Bothalia

’N TYDSKRIF VIR PLANTKUNDiGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 17,2 Oct./Okt. 1987

rilen^Ur^

PUBLICATIONS OF THE BOTANICAL RESEARCH INSTITUTE

PUBLIKASIES VAN DIE NAVORSINGSINSTITUUT VIR PLANTKUNDE

Obtainable from the Division of Agricultural Information, De-

partment of Agriculture and Water Supply, Private Bag X144,

Pretoria 0001, Republic of South Africa. A current price list of

all available publications will be issued on request.

BOTHALIA

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

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

house journal of the Botanical Research Institute is devoted to

the furtherance of botanical science. The main fields covered

are taxonomy, ecology, anatomy and cytology. One or two parts

of the journal and an index to contents, authors and subjects are

published annually.

Verkrygbaar van die Afdeling Landbou-inligting, Departement

van Landbou en Watervoorsiening, Privaatsak X144, Pretoria

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

bare publikasies kan aangevra word.

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

Minister en Minister van Landbou van die Unie van Suid-Afrika.

Hierdie lyfblad van die Navorsingsinstituut vir Plantkunde isgewy

aan die bevordering van die wetenskap van plantkunde. Die hoof-

gebiede wat gedek word, is taksonomie, ekologie, anatomie en

sitologie. Een of twee dele van die tydskrif en ’n indeksvan die

inhoud, outeurs en onderwerpe verskyn jaarliks.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The memoirs are individual treatises usually of an ecological ’n Reeks van losstaande omvattende verhandelings oor vemaam-

nature, but sometimes dealing with taxonomy or economic lik ekologiese, maar soms ook taksonomiese of plantekonomiese

botany. onderwerpe.

THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA

This serial presents colour plates of African plants with ac-

companying text. The plates are prepared mainly by the artists

at the Botanical Research Institute. Many well-known botanical

artists have contributed to the series, such as Cythna Letty (over

700 plates), Kathleen Lansdell, Stella Gower, Betty Connell,

Peter Bally and Fay Anderson. The Editor is pleased to receive

living plants of general interest or of economic value for

illustration.

Two parts of ten plates each are published annually. A volume

consists of four parts. The publication is available in English and

Afrikaans.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaan-

de teks. Die skilderye word meestal deur die kunstenaars van die

Navorsingsinstituut vir Plantkunde voorberei. Talle bekende bota-

niese kunstenaars het tot die reeks bygedra, soos Cythna Letty

(meer as 700 plate), Kathleen Lansdell, Stella Gower, Betty Con-

nell, Peter Bally en Fay Anderson. Die Redakteur verwelkom

lewende plante van algemene belang of ekonomiese waarde vir

afbeelding.

Twee dele, elk met tien plate, word jaarliks gepubliseer. ’n Volume

bestaan uit vier dele. Die publikasie is beskikbaar in Afrikaans en

Engels.

FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA

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

Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, South

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

beskrywings van families, genusse, spesies, infraspesifieke taksons,

sleutels tot genusse en spesies, sinonimie, literatuur, verwysings

na enkele eksemplare, asook beknopte taksonomiese en ekologiese

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:

A. A. Balkema Marketing, Box/Posbus 317, Claremont 7735, RSA.

PALAEOFLORA OF SOUTHERN AFRICA / PALAEOFLORA VAN SUIDELIKE AFRIKA.

A palaeoflora on a pattern comparable to that of the Flora of

southern Africa. Much of the information is presented in the

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 fotograflese 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

South Africa

Republiek van

Suid-Afrika

BOTH ALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 17,2

Editor/Redakteur: O. A. Leistner

Editorial Board/Redaksieraad

D. F. Cutler

B. de Winter

D. J. B. Killick

O. A. Leistner

P. H. Raven

J. P. Rourke

M. J. Werger

Royal Botanic Gardens, Kew, UK

Botanical Research Institute, Pretoria, RSA

Botanical Research Institute, Pretoria, RSA

Botanical Research Institute, Pretoria, RSA

Missouri Botanical Garden, St Louis, USA

National Botanic Gardens, Kirstenbosch, RSA

University of Utrecht, Utrecht, Netherlands

ISSN 0006 8241

Published by the Botanical Research Institute, Department of Agriculture and Water Supply, Private Bag X101,

Pretoria 0001, South Africa

Uitgegee deur die Navorsingsinstituut vir Plantkunde, Departement van Landbou en Watervoorsiening,

Privaatsak X101, Pretoria 0001, Suid-Afrika

1987

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume1717unse_0

CONTENTS - INHOUD

Volume 17,2

1. Studies in the Ericoideae. V. The genus Coilostigma (Ericaceae). E. G. H. OLIVER 163

2. Synopsis of the genus Salix (Salicaceae) in southern Africa. K. L. IMMELMAN 171

3. Notes on African plants:

Acarosporaceae. A new yellow Acarospora (Lichenes) from the Waterbcrg, South West

Africa/Namibia. F. BRUSSE 179

Cucurbitaceae. Orthographicambiguityclarified. B. D. SCHRIRE 181

Fabaceae. A new species of Rhynchosia from the Richtersveld. G. GERMISHUIZEN 181

Lichenes. Psathyrophlyctis, a new lichen genus from southern Africa. F. BRUSSE 182

Liliaceae. Notes on Kniphofia. L. E. CODD 185

Polygonaceae. A new variety of Oxygonum datum. G. GERMISHUIZEN 185

Trapeliaceae. A new species of Trapelia (Lichenes) form southern Africa. F. BRUSSE 187

Zygophyllaceae. A new species of Zygophyllum from southern Africa. E. RETIEF 189

4. Taxonomy of the genus Ehrharta (Poaceae) in southern Africa: the Villosa group. G. E. GIBBS

RUSSELL 191

5. Leaf anatomy of the genus Ehrharta (Poaceae) in southern Africa: the Villosa group. R. P. ELLIS . 195

6. PhytogeographyofthesubtribeLeipoldtiinae(Mesembryanthemaceae). H. E. K. HARTMANN .... 205

7. Preliminary floristic analysis of the major biomes in southern Africa. G. E. GIBBS RUSSELL 213

8. ’n Kontrolelys van varings en blomplante van die Wonderkloofnatuurreservaat, Transvaal, Suid-

Afrika. J. P. KLUGEenC. DYER 229

9. A checklist of vascular plants of the Amatole Mountains, eastern Cape Province/Ciskei. P. B.

PHILLIPSON 237

10. Miscellaneous notes:

Chromosomestudieson African plants. 5. J. J. SPIESandH. DUPLESSIS 257

11. Obituary: Rolf Dahlgren (1932-1987). A. E. VAN WYK 261

12. Obituary: John Frederick Vicars Phillips (1899-1987). EMSIE DUPLESSIS 267

13. New taxa, new records and name changes for southern African plants. G.E. GIBBS RUSSELL,

C. REID,L. FISH, G. GERMISHUIZEN, M. VAN WYK, J. VAN ROOY and STAFF 269

14. ReviewoftheworkoftheBotanicalResearchlnstitute 1986/1987 277

15. Book reviews 297

Bothalia 17,2: 163-170 (1987)

Studies in the Ericoideae (Ericaceae). V. The genus Coilostigma

E. G. H. OLIVER*

Keywords: Cape Province, Coilostigma, Ericaceae, Ericoideae, morphology, phytogeography, pollination biology, taxonomy

ABSTRACT

A revision of the genus Coilostigma Klotzsch is presented which recognizes only two species, C. zeyherianum

Klotzsch and C. glabrum Benth. The genus belongs to the Ericaceae — Ericoideae and is endemic in the southern

and eastern Cape Province. Fundamental differences in the ovary complement have necessitated the recognition of

two subgenera, Coilostigma and Anomalosepala E. G. H. Oliver. Aspects covered include history, morphology,

phytogeography, pollination biology and taxonomy.

UITTREKSEL

Hierdie is ’n hersiening van die genus Coilostigma Klotzsch waarin net twee spesies, C. zeyherianum Klotzsch en

C. glabrum Benth., erkenning geniet. Die genus behoort tot die Ericaceae — Ericoideae en is endemies in die

suidelike en oostelike dele van die Kaapprovinsie. Basiese verskille van die vrugbeginsel-komplement noodsaak

die erkenning van twee subgenera, Coilostigma en Anomalosepala E. G. H. Oliver. Aspekte wat bespreek word

sluit in geskiedenis, morfologie, fitogeografie, bestuiwingsbiologie en taksonomie.

HISTORICAL OUTLINE

The genus Coilostigma was described by Klotzsch

(1838) in his major revision of the Ericoideae. He

based it on his three new species, C. tenuifolium, C.

zeyherianum and C. dregeanum , all from the eastern

Cape Province, and all possessing an unequal calyx.

Bentham (1839), in his revision of the family for

De Candolle’s Prodromus, retained the genus but

redefined it to incorporate his new species, C. gla-

brum, and Klotzsch’s monotypic genus Thamnium,

T. puberulum. He formed two sections within the

genus, namely Eucoilostigma and Thamnium. The

inclusion of the superficially similar Thamnium pu-

berulum ignored the feature of the unequal calyx for

Coilostigma. Bentham (1876) took a conservative

view of the family in Bentham & Hooker’s Genera

and sunk both Coilostigma and Thamnium under the

genus Scyphogyne Brongn. Drude (1897) followed

Bentham’s latter treatment of the genera.

In Flora capensis. Brown (1906) retained the cir-

cumscription of the genus as applied by Bentham

(1839) but ignored the sectional subdivision. He re-

moved the discordant C. puberulum to Thoraco-

sperma. All these changes were retained by Phillips

(1926) in the first edition of his Genera. Later, how-

ever (Phillips 1944), he proposed a completely dif-

ferent classification of the family in South Africa,

implemented in the second edition of his Genera

(Phillips 1951). He reduced the number of genera to

only eight, including Erica L. This action in some

cases placed totally unrelated genera together. He

placed Coilostigma under Salaxis Salisb.

MORPHOLOGY

The plants of Coilostigma are typical ericoid

woody shrublets with one species, C. zeyherianum,

being single-stemmed and the other, C. glabrum, a

multi-stemmed coppicing shrublet. Most of the other

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

vegetative characters are very similar in both

species.

The bract is totally recaulescent in C. zeyherianum

(Figure 4,1), only very rarely partially so, and forms

part of the calyx as the large abaxial member in what

is referred to as an unequally 4-lobed calyx. In C.

glabrum, on the other hand, the bract is variable in

position and with the calyx exhibits a remarkable

diversity of form. It is variably recaulescent within

any one inflorescence, with the lowest whorl of flow-

ers having a partially recaulescent bract and the up-

per whorl a totally recaulescent bract. When the

bract is partially recaulescent the calyx can be either:

(1) 4-lobed with two normally sized lateral lobes

(sepals) and very reduced ad- and abaxial lobes (se-

pals) (Figure 5,2), or

(2) 4-lobed with two lateral lobes (sepals) and an

adaxial lobe (sepal) and a larger less fused abaxial

lobe (bract) (Figure 5,1), or

(3) 3-lobed with two lateral lobes (sepals) and a

slightly reduced adaxial (sepal) and no abaxial lobe

(bract or sepal), or

(4) 2-lobed with only two normally-sized lateral

lobes (sepal), the ad- and abaxial lobes (bract and

sepal/s) being absent.

As in all species of the Ericoideae with an unequal

calyx the bracteoles are totally absent in all flowers.

This is also the case where the bract is only partially

recaulescent.

The pollen in both species occurs as single tricol-

porate grains with scabrate sculpturing in C. zeyhe-

rianum and no sculpturing in C. glabrum (Figure 1).

The ovary displays two distinct types. In C. gla-

brum it is 4-, rarely 3-locular, regular in shape and

hairy with a single ovule in each locule. In C. zeyhe-

rianum the ovary is only 2-locular, flattened and gla-

brous.

As a result of ovary characters the fruit is very

different in shape in the two species. In C. glabrum it

164

Bothalia 17,2 (1987)

FIGURE 1. — Pollen of the species of Coilostigma. 1, C. zeyherianum (Oliver 7949); 2, C. glabrum (Oliver 8801). Scanning

electron micrographs, left hand x 1000, right hand x 3000.

is regular or slightly irregular with usually only one

of the developed seeds being fertile. In C. zeyheria-

num the fruit is mostly irregular due to the develop-

ment of only one fertile seed and is only occasionally

2-seeded, and regularly complanate. In both species

the seed is extremely difficult to remove from the

fruit in the dry state. This indicates that the whole

flower is shed as the propagule and must disintegrate

or be wetted prior to germination.

The seeds are similar in both species and are typ-

ically ericoid in form, i.e. spherical with a hard re-

ticulate testa (Figure 2). This type of seed is nor-

mally associated within the Ericoideae with a dehis-

cent capsule as in the genera Erica L., Blaeria L.,

Philippia Klotzsch and Ericinella Klotzsch. The in-

dehiscent fruit and ericoid seed in this genus is

shared with genera such as Coccosperma Klotzsch,

Thamnus Klotzsch and Platycalyx N.E. Br. and can

be regarded as a stage in the evolution of the inde-

hiscent soft-seeded fruits found in the more ‘ad-

vanced’ genera within the subfamily.

GENERIC DELIMITATION AND RELATIONSHIPS

The genus Coilostigma, as recognized in this re-

vision, is characterized by the possession of a totally

recaulescent bract forming an unequally 4-lobed ca-

lyx, a 4-lobed corolla, four free stamens, bilobed

anthers, single pollen grains, a 2-, 3- or 4-locular

ovary with a single pendulous ovule in each locule,

an indehiscent dry berry and seeds with a hard testa.

With this circumscription Coilostigma is somewhat

isolated within the Ericoideae. The 2-locular 1-

seeded ovary of C. zeyherianum is a character found

in a number of genera, namely Grisebachia

Klotzsch, Eremia D. Don (pro parte), Sympieza

Klotzsch, Platycalyx, Simocheilus Klotzsch, Acroste-

mon Klotzsch and Arachnocalyx Compton, all how-

ever with an equal calyx. Only in Platycalyx is there

a possibility of a close relationship in that the fruiting

stage is a similar dry berry with hard-walled seeds. In

the other genera the evolutionary reduction has re-

sulted in soft-walled seeds.

The 4-celled 1 -seeded ovary in C. glabrum is a

character which the subgenus Anomalosepala shares

with Philippia and Ericinella, both of which have an

unequal calyx and hard-walled seeds but with many

seeds per locule in a dehiscent capsule. It is also

shared with the equal-calyxed genera Erica, Blaeria,

Eremia (pro parte ) and Thoracosperma Klotzsch.

The first two have numerous seeds per locule in a

dehiscent capsule. Eremia has a very similar ovary

and fruit but flowers totally different in appearance

apart from the equal calyx. Thoracosperma on the

other hand has rather similar looking flowers, espe-

cially in T. puberulum (cf. Historical Outline), but

with a fruit that contains soft, thin-walled seeds.

There is therefore no clear-cut relationship for this

genus within the subfamily. It has probably been

derived from some Erica-Blaeria-Philippia ancestral

stock. A detailed analysis of relationships and pos-

sible evolutionary paths will be published when the

revision of all the ‘minor’ genera of the Ericoideae

has been completed.

Bothalia 17,2 (1987)

165

FIGURE 2. — Seeds of the species of Coilostigma. 1, C. zeyherianum var. zeyherianum (Oliver 7949) ; 2, C. zeyherianum var.

tenuifolium (Oliver 7948); 3, C. glabrum (Oliver 8817). Scanning electron micrographs, left hand x 40, right hand x 200.

PHYTOGEOGRAPHY

The genus Coilostigma is endemic in the southern

and eastern parts of the Cape Province (Figure 3)

corresponding to the Langeberg and South Eastern

Phytogeographical Centres proposed by Weimarck

(1941). This falls within the limits of the Cape Floral

Region (Weimarck 1941; Goldblatt 1978; Oliver et

al. 1983). The extended distribution of C. zeyheria-

num further eastwards to areas near Alexandria and

Grahamstown makes the genus the only endemic

one in the Cape to extend beyond the strict limits of

the Cape Floral Region as defined by Goldblatt

(1978). The Grahamstown area can be regarded as a

depauperate relictual extension of the Cape Flora

proper (Bond & Goldblatt 1984).

The genus is one of only two in the Ericoideae in

the Cape Floral Region with an eastern distribution,

all the others having their main centres in the south-

western Cape. The other eastern genus is the mono-

typic Thamnus. Thoracosperma is a southern genus

only just represented in the south-western Cape by

T. puberulum.

The disjunction between the distribution ranges of

the two species (Figure 3), which is quite consider-

able in terms of Ericoideae in the Cape Floral Re-

gion, remains inexplicable. C. glabrum is very local-

ized in one population on the northern drier slopes

of the low range of hills just south of the Langeberg

near Riversdale. Here it receives a fairly high annual

rainfall of ± 800 mm which can be throughout the

166

Bothalia 17,2 (1987)

year but is mainly in the winter months. C. zeyheria-

num is mainly concentrated on the coastal plains in

the Humansdorp/Port Elizabeth area where it grows

on sandy flats with an annual rainfall of 700-900 mm

falling throughout the year. The outlying population

on the coast near the mouth of the Boesmansrivier

could well occur on sand as well as the inland popu-

lation just east of Grahamstown because the latter

area has flora with some coastal affinities (Jacot

Guillarmod pers.comm.).

POLLINATION BIOLOGY

In the light of the findings of Rebelo, Siegfried

and Oliver (1985) most of the floral features of the

two species are consistent with the anemophilous

syndrome: 1, the stigma is enlarged, subinfundibuli-

form to peltate; 2, the stamens are exserted (ex-

serted stamens were, however, found to be more

generally important in ornithophilous and entomo-

philous species within the subfamily); 3, the pollen

grains are small, non-sticky and easily shed; 4, there

is no sign of development of nectaries below the

ovary, and 5, the corolla in C. zeyherianum is incon-

spicuous, pale yellow, soon turning brown. How-

ever, the corolla in C. glabrum is dark pink, a colour

that could be expected to act as an attractant to in-

sects.

This situation is similar to that existing in Erici-

nella multiflora Klotzsch in which the pollen could

be the reward for any visiting insects attracted by the

colour of the flowers.

It is surprising that the typically anemophilous

genera, namely Philippia, Salaxis, Coccosperma,

Scyphogyne Brongn. and Nagelocarpus Bullock all

have an unequal calyx (totally recaulescent bract)

but, in contrast to our genus, they have pollen grains

in tetrads.

The occurrence of anemophily in the genus was

verified in the field when small clouds of pollen were

seen to be emitted from the plants when disturbed.

COILOSTIGMA

Coilostigma Klotzsch in Linnaea 12: 234 (1838);

Benth.: 708 (1839); N.E. Br.: 327 (1906); E. G. H.

Oliver: 437 (1975).

Salaxis sensu Phillips: 71 (1944), pro parte; Phil-

lips; 561 (1951), pro parte.

TYPE: C. zeyherianum Klotzsch (lectotype cho-

sen here).

Perennial woody shrublets, single- or multi-

stemmed, up to 1 m tall. Branches lacking infrafoliar

sterigmata or ridges. Leaves 3-nate, erect imbricate,

ericoid, sulcate, linear. Inflorescence of 1-3, rarely

4, whorls of 3-nate small flowers at the ends of

branches (mesoblasts) and lateral absolute or partial

brachyblasts scattered along the mesoblasts, occa-

sionally clustered towards the ends into compound

heads; pedicel short, relative to the flower. Bract

partially recaulescent and foliaceous in the lowest

flowers to fully recaulescent in any one inflorescence

or always fully recaulescent; bracteoles absent. Ca-

lyx unequally (2)3(4)-lobed excluding or including

the totally recaulescent bract as the abaxial member,

lateral lobes usually slightly larger, reduced abaxial

sepal sometimes present, adaxial sepal sometimes

reduced or absent, sepals Vs-lA the length of

the corolla. Corolla 4-lobed, tubular to narrowly

ovoid to urceolate, glabrous or hirsute, pale yellow

to white, or pink; lobes short, erect to slightly

spreading. Stamens 4, free, exserted or included by

abortion; filaments linear; anthers bilobed, muti-

cous, dorsifixed near the base, thecae oblong, pore

Vio -Vi the length of the theca; pollen grains

single, tricolporate. Ovary 2(3)4-locular with a sin-

gle pendulous ovule per locule, transversely broadly

obovate, complanate or globose, glabrous or pube-

rulous; style filiform, exserted; stigma broad, peltate

to subinfundibuliform with 2(3) or 4 stigmatic pro-

cesses. Fruit an indehiscent berry with a thin, rather

dry, leathery pericarp, irregularly obovoid with only

one seed developed from a 2-locular ovary or glob-

Bothalia 17,2 (1987)

167

ose from a 4-locular ovary; seeds ovoid to spherical

with a hard reticulate testa.

A genus of only two species, endemic in the south-

ern and eastern Cape Province, southern Africa.

The name is derived from the Greek coilos ‘hollow’

and stigma.

The two species exhibit distinct fundamental dif-

ferences in ovary characters necessitating recognit-

ion at subgeneric level.

KEY TO THE SUBGENERA AND SPECIES

Ovary 2- rarely 3-locular, glabrous Subgenus Coilostigma

1. C. zeyherianum

Ovary 4- rarely 3-locular, puberulous ... Subgenus Anomalosepala

2. C. glabrum

Subgenus Coilostigma

Coilostigma Klotzsch in Linnaea 12: 234 (1838);

N.E. Br.: 327 (1906), pro parte; E. G. H. Oliver:

437 (1975).

Coilostigma sect. Eucoilostigma Benth.: 708

(1839).

Scyphogyne sect. Coilostigma Benth.: 594 (1876).

Type: C. zeyherianum Klotzsch.

Shrub single-stemmed; bract totally recaulescent;

calyx unequally 4-lobed with the recaulescent bract

as the abaxial lobe and 3 sepals; ovary 2-locular;

fruit irregularly obovoid, complanate.

1. Coilostigma zeyherianum Klotzsch in Lin-

naea 12: 234 (1838); N.E. Br.: 328 (1905). Type: In

montibus ‘Van Stadensriviersberge’ Ecklon &

Zeyhers.n. (Bf, BOL!, E!, K!, LD!, MEL!, P!, S!,

UPS!, W!, Z!); idem as 296 (G!, MO!, W!). Lecto-

type (chosen here): Ecklon & Zeyhers.n. (BOL).

Erect, single-stemmed perennial shrub up to 1 m

tall. Branches subflexuose, puberulous, without ster-

igmata, bark splitting irregularly with age. Leaves

erect, imbricate, 1, 5-3,0 mm long, linear acute to

obtuse, rounded below, flat above, glabrous, edged

with a few sessile glands and some hairs; petiole 0,5

mm long, appressed, glabrous, edged with sessile

glands. Flowers 1— 6(9)( 12) at the ends of the

branches (mesoblasts) and lateral brachyblasts scat-

tered along the branches, occasionally clustered to-

wards the ends of the branches; pedicel 0,3-0, 6 mm

long, glabrous or puberulous. Bract totally recaules-

cent as abaxial lobe of the calyx. Calyx unequally 4-

lobed, joined at the base, large lobe abaxial, 0, 7-2,0

mm long and foliaceous, ± Vs as long as to equal

the length of the corolla, occasionally longer, other

lobes 0,4— 0,6 mm long, oblong to narrowly deltoid,

the laterals slighter larger and the adaxial sometimes

reduced, all glabrous to puberulous at the base and

ciliate. Corolla 1,5-2, 5 mm long, tubular to narrowly

ovoid to urceolate, tangentially complanate in the

fruiting stage, glabrous or hirsute, dirty pale yellow

to brown; lobes erect or very slightly spreading,

± V& the length of the tube. Stamens exserted or

by abortion included; filaments 1, 7-2,0 mm long,

linear, glabrous; anthers 0,5-1 ,0 mm long, thecae

oblong, dorsifixed near the base, muticous, sparsely

strigulose, pore ± Vio the length of the theca.

Ovary 2-locular, 0,5 x 0,6 mm, transversely broadly

obovate, complanate, glabrous; style filiform, ± 2,5

mm long, exserted; stigma 0,6 mm broad, peltate to

subinfundibuliform with 2 stigmatic processes. Fruit

irregularly obovoid, 0,7-0, 8 mm long; seeds ovoid to

spherical, 0,6-0, 8 mm long/diam., reticulate, cells ±

angular-circular, ridges straight. Figure 4.

A species forming erect shrublets, occurring on

sandy plains near the coast from south of Humans-

dorp eastwards to Kenton-on-Sea flowering from

January to December depending on the locality.

A taxon containing two distinct varieties which

occur growing together in most populations.

FIGURE 4. — C. zeyherianum. 1, flower; 2, bract (abaxial seg-

ment of the calyx); 3, lateral sepal; 4, anther, front, side

and back views; 5, ovary; 6, leaf; all drawn x 25 from

Oliver 7949 (STE); 7, sepals, lateral and adaxial; drawn x

25 from Ecklon & Zeyher s.n. (BOL).

KEY TO THE VARIETIES

Flowers glabrous la. var. zeyherianum

Flowers pubescent lb. var. tenuifolium

la. var. zeyherianum

Coilostigma zeyherianum Klotzsch: 234 (1838); N.E. Br.: 328

(1905). Lectotype (chosen here): Ecklon & Zeyher s.n. (BOL).

C. dregeanum Klotzsch: 235 (1838); N.E. Br.: 1127 (1909).

Type: P.b.sp., Drege 7753 (Bf, K, fragm.!; BOL, fragm.!); idem

s.n. (G!, G-DC!).

Flower with completely glabrous corolla.

Vouchers: Oliver 7936 (GRA, NY, P, PRE, S, STE); 7946

(BM, BOL, E, K. NBG, MO, PRE, STE).

lb. var. tenuifolium (Klotzsch) E. G. H. Oliver,

comb, et stat. nov. Types: In planitie inter ‘Kraka-

168

Bothalia 17,2 (1987)

kamma’ et montes ‘Vanstadensriviersberge’, Ecklon

& Zeyher s.n. (Bt, E!, S!); idem as 294 (G!,

GOET!, LD!, M!, MEL!, MO!, S!, W!, UPS!, Z!);

in sylvis 'Olifantshoek’ prope flumen ‘'Bosjesmansri-

vier’, Ecklon & Zeyher s.n. (Bt, BOL!). Lectotype

(chosen here): Ecklon & Zeyher s.n. (S).

Flower with pubescent to hirsute corolla, other-

wise as in the typical variety.

Vouchers: Oliver 7935 (GRA, NY, P, PRE, S, STE); 7945

(BM, BOL, E, K, NBG, MO, PRE, STE).

Klotzsch (1838) described three species, C. tenui-

folium, C. zeyherianum and C. dregeanum, when he

created the genus Coilostigma. These he based on

differences in corolla hairiness, sepal shape and

branch thickness. An examination of all subsequent

collections has shown the sepal shape and branch

characters to be continuously variable over the

whole distribution range and therefore unreliable for

taxonomic delimitation.

There is discontinuity in the indumentum of the

corolla where material can easily be placed in gla-

brous or pubescent to hirsute groups. During field

investigations of five widely separated populations in

the Port Elizabeth to Humansdorp area, I found that

both forms did occur together to a varying degree in

each population and that no intermediates existed. I

consider this type of discontinuity worthy of recogni-

tion at varietal level.

Even though the species is recorded as widespread

on the flats it is today by no means common. Its

habitat is being inundated by alien vegetation, in

particular the Port Jackson Willow, Acacia saligna

(Labill.) Wendl., or it is being destroyed by housing

estates or farming practices. In the Port Elizabeth

area plants are very difficult to find nowadays in

places where they must have been abundant in the

past. A small population could survive in the reserve

alongside the railway line in Walmer.

The best populations I found occurred on the flats

west of the mouth of the Slangrivier south-west of

Humansdorp, but here the recently applied agricul-

tural practice of bush-cutting for pasturage is deci-

mating the species. Only on the outcrops of low sta-

ble dunes or in the dune slacks too small to man-

oeuvre in, does the species still manage to survive

and form almost pure stands.

The populations near the mouth of the

Boesmansrivier have undoubtedly disappeared as

the sandy habitat is also ideal for farming in the area.

Reported populations at Slaaikraal and Coldsprings

west of Grahamstown could not be located. It seems

unlikely that the species should occur so far inland,

but Dr A. Jacot Guillarmod assures me that vegeta-

tion elements with coastal affinities do occur in that

area.

Subgenus Anomalosepala E. G. H. Oliver, sub-

gen. nov. a subgen. typico frutice caulibus multis,

bractea perfecte vel partim recaulescenti, calyce in-

aequaliter (2)3(4)-lobato; ovario (3)4-loculari,

fructu regulariter globoso differt.

Coilostigma sect. Thamnium (Klotzsch) Benth.:

708 (1839), pro parte.

Scyphogyne sect. Thamnium (Klotzsch) Benth.:

594 (1876), pro parte.

Coilostigma N.E. Br.: 327 (1906), pro parte; E.

G. H. Oliver: 437 (1975), pro parte.

Type: C. glabrum Benth.

Shrub multi-stemmed; bract partially to totally re-

caulescent in any one inflorescence; calyx unequally

(2)3(4)-lobed; ovary (3)4-locular; fruit globose,

regular.

2. Coilostigma glabrum Benth. in De Candolle,

Prodroinus 7: 708 (1838); N.E. Br.: 328 (1905).

FIGURE 5. — C. glabrum. 1, flower; 2, lower part of flower showing partially recaulescent bract and very reduced ab- and adaxial

sepals; 3, lower part of flower showing reduced adaxial sepal; 4, bract (abaxial segment ot the calyx); 5, sepals, lateral and

adaxial; 6, reduced adaxial sepals; 7, anther, front, side and back views; 8, ovary; 9, leaf. All drawn x 25 from Oliver 8801

(STE).

Bothalia 17,2 (1987)

169

Type: In Cape Colony, Burchell 6875 (K, holo.!;

BOL, fragm.!).

Erect multi-stemmed shrublet up to 500 mm tall,

stems arising from a woody rootstock. Branches

straight, fastigiate, slender, sparsely and shortly pu-

berulous, sterigmata absent; bark grey, splitting ir-

regularly. Leaves erect, imbricate, 2, 0-3,0 mm long,

linear to linear-lanceolate, obtuse, sulcate, rounded

below, flat above, glabrous, edged with a few sessile

glands; petiole appressed, 0,4-0, 6 mm long, ciliate

with short hairs and sessile glands. Flowers 1-9 at the

ends of branches (mesoblasts) and of lateral very

short brachyblasts often crowded towards the ends

of the branches or subverticillate; pedicel very short,

0, 1-0,4 mm long, puberulous. Bract partially recau-

lescent, 0,9-1, 2 mm long, subfoliaceous, oblong and

basal in position in the lowest flowers in any one

inflorescence to fully recaulescent, subequal and

joined to the lateral calyx lobes in the upper flowers,

glabrous, ciliate with short hairs and sessile glands.

Calyx (2)3(4)-lobed slightly joined, all equal or the 2

laterals 0,7-0, 9 mm long, narrowly ovate and the ad-

and abaxial lobes variously reduced, 0, 1-0,8 mm

long, narrowly ovate to broadly deltoid or absent, all

ciliate with short hairs and sometimes sessile glands,

glabrous but puberulous at the base, the large lobes

acute with slightly sulcate apex. Corolla 1,8-2, 4 mm

long, tubular to narrowly campanulate to narrowly

ovoid, bulging out adaxially when the sepal is ab-

sent, glabrous, pale to dark pink, lobes erect to

slightly spreading, ± Ve the length of the tube.

Stamens exserted, occasionally included by abortion;

filaments linear, 1,2 mm long, often subsigmoid just

below the anther; anthers muticous rarely minutely

decurrent-aristate, glabrous, oblong, occasionally

cuneate; thecae oblong, obtuse, 0,8-1, 2 mm long,

dorsifixed near the base, occasionally prognathous

at the base; pore ± Vi the length of the theca.

Ovary 4(3)-locular with a single pendulous ovule in

each locule, globose, 0,5 x 0,6 mm, densely puberu-

lous; style exserted, filiform, 2, 5-3,0 mm long, gla-

brous; stigma subinfundibuliform, 0,5 mm wide, gla-

brous. Fruit globose, 1,0 x 0,9 mm, puberulous;

seeds broadly ellipsoid, ± 0,8-0, 9 mm long, testa

reticulate, cells ± elongate, ridges straight. Figure 5.

A species forming compact coppicing shrublets up

to 500 mm tall; confined to a few silcrete hills near

Garcia’s Pass in the Riversdale area of the southern

Cape Province and flowering from October to Janu-

ary.

This species was until recently only known from

the type collection made in November 1814 by Bur-

chell. The population, consisting of only 24 plants on

the Kleinberg north-west of Riversdale, corresponds

very closely to the more exact locality on Burchell’s

specimens ‘between Kleine Vet River and foot of

Langeberg’. The species appears to be very rare.

The row of hills forming the ridge just south of the

Langeberg Mountains could well have additional

populations of C. glabrum on it. However, the

plants are rather inconspicuous and would seem to

flower most profusely a year or two after a fire. Be-

ing copious resprouters the plants grow quickly and

flower sooner than the surrounding reseeders and

are then more conspicuous.

The species is remarkable in the whole subfamily

for the degree of variability in the form and arrange-

ment of the calyx. Even within a single inflorescence

the calyx may vary considerably. In the genus Sym-

pieza the calyx may vary from 2-4-lobed in a single

inflorescence, but in that genus there is no partially

to fully recaulescent bract to complicate the issue.

The above variability within the calyx is confined

to C. glabrum and does not occur in the commoner

and more widespread C. zeyherianum.

Vouchers: Oliver 7548 (BOL, K, MO, NBG, PRE, S, STE);

8801 (BM, E, G, NY, P, PRE, S, STE, UPS, W).

ACKNOWLEDGEMENTS

Curators of the following herbaria are thanked for

the loan of specimens: BOL, K, MEL, MO, NBG,

P, PRE, S, SAM. Discussions with A. Jacot Guillar-

mod of GRA are appreciated. Electron microscope

investigations were carried out using the ISI 100A at

the Fruit and Fruit Technology Research Institute,

Stellenbosch, with the assistance of C. Swart and P.

van der Merwe.

REFERENCES

BENTHAM, G. 1839. Ericaceae. In A. P. De Candolle, Prodro-

mus systematis naturalis regni vegetabilis 7: 580-733. Paris.

BENTHAM, G. 1876. Ericaceae. In G. Bentham & J. D.

Hooker, Genera ptantarum 2: 577-604. Reeve, London.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora, a

descriptive catalogue. Journal of South African Botany,

Supplementary Volume 13, pp. 455.

BROWN, N. E. 1906. Ericaceae. In W. T. Thistleton-Dyer, Flora

capensis 4,1: 337-418. Reeve, Ashford.

DE CANDOLLE, A. P. 1838. Prodromus systematis naturalis

regni vegetabilis 7. Paris.

DRUDE, O. 1897. Ericoideae-Ericaceae. In H. G. A. Engler &

K. A. E. Prantl, Die Naturlichen Pflanzenfamilien 4,1:

57-65. Engelmann, Leipzig.

GOLDBLATT, P. 1978. An analysis of the flora of southern

Africa: its characteristics, relationships and origins. Annals

of the Missouri Botanical Garden 65: 369-436.

KLOTZSCH, J. F. 1838. Ericearum, genera et species. Linnaea

12: 211-247.

OLIVER, E. G. H. 1975. Ericaceae. In R. A. Dyer, The genera

of southern African flowering plants 1: 429-439. Botanical

Research Institute, Pretoria.

OLIVER, E. G. H., LINDER, H. P. & ROURKE, J. P. 1983.

Geographical distribution of present-day Cape taxa and

their phytogeographical significance. Bothalia 14: 427-440.

PHILLIPS, E. 1926. The genera of South African flowering

plants, 1st edn, 457-465. Division of Botany and Plant

Pathology, Pretoria.

PHILLIPS, E. 1944. Notes on the minor genera of the Ericaceae.

Journal of South African Botany 10: 69-73.

PHILLIPS, E. 1951. The genera of South African flowering

plants, 2nd edn, 558-562. Division of Botany and Plant

Pathology, Pretoria.

REBELO, A. G., SIEGFRIED, W. R. & OLIVER, E. G. H.

1985. Pollination syndromes of Erica species in the south-

western Cape. South African Journal of Botany 51:

270-280.

WEIMARCK, H. 1941. Phytogeographical groups, centres and

intervals within the Cape Flora. Lunds Universitets Ars-

skrift 37: 1-143.

SPECIMENS EXAMINED

Anderson 279 (1) GRA.

Batten s.n. (1) NBG; Blake s.n. (1) STE; Bodkin sub Bolus 6693

(1) BOL, PRE; H. Bolus 9803 (1) BOL, PRE; L. Bolus in BOL

27094 & 27905 (1) BOL; Burchell 6875 (2) BOL, K.

170

Bothalia 17,2 (1987)

Drege 7753 (1) K, S; s.n. (1) G-DC.

Ecklon & Zeyher 294 (1) G, LD. M, MEL, MO, S, UPS, W, Z;

296 (1) G, MEL, MO, W; s.n. (1) BOL, E, GRA, MEL, K, P,

PRE, S, W, Z.

Fourcade 2151 (1) BOL, K, NBG, STE; 2152 (1) BOL, K.

Long 301 (1) GRA, K, PRE; 302 (1) K, PRE; 303 (1) BOL, K;

Lynes 99 (1) BM; s.n. (1) BM.

Mund s.n. (1) BOL.

Oliver 4492 (1) PRE, STE; 4494 (1) (STE); 7548 (2) BOL, K,

MO, NBG, PRE, S, STE; 7935 & 7936 ( 1) GRA, NY, P, PRE, S,

STE ; 7939 ( 1 ) STE ; 7940 ( 1 ) PRE , STE ; 7942 ( 1 ) PRE , STE ; 7943

(1) STE; 7945 & 7946 (1) PRE, STE; 7948 & 7949 (1) BM, BOL,

E, K, NBG, MO, PRE, STE; 8801 (2) B, BM, E, G, MO, NY, P,

PRE, S, STE, UPS, W; 8802 (2) STE; 8817 (2) PRE, STE.

Paterson 1145 (1) GRA, Z; 1156 (1) GRA; 2290 (1) Z; Pole

Evans 18277 (1) PRE.

Rogers 28662 (1) GRA.

Sim 36 (1) BOL.

Trash 9 (1) GRA.

West 279 (1) BOL.

Zeyher 719 (1) BM, BOL, K, PRE; 3236 (1) GRA, MEL, P,

PRE; 3321 (1) GRA, MEL, P, W; s.n. (1) BOL, K.

Bothalia 17,2: 171-177 (1987)

Synopsis of the genus Salix (Salicaceae) in southern Africa

K. L. IMMELMAN*

Keywords: phytogeography, Salicaceae, Salix, southern Africa, taxonomy

ABSTRACT

One species of Salix, S. mucronata Thunb. (=S. subserrata Willd.), with five subspecies, is recognized as

indigenous to the southern African region. Problems of delimitation in the genus in southern Africa are discussed,

and a key to the indigenous and exotic taxa is presented. The synonymy of the subspecies is presented, with leaf

silhouettes and a distribution map of each. The following new combinations are made: S. mucronata subsp. hirsuta

( hunb.) Immelman, S. mucronata subsp. capensis (Thunb.) Immelman, S. mucronata subsp. woodii (Seemen)

Immelman and S. mucronata subsp. wilmsii (Seemen) Immelman.

UITTREKSEL

Een Salix- spesie, 5. mucronata Thunb. ( =S . subserrata Willd.), met vyf subspesies, word erken as inheems in

die suider-Afrikaanse gebied. Probleme met omgrensing in die genus in suidelike Afrika word bespreek, en daar is

n sleutel tot die inheemse en uitheemse taksons. Die sinonimie van die subspesies word gegee, met blaarsilhoeette

en n verspreidingskaart van elk. Die volgende nuwe kombinasies word gemaak: S. mucronata subsp. hirsuta

(Thunb.) Immelman, S. mucronata subsp. capensis (Thunb.) Immelman, S. mucronata subsp. woodii (Seemen)

Immelman en S. mucronata subsp. wilmsii (Seemen) Immelman.

INTRODUCTION

Identification of taxa within the genus Salix in

southern Africa is difficult for the following reasons:

1, all species are dioecious; 2, spring leaves, which

are present when the flowers first appear, differ con-

siderably from summer leaves, and are similar in all

the southern African material; 3, flowers and fruits

are similar in all the taxa; 4, leaf and pubescence

characters are highly variable within the taxa, so that

it is difficult to discern a pattern.

METHODS

Specimens were borrowed from all major South

African herbaria, as well as from Windhoek and Ha-

rare. The following macromorphological characters

were examined: leaf size, shape and margins; petiole

length; pubescence of leaves and twigs; and struc-

ture of flowers and fruits. The glands in male and

female flowers were examined. The number of sta-

mens in the male flowers was counted, because this

character has been often used to distinguish species

of Salix in other parts of the world. Adaxial and

abaxial leaf surfaces, as well as pollen and seeds,

were viewed with the SEM.

RESULTS

No taxonomically useful results were obtained

from either the SEM examinations or the stamen

counts. The number of stamens varied from 3-12 per

flower, and varied even within the same inflores-

cence. The glands of both male and female flowers

varied considerably, with 2 to numerous glands in

the male flower and a glandular ring in the female.

This ring varied from entire to deeply lobed, the

number and size of the lobes being irregular, but it

* Botanical Research Institute, Private Bag X101, Pretoria 0001.

did not present any pattern. Fruits and flowers of all

southern African specimens were found to be essen-

tially similar.

Leaf length, breadth, proportions and margin, pu-

bescence of both leaves and twigs, and petiole

length, however, did differ between taxa, and these

differences were correlated with distribution. In

northern SWA/Namibia, the south-western Cape

and the northern Transvaal, the summer leaves are

large, relatively broad, glabrous or sometimes with

grey canescence, and have either entire or toothed

margins. Around the Olifants River (south-western

Cape) the leaves are similar in shape to the above

but have a dense covering of long silvery trichomes.

Specimens from the rest of the Cape Province, in-

cluding the Orange River system, and extending

along the Vaal into the Transvaal and Orange Free

State, and into Lesotho, have short, relatively broad

leaves, always toothed and glabrous, with short pe-

tioles and glabrous twigs. Plants from Natal and

most of the Transvaal (excluding the Vaal and its

tributaries), have long narrow leaves, always

toothed, with longer petioles and (usually) grey-can-

escent twigs. Plants with larger, broader leaves,

nearly always with entire margins, occur in the

Transvaal Lowveld and escarpment from Swaziland

to the northern Transvaal border and into Zim-

babwe.

DISCUSSION

It was difficult to distinguish clearly any of the

taxa morphologically, except the Olifants River

taxon which has densely silvery-pubescent leaves.

Most characters showed at least some overlap and,

in the case of the Transvaal Lowveld taxon, the

characters were especially variable. However, it is

possible to recognize most specimens as belonging to

one or other of several taxa and, in addition, the

172

Bothalia 17,2 (1987)

variation is geographically correlated. For these

reasons it was decided not to place all indigenous

plants in a single highly polymorphic taxon. Burtt

Davy (1922) in his study of the genus in the whole of

Africa, was the first to suggest that each species or

variety of Salix was restricted to a certain drainage

basin(s). In southern Africa at least, this conclusion

can be accepted, though distributions of the taxa do

overlap to some degree in the south-western Cape

and to a larger extent in the Transvaal. Each taxon

shows a well defined range and for this reason, al-

though the morphological differences are small, the

taxa are recognized at subspecific rather than var-

ietal level.

It is not certain whether the variability and inter-

grading of characters is due to inter-taxon hybridiza-

tion since no attempt has been made to hybridize the

indigenous taxa under controlled conditions. It is

possible that S. mucronata subsp. wilmsii is a hybrid

between subsp. woodii and subsp. mucronata. It is in

many ways intermediate between these two subspe-

cies, it occurs within the range of both, and it ex-

hibits a great range of variation within the subspe-

cies. Its variation, however, may also be ascribed to

inter-taxon diversity. Therefore, until experimental

studies have been done, it appears best to maintain it

as a separate taxon. The taxa are difficult to identify

because the differences between them are both small

and variable. A further difficulty results from the

fact that there is a great difference on each plant

between the spring (immature) and summer (ma-

ture) leaves. It is preferred to call them spring and

summer leaves respectively, as ‘immature’ implies

further development, whereas the leaves maintain

their differences even when growth is complete. The

spring leaves are smaller, broader in proportion to

their length, often obovate, the margins are entire

and the apices may be rounded rather than acute.

The transition to summer leaves is gradual, but is

usually complete by November. In the following

key, only specimens collected from November to

May have been considered. For specimens collected

at other times of the year the locality may present

the only clue to identity. Unfortunately, as stated

earlier, no flower or fruit characters were found to

distinguish taxa. This made the identification of type

specimens difficult, as a number were collected in

spring and in some cases no locality was given.

It is often stated that the southern African taxa do

not have stipules. Examination of fresh material of

subsp. woodii, however, showed that this was not

strictly true, as vestigial stipules are present on

young twigs. These are ± 1 mm long, thick in tex-

ture, and are later deciduous; in dry herbarium ma-

terial they are not noticeable. S. mucronata subsp.

mucronata does have stipules further north in its

range, but not within southern Africa.

A few exotic species appear to have become natu-

ralized in southern Africa, and are included in the

key. S. babylonica (Weeping Willow) (Figure 1.13)

is widespread and well known, while the other

species, from the few specimens available, appear to

be more restricted. No specimens have been seen

which appear to represent hybrids between the in-

digenous and exotic taxa. More collecting of exotic

willows is needed, together with information about

whether they are cultivated, naturalized or spread-

ing. Only female plants of the exotic species are

present in this country, and none of the specimens

seen were male.

KEY TO SPECIES AND SUBSPECIES

la Tree with branches hanging vertically; leaves tapering to a long acuminate apex (whip-tip); female

flowers sessile 2. 5. babylonica L. (Figure 1.13)

lb Trees or shrubs, branches may droop but not hanging vertically; leaves acute or acuminate but not

tapering to a long ‘whip-tip'; female flowers and fruits sessile or pedicellate:

2a Branches ascending; stipules usually present, foliose or dentate, may be caducous; female flowers

sessile (may be shortly pedicellate in 5. lasiandra )\

3a Basal 1-3 leaves on flowering shoots with long silky golden-white hairs on margin; female flowers

with a single gland at base of ovary 4. 5. alba L. (x 5. fragilis L. ?)

3b Basal leaves on flowering shoots without long hairs on margin; female flowers with 2 glands at

base of ovary:

4a Petiole with a pair of glands on upper side near junction with lamina; style at least 0,5 mm

long; twigs not brittle at nodes 5.5. lasiandra Benth.

4b Petiole without glands; style very short or absent; twigs brittle at nodes 3.5. fragilis Benth.

2b Branches drooping; stipules, if present, minute and subulate, not dentate; female flowers and fruits

sessile:

5a Leaves and branches densely silvery hirsute; found on the Olifants River (SW Cape) and its

tributaries lb. 5. mucronata subsp. hirsuta (Figure 1.4)

5b Leaves and branches glabrous or grey-canescent; not found on the Olifants River:

6a Leaf lamina (8-) 14-23 mm wide, 3-5 (-7) times as long as wide; northern SWA/Namibia, SW

Cape and occasionally N Transvaal la. 5. mucronata subsp. mucronata (Figure 1. 1-1.3)

6b Leaf lamina 5-20 mm wide, usually more than 5 times as long as wide; rarely in SW Cape,

never in northern SWA/Namihia:

7a Summer leaves usually shorter than 65 mm; petioles 2-5 mm long; twigs always glabrous;

mainly on Orange River and its tributaries, including the Vaal, and on rivers of the

southern and eastern Cape as far north as Port Shepstone in Natal, occasionally found

elsewhere lc. 5. mucronata subsp. capensis (Figure 1.5, 1.6)

Bothalia 17,2 (1987)

173

7b Summer leaves usually longer than 60 mm; petioles 4-14 mm long; twigs grey-canescent to

puberulous, sometimes glabrous; mainly on Limpopo and Olifants (Transvaal) Rivers

and their tributaries, and on Natal rivers as far south as Port Shepstone;

8a Summer leaves usually entire, 10-20 mm wide; branches and young leaves often very

densely grey-canescent; along Transvaal escarpment and into lowvcld as far south

as Swaziland , occasional specimens from other places

le. S. mucronata subsp. wilmsii (Figure 1.10-1.12)

Xb Summer leaves always toothed, 6-14 mm wide; branches and young leaves less densely

canescent, may be puberulous or sometimes glabrous; along whole of drainage

basin of Limpopo, Olifants (Transvaal) and Maputo Rivers, and Natal rivers as far

south as Port Shepstone Id. 5. mucronata subsp. woodii (Figure 1.7-1. 9)

Salix mucronata Thunb., Prodromus plantarum

capensium 6 (1794); Willd. 4: 685 (1806); Thunb. :

31 (1807); Burtt Davy: 70 (1922) p.p.

(a) subsp. mucronata

Type: No type designated by Thunberg, single

specimen in herb. Thunb. annotated ‘S mucronata’

in his hand, Thunberg (sheet 23065, UPS, lecto.,

here designated; microfiche in PRE!).

S. subserrata Willd.: 671 (1806); Milne-Redhead: 474 (1936);

Maire: 50, fig. 1129 (1961); Friedrich-Holzhammer: 14 (1967).

Type: Egypt, near Cairo, Bulak, no collector (sheet 18137, B-

WILLD; microfiche in PRE!).

S. aegyptica sensu Thunb.: 30 (1807), non Willd. (1806). Syn-

types: Cape Province, near rivers at Roodesand, Sept. -Oct.,

Thunberg s.n. (sheets 22885, 22886, 22887, UPS; microfiche in

PRE!).

S. safsaf Forssk. ex Trautv.: 6, t. 2 (1836); Forssk.: 76 (1775), as

5. safsaf baelledi, nom. nud.; Anderss.: 196 (1868); Boiss.: 1183

(1897); Skan: 318 (1917); Burtt Davy: 432 (1932) (may refer to

subsp. wilmsii ?). Type: Egypt, Herb. Sieber (LE, holo.; K, iso.

fide Wilmot-Dear in litt.).

The unusual distribution of subsp. mucronata,

which occurs in northern SWA/Namibia and in the

south-western Cape, suggests that it may once have

occurred in the intervening area, and that it has since

died out there due to the drying up of perennial

rivers in much of SWA/Namibia and Namaqualand.

There is no noticeable difference between the SWA-

/Namibian and Cape populations that might justify

describing a new subspecies. Subsp. mucronata is

widely distributed in Africa, entering the area under

consideration also in the northern Transvaal, with

one record from the eastern Transvaal. Figures

1.1-1. 3 & 2A.

Within the subspecies as a whole there is a great

range of variation, from specimens with glabrous

branches and entire glabrous lanceolate leaves, to

(in various combinations) broadly elliptic leaves with

regularly to irregularly serrate margins, and slender

or stout stems with dense canescence. None of the

specimens from our area have stipules, though they

do occur in the subspecies elsewhere in Africa.

Burtt Davy (1922) seems to have had a mixed con-

cept of S. mucronata. His S. mucronata var. mucro-

nata comprises elements of both subsp. mucronata

(as delimited here) and of subsp. capensis. The

range of distribution given on Burtt Davy’s map in-

cludes that of both subspecies, while the synonyms

cited comprise S. aegyptica Thunb. (S. mucronata

subsp. mucronata), ‘S. capensis auct. non Thunb.’ ,

and S. mucronata var. integra (placed here under

‘taxa insufficiently known’). His illustrations of the

leaves are clearly those of S. mucronata subsp. mu-

cronata. Burtt Davy therefore probably had speci-

mens of both subsp. mucronata and subsp. capensis

in mind when delimiting his S. mucronata var. mu-

cronata.

Vouchers: Hemm 452 (J); Leistner, Oliver, Steenkamp &

Vorster 110 ; Marloth 4283, 11843; Merxmuller & Giess 30494; Van

Wyk, Relief & Herman 6737.

(b) subsp. hirsuta (Thunb.) Immelman, comb.

nov.

Salix hirsuta Thunb., Prodromus plantarum capensium 6

(1794); Thunb.: 31 (1807); Willd.: 695 (1806); Fries: 120 (1856);

Krauss: 88 (1844); Skan: 579 (1912); Adamson & Salter: 311

(1950). S. capensis var. hirsuta (Thunb.) Anderss.: 14 (1867);

Anderss.: 198 (1868); Sim: 329 (1907). Type: No type designated

by Thunberg, single specimen in herb. Thunb. annotated ‘S. hir-

suta' (sheet 23028, UPS, lecto., here designated; microfiche in

PRE!). Figures 1.4 & 2B.

Vouchers: Boucher 1985; Hanekom 1272; Hugo 746; Van Jaars-

veld 4496.

(c) subsp. capensis (Thunb.) Immelman, comb.

nov.

Salix capensis Thunb., Flora capensis 31 (1807); Harvey: 347

(1838); Anderss.: 197 (1868) excl. vars. mucronata, hirsuta; Sim:

328, t. 146 (1907); Skan: 576 (1912) excl. var. mucronata et syn. S.

aegyptica; Marloth: 130, fig. 73 (1913); Burtt Davy: 69 (1922);

Burtt Davy: 432 (1932); Friedrich-Holzhammer: 14 (1967); Jacot

Guillarmod: 161 (1971). Syntypes: Cape Province, near rivers in

mountains near Hantam, Thunberg s.n., (sheets 22958, 22959,

22960, UPS, microfiche in PRE!).

5. gariepina Burch.: 317, t. 6 (1822); Pappe: 35 (1862); Burtt

Davy: 338 (1921). S. capensis var. gariepina (Burch.) Anderss.: 13

(1867); Anderss.: 197 (1868); Sim: 328 (1907); Skan: 576 (1912);

Burtt Davy: 432 (1932). Type: Cape Province, Prieska, banks of

Orange River, Burchell 1637 (K!).

S. crateradenia Seemen: 9 (1899); Skan: 578 (1912). Type: Bo-

tswana, Passarge 41 of 1896 (not located).

S. mucronata var. mucronata sensu Burtt Davy p.p. (excl. east-

ern element): 71 (1922). S. mucronata var. caffra Burtt Davy: 71

(1922). Isotypes: Cape Province, Eastern Districts, Cooper 48

(BM!, K!).

Burtt Davy’s concept of S. capensis is narrower

than that adopted here for subsp. capensis, and I

include taxa he accepts as separate. These are S.

crateradenia from the northern Cape and S. capensis

var. caffra from the eastern Cape. S. capensis var.

integra, possibly from the eastern Cape, may also

belong here but is mentioned under ‘taxa insuffi-

ciently known’ below. S. mucronata var. mucronata

sensu Burtt Davy pro parte, excluding those speci-

mens from the southern and eastern Cape, must also

be included. Figures 1.5, 1.6; 2C.

The type of S. crateradenia (from the northern

Cape) has not been located but, judging from See-

men’s description, it is almost certainly S. mucronata

subsp. capensis. He commented that it is close to S.

capensis, but distinguished it by its well defined style

174

Bothalia 17,2 (1987)

FIGURE 1. — 1-3, Salix mucronuta subsp. mucronata, summer leaves. 1, Botswana, Chobe River, Miller B946; 2, SW Cape,

Eerste River, Boucher 3509; 3, N Transvaal, Tate Vondo Reserve, Hemm 452. 4, S. mucronata subsp. hirsuta, summer leaf,

W Cape, Cederberg, Uitkyk Pass, Goldblatt 3278. 5-6, S. mucronata subsp. capensis, summer leaves. 5, Orange Free State,

Willem Pretorius Reserve, Bourquin 888 ; 6, N Cape, Barkley West, Hafstrom H. 961. 7-8, S. mucronata subsp. woodii,

summer leaves. 7, Natal, Vants Drift, Letty 486 sub Codd s.n.; 8, Transvaal, Pienaars River, Repton 435. 9, intermediate

between subsp. woodii and subsp. wilmsii ?, summer leaf, Transvaal, Krugersdorp, near Skeerpoort River, Codd 10096.

10-12, Salix mucronata subsp. wilmsii, summer leaves. 10, Transvaal, Lydenburg, Lowveld Botanic Garden, Buitendag 997 ;

1 1, Transvaal, Kruger National Park, Sigaas, Van der Schijff 357; 12, Transvaal, Kruger National Park, near Punda Maria,

Codd 5558. 13, S. babylonica, summer leaf. Cape, near Cape Town, Ecklon 713. All in PRE, x 1.

Bothalia 17,2 (1987)

175

FIGURE 2. A, Salix mucronata subsp. mucronata', B, S. rnucronata subsp. hirsuta ; C, S. mucronata subsp. capensis\ D, S.

mucronata subsp. woodii; E, S. mucronata subsp. wilmsii. 1, Orange River drainage basin; 2, Olifants River (Cape) drainage

basin; 3, Limpopo River drainage basin; 4, Olifants River (Transvaal) drainage basin; 5, Maputo River drainage basin, 6,

Natal rivers; 7, Cape rivers.

176

Bothalia 17,2 (1987)

and the entire glands in the female flower. Styles

have not been found to vary significantly in this

study, and the variation in glands was found to be

considerable, without any discrete ranges of varia-

tion. Burtt Davy states that the species comes from

the headwaters of the Kuruman River, which is in

the northern Cape, and is a tributary of the Orange

River. A population of Salix has been found at this

approximate locality by Mr A. Gubb, of the McGre-

gor Museum, Kimberley, who states that he does not

consider it to differ from S. mucronata subsp. capen-

sis (pers. comm.).

Some specimens which might be intermediate be-

tween subsp. capensis and subsp. woodii have been

seen. These are Green 88, Leendertz 3752, Muller

1053 and Sutton 884 (all at PRE). These occur near

the boundaries of the two subspecies, both between

the Orange and Limpopo drainage basins and the

Orange River drainage basin and the Natal rivers.

Vouchers: Hafstrom H 961', Merxmiiller 2270; Moffett 627;

Oliver 3121; Van der Westhuizen 44178.

(d) subsp. woodii (Seemen) Immelman, comb.

nov.

Salix woodii Seemen in Botanische Jahrbiicher 21 Beiblatter 53

(1896); Wood: 121 (1907); Skan: 577 (1912); Bews: 79 (1921);

Burtt Davy: 40 (1921); Burtt Davy: 432 ( 1932); Jacot Guillarmod:

161 (1971); Compton: 172 (1976). Type: Natal, Tugela, near Col-

enso, Wood 4970 (not found).

For a discussion of specimens intermediate be-

tween this subspecies and subsp. capensis, see under

that subspecies. It also appears to grade into subsp.

wilmsii.

Although the type has not been found, the de-

tailed description allows it to be confidently identi-

fied as this subspecies. The leaves are said by See-

men to be 90 x 11 mm, with the margin having small

sharp teeth, and the shape narrowly lanceolate to

linear, which can only be subsp. woodii. The fact

that the type comes from Natal, from the Tugela

River near Colenso, confirms this identification. Fig-

ures 1.7-1. 9 & 2D.

Vouchers: Acocks 10120; Codd 10095; Duggan & Henderson

24; Killick & Marais 2130; Letty 481; Rogers 2736 (GRA).

(e) subsp. wilmsii (Seemen) Immelman, comb.

nov.

S. wilmsii Seemen in Botanische Jahrbiicher 27 Beiblatter 64: 9

(1900); Burtt Davy: 40 (1921); Burtt Davy: 432 (1932). S. woodii

var. wilmsii (Seemen) Skan: 578 (1912). Type: Transvaal, Lyden-

burg, Wilms F.S.A. 1350 (PRE, lecto., here designated!; BOL!).

S. wilmsii x safsaf, Burtt Davy: 432 (1932). Type: Transvaal,

Lydenburg, Grootfontein River, foot of Burgers Pass, Davy H

1559 (PRE!). Burtt Davy also cites: Transvaal, Barberton, Pole

Evans H 2965 (PRE!), and queries whether it is the reciprocal

cross.

S. wilmsii x woodii, Burtt Davy: 432 (1932). Isotypes: Trans-

vaal, Barberton, Galpin 1278 (GRA!, PRE!).

Specimens of this subspecies have been seen

which are very close to subsp. mucronata, e.g.

Hardy 401 and Hemm 452. In its typical form, subsp.

wilmsii is easily distinguishable, but it may approach

subsp. woodii and subsp. mucronata in appearance.

Only further research can establish whether the

taxon is simply very variable or whether it has under-

gone introgressive hybridization with these other

subspecies. As mentioned in the Discussion, subsp.

wilmsii itself may also be a hybrid between subsp.

woodii and subsp. mucronata. In its ‘pure’ form,

subsp. wilmsii has stout, densely canescent twigs and

large, broadly lanceolate, entire leaves, which are

densely grey-canescent when young. Figures

1.10-1.12 & 2E.

Vouchers: De Winter 7685; Gerstner 5492; Hardy 957; Prior 33;

Theron 3569.

Taxa insufficiently known

S. mucronata Thunb. var. integra Burtt Davy in

Journal of Ecology 10: 70 (1922). Type: Cape Pro-

vince, Camdeboo, on the flats and at the river near

the Camdeboosberg, 2000-3000 ft, Drege s.n. (K!,

S!). The Stockholm specimen is S. mucronata subsp.

capensis but the one from Kew is subsp. hirsuta, with

large entire leaves and dense silvery pubescence on

the young leaves and twigs. No other specimen re-

sembling this has been seen from the area, and it is

possible that the Kew specimen has been misla-

belled. Drege did collect at the Olifants River,

where subsp. hirsuta occurs.

S. woodii var. grandifolia Burtt Davy, Flowering

Plants and Ferns of the Transvaal 2: 432 (1832),

nom. nud. Specimens cited: Davy 10614; Legat H

4331; Robertson 1474 (none of these found). Burtt

Davy speculates whether this is not a hybrid between

S. woodii and S. wilmsii.

S. woodii Seemen x safsaf Forssk. ex Trautv.?,

Burtt Davy, Flowering Plants and Ferns of the

Transvaal 2: 432 (1932). Type: Transvaal, Louis Tri-

chardt, 3100 ft, Rogers 21690 (not found). From the

description this could be subsp. woodii.

ACKNOWLEDGEMENTS

I would like to thank Mrs S. Perold for her help

with the SEM work for this study, and Miss M. Wil-

mot-Dear for identifying the specimens of exotic

Salix species sent to her at Kew.

REFERENCES

ADAMSON, R. S. & SALTER, T. M. 1950. Flora of the Cape

Peninsula. Juta, Cape Town.

ANDERSSON, N. J. 1867. Monographia Salicum. Norstedt,

Stockholm.

ANDERSSON, N. J. 1868. Salicaceae. In A. P. de Candolle,

Prodromus systematis naturalis regni vegetabilis 16,2:

190-331. Masson, Paris.

BEWS, J. 1921. An introduction to the flora of Natal. City Print-

ing Works, Pietermaritzburg.

BOISSIER, P.E. 1897. Flora orientals 4. Eumdem, Geneve.

BURCHELL, W. J. 1822. Travels in the interior of southern

Africa 1. Longman, London.

BURTT DAVY, J. 1921. New or noteworthy South African

plants. Kew Bulletin 1921: 338-340.

BURTT DAVY, J. 1922. The distribution and origin of Salix in

South Africa. Journal of Ecology 10: 62-86.

BURTT DAVY, J. 1932. Flowering plants and ferns of the Trans-

vaal, with Swaziland 2. Longmans, Green, London.

COMPTON, R. H. 1976. An annotated checklist of the flora of

Swaziland. National Botanic Gardens, Kirstenbosch.

FORSSKAl, P. 1775. Flora aegyptiaco-arabica. Mailed, Hau-

niae.

Bothalia 17,2 (1987)

177

FRIEDRICH-HOLZHAMMER, M. 1967. Salicaceae. In H.

Merxmiiller, Prodromus einer Flora von Siidwestafrika 14.

Cramer, Lehre.

FRIES, E. 1856. Anmarkningar ofver de Kapska Pilarterna. Of-

versigt af Kungelsk Vetenskapsakademiens forhandlingar 8:

119-122.

HARVEY, W. H. 1838. The genera of South African plants. Ro-

bertson, Cape Town.

JACOT GUILLARMOD, A. 1971. Flora of Lesotho. Cramer,

Lehre.

KRAUSS, F. 1844. Pflanzen des Cap- und Natal-landes. Flora 27.

MAIRE, R. 1961. Salicaceae. Flore de lAfrique du Nord 7:

47-71. Lechevalier, Paris.

MARLOTH, R. 1913. The Flora of South Africa 1. Darter, Cape

Town.

MILNE-REDHEAD, E. 1936. Tropical African plants XIV. Kew

Bulletin 1936: 474.

PAPPE, L. 1862. Silva capensis. Ward, London

SEEMEN, K. O. von. 1899. Zwei neue Weidenarten aus Siid-

Afrika. Botanische Jalirbiicher 27, Beiblatter 64.

SIM, T. R. 1907. The forests and the forest flora of the colony of

the Cape of Good Hope. Taylor & Henderson, Aberdeen.

SKAN, S. A. 1912. Salicaceae. In W. T. Thistleton-Dyer, Flora

capensis 5,2: 575-579. Reeve, London.

SKAN, S. A. 1917. Salicaceae. In D. Prain, Flora of tropical

Africa 6,2: 316-324. Reeve, London.

THUNBERG, C. P. 1794. Prodromus plantarum capensium Up-

psala.

THUNBERG, C. P. 1807. Flora capensis. Cotta, Stuttgart.

TRAUTVETTER, E. R. von. 1836. Salicetum, Petropoli.

WILLDENOW, C. L. 1806. Caroli a Linne Species Plantarum,

Editio quarta 4. G. C. Nauk, Berlin.

WOOD, J. M. 1907. A handbook to the flora of Natal. Bennet,

Durban.

Bothalia 17,2: 179-190 (1987)

Notes on African plants

VARIOUS AUTHORS

ACAROSPORACEAE

A NEW YELLOW ACAROSPORA (LICHENES) FROM THE WATERBERG,

SOUTH WEST AFRICA/NAMIBIA

Acarospora elegans Brusse sp. nov., thallo ut in

A. oxytona (Ach.) Massal. sed excipulo tenuiore

(5-10 pm), et ascosporis latioribus.

Thallus citrinus, crustosus et effiguratus, saxicola,

usque ad 20 mm diametro. Lobi marginales usque ad

2,5 mm longi, 0,3-1 ,0 mm lati. Areolae irregu-

lares, 0, 2-1,0 mm diametro, sessiles vel subpeltatae.

Fissurae 0, 1-0,2 mm latae, 0, 2-1,0 mm profundae.

Superficies opaca, laevis vel undulata, plana vel con-

vexa, citrina. Cortex superior 25-45 pm crassus (in

lateribus et in pagina inferior, 10-15 pm crassus),

paraplectenchymatus, cellulis 3-8,5 pm diametro.

Stratum gonidiale 60-80 pm crassum, algis Treboux-

iis, 5-15 pm diametro. Medulla alba, 70-300 pm

crassa. Pagina inferior pallida vel brunnescens.

Apothecia usque ad 1,5 mm diametro, emergentia

vel sessilia; pagina hymenii pallida. Excipulum thal-

linum in lateribus circa 120 pm crassum. Excipulum

hyalinum, J — , periclinate prosoplectenchymatum,

5-10 pm crassum, sed crassius ad paginam. Hypo-

thecium 15-50 pm crassum, hyalinum, J+ caeru-

leum. Hymenium hyalinum, 80-90 pm altum, J +

caeruleum. Paraphyses fere simplices vel anastomo-

santes, in gelatina arete inclusae, septatae, lumini-

bus 1,2-1 ,3 pm crassis, gelatinis J+ caeruleis. Asci

clavati vel acuminate clavati, 60-75 x 14-19 pm,

parietibus incrassatis, J- (Figure 2). Ascosporae nu-

merosae, hyalinae, ellipsoideae, simplices, 4-5 x

1,8-2, 8 pm. Pycnidia globosa, hyalina, 230-290 pm

profunda, 16CU200 pm lata, cum algis cincta, parieti-

bus hyalinis, circa 10 pm crassis, periclinate proso-

plectenchymatis, cavitatibus convolutis. Pycnidio-

sporae hyalinae, longe ellipsoideae, 2, 0-3, 5 x

1,0-1 ,2 pm. Thallus acidum rhizocarpicunr conti-

nens.

TYPE. — South West Africa/Namibia, 2017

(Waterberg): Waterberg campsite trial, common on

S facing sandstone cliffs (-CA), F. Brusse 4218,

1984.03.22 (PRE, holo.; COLO, LD, iso.) Figure 1.

Thallus as in A. oxytona , but the exciple thinner

(5-10 pm), and the ascospores broader.

Thallus yellow, effigurate-crustose, saxicole, up to

20 mm diam. (larger by confluence). Lobes margi-

nal, up to 2,5 mm long, 0,3-1 ,0 mm broad. Areoles

irregular, 0,2-1 ,0 mm across, sessile to subpeltate.

Fissures 0, 1-0,2 mm wide, 0,2-1 ,0 mm deep. Upper

surface opaque, smooth to undulate, plane to con-

vex, yellow. Upper cortex 25-45 pm thick (10-15 pm

thick on sides and lower surface), paraplectenchy-

matous, cells 3-8,5 pm diam. Algal layer 60-80 pm

thick; algae Trebouxia , 5-15 pm diam. Medulla

white. 70-300 pm thick. Lower surface pale to

brownish. Apotliccia up to 1,5 mm diam., emergent

to sessile; hymenial surface pale. Thalline exciple

about 120 pm thick on sides. Exciple hyaline, J — ,

periclinally prosoplectenchymatous, 5—10 pm thick,

but thicker at surface. Hypothecium 15-50 pm thick,

hyaline, J+ blue. Hymenium hyaline, 80-90 pm

high, J+ blue. Paraphyses largely simple, but often

anastomosed, strongly gelled, septate, lumens

1,2-1, 3 pm thick, gel J+ blue. Asci clavate or acumi-

nate-clavate, 60-75 x 14-19 pm, walls thick (par-

ticularly at apex), J— (Figure 2). Ascospores numer-

ous, hyaline, ellipsoid, simple, 4—5 x 1,8-2, 8 pm.

Pycnidia globose, hyaline, 230-290 pm deep,

160-200 pm wide, surrounded by algae; walls hya-

line, about 10 pm thick, periclinally prosoplecten-

chymatous; cavity convoluted. Pycnidiospores hya-

line, long-ellipsoid, 2, 0-3, 5 x 1,0-1, 2 pm. Chemis-

try. Rhizocarpic acid in the cortex.

This new species resembles Acarospora oxytona

(Ach.) Massal. in thallus habit and lobe anatomy,

but differs in the thickness of the exciple and in the

ascospore size. The exciple is only 5-10 pm thick in

A. elegans , whereas it is 30-50 pm thick in A. oxy-

tona (Magnusson 1929). The ascospores are some-

what broader in the former species, reaching 2,8 pm

in equatorial diameter, and are ellipsoid. A. oxytona

ascospores only reach 2 pm in equatorial diameter,

being narrowly ellipsoid. The hypothecium of A.

oxytona also tends to be thicker and is darker blue in

Lugol’s iodine solution as well.

The lobe surfaces (Figure 3) of A. elegans are

never scabrous as they often are in A. oxytona (Fig-

ure 4), however, and the latter is often an alpine

lichen in the northern hemisphere. A. elegans is

abundant on south facing vertical or near vertical

sandstone cliffs in the Waterberg of South West

FIGURE 1. — Acarospora elegans Brusse, habit. F. Brusse 4218,

holotype. Scale in mm.

180

Bothalia 17,2 (1987)

FIGURE 2. — Acaros-

pora elegans

Brusse, ascus

and paraphysis.

F. Brusse 4218,

holotype. Bar =

10 pm.

Africa/Namibia. A photograph of this lichen from a

distance, is given by McDonald (1986), where the

yellow areas on the rocks are mainly or wholly this

new lichen. This is an unusually shaded place for an

Acarospora , particularly a yellow one, which usually

competes best in dryer places. In this regard, the

new species is similar to A. oxytona, because the

FIGURE 3. — Acarospora elegans Brusse, scanning electron

micrograph of upper surface. F. Brusse 4218, holotype.

Bar = 8,9 pm.

FIGURE 4. — Acarospora oxytona (Ach.) Massal., scanning

electron micrograph of upper surface. Hora s.n. (Arnold

Lichen Exsiccati 1159a). Bar = 9,2 pm.

latter also grows on steep cliff faces and under rock

overhangs (Magnusson 1929; Santesson 1984).

The cortical chemistry is usual for a yellow Acaro-

spora, containing the bright yellow pigment rhizo-

carpic acid. The medulla contains acetone extracta-

ble substances which show up as faint spots on TLC

plates (Culberson 1972; Culberson & Johnson 1982)

in longwave UV light. These substances are identical

to those found in A. oxytona, the extract of which

was co-chromatographed with that of A. elegans.

However, A. oxytona contains acaranoic and acare-

noic acids (Sarma & Huneck 1968; Follmann &

Huneck 1971; Huneck & Hofle 1980) in addition,

which could not be detected in A. elegans.

Several other effigurate yellow Acarosporae are

known from southern Africa (Magnusson 1933), but

these all have cylindrical ascospores, which are only

1 or 1,5 pm wide. Two of these (A. austroafricana

(Zahlbr.) H. Magn. and A. finckei Zahlbr.) are Na-

mib Desert lichens, known from schistose rocks. The

remaining species, A. calviniensis Magn., is a thin-

ner lichen, reaching 0,25 mm in thickness, and with

areoles only up to 0,5 mm across. The pycnidio-

spores of the last lichen are also shorter, reaching 1,7

pm long.

A. elegans is presently known only from the

Waterberg in South West Africa/Namibia, on steep

sandstone rock faces with a southern aspect.

Specimens of Acarospora oxytona (Ach.) Massal.

examined:

CZECHOSLOVAKIA. — 4916: An Gneissfelsen bei Rossatz

[Rossitz] im Donauthale, Baumgartner s.n., 1898 (Arnold, Li-

chenes Exsiccati 1159c\ PRE CH3216). 5014: An Quarzfelsen der

Scharka bei Prag, Hora s.n., Friihjahr 1886 ( Arnold , Lichenes

Exsiccati 1159a\ PRE CH3213).

RUMANIA. — 4422: Hungaria austro-orientalis (Banatus), in

rupibus calcareis ad Mehadiam, Lojka s.n. (Flora Exsiccata Aus-

tro-Hungarica 1951; PRE CH606 ). 4522: An senkrechten Quarz-

trachytwanden unter dem Gipfel des Berges Treszkovacz bei Sa-

vinicza an der unteren Donau, Comitat Krasso-Szoreny [Caras-

Severin] in Ungarn, Lojka s.n., 25. April 1886 Arnold Lichen

Exsiccati 1159b; PRE CH3214).

Bothalia 17,2 (1987)

181

ACKNOWLEDGEMENTS

I would like to thank Mr Trygve Cooper, the

senior ranger at the Waterberg Game Park, for his

friendly assistance during a short collecting stop

there.

REFERENCES

CULBERSON, C.F. 1972. Improved conditions and new data for

the identification of lichen products by a standardized thin-

layer chromatographic method. Journal of Chromatogra-

phy 72: 113-125.

CULBERSON, C.F. & JOHNSON, A. 1982. Substitution of

methyl tert. -butyl ether for diethyl ether in the standard-

ized thin-layer chromatographic method for lichen pro-

ducts. Journal of Chromatography 238: 483-487.

FOLLMANN, G. & HUNECK, S. 1971. Mitteilungen iiber

Flechteninhaltsstoffe LXXXVIII. Zur vergleichenden

Phytochemie der Krustenflechtenfamilie Acarosporaceae.

Philippia 1: 65-79.

HUNECK, S. & HOFLE, G. 1980. Structure of acaranoic and

acarenoic acids. Phytochemistry 19: 2713-2715.

MAGNUSSON, A.H. 1929. A monograph of the genus Acaro-

spora. Kungliga Svenska Vetenskapsakademiens Hand-

lingar, Ser. 3, 7, 4: 1-400.

MAGNUSSON, A.H. 1933. Supplement to the monograph of the

genus Acarospora. Annales de Cryptogamie Exotique 6:

13-48.

McDONALD, E. 1986. Lichens of southern Africa. South Afri-

can Panorama 31,7:13, fig. 4.

SANTESSON, R. 1984. The lichens of Sweden and Norway:

1-333. Swedish Museum of Natural History, Stockholm.

SARMA, K.G. & HUNECK, S. 1968. Uber Flechteninhalts-

stoffe, 52. Mitteilung: Ober die Inhaltsstoffe einiger Flech-

ten aus dem Himalaja. Die Pharmazie 23: 583-584.

F. BRUSSE*

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

CUCURBITACEAE

ORTHOGRAPHIC AMBIGUITY CLARIFIED

Acanthosicyos Welw. ex Hook. f. was published in

Benth. & Hook.f., Genera plantarum 1: 824 in Sep-

tember 1867 and although Welwitsch described the

genus it only appeared in Transactions of the Lin-

nean Society 27:30, two years later.

The type species, A. horrida was published as a

descriptio generico-specifica in accordance with

Art. 42.1 (Sydney Code) but this was an orthogra-

phic error. Art. 75 in Ex. 1 of recommendation

75 A. 1 is quite specific that the classical gender of

-sicyos is masculine and this should be retained in

the epithet as A. horridus.

The name has been published correctly both in

Index Kewensis and by Jeffrey (1962) in Kew Bull-

etin 15,3:340; but it has been consistently misused in

the South African literature.

B.D. SCHRIRE*

* Botanical Research Institute, Private Bag X101, Pretoria 0001.

Presently: SALO, Royal Botanic Gardens, Kew, Richmond, Sur-

rey TW9 3AE, England.

FABACEAE

A NEW SPECIES OF RHYNCHOSIA FROM THE RICHTERSVELD

Rhynchosia emarginata Germishuizen, sp.

nov., R. candidae affinis sed floribus longioribus,

foliolis emarginatis, infra punctis luteis glandulari-

bus differt.

TYPE. — Cape Province, 2816 (Oranjemund):

Richtersveld, Numees, west side of ridge north of

Numees Camp (-BD), McDonald 703 (PRE, holo.;

STE-U, iso.). Figure 5.

Robust, erect, much branched woody shruh up to

0,75 m tall. Branches and branchlets densely white

adpressed tomentose, interspersed with conspicuous

yellow resinous glands; old stems glabrescent with

bark flaking off in irregular pieces. Stipules free, ad-

nate to base of petiole, up to 3,5 mm long, white

adpressed tomentose on the outside, glabrous in-

side. Leaves trifoliolate; leaflets broadly obovate,

(4-) 5-11,2 (-12) x 3,5-8 (-9) mm, emarginate, ob-

tuse at base, with dense, adpressed strigose white

hairs on the upper surface, sparsely white adpressed

strigose mainly along the veins interspersed with

conspicuous yellow resinous gland dots on lower sur-

face. Petiole up to 11,5 mm long, white adpressed

tomentose, interspersed with yellow resinous glands.

Inflorescences axillary, few-flowered racemes up to

65 mm long, including a 25-45 mm long peduncle,

white adpressed hairy, interspersed with yellow

resinous glands; bracts caducous. Flowers up to

17 mm long. Calyx 8 mm long, 5-lobed; lobes un-

equal, 4-5 mm long, the upper pair connate, white

adpressed strigose, interspersed with yellow resinous

glands. Standard yellow with conspicuous purple

veins on the outside, obovate, 15 x 10 mm, eared at

the base, glabrous. Wing petals yellow, oblong,

eared, 13 x 3 mm, sculpturing absent. Keel blades

up to 17 mm long, yellow, purple at the apex, shal-

lowly pocketed. Stamens up to 17 mm long, 9 fused

into a staminal sheath and the 1 vexillar stamen free;

anthers uniform, 1 mm long. Ovary ovate, strigose.

Style up to 18 mm long, filiform, slightly thickened

and incurved in the upper third; stigma capitate,

hairy. Fruit a rimmed, flat, slightly falcate 2-seeded

pod, 30 x 8 mm, straw-coloured, often streaked

with purple, densely tomentose when young becom-

ing sparsely tomentose, interspersed with yellow

resinous glands later.

CAPE. — 2816 (Oranjemund): Richtersveld. Numees (-BD),

Jurgens 52; Jurgens 10114; McDonald 753; west side of ridge

182

Bothalia 17,2 (1987)

FIGURE 5.— Holotype of Rhynchosia emarginata Germishui-

zen.

14” 16” 18” 20”

FIGURE 6. — Distribution map of Rhynchosia emarginata

Germishuizen.

fied as Rhynchosia schlechteri Bak.f.and R. viscidula

Steud. R. emarginata can be distinguished from the

two above species by having emarginate obovate

leaflets adpressed hairy on the undersurface, inter-

spersed with conspicuous yellow resinous glands and

the absence of yeljow. glandular hairs with bulbous

bases on the stems, petioles ahd undersurface of the

leaves that are readily found on the above-men-

tioned species. In appearance, R. emarginata is clos-

est to R. emarginata but it differs in having longer

flowers, emarginate leaflets and yellow resinous

glands on the undersurface of the leaflets.

north of Numees Camp (-BD), McDonald 703. 2817 (Vioolsdrif):

head of Helskloof, Hottentotsparadysberg (-AA), Thompson &

Le Roux 146.

R. emarginata is found in the Richtersveld in

north-western Cape Province just south of the

Orange River (Figure 6) growing amongst rocks on

steep slopes or in dry streambeds. The first record of

the species in the PRE Flerbarium was collected by

Thompson and Le Roux during August 1977 from

Helskloof. It flowers during August and September.

All the material cited above was incorrectly identi-

ACKNOWLEDGEMENTS

I would like to thank Dr H.F. Glen for translating

the diagnosis into Latin and Mr E.G.H. Oliver of

the Stellenbosch Herbarium for sending their ma-

terial of this species on loan and Mrs W. Roux for

drawing the map

G. GERMISHUIZEN*

* Botanical Research Institute, Private Bag X101, Pretoria

0001.

LICHENES

PSATHYROPHLYCTIS, A NEW LICHEN GENUS FROM SOUTHERN AFRICA

Psathyrophlyctis serpentaria Brusse , gen. et sp.

nov.

Thallus crustosus, terricola, usque ad 60 mm di-

ametro, 0,5-1 ,0 mm crassus, olivaceus, omnino in

sorediis dissolutus. Soredia 20-40 pm diametro. Al-

gae chlorococcaleae, 4,5-10 pm diametro. Apothecia

lecideina, atra, convexa, laeves vel verrucata, usque

ad 1 ,5 mm diametro. Excipulum stramineum vel pal-

lide brunneum, radiatim et anticlinate scleroplecten-

chymatum (Figures 8e & 9), 60-120 pm crassum.

Hypothecium brunneum, collenplectenchymatum

(Figure 8a), 30-45 pm crassum, cellulis 4-8 pm di-

ametro. Hymenium hyalinum vel stramineum, circa

100 pm altum, interruptum. Asci clavati vel late cla-

vati, cum tholis et parietibus externis J+ caeruleis

(Figure 10). Ascosporae unicae vel binae, hyalinae,

muriformes, septis incrassatis, J — , 35-45 x 17-18,5

pm, loculis 10-14-seriatis, locellis 4-5-seriatis. Pyc-

nidia non visa. Thallus materiam ignotam continens

(Table 1).

TYPE. — Cape Province, 3321 (Ladismith): 17

km N of Riversdale, Garcia’s Pass, SW side of Ka-

Bothalia 17,2 (1987)

183

FIGURE 7. — Psathyrophlyctis serpentaria Brusse, habit.

F. Brusse 3612, holotype. Scale in mm.

reekop, in the Langeberg, on soil near rock on a

steep S slope (-CC), F. Brusse 3612, 1981.05.10

(PRE, holo.). Figure 7.

Thallus crustose, on soil, up to 60 mm across,

0, 5-1,0 mm thick, olive green, completely dissolved

into soredia. Soredia 20-40 pm diam. Algae chloro-

coccalean, 4,5-10 pm diam. Apothecia lecideine,

black, convex, smooth to warted, up to 1,5 mm

across. Excipulum stramineous or pale brown (some

of the interstitial spaces are dark brown and are indi-

cated in Figure 8 by heavy lines), radiately and anti-

clinally scleroplectenchymatous (Figures 8e & 9),

60-120 pm thick. Hypothecium brown, collenplec-

tenchymatous (Figure 8a), 30-45 pm thick, cells

4-8pm diam. Hymenium hyaline to stramineous,

about 100 pm high, interrupted by areas of sterile

hymenium with more branched paraphyses and

more deeply brown colour. Asci clavate to broadly

clavate (mature asci), one- or two-spored, tholus

and outer wall J+ blue (Figure 10). Ascospores hya-

line, muriform, septa thickened but J — , 35-45 x

17-18,5 pm, 10-14-loculate, 4— 5-locellate. Pycnidia

not seen. Chemistry : an unknown substance present

(Table 1), which is invisible in daylight and longwave

ultra-violet light.

TABLE 1. — Thin-layer chromatographic data for the unknown

substance in Psathyrophlyctis serpentaria (Culberson 1972;

Culberson & Johnson 1982)

A = benzene:dioxane:glacial acetic acid (180:45:5 by volume)

B = n-hexane:methyl rerf.-butyl ether:formic acid (140:72:18 by

volume)

C = toluene:glacial acetic acid (200:30 by volume)

FIGURE 8.— Psathyrophlyctis

serpentaria Brusse, section

of apothecium edge, show-

ing exciple (e), hypothe-

cium (a), and hymenium

(h). F. Brusse 3612, holo-

type. Bar = 100 pm.

184

Bothalia 17,2 (1987)

FIGURE 9. — Psathyrophlyctis serpentaria Brusse, scanning elec-

tron micrograph of the exciple, showing its sclerotized tis-

sue. F. Brusse 3612, holotype. Bar = 6 pm.

FIGURE 10. — Psathyrophlyctis serpentaria Brusse, asci and par-

aphyses. The ascus on the right is immature. F. Brusse

3612, holotype. Bar = 10 pm.

This new lichen superficially resembles a Phlyctis,

because of the J+ strongly blue, thick outer wall,

and hyaline muriform spores (Poelt 1969). However,

the ascospores of Phlyctis are thin-walled, and the

tholus is J— and not well developed (Poelt 1973).

The exciple is also truly paraplectenchymatous,

whereas this lichen only appears to be paraplecten-

chymatous, due to the dark brown interstices. The

exciple of Psathyrophlyctis is in fact scleroplecten-

chymatous, as can be seen from a scanning electron

micrograph of a section (Figure 9), and also from

sections in lactophenol cotton-blue stain, under the

light microscope (Figure 8).

The interrupted hymenium, and the sclerotized

exciple tissue reminds one of certain genera in the

Graphidaceae, such as Medusulina , but here the of-

ten elongated apothecia are seated in stromata and

the interruptions are of stromal tissue, not sterile

hymenial tissue. The ascospores of Medusulina are

thick-walled as well, but as is typical of this family,

the walls become brown and sometimes even blue-

brown in Lugol’s iodine solution (Redinger 1933).

The paraphyses of the Graphidaceae are generally

simple or only sparingly branched and Trentepohlia

is a common photobiont.

Psathyrophlyctis may be related to Phlyctidia

Mull. Arg. ex Zahlbr. (1907), but the ascospores are

only transversely septate in the latter (Zahlbruckner

1907, 1926).

As stated in the description, the thallus of this

lichen is completely sorediate and is quite thick, and

resembles the thallus of Lecidea crassa Nees ex Stiz.

somewhat. However the thallus of Lecidea crassa is

yellow, due to usnic acid, and also contains a series

of terpenes. Psathyrophlyctis serpentaria has an olive

green thallus, and looks yellowish in places under a

dissecting microscope, but no yellow pigments were

detected in an acetone extract of this lichen. Only

two substances were detected by TLC (Culberson

1972; Culberson & Johnson 1982) in P. serpentaria ,

one was a small grey terpene spot, and the other was

an evenly pale orange spot, after sulphuric acid and

heat treatment. The latter substance seems to be

new, and the TLC data for it is shown in Table 1.

This species is presently known only from the type

locality, the seaward slopes of the Langeberg Range

near Riversdale, on soil near rock. The slopes are

heavily covered with fynbos vegetation, and this may

explain its occurrence near rock, where some light

and perhaps some fire protection may be available.

REFERENCES

CULBERSON, C.F. 1972. Improved conditions and new data for

the identification of lichen products by a standardized thin-

layer chromatographic method. Journal of Chromatogra-

phy 72:113 -125.

CULBERSON, C.F. & JOHNSON, A. 1982. Substitution of

methyl tert. -butyl ether for diethyl ether in the standard-

ized thin-layer chromatographic method for lichen prod-

ucts. Journal of Chromatography 238: 483-487.

POELT, J. 1969. Bestimmungsschliissel europaischer Flechten.

Cramer, Lehre.

POELT, J. 1973. Classification. In V. Ahmadjian & M.E. Hale,

The Lichens: 599-632. Academic Press, New York, San

Francisco, London.

REDINGER, K. 1933. Die Graphidineen der ersten Regnell-

’schen Expedition nach Brasilien 1892-94. 1. Glyphis, Me-

dusulina und Sarcographa , Arkiv for Botanik 25A, 13:1-20.

ZAHLBRUCKNER, A. 1907. Specifier Teil. In A. Engler & K.

Prantl, Die natiirlichen Pflanzenfamilien , Teil 1, Abteilung

1: 49-249. Engelmann, Leipzig.

ZAHLBRUCKNER, A. 1926. Spezieller Teil. In A. Engler & K.

Prantl, Die natiirlichen Pflanzenfamilien , edn 2, vol. 8:

61-270. Engelmann, Leipzig.

F. BRUSSE*

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

Bothalia 17,2 (1987)

LILIACEAE

NOTES ON KNIPHOFIA

] 85

Kniphofia bruceae (Codd) Codd , stat. nov.

K. praecox Bak. subsp. bruceae Codd in Bothalia 9: 449 (1968),

Type: Cape, near Komga, Bruce 604 (PRE, holo.).

The plant described as K. praecox Bak. in Saund-

er’s Refugium Botanicum, 1. 169 (1870) came from

the garden of Mr Saunders and is recorded as having

been collected by Thomas Cooper at the Cape. An

examination of the type specimen in K, however, led

to the conclusion that the plant was of hybrid deri-

vation. As mentioned in Bothalia 9: 445 (1968), it

resembles specimens of unknown origin cultivated in

Europe since the early part of the nineteenth century

and which are preserved in several European her-

baria (e.g. BM, G, L) under names such as ‘Aletris

uvaria’, ‘Tritomanthe uvaria’ etc. They differ from

Kniphofia uvaria (and related robust species such as

K. linearifolia) in having longer and more acuminate

bracts and cannot be exactly matched by specimens

collected in the wild state. Similar hybrids are now in

cultivation throughout the world and two forms, a

summer-flowering and a winter-flowering red-hot

poker, are widely grown in South Africa and are

occasionally found as garden escapes.

It was further concluded that the narrow bracts

were contributed by a little-known robust Kniphofia

which has been recorded from three small disjunct

areas: (a) near Komga, (b) between Plettenberg Bay

and Knysna and (c) near Kouga in the Willowmore

District. Because of its affinity with and possible par-

entage of K. praecox , this Kniphofia was described

as K. praecox subsp. bruceae in 1968. However, it

may be separated from K. praecox on the basis of

several characters, for example the narrower, even

more acuminate bracts, longer pedicels, the some-

what shorter perianth which tends to be constricted

above the ovary, and the well exserted stamens. It is,

therefore, considered that a more satisfactory treat-

ment would be to accord separate species rank to the

wild plant.

In view of its rarity and limited distribution, it may

be questioned if K. bruceae could have been intro-

duced to Europe before or near the beginning of the

nineteenth century. Fortunately, such an introduc-

tion can be confirmed by the illustration in Jacquin’s

Fragmenta t.4 (probably 1800), reproduced in Bo-

thalia 9: 381 (1968). Although entitled ‘Veltheimia

uvaria’, it is not that species, but is an excellent

match of the type of K. bruceae from near Komga.

Note, for instance, the shape of the raceme, the

drawing of a separate flower showing the constric-

tion above the ovary and the well exserted stamens,

the acuminate bracts at the base of the raceme, and

the portion of an old raceme with numerous persist-

ent pedicels, all of which are characteristic of K. bru-

ceae (illustrated in Bothalia 9: 450, 451, 1968). Al-

though Jacquin does not record the origin of his

plant, there can be little doubt that it must have been

collected by Georg Scholl, collector for the

Schonbrunn Gardens, who spent from 1786 to 1799

in South Africa, accompanied during 1786 to Feb.

1787 by Franz Boos. Details of his itineraries are not

known but it may be accepted that his travels took

him eastwards as far as Kaffraria.

In Bothalia 9: 448 (1968), the few known speci-

mens from the Knysna - Willowmore areas were

included in K. praecox subsp. praecox. They are now

considered to belong rather in K. bruceae and, al-

though they differ somewhat from the Komga speci-

mens, they also differ among themselves, and more

material from these areas is desirable. Since the 1968

revision, only one additional specimen, Comins 1552

from Kei Road, a good match of the type of K. bru-

ceae, has been seen.

L.E. CODD*

* Botanical Research Institute, Private Bag X101,

Pretoria 0001.

POLYGONACEAE

A NEW VARIETY OF OXYGONUM ALATUM

Oxygonum alatum Burch., Travels in the in-

terior of southern Africa 1: 548 (1822). Type: South

Africa, Griqualand West, Burchell 2074 (K, holo.!;

PRE!).

Annual herb, up to 0,5 m tall, much branched

from the base. Stems prostrate, semi-erect or erect,

longitudinally ridged, reddish purple at the base,

sparingly or densely covered with white cup-shaped

or elongated scales. Ocrea membranous, funnel-

shaped, 6-9 mm long, greenish-white, covered with

cup-shaped or elongated scales, lobed; lobes tri-

angular, ending in a red or white rigid seta, 2-3,5

mm long. Leaves simple, fleshy, lanceolate, ovate or

rhomboid, (14—) 17—85 (-125) x (2-) 4—25 mm, en-

tire or shallowly to deeply lobed, the lobes usually

further divided, acute, narrowing and tapering to a

short petiole, glabrous or covered with white cup-

shaped or elongated scales especially along the mid-

rib and margins. Inflorescence a long lax thyrse with

fascicles of up to 4 flowers in the axils of ovate mem-

branous bracts. Flowers hermaphrodite and male.

Perianth pink or white, 5-cleft, the outer two seg-

ments keeled and mucronate, up to 7 mm long, gla-

brous or with a few cup-shaped or elongated scales.

Stamens 8, inserted on the inner perianth; anthers

bluish, 1 mm long; filaments white, 4 mm long, with

a ring of cilia one-eigth from the base. Styles 3,

joined for a third of their lengths; stigmas capitate.

Fruit a nut, winged at the three angles, sometimes

with small spreading prickles a third from the base,

red or yellow-brown, glabrous or with a few scat-

tered cup-shaped or elongated scales, 10-13 x

9-11 mm.

186

Bothalia 17,2 (1987)

In semi-arid grassland and open woodland on red

sandy soils, often locally common in disturbed places

and under trees, in SWA/Namibia, Botswana, the

western Transvaal and northern Cape Province.

Two varieties are recognized and are separated on

the presence or absence of cup-shaped or elongated

scales.

a. var. alatum

Oxygonum alatum Burch., Travels in the interior of southern

Africa 1: 548 (1822); Meisn.: 38 (1856); Hook, f.: 14 (1880);

Engl.: 6 (1888); Schinz: 57 (1896); Hiern: 902 (1900); Schinz: 870

(1901); Dinter: 58 (1909); C.H. Wr.: 460 (1912); Bak. & C.H.

Wr.: 99 (1909); Burtt-Davy: 167 (1926); R.A. Grah.: 165 (1957);

Tikovsky & Merxm.: 3 (1969). Type: South Africa, Griqualand

West, Burchell 2074 (K, holo.!; PRE).

O. alatum var. incisum Sond.: 98 (1850), ex descr.

O. alatum var. integrifolium Sond.: 98 (1850), ex descr., excl.

syn. O. dregeanum Meisn.

O. alatum var. marlothii Engl.: 6 (1888); C.H. Wr.: 460 (1912).

Type: Cape (British Bechuanaland), Kuruman, Marloth 1016

(BOL, iso.!; GRA!, K!, SAM!).

Only cup-shaped scales are present scattered over

the whole plant. Figure 11. Distribution and ecology

more or less as for the species. Figure 12.

Vouchers: Germishuizen 2624 (PRE); 2674 (PRE, WIND);

Leistner 1366 (NBG, PRE); Lewis 4204 (PRE, SAM, STE-U);

Maguire 2225 (BOL, NBG, PRE, WIND).

FIGURE 11. — Oxygonum alatum var. alatum

(Germishuizen 2624 , PRE). Cup-shaped scales

on abaxial leaf surface, x 110.

b. var. longisquamatum (Germishuizen, var.

nov., a varietate typico (solum) squamis elongatis

differt.

TYPE. — SWA/Namibia, near Oshandi 25 km

SE of Oshikango, R.J. Rodin 9104 (WIND, holo.).

Figure 13.

The whole plant is covered with both cup-shaped

and elongated scales in varying density. Figure 14.

Found in northern South West Africa/Namibia

and northern Botswana. Figure 15.

SWA/NAMIBIA. — 1715 (Ondangua): Ogongo Agricultural

College (-CB), E.J. van Jaarsveld 2862 (NBG). 1716 (Enana):

Oshandi, 25 km SE of Oshikango (-AC), R.J. Rodin 9104

(WIND); Oshigambo (-CC), 5. Soini s.n. (PRE, WIND). 1920

(Tsumkwe): 6 km E of Tsumkwe (-DA), W. Giess, J.S. Watt &

J. Snyman 11024 (WIND); 16 km E of Tsumkwe (-DA), W.

Giess, J.S. Watt & J. Snyman 11171 (WIND).

BOTSWANA. — 1823 (Siambusso): Molapo, 2 km NW of James

Camp (-CB), D.T. Williamson 81 (PRE). 1926 (Tsekanyani):

Nata area at Nata River Delta (-CD), J.F. Ngoni 488 (PRE).

2124 (Rakops): Maun-Rakop, 16 km W of Rakop (-AB), H.J.

van Rensburg B4099 (PRE); Kalatraverse, 12 km W of eastern

boundary of Central Kalahari Game Reserve (-AC), A.R. Kreu-

len 644 (PRE). 2125 (Lothlekane): Orapa (-AD), O. Kerfoot

7752 (PRE). 2126 (Tlada Mabeli): Tlada Mabeli-Masu area, near

Soa Pan (-AA), J.F. Ngoni 308 (PRE).

The epithet longisquamatum refers to the elong-

ated scales scattered over the whole plant, which can

be seen with the naked eye.

FIGURE 12. — Distribution of Oxygonum alalum var. alatum in

southern Africa.

FIGURE 13. — Holotype of Oxygonum alatum var. longisqua-

matum Germishuizen.

Bothalia 17,2 (1987)

187

FIGURE 14. — Oxygonum alatum var. longisquama-

tum Germishuizen ( Giess 11171, WIND).

Elongated scales present on stem surface, x

230.

FIGURE 15. — Distribution of Oxygonum alatum var. longis-

quamatum in southern Africa.

ACKNOWLEDGEMENTS

I would like to thank Mrs S. Perold for helping

with the SEM work, Mrs A. Romanowski for the

photographs and Dr H.F. Glen for translating the

diagnosis into Latin.

REFERENCES

BAKER, J.G. & WRIGHT, C.H. 1909. Polygonaceae. In W.T.

Thiselton-Dyer, Flora of tropical Africa 6,1: 99-103.

Reeve, Kent.

BURCHELL, W.J. 1822. Travels in the interior of southern Africa

1: 548. Longman, Hurst, Rees, Orme & Brown, London.

BURTT DAVY, J. 1926. Polygonaceae. A manual of the flow-

ering plants and ferns of the Transvaal with Swaziland,

South Africa 1: 167-168. Longmans, Green, London.

DINTER, K. 1909. Deutsch-Sudwest-Afrika Flora forst- und land-

wirtschaftliche Fragmente : 58. Weigel, Leipzig.

ENGLER, A. 1888. Plantae marlothianae. Ein Beitrag zur

Kenntnis der Flora Siidafrikas. Botanische Jahrbucher

10:6.

GRAHAM, R.A. 1957. A revision of Oxygonum. Kew Bulletin

1: 145-172.

HIERN, W.P. 1900. Polygonaceae. Catalogue of the African

plants collected by Dr Friedrich Welwitsch in 1853-1861 :

902. London.

HOOKER, J.D. 1880. Hooker’s icones plantarum 14: 14, t. 1321.

Williams & Norgate, London.

MEISNER, C.F. 1856. Polygonaceae. In A. de Candolle, Pro-

dromus systematis universalis regni vegetabilis 14: 1-186.

Paris.

SCHINZ, H. 1896. Polygonaceae. Die Pflanzenwelt Deutsch-

Siidwest-Afrikas. Bulletin de I’Herbier Boissier 4, App. 3:

56-57.

SCHINZ, H. 1901. Polygonaceae. Bulletin de I'Herbier Boissier

ser. 2,1: 869-872.

SONDER, O.W. 1850. Polygonaceae. Linnaea 23:98.

TIKOVSKY, H. & MERXMULLER, H. 1969. Polygonaceae. In

H. Merxmiiller, Prodromus einer Flora von Siidwestafrika

23: 1-10. Cramer, Lehre.

WRIGHT, C.H. 1912. Polygonaceae. In W.T. Thiselton-Dyer,

Flora capensis 5,1: 459-482. Reeve, London.

G. GERMISHUIZEN

TRAPELIACEAE

A NEW SPECIES OF TRAPELIA (LICHENES) FROM SOUTHERN AFRICA

Trapelia chiodectonoides Brusse, sp. nov.

Thallus crustosus, saxicola, cretaceus vel pallide

cinereus, pannum grandem continuum circa 200 mm

diametro efficiens, rimosus vel rimoso-areolatus,

areolis 0, 2-4,0 mm diametro, vulgo circa 1 mm di-

ametro, laevis, 0,2-1 ,0 mm crassus, rimis usque ad

0,1 mm latis. Superficies laevis, subnitida, isidiis so-

rediisque destituta. Cortex superior 10-17 pm cras-

sus, paraplectenchymatus, cellulis 4—8 pm diametro.

Stratum gonidiale 35-50 pm crassum; algae ad Chlo-

rococcales pertinentes, 5-15 pm diametro. Medulla

alba, C+ rubra. Apothecia rubiginosa, usque ad

0,5 mm diametro, lecanorina sed marginibus mox

fatiscentibus, dein lecideina, in maculis circularibus,

2-5 mm diametro, super areolis grandibus et pustu-

latis aggregata. Excipulum thallinum evanescens.

Excipulum pallide porphyreum, in lateribus 20 pm

crassum, infra degenerum. Hypothecium hyalinum,

40-100 pm crassum, cellularum isodiametrarum

compactarum, 3-7 pm diametro compositum. Hy-

menium hyalinum, 90-110 pm altum, J+ pallide cae-

ruleum. Paraphyses septatae, ramosae, liberae,

1,1-1 ,3 pm crassae. Asci cylindrici vel fusiformo-cyl-

indrici, 75-100 x 15-22 pm, tholis umbonatis, J +

perpallide caeruleis, fere hyalinis (Figura 17). Asco-

sporae octonae, hyalinae, simplices, ellipsoideae,

16-21 x 8-10,5 pm. Pycnidia non visa. Thallus aci-

dum gyrophoricum continens.

TYPE. — Natal, 2829 (Harrismith): Cathedral

Peak Reserve, Doreen Falls area, near lower falls,

on steep E slope, on SE face of Clarens sandstone

boulder in grassland, alt. 1 680 m (-CC), F. Brusse

4540 , 1986.01.22 (PRE, holo.; BM, LD, iso.). Fig-

ure 16.

Thallus crustose, saxicolous, ashy or chalky whit-

ish, growing in large continuous patches up to

188

Bothalia 17,2 (1987)

FIGURE 16. — Trapelia chiodectonoides , habit. F. Brusse 4540,

isotype. Scale in mm.

FIGURE 17. — Trapelia chiodecto-

noides, ascus and paraphyses.

F. Brusse 4540, holotype. Bar

= 10 pm.

200 mm across, rimose to rimose-areolate, areoles

0, 2^4,0 mm across, commonly 1 mm across, smooth,

0,2-1 ,0 mm thick, cracks up to 0,1 mm wide. Surface

smooth, subnitid, not isidiate or sorediate. Upper

cortex 10-17 pm thick, paraplectenchymatous, cells

4—8 pm diam. Algal layer 35-50 pm thick; algae

Chlorococcalean, 5-15 pm diam. Medulla white, C+

red. Apothecia brownish red, up to 0,5 mm diam.,

lecanorine but margins soon disintegrating, becom-

ing lecideine, aggregated into circular spots, 2-5 mm

across, on large pustular areoles (Figure 16). Thai-

line exciple evanescent. Exciple pale reddish brown,

20 pm thick on flanks, degenerate below. Hypothe-

cium hyaline, 40-100 pm thick, composed of com-

pacted isodiametric cells, 3-7 pm diam. Hymenium

hyaline, 90-110 pm high, J+ pale blue. Paraphyses

septate, branched, free, 1,1-1 ,3 pm thick. Asci

eight-spored, cylindrical to fusiform-cylindrical,

75-100 x 15-22 pm, tholus bossed, J+ very pale

blue almost hyaline (Figure 17). Ascospores hyaline,

simple, ellipsoid, 16-21 x 8-10,5 pm. Pycnidia not

seen. Chemistry, gyrophoric acid present.

Etymology: the specific epithet is derived from the

Greek generic name Chiodecton and the Greek suf-

fix, - oides , which indicates resemblance, because the

aggregated apothecia superficially resemble those of

some Chiodecton species. The apothecia of Chiodec-

ton are, however, very different, being embedded in

stromatic tissue, which is often sclerotic.

The apothecial habit of this new species is unique

in Trapelia, being an unknown trait in this genus

until now (Coppins & James 1984; Hertel 1969,

1970, 1977). The thallus is also strongly developed

and coherent, a condition not common in this genus

(Coppins & James 1984; Hertel 1969, 1977).

The internal anatomy and dimensions of T. chio-

dectonoides apothecia resemble those of the wide-

spread T. coarctata (Sm. & Sow.) Choisy rather

closely, but seem slightly smaller in all respects.

T. coarctata apothecia are not aggregated into sub-

pustular spots, and the hymenial surface is normally

brown and larger. The upper layer of the thallus of

T. coarctata is undifferentiated (Hertel 1977),

whereas T. chiodectonoides has a distinctly paraplec-

tenchymatous upper cortex.

The chemistry of this new species is typical for a

member of the Trapeliaceae, most members tested

producing gyrophoric acid as the only constituent

(Hertel & Leuckert 1969).

Trapelia chiodectonoides is presently known only

from the type locality, the Doreen Falls area of the

Cathedral Peak Reserve in the Natal Drakensberg,

on Clarens (Cave) sandstone.

REFERENCES

COPPINS, B.J. & JAMES, P.W. 1984. New or interesting British

lichens V. The Lichenologist 16,3: 241-264.

HERTEL, H. 1969. Die Flechtengattung Trapelia Choisy. Herzo-

gia 1: 111-130.

HERTEL, H. 1970. Trapeliaceae — eine neue Flechtenfamilie.

Vortrdge aus dem Gesamtgebiet der Botanik (n.s.) 4:

171-185.

HERTEL, H. 1977. Gesteinsbewohnende Arten der Sammelgat-

tung Lecidea (Lichenes) aus Zentral-, Ost- und Siidasien.

Eine erste Ubersicht. Khumbu Himal 6,3 : 145-378.

HERTEL, H. & LEUCKERT, C. 1969. Ober Flechtenstoffe und

Systematik einiger Arten der Gattungen Lecidea, Placop-

sis und Trapelia mit C+ rot reagierendem Thallus. Willde-

nowia 5: 369-383.

F. BRUSSE*

* Botanical Research Institute, Private Bag X101,

Pretoria 0001.

Bothalia 17,2 (1987)

ZYGOPHYLLACEAE

A NEW SPECIES OF ZYGOPHYLLUM FROM SOUTHERN AFRICA

189

Zygophyllum macrocarpon Retief, sp. nov., Z.

foetido Schrad. & Wendl. affinis sed fructibus adum-

bratione oblongis non subglobosis alabastris dense

pubescentibus differt.

TYPE. — Cape, 2817 (Vioolsdrif): Richters-

veld, Kodaspiek, main ridge south east of beacon

and up to summit (-AA), Oliver, Tolken & Venter

417 { PRE, holo.; K, MO).

A glabrous succulent shrub up to 1,8 m high.

Stems woody, sparsely branched. Leaves fleshy, op-

posite, petiolate, bipinnate, broadly ovate, variable

in size, 27-60 x 24-58 mm, apex rounded, base as-

ymmetrical, margin entire; petioles 5-10 mm long.

Interpetiolar stipules ovate to broadly ovate, 2-5 x

2-3 mm. Flowers 2 together in the axil of a leaf.

Sepals 5, persistent, ovate, ±5x3 mm, connate at

the base, margins hairy. Petals 5, obovate, 15-20 x

8-12 mm, yellow with purplish brown or red mark-

ings in the throat. Disc fleshy, 10-angled. Stamens

10, inserted at the disc base; filaments terete, ±

7-7,5 mm long; anthers ±1,5 mm long; appendages

undivided, margin fimbriated, 0,25 times as long as

the filament. Ovary sessile on the disc. 4-5-locular

with several pendulous ovules in each loculus; style

terete, ± 5 mm long; stigma minute. Fruit a 4-5-

angled capsule, 5-locular with several seeds, oblong,

35-50 x 71-21 mm. Seeds crugtaceous, ± 5 mm

long, brownish black. Figure 18.

SWA/NAMIBIA.— 2716 (Witputz): Udabib Mountains (-BB),

Muller 799 (PRE); Aurus Mountain (-CA), Muller 736 (PRE);

Farm Spitzkopp (-DC), Giess 13045 (PRE).

CAPE. — 2816 (Oranjemund): Sendelingsdrif (-BB), Van der

Westhuizen 133180 (PRE); head of Helskloof, Hottentotparadys

Mountain (-BD), Thompson & Le Roux 121 (PRE, STE). 2817

(Vioolsdrif): Richtersveld, Kodaspiek, main ridge south east of

beacon and up to summit (-AA), Oliver, Tolken & Venter 417 (K,

MO, PRE).

Z. macrocarpon is found on mountain slopes in

the Richtersveld and the southern part of South

West Africa/Namibia. The species grows in well

drained stony clayey or sandy loamy soil. According

to Giess (1971) in South West Africa/Namibia this

species occurs in Desert and Succulent Steppe (Win-

ter rainfall area). In the Cape Z. macrocarpon is

found in two different veld types, namely, in Succu-

lent Karoo and Namaqualand Broken Veld (Acocks

1975). These veld types are dominated by succu-

lents, mainly species of Mesembryanthemaceae to-

gether with a few trees or large shrubs. According to

herbarium records Z. macrocarpon flowers and

fruits from July to September.

Z. macrocarpon is most closely related to Z. foeti-

dum Schrad. & Wendl. The leaves of these two

species are broadly ovate and prominently petiolate.

The flowers are also similar in structure and size. Z.

foetidum differs from Z. macrocarpon in having

hairy flowerbuds. The fruits of Z. macrocarpon are

oblong in outline while in Z. foetidum they are sub-

globose. These two species also differ in their distri-

bution. Z. foetidum occurs from south of Springbok

FIGURE 18. — Zygophyllum macrocarpon. I, flowering branch,

x 1; 2, fruiting branch, x 1 (Oliver, Tolken & Venter 417).

190

Bothalia 17,2 (1987)

to the eastern Cape, whereas Z. macrocarpon is

found north of Springbok.

The specific epithet ‘macrocarpon’ refers to the

size of the fruit. These are large, oblong, and 5-

angled which, together with the conspicuously large

bipinnate leaves and flowers, distinguish Z. macro-

carpon from other southern African species of Zygo-

phyllum. The same fruit form is found in Z. debile

Cham. & Schlechtd. These fruits are, however,

small, 13-20 mm long, compared with Z. macrocar-

pon of which the fruits are 35-50 mm long.

Z. foetidum Schrad. & Wendl., Z. leptopetalum E.

Mey. ex Sond. and Z. meyeri Sond. also have large

bipinnate leaves but differ from Z. macrocarpon in

that their fruits are subglobose in outline. The petals

of Z. macrocarpon are 15-20 mm long, and there-

fore large in comparison with, for example, those of

Z. dregeanum which are 3-4 mm long.

The flowers of Z. fulvum are similar in size to

those of Z. macrocarpon , but differ in having leaflets

that are elliptic in shape and not broadly ovate as in

the case of Z. macrocarpon.

Based on the fruit dehiscence, the genus Zygo-

phyllum can be divided into two subgenera, namely,

Agrophyllum Endl. and Zygophyllotypus Van

Eluysteen. The dehiscence of the subgenus Agro-

phyllum is septicidal, whereas loculicidal dehiscence

is found in the subgenus Zygophyllotypus. The

species of this latter subgenus, indigenous in south-

ern Africa, are grouped in different sections. Z.

macrocarpon is placed in the section Capensia Engl,

to which the species Z. foetidum also belongs.

REFERENCES

ACOCKS, J.P.H. 1975. Veld types of South Africa. Memoirs of

the Botanical Survey of South Africa No. 40.

GIESS, W. 1971. A preliminary vegetation map of South West

Africa. Dinteria 4: 8-9.

E. RETIEF*

* Botanical Research Institute, Private Bag X101,

Pretoria 0001.

Bothalia 17,2: 191-194 (1987)

Taxonomy of the genus Ehrharta (Poaceae) in southern Africa: the

Villosa group

G. E. GIBBS RUSSELL*

Keywords: Fynbos, Capensis, Ehrharta, Poaceae, Succulent Karoo, taxonomy

ABSTRACT

The Villosa species group in the genus Ehrharta Thunb. is differentiated morphologically by very large, pro-

fusely hairy, bearded and aristate spikelets and by a suffrutescent habit, with culms woody at the base and with

reduced leaf blades. The Villosa group is composed of two species, one with a variety: E. thunbergii Gibbs Russell,

nom. nov., E. villosa Schult. f. var. villosa and E. villosa var. maxima Stapf. Members of the group occur on sandy

soils in the Succulent Karoo and Fynbos Biomes, along the west coast in Strandveld and. on the southern coast as

far east as the Fish River. Morphologically, the group appears to be related to the Calycina and Capensis groups.

UITTREKSEL

Die Villosa-groep in die genus Ehrharta Thunb. word morfologies onderskei deur baie groot, dig behaarde,

bebaarde en skerppuntige blompakkies asook halfstruikagtige habitus, met halms houtagtig aan die basis en met

gereduseerde blaarlaminas. Die Villosa-groep sluit twee spesies in, een met ’n varieteit: E. thunbergii Gibbs

Russell, nom. nov., E. villosa Schult. f. var. villosa en E. villosa Schult. f. var. maxima Stapf. Lede van die groep

kom voor op sanderige gronde in die Sukkulente Karoo- en Fynbosbiome, langs die weskus in die Strandveld en

aan die suidelike kus so ver oos as die Visrivier. Morfologies vertoon die groep verwantskap met die Calycina- en

Capensis-groepe.

INTRODUCTION

Previous papers in this series have outlined the

seven species groups of Ehrharta Thunb. in southern

Africa (Gibbs Russell & Ellis 1987), and dealt in

detail with the morphology and anatomy of the taxa

in the Setacea group (Gibbs Russell 1987; Ellis

1987a). In the Villosa group, as in all the groups

except the Setacea group, the first and second sterile

lemmas are of similar size and ornamentation (Fig-

ure 1). The Villosa group is distinguished morpholo-

gically by large spikelets 10-18 mm long that have

sterile lemmas with conspicuously bearded bases,

profusely hairy sides and mucronate to aristate tips;

and by a suffrutescent habit with culms woody at the

base and with reduced leaf blades. Besides these

characters of habit and spikelet, the two species

share features in leaf anatomy (Ellis 1987b). Within

the group, the species differ mainly in spikelet size,

relative glume length, rhizome structure, and in

habitat and distribution, as well as in anatomical

characters (Ellis 1987b). All the taxa are robust and

have long, stout rhizomes. They occur only in sandy

soil, and are the only species of Ehrharta to grow on

seaside dunes.

Because of the spikelet similarities, Chippindall

(1955) questioned the level of treatment of the three

taxa, and only described E. villosa in detail, includ-

ing within it some of the distinctive vegetative

characters of E. thunbergii. However, previous

treatments from Schultes (1830) to Stapf (1900) dis-

tinguished two taxa (E. villosa Schult. f. and E. gi-

gantea Thunb.) on spikelet and rhizome characters

similar to those used here.

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

It is unfortunate that a new name was required for

the species previously known as E. gigantea Thunb.

For 194 years, since 1794, the epithet ‘gigantea' has

been applied to the smaller, inland specimens of the

Villosa group. However, this name is based on the

same type, the specimen in the Thunberg herbar-

ium, as Linnaeus the Younger’s (1781) Aira villosa,

and therefore, as a superfluous name, was illegiti-

mate when published. The epithet ‘villosa’ cannot be

taken up, however, because the new combination

was not made in Ehrharta before Schultes (1830)

published his own Ehrharta villosa, based on a dif-

ferent type. No other name has been applied to the

species, so a new name is necessary. It is fortunate

that ‘villosa’ was already occupied in Ehrharta, or

the species previously known as E. gigantea would

have had to become E. villosa , and E. villosa would

have required a new name, an even more unsatisfac-

tory situation.

FIGURE 1. — Spikelet of E. villosa var. villosa (Crook 2260,

PRE): a, whole spikelet; b, glumes; c, first sterile lemma;

d, fertile lemma; e, second sterile lemma; all x 2.

192

Bothalia 17,2 (1987)

METHODS

The descriptive data for the species and infraspeci-

fic taxa has been recorded through the DELTA

computer system, originated by Watson and Dall-

witz (Dallwitz 1984, Watson & Dallwitz 1980, 1985)

and used previously at species level by Webster

(1983). The system has the advantage of recording

comparable character states for each taxon in a form

that can be used to generate descriptions, keys and

an online identification facility, and that can be used

directly for classificatory studies. Because of this

change in methodology, the descriptions given from

now on include more characters than were pre-

viously recorded, and are strictly comparable.

KEY TO SPECIES IN THE VILLOSA GROUP

Glumes 1-f as long as spikelet, 5-nerved, upper glume to

8 mm long; spikelets 8-10 mm long; rhizomes

densely covered with hairy cataphylls, internodes

often sub-bulbous 1. E. thunbergii

Glumes f as long to about equalling spikelet, 5-9-nerved,

upper glume 8-13 mm long; spikelets (10—)

11-18 mm long; rhizomes naked, neither sub-bul-

bous nor bulbous 2. E. villosa

1. Ehrharta thunbergii Gibbs Russell, nom.

nov. Type: Thunberg, (sheet 8851, UPS, holo.-PRE,

microfiche!).

Aira villosa L. f., Supplementum plantarum: 109 (1781). Melica

gigantea Thunb.: 21 (1794). Ehrharta gigantea (Thunb.) Swartz:

58 (1802); Thunb.: 339 (1818); Schrader; 2079 (1821); Thunb.:

336 (1823); Schult.: 1375 (1830); Trinius: 16 (1839); Nees: 216

(1841); Steudel: 5 (1855); Stapf: 680 (1900); Chippindall: 45

(1955). Type; Thunberg, (sheet 8851, UPS.-PRE microfiche!).

Ehrharta gigantea (Thunb.) Swartz var. neesii Stapf: 680 ( 1900).

Type: Drege, near Riebeecks Castle, among shrubs (K, holo.!).

Ehrharta gigantea (Thunb.) Swartz var. stenophylla Stapf: 881

(1900). Type: Schlechter 9058 (K, holo.!).

Perennial, tufted, erect, long-rhizomatous, ro-

bust, suffrutescent. Rhizomes often with internodes

sub-bulbous, clad with imbricate, thickened, hairy

cataphylls. Culms several, to 1 ,5 m tall, 3 mm across,

woody, solid, sometimes geniculate at lower nodes,

crowded, branched at base, sometimes branching in

fascicles above, nodes usually black, rarely with as-

cending hairs, the lowest internode sometimes sub-

bulbous, the ‘bulb' pale orange, polished, shining.

Young shoots intravaginal. Leaves mostly basal, per-

sistent, culm leaves usually with blades reduced, of-

ten auriculate from mouth of sheath, the auricles

rarely accrescent, to 10 mm long, with bristly edges;

basal sheaths loose, papery, often splitting into

fibres, grey or whitish; culm sheaths not overlap-

ping; ligule a fringed membrane 0,5 mm long; blades

persistent, linear, to 5 mm across, flat or rolled,

gradually tapering at the tip, erect or spreading, her-

baceous, usually glabrous, but sometimes hairy. In-

florescence a fascicled panicle, narrow, often sin-

uous, to 160 mm long, somewhat overtopping

leaves, exserted from uppermost leaf sheath, of nu-

merous spreading spikelets. Spikelets pedicellate on

filiform curled pedicels, laterally compressed,

8-10 mm long, 2-3 mm across above glumes.

Glumes keeled, subequal, 5-3 as long as the spikelet,

translucent, sometimes purple-tinged; lower glume

to 6 mm long, 5-nerved, acute; upper glume to 8 mm

long, 5-nerved, acute. Florets with lemmas decidedly

firmer than the glumes, keeled. Sterile lemmas

slightly rounded on sides, similar in shape and tex-

ture. First sterile lemma §-f length of second sterile

lemma, with keel and margins parallel; base substi-

pitate, conspicuously bearded; sides with profuse

long spreading white hairs, otherwise smooth and

unornamented, dull; tip abruptly aristate from keel,

aristae to 3 length of lemma, usually dark purple.

Second sterile lemma similar to first sterile lemma,

but larger and distinctly stipitate. Fertile floret

shorter than second sterile lemma, lemma differing

from sterile lemmas, strongly laterally compressed,

sides apparently nerveless, sparsely hairy, tip trun-

cate; palea f or more as long as lemma, keeled,

1-nerved. Lodicules 2, membranous, 2-lobed. Sta-

mens 6, anthers 5 mm long, white. Stigmas white.

Caryopsis not seen.

This species is distinguished from other Ehrharta

species by the large hairy spikelets with short, trans-

lucent glumes and the sub-bulbous rhizomes with

hairy cataphylls. It is the most widespread and abun-

dant of the members of the Villosa group, and its

distribution is shown in Figure 2. Unlike most Ehr-

hartas, which tend to have an ‘eastern’ or ‘western’

distribution, its range extends both northwards

through Fynbos, Strandveld, and Succulent Karoo

to the southern border of South West Africa/Nami-

bia and eastward along the coastal ranges as far as

Mossel Bay. However, it appears to be less common

in the eastern part of the range.

Stapf’s two varieties are not recognized because

they intergrade with the more common and wide-

spread form, and because they are not distinguished

by a separate range or habitat. However, each of

these forms shows an interesting link to the Capensis

group. ‘Bulbous’ basal internodes (var. stenophylla)

are characteristic of most species in the Capensis

group, and the specimens with many-noded genicu-

late culms with small flat leaf blades (var. neesii ) are

similar in habit to the only non-bulbous member of

the Capensis group, E. barbinodis. There is in ad-

dition a form with broad and often hairy leaves (for

example, Gibbs Russell 5619 ) that occurs from Cal-

vinia to Lambert’s Bay and Piketberg. A few speci-

mens intermediate to E. villosa var. villosa are dis-

cussed under that variety.

E. thunbergii grows in sandy or sandy gravel soils,

mostly on hill slopes, but also in coastal sand and

14° 16° 18° 20° 22° 24° 26°

FIGURE. 2. — Distribution of E. thunbergii.

Bothalia 17,2 (1987)

193

occasionally beside watercourses. Flowering occurs

from September to December.

Vouchers: Acocks 23393; Andreae 1314; Boucher 4724; Lieben-

berg 6551; Schlechter 10208.

2. Ehrharta villosa Schult. f. in Systema vegeta-

bilium 7,2: 1374 (1830). Trinius: 16 (1839); Nees:

213 (1841); Steudel: 5 (1855); Stapf: 681 (1900);

Chippindall: 45 (1955); Smook & Gibbs Russell: 55

(1985). Type: Ecklon, Promont. b. spei, in arenosis

maritimis (P, holo.).

Perennial, tufted, erect, long-rhizomatous, ro-

bust, suffrutescent. Rhizomes woody, naked. Culms

several, to 1,5 m tall, woody, solid, branched at

base, branching in fascicles above, nodes glabrous,

lowest internode never bulbous. Young shoots intra-

vaginal. Leaves not basally aggregated, culm leaves

with blades reduced, often auriculate from mouth of

sheath, the auricles not accrescent; basal sheaths

loose, papery, grey or whitish; culm sheaths not

overlapping; ligule a fringed membrane 0,5 mm

long; blades deciduous or persistent on basal sheaths

and culm sheaths, linear, to 8 mm across, but rolled

and appearing setaceous, gradually tapering at the

tip, herbaceous, glabrous. Inflorescence a fascicled

panicle, narrow, sometimes sinuous, 40-260 mm

long, somewhat overtopping leaves, exserted from

or sometimes closely subtended by uppermost leaf

sheath, of numerous spreading spikelets. Spikelets

pedicellate on filiform curled pedicels, laterally com-

pressed, (10—) 11-18 mm long, 3-4 mm across above

glumes. Glumes keeled, more or less equal, j as long

to slightly longer than rest of spikelet, opaque white,

sometimes purple-tinged; lower glume 8-13 mm

long, 5-9-nerved, acute; upper glume 9-18 mm long,

5-9-nerved, acute. Florets with lemmas decidedly

firmer than the glumes, keeled. Sterile lemmas

slightly rounded on sides, similar in shape and tex-

ture. Sterile lemmas similar to E. thunbergii, but

first sterile lemma with tip abruptly mucronate or

aristate from keel, purple or pale. Second sterile

lemma similar to first sterile lemma, but larger and

distinctly stipitate. Fertile floret similar to E. thun-

bergii but anthers 6-8,5 mm long, brownish yellow.

Caryopsis 5,5 mm long, ovate, flattened.

This species grows on sea dunes from Lambert’s

Bay to Port Alfred, and is our only indigenous grass

that behaves as a ‘marram’. It is distinguished from

E. thunbergii by its naked rhizomes and longer

spikelets with comparatively longer glumes.

KEY TO VARIETIES

Inflorescence exserted from uppermost leaf sheath, the

sheath usually not inflated; upper glume 9-13 mm

long; culms to 3 mm across 2a. E. villosa var. villosa

Inflorescence closely subtended or enveloped by inflated

uppermost leaf sheath; upper glume (10-)

13-18 mm long; culms to 5 mm across

2b. E. villosa var. maxima

2a. var. villosa. Stapf: 681 (1900); Chippindall:

45 (1955); Smook & Gibbs Russell: 55 (1985).

Culms robust, to 3 mm across. Inflorescence inter-

rupted, to 150 mm long, usually exserted from up-

permost leaf sheath, the sheaths usually not inflated.

Spikelets 11-14 mm long, to 3 mm across laterally

above glumes. Glumes I as long to slightly shorter

than rest of spikelet; lower glume 8—12 mm long;

upper glume 9-13 mm long. Florets with sterile lem-

mas mucronate or aristate. Anthers 7,5-8 mm long.

Variety villosa is distinguished from var. maxima

by its exserted inflorescences, smaller stature and

somewhat smaller spikelets. The two varieties are

sympatric throughout most of their distribution (Fig-

ures 3 & 4), although this one extends farther north

on the west coast and is more common in the south-

western Cape.

A few intermediate specimens occur, always near

the sea but not on seaside dunes, that have small

glumes and spikelets like E. thunbergii, but naked

rhizomes like E. villosa : Ellis 4640 from Lamberts

Bay; Gibbs Russell 5670 from the Cape Peninsula;

and Liebenberg 4015 from Mossel Bay. Most of the

specimens from Mossel Bay, to the east of the range

of both this variety and of E. thunbergii, appear to

be of this intermediate type, although these inter-

mediates are few and sporadic elsewhere. Inland,

rare intermediate specimens show a different pattern

of variation. Ellis 4693 , from the roadside at Cloete’s

Pass, has the hairy rhizome cataphylls of E. thunber-

gii and the larger spikelets with long glumes of E.

villosa. Emdon 200, from disturbed Fynbos, has

large glumes that are translucent.

FIGURE. 3. — Distribution of E. villosa var. villosa. Interme-

diates to E. thunbergii are shown by triangles, A . Interme-

diates to E. villosa var. maxima are shown by squares, □.

Where symbols would be superimposed, those for interme-

diates are shown apparently in the sea directly below or to

the left of the correct quarter degree.

FIGURE. 4. — Distribution of E. villosa var. maxima.

194

Bothalia 17,2 (1987)

The intermediates, all mapped in Figure 3 with E.

villosa var. villosa, occur either at the extremeties of

the distribution of this variety, or in disturbed habi-

tats, and all occur within the range of E. thunbergii.

Furthermore, no intermediates are known from the

part of the range of E. thunbergii that lies outside

that of E. villosa. Therefore, the hypothesis that

most likely accounts for the intermediates is that

they are the result of hybridization between the two

taxa.

Variety villosa grows most commonly on seaside

dunes, but has been collected as far as 1 km inland

on the limestone ridges at De Hoop, although still in

sandy soil. It flowers from October to December.

Vouchers: Bohnen 4541 ; Cleghorn 3122; Crook 2260; Marloth

3046; Smith 4649.

2b. var. maxima Stapf in Flora capensis 7: 681

(1900). Chippindall: 45 (1955); Smook & Gibbs Rus-

sell: 55 (1985).

Culms very robust, to 5 mm across. Inflorescence

dense, to 260 mm long, closely subtended or envel-

oped by uppermost inflated leaf sheath. Spikelets

(10-) 12-18 mm long, to 4 mm across laterally above

glumes. Glumes slightly shorter to slightly longer

than rest of spikelet; lower glume 9-13 mm long;

upper glume (10—) 13-18 mm long. Florets with first

sterile lemma tip mucronate. Anthers 6-8,5 mm

long.

The distribution of this variety is shown in Figure

4. It extends as far east as Port Alfred, where it is the

only Ehrharta on the dunes, but extends no farther

than the Cape Peninsula. It is less common than var.

villosa in the western part of its range, where it is

known from a few specific sites, such as Robben

Island. It is recognized at varietal rather than sub-

specific rank because of the close similarity between

the two taxa in morphology, anatomy, distribution

and habitat.

Variety maxima occurs only on seaside dunes, and

flowers from September sporadically to March.

Vouchers: Boucher 1689; Britten 778; Theron 1108; Tyson

Herb. Marl. No. 8598; UPE Staff 158.

CONCLUSION

This closely related group of species shows a dine

in size, from the smaller, aristate forms of E. thun-

bergii on Namaqualand hillsides, to the robust,

short-mucronate, dune-binding E. villosa var. max-

ima at Port Alfred. E. villosa var. villosa , which is

concentrated geographically in the middle part of

the distribution of the group, also has morphological

and anatomical (Ellis 1987b) intermediates to both

the other taxa. The rhizome differences between the

species may be related to habitat differences. In the

shifting dune sands there are no cataphylls and no

bulb-like structures.

Relationships of the Villosa group to other species

groups appear somewhat different when examined

morphologically and anatomically. Morphologically,

the Villosa group is linked to the Calycina group by

the hairy spikelets with long glumes, and to the Ca-

pensis group by the large spikelets with short-awned

or mucronate and stipitate sterile lemmas, and by

the ‘bulbous’ internodes. Ellis (1987b) reports ana-

tomical links to the Calycina group, which is similar

anatomically to the Capensis group. However, the

suggested anatomical link to the Ramosa group is

shown morphologically in only one species in the

Ramosa group, which has long glumes and suffrutes-

cent culms.

ACKNOWLEDGEMENTS

I thank the Directors of the following herbaria for

loans of specimens BOL, J, JF, K, NBG, SAM and

STE, as listed in detail previously (Gibbs Russell

1987). Also, R. P. Ellis and W. D. Clayton for prof-

itable discussions, L. Watson and M. J. Dallwitz for

access to the DELTA computer system, G. Condy

for the spikelet drawing, and especially W. Roux for

technical assistance and preparation of maps.

REFERENCES

CHIPPINDALL, L. 1955. A guide to the identification of grasses

in South Africa. In C. Meredith, The grasses and pastures

of South Africa. Central News Agency, Cape Town.

DALLWITZ, M. J. 1984. Automatic typesetting of computer-

generated keys and descriptions. In R. Allkin & F.A.

Bisby, Databases in systematics. Systematics Association

Special Volume No. 26. Academic Press, London.

ELLIS, R. P. 1987a. Leaf anatomy of the genus Ehrharta (Poa-

ceae) in southern Africa: the Setacea group. Bothalia 17:

75-89.

ELLIS, R. P. 1987b. Leaf anatomy of the genus Ehrharta (Poa-

ceae) in southern Africa: the Villosa group. Bothalia 17:

195-204.

GIBBS RUSSELL, G. E. 1987. Taxonomy of the genus Ehrharta

(Poaceae) in southern Africa: the Setacea group. Bothalia

17: 67-73.

GIBBS RUSSELL, G. E. & ELLIS, R. P. 1987. Species groups

in Ehrharta (Poaceae) in southern Africa. Bothalia 17:

51-65.

LINNAEUS, C. 1781. Supplementum plantarum. Impensis Or-

phanotrophei, Brunsvigae.

NEES AB ESENBECK, C. G. 1841. Florae africae australioris.

Prausnitzianis, Cracow.

SCHRADER, H. A. 1821. Analecta ad floram capensem. Sect.

II, Cyperaceae. Gramineae. Gottingische gelehrte Anzei-

gen 1821: 2065-5079.

SCHULTES, J. A. 1830. Systema vegetabilium , Vol. 7, Part 2.

Cottae, Stuttgart.

SMOOK, L. & GIBBS RUSSELL, G. E. 1985. Poaceae. In

G. E. Gibbs Russell, C. Reid, J. van Rooy & L. Smook,

List of species of southern African plants, edn 2, part 1.

Memoirs of the Botanical Survey of South Africa No. 51.

STAPF, O. 1900. Gramineae. In W. T. Thiselton-Dyer, Flora

capensis 7: 310-750. Reeve, London.

STEUDEL, E. G. 1855. Synopsis plantarum graminearum.

Metzler, Stuttgart.

SWARTZ, O. 1802. The botanical history of the genus Ehrharta.

Transactions of the Linnean Society 6: 264-270.

THUNBERG, C. P. 1794. Prodromus plantarum capensium. Ed-

man, Uppsala.

THUNBERG, C. P. 1818. Flora capensis 2: 339. Hafniae, Copen-

hagen.

THUNBERG, C. P. 1823. Flora capensis. 2nd edn, J. A. Schultes

(ed.). Cottae, Stuttgart.

TRINIUS, C. B. 1839. Phalaridea. Memoires de I'Academie

Imperiale des Sciences de St. Petersbourg, ser. 6, 5: 12-26.

WATSON, L. & DALLWITZ, M. J. 1980. Australian grass gen-

era, anatomy, morphology and keys. Australian National

University, Research School of Biological Sciences, Can-

berra.

WATSON, L. & DALLWITZ, M. J. 1985. Australian grass gen-

era, anatomy, morphology, keys and classification, 2nd

edn. Australian National University, Research School of

Biological Sciences, Canberra.

WEBSTER. R. D. 1983. A revision of the genus Digitaria Haller

(Paniceae: Poaceae) in Australia. Brunonia 6: 131-216.

Bothalia 17,2: 195-204 (1987)

Leaf anatomy of the genus Ehrharta (Poaceae) in southern Africa:

the Villosa group

R. P. ELLIS*

Keywords: Ehrharta, E. thunbergii, E. villosa, leaf anatomy, Poaceae, stomatal flanges, systematics

ABSTRACT

The leaf blade anatomy of Ehrharta villosa Schult. f. var. villosa, var. maxima Stapf and E. thunbergii Gibbs

Russell is described and illustrated. These three taxa, constituting the Villosa species group, share a diagnostic leaf

anatomy distinguished by the absence of a distinct midrib, adaxial semi-radiate mesophyll with the abaxial chloren-

chyma palisade-like in arrangement, rectangular long cells and the stomatal apertures which are overlapped by

four cuticular flanges projecting from the two adjacent interstomatal cells. These combined attributes characterize

this species group, and the stomatal flanges are unique to this group in the genus Ehrharta Thunb. Microhairs are

absent in E. villosa but are present in E. thunbergii which also possesses abaxial prickles and plentiful, rounded

silica bodies not associated with cork cells as in E. villosa. These two taxa can, therefore, be separated anatom-

ically. Nevertheless, they share many features and are undoubtedly closely related and their classification in the

same species group is substantiated by the anatomical evidence presented in this paper.

U1TTREKSEL

Die blaarskyfanatomie van Ehrharta villosa Schult. f. var. villosa en var. maxima Stapf en E. thunbergii Gibbs

Russell word beskryf en geillustreer. Hierdie drie taksons, wat die Villosa-spesiegroep verteenwoordig, vertoon ’n

diagnostiese blaaranatomie, gekenmerk deur die afwesigheid van ’n duidelike hoofaar, semi-radiale adaksiale

mesofil met die abaksiale chlorenchiem palisade-agtig gerangskik, reghoekige langselle en die huidmondjie-ope-

ninge wat oorvleuel word deur vier kutikulere krae wat vanaf die twee aangrensende selle strek. Die kombinasie

van kenmerke onderskei hierdie spesiegroep, en die huidmondjie-krae is uniek by hierdie groep in die genus

Ehrharta Thunb. Mikrohare is afwesig by E. villosa maar aanwesig by E. thunbergii wat ook abaksiale stekelhare

en volop ronde silikaliggaampies, wat nie met kurkselle geassosieer is soos by E. villosa nie, besit. Hierdie twee

taksons kan dus anatomies onderskei word maar het nietemin baie kenmerke gemeen en is ongetwyfeld nouver-

want aan mekaar en hul klassifikasie in dieselfde spesiegroep word ondersteun deur anatomiese gegewens wat hier

aangebied word.

INTRODUCTION

The species of the Villosa group of the genus Ehr-

harta Thunb. are distinguished morphologically by

their large spikelets with profusely hairy, conspicu-

ously bearded and mucronate sterile lemmas (Gibbs

Russell & Ellis 1987). The leaf blades are reduced

and rolled and the culms are suffrutescent, some-

times with swollen or tuberous bases. Creeping,

underground rhizomes occur in all taxa.

Taxa included in this group are Ehrharta villosa

Schult. f. var. villosa and var. maxima Stapf, and E.

thunbergii Gibbs Russell (= E. gigantea Thunb.).

Chippindall (1955) considered E. villosa var. villosa

and E. thunbergii to be conspecific, whereas Smook

& Gibbs Russell (1985) synonomize E. villosa var.

maxima and E. thunbergii. In the present treatment

E. thunbergii is considered as a separate species fol-

lowing Gibbs Russell (1987) and consequently, three

taxa are assigned to the Villosa species group.

The leaf blade anatomy of taxa belonging to this

species group has received very little attention from

previous workers. Metcalfe (1960) gives a full de-

scription of E. villosa var. maxima and Engelbrecht

(1956) also describes the leaf anatomy of E. villosa

based on a representative sample of 18 specimens, 8

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

identified as E. villosa and 10 as E. thunbergii (= E.

gigantea) but considered as a single species.

This paper describes and illustrates the leaf blade

anatomy of the taxa of the Villosa group and dis-

cusses the affinities of these taxa and of the species

group by reference to this anatomical evidence. By

implication the anatomical data is compared and

contrasted with the morphological data as it is reflec-

ted in the classification of the group (Gibbs Russell

1987). The herbarium voucher specimens used in

this anatomical study were included in the sample on

which the above taxonomic conclusions were based.

The methodology is described in Gibbs Russell &

Ellis (1987) and the format of the paper follows that

of the first paper in the series (Ellis 1987a).

LEAF ANATOMY OF THE SPECIES OF THE VILLOSA

GROUP

E, villosa Schult. f.

Transverse section

The leaf blade is loosely inrolled (Figures 1.1, 2.1,

2.3, 3.1, 3.3) without a distinguishable keel, the

median vascular bundle being structurally identical

to the lateral first order bundles (Figures 1.1, 2.3,

3.3). Successive first order bundles are separated by

2-3 third order bundles except laterally where only a

single smaller bundle is located between successive

first order bundles.

196

Bothalia 17,2 (1987)

FIGURE 1. — Leaf anatomy of Ehrharta villosa var. maxima, Ellis 601\ 1-2, leaf in transverse section: 1, vascular bundle

arrangement and absence of keel (n), scale bar = 20 pm; 2, anatomical detail with sunken abaxial stomata (s) and mesophyll

with adaxially located chlorenchyma cells radiately arranged and abaxial cells palisade-like, scale bar = 10 pm. 3-4, abaxial

epidermis: 3, epidermal zonation with costal (darkly staining) and intercostal zones, scale bar = 10 pm; 4, short intercostal

long cells and flanged interstomatal cells with flanges projecting over the sunken stomata (s), scale bar = 5 pm.

Rounded adaxial ribs are associated with all the

vascular bundles (Figures 1.2, 2.2, 2.4, 3.2, 3.4),

those of the first order bundles being slightly larger.

Shallow, but rather narrow, furrows are present be-

tween all the ribs.

The mesophyll tissue is unusual in that it is semi-

radiate in arrangement, particularly the adaxially

situated cells located in the ribs (Figures 1.2, 2.2,

2.4), but the arrangement of the abaxial layers of

chlorenchyma cells is palisade-like (Figure 2.2, 2.4)

and may be conspicuous due to denser chloroplast

concentrations (Figure 3.4). The chlorenchyma cells

are relatively large, somewhat variable in shape but

tightly packed so that no large intercellular air

spaces are visible in transection (Figures 2.2, 2.4).

The chloroplasts are evenly but densely distributed

throughout all the chlorenchyma cells.

Abaxial epidermis

Costal and intercostal zones are clearly differen-

tiated (Figure 1.3, 2.5) due to differential staining,

although the epidermal cells of these two zones do

not necessarily differ greatly in structure (Figure

2.6). The costal zones lack stomata and consist of

narrower cells (Figure 1.4). The intercostal long cells

are rather short, and rectangular with slightly undu-

lating walls. The cells of the central files of each zone

may tend to be longer and wider than the lateral cells

(Figure 2.5). These larger cells are sometimes also

evident in the leaf sections (Figure 1.2, 2.4).

Stomata are common in 3-5 files in each intercos-

tal zone (Figure 1.3, 2. 5). They are clearly sunken

well below the level of the rest of the epidermis with

the guard and subsidiary cells being overlapped by

four distinct cuticular flanges extending over the sto-

matal aperture from the adjacent interstomatal long

cells (Figures 1.2, 2.2, 2.4). In surface view a distinct

cross-shaped aperture is formed by these flanges,

below which the stomatal apparatus is located (Fig-

ure 1.4, 2.6). SEM studies reveal that the flanges are

papilla-like (Figures 4. 1-4.4).

Costal silica bodies are not well differentiated and

are usually small, rounded and intimately associated

with an enfolding cork cell (Figure 1.4, 2.6). In less

typical specimens, however, the silica bodies may be

much more evident and numerous (Figures 6.1, 6.3).

Prickles are absent but prickles are present on the

adaxial costal zones which are equivalent to the ribs

as seen in transverse section (Figures 1.2, 2.4, 3.2).

No microhairs were seen either with the light or the

scanning electron microscope (Figures 1.4, 2.6,

4.1-4. 4).

Specimens examined

E. villosa var. villosa

CAPE. — 3218 (Clanwilliam): Lamberts Bay (-AB), Ellis 4640

(atypical tending toward E. thunbergii) . 3318 (Cape Town): Dar-

ling Dist., Yzerfontein (-AC), Ellis 1686. 3420 (Bredasdorp):

Bredasdorp Dist., De Hoop Nature Reserve (-AD), Ellis 1284,

4665. 3421 (Riversdale): Mossel Bay Dist., Albertinia (-BA).

Ellis 1651 (atypical tending toward E. tlumbergii).

Bothalia 17,2 (1987)

197

FIGURE 2. — Leaf anatomy of Ehrharta villosa var. villosa specimens resembling var. maxima in structure. 1-2, Ellis 1686, leaf

blade transection: 1, loosely inrolled blade without a keel, scale bar = 20 pm; 2, detail of the chlorenchyma showing dense

abaxial palisade-like cells; note sunken stomata (s) with overlapping flanges, scale bar = 5 pm. 3-4, Ellis 1284, transection: 3,

inrolled leaf, median vascular bundle (n) only, scale bar = 20 pm; 4, anatomical detail showing sunken guard cells and radiate

arrangement of the chlorenchyma, scale bar = 5 pm. 5, Ellis 1284, abaxial epidermis with costal zones and intercostal zones

with stomatal files, scale bar = 10 pm. 6, Ellis 1686, abaxial epidermis with detail of stomatal flanges (s) and costal zones,

scale bar = 5 pm.

E. villosa var. maxima

CAPE. — 3325 (Port Elizabeth): Port Elizabeth, Swartkops

Beach (-DC), Ellis 601.

Comments

E. villosa possesses the characteristic leaf anatomy

of the Villosa group being distinguished by the ab-

sence of a keel or midrib, the palisade-like abaxial

mesophyll, the flanged stomata, and the rectangular

long cells. The anatomy of var. villosa and var. max-

ima is very similar and these two taxa appear to show

close affinities, being indistinguishable on leaf blade

anatomy, a fact which appears to corroborate their

separation at only the varietal level.

Although the var. maxima anatomical sample

used in this study is inadequate, the specimen exam-

ined (Ellis 601) conforms in all respects to the de-

scription given by Metcalfe (1960) for material from

Western Australia even though his microtechnique

procedures did not allow a detailed examination of

the mesophyll. These two specimens reveal that the

leaf anatomy of var. maxima conforms very closely

with that of var. villosa, with some specimens of the

latter being virtually indistinguishable from var.

maxima in leaf anatomy (Figures 2. 1-2.6).

E. villosa var. villosa is a rather variable taxon

anatomically. Some specimens of var. villosa corre-

spond very closely in leaf size and thickness to the

relatively large leaves of var. maxima, as a compari-

198

Bothalia 17,2 (1987)

FIGURE 3. — Transectional leaf anatomy of Ehrharta villosa var. villosa. 1-2, Ellis 4665 : 1, inrolled outline with median bundle (n)

only, scale bar = 20 |im; 2, anatomical detail showing rather angular chlorenchyma cells with radiate adaxial layers and

palisade-like abaxial layers, scale bar = 10 pm. 3-4, Ellis 4640: 3, inrolled outline without additional parenchyma in associa-

tion with the median bundle (n), scale bar = 20 pm; 4, interference contrast illumination of detail of chlorenchyma cell

arrangement, scale bar = 10 pm.

son of Figures 1.1, 1.2 and 2.1^1 shows. Others,

however, resemble E. thunbergii with thinner leaves

(Figures 3.3, 3.4). A similar trend is also evident in

the epidermal structure, with Figures 2.5, 2.6 resem-

bling the var. maxima condition, whereas Figures

6.1, 6.3 approximate closely some of the E. thunber-

gii specimens. E. villosa var. villosa, therefore, is

intermediate in leaf anatomy between var. maxima

and E. thunbergii and the interface between these

two taxa is not very distinct.

The intermediate nature of var. villosa is also evi-

dent in its spikelet size and habitat requirements and

several specimens have proved difficult to assign to

either var. villosa or E. thunbergii on morphological

criteria. This is particularly the case if the rhizome

characters are not evident. But E. villosa is a species

of deep, loose sand of the lowland fynbos and only

occurs at higher altitudes where drift sand occurs as

a result of wind or water deposition.

The clinal variation in anatomical structure in var.

villosa appears to be a reflection of these habitat

gradients. Those specimens most resembling var.

maxima are all from coastal dune habitats (Figures

2. 1-2.6) to which var. maxima appears to be con-

fined. With increasing altitude and distance from the

sea the var. villosa specimens (Figures 3. 1-3.4, 6.1,

6.3) tend to merge with E. thunbergii, which is a

species of higher altitudes, heavier soils and the

mountain fynbos.

E. thunbergii Gibbs Russell

Transverse section

Blade loosely to rather tightly inrolled (Figures

5.3, 5.5). A slight keel may sometimes be devel-

oped, as evidenced by the presence of additional

colourless parenchyma associated with the median

vascular bundle (Figures 5.3, 5.5). This development

is not equally evident in all specimens and several

have the median bundle structurally identical to the

lateral first order bundles, without additional paren-

chyma (Figure 5.1). One or two third order bundles

occur between consecutive first order bundles.

Adaxial ribs are slight but rounded (Figure 5.2) or

may be more conspicuous but then abaxial intercos-

tal ribs alternate with the adaxial costal ribs (Figures

5.4, 5.5). Adaxial furrows are shallow and wider

than in E. villosa.

The mesophyll is rather variable but all specimens

conform to the general pattern so characteristic of

this group. Examples with semi-radiate chloren-

chyma with an abaxial palisade-like layer are illus-

trated in Figures 5.2, 7.1 and 7.3 and correspond

closely to the E. villosa specimens illustrated in Fig-

ures 3.2 and 3.4. Other E. thunbergii specimens,

with thinner leaves and fewer chlorenchyma cell lay-

ers differ slightly from this pattern (Figure 5.4). The

chlorenchyma cells themselves remain rather large,

somewhat angular and tightly packed with very small

intercellular air spaces (Figures 5.2, 5.4).

Bothalia 17,2 (1987)

199

FIGURE 4. — Abaxial epidermal ultrastructure of representatives of the Villosa group. 1-4 Ehrharta villosa var. villosa. 1-2, Ellis

4640 : 1, thickened epidermal cells, no microhairs and flanged stomata, x 200; 2, detail of the four papilla-like flanges

overarching the stomatal apparatus with the guard cells visible below this aperture, x 1000. 3-4, Ellis 4665: 3, thick cuticle,

sunken stomata and microhairs absent, x 200; 4, guard cells beneath the overarching papillate flanges, x 1000. 5-8, Ehrharta

thunbergii. 5-6, Ellis 4648: 5, distinct costal zone with raised, round silica bodies and prickles; intercostal zone with micro-

hairs and files of flanged stomata, x 200; 6, detail of microhair with tapering distal cell and flanges obscuring adjacent stoma,

x 1000. 7-8, Ellis 4626 illustrating anatomical variation in E. thunbergii: 7, costal prickles, intercostal microhairs and flanged

stomata, but note the diamond-shaped intercostal long cells, x 200; 8, microhair with tapering distal cell, x 1000.

200

Bothalia 17,2 (1987)

FIGURE 5. — Leaf anatomy of Ehrharta thunbergii. 1-5, leaf transections; 6-8, abaxial epidermides. 1-2, Ellis 1152: 1, median

bundle not structurally distinct, scale bar = 20 pm; 2, chlorenchyma cell detail and arrangement typical of that of the Villosa

group, scale bar = 5 pm, 3-4, Ellis 708: 3, inrolled blade with slight keel (n), scale bar = 20 pm; 4, detail showing less

pronounced radiate and palisade chlorenchyma arrangement and abaxial ribs, scale bar = 5 pm. 5-6, Ellis 1145: 5, leaf

outline showing small but definite keel (n), scale bar = 20 pm; 6, elongated intercostal long cells, stomata obscured by flanges

(s), scale bar = 5 pm. 7, Ellis 708 with elongated stained and unstained intercostal long cells with slightly undulating walls,

microhairs (n) and flanged stomata (s), scale bar= 5 pm. 8, Ellis 1152: note silica bodies, costal and intercostal prickle hairs

(p), stained and unstained long cells, microhairs (n) and flanged stomata, scale bar = 5 pm.

Bothalia 17,2 (1987)

201

Abaxial epidermis

Costal and intercostal zones are always distin-

guishable (Figures 5. 6-5. 8, 6.2, 6.4, 7.2, 7. 4-7. 6).

Cell size and shape differ markedly between these

two zones on all the specimens examined. The inter-

costal long cells are often much more elongated than

in E. villosa but this character is variable with Fig-

ures 5.6, 5.7 and 6.4 representing the two extremes

encountered in this species. The long cell shape is

usually rectangular but may be diamond-shaped

(Figure 7.4). The markedly elongated long cells may

also stain with safranin (Figures 5.7, 5.8).

Stomata occur in 2-3 files on either side of each

costal zone but are absent from the central files of

the intercostal zones. These stomata are always

sunken and overlapped by cuticular flanges although

these are not always easily visible with the light

microscope (Figures 5.6, 5.7). The specimens with

thinner leaves and elongated long cells have less con-

spicuous flanges associated with more superficial

stomata. Those specimens tending toward E. villosa

in leaf anatomy have this characteristic well devel-

oped (Figures 6.2, 6.4), as do the specimens showing

similarities with the Calycina group (Figure 7.4) or

the Ramosa group (Figures 7.5, 7.6). Although vari-

able, this attribute is evident on all the specimens

studied and is confirmed by the SEM (Figures

4.5-4. 7).

Costal silica bodies are generally well differen-

tiated, being conspicuous and rounded and alternat-

ing along the costal files (Figures 5.8, 6.4, 7.2). Cres-

cent-shaped, enfolding cork cells do not appear to

occur in this species. Abaxial costal prickles are

common and were observed on all specimens with

two specimens (Ellis 1152 and 5102) even possessing

large intercostal prickles associated with the stoma-

tal bands (Figure 5.8). Microhairs, although very

small, were detected on all specimens, even those

resembling E. villosa in other anatomical character-

istics. Ultrastructurally these hairs are seen to have a

tapering distal cell (Figures 4.6, 4.8).

Specimens examined

CAPE. — 3118 (Vanrhynsdorp): Vanrhynsdorp Dist., Gifberg

(-DD), Ellis 5102. 3119 (Calvinia): Nieuwoudtville Dist., Van

Rhyn's Pass (-AC), Ellis 1145, 4626. 3218 (Clanwilliam): Clanwil-

liam Dist., Pakhuis Pass (-BB), Ellis 1700 ; Langvlei Valley, Sand-

berg Station (-BC), Ellis 4642\ Piketberg Dist., Versveld’s Pass

(-DD), Ellis 5130. 3219 (Wuppertal): Cedarberg, Pakhuis Pass

(-AA), Ellis 708, 1152, 4633, 4635\ Kouebokkeveld, Skurweberg.

Op-die-Berg (-CD), Ellis 4648. 3318 (Cape Town): Malmesbury

Dist., Paardeberg (-DB), Boucher 4724. 3321 (Ladismith):

Langeberge, Cloete’s Pass, Bergkloof (-DC), Ellis 4693.

Comments

The diagnostic anatomical attributes of the Villosa

group are all present in E. thunbergii although they

may be somewhat modified on some specimens.

FIGURE 6. — A comparison of the abaxial leaf epidermis of Ehrharta villosa ( 1 & 3) and Ehrharta thunbergii (2 & 4). 1, E. villosa,

Ellis 4640, showing short intercostal long cells with slightly sinuous walls; microhairs and prickles absent, scale bar = 10 urn.

2, E. thunbergii, Ellis 4635, with very sinuous long cell walls, intercostal microhairs and costal prickles, scale bar = 5 pm. 3,

E. villosa , Ellis 1651, short, slightly sinuous long cells, flanged stomata, scale bar = 10 urn. 4, E. thunbergii. Ellis 4648,

prominent, round costal silica bodies and prickles. Intercostal microhairs (n) and stomata (s) overarched by flanges from

interstomatal long cells, scale bar = 5 pm.

202

Bothalia 17,2 (1987)

FIGURE 7. — Anatomical variation in Ehrharta thunbergii. 1-2, Ellis 1700 resembling E. villosa: 1, transverse section with radiate

and palisade-like mesophyll cells, scale bar = 10 pm; 2, abaxial epidermis with rectangular, sinuous-walled long cells but

stomatal flanges not conspicuous, scale bar = 5 pm. 3-4 Ellis 4626 resembling the Calycina group: 3, normal transverse

section but note the enlarged abaxial epidermal cells in the centres of the intercostal zones, scale bar = 10 pm; 4, central

intercostal long cells markedly elongated and diamond-shaped but stomata retain characteristic flanges, scale bar = 5 pm.

5-6, Ellis 4693 resembling the Ramosa group: 5, conspicuous intercostal short cell pairs separate successive long cells, scale

bar = 10 pm; 6, nucleate epidermal long and short cells but stomata flanged, scale bar = 5 pm.

Thus only a median vascular bundle is normally

present but in a few specimens additional colourless

parenchyma is associated with the median bundle,

which, by definition, constitutes a slight keel. The

semi-radiate adaxial, and palisade-like abaxial meso-

phyll, so characteristic of this group, is evident in

most specimens. However, in a few, particularly

those with thinner leaves and with abaxial intercostal

ribs, this pattern may be modified slightly. In all

specimens the stomata are sunken and overlapped

by four papillate epidermal flanges. However, in

those specimens with elongate intercostal long cells

the stomata may be almost flush with the level of the

epidermis and the flanges are tiny. These diagnostic

features are common to all taxa of the Villosa group

and serve to unite E. villosa and E. thunbergii in a

group separated from all the other species of Ehr-

harta.

In addition, several characters serve to separate

E. thunbergii from E. villosa , although this distinc-

tion is not very clear-cut. Examples are the presence

of microhairs and abaxial prickles, both of which are

lacking in E. villosa. The costal silica bodies of E.

thunbergii are also well differentiated and plentiful

and alternate with costal short cells. They are not

associated with cork cells as in E. villosa. These two

taxa can, therefore, be distinguished anatomically.

Yet other attributes intergrade between the taxa

of this species group, and the leaf anatomy of the E.

thunbergii specimens studied shows a certain degree

of variation. A distinct gradation is evident from

Bothalia 17,2 (1987)

203

those specimens closely resembling E. villosa (Fig-

ures 6.2, 6.4, 7.1, 7.3) to the extreme type with thin-

ner leaves and elongated long cells (Figures

5. 3-5. 7). The interface with E. villosa is indistinct. A

continuum is discernible from those specimens re-

sembling E. villosa to the extreme specimens which

may display characteristics of some of the other Ehr-

harta species groups, the Calycina group in particu-

lar. Calycina type features observed are the fusiform

intercostal long cells as in Figure 7.4, the tendency to

stain with safranin (Figures 5. 6-5. 8) and the inter-

costal abaxial ribs (Figure 5.4) or the inflated central

cells of the intercostal zones as illustrated for E. vil-

losa (Figures 2.4, 2.5). A single specimen, Ellis

4642, although not illustrated, resembles E. calycina

particularly closely, even having straight-walled fusi-

form long cells and intercostal macrohairs which

were not observed on any other E. thunbergii speci-

men. However, flanged stomata indicate the true

identity of this specimen.

One other interesting and deviant specimen is

Ellis 4693 (Figures 7.5, 7.6) which shows similarities

with the Ramosa group of species. The sinuous, rect-

angular long cells, all separated by conspicuous

cork/silica cell pairs and the irregular, dumbbell-

shaped silica bodies, are reminiscent of the Ramosa

group and were not seen in any other E. thunbergii

specimens. However, this specimen also has dis-

tinctly flanged stomata.

The anatomical sample examined in this study is

heavily biased toward the north-western parts of the

distribution range of E. thunbergii. Those specimens

from high altitudes in the extreme north at Van

Rhyn’s Pass (Ellis 1145, Figures 5.5, 5.6; Ellis 4626,

Figures 7.3, 7.4) show anatomical similarities with

E. calycina. A specimen (Ellis 4642) from lower alti-

tude in the strandveld at Langvlei resembles E. caly-

cina very closely indeed. On the other hand, few

specimens from the east have been classified as E.

thunbergii (these being mainly identified as E. vil-

losa) and Ellis 4693 from Cloete’s Pass in the eastern

Langeberge resembles the Ramosa group in certain

respects. These observations may reflect transitions

to these other Ehrharta species groups but a much

more representative sample must be studied before

this can be confirmed. Nevertheless, this does serve

to demonstrate that the Villosa group is not discrete,

and that characteristics of some other groups are

evident, as is the case throughout the genus.

These observations are largely in agreement with

the findings of Engelbrecht (1956) and the few ex-

ceptions noted will be briefly discussed. For the ma-

jority of specimens the epidermis is described as be-

ing homogenous with costal and intercostal zones

not being distinguishable (Engelbrecht 1956). In the

present study the condition is described where these

zones are structurally identical, as in E. villosa var.

maxima for example, but are distinguishable on ac-

count of their differential staining. Different staining

procedures, therefore, may account for this appar-

ently superficial difference between the findings of

these two studies. Engelbrecht (1956) does record

the absence of microhairs and prickles associated

with the homogenous type of epidermis (which ap-

pears to be homologous with E. villosa ) whereas the

epidermis with distinct epidermal zonation was asso-

ciated with the presence of these hairs. This correla-

tion was observed in the present study and is consid-

ered to be a specific difference between E. villosa

and E. thunbergii but Engelbrecht (1956) did not

attribute any taxonomic significance to it. He also

records cuticular stomatal flanges for all the speci-

mens he examined and notes the uniqueness of this

feature in the genus.

DISCUSSION AND CONCLUSIONS

The three taxa of the Villosa group, share a dis-

tinctive leaf anatomy characterized by a unique com-

bination of attributes as well as similar vegetative

morphology and a specific habitat. These distin-

guishing features correlate with the diagnostic large,

hairy spikelets, and their assignment to the same

small species group appears to be fully justified by

the anatomical as well as the morphological evidence

(Gibbs Russell 1987). This group also appears to

represent a natural grouping.

The leaf anatomy is characterized by the absence

of a keel, palisade-like mesophyll abaxially located,

rectangular long cells and stomatal apertures which

are overlapped by four cuticular flanges projecting

from the two adjacent interstomatal long cells. This

latter feature is unique to this species group in the

genus Ehrharta.

Although Engelbrecht (1956) studied only uni-

fixed leaf blade material he noted that the form of

the cells of the abaxial chlorenchyma layer differed

from the remainder, an observation confirmed in

this study. However, he also reports cell wall invagi-

nations as being present and characteristic of E. vil-

losa. These invaginations were not observed on all

chlorenchyma cells, however, but appeared to be

confined to those cells adjacent to the vascular bun-

dles or adjoining the adaxial epidermis. This obser-

vation was not confirmed in the present study, in

which field-fixed material was examined, and ap-

pears to be an artefact probably resulting from im-

perfect rehydration of the mesophyll tissue.

Engelbrecht (1956) recognized two basic groups of

species in Ehrharta — one with invaginated chloren-

chyma and one without. E. villosa is placed in the

group with invaginations together with taxa of the

Setacea and Ramosa species groups as here consti-

tuted (Gibbs Russell & Ellis 1987). The present find-

ings are in disagreement with Engelbrecht's (1956)

grouping, as the Setacea group is the only group in

which arm cells were observed (Ellis 1987) and the

Setacea and Villosa groups are not considered to be

closely related.

Although he examined a large sample, Engel-

brecht (1956) was unable to distinguish E. villosa

and E. thunbergii either anatomically or morpholo-

gically and concluded that they do not represent two

separate species. The present study is not in full

agreement with this conclusion as E. villosa and E.

thunbergii were found to differ in several respects

such as the presence of microhairs and prickles as

well as differences in silica bodies. Although these

differences appear to be consistent and diagnostic, it

must be remembered that the interface between

204

Bothalia 17,2 (1987)

these two species is not distinct as far as most other

characters are concerned and a continuum is evident

between them without clear character disjunctions.

E. villosa and E. thunbergii, therefore, intergrade to

a certain extent and, although their extremes are

anatomically quite distinct, a small proportion of

specimens are somewhat intermediate. The decision

to consider these two taxa as being conspecific

(Chippindall 1955; Engelbrecht 1956), therefore,

has some merit. However, the placing of E. thunber-

gii in synonomy under E. villosa results in a very

variable, polymorphic entity with a wide ecological

tolerance. The recognition of three taxa seems to be

a more practical solution which probably reflects the

natural situation more accurately. However, a cline

undoubtedly exists from E. villosa var. maxima

through var. villosa to E. thunbergii with each of

these taxa occupying slightly different habitats and

differing in morphology and leaf anatomy.

The relationships of the Villosa group to the rest

of the genus are not very clear* from anatomical evi-

dence alone. The group does not occupy such an

isolated position within the genus as does the Seta-

cea group (Ellis 1987) which possesses such taxono-

mically significant diagnostic features as arm cells

and distinct microhairs and silica bodies. There are

also no anatomical intermediates between the Seta-

cea group and any of the other species groups. Al-

though the Villosa group is readily diagnosed by its

flanged stomata, this feature cannot be accorded the

high taxonomic value that arm cells and microhair

and silica body shape have in the classification of the

Poaceae, because it is encountered independently in

different subfamilies.

In addition, several E. thunbergii specimens dis-

play strong Calycina group attributes in their leaf

anatomy, and both these groups have very similar

microhairs. The Villosa and Calycina groups also

share very similar hairy spikelets, which differ

mainly in size and profuseness of vesture, but occur

in no other Ehrharta species group. The indications

are, therefore, that the Villosa group is more closely

related to the Calycina group than to any of the

other groups. However, as is common in this genus,

a reticulate pattern of relationships can be expected

and Ramosa group characteristics were also ob-

served on a few specimens.

The Villosa group, although distinct in morpho-

logy, anatomy and ecology, does show certain affini-

ties with the Calycina group and undoubtedly be-

longs to the genus Ehrharta. This group, therefore,

appears to be a specialized perennial line with strong

underground rhizomes and suffrutescent culms

which has become adapted to a sandy habitat.

ACKNOWLEDGEMENTS

The author is grateful to Mrs H. Ebertsohn for

technical assistance, to Mrs S. Perold for the SEM,

work, to Mrs A. Romanowski for the photography

and Mrs S. S. Brink for typing the manuscript.

Voucher specimens were identified by the staff of

the National Herbarium (PRE) and revised by Dr

G. E. Gibbs Russell.

REFERENCES

CHIPPINDALL, L. K. A. 1955. In D. Meredith, The grasses and

pastures of South Africa. C NA, Johannesburg.

ELLIS, R. P. 1987a. Leaf anatomy of the genus Ehrharta (Poa-

ceae) in southern Africa: the Setacea group. Bothalia 17:

75-89.

ENGELBRECHT, A. H. P. 1956. 'n Morfologiese studie van die

genus Ehrharta Thunb. M.Sc. thesis. University of Pre-

toria.

GIBBS RUSSELL, G. E. 1987. Taxonomy of the genus Ehrharta

(Poaceae) in southern Africa: the Villosa group. Bothalia

17: 191-194.

GIBBS RUSSELL, G. E. & ELLIS, R. P. 1987. Species groups

in Ehrharta (Poaceae) in southern Africa. Bothalia 17:

51-65.

METCALFE, C. R. 1960. Anatomy of the Monocotyledons. 1.

Gramineae. Clarendon Press, Oxford.

SMOOK, L. & GIBBS RUSSELL, G. E. 1985. Poaceae. In G.

E. Gibbs Russell, C. Reid, J. van Rooy & L. Smook, List

of species of southern African plants, edn 2, part 1. Mem-

oirs of the Botanical Survey of South Africa No. 51.

Bothalia 17,2: 205-212 (1987)

Phytogeography of the subtribe Leipoldtiinae (Mesembryanthema-

ceae)

H. E. K. HARTMANN*

Keywords: ecology, Leipoldtiinae, Mesembryanthemaceae, phytogeography, winter rainfall

ABSTRACT

An investigation of distribution and species frequency of the nine genera of the subtribe Leipoldtiinae shows

that two centres of diversity can be distinguished. These coincide more or less with the ‘Gariep centre’ in the north

and the ‘Vanrhynsdorp centre’ in the south (both sensu Nordenstam 1969). Speciation seems to have occurred in

both centres. The subtribe may have evolved in an arid winter rainfall area which could have been situated outside

its present distribution area. The centres of distribution coincide with those observed in both subfamilies of the

Mesembryanthemaceae.

UITTREKSEL

’n Ondersoek na verspreiding en spesie-frekwensie van die nege genusse van die subtribus Leipoldtiinae toon

dat twee sentrums van diversiteit onderskei kan word. Hierdie twee kom min of meer ooreen met die 'Gariepsen-

trum’ in die noorde en die ‘Vanrhynsdorpsentrum’ in die suide (albei sensu Nordenstam 1969). Dit wil voorkom

asof spesiasie in albei sentrums plaasgevind het. Die subtribus kon moontlik in ’n dorre winterreengebied wat buite

die huidige verspreidingsgebied gelee kon gewees het, ontwikkel het. Die verspreidingsentrums kom ooreen met

die wat in albei subfamilies van die Mesembryanthemaceae waargeneem is.

INTRODUCTION

The subtribe Leipoldtiinae of the Mesembryan-

themaceae comprises nine genera with a total of 81

species. Members of the subtribe can be recognized

by their rather hard xeromorphic leaves (the outer

epidermal wall always contains calcium oxalate crys-

tals) and their multilocular capsules (Figure 1) with

valve wings, closing bodies and complete covering

membranes. These membranes are distally recurved

and radially traversed by a trace of spongy tissue

which distally forms an additional closing device

(closing bulge, ledge or rod, Hartmann 1983c). Dis-

semination depends directly on the described inter-

nal structure (Funktionstyp 1 after Hartmann 1983c)

and is effected in most taxa only by rain drops.

This is the first analysis of its kind within the Me-

sembryanthemaceae because this is the first group of

genera for which adequate data are available. The

phytogeography of the subtribe is of particular inter-

est for two reasons: the distribution area of the Lei-

poldtiinae coincides with the centre of distribution of

the entire family; and within this area, members of

the subtribe form a major part of the vegetation.

METHODS AND MATERIAL

The investigation is based on numerous field stu-

dies carried out between 1977 and 1986, about 6 000

relevant collections and about 800 additional herbar-

ium sheets. Ecological data are derived partly from

literature and partly from own observations; rainfall

data were provided by the Weather Bureau, Pre-

toria. Information concerning morphology, anat-

omy, evolution and detailed distribution are taken

* Institut fiir Allgemeine Botanik, Universitat Hamburg, Ohn-

horststrasse 18, D-2000 Hamburg 52, West Germany.

from the relevant genus monographs (see Refer-

ences).

RESULTS

Distribution of the subtribe

The subtribe is confined to the south-western

Cape. Only one genus (Cephalophyllum) occurs in

the entire range of the subtribe (see Figure 2), the

other genera are restricted to smaller areas. The fre-

quency distribution of genera is consequently un-

even (Figure 3). Two main centres can be distin-

guished, a southern one, which will be referred to as

‘Vanrhynsdorp centre’ (sensu Nordenstam 1969),

and a northern one, which agrees largely with the

‘Gariep centre’ (sensu Nordenstam 1969). These

centres stand out even more prominently when

species distribution (Figure 4) is considered, and

their significance will be discussed below.

FIGURE 1. — Open fruit of Leipoldtiinae, schematic. ACD =

additional closing device; C = columella; CB = closing

body; CM = covering membrane; EA = expanding area;

EC = expanding keel; RCM = recurved distal portion of

covering membrane; RT = radial trace of spongy tissue; V

= valve; VW = valve wing.

206

Bothalia 17,2 (1987)

FIGURE 2. — Distribution areas of Cephalophyllum (thin lines)

and Argyroderma (bold line). Horizontal lines = Cephalo-

phyllum subgenus Cephalophyllum ; vertical lines = Ce-

phalophyllum subgenus Homophyllum; overlapping areas

dotted.

FIGURE 3. — Frequency of genera of Leipoldtiinae (N = 9) per

30' x 30' square. Two centres can be distinguished: ‘Gar-

iep centre’ (bold figures between latitudes 27° and 31°);

‘Vanrhynsdorp centre’ (bold figures between latitudes 31°

and 32°). Bold lines delineate the four main rainfall areas

of the region.

The eastern limit of the distribution area of the

Leipoldtiinae coincides for most of its extension with

the eastern boundary of the winter rainfall area (de-

fined here as the region receiving at least 60% of its

annual precipitation between April and September,

Figure 3). Only in the south, three genera (Cephalo-

phyllum, Jordaaniella and Leipoldtia) extend consid-

erably into the March/November or all season rain-

FIGURE 4. — Frequency of species of Leipoldtiinae (N = 81)

per 30' X 30' square. The same two centres as in Figure 3

stand out (bold figures).

fall areas. Only two species out of the 81 have their

centre of distribution outside the winter rainfall re-

gion: Cephalophyllum subulatoides (Haw.) N.E. Br.

(southern Little Karoo, 33°30'S, 21°-23°E) and

Cheiridopsis caroli-schmidtii (Dinter & Berger)

N.E. Br. (near Aus, 27°S, 16°E). The amount of

annual precipitation decreases in the distribution

area from south to north, and less markedly from

west to east (Figure 5).

Phytochorologically, the main concentration of

taxa falls into the Western Cape Domain of the Ka-

roo-Namib Region as defined by Werger (1978),

with extensions into the Namib Domain in the north,

into the Karoo Domain in the north-east and even

into Capensis in the south (Figure 4). Only a few

species have their centre of distribution in Capensis

sensu Werger (1978), namely Cephalophyllum diver-

siphyllum (Haw.) H. E. K. Hartm., C. loreum (L.)

Schwantes, and C. subulatoides (Haw.) N.E. Br.

Distribution of genera

Cephalophyllum

The largest genus of the subtribe, Cephalophyllum

N.E. Br., with 30 species, has the largest distribution

area (Figure 2) and grows in a wide range of ecologi-

cal situations, from coastal to high mountain habi-

tats, from monotypic open succulent associations to

undergrowth in fynbos, and from the highest to the

lowest rainfall regimes in different seasons. The two

subgenera have different centres of distribution (Fig-

ure 2): subgenus Cephalophyllum dominates in the

south and subgenus Homophyllum in the north, with

a wide corridor in which they overlap.

A species frequency analysis (based on numbers

of species per 30' x 30' square) reveals that the high-

est number of species occurs in the Vanrhynsdorp

centre (Figure 6), which lies in the zone of overlap of

the subgenera (Figure 2). A second centre of species

Bothalia 17,2 (1987)

207

FIGURE 5. — Frequency of species of Argyroderma (N = 10)

per 30' x 30' square and selected isohyets (bold line = 100

mm; medium line = 400 mm; thin line = 200 mm).

frequency lies in the north and coincides with the

Gariep centre of the subtribe (Figure 3). These

centres cannot be correlated directly with the subge-

nera, as the centre of subgenus Cephalophyllum is

situated near 33°S, 20°E (Figure 6 and Hartmann in

press). In spite of the high species numbers per

square, sympatry in Cephalophyllum is rare and

speciation has been predominantly allopatric (Hart-

mann in press). Twelve species (40%) can be consid-

ered endemic: they occur in one to three 30' squares

only (Figure 6). They are found near the two species

centres but their frequency is low (1-3 per 30'

square).

FIGURE 6. — Frequency of species of Cephalophyllum (N = 30)

per 30' x 30' square. Small figures indicate frequency of

endemic species.

Consequently, Cephalophyllum can be described

as a genus with mostly widespread yet vicariant

species, which have become adapted to a wide range

of edaphic and climatic conditions: the genus occurs

in 13 different vegetation types as defined by Acocks

(1975) (Hartmann in press). The extended distribu-

tion area reflects the high degree of adaptability of

the genus even today, supporting the hypothesis that

the genus is at present in an active evolutionary

phase (Hartmann in press).

Argyroderma

Closely related to Cephalophyllum is the genus

Argyroderma N.E. Br. (Hartmann 1983c, in press)

with ten species (Hartmann 1978a), restricted in dis-

tribution to the Knersvlakte in the Vanrhynsdorp

centre (Figure 2), with a maximum of six species per

30' square (Figure 5). In contrast to Cephalophyl-

lum, Argyroderma therefore presents a genus of nar-

row distribution which is due to the strict adaptation

to the unique edaphic mosaic of the Knersvlakte.

Leipoldtia

With only eight species, Leipoldtia L. Bol. is one

of the smaller genera of the subtribe, yet it is one of

wide distribution (Figure 7). The predominantly

shrubby plants often grow in conspicuous patches,

mostly on gravelly and flat sedimentary soils. In con-

trast to the wide occurrence of subgenus Leipoldtia,

the subgenera A ureae and Cephalophylloides inhabit

only restricted areas overlapping with that of subge-

nus Leipoldtia (Figure 7). Only one species (L. fru-

tescens, of the subgenus Aureae) grows in coastal

plains (Strandveld sensu Acocks 1975). All other

taxa inhabit higher altitudes, mainly in Namaqua-

land Broken Veld, Western Mountain Karoo and

Karroid Broken Veld of the Little Karoo.

FIGURE 7. — Distribution area of Leipoldtia. Horizontal lines =

subgenus Leipoldtia; vertical lines = subgenus Cephalo-

phylloides; dots = subgenus Aureae.

208

Bothalia 17,2 (1987)

FIGURE 8. — Frequency of species of Leipoldtia (N = 8) per 30'

x 30' square. H = occurrence of Hallianthus; V = occur-

rence of Vanzijlia in the square.

The highest species frequencies are found within

the southern Gariep centre (Figure 8), where in cer-

tain localities up to three species occur sympatri-

cally, each fitting into an ecological niche slightly

different from those of the others (Hartmann & Rust

in prep.). Since the same species can inhabit ob-

viously different habitats in distant areas, a fine

regulation of adaptation in regard to competition

and isolation can be expected, but this is not yet

understood (Rust unpublished).

Although 50% of all species of Leipoldtia occur in

one 30' square, it cannot be simply assumed that this

area presents a centre of origin. The scattered distri-

bution of endemic species (Figure 8), and the varied

character expressions in geographically separated

areas of one species, point more towards an active

phase of speciation by means of differentiation of

semi-isolated, more or less marginal populations.

The high species frequency in the Gariep centre

could be the result of secondary invasions combined

with the stabilization of hybrid derivatives as ecolo-

gically well adapted forms which have become iso-

lated (Rust unpublished).

Hallianthus and Vanzijlia

Two monotypic genera with allopatric distribution

are related to the genus Leipoldtia (Hartmann

1983c): Hallianthus H. E. K. Hartm. and Vanzijlia

L. Bol. Hallianthus grows predominantly at altitudes

above 300 m (Hartmann 1983b) in Namaqualand

Broken Veld (sensu Acocks 1975) in associations of

succulent shrubs. It extends northwards into the

Gariep centre (Figure 8). Vanzijlia , as a mainly

coastal genus, occurs around and in the Vanrhyns-

dorp centre (Figure 8). Most populations have been

found in shrubby succulent associations of the

Strandveld (sensu Acocks 1975), few inland in Suc-

culent Karoo (Hartmann 1983a).

Cheiridopsis and Odontophorus

The genus Cheiridopsis N.E. Br. with 23 species,

represents the third of the widespread genera of the

subtribe. Like the genus Odontophorus N.E. Br., it

can be recognized by its papillose leaf surface (Hart-

mann 1983c). As in Cephalophyllum, the two larger

subgenera (subgenus Cheiridopsis and Aequifoliae)

inhabit largely different geographical areas (Figure

FIGURE 9. — Distribution of Cheiridopsis and Odontophorus

(thin contour between latitude 28°55' and 30°10'). Hori-

zontal lines = C. subgenus Aequifoliae; vertical lines = C.

subgenus Cheiridopsis; dots = area in which subgenera

Aequifoliae and Cheiridopsis overlap; bold contour = C.

subgenus Odontophoroides.

FIGURE 10. — Frequency of species of Cheiridopsis per 30' x

30' square (N = 23). Small figures indicate number of

endemic species; O = Odontophorus; one symbol repre-

sents one species per square.

Bothalia 17,2 (1987)

9) with an intervening zone in which they overlap.

The third, smaller, subgenus Odontophoroides, as

well as all species of the closely related genus Odon-

tophorus, also occur in and near to this area of over-

lap. Consequently the number of species per 30'

square is high (Figure 10) and contributes substan-

tially to the formation of the southern Gariep centre

(Figure 4). Furthermore, most of the endemic

species of Cheiridopsis (occurring in at most three

adjacent 30' squares) are found here (Figure 10).

In and around the Gariep centre, sympatry of dif-

ferent species of Cheiridopsis is frequent, but in

most localities different, well defined microhabitats

are colonized (Hartmann & Dehn 1987). The three

species of Odontophorus occur allopatrically in dis-

junct areas (Hartmann 1976) as do the six species of

subgenus Odontophoroides (Hartmann & Dehn

1987).

Cheiridopsis and Odontophorus constitute a genus

pair similar in principle to Cephalophyllum/ Argyro-

derma in so far as the genus of restricted distribution

occurs in a centre of high species frequency of the

widespread genus, but they differ in so far as Cheiri-

dopsis/Odontophorus only possess one centre of di-

versity.

Another characteristic of the Cheiridopsis/ Odon-

tophorus complex is its nearly exclusive distribution

along the eastern boundary of the area of the sub-

tribe at altitudes above 300 m. Only three species, C.

brownii Schick & Tischer and C. robusta (Haw.)

N.E. Br. in the north and C. rostrata (L.) N.E. Br. in

the south, occur on lower coastal plains. Neverthe-

less, the ecological amplitude of the genus is consid-

erable. It occurs in eight veldtypes of Acocks (1975),

with one species, C. cigarettifera (Berger) N.E. Br.

in the south, growing in six different vegetation

types (Hartmann & Dehn 1987). The adaptation to

shrubby Renosterveld is particularly remarkable

since this vegetation type is rather remote from Ka-

roo types of vegetation.

Jordaaniella and Fenestraria

A distribution pattern different from the afore-

mentioned ones is found in the genera Jordaaniella

H. E. K. Hartm. and Fenestraria N.E. Br. (Figure

11), which are distinguished from the other genera

of the subtribe by a considerable reduction of the

closing body (Hartmann 1982, 1983c, 1984).

The plants grow exclusively on littoral aeolian

sands, the distribution coinciding with Strandveld

(sensu Acocks 1975) in the coastal fogbelt (sensu

Nagel 1962) along the west coast. The occurrence

along the south coast agrees well with the ‘south

coast Strandveld’ as delineated by Moll et al. (1984),

although the easternmost population exceeds these

boundaries (Figure 11).

Only five species are recognized in the complex:

one in Fenestraria and four in Jordaaniella (Figure

11). Sympatry is restricted to a section along the

west coast (28°-30°30'S) with a maximum of three

species near 30°S, 17°E. Both endemic species (J.

clavifolia (L. Bol.) H. E. K. Hartm. and J. longifolia

209

(L. Bol.) H. E. K. Hartm.) occur in this region, but

it is doubtful whether the centre of origin lies here

(Hartmann 1984).

Clearly, the limited distribution areas of both gen-

era are due to their very narrow ecological ampli-

tudes leaving limited scope for speciation (Hart-

mann 1984).

FIGURE 11. — Distribution of Fenestraria (solid bars) and Jor-

daaniella (open circles = J. cuprea; open triangles = J.

clavifolia; full triangles = J. longifolia; full dots = J. du-

bia).

DISCUSSION

Interpretation of the phytogeographical data will

concentrate on two different aspects: firstly the com-

position, development and meaning of the two geo-

graphical centres, and secondly the distribution of

the genera and their possible evolution. In the pro-

cess, ideas will be developed on the origin and the

history of the Leipoldtiinae as a whole.

High species numbers in restricted geographical

areas mostly reflect wide ecological diversity, and

two processes can be distinguished: 1, specialization

in situ, which can be recognized by a high percentage

of endemic taxa; 2, overlapping of distribution areas

of species which have their main (mostly wide) ex-

tension outside the centre of diversity; in the centre

itself they colonize the ecologically and/or geo-

graphically extreme habitats to which they are

adapted.

In the first case close relationship between the

taxa is common, whereas in the second, relationship

is irrelevant, the effect being simply accumulative.

In both centres of multiplicity of the Leipoldtiinae, a

core of closely related species, including several en-

demics, is found (Figures 2, 5-11). In the Gariep

centre, the genera Cephalophyllum (Figure 6) and

Cheiridopsis (Figure 10) represent the major portion

(60-70%) of the occurring species. In the Vanrhyns-

dorp centre, species of Cephalophyllum (Figure 6)

210

and Argyroderma (Figure 5) contribute largely to

the formation of the centre (up to 14 out of 17

species in one 30' square). Of the additional taxa

occurring in these centres, some clearly reach the

limit of their distribution here: Vanzijlia and Hal-

lianthus in the Vanrhynsdorp centre (Figure 8); and

Hallianthus (Figure 8), J ordaaniella and Fenestraria

(Figure 11) in the Gariep centre.

Obviously, in both centres speciation and overlap-

ping of distribution areas contribute to the high de-

gree of diversity, and it can be suggested that several

subsequent events contributed to bring about the

present situation. A similar case has been reported

in the eastern Cape, for which Gibbs Russell & Ro-

binson (1981) stress the accumulative effect of differ-

ent phytochoria meeting, whereas Cowling (1982)

points out certain groups with high endemism.

Both centres of the Leipoldtiinae exhibit extreme

variation in geology and soils, often in island pattern

distribution, a condition considered favourable for

speciation of semi-isolated populations. In addition,

the dense mosaic of different ecological niches in

both centres permits short-distance migration under

changing climatic conditions, thus allowing persist-

ence of taxa within the area.

Consequently, it is not surprising that both phyto-

geographical centres have been traced in other

groups as well (e.g. Nordenstam (1969) who named

the Vanrhynsdorp centre, and circumscribed a ‘Gar-

iep centre’ in the north). Of the two, the less promi-

nent Vanrhynsdorp centre seems the more uniform,

its undulating surface suggesting differentiation

mainly in edaphic factors (e.g. Nordenstam 1969).

Yet, on close investigation a wealth of ecological

niches is found and is reflected in the vegetation

(Jurgens 1986).

The Gariep centre has been accepted more readily

as a phytogeographical centre because its geological

differentiation is complimented by a large relief for-

mation into which the extensions of the surrounding

widespread plains reach (e.g. Tolken pers. comm.;

Nordenstam 1969; Goldblatt 1976; Moffett 1979).

The Gariep centre of the Leipoldtiinae covers a

wider area (Figures 3 & 4) than the one circum-

scribed by Nordenstam (1969) and receives varying

amounts of annual precipitation. It is not surprising,

therefore, that not a single species occurs over the

entire range. Distinct subcentres of genera can be

seen: In Cephalophyllum (Figure 6), species num-

bers decrease from northwest to southeast, in Cheiri-

dopsis (Figure 10) an increase occurs in the same

direction, and Leipoldtia has its highest species fre-

quency in the two central 30' squares (Figure 8).

Yet, it seems impossible to subdivide the centre into

smaller entities, and it seems possible that within the

area migration movements could have taken place.

Six species of five genera occur in both phytogeo-

graphical centres: Cephalophyllum inaequale L.

Bol., Cephalophyllum pillansii L. Bol., Hallianthus

planus (L. Bol.) H. E. K. Hartm., Jordaaniella cu-

prea (L. Bol.) H. E. K. Hartm., Leipoldtia schultzei

(Schltr. & Diels) Friedrich and Vanzijlia annulata

(Berger) L. Bol. As can be expected, all of them

have their main distribution areas outside the

Bothalia 17,2 (1987)

centres, thus contributing to the ‘overlapping effect’

(see above).

All nine genera have different distribution pat-

terns, and three main types can be distinguished: A

— restricted distribution, B — wide distribution with

one distinct centre, C — wide distribution without a

distinct centre.

The genera Argyroderma (Figure 5) and Odonto-

phorus (Figure 9) represent examples of pattern A

and their restricted geographical occurrence is corre-

lated with marked ecological adaptations (Hartmann

1976, 1978a).

Edaphic factors seem to play important roles in

both genera, indicating that speciation may have oc-

curred in the present areas of distribution. This

would suggest that appropriate climatic conditions

have prevailed for a considerable period allowing

extensive radiation processes to take place.

A genus of wide distribution with one distinct

centre of diversity (pattern B) is Cheiridopsis, where

highest species numbers and occurrence of all three

subgenera coincide (Figures 9 & 10). Based mainly

on morphological data, Hartmann & Dehn (1987)

suggest that the centre of origin of the genus (and of

the closely related genus Odontophorus) lies here,

speciation having been promoted by the varied eco-

logical conditions within the southern Gariep centre

(see above). Marginal speciation (via semi-isolated

populations) in extreme habitats seems to have been

rare and has been recorded for Cheiridopsis caroli-

schmidtii (Dinter & Berger) N.E. Br. in the north on

weathered gneiss in the March rainfall area, and

Cheiridopsis rostrata (L.) N.E. Br. in the south-west

on sandy soils in West Coast Strandveld (sensu Moll

et al. 1984). Leipoldtia can be taken as another ex-

ample of distribution pattern B, but the centre is less

prominent and marginal speciation (Figure 8) more

frequent — both smaller subgenera colonize the bor-

der areas. The genus could therefore also be seen as

an example of pattern C, which lacks a distinct

centre. The predominantly shrubby habit and gener-

ally low expression of succulence in leaf, stem and

root indicate that Leipoldtia may have evolved

under slightly wetter conditions than it experiences

today, perhaps even outside its present distribution

area.

The widespread monotypic genera Fenestraria

(Figure 11), Hallianthus and Vanzijlia (Figure 8) na-

turally lack centres of species richness, and even sur-

veys for character multiplicity (Hartmann 1982,

1983a, 1983b) have not yielded positive results. Dif-

ferentiation patterns can be correlated with geo-

graphical distribution (flower characters in Fenestra-

ria, Hartmann 1982) or ecological factors (germi-

nation in Vanzijlia, Hartmann 1983a) or they can

occur incidentally (flower characters in Hallianthus

with island-type distribution, Hartmann 1983b).

Adaptation to rocky habitats restricts the distribu-

tion of Hallianthus. The genus could well have oc-

curred over wider conjunct areas at a time when

sedimentary plains covered less ground.

Assessment of the widespread genus Jordaaniella

is complicated by the strict adaptation to coastal

habitats (Figure 11) over several climatic ranges. Ex-

Bothalia 17,2 (1987)

211

tensive migration along the coast could have taken

place, and although three of the four species occur in

close proximity (near 30°S, 17°E, Figure 11) it does

not seem justified to conclude that this region is the

centre of origin (see also Hartmann 1984). Never-

theless, the creeping habit and prolonged germi-

nation indicate that the genus probably originated in

a coastal habitat.

The large genus Cephalophyllum exhibits a com-

plex distribution pattern which cannot be assigned to

one of the types mentioned above. Following the

character analyses of Hartmann (1978b, in press),

the most primitive capsule types and least special-

ized growth forms occur in species in the Vanrhyns-

dorp centre, where the highest species frequencies of

the genus are found as well (Figure 6). But the num-

ber of endemics is low, suggesting that the present

species richness is mainly due to overlapping of dis-

tribution areas.

In contrast, the species of the Gariep centre show

a high degree of uniformity in capsule morphology,

differing mainly in habitat and seed characters, so

that this region can be considered to represent a

centre of origin for part of the genus (Hartmann in

press).

In contrast to Cheiridopsis (above), relations be-

tween the two subgenera cannot be elucidated from

distribution data alone (Figure 2). This is mainly due

to the more or less strict allopatric occurrence of the

nine species of Cephalophyllum subgenus Cephalo-

phyllum, in which highest frequencies per 30' square

(three species, Hartmann in press) are attained

mainly by overlapping of wide distribution areas

(only two species of restricted distribution are

known, one of them isolated in the south-west, see

Figure 6). Speciation in this subgenus seems to have

followed predominantly the parapatric model (sensu

Wiley 1981) in which narrow contact zones existed

between diverging populations.

Some speculations can be made on the phytogeo-

graphical development of Cephalophyllum. The pre-

decessor of this genus (and Argyroderma?) could

have existed in the Vanrhynsdorp centre, but a ma-

jor part of the genus would have occurred in (or

migrated to ?) adjacent areas. Differentiation of the

two subgenera possibly took place outside the pres-

ent distribution range, probably followed by a phase

of extensive geographical spreading — Cephalophyl-

lum subgenus Cephalophyllum mainly into the wet-

ter south, subgenus Homophyllum mainly into the

drier north of the present distribution area. While

parapatric speciation may have prevailed in subge-

nus Cephalophyllum, radiation of a considerable

part of the subgenus Homophyllum could have

taken place in the Gariep centre. Distribution pat-

tern B (above) can be applied to this species group.

At first sight it seems improbable that a common

pattern for the biogeographical development of the

Leipoldtiinae can be found. But the extensive mesh-

ing of character states, as demonstrated for the sub-

tribe by Hartmann (1983c), clearly indicates that it

constitutes a monophyletic group. The area of origin

cannot be deduced from present distribution data

and may well have lain outside the existing range.

Yet, it was probably not too far away because speci-

ation processes are generally slow and many taxa of

restricted occurrence (e.g. species of Cheiridopsis

subgenus Odontophoroides) and/or with narrow eco-

logical amplitude (e.g. Argyroderma, Fenestruria)

can hardly be imagined to have migrated far.

With respect to ecology, the present data suggest

that a winter rainfall regime with rather low annual

amounts of precipitation (<200 mm, Figure 5) could

have prevailed in the area of origin. The few extant

exceptions could be seen as ecological fugitives. It is

difficult though to localize an adequate region in the

past, as opinions on past climates in South Africa are

highly controversial (e.g. Van Zinderen Bakker Sr

1978; Deacon 1983), in particular with reference to

the extent of the winter rainfall area (e.g. Van Zin-

deren Bakker Sr 1976, 1978; Lancaster 1979) in the

late Pleistocene. In any case, it is accepted that

higher precipitation prevailed in the west than at

present (Kent & Griebnitz 1985). This leads some

authors to the conclusion that the Namib is of rather

recent origin (e.g. Tankard & Rogers 1978; Axelrod

& Raven 1978) while others (e.g. Lancaster 1979)

assume that arid ‘isolated refuges along the Namib

coast’ or widespread deserts (e.g. Sarnthein 1978)

existed during the wet phases of the late Pleistocene.

Van Zinderen Bakker Sr (1978) also states that the

Namib is ‘ancient’ and has only marginally been in-

fluenced by higher rainfall, retaining its ‘hyper-arid

centre’ since the late Tertiary. An area like this

could present an ideal cradle for speciation, permit-

ting successive peripheral isolation of populations

while re-migration could occur as well.

It seems possible therefore, that the predecessors

of the extant genera of Leipoldtiinae originated on

the fringes of an ancient Namib, probably under

slightly higher rainfall regimes than today (based on

the assumption that the less pronounced xeromor-

phic character expressions in Leipoldtia, Cephalo-

phyllum and Jordaaniella reflect the more primitive

stages). The existence of strictly coastal taxa (Jor-

daaniella, Fenestraria) suggests that similar habitats

were already available during early speciation pro-

cesses.

The origin of the Leipoldtiinae as a whole remains

even more obscure than the diversification within

the subtribe. By its unique morphological character

set, the taxon is rather isolated within the family,

although the xeromorphic leaf characters indicate

that the closest relatives must be sought among the

xeromorphic genera of the subfamily Ruschioideae.

The main frequency centre of these genera lies

around the mouth of the Orange River, coinciding

with the Gariep centre of the Leipoldtiinae. A less

pronounced centre is found in the Little Karoo, and

a third minor one in the Knersvlakte (Vanrhynsdorp

centre), but genera of xeromorphic Ruschioideae

occur as far east at 29°E, and as far north as 26°S in

almost every degree square. Leipoldtiinae colonize a

restricted area within the range of the Ruschioideae,

thus emphasizing the relative isolation of the group.

They contribute substantially to the formation of the

western frequency centres (up to 50% of genera per

30' square), and present a typical ‘arid winter rain-

fall’ distribution.

212

Bothalia 17,2 (1987)

The possibility cannot be excluded that this cli-

matic regime presents the centre of origin of the

xeromorphic members of the Ruschioideae, but in

the absence of information from less derived sub-

tribes no evidence can be offered for or against the

suggestion.

With reference to the entire family, it may be im-

portant to note that distribution patterns of both

subfamilies coincide almost completely, and that

non-xeromorphic genera have their centres of fre-

quency in the winter rainfall region as well. It can

therefore be suspected that adaptation to winter

rainfall with lower precipitation amounts could pre-

sent a primary step of evolution within Mesembryan-

themaceae, but the phytogeography of the Leipoldt-

iinae cannot be used as proof because of their

specialized derived character set.

ACKNOWLEDGEMENTS

These studies were made possible through sub-

stantial financial support from the Deutsche

Forschungsgemeinschaft and additional grants from

the University of Flamburg, the National Botanic

Gardens of South Africa and the Council for Scien-

tific and Industrial Research. To all of them I extend

my sincere thanks.

For further support I am grateful to Mr M. B.

Bayer (formerly Karoo Garden, Worcester), the

Department of Botany of the University of Stellen-

bosch, and the staff of the Compton Herbarium, the

Botanical Research Unit (Stellenbosch) and the Bo-

tanical Research Institute (Pretoria). I would also

like to thank all friends in South Africa for encour-

aging help and all the members of the Hamburg

study group ‘Mesembryanthemaceae’, who have

been engaged in the project at some stage or other.

Thanks are also due to the directors of the herbaria,

B, BM, BOL, K, M, NBG, SAM and UPP for gen-

erous and long-term loans of valuable material.

REFERENCES

ACOCKS, J. P. H. 1975. Veld types of South Africa, edn 2.

Memoirs of the Botanical Survey of South Africa No. 40.

AXELROD, D. I. & RAVEN, P. H. 1978. Late Cretaceous and

Tertiary vegetation history of Africa. In M. J. A. Werger,

Biogeography and ecology of southern Africa. Junk, The

Hague.

COWLING, R. M. 1982. Patterns of plant endemism in the

south-eastern Cape. The Naturalist 27 ,1:17-20.

DEACON, H. J. 1983. Another look at the Pleistocene climates

of South Africa. South African Journal of Science

79:325-328.

GIBBS RUSSELL, G. E. & ROBINSON, E. R. 1981. Phytogeo-

graphy and speciation in the vegetation of the eastern

Cape. Bothalia 13:467-472.

GOLDBLATT, P. 1976. The genus Moraea in the winter rainfall

region of southern Africa. Annals of the Missouri Botanical

Garden 63:657-682.

HARTMANN, H. 1976. Monographic der Gattung Odontopho-

rus N.E. Br. (Mesembryanthemaceae Fenzl). Botanische

Jahrbiicher 97:161-225.

HARTMANN, H. 1978a. Monographic der Gattung Argyro-

derma N.E. Br. (Mesembryanthemaceae Fenzl). Mittei-

lungen aus dem Institut fur Allgemeine Botanik, Hamburg

15:121-235.

HARTMANN, H. 1987b. Zur Kenntnis der Gattung Cephalo-

phyllum N.E. Br. Botanische Jahrbiicher 99:264—302.

HARTMANN, H. 1982. Monographien der Subtribus Leipoldtii-

nae. III. Monographic der Gattung Fenestraria (Mesem-

bryanthemaceae). Botanische Jahrbiicher 103:145-183.

HARTMANN, H. 1983a. Monographien der Subtribus Leipoldt-

iinae. IV. Monographic der Gattung Vanzijlia (Mesem-

bryanthemaceae). Botanische Jahrbiicher 103:499-538.

HARTMANN, H. 1983b. Monographien der Subtribus Leipoldt-

iinae. V. Monographic der Gattung Hallianthus (Mesem-

bryanthemaceae). Botanische Jahrbiicher 104:143-169.

HARTMANN, H. 1983c. Untersuchungen zum Merkmalsbe-

stand und zur Taxonomie der Subtribus Leipoldtiinae (Me-

sembryanthemaceae). Bibliotheca Botanica 136:1-67.

HARTMANN, H. 1984. Monographien der Subtribus Leipoldtii-

nae. VI. Monographic der Gattung Jordaaniella (Mesem-

bryanthemaceae). Botanische Jahrbiicher 104:321-360.

HARTMANN, H. in press. Monographien der Subtribus Lei-

poldtiinae. VIII. Monographic der Gattung Cephalophyl-

lum (Mesembryanthemaceae). Mitteilungen aus dem Insti-

tut fur Allgemeine Botanik Hamburg 22.

HARTMANN, H. & DEHN, M. 1987. Monographien der Sub-

tribus Leipoldtiinae. VII. Monographic der Gattung Chei-

ridopsis (Mesembryanthemaceae). Botanische Jahrbiicher

108:567-663.

JURGENS, N. 1986. Untersuchungen zur Okologie sukkulenter

Pflanzen des siidlichen Afrika. Mitteilungen aus dem Insti-

tut fur Allgemeine Botanik Hamburg 21:139-365.

KENT, L. E. & GRIBNITZ, K.-H. 1985. Freshwater shell depos-

its in the north-western Cape Province: further evidence

for a widespread wet phase during the late Pleistocene in

southern Africa. South African Journal of Science

81:361-370.

LANCASTER, I. N. 1979. Evidence for a widespread late Pleis-

tocene humid period in the Kalahari. Nature 279:145-146.

MOFFETT, R. O. 1979. Sarcocaulon and the Gariep: products of

a hostile environment. Paper presented to the 5th Annual

Congress of the South African Association of Botanists,

Stellenbosch: 24-27.

MOLL, E. J., CAMPBELL, B. M., COWLING, R. M., BOSSI,

L., JARMAN, M. L. & BOUCHER, C. 1984, A descrip-

tion of major vegetation categories in and adjacent to the

Fynbos biome. South African National Scientific Program-

mes Report No. 83.

NAGEL, J. F. 1962. Fog precipitation measurements on Africa’s

south-west coast. Notos 11:51-60.

NORDENSTAM, B. 1969. Phytogeography of the genus Fu-

ry ops (Compositae). A contribution to the phytogeogra-

phy of southern Africa. Opera Botanica No. 33.

SARNTHEIN, M. 1978. Sand deserts during glacial maximum

and climatic optimum. Nature 272:43-46.

TANKARD, A. J. & ROGERS, J. 1978. Late cenozoic pa-

laeoenvironment on the west coast of southern Africa.

Journal of Biogeography 5:319-337.

VAN ZINDEREN BARKER Sr, E. M. 1976. The evolution of

Late-Quarternary palaeoclimates of southern Africa. Pa-

laeoecology of Africa 9:160-202.

VAN ZINDEREN BARKER Sr, E. M. 1978. Quarternary vege-

tation changes in southern Africa. In M. J. A. Werger,

Biogeography and ecology of southern Africa. Junk, The

Hague.

WERGER, M. J. A. 1978. Biogeographical division of southern

Africa. In M. J. A. Werger, Biogeography and ecology of

southern Africa. Junk, The Hague.

WILEY, E. O. 1981. Phylogenetics. Wiley, New York.

Bothalia 17,2: 213-227 (1987)

Preliminary floristic analysis of the major biomes in southern

Africa

G. E. GIBBS RUSSELL *

Keywords: biomes, Desert, Fynbos, Grassland, Karoo, Nama-Karoo, Savanna, species diversity. Succulent Karoo

ABSTRACT

Over 24 000 plant taxa are known to occur in the southern African flora, which is extraordinarily rich on a

species/area basis. Lists of species and infraspecific taxa recorded for the six major biomes, Fynbos, Savanna,

Grassland, Nama-Karoo, Succulent Karoo and Desert, were obtained from the PRECIS specimen database.

These lists were analysed by numbers of unique and shared species and infraspecific taxa, by differential occur-

rence and life forms of large genera, and by differential occurrence of families. Each biome is floristically distinct

except Nama-Karoo. The biomes form two main groupings, those with winter rainfall and those with summer

rainfall. Succulent Karoo is most similar to Fynbos and Nama-Karoo is most similar to Savanna.

UITTREKSEL

Dit is bekend dat meer as 24 000 planttaksons in die suider-Afrikaanse flora voorkom, wat op 'n spesies/area-

grondslag buitengewoon ryk is. Lyste van spesies en infraspesifieke taksons van die ses hoofbiome, Fynbos,

Savanne, Grasveld, Nama-Karoo, Sukkulente Karoo en Woestyn, is vanaf die PRECIS-eksemplaardatabasis

verkry. Hierdie lyste is ontleed in terme van unieke en gemeenskaplike spesies en infraspesifieke taksons, differen-

siele voorkoms en lewensvorme van groot genusse, en die differensiele voorkoms van families. Elke bioom

behalwe Nama-Karoo, is floristies kenmerkend. Die biome vorm twee hoofgroeperings, die met winterreenval en

die met somerreenval. Sukkulente Karoo toon die meeste ooreenkoms met Fynbos en Nama-Karoo toon die

meeste ooreenkoms met Savanne.

CONTENTS

Introduction 213

Methods 214

Results and discussion 216

Area, taxa and specimens 216

Comparison of biomes by numbers of species

and infraspecific taxa 216

Sorenson’scoefficientsofsiinilarity 216

Percentagesofuniqueandsharedtaxa 217

Comparison of biomes by important families

and large genera 217

Differential occurrence of important families 217

Centres of diversity of large genera 219

Life forms and centres of diversity of large

genera 220

Floristic characteristics and relationships of

the biomes

Fynbos 221

Savanna 221

Grassland 221

Nama-Karoo 221

Succulent Karoo 221

Desert 221

Relationships 222

Conclusions 222

Acknowledgements 222

References 222

Appendix 1 223

Appendix 2 225

Appendix 3 226

Appendix 4 227

* Botanical Research Institute, Private Bag X101, Pretoria 0001.

INTRODUCTION

The southern African flora is extremely species-

rich in terms of species/area ratios, with 0,0081

species/km2 overall (Figure 1). This value is higher

than those recorded for humid tropical floras such as

Brazil (0,0044) and Asia (0,0041) (Gibbs Russell

1985b). The winter rainfall Cape Floral Kingdom is

well known to be extremely species-rich (Goldblatt

1978). However, even when the Cape flora is ex-

cluded from calculation, the species/area ratio for

the rest of the southern African flora (0,0061) is still

considerably higher than that of the humid tropics,

and nearly twice that of Australia (0,0032), which

also includes both tropical and temperate areas.

These species/area ratios indicate in a superficial

way that the remarkable species richness of the

southern African flora is not restricted to the Cape

Floral Kingdom. The aim of this study is to investi-

gate the floristic richness of the major biomes and to

explore floristic relationships between these biomes

using distribution data for families, genera and

species.

At the present time, the PRECIS (Pretoria Na-

tional Herbarium Computerized Information Sys-

tem) specimen database is by far the most compre-

hensive source of information on the distribution of

plant taxa in southern Africa. Although PRECIS has

certain limitations (see Methods), this preliminary

study forms a base against which more detailed stu-

dies of particular biomes can be put in context, and

which will allow the generation of hypotheses to

guide future studies. A re-evaluation should be done

when more complete checklists, based on co-opera-

tive herbarium studies and intensive field work, have

been compiled for all the biomes.

214

Bothalia 17,2 (1987)

FIGURE 1. — Species/area ratios

for large regions. The num-

ber of species and areas in

km2 for each region follow

Gibbs Russell (1985b).

METHODS

This study is based on checklists compiled from

PRECIS for quarter degree latitude and longitude

grids representing the major biomes for southern

Africa. The biomes adopted were determined by

superimposing five recent treatments of southern

African vegetation using floristic, structural and en-

vironmental criteria (Werger 1978; Scheepers 1982

based on Acocks 1975; White 1983; Huntley 1984;

Rutherford & Westfall 1986). The resulting compos-

ite map showed six major regions that were recog-

nized as entities, even though none of the studies

agreed on exact boundaries. Elimination of all areas

of disagreement, and of areas smaller than a quarter

degree, yielded the regions accepted as the core bi-

omes for this investigation (Figure 2). Important en-

vironmental characteristics of the core biomes are

shown in Table 1.

The lack of agreement between the treatments

occurred at three levels: 1, exact boundaries at quar-

FIGURE 2. — Quarter degree grids searched in PRECIS for each biome.

Bothalia 17,2 (1987)

215

TABLE 1. — Characteristics of the biomes

* Rutherford & Westfall (1986)

t Huntley (1974)

ter degree scale; 2, areas of transition between kar-

roid and savanna regions; and 3, areas of compli-

cated vegetation relationships, such as the eastern

Transvaal, Natal and the eastern Cape. The two

‘karroid’ biomes accepted here were recognized at

the highest level of classification only by Rutherford

& Westfall (1986), on grounds of differences in

dominant plant life form and environmental condi-

tions. Their Succulent Karoo Biome roughly coin-

cided with the Western Cape Domain of the Karoo-

Namib Region defined by Werger (1978) at a sec-

ondary level of classification on phytochorological

grounds. The other three vegetation studies treated

the entire karroid vegetation as a single entity at the

highest level. In this investigation. Succulent Karoo

was separated from Nama-Karoo, and a secondary

aim of the study was to examine the floristic relation-

ship between them.

Besides the six core biomes adopted for this study,

all the treatments recognized the high altitude vege-

tation of the Drakensberg, and the forests of the

southern Cape. However, these small irregularly

shaped areas were not accessible to computer search

at the scale of quarter degree grid reference, and

could therefore not be included.

The PRECIS specimen database records label in-

formation for ± 610 000 specimens in the National

Herbarium (PRE). The overall operation and imple-

mentation of PRECIS have been reported several

times (Gibbs Russell & Gonsalves 1984; Gibbs Rus-

sell 1985a). More recently, new programming has

allowed compilation from the database of checklists

of plant species and infraspecific taxa from any com-

bination of quarter degree grids. Several special pro-

grammes were written to compare the checklists by

providing lists of unique taxa, lists of shared taxa,

and a matrix of all taxa with the biomes from which

they were recorded.

The total numbers of unique and shared taxa, ob-

tained by employing these programmes, were used

to calculate Sorenson’s (1948) coefficients of similar-

ity, and percentages of unique taxa and taxa shared

between biomes. The ranking of families for each

biome, the identification of widespread taxa, and the

determination of centres of diversity for ‘large’ gen-

era with 10 or more species and infraspecific taxa

were obtained by manual searches of printout. A

biome was considered to be a centre of diversity for

a genus if it contained 50% or more of the taxa re-

ported for the genus. In a few cases, slightly less than

half (to 45%) was accepted in biomes of low collect-

ing intensity. Life forms follow the definitions of

Raunkiaer (1934) as stated by Rutherford & West-

fall (1986), but with the inclusion of ‘Succulent’, and

were determined from Dyer (1975, 1976) and her-

barium specimens. At all stages of work, doubtful

records encountered on PRECIS listings were

checked in PRE.

An inherent weakness in the method used is the

uneven collecting intensity for the different biomes.

Gibbs Russell et al. (1984) showed that the collecting

intensity represented in PRECIS for the eastern and

southern mesic areas is far higher than for the west-

ern arid areas. Therefore, the checklists used here

undoubtedly differ in completeness, and it must be

emphasized that these results are preliminary. Table

2 illustrates the differences in collecting intensity be-

tween the biomes by comparing the specimens and

the taxa per km2 as well as the specimens per taxon

recorded in PRECIS for each biome. Although Fyn-

bos, and to a lesser degree Grassland, appear to be

better collected than the other biomes on a speci-

mens/km2 or taxa/km2 basis, Savanna in fact exhibits

more ‘repeat’ collections than either. However, it is

apparent that mesic Fynbos, Savanna and Grassland

are better collected than arid Nama-Karoo, Succu-

lent Karoo and Desert.

PRECIS is known to have errors in about 7% of

specimen identifications and quarter degree grid re-

ferences. Until these errors can be corrected, an

ongoing process in system management, results must

be used with discretion. In this study, identifications

directly from PRECIS are used only at the level of

family and genus, while at the level of species and

infraspecific taxa, only total numbers, and not iden-

tifications, are used unless the records were checked

216 Bothalia 17,2 (1987)

TABLE 2. — Collecting intensity reported from PRECIS for each biome. Area was determined from the number of quarter degree

grids searched (and ‘average’ quarter degree covers 666 km2)

in PRE. For the same reason, distribution is given

only at biome level, and not to individual quarter

degree grids.

Despite the limitations imposed by differences in

collecting intensity and by the accuracy of individual

PRECIS records, at the present time PRECIS is the

most reliable and complete source of information

about the distribution of taxa throughout the south-

ern African flora. Publication of these preliminary

results is therefore considered worthwhile.

Throughout the study, the number of species and

infraspecific taxa, rather than species alone, were

used in comparisons because of taxonomic uncer-

tainty about the correct level of treatment for many

of these entities, as explained in detail in Gibbs Rus-

sell (1985b). For the sake of brevity, the term ‘taxa’

in this context is used in place of the longer phrase

‘species and infraspecific taxa’.

RESULTS AND DISCUSSION

Area, taxa and specimens

The area, taxa and specimens covered in this study

are summarized in Table 3. The five recent vegeta-

tion treatments used to determine the biomes for

this study agreed on about 40% of the total area of

southern Africa at a scale available for computer

search. About 60% of all southern African taxa re-

presented in PRECIS were reported from the area

designated. Certain taxa were not included in the

study for the following reasons: 1, they are known

only outside the areas of the core biomes; 2, they are

not represented in PRECIS; or 3, they are repre-

sented in PRECIS, but the distribution is not re-

corded as a quarter degree grid. Only about 25% of

the specimens in PRECIS are reported in the study.

This low figure results from the uneven collecting

intensity in the National Herbarium mentioned

above.

TABLE 3. — Total size of sample reported for all biomes

Number of specimens 146 298

Out of 610000 in PRECIS 24%

Out of ±2 000 000 in southern African herbaria 7%

Number of taxa 14 391

Out of 24 000 in southern Africa 60%

Area covered (1611 quarter degree grids

@666 km2 per quarter degree) 1 072 926 km2

Out of 2 573 000 km2 for southern Africa 42%

Comparison of biomes by numbers of species and

infraspecific taxa

Widely differing numbers of taxa have been re-

corded for the six biomes, and the differences in

taxon numbers are not related to the area sampled

(Table 2). Fynbos has the most taxa although it is the

smallest in area. Savanna, which covers by far the

largest area, has about 1 500 fewer taxa than Fynbos.

Similarly, Grassland has about 1 700 more taxa than

Nama-Karoo, although Nama-Karoo covers about

twice the area of Grassland. Nama-Karoo and Suc-

culent Karoo have similar numbers of taxa, but

Nama-Karoo covers about four times the area of

Succulent Karoo. The number of taxa recorded for

Desert is extremely low even though its area is

slightly larger than that of Fynbos.

The checklists for each biome were compared

both by Sorenson’s (1948) coefficient of similarity,

and by percentage comparisons within each biome.

Sorenson’s coefficients give comparable values for

checklists of different length. Low Sorenson’s coef-

ficients signify low similarity between lists of taxa,

while higher values show a greater similarity. The

percentage comparisons show the proportion of taxa

within each biome that are unique and that are

shared with other biomes.

Sorenson’s coefficients of similarity

The Sorenson’s coefficients of similarity between

the six major biomes are shown in Figure 3. The

values for the coefficients are generally low (30 or

less), indicating that each biome has its own flora

which is quite distinct from that of the others. The

exception is the coefficient between Savanna and

Grassland, which is considerably higher than any

other.

For Savanna, the highest Sorenson’s coefficients

occur with Grassland and with Nama-Karoo, and the

values are low (less than 20) for the other biomes.

Grassland, which has the strongest similarity to

Savanna, has very low Sorenson’s coefficients with

Desert and with Succulent Karoo, and somewhat

higher values with Fynbos and Nama-Karoo. Desert

has very low values with all biomes except Nama-

Karoo. Fynbos has low Sorenson’s coefficients with

all biomes except Succulent Karoo. Succulent Karoo

and Nama-Karoo show opposite relationships. Ex-

cluding the Sorenson’s coefficient between the two

‘karroid’ biomes, Succulent Karoo has its highest

value with Fynbos, and very low values with Desert,

Bothalia 17,2 (1987)

217

FIGURE 3. — Number of taxa and Sorenson’s coefficients of

similarity for the biomes. The number in each large circle is

the number of species and infraspecific taxa reported for

the biome. The number in each small circle is the Soren-

son’s coefficient of similarity between pairs of biomes.

Savanna and Grassland, whereas Nama-Karoo has

its highest value with Savanna, high values with De-

sert and Grassland, and a low value with Fynbos.

Percentages of unique and shared taxa

The percentages of taxa unique to each biome and

shared between biomes are shown in Figure 4. The

biomes vary greatly in percentages of unique taxa.

Fynbos has the highest percentage (which is consist-

ent with a value of 68% given by Bond & Goldblatt

(1984)), and Savanna is also well above the others.

Grassland and Succulent Karoo are similar, and De-

sert and Nama-Karoo have similar and very low per-

centages of unique taxa.

FIGURE 4. — Number and percentage of species and infraspe-

cific taxa unique to and shared between biomes. In each

large circle, the upper number is the number of unique

taxa, the lower number is the total number of taxa for the

biome, and the percentage is the percentage of taxa unique

for the biome. The number in each small circle is the per-

centage of taxa shared between pairs of biomes, and the

number on the line connecting a pair is the Sorenson’s

coefficient of similarity.

The percentage of taxa shared between the bi-

omes amplifies the relationships shown by the So-

renson’s coefficients. The few apparent contradic-

tions result from comparing taxon lists of very dif-

ferent length: where one list is long and the other

short, the percentage of shared taxa differs markedly

from the Sorenson’s values.

The close floristic relationships between Savanna

and Grassland and Savanna and Nama-Karoo showr

by the Sorenson’s coefficients are borne out by the

high percentage of Grassland and Nama-Karoo taxe

that is shared with Savanna. Savanna itself shares

most taxa with Grassland, shares the same percent-

age of its taxa with Nama-Karoo as with Fynbos, and

shares a very low percentage of its taxa with Suc-

culent Karoo and Desert. Grassland shares a very

high percentage of its taxa with Savanna, and is simi-

lar to Savanna in that its lowest percentage of shared

taxa is with Succulent Karoo and Desert, but Grass-

land shares a considerably higher percentage of taxa

with Fynbos than with Nama-Karoo. Desert, which

because of its small flora shows very low Sorenson’s

values with all biomes except Nama-KarOo, shares

about the same very high percentage of its taxa with

Savanna as with Nama-Karoo. Desert has a very low

percentage of unique taxa, and shares more than

20% of its taxa with Grassland, with Succulent Ka-

roo and with Fynbos. In contrast, Fynbos, which has

a high percentage of unique taxa, does not share

more than 20% of its taxa with any other biome. The

close relationships of Succulent Karoo to Fynbos

and of Nama-Karoo to Savanna, already indicated

by Sorenson’s coefficients, are borne out by the high

percentage of Succulent Karoo taxa shared with

Fynbos, and the high percentage of Nama-Karoo

taxa shared with Savanna. Both of the ‘karroid’ bi-

omes share the lowest percentage of taxa with De-

sert and an intermediate percentage with Grassland.

Comparison of biomes by important families and

large genera

Differential occurrence of important families

Forty-six families comprise 1% or more of the

taxa in at least one biome. In each biome there are

between 22 and 28 families that comprise 1% or

more of the taxa, and that together account for be-

tween 55 and 60% of the total number of taxa. Each

of these families can be used to distinguish and/or

link the biomes (Table 4).

In order to compare the biomes in this way, these

important families are ranked for every biome by

number of taxa from largest (rank of 1) to smallest

(rank of 22 to 28). Ranking is necessary for com-

parison at family level because the biomes differ so

greatly in number of taxa. A family well represented

in a species-poor biome may in fact have fewer taxa

in that biome than the same family has in a biome

with a rich flora, even though the family is a negligi-

ble component of the more species-rich biome

(Gibbs Russell 1975, 1985b). The families are

ranked in three groups in the discussion: the largest

families (1-3 in bold type in Table 4); the next rank

(4—10 in italics in Table 4); and the lowest rank (from

218

Bothalia 17,2 (1987)

11 onwards in roman type in Table 4). The biomes

are characterized by the presence, absence or differ-

ence in rank of certain large families, and the occur-

rence of some families can be linked to simple en-

vironmental parameters characteristic of certain

combinations of biomes.

The seven plant families that comprise 1% or

more of the taxa in all of the biomes are shown in

Table 4a. Three families, Asteraceae, Poaceae and

Fabaceae are the three largest in all biomes (with the

exception of Poaceae in Succulent Karoo and Fyn-

bos), and either Asteraceae or Poaceae is the largest

family in all biomes. Asteraceae and not Poaceae is

the largest family in Grassland.

The six biomes are briefly discussed in turn below:

Fynbos (Table 4b) is distinguished by eight fami-

lies that are important in no other biome. Of these,

Ericaceae is one of the three largest families, and

TABLE 4. — Families represented by more than 1% of the total number of taxa in any one of the biomes. Sv, Savanna; G, Grassland;

D, Desert; N-K, Nama-Karoo; SK, Succulent Karoo; F, Fynbos. The number in the matrix is the rank according to number

of taxa in the family in a given biome, with T’ signifying the largest family in the biome

Sv G D N-K SK F Sv G D N-K SK F

4a. Families comprising more than 1% of the total number of

taxa in all biomes

No families form more than 1% of flora only in Nama-Karoo.

No families have high rank only in Nama-Karoo.

No families are absent only from Nama-Karoo.

4f. Families that distinguish Succulent Karoo

High rank of :

Iridaceae 17 10 14 2 4

Crassulaceae 14 12 9 19

Geraniaceae 23 16 10 16

4g. Families that distinguish Desert

Presence of:

Pedaliaceae 17

Burseraceae 20

High rank of:

Bothalia 17,2 (1987)

219

Restionaceae, Rutaceae and Proteaceae among the

ten largest families in Fynbos only. In contrast, As-

clepiadaceae is not important, and only in Fynbos is

Scrophulariaceae not one of the ten largest families.

Savanna (Table 4c) is distinguished by one import-

ant family, Verbenaceae, one family, Rubiaceae,

that ranks among the ten largest in no other biome,

and one family, Mesembryanthemaceae, that does

not occur among the important families. Grassland

(Table 4d) is distinguished by the high rank of Orchi-

daceae and Lamiaceae, which are among the ten

largest families only in this biome, and only here are

Sterculiaceae and Aizoaceae absent from the im-

portant families. Nama-Karoo (Table 4e) is the only

biome which is not distinguished from the others by

differential occurrence of families. Succulent Karoo

(Table 4f) is distinguished by the high rank of Irida-

ceae, which is one of the three largest families, and

Crassulaceae and Geraniaceae, which are among the

ten largest families only in this biome. Desert (Table

4g) is distinguished by the occurrence of Pedaliaceae

and Burseraceae as important families, by the occur-

rence of Chenopodiaceae and Capparaceae among

the ten largest families, and by the absence of Irida-

ceae as an important family.

A number of families indicate floristic relation-

ships between biomes with different rainfall seasona-

lity and amount. The four summer rainfall biomes

(Table 4h) are variously linked by 11 families that do

not occur as an important component of the winter

rainfall biomes. Winter rainfall biomes (Table 4i)

are linked by the occurrence of three families, Pro-

teaceae, Oxalidaceae and Campanulaceae, that are

not important in summer rainfall areas, and one

family, Poaceae, that ranks first or second in sum-

mer rainfall biomes, but has a lower rank in the win-

ter rainfall areas.

In contrast to the above groupings based on rain-

fall seasonality, other families link biomes with simi-

lar amounts of rainfall. The arid biomes are linked

by four families (Table 4j). Chenopodiaceae and Zy-

gophyllaceae are important, and Aizoaceae and Me-

sembryanthemaceae are among the ten largest fami-

lies only in the arid biomes. Finally, a group of six

families, all with low ranking, weakly links the sum-

mer rainfall biomes Grassland and Nama-Karoo to

the winter rainfall biomes (Table 4k). Savanna is not

linked to the winter rainfall biomes at family level.

Centres of diversity of large genera

The large genera (with 10 or more taxa) with

centres of diversity in one, two or three biomes are

listed in Appendices 1-3. The large genera with no

apparent centre of diversity are listed in Appendix 4.

Figure 5 summarizes this information by showing the

numbers and percentages of large genera with

centres of diversity within and shared between the

biomes.

Only in the case of Fynbos and Savanna are more

than half the large genera centred in a single biome,

whereas each of the other four biomes shares more

than half its large genera with another biome. The

highest number of large genera have their centre of

FIGURE 5. — Number and percentage of large genera (10 or

more taxa) with centres of diversity in each biome and

shared between biomes. In each large circle, the upper

number is the number of large genera with a centre of

diversity only in that biome, the lower number is the total

number of large genera with a centre of diversity in that

biome, and the percentage is the percentage of large gen-

era with a centre of diversity only in that biome. The num-

ber in each small circle is the percentage of genera shared

between pairs of biomes, and the number on the line be-

tween a pair is the number of genera with shared centres of

diversity. Absence of linkage lines indicates no large gen-

era in common.

diversity in Fynbos, and 72% of these genera are

centred only in Fynbos. Fynbos is a shared centre of

diversity for similar numbers of genera with Suc-

culent Karoo and with Grassland, and for low num-

bers with Savanna and with Nama-Karoo. No genera

have centres of diversity in both Fynbos and Desert.

In Savanna, as in Fynbos, over half the large genera

have centres of diversity in no other biome, and Sav-

anna also shares genera with centres of diversity in

four other biomes. No genera have centres of diver-

sity in both Savanna and Succulent Karoo. For

Grassland, over half the large genera share their

centres of diversity with Savanna, and nearly a quar-

ter share their centres of diversity with Fynbos.

Grassland shares large genera only with Savanna

and Fynbos, and no genera have centres of diversity

in both Grassland and Nama-Karoo, Grassland and

Succulent Karoo or Grassland and Desert. A very

low percentage of large genera have their centre of

diversity in Nama-Karoo alone. Over two-thirds of

large genera in Nama-Karoo share their centre of

diversity with Savanna, and Nama-Karoo shares

genera with centres of diversity in all biomes except

Grassland. In Succulent Karoo, a very high percent-

age of large genera shares a centre of diversity with

Fynbos, and a low percentage shares a centre of di-

versity with Nama-Karoo. Succulent Karoo shares

large genera only with Fynbos and Nama-Karoo,

and no genera have centres of diversity in both Suc-

culent Karoo and Savanna, Succulent Karoo and

Grassland or Succulent Karoo and Desert. Only two

genera have their diversity centred in both of the

‘karroid' biomes, and this is the lowest percentage of

shared large genera for either Nama-Karoo or Suc-

culent Karoo. Only three large genera have a centre

of diversity in Desert, and all three are shared with

of life forms reported

220

Bothalia 17,2 (1987)

Savanna or with Nama-Karoo. No genera have

centres of diversity in both Desert and Grassland,

Desert and Succulent Karoo, or Desert and Fynbos.

Life forms and centres of diversity of large genera

Figure 6 shows life form spectra for large genera

with centres of diversity either only in one or in more

than one biome. The basis for plant classification is

that floral characters are conservative at family and

genus level, whereas vegetative characters can be

variable between members of a higher category.

Raunkiaer’s life forms indicate broad basic differ-

ences in vegetative states, depending on the position

of the perennating bud, and indicate differences in

utilization of resources. The fact that a genus has

many species and infraspecific taxa in a certain bi-

ome suggests that the adaptations displayed by the

taxa are compatible with the environment of that

biome. Thus the differences in characteristics of the

genera, as illustrated by life forms, can show conver-

gent adaptations in a number of separate evolution-

ary lines to the conditions in the biome. However, a

centre of diversity for a genus in a particular biome

does not imply that speciation occurred either in that

biome, or under current environmental conditions.

The biomes are characterized by differences in the

life forms reported in the large genera. In Fynbos,

chamaephytes are the most commonly reported life

form in the large genera. In Savanna, phanerophytes

are reported more frequently than in any other bi-

ome. For Grassland, nearly half the life forms re-

ported are hemicryptophytes. Grassland differs from

Savanna by having fewer phanerophytes and cham-

aephytes (the woody component), and from Nama-

Karoo by having fewer cryptophytes. Nama-Karoo

is similar to Grassland, but with more cryptophytes

reported among the few genera with their centre of

diversity in Nama-Karoo only. Succulent Karoo is

remarkable because it has similar values for chamae-

phytes, hemicryptophytes and cryptophytes. The

comparative value for cryptophytes is far higher than

for any other biome, and phanerophytes are not re-

ported at all. The life form spectrum for Desert may

be misleading because it is based on three genera

only, and therefore it is not considered further.

FIGURE 6. — Life form spectra of large genera (10 or more taxa)

for each biome. Life forms of genera with centres of diver-

sity only in a particular biome are shown by stripes, and life

forms in all genera with centres of diversity in a particular

as well as in other biomes are shown by stippling. Life

forms are indicated by the following symbols: P = phane-

rophytes; Ch = chamaephytes; H = hemicryptophytes;

Cr = cryptophytes; T = therophytes; S = succulents.

The differences in occurrence of each of the life

forms in the biomes can also be examined. Phanero-

phytes appear only in genera with a centre of diver-

sity in Fynbos or Savanna. Chamaephytes and hemi-

cryptophytes show a basic difference between the

summer and the winter rainfall biomes. Chamae-

phytes are reported most often in winter rainfall

Fynbos and Succulent Karoo. Hemicryptophytes are

the most abundant life form in the summer rainfall

Savanna, Grassland, Nama-Karoo and Desert.

Cryptophytes occur in low numbers in all the bi-

omes, but are reported often only in genera with

their centre of diversity in Succulent Karoo, and to a

lesser extent, Nama-Karoo. Therophytes are re-

ported in all biomes, but are less frequently re-

ported in genera of which the centre of diversity is

confined to a single biome, and are more frequently

reported in genera with centres in more than one

Bothalia 17,2 (1987)

221

biome. Succulents are reported in all biomes, but the

mesic biomes Fynbos, Savanna and Grassland, have

succulents reported in genera with centres in each

one, while the more arid Succulent Karoo and

Nama-Karoo have succulents reported only in gen-

era with centres of diversity in more than one biome.

Floristic characteristics and relationships of the

biomes

Fynbos

Fynbos has the largest number of taxa, the highest

percent of unique taxa, the largest number of im-

portant families that do not occur in any other bi-

ome, and the greatest number of centres of diversity

for large genera. At species level, the Sorenson’s

coefficient of similarity and the percentage of shared

taxa show that Fynbos is most closely related to Suc-

culent Karoo, the other winter rainfall biome. At the

generic level, Fynbos shares more centres of diver-

sity for large genera with Succulent Karoo than with

any other biome. At the family level, Fynbos is

linked only to Succulent Karoo by four important

families. The less-marked relationship between Fyn-

bos and Grassland will be discussed under Grassland

below.

Savanna

Savanna is second to Fynbos in number of taxa,

percentage of unique taxa, and in number of centres

of diversity for large genera. However, Savanna is

distinguished at family level by only three families,

while it is linked to the other summer rainfall biomes

by eight families. The closest relationship of Sav-

anna is to Grassland, as shown by the very high So-

renson’s coefficient of similarity and the percentage

of shared taxa, the number of large genera with

centres of diversity in both Savanna and Grassland,

and the six families that link them, three of which

are important only in Savanna and Grassland. A

weaker relationship between Savanna and Nama-

Karoo is shown by a high Sorenson’s coefficient and

the percentage of shared taxa, a considerable num-

ber of large genera with centres of diversity in both

Savanna and Nama-Karoo, and by four families that

link them.

Grassland

A moderately large number of taxa is reported for

Grassland, which is distinguished by four families.

Its relationship with Savanna is the closest demon-

strated in this study, as discussed above. Grassland

shows similar moderate Sorenson’s coefficients with

both Nama-Karoo and Fynbos, but other compari-

sons show that Grassland is in fact more closely re-

lated to Fynbos than to Nama-Karoo. The percent-

age of Grassland taxa shared with Fynbos is far

higher than the percentage shared with Nama-Ka-

roo, and a number of large genera, nearly all hemi-

cryptophytes, have centres of diversity in both

Grassland and Fynbos, while no large genera have

centres of diversity in both Grassland and Nama-Ka-

roo. At family level. Grassland is linked to Nama-

Karoo only by families that also link it to Savanna

(Table 4h) or to Fynbos (Table 4k), while it is linked

independently to Fynbos by two families (Table 4k).

Nama-Karoo

Nama-Karoo is not well defined floristically in this

study. At species level, its number of taxa is low,

particularly with respect to its large area, and the

percentage of unique taxa is very low, hardly higher

than that of Desert. Nama-Karoo is the only biome

for which all Sorenson’s coefficients except one (to

Fynbos) are greater than 20. Over half of Nama-Ka-

roo taxa are shared with Savanna, about a third are

shared with Grassland and another third with Fyn-

bos. At generic level, few large genera have a centre

of diversity in Nama-Karoo, and of these, more have

a shared centre of diversity with Savanna, with Fyn-

bos or with Desert than are centred in Nama-Karoo

alone. At family level, Nama-Karoo is the only bi-

ome that cannot be defined by differential occur-

rence of important families. It is linked to all the

other summer rainfall biomes, and also to the winter

rainfall Succulent Karoo through the arid biomes.

Succulent Karoo

The number of taxa reported for Succulent Karoo

is similar to that of Nama-Karoo, but the area cov-

ered is about a quarter as large, and Succulent Ka-

roo has more unique taxa. It is distinguished from

other biomes by three important families. Succulent

Karoo is shown by Sorertson’s coefficients, by per-

centage of shared taxa and by centres of diversity of

large genera to be related floristically both to Fynbos

and Nama-Karoo. The much higher values in every

case show that the relationship is strongest to Fynbos

(see Fynbos above). Over half the Succulent Karoo

taxa and over three quarters of the large genera are

shared with Fynbos. At family level, the strong links

of Succulent Karoo to Fynbos are shown by four

families that are important only in these two biomes,

whereas at family level Succulent Karoo is linked to

Nama-Karoo only through the group of families that

links the three arid biomes.

Desert

A very small number of taxa are reported for De-

sert, and the percentage of unique taxa is lower than

for any other biome. There are no large genera with

a centre of diversity in Desert alone. However, De-

sert is distinguished by four important families. Re-

lationships of the Desert flora are shown by Soren-

son’s coefficients and by the percentage of shared

taxa, to be highest with Savanna and with Nama-Ka-

roo, and it is only with these two biomes that Desert

shares centres of diversity for large genera. In ad-

dition, Desert is linked to Nama-Karoo by ten fami-

lies, two of which are important only in Desert and

Nama-Karoo, and it is linked to Savanna by four

families, one of which is important only in Desert

and Savanna. Desert is also linked to the arid but

winter rainfall Succulent Karoo by four families.

222

Bothalia 17,2 (1987)

Relationships

The distribution of species, genera and families

and the life form spectra shows that the biomes fall

floristically into two groups, which correspond to the

summer rainfall region (Savanna, Grassland, Nama-

Karoo, Desert) and the winter rainfall region (Fyn-

bos. Succulent Karoo). The present analysis of

14 000 taxa therefore supports and extends the ‘win-

ter rainfall biome’ concept first put forward on the

basis of a few genera by Bayer (1984). A detailed

study of grass subfamily distributions also shows a

similar basic division, with Chloridoideae and Pan-

icoideae most abundant in summer rainfall areas and

Arundinoideae most abundant in winter rainfall

areas (Gibbs Russell 1986).

Nama-Karoo and Succulent Karoo, which have

previously been placed together at highest level in all

vegetation studies except that of Rutherford &

Westfall (1986), are not closely related floristically.

Nama-Karoo is more closely related to Savanna than

to Succulent Karoo, and Succulent Karoo is more

closely related to Fynbos than to Nama-Karoo.

Within the summer rainfall group, at species level,

the strongest relationship is between Savanna and

Grassland, with a weaker relationship between Sa-

vanna and Nama-Karoo. The same relationships are

shown at generic level, and the distinctness of Nama-

Karoo from Grassland and of Desert from Succulent

Karoo is emphasized. At family level, particular fa-

milies link and demarcate the summer rainfall bi-

omes and the winter rainfall biomes, but another

group of families complicates this simple difference

by linking the arid biomes of both summer and win-

ter rainfall regimes.

Secondary links connect the two major groups

through Nama-Karoo, which lies between the other

biomes geographically. Nama-Karoo is ill-defined as

an entity, and is strongly linked at species, genus and

family level to Savanna and Desert; it is more

weakly linked at species and family level to Succu-

lent Karoo and at genus level to Fynbos. Grassland,

which is very strongly allied to Savanna, shows a

secondary link to Fynbos, independent of Nama-Ka-

roo, at species, genus and family level.

CONCLUSIONS

At the highest level of floristic comparison the

winter rainfall biomes and the summer rainfall bi-

omes form two separate groups. Within these

groups, each biome is floristically distinct at the level

of species and infraspecific taxa, whether measured

by Sorenson’s coefficient of similarity or by percent-

age of shared taxa, and each biome (except Desert)

is rich in taxa. Each is a centre of diversity for certain

large genera, and the life form spectrum for these

genera is different for each biome. Each (except

Nama-Karoo) is distinguished by differences in the

occurrence of important plant families.

The floristic distinctness of the biomes, coupled

with high taxon numbers, implies that each should

be studied and managed as a separate entity. Be-

cause of the high numbers of species and infraspe-

cific taxa, it is unlikely that conservation of limited

areas in nature reserves will protect a large propor-

tion of the taxa in any one biome.

This study is hampered by the dearth of specimen

records from arid areas, and for this reason it may be

criticized for being too preliminary. However, the

trends indicated should serve as stimulus to more

precise analyses. Unfortunately precision can only

be achieved when primary data are available to com-

pile more complete and accurate checklists. This

should be done through bringing together records

from many herbaria and from literature, and most

important, through rationally planned specimen col-

lecting designed to cover all biomes adequately.

The conclusions are based on plant distributions

as they are now known, that result from interactions

over a long geological, climatological and evolution-

ary history. It is not apparent to what extent these

distributions have been influenced by present or past

environments. However, listing and comparing the

taxa in each biome is the first step in unravelling the

events that have led to the formation of its character-

istic flora. PRECIS has given us a preliminary look

that will allow the generation of hypotheses for more

rigorous testing using stronger data sets and more

refined techniques.

ACKNOWLEDGEMENTS

The existence of PRECIS is due to the fore-

sightedness and tenacity of Dr B. de Winter, who

stood by the system from its beginning. J. C. Mog-

ford searched PRECIS for the basic species lists, and

B. C. de Wet wrote computer programs to generate

subsidiary lists. R. H. Westfall, C. Hilton-Taylor,

E. J. Moll and R. M. Cowling discussed particular

points. W. Roux provided technical assistance and

B. B. Gibbs assisted with mapping.

REFERENCES

ACOCKS, J. P. H. 1975. Veld types of South Africa, edn 2.

Memoirs of the Botanical Survey of South Africa No. 40.

BAYER, M. B. The Cape Flora and the Karoo. Veld and Flora

70: 17-19.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora

— a descriptive catalogue. Journal of South African Bot-

any Supplement 13.

BRENAN, J. P. M. 1978. Some aspects of the phytogeography of

tropical Africa. Annals of the Missouri Botanical Garden

65: 437-478.

DYER, R. A. 1975. The genera of southern African flowering

plants, Vol. 1. Dicotyledons. Department of Agricultural

Technical Services, Pretoria.

DYER, R. A. 1976. The genera of southern African flowering

plants, Vol. 2. Monocotyledons. Department of Agricul-

tural Technical Services, Pretoria.

GIBBS RUSSELL, G. E. 1975. Comparison of the size of various

African floras. Kirkia 10: 123-140.

GIBBS RUSSELL, G. E. 1985a. PRECIS, the National Her-

barium’s computerized information system. South African

Journal of Science 81: 62-65.

GIBBS RUSSELL, G. E. 1985b. Analysis of the size and compo-

sition of the southern African flora. Bothalia 15: 613-630.

GIBBS RUSSELL, G. E. 1986. Significance of different centres

of diversity in subfamilies of Poaceae in southern Africa.

Palaeoecology of Africa 17: 183-192.

GIBBS RUSSELL, G. E. & GONSALVES, P. 1984. PRECIS —

a curatorial and phytogeographic system. In R. Allkin &

F. A. Bisby, Databases in systematics, the Systematics As-

sociation Special Volume No. 26. Academic Press, Lon-

don.

Bothalia 17,2 (1987)

223

GIBBS RUSSELL, G. E., RETIEF, E. & SMOOK, L. 1984.

Intensity of plant collecting in southern Africa. Bothalia

15: 131-138.

GOLDBLATT, P. 1978. Analysis of the flora of southern Africa:

its characteristics, relationships and origins. Annals of the

Missouri Botanical Garden 65: 369-436.

HUNTLEY, B. J. 1984. Characteristics of southern African bi-

omes. In P. de V. Booysen & N. M. Tainton, Ecological

effects of fire in South African ecosystems. Springer Verlag,

Berlin.

RAUNKIAER, C. 1934. The life forms of plants and statistical

plant geography. Oxford University Press, Oxford.

RUTHERFORD, M. C. & WESTFALL, R. H. 1986. The bi-

omes of southern Africa — an objective categorization.

Memoirs of the Botanical Survey of South Africa No. 54.

SCHEEPERS, J. C. 1982. The status of conservation in South

Africa. Journal of the South African Biological Society 23:

64-71.

SORENSON, T. 1948. A method for establishing groups of equal

magnitude in plant sociology based on similarity of species

content. Kongelige Dansk Videnskabernes Selskab 5: 1-34.

WERGER, M. J. A. 1978. Biogeographical division of southern

Africa. In M. J. A. Werger, Biogeography and ecology of

southern Africa. Junk, The Hague.

WHITE, F. 1983. Vegetation of Africa. UNESCO, Paris.

APPENDIX 1. — Large genera (10 taxa or more) with centre of

diversity in one biome. * = genera reported only from a

single biome. Life forms are abbreviated: P, phanerophyte;

Ch, chamaephyte; H, hemicryptophyte; Cr, cryptophyte;

T, therophyte; S, succulent

a. Large genera with centre of diversity reported for Fynbos only

224

Bothalia 17,2 (1987)

Bothalia 17,2 (1987)

Fabaceae

APPENDIX 2. — Large genera (10 taxa or more) with centres of

diversity in two biomes. Life forms are abbreviated: P, pha-

nerophyte; Ch, chamaephyte; H, hemicryptophyte; Cr,

cryptophyte; T, therophyte; S, succulent

a. Large genera with centre of diversity reported for Savanna and

Grassland

Total no. % of reported

Family and genus of taxa taxa in: Life form

reported Savanna Grassland

Aspleniaceae

225

b. Large genera with centres of diversity reported for Savanna

226

Bothalia 17,2 (1987)

c. Large genera with centres of diversity reported for Savanna

and Fynbos

Total no. % of reported

Family and genus of tax a taxain: Life form

reported Savanna Fynbos

Dicranaceae

d. Large genera with centres of diversity reported for Fynbos

and Grassland

Total no. % of reported

Family and genus of taxa taxa in: Life form

reported Fynbos Grassland

Scrophulariaceae

f. Large genera with centres of diversity reported for Fynbos

and Nama-Karoo

Total no.

Family and genus of taxa

reported

% of reported

taxain: Life form

Fynbos Nama-

Karoo

Chenopodiaceae

g. Large genus with centres of diversity reported for Nama-

Karoo and Succulent Karoo

Total no. % of reported

Family and genus of taxa taxain: Life form

reported Nama- Succulent

Karoo Karoo

Asteraceae

Eriocephalus 19 63 58 Ch

h. Large genus with centres of diversity reported for Nama-

Karoo and Desert

Total no. % of reported

Family and genus of taxa taxain: Life form

reported Nama- Desert

Karoo

Mesembryanthemaceae

Psilocaulon 19 47 32 H, T

APPENDIX 3. — Large genera (10 taxa or more) with centres of

diversity in three biomes. Life forms are abbreviated: P,

phanerophyte; Ch, chamaephyte; H, hemicryptophyte;

Cr, cryptophyte; T, therophyte; S, succulent

a. Large genera with centres of diversity reported for Savanna,

Grassland and Fynbos

Total no. % of reported

Life form

H

P, Ch

Ch, H, T

Bothalia 17,2 (1987)

227

b. Large genera with centres of diversity reported for Savanna,

Nama-Karoo and Desert

Total no. % of reported

Family and of taxa taxain:

genus reported Savan- Nama- Desert 1 e orrn

na Karoo

Poaceae

c. Large genus with centres of diversity reported for Fynbos,

Succulent Karoo and Nama-Karoo

Zygophyllaceae

APPENDIX 4. — Large genera (10 taxa or more) with no apparent

centre of diversity. Life forms are abbreviated: P,phanero-

phyte; Ch, chamaephyte; H,hemicryptophyte; Cr.crypto-

phyte; T, therophyte; S, succulent

d. Large genus with centres of diversity reported for Fynbos,

Savanna and Nama-Karoo

Total no. %of reported

Family and of taxa taxain: Life forr

genus reported Fyn- Savan- Nama-

bos na Karoo

Geraniaceae

Monsonia 10 50 50 50 Ch,H,T,S

Bothalia 17,2: 229-235 (1987)

’ii Kontrolelys van varings en blomplante van die Wonderkloof-

natuurreservaat, Transvaal

J. P. KLUGE* en C. DYER**

Trefwoorde: checklist, Dicotyledoneae, Monocotyledoneae, Pteridophyta, South Africa, Transvaal

UITTREKSEL

'n Sistematiese lys van varings en blomplante word weergegee. Die verhoudinjSs tussen die aantal families,

genusse en spesies van Pteridophyta, Monocotyledoneae en Dicotyledoneae word aangetoon. Families wat meer

as 1% tot die aantal spesies van die gebied bydra, en genusse wat meer as drie spesies bevat, word getabuleer.

ABSTRACT

A checklist of ferns and flowering plants of the W onderkloof Nature Reserve — A systematic list of ferns and

flowering plants of the Wonderkloof Nature Reserve is given. The relationships between the numbers of families,

genera and species of Pteridophyta, Monocotyledoneae and Dicotyledoneae are indicated. Families contributing

more than 1% to the total number of species of the area, and genera containing more than three species are

tabulated.

Die Wonderkloof-natuurreservaat is tussen Nel-

spruit en Lydenburg, ongeveer 50 kilometer van

Nelspruit in 'n noordwestelike rigting gelee en kom

tussen 25° 15'-30' suiderbreedte en 30° 30'-45' oos-

terlengte voor. Die natuurreservaat is op die plaas

Waterval 269 JT gelee, vorm ’n deel van die Uit-

soekstaatsbos en beslaan 523 ha.

Die reservaat is teen die Transvaalse Drakensberg

in die noordoostelike bergsuurveld (Acocks 1975)

gelee en word gedreineer deur ’n ooswaarts vloei-

ende sytak van die Houtboschloop. Die relief in die

reservaat is baie sterk en die hoogte bo seespieel

wissel van 1 000 tot 1 300m bo seespieel.

Die geologiese formasie bestaan uit die Transvaal

Sisteem en word op die reservaat deur die Pretoria

Serie verteenwoordig. Die kranse, wat ook die

boonste grens van die reservaat vorm, bestaan uit

die boonste Nooitgedacht-kwartsiet. Direk onder

die kranse kom die boonste skalie, voor of ’n gang

van diabaas wat deur die reservaat voorkom en wat

vir ’n kort ent direk onder die kranse verskyn. Die

kranse teen die suidfront-helling word deur die on-

derste Nooitgedacht-kwartsiet gevorm. Die laerlig-

gende oostelike deel van die reservaat bestaan uit

skalie met sandsteenlae.

METODE

Planteksemplare is gedurende die periode

1974-1979 deur die senior outeur en mnr D. G. El-

lan-Puttick versamel. Een stel eksemplare is aan die

Nasionale Herbarium in Pretoria gestuur vir identifi-

kasie en benaming terwyl ’n tweede stel in die Her-

* D. R. de Wet Bosbounavorsingstasie, Privaatsak X520, Sabie

1260. Tans: Laeveld Botaniese Tuin, Posbus 1024, Nelspruit

1200.

** D. R. de Wet Bosbounavorsingstasie, Privaatsak X520, Sabie

1260. Tans: Suid-Afrikaanse Navorsingsinstituut vir Bosbou, Pos-

bus 727, Pretoria 0001.

barium van die Departement van Bosbou by die

D. R. de Wet Bosbounavorsingstasie gehuisves

word. H. le Roux se eksemplare is in die Herbarium

van die Laeveldse Botaniese Tuin.

RESULT ATE EN BESPREKING

’n Totaal van 459 spesies is opgeteken. Die aantal

families, genusse en spesies van die Pteridophyta,

Monocotyledoneae en die Dicotyledoneae word in

Tabel 1 aangegee.

Daar is 24 families, dit is 24% van die totale aantal

families, waarvan die spesies meer as 1% van die

totale aantal spesies verteenwoordig. In Tabel 2

word die families volgens grootte gelys. Die aantal

genusse teenwoordig in die families word ook aange-

dui.

Die 17 genusse wat meer as 3 spesies behels, word

in Tabel 3 gelys.

KONTROLELYS

Die versamelnommer volg elke takson. Versamel-

nommers van die senior outeur is met ’n K gemerk

terwyl die van D. G. Ellan-Puttick met ’n D gemerk

is.

In die kontrolelys is die Pteridophyta gerangskik

volgens Anthony & Schelpe (1985) en die Angio-

spermae volgens Dyer (1975, 1976). Die spesies is

alfabeties gerangskik binne elke genus.

VERWYSINGS

ACOCKS, J. P. H. 1975. Veld Types of South Africa. Memoirs of

the Botanical Survey of South Africa No. 40.

ANTHONY, NICOLA C. & SCHELPE, E. A. C. L. E. 1985. A

checklist of Pteridophytes of the ‘Flora of southern Africa’

region. Bothalia 15: 541-544.

DYER, R. A. 1975. The genera of southern African flowering

plants. Vol. 1. Department of Agricultural Technical Ser-

vices, Pretoria.

DYER, R. A. 1976. The genera of southern African flowering

plants. Vol. 2. Department of Agricultural Technical Ser-

vices, Pretoria.

230

Bothalia 17,2 (1987)

TABEL 1. — Aantal families, genusse en spesies van Pteridophyta, Monocotyledoneae en Dicotyledoneae van die Wonderkloof-

natuuneservaat

TABEL 3. — ’n Oorsig van die genusse met meer as drie spesies

PTERIDOPHYTA

SELAGINELLACEAE

Selaginella dregei (Presl) Hieron.

OPHIOGLOSSACEAE

Ophioglossum reticulatum L., K 2351

OSMUNDACEAE

Osmunda regalis L., D 25

SCHIZAEACEAE

Anemia dregeana Kunze, K 1275

GLEICHENIACEAE

Dicranopteris linearis ( Bunn . f.) Underw., K 1146

HYMENOPHYLLACEAE

Hymenophyllum capense Schrad., K 1219

CYATHEACEAE

Cyathea dregei Kunze, K 2022

DENNSTAEDTIACEAE

Pteridium aquilinum (L.) Kuhn, K 1657

ADIANTACEAE

Adiantum poiretii Wikstr. var. poiretii, K 1318

Cheilanthes

concolor (Langsd. & Fisch.) Schelpe & N.C. Anthony, K 1019,

1416

multifida Swartz, K 1210

Pellaea

calomelanos (Swartz) Link, K 1611

pectiniformis Bak., K 2010

viridis (Forssk.) Prantl, K 1608

POLYPODIACEAE

Polypodium polypodioides (L.) Hitchc. subsp. ecklonii (Kunze)

Schelpe, K 1633

Pleopeltis macrocarpa (Willd.) Kaulf, K 1216

DAVALLIACEAE

Arthropteris monocarpa ( Cordem .) C. Chr., K 1320

Oleandra distenta Kunze, K 1018

Bothalia 17,2 (1987)

231

aspleniaceae

Asplenium splendens Kunze, D 55

THELYPTERIDACEAE

Thelypteris gueinziana (Mett.) Schelpe, K 121 7, 1632

BLECHNACEAE

Blechnum giganteum (Kaulf.) Schlechtd., K 1218

GYMNOSPERMAE

ZAMIACEAE

Encephalartos humilis Verdoom *

ANGIOSPERMAE — MONOCOTYLEDONEAE

POACEAE

Imperata cylindrica (L.) Raeuschel, D 88

Eulalia villosa (Thunb.) Nees, D 278, K 1665

Schizachyrium sanguineum (Retz.) Alst., K 1670

Hyparrhenia

filipendula (Hochst.) Stapf \a.i. filipendula, K 1153, 1662

variabilis Stapf, K 1154

Monocymbium ceresiiforme (Nees) Stapf, K 1 739

Trachypogon spicatus (L. f.) Kuntze, D 282

Themeda triandra Forssk., D 281

Digitaiia diagonalis (Nees) Stapf, K 1224

Brachiaria bovonei (Chiov.) Robyns, K 1410

Paspalum scrobiculatum L., K 1411

Oplismenus hirtellus (L.) Beauv., K 1215

Panicum

dregeanum Nees, D 314

sp. cf. fulgens Stapf, D 284

Panicum natalense Hochst., K 1 738

Setaria

megaphylla (Steud.) Dur. & Schinz, K 1211

sphacelata (Schumach.) Moss, K 1671

Rhynchelytrum repens (Willd.) C.E. Hubb., D 286

Trichopteryx dregeana Nees, K 1266

Loudetia

densispica (Rendle) C.E. Hubb., D 280

simplex (Nees) C.E. Hubb., D 283

Phragmites australis (Cav.) Steud., K 2118

Aristida junciformis Trin. & Rupr. subsp. junciformis, D 277

Sporobolus

centrifugus (Trin.) Nees, K 1 734

stapfianus Gand., K 1412

Eragrostis

capensis (Thunb.) Trin., K 1413

sp. cf. racemosa (Thunb.) Steud., D 285

superba Peyr., D 316

Ctenium concinnum Nees, D 279

CYPERACEAE

Cyperus

obtusiflorus Vahl var. flavissimus Boeck., D 122

rupestris Kunth, D 232

Pycreus pelophilus (Ridley) C.B. Cl., D 5

Mariscus

dregeanus Kunth, K 1628

uitenhagensis Steud., D 172, K 1406

Kyllinga

alba Nees, D 58

melanosperma Nees, K 1610

Schoenoplectus corymbosus (Roth, ex Roem. & Schult.) J. Raynal,

D 148

Fimbristylis bisum bellata (Forssk.) Bub., D 41

Bulbostylis burcheUii (Fic.&Hiem) C.B. Cl., D 256, K 1656, 1 740

Coleochloa setifera (Ridley) Gilly, D 312

Carex spicato-paniculata C.B. Cl., K 1616

XYR1DACEAE

Xyris

capensis Thunb., D 3

rehmannii Nilss., K 1151

COMMELINACEAE

Commelina

africana L., D 175, K 1155

erecta L., D 176

Aneilema hockii De Wild., K 1 751

Cyanotis

foecunda Hassk., D 304

speciosa (L.f.) Hassk., D 194

JUNCACEAE

Juncus lomatophyllus Spreng., D 147

L1L1ACEAE

Androcymbium melanthioides Willd. forma subulatum Bak.,

K 166 7

Anthericum

angulicaule Bak., D 66

galpinii Bak., D 257, K 1653

Eriospermum luteo-rubrum Bak., D 306

Aloe

arborescens Mill. *

petricola Pole Evans*

Agapanthus inapertus Beauv., K 1626

Tulbaghia acutiloba Harv., K 1404

Dipcadi gracillimum Bak., K 1405

Ledebouria

revoluta (L.f.) Jessop, D 189

sp., D 309

Protasparagus

aethiopicus (L.) Oberm., K 1305

laricinus (Burch.) Oberm., K. 1396

Smilax kraussiana Meisn., D 241

AMARYLLIDACEAE

Clivia caulescens R.A. Dyer, D 92

Nerine rehmannii (Bak.) L. Bol., D 267, 301

HYPOXIDACEAE

Hypoxis

filiformis/Jafc., K 1402

multiceps Buchinger, D 166

rigidula Bak. D 49, K 141 7

VELLOZIACEAE

Xerophyta

equisetoides Bak., D 65

viscosa Bak., K 1418

DIOSCOREACEAE

Dioscorea

cotinifolia Kunth, D 236

rupicola Kunth, K 1 743

IR1DACEAE

Moraea spathulata (L. f.) Klatt, D 244

Dietes iridioides (L.) Sweet ex Klatt, D 96

Aristea sp. cf. woodii ./V.E. Br., D 269

Crocosmia aurea Planch., D 325

Gladiolus

crassifolius Bak., D 8

elliotii Bak., K 1652

exiguus G.J. Lewis, D 275

longicollis Bak., D 202

vemus Oberm., K 1327

Anomatheca laxa (Thunb.) Goldbl., D 231

ORCHIDACEAE

Holothrix orthoceras (Harv.) Reichb. f, D 9

Habenaria falcicomis (Lined, ex Lindl.) H.Bol. var. caffra (Schltr.)

Renz & Schelpe, H. le Roux

Disperis micrantha Lindl., K 1213

Polystachya

albescens Ridley subsp. imbricata (Rolfe) Summerh., K 1609

cultriformis (Thouars) Spreng., K 1634

Eulophia angolensis (Reichb. f.) Summerh., K 1636

Mystacidium venosum Harv. ex Rolfe, K 1325

232

Bothalia 17,2 (1987)

DICOTYLEDONEAE

P1PERACEAE

Peperomia retusa (L. f.) A. Dietr., K 1236

ULMACEAE

Celtis africana Bunn, f, K 1399

MORACEAE

Ficus

glumosa (Miq.) Del., K 1737

ingens (Miq.) Miq., K 1229

salicifolia Vahl, K 1420

sycomorus L., K 1419

thonningii Blume, D 75, K 1421

URTICACEAE

Laportea peduncularis (Wedd.) Chew, K 1 748

Australina acuminata Wedd., K 174 7

PROTEACEAE

Faurea

saligna Harv. , D 219

speciosa (Welw.) Welw., D 76

Protea gaguedi Gmel., K 1156

SANTALACEAE

Osyridicarpos schimperianus (Hochst. ex A. Rich.) DC., K 1621

Thesium

sp. cf. resedoides/1. W. Hill, D 79

sp. cf. utile A. W. Hill, K 1622

OLACACEAE

Ximenia caffra Sond., K 1613

POLYGONACEAE

Polygonum

meisnerianum Cham. & Schlechtd., K 1663

pulchrum Blume, D 29

salicifolium Willd., D 31

Oxygonum

sp. cf. dregeanum Meisn. var. dregeanum, D 187

dregeanum Meisn. var. strictum (C.H. Wr.) R. Grab., D 128

AMARANTHACEAE

Pupalia lappacea (L.) Juss., K 1273, 1 733

CARYOPHYLLACEAE

Silene burchellii Otth, D 261

RANUNCULACEAE

Knowltonia transvaalensis Szyszyl., D 220

ANNONACEAE

Annona senegalensisPers., D 228

LAURACEAE

Cryptocarya woodii Engl., K 1270

CAPPARACEAE

Cleome monophylla L., D 288

CRASSULACEAE

Kalanchoe

paniculata Harv., K 1635

rotundifolia (Haw.) Haw., D 53

Crassula

alba Forssk. var. alba, D 10, K 1308

globularioides Britt, subsp. argyrophylla (Schonl. & Bak. f.)

Toelken, D 109

obovata Haw. var. obovata, K 1668

schimperi Fischer & C.A. Mey. var. lanceolata (Eckl. & Zeyh.)

Toelken, K 1223

vaginata Eckl. & Zeyh., K 1265

ESCALLONIACEAE

Choristylis rhamnoides Harv., K 28, 1276

ROSACEAE

Rubus sp., D 177

Cliffortia strobilifera Murray, K 1323

Parinari capensis Harv., Dill

FABACEAE

Acacia

ataxacantha£)C., D 215

caffra (Thunb.) Willd., K 23, 1148

karroo Hayne, K 1149

Elephantorrhiza elephantina (Burch.) Skeels, D 149

Bauhinia galpinii N.E. Br., K 1615

Cassia floribunda Cav., K 1605

Lotononis

corymbosa Benth., D 274

eriantha Benth. var. eriantha, D 197

Pearsonia

obovata (Schinz) Polhill, D 21, 260

sessilifolia (Harv.) Duemmer subsp. filifolia (H. Bol.) Polhill,

K 1226

sessilifolia (Harv.) Duemmer subsp. marginata (Schinz) Polhill,

D 11, 22, 112

Crotalaria

capensis Jac q., D 230

recta Steud. ex A. Rich., K 1 744

Argyrolobium

speciosum Eckl. & Zeyh., D 229

tomentosum (Andr.) Druce, D 19

Lotus discolor E. Mey., D 169

Indigofera

adenoides Bak. f., D 36

hilaris Eckl. & Zeyh., D 84

masoniae N.E. Br., K 1629

melanadenia Benth. ex Harv., K 1263

pseudo-indigofera (Merxm.) J.B. Gillett, D 300, K 1654

sanguinea N.E. Br., K 56

swaziensis H. Bol. var. swaziensis, D 208

Tephrosia

cordata Hutch. & Burtt Davy, D 320, K 1259

longipes Meisn. var. lurida (Sond.) J.B. Gillett, D 186, 253

polystachya E. Mey. var. latifolia Harv., K 1221

retusa Burtt Davy, D 254

Aeschynomene micrantha DC., D 247

Stylosanthes fruticosa (Retz.) Alston, D 37

Zornia linearis E. Mey., D 1 74, 252

Desm odium

dregeanum Benth., K 1222

repandum (Vahl) DC., D 16

Pseudarthria hookeri Wight & Am., D 255

Alysicarpus zeyheri Harv., D 246

Dalbergia armata E. Mey., K 2117

Pterocarpus angolensis DC., D 117

Abrus laevigatus E. Mey., D 167, K 1207, 1220

Dumasia viUosa DC. var. villosa, D 327

Erythrina lysistemon Hutch., K 1943

Mucuna coriacea Bak. subsp. irritans (Burtt Davy) Verde., K 1943

Rhynchosia

komatiensis Harms, D 64

monophylla Schltr., D 60, 289

reptabunda N.E. Br., D 168

totta (Thunb.) DC., D 73

Eriosema

angustifolium Burtt Davy, D 191

burkei Benth., D 123, 1 78

ellipticifolium Schinz, K 2114

psoraleoides (Lam.) G. Don, D 222

Flemingia grahamiana Wight & Am., D 93

Vigna angustifoliolata Verde., D 173, K 1408

Sphenostylis angustifolia Sond., D 113

GERANIACEAE

Geranium omithopodium Eckl. <£ Zeyh., D 52

Monsonia attenuata Harv., D 268

Pelargonium

alchemilloides (L.) L ’Herit., D 105

luridum (Andr.) Sweet, D 103

PITTOSPORACEAE

Pittosporum viridiflorum Sims, D 97

OXALIDACEAE

Oxaiis obliquifolia Steud. ex A. Rich., K 1414

Bothalia 17,2 (1987)

233

LINACEAE

Linum thunbergii Eckl. & Zeyh., D 211

RUTACEAE

Zanthoxylum thorncroftii (Verdoom) Waterm., K 1143

Vepris reflexa Verdoom, K 1235

Clausena anisata (Willd.) Hook. f. ex Benth., K 1214

MELIACEAE

Ekebergia pterophylla (C. DC.) Hofmeyr, D 120

POLYGALACEAE

Polygala

albida Schinz, D 321

amatymbica Eckl. & Zeyh., D 115

hottentotta Presl, D 259

uncinata E. Mey. ex Meisn., D 251, 296

EUPHORB1ACEAE

Phyllanthus

maderaspatensis L., D 258

nummulariifolius Poir., K 2116

Bridelia micrantha (Hochst.) Baill., K 2122

Acalypha

glandulifoba Buchinger ex Meisn., D 1 79, 192, 311

wilmsii Pax ex Prain & Hutch., D 106

Tragia rogersii Prain, D 181, K 1397

Jatropha

latifolia Pax var. latifolia, K 16 74

latifolia Rax var. swazica Prain, D 129

Clutia

affinis Sond., K 1617

monticola S. Moore, D 61

pulchella L., D 136, K 1607

Euphorbia

ingens E. Mey. ex Boiss. *

pseudotuberosa Pax, D 104

ANACARDIACEAE

Harpephyllum caffrum Bernh., K 1277

Lannea

discolor (Sond.) Engl., K 1 735

edulis (Sond.) Engl., D 133

Protorhus longifolia (Bemh.) Engl., K 1278

Rhus

chirindensis Bak. f. forma legatii (Schonl.) R. & A. Fernandes,

K 1230

dentata Thunb., D 152

montana Diels var. gerrardii (Han. & Engl.) D. Fernandes,

D 235

pentheri Zahlbr., K 1234, 1304

pyroides Burch., K 1147, 1664

rehmanniana Engl., K 1274

rogersii Schonl., D 170

AQUIFOLIACEAE

Ilex mitis (L.) Radik., K 1630

CELASTRACEAE

Maytenus

acuminata (L. f.) Loes., K 1268

heterophylla (Eckl. <£ Zeyh.) N.K.B. Robson, K 2115

peduncularis (Sond.) Loes., K 1625

undata (Thunb.) Blakelock, K 1231

Pterocelastrus echinatus N.E. Br., D 98, 214

Rhynea phyllicaefolia (DC.) Hilliard & Burtt, D 27

ICACINACEAE

Cassinopsis ilicifolia (Hochst.) Kuntze, D 293

Apodytes dimidiatair. Mey. ex Am. subsp. dimidiata, K 1279

SAPINDACEAE

Pappea capensis Eckl. & Zeyh., K 1228

Hippobromus pauciflorus (L. f.) Radik., K 1227

VITACEAE

Rhoicissus

tomentosa (Lam.) Wild & Drumm., K 1612

tridentata (L.f.) Wild & Drumm., K 2119

Cyphostemma

humile (N.E. Br.) Desc. subsp. dolichopus (C.A. Sm.) Wild &

Drumm., D 240

woodii (Gilg & Brandt) Desc., K 1415

TILIACEAE

Corchorus asplenifolius Burch. , D 182

Grewia occidentalis L., D 156

Triumfetta welwitschii Mast. var. hirsuta (Sprague & Hutch.) Wild,

D 90

MALVACEAE

Sida

dregei Burtt Davy, D 221

hoepfneri Guerke, D 35

Pavonia

burchellii (DC.) R.A. Dyer, K 1233

columella Cav., D 328

Hibiscus

pedunculatus L.f., D 320

schinzii Guerke, D 307

STERCULIACEAE

Dombeya

pulchra N.E. Br., K 22, 1 745

sp. cf. rotundifolia (Hochst.) Planch, var. rotundifolia, D 81

Hermannia

grandifolia N.E. Br., D 33

sp., K 1736

staurostemon K. Schum., D 35A, 212

Waltheria indica L., D 295

OCHNACEAE

Ochna

arborea Burch, ex DC. var. oconnorii (Phill.) Du Toit, D 310

confusa Burtt Davy & Greenway, D 132

natalitia (Meisn.) Walp., D 153

serrulata (Hochst.) Walp., D 134

CLUSIACEAE

Hypericum

aethiopicum Thunb. subsp. sonderi (Bred.) N.K.B. Robson,

D 85

lalandii Choisy, D 266

revolutum Vahl, D 151

FL ACOU RTIACE AE

Dovyalis lucida Sim, K 1212

Rawsonia lucida Harv. & Sond. *

PASSIFLORACEAE

Basananthe sandersonii (Harv.) De Wilde, D 249

Adenia gummifera (Harv.) Harms var. gummifera, K 1 741

Passiflora edulis Sims, K 2120

BEGONIACEAE

Begonia sutherlandii Hook, f, D 317

THYMELAEACEAE

Passerina montana Thoday, D 146

RHIZOPHORACEAE

Cassipourea genardii (Schinz) Alston, K 1659

COMB RET ACE AE

Combretum

kraussii Hochst., K 2013

zeyheri Sond., K 1 745

MYRTACEAE

Psidium guajava L., K 2121

Eugenia natalitia Sond., K 1269

Syzygium cordatum Hochst., D 121

Heteropyxis canescens Oliv., K 1306

BALSAMINACEAE

Impatiens sylvicola Burtt Davy & Greenway, K 1631

RHAMNACEAE

Ziziphus mucronata Willd. Subsp. mucronata, K 1624

Helinus integrifolius (Lam.) Kuntze, D 242

234

Bothalia 17,2 (1987)

MELASTOMATACEAE

Antherotoma naudinii Hook, f., D 40

Dissotis canescens (E. Mey. ex Grah .) Hook, f., D 7

ONAGRACEAE

Epilobium salignum Hausskn., K 1319

Oenothera rosea L ’Herit. ex Ait., D 213

ARALIACEAE

Cussonia spicata Thunb., K 1400

APIACEAE

Centella asiatica (L.) Urb., K 2012

Heteromorpha involucrata Corn., D 319

Annesorrhiza flagellifolia Burtt Davy , D 118

ERICACEAE

Erica drakensbergensis Guth. & Bol., D 108

MYRSINACEAE

Maesa lanceolata Forssk., D 225

SAPOTACEAE

Bequaertiodendron magalismontanum (Sond.) Heine (5 J.H.

Hems!., D 77

EBENACEAE

Euclea crispa (Thunb.) Guerke subsp. crispa, D 91

Diospyros

galpinii (Hiem) De Winter, D 126

whyteana (Hiem) F. White, K 2014

OLEACEAE

Schrebera alata (Hochst.) Welw., K 1 742

Jasminum streptopus E. Mey. var. transvaalensis (S. Moore)

Verdoom, D 233

LOGANIACEAE

Strychnos madagascariensis TW. K 1614

Nuxia congesta R. Br. ex Fresen., D 82

Buddleja salviifolia (L.) Lam., D 80

PERIPLOCACEAE

Cryptolepis oblongifolia (Meisn.) Schltr., D 125, 315

Raphionacme hirsuta (E. Mey.) R.A. Dyer ex Phill., D 119

ASCLEPIADACEAE

Schizoglossum pachyglossum Schltr. var. productum N.E. Br.,

K 1666

Pachycarpus transvaalensis N.E. Br., D 201

Asclepia s

aurea (Schltr.) Schltr., D 195

physocarpa (E. Mey.) Schltr., K 1606

Sarcostemma viminale (L.) R. Br., K 1661

Brachystelma macropetalum (Schltr.) N.E. Br., K 1672

Ceropegia

meyeri Decne., K 1150

racemosa N.E. Br. subsp. setifera (Schltr.) Huber, K 1267

Riocreuxia torulosa Decne. , D 237

Sphaerocodon natalense (Meisn.) Hook, f, D 223

CONVOLVULACEAE

Merremia tridentata (L.)Hall.f. subsp. angustifolia (Jacq.) Ooststi

D 38

Ipomoea

atherstonei Bak., D 162

bolusiana Schinz, D 100

sp. cf. ommaneyi Rendle, D 161

plebeia R. Br. subsp. africana/1. Meeuse, D 4

Turbina oblongata (E. Mey. ex Choisy) A. Meeuse, D 150, 160

BORAGINACEAE

Trichodesma physaloides (Fenzl) A. DC., D 292

Cynoglossum lanceolatum Forssk., D 243

VERBENACEAE

Verbena bonariensis L., D 6

Lantana rugosa Thunb., D 87, 157

Lippia wilmsii Pearson, D 101, 297

Clerodendrum

glabrum E. Mey., K 1232, 1623

triphyllum (Harv.) Pearson, K 201 9

LAMIACEAE

Acrotome hispida Benth., D 114, K 1409

Stachys

grandifolia E. Mey. ex Benth., D 326

natalensis Hochst. var. natalensis, K 1 750

Aeollanthus rehmannii Guerke, D 308

Pycnostachys

reticulata (E. Mey.) Benth., K 1260

urticifolia Hook., D 224

Plectranthus

fruticosus L ’Herit., K 1264

laxiflorus Benth. D 32

rubropunctatus Codd, K 1209

verticillatus (L. f.) Druce, D 17

Iboza riparia (Hochst.) N.E. Br., K 1328

Hemizygia transvaalensis (Schltr.) Ashby, D 47, 83, K 24

Becium obovatum (E. Mey. ex Benth.) N.E. Br., D 71, 116

Orthosiphon senatus Schltr., D 154, K 1619

SOLANACEAE

Physalis peruviana L., D 143

Solanum retroflexum Dun., K 1 749

SCROPHULARIACEAE

Halleria lucida L., D 99

Phygelius aequalis Harv. ex Hiem, D 137

Bowkeria cymosa MacOwan, K 1262

Sutera grandiflora (Galpin) Hiem, D 14

Ilysanthes wihnsii Engl., D 2

Graderia su bin tegra Mast., D 72

Sopubia simplex (Hochst.) Hochst., D 42, 139, 276

Striga

asiatica (L.) Kuntze, D 34, 302

bilabiata (Thunb.) Kuntze, D 216

SELAGINACEAE

Hebenstretia

integrifolia L., D 135

sp., D 107

Selago elata Rolfe, K 1261

Tetraselago natalensis (Rolfe) Junell, D 265

PEDAL1ACEAE

Ceratotheca triloba (Bernh.) Hook, f, D 13

GESNERIACEAE

Streptocarpus cyaneus S. Moore, K 1208

ACANTHACEAE

Thunbergia

atriplicifolia Nees, D 63

natalensis Hook. D 155

neglecta Sond., D 227

Crabbea hirsuta Harv., D 329

Barleria ovata E. Mey. ex Nees, D 263

Sclerochiton harveyanus Nees, D 324

Crossandra greenstockii S. Moore, D 238

Dicliptera clinopodia Nees, D 39

Hypoestes

aristata R. Br., K 131 7

phaylopsoides S. Moore, D 18

Justicia anagalloides T. Anders., D 180, K 1407

PLANTAGINACEAE

Plantago major L., K 1321

RUBIACEAE

Kohautia

lasiocarpa Klotzsch, D 217

virgata (Willd.) Brem., D 313

Oldenlandia herbacea (L.) Roxb., D 1, 44, 305, K 1655

Breonadia salicina (Vahl) Hepper & Wood, K 2021

Cephalanthus natalensis Oliv., D 140

Rothmannia globosa (Hochst.) Keay, D 322

Tricalysia

capensis (Meisn.) Sim, D 95

lanceolata (Sond.) Burtt Davy, K 1658

Bothalia 17,2 (1987)

235

Pentanisia

angustifolia (Hochst.) Hochst., D 205

prunelloides (Klotzsch ex Eckl. £ Zeyh.) Walp., D 62

Vangueria infausta Burch., D 171

Canthium

gueinzii Sond., D 94, 142, K 25

inerme (L. f.) Kuntze, K 1620, 1660

Fadogia tetraquetra Krause, D 299

Pavetta

assimilis Sond. var. assimilis, K 1307

gardeniifolia Hochst. ex A. Rich., K 1627

cooperi Harv. £ Sond., D 165

edentula Sond., D 239

Psychotria capensis (Eckl.) Vatke, D 74

Anthospermum

bispidulum E. Mey. ex Harv. £ Sond., D 294(303, K 1237

rigidum Eckl. £ Zeyh., D 245

Richardia brasiliensis Gomez, D 318

D1PSACACEAE

Cephalaria pungens Szabo, K 1142

Scabiosa columbaria L., D 48

CUCURBITACEAE

Momordica foetida Schum. £ Thonn., D 287

Coccinia

adoensis (Hochst. ex A. Rich.) Cogn., D 226

palmata (Sond.) Cogn., K 1152

CAMPANULACEAE

Wahlenbergia zeyheri Eckl. £ Zeyh., D 246

Lightfootia paniculata Sond., D 78

LOBELIACEAE

Lobelia decipiens Sond., D 138, 262, K 1401

ASTERACEAE

Vernonia

galpinii Klatt, D 248

hirsuta (DC.) Sch. Bip., D 89

natalensis Sch. Bip., D 203

neocorymbosa Hilliard, K 1271

poskeana Vatke £ Hildebr., D 51

stipulacea Klatt, K 2126

sutherlandii Harv., K 29

Stomatanthes africanus (Oliv.£Hiem) R.M.King£H. Robinson,

D 130

Mikania natalensis DC., K 1949

Aster

comptonii Lippert, D 234

lydenburgensis Lippert, D 45, 69

Felicia mossamedensis (Hiem) Mendonga, D 46, 70

Nidorella auriculata DC. subsp. auriculata, D 207

Conyza pinnata (L. f.) Kuntze, D 198

Blumea alata (D. Don) DC., D 291

Helichrysum

aureum (Houtt.) Merr., D 20

cephaloideum DC., D 144, K 1669

kraussii Sch. Bip., K 1951

nudifolium (L.) Less. var. nudifolium, D 210, K 1144

oxyphylium DC., D 86

pilosellum (L. f.) Less., D 59

rugulosum Less., D 206, K 1225

splendidum (Thunb.) Less., D 127

harveyanum Wild, D 273

Athrixia phylicoides DC., D 26

Geigeria burkei Harv. subsp. burkei var. burkei, D 28

Callilepis laureola DC., D 102

Anisopappus smutsii Hutch., K 1326

Acanthospermum australe (Loefl.) Kuntze, D 272

Inezia integrifolia (Klatt) Phill., D 183

Schistostephium crataegifolium (DC.) Fenzl ex Harv.,p 15, 24

Crassocephalum picridifolium (DC.) S. Moore, K 1324

Senecio

bupleuroides DC., D 271

coronatus (Thunb.) Harv., D 131

deltoideus Less., K 1946

inornatus DC., K 1303, 1322

glaberrimus DC., D 218

microglossus DC., D 163, K 16 73

pterophorusDC., D 30, 54

striatifoliusDC., K 1145

tamoides DC., K 26

Euryops pedunculatus N.E. Br., K 1272

Osteospermum jucundum (Phill.) T. Norl., D 68

Berkheya insignis (Harv.) Thell, D 124

Dicoma

anomala Sond. subsp. cirsioides (Harv.) Wild, D 23

zeyheri Sond., D 250

Gerbera

ambigua (Cass.) Sch. Bip., D 57, 60

jamesonii H. Bol. ex Hook. f.,D 67

Bothalia 17,2: 237-256 (1987)

A checklist of vascular plants of the Amatole Mountains, eastern

Cape Province/Ciskei

P. B. PHILLIPSON*

Keywords: Amatole Mountains, checklist, Ciskei, eastern Cape Province, southern Africa, vascular plants, Winterberg

ABSTRACT

A checklist of vascular plants of the Amatole Mountains is presented. The physical environment, climate and

vegetation of the study area and the history of its botanical exploration are described. The mountains form part of

the Winterberg Range in the eastern Cape/Ciskei region of south-eastern Africa, and cover an area of approxi-

mately 900 km2. The altitude ranges from about 700 m to 2 000 m above sea level, and the topography is very

varied. The climate is warm temperate and supports various vegetation types including forest, sclerophyllous

shrubland, grassland and marshland. The checklist records the occurrence of 1 215 taxa. The largest families and

genera in the area contain predominantly grassland herbs. Many of the characteristic families of the Cape Floristic

Region and of the arid areas of southern Africa are poorly represented in the Amatole Mountains.

UITTREKSEL

’n Kontrolelys van vaatplante van die Amatoleberge word verskaf. Die fisiese omgewing, klimaat en plantegroei

van die studiegebied en die geskiedenis van die plantkundige verkenning daarvan, word beskryf. Hierdie berge

vorm deel van die Winterberg-reeks in die Oos-Kaap/Ciskei-gebied van suidoostelike Afrika en beslaan ongeveer

900 km2. Die hoogte bo seespieel strek vanaf ongeveer 700 m tot 2 000 m en die topografie is baie varierend. Die

klimaat is warm gematig en onderhou verskeie plantegroeitipes waaronder woud, sklerofiele struikveld, grasveld

en vleiland. Daar is 1 215 taksons op die kontrolelys aangeteken. Die grootste families en genusse in die gebied

bevat oorwegend grasveldkruide. Baie van die kenmerkende families van die Kaapse Flora-gebied en van die

dorre gebiede van die suidelike Afrika is swak verteenwoordig in die Amatoleberge.

INTRODUCTION

The Amatole Mountains in the south-east of

southern Africa are of particular botanical and eco-

logical interest for a number of reasons. The Moun-

tains lie in a region where six major African phyto-

choria meet. These are the Indian Ocean Coastal

Belt, the Sudano-Zambezian Region, the Karoo-Na-

mib Region, the Cape Region and the Afro-mon-

tane archipelago with its associated Afro-alpine

areas (Werger 1978). The Mountains receive a rela-

tively high rainfall, and act as an important drainage

sponge for the neighbouring lower lying semi-arid

areas. The indigenous forests and marshlands are

believed to be particularly important in this respect.

State forestry plantations and residential areas are

responsible for the introduction of many exotic

species, some of which have become naturalized.

The establishment of forest plantations and the en-

croachment of exotics into natural vegetation have

had a very significant effect on the indigenous flora

and on the ecology of the area as a whole.

The Amatole Mountains have been known to bot-

anists since the early nineteenth century, and have

gradually become relatively thoroughly explored.

They have long been recognized as an area with a

high species diversity, but little precise information

has been available about their flora.

The aim of this work is to provide a reliable and

comprehensive checklist of the vascular plants of the

* Formerly: Department of Botany, University of Fort Hare,

Alice, Ciskei.

Amatole Mountains which can serve as a basic refer-

ence for taxonomic, floristic and ecological research,

and for educational purposes in the future.

STUDY AREA

Physical Environment

The name Amatole (or Amatola) Mountains has

been applied to a vaguely-defined section of the

Winterberg Range, centred on the well known

Hogsback Ridges. In the present study they are de-

fined as bounded by the Kat and Esk Rivers in the

north-west and the Thomas and Keiskamma Rivers

in the east. The bottom edge of the escarpment

forms the southern edge, while an arbitrary line

across the African surface plateau connecting the

Esk and Thomas Rivers forms the northern bound-

ary (Figure 1). Defined in this way the Amatole

Mountains form an area of about 900 km2, lying

within the latitudes 32° and 33° S, and longitudes 26°

and 28° E.

The Amatole Mountains are part of a long outly-

ing spur of the high interior plateau of southern

Africa. The spur extends south-east and then east

from the Great Escarpment, gradually loosing

height and disappearing near the town of Stutter-

heim. It has been breached by the Fish River, which

separates the Amatoles and other mountains of the

Winterberg Range to the east from the Bankberg,

Bruintjeshoogte and Tandjesberg Ranges to the

west.

The Winterberg Range has a steep, irregular es-

carpment facing south, which rises above the ‘Post-

African’ erosion surface. To the north of the escarp-

32°45'S

238

Bothalia 17,2 (1987)

27u00 1 E

Alice

10 km

26 45 ' E

Keiskammahoek

A 20

KEY

M-«- Bottom edge of

escarpment

Study area: Amatol e Mts

a Mountain peaks.

FIGURE 1. — Map of the study area showing the main mountain peaks, rivers and settlements. Amatole Mountains, main peaks:

Elandsberg 1(2 017 m), 2(1 960 m), 3(1 877 m); unnamed, 4(1 800 m); Gaika’s Kop 5(1 963 m); Tor Doone 6(1 565 m);

Menziesberg, 7(1 645 m); Pefferskop, 8(1 086 m); Juanasberg, 9(1 411 m), Tyume Peak 10(1 481 m); Hogsback Peaks,

11(1 836 m), 12(1 826 m), 13(1 937 m); Iron Rock, 14(1 089 m); Gxulu Kop, 15(1 427 m); Mount Macdonald, 16(991 m);

Geju Mountain, 17(1 868 m); Cata Mountain, 18(1 641 m); Mount Thomas 19(1 616 m). Pirie Mountains, main peaks: Mount

Kempt, 20(1 420 m); Pirie Mountain, 21(1 270 m).

ment, the land, which forms part of the older ‘Afri-

can’ erosion surface, slopes gradually into the basins

of the Fish and Kei Rivers. Remnants of the still

older ‘Gondwana’ erosion surface remain as scat-

tered peaks, ridges and mesas above the African

surface plateau.

The escarpment of the Amatole Mountains falls

from the African surface plateau at about 1 500 m to

between 700 and 1 000 m, broken in many places by

an intermediate platform at about 1 250 m. Above

the plateau the highest mountain peaks reach about

2 000 m. The most prominent of the peaks are shown

in Figure 1.

The Winterberg Range consists of dolerite sheets,

dikes and sills intruded into weak shales and sand-

stones of the Karoo sequence (Beaufort Group).

The resistant dolerite has strongly influenced the

landscape, helping to maintain the prominent es-

carpment and the Gondwana remains, and also caus-

ing a number of less important features. The geo-

morphology of the Hogsback area was described in

more detail by Agnew (1958).

Climate

The climate of the Amatole Moutains is warm

temperate, characterized by high rainfall, cold win-

ters and moderately warm summers relative to other

parts of southern Africa.

Moist air from the south is forced upwards by the

escarpment and results in rainfall of between 750

and 1 500 mm per annum, increasing from lower to

higher altitudes (Story 1952). Behind the escarp-

ment and on north-facing slopes the rain shadow

decreases the rainfall, while local topographic effects

may increase it in certain places. Rainfall is more

evenly spread throughout the year than is common

in many parts of southern Africa, although it is high-

est in summer, with a peak in February. On the es-

carpment and south-facing mountain slopes fog is

common, and mist gauges have recorded 20 - 30%

gains over the standard precipitation (S. Russell

pers. comm.).

At the Hogsback settlement (about 1 250 m) the

annual average temperature is about 15 °C, with ab-

solute maxima and minima of 40° in summer and -6°

32°45 1 S 32°45 ' S

Bothalia 17,2 (1987)

239

in winter respectively, with some snow falling in

most years. Obviously altitude and topography influ-

ence temperatures, and very much colder conditions

are found at the summits of the main peaks.

Vegetation

The vegetation of the Amatole Mountains is pre-

dominantly Afro-montane in affinity (White 1978,

1983), and many typical Afro-montane species reach

their southern limit in this region. In addition the

Mountains contain a significant Cape element in

their flora, together with many SE African endem-

ics. They fall within an area of Highland and Dohne

Sourveld in the vegetation classification of Acocks

(1975).

The high rainfall in the Amatole Mountains is able

to support well developed high forest. These forests

are floristically rich, containing both evergreen and

deciduous species, with large specimens of Podocar-

pus falcatus being particularly prominent. Well pre-

served forest is present on most of the escarpment

slopes, and in some areas of the intermediate plat-

form. On the mountain peaks, above the plateau,

some forest/woodland patches are found in sheltered

areas, mainly in south-facing positions. These

patches are poor in species and are probably above

the altitudinal limit for many forest tree species in

this region.

The plateau areas and mountain slopes are largely

grass-covered. This ‘sour’ grassland has provided

good grazing land for domestic stock, and this has

given rise to the Mountains’ name. Amatole (or

Amatola) is derived from the Xhosa word amathole,

the plural of ithole, meaning calves. Oral history de-

scribes the large herds of cattle owned by pastoralists

in the days of Paramount Chief Sandile. These herds

thrived on the rich grassland and produced many

calves, and the area eventually became known as the

‘mountains of calves’ (iintabe zakwamathole) (Pahl

pers. comm.). Excessive grazing by domestic stock

can severely reduce grass cover, and this may lead to

soil erosion. Serious erosion has occurred in some

localities, particularly near the base of the escarp-

ment. At higher altitudes bare ground does not ap-

pear prone to excessive erosion, and is usually colo-

nized by unpalatable herbs and small shrubs which

help to stabilize the soil surface. Helichrysum argy-

rophyllum, in particular, covers large areas that have

been overgrazed, and once it is established, re-

growth of grasses appears to be very slow. Grassland

in the Amatole Mountains contains a large number

of herbs which are highly responsive to fire, being

inconspicuous in moribund grassland, but flowering

prolifically after burning.

In many places the vegetation is dominated by

sclerophyllous shrubs growing up to about 3 m tall.

Such areas are relatively poor in species, with Clif-

fortia spp., Erica brownleeae, Passerina spp. and

Stoebe spp. predominating. These communities are

often referred to as ‘false-macchia’ to distinguish

them from the true macchia (fynbos) of the Cape

region and can colonize grassland areas very rapidly.

It is not certain whether they represent a climax

stage, or merely a step in the succession from grass-

land to forest. In many localities old stands of false-

macchia may contain forest pioneer species such as

Buddleja salvifolia, Halleria lucida and Rapanea

melanophloeos, and probably in the more sheltered

localities the succession could procede to forest,

while in the more exposed areas a mixed false-mac-

chia/woodland may result. The false-macchia com-

munities are very sensitive to fire, and controlled

burning programmes have been employed success-

fully in eliminating sclerophyllous shrubs and pro-

moting the re-establishment of pasture grasses.

Overgrazing pratices may encourage the encroach-

ment of false-macchia. The agricultural management

of vegetation in the Amatole Mountains has been

discussed by Trollope (1973).

The summits of the highest peaks reach into the

Sub-alpine Belt of the Afro-alpine Region (Killick

1978), where a montane moorland vegetation oc-

curs. On rocky areas this consists of small sclero-

phyllous shrubs, growing to a height of about 0,5 m,

often with scattered small trees of Protea subvestita.

These shrubs may include Arrowsmithia styphe-

lioides, Chrysocoma tenuifolia, Cliffortia paucistami-

nea, Erica spp., Euryops dyeri, Muraltia spp. and

Passerina montana, with Restio sejunctus and Tham-

nocalamus tesselata also common. Elsewhere on the

summits grassland may occur with Agrostis spp.,

Aristida junciformis subsp. galpinii and Festuca spp.

predominating. Little is known about the ecology of

the summit vegetation.

Depressions in level areas may support patches of

marshland. These are usually dominated by species

of Cyperaceae, although the invasion of some mar-

shes near Gaika’s Kop by Phragmites australis has

been noted.

Man has certainly had a significant influence on

the vegetation of the Amatole Mountains over a

long period of time. Since about 1850 forestry activi-

ties and the development of the Hogsback residen-

tial/recreational area have caused the introduction of

a large number of exotic species. A number of eco-

nomically important forest trees have been planted

in the region, and some of these have become natu-

ralized. The Hogsback settlement is well known as a

place where temperate garden plants will flourish,

and some of these have also become naturalized.

Most of the naturalized exotics in the Amatole

Mountains are confined to disturbed ground near

plantations and cultivated ground, and most have

failed to encroach significantly on the natural vege-

tation. Important exceptions are Acacia mearnsii,

Pinus spp., Rubus fruticosus and certain grasses, no-

tably Stipa trichotoma , which have become wide-

spread.

HISTORY OF BOTANICAL EXPLORATION IN THE

AMATOLE MOUNTAINS

The earliest known preserved botanical specimens

from the Amatole Mountains are those collected by

C. F. Ecklon and C. L. P. Zeyher in 1831/2. Their

contemporary, J. F. Drege, and earlier collectors

such as W. J. Burchell and C. P. Thunberg collected

in neighbouring areas, but did not actually visit the

Amatole Mountains (Gunn & Codd 1981). Ecklon

and Zeyher collected on ‘Schumiberg’, the mountain

240

Bothalia 17,2 (1987)

now known as Juanasberg. While collecting in the

area they probably stayed at Tyumie Mission, which

had been founded by Rev. J. Brownlee in 1820.

Brownlee was himself a keen amateur botanist, but

he only started preserving plant specimens after

moving to King William’s Town in 1825 (Gunn &

Codd 1981). During this early period the Amatole

Mountains must have been very inaccessible, and

further botanical exploration was probably pre-

vented by the series of frontier wars which continued

until 1847. In 1860 T. Cooper visited the area, and

collected extensively, particularly on Elandsberg.

Soon a number of forest stations were set up along

the Winterberg, and gradually the mountains be-

came more accessible (Sim 1907). As botanical ex-

ploration of southern Africa proceeded, many im-

portant collectors of the late nineteenth century visi-

ted parts of the Winterberg, but little material was

collected in the Amatole Mountains. The Pirie

Mountains in the east, and the Boschberg to the west

of the Winterberg were extensively explored by T.

R. Sim and P. Macowan respectively. J. Buchanan

collected grasses and ferns in the Amatole Moun-

tains while he was stationed at Lovedale Mission

from 1876 to 1877, and Sim and W. G. Bennie also

made some collections in the area (Sim 1915; Gunn

& Codd 1981). However, in general the flora of the

area remained poorly known.

In the early part of the twentieth century G. Rat-

tray made extensive collections in the Amatole

Mountains, mainly at Hogsback, and some other im-

portant collectors visited the area at this time. In

1934 systematic botanical exploration of the Moun-

tains commenced with the appointment of M. H.

Giffen as the lecturer in Botany at the nearby South

African Native College (now the University of Fort

Hare). Giffen collected extensively during the 1930’s

and 1940’s but unfortunately did not distribute du-

plicate specimens. His main interest was in diatoms

and his collections of other plant groups did not re-

ceive the attention they deserved (M.H. Giffen pers.

comm.).

In 1947 R. Story commenced a botanical survey of

the Keiskammahoek District, which includes a por-

tion of the Amatole Mountains, but also includes

part of the Pirie Mountains and some lower lying

areas (Story 1952). This work is important, not only

with respect to the specimens collected, but also in

providing a basic ecological account of the District.

Since the mid-1950’s regular student field excur-

sions to the Hogsback area have been organized by

A. R. A. Noel, A. Jacot Guillarmod and R. A.

Lubke of Rhodes University, Grahamstown, which

formed the basis of an unpublished checklist. Speci-

mens collected on these visits are housed at RUH,

but have not been critically identified. Since 1975

staff and students at the University of Fort Hare

have continued collecting in the Amatole Moun-

tains, and a number of ecological research projects

have been carried out. Field work undertaken for

the present study was concentrated on the more

poorly collected localities, habitats and taxa, in or-

der to give a more thorough coverage of the flora of

the area. The Giffen collections were finally identi-

fied and labelled and his duplicate specimens and

other collections were distributed by G. E. Gibbs

Russell. By July 1986 nearly 3000 specimens of vas-

cular plants from the Amatole Mountains had accu-

mulated in the Herbarium of the Department of

Plant Sciences at the University (UFH), and these

form the basis of the present checklist.

The Amatole Mountains have been visited by

many botanists from other institutions during the

past 20 to 30 years, and they have become botan-

ically well explored in comparison with many areas

of southern Africa.

CHECKLIST

The present checklist of vascular plants was com-

piled from a number of sources. The specimens at

UFH have been examined and identified using the

relevant Floras and monographs. In problematic

cases comparison of material with authentically de-

termined specimens in other herbaria has been

made. In many cases, specimens of taxa currently

under revision have been seen by or discussed with

the taxonomists concerned. Specimens at UFH thus

form the basis of the checklist. Duplicate material

has been distributed to many other herbaria, but the

main duplicate sets are held at K, MO and PRE.

Records of additional taxa have been obtained

directly from taxonomic literature where a locality

within the study area has been specifically men-

tioned. An extensive search of the available litera-

ture was made for such records, and these have been

cited in full in the checklist.

No attempt has been made to systematically

search for material from the Amatole Mountains in

other herbaria, however a comparison was made

with the computerized record (PRECIS) of the col-

lections at PRE. Duplicates of the historically im-

portant Cooper, Sim and Rattray collections, to-

gether with many recent collections, are listed and

these were checked against the present checklist. In

a sample section of about a sixth of the PRECIS

record no additional taxa were found.

In the course of field work for the present study a

few sight records of additional taxa were made, and

these are given in the checklist.

Nomenclature follows the list of accepted taxa at

PRE (Gibbs Russell et al. 1984, 1985), except where

more recent treatments are available. Every effort

has been made to ensure that the checklist was taxo-

nomically and nomenclaturally up-to-date on com-

pletion in December 1986.

DISCUSSION

The checklist includes 1 215 taxa, 65 pterido-

phytes, 4 gymnosperms, 328 monocots and 818 di-

cots. The largest families (with over 50 taxa) are the

Asteraceae (208 taxa, 17%), Poaceae (83 taxa, 7%),

Fabaceae (67 taxa, 6%), Cyperaceae (62 taxa, 5%),

Liliaceae (54 taxa, 4%) and Orchidaceae (53

taxa,4%). The two largest genera are Senecio (50

taxa, 4%) and Helichrysum (49 taxa, 4%), both

members of the Asteraceae. No other genus has

Bothalia 17,2 (1987)

241

more than 14 recorded taxa, but the following have

10 or more: Argyrolobium, Asplenium, Cheilanthes,

Crassula, Disa, Erica, Geranium, Hypoxis, Indigo-

fera, Pelargonium, Rhus, Stachys and Wahlenbergia.

Most of the largest families and genera are groups

containing predominantly grassland herbs, notably

the Asteraceae and the Poaceae, and this reflects the

high diversity of species found in this habitat in the

Amatole Mountains. There is also a large number of

species of Cyperaceae, a family characteristic of

marshland habitats. Taxa typical of forest habitats

are only represented, in these figures, by the rela-

tively small genera Asplenium and Cheilanthes (both

Pteridophytes), and the sclerophyllous vegetation

only by the genus Erica.

Comparison of the figures given above with those

given by Goldblatt (1978) for the whole of southern

Africa show some similarities in the relative propor-

tions of many taxa. The most noticeable differences

in the flora of the Amatole Mountains is the low

number of species of Ericaceae, Proteaceae and

Restionaceae, characteristic families of the Cape

Floral Region, and of Mesembryanthemaceae (in-

cluded in Aizoaceae by Goldblatt), which are con-

centrated in arid areas.

Some specimens belonging to genera which are

currently in a state of taxonomic confusion or cur-

rently under revision could not be identified, these

include Alchemilla, Erica, Harveya, Hypoxis, Indi-

gofera, Lotononis, Pentaschistis, Senecio and Wah-

lenbergia. Other genera contain probable new

species, these include Cliffortia, Cineraria, Conium,

Crassula, Fuirena, Helichrysum, Passerina, Pentzia,

Stoebe and Watsonia. Some naturalized exotics re-

corded in the Amatole Mountains, but lacking

voucher specimens, were not identified to species

level, and the record of Othonna sp. is based on an

unsubstantiated literature reference. In some cases,

specimens of species divided into varieties or subspe-

cies could not be assigned to these taxa with cer-

tainty.

ACKNOWLEDGEMENTS

I am very grateful to many people who have con-

tributed in some way to the completion of this study.

I wish to thank the students, colleagues and friends

who have accompanied me on field trips, especially

Liz Brown and Ftilde van Vlaenderen, the taxonom-

ists who have discussed particular taxa in the study

area and who have helped with identifications, espe-

cially O. M. Hilliard and B. L. Burtt, the directors

and curators of E, GRA, K, KEI, PRE, RUH and

UFH who have allowed me to consult their collec-

tions and all those who have made botanical collec-

tions in the Amatole Mountains.

REFERENCES

ACOCKS, J. P. H. 1975. Veld types of South Africa, 2nd edn.

Memoirs of the Botanical Survey of South Africa No. 40.

AGNEW, S. 1958. The landforms of the Hogsback area in the

Amatole Range. Fort Hare Papers 2: 1-14.

ANTHONY, N. C. 1984. A revision of the southern African

species of Cheilanthes Swartz and Pellaea Link (Pterida-

ceae). Contributions of the Bolus Herbarium 11.

ANTHONY, N. C. & SCHELPE, E. A. C. L. E. 1985. A check-

list of pteridophytes of the Flora of southern Africa region.

Bothalia 15: 541-544.

ARCHIBALD, E. E. A. 1967. The genus Dioscorea in the Cape

Province west of East London. Journal of South African

Botany 33: 1-46.

CODD, L. E. 1975. Plectranthus (Labiatae) and allied genera in

southern Africa. Bothalia 11: 371-442.

CODD, L. E., DE WINTER, B. & RYCROFT, H. B. (eds).

1963. Flora of southern Africa. Government Printer, Pre-

toria.

DE DALLA TORRE, C. G. & HARMS, H. 1963. Genera

siphonogamarum. Wissenschaftliche Neudrucke, Wiesba-

den.

DE VOS, N. P. 1972. The genus Romulea in South Africa. Jour-

nal of South African Botany. Supplementary volume No.

9: 1-307.

GIBBS RUSSELL, G. E. & THE STAFF OF THE NATIONAL

HERBARIUM, 1984. List of species of southern African

plants. Memoirs of the Botanical Survey of South Africa

No. 48.

GOLDBLATT, P. 1978. An analysis of the flora of southern

Africa: its characteristics, relationships, and origins. An-

nals of the Missouri Botanical Garden 65: 369-426.

GIBBS RUSSELL, G. E., REID, C., VAN ROOY, J. &

SMOOK, L. 1985. List of species of southern African

plants, edn 2, part 1. Memoirs of the Botanical Survey of

South Africa No. 51.

GUNN, M. & CODD, L. E. 1981. Botanical exploration of south-

ern Africa. Balkema, Cape Town.

HALL, A. V. 1965. Studies of the South African species of Eulo-

phia. Journal of South African Botany. Supplementary vol-

ume No. 5: 1-248.

HALL, A. V. 1982. A revision of the southern African species of

Satyrium. Contributions from the Bolus Herbarium 10:

1-137.

HEDGE, I. C. 1974. A revision of Salvia in Africa. Notes from

the Royal Botanical Garden Edinburgh 33: 1-121.

HILLIARD, O. M. 1977. Compositae in Natal. University of

Natal Press, Pietermaritzburg.

HILLIARD, O. M. 1983. Tribe Inuleae, subtribe Gnaphaliinae.

In O. A. Leistner, Flora of southern Africa 33, part 7,

fascicle 2 (first part): 1-325.

HILLIARD, O. M. & BURTT, B. L. 1984. A revision of Diascia

section Racemosae. Jounal of South African Botany 50:

269-340.

HILLIARD, O. M. & BURTT, B. L. 1985. Conium (Umbellife-

rae) in southern Africa. South African Journal of Botany

51: 465-474.

HILLIARD, O. M. & BURTT, B. L. 1986. Notes on some plants

of southern Africa chiefly from Natal: XII. Notes from the

Royal Botanical Garden Edinburgh 43: 189-228.

JACOBSEN, W. B. G. 1978. Some problems of South African

Pteridophyta. Journal of South African Botany 44: 157—

185.

JACOBSEN, W. B. G. 1983. The ferns and fern allies of southern

Africa. Butterworths, Durban, Pretoria.

KILLICK, D. J. B. 1969. The South African species of Myrica.

Bothalia 10: 5-17.

KILLICK, D. J. B. 1978. The Afro-alpine Region. In M. J. A.

Werger, Biogeography and ecology of southern Africa 1:

515-560. Junk, The Hague.

KUKKONEN, I. 1983. The genus Schoenoxiphium ( Cyperaceae ).

A preliminary account. Bothalia 14: 819-823.

KUPICHA, F. K. 1984. Studies on African Asclepiadaceae. Kew

Bulletin 38: 599-672.

LANGENEGGER, J. E. 1976. Dovyalis. In J. H. Ross, Flora of

southern Africa 22: 84-90.

LINDER, H. P. 1981a. Taxonomic studies in Disinae. III. A re-

vision of Disa Berg, excluding sect. Micranthae Lindl.

Contributions from the Bolus Herbarium 9: 1-370.

LINDER, H. P. 1981b. Taxonomic studies in the Disinae. VI. A

revision of the genus Herschelia. Bothalia 13: 365-388.

LINDER, H. P. 1985. Conspectus of African species of Restiona-

ceae. Bothalia 15: 387-503.

MARAIS, W. 1970. Cruciferae. In L. E. Codd, B. De Winter,

D. J. B. Killick & H. B. Rycroft. Flora of southern Africa 13.

NORDENSTAM, B. 1968. The genus Euryops. Part 1. Taxo-

nomy. Opera Botanica No. 20.

OBERMEYER, A. A. 1973. A note on Laurembergia repens.

Bothalia 11: 116-117.

242

Bothalia 17,2 (1987)

OBERMEYER, A. A. 1978. Omithogalum : a revision of the

southern African species. Bothalia 12: 323-376.

PUFF, C. 1978. The genus Galium L. (Rubiaceae) in southern

Africa. Journal of South African Botany 44: 203-279.

REID, C. & DYER, R. A. 1984. A review of the southern African

species of Cyrtanthus. American Plant Life Society, Cali-

fornia.

ROESSLER, H. 1959. Revision der Arctotidae — Gorteriinae

(Compositae). Mitteilungen aus der Botanischen Staats-

sammlung, Munchen 3: 71-500.

SIM, T. R. 1907. The forests and forest flora of the colony of the

Cape of Good Hope. Taylor & Henderson, Aberdeen.

SIM, T. R. 1915. The ferns of South Africa. 2nd edn. University

Press, Cambridge.

SNIJMAN, D. 1984. A revision of the genus Haemanthus L.

(Amaryllidaceae). Journal of South African Botany Sup-

plementary volume 12: 1-139.

STIRTON, C . H. 1984. Notes on the genus Rubus in southern

Africa. Bothalia 15: 101-106.

STORY, R. 1952. A botanical survey of the Keiskammahoek

District. Memoirs of the Botanical Survey of South Africa

No. 30.

TOLKEN, H. R. 1977. A revision of the genus Crassula in south-

ern Africa. Contributions from the Bolus Herbarium 8:

1-595.

TROLLOPE, W. S. W. 1973. Fire as a method of controlling

Macchia (Fynbos) vegetation on the Amatole Mountains

of the eastern Cape. Proceedings of the Grassland Society

of southern Africa 8: 35-41.

VAN DER WALT, J. J. A. 1973. The South African species of

Commiphora. Bothalia 11: 53-102.

WEIMARCK, H. 1949. A revision of the genus Alepidea. Botan-

iska Notiser 1949, 4: 217-268.

WERGER, M. J. A. 1978. Biogeographical division of southern

Africa. In M. J. A. Werger, Biogeography and ecology of

southern Africa 1: 147-170. Junk, The Hague.

WHITE, F. 1978. The Afro-montane Region. In M. J. A.

Werger, Biogeography and ecology of southern Africa 1:

463-513. Junk, The Hague.

WHITE, F. 1983. Vegetation of Africa. UNESCO, Paris.

CHECKLIST

The checklist is divided into four sections: section 1 deals with the Pteridophyta, each genus prefixed with ‘P’ and numbered

according to the sequence in Anthony & Schelpe (1985); sections 2, 3 and 4, which deal with the Gymnospermae, Angiospermae-

Monocotyledoneae and Angiospermae-Dicotyledoneae respectively, are numbered according to the system used in the Flora of

southern Africa (Dyer 1975, 1976). This is based on that of De Dalle Torre & Harms (1963), except the Poaceae which axe number-

ed according to the Kew system, each genus prefixed with ‘K\ In the checklist the voucher specimens cited are specimens at UFH,

unless otherwise stated or unless a literature citation is given. The following abbreviations for collectors’s names are used: Br =E.D.

Brown, Fu =H.D. Furness, GR =G.E. Gibbs Russell, Gi = M.H.Giffen, Hu = A. Hutchings, Ph = P.B. Phillipson and Tu = M.Tusenius.

PTERIDOPHYTA

LYCOPODIACEAE

P2 Lycopodium

clavatum L. Fu & Ph 146; GR 3481; Gi 36, 235, 582; Ph 408;

Rayment s.n.

gnidioidesZ,. f. Gi 236, 1050, 1569, s.n.; Grierson s.n.; Ph 593.

saururus Lam. Fu & Ph 42; Ph & Hu 120.

verticillatum L. f. Gi 23 1, s.n.; Rayment s.n.

SELAGINELLACEAE

P3 Selaginella

caffrorum (MildeJ Hieron. Gi 124; Ph 830.

kraussiana (Kunze) A. Br. ex Kuhn Bryant s.n.; Gi 807; Mgu-

dlwa 29.

OPHIOGLOSSACEAE

P6 Ophioglossum polyphyllum A. Br. Gi 1502.

GLEICHENIACEAE

P10 Gleichenia polypodioides (L.) J.E. Sm. Gi s.n.; Ph 343.

SCHIZAEACEAE

P12 Schizaea pectinata (L.) Swartz Rayment s.n.

P14 Mohria caffrorum (L.) Desv. Fu & Ph 214; Gi 805, s.n.

CYATHEACEAE

P19 Cyathea capensis (L. f.) J.E. Sm. Gi 604, 779, s.n.

HYMENOPHYLLACEAE

P20 Trichomanes pyxidiferum L. var. melanotrichum (Schlechtd.)

Schelpe GR 3828; Gi 774.

DENNSTAEDTIACEAE

P23 Histiopteris incisa (Thunb.) J. Sm. Ph 1505.

P24 Pteridium aquilinum (L.) Kuhn subsp. aquilinum Fu & Ph

223; GR 3484; Gi s.n.

P26 Hypolepis sparsisora (Schrad.) Kuhn Gi s.n.

ADIANTACEAE/PTERIDACEAE

P32 Adiantum

capillus-veneris L. Gi 590; Ph 877.

poiretii Wikstr. var. poiretii Gi 554, s.n.

P33 Pteris

cretica L. Gi 553, 798, 799, 1472.

dentata Forssk. GR 3810; Gi s.n.

P34 Cheilanthes

bergiana Schlechtd. Gi 552, s.n.; Ph 115.

capensis (Thunb.) Swartz, Chumie Forest, Young sub TRV 190

(PRE), Tor Doone, Giffen 1278a (PRE) (Anthony 1984:

102).

concolor (Langsd. & Fisch.) R. & A. Tryon Gi 3.

eckloniana (Kunze) Mett. Gi 796, 845; Ph 1040.

hirta Swartz GR 3042; Gi 1457, s.n.; Ph 832.

multifida (Swartz) Swartz subsp. multifida Gi 1463.

quadripinnata (Forssk.) Kuhn Gi 783, s.n.; Ph 1006.

viridis (Forssk.) Swartz var. glauca (Sim) Schelpe & N.C.

Anthony Gi s.n.

viridis (Forssk.) Swartz var. macrophylla (Kuntze) Schelpe &

N.C. Anthony Gi 791, 793, 794, s.n.; Ph 113.

viridis (Forssk.) Swartz var. viridis GR 3015; Gi s.n.

P35 Pellaea calomelanos (Swartz) Link Gi s.n.

POLYPODIACEAE

P42 Polypodium

polypodioides (L.) Hitchc. subsp. ecklonii (Kunze) Schelpe

GR 3815, 381 7; Gi 802, 803, s.n.

vulgare L. Gi 800, 801, 1341, s.n., Ph 824.

P43 x Pleopodium simianum Schelpe & N.C. Anthony Gi 511.

P44 Pleopeltis

macrocarpa (Bory ex Willd.) Kaulf. Gi s.n.

schraderi (Mett.) Tardieu Fu & Ph 43; Gi 61, s.n.

ASPLENIACEAE

P52 Asplenium

aethiopicum (Bunn, f.) Becherer Gi 2, 787, 1462, s.n.

boltonii Hook, ex Schelpe GR 3829; Gi 786, s.n.

erectum Bory ex Willd. var. erectum GR 3820; Gi 551, 785,

s.n.

lunulatum Swartz Gi 528, s.n.

monanthesZ,. Gi 784, s.n.

platyneuron (L.) Oakes Gi 1473, s.n.

protensum Schrad. Gi s.n.

rutifolium (Berg.) Kunze GR 3827; Gi s.n.

stoloniferum Bory GR 3826; Gi s.n.

Bothalia 17,2 (1987)

243

theciferum (H.B.K.) Mett. vai. concinnum (Schrad.) C. Chr. Gi

789.

trichomanes L. Ph 878.

varians Wall, ex Hook. & Grev. subsp. fimbria turn (Kunze)

Sc he Ip e Gi s.n.

P53 Ceterach cordatum (Thunb.J Desv. Gi 42.

THELYPTERIDACEAE

P54 Thelypteris

bergiana (Schlechtd.) Ching Gi 471, 555, s.n.; Ph 91 7.

confluens (Thunb.) Morton Ph 1284.

pozoi (Lagasca) Morton Gi 472, 775, s.n.

ATHYRIACEAE

P61 Cystopteris fragilis (L.) Bernh. Gi 138, s.n.

LOMARIOPSIDACEAE

P62 Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe Gi

759, 804, s.n.; Ph 1270.

ASPIDIACEAE/DRYOPTERIDACEAE

P66 Dryopteris inaequalis (Schlechtd.) Kuntze Gi s.n.; Ph 939.

P67 Cyrtomium caryotideum (Wall, ex Hook. & Grev.) Presl var.

micropterum (Kunze) C. Chr. Gi 777, 780, s.n.

P68 Polystichum

luctuosum (Kunze) T. Moore GR 3832; Gi 843, s.n.

monticola Schelpe & N.C. Anthony Fu & Ph 49; Gi s.n.

pungens (Kaulf.) Presl Gi 776, 1426a.

transkeiense Jacobsen, Hogsback, Jacobsen 4546 (Jacobsen

1978: 170).

P69 Arachnoides foliosa (C. Chr.) Schelpe, Hogsback, Jacobsen

4544 (Jacobsen 1983: 450).

P70 Rumohra adiantiformis (G. Forst.) Ching Gi s.n.

BLECHNACEAE

P75 Blechnum

australe L. var. australe Gi 781, 1464, s.n.; Ph 337.

capense (L.) Schlechtd. Ph 955.

giganteum (Kaulf.) Schlechtd. Gi 782, 1436, s.n.

tabulare (Thunb.) Kuhn Gi s.n.

GYMNOSPERMAE

PODOCARPACEAE

13 Podocarpus

falcatus (Thunb.) R. Br. ex Mirb. Fu s.n.; Fu & Ph 250; GR

3825, 3835; Gi 75.

latifolius (Thunb.) R. Br. ex Mirb. Gi 870; Ph 571.

PINACEAE

22 Pinus spp. Seen naturalized in many places.

CUPRESSACEAE

38 Widdringtonia nodiflora (L.) Powrie Gi 234.

ANGIOSPERMAE — MONOCOTYLEDONEAE

POTAMOGETONACEAE

58 Potamogeton pusillusZ. Fu & Ph 197.

POACEAE

K2 Coix lacryma-jobi L. Ph 135.

K10 Ischaemum fasciculatum Brongn. Ph 1486.

K28 Elionurus muticus (Spreng.) Kunth Fu & Ph 100; GR

3414, 3453, 3490a; Ph & Hu 93.

K29 Coelorhachis capensis Stapf Ph 1323.

K38 Miscanthus capensis (Nees) Anderss. var. capensis Fu & Ph

299; Gi 660, s.n.

K53 Eulalia villosa (Thunb.) Nees Ph 1016.

K71 Andropogon appendiculatus Nees Br 42, 159; Fu et al. 10,

11; GR 34 77; Gi 1280; Ph & Hu 59.

K72 Cymbopogon nardus (L.) Rendle Fu & Ph 222, 300; GR

3133, 3540; Gi 1284, s.n.

K73 Hyparrhenia hirta Stapf Gi s.n.

K78 Trachypogon spicatus (L. f.) Kuntze Br 10, 15; Fu & Ph

249; Gi 1283.

K80 Heteropogon contortus (L.) Roem. & Schult. GR 3445; Ph

979.

K83 Themeda triandra Forssk. Br 18, 33; Fu & Ph 244; GR

3452.

K89 Digitaria

diagonals (Nees) Stapf var. diagonalis Ph 1469.

sanguinalis (L.) Scop. Mahlobo 9.

setifolia Stapf Fu & Ph 212; GR 3492a.

temata (A. Rich.) Stapf Gi s.n.

K94 Alloteropsis semialata (R. Br.) Hitchc. subsp. eckloniana

(Nees) Gibbs Russell GR 3447; Ph 1193.

K104 Brachiaria eruciformis (J.E. Sm .) Griseb. Ph 1050.

K107 Paspalum scrobiculatum L. Gi s.n.

K112 Echinochloa crus-galli (L.) Beauv. Gi s.n.

K115 Oplismenus hirtellus (L.) Beauv. Gi 659, 1442, s.n.

K1 16 Panicum

aequinerve Nees Fu & Ph 225; GR 3416, 3489a; Gi s.n.

deustum Thunb. GR 3130; Gi 163, 512; Ph 105.

ecklonii Nees GR 3411, 3454; Ph 1155.

hymeniochilum Nees Ph 1324.

K128 Setaria

sphacelata (Schumach.) Moss var. sericea (Stapf) Clayton Ph

957.

sphacelata (Schumach.) Moss var. sphacelata Ph 1057.

K132a Rhynchelytrum nerviglume (Franch.) Chiov. Ph 964.

K139 Pennisetum

clandestinum Chiov. Ph 394.

macrourum Trin. Br 19; Fu 681; Fu & Ph 26, 1 77, 211.

thunbergii Kunth Br 01, 1 7, 24, 113; Fu & Ph 78, 80, 82, 121,

135, 147.

K160 Ehrharta

calycina J.E. Sm. var. calycina Fu & Ph 357.

erecta Lam. var. erecta Mdzeke 21; Ph 931.

K163 Phalaris

angusta Nees ex Trin. Ph 506.

arundinacea L. Fu & Ph 1 70.

K164 Anthoxanthum

ecklonii (Nees ex Trin.) Stapf Dyer 339 (GRA); Rattray 204

(GRA).

odoratum L. Ph 906.

K173 Arundinella nepalensis Trin. Br 40; Fu 756; Fu et al. 44;

Gi s. n. ; Ph 1058.

K174 Tristachya leucothrix Nees Br 34, 43; Fu et al. 12; Fu &

Ph 330; GR 3409; Ph & Hu 62.

K192 Holcus lanatusi. Fu&Phl78.

K197 Helictrotrichon hirtulum (Steud.) Schweick. GR 3518; Ph

1028.

K204c Merxmuellera drakensbergensis (Schweick.) Conert Ph &

Hu 1.

K205 Pentaschistis

setifolia (Thunb.) McClean Br 16; Fu & Ph 213.

tysonii Stapf Fu & Ph 97; GR 3507a.

sp. Fu & Ph 358.

K214 Phragmites australis (Cav.) Steud. Fu 759; Fu & Ph 1 75.

K243 Agrostis

barbuligera Stapf \ ar. barbuligera Br 09; Fu 724; GR 3418.

bergiana Trin. var. bergiana Mdzeke 1 7.

K262 Aristida junciformis Trin. & Rupr. subsp. galpinii (Stapf)

De Winter Br 20, 38, 157; Fu et al. 48; Gi s.n.; Mdzeke

06.

K263 Stipa

dregeana Steud. var. elongata (Nees) Stapf Ph 797.

trichotoma A'ees Fu & Ph 31 7.

K283 Sporobolus centrifugus (Trin.) Nees GR 3487a ; Gi 1282;

Ph & Hu 87.

K286 Eragrostis

caesia Stapf Gi s.n.

capensis (Thunb.) Trin. Br 134; Fu et al. 45; GR 3446; Ph

978; Ph & Hu 60.

244

Bothalia 17,2 (1987)

curvula (Schrad.) Nees GR 3444; Ph 1487.

planiculmis Nees Fu 752.

racemosa (Thunb.) Steud. Ph & Hu 96; Ph 1257.

K294 Microchloa caffraTVees Gi s.n.

K296 Cynodon dactylon (L.) Pers. Fu & Ph 281; Gi s.n.

K298 Harpochloa falx (L. f.) Kuntze Br 158; Fu & Ph 58; GR

3438; Gi 1279a; Ph & Hu 67.

K371 Fingerhuthia sesleriifonnis Nees Br 12; Fu 722; Fu et al.

07; Fu & Ph 81, 96.

K374 Koeleria capensis (Steud.) Nees Fu & Ph 99; GR 3412; Ph

& Hu 86.

K386 Melica racemosa Thunb. GR 3519.

K399 Lasiochloa longifolia (Schrad.) Kunth Ph 1135; Ph & Hu

162.

K400 Stiburus alopecuroides (Hack.) Stapf Br 77.

K404 Briza

maxima L. Ph 1525.

minor L. Ph 50 7.

K407 Poa

annua L. Mdzeke 20.

binata Nees Ph 134 7.

heterogam a Hack. Ph 1169.

pratensis L. Fu & Ph 328.

K417 Festuca

caprina Nees var. caprina Br 118, 153; Fu & Ph 98.

capnna Nees var. irrasa Stapf GR 3408.

costata Nees var. costata Br 1 23; Ph & Hu 89.

longipes Stapf Ph 791.

K418 Vulpia

bromoides (L.) S.F. Gray Fu et al. 39.

myuros (L.) C.C. Gmel. Fu & Ph 321.

K428 Bromus

molliformis Lloyd Fu & Ph 320.

speciosus Nees Fu & Ph 132; GR 3413, 3494a; Ph & Hu 92.

unioloides H.B.K. Mdzeke 28.

K432 Brachypodium flexum Nees GR 3024; Gi s.n.; Russell

2354.

K433 Lolium multiflorum Lam. Fu & Ph 343.

K457 Thamnocalamus tesselata (Nees) Soderstrom & Ellis Gi

280; Ph 434.

CYPERACEAE

454 Ascolepis capensis (Kunth) Ridley Br 109, 135, 147; Fu

735; Fu et al. 53; Fu & Ph 1 73, 332.

456 Carpha

bracteosa C.B. Cl. Fu & Ph 182, 347.

glomerata (Thunb.) Nees Ph 935.

459 Cyperus

albostriatus Schrad. GR 3023; Gi 1589.

difformis L. Ph 5 72.

obtusiflorus Vahl var. flavissimus Boeck. Ph 99, 1298.

pulcher Thunb. Ph 110, 950.

schlechteri C.B. Cl. Ph 1165.

semitrifidus Schrad. Ph 1005, 1164; Ph & Hu 165.

tenellusZ,. var. tenellus Gi 1596; Ph 344.

459a Pycreus

betschuanus (Boeck.) C.B. Cl. Fu & Ph 122, 157, 200.

cooperi C.B. Cl. Br 25, 140; Fu et al. 14; Fu & Ph 123, 156,

169, 1 79, 205.

macranthus C.B. Cl. Fu & Ph 158, 207.

mundtii Nees Fu & Ph 163, 187.

nitidus (Lam.) J. Raynal Fu & Ph 102, 164, 1 71; Ph 237, 238.

oakfortensis C.B. Cl. Fu & Ph 28.

unioloides (R. Br.) Urb. Fu et al. 06, 51.

459c Mariscus

congestus (Vahl) C.B. Cl. GR 3007; Gi 1593; Ph 156, 239,

1035.

grantii C.B. G. Ph 1254.

owanii (Boeck.) C.B. Cl. Ph 398.

tabularis (Schrad.) C.B. Cl. Ph 1299.

thunbergii (Vahl) Schrad. Fu & Ph 192.

462 Kyllinga

elatior Kunth Ph 109.

erecta Schumach. Fu & Ph 186.

melanosperma Nees Ph 1296.

pauciflora Ridley Br 26, 136, 168; Fu & Ph 159; Ph 242.

pulchella Kunth Ph 1188.

465 Ficinia

bergiana Kunth Ph 921.

cinnamomea C.B. Cl. Gi 3509a; Ph 944.

fascicularis Nees Gi 1597, s.n.; Ph 426.

stolonifera Boeck. Ph & Hu 76.

tristachya (Rottb.) Nees GR 3469.

467 Fuirena

pubescens (Poir.) Kunth Fu et al. 54; Fu & Ph 208; GR 3510a.

sp. Ph 1498.

468 Scirpus

falsus C.B. Cl. Ph & Hu 17.

ficinioides Kunth Br 02, 30, 87, 101, 103; Fu et al. 29; Fu &

Ph 79, 335.

inanis (Thunb.) Steud. Fu & Ph 125.

468a Schoenoplectus paludicola (Kunth) Palla ex J. Raynal Ph

1328.

468b Isolepis

cemua (Vahl) Roem. & Schult. Fu 692; Fu et al. 40; Fu & Ph

77, 189.

costata (Boeck.) A. Rich. Br 16 7; Fu 693, 719; Fu et al. 03,

41, 42; Fu & Ph 124; GR 3003.

fluitans (L.) R. Br. Fu & Ph 01, 134; Ph 1225.

ludwigii Kunth Fu & Ph 162.

natans (Thunb.) Dietr. Fu & Ph 29, 131, 201, 209.

sepulcralis Steud. GR 3513a; Ph 493.

469 Eleocharis acutangula (Roxb.) Schult. Fu & Ph 193.

471 Fimbristylis complanata (Retz.) Link Ph 236, 1203.

471a Bulbostylis

humilis (Kunth) C.B. Cl. Fu & Ph 7; Ph 1350.

schoenoides (Kunth) C.B. Cl. Fu & Ph 83, 103, 333.

471b Abildgaardia ovata (Burm. f.) Krai Ph 1310.

492 Rhynchospora brownii Roem. & Schult. Fu 728, 733; Fu et

al. 5; Fu & Ph 14, 160.

494 Tetraria

cuspidata (Rottb.) C.B. Cl. Fu & Ph 118, 248; GR 3514a; Ph

562.

macowaniana B.L. Burtt Fu <6 Ph 234; Ph 55 7.

521 Schoenoxiphium

lehmannii (Nees) Steud. Gi 513, 1588; Ph 930.

perdensum Kukkonen, Keiskammahoek, near Ghulu Kop,

4,000 ft Dyer 245a (K) (Kukkonen 1983 : 822).

rufum Nees Ph & Hu 112; Robinson s.n.

sparteum (Wahlenb.) C.B. G. Ph & Hu 16, 83, 166.

sp. aff. S. schweickerdtii Merxm. & Podlech Ph 430; Ph & Hu

18.

525 Carex

acutiformis Ehrh. Br 98; Fu 688, 745, 746; Fu et al. 52; Fu &

Ph 86.

clavata Thunb. Fu & Ph 128, 1 72; Gi 1586.

petitiana/l. Rich. Ph 940.

schlechteri Nelmes Ph 788.

zuluensis C.B. G. Ph 11 70.

ARACEAE

748 Zantedeschia

aethiopica (L.) Spreng. GR 3500; Gi 885.

albomaculata (Hook.) Baill. subsp. albomaculata Ph 1098.

RESTIONACEAE

804 Restio sejunctus Mast. Ph 323.

804c Ischyrolepis distracta (Mast.) Linder, Gaika’s Kop (Linder

1985: 404).

804j Calopsis paniculata (Rottb.) Desv. Gi 101 7; Ph 411.

804p Hydrophilus rattrayi (Pillans) Linder, Hogsback (Linder

1985: 484).

807 Elegia asperiflora (Nees) Kunth var. lacerata (Pillans) Pillans

Fu&Ph 16,1 76; Gi s.n.

Bothalia 17,2 (1987)

245

XYRIDACEAE

826 Xyris capensis Thunb. Fu 734; Fu et al. 02; Fu & Ph 30; Gi

273.

ER10CAULACEAE

828 Eriocaulon dregei Hochst. var. dregei Br 29, 137; Fu 736;

Fu et al. 09; Fu & Ph 4; Ph 241.

COMMELINACEAE

896 Commelina

africanaZ,. var. africana Br21;Ph 142.

africana'Z. var. krebsiana (Kunth) C.B. Cl. Gi s.n.

JUNCACEAE

936 Juncus

bufonius L. Fu & Ph 202; Ph 499.

capensis Thunb. GR 3511a.

dregeanus Kunth Br 110, 126; Fu & Ph 3, 85, 101, 188; GR

3512a; Ph 494.

effusus L. Fu 714, 743, 744; Fu et al. 30; Fu & Ph 130, 190;

Ph 812.

exsertus Buchen. Fu & Ph 1 91.

lomatophyllus Spreng. Br 139; Fu 682, 723; Fu & Ph 08; GR

3002.

oxycarpus Kunth Fu et al. 49; Fu & Ph 27, 129; GR 3504a;

Ph 496.

punctorius L. f. Fu & Ph 165.

LILIACEAE

969 Androcymbium longipes Bak. Gi 640; Ph 1348.

972 Wurmbea elatior B. Nord. Br 13; Ph 1074.

985 Bulbine

abyssinica A Rich. Ph 1290.

lagopus (Thunb.) N.E. Br. Ph 884, 1333.

985a Trachyandra saltii (Bak.) Oberm. var. saltii GR 3441; Ph &

Hu 51.

989 Anthericum angulicaule Bak. Ph 1332.

990 Chlorophytum

bowkeri-Bck. Ph 1191.

comosum (Thunb.) Jacq. Gi 620, 720, 1542.

1002 Caesia contorta (L. f.) Dur. & Schinz GR 3471; Ph 770.

1011 Bowiea volubilis Harv. ex Hook. f. Ph & Hu 100.

1012 Eriospermum natalense Bak. Gi 1106; Ph & Hu 98.

1024 Kniphofia

baurii Bak. Phll04.

linearifolia Bak. Ph 1319.

northiae Bak. Gi 564.

parviflora Kunth Ph 1277.

triangularis Kunth subsp. triangularis Fu & Ph 199; Gi 1352.

uvaria (L.) Hook. Ph 1078.

1026 Aloe

arborescens Mill. Ph 833.

aristata Haw. Ph 1138.

ecklonis Salm-Dyck Gi 1322.

ferox Mill. Seen at lower altitudes near Mitchell’s Pass,

maculata All. Fu & Ph 313.

1046 Agapanthus

campanulatus Leighton subsp. campanulatus Ph 1266.

praecox Willd. subsp. orientalis (Leighton) Leighton GR 3507;

Gi 1247.

1047 Tulbaghia acutiloba Harv. Gi s.n.; Ph & Hu 101.

1072 Lilium formosanum (Bak.) Wallace Ph 1044.

1079 Albuca

caudata Jacq. Gi 210; Ph & Hu 69.

fastigiata (L. f.) Dry and. Ph 1182.

nelsonii N.E. Br. Ph & Hu 66.

setose Jacq. GR 3461; Ph 1139.

1080 Urginea tenella Bak. Ph & Hu 55.

1082 Drimia elata Jacq. Gi 1650.

1086 Scilla nervosa (Burch.) Jessop Fu & Ph 139.

1088 Eucomis

autumnalis (Mill.) Chitt. subsp. autumnalis GR 3491a.

comosa (Houtt.) Wehrh. var. comosa Fu & Ph 166; Gi 454.

1089 Omithogalum

conicum Jacq. subsp. conicum Ph 941.

graminifolium Thunb. Ph & Hu 40.

juncifolium Jacq. Fu & Ph 11 7; Gi 281, 1165; Ph 943; Ph &

Hu 39.

longibracteatum Jacq. Ph 887.

paludosum Bak., Elandsberg, Cooper 219 (K) (Obermeyer

1978: 350).

tenuifolium Delaroche subsp. tenuifolium Fu et al. 16; Fu &

Ph 340; GR 3467; Gi 1165a.

1090a Ledebouria floribunda (Bak.) Jessop Ph 968; Ph & Hu

1079.

1101 Massonia echinata L. f. Fu 680.

1113 Protasparagus

aethiopicus (L.) Oberm. Fu & Ph 263; Gi 849; Ph 884.

africanus (Lam.) Oberm. Ph 1136.

densiflorus (Kunth) Oberm. Ph 1102; Ph & Hu 64.

denudatus (Kunth) Oberm. Gi s.n.

laricinus (Burch.) Oberm. Ph 1103; Tu 138.

macowanii (Bak.) Oberm. Ph 1106.

subulatus (Thunb.) Oberm. Gi 848.

virgatus (Bak.) Oberm. Gi 453.

1113a Myrsiphyllum

asparagoides (L.) Willd. Fu & Ph 45; Gi 621, g799, 11)5;

Robinson 1065.

ramosissimum (Bak.) Oberm. Nete 21; Ph & Hu 5; Tyibilika

44.

1 147 Behnia reticulata (Thunb.) Didr. Gi 494, 530, 645.

AMARYLLIDACEAE

1167 Haemanthus albiflos Jacq., Menziesberg, Acocks 1114 (K,

PRE); Hogsback Forest Reserve, Dahlstrand 2935 (GRA,

STE) (Snijman 1984: 63).

1167a Scadoxus puniceus (L.) Friis & Nordal Ph 945.

1175 Nerine undulata (L.) Herb. GR 3027a; Gi 449, 1523;

McGillivray 48; Ph 559, 1486.

1191 Cyrtanthus

breviflorus Harv., Umgaika Kop, Cooper 255 (Reid & Dyer

1984: 14).

huttonii Bak. Gi s.n.; Ph 1113.

tuckii Bak. var. viridilobus Verdoom Ph 948.

HYPOXIDACEAE

1230 Hypoxis

angustifolia Lam. Fu & Ph 342.

costata Bak. GR 3456.

filiform is Bak. Br 107; Fu et al. 4 7; Fu & Ph 12, 95.

flanaganii Bak. Ph 1011.

hemerocallidea Fisch. & Mey. GR 3482.

multiceps Buchinger Ph & Hu 91.

rigidula Bak. var. rigidula Ph & Hu 80.

setosa Bak. Fu & Ph 344.

woodii Bak. Ph 1194, 1496.

sp. Br 161; Fu & Ph 322; GR 3483; Ph 993.

DIOSCOREACEAE

1252 Dioscorea

retusaMwr. Ph 1192, 1316.

rupicola Kunth Gi 995, 1105.

stipulosa Uline ex Kunth Gi 1195, 1300; Ph 1227.

sylvatica (Kunth) Eckl. , Hogsback Pass, Archibald 7556 (GRA)

(Archibald 1967: 36).

IRIDACEAE

1260 Syringodea concolor (Bak.) De Vos Gi s.n.

1261 Romulea

atrandra G.J. Lewis var. lewisiae De Vos Ph 879.

autumnalis L. Bol., Summit of Hog’s Back, Scully 346 (SAM)

(De Vos 1972: 205).

macowanii Bak. var. macowanii Ph 826.

1265 Moraea

elliottii Bak. Fu & Ph 120.

huttonii (Bak.) Oberm. Gi 196; Makunga M31; Ph 400.

246

Bothalia 17,2 (1987)

muddii N.E. Br. Ph 1268.

reticulata Goldbl. GR 3019; Gi s.n., 443; Ph 1054.

stricta Bak. Ph & Hu 36.

1265a Dietes iridoides (L.) Sweet ex Klatt Ph 954.

1295 Aristea

anceps Eckl. ex Klatt Ph & Hu 73.

cognata N.E. Br. ex Weim. Fu & Ph 91; Ph & Hu 72.

ecklonii Bak. Makunga M42.

montana Bak. Ph 1273.

schizolaena Harv. Fu & Ph 233; Gi 4 77, gl262; Ph & Hu 71.

1299 Schizostylis coccinea Backh. & Harv. Fu 684; GR 3136;

Gi 165; Makunga M81.

1301 Hesperantha

huttonii (Bak.) Hilliard & Burtt Gi s.n. Ph 163.

longituba (Klatt) Bak. Gi s.n., Fu & Ph 38.

pulchra ZtoA:. Ph 561.

sp. Br 88 b.

1302 Ixia orientalis L. Bol. Br 106, 149, 152; Fu & Ph 88;

Makunga M70.

1303 Dierama

igneum Klatt Fu & Ph 133; Ph 987.

pendulum (L. f.) Bak. Gi 726a; Ph 1343.

pulcherrimum (Hook f.) Bak. GR 3018.

1306 Tritonia

disticha (Klatt) Bak. subsp. rubrolucens (R.C. Post.) De Vos

Makunga M64; Ph & Hu 25.

lineata (Salisb.) Ker-Gawl. var. lineata Ph 387.

1311 Gladiolus

dalenii Van Geel Ph 1229.

ecklonii Lehm. subsp. ecklonii Makunga M42.

longicollis Bak. var. longicollis Br 1 08; Ph & Hu 2.

ochroleucus Bak. var. macowanii (Bak.) Oberm. Page &

Tapson 20.

ochroleucus Bak. var. ochroleucus Ph 1219.

1315 Watsonia

longifolia J.W. Mathews & L. Bol. GR 3545; Gi 118, 726;

Makunga M13;Ph & Hu 157, 158.

pillansii L. Bol. Gi 490.

sp. Fu & Ph 181; Ph & Hu 21, 58.

1316a Anomatheca laxa (Thunb.) Goldbl. Gi 142, s.n.; Ph 106.

ORCHIDACEAE

1407 Stenoglottis fimbriata Lindl. Gi s.n.

1408 Holothrix

orthoceras (Harv.) Reich b. f. Gi 1536, s.n.

scopularia (Lindl.) Reichb. f. Fu & Ph 110; Gi 215.

1414 Huttonaea pulchra Harv. Gi 604, 613, s.n.

1422 Habenaria

falcicomis (Burch, ex Lindl.) H. Bol. var. caffra (Schltr.) Renz

& Schelpe Gi 605; Ph 1306.

laevigata Lindl. subsp. laevigata Gi 607.

malacophylla Reichb. f. Gi 1538.

1422b Bonatea cassidea Sond. Gi 86; Ph 904.

1429 Neobolusia tysonii (H. Bol.) Schltr. Ph 1291.

1430 Satyrium

bracteatum (L. f.) Thunb., Gaika’s Kop, Rattray s.n. (BOL)

(Hall 1982: 122).

cristatum Sond. var. cristatum, Hogsback Mtn, Rattray s.n.

(BOL) (Hall 1982: 94).

hallackii H. Bol subsp. ocellatum (H. Bol.) A. V. Hall, Hogsback

Mtn, Rattray 15780 (BOL, PRE), Rattray 86 (GRA)

(Hall 1982: 57).

ligulatum Lindl., Hogsback, Barker 1488 (NBG) (Hall 1982:

46).

longicauda Lindl. var. jacottetianum (Kraenzl.) A. V. Hall Gi

216, 617, 1315.

longicauda Lindl. var. longicauda Ph 612, 618, 1167.

parviflorum Swartz Gi 108; Ph 1267.

143 1 Schizochilus zeyheri Sond. Gi 21 7, 603; Ph 1282.

1433 Brownleea

coerulea Harv. ex Lindl. Gi 444, 602; Ph 131 7.

m acroceras Sond. Ph 1265.

parviflora Harv. ex Lindl. Gi 158.

recurvata Sond. Gi 608.

1434 Disa

aconitoides Sond. subsp. aconitoides Gi 1314.

chrysostachya Swartz Gi 214.

crassicornis Lindl. GR 3005; Gi 1 09.

pulchra Sond. Ph & Hu 56 (KEI).

sagittalis (L. f.) Swartz GR 3473; Gi 1162; Ph & Hu 44, 122;

Robinson s.n.

sanguinea Sond. , Amatole Mts (Linder 1981a: 70).

scullyi H. Bol., Menziesberg, Scully s.n. (BM, BOL, K); Hogs-

back, Batten s.n. (BOL) (Linder 1981a: 119).

stricta Sond. Gi614.

thodei Schltr. ex Kraenzl., Gaika’s Kop (Linder 1981a: 115).

tysonii H. Bol., Quarter degree square 3227CA, land over 1500

m (from distribution map, Linder 1981a: 74).

versicolor Reichb. f. Gi 245, 131 9.

1435 Herscheliavenusta (H. Bol.) Kraenzl., Hogsback Mtn, Rat-

tray s.n. (BOL) (Linder 1981b: 378).

1436 Monadenia brevicomis Lindl. Gi 606.

1437 Disperis

lindleyana Reichb. f. Gi 609.

macowanii H. Bol. Ph 1345.

micrantha Lindl. Gi s.n.

stenoplectron Reichb. f. Gi s.n.

wealii Reichb. f. Gi 131 7.

1440 Corycium

dracomontanum Parkman & Schelpe Gi 1163; Ph & Hu 163.

magnum (Reichb. f.) Rolf e Gi615.

1565 Polystachya

ottoniana Reichb. f. GR 3834; Gi 19, 556, 610.

pubescens Reichb. f. Gi 889.

1648 Eulophia

aculeata (L. f.) Spreng. subsp. aculeata GR 3474; Ph & Hu

131.

aculeata (L. f.) Spreng. subsp. huttonii (Rolfe) A.V. Hall Gi

1164, 1166, 1246; Ph & Hu 164.

foliosa (Lindl.) H. Bol., Chumie Peak, Scully 172 (BOL) (Hall

1965: 228).

macowanii Rolfe, Chumie Peak, Scully 173 (BOL) (Hall 1965:

232).

meleagris Reichb. f. Ph & Hu 132.

ovalis Lindl. subsp. ovalis Ph & Hu 65.

1828 Angraecum

conchiferum Lindl. Gi 770.

sacciferum Lindl. Gi 601.

1837 Mystacidium

flanaganii (H. Bol.) H. Bol. Ph 114.

gracile ( Reichb . f.) Harv. Gi 20, 611.

ANGIOSPERMAE — DICOTYLEDONEAE

PIPERACEAE

1866 Peperomia

retusa (L. f.) A. Dietr. Ph 1114.

tetraphylla (G. Forst.) Hook. & Am. GR 3824; Gi 44, 893.

SALICACEAE

1873 Salix

capensis Thunb. subsp. capensis Gi 249.

sp. Tu 118.

MYRICACEAE

1874 Myrica

brevifolia E. Mey. ex DC. Fu & Ph 235; Gi 194; Ph 425.

serrata Lam., Hogsback, Rattray 303 (PRE) (Killick 1969: 9).

BETULACEAE

1887 Betula sp. GR 3813, 3814.

FAGACEAE

1893 Quercus robur L. Robinson 1058.

Bothalia 17,2 (1987)

247

ULMACEAE

1898 Celtis africana Burm. f. Tu 183.

1 906 Chaetacme aristata Planch. Ph 786.

MORACEAE

196 1 Ficus

burtt-davyi Hutch. Fu & Ph 312.

sur Forssk. Fu s.n.; Gi s.n.

URTICACEAE

1980 Laportea peduncularis (Wedd.) Chew Gi 112, 400, 560,

869, 1019.

2007 Parietaria micrantha Ledeb. Ph 1340.

2013 Droguetia iners Forssk. subsp. iners Gi 1458, 1537.

2014a Didymodoxa caffra (Thunb.) Friis & Wilmot-Dear Gi

868, 1545.

PROTEACEAE

2035 Protea

simplex Phill. Gi 1160; Lyle 79.

subvestita N.E. Br. Gi 1353; Ph 567; Tu 250.

LORANTHACEAE

2074a Tapinanthus prunifolius (E. Mey. ex Harv.) V. Tieghem

Gi 1065.

VISCACEAE

2093 Viscum obscurum Thunb. Fu & Ph 255.

SANTALACEAE

2104 Colpoon compressum Berg. Gi s.n.

2116 Osyridocarpus schimperianus (Hochst. ex A. Rich.) DC. Gi

478.

2118 Thesium

pallidum DC. Gi 230, 1261.

triflorum Thunb. ex L. f. Gi 581.

POLYGONACEAE

2195 Rumex

angiocarpus Murb. Br 86; Fu 747; Gi 1226, 1276.

crispusZ,. Ph 1287.

dregeanus Meisn. var. dregeanus Ph 1166.

lanceolatus Thunb. Ph 1187.

sagittatus Thunb. Gi 498; McGillivray 5; Ph 1036.

steudelii Hochst. Ph 1223, 1289.

woodii N.E. Br. Fu & Ph 136, 316.

2201 Polygonum

lapathifolium L. var. maculatum (S.F. Gray) T.-Dyer & Trin.

Ph 1025.

meisnerianum Cham. & Schlechtd. Fu & Ph 196; Gi s.n.

persicaria L. GR 3000.

salicifolium Willd. Ph 1315.

CHENOPODIACEAE

2223 Chenopodium multifidum L. Ph 1046.

AMARANTHACEAE

2312 Cyathula uncinulata (Schrad.) Schinz Gi 863; Ph 1471.

2328 Achyranthes sicula (L.) All. Gi 544, 864.

AIZOACEAE

2379 Psammotropha mucronata (Thunb.) Fenzl var. mucronata

Br 156; Gi406, 1227.

PHYTOLACCACEAE

2380 Phytolacca

heptandra Retz. Ph 1301.

octandra L. Ph 264.

MESEMBRYANTHEMACEAE

2405 Aptenia cordifolia (L. f.) Schwant. Gi 1056.

CARYOPHYLLACEAE

2429 Stellaria media (L.) Vill. Ph 887.

2430 Cerastium

capense Sond. Br 100, 104, 145; Fu & Ph 331; Ph 926.

indicum Wight & Am. Ph 1482.

2490 Silene

burchellii Otth var. burchellii Br 127; Ph 1281.

undulata>l/f. Gi 1081.

vulgaris (Moench) Garcke subsp. macrocarpa (Marsden) Jones

& Turrill Ph 1131.

2502 Dianthus

basuticus Burtt Davy subsp. basuticus var. basuticus Ph & Hu

168.

crenatus Thunb. Ph & Hu 150.

RANUNCULACEAE

254 1 Anemone caffra Eckl. & Zeyh. Ph 106 7; Ph & Hu 138.

2541a Knowltonia cordata H. Rasm. Ph 913.

2542 Clematis brachiata Thunb. Gi 1250; Mgudlwa 40; Ph 551;

Tu 193.

2546 Ranunculus

baurii Macowan Ph 432.

meyeri Harv. Fu & Ph 2, 50, 334; Gi s.n.

multifidus Forssk. Br 102; Fu & Ph 75; GR 3497; Gi 441,

592, 593; Ph 423.

2548 Thalictrum rhynchocarpum Dill. & Rich. Gi 46 7.

MENISPERMACEAE

2574 Cissampelos torulosa E. Mey. ex Harv. Fu & Ph 256; Gi

1504; Ph 112.

TRIMENIACEAE

2759a Xymalos monospora (Harv.) Baill. Gi 73, g852 s.n.; Ph

869, 870.

LAURACEAE

2813 Cryptocarya woodii Engl. Gi 99.

PAPAVERACEAE

2853 Papaver aculeatum Thunb. Fu £ Ph 87, 141.

FUMARIACEAE

2858a Phacocapnos pruinosus (E. Mey.) Bemh. Ph & Hu 167.

2861 Fumaria muralis Sond. ex Koch subsp. muralis Ph 1526.

BRASSICACEAE

2875 Heliophila

elongata (Thunb.) DC. Gi 284, 1245, s.n.; Ph 1097; Ph & Hu

34.

rigidiuscula Sond. Fu & Ph 108, 356; Gi 111 7.

2883 Lepidium ecklonii Schrad., Hogsback, Jacot Guillarmod

4523 (Marais 1970: 93).

2965 Rorippa nasturtium-aquaticum (L.) Hayek GR 3823.

2966 Cardamine africana L. Ph 903.

CAPPARACEAE

3112 Maerua racemulosa (A. DC.) Gilg & Ben. Gi 534.

DROSERACEAE

3136 Drosera aliciae R. Hamet Fu & Ph 18; Gi 244, 272, 557;

Ph & Hu 123.

CRASSULACEAE

3 164 Cotyledon orbiculata L. var. oblonga (Haw.) DC. Ph 897.

3 168 Crassula

cordata Thunb. Gi 476.

dependens H. Bol. Fu 697; Fu & Ph 31.

natans Thunb. var. natans Fu & Ph 204; Ph 1141.

nemorosa (Eckl. & Zeyh.) Walp. Gi 1029a; Ph 907.

nudicaulis L. var. nudicaulis Br 90; Gi 1332.

obovata Haw. var. obovata Gi 349.

orbicularis L. Ph 884.

248

Bothalia 17,2 (1987)

pellucida L. subsp. maiginalis (Dryand. in Ait.) Tolken Br 82;

Gi 350, 1340; Russell 2351.

sarcocaulis Eckl. & Zeyh. subsp. sarcocaulis Gi 1577a, s.n.

sediflora (Eckl. & Zeyh.) Endl. ex Walp. var. amatolica (Schonl.)

Tolken, Cata Ridge, Dyer 356 (GRA, K. LU, PRE)

(Tolken 1977: 374).

spathulata Thunb. Gi 1018, 1540.

vaginata Eckl. & Zeyh. Ph 1065.

vaiOantii (Mild.) Roth. Ph 1483; Ph & Hu 145.

sp. Ph 564; Ph & Hu 53.

ESCALLONIACEAE

3241 Choristylis rhamnoides Harv. Gi 320, 1444, s.n.

PITTOSPORACEAE

3252 Pittosporum viridiflorum Sims Gi 98, s.n.; Ph 759; Tu

235.

HAMAMELIDACEAE

3311 Trichocladus ellipticus Eckl. & Zeyh. ex Walp. Fu s.n.; Gi

97, 890, s.n.; Ph 778, 1122.

ROSACEAE

3353 Rubus

fruticosus L. Gi 1348, 1351 ; Tu 91.

immixtus C.E. Gust. Fu & Ph 217; GR 3489; Gi 111, 1152;

Tu 106.

ludwigii Eckl. & Zeyh. subsp. ludwigii, Hogsback, Rattray s.n.

(PRE) (Stilton 1984: 103).

phoenicolasius Maxim. Ph 937.

pinnatus Willd. Gi 2 75; Ph 938.

rigidus Sm. Gi 185, 343; Ph 936.

3355 Duchesnea indica (Andr.) Focke GR 3026; Mahlobo 24;

Mgudlwa 23.

3365 Geum capense Thunb. Gi 341; Ph 390.

3375 Alchemilla

capensis Thunb. Ph 512.

elongata Eckl. & Zeyh. Fu & Ph 168.

hirsuto-petiolata (De Wild.) Rothm. Gi 762; Ph & Hu 135.

sp. 1 Ph & Hu 28.

sp. 2 Fu 696, 755; Gi 1271; Ph 1320.

3316 Agrimonia procera Wallr. Br 1 76; Gi 402, 1026; Ph 1489.

3379 Leucosidea sericea Eckl. & Zeyh. Gi 538; Ph 410, 428;

Robinson 1001, 1064; Tu 207, 215.

3388 Cliff ortia

linearifolia Eckl. & Zeyh. Fu & Ph 305; Tu 92, 94, 95, 102,

103, 119, 126, 194, 221.

paucistaininea Weim. Fu & Ph 215, 242; GR 3493a; Gi 342;

Tu 222, 229, 240, 244.

serpyllifolia Cham. & Schlechtd. Ph 508; Tu 214.

strobilifera Murray Tu 254.

sp. (C. eriocephalina Cham, sensu Story 1952: 152) Story 3318

(GRA).

3389 Rosa odorata Sweet Ph 916, 1043.

3396 Prunus

africana (Hook, f.) Kalkm. Tu 197.

spp. Introduced species have become naturalized, mainly along

roads.

FABACEAE

3446 Acacia

karroo liayne Tu 269.

mearnsii De Wild. Ph 899.

melanoxylon R. Br. Ph 892.

3468 Entada spicata (E. Mey.) Druce Fu s.n.; Gi 1058; Tu 190.

3506 Schotia latifolia Jacq. Fu s.n.; Fu & Ph 267; Gi 532, 1157.

3536 Cassia capensis Thunb. var. capensis Gi 1061, 1149; Ph 95.

3607 Calpumia

aurea (Ait.) Benth. subsp. sylvatica (Burch.) Brummitt Fu s.n.;

Gi 527, g889; Tu 192.

floribunda Harv. Gi 161, 1454.

3608 Virgilia divaricata Adamson Tu 246.

3657 Lotononis

camosa Benth. Gi 11 72; Ph 959, 1094, 1230; Ph & Hu 141.

cytisoidesitofc. var. cytisoides Br 180; Fu & Ph 60; GR 3516.

sp. Ph 1218, 1300.

3662 Aspalathus

frankenioides DC. Fu & Ph 238; Ph 1280.

simii//. Bol. subsp. katbergensis R. Dahlg. Ph 969.

3663 Buchenroedera

holosericeaftent/;. GR 3538; Ph 793, 996, 1221; Ph & Hu 14.

multiflora Eckl. & Zeyh. GR 3505a; Ph 418; Tu 249.

tenuifolia Eckl. & Zeyh. var. tenuifolia Ph 981; Tu 260.

3665 Melolobium alpinum Eckl. & Zeyh. Ph & Hu 149.

3673 Argyrolobium

baptisioides Walp. Br 1 70; GR 3499a.

crassifolium Eckl. & Zeyh. Tu 259.

longifolium Walp. Ph 1163, 1237.

molle Eckl. & Zeyh. Gi 1142; Ph 98.

pilosum Harv. GR 3470, 3506a; Gi 1326; Ph 1161.

speciosum Eckl. <6 Zeyh. Gi s.n.; Ph & Hu 50.

stipulaceum Eckl. & Zeyh. Ph 1216.

sutherlandii Harv. Ph 1162.

tomentosum (Andr.) Druce Gi 334, 763, 979.

tuberosum Eckl. & Zeyh. Fu et al. 46; Ph 1295, 1349.

3681 Ulex europaeus L. Gi 34, s.n.

3688 Medicago lupulina L. Ph 920.

3690 Trifolium

burchellianum Ser. subsp. burchellianum GR 3494, 3502a; Gi

719, 1146; Ph 395.

repens L. GR3816.

3702 Indigofera

alpina Eckl. & Zeyh. Br 116; Ph 41 7.

amatolensis Phillip son Ph 427; Ph & Hu 9, 139.

cuneifolia Eckl. & Zeyh. Gi 67, 335, 487, 1126, 1252; Ph 334;

Robinson 1061; Tu 209.

evansii Schltr. Gi 1530; Ph 404.

hedyantha Eckl. & Zeyh. Br 166; Gi 1230; Ph & Hu 129.

mollis Eckl. & Zeyh. Ph & Hu 153.

monostachya Eckl. & Zeyh. Br 115; Fu & Ph 71, 353, 354.

stricta L. f. Gi 1 098, 164 7; Ph 146.

woodii H. Bol. Gi 239; Ph 1015.

sp. Ph 403.

3703 Psoralea pinnata L. Fu & Ph 220; GR 3495a, 3821; Gi

336, 493, g840, 1253, 1646; Ph 164, 340, 821.

3703c Otholobium

caffrum (Eckl. & Zeyh.) C.H. Stirton Fu & Ph 229; Gi 233,

1310; McGillivray 15.

stachyerum (Eckl. & Zeyh.) C.H. Stirton GR 3498a.

3718 Tephrosia

capensis (Jacq.) Pers. var. capensis Ph 1018.

grandiflora (Ait.) Pers. Gi 270; Ph 148, 956.

macropoda E. Mey. Ph 1238.

margin ella H.M. Forbes Gi 1240.

polystachya E. Mey. Gi 162; Ph 1 039.

3754 Sutherlandia frutescensi?. Br. Ph 1134.

3756 Lessertia

llexuosa E. Mey. Fu dl Ph 140.

harveyana L. Bol. Gi 1099.

perennans DC. var. perennans Gi 1113; Ph <6 Hu 125.

3807 Desm odium repandum (Vahl) DC. Gi 837; Ph 1038.

3810 Alysicaxpus rugosus (Willd.) DC. subsp. perennirufus

J. Leonard Gi 1070; Ph 1 180.

3852 Vicia spp. Seen at various localities as weeds of cultiva-

tion.

3897 Rhynchosia

angulosa Schinz Ph 1309.

argentea Harv. Gi 1107, 1190, 1313.

caribaea (Jacq.) DC. Ph 1029.

cooperi (Harv. ex Bak. f.) Burtt Davy Ph 1189, 1302.

hannsiana Schltr. ex Zahlbr. var. burchellii Burtt Davy Ph

1312.

3898 Eriosema acuminata (Eckl. & Zeyh.) C.H. Stirton Ph

1019, 1184, 1314.

Bothalia 17,2 (1987)

249

3905 Vigna vexillata (L.) A. Rich. Gi 1 064; Ph 139.

3909 Lablab purpureus (L.) Sweet subsp. uncinatus Verde. Gi

160, 81 7, g892.

3910 Dolichos

angustifolius Eckl. & Zeyh. Ph 1311.

linearis E. Mey. Ph 549, 1213.

GERANIACEAE

3924 Geranium

amatolicum Hilliard & Burtt Gi 311; Ph 933, 953, 1000; Ph &

Hu 20, 118; Robinson 1062.

baurianum Knuth Br 105, 111; Fu & Ph 70; GR 3486; Gi 21;

Ph 422, 999.

caffrum Eckl. & Zeyh. Gi 1133.

contortum Eckl. & Zeyh. Br 148; Gi 226.

discolor Hilliard & Burtt Fu & Ph 54, 228; GR 3505; Gi 193,

1112, 1306; Ph 976.

harveyi fln'q. Phll44.

molle L. Ph & Hu 133.

multisectum N.E. Br. Ph 799; Ph & Hu 146.

ornithopodon Eckl. & Zeyh. Gi 505, 918, 1094; Ph 932.

schlechteri Knuth Fu 694.

wakkerstroomianum Knuth Gi303, 1309; Ph 1007.

3925 Monsonia emarginata (L. f.) L 'Herit. Gi 301, 1143.

3927 Erodium cicutarium (L.) L ’Herit. ex Ait. Ph & Hu 134.

3928 Pelargonium

alchemilloides (L.) L’Herit. Gi 95; Ph 143, 1068.

althaeoides (L.) L ’Herit. Ph & Hu 125.

caffrum (Eckl. & Zeyh.) Harv. Ph & Hu 137.

cordifolium (Cav.) Curt. Fu & Ph 224; Gi308, 1121; Ph 419;

Robinson 1060.

grossularioides (L.) L ’Herit. Gi 302; Ph 10 73.

iocastum (Eckl. & Zeyh.) Steud. Gi 1183, 1329, s.n.; Ph & Hu

114.

multicaule Jacq. subsp. multicaule Gi 1136; Ph 971.

peltatum (L.) L ’Herit. Gi 139.

ranunculophyllum (Eckl. & Zeyh.) Bak. Ph 1476, 1540.

reniforme Curtis Ph 550.

schizopetalum Sweet Gi 1119; Ph & Hu 57.

sidifolium (Thunb.) Knuth Ph & Hu 142.

zonale (L.) L’Herit. Gi310, 867, 1100; Ph 151, 392; Tu 204.

OXALIDACEAE

3936 Oxalis

bifurca Lodd. var. bifurca Ph & Hu 151.

comiculata L. Ph 924.

semiloba Sond. Br 125; Fu & Ph 341; GR 3524; Ph 1029, 1228

(double flowers), 1275; Ph & Hu 70.

smithiana Eckl. & Zeyh. Br 88a; GR 3422; Ph 1064; Ph & Hu

37.

tragopoda Salter Gi 1460.

LINACEAE

3943 Linum

bienne Mill. Fu & Ph 1 83.

thunbergii Eckl. & Zeyh. Br 162, 184; Fu & Ph 116, 142; Gi

241, 300, 1074, 1234.

RUTACEAE

3991 Xanthoxylum

capense (Thunb.) Harv. Fu & Ph 288; Gi 1060.

davyi (Verdoom) Waterm. Gi 1439; Ph 900a, 1462.

4035 Calodendrum capense (L. f.) Thunb. Ph 1048.

4038 Agathosma ovata (Thunb.) Pillans Ph 895.

4076 Vepris lanceolata (Lam.) G. Don Fu s.n.; Gi 895.

4091 Clausena anisata (Willd.) Hook. f. ex Benth. Gi 71.

BURSERACEAE

4151 Commiphora woodii Engl. Pefferskop, Acocks 11900 (Van

der Walt 1973: 85).

PTAEROXYLACEAE

4157 Ptaeroxylon obliquum (Thunb.) Radik. Ph 1117.

MEL1ACEAE

4193 Ekebergia capensis Sparrm. Ph 1119.

POLYGALACEAE

4273 Polygala

confusa Macowan Gi 291; Ph 107.

fruticosa Berg. Ph 967.

hispida Burch. Fu & Ph 114; Gi 242, 287, 1088, 1304; Ph

393; Ph & Hu 22.

hottentotta Presl Br 1 73; Fu et al. 13; Fu & Ph 115.

myrtifolia L. Gi 1565; Ph 894.

ohlendorfiana Eckl. & Zeyh. Br 151; Fu & Ph 55; Gi g718; Ph

391, 986.

refracta DC. Ph 1263.

uncinata E. Mey. ex Meisn. Fu & Ph 89.

virgata Thunb. Ph 582.

4278 Muraltia

alticola Schltr. Ph 439.

macroceras DC. GR 3492; Gi 195, 247, 1124, s.n.; Ph 414.

saxicola Chod. Ph 436.

4279 Nylandtia spinosa (L.) Dumort. Ph 1126.

EUPHORB1ACEAE

4299 Phyllanthus incurvus Thunb. Ph 1232.

4348 Croton rivularis E. Mey. Gi 1468; Ph 835.

4370 Adenocline

acuta (Thunb.) Baill. Gi 1443, 1535.

pauciflora Turcz. Fu & Ph 359; Gi 232, g748.

4372 Leidesia obtusa (Thunb.) Muell. Arg. Gi 1546.

4407 Acalypha

ecklonii Baill. GR 301 0; Ph 1183.

peduncularis E. Mey. ex Meisn. Fu & Ph 230, 352; Gi 992,

1024, 1028, 1154,1181, 1264.

4416a Ctenomeria capensis (Thunb.) Harv. ex Sond. Gi 164,

883, 886, 976, 1506.

4424 Ricinus communis L. Ph 1049.

4448 Clutia

af finis Sond. GR 3822; Gi 41, 70, 412, 768, g886; Mahlobo

41; Ph 150.

alatemoides L. (var. not determined) Fu & Ph 345; Gi 416,

769; Ph 407; Tu 208.

discep tata Train Gi 116 9.

heterophylla Thunb. Fu & Ph 296; Gi 1111; Ph 1201; Ph &

Hu 77.

hirsuta E. Mey. ex Sond. var. hirsuta Ph 962.

katharinae Pax Fu & Ph 39; Ph 558.

natalensis Bemh. ex Krauss Tu 168, 169, 1 71.

pulchella L. var. pulchella Fu & Ph 268; Ph 503; Tu 255.

4498 Euphorbia

epicyparissias E. Mey. ex Bioss. var. epicyparissias Fu 35; Gi

40, 765, 766; Ph 409; Tu 231.

kraussiana Bemh. (var. not determined) Gi 764; Mahlobo 42.

pulvinata Marloth. Seen among rocks on Gaika’s Kop and

neighbouring peaks.

sclerophylla Boiss. Fu & Ph 61; GR 3429.

striata Thunb. Fu & Ph 109.

4498b Chamaesyce inaequilatera (Sond.) Sojak Ph 1047.

ANACARDIACEAE

4562 Harpephyllum caffrum Bemh. Fu s.n.; Tu 188.

4594 Rhus

chirindensis Bak. f. Fu s. n. ; Fu & Ph 252; Gi 579, 994, 1 059.

dentata Thunb. Fu & Ph 284, 294; Gi g882, 991; Ph 548; Tu

141.

dentata Thunb. x R. divaricata Eckl. & Zeyh. Tu 136.

discolor E. Mey. ex Sond. Gi 1308; Ph 545.

fastigiata Eckl. & Zeyh. Fu & Ph 311; Ph 244.

fastigiata Eckl. & Zeyh. x R. rehmanniana Engl. Fu & Ph 310;

Gig884.

incisa L. f. var. effusa (Presl) R. Fernandes Ph 794.

krebsiana Presl ex Engl. Tu 270 (possibly crossed with R. di-

varicata Eckl. & Zeyh.).

250

Bothalia 17,2 (1987)

pallens Eckl. & Zeyh. forma pallens Robinson 1071.

pyroides Burch, vai. gracilis (Engl.) Burt t Davy Gi 1241, 1294,

1305, 1349.

pyroides Burch, var. pyroides Gi 323, 324; Ph 552; Russell

2339; Tu 117, 120, 121, 129.

rehmanniana Engl. Fu & Ph 308; Gi g883.

tomentosaZ,. Ph 333; Robinson 1070, 1501, s.n.; Tu 239.

AQUIFOLIACEAE

4614 Ilex mitis (L.) Radik. Fu s.n.; Gi 136.

CELASTRACEAE

4626 Maytenus

acuminata (L. f.) Loes var. acuminata Ph 761, 777, 868; Tu

210.

heterophylla (Eckl. & Zeyh.) N.K.B. Robson Fu & Ph 264; Ph

785, 795, 873; Tu 202.

nemorosa (Eckl. & Zeyh.) Marais Gi 496, 535, g851, 892; Ph

781, 881, 887; Tu 213.

peduncularis (Sond.) Loes. Ph 872, 882; Tu 216.

undata (Thunb.) Blakelock Ph 764.

4628 Putterlickia verrucosa (E. Mey. ex Sond.) Szyszyl. Gi

1102; Ph 896.

4630 Pterocelastrus tricuspidatus (Lam.) Sond. Gi 316.

4641 Cassine

aethiopica Thunb. Fu s.n.; Ph 901.

papillosa ( Hochst .) Kuntze Fu s.n.; Fu & Ph 265; Gi 773; Ph

883, 900b, 1125.

tetragona (L. f.) Loes Ph 871.

ICACINACEAE

4671 Cassinopsis ilicifolia (Hochst.) Kuntze Fu s.n.; Ph 1124,

1318.

4686 Apodytes dimidiata E. Mey. ex Am. subsp. dimidiata Tu

205.

SAPINDACEAE

4734 Allophylus decipiens (Sond.) Radik. Fu s.n.; Fu & Ph 271;

Ph 779; Tu 184.

4836 Hippobromus pauciflorus (L. f.) Radik. Gi 533, 1036; Tu

178.

MELIANTHACEAE

4854 Melianthus

dregeanus Sond. subsp. dregeanus Gi g841, 1526; Ph 415; Tu

245.

major L. Gi s.n.

BALSAMINACEAE

4856 Impatiens hochstetteri Warb. subsp. hochstetteri GR 3128;

Gi 312; Mgudlwa 25; Ph 162.

RHAMNACEAE

4861 Ziziphus mucronata Willd. subsp. mucronata Fu & Ph 291;

Ph 104.

4874 Scutia myrtina (Burm. f.) Kurz Fu s.n.; Fu & Ph 295; Gi

1063; Mahlobo 12; Tu 203.

4875 Rhamnus prinoides L’Herit. Fu s.n.; Gi 68, 251, 1520; Ph

553.

4880 Noltia africana (L.) Reichb. f. Fu & Ph 285; Gi 1585; Ph

589.

4886 Phylica galpinii Pillans Gi 276; Ph 566.

4905 Helinus integrifolius (Lam.) Kuntze Fu & Ph 254; Gi

1084; Ph 796.

VITACEAE

4917 Rhoicissus

digitata (L. f.) Gilg & Brandt Tu 189.

microphylla (Turcz.) Gilg & Brandt Ph 554, 1342; Tu 137,

142, 149.

revoilii Planch. Gi g881; Ph 763.

tridentata (L. f.) Wild & Drum, subsp. cuneifolia (Eckl. &

Zeyh.) N.R. Urton Fu & Ph 258; Gi 573.

4918a Cyphostemma cirrhosa (Thunb.) Desc. Gi 578, 1067; Ph

147.

T1LIACEAE

4957 Sparrmannia ricinocarpa (Eckl. & Zeyh.) Kuntze Gi 271.

4966 Grewia occidentals L. Fu & Ph 274; Tu 1 91.

MALVACEAE

4983 Abutilon sonneratianum (Cav.) Sweet Ph 159.

4986 a Anisodontea scabrosa (L.) Bates Ph 511.

4998 Sida

rh om bifolia L. Ph 1033.

temata L. f. Gi 1541; Russell 2356.

5007 Pavonia columella Cav. Gi g844; Ph 1042.

5013 Hibiscus

aethiopicus L. var. aethiopicus Gi 1 083.

trionum L. Gi 1086; Ph 386.

STERCULIACEAE

5056 Hermannia

erodioides (Burch, ex DC.) Kuntze Ph 820, 914.

geniculata Eckl. & Zeyh. Ph 1475.

velutina DC. Gi 29 7; Ph 590; Tu 186.

OCHNACEAE

5112 Ochna serrulata (Hochst.) Walp. Fu s.n.; Fu & Ph 262; GR

3820; Gi 314; Ph 875.

CLUSIACEAE

5168 Hypericum

aethiopicum Thunb. subsp. aethiopicum GR 3428.

lalandii Choisy Br 27; Fu & Ph 143; Gi 299, 731, 1077.

natalense Wood & Evans Gi 1139.

VIOLACEAE

5271 Hybanthus capensis (Thunb.) Engl. Ph 1523.

5274 Viola arvensis Murray Ph 984; Ph & Hu 23.

FLACOURTIACEAE

5296 Kiggelaria africana L. Gi 285; Tu 131, 206.

5304 Scolopia

mundii (Eckl. & Zeyh.) Warb. Fu s.n.; Gi 891; Ph 760, 784.

zeyheri (Nees) Harv. Fu s.n.; Tu 268.

5315 Trimeria

grandifolia (Hochst.) Warb. Fu & Ph 260; Gi s.n.

trinervis Harv. Gi 1159; Ph 772, 798; Tu 185.

5328 Dovyalis

lucidaSim, Hogsback, Story 369 (Langenegger 1976: 90).

zeyheri (Sond.) Warb. Ph 789; Tu 180.

ACHARLACEAE

5374 Ceratiosicyos laevis (Thunb.) A. Meeuse Gi 1022, 1441,

1654; Ph 502.

5375 Acharia tragodes Thunb. Ph 1197.

CACTACEAE

5417 Opuntia spp. (seen occasionally in disturbed places).

THYMELAEACEAE

5435 Gnidia

baurii C.H. Wr. GR 3440; Ph 563.

nodiflora Meisn. Gi 1109; Ph 389.

phaeotricha Gilg Gi 248; Ph & Hu 63, 95.

pulchella Meisn. Fu & Ph 286; Gi 250, 485; Ph 243; Tu 23 7.

sericea L. var. sericea Fu 6 77; Gi 415, 486; McGillivray 23; Ph

341, 405.

5436 Struthiola parviflora Bartl. ex Meisn. Ph 1202.

5438 Englerodaphne pilosa Burtt Davy Fu 58; Gi 208, 559.

5461 Passerina

filiform is L. Ph 1200.

montana Thoday Ph 1140; Tu 1 70.

vulgaris Thoday Gi s.n.; Tu 21 7.

Bothalia 17,2 (1987)

251

sp. Between the Wolf and Mnyameni Rivers (Story 1952: 99).

5465 Dais cotinifolia L. Ph 736.

LYTHRACEAE

5476 Lythrum hyssopifolium L. Gi 1346.

MYRTACEAE

5578 Eugenia zeyheri Harv. Fu&Ph251.

5598 Eucalyptus sp. (seen naturalized, mainly near plantations).

ONAGRACEAE

5795 Epilobium capense Buck, ex Hochst. Br 23, 160, 169,

1 78; Fu et al. 15; Fu & Ph 226; Gi 221, 11 78.

Epilobium

hirsutum L. Gi 975; Ph 1045.

tetragonum L. subsp. tetragonum Br 133, 1 75; Fu 71 7.

5804 Oenothera rosea/l/i. Ph 397, 925.

HALORAGACEAE

5833 Laurembergia repens Berg, (intermediate between sub-

species, see Obermeyer 1973) Fu & Ph 6; Ph 1325.

5836 Gunnera perpensa L. Gi 1 042, 1 244; Ph 396.

ARALIACEAE

5872 Cussonia

paniculata Eckl. & Zeyh. Gi 1161; Ph & Hu 66; Tu 140, 148.

spicata Thunb. Mgobozi 21; Tu 1 76.

APIACEAE

5894 Centella

asiatica (L.) Urb. GR 3404; Gi 362, 469, g887; Ph 336.

glabrata L. var. natalensis Adamson GR 3472; Gi 363, 1237.

5918 Sanicula elata Buch.-Ham. GR 3811; Gi 141, 366, s.n.

5922 Alepidea

acutidens Weim., Nyameni Location, near KeiskamaHoek, 3 500

ft, Schonland 4461 (GRA, PRE) (Weimarck 1949: 245).

amatymbica Eckl. & Zeyh. var. amatymbica GR 3502.

amatymbica Eckl. & Zeyh. var. aquatica (Kuntze) Weim. Fu et

al. 36; GR 224.

capensis (Berg.) R.A. Dyer var. capensis GR 3443, 3488a; Gi

506; Ph 1463, 1468.

macowanii Duemmer Ph 1269.

pilifera Weim. Fu & Ph 149, 349; Ph & Hu 52.

senataEckl.&Zeyh. var. cathcartensis (Kuntze) Weim. Gi 1260.

seriata Eckl. & Zeyh. var. serrata Gi 220, 1327; Ph & Hu 74.

5970 Conium

chaerophylloides (Thunb.) Sond. Fu & Ph 346.

fontanum Hilliard & Burtt var. alticola Hilliard & Burtt Ph

1001.

fontanum Hilliard & Burtt var. fontanum, Thomas Mtn, Hilliard

& Burtt 14798 (E, NU) (Hilliard & Burtt 1985: 472).

sp. Hogsback Forest Reserve, Hilliard & Burtt 10992 (E, NU)

(Hilliard & Burtt 1985: 472).

5990 Lichtensteinia interrupta (Thunb.) E. Mey. Ph 1186.

5992 Heteromorpha arborescens (Spreng.) Cham. & Schlechtd.

Tu 267.

5994 Bupleurum mundtii Cham. & Schlechtd. GR 3508; Ph

1014; Ph & Hu 44 (KEI).

6004a Ciclospermum leptophyllum (Pers.) Sprague Ph 1032.

6017a Sonderina humilis (Meisn.) Wolff Ph 905, 1033.

6033 Pimpinella caffra (Eckl. <6 Zeyh.) D. Dietr. Ph 825, 1061,

1283.

6038 Sium repandum Welw. ex Hiem Fu & Ph 195.

6045 Polemannia

grossulariifolia Eckl. & Zeyh. Gi 1296, s.n.; Tu 153.

montana Schltr. & Wolff, Gaika’s Kop, Hilliard & Burtt 18805

(E, NU) (Hilliard & Burtt unpublished).

6078 Annesorrhiza schlechteri Wolff Fu 20; Fu & Ph 25, 106.

6078b Stenosemis angustifolia Sond. Ph 1021.

6116 Peucedanum

caffrum (Meisn.) Phill. Ph & Hu 43.

capense (Thunb.) Sond. var. lanceolatum (E. Mey. ex Meisn.)

Sond. Gi 69, 368; Ph & Hu 30; Tu 238.

6142 Daucus carota L. Tu 100.

CORNACEAE

6156 Curtisia dentata (Burnt, f.) C.A. Sm. Gi g849.

ERICACEAE

6237 Erica

alopecurus Harv. var. alopecurus Gi s.n.; Ph 437, 570.

brownleeae H. Bol. Fu & Ph 20, 216, 241; GR 3496a; Gi 25,

1321; Ph & Hu 111.

caespitosa Hilliard & Burtt Fu & Ph 112, 239; Ph & Hu 12; Ph

1256.

caffra L. Gi 698a, s.n.

caffrorum H. Bol. var. caffrorum Ph 435, 441; Ph & Hu 8.

frigida H. Bol. Ph & Hu 6.

leucopelta Tausch. var. leucopelta Gi s.n.; Ph & Hu 147; Tu

251.

maesta H. Bol. Fu & Ph 40, 41; Gi 22; Ph & Hu 33.

woodii H. Bol. subsp. woodii Ph 1220.

woodii H. Bol. subsp. platyura Hilliard & Burtt, Hogsback, Rat-

tray 216 (PRE) (Hilliard & Burtt 1986).

sp. Ph 1142.

6241 Ericinella multiflora Klotzsch Fu & Ph 221; Gi 1320; Ph

& Hu 110; Tu 248, 252.

MYRSINACEAE

6283 Maesa alnifolia Harv. Fu & Ph 261; Gi 65, 401, 531, 937.

6313 Myrsine africana L. GR 3021; Gi 146, 537; Ph 342, 424;

Tu 139, 226.

6314 Rapanea melanophloeos (L.) Mez Fu s.n.; Gi 74; Tu 199.

PRIMULACEAE

6330 Lysimachia nutans Nees Gi 231, 771, 1135; Ph & Hu 11 7.

6338 Anagallis huttonii Harv. Br 1 77; Fu 690, 720; Fu 04; Fu &

Ph 24.

PLUMBAGINACEAE

6343 Plumbago auriculata Lam. Ph 102.

SAPOTACEAE

6386 Mimusops obovata Sond. Fu & Ph 259; Ph 880.

EBENACEAE

6404 Euclea

crispa (Thunb.) Guerke var. crispa Fu & Ph 289; Ph 773; Tu

187, 232, 233.

schimperi A DC. var. schimperi Ph 1120.

undulata Thunb. var. undulata, Elandsberg (Story 1952: 93).

6406 Diospyros

austro-africana De Winter var. microphylla (Burch.) De Winter

Ph 547; Tu 143, 144, 147.

dichrophylla (Gand.) De Winter Gi 1528; Tu 262.

lycioides Desf. subsp. sericea (Bernh.) De Winter Fu & Ph 292.

scabrida (Harv. ex Hiem) De Winter var. cordata (E. Mey. ex A.

DC.) De Winter Ph 889.

simii (Kuntze) De Winter Ph 776, 790.

villosa L. var. villosa Fu & Ph 253; Gi 978; Tu 201.

whyteana (Hiem) F. White Ph 1116; Tu 200.

OLEACEAE

6428 Chionanthus foveolata (E. Mey.) Steam subsp. foveolata

Fu s.n.; Gi g842.

6434 Olea

capensis L. subsp. macrocarpa (C.H. Wright) Verdoorn Fu s.n.

europaeaZ,. subsp. africana (Mill.) P.S. Green Tu 271.

LOGANIACEAE

6473 Buddleja

auriculata Benth. Ph 828, 831.

dysophylla (Benth.) Radik. Gi 413; Tu 1 75.

252

Bothalia 17,2 (1987)

saligna Willd. Tu 1 73.

salvifolia (L.) Lam. Fu & Ph 46, 231; Gi 475, 536, 705; Tu

127, 128,172.

GENTIANACEAE

6481 Sebaea

hymenosepala Gilg Fu & Ph 94; Gi 1141, 3504.

longicaulis Schinz Gi 1021, s.n.; Ph 141, 433.

macrophylla Gilg Gi 4 70, s.n.; McGillivray 30.

repens Schinz Br 78; Ph 438.

sedoides Gilg var. confertiflora (Schinz) Marais Fu & Ph 1 7; Ph

1271.

thorn asii S. Moore Fu & Ph 48.

6503 Chironia krebsii Griseb. Fu & Ph 11, 92; Gi 1116; GR

3515a.

APOCYNACEAE

6559 Carissa

bispinosa (L.) Desf. var. acuminata (E. Mey.) Codd Gi 96, 930;

Ph 111.

bispinosa (L.) Desf. var. bispinosa Fu & Ph 257.

6688 Strophanthus speciosus (Ward & Harv.) Reber Fu & Ph

270; Ph 829, 1154.

ASCLEPIADACEAE

6777 Xysmalobium

involucratum Decne. Ph 960; Ph & Hu 61.

orbicular eD.Dietr. Ph 1160.

parviflorum Harv. ex Scott Elliott Ph 12 74; Ph & Hu 4 7.

prunelloides Turcz. Fu & Ph 355; Ph 977; Ph & Hu 97.

stockenstromense Scott Elliott Ph & Hu 152.

undulatum (L.) Ait. f. Ph 1195.

6778 Schizoglossum

atropurpureum E. Mey. subsp. tridentatum (Schltr.) Kupicha

Gi 243; Ph 1307; Ph & Hu 102.

cordifolium E. Mey. Ph & Hu 140.

hamatum E.Mey., Hogsback Mtn, Rattray 269 (BOL); Rattray

15764 (BOL) (Kupicha 1984: 605).

6778a Aspidoglossum gracile (E. Mey.) Kupicha Ph 1262.

6779 Fanninia caloglossa Harv. GR 3484a; Ph 1157.

6181a Pachycarpus concolor E. Mey. Ph 1231.

6791 Asclepias

diploglossa (Turcz.) Druce Gi s.n.; Ph 998; Ph & Hu 148.

fru ticosa L. Gi 1 038.

gibba (E. Mey.) Schltr. Gi 1156; Ph 775.

peltigera (E. Mey.) Schltr. Ph 1168.

physocarpa (E. Mey.) Schltr. Gr 3009; Gi 1057, 1078; Ph 958.

stellifera Schltr. Ph 1133.

6834 Cynanchum ellipticum (Harv.) R.A. Dyer Fu & Ph 269b;

Gi 525.

6849 Sarcostemma viminale (L.) R. Br. Ph 1495.

6860 Secamone

alpinii Schultes Gi g839.

filiformis (L. f.) J.H. Ross Fu & Ph 269; Gi 1097.

6861 Sisyranthus barbatus (Turcz.) N.E. Br. GR 3480; Ph & Hu

99.

6868 Anisotoma cordifolia Fenzl Gi s.n.; Ph & Hu 85.

6875 Riocreuxia torulosa Decne. Gi 228, 1134, 1183a.

6899 Tylophora

cordata (Thunb.) Druce Gi 1104.

flanaganii Schltr. Phlll8.

CONVOLVULACEAE

6968 Cuscuta

appendiculata Engelm. Gi 225.

cassytoides Nees ex Engelm. Gi s.n.

6993 Convolvulus farinosus L. Gi 1080; Ph 1308.

7003 Ipomoea purpurea (L.) Roth Ph 1330.

BORAGINACEAE

7043 Ehretia rigida (Thunb.) Druce Tu 1 79.

7064 Cynoglossum

enerve Turcz. Ph 1066, 1185.

hispidum Thunb. Gi 1027.

lanceolatum Forssk. Hughes and Mjwara 32; Ph 1492.

spelaeum Hilliard & Burtt Gi 722a; Ph 1 294.

7100 Myosotis

sylvatica Hoffm. Fu <& Ph 184.

semiamplexicaulis DC. Ph 1004.

7109 Lithospermum papillosum Thunb. Gi 1144, 1170; Ph 92.

7118 Echium sp. Seen as a weed in ploughed land.

VERBENACEAE

7138 Verbena

bonariensis L. Ph 1 031; Russell 2348.

venosa Gill. & Hook. Ph 91.

7153 Priva meyeri Jaub. & Spach var. meyeri Ph 1236.

LABIATAE

7211 Ajuga ophrydis Burch, ex Benth. Fu {£ Ph 315; GR 3419.

7212 Teucrium trifidum Retz. Ph 1235.

7238 Marrubium vulgare L. Ph 1303.

7264 Leonotis

leonurus (L.) R. Br. var. leonurus Fu & Ph 290; Gi 908, 1518;

McGillivray 34; Tu 265.

ocymifolia (Burm. f.) Iwarsson var. ocymifolia Fu & Ph 297;

Ph 1214.

ocymifolia (Burm.f.) Iwarsson var. raineriana (Visiani) Iwarsson

Gi 1438, 1519; Mahlobo 20, Ph 1041.

7281 Stachys

aethiopica L. Ph 1100, 1544; Ph & Hu 169.

caffra E. Mey. ex Benth. Gi s.n.

flexuosa Skan GR 351 7;Ph 1478; Ph & Hu 11, 161.

graciliflora Presl Gi 574, g808, 1475, 1547; Ph 108, 1472.

grandiflora E. Mey. ex Benth. Ph 934, 1543, 1546; Ph & Hu

160.

humifusa Burch, ex Benth. Ph 1326.

malacophylla Skan Ph 1479.

sp. 1 Ph 963, 1234, 1464.

sp. 2 Ph 1547.

sp. 3 Ph 1002, 1259; Ph & Hu 41, 159.

7290 Salvia

aurita Thunb. var. aurita, Tyumie Berg, Ecklon (Hedge 1974:

67).

aurita Thunb. var. galpinii (Skan) Hedge Gi g896, 14 76.

repens Burch, ex Benth. var. repens Gi 1140, 1511; Ph 97,

160; Ph & Hu 103.

verbenaca L. Ph 1129.

7328 Mentha

aquatica L. Br 1 74; Fu 739, 742; Gi 906.

longifolia (L.) Huds. subsp. capensis (Thunb.) Briq. Fu & Ph

76.

7350 Plectranthus

ambiguus (H. Bol.) Codd Ph 1250.