL); 302a & b (S);

near Eerste River (-BA), Van Brake! 5019 (BOL). 3419 (Caledon):

Greyton, kloof, on earth bank (- BA), S. M. Perold 1171a (PRE).

Europe

BRITAIN . — Pearson 290 (C); April 1877 CH542, June 1879

CH473 (PRE); May 1885 (S); April 1881 (UPS).

PORTUGAL. — Casas et al.2,4, 10, 11 (LISU); Sergio et al. 26.1,

28.3 (LISU).

SPAIN.— S. Arnell 6.5.1960 (S); S. Arnell 3.5.1960 (UPS); Casas

12.4.57, 16.4.1957, 4.4.1958, 25.9.76, 3.1.1984 , 2.5.1984 (BCB);

Cros etal. (BCB); Volk 75-584, 82-983 (M).

FRANCE. — D' A rice Fevr. ’84 (UPS)', D’Arice Mars ’89 (UPS);

Boulay Avril '96 (UPS); Crozals 311902 (ex Schiffner) (W, S); De

Sloover 25.130 (PRC); Hebrard 1-4382, 1-5283, 3-14586 (Private

Herb. J. P. Hebrard )\K. Preis 27.5 .37; JirtVana4.4.1977 (PRC).

ITALY.—//. Gams 3.4.37 (GB, S); 15.2.1952 (S); Levier (Bryo-

theca Italica; ex herb. Guzelberg; G, S, UPS); Milde 2 Aug. ’97 (UPS);

Oliver April 1867 (UPS).

YUGOSLAVIA. — Baumgartner 5.4.1908, 15.3.1910, 28.6.1911,

19.4.1913 (W); Glowacki 26.7.1896 (S); Latzel 10 Febr. 1908:

Schiffner 10.4.1909, 12.4.1909 ; Weiss29Jan. 1867 ( W).

GREECE. — Volk 85-802, 86-948 (M).

NORTH AFRICA. — Trabut, March 1908, 1171 (1939) (S).

Macaronesia

CANARY ISLANDS.— S. Arnell 28.2.1959, 5.3.1959 (S); Losada

& Rivero (TFC).

MADEIRA ARCHIPELAGO— M. Nobrega 20.1.1954, 21.1.

1954, 4. 11. 1953 (S).

United States of America

CALIFORNIA.—//. N. Bolander 26: Brandeger 1892 (NY); A. M.

Carter 252 (PRC); Howe 165 (NY); April 1894 (Ohio State Univ.);

Feb. 8, 1896 (W); E. W . Koch s.n.: Dale M . J . Mueller 6905, Walter

R. Shaw 1901; no collector’ s name, 1704 (in part), 1729: no collector’ s

name or number (NY).

MISSOURI. — J. A. Steyermark428 (NY).

PENNSYLVANIA. — T. C. Porter s.n. (NY).

TEXAS.—/7. McAllister, March 1929 :M. S. Young ( NY).

Australia

NEW SOUTH WALES. — Na-Thalang 54, 72. (SYD).

SOUTH AUSTRALIA. — Na-Thalang 126 (SYD).

R. trabutiana Steph. incorrectly determined as R. ni-

grella

PORTUGAL. — Casas et al. 2, /O(LISU).

CANARY ISLANDS. — S. Arnell 4.3.1959, 7.3.1959 (S); 28.12.

1951 (UPS).

R. atromarginata Lev. incorrectly determined as R.

nigrella

SPAIN.— S. Rungly 2931 (C).

FRANCE. — S. Arnell 16.7.53 (UPS).

R. congoana Steph. incorrectly determined as R. capen-

sis Steph.

TANZANIA.— G. Een 14.3.1967. (Herb. Een, S).

R. atropurpurea Sim incorrectly determined as R. ni-

grella var. austro-africana

ZIMBABWE. — Sim July 1920 (PRE, UPS).

SWA/NAMIBIA. — Volk 2766 (PRE).

TRANSVAAL. — Sim Jan. 1918 (PRE).

Bothalia 18,1:51-56(1988)

Two closely related species of Caloplaca (Teloschistaceae, Lichenes)

from the Namih Desert

I. KARNEFELT*

Keywords: anatomy, Caloplaca elegantissima, C. namibensis, sp. nov., Lichenes, Namib Desert. Teloschistaceae

ABSTRACT

The anatomical and reproductive adaptations of two closely related lichen species, Caloplaca elegantissima (Nyl.)

Zahlbr. and C. namibensis Kamef., sp. nov., occurring in the outer Namib fog desert, are discussed. Both species belong

to the crustose forms, frequently found in the remarkably rich lichen communities, which largely depend on fog precipita-

tion for their water supply. Both species are endemic to the Namib Desert. They are mainly distributed in South West

Africa/Namibia but also extend into south-western Angola. The asexual isidiate species, C. namibensis Kamef., is

described as new.

UITTREKSEL

Die anatomiese en voortplantingsaanpassings van twee naverwante ligeenspesies, Caloplaca elegantissima (Nyl.)

Zahlbr. en C. namibensis Kamef., sp. nov., wat in die buitenste Namib-miswoestyn voorkom, word bespreek. Albei

spesies behoort tot die korsagtige vorms wat dikwels aangetref word in die buitengewoon ryk ligeengemeenskappe wat

hoofsaaklik van die misneerslag afhanklik is vir water. Albei spesies is endemies in die Namibwoestyn. Hulle kom

hoofsaaklik in Suidwes-Afrika/Namibie voor maar is ook tot in suidwestelike Angola versprei. Die ongeslagtelike isidiate

spesie C. namibensis Kamef. word nuut beskryf.

INTRODUCTION

The Namib is a narrow belt of desert more than 2 000

km long, stretching northwards from the Olifants River

in the Cape Province (RSA) to Mossamedes in Angola.

The central Namib Desert is characterized by the world's

most magnificent sand dunes. The gravel desert, occupy-

ing most of the landward parts of the central and northern

Namib Desert, is usually almost devoid of higher plants.

The few which occur are mainly succulents with specia-

lized anatomical and physiological adaptations to survive

in the harsh environment (Walter 1973; White 1983).

Ephemeral species appear only after extremely rare rain

showers.

The desert is, however, not entirely devoid of vegeta-

tion. Rock outcrops, stones and pebbles are frequently

covered with colourful lichens. Even the seemingly bar-

ren soil can be covered with fruticose lichens over large

areas, similar to the lichen cover which is a familiar sight

in arctic or subarctic regions in the northern hemisphere.

Kappen (1982) estimated the biomass in one of these

areas at 250 g/m2. The existence of this spectacular

lichen vegetation is entirely due to the effect of frequent

fogs. These fogs are caused by the cold Benguela current

which flows northwards from the southern Atlantic

Ocean along the south-western coast of Africa. Fog-

induced lichen communities also occur in the coastal

deserts of Peru and Baja California (Thomson & litis

1968; Rundel etal. 1972).

Lichens occurring in these coastal desert areas can

evidently absorb sufficient amounts of moisture from fog

to allow photosynthesis for successful reproduction and

dispersal. Lange et al. (1970a, 1970b) studied the effect

of dew on lichen communities in the Negev Desert.

* Department of Systematic Botany, University of Lund, Ostra

Vallgatan 18, 223 61 Lund, Sweden.

MS. received: 1986.10.13.

According to them the early morning dew provided suffi-

cient moisture for about 3 h of photosynthesis. Apart

from the effect of fog, the annual precipitation in the

central Namib is not more than 30 mm, and it may not

even rain every year (Kappen 1982).

More or less well known lichens occurring in the

desert are Combea mollusca (Ach.) Nyl., Santessonia

namibensis Hale & Vobis, Teloschistes capensis (L. f.)

Miill. Arg., Xanthomaculina convoluta (Hue) Hale,

Xanthomaculina hottentotta (Ach.) Hale and Xanthoria

marlothii Zahlbr. All these species have various anatom-

ical adaptations which aid in the reduction of light inten-

sity and evaporation. Biidel & Wessels (1986) discussed

the remarkable anatomical adaptions of Xanthomaculina

convoluta (as Parmelia hueana Gyeln.). It is presumed

that heavily pigmented layers, as encountered in Xantho-

dactylon flammeum (L. f.) Dodge, could be effective in

reducing light intensity. Members of this species are less

deeply pigmented in populations further south, in the

Cape Province.

Teloschistes capensis (L. f.) Miill. Arg., forms exten-

sive populations in areas north of Swakopmund, and has

a deeper pigmentation than similar populations found in

the Cape. Individuals which occur in the central Namib

are furthermore covered by a rather dense tomentum,

which is much sparser in the Cape. Presumably this

dense tomentum favours the absorption of water derived

from the coastal fog.

Caloplaca eudoxa (Miill. Arg.) Zahlbr. is another

remarkable endemic member of the Namib lichen flora.

The species is characterized by a subfruticose growth

form and thick scleroplectenchymatous hyphae which

cover the photobiont (Poelt & Pelleter 1984). The two

Caloplaca species discussed in this paper, C. elegantis-

sima (Nyl.) Zahlbr. and C. namibensis Kamef., have

52

Bothalia 18,1 (1988)

similar anatomical adaptations. They are among the most through a Leitz, Ortholux microscope. Secondary corn-

abundant and spectacular crustose lichen species pounds present in the material were determined by

endemic to the Namib. means of standardized TLC methods (Culberson 1972).

MATERIAL AND METHODS

THE SPECIES

The material which formed the basis of this study

came mainly from my own collections made during a

visit to the central Namib fog desert in January 1986. I

have also studied a few collections on loan from B, BM,

G, H, LISU and S. The macrophotographs were taken

with an Olympus OM-2 camera, through a Zeiss dissect-

ing microscope.

For anatomical studies, the material was sectioned on

a Kryomat, Leitz freezing microtome. The sections were

embedded in lactophenol cotton-blue, studied, and

photographed by means of an Olympus OM-2 camera.

1. Caloplaca elegantissima (Nyl.) Zahlbr., Cata-

logus lichenum universalis 7: 238 (1931). Lecanora ele-

gantissima Nyl.: 510 (1868). Type: SWA/Namibia,

2114 (Omaruru): Lagunenberg Mountain, north-east of

Mile 72 (— CC), Karnefelt 8605—26 (LD, neo., here

designated).

Caloplaca diploplaca Zahlbr.: 268 (1932). Type: SWA/Namibia,

1927, Kriege sub P. van der Byl 709 (STEU, lecto. , selected here).

Caloplaca indurata Wirth & Vezda: 1 (1975). Type: Africa austro-

occidentalis, Swakopmund, 15 km ad orientem urbis Swakopmund,

Sept. 1974, Volk, in Vezda, Lich. sel. exs. 1346 (STU, holo.; iso.

distributed in Vezda, Lich. sel. exs. 1346, seen in H, LD, S).

FIGURE 1 . — a, C. namibensis, cross section, dark pigmented layer and thick cortical layer covering the darkly stained photobiont, Karnefelt

8602-56 (LD), bar = 100 /xm; b, C. namibensis, cross section, Karnefelt 8602-56 (LD), bar = 100 /xm; c, C. namibensis, cross section

showing structure of cortical layer, Karnefelt 8603a-5 (LD), bar = 10 pm; d, C. elegantissima, cross section showing structure of cortical

layer, Desmond 25276 (LD), bar = 10 pm; e, C. elegantissima, cross section of specimen showing the internal pigmented layer and thick

cortical layer covering the basally located photobiont, overlying a medullary layer with crystals, Vezda, Lich. sel. exs. 1346 (LD), bar =

100 /xm; f, C. elegantissima, cross section showing the internal pigmented layer, Vezda, Lich. sel. exs. 1346 (LD), bar = 10 /xm.

Bothalia 18,1 (1988)

53

Thallus radiate, (5— )20— 50(— 100) mm across, the

smallest occurring on pebbles, the largest on rather large

rocks, composed of cartilaginous convex lobes 0,5-1 ,8

mm broad, 5-30 mm long, marginally distinctively effi-

gurated, weakly dichotomously branched, secondarily

becoming raised and unattached; central portions com-

posed of short, 1-3 mm long, irregularly arranged,

rather sparse lobes; short accessory lobules occasionally

developed at the margins of the lobes; scarlet, orange-red

to pale orange, larger individuals usually lighter pig-

mented at margins.

Epinecral zone 20-45 pm thick, prosoplectenchyma-

tous, hyphae thick-walled (Figure Id, e). Cortex hya-

line, 150-230 pm thick, composed of strongly gelati-

nized hyphae, the upper 15-20 /xm, orange-yellowish

pigmented (Figure Id, e, f). Photobiont green and sphe-

rical, 5,5—12 pm in diameter, occurring in clusters

35-75 jam large, mainly embedded in a layer of

strongly gelatinized hyphae. Medulla 200-250 pm

thick, composed of more lax hyphae, partly embedding

the clustered photobiont, often containing numerous

small granules, fluorescent when examined with inter-

ference contrast microscopy.

Apothecia scattered, limited mainly to central thallus

portions, but may also develop marginally on the peri-

pheral lobes, sessile, becoming slightly raised with

maturity, 0,5— 1 ,5 mm across, margin entire or occasio-

nally irregular, several smaller apothecia occasionally

fused; disc at first cupular or plane, becoming strongly

convex, usually darker or of the same colour as the

lobes. Excipulum up to 100 pm thick. Hymenium

75—100 pm thick, covered by a thin epihymenial layer.

Asci 45-50 x 9—12 pm. Ascospores broadly ellipsoi-

dal, 10—12 x 8—9 pm, septum ± 2 pm thick. Para-

physes 75—100 x 0,75 pm. Hypothecium up to 300 jum

deep, hyaline. Photobiont concentrated in clusters below

the hypothecium. Conidiomata not observed.

SWA/NAMIBIA. — 2113 (Cape Cross): Skeleton Coast Park,

marble ridge north of entrance gate at Ugabmund (-BA), Karnefelt

8609-5 ; 8609-6- 8609-22 ; 8609-24 ; 8609-30: 8609-39 (LD). 2114

(Omaruru): Lagunenberg Mountain, north-east of Mile 72 (-CC),

Karnefelt 8605-19: 8605-20: 8605-26 (LD); gravel flats east of Cape

Cross (-CA), Karnefelt 8610-29: 8610-37: 8610-38 (LD); Wessels

5132: 5134a (UNIVERSITY OF THE NORTH); 25 km north of Hen-

ties Bay Road junction, black ridge (-CA), Nordenstam & Lundgren

2317 (S). 2214 (Swakopmund): gravel flats east of Swakopmund

(-DA), Karnefelt 8602-33: 8602-34: 8602-19: 8602-52 (LD); 15 km,

in Sept.-oriente a urbe Swakopmund (-DA), 1974, Volk in Vezda,

Lich. sel. exs. 1346 as C. indurata (H, LD, S); 35 km north of Swa-

kopmund (-AD), Nordenstam & Lundgren 2322 (S); Namib Desert

(-??), Desmond 25276 (LD).

C. elegantissima contains several anthraquinones,

which were difficult to separate with the TLC method

used. Extracts of C. elegantissima and C. namibensis

were examined together on the same TLC plate, but the

anthraquinones could not be distinguished due to the

weakness of the spots (the anthraquinones are difficult to

Extract from the cortex). Steiner & Hauschild (1970)

reported the presence of emodin (traces), parietin, telo-

schistin, xanthorin, erythroglaucin, fallacinal and parie-

tinic acid in C. elegantissima. No material was cited, but

it is presumed that the fertile species was investigated as

it is the most conspicuous of the two species.

Typification

The original description of Lecanora (Placodium)

elegantissma was presumably based on composite mate-

rial involving both species discussed in this article. The

short morphological diagnosis described a narrow-lobed

specimen resembling Placodium elegans [=Xanthoria

elegans (Link) Th. Fr. ] , but no isidia or similar struc-

tures were mentioned (Nylander 1868). The diagnosis,

however, included a description of apothecia and asco-

spores and the original spore size mentioned corresponds

well with my own results based on examined asci of C.

elegantissima.

The fragment (annotated ‘vestigio’) in the Nylander

herbarium at Helsinki, does not belong to C. elegantis-

sima but to C. namibensis. It therefore cannot possibly

be used as the type of C. elegantissima. The isidia have,

however, been tom off during more than 100 years of

storage, leaving distinctive marks at their previous loca-

tion (H-NYL 30513).

Nylander cited no material other than Welwitsch’s

collections from Mossamedes (erroneously spelled

Monamedes). In a footnote he discussed a new species

Placodium flavorubens Nyl. [ =Caloplaca flavorubens

(Nyl.) Zahlbr.] occurring on coastal rocks in southern

Africa, which presumably is identical to C. sublobulata

(Nyl.) Zahlbr. Since C. elegantissima and C. namibensis

occur in the same habitats and occasionally also cover

the same stones, Welwitsch most probably collected

both species on a few of the stone samples. On Wel-

witsch’s request, Nylander eventually studied these

samples while in Paris. He must have received a mixed

collection of mainly C. namibensis, which also included

a few lobes and apothecia of C. elegantissima.

Other possible type material from Welwitsch’s origi-

nal collection is kept at BM and LISU (see Vainio 1901 ).

The material I received on loan from these herbaria, no.

45 Welwitsch Iter Angolense, Mossamedes, near Cabo

Negro, unfortunately turned out to belong to the same

species described here as new, and characterized by

rather narrow lobes and small isidia or fragments of

isidia. Vainio (1901) also cited nos 48 and 49 as pr. p.

under his Placodium elegantissimum, but these numbers

are mainly a brown Parmelia, and it is not likely that

Nylander used this material for his original description.

Since all known type material of C. elegantissima

must be superseded as being in serious conflict with the

protologue (Art. 8 of the I.C.B.N.), it is either possible

to choose a neotype, or to have the name rejected in

accordance with Art. 69 of the I.C.B .N. In fact, I did not

see any material in the original collections which corre-

sponds in any way to C. elegantissima, other than the

type material of much younger epithets such as C. diplo-

placa Zahlbruckner (1932) and C. indurata Wirth &

Vezda (1975).

The provisons of Art. 69 (a name must be rejected if it

has been widely and persistently used for a taxon not

including its type), is not really relevant in this case. 1

have therefore decided to select a neotype from my own

collections of C. elegantissima.

54

Bothalia 18,1 (1988)

FIGURE 2. — a, Caloplaca elegantissima, well developed fertile specimen on dolerite, Karnefelt 8605-26 (LD); b, C. elegantissima, specimen

dominated by rather short central lobes, on dolerite, Karnefelt 8605- 39 (LD); c, C. elegantissima, specimen on quartzite pebbles,

Karnefelt 8605-19 (LD); d, C. elegantissima close-up of specimen on quartzite pebble; accessory lobules seen in c & d, Karnefelt 8605-20

(LD); e, C. namibensis, close-up of specimen with rather long densely arranged marginal lobes, on quartzite, Karnefelt 8602-36 (LD); f,

C. namibensis, specimen on quartzite pebbles, the central portions covered with minute isidia, Karnefelt 8610-2 (LD). All bars = 10 mm.

2. Caloplaca namibensis Kdrnef. , sp. nov.

ICaloplaca diploplaca Zahlbr. var. gracilior Zahlbr., Annales de

cryptogamie exotique 5: 269 (1932). Type: SWA/Namibia [Deutsch-

Sudwestafrika], Haifischbucht, Fincke (W. not seen).

Thallus Caloplacae elegantissimae similis sed lobis

marginalibus densioribus et angustioribus, partibus lobo-

rum centralium saepe magis areolatis et isidiis sat nume-

rosis instructis differt.

TYPE.— SWA/NAMIBIA, 2114 (Omaruru): gravel

flats east of Cape Cross (— CA), Karnefelt 8610-2 (LD,

holo.).

Thallus radiate, (5 — ) 10 — 30(— 60) mm across, the

smallest occurring on pebbles, occasionally up to 60 mm

across on larger stones, composed of cartilaginous,

rather thin, densely placed lobes, 0,2— 0,5 mm wide.

broadest at the lobe tips; marginal lobes usually 5-15

mm long, distinctively effigurate, weakly dichotomously

branched, secondarily becoming raised and unattached;

central portions composed of short, irregularly arranged

or occasionally more areolated lobes, covered with

simple or coralloid, rather small isidia; short accessory

lobules occasionally develop at the margins of the lobes;

orange to pale orange, the central portions usually pale

brownish due to the accumulation of numerous small

sand grains.

Epinecral zone 10-20 ^tm thick, prosoplectenchyma-

tous, hyphae thick-walled (Figure la, c). Cortex 1 20—

230 (urn thick, composed of strongly gelatinized hyphae,

hyaline but orange-yellowish in a thin layer above (Fig-

ure la, b, c). Photobiont green and spherical, 5,5-12

pm in diameter, occurring in clusters, 35—85 pm large,

Bothalia 18,1 (1988)

55

mainly embedded in a layer of gelatinized hyphae. Me-

dulla 200—300 gm thick, composed of more lax hy-

phae, partly embedding the clustered photobiont, often

containing small granules, fluorescent when examined

by means of interference contrast microscopy. Ascomata

and conidiomata not observed.

Caloplaca namibensis contains several anthraqui-

nones which were difficult to separate by means of TLC

(see discussion of C. elegantissima). I have not seen the

type material of Caloplaca diploplaca Zahlbr. var. gra-

cilior Zahlbr. The description, however, indicates that

this taxon belongs to C. namibensis (see Zahlbruckner

1932).

ANGOLA. — 1213: Benguela, Welwitsch s.n., 1859, (H-NYL

30513). 1511: Mossamedes, between Caroca and Cazimba at Cabo

Negro, Welwitsch, Iter Angolense 45 (BM, LISU).

SWA/NAMIBIA. — 21 14 (Omaruru): gravel flats east of Cape Cross

(-CA), Kdrnefelt 8610-2 ; 8610-4 ; 8610-12 ; 8610-15; 8610-18- 8610-

20; 8610-23 ; 8610^42 (LD). 2214 (Swakopmund): gravel flats east of

Swakopmund (-BA), Kdrnefelt 8602-4 ; 8602-7 ; 8602-12 ; 8602-2 1;

8602-23-8602-25 ; 8602-32; 8602-35; 8602-36; 8602-50; 8602-53;

8602-54; 8602-56 (LD); Wessels 5082; 5085; 5087 (UNIVERSITY OF

THE NORTH); Omaruru flumen (— AB), Melander 3 (H); 22 km east

of Walvis Bay (—DC), Nordenstam & Lundgren 2311 (S); between

Swakopmund and Cape Cross (—??), Mattick 6812 (B). 2315

(Rostock): Namib Naukluft Park, Gobabeb, coastal desert habitat

(-CA), Kdrnefelt 8603a-3-8603a-5;8603a-9(LD). 2615 (Luderitz):

Angra Pequena (— CA), 1888 Schinz (G).

Characteristics and differences

C. elegantissima is characterized by slightly sepa-

rated, peripherally effigurated, cartilaginous, strikingly

scarlet or orange-reddish lobes (Figure 2a). The central

part of the thallus, especially in large well developed

individuals, usually consists of short irregularly arranged

lobes (Figure 2b), and the species is often fertile.

C. elegantissima differs from the closely related

C. namibensis, by having slightly broader (0,5- 1,8

mm), and more separated lobes, whereas C. namibensis

has more closely spaced (contiguous) lobes, 0,2-0, 5

mm wide, with almost no interspaces (Figure 2e, f).

However, the main difference between the two species is

that C. namibensis is isidiate, and C. elegantissima is

not. When growing side by side, as is often the case on

the gravel plains, C. elegantissima appears as the more

robust of the two species. The specific epithet elegantis-

sima, therefore describes the isidiate species better than

the sexual one.

Habitat, ecology and reproduction

C. elegantissima and C . namibensis are two of the

most common species in the Namib fog desert. As with

most other species which occur in that region of the

Namib Desert, they seem to be perfectly adapted to the

extreme local environmental conditions.

Two other rather common crustose species, C. volkii

Wirth & Vezda (1975) and Lecidella crystallina

Wirth & Vezda (1975), normally occur directly on the

ground. C. elegantissima and C. namibensis, on the

other hand, grow on rocks of various sizes. Individuals

on small pebbles, obviously a very common occurrence,

can cover the substrate completely (Figure 2c, d).

C. elegantissima, in particular, seems to occur rather

frequently on small pebbles, ± 5 mm in size, whereas

C. namibensis is more common on stones, 10 to 30 mm

in size. The most luxuriant and richly fertile specimens

of C. elegantissima were found on dolerite outcrops

composed of larger stones and rocks. This observation

may be correlated with the effect of drifting fog, which

condenses more rapidly on cooling rock outcrops (Kap-

pen 1982).

Sexual reproduction is presumably an important mode

of dispersal for the frequently richly fertile C. elegantis-

sima, whereas asexual dispersal by means of isidia must

be an important mode of reproduction forC. namibensis,

producing extensive clones. However, accessory lobules

are frequently developed in both species, especially on

smaller individuals growing on pebbles (Figure 2c, d).

Fragmentation of the frequently raised lobes could also

be interpreted as an adaptive arrangement for clonal dis-

persal in this regionally successfully distributed species

pair.

Anatomical adaptations

The most typical environmental adaptation of these

species is the anatomical structure, which is character-

ized by extremely thick external tissue (Figure 1). This

FIGURE 3. — Distribution of Caloplaca elegantissima. Arrow indi-

cates localities in southern Angola.

FIGURE 4. — Distribution of Caloplaca namibensis. Arrow indicates

localities in southern Angola.

56

Bothalia 18,1(1988)

tissue, distinctively stratified into epinecral and cortical

layers, is 185 to 295 jtim thick in C. elegantissima and

130 to 250 fim thick in C. namibensis. The tissue forms

a solid cover over the basally located photobiont (Figure

lc, d). A thick cortical layer acts mainly in reducing the

intensity of light (Kappen 1973; Vogel 1955), in reduc-

ing evaporation loss and also in permitting direct absorp-

tion of water vapour from dry air (Galun 1963; Rogers

1977).

In C. elegantissima, the pigmented cortical zone is

generally better developed than in the closely related

species (Figure le, f). In addition, the cortex in C. el-

egantissima is composed of more strongly gelatinized

hyphae, than that of C. namibensis.

Distribution

C. elegantissima and C. namibensis are endemic to

the outer Namib Desert, and are known to occur from

south-western Angola in the north, to the Liideritz region

in the south (Figures 3 & 4). It is suspected, however,

that the area of both species may extend to the southern

boundary of the Namib Desert. Closely related species

from other coastal fog deserts of the world are not

known.

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VOGEL, S. 1955. Niedere ‘Fensterpflanzen’ in der siidafrikanischen

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Bothalia 18,1:57-77(1988)

The Cheilanthes hirta complex and allied species (Adiantaceae/Pteri-

daceae) in southern Africa

W. B. G. JACOBSEN* and N. H. G. JACOBSEN**

Keywords: Adiantaceae, Cheilanthes hirta Swartz complex, evolution, key, phytogeography, Pteridaceae, southern Africa, spore morphology

ABSTRACT

The very variable complex of plants until now ascribed to Cheilanthes hirta Swartz, together with some derived or

allied species, is investigated. The type of the species as established by N. C. Anthony (1984), is accepted. Three new

varieties are distinguished: Cheilanthes hirta Swartz var. brevipilosa W. & N. Jacobsen, var. inferacampestris W. & N.

Jacobsen and var. nemorosa W. & N. Jacobsen. One new form of Cheilanthes hirta var. brevipilosa W. & N. Jacobsen is

recognized: forma waterbergensis W. & N. Jacobsen. Var. laxa Kunze (1836) is given the new status of forma: Chei-

lanthes hirta var. brevipilosa W. & N. Jacobsen forma laxa (Kunze) W. & N. Jacobsen. Three allied species are discussed

and included in the key to all taxa mentioned. Taxa are described and information given includes notes on distribution and

ecology. Special attention is paid to spore morphology. Most taxa are illustrated by a habit photo, and all by line drawings

and scanning electron micrographs (SEM) of the spores. Possible evolutionary conclusions, particularly on the nature of the

spores and on frond dimensions in relation to geographical distribution and climate are discussed.

UITTREKSEL

Die hoogs varierende plantkompleks wat tot nou toe by Cheilanthes hirta Swartz ingesluit is, tesame met sommige

afgeleide of verwante spesies, word ondersoek. Die tipe van die spesie soos deur N. C. Anthony vasgestel, word aanvaar.

Drie nuwe varieteite word onderskei: Cheilanthes hirta Swartz var. brevipilosa W. & N. Jacobsen, var. inferacampestris

W. & N. Jacobsen en var. nemorosa W. & N. Jacobsen, lien nuwe vorm van Cheilanthes hirta var. brevipilosa W. & N.

Jacobsen word erken: forma waterbergensis W. & N. Jacobsen. Var. laxa Kunze (1836) word hier die nuwe status van

forma gegee: Cheilanthes hirta var. brevipilosa W. & N. Jacobsen forma laxa (Kunze) W. & N. Jacobsen. Drie verwante

spesies word bespreek en by die sleutel tot alle genoemde taksons ingesluit. Taksons word beskryf en inligting wat verskaf

word, sluit aantekeninge oor verspreiding en ekologie in. Spesiale aandag word aan spoormorfologie gegee. Die meeste

taksons word ge'illustreer met ’n foto van die groeiwyse, en almal met lyntekeninge en aftaselektronmikrogawe (SEM) van

die spore. Moontlike evolusionere gevolgtrekkings, veral oor die aard van die spore en oor blaarafmetings met betrekking

tot geografiese verspreiding en klimaat, word bespreek.

CONTENTS

Introduction and aims 57

History and synonymy 58

Material and methods 59

Results 59

Key to taxa 60

Description of taxa 60

1. C.contracta 60

2. C. hirta var. hirta 61

3. C. hirta v ar. brevipilosa 64

4. C. hirta var. brevipilosa forma laxa 65

5. C. hirta var. brevipilosa forma waterberg-

ensis 68

6. C. hirta var. nemorosa 69

7. C. hirta var. inferacampestris 71

8. C. hyaloglandulosa 72

9. C. nielsii 72

Discussion

Spores and possible evolution 73

Morphology, phytogeography and climate 74

Acknowledgements 76

* P.O. Box 1178, White River 1240.

** Nature Conservation Division, Private Bag X 209, Pretoria 0001 .

MS. received: 1986.04.07.

References 76

Specimens examined 77

INTRODUCTION AND AIMS

Amongs the 24 currently accepted species of Chei-

lanthes Swartz (Anthony 1984) in southern Africa, the

widespread C. hirta Swartz contains in its broader con-

cept a number of rather different looking plants. In this

respect it resembles the equally widespread C. viridis

(Forssk.) Swartz complex which has been split into a

number of varieties or separate species (Anthony 1984).

This has so far not been done in the case of C. hirta.

Since the first naming of the species by Swartz as Adian-

tum caffrorum in 1801 and its description by him in 1806

as Cheilanthes hirta, it has been referred to under a

number of different specific epithets in different genera

and with varying breadth of concept until as recently as

1983 (Jacobsen) and 1984 (Anthony).

The present paper deals with the varying forms, varie-

ties and allied species which were at one time or another

included in the species concept. The authors realize that,

similar to the C. viridis complex, the various forms of C.

hirta will propagate partly sexually and partly apoga-

mously with the possibility of the more xerophytic ones

being apogamous. Cytological studies might clarify in

what way polyploidy and hybridization affect their evo-

lution. This is at present being done with a number of

cheilanthoid ferns of Europe, the Mediterranean and

58

Bothalia 18,1 (1988)

Macaronesia and it has led to the creation of a great

number of hybrids, each with its own name. The authors

are not equipped to do this, but they have made a

thorough examination of morphological features, includ-

ing those of the spores, and they conclude that their

findings warrant separation of the C. hirta complex into

a number of taxa.

HISTORY AND SYNONYMY

The typification of the species presents difficulties. It

could either be based on a collection by Dr Groendal in

Herb. Gastroemi of a plant from Mauritius in the

Riksmuseet, Stockholm, or on a collection by Thunberg

from the Cape of Good Hope in Herb. Thunberg,

Uppsala. Certain entries by Wikstroem on the former

sheet seem to indicate that this is indeed the specimen on

which Swartz (1801: 85) based his Adiantum caffrorum

and on which he himself changed the name to Chei-

lanthes hirta (Swartz 1806: 128, 329). Anthony (1984)

accepts the sheet from Mauritius as the type of the spe-

cies. Figure 1 .

Professor N. Lundqvist of the Cryptogamic Section of

the Riksmuseet (personal correspondence) is of the opi-

nion that Swartz typified both his Adiantum caffrorum

and his Cheilanthes hirta with material from South

Africa and that the sheet from Mauritius is not the type.

He also mentioned that it was not marked thus by Prof.

Schelpe, who quotes it as the type (1969: 72). The sheet

in the Thunberg Herbarium is, according to Lundqvist,

not the type, as Swartz had not seen it. He did not com-

ment on the possible locality of the type sheet. Thus the

question of the whereabouts of the type sheet is not

solved. It appears best, however, to follow Anthony

(1984) and to accept the sheet from Mauritius at least as

the lectotype. The specific epithet hirtum or hirta based

on Swartz’ material was combined by various authors

with various genera, which are now included in Chei-

lanthes.

Kunze recognized the variability of the species in

1836 and distinguished four forms: a contracta, P inter-

media, y laxa and 5 parviloba. The types of these names

were housed in his herbarium in Leipzig and were

destroyed during the last war in air attacks on the city. In

later works, beginning with Pappe & Rawson (1858),

and including those of Sim (1915), Jacobsen (1983) and

b

FIGURE 1.— a, a form of C.

hirta, collected 30.4. 1905 at

Koedoespoort near Pretoria,

Leendertz 622, X 0,5; b,

photostat of the lectotype of

Cheilanthes hirta Swartz in

Herb. Gastroemi, Riksmu-

seet, Stockholm, x 1.

Bothalia 18,1 (1988)

59

Anthony (1984), the forms of Kunze are referred to as the correct name of the plant is now C. hirta Swartz var.

varieties, a procedure in agreement with Article 35.3 of brevipilosa W. & N. Jacobsen forma laxa (Kunze) W. &

the Code. Anthony (1984) and Roux (1986) have investi- N. Jacobsen. The photostat of an isosyntype of C. hirta

gated the typification of Kunze’s forms. They have Swartz var. laxa Kunze shows six very damaged fronds,

established the following: collected by Drege and now housed at the Museum in

Leiden under Morton 751. The locality is not known.

1 , var. contractu is synonymous with C. contractu The habit of these Plants is similar t0 the 'ectotype above

(Kunze) Mett. , a species on its own; and the Plants are considered to be a good match of it.

2, for var. intermedia, Anthony (1984) selects as lec-

totype a collection by Drege from the Witberg in the

southern Cape, now in B under No. 000414 and deter-

mined by Kunze. A duplicate of this collection is in

SAM. Roux (1986) also studied material from B but he

states that he could not find any material listed by Kunze

in his text and he concludes that the variety is untypified;

3, for var. laxa, Anthony (1984) selects a collection

by Drege from near Bokpoort and Nieuweveld as lecto-

type in B under No. 000413, with a duplicate of this

collection in SAM. Roux notes that the specimen Drege

s.n. (BM) collected near Enon and Bontjes River,

mounted on one sheet with some plants collected by

Cooper can be identified with var. laxa, but they are not

cited by Kunze and cannot be considered as syntypes;

4, var. parviloba is cited by Roux (1986) as synony-

mous with Cheilanthes parviloba Swartz.

The authors of this paper have investigated the dupli-

cates of the lectotypes of var. intermedia and var. laxa in

SAM as well as photostats of a probable syntype of var.

intermedia and of an isosyntype of var. laxa . The lecto-

type of C. hirta var. intermedia is a frond without a

rhizome, with a thick stipe and a contracted, narrowly

linear lamina. Stipe, lamina and vestiture coincide with

what the authors consider to be C. hirta Swartz var. hirta

and they sink var. intermedia under it.

The photostat of a probable syntype of var. intermedia

shows a small whole plant and a large stipe and lamina

on the left and right sides of the sheet respectively, col-

lected by Drege, Ecklon & Zeyher, s. n., s. d. The sheet

is housed at the Museum at Leiden under Morton 750.

The plant on the left is from the Paarl Mountain and the

frond on the right from the Sneeuwbergen. They are

obviously different species, the latter is Cheilanthes con-

tractu (Kunze) Mett. ex Kuhn, the former in habit and

size similar to what the authors describe as C. hirta

Swartz var. brevipilosa W. & N. Jacobsen forma laxa

(Kunze) W. & N. Jacobsen (see below).

The lectotype chosen by Anthony (1984) for C. hirta

var. laxa consists of a rather damaged plant with thin

stipes and rhachises and rather crumpled pinnae as well

as part of a stipe and lamina of similar poor preservation.

When Kunze described the forms of the C. hirta com-

plex he had only material from the southern and eastern

Cape at his disposal. The authors of this paper worked

with much more material, including specimens from the

Transvaal, the Orange Free State, Lesotho, Natal and the

northern Cape Province. On account of shape, character

of stipe and lamina and vestiture, they consider the lecto-

type to represent a form of their var. brevipilosa , so that

Pappe & Rawson (1858: 252) described as Chei-

lanthes glandulosa a plant collected by the missionary R.

Moffat, possibly from a locality near Kuruman. Accord-

ing to their text this appears to be a lax form of C. hirta.

The whereabouts of the type are not known.

Anthony (1984) incorporated C. nielsii Jacobsen

(1983) in her concept of C. hirta as a shade form.

MATERIALS AND METHODS

The authors have studied the large collection in the

National Herbarium at Pretoria as well as their own col-

lections. The following features were used to distinguish

taxa: rhizome and basal stipe scales, lamina configura-

tion, length to width ratio of lamina and largest pinna,

vestiture of stipe, rhachis and lamina, angle setting of

pinnae to rhachis, and sculpturing of the spores by means

of electron microscope scanning. The latter was carried

out by the Botanical Research Institute, Pretoria.

RESULTS

The authors have come to the conclusion that the com-

plex of forms should be subdivided, that the obviously

allied and consistently deviating forms without interme-

diates should be considered separate species and that the

remaining forms of C. hirta sensu lato should be split

into a number of forms or varieties.

As an allied species which might have evolved from a

common ancestor, Cheilanthes parviloba (Swartz)

Swartz was considered a separate species by its author,

who had originally named it Adiantum parvilobum in

1800, but transferred it to Cheilanthes in 1806. Though

later included as a variety of C. hirta by Kunze (1836),

Pappe & Rawson (1858) and others up to Christensen's

Index ftlicum (1906), Sim (1915) accepted it again at

species level and this view is supported by the present

authors. Due to its glabrous and viscid lamina surface it

is immediately distinct from all other more or less hirsute

members of the complex. It is therefore not discussed

any further in this study.

The general essential features of all forms of the com-

plex are set out by Jacobsen (1983: 261-263) and by

Anthony (1984: 68-69). An important diagnostic fea-

ture is the presence of patent or adpressed multicellular,

often gland-tipped hairs on stipe, rhachis and lamina

surfaces. As gradations occur between a number of the

taxa, separation at species level has been possible only in

a few instances. Three of the hirsute members are sepa-

rated at specific level, and four at varietal level. One of

the varieties is split into two formae. The main differ-

ences are shown in the following key.

60

Bothalia 18,1 (1988)

KEY TO THE TAX A

la Fronds 2-pinnatifid to 2-pinnate, sometimes 3-pinnatifid at the extreme base, elliptic, not exceeding 175 mm in

length, frail; rhizome very short, erect, almost like that of an annual herb 9. C. nielsii

lb Fronds 3-4-pinnatifid:

2a Lamina densely covered with translucent, hyaline glands 8. C. hyaloglandulosa

2b Lamina mainly hirsute, hyaline glands occasionally present;

3a Hairs on stipe and rhachis twisted and adpressed; fronds mostly strongly contracted, lamina on long stout

stipes 1. C.contracta

3b Hairs on stipe and rhachis mainly patent:

4a Setting of pinnae to rhachis mainly at 45°-57°, pinnae crowded towards apex, pinnules more or less

inrolled, hairs on upper surface of lamina long, thin, sparse, undersurface with sparse hairs along

costae and costules only 2. C. hirta var. hirta

4b Setting of pinnae to rhachis mainly at more than 50°:

5a Hairs on lamina short, thick, often glandular, varying in length:

6a Hairs very short, glassy, occasionally glandular on upper side of lamina, very sparse, short, gland-

tipped hairs on undersurface; fronds narrowly lanceolate to linear, often contracted, with pinnules

strongly inrolled 3. C. hirta var. brevipilosa

6b More densely hirsute on both surfaces; fronds lanceolate, not contracted, pinnae spreading; plants

small; stipe thin, fragile, reddish to purplish;

7a Hairs on rhachis 1,0— 1,8 mm long, dense, mainly gland-tipped, 4-5-celled

4. C. hirta var. brevipilosa forma laxa

7b Hairs on rhachis mostly thin and fine, clear, hyaline, 0,4- 1 ,4 mm long, mostly 2-3-celled, with

characteristic pyramidal basal cell 5. C. hirta var. brevipilosa forma waterbergensis

5b Hairs on lamina long, thin, flexuous, only occasionally with a small glandular tip:

8a Fronds finely divided, up to 4-pinnatifid, ovate-lanceolate, broadest below the middle of the lamina;

hairs on rhachis 1-2 mm long, fine and twisted, 5-6-celled 6. C. hirta var. nemorosa

8b Fronds lax, 3-pinnatifid, lanceolate, broadest in middle of the lamina; hairs on rhachis 1,5 -2, 8 mm

long, very dense in basal part, 6— 8( 15)-celled 7. C. hirta var. inferacampestris

DESCRIPTION OF TAX A

1. Cheilanthes contracta {Kunze) Mett. ex Kuhn ,

Filices Africanae: 70 (1868); N. C. Anthony 11: 60, t.

12, fig. 14 A'-D' (1984). Type: Cape Province, Zwart-

kops River, Drege (LZ, syn.t; BM lecto.; K, L-BOL

photo.; S, SAM). Figures 2b, 3, 4a— c.

Cheilanthes hirta var. contracta Kunze 10: 539 (1836); Pappe &

Rawson: 35 ( 1858); Sim: 85 ( 1892); Sim: 229, t. Ill, fig. 1(1915),

pro parte. Myriopteris contracta (Kunze) Fee: 149 ( 1852).

Cheilanthes hirta var. contracta sensu Sim in Jacobsen: 262 (1983),

pro parte.

The species (Figures 2b, 3) has recently been

described by Anthony (1984) so that only a few pertinent

morphological features need be stressed here, which dis-

tinguish the species from all other taxa of the complex:

the long creeping rhizome; the rhizome scales which are

darker than those of other members of the complex; the

dark green coloration of the ventral surface of the lamina

(only common with C. hirta var. hirta) and especially

the twisted 3 — 4( — 6)-cellular hairs on stipe, rhachis and

lamina surfaces, adpressed downwards along the rhachis

(Figure 3c). The pinnae are mostly 12-15 mm apart,

9-28 x 2,5—10 mm, with up to 10 pinnules, spreading

at an angle of 38°-52° (average 46°) from the rhachis, bu

curving upwards to run parallel or even towards it and

touching the pinnae above (Figure 2 b). The pinnules

may be very close together or more often well separated

and alternate on the costae, about 4x2 mm where

measurable, but often too contracted, especially when

mature or drying out; the sori are dark brown, about 0,5

mm in diameter. The spores are globular, about 50 gm

in diameter, rugulose with anastomosing rugae and a

prominent trilete ridge (Figure 4a— c).

Vouchers: Acocks 19081 (PRE); Esterhuysen 25808 (PRE); Lieben-

berg 7253 (PRE); Lomw 2 (PRE); Taylor 5915 (PRE).

Distribution

The species is essentially southern, but has recently

been found to occur in the eastern Transvaal, where it

overlaps in its distribution with that of C. hirta var. hirta

and of var. brevipilosa. In the Cape Province it extends

from the Kamiesberg in Namaqualand along the Cape

mountain ranges to the south-western, southern and

south-eastern parts of the province, approximately as far

as the Grahamstown area. Northwards it is found as far

as the Swartberg. The species is quite frequent in the

Cape, but rare in the Transvaal (Figure 5).

Ecology

C. contracta usually grows in shallow, sandy lithosols

amongst rocks or even on sheetrock mats. Quartzite soils

seem to be preferred. Aspect is of no importance, but

plants at the foot of large boulders in half or part shade

tend to be larger. In arid fynbos the stipe may be quite

long, frequently as long as the lamina. In spite of the

creeping rhizome the fronds are mostly only ± 3 mm

apart, the stipes irregularly sprouting, young ones indis-

criminately mixed with mature ones, so that large clus-

ters of fronds develop.

The altitudinal range is large, according to Anthony (/.

c.) between 60 and 1 830 m in the Cape, but apparently

not below 1 000 m in the Transvaal. An herbarium sheet

Bothalia 18,1 (1988)

61

FIGURE 2. — a, Cheilanthes hirta Swartz var. hirta.

Transvaal, 2530 (Lydenburg): on Old Kruger

Road at Waterval Onder, amongst grass and

herbs in sunlight (-CA), 13.1. 1978, Jacobsen

4907, x 0,7; b, Cheilanihes contracta (Kunze)

Mett. ex Kuhn. Cape, 3323 (Willowmore):

west end of Baviaanskloof, (-BD), Jacobsen

5181 , x 0,7.

from near Sabie bears the remark 'xerophytic on quartz-

ite ridge’. This refers probably to the Pretoria Series

quartzite. The other known Transvaal record is from

Kaapsche Hoop, which lies well above 1 000 m. Condi-

tions in the Cape are mostly fairly arid, the annual rain-

fall varying between 200 and 800 mm. The Transvaal

localities, which fall in the summer rainfall area, receive

more than 1 000 mm per year.

FIGURE 3. — Cheilanthes contracta (Kunze) Mett. ex Kuhn. Cape,

3321 (Ladismith): Sevenweekspoort ( — AD), Jacobsen 5177. a,

outline of frond, X 0,6; b, pinnule, x 8; c, portion of rhachis

(characteristic: the basiscopically adpressed long, multicellular

hairs), X 32; d, rhizome scale, x 48; e & f, stipe and rhachis hairs

respectively, x 280.

2. Cheilanthes hirta Swartz var. hirta. Chei-

lanthes hirta Swartz: 128, 329 (1806). Type: Mauritius,

Groendal (S, lecto.-Herb. Jacobsen, photo.!). Figures

2a, 4d-f, 6,7.

Adiantum hirtum (Swartz) Poir. in Lam.: 142 (1810). Notholaena

hirta (Swartz) J. Sm.: 50 (1841). Myriopteris hirta (Swartz) J. Sm.:

174(1866).

Notholaena capensis Sprengel: 32 (1828). Type: Cape Province.

Uitenhage, Zeyher 275 (B, iso.).

Cheilanthes hirta var. intermedia Kunze 10: 539 (1836); Roux: 370

(1986). C. hirta var. intermedia (Kunze) Pappe & Rawson: 35 (1858);

Anthony 11: 69(1984). (No type indicated; see discussion of lectotype

under History and synonymy above).

Cheilanthes hirta var. contracta sensu Sim: 229, t. Ill, figs 1 & 1

beta (1915), pro parte; Tardieu 1: 137, t. 19, figs 1-3 (1958); Jacob-

sen: 262, pro parte, fig. 66 ( 1983).

leones: Tardieu-Blot: t. 19 ( 1958); Jacobsen: 101, fig. 66 ( 1983).

The lectotype is a rather contracted plant, of which

only the upper part is preserved, not dissimilar to certain

specimens from South Africa (Figure la, b). The conclu-

sion that the South African populations agree with the

lectotype is based on investigation of a photo of the type

sheet and on the following observations: what is here

considered to be var. hirta is, apart from var. inferacam-

pestris, the only eastern representative of the complex,

reaching the coast of South Africa in Transkei (Figure

7). Tardieu-Blot (1958) describes under Cheilanthes

hirta Swartz var. contracta Kunze a plant with con-

62

Bothalia 18,1 (1988)

FIGURE 4. — Cwilanthes contracta (Kunze) Mett. ex Kuhn, Jacobsen 5181: a, several spores, X 380; b, single

spore, distal face, x 1160; c, single spore, proximal face, X 1040. Cheilanthes hirta Swartz var. hirla,

Jacobsen 4907: d, single spore, side view, x 930; e, single spore, distal face, X 730; f, single spore, proximal

face, x 730. Cheilanthes hirla Swartz var. brevipilosa W. & N. Jacobsen: g, N. Jacobsen 4522 , single spore,

proximal face, x 1200; h, Jacobsen 5267, single spore, distal face, x 870.

Bothalia 18,1 (1988)

63

FIGURE 6. — Cheilanthes hirtu Swartz var. hirta. Transvaal, 2530

Lydenburg (— CB), Jacobsen 4907. a, outline of frond, x 0,3; b,

portion of rhachis with right pinna, x 2; c, portion of rhachis, x

5; d, scale at base of stipe, x 12; e, hair of stipe, x 35; f, hair of

centre of rhachis, x 35; g, hair of underside of pinnule, x 46; h,

hair of upper surface, x 73.

traded, elongate fronds, with long, white, patent hairs

and with globular trilete spores with convex faces and

cristate-reticulate sporoderm. The senior author saw a

contracted, elongate plant with long white hairs on the

surfaces which he identified as a form of Cheilanthes

hirta on Reunion in 1978. No other forms of the variable

species are recorded from these localities. It is accepted

therefore that the variety described here agrees with the

Mauritian plant.

Rhizome creeping or shortly ascending with narrowly

linear, subulate, flexuous scales, reddish brown with

median dark stripe in the lower part, where they are 3-5

x 0,2-0, 5 mm, uniformly reddish brown higher up,

8-11 x 0,5 mm. Fronds erect, tufted or closely spaced.

FIGURE 7. — Distribution of Cheilanthes hirta var. hirta.

slightly contracted to densely crumpled, pinnae slightly

to tightly inrolled and twisted out of the plane of the

frond. Stipe terete, stout, reddish brown to very dark

brown, dull or shiny, 1—2 mm in diameter, basal scales

6-13 mm long, linear to narrowly lanceolate (Figure

6d), crowded at base, diminishing in frequency upwards

and absent from about 2/3 of the length of the stipe, be-

coming gradually replaced by grey to reddish patent

hairs, 2,0— 2,5 mm long (Figure 6e), protruding from a

coat of de flexed or patent shorter hairs. Lamina very

narrowly lanceolate (Figures 2a, 6a), 90-410 x 5-55

mm, averaging 200 x 24 mm, 2 to 3-pinnate; apex

rounded to acute with pinnatifid terminal segment. Pin-

nae triangular with 6 to 7 pairs of pinnules, lowest of

these often strongly basiscopically developed, pinnate to

bipinnatifid at base, set at an average angle of 40°— 57°

(rarely up to 79°) to the rhachis, and then strongly bent

upwards; 2 to 5 pairs of pinnae reduced towards base,

lowest one 3-8 mm long, 15-28 mm distant from near-

est above; centre pinnae (9—) 12—42 x (3—) 7—12 mm,

averaging 23 x 9 mm, 7-15 mm distant from each

other; top pinnae crowded or up to 8 mm distant from the

terminal segment; costae and larger costules winged.

Pinnules triangular to oblong, lobed, not hairy at the

margin, but with sparse, long, thin, simple or Finely

gland-tipped, 3-5-celled hairs 0,75—1,5 mm long on

upper surface and along the costules on the undersurface

(Figure 6g, h), elsewhere practically glabrous. Rhachis

round, shiny or dull greyish to reddish brown, casta-

neous or dark brown, generally shaggy with patent red-

dish, occasionally gland-tipped 3-7-celled hairs (Figure

60- Sporangia mostly barely visible under the inrolled

lobes of the pinnules. Spores averaging 64 g, m in dia-

meter [Std Dev. (standard deviation) ± 2,71], rounded

though faintly triangular in outline with convex faces,

trilete ridge not or weakly developed, reticulate-cristate,

occasionally rugulose-cristate on a smoothly granulose

base (Figure 4d- f).

Vouchers: Fourie U7>21 (PRE); Jacobsen 4521 (PRE); Lambrechts

45 (PRE); Leendertz 4047 (PRE); Theron 2469 (PRE).

Distribution

Apart from the type locality in Mauritius, the presence

of the variety has been confirmed in Reunion, in Mada-

gascar, Zimbabwe and southern Africa. Here it is known

from the Transvaal Highveld, northern and western

64

Bothalia 18,1 (1988)

Transvaal, Swaziland, the Orange Free State, Lesotho,

Natal, Transkei and the eastern and southern Cape, with

one occurrence as far west as the Somerset West area

(Figure 7). The variety is frequent in the Transvaal,

Natal, Transkei and the eastern Cape, rare in the OFS,

Upper Karoo and southern Cape.

Ecology

The variety grows mostly in shallow sandy to loamy

lithosols in rocky ground, amongst boulders and out-

crops, often at the foot of boulders or cliffs, usually

amongst some herbs and grass, but also in open bush

country in sunlight or preferably in part shade. The alti-

tude ranges from almost sea level in Natal, Transkei and

the Cape to more than 2 000 m (almost 3 000 m in Leso-

tho in a deviating, dwarf, highly contracted form) at 400

to 1 000 mm annual rainfall, however, in the wetter

areas not in forests, but always in open well drained

positions. In the winter rainfall area of the Cape the

plants dry and shrivel during the hot summer months,

while on higher ground in the Free State, Lesotho and

Transvaal they fade during the winter to develop new

shoots at the onset of the summer rains. They may be

evergreen, but are not soriferous during the winter in the

coastal areas of higher rainfall.

Notes

It had initially been thought, that the fronds of C. hirta

var. hirta were always highly contracted, corresponding

to Kunze’s (1836) var. contracta, but detailed investiga-

tions have shown that the essential differences between

the taxa of the complex lie in the type of vestiture of

rhachis and lamina and the spore characters. Contracted

habit as well as more spread out fronds may occur in C.

contracta, C. hirta var. hirta and var. brevipilosa. All

three are well separated by the nature of the rhachis and

surface hairs as well as by differences in the nature and

size of the spores.

3. Cheilanthes hirta Swartz var. brevipilosa W. &

N. Jacobsen, var. nova. Type: Transvaal, Mondeor,

Johannesburg Jacobsen 5254 (PRE, holo.; Herb. Jacob-

sen, iso.) Figures 4g, h; 8 & 9.

A varietatibus et formis ceteris speciei pilis brevissi-

mis glandulosis in superficiebus ambabus laminae dif-

fert.

Rhizome creeping with broadly based lanceolate, acu-

minate, light brown to reddish scales, 5-9 x 0,5- 1,2

mm, basal ones shorter with black median stripe. Fronds

erect, tufted, closely spaced. Stipe terete, dark brown,

dull or shiny, 1, 2-2,0 mm in diameter, stout, basal

scales as those on rhizome, acuminate to linear, 5— 10 x

O, 4-0, 8 mm, becoming less and smaller upwards and

replaced about half way up by patent, twisted or shaggy

light brown or reddish hairs up to 1 mm long protruding

from a shaggy coat of shorter hairs. Lamina linear-lan-

ceolate to lanceolate, often broadest above the middle

(Figure 9a), 3-pinnatifid to 3-pinnate, slightly contracted

to densely crumpled (Figure 8) with the pinnae flat or

inrolled and twisted out of the plane of the frond; apex

rounded or subacute with a tiny pinnatifid terminal seg-

ment 2-5 mm long. Pinnae mostly spreading at 63°-79°

from the rhachis, but often at more acute angles, es-

pecially in contracted forms, otherwise pinnae and pin-

nules as in var. hirta, except for very short, glassy, occa-

FIGURE 8. — Cheilanthes hirta Swartz var. brevipilosa W. & N.

Jacobsen. Transvaal, 2527 (Rustenburg): Rustenburg Nature

Reserve, on rocky ground, in full sunlight, 1 460 m (-CC),

Jacobsen 4522. This is an extremely contracted form as the

variety is usually more open in habit than C. hirta var. hirta, X 1 .

sionally glandular hairs, arranged flabelliform along the

costules of the pinnules on the upper side (Figure 90,

and very sparse, short, gland-tipped hairs on the under-

side which may be almost glabrous. Rhachis densely

covered by short and adpressed, 1 — 3-celled gland-

tipped hairs 0,2-0, 5 mm long, with occasional patent

longer hairs of this type, up to 5-celled and 1 ,8 mm long

(Figure 9d, e). Spores averaging 48 /xm in diameter (Std

Dev. ± 1,34), strictly triangular, trilete ridge prominent,

essentially strongly and coarsely rugulose-cristate,

especially on distal face, but with isolated short rugae

and rare verrucae on proximal face (Figure 4g, h).

Vouchers: Jacobsen 5069 (PRE); Reid 462 (PRE); Repton 5335

(PRE); Thode A7877 (PRE); Westfall 905 (PRE).

Distribution

Known so far from South Africa and SWA/Namibia.

In the former mainly in a strip trending north-south from

the northern, eastern and western Transvaal through the

Orange Free State and western Lesotho to the eastern

Cape, with an isolated outlier in the northern Cape Pro-

vince south of Vryburg and three north-south orientated

occurrences in southern SWA/Namibia (Figure 10). A

find allotted to this variety in the Natal Drakensberg is

doubtful and may be a hybrid of var. hirta and var.

nemorosa (Jacobsen 4628). The variety overlaps in its

distribution to a large extent with that of var. hirta, but

seems to be absent from Natal and the Transkei.

Bothalia 18,1 (1988)

65

FIGURE 9. — Cheilanthes hirta

Swartz var. brevipilosa W. &

N. Jacobsen. Transvaal 2627

(Potchefstroom): Mondeor,

Johannesburg (-BB), Jacobsen

5267: a, outline of frond, X 0,6;

b, portion of rhachis with right

pinna, X 4; c, portion of rha-

chis, x 15; d, short glandular

hair of rhachis, x 123; e, multi-

cellular hair of rhachis, X 123;

f, (same locality, but Jacobsen

5254) terminal segment of pinna

with characteristic very short

glandular hairs in fan-like orien-

tation, x 23.

Ecology

Ecologically its requirements are similar to those of

var. hirta. The altitudinal range also tends to be alike,

probably from about 100-2 000 m. Highly alpine forms

are missing. On the whole the variety prefers more xero-

phytic localities, yet has a broad moisture tolerance

range, and occurs in both summer and winter rainfall

areas with an annual precipitation ranging from as low as

100 mm to about 1 000 mm. It is common in the

Transvaal and frequent in the OFS, but becomes rarer in

the eastern Cape Province except for the Hogsback-Keis-

kamma area.

Notes

The variety is distinguished by its smaller, triangular

spores from var. hirta , which has large, rounded spores.

The original concept of a taxonomic separation on

account of the generally more open and spreading lamina

in var. brevipilosa could not be upheld, as some fairly

contracted forms with triangular spores (N. Jacobsen

4522, Figure 8) were seen. A subsequent study disclosed

the different type of vestiture, which forms a most

important additional criterion for differentiation and

which was used in the key to the taxa of the complex.

4. Cheilanthes hirta Swartz var. brevipilosa W. N.

Jacobsen forma laxa ( Kunze ) W. & N. Jacobsen, stat.

nov. Type: near Bokpoort and Nieuweveld, Drege s. n.

(B, lecto; SAM!; BOL, photo.). Figures 11a, 12 &

13a-c.

Cheilanthes hirta Swartz var. laxa Kunze in Linnaea 10: 540 ( 1836).

?Cheilanthes glandulosa Pappe & Rawson: 35 (1858). Type: ?Cape,

nr Kuruman, Moffat s. n. (not found).

leones: Anthony: t. 14A, B (1984).

Rhizome erect, small, with closely tufted fronds with

scales lanceolate, acute, 4-5 x 0,6— 0,8 mm, lower

ones light brown with dark median stripe, upper ones

light brown to foxy red, broader and larger. Stipe terete,

reddish brown, more rarely dark brown to almost black,

thin, reaching only rarely 1 mm in diameter, basal scales

lanceolate-acuminate, 4 x 0,6 mm, pale brown to

brown, replaced higher up by patent, occasionally gland-

tipped, light brown hairs up to 2 mm long. Lamina lan-

ceolate to oblong (Figures 11a, 12a), broadest in the

middle, 3-pinnatifid to 3-pinnate, 60-250 x 15-50

mm, averaging 140 x 32 mm, about 4/ times longer

than broad, apex rounded, pinnatifid, terminal segment

7-8 mm long. Pinnae triangular, mostly with 3-4 pairs

of pinnules, spreading at 70°— 90° from the rhachis, 0-3

pairs slightly reduced towards base, lowest 8-15 x

6-10 mm, 13-27 mm distant from next above; centre

pinnae 8-32 x 5—14 mm, 7—15 mm apart, apex

rounded, slightly basiscopically developed at the base

(Figure 12b); top pinnae crowded or up to 5 mm distant

from terminal segment. Pinnules oblong, lobed to pinna-

tifid, apex obtuse, terminal segment unequally pinnati-

fid, decurrent, some simple or gland-tipped hairs on the

66

Bothalia 18,1 (1988)

FIGURE 11. — a, Cheilanthes hirta

Swartz var. brevipilosa W. &

N. Jacobsen forma laxa (Kunze)

W. & N. Jacobsen. Cape, 3224

(Graaff-Reinet): Karoo Nature

Reserve, under shrubs in shade

amongst pillars of dolorite

( — BB), Jacobsen 5185, X 0,6;

b, Cheilanthes hirta Swartz var.

brevipilosa W. & N. Jacobsen

forma waterbergensis W. & N.

Jacobsen. Transvaal, 2328 (Bal-

timore): farm Ketting 368 LR,

on south-facing slope, in cre-

vices of sandstone cliff, 1 085

m (-BD), Jacobsen 5121

(holotype), x 0,6.

upper side (Figure 1 20, fairly thick, simple or gland-

tipped 3-5-celled hairs on margins and underside (Fig-

ure 12e, g). Rachis and costae round, dull or shiny, red

to reddish brown with a dense coat of often gland-tipped,

patent, red, (2) 4-5 (lO)-celled hairs, 1-2 mm long

(Figures 12c, h). Sporangia discrete, marginal, under

the lobes of the pinnules. Spores about 46 /u,m in diame-

ter (Std Dev. ± 1,10), triangular in outline with a faintly

prominent trilete ridge, reticulate-cristate, the trilete

ridge formed by reticulating cristae (Figure 13a-c).

Vouchers: Allardice 1504 (PRE); Giffen FH 1838 (PRE); Henrici

3966 (PRE); Jacobsen 5 185 (PRE); Oliver & Mueller 6455 (PRE).

Distribution

The form extends from the south-western Transvaal

into the northern Cape Province and south-eastern SWA /

Namibia and southwards in a broad strip through the

central Karoo, the OFS and part of Lesotho to the south-

eastern Cape. A few finds of the form have been made in

central SWA/Namibia (Figure 14).

FIGURE 12. — Cheilanthes hirta Swartz var. brevipilosa W. & N.

Jacobsen forma laxa (Kunze) W. & N. Jacobsen. Cape, 3224

(Graaff-Reinet): Karoo Nature Reserve ( - BB), Jacobsen 5185: a,

outline of frond, x 0,5; b, portion of rhachis with right pinna, X

3; c, portion of rhachis, x 13; d, basal stipe scale with sclerotic

brown centre, X 38; e, glandular hair of undersurface with russet-

coloured interstices, x 102; f, unicellular clear hair of upper

surface, x 222; g, clear hair of undersurface, x 102; h, rhachis

hair, x 102.

Bothalia 18,1 (1988)

67

FIGURE 13. — Cheilanthes hirta Swartz var. brevipilosa W. & N. Jacobsen forma laxa (Kunze) W. & N. Jacobsen,

Allardice 1504 : a, several spores, X 380; b, single spore, distal face, x 1040; c, single spore, proximal face, x

1040. Cheilanthes hirta Swartz var. brevipilosa W. & N. Jacobsen forma waterbergensis W. & N. Jacobsen,

Jacobsen 5137: d, several spores, x 380; e, single spore, distal face, x 1050; f, single spore, proximal face, x

1050. Cheilanthes hyaloglandulosa W. & N. Jacobsen, N. Jacobsen 5255: g, single spore, proximal face, x 925;

h, single spore, distal face, x 880.

68

Bothalia 18,1 (1988)

FIGURE 14. — Distribution of Cheilanthes hirta var. brevipitosa for-

mae laxa, • ; and waterbergensis, 4 .

Ecology

It is a xerophytic representative of the C. hirta com-

plex, growing mostly on stony slopes or amongst boul-

ders or in rock crevices (very often of dolerite) under

fairly arid conditions, never on north aspects, rarely on

west-facing slopes, but frequently on east and south

aspects. It shelters usually under boulders or overhang-

ing krantzes or, if sufficient scrubby vegetation is avail-

able, as for instance in the Karoo Nature Reserve at

Graaff-Reinet or in the south-eastern Cape, under

shrubs, sometimes belonging to the genus Euphorbia.

The habit of the plants is therefore not xeromorphic, but

mesomorphic, with a small rhizome and frail stipes and

rhachis.

The altitudinal range extends from practically sea

level in the south-eastern Cape to about 2 000 m in west-

ern Lesotho in areas with the rainfall ranging from 200 to

± 1 000 mm annually. Phenologically it follows the

pattern of C. hirta var. hirta and var. brevipilosa. It is

occasional to rare and mostly isolated.

Notes

C. hirta var. brevipilosa forma laxa probably corre-

sponds with C. glandulosa of Pappe & Rawson (1858),

the type of which was collected in Griqualand in 1857 by

Moffat, but is now lost. According to the description it

had purple stipes and rhachises with glandular hairs.

Pappe & Rawson (1858: 36) remark that this species was

‘very like C. hirta var. laxa of Kunze (1836)’, which is

obviously a lax variety of the species. It appears there-

fore that Baker, cited in Anthony: 70 (1984), erred in

comparing a contracted form with the type specimen of

C. glandulosa in Herb. Rawson.

C. hirta var. brevipilosa forma laxa differs from all

other taxa of the complex, except C. hirta var. brevipi-

losa forma waterbergensis and C. nielsii, in the smaller

size and fragile stipe. It differs from C. hirta var. brevi-

pilosa forma waterbergensis in the 1-2 mm long hairs on

the rhachis which are without long basal pyramidal cells,

and from C. nielsii in the tripinnate fronds.

5. Cheilanthes hirta Swartz var. brevipilosa VP. &

N. Jacobsen forma waterbergensis VP. & N. Jacobsen,

forma nova. Type: north-western Transvaal, farm Ret-

ting 368 LR, N. Jacobsen 5121 (PRE, holo.), Figures

lib, 13d— f& 15.

A taxis affinibus praeter C. hirtam var. brevipilosam

forma laxam et C. nielsii, statura parva et stipite gracili

differt. A C. hirta var. brevipilosa forma laxa cellulis

basalibus pyramidalibus pilorum longiorum rhachidis et

a C. nielsii fronde tripinnata differt.

All morphological features coincide with those of C.

hirta var. brevipilosa forma laxa, but pinnae mostly with

5 pairs of pinnules. Pinnules triangular to oblong, deeply

lobed, terminal segment lobed, apex rounded, margins

not hairy (Figure 15b), but hirsute on both surfaces with

short 1— 3-celled, gland-tipped hairs (Figures 15f, g).

Rhachis well covered with mostly thin and fine hairs,

often gland-tipped, mostly 2- or 3-celled, but also with

some longer ones with up to 6 cells, with a characteristic

pyramidal long basal cell (Figures 15d, e). Spores about

51 p.m in diameter (Std. Dev. ± 0,57), triangular with

faintly prominent trilete ridge; spore surface forming a

network of irregular rugulose cristae, sparse on proximal

face and on the trilete ridge.

Vouchers: Burn Davy 1670 (PRE); Galpin M 700 (PRE); Jacob-

sen 5137 (PRE); Rodin 4127 ( PRE).

FIGURE 15. — Cheilanthes hirta Swartz var. brevipilosa W. & N.

Jacobsen forma waterbergensis W. & N. Jacobsen. Transvaal,

2328 (Baltimore): Koedoesrand, farm Wemmersvlei 185 LR

( — AC), Jacobsen 5137 : a, outline of frond, x 0,7; b, portion of

rhachis with left pinna, x 3; c, portion of rhachis, X 27; d,

rhachis hair, x 136; e, stipe hair, x 136; f & g, very short hairs of

the lamina surfaces, x 406 (both d & e show the characteristic

pyramidal basal cell of these hairs).

Distribution

This form occurs to the north of the area of forma laxa

in the western and north-western Transvaal with the

northern limit north of the Waterberg. There are isolated

finds from the Loskop Dam and from near Pretoriuskop

in the Kruger National Park (see Figure 14).

Ecology

Forma waterbergensis is a xerophytic form of the drier

areas of the Transvaal, where the annual rainfall does not

Bothalia 18,1 (1988)

69

exceed 600 mm. Scarcity of material prevents an assess-

ment of the altitudinal range. It may be between 700 and

1 100 m. Similar to forma laxa, the plants prefer south

aspects, growing in shallow lithosols of sandstones, fel-

site or granite amongst and in shelter of boulders or in

rock crevices. The form is rare and has been collected

only a few times.

Notes

The form was separated from the similar and wide-

spread forma laxa on account of the different nature of

the hairs of the rhachis. It differs from C. hirta var.

brevipilosa in the pyramidal basal cells of the rhachis

hairs; in its small size and in the frail stipe and rhachis,

which are very different from the robust stipe of the

variety. The pinnae spread from the rhachis at almost 90°

while those of var. brevipilosa are always ascending.

6. Cheilanthes hirta Swartz var. nemorosa W. &

N. Jacobsen, var. nova. Type: eastern Transvaal, Mount

Sheba Nature Reserve, Jacobsen 4443 (PRE, holo.;

Herb. Jacobsen, iso.) Figures 16, 17 & 18a-c.

A taxis affinibus aliis, praeter aliquas formas mon-

tanas, statura magna, fronde usque ad quadripinnatifida,

ovato-lanceolata, latissime infra medium et sporis dense

subtiliterque reticulato-cristatis differt.

Rhizome robust, erect, with densely tufted fronds and

with linear, subulate, light brown to reddish brown

scales 5— 9 x 0,2— 0,5 mm. Stipe terete, dark reddish

brown to almost black, dull or shiny, stout, up to 2 mm

in diameter, basal scales lanceolate to very narrowly

linear, acuminate to subulate, 5 — 10 x 0,2— 1,2 mm

(Figure 17d), foxy red to brown, higher up replaced by

U

FIGURE 16. — Cheilanthes hirta Swartz var. nemorosa W. & N.

Jacobsen. Transvaal, 2430 (Pilgrim’s Rest): Mount Sheba Nature

Reserve, on way to Gola Gola Waterfalls, in light scrubby forest,

1 660 m (-DB) Jacobsen 4443 (isotype), x 0,4.

FIGURE 17. — Cheilanthes hirta Swartz var. nemorosa W. & N.

Jacobsen. 2730 (Vryheid): Kwamadhlangampisi Mountain, near

Dirkiesdorp, Transvaal ( — AB), Jacobsen 5113: a, outline of

lamina, X 0,2; b, portion of rhachis with right pinna, X 3; c,

portion of rhachis, X 10; d, scale at base of stipe, x 38; e, rhachis

hair, X 85.

patent, occasionally gland-tipped light brown to fox-red

hairs up to 2 mm long. Lamina ovate-lanceolate to nar-

rowly lanceolate, broadest below the middle (Figures 16,

17a), long tapering towards the apex, 3-pinnate to 4-pin-

natifid, finely divided, apex rounded to acute with a

minute terminal segment 3-5 mm long, base tapering

with 1-4 pairs of pinnae reduced. Pinnae triangular to

oblong, spreading at 61 °— 80° from the rhachis, in large

specimens the lowest 20—28 x 5-20 mm, 35-44 mm

distant from next above, largest below middle of frond

40-95 x 14—25 mm, 19—23 mm from each other,

uppermost pinnae 1—3 mm from the terminal segment.

Pinnules in (4)5-9 pairs, oblong, deeply pinnatifid to

pinnate to a winged costule, upper ones, elliptic, slightly

lobed (especially in the high altitude forms), lobes

oblong-cuneate or rounded, apex obtuse (Figure 17b),

terminal segment pinnatifid with apex rounded, margins

and both surfaces with long, 4— 5-celled non-glandular

hairs. Rhachis round, shiny below, matt above, casta-

neous to dark brown with mostly non-glandular, 5 or

6(9)-celled, usually with fine and twisted hairs up to 2

mm long (Figure 17c, e). Spores on average 53 gm in

diameter (Std Dev. ± 2,29), globular to faintly triangu-

lar in outline, trilete ridge not or only very faintly

developed, loosely to densely and finely reticulate-

cristate, rarely cristate-rugulose (Figure 18a-c).

Vouchers: Buitendag 1246 (PRE); Compton 30444 (PRE); Jacobsen

51 13 (PRE); Strey 6167 (PRE), Wager 4595 (PRE).

Distribution

The variety is confined to the high rainfall areas of

South Africa, following essentially the eastern escarp-

ment and the mountains of the southern Cape. The distri-

bution is somewhat patchy with isolated areas along the

Soutpansberg-Woodbush escarpment, a long strip from

The Downs in eastern Transvaal through Swaziland to

the Louwsburg area and some isolated occurrences in the

Witbank-Belfast and Ermelo areas. After a gap in the

70

Bothalia 18,1 (1988)

FIGURE 18. — Cheilanthes hirta Swartz var. nemorosa W. & N. Jacobsen, Jacobsen 4443: a, several spores, X

380; b, single spore, distal face, x 890; c, single spore, proximal face, x 840. Cheilanthes hirta Swartz var.

inferacampestris W. & N. Jacobsen, V. d. Schijff 3990: d, single spore, proximal face, X 1 100; e, single spore,

distal face, x 1 100. Cheilanthes nielsii Jacobsen, Jacobsen 5125: f, several spores, X 380; g, single spore,

proximal face, x 1500; h, single spore, distal face, x 1500.

Bothalia 18,1 (1988)

71

FIGURE 19. — Distribution of Cheilanthes hirta var. nemorosa.

Natal Midlands it is found again all along the OFS/Leso-

tho border, along the Drakensberg and into southern

Natal and Transkei, ending in the Hogsback area of the

eastern Cape. The southern Cape area extends from the

Suurberg to the Grootvadersbos (Figure 19).

Ecology

The variety grows as a hemicryptophyte in shallow

lateritic, humiferous soils of fersiallitic to ferallitic

nature, mostly amongst boulders on mountain slopes or

in gullies and ravines, as long as there is a certain

amount of shade. It does not enter the wet forest of its

environment, but prefers forest edges and scrub forest in

areas of 800 to more than 1 200 mm annual rainfall. It

was thought not to occur at greater altitudes than about

1 800 m, but a dwarf, typical montane form occupies

elevations from 1 800 to about 2 300 m in the Witzie-

shoek-Sentinel area, either on scree slopes in the shelter

of ericoid scrub or sometimes in company with Polysti-

chum alticola Schelpe & N. C. Anthony or chasmophy-

tic on basalt cliffs, again with P. alticola or Cheilanthes

eckloniana (Kunze) Mett. These plants are usually be-

tween 150 and 200 mm, rarely up to 300 mm tall and

about 50 mm broad.

Note

C. hirta var. nemorosa is morphologically an easily

recognized type and its spores are rather different from

the other members of the C. hirta complex. This might

have justified specific rank, but some intermediates be-

tween it and var. brevipilosa or var. inferacampestris

respectively were seen, so that varietal status is prefer-

able.

7. Cheilanthes hirta Swartz var. inferacampestris

VF. & N. Jacobsen, var. nov. Type: Transvaal, Kruger

National Park, V. d. Schijff 3990 (PRE, holo.). Figures

18d, e; 20 & 21.

A varietate typica pinnis patentibus laxis differt; a

taxis ceteris complexi pilis longioribus laminae differt.

Rhizome black, robust, erect, with closely tufted

fronds; scales practically absent except for a few very

small, almost black scales low on the rhizome. Stipe

terete, 1 mm in diameter, dark brown, dull, firm; basal

scales 5 x 0,6 mm, lanceolate, light brown, replaced

upwards by patent light brown hairs up to 2 mm long

(Figure 21d). Lamina slender, not contracted, delicately

3-pinnatifid to 3-pinnate, lanceolate to narrowly oblong,

broadest in the middle (Figures 20, 21a), 180-360 x

30-70 mm, apex acute with a small terminal segment,

base with only a few pairs of pinnae slightly reduced.

Pinnae spreading at 60°-78° from the rhachis, lowest 28

mm long, 35 mm distant from next above, largest

coarsely serrate to bi-pinnatifid, 30-33 x 20-23 mm,

18 mm apart, triangular, subacute to obtuse (Figure

21b), upper ones crowded, about 4 mm distant from the

terminal segment, which is pinnatifid or lobed. Pinnules

elliptic in outline, apex lobed, margins and surfaces with

scattered, straight, long, 4 or 5-celled hairs (Figure 2 le),

similar to those on the rhachis. Rhachis cylindrical, cas-

taneous or greenish brown in the upper part, hairs dense

(Figure 21c), patent, (1-) 1,5 — 2, 8 mm long, simple or

only occasionally glandular, 6-8 ( — 1 5)-celled. Spores

41 pm in diameter (Std Dev. ± 0,97), globular to

rounded triangular, trilete ridge weakly developed, con-

sisting of anastomosing cristae; distal face loosely, but

strongly cristate, proximal face sparsely so, often only

with some isolated rugose cristae on a rugulose base, so

that the spores appear echinate with blunt projections in

profile (Figure 18d, e).

Vouchers: Buitendag 1060 (PRE); Codd & De Winter 4895 (PRE);

Van der Schijff 3745 (PRE); Vassilatos & Mantell 805 (PRE); Wager

1622c ( PRE).

FIGURE 20. — Cheilanthes hirta Swartz var. inferacampestris W. &

N. Jacobsen. Transvaal, 2531 (Komatipoort): Lebombo Moun-

tains, Crocodile Bridge section of the Kruger National Park

(-BD), H. P. v. d. Schijff 3990 (holotype), x 0,25. The tall

growth and very lax habit are characteristic.

72

Bothalia 18,1 (1988)

FIGURE 21. — Cheilanthes hirta Swartz var. inferacampestris W. &

N. Jacobsen. Kruger National Park, H. P. v. d. Schijff7743: a,

outline of lamina, x 0,25; b, portion of rhachis with right pinna,

x 2; c, portion of rhachis, x 13; d, stipe hair, x 70; e, lamina

hair, x 70; f, glandular rhachis hair, x 70.

Distribution

Var. inferacampestris is found from the eastern

Transvaal (southern Kruger National Park) through

eastern Swaziland, Zululand and Natal, to the Port

Shepstone area. One isolated specimen was seen from

the Devuli Ranch in the Bikita District of Zimbabwe

(Figure 22).

Ecology

The variety, although confined to low-lying hot areas

is by no means typical of xerophytic habitats, but is

usually found in sheltered shady localities, on south-

facing rocky slopes with humiferous loamy soil. It may

be found in scrub forest or in kloofs and ravines, in the

shade of large boulders or amongst undergrowth. The

annual rainfall varies between 600 mm (where the

variety grows only in moist localities) and ± 1 000 mm.

The altitudinal range is between 50 and 600 m.

FIGURE 22. — Distribution of Cheilanthes hirta var. inferacampestris,

• ; C. nielsii , ■ ; and C. hyaloglandulosa, A .

8. Cheilanthes hyaloglandulosa W. & N. Jacob-

sen in South African Journal of Botany 51: 145—148

(1985). Type: Transvaal, farm Wanhoop 78 JT, N.

Jacobsen 5255 (PRE, holo.). Figures 13g, h; 23.

The species has been described recently, but it was felt

that some supporting drawings and spore micrographs

might be informative. As the spores have not yet been

described a few words are necessary. Spores average 60

/xm in diameter, are roughly triangular with prominent

cristate trilete ridge, the surface is loosely cristate with

broad anastomosing cristae, up to 12 gm high on a

smooth base.

Vouchers: Jacobsen 5286 (PRE); Mogg 4894 (PRE).

Distribution and ecology

The species is known only from a few localities above

1 800 m altitude in the eastern Transvaal escarpment

(Figure 22), where they grow in shallow lithosols over

quartzite, along rocky shelves and amongst boulders on

east and north-facing slopes in full sunlight. The fronds

wither away during the cold season.

9. Cheilanthes nielsii W. B. G. Jacobsen , The

ferns and fern allies of southern Africa: 252 (1983).

Type: north-western Transvaal, Potgietersrus District,

farm Goedgelegen 194 LR, Koperkop, Jacobsen 5126

(PRE, holo.). Figures 1 8f — h; 24.

Cheilanthes hirta sensu N. C. Anthony: t. 13F (1984).

leones: Jacobsen: fig. 182a, b (1983).

This species was fully described by Jacobsen (1983)

and merely a few line drawings of the species as well as

micrographs of the spores are presented here. As the

latter have not yet been described this is done in this

paper.

Transvaal, 2530 (Lydenburg): Farm Wanhoop 78 JT, Dullstroom

District (-AA), Jacobsen 5255: a, outline of frond, x 0,4; b,

portion of rhachis with right pinna, X 4; c, portion of rhachis, X

10; d, basal rhachis scale, x 13; e & f, rhachis hairs, x 216; g,

stalked gland from the upper surface of the lamina, X 216.

Bothalia 18,1 (1988)

73

FIGURE 24. — Cheilanthes nielsii W.B.G. Jacobsen. Transvaal, 2328

(Baltimore): farm Goedgelegen 194 LR, Koperkop (-AD), Ja-

cobsen 5126 : a, outline of frond, x 0,9; b, portion of rhachis and

left pinna, x 5; c, portion of rhachis, X 14; d, hair at base of

stipe, X 1 14; e, hair at edge of pinnule, X 207.

Spores average 36 /u-m in diameter (Std Dev. ± 0,52),

are globular, the trilete ridge is not visible due to densely

anastomosing, well developed cristae about 3 /urn high

(Figure 1 8f — h).

Vouchers: Jacobsen 5125 (K); 5207 (PRE); 5251 (PRE); Rang s.n.

(PRE).

Distribution

The species is so far known with certainty from the

northern and north-western Transvaal (because of poor

preservation the few records from near Pretoria are

dubious) and from a few isolated occurrences in the

northern Cape and from one place in Botswana. A record

from SWA/Namibia is doubtful, again due to poor

preservation of the specimen (see Figure 22).

Ecology

The species inhabits very dry situations, growing on

rocky hillsides or hilltops mostly in full sunlight amongst

rock outcrops, on ledges and at boulder bases on north

and west aspects, very much rarer on SSW-facing slopes

and then in partial shade. Rainfall requirements are in the

400-600 mm range annually and the altitude lies

between 700 and 1 100 m.

Notes

C. nielsii is included in this study merely because its

indumentum has a certain similarity to that of the mem-

bers of the C. hirta complex. Anthony (1984) declares

that it is indeed a shade form of C. hirta. In view of the

data given above, this view cannot be upheld. Anthony

also states that the spores are indistinguishable from

those of C. hirta. While it is true that some members of

the C. hirta complex have similarly strongly cristate

spores, as for instance C. hirta var. hirta , others within

the complex have deviating spores. The spores of C.

nielsii are in fact not like any of the other members of the

complex as they are very small, having an average size

of 36 /xm with very little variation. The smallest spores

found within the complex are those of C. hirta var. infe-

racampestris with an average of 41 pun, while those of

var. hirta average 64 p.m. Morphologically C. nielsii is

entirely different from var. inferacampestris, a form

always growing in shade under fairly dry climatic con-

ditions. As pointed out by Jacobsen (1983) the plants

have morphologically a greater affinity to C. inaequalis

(Kunze) Mett. var. buchananii (Bak.) Schelpe and to C.

leachii (Schelpe) Schelpe as well as to C. capensis

(Thunb.) Swartz than to C. hirta. Although the possibil-

ity of a derivation from the latter cannot be ruled out, it is

felt that the specific status should be maintained because

of the consistent morphological differences and lack of

intermediates.

DISCUSSION

Spores and possible evolution

The separation of a complex of species on the basis of

spores alone is debatable and the authors quote C. Wood

(1973), that ‘great caution must obviously be exercised

when we assume that a character such as spore mor-

phology is of prime importance in either taxonomy or

phylogeny, when so little is known concerning the

methods by which such sculpturing is produced’ (this

refers to the sculpturing of the sporoderm). He found that

there was a great variation in the spores of the Thelyp-

teridaceae. Bearing this in mind, as well as the tendency

of xerophytic ferns to be apogamous and to produce

hybrids, the authors have put emphasis on morphological

characters, which are of immediate use to the botanist in

the field, and used the nature of the spores only in a

supporting function. Study of the spores has, however,

revealed the existence of a distinct and separate taxon

within the group of rather contracted members of the

complex. Later morphological studies confirmed that the

taxon differed also substantially in other respects. This

variety was named Cheilanthes hirta var. brevipilosa.

The size of the spores of members of the complex

varies greatly. The type variety, var. hirta, shows an

average of 64 pun, whereas spores of other members are

as small as 41 pun (var. inferacampestris) and those of

C. nielsii are consistently around the 36 pim mark. This

agrees with R. M. & A. F. Tryon’s (1973) findings for

the entire cheilanthoid group. The large size of the

spores of C. hirta var. hirta seems to point to an ancient

type and its occurrence on Madagascar, Reunion and

Mauritius may indicate that the variety existed already

during the Miocene, if not the Oligocene periods. The

rather large standard deviation (Std Dev.) of ± 2,71

confirms the noticeable irregularity in size and ornamen-

tation, which would indicate apogamy and tendency to

form hybrids.

Var. brevipilosa likewise seems to have separated

from the ancestral form at a fairly early age as it is pro-

bably diploid (Prof. T. Reichstein, pers. comm.), but

apparently originated later than the Miocene period. The

spores are of average size (48 ju,m) with relatively little

variation (Std Dev. ± 1,34). Var. nemorosa with its

relatively large spores (average 53 p. m) and greater

variability — the standard deviation here is ± 2,29 — is

74 Bothalia 18,1 (1988)

TABLE 1. — Measurements of laminae and pinnae (in mm) of the Cheilanthes hirta complex and related species. Taxa are arranged

from mesophytic (top) to xerophytic

Lamina

c

S

si

_i

o

cS

*

s

o

CT)

OS

£

_o

ca

i

<D

Largest pinna

probably also an older segregate. The sporoderm is

generally consistently finely reticulate. With its lax

growth and its spreading, finely divided pinnae, var.

nemorosa is easily distinguished from all other members

of the complex, even when unusually small, as in the

montane forms.

The spores of var. inferacampestris are much smaller,

averaging 41 pxn. The variety has probably evolved

from var. nemorosa in more recent times. This assump-

tion is merely based on morphological similarity, as the

sporoderm of the varieties is entirely different, being

loosely and strongly cristate in inferacampestris and

finely reticulate to reticulate-rugulose in var. nemorosa.

The spores of the two formae of var. brevipilosa with

their distinctly triangular shape, prominent trilete ridge

and rugulose-cristate sporoderm, are similar to the typi-

cal forma, which was the main reason for considering

them as derivatives of it — even though their small and

delicate, widely spreading fronds have little morpho-

logical affinity with it. The spores vary very little, hav-

ing a standard deviation of only ± 1,10 for forma laxa

and ±0,57 for forma waterbergensis.

C. contracta has entirely different spores and is there-

fore thought to have originated from a non-related tribe,

which developed in the southern Cape, and in a later

period extended its range to the Transvaal. Spore varia-

tion is minimal. C. hyaloglandulosa has large, strongly

cristate spores (± 60 pm), similar to var. hirta and has

probably evolved from it, but is considered to be a sepa-

rate species because of its dense cover of unicellular

glands on several-celled stalks. The tiny spores of C.

nielsii are indeed superficially similar in their cristate,

round sporoderm to those of C. hirta var. hirta, but they

are almost only half as large. This fact and the consistent

differences in habit exclude the plants from the C. hirta

group. C. nielsii is therefore considered a distinct spec-

ies, even though it may have derived from the C. hirta

group. Cristate sporoderms are, according to Tryon &

Tryon (1973), predominant in the cheilanthoid ferns,

even if this is .less pronounced amongst the South Afri-

can species. Yet the spores of C. marlothii (Hieron.)

Schelpe ( =Notholaena marlothii Hiem), a plant of very

different habit, are very similar to those of certain types

of the C. hirta complex [compare Anthony (1984), Plate

2, C-D with Plate 13, C— E].

Morphology, phytogeography and climate

The length and width of the lamina and of the largest

pinnae were measured as well as the angle of the pinnae

to the rhachis. These parameters show some correlation

with the climatic conditions under which the plants

grow. The results are summarized on Table 1 and the

Bothalia 18,1 (1988)

75

1 S3 4 B B 7 a 9 ID 11 18 13 W 18 16 T7 18 IS BO

FIGURE 25. — Fiistograms of the ratio length to width of lamina of the

members of the Cheilanthes hirta complex and some allied spe-

cies. A, C. nielsiir B, C. hirta var. brevipilosa formae laxa and

waterbergensis ; C, C. contractor D, C. hirta var. brevipilosar E,

C. hirta var. hirtar F, C. hyaloglandulosar G, C. hirta var. infera-

campestrisr H, C. hirta var. nemorosa.

ratios of length to width of lamina and largest pinnae are

depicted on the frequency histograms in Figures 25, 26

& 27.

It becomes evident that most of the members of the

complex are typically associated with a certain type of

climate. This is discussed in the sections on the ecology

of the various taxa. The succession from east to west,

i.e. from mesophytic to meso-xerophytic to xerophytic is

reflected in the following sequence of the taxa:

C. hirta var. nemorosa-C . hirta var. inferacampes-

tris—C. hyaloglandulosa-C . hirta var. hirta-C. con-

tracta—C. hirta var. brevipilosa-C . hirta var. brevipi-

losa formae laxa/waterbergensis-C . nielsii.

The morphological changes parallel to this succession

are: the lamina becomes narrower from the mesophytic

environment towards the meso-xerophytic environment

but broadens again as the xerophytic end is approached.

The ratio maxima of length to width of the lamina moves

from 2-3 times as long as wide ( C . hirta var. nemorosa)

to 20 times (C. hirta var. hirta) and decreases to 2-3

times (C. nielsii). The largest pinnae become shorter

with the change to xerophytic conditions, being 2-6

FIGURE 26. — Ftistogram of the ratio length to width of

largest pinnae of the members of the Cheilanthes

hirta complex and some allied species. A, C. niel-

siir B, C. hirta var. brevipilosa formae laxa and

waterbergensis ; C, C. contractor D, C. hirta var.

brevipilosa ; E, C. hirta var. hirtar F, C. hyaloglan-

dulosar G, C. hirta var. inferacampestrisr Ft, C.

hirta var. nemorosa.

times as long as wide in C. hirta var. nemorosa and only

1-2 times as long as wide in C. nielsii.

The angle of the pinnae to the rhachis becomes steeper

towards the meso-xerophytic zone, but flattens out again

towards the xerophytic end, averaging about 73° in C.

hirta var. nemorosa, decreasing to between 46° and 55° in

76

250~

Bothalia 18,1 (1988)

200.

150.

100.

SO.

O -

_ 0

E

E

73o 25o i5o i5o 5S0 600

"7J0

FIGURE 27. — Diagram of plotted

measurements of lamina para-

meters of the members of the

Cheilanthes hirta complex and

some allied species: top, lamina;

bottom, largest pinnae. — O — ,

C. nielsii ; — ■ — , C. hirta var.

brevipilosa formae laxa and

waterbergensis', — • — , C. hirta

var. brevipilosa ; , C.

hirta var. hirta: — , C. hyalo-

glandulosa: — — , C. hirta

var. inferacampestris: — A—,

C. hirta var. nemorosa.

C. contracta, C. hirta var. hirta and some forms of C.

hirta var. brevipilosa , but reaching 74° in the xerophytic

forms of the latter and 76° in C. nielsii.

The size and division of fronds decreases generally

from wet to dry conditions.

The relatively tall mesomorphic habit with finely

divided fronds which allows for maximum light expo-

sure in C. hirta var. inferacampestris and C. hirta var.

nemorosa gives way to the elongate meso-xeromorphic

forms of the C. hirta var. hirtalC. hirta var. brevi-

pilosa/C. contracta group with upwards directed, often

contracted pinnae suited to cope with water loss.

Towards the xerophytic end, however, no further con-

traction takes place and the formae laxa and waterber-

gensis escape desiccation by growing only in the shade

of rocks, herbs or scrub on south-facing slopes, or

wherever they are sufficiently protected from desicca-

tion. They are small and delicate and not at all xeromor-

phic. C. nielsii, the most brittle of all, grows on fully

exposed sites, in sunlight and frequently under very hot

conditions. It is, however, entirely seasonal, being green

only when the rainy season has fully set in and shrivel-

ling rapidly when it gets too dry. In this respect and with

its small rhizome and generally small size it resembles

the small therophytes of arid areas. Its deviating position

from the other members of the complex is well demon-

strated in Figure 27, where its lamina ratios plot outside

the general trend and where the ratios of the largest pin-

nae, although beginning within the trend, leave it at a

steeper angle.

ACKNOWLEDGEMENTS

We acknowledge with thanks the assistance of Miss J.

Matthews in preparing the photographs of the illustra-

tions. Mrs S. M. Perold and the Director of the Botanical

Research Institute are gratefully thanked for the SEM

photos of the spores as well as permitting the authors to

examine the specimens housed there. N. H. G. Jacobsen

thanks the Director of Nature Conservation for permis-

sion to publish this article.

REFERENCES

ANTHONY, N. C. 1984. A revision of the southern African species of

Cheilanthes Swartz and Pellaea Link (Pteridaceae). Contributions

from the Bolus Herbarium 1 1 .

CHRISTENSEN, C. 1906. Index ftlicum. Hafniae.

FEE, A. L. P. 1852. Genera ftlicum. Polypodiacees. In Memoires sur

les families des fougeres 5. Strasbourg.

JACOBSEN, W. B. G. 1983. The ferns and fern allies of southern

Africa. Butterworths, Durban, Pretoria.

JACOBSEN, W. & N. 1985. A new species of Cheilanthes (Pterido-

phyta, Adiantaceae) from the eastern Transvaal. South African

Journal of Botany 51: 145-148.

KUHN, M. 1868. F dices africanae . Engelmann, Lipsiae.

KUNZE, G. 1836. Plantarum acotyledonearum Africae Australioris

recensio nova, 1. Filices. Linnaea 10: 481-750.

LAMARCK, J. B. A. P. M. DE 1810. Encyclopedie methodique Bota-

nique, Supplement 1. Paris.

PAPPE, K. W. L. & RAWSON, W. 1858. Synopsis ftlicum Africae

Australis: 35. Cape Town.

ROUX, J. P. 1986. A review and typification of some of Kunze’s

newly described South African Pteridophyta published in his Aco-

tyledonearum Africae Australioris recensio nova. Journal of the

Linnean Society 92: 343-38 1 .

SCHELPE, E. A. C. L. E. 1969. Review of tropical African Pterido-

phyta. Contributions from the Bolus Herbarium 1 .

SIM,T. R. 1892. The ferns of South Africa. Juta, Cape Town.

SIM, T. R. 1915. The ferns of South Africa , edn 2, University Press,

Cambridge.

SMITH, J. 1842. An arrangement and definition of the genera of ferns,

with observations on the affinities of each genus. In W. J.

Hooker, The Journal of Botany 4. Longman, London.

SMITH, J. 1866 Ferns. British and foreign. Hardwicke, London.

SPRENGEL, C. 1828. Linnaei systema vegetabilium , edn 16. Got-

tingen.

SWARTZ, O. 1801. Adiantum caffrorum. Journal fiir die Botanik 2:

85. Schrader, Gottingen.

SWARTZ, O. 1806. Synopsis ftlicum. Kiel.

TARDIEU-BLOT, M. L. 1958. Polypodiacees 1. In H. Humbert,

Flore de Madagascar et des Comores. Firmin Didot, Paris.

TRYON, R. M. & TRYON, A. F. 1973. Geography, spores and

evolutionary relationships in the Cheilanthoid ferns. In Jermy et

al. , The phytogeny and classification of the ferns. Botanical Jour-

nal of the Linnean Society 67, Supplement 1: 145-153.

Bothalia 18,1 (1988)

77

WOOD, C. 1973. Spore variation in the Thelypteridaceae. In Jermy el

al. , The phytogeny and classification of the ferns. Botanical Jour-

nal of the Linnean Society 67, Supplement 1: 191-202.

SPECIMENS EXAMINED

All specimens listed are in the National Herbarium, Pretoria, unless

stated otherwise. Those marked (Jac) are in the senior author's private

herbarium.

Acocks 270 (4); 9160, 9825, 10810 (2); 19081 (1). Allardice 1504 (4).

Archibald 5904, 5926 (5).

Balkwill & Cron 239 (2). Bamps 7017 (2). Barnard 2367 (2). Baur

136 (6). Bosman 2877 (3); s.n. (6). Bottomley 2779, 2878 (2). Bottom-

ley, Watson & Mogg2072 (3). Bradfteld 132, 132a (3). Brandmueller

1987 (3); s.n. (2). Bredenkamp 449 (3). Bremekamp & Schweickerdt

386 (6). Brown (6 Shapiro 235 (2). Brueckner 385 (4). Buitendag 1060

(7); 1246 (6). Burrows 2121 (4). Burn Davy 1670 (5).

Codd & De Winter 4895 (7). Coetzee 978 (3). Comins 810 (2). Comp-

ton 30069 (7); 30444 (6). Culverwell 0050, 0664 (7).

Dahlstrand 2584 (4). Daly 54 (2); 725 (1). De Beer 177 (2). De Jonc-

heere 292 (2); 294 (6); 303 (7). Dieterlen 264a (2); 264b (6); CH 6721

(3). Dyer 1698 (2).

Elliott 1506c (3). Emdon 85 (1). Esterhuysen 2053, 13157 (4); 15517

(6); 17082, 25808, 25917 (1); 26070 (6).

Fisher & Schweickerdt 540 (7). Flanagan 192c (2). Fourie 117121 , 791

(2); 903 (6).

Galpin M 700 (5). Gibbs Russell 3042 (2); 3127 BA, 4570 ( 3). Giffen

FH 1838 (4); FH 1854, s. n. (3). Grant 3180 (2). Griffin s. n. (4).

Guillarmod 176 (2); 808, 830 (3).

Hafstroem 858 (4); 921 (9); 1187 (4); 5117 (6). Hafstroem & Acocks

1950 (6). Hanekom 1432 (1). Hardcastle 2 (6); 274 ( 2). Henrici 1733

(2), 3966 (4). Hilliard & Burn 1713 (6). Holub s. n. (2). Huntley 1915

(2).

Jacobsen 5165, 5177, 5181, (1) Jac; 4269, 4521, 4525, 4907, 5062,

5318, (2) Jac; 4522, 4628, 5069, 5262, 5267, 5502 (3) Jac; 5254 (3)

PRE, Jac; 5185 (4); 5121, 5137 (5); 2309, 4337, 4369, 5113, 5638,

5657, 5672 (6) Jac; 4443 (6); 5255 (8) FR, Jac, PRE; 5286 (8) PRE,

Reichstein; 5125 (9) K; 5126, 5199, 5207, 5251 (9). Jacobsz 413 (3).

Jenkins 910c, 914c (2). Johnson 643 (5). Jooste 193 (4).

Kluge 2179 (6). Kotsokoane 248 ( 6). Krijt2311 (3).

Lambrechts 45 (2). Lang 5112 (6). Leach & Bayliss 12671 (2). Leen-

dertz 856c, 980e, 1076, 2959 (2); 800 (3). Leeuwenberg 10912 (3).

Liebenberg 7263 (1). Lindstedt 63 ( 4). Louw2 (3).

Marloth 761 (2); 1077 (4); 4734, 6915 (1). Marsh s. n. (1). Mauve,

Reid & Wikner 112 (1). McLoughlin 732 (6). Meyer in Herb. Giess

10775 (4). Mitchell s. n. (1). Mogg 600 (3); 1611 (2); 2282 (4); 2492

(2) ; 4210 (6); 4640 (3); 4894 (8); 7120 (2); 8882 (3); 11776, 15371,

18563 (2); 20729 (3). Morris 763 (7). Moss & Rogers 1169 (3). Mott

450 (2). Mueller 967 (4); 1559 (3). Murray 525 (3).

Nelson 464a (6).

Oliver <6 Mueller 6455 (4).

Paterson 794, 1734c (2). Pegler 269 (6). Peplon 6007 (7). Pocock S

181 (1). Potgieter 21830 (3). Rang s. n. (9). Rauh & Schlieben 9762

(3) . Reid 124, 170 (2); 462, s. n. (3). Repton 5335 (3). Roberts 3228

(ft). Rodin 1052 (2); 4127 (5). Rogers 990c (2); s. n. (3). Rogers &

Moss 3583 (3). Rose & Innes 148 (2).

Scharf 1164 (1). Scheepers 434 (6), 1080 (2), s. n. (3). Schelpe 5912

(4) , 5919 (1). Schlechter 16c (1), 995c (2). Schlieben 11045 (4).

Schmitz 400 (6); 4204, 6471 (3); 6506 (6). Schweickerdt s. n. (2). Sim

193c, 194c (2); 345 s. n. (3); 1244c, 3678 (2). Smith 4408 (2). Smook

942 (2); 2019 (3). Smook & Harding 703 (4). Snijman 9 (3); 334 (1).

Siam 83, 177 (3); 178 (4). Story 3390 (2). Strey 4254 (2); 6167, 7918

(6).

Taylor 5653 (3); 5915 (1); 7995 (6); 8009 (2). Theiler s. n. (2). Theron

856 (5); 2469 (2); 6475 (4). Thode A1877 (3). Tyson 136 (4).

Van der Schijff53 (5); 604 (7); 3745, 3990 (7); 5338 (2); 5560 (6). Van

der Schijff & Schweickerdt 5773 (1). Van Hoepen 1675 (2). Van Wyk

s. n. (2). Vassilatos & Mantell 805 (7). Venter 1118 (2).

Wager 33 (3); 198 (6); 1480 (1); 1622 (7); 4595, s. n. (6). Wasserfall

12 (2). Watson s. n. (3). Werdermann & Oberdieck 1328 (9). Werger

141 (3). Wesf j. n. (6). Williams 793c ( 6). Wilman 1507c (4).

Zeyde 159 (2).

'

Bothalia 18,1:79-88(1988)

Solarium (Solanaceae) in Ghana

Z. R. BUKENYA* andJ. B. HALL**

Keywords: food crops, Ghana, indigenous taxa, key, medicinal plants, ornamental plants, Solanum, Solanaceae

ABSTRACT

Of the 22 species, subspecies and varieties in the genus Solanum L. (Solanaceae) that occur in Ghana, about 15 are

indigenous. In Ghana serveral members of the genus are utilized as food crops while others are put to medicinal and

ornamental use. Up-to-date and detailed descriptions for all the Solanum taxa occurring in Ghana and a key to the species

are provided.

UITTREKSEL

Van die 22 spesies, subspesies en varieteite in die genus Solanum L. (Solanaceae) wat in Ghana voorkom, is ongeveer

15 inheems. In Ghana word verskeie lede van die genus as voedselgewasse benut terwyl ander as medisinale en sierplante

aangewend word. Die jongste en uitvoerige beskrywings van al die Solanum- taksons wat in Ghana voorkom en ’n sleutel tot

die spesies word verskaf.

CONTENTS

Introduction 78

Materials and methods 80

Descriptions of Solanum species 81

Solanum L 81

Key to species 81

Subgenus Solanum 82

Section Solanum 82

1.5. nigrum L 82

la. subsp. nigrum 82

2. S. americanum Mill 82

3. S. scabrum Mill 82

Section Afrosolanum Bitter 82

4a. S. terminate Forssk. subsp. welwitschii

(C.H.Wr.) Heine 82

4b. S. terminate Forssk. subsp. inconstans

(C.H.Wr.) Heine 83

Subgenus Brevantherum (Seithe) D’ Arcy 83

Section Brevantherum D’Arcy 83

5. S. erianthum D. Don 83

Subgenus Leptostemonum (Dun.) Bitter 83

Section Acanthophora Dun 83

6. S. mammosum L 83

7. S. aculeatissimum Jacq 83

8. S. capsicoides All 84

Section Ischyracanthum Bitter 84

9. S. arundo Mattei 84

Section Melongena 84

10. S. melongena L 84

11.5. incanum L 84

12. 5. macrocarpon L 84

13.5. wrightii Benth 85

Section Oliganthes Bitter 85

14. S. anguivi Lam 85

15. 5. aethiopicumL 86

16. S. gilo Raddi 86

Section Torva Nees 86

* Senior Lecturer, Botany Department, Makerere University, P.O.

Box 7062, Kampala, Uganda.

** Formerly: Senior Lecturer, Botany Department, University of

Ghana. (Deceased).

MS. received: 1987.02.02.

17. S. torvum Swartz 86

1 8 . S. anomalum Thonn 86

Subgenus Potatoe (G. Don) D’Arcy 87

Section Potatoe 87

19. S. tuberosum L 87

Section Jasmino solanum Bitter ex Seithe 87

20. S. seaforthianum Andr. var. disjunctum

Schulz 87

Discussion and conclusions 87

Acknowledgements 88

References 88

INTRODUCTION

The genus Solanum L. contains about 2 000 species of

which about 15 are indigenous to Ghana. The rest have

been introduced and are now cultivated crops, viz. 5.

scabrum Mill., S. melongena L. and S. tuberosum L.

Others are introduced ornamentals, including 5. mam-

mosum L., S. capsicoides All. and S. wrightii Benth.,

whereas 5. americanum Mill, and S. arundo Mattei are

weeds of recent introduction and a species introduced

into Legon Botanical Garden, respectively.

The genus is of great economic importance, with seve-

ral species that are food crops, yielding edible leaves,

fruits and tubers. Other species are medicinal plants,

ornamentals or weeds of cultivation. The genus is widely

distributed throughout the world.

Since the genus Solanum was described by Linnaeus

in 1753, it has been reclassified innumerable times, and

a multitude of species, subspecies and varieties has been

named. The major revision of Solanum in Africa was

done by Bitter (1913, 1917, 1921, 1923), utilizing

mainly collections from German expeditions. He erected

a partial classification of Solanum. Despite some limita-

tions, especially the validity of some of his varieties

which are based on minor variations and are of very local

occurrence. Bitter’s work is the most authoritative avail-

able today on African Solanum.

D’Arcy (1972) provided a modem classification of the

genus Solanum into subgenera, sections and series and

his classification is widely followed today.

80

Bothalia 18,1 (1988)

Although the above major works and others attempted

to streamline the taxonomy of Solatium, the genus is a

difficult one and is complicated by several factors. These

factors include the difficulty of associating the names of

Solatium used by earlier taxonomists with the plants of

today due to early descriptions being brief, often vague

and frequently lacking in characters now considered to

be diagnostic. Again some of the early names, for exam-

ple many of the names of Linnaeus and those before him,

were not typified, or if so, the type is not easy to discern

(Hepper 1979).

Another problem is the occurrence of polyploid series

within taxa (Edmonds 1977), for example tetraploids and

hexaploids within the S. nigrum complex. These may

provide a barrier to hybridization between morphologi-

cally similar plants leading to cytoraces difficult to dif-

ferentiate using classical methods.

There is also considerable phenotypic plasticity within

species and hybridization between closely related spe-

cies. Hybridization followed by inbreeding may result in

formation of new populations different from either pa-

rent. This is particularly true for the cultivated species. A

large number of ‘microspecies’ or ’semispecies’ (Grant

1971 ) occur in section Solatium and it is problematical to

decide how many of these deserve taxonomic recogni-

tion.

Today there is no single up-to-date work covering the

classification of the entire genus in West Africa. Gbile

(1979) made descriptions of Solatium species in Nigeria.

His work was however based only on the herbarium

specimens kept at the Forest Herbarium Ibadan (FHI).

He therefore did not treat all the Solatium species occur-

ring in Nigeria. In Ghana itself not much work has been

done on Solatium. Some data on Solammi are found in

Floras, including the Flora of tropical Africa (Wright

1906) and the Flora of west tropical Africa (Heine

1963). In the Flora of tropical Africa, species were

viewed in a narrow sense and interspecific variation

seems to have been ignored. Species delimitation was

based on inadequate material and on few morphological

characters.

Wright (1975), Nsowah (1969) and Hossain (1973),

studied a few species of Solatium in Ghana. Wright

(1975), however, based his study on specimens collected

from one district (Cape Coast), hence the complex range

of variation of the species he studied, particularly S.

nigrum s.L, could not be appreciated. Nsowah (1969)

unfortunately did not identify with care the several

strains of garden eggs he studied.

Despite the existence of the FWTA edn 2 (Heine

1963), many workers, especially in horticulture and ge-

netics, continue to misidentify Solatium spp. For exam-

ple Epenhuijsen (1974) misapplied the names S. gilo

Raddi to S. macrocarpon L., S. iticanum L. to S. gilo

and S. scabrum to 5. nigrum L.; Nsowah (1969) misap-

plied S. integrifolium Poir to S. gilo.

Such misidentifications occur because, unfortunately,

the FWTA, edn 2 does not provide full descriptions for

the West African Solanum spp. but only gives a key to

the species. With a key alone one is likely to go wrong in

identification of critical species and the key in this Flora

does not, in some cases, bring out differences between

closely related species. This flora, like the treatment of

Solanum by Gbile (1979), does not give consideration to

experimental work on cultivated species, e.g. S. gilo

Raddi, S. aethiopicum L. and S. macrocarpon L., hence

their variability was not appreciated. Critical species like

S. nigrum complex were treated only in the broad sense.

We have therefore attempted to provide detailed, up-

to-date descriptions of all the Solanum spp. occurring in

Ghana.

MATERIALS AND METHODS

The gross morphological studies were based on about

200 specimens. A large number of these specimens were

those already existing at the Legon herbarium, Ghana,

whereas others were freshly collected by the authors

from various regions of Ghana.

Information on all the specimens was recorded and the

morphological characters of the species scored. For

TABLE 1. — Infrageneric classification of Ghana Solanum spp. based on

D’Arcy (1972)

Bothalia 18,1 (1988)

some of the species, viz. S. nigrum s.l., S. americanum,

S. scabrum, S. melongena, S. macrocarpon complex, S.

anguivi Lam., S. aethiopicum L. and S. gilo, various

accessions were collected from different localities in

Ghana and were grown in an experimental plot at Legon.

The morphological characteristics of the plants belong-

ing to the above species were observed throughout the

various developmental stages and measurements of, for

example, flower and fruit characters were based on ma-

ture live plants. Specimens for the herbarium were ob-

tained from these plants.

After the study all the specimens were deposited at the

Legon herbarium. Some of the specimens studied are

cited under the species descriptions. The infrageneric

81

classification of Ghana Solatium spp. (Table 1 ) is based

on D’Arcy (1972).

DESCRIPTIONS OF SOLANUM SPECIES

Solanum L., Species plantarum: 184-188 (1753);

L.: 85 (1754); D’Arcy: 85(1973).

The genus is recognized by the 5-parted calyx and by

long, often connivent anthers dehiscing by terminal

pores and with short filaments. Other useful characters

for recognition are the frequently rotate, 5-lobed corolla,

the fruit being a berry with flattened seeds; often stellate

pubescence and prickles; the often extra-axillary, usually

cymose inflorescence.

KEY TO SPECIES

la Plants climbers:

2a Leaves deeply lobed 20. S . seaforthianum

2b Leaves more or less entire:

3a Fruits spindle-shaped; inflorescence lateral, up to 7-flowered 4b. S. terminate subsp. inconstans

3b Fruits globose; inflorescence terminal, spiciform, up to 40-flowered 4a. S. terminate subsp. welwitschii

lb Plants non-climbers:

4a Leaves compound; stolons tuberous 19. S. tuberosum

4b Leaves simple; tuberous stolons absent:

5a Fruits red when mature:

6a Leaves glabrous when mature, apart from a few scattered stellate hairs:

7a Flowers solitary or inflorescence with less than 10 flowers; corolla 5-6 mm long; fruit about 15 mm

diam 15. S. aethiopicum

7b Flowers up to 30 per inflorescence, axillary and subfasciculate; fruits about 6 mm diam. .. 18. 5. anomalum

6b Leaves stellate-pubescent;

8a Corolla about 8 mm long; inflorescence 3-20-flowered; fruits 7- 15 mm long 14. S. anguivi

8b Corolla about 15 mm long; flowers usually solitary, occasionally inflorescence 2( — 3 or morel-

flowered; fruits 20-70 mm long 16. S. gilo

5b Fruits more or less yellow or blackish when ripe:

9a Inflorescence umbelliform; hairs simple or absent; fruit blackish when mature:

10a Corolla about 7 mm long; fruits about 17 mm diam 3. 5. scabrum

10b Corolla 3-4 mm long; fruits 7— 10 mm diam.:

1 la Pollen diam. 17,7- 19,2 gm; seeds 1 x 1 mm; 2n=24 2. 5. americanum

lib Pollen diam. 25,1 — 28 /am; seeds 2x 1 mm; 2n = 72 la. S. nigrum s. str.

9b Inflorescence various, never umbellate; some hairs stellate; fruits yellow:

12a Fruits pubescent 5. S.erianthum

12b Fruits glabrous:

13a Small tree; upper surface of leaves with simple hairs, lower surface with stellate hairs 13. S. wrightii

13b Shrubs; hairs on both surfaces of leaves either stellate or simple, occasionally simple hairs on

upper surface, simple hairs mixed with stellate hairs on lower surface:

14a Leaves very small, about 35x25 mm, subsessile; prickles very much decurved 9. S. arutuio

14b Leaves about 170x80- 150 mm, petiolate; prickles not decurved or very slightly decurved:

15a Fruits less than 10 mm diam.; inflorescence up to 40-flowered, corymbose 17. S.torvum

15b Fruits more than 10 mm diam.; inflorescence less than 15-flowered, never corymbose:

16a Fruit length less than width r 12. S. macrocarpon

16b Fruit length greater than (rarely equal to) width:

17a Prickles almost always absent; fruits usually oval; flowers purple 10. S. melongena

17b Prickles always present:

18a Fruit with terminal nipple or mammilla 6. S.mammosum

18b Fruit without terminal nipple or mammilla:

19a Hairs on leaves all uniformly pilose; seeds winged 8. S. capsicoides

19b Hairs and seeds not as above;

20a Hairs on both surfaces of leaves all stellate; flowers purple 11. 5. incanum

20b Hairs on upper surface of leaves simple, lower with rare stellate hairs mixed with

simple hairs; flowers white 7. S. aculeatissimum

82

Bothalia 18,1 (1988)

Subgenus Solatium D'Arcy in Annals of Missouri

Botanical Garden 60: 733 (1973).

Leaves subentire or shallowly lobed, often membrana-

ceous, indumentum simple, hairs rarely branching.

Prickles absent. Flowers mostly small, congested.

Corolla mostly deeply lobed; filaments often pubescent.

Anthers short, dehiscing introrsely by large, often

oblique terminal pores and sometimes ultimately by

longitudinal slits in the upper portion. Ovary glabrous.

Fruit rather small, 7—15 mm diam.

Section Solanum D'Arcy in Annals of Missouri

Botanical Garden 60: 733 (1973).

1. Solanum nigrum L., Species plantation: 186

(1753); Heine: 335 (1963); Hepper: 12 (1976); Gbile:

1 17 (1979). All the above authors have taken S. nigrum

in a broad sense.

Herb, about 1 m high. Leaves elliptic, 60— 100 x

40—70 mm; lamina glabrous when mature, margin entire

or toothed, repand or with 2-3 pairs of short lobes;

petiole 10-30 mm long. Inflorescence 6-8-flowered;

peduncle ± 10 mm long; pedicels ± 5 mm long. Corolla

3-4 mm long. Fruits 5-8 mm diam., shiny black when

ripe; fruit stalk ± 10 mm long. Seeds ± 1 x2 mm.

The name S. nigrum has been used in West Africa in a

broad sense until the work of Edmonds (Gray 1968; Ed-

monds 1971, 1972). Edmonds identified some of the

specimens we sent to her from the Ghana herbarium

(Legon), and isolated three Ghanian taxa from the com-

plex. These are S. nigrum s. str., S. americanum Mill.,

and S. scabrum Mill. The latter is easy to isolate from

the other taxa on account of its fruit size (more than 10

mm diam.). The rest of the complex of species awaits a

thorough analysis before tens of specimens in the Ghana

herbarium can be placed in the correct taxa. S. nigrum

complex is widely distributed in Ghana.

Vouchers: Western Region: Ankobra Ferry nr Axim, fl. & fr. July

1969, Enli GC 39163; Busua Bay, fl. & fr. February 1956, Morton

A 1796. Central Region: Bando nr Asebu, fr. November 1962, Hall

2411. Eastern Region: Mampong Scarp, Akwapim, fl. & fr. June

1953, Morton s.n.; Mt Ejuanema, Kwahu, fl. & fr. December 1957,

Adams 5125. Volta Region: Adzide-Keta nr Keta Lagoon, fl. & fr.

August 1957, Akpabla 1921 ; Amedzofe, fr. June 1958 , Morton A3418.

Greater Accra Region: Legon Hill, fl. & fr. May 1959, Morton A3675

Achimota, fl. & fr. February 1926, Irvine 93.

la. subsp. nigrum.

Edmonds: 141-178(1977).

Herb about 1 m high, with abundant simple hairs

when young, subglabrescent when mature. Stem robust.

Leaves usually bearing simple hairs on both surfaces and

margin, margin repand or crenate with teeth ± 5 mm

long; petiole 20—35 mm long. Inflorescence lateral, um-

bellate cymes, 6-8-flowered. Corolla ± 4 mm long.

Fruit ± 7 mm diam., shiny black when ripe. Seeds ±

1x2 mm. 2n=72 (Edmonds 1977). Pollen diam.

25,1-28 gm. This subspecies and S. nigrum as a whole

constitute a Eurasian taxon. However, it has spread to all

the other continents.

Voucher: Greater Accra Region: by the railway station, Achimota,

fl. & fr. February 1954, Morton 25359.

2. Solanum americanum Mill, in Gardener’s dic-

tionary edn 8: (1768); D’Arcy: 735 (1973); Edmonds:

141-178(1977).

Herb about 1 m high; stem with simple hairs when

young, glabrous when mature. Leaves 50-80x30—50

mm; mature lamina glabrous, margin with two pairs of

lobes, up to 10 mm long; petiole up to 20 mm long.

Inflorescence lateral, umbellate cymes, 5— 6-flowered;

peduncle ± 10 mm long; pedicel ± 7 mm long. Corolla

± 3 mm long; calyx ± 1 mm long. Fruit up to 9x10

mm, shiny black when ripe; fruit stalk 10-13 mm long;

fruiting calyx 1-2 mm long. Seeds lxl mm. 2n=24

(Edmonds 1977). Pollen diam. 17,7- 19,2 /am.

S. americanum is an introduced weed, collected from

the coast. The extent of its distribution will be fully

known only after a thorough study of the S. nigrum com-

plex has been made. It is of S American origin, probably

recently introduced into Africa.

Vouchers: Central Region: Cape Coast (Shore) nr Castle, fr. June

1979, Hall & Bukenya GC 47120; Greater Accra Region: Legon Bo-

tanical Garden (experimental plot), fl. & fr. February 1980, Bukenya

S19 (grown from seed of GC 47120).

3. Solanum scabrum Mill, in Gardener’s dictio-

nary edn 8: (1768); Edmonds: 141 — 178 (1977).

Bushy subshrub 1 - 1 ,5 m high; stem with simple hairs

when young, glabrous when mature. Leaves up to

250x200 mm; lamina glabrous; petiole up to 70 mm

long. Inflorescence lateral, umbellate cymes, 6 — 10-

flowered; peduncle 20-25 mm long; pedicel ± 5 mm

long. Corolla ± 7x1,5 mm. Fruit 13x17 mm, dull

purple-black when ripe; fruit stalk 30-40 mm long;

fruiting calyx about 2 mm long. Seeds 1x2 mm. 2n=72

(Edmonds 1977).

In Ghana S. scrabrum is known only in cultivation. It

is grown for its edible leaves. The fruits, though eaten in

Europe, where it is known as ‘garden huckleberry’, do

not seem to be eaten in Ghana. It is of African origin,

and has spread to America and other continents.

Vouchers: Greater Accra Region: Avenor, fl. & fr. July 1979, Hall

<& Bukenya GC 47130; Legon Botanical Garden (experimental plot),

fl. & fr. February 1980, Bukenya S44 (seed from GC 47130); Akligo,

fl. & fr. November 1967, GC 37361 .

Section Afrosolanum Bitter in Botanische

Jahrbucher 54: 420-1, 440 (1917); D’Arcy: 266, 274

(1972).

4a. Solanum terminale Forssk. subsp. welwitschii

(C. H. Wr.) Heine in Kew Bulletin 14: 248 (1960);

Heine: 331 (1963); Gbile: 118(1979).

S. welwitschii C.H. Wr.: 126 ( 1894); C.H. Wr.: 213 ( 1906); Bitter:

478 (1917); Hutch. &Dalz.: 206(1931).

Slender woody climber. Leaves up to 170 x 80 mm;

petiole up to 30 mm long. Inflorescence terminal, spici-

form ; flowers up to 40, in cymules, subsessile on axis.

Corolla 8-10 mm long with very few simple hairs on

outer surface, glabrous on inner surface. Fruits globose,

± 10 mm diam.; fruit stalk up to 6 mm long. Seeds ±

1 ,5 x 2,5 mm, smooth.

Subsp. welwitschii is fairly well distributed in secon-

dary forest.

Vouchers: Western Region: nr Bibiani, 11. & fr. December 1953,

Adams 2101. Central Region: nr Manso, Akim, fr. December 1933,

Irvine 2086. Eastern Region: Kade, fl. & fr. February 1955, Morton

8387; Atewa F.R., fr. January 1972, Lock & Hall GC 43502. Volta

Region: Togo Plateau, fl. April 1953, Morion 9197.

Bothalia 18,1 (1988)

83

4b. Solanum terminate Forssk. subsp. inconstans

(C.H. Wr.) Heine in Kew Bulletin 14: 247 (1960);

Heine: 331 (1963); Gbile: 1 18(1979).

5. inconstans C.H. Wr.: 127 (1894); Bitter: 482 (1917); Hutch. &

Dalz.: 207(1931).

S. togoense Dammer: 59 (1905).

5. suberosum Dammer: 182 (1906).

Slender woody climber. Leaves up to 100 x 50 mm;

petiole up to 18 mm long. Inflorescence subracemose,

leaf-opposed or in leaf axil, 2-7-flowered; peduncle up

to 35 mm long; pedicel ± 15 mm long. Corolla 8—10

mm long, with simple hairs on the outer surface, gla-

brous on the inner surface. Fruits spindle-shaped, ± 25

x 10 mm; fruit stalk ± 28 mm long. Seeds 1,5 x 2 mm,

embedded in a cushion of hairs ± 1 mm long.

Subsp. inconstans is a rare taxon, found in disturbed

forest. Both subspecies of S. terminale are restricted to

tropical Africa.

Vouchers: Eastern Region: Kade F.R., fl. October 1972, Hall GC

43831 . Volta Region: Togo Plateau F.R., fl. & fr. June 1972, Abbiw

GC 43712.

Subgenus Brevantherum (Seithe) D’Arcy in An-

nals of Missouri Botanical Garden 59: 267-274 (1972);

D’Arcy: 713 (1973).

Plants unarmed. Hairs branched or stellate. Leaves

mostly entire or nearly so. Anthers stout, opening by

large terminal pores and sometimes ultimately by longi-

tudinal slits. Ovary glabrous to tomentose. Fruits rather

small, 7-20 mm diam.

Section Brevantherum D'Arcy in Annals of Mis-

souri Botanical Garden 60: 716 (1973).

5. Solanum erianthum D. Don, Prodromus florae

Nepalensis: 96 (1825); D’Arcy: 717 (1973); Gbile: 118

(1979).

S. verbascifolium auct. non L.: Irvine: 733 (1961); Heine: 335

(1963).

Shrub or small tree 4-7 m high; young stem heavily

covered with sessile stellate hairs. Leaves elliptic-ovate,

entire, up to 230 x 180 mm; apex acute, base truncate,

oblique or attenuate; both surfaces of leaves bear a heavy

covering of stellate hairs with 5-15 arms raised on se-

tae; petiole about 35 mm long. Inflorescence terminal,

paniculate with numerous flowers. Flowers perfect; pe-

dicel ± 5 mm long, bearing more or less sessile stellate

hairs with ± 15 arms. Corolla 7-9 mm long, bearing

sessile stellate hairs on the outer surface, and veins on

inner suface. Calyx on both sides bearing sessile stellate

hairs. Fruit spherical, pubescent, ± 8 mm diam., green

when young, yellow when ripe; fruit stalk ± 10 mm

long. Seeds 1 x 1 mm.

5. erianthum is a widespread tropical weed. S. erian-

thum and 5. torvum form the most abundant shrub seed

stock in Ghanian forest soils, with the greatest abun-

dance in the Moist Semideciduous, Dry Semideciduous

and Southern Marginal vegetation types. When forests

are cut the seeds germinate and these species form the

most abundant secondary species. Fruits of S. erianthum

are believed to be poisonous; leaves are used for healing

‘Craw-Craw’.

The closely related species, 5. mauritianum Scop, and

S. umbellatum Miller are established elsewhere in Africa

(R. N. Lester pers. comm.). These two species however

have not yet been recorded for Ghana.

Vouchers: Western Region: Krobo, fr. May 1952, Boughey, s. n.

Eastern Region: Hweehwee nr Akwaseho, fl. June 1979, Hall & Buken

ya GC 47105; Kade F.R., Owusu, fr. August 1970, Annan & Aryeetey

GC 53378. Brong Ahafo Region: Goaso-Ntotroso, fr. April 1968,

Andoh-Ampah 53628. Volta Region: Akpafu-Mempeasem, fr. January

1950, Noamesi 64.

Subgenus Leptostemonum (Dun.) Bitter in Bota-

nische Jahrbiicher 55: 69-89 (1919); D’Arcy: 684

(1973).

Indumentum often stellate; prickles usually present.

Anthers mostly slender, tapering to the tip and opening

by small terminal pores or, if stout, they narrow abruptly

to a small tip and often open also by longitudinal slits

near the base, dehiscing introrsely or extrorsely by out-

ward bending of the tips. Ovary glabrous. Fruits often

large, (7-) 10-80 (-90-130) mm diam.

Section Acanthophora Dun., Histoire des Sola-

num: 131 - 132 (1813); D’Arcy: 709 (1973).

6. Solanum mammosum L., Species plantarum:

187 (1753); D’Arcy 712 (1973); Nee: 576 (1979).

Shrub about 1,5 m high; stem densely clothed with

simple hairs ± 2 mm long, and slightly decurved

prickles that are 5 mm long, base 2 mm broad. Leaves ±

110 x 90 mm, lobed or doubly-lobed, to about V: width

of leaf; lobes triangular; prickles on midrib straight, ±

17 mm long, base 1 mm broad, on primary lateral veins

3-8 mm long; hairs on both surfaces of leaves mainly

simple, on lower surface some stellate hairs mixed with

simple hairs; petiole up to 70 mm long with simple hairs

and straight prickles ± 13 mm long, base ± 1 mm

broad. Inflorescence 3-4-flowered; pedicel ± 7 mm

long. Corolla violet or blue, ± 12 mm long. Fruit up to

50 mm wide, bearing a terminal nipple or mammilla, 5

mammillae or protuberances often present at base.

S. mammosum is an introduced plant rarely seen, see-

mingly not naturalized. It is of South American origin,

naturally growing in disturbed habitats. It has spread to

other parts of the world as an ornamental plant. Its fruits

are poisonous.

Voucher: Ghana, Greater Accra Region: school nursery, Achimota,

fl. March 1970, Doe-Lawson GC 39999.

7. Solanum aculeatissimum Jacq., leones planta-

rum: 45 (1781); C.H. Wr.: 228 (1906); Bitter: 148

(1923); Dalziel: 432 (1937); Heine: 535 (1963); Gbile:

115(1979).

Shrub ± 1,5 m high; stem densely pubescent, with

simple hairs 0,1 — 1 mm long, and decurved prickles up

to 12 mm long with base 0,5 mm broad. Leaves ± 180 x

160 mm; lobed or doubly lobed to / width of leaf, with

3—4 pairs of major triangular lobes; simple pilose hairs

on both surfaces of leaves, rare stellate hairs on under-

side; prickles on midrib and petiole, slender, ± 15 mm

long, base ± 2 mm broad, on primary lateral veins,

slender, ± 7 mm long, base 0,8 mm broad. Inflores-

cence 3-6-flowered; petiole, 4-10 mm long. Corolla

white, ± 12 mm long; peduncle 0-0,5 mm long; pedicel

± 1 1 mm long. Fruit globose, ± 25 mm diam. Seeds 3

x 4 mm.

84

S. aculeatissimum occurs wild in secondary forest, but

is rare in Ghana. Ghana material has shorter hairs and

less abundant prickles than specimens from the moun-

tains of Sierra Leone and Cameroun. The species is trop-

ical African, found in the tropical zone and in South

Africa, but has close relationship to several South Amer-

ican species. All Ghana specimens come from the West-

ern Region.

Vouchers: Western Region: forest clearing, Awaso Sefwi, fl. & fr.

December 1953, Adams 2080\ nr Ebin timber Rd, June 1956, Enti FH

6440.

8. Solatium capsicoides All. in Melanges de Philo-

sophic et de Mathematique: 64 (1774); Nee; 574 (1979).

S. ciliatum Lam.: 21 (1799); Heine: 334 (1963).

Shrub ± 1,5 m high; stem bearing sparse simple hairs

4-5 mm long, and prickles 2— 10 mm long with base up

to 2 mm broad. Leaves ± 150x135 mm, lobed or

doubly-lobed to /—/ width of leaf; lobes 2-3 pairs,

ovate; both surfaces with sparse uniform simple pilose

hairs; margin ciliate; prickles on midrib ± 13 mm long,

base 2 mm broad, on primary and secondary lateral veins

2-7 mm long; petiole up to 50 mm long with simple

hairs and prickles 14 mm long with base 2 mm broad.

Inflorescence ± 4-flowered; pedicel ± 10 mm long.

Corolla white or pale mauve, ± 12 mm long. Fruit

globose, ± 45 mm diam. Seeds ± 1,5x2 mm, sur-

rounded by a wing ± 2 mm broad.

S. capsicoides has limited distribution in Ghana; appa-

rently introduced into botanical gardens as an ornamental

plant; it does not seem to be naturalized. Its origin is

South American (Brazil) and it has successfully spread to

all continents as a weed with preferred habitat being

drier, open lowlands.

Vouchers: Eastern Region: outside Botanical Garden Aburi, fl. & fr.

January 1957, Akpabla 1816. Greater Accra Region: Legon, fl. & fr.

October 1961, Morton A5008.

Section Ischyracanthum Bitter in Feddes Reperto-

rium, Beihefte 16; 142(1923); D’Arcy: 269, 275 (1972).

9. Solanum arundo Mattei in Bollettino del R.

Orto botanico, Palermo: 188(1908).

Shrub 1-2 m high; stem bearing stellate hairs;

prickles strongly decurved, ± 8 mm long, base up to 5

mm broad. Leaves very small with 2 pairs of lobes ± 8

mm long, ± 35 mm wide; base attenuate; small stellate

hairs with 6—8 more or less equal arms on both surfaces;

prickles on leaves straight, up to 12 mm long, base 3 mm

broad; petiole ± 3 m long. Inflorescence racemose, op-

posite a leaf or slightly below; flowers up to 10 or more;

pedicels ± 6 mm long, with stellate hairs. Corolla vio-

let, ± 13 mm long, with stellate hairs on the inner sur-

faces, on veins and on the outer surface. Fruits spheri-

cal, ± 13x12 mm, with more or less decurved stalks ±

14 mm long; young fruits variegated dark and pale green;

ripe fruits brownish. Seeds 2x2 mm.

S. arundo was probably introduced into Legon Bota-

nical Garden from northern Kenya; it is not naturalized.

Its area of concentration is in the drier parts of east tropi-

cal Africa. It has also been recorded from the west coast

of India.

Bothalia 18,1 (1988)

Voucher: Ghana, Greater Accra Region: Legon Botanical Garden,

fl. & fr. October 1958, Morton A3442.

Section Melongena Dun., Histoire des Solanum:

208-218 (1813); D’Arcy: 698 (1972).

10. Solanum melongena L., Species plantarum:

186 (1753); Wright: 242 (1906); Bitter: 292 (1923);

Heine: 322 (1963); D'Arcy: 704 (1973); Hepper: 122

(1976); Khan: 630 (1979).

S. edule Schumach. & Thonn.: 125 (1827); Junghans: 91 (1962).

Shrub ± 1,5m high; stem with stellate hairs of 8 - 10

unequal arms. Leaves ±150x100 mm, bearing on both

surfaces short-stalked stellate hairs, margin with 2—3

pairs of lobes up to 30 mm long; petiole about 40 mm

long. Flowers usually solitary or inflorescence 2—3-

flowered; pedicel ± 25 mm long. Corolla 20-22 mm

long, purple, petal tips apiculate, incurved. Calyx about

10 mm long, normally not prickly but prickles 2—3 mm

long may occur. Fruit ovoid or globose, 60— 130x

30—100 mm, green, with white patches, white, or

purple when young, orange-yellow to brownish when

ripe; fruit stalk 20-80 mm long, decurved. Seeds 3x4

mm.

S. melongena is cultivated for its fruits which are

eaten in stew and soup. It is a significant source of in-

come to horticultural farmers especially in the coastal

parts of southern Ghana, the species is believed to have

originated in Asia (Indo-Burma). It is cultivated on all

continents for its edible fruits.

Vouchers: Central Region: Kromantse, fr. June 1979, Hall &

Bukenya GC 47121. Eastern Region: N of Mankrong (Affram), fr.

May 1961, Morton A4193. Greater Accra Region: Nungua, fl. July

1979, Hall & Bukenya GC 47135 ; Achimota, fl. Aug. 1927, Inine

775.

11. Solanum incanum L., Species plantarum: 188

(1753); C.H. Wr.: 238 (1906); Bitter: 200 (1923); Dal-

ziel: 433 (1937); Heine; 332 (1963); Gbile: 1 18 (1979).

Shrub 1 - 1 ,5 m high; stem with stellate floccose hairs

with 8-10 more or less equal arms; prickles ± 4 mm

long, base ± 1 mm broad. Leaves ± 160x120 mm,

margin with 3 pairs of short lobes, middle lobe up to 20

mm long, both surfaces with stellate hairs; prickles on

midrib ± 3 mm long, base ± 0,5 mm broad; petiole up

to 70 mm long. Inflorescence up to 5-flowered; pedicel

± 10 mm long. Corolla violet, ± 18 mm long. Calyx,

especially on the lowermost flower, very prickly,

prickles ± 2 mm long. Fruits globose, ± 20 mm diam.,

green with light green patches when young, yellow when

ripe.

S. incanum is a rare species in Ghana. It has not been

collected since 1956. It is common in drier parts of tropi-

cal Africa and India. In Ghana it has been collected from

one locality.

Voucher: Greater Accra Region: Nungua beach, roadside, fl. May

1956, Morton 2086.

12. Solanum macrocarpon L., Mantissa Altera:

205 (1771); C.H. Wr.: 214 (1906); Bitter: 195 (1923);

Burkill: 333 (1925); Dalziel: 343 (1937); Heine: 224

(1963).

S. dasyphyllum Schumach. & Thonn.: 126 (1827); Bitter: 188

( 1923); Heine: 334 ( 1963), syn. nov.

Bothalia 18,1 (1988)

85

S. duplosinuatum Klotzch: 233 (1862); Wright: 243 (1906); Hutch.

&Dalz.: 207 (1931).

5. thonningianum Jacq.: 123 (181 1-16); Junghans: 91 (1962).

S. atropo Schumach. & Thonn.: 124 ( 1827).

S. zanoni Gouan (1773)Gouan: 7 (1773).

Subshrub or shrub, 0,5- 1 ,5 m high; stem terete, gla-

brous or with stellate hairs more or less sessile or on

robust setae up to 2 m long. Stem not prickly or with

straight, robust prickles up to 6 mm long, base up to 3

mm broad. Leaves 150—460 x 80-300 mm, entire or

with short lobes ± 10 mm long to deeply doubly-lobed

with major lobes up to 70 mm long; young leaves bear on

upper surface (especially on veins) simple, branched or

stellate hairs on setae up to 2 mm long, either singly or in

combination; lower surface with branched, stellate hairs

on robust setae 0,5—2 mm long and/or more or less

sessile stellate hairs; leaf margin of young leaves with

stellate hairs on robust setae up to 2 mm long or short-

stalked glandular hairs; mature leaves glabrous or with

simple hairs and stellate hairs on setae up to 2 mm long,

upper surface with sessile stellate hairs; lower surface

with stellate hairs on setae up to 2 mm long or sessile;

leaf margin with stellate hairs on setae up to 2 mm long,

sessile stellate hairs, branched or simple hairs; prickles

present or absent on leaves, when present, principally on

midrib and lateral veins, on mature leaves prickles on

midrib and primary lateral veins straight, up to 13 mm

long, base ± 3 mm broad, on secondary and tertiary

lateral nerves up to 7 mm long, base 1 mm broad; leaf

subsessile or with petiole up to 70 mm long. Inflores-

cence lateral, racemose, 3- 12-flowered, the lowermost

flower or flowers hermaphrodite, bigger than the rest,

distal flowers with short styles, functionally male, nor-

mally 1 — 2( — 3) hermaphrodite plus 1—3 functionally

male flowers present. Corolla infundibuliform-rotate or

campanulate-rotate, 20-35 mm long, white, light purp-

lish or blue, the outer surface with simple, sessile stellate

hairs or short-stalked or sessile glandular hairs, inner

surfaces glabrous. Calyx not prickly or with prickles up

to 10 mm long; outer surface with sessile stellate hairs,

stellate hairs on setae up to 2 mm long, simple hairs or

short-stalked glandular hairs; fruiting calyx often accres-

cent, 15-1 10 mm long. Ovary glabrous or with short-

stalked or sessile glandular hairs. Fruit depressed glo-

bose, 20-60 x 30-100 mm, green, ivory or purplish

white with dark stripes; when ripe yellow to brownish;

fruit stalk erect or decurved, 10—40 mm long. Seeds

3—3,5 x 2— 3 mm.

The S. macrocarpon complex is extremely variable.

In Ghana it is cultivated mainly for its leaves, especially

of plants with glabrous leaves. Leaves that are prickly

have to be de-prickled before cooking. The fruits are also

eaten. In other parts of Africa, e.g. E Africa, it is less

cultivated. It is a native species of Africa from where it

has been introduced to other parts of the world. It occurs

in almost every region of Ghana.

Vouchers: Western Region: Nkwanta, Ankasa G.P.R., fl. & fr.

January 1980, Hall GC 471680. Ashanti Region: Abodon, fr. June

1979, Hall & Bukenya GC 4711. Central Region: Anyam Main, fl. &

fr. June 1979, Hall & Bukenya GC 47122. Eastern Region: Hweehwee

nr Akwaseho, fl. & fr. June 1979, Halt & Bukenya GC 47103 ; Kade,

fl. & fr. January 1980, Hall GC 471679. Brong Ahafo Region: Bui

Nat. Park, fl. & fr. July 1976, Hall & Swaine GC 46122. Greater

Accra Region: Achimota, fl. May 1958, Akpabla 1783: Sapreiman, fl.

& fr. July 1979, Hall & Bukenya GC 47128.

13. Solatium wrightii Benth. in Flora Hongkong-

ensis: 243 (1861); Bitter: 180(1923); Heine: 248 (1960);

Irvine: 733 (1961); Heine: 335 (1963); Gbile: 119

(1979).

Small tree up to 5 m high; stem bearing prickles, and

stellate hairs on setae ± 1 mm long. Leaves ± 250 X

200 mm, with 2—3 pairs of prominent lobes up to 80 mm

long; upper surface bearing simple hairs ± 1 mm long,

lower surface with stellate hairs with 5 — 8 unequal arms

on setae ± 0,5 mm long; prickles on midrib 10-20 mm

long, base 4 mm broad; petiole ± 60 mm long; prickles

on petiole up to 20 mm long, base 4 mm broad. Inflo-

rescence ± 6-flowered; pedicel ± 20 mm long. Corolla

blue to violet, up to 30 mm long. Fruit globose, green

when young with light green patches, yellow when ripe.

S. wrightii is a decorative introduced tree; a native of

Bolivia, it has been introduced to other tropical areas of

the world.

Vouchers: Ghana, Greater Accra Region: Cadbury House 8, Achi-

mota, fl. 1935, Bannerman s. n .; Hort. Nursery, Achimota, fl. January

1958, Akpabla I860.

Section Oliganthes Bitter in Feddes Repertorium,

Beihefte 16: 1 (1923); D’Arcy: 272(1972).

14. Solanum anguivi Lam. in Tableau Encyclope-

dique et Methodique: 23 (1794); Hepper: 287—292

(1978).

5. indicum L. subsp. distichum (Schumach. & Thonn.) Bitter 13

(1923); Heine: 333 (1963); Irvine 73 (1961); Hepper: 121 (1976);

Gbile: 116(1979).

S. indicum auct. non L.

S. anomalum auct. non Thonn.

Shrub 1,5—3 m high; stem bearing small, sessile

stellate hairs with 4-8 more or less equal arms; some-

times prickly. Leaves elliptic-ovate, 100—200 x

50-100 mm, sinuate to distinctly lobed, 2—4 pairs of

lobes, lobes 20-30 mm long, apex acute to obtuse, base

oblique, occasionally truncate or subcordate; leaves bear

on both surfaces but nore so on the lower, more or less

sessile stellate hairs with 6- 10 more or less equal arms;

petiole 20—60 mm long, with dense stellate hairs. Inflo-

rescence a raceme-like cyme, occasionally flowers soli-

tary, extra-axillary, many-flowered, 5 to more than 15,

flowers mostly hermaphrodite, occasionally the distal

flowers with short styles, functionally male; peduncle

0-3—6 mm; pedicel 10-15 mm long, bearing stellate

hairs. Corolla ± mm long, white, occasionally with

light purple veins on the outer surface; with stellate hairs

outside, more or less glabrous on inner surface. Fruit

7—12 x 8-12 mm, mostly globose, smooth, green

when young, red when ripe; fruit stalk 8-15 mm long,

usually erect, occasionally horizontal or decurved. Seeds

1 ,5—2,1 mm long by 1 ,9-2,9 mm wide.

S. anguivi is widely distributed in Ghana, either wild

or as a semicultivated plant for the fruits which are bitter

and cooked in stew. The soup is taken before a meal as

an appetiser. The species is widely distributed through-

out Africa as a semicultivated plant or as a weed in

varying ecological situations. In Ghana it is found in all

the regions.

Vouchers: Western Region: Tonton F. R. (cleared area), fr. May

1956, Enti FH 6095. Ashanti Region: Abodon, fl. & fr. June 1979,

Hall & Bukenya GC 47113', Drobo, fl. & fr. April 1958, Morton

A3228. Eastern Region: Aburi, fl. & fr. April 1899. Brown 320: Ajena

86

Bothalia 18,1 (1988)

to Gyakiti Rd, fl. & fr. October 1953, Morton s. n. Brong Ahafo

Region: nr Menje, fl. & fr. December 1956, Morton A2609. Volta

Region: Akpafu Mempeasem, fl. & fr. 1956, Noamesi 68. Greater

Accra Region: Achimota, fl. & fr. June 1932, Irvine 1770', fl. & fr.

March 1954, Morton GC 25400.

15. Solanum aethiopicum L., Amoenitates acade-

micae: 307 (1759); Dalziel: 432 (1937); Heine: 332

(1963); Gbile: 115(1979).

Subshrub, about 0,6 m high; stem glabrous. Leaves

ovate, about 120 x 90 mm, apex acute, base oblique,

attenuate or truncate; margin repand; young leaves bear

on both surfaces small, sessile stellate hairs with 5-8

more or less equal arms; mature leaves subglabrous; pe-

tiole 50—60 mm long. Inflorescence 2— 3-flowered, ses-

sile or flowers solitary, lateral; flowers hermaphrodite;

pedicel ± 7 mm long with scattered stellate hairs. Co-

rolla white, 5-6 mm long, glabrous; flower buds with

dense stellate hairs; style also with stellate hairs. Calyx

as long as the corolla tube. Fruit globose ± 18 mm

diam., green with dark green stripes when young, shine

red when ripe; fruiting calyx ± 7 mm long; stalk up to 12

mm long. Seeds 2,8 x 3 mm.

S. aethiopicum is occasionally cultivated by immi-

grants from Togo in southern Ghana for its leaves which

are used as a vegetable. In other parts of tropical Africa,

it is more frequently cultivated. Lester & Niakan (1986)

treat it as Solanum aethiopicum Shumach group. In

Ghana it was collected from one locality.

Vouchers: Greater Accra Region: Sarpeiman, fl. July 1979, Hall &

Bukenya GC 47131' fl. & fr. July 1979, Hal! GC 47129.

16. Solanum gilo Raddi in Atti della Societa Ita-

liana delle Scienze Modena: 31 (1 920); Bitter: 48 ( 1 923);

Hepper: 121 (1976); Gbile: 116(1979).

S. integrifolium auct. non Pior.: Nsowah: 61 (1969).

Shrub 1-2 m high; stem bearing tiny sessile stellate

hairs of 4-5 more or less equal arms. Leaves elliptic-

ovate, 100-240 x 70-180 mm, margin sinuate-repand

to distinctly lobed with 2-4 pairs of lobes up to 50 mm

long, apex acute, base oblique; both surfaces but es-

pecially the lower, with more or less sessile stellate hairs

with 7—9 unequal arms; petiole 15-80 mm long;

prickles may occasionally be present on early leaves.

Flowers usually solitary or two next to each other, rarely

2— 5 or more arising from a common stalk, extra-axillary

and hermaphrodite. Corolla ± 15 mm long, white, occa-

sionally light purple; petal tips apiculate, incurved; inner

and outer surface of corolla with stellate hairs. Fruit

ellipsoid, globose or oval, 13-70 x 13-50 mm,

smooth or with shallow longitudinal grooves, usually

solitary, occasionally two or more together, green, white

or purple when young, red when mature; fruit stalk

16-40 mm long, mostly decurved. Seeds 3x4 mm.

S. gilo is by far the most widely cultivated species of

Solanum throughout the forest zone of Ghana. Its fruits

are used in soup or stew preparation. It is preferred to S.

melongena because it has firmer flesh than the latter.

Where these two species are grown together sterile natu-

ral hybrids occur.

The type specimen was collected from Brazil, but the

species must have been introduced into South America

from Africa. The Brazilian vernacular name, ‘gilo’, was

adopted as the specific epithet. The same vernacular

name has been recorded for the species in Angola, sug-

gesting that Angola may well have been the source of the

Brazilian plants. S. gilo has subsequently been intro-

duced to other parts of the world. Lester & Niakan

(1986) treat it as Solanum aethiopicum L. Gilo group.

Vouchers: Ashanti Region: Abodon, fl. & fr. June 1979, Hall &

Bukenya GC 47116. Eastern Region: Hweehwee nr Akwaseho, fl. &

fr. June 1979, Hall & Bukenya GC 47127. Volta region: Ashiagborvi,

nr Tadzevu, V/R, fl. & fr. August 1979, Hall GC 47150. Greater

Accra Region: Legon, fl. & fr. December 1979, Hall & Bukenya GC

47116, GC47117.GC47118.

Section Torva Nees in Transactions of the Linnean

Society: 51 (1834); D’Arcy: 273 (1972).

17. Solanum torvum Swartz, Prodromus: 47

(1788); Wright: 231 (1906); Bitter: 252 (1921); Dalziel:

435 (1937); Irvine: 732 (1961); Heine 335 (1963);

D’Arcy: 708 (1973); Hossain: 293 (1973); Gbile: 118

(1979).

Shrub about 2 m high; stem bearing stellate hairs and

prickles up to 8 mm long and 7 mm broad. Leaves up to

160 x 150 mm; margin subentire or distinctly lobed with

2-3 pairs of lobes; lobes up to 50 mm long; base ob-

lique, attenuate or subcordate; young leaves with dense,

stalked stellate hairs on both surfaces; mature leaves on

upper surface with more or less sessile, stellate hairs,

4-6 armed with one arm much longer than the rest;

lower surface with stalked stellate hairs of 8-9 more or

less equal arms; petiole up to 70 mm long. Inflorescence

a corymbose cyme; lateral, distal flowers with short

styles, functionally male; peduncle ± 3 mm long; pedi-

cel ± 7 mm long, bearing glandular hairs. Corolla

white, 10-12 mm long, outer surface bearing sessile

stellate hairs, 6—7-armed with one arm much longer

than the rest, glabrous within. Calyx ± 4 mm long, lobes

triangular, bearing glandular and sessile, 7 -8-armed

stellate hairs on the outer surface, glabrous on the inner

surface. Fruit globose, 10-15 mm diam., yellow when

ripe; fruit stalk up to 20 mm long.

5. torvum is a common weed throughout the tropics.

Its fruits are collected in some parts of Ghana and eaten

in soup and stew. S. torvum has a wide distribution in

Ghana.

Vouchers: Ashanti Region: Abodon, fr. June 1979, Hall & Bukenya

GC 47112', Ashanti, fl. & fr. July 1953, Antenson s. n. Central Region:

Abura nr Cape Coast, fl. September 1962, Hall 2343. Eastern Region:

Hweehwee nr Akwaseho, fr. June 1979, Hall & Bukenya GC 47 101 ;

Bosuso, fl. & fr. January 1939, Akpabla 447. Volta Region: nr Bishop

Herman College Kpandu, fl. & fr. December 1973, Veldkamp 6086.

Greater Accra Region: Legon Hill, fl. & fr. October 1955, Adams

3371.

18. Solanum anomalum Thonn., Beskrivelse af

guineiske planter: 126 (1827); Bitter: 273 (1921); Dal-

ziel: 443 (1937) Irvine: 732 (1961); Heine: 332 (1963);

Gbile: 115(1979).

Shrub; stem glabrous; prickles about 5 mm long with

base 4 mm broad. Leaves sinuate, up to 170 x 80 mm;

base attenuate; young leaves bear on both surfaces a

dense covering of small sessile stellate hairs with 4—6

more or less equal arms; mature leaves glabrous; petiole

30—60 mm long. Inflorescence subfasciculate, in leaf

axils; peduncle ± 2 mm long; pedicel ± 6 mm long,

bearing small sessile stellate hairs. Flowers hermaphro-

dite. Corolla white, 5-6 mm long, with stellate hairs on

the outer surface, inner surface glabrous. Calyx ± 5 mm

Bothalia 18,1 (1988)

87

long, bearing small sessile stellate hairs on the outer

surface, glabrous within. Fruit spherical, ± 6 mm

diam., green when young, red when ripe. Seeds 2 x

2,5-3 mm.

S. anomalum occurs wild in thickets and secondary

forest in the drier parts of the forest zone. It seems to be

restricted to West Africa.

Vouchers: Western Region: Busua Bay, fl. & fr. May 1956, Morton

A2I13. Ashanti Region: Sunyani Sec. forest, fr. June 1956, Bossman

Osei 42. Central Region: Cape Coast, Hall 548. Eastern Region: Aburi

Scarp, fl. & fr. November 1955, Morton A1755 . Volta Region: Volta

Gap, fl. & fr. October 1953, Morton s. n. Greater Accra Region:

Legon, fr. August 1979, Hall GC 47134.

Subgenus Potatoe (G. Don) D’Arcy, in Annals of

Missouri Botanical Garden 59: 272 (1972); D’Arcy: 750

(1973).

Herbs, rarely woody, often glandular-pubescent and

aromatic, unarmed; hairs simple. Leaves mostly com-

pound or deeply lobed, but simple, entire leaves often

present at certain stage. Inflorescence mostly paniculate;

peduncles once or temately branched, often pendulous;

pedicels mostly articulating near the base or above it.

Fruits 10—20 mm diam.

Section Potatoe

19. Solanum tuberosum L., Species plantarum:

185 (1753); D’Arcy: 752 (1973).

Herb ± 0,5 m high; stem with shallow grooves when

dry, and simple hairs. Stolons tuberiferous. Leaves com-

pound, interceptedly imparipinnate; major leaflets 5-7,

alternating with ± 3 pairs of minor leaflets. Leaflets

elliptic to oblong-elliptic, ± 50 x 25 mm; simple hairs

on both surfaces; subsessile, margin entire; petiole up to

60 mm long. Inflorescence paniculate, with up to 10

flowers; pedicel 12-15 mm long with simple hairs, ter-

minal on long peduncle ± 80 mm long. Flowers her-

maphrodite. Corolla white, 10—12 mm long. Calyx ± 7

mm long, with simple hairs on the outer surface, gla-

brous within. Fruits globose, about 7 mm diam.

S. tuberosum, the Irish potato, is a mainly temperate

crop introduced to Ghana at about the time of the Second

World War. First introduced at Mampong (Ashanti Re-

gion), its cultivation was only slowly taken up by the

inhabitants of the area. Potato farming was therefore

taken over by the Ministry of Agriculture, which grew it

on big farms. The project was however abandoned. The

crop spread to other areas. The present major potato

growing areas are Pepease (Eastern Region) and Amed-

zofe (Volta Region), where it is grown early in the major

rainy season by individual farmers and co-operatives.

Potato is grown for its stem tubers, 90% of which are

consumed by foreigners.

5. tuberosum had its origin in the Andes (Bircher

1960). It was introduced to Europe, from where it spread

to other parts of the world. The Ghana herbarium has not

many collections of S. tuberosum.

Voucher: Volta Region: Kpoeta, fl. August 1946, Sape s. n.

Section Jasminosolanum Bitter ex Seithe in Bota-

nische Jahrbiicher 81,3: 291 (1962); D’Arcy: 757

(1973).

20. Solanum seaforthianum Andr. var. disjunc-

tum O. E. Schulz in Symbolae Antillanae: 169 (1909);

Bitter: 309 (1923); Irvine: 732 (1961); Heine: 332

(1963); D’Arcy: 758 (1973); Gbile: 1 17 (1979).

Woody climber; stem terete, glabrous. Leaves com-

pound, imparipinnate to deeply pinnatifid with about

7-9 leaflets or lobes; leaflets up to 50 x 30 mm, ellip-

tic; lower leaflets with a more or less winged petiolule up

to 5 mm long; upper leaflets (i.e. lobes) webbed to-

gether; lamina glabrous; margin ciliolate with sparse

simple hairs; petiole up to 50 mm long. Inflorescence

glabrous, mostly terminal or lateral, paniculate with

about 20 flowers; pedicel up to 7 mm long. Flowers

hermaphrodite. Corolla blue to violet, up to 10 mm

long. Fruits spherical, ± 10 mm diam., red when ma-

ture; up to 20 from an inflorescence; fruit stalk 10-14

mm long. Seeds 2x2 mm.

S. seaforthianum is an introduced decorative climber

seemingly not yet naturalized. Its native home is central

America and the West Indies. It has spread to many parts

of tropical Africa where it has been introduced for deco-

rative purposes. The species has been recorded in

Ghana.

Vouchers: Greater Accra Region: Achimota, fl. February 1931, Ear-

won //; fl. & fr. April 1937, Onyeama 17', fl. March 1954,

Ante son s. n.

DISCUSSION AND CONCLUSIONS

In Ghana, the most widespread Solanum species are S.

torvum, S. aethiopicum Gilo group, S. nigrum s. I. and

S. terminate subsp. welwitschii. On the other hand, the

S. aethiopicum Shumach group, S. scabrum, S. inca-

num, S. capsicoides and S. aculeatissimum have res-

tricted occurrence.

Although some species are constant, for example S.

torvum, S. erianthum and S. aethiopicum Gilo group,

others are not. In cultivation, the S. macrocarpon com-

plex, for instance shows great variability. Bukenya &

Hall (1987) identified, named and described six cultivars

of the S. macrocarpon complex in Ghana. Bukenya 1980

(in unpublished M.Sc. thesis) studied experimentally the

S. macrocarpon complex and proposed to sink S. dasy-

phyllum under S. macrocarpon. In our description of the

S. macrocarpon complex, the former is therefore listed

as a synonym of the latter. The classification and nomen-

clature of the infraspecific categories of the S. macrocar-

pon complex need to be revised.

A detailed study of the S. nigrum complex is required

before the numerous specimens belonging to this com-

plex of species, available at the Legon herbarium, can be

sorted out.

Experimental work on the difficult species of the ge-

nus has been recommended by Bitter (1923), Hawkes et

al. (1979) and Edmonds (1979) as necessary to contrib-

ute to the improvement of the taxonomy of Solanum.

Cross breeding experiments have thrown some light on

the taxonomy of the genus. Omidiji (1979) and Lester &

Niakan (1986) for example, obtained fully fertile hybrids

from crosses involving S. gilo and S. aethiopicum. The

chemotaxonomic study by Pearce & Lester (1979) points

to a very close relationship between S. melongena and 5.

anguivi. This has raised doubts as to the specific limits

within the above sets of taxa.

88

We agree with Heine (1963) that a modem revision of

the entire genus is long overdue.

ACKNOWLEDGEMENTS

We wish to extend our gratitude to the Uganda Gov-

ernment for sponsoring the study, and the University of

Ghana, Legon for providing the facilities for the study.

We are very grateful to Prof. R. H. Lester of Birming-

ham for his detailed comments and amendments.

Thanks go to Dr J. M. Edmonds of Cambridge for

identifying some of the specimens belonging to the S.

nigrum complex. We are also grateful to all individuals

who provided us with specimens and any form of assis-

tance during this study.

REFERENCES

BENTHAM, G. 1861 . Flora Hongkongensis: a description of the flow-

ering plants and ferns of the island of Hong Kong. Lovell Reeve,

London.

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tian Bookshops, Cairo.

BITTER, G. 1913. Solana Africana 1. Botanische Jahrbiicher 49:

560-569.

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416- 506.

BITTER, G. 1919. The papuasischen Arten von Solanum. Botanische

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BITTER, G. 192L Solana Africana 3. Botanische Jahrbiicher 57:

248-286.

BITTER, G. 1923. Solana Africana 4. Feddes Repertorium, Beihefte

16: 1-320.

BUKENYA, Z. R. & HALL, J. B. 1987. Six cultivars of Solanum

macrocarpon L. (Solanaceae) in Ghana. Bothalia 17: 91-95.

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333-341.

DALZIEL, J. M. 1937. The useful plants of west tropical Africa.

Crown Agents, London.

DAMMER, U. 1905. Solanaceae africanae. Botanische Jahrbiicher

38: 57-60.

DAMMER, U. 1906. Solanaceae africanae I. Botanische Jahrbiicher

38: 176-195.

D’ARCY, W. G. 1972. Solanaceae studies 2. Typification of subdivi-

sions of Solanum. Annals of the Missouri Botanical Garden 59:

262-278.

D’ARCY, W. G. 1973. Solanaceae. In R. E. Woodson Jr, R. W.

Schery & collaborators. Flora of Panama. Anrnls of the Missouri

■ Botanical Garden 60: 573—780.

DON, D. 1825. Prodromus florae Nepalensis; sive enumeratio vegeta-

bilium, quae in itinere per Nepaliam ...., Londini.

DUNAL, M. F. 1813. Histoire Naturelle, Medicinale et Economique

des Solanum. Paris, Strasbourg, Montpellier.

EDMONDS, J. M. 1971. Solanum L. In W. T. Steam, Taxonomy and

nomenclatural notes on Jamaican gamopetalous plants. Journal of

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EDMONDS, J. M. 1972. A synopsis of the taxonomy of Solanum sect.

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EDMONDS, J. M. 1977. Taxonomic studies on Solanum sect. Sola-

num (Maurella). Botanical Journal of the Linnean Society 75:

141-178.

EDMONDS, J. M. 1979. Taxonomic studies of Solanum sect. Sola-

num (Maurella). Linnean Society Symposium 7: 529-548.

EPENHU1JSEN, V. M. VAN. 1974. Growing native vegetables in

Nigeria. FAO, Rome.

GBILE, Z.O. 1979. Solanum in Nigeria. Linnean Society Symposium

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GRAY, J. M. 1968. The taxonomy of the Maurella section of the genus

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HAWKES, J. G., LESTER, R. N. & SKELDING, A. D. (eds). 1979.

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Solanum melongena L. Linnean Society Symposium 7: 615—627.

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Bothalia 18, 1:89-96(1988)

Notes on African plants

VARIOUS AUTHORS

ACAROSPORACEAE

L1THOGLYPHA, A NEW LICHEN GENUS FROM CLARENS SANDSTONE

Lithoglypha Brusse, gen. nov.

Thallus endolithicus (vel crustosus). Algae ad Chloro-

coccales pertinentes. Apothecia primuni cleistothecifor-

mia dein pseudolecideina. Excipulum atro-porphyreum,

inferne prosoplectenchymatum, superne paraplectenchy-

matum. Epithecium atro-porphyreum, paraplectenchy-

matum, evanescens. Hypothecium inspersum.

Hymenium hyalinum, 50—100 pm altum, J+ caeruleum

vel vinoso-rubescens. Paraphyses septatae, cylindricae,

conglutinatae, simplices vel ramosae et anastomosantes.

Asci clavati, tholis J— vel pallide caeruleis, gelatinis

extemis J+ caeruleis vel vinoso-rubescentibus (Figure

1). Ascosporae numerosae (± 100 vel plures), hyalinae,

ellipsoideae, simplices, parvae, 2—8 x 1—3 pm. Pycni-

dia globosa vel pyriformia, circa 150 pm diametro. Pyc-

nidiosporophora hyalina, cylindrica, septata, simplicia

vel ramosa. Pycnidiosporae acrogenae, hyalinae,

aciculares, 3,5-8 x 0,8 gun.

TYPE. — Lithoglypha aggregata Brusse

Thallus endolithic (or crustose). Algae belonging to

the Chlorococcales. Apothecia first cleistotheciform then

pseudolecideine. Excipulum dark reddish brown, proso-

plectenchymatous below, becoming paraplectenchyma-

tous above. Epithecium dark reddish brown, para-

plectenchymatous, evanescent, but vestiges always

remain at edges of hymenium. Hypothecium inspersed.

Hymenium hyaline, 50- 100 pm high, J+ blue to wine-

red. Paraphyses septate, cylindrical, conglutinate,

simple to branched and anastomosed. Asci clavate,

tholus J- or pale blue, external gel J+ blue or wine-red

(Figure 1). Ascospores numerous (± 100 or more),

hyaline, ellipsoid, simple, small, 2—8 x 1—3 gim.

Pycnidia globose or pear-shaped, about 150 pm diam.

Pycnidiosporophores hyaline, cylindrical, septate,

simple to branched. Pycnidiospores acrogenous, hyaline

needles, 4-8 x 0,8 pm.

Etymology: ‘lithos’ (Greek) = rock, ‘glyphe’ (Greek)

= writing on stone tablet or an engraving.

Lithoglypha aggregata Brusse, sp. nov.

Thallus endolithicus. Algae ad Myrmeciam perti-

nentes, cellulis 5—17,5 gm diametro. Apothecia breviter

pseudolirellina, usque ad 0,4 mm lata, aggregata,

fasciculis ad 6 mm diametro. Excipulum 15 — 20 pm

crassum, atro-porphyreum. Epithecium circa 15 pm

crassum, atro-porphyreum, evanescens. Hypothecium

hyalinum vel stramineum, obconicum, usque ad 50 pm

crassum. Hymenium hyalinum, 65-80 pm altum, J +

caeruleum. Paraphyses septatae, cylindricae, congluti-

natae, simplices vel ramosae et anastomosantes, lumini-

bus 0,8— 1 ,3 pm crassis. Asci clavati, 60— 67 x 14-26

pm, tholis J+ pallide caeruleis (Figure 1). Ascosporae

numerosae, hyalinae, ellipsoideae, simplices, 4—7 x

1, 3-2,4 pm. Pycnidia globosa vel pyriformia,

100- 180 pm profunda, 70—1 10 pm lata. Pycnidiospo-

rae acrogenae, hyalinae, aciculares, 3,5— 7,5 x 0,8 pm.

Thallus sine materiis chimicis.

TYPE. — Natal, 2929 (Underberg: 65 km SW of

Estcourt, Giant’s Castle Game Reserve, about half-way

along Giant’s Hut trail, along umChezi (Bushman’s)

River, on Clarens Sandstone on NE slope, in full sun,

alt. 1 950 m (— BC), F. Brusse 4558, 1986.01 .23 (PRE,

holo.; BM, COLO, LD, iso.). Figure 2.

Thallus endolithic. Algae, Myrmecia, 5-17,5 pm

diam. Apothecia shortly pseudolirelline to sometimes

round, up to 0,4 mm wide, aggregated, clusters to 6 mm

across. Exciple 15 — 20 gm thick, dark reddish brown.

Epithecium about 15 gm thick, dark reddish brown,

evanescent (except at edges). Hypothecium hyaline to

stramineous, obconical, up to 50 gun thick. Hymenium

hyaline, 65 — 80 pm high, J + blue. Paraphyses septate,

90

Bothalia 18,1 (1988)

cylindrical, conglutinate, simple to branched and anasto-

mosed, lumens 0,8— 1 ,3 /zm thick. Asci clavate, 60-67

x 14-26 /zm, tholus J + pale blue (Figure 1).

Ascospores numerous, hyaline, ellipsoid, simple, 4-7

x 1, 3-2,4 gm. Pycnidia globose to pear-shaped,

100—180 /zm deep, 70-110 gm wide. Pycnidiospores

acrogenous, hyaline needles, 3, 5-7, 5 x 0,8 /zm.

Chemistry: no lichen substances found by TLC.

FIGURE 2. — Lithoglypha aggregata Brusse, habit. F. Brusse 4558 ,

holotype. Scale in mm.

This new genus resembles Sarcogyne Fw. most

closely, but differs in the pycnidiosporogenous cells and

pycnidiospores which are needle-shaped instead of very

small ellipsoid, as in Sarcogyne (Poelt 1969: Poelt &

V£zda 1981 ).

The genus Sarcogyne has recently been subdivided

into Polysporina V£zda (1978; Poelt & V£zda 1981),

but they may be based on the same type, depending on

the clarification of the status of Sarcogyne corrugata

Fw., the type of Sarcogyne. Zahlbruckner (1927)

regarded S. corrugata as a synonym of S. simplex

(Davies) Nyl., the type of Polysporina V£zda (1978).

The genus Myriosperma Naeg. in Hepp, is illegitimate

(Art. 42b, Voss et al. 1983), and is therefore not avail-

able for those species with uncarbonized exciples and

epithecia. Nevertheless, Lithoglypha differs from both

these groups of Sarcogyne, by its pycnidiospores, which

are needle-shaped instead of very small ellipsoid (Poelt

1969; Poelt & V£zda 1981).

Besides the clear-cut difference in the pycnidiospores,

L. aggregata resembles S. simplex, also known from

non-calcareous rocks, but the apothecia show little ten-

dency to become fused into clumps. S. simplex, thus far

known only from the northern hemisphere, also has sig-

nificantly thinner ascospores. A species with a similar

apothecial habit is 5. cyclocarpa (Anzi) Stnr., but this is

an obligately calcicolous lichen, with a well developed

epilithic thallus, with thinner ascospores and which is

also known only from the northern hemisphere — only

Europe and western Asia in fact (Magnusson 1935,

1936, 1937; Poelt 1969; Poelt & VSzda 1981; Zahl-

bruckner 1927).

Another species with a tendency to produce clustered

apothecia is Sarcogyne austroafricana (Zahlbr.) Magn.,

but this is a true Sarcogyne with very small ellipsoid

pycnidiospores, 2—2,5 x 1 /xm in size (not present in

original description). The apothecia of S. austroafricana

are also larger [about the size of S. clavus (Ram.) Krem-

pelh.] and the clusters are composed of only a few apo-

thecia that are only slightly distorted. The hypothecia of

S. clavus and S. austroafricana are a dirty brown (Mag-

nusson 1936, 1937; Zahlbruckner 1926), whereas that of

L. aggregata is at most stramineous.

The photobiont of Lithoglypha aggregata is a species

of Myrmecia, which is also typical for Sarcogyne

(Tschermak-Woess 1978).

This new lichen is presently known only from expo-

sures of the Clarens Formation, otherwise known as

Cave Sandstone, more or less encircling Lesotho. The

two known records come from both sides of Lesotho but

in the northern half, which may indicate its absence in

the south-west.

O.F.S. — 2828 (Bethlehem): Clarens, on rocks (-CB), J. E. van der

Plank 1896, 1943.07. (PRE).

REFERENCES

MAGNUSSON, A. H. 1935. On the species of Biatorella and Sarco-

gyne in America. Annales de Cryptogamie Exotique 7: 115-146.

MAGNUSSON, A. H. 1936. Acarosporaceae und Thelocarpaceae.

Rabenhorst’s Kryptogamen-Flora, edn2, 9,5, 1: 1-318.

MAGNUSSON, A. H. 1937. Additional notes on Acarosporaceae.

Acta Horti Gotoburgensis 12: 87- 103.

POELT, J. 1969. Bestimmungsschliissel europaischer Flechten, pp. 71

+ 757. Cramer, Lehre.

POELT, J. & VIiZDA, A. 1981. Bestimmungsschliissel europaischer

Flechten, Erganzungsheft 2: 248-249. Cramer, Vaduz.

TSCHERMAK-WOESS, E. 1978. Myrmecia reticulata as a phyco-

biont and free-living — free-living Trebouxia — the problem of Ste-

nocybe septata. The Lichenologist 10: 69-79.

VEZDA, A. 1978. Neue oder wenig bekannte Flechten in der Tsche-

choslowakei. II. Folia Geobotanica et Phytotaxonomica 13:

397-420.

VOSS, E. G. etal.(t ds) 1983. International Code of Botanical Nomen-

clature. Regnum Vegetabile 111:1 -472.

ZAHLBRUCKNER, A. 1926. Afrikanische Flechten (Lichenes).

Botanische Jahrbiicher 60: 468-552.

ZAHLBRUCKNER, A. 1927. Catalogus lichenum universalis 5:

9-27. Bomtraeger, Leipzig.

F. BRUSSE

MS. received: 1987.07.13.

ADIANTACEAE/PTERIDACEAE

DORYOPTERIS PILOSA, A NEW RECORD FOR SOUTH AFRICA

On the 25th October 1980, the junior author collected

the fern specimen Jacobsen 5252 in the Penge area of the

eastern Transvaal (Quarter degree sheet 2430 AD at an

altitude of 840 m) in the shade of dry scrub, where the

plant grew associated with Ceterach cordatum (Thunb.)

Desv. The plant is hairy, 260 mm tall and displays strong

dimorphism, with long fertile fronds and short sterile

ones (Figure 3). At that time it was examined only

Bothalia 18,1 (1988)

casually by the senior author and was filed away

amongst plants ascribed to the Doryopteris concolor

(Langsd. & Fish.) R. & A. Tryon complex.

In January 1987, Mrs Susan Strauss of the Fern

Society of Southern Africa sent a fragmentary specimen

of a hairy fern to the senior author, consisting of a long-

stalked fertile frond, 150 mm high, and a very short 50

mm high sterile frond. The plant obviously belonged to

the genus Doryopteris. It was found in the Thabazimbi

District, in the Kransberg area on the farm Geelhout-

bosch 269 KQ (2427 BC) at an altitude of 1 350 m. The

little clump of plants grew together with Cheilcmthes

concolor , Ceterach cordatum and Anemia simii Tard.

On checking through the senior author’s herbarium it

was found that Jacobsen 5252 and Susan Strauss s.n. are

specimens of a species distinct from Cheilanthes conco-

lor. They were identified as Doryopteris pilosa (Poir.)

Kuhn. This species is described and illustrated in Tar-

dieu-Blot (1958): 146, Figure 21, 3-5. As it has so far

not been described in the comprehensive literature on

southern African ferns (Jacobsen 1983; Schelpe &

Anthony 1986), its synonyms and a short description are

inserted here (mainly after Tardieu-Blot, 1958).

FIGURE 3. — Doryopteris pilosa from Penge, eastern Transvaal, x

0,4.

Doryopteris pilosa (Poir.) Kuhn in C. C. von der

Decken’s Reisen in Ost Afrika in 1859-1861, 3,3:63

(1879). Pteris pilosa Poir. in Lam.: 717 (1804). Pellaea

pilosa Hook.: t. 1 12A (1858). Allosorus pilosus Kuntze:

806 (1891). Cheilanthes heterophylla Willd. in Kaulf.:

120 (1824). Pellaea hirtula C.Chr.: 196 (1925).

91

Rhizome short, with tufted fronds and linear clathrate

scales, composed of elongate cells. Sterile fronds with

very short stipe 20—30 mm long, bearing pale, thin and

narrow, reddish scales; lamina 30—40 mm long and

broad, triangular, hirsute on both surfaces, palmately

lobed, with the basal lobe basiscopically developed and

sinuately lobed, upper lobes entire, rounded. Stipe of

fertile fronds 180—200 mm long, dark brown, shiny,

scaly at the base, but increasingly beset upwards with

stringy, adpressed matted hairs; lamina pedate-trian-

gular, 60—120 mm long, 2-pinnate to 3-pinnatifid, ulti-

mate lobes incised to % of their length, the lowest basal

pinnules strongly basiscopically developed, the upper

pinnae decurrently lobed; lower surface pilose with

whitish or reddish hairs and narrowly lanceolate, caudate

scales on costae and rhachis; costae dark brown and

shiny for half the length of the pinnae. Sori continuous

with a scarious, narrow, slightly fringed indusium; spo-

rangia long-stalked (fide Tryon 1942); spores globular

tetrahedral with convex faces and short, irregularly

reticulate cristae (fide Tardieu-Blot 1958).

In view of some recent nomenclatural changes within

the genus Doryopteris , which resulted in the transfer of

some species to the genus Cheilanthes , the authors have

paid particular attention to the characteristics of this

species. They are of the opinion that due to the strong

dimorphism, the pedate, non-multipinnate structure of

the fertile leaf (Tryon & Tryon 1982) and the long-

stalked sporangia (Tryon 1942), the species should

remain within the genus Doryopteris.

The species was so far known only from Madagascar

(Tardieu 1958), Mauritius, Reunion (Tardieu-Blot

1960), the Congo Republic and Zaire (Tardieu-Blot

1953). The two widely separated localities in the

Transvaal are new records (Figure 4). The distribution

pattern is entirely disjunct and the ecology is somewhat

puzzling. The locality in the Congo Republic is given as

near Djambala, about 180 km north of Brazzaville, i.e.

in an area of tropical rain forest with an annual rainfall of

1 200— 1 600 mm, whereas all the other occurrences are

apparently in drier areas. Tardieu-Blot (1958) however,

mentions ‘foret tropophile et bush xerophile’ for the lo-

calities in the relatively dry south-west of the Malagasy

FIGURE 4. — The distribution of Doryopteris pilosa.

92

Bothalia 18,1 (1988)

Republic. Those on Reunion and Mauritius are concen-

trated on the drier south-western rain shadow side of the

islands, where the annual rainfall is between 475 and 890

mm (White 1983), with a very distinct dry season.

The South African localities seem to have a climate

similar to that of south-western Madagascar (400-500

mm annually). The plants grow on south facing rocky

slopes, in the shade or half-shade of scrub. Their appear-

ance is xeromorphic.

The very distinct disjunct distribution pattern is not

easily understood. It is comparable to that of other ferns

such as Microsorium pappei (Mett. ex Kuhn) Tard.

(Madagascar, Tanzania, Zimbabwe, Natal), Asplenium

christii Hieron. (Kenya/Tanzania, Zimbabwe/Mozam-

bique and Zululand), Asplenium platyneuron (L.) Oakes

(North America, South Africa), Elaphoglossum con-

forme (Swartz) J. Sm. (St Helena, Liberia, tropical east

African mountains, south-western Cape) and Doryop-

teris callolepis (East African mountains, Zimbabwe,

south-western Cape), to name but a few.

In a number of occurrences very specific edaphic

and/or climatic conditions may be responsible, other

records may represent relics of a former, wider distribu-

tion. The discovery of exotic invaders in a number of

oases with totally sporadic distribution patterns, how-

ever, seems to point more and more to completely

chance settlement, facilitated by the minute spores

readily distributed by wind. Such sporadic distribution

records are known for Lunathyrium japonicum (Thunb.)

Kurata, Adiantum hispidulum Swartz, Macrothelypteris

torresiana (Gaud.) Ching and others. In many instances

more complete records, supported by new finds, may

close such distribution gaps or at least make the explana-

tion of these sporadic occurrences more credible.

ACKNOWLEDGEMENTS

The assistance of Miss J. Matthews with the illustra-

tions is gratefully acknowledged. N. H. G. Jacobsen

thanks the Director of Nature Conservation, Transvaal,

for permission to publish this paper.

REFERENCES

CHRISTENSEN, C. F. A. 1925. Pellaea hirtula. Supplement (ad

Bonapart: Fougeres de Madagascar). Hagerup, Copenhagen.

HOOKER, W. J. 1858. Species ftlicum. Pamplin, London.

JACOBSEN, W. B. G. 1983. Ferns and fern allies of southern Africa.

Butterworths, Durban, Pretoria.

KAULFUSS, G. F. 1824. Enumeratio ftlicum. Leipzig.

KUHN, F. A. M. 1879. Doryopteris pilosa. In C. C. von der Decken,

Reisen in Ost Africa in 1859 - 1861 . Leipzig & Heidelberg.

KUNTZE, C. E. O. 1891 . Revisio generum plantarum 2. Leipzig.

LAMARCK, J. B. A. P. M. DE 1804. Encyclopedic methodique.

Botanique 5. Paris.

SCHELPE, E. A. C. L. E. & ANTHONY, N. C. 1986. Flora of

southern Africa. Pteridophyta. Botanical Research Institute, Pre-

toria.

TARDIEU-BLOT, M. L. 1953. Les pteridophytes de l’Afrique Inter-

tropicale Francaise. Memoires de I'Institut francais d’Afrique

noire 28:1-241.

TARDIEU-BLOT, M. L. 1958. Polypodiacees I. In H. Humbert,

Flore de Madagascar et des Comores. Firmin Didot, Paris.

TARDIEU-BLOT, M. L. 1960. Les fougeres des Mascareignes et des

Seychelles. Notulae Systematicae 16, 1 — 2: 151—201.

TRYON. R. M. 1942. A revision of the genus Doryopteris. Contribu-

tions from the Gray Herbarium of Harvard University 143: 3 — 80.

TRYON. R. M. & TRYON, A. F. 1982. Ferns and allied plants.

Springer Verlag, Stuttgart.

WHITE, F. 1983. The vegetation of Africa. UNESCO, Paris.

W. B. G. JACOBSEN* and N. H. G. JACOBSEN**

* P.O. Box 1178, White River 1240.

** Division of Nature Conservation, Private Bag X209, Pretoria 000 1 .

MS. received: 1987.06.02.

LAMIACEAE

RESUSCITATION OF SYNCOLOSTEMON RAMULOSUS E. MEY. EX BENTH.

Three closely related species of Syncolostemon , S.

densiflorus Benth., S. ramulosus E. Mey. ex Benth. and

S. rotundifolius E. Mey. ex Benth., were described by

Bentham ( 1838), all based on Drege specimens collected

in the Transkei during Dr Andrew Smith's expedition to

Natal in 1832.

In my treatments of the genus (Codd 1976, 1985), S.

densiflorus and S. rotundifolius were recognized as dis-

tinct species, well documented by modern material, but

5. ramulosus remained something of a mystery. In the

shape of the calyx teeth (probably the most reliable char-

acter for separating the three species, as indicated

below), S. ramulosus is somewhat intermediate between

the other two and, being nearer to S. densiflorus in this

respect, was placed under the latter species as a syno-

nym.

In recent years, as a result of thorough collecting in

the area adjoining the Umtamvuna River by Mr A. T. D.

Abbott, who farms near Port Edward, and Prof. A. E.

van Wyk of Pretoria University, it has become apparent

that this view is incorrect. My awareness of this was

alerted by two specimens collected in April 1986 by Mr

Abbott on the farm Clearwater near Port Edward, Nos

3101 and i/02, which he maintained were distinct spe-

cies, though both would run to S. rotundifolius in my

key. On closer examination. No. 3101 was found to

agree with the type material of S. rotundifolius whereas

No. 3102 matched the type material of S. ramulosus.

Certain other small differences were noted and it was

realised that the following specimens, previously in-

cluded in S. rotundifolius , were in fact 5. ramulosus ,

which should be regarded as a distinct species: Acocks

1333 f, Nicholson 1419 (from near Port Edward, Natal);

and Strey 8638 from Umtentu, Transkei). The following

are the main distinguishing characters.

S. rotundifolius. Shrub 0,8- 1,8 m tall. Leaf blade

obovate to subrotund, 6-15 x 4-12 mm. Inflorescence

Bothalia 18,1 (1988)

93

medium-dense to lax; verticils 2-flowered. Calyx puber-

ulous to shortly glandular-pubescent; teeth deltoid, sub-

equal, 1,5-2 mm long. Corolla bright mauve or

magenta-pink to pink, 20-27 mm long, 15 mm wide at

the mouth; lower lip 8-10 mm long. Stamens exserted

by 10— 15 mm.

S. ramulosus. Shrub 0,6— 1,2 m tall. Leaf blade obo-

vate to rotund, 5-7 x 3,5-6 mm. Inflorescence

medium-dense; verticils (2-) 4-6-flowered. Calyx

glandular-hispidulous; teeth lanceolate-subulate 2-2,5

mm long, the uppermost tooth usually slightly shorter

and broader than the lower 4. Corolla lilac to pale

pinkish mauve or occasionally white, 16—20 mm long, 5

mm wide at the mouth; lower lip 3 mm long. Stamens

exserted by 5—6 mm.

S. densiflorus. Shrub 1—2,2 m tall. Leaf blade

broadly elliptic to broadly ovate or rotund, 5-15 x

4- 10 mm. Inflorescence dense; verticils 4-6-flowered,

Calyx glandular-hispidulous; teeth lanceolate-subulate,

2,5 — 3 mm long, the uppermost tooth usually shorter

than the lower 4. Corolla crimson to pink or rarely

whitish, 18 — 23 mm long, 8—12 mm wide at the mouth;

lower lip 5 mm long. Stamens exserted by 8— 10 mm.

REFERENCES

BENTHAM, G. 1838. In E. H. F. Meyer , Commentariorum de plantis

Africae australioris 2: 231 . Konigsberg.

CODD, L. E. 1976. The genus Syncolostemon. Bothalia 12: 21-27.

CODD, L. E. 1985. Syncolostemon. Flora of southern Africa 28,4:

185-191.

L. E. CODD

PORPIDIACEAE

A NEW SPECIES OF PORPIDIA FROM THE DRAKENSBERG

Porpidia afromontana Brusse, sp. nov.

Thallus crustosus, basalticola, fuscus, areolatus, ad 80

mm diametro, 150—800 yum crassus. Areolae 0,1 — 1,0

mm diametro, fuscae, marginibus albidis, non soredia-

tis. Prothallus ater. Stratum epinecrale hyalinum,

10—45 yu m crassum. Cortex superior 15 — 35 yum cras-

sus, paraplectenchymatus, cellulis 3, 5 -8,0 pm diame-

tro, hyalinus, sed supemis 5-8 pm badius. Stratum

gonidiale 40-60 pm crassum; algae 5-27 pm diame-

tro, Chlorococcaleae. Medulla alba, 50—650 pm crassa.

Apothecia atra, 0,2-0, 8 mm diametro, immersa, plana

vel concava. Excipulum reductum, 5—15 pm crassum,

interne et infeme hyalinum, supeme viridi-atrum et para-

plectenchymatum. Hypothecium hyalinum vel strami-

neum, 40—70 yum crassum, paraplectenchymatum, cel-

lulis 3-5 yum diametro. Hymenium hyalinum, 100 pm

altum; epihymenium atrum, 5—10 pm crassum, granu-

lare. Paraphyses graciles, septatae, ramosae et anasto-

mosantes, 1, 2-2,0 pm crassae, capitatae, capitibus

2,0— 3,0 pm crassis. Asci clavati, 80—90 x 25-30 pm,

tholis J+ perpallide caeruleis, cylindris axialibus J +

caeruleis (Figure 5). Ascosporae sextae vel octonae,

hyalinae, simplices, late ellipsoideae, halonatae, 17—22

x 8 — 1 1 ,5 pm. Pycnidia non visa. Thallus acidum con-

fluenticum continens.

TYPE. — Cape, 3028 (Matatiele): 65 km N of

Maclear, summit of Naude’s Nek, on basalt rocks on

summit plateau, in full sun, alt. 2 500 m (-CA). F.

Brusse 4590, 1986.01.26 (PRE, holo.; BM, COLO, E,

LD, UC, UPS, iso.). Figure 6.

Thallus crustose, on basalt, brown, areolate, to 80 mm

across, 150-800 gm thick. Areoles 0,1 -1,0 mm

across, brown, margins whitish, not sorediate. Prothal-

lus black. Epinecral zone hyaline, 10—45 yum thick.

Upper cortex 15-35 yum thick, paraplectenchymatous,

cells 3, 5-8,0 yum diam., hyaline, but upper 5-8 yum

reddish brown. Algal layer 40-60 yum thick; algae

5-27 yum diam., Chloroccocalean. Medulla white (or

rusty-coloured due to mineral inclusions from weather-

ing of basalt), 50—650 yum thick. Apothecia black.

FIGURE 5. — Porpidia afromontana Brusse, ascus and paraphyses. F.

Brusse 4590, holotype. Bar = 10 yxm.

94

Bothalia 18,1 (1988)

FIGURE 6. — Porpidia afromontana Brusse, habit. F . Brusse 4590 ,

holotype. Scale in mm.

0,2-0, 8 mm across, immersed, flat or concave. Exciple

reduced (developed only on flanks), 5-15 /xm thick,

hyaline within and below, greenish black and paraplec-

tenchymatous above. Hypothecium hyaline or strami-

neous, 40-70 pm thick, paraplectenchymatous, cells

3-5 pm diam. Hymenium hyaline, 100 pm high; epihy-

menium black, 5-10 pm thick, granular. Paraphyses

slender, septate, branched and anastomosed, 1, 2-2,0

pm thick, capitate, heads 2-3 pm thick. Asci clavate,

six or eight-spored, 80-90 x 25-30 pm, tholus J +

very pale blue, with J+ blue axial cylinder (Figure 5).

Ascospores hyaline, simple, broadly ellipsoid, halonate,

17—22 x 8—11,5 pm. Pycnidia not seen. Chemistry:

confluence acid present.

Etymology: afromontana (Latin) = pertaining to, or

from African mountains, hence afromontane.

This new species is a typical Porpidia except for the

pale hypothecium, which makes it assignable to Poeltia-

ria Hertel (1984). However, this genus is based on a

pale, rather than a dark hypothecium, as is typical for

Porpidia, but this in itself, cannot be significant at the

generic level. The brown pigments produced in lichens

are probably all the products of oxidative coupling of

phenols derived from polyketides, similar to the 1,8-

dihydroxynaphthalene oxidative coupling demonstrated

in some ascomycetes and ascomycete anamorphs

(Wheeler 1983). The amount of genetic material (DNA)

involved in this pigment production is too limited to be

of significance at the generic level, and a single locus is

probably involved in its production. It is also possible

* Porpidia urbanskyana (Zahlbr.) Brusse, comb. nov. Basionym:

Lecidea urbanskyarui Zahlbr., Deutsche Siidpolar-Expedition

1901-1903 8: 38 (1906). Synonym: Poeltiaria urbanskyana (Zahlbr.)

Hertel, Beihefte zur Nova Hedwigia 79: 432 (1984).

** Porpidia turgescens (Koerb.) Brusse, comb. nov. Basionym: Leci-

della turgescens Koerb., Abhandlungen der Schlesischen Gesellschaft

fur vaterlandische Cultur 2: 34 (1862). Synonym: Poeltiaria turgescens

(Koerb.) Hertel, Beihefte zur Nova Hedwigia 79: 431 (1984).

that the phenomenon of a dark hypothecium, is simply a

question of control mechanisms, which control the ex-

pression of the brown pigment in a particular anatomical

region (in this case the hypothecium). For instance,

brown pigments are often expressed in cortical,

excipular, epihy menial, and hypothecial tissue, but not

in medullary and rarely in hymenial tissue.

Porpidia afromontana is closest to Porpidia

urbanskyana (Zahlbr.) Brusse*, but has a much thinner

exciple (5—15 pm) than the latter (40—60 pm\ Hertel

1984). The apothecia of P. afromontana are also sunken

in the areoles, whereas those of P. urbanskyana are

adnate and larger (to 1,2 mm across). The thallus of P.

afromontana is also much thicker (to 800 pm thick),

brown-pigmented, and contains confluentic acid,

whereas that of P. urbanskyana is thin (rarely thicker

than 100 pm), is grey-coloured (lacks brown pigments in

the upper cortex), and contains traces of stictic acid

(Hertel 1984).

The species of Porpidia with brown thalli have

recently been thoroughly revised by Schwab (1986) for

central and northern Europe, but these lichens all have

brown hypothecia, a character that is significant at the

species level. Porpidia has also received attention in

central, southern and eastern Asia, including the Hima-

laya mountain range (Hertel 1977).

Confluentic acid is well known in Porpidia, being

known in P. flavocoerulescens (Homem.) Hertel &

Schwab, P. tuberculosa (Sm.) Hertel & Knoph, P. tur-

gescens (Koerb.) Brusse** (with 2'-0-methylperlatolic

acid) and is also known in Xenolecia spadicomma (Nyl.)

Hertel, a lichen with a similar apothecial structure to P.

afromontana. Since pycnidiospores were not found in P.

afromontana, it is not possible to exclude the possibility

that it belongs to Xenolecia Hertel (1984).

At present Porpidia afromontana is known only from

the type locality, the summit of Naude’s Nek, at 2 500 m

altitude, in the Drakensberg, on basalt.

REFERENCES

HERTEL, H. 1977. Gesteinsbewohnende Arten der Sammelgattung

Lecidea (Lichenes) aus Zentral-, Ost- und Sudasien, Eine erste

Ubersicht. Khumbu Himal 6,3: 200-225.

HERTEL, H. 1984. Uber saxicole, lecideoide Flechten der Subantark-

tis. Beihefte zur Nova Hedwigia 79: 399-499.

SCHWAB, A. J. 1986. Rostfarbene Arten der Sammelgattung Lecidea

(Lecanorales). Revision der Arten Mittel- und Nordeuropas. Mit-

teilungen aus der Botanischen Staatssammlung, Miinchen 22:

221-476.

WHEELER, M. H. 1983. Comparisons of fungal melanin biosynthesis

in ascomycetous, imperfect and basidiomycetous fungi. Transac-

tions of the British Mycological Society 81, 1 : 29-36.

F. BRUSSE

MS. received: 1987.08.13.

SC HIZODISC US , A NEW PORPIDIOID LICHEN GENUS FROM THE DRAKENSBERG

Schizodiscus Brusse, gen. nov.

Thallus crustaceus, saxicola. Cortex superior para-

plectenchymatus, cellulis 2,5— 4,0 pm diametro. Algae

Chlorococcaleae. Apothecia atra, immersa. Excipulum et

hypothecium ut in Lecidea. Paraphyses simplices vel

furcatae, conglutinatae, septatae. Asci clavati, non-halo-

nati, tholis J+ caeruleis ut in iconem (Figura 7) illu-

stratis. Ascosporae octonae, hyalinae, simplices, ellip-

soideae, non-halonatae. Pycnidia immersa, globosa vel

oblonga. Pycnidiosporae acrogenae, bacillares, hyali-

nae, rigidae, rectae.

Bothalia 18,1 (1988)

95

TYPE. — Schizodiscus afroalpinus Brusse

Thallus crustose, saxicolous. Upper cortex paraplec-

tenchymatous, cells 2, 5-4,0 pm diam. Algae Chloro-

coccalean. Apothecia black, immersed. Exciple and

hypothecium as in Lecidea. Paraphyses simple or

branched, septate, conglutinate. Asci clavate, eight-

spored, non-gelatinized, tholus J+ blue as in Figure 7.

Ascospores hyaline, simple, ellipsoid, non-halonate.

Pycnidia immersed, spherical to globose. Pycnidio-

spores acrogenous, straight, hyaline, rigid rods.

Etymology: the generic name is derived from the

Greek ‘schizo’ meaning split and ‘discos’ meaning disc,

in allusion to the apothecia which develop fissures and

cracks with age.

FIGURE 7. — Schizodiscus afroalpinus Brusse,

ascus and paraphysis. F. Brusse 4523 , holo-

type. Bar = 10 jum.

Schizodiscus afroalpinus Brusse, sp. nov.

Thallus crustosus, saxicola, caesio-cinereus, usque ad

40 mm diametro, 0,2-1 mm crassus, continuus et

rimosus, vel rimoso-areolatus, vel interdum dispersus,

areolis (0,2-) 0, 5-4,0 mm diametro, planis vel inter-

dum convexis, isidiis sorediisque destitutis. Cortex supe-

rior 17-20 pm crassus, paraplectenchymatus, cellulis

2, 5-4,0 pm diametro. Stratum gonidiale 50-70 gm

crassum, algis chlorococcaleis, 5—15 pm diametro.

Medulla alba, J — , 150-800 pm crassa. Apothecia atra,

innata, interdum emergentia, plana, lecideina, usque ad

1,5 mm diametro, plerumque cum fissuris cincta,

primum integra deinde fissurata. Excipulum infeme

brunneum, circa 10 pm crassum, periclinate prosoplec-

tenchymatum, supeme circa 20 pm crassum, atrovirens,

paraplectenchymatum. Hypothecium stramineum vel

pallide badium, 50—100 pm crassum, granulis insper-

sum, paraplectenchymatum, cellulis 3,0— 5,5 //.m

diametro, supeme (subhymenium) cum guttulis insper-

sum. Hymenium hyalinum (vel pallide viride), 70-80

pm altum, J+ caeruleum, epihymenio fusco-brunnes-

centi. Paraphyses simplices vel furcatae, septatae,

ecapitatae, conglutinatae, lumenibus 1 — 1,5 pm crassis,

gelatinis J+ pallide caeruleis. Asci clavati vel subcylin-

drici, non-halonati, tholis J+ caeruleis ut in iconem

(Figura 7) illustratis. Ascosporae octonae, hyalinae, sim-

plices, ellipsoideae, ehalonatae, 10,0-17,5 x 6, 0-7, 5

pm. Pycnidia (in typo non visa) immersa, globosa vel

oblonga, saepe aggregata, 100-120 pm profunda,

70—100 /urn lata. Pycnidiosporae acrogenae, bacillares,

hyalinae, rectae, rigidae, 6,5-15 x 0,8 pm. Thallus

acidum 2'-0-methylperlatolicum solum continens.

TYPE. — 2828 (Bethlehem): 31 km S of Phuthadi-

tjhaba (Witsieshoek), summit of Western Buttress

(Mont-aux-Sources), on low basalt exposures near soil

near seepage areas, on gentle S slope on summit plateau,

alt. 3 080 m (-DB), F. Brusse 4523, 1986.01 .21 (PRE,

holo.; BM, LD, iso.). Figure 8.

FIGURE 8. — Schizodiscus afroalpinus Brusse, habit. F. Brusse 4523 ,

holotype. Scale in mm.

Thallus crustose, saxicolous, pale blue-grey, up to 40

mm across, 0,2- 1 mm thick, continuous and rimose, to

rimose-areolate, to sometimes dispersed, areoles (0,2-)

0, 5-4,0 mm across, flat or sometimes convex, lacking

isidia and soredia. Upper cortex 17-20 pm thick, para-

plectenchymatous, cells 2, 5-4,0 yam diam. Algal layer

50-70 yu,m, algae chlorococcalean, 5-15 pm diam.

Medulla white, J-, 150-800 pm thick. Apothecia

black, innate sometimes emergent, flat, lecideine, up to

1,5 mm diam., often surrounded by a fissure, at first

entire then becoming fissured (starting from the interior,

not from the margins). Exciple brown, periclinally pro-

soplectenchymatous and about 10 pm thick below, be-

coming dark green, paraplectenchymatous and about 20

pm thick above (at the flanks of the hymenium). Hypo-

thecium stramineous or pale reddish brown, 50- 100 pm

thick, granular inspersed, paraplectenchymatous, cells

96

3,0— 5,5 /x m diam., becoming inspersed with ‘oil-drop-

lets’ above (in the subhymenial zone). Hymenium hya-

line (or pale green in thick sections), 70-80 /xm high,

J+ blue, epihymenium fuscous brown. Paraphyses

simple to branched, septate, ecapitate, conglutinate, lu-

mens 1 — 1,5 /xm thick, gelatin J+ pale blue. Asci clavate

to subcylindrical, eight- spored, non-halonate (no gelati-

nous sheath or excess present), tholus J+ blue as illus-

trated in Figure 7. Ascospores hyaline, simple, ellipsoid,

non-halonate, 10,0—17,5 x 6,0— 7,5 /xm. Pycnidia (not

seen in holotype) immersed, globose or oblong, often

aggregated (sometimes in darkened verrucae), 100-120

/xm deep, 70—100 /xm wide. Pycnidiospores acroge-

nous, hyaline, rigid, straight rods or needles, 6,5-15 x