Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 17,1 July/Julie 1987

BOTANICAL RESEARCH INSTITUTE • DEPARTMENT OF AGRICULTURE AND WATER SUPPLY • REPUBLIC OF SOU ,FWCA

NAVORSINGSINSTITUUT VIR PLANTKUNDE • DEPARTEMENT VAN LANDBOU EN WATERVOO RSIENING • REPl'BLiEK VAN >AFPIKA

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. Two parts of the

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

ed 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 virPlantkundeisgewy

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 indeks van die

inhoud, outeurs en onderwerpe verskyn jaarliks.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The memoirs are individual treatises usually of an ecological

nature, but sometimes dealing with taxonomy or economic

botany.

’n Reeks van losstaande omvattende verhandelings oor vemaam-

lik ekologiese, maar soms ook taksonomiese of plantekonomiese

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 tire 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 fotografiese plate waarop fossiele populasies

afgebeeld word. Reeds beskikbaar:

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

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

Obtainable from/Beskikbaar van: A. A. Balkema Marketing, Box/Posbus 3 1 7, Claremont 7735, RSA.

Republic of

South Africa

Republiek van

Suid-Afrika

BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 17,1

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 000 1 , South Africa

Uitgegee deurdie Navorsingsinstituut vir Plantkunde, Departement van Landbou en Watervoorsiening,

Privaatsak X 1 0 1 , Pretoria 0001 , Suid-Afrika

1987

Digitized by the Internet Archive

in 2016

https ://arch i ve . o rg/detai Is/bothal iavo Iume1717unse

CONTENTS — INHOUD

Volume 17,1

1 . The Indigoferafilifolia complex (Fabaceae) in southern Africa. J. K. JAR VIE and C. H. STIRTON 1

2. A synopsis of Tephrosia subgenus Barbistyla (Fabaceae) in southern Africa. B . D. SCHRIRE 7

3. A new species and a new combination of Asclepias (Asclepiadaceae) in southern Africa. A. NICHOLAS ... 17

4. Two new brown subcrustose Parmelia species from southern Africa (lichenized Ascomycetes).

F. BRUSSE 25

5 . Notes on African plants:

Asclepiadaceae. Notes on Asclepias diploglossa, A. cognata and A.flava. A. NICHOLAS 29

Erysiphaceae. Two new Oidium species from the Transvaal. G. J. M. A. GORTER and A. EICKER... 32

Fabaceae. A new species of Indigofera from Natal and Transkei. G. GERMISHUIZEN 33

Lichinaceae. A new species of Gonohymenia from Etosha Pan limestone. F. BRUSSE 35

Lichinaceae. A new species of Thyrea from Otavi dolomite (Damara System). F. BRUSSE 37

Polygonaceae. A new species of Oxygonum from Natal. G. GERMISHUIZEN 40

Teloschistaceae. A new species of Caloplaca from southern Africa. I. KARNEFELT 41

6. Cetraria (Parmeliaceae) and some related genera on the African continent. I. KARNEFELT 45

7. Species groups in the genus Ehrharta (Poaceae) in southern Africa. G. E. GIBBS RUSSELL and

R. P. ELLIS 51

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

RUSSELL 67

9. Leaf anatomy of the genus Ehrharta (Poaceae) in southern Africa: the Setacea group. R. P. ELLIS 75

10. Six cultivars of Solarium macrocarpon (Solanaceae) in Ghana. Z. R. BUKENYA and J. B. HALL 91

11. Bibliography of Fusarium (Fungi: Hyphomycetes) in South Africa, 1945-1985. W. F. O. MARASAS,

SANDRA C. LAMPRECHT, P. S. VAN WYK and R. Y. ANELICH 97

12. The genus Rubus (Rosaceae) in South Africa. IV. Natural hybridization. J. J. SPIES, C. H. STIRTON and

H. DUPLESSIS 105

13. Ordination by detrended correspondence analysis (DC A) of the vegetation of Swartboschkloof,

Jonkershoek, Cape Province. D. J. MCDONALD 121

14. Miscellaneous notes:

Chromosome studies on African plants. 3. J. J. SPIES and H. DU PLESSIS 131

Chromosome studies on African plants. 4. J. J. SPIES and A. JONKER 135

Performance of a laboratory-constructed anemometer under summer field conditions on a Mountain

Fynbos experimental site. G. W. DAVIS 136

1 5 . Review of the work of the Botanical Research Institute ,1985/1986 139

16. Book reviews 155

1 7 . Guide for authors to Bothalia 157

Bothalia 17,1: 1-6(1987)

The Indigo fera fili folia complex (Fabaceae) in southern Africa

J. K. JAR VIE* and C. H. STIRTON**

Keywords: Fabaceae, Indigoferafilifolia complex, new species, southern Africa, taxonomy

ABSTRACT

The Indigoferafilifolia Thunb. complex, previously known as a widespread polymorphic species, is segregated into

three species. /. filifolia is retained as a variable species and two allied species, /. gifbergensis C. H. Stirton & J. K. Jarvie

and /. ionii J. K. Jarvie & C. H. Stirton are described as new.

UITTREKSEL

Die Indigofera filifolia Thunb. -kompleks, voorheen bekend as ’n wydverspreide polimorfiese spesie, word in drie

spesies verdeel. /. filifolia word as ’n veelvormige spesie behou en twee verwante spesies, /. gifbergensis C. H. Stirton &

J. K. Jarvie en /. ionii J. K. Jarvie & C. H. Stirton word as nuut beskryf.

INTRODUCTION

The pantropical genus Indigofera comprises some 800

species. It forms a large and distinctive part of the tribe

Indigofereae which also contains two closely related

genera Cyamopsis and Rhynchotropis along with an

anomalous Madagascan genus Phylloxyon (Polhill

1981).

Indigofera was once the main source of indigo dye but

its importance dwindled as cheaper synthetic analogues

were developed to replace the plant source of the dye.

There is, however, a renewed interest in the genus and it

has begun to be of some value as a source of forage

legume (De Kort & Thijsse 1984).

The last major revision of Indigofera in Africa was by

Gillett (1958), who recognized 278 species. He divided

the African representatives into 5 subgenera, 5 sections

and 19 subsections. As this revision was a precursor to a

revision of the genus for the Flora of Tropical East

Africa he treated only 70 species in any detail. The work

is essentially regional and ignores to a considerable ex-

tent areas such as southern Africa.

Our overall knowledge of southern African species of

Indigofera is still dependent on the 1862 revision by

William Harvey. This account was subsequently added

to and modified by the following workers: Baker (1905),

Brown (1925, 1926), Dinter (1932), Fourcade (1932),

Bremekamp (1932), Guthrie, Pillans & Salter (1939),

Dyer (1944), Suessenguth & Merxmiiller (1951), Merx-

miiller & Schreiber (1957), Schreiber (1970) and Stirton

(1982).

The present investigation focuses on the widespread

polymorphic species I. filifolia in South Africa. Gillett

(1958) placed I. filifolia Thunb. as the sole member of

his section Juncifolieae. His diagnosis is limited to the

presence of petioles many times longer than the small

terminal leaflet, persistent when the latter falls off; fruit-

ing pedicels erect and spreading; and endocarp spotted

with seeds ± separated.

We are indebted to Mr Ion Williams for drawing our

attention to a small aphyllous plant which did not appear

to fit the description of I. filifolia. He felt certain that it

represented a new species. We confirm this and name it

I. ionii in his honour. As part of our study we also looked

at all the available South African aphyllous specimens of

Indigofera that seemed pertinent to the I. filifolia group.

We conclude that what had been known as I. filifolia

Thunb. in herbaria should be separated into three spec-

ies: I. filifolia Thunb., I. ionii J. K. Jarvie & C. H.

Stirton, and I. gifbergensis C. H. Stirton & J. K. Jarvie.

KEY TO SPECIES

la Leaflets 3-5, terminal leaflet present, narrowly obovate to obovate, base obtuse; petiole 14-25 mm long; stipules

narrowly triangular; flowers 5-6(7) mm long:

2a Dense, erect shrub up to lm tall; leaflets recurved mucronate, obovate, biramous hairs present on lower sur-

face but restricted to the margin; inflorescence 15-30-flowered; flowers brick-red; calyx-teeth finger-like,

sinus between vexillar teeth narrow; standard broadly elliptic; peg on outer surface of keel blade pointing

downwards; scales absent from base of anthers; upper parts of Filaments uniformly thickened; seeds dark

chestnut or khaki and covered in black blotches and stripes, elliptic 1. gifbergensis

2b Small lax wiry herb, sparsely branched; leaflets with straight mucro, narrowly obovate, biramous hairs

sparsely but evenly distributed over lower surface; inflorescence 6-10-flowered; flowers pink; calyx teeth tri-

angular, sinus between vexillar teeth broad; standard very broadly ovate; peg on outer surface of keel blade

pointing upwards; scales present at base of anthers; upper parts of filaments alternately thick and thin; seeds

olive-green, oblong 2. ionii

lb Leaflets 6-8, upper leaflets absent or reduced to scales, elliptic, base cuneate; petiole 40-60 mm long; stipules

filiform; flowers (7,5) 8,0-10,5 mm long 3. filifolia

* Department of Botany, Plant Sciences Laboratories, University of

Reading, Whiteknights, Reading RG6 2AS, England; currently at Pus- ** B. A. Krukoff Botanist for Legume Research, The Herbarium,

litbangtri. Project ATA-71, Jl. Cimmanggu No. 1, Bogor, Java, Indo- Royal Botanical Gardens, Kew, Richmond, Surrey, TW9 3AE, Eng-

nesia. land, UK.

2

Bothalia 17,1 (1987)

QUICK-KEY CHARACTERS

1 . Flowers brick-red /. gifbergensis

Flowers pink /. ionii, /. filifolia

2. Leaflets with straight mucro I. ionii, [.filifolia

Leaflets recurved-mucronate I. gifbergensis

3. Anthers tufted; short filaments thicker than long filaments

I. ionii, I. filifolia

Anthers without tufts, filaments uniformly thin ... /. gifbergensis

4. Peg on outer surface of keel blade pointing downwards ... /. ionii

Peg on outer surface of keel blades pointing upwards

/. gifbergensis, I. filifolia

5. Fruits compressed I. ionii

Fruits terete I. gifbergensis, I. filifolia

6. Seeds dark chestnut or khaki, blotched I . gifbergensis

Seeds olive-green I. ionii

Seeds chocolate-brown I. filifolia

1 . Indigofera gifbergensis C. H. Stirton & J. K.

Jarvie, sp. nov. subsectioni Juncifolieae Gillettii prox-

ima.

Frutex erectus lignosus ad 1 m altus, valde ramosus,

caulibus veterioribus glabris, junioribus appresso-strigu-

losis glabrescentibus. Stipulae conferruminatae, basi

petioli adnatae, ad 4 mm longae, anguste triangulares.

Folia regularia, imparipinnata. Foliola obovata, 6-8 x

3.0- 3 ,5 mm, basi obtusa, recurvato-mucronata. Inflo-

rescentiae axillares, racemosae, laxae, 15-30-florae,

sanguineae. Flores 5-6(7) mm longi, unoquoque bractea

caduca anguste triangulari 2, 0-2, 3 mm longa subtenti.

Tubus calycis 1,5 mm longus; lobi calycis omnes

aequales triangulares. Vexillum late ellipticum, 5-6 x

4. 0- 4, 5 mm. Petala alarum 5, 5-6,0 x 2, 0-2, 5 mm.

Laminae carinae 6 x 2-3 mm, manifeste saccatae. An-

therae late apiculatae, ad 0,7-0, 8 longae, squamis basa-

libus absentibus. Pistillum 5 mm longum, subtiliter fim-

briatum. Fructus 25-30 x 2,5-3 ,0 mm, dehiscens; septa

doliiformia, semina secedenta. Semina 2, 5-2, 7 x

1,7-1 ,8 mm, badia, maculis lineisque purpurascentibus.

TYPE. — Cape, Pilaarsberg, north of Pakhuis Pass,

9.1967, Kerfoot 5922 (NBG, holo.!).

Erect shrub up to 1 m tall, dense; woody below, half-

herbaceous above; branches set close, stiff; appressed

strigulose when young but glabrous when mature. Sti-

pules fused, adnate to base of petiole, up to 4 mm long,

acute, glabrous. Leaves scarce, imparipinnate. Leaflets

obovate, 3-5, 6-8 x 3, 0-3, 5 mm, base obtuse, apex

recurved mucronate, glabrous above, covered in few

flattened hairs around margin below. Petiole 14-15 mm

long; petiolules 0,5 mm long. Inflorescences axillary,

racemose, lax, 15-30-flowered; peduncle 60-140 mm

long. Flowers 5-6 (7) mm long, red, each subtended by

a 2, 0-2, 3 mm long, narrowly triangular, rapidly cadu-

cous bract; pedicel < 1 mm long. Calyx 2-3 mm long;

tube 1,5 mm long; teeth equal, 1,5 mm long, triangular,

sparsely fringed with flattened biramous hairs. Standard

broadly elliptic, 5-6 x 4, 0-4, 5 mm; claw short; auricles

absent; apex rounded; appendages absent. Wing petals

5, 5-6,0 x 2, 0-2, 5 mm, upper part flattened near the

margin, raised above but equal in length to the keel

blades, few scattered hairs present. Keel blades 6 mm

long, 2-3 mm wide at maximum, fringed along upper

margin. Androecium 5 mm long; vexillar stamen free;

anthers uniform, 0,7-0, 8 mm long, broadly apiculate,

anthers on long filaments basifixed, anthers on short

filaments dorsifixed; scales absent. Pistil 5, 0-5, 5 mm

long, glabrous, narrowly oblong, sessile, 7-8-ovuled;

style upcurved, height of curvature 2 mm; stigma capi-

tate. Fruits narrowly oblong, 25-30 x 2, 5-3,0 mm,

chestnut-brown, dehiscent; inner surface shiny white;

septa up to 2 mm wide, barrel-shaped, separating the

seeds, white with red flecks when in dry state. Seeds

elliptic, 2, 5-2, 7 x 1 ,7-1,8 mm, dark chestnut or khaki,

covered in large black blotches and stripes with small

speckles in between; hilum exserted. Figure 1 .

Indigofera gifbergensis is restricted to mountainous

regions from Kobe Pass in the north to Cedarberg Sneeu-

kop in the south (Figure 2).

This mountain fynbos endemic flowers from late

August to November. It occurs between 600-1 650 m.

CAPE. — 3118 (Vanrhynsdorp): Gifberg (-DC), 10.1953, Esterhuy-

sen 22084 (BOL); 9. 1948, Acocks 12891 (K); 8. 1948, Compton 20838

(NBG, STE); just before bridge over Snorkfontein River (-DC),

2.1979, Snijman 101 (MO, NBG, PRE); summit of Kobe Pass (— DB),

10.1973, Hall 4158 (NBG, PRE, STE). 3119 (Calvinia): Lokenberg,

WSW of Calvinia (-CA), 8.1953, Acocks 16882 (PRE). 3218 (Clan-

william): Langberg (-BA), 12.1951, Bond 1393 (NBG); 12.1941,

Esterhuysen 7335 (BOL); Pilaarsberg, north of Pakhuis Pass (-BB),

9.1967, Kerfoot 5922 (NBG). 3219 (Wuppertal): Heuningvlei Forest

FIGURE 1. — Indigofera gifbergen-

sis 1, flower, x 5,5; 2, calyx

opened out, x 6,8; 3, standard

opened out, x 8,2; 4, inner face

of wing petal, x 8,2; 5, outer

face of keel petal showing posi-

tion and shape of the peg, x

8,2; 6, androecium flattened

out, x 8,2; 7, pistil, x 8,2; 8,

fruit with section cut away to

show seeds and septa, x 2; 9,

seed, x 6, 8. 1-7: Kerfoot 5922.

8-9: Leipoldt 3366.

Bothalia 17,1 (1987)

3

1500m

FIGURE 2. — Distribution of Indigofera gifbergensis, ▼ ; I.filifolia, • in southern Africa.

Station along road to Groenberg (-AA), 10.1967, Emdon 143 (PRE,

STE); Pakhuis Mountain (-AA), 12.1940, Leipoldt 3366 (BOL);

1 1 . 1929, Thode A2112 (PRE); 9. 1940, Compton 9598 (NBG); 9. 1937,

Compton 6939 (NGB); 12.1981, Stirton 10196 (STE); Sneeukop,

Cedarberg (-CD), 2.1936, Compton 6207 (NBG). Without precise

locality: 8.1936, Thorne s.n (SAM 52691).

This distinctive species is easily separated from I. fili-

folia and /. ionii by its stiff compact habit, small brick-

red flowers, the absence of tufts on anthers and its

blotched seeds.

2. Indigofera ionii J. K. Jarvie & C. H. Stirton,

sp. nov. subsectioni Juncifolieae Gillettii proxima. /.

filifoliae Thunb. et I. gifbergensis C. H. Stirton & J. K.

Jarvie affinis sed ab ambabus habitu humiliore, semini-

bus olivaceis, leguminibus compressis, sacculo carinae

deorsum curvato, differt.

Herba perennis decumbens usque 0,8 m alta. Stipulae

liberae ad caulem appressae, glabrae. Folia juventute

praesentia, imparipinnata, petiolata. Foliola anguste

obovata, 3 (4 vel 5), 8-10 x 3-5 mm, lateralia minora,

basi obtusa, apice mucronata, supra glabra, subtus

sparse aequaliter pubescentia. Petioli 15-25 mm longi.

Inflorescentiae axillares, 6- 10-florae, racemosae, laxae,

40-70 (80) mm longae. Flores 5-6 (7) mm longi, uno-

quoque bractea caduca anguste triangulari 2,5-3 ,0 mm

longa subtenti; pedicellus filiformis, 2-3,5 mm longus.

Calyx 3, 0-3, 5 mm longus; tubus 1,5-1 ,6 mm longus,

glaber, dentibus 1,5 mm longis, ± aequalibus, triangula-

ribus. Vexillum 5-7,0 mm longum et latum, transverse

late ovatum; auriculae et calli absentes. Alae 6,8-7 ,0 x

2,5 mm, margine inferiore involutae; extremitas infera

laminae incisa. Petala carinae 6,5-7 ,0 x 2, 0-2, 5 mm,

apice pubescentia. Androecium 6, 0-6, 5 mm longum;

stamine vexillari libero; antherae uniformes, <0,5 mm

longae, ovatae, apiculatae, squamis basalibus praesenti-

bus; filamenta antherarum basifixarum longa, tenuia;

filamenta antherarum medifixarum, breviora, crassiora.

Pistillum 5-6 mm longum, compressum, anguste ob-

longum, 8-ovulatum. Fructus 3, 0-3, 5 mm longus, com-

pressus, dehiscens; septa <1 mm longa. Semina ob-

longa, 2-3 x 2 mm, olivacea.

TYPE. — Cape, Vogelgat, nearHermanus,/. Williams

3519 (K,holo.!; NBG, iso.!).

Small wiry erect or sprawling herb up to 0,8 m tall;

sparsely branched, mostly near the base. Stipules free,

2,5-3 ,0 mm long, appressed to stem, glabrous. Leaves

present during early stages of growth, imparipinnate,

petiolate. Leaflets 3, occasionally 4 or 5, narrowly obo-

vate, 8-10 x 3-5 mm; terminal largest; base obtuse,

apex mucronate, straight, glabrous above, sparsely and

evenly scattered with biramous hairs beneath. Petiole

1 5—25 mm long; petiolule < 1 mm long. Inflorescences

axillary, 6-10-flowered, racemose, lax, 40-70 (-80)

mm long. Flowers 5-6 (7) mm long, each subtended by

a 2,5-3 ,0 mm long, narrowly triangular caducous bract;

pedicel filiform, 2, 0-3, 5 mm long, elongating and thick-

ening during fruit formation. Calyx 3, 0-3, 5 mm long,

glabrous; tube 1,5-1, 6 mm long; teeth 1,5 mm long,

more or less equal, triangular; sinus between vexillar

teeth broad. Standard 5, 0-7,0 mm long and wide, very

broadly ovate, claw very short, venation fine, branched

from three traces at base, apex slightly emarginate;

auricles and appendages absent. Wing petals 6, 8-7,0 x

2,5 mm, infolded along lower margin; auricle inflated

and flattened; with scattered hairs along the margin to-

4

Bothalia 17,1 (1987)

FIGURE 3. — Indigofera ionii. 1,

calyx opened out, x 6,6; 2,

standard opened out, x 8; 3,

inner face of wing petal, X 8; 4,

outer face of keel petal showing

position and shape of the peg, x

8; 5, androecium flattened out,

X 8; 6, pistil, X 8; 7, fruit with

section cut away to show seeds

and septa, x 1,3; 8, septum, x

10; 9, seeds, LHS seed from

ends of the fruit and RHS seed

from the centre of the fruit, x

8,6. Voucher: Williams 3519.

wards the apex; lower end of blade notched. Keel petals

6,5-7 ,0 x 2,0-2, 5 mm, apex pubescent. Androecium

6, 0-6, 5 mm long; vexillar stamen free; anthers uniform,

< 0,5 mm long, ovate, apiculate, tufted at base; basi-

fixed anthers on long thin filaments; medifixed anthers

on short stout filaments. Pistil 5-6 mm long, com-

pressed, <0,5 mm long, glabrous, straight, narrowly

oblong, 8-ovuled; style upcurved, height of curvature

1,5 mm. Fruit 30-35 mm long, compressed, surface

coarsely rippled, margins distinctly ribbed, dehiscent;

septa < 1 mm long, white with small orange flecks.

Seeds oblong, 2-3 x 2 mm, olive-green. Figure 3.

Indigofera ionii is restricted to the mountains of the

south-western Cape (Figure 4). This fugitive fynbos

species favours damp areas, usually on tussocks in river

beds, vleis and marshes. It flowers sporadically from

August to April, with peaks between September and Jan-

uary. It occurs from 500-700 m.

CAPE. — 3418 (Simonstown): plateau between Constantiaberg and

Noordhoek (-AB), Levyns 6848 (BOL); west of Grootkop (-AB),

1.1940, Salter 8305 (BOL); Noordhoek Mountain (-AB), 11.1942,

Barker 2091 (NBG); marsh west of Klaasjagersberg (-AB); 1.1936,

Salter 5751 (BOL, BM, NBG); 8.1938, Salter 7606 (BM); Constantia-

berg (-AB), 3.1940, Bond 192 (NBG); 9.1922, Schlechter 1469

(BOL); 12.1943, Compton 15463 (NBG); Redhill (-AB), 1.1940, Le-

wis 709 (NBG); source of Prinskasteel River (-AB), 1.1948, Salter

8302 (BOL); stream south of Constantiaberg (-AB), 1 1 . 1896, Wolley-

Dod 1914 (K); Cape Point (-AD), 8.1944, Compton 15833 (NBG);

near Smitswinkel (-AD), Levyns 6850 (BOL); Compton s.n. (BOL);

Booiskraal near Brightwater, Cape of Good Hope Nature Reserve

(-AD), 12.1981, Grobbelaar 2647 (PRE); beacon near Smith’s farm

(-AD), 11.1939, Salter 8279 (BOL); Modderdam (-BA), 12.1938,

Salter 7874 (BM, BOL, PRE); 12.1938, Salter 7875 (BOL); Muizen-

berg Mountain (-BB), 12.1948, Compton 21210 (NBG); Rooi Els

Flats (-BD), 1 1 . 1945, Leighton 1448 (BOL); flats south west of Klein

Hangklip (-BD), 9.1969, Boucher 566 (STE); near Somersfontein

(-BD), 9.1969, Boucher 711 (STE). 3419 (Caledon): flats near Klein-

mond (-AC), De Vos 855 (STE); Palmiet Valley near Albertyn’s house

(-AC), 10.1948, De Vos 1045 (STE); Vogelpool, Vogelgat, near Her-

manus (-AD), 3.1984, Williams 3543 (K, NBG); Williams 3519 (K,

NBG); 9.1984, Stirton 10762 (K); marshy slopes at head of Boskloof

between Steenbras and Kogelberg (-BD), 2.1975. Esterhuysen s.n.

(BOL). Without precise locality: Forbes s.n. (K).

Bothalia 17,1 (1987)

5

3. Indigofera fllifolia Thunb., Prodromus planta-

rum capensium: 132 (1800); Thunb.: 595 (1823). Type:

Cape, ‘in collibus infra Tafelberg latere orientali’, Thun-

berg s.n. (UPS-Thunb. 17338, holo., seen on micro-

fiche).

Lebeckia contaminata Ker: 104 (1816) non Thunb. (1800). L. con-

taminata Ait.f. : 261 (1812) non Willd. (1803). Indigofera filifolia Ker:

notes 2 (1817). Spartium contaminatum Ait.: 10 (1789) non L. (1771).

Type: Cape of Good Hope, Masson s.n. (BM!, holo.!).

Lebeckia nuda Sims: t. 2214 (1821). Indigofera nuda (Sims) G.

Don: 301 (1850); G. Don: 209 (1832). Type: Anonymous s.n. Raised

from Cape seeds in 1814 by Messrs. Whitley, Brame & Milne at the

Fulham Nursery and then sent to Sims in October 1819 as Lebeckia

contaminata (BM, holo.!).

Indigofera aphylla Breit. ex Link: 251 (1822); Breit.: 209 (1817).

Type: ‘Kap der guten Hoffnung (Cape of Good Hope), cultivated in the

Breiter Gardens in Leipzig and later by Link in Berlin, collector un-

known (B, holo., destroyed).

Indigofera juncea DC.: 225 (1825); Mordant de Launay: 227, t. 227

(1820); Drapiez: 452, t. 452 (1834). Type: cultivated from seed sent

from England, 24 Jul. 1818, Cels s.n. (G, holo.!). Figure 5.

FIGURE 5. — Painting of Indigofera juncea DC. (Cels s.n.) = I. filifo-

lia.

Tree-like shrub up to 2 m tall, open habit, woody;

branches sparsely appressed strigulose when young but

glabrous when mature. Stipules free, filiform, strigulose,

up to 2 mm long, parallel to axis of branch. Leaves

common on younger plants, petiolate, basal pair largest,

upper pairs being reduced to scales. Basal leaflets 6-8,

elliptic, rarely narrowly obovate, 15-20 x 8-10 mm;

base cuneate; apex mucronate, straight; glabrous above,

sparsely and evenly scattered with appressed biramous

hairs beneath. Petiole 40-60 mm long; petiolules up to 2

mm long. Inflorescences axillary, clustered at ends of

branches forming a pseudocompound structure, race-

mose, lax, up to 50-flowered. Flowers (7,5) 8,0-10,5

mm long, each subtended by a short 1-2 mm long, nar-

rowly triangular, rapidly caducous bract; pedicel 4, 0-4, 3

mm long. Calyx 4, 0-4, 5 mm long; tube 1,7-3 ,4 mm

long; teeth unequal, 1, 1-3,0 mm long; triangular sinus

between vexillar pair very broad, hairs few, scattered in

the sinuses of the teeth. Standard (6,1) 8, 4-9, 2 (10,2) x

(5) 6, 3-7, 6 mm, orbicular; claw very short; venation

fine, branching from three traces at the base; apex

slightly hooded, auricles and appendages absent. Wing

petals 9-10 X 2, 5-3,0 mm, constricted in the middle;

auricle inflated and flattened. Keel petals 9 x 3, 0-3, 5

mm; apex pubescent. Androecium 9-10 mm long; vexil-

lar stamen free; anthers uniform, 1 mm long, apiculate

and tufted, anthers on long filaments basifixed, those on

short filaments dorsifixed; scales present. Pistil 6, 6-9,0

(9,9) mm long, terete, < 0,5 mm wide, glabrous,

arched, narrowly oblong, 8-14-ovuled; style upcurved,

height of curvature 2 mm. Fruit 40-55 x 4 mm, dark

brown, dehiscent. Seeds 2, 6-2, 7 mm long and wide,

chocolate-brown, hilum protruding, transversely oblong.

Figure 6.

Indigofera filifolia is a widespread fynbos endemic

occurring from the Cape Peninsula eastwards to Heidel-

berg (Figure 2). It favours damp sites and is especially

common along watercourses. The plants flower through-

out the year but with a peak from February to March.

This species coppices strongly after bums.

CAPE. — 3318 (Cape Town): valley behind Table Mountain (-CD),

February, Smuts 1083 (PRE); 3.1920, Andreae 296 (STE); north of

Window stream (-CD), 4.1956, Esterhuysen 25652 (BOL, MO); St

James, Cape Peninsula (-CD), 4.1914, St. Clair Capron s.n. (PRE);

summit of Table Mountain (-CD), 3.1899, Schoenberg 4867 (PRE);

3.1922, Wilson 130 (STE); 3.1917, Page s.n. (PRE); 4.1907, Kensit

68 (PRE); 2. 1929, Gillett3527 (STE); Kensit 68 (PRE); Prior s.n. (K);

4.1881, Bolus 4778 (K, BM, BOL); 3.1947, Barker 4445 (NBG);

3.1952, Salter s.n. (STE); 3.1910, Worsdell s.n. (K); 4.1929, Hut-

chinson & Smuts 3190 (BM, K); Rondebosch (-CD), July, Drege s.n.

(K). 3319 (Worcester): Franschhoek Pass (-CC), 3.1934, Smuts &

Gillett s.n. (STE). 3320 (Montagu): Swellendam Forest Reserve

(-CD), 1.1969, Marsh 1110 (STE); below 12 o’clock Peak (-CD),

5.1963, Taylor 4752 (PRE, STE); Barrydale Pass (-DC), Van Breda

1518 (K, PRE); Tradouw Pass (-DC), 3.1954, Balfour Browne 18

(BM). 3321 (Ladismith): Garcias Pass (-CC), 4.1935, Esterhuysen

32484 (BOL); Van Breda 1101 (K); 11.1814, Burchell 6974 (K).3322

(Oudtshoom): Montagu Pass (-CD), 2.1944, Fourcade 6367 (STE);

12.1949, Martin 76 (NBG); 6.1927, Fourcade 3245 (STE. PRE);

1 .1897, Guthrie s.n. (NBG); lower Montagu Pass, between Toll House

and bridge, 6.1943, Fourcade 6020 (STE); on Postberg near George

(-CD), 8.1814, Burchell 5952 (K, BOL, PRE); 7.1912, Rogers 4321

(BOL); Prior s.n. (K); 3.1928, Maude s.n. (K). 3323 (Willowmore):

Coldstream (-DC), Galpin 9565 (K). 3418 (Simonstown): Orange

Kloof River (-AB), 2.1896, Wolley-Dod 820 (BOL); 5.1953; Salter

9558 (BM, MO); above Kalk Bay (-AB), 3.1896, Wolley-Dod 1007

BM, BOL); 4.1974, Goldblatt 1399 (MO, PRE); Constantia Valley

(-AB), 4. 1886, Thode 9 (PRE); 1 1 . 1938, Salter 7852 (BOL); 3.1939.

Salter 8032 (BOL); Silvermine Plateau (-AB), 4.1974, Goldblatt 1585

(MO, NBG, PRE); 3.1932, Salter 2059 (BM, BOL); 3.1936, Salter

5947 (BM); Simonstown (-AB), 5.1884, Marloth 94 (PRE); 3.1935,

Minicki s.n. (NBG); 3.1918, Pillans 3488 (PRE); 8.1865, Wright s.n.

(K); Hout Bay (-AB), Prior s.n. (K); 4. 1955, Baker 657 (BM. NBG);

Blaauberg (-AB), Drege s.n (K); Schusters Kraal (-AB), 3.1941,

Penfold s.n. (NBG); flats at Smitswinkel vlei (-AD), 2.1961, Thomas

s.n. (STE); Boys Kraal Rivier Vlei (-AD), 3.1946, Compton 17940

(NBG); Bettys Bay (-BD), 3.1972, Vogts s.n. (STE); Palmiet Kloof,

east of bridge (-BD), 3.1948, De Vos 1299 (STE); 7.1950, Parker

4491 (BOL, K, NBG); 7.1975, Stirton 5013 (PRE); Ebersohn 243

(NBG); Palmiet River Mountains (-BD), 1.1947, Stokoe s.n. (NBG);

Rooi Els (-BD), 3.1948, Parker 4319 (BOL. K. NBG); Elephant Rock

Mountain (-BD), 3.1971, Boucher 1464 (K, PRE, STE); Platbos Pass

(-BD), March, Boucher 1176 (K, PRE, SRGH, STE). 3419 (Caledon):

Elgin (-AA), 4.1946, Stokoe s.n. (SAM 59572); Nuwekloof, Houw-

hoek Mountains (-AA), 3. 1815. Burchell 8052, 8134 (K); on the sides

of the Bot River, near Houwhoek (-AA), Bowes s.n. (BM); Swartberg

Mountain near Caledon (-AB), 9.1904, Bolus s.n. (BOL); Palmietberg

(-AC), 3.1983, Van Wyk 1142 (PRE); Hermanus (-AC), 4.1949, De

6

Bothalia 17,1 (1987)

FIGURE 6. — Indigofera filifolia. 1,

calyx opened out, X 6,8; 2,

standard opened out, x 8,2; 3,

inner face of wing petal, X 8,2;

4, outer face of keel petal show-

ing position and shape of the

peg, x 8,2; 5, androecium flat-

tened out, x 8,2; 6, pistil, x

8,2; 7, fruit with section cut

away to show seeds and septa,

x 1,7; 8, septum, x 6,8; 9,

seed, x 6,8. Voucher: Williams

3542.

Waal s.n. (BOL, NBG); 4.1946, Compton 17963 (BOL, NBG);

4.1980, Orchard 545 (K, MO, PRE, STE); Cascade Pool, Vogelgat

(-AD), 4.1984, Williams 3542 (K); Sues Pool, Vogelgat (-AD),

4.1979, Williams 2739 (MO, NBG, PRE); Vogelkloof, 4. 1950, Martin

322 (STE); Maanschijnkop (-AD), 4.1978, Williams 2465 (MO,

NBG); Babylon’s Tower (-AD), 2.1941, Esterhuysen 4974 (BOL);

near Stanford, foot of Kleinrivier Mountains (-AD), 4.1952, Stokoe

s.n. (NGB, STE); Boesmanskloof (-BA), 4.1983, Forsyth 24 (PRE);

Genadendal, Baviaanskloof (-BA), 3.1933, Gillett 840 (BOL, STE);

Riviersonderend (-BB), 4.1950, Middlemost 1534 (NBG); Wynand

Louw’s Bos (-BD), 3.1963, Van der Merwe 1357 (PRE, STE); Koude-

bergkloof near Nuwepos (-DA), 4.1975, Oliver 5845 (PRE, STE).

3420 (Bredasdorp): Swellendam Mountain (-AB), 2.1941, Compton

10600 (NBG); 4.1926, Smith 2720 (PRE); 3.1930, Thode 2318 (K);

1.1815, Burchell 7401 (K); Hermitage Kloof (-AD), 12.1958, Stokoe

s.n. (NBG). 3421 (Riversdale): Lemoenshoek Mountain near Heidel-

berg (-AA), 3.1948, Stokoe s.n. (SAM 61559). Without precise local-

ity: Cape, Harvey, s.n. (BM, K); Cape Flats, February, Ecklon &

Zeyher s.n. (SAM 32330); Bowie s.n. (BM); Langeberg Mountains,

4. 1923, Muir 2528 (PRE).

ACKNOWLEDGEMENTS

The authors wish to thank the following: the curators

and staff of BM, BOL, K, MO, NBG, PRE and STE for

their help and loan of much valuable material; Dr B. V.

Verdcourt and Miss M. Wilmot-Dear for kindly provid-

ing the Latin translations; Dr Ion Williams for his hospi-

tality and kindness to the second author and for his accu-

rate field notes and comprehensive collections of /. ionii.

Finally the first author would like to thank Drs Barbara

Pickersgill, J. Barnett and R. Wadsworth (Reading Uni-

versity); Dr I. C. Trueman (Wolverhampton Polytech-

nic); Brenda Timmons, Chloe Bennett and Lani Rubin

for help and advice.

REFERENCES

AITON, W. 1789. Spartium contaminatum. Hortus kewensis 3: 10.

AITON, W. T. 1812. Lebeckia contaminata. Hortus kewensis 4: 261 .

BAKER, E. G. 1905. Report on some South African species of Indigo-

fera in the Albany Museum Herbarium. Record of the Albany

Museum 1: 279-281.

BREITER, C. A. 1817. Indigofera aphylla. Hortus breiterianus: 209.

Leipzig.

BREMEKAMP, C. E. B. 1932. New or otherwise noteworthy plants

from the northern Transvaal. Annals of the Transvaal Museum 15:

233-264.

BROWN, N. E. 1925. New species of Indigofera from the Transvaal

and Swaziland. Kew Bulletin 1925: 142-159.

BROWN, N. E. 1926. Indigofera. In J. B. Burtt Davy, Manual of

flowering plants and ferns, Transvaal 2: 44-^15.

DE CANDOLLE, A. P. 1825. Indigofera juncea. Prodromus systema-

tis 2: 225. Paris.

DE KORT, I. & THUSSE, G. 1984. A revision of the genus Indigo-

fera (Leguminosae — Papilionoideae) in south-east Asia. Blumea

30: 89-151.

DENTER, K. 1932. Diagnosen neuer sudwestafrikanischer Pflanzen.

Feddes Repertorium 30: 198.

DON, G. 1832. Indigofera nuda. A general system of gardening and

botany 2: 209. London.

DON, G. 1850. Indigofera nuda. In J. C. Loudon, Hortus britannicus :

301. London.

DRAPEEZ, P. A. J. 1834. Indigofera juncea. Herbier de T Amateur de

Fleurs 7: 452. t. 452.

DYER, R. A. 1944. Indigofera burkeana. Flowering Plants of South

Africa: 939.

FOURCADE, H. G. 1932. Contributions to the flora of the Knysna

and neighbouring divisions. Transactions of the Royal Society of

South Africa 21: 75-102.

GILLETT, J. B. 1958. Indigofera (Michrocharis) in tropical Africa

with the related genera Cyamopsis and Rhynchotropis . Kew Bulle-

tin, Additional Series 1: 1-166.

GUTHRIE, L., PILLANS, N. S. & SALTER, T. M. 1939. Plantae

novae Africanae. Journal of South African Botany 5: 66-73.

HARVEY, W. H. 1862. Leguminosae. In W. H. Harvey & O. W.

Sonder, Flora capensis 2: 163-203. Reeve, London.

KER, J. B. 1816. Lebeckia contaminata. Botanical Register 2: 104.

KER, J. B. 1817. Indigofera filifolia. Botanical Register 3: notes 2.

LINK, H. F. 1822. Indigofera aphylla. Enumeratio plantarum horti

regii bot. berolinensis 2: 251. Berol.

MERXMULLER, H. & SCHREIBER, A. 1957. Einige neue Legumi-

nosen aus Siidwestafrika. Bulletin du Jardin botanique de L’Etat,

a Bruxelles 27: 269-274.

MORDANT DE LAUNAY, J. C. M. & LOISELEUR-DESLON-

CHAMPS, J. L. A. 1820. Indigofera juncea. Herbier General de

I’ Amateur 4: 227 , t. 227. Paris.

POLHILL, R. M. 1981. Indigofereae (Benth.) Rydb. In R. M. Polhill

& P. H. Raven, Advances in legume systematics: 289-291. Royal

Botanic Gardens, Kew.

SCHREIBER, A. 1970. Einige neue Fabaceen aus Siidwestafrika. Mit-

teilungen aus der Botanischen Staatssammlung Munchen 8:

137-145.

SIMS, J. 1821 . Lebeckia nuda. Curtis’s Botanical Magazine: t. 2214.

STIRTON, C. H. 1982. A new species of Indigofera from the south-

west Cape. Bothalia 14: 69-82.

SUESSENGUTH, K. & MERXMULLER, H. 1951. Species et varie-

tates novae vel rarae in Africa australi et centrali lectae. Mittei-

lungen aus der Botanischen Staatssammlung Munchen 3: 89.

THUNBERG, C. P. 1800. Indigofera filifolia. Prodromus plantarum

capensium. 2: 132.

THUNBERG, C. P. 1823. Indigofera filifolia. Flora capensis, edn 2.

Copenhagen.

Bothalia 17,1:7-15 (1987)

A synopsis of Tephrosia subgenus Barbistyla( Fabaceae) in southern

Africa

B. D. SCHRIRE*

Keywords: Fabaceae, key, Millettieae, new combinations, new species, Papilionoideae, southern Africa, subgenus Barbistyla, synopsis,

Tephrosia, T. brummittii

ABSTRACT

A synopsis of the southern African species of Tephrosia Pers. subgenus Barbistyla Brummitt is given. Twenty-one

species are recognized. A new species, T. brummittii B. D. Schrire, is described and six new combinations are presented.

The key is partly descriptive to provide diagnoses of the taxa and includes information on distributions. Vouchers are noted

from at least three different herbaria to aid identification. Detailed notes are provided on atypical collections and taxa that

require further investigation. Typification has been done as far as necessary to ensure that the names used are correct.

UITTREKSEL

’n Samevatting van die suider-Afrikaanse spesies van Tephrosia subgenus Barbistyla Brummitt word gegee. Een-en-

twintig spesies word erken. ’n Nuwe spesie, T. brummittii B. D. Schrire, word beskryf en ses nuwe kombinasies word

voorgestel. Die sleutel is gedeeltelik beskrywend om diagnoses van die taksons te verskaf en bevat inligting oor versprei-

ding. Kontrole-eksemplare is van ten minste drie herbaria aangeteken om identifikasie te vergemaklik. Gedetailleerde notas

word verskaf oor atipiese versamelings en oor taksons wat verdere ondersoeke vereis. Tipifikasie is gedoen sover dit nodig

was om seker te maak dat die name wat gebruik is, korrek is.

INTRODUCTION

Tephrosia is a pantropical genus in the tribe Millet-

tieae (Papilionoideae) comprising over 400 species

mostly from Africa. An overview of the nomenclatural

history, systematic position, taxonomy and morphology

of the genus is given by Bosman & De Haas (1983) and

the most recent reviews of the tribe are by Geesink

(1981, 1984).

In the Flora of southern Africa region it is estimated

that there are 52 species eight subspecies and 10 varie-

ties. There are at least three interspecific hybrids with the

likelihood of more being found in the future.

My treatment of the genera closely allied to Tephrosia

follows that by Brummitt (1980), i.e. Requienia and

Ptycholobium are retained as separate genera whereas

Lupinophyllum is sunk in Tephrosia.

Despite recent criticism by Geesink (1984) there is

good reason for retaining the two subgeneric divisions in

Tephrosia (Brummitt 1980). Although precise characters

such as arrangement and position of hairs either along

the style or just below the stigma, or the degree of style

twisting, are variable, the two subgenera are character-

ized by flowers with different pollination strategies.

The salient features of each subgenus are summarized

in the key below.

In subgenus Barbistyla, 21 species are recognized; a

number of H.M.L. Forbes’s (1948) species are reduced

to synonomy or to subspecific status. The subgenus has a

high degree of endemism in southern Africa with the T.

natalensis, T. macropoda, T. albissima and T. grandi-

flora complexes almost all radiating exclusively within

the Flora of southern Africa area. The three latter groups

appear to be closely related to one another. It is however

only these species in the subgenus that are endemic to us.

The T. longipes complex and some of the other isolated

species are more tropical in origin and they represent

southerly extensions into the FSA area. A number of

other areas of diversity for the subgenus occur in tropical

Africa.

Selection pressures for evolutionary divergence at the

generic level appear to have been centred on the breeding

system. The subgeneric categories have different flower

structures and this is reflected in their different polli-

nation strategies. Within the subgenus Barbistyla, inflo-

rescence characters (bracts) and fruit characters (pod size

and seed position) separate the T. grandiflora allies from

those of T. longipes. Inflorescence specialization has

been particularly marked in the T. grandiflora alliance.

Flowers (4-) 6-13 (-18) mm long (from calyx across the keel), orange or pink; wings and keel distally curving up-

wards (the keel often appearing L-shaped); standard somewhat folded over the keel basally and curving upwards

with the distal half reflexed; style glabrous, with or without a tuft of hairs just below the stigma .... subgenus Tephrosia

Flowers (1 1— ) 14-24 (-30) mm long (from calyx across the keel), pink or mauve; wings and

keel forwardly directed, obtriangular; standard generally straight, either folded over the keel or fully reflexed

away from it; style pubescent, with or without a tuft of hairs just below the stigma subgenus Barbistyla

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

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

Richmond, Surrey TW9 3AE, England, UK.

8

Bothalia 17,1 (1987)

An interpretation of the above might be that in the

early stages of the evolution of Tephrosia the floral char-

acters were fairly labile, and only after the build up of

various constraints, did two major alternative strategies

become dominant. The multifunctional-structural flower

had now become constrained by the accumulated adapt-

ions and interactions (burden) that made it successful.

This became a fixed feature around which the other parts

of the breeding system could interact. Further selection

could be channelled in either of two directions: speciali-

zation in the ovules and ovary and ultimately fruit and

seed dispersal, or specialization in inflorescences acting

on floral development, presentation and protection.

Further examination of these hypotheses could add to

our knowledge of the radiation and interrelationships

within the genus Tephrosia as a whole.

This synopsis is a preliminary account of the subgenus

Barbistyla aimed at providing names and a means of

identification for a difficult and variable group of plants.

Hybridization is so far only known from isolated cases in

disturbed areas, between closely related species.

The genus appears to comprise a number of actively

evolving complexes in which peripheral variation is dif-

ficult to assess, and much of the underlying genetic va-

riation is not readily quantifiable in morphological

terms. A full revision of the genus for the Flora of

southern Africa is in preparation.

KEY TO SPECIES

(Note: 1: b = length/breadth)

la Stipules and flower bracts triangular to narrowly triangular or subulate, ± of equal size, stipules never

auriculate at the base or with reflexed margins:

2a Slender, procumbent to erect perennial herbs with many flexuous stems; leaflets 9-13, very narrowly

elliptic to narrowly oblanceolate or oblong, 6-25 (28) mm long, often discolorous; pseudoracemes laxly

flowered; pods linear, 35-55 mm long; seeds 6-9; from E Transvaal and Swaziland 1 T. retusa

2b Robust, decumbent to erect suffrutices, subshrubs or annuals:

3a Pseudoracemes compact and densely congested, on short peduncles 2-30 (-40) mm long, inflorescences

often but not always more or less exceeded by the leaves; pods grey to brown villous:

4a Leaves ( 1— )3— <— 5)-foliolate , stipules narrowly triangular; leaflets abaxially grey sericeous; from SWA/

Namibia and Botswana 2.T. cephalantha var. decumbens

4b Leaves (6-) 7-10 (— 1 1 )-foliolate, stipules triangular; leaflets abaxially appressed pubescent not

sericeous; from NE Transvaal 3. T. aequalata subsp. australis

3b Pseudoracemes elongated and laxly flowered, up to 300 mm long excluding the peduncle or, if ±

congested, then at the ends of peduncles raised well above the leaves; pods glabrous to thinly

pubescent:

5a Robust herbaceous-stemmed suffrutices or annuals; pods linear, l:b ± 12:1; seeds orbicular or with

longer dimension along the longitudinal axis of the pod:

6a Leaflets linear to linear-elliptic, l:b ± 12:1 with acuminate apices: leaflets in 1-7 (8) pairs, (20-)

30-100 (— 120)X( 1— ) 2-7 (-14) mm; widespread in S Botswana, Transvaal, OFS, Natal and

N Cape 4. T. longipes

6b Leaflet shape other than above, l:b ± 6: 1 or less with acute, rounded, truncate or emarginate apices:

7a Youngest foliage densely white or yellowish pubescent and conspicuously dark mucronate;

from SWA/Namibia, N Botswana, N and NE Transvaal:

8a Leaflets narrowly to very nanowly elliptic with acute apices, (8-) 15-35 (-45) X (2-) 3-7 (-12)

mm; pods with 10-12 seeds; from N Transvaal 5. T. euchroa

8b Leaflets elliptic-oblong, oblanceolate or narrowly oblong with rounded, truncate or emarginate

apices, (10—) 35-60 (-80) X 7-12 (-15) mm; pods with 12-16 seeds; from SWA/Namibia,

N Botswana and NE Transvaal:

9a Stems with conspicuous spreading brown to grey hairs up to 2,5 mm long:

10a Annual slender herb with a long thin taproot; stems not strongly decumbent; leaves few in

number, restricted to the lower part of the plant; leaflets in 1-4 (5) pairs; from

N Botswana 6. T. euprepes

10b Perennial with woody rootstock or, if annual then with robust, strongly decumbent stems;

leaflets in 4-9 pairs:

11a Stems erect, conspicuously long brown pilose; petiole and rhachis together

(90—) 120-180 (-200) mm long; calyx with long spreading hairs; from

N Botswana 7a. T. caerulea subsp. caerulea

lib Stems usually decumbent, to 1 m or more long, rarely erect; moderately to densely

yellowish or grey pilose; petiole and rhachis together 40-1 10 (-120) mm long; calyx

shortly pubescent; from NE Transvaal 8. T. reptans var. reptans

9b Stems with appressed or ascending grey hairs; from SWA/Namibia, N Botswana

7b. 7". caerulea subsp. otaviensis

7b Youngest foliage conspicuously dark brown in dried material and essentially glabrous or sparsely to

moderately long pilose; from Natal midlands, N Natal, adjacent S Transvaal and Swazi-

land:

12a Pseudoracemes elongated with at least three distinct flowering nodes along the rhachis; calyx

teeth lanceolate to linear, equal to or longer than the tube; from Howick and Pietermaritzburg

Districts of Natal midlands, Utrecht to Paulpietersburg Districts of N Natal, adjacent

S Transvaal and Swaziland 9a. T. natalensis subsp. natalensis

12b Pseudoracemes in pseudoheads at the ends of the peduncles, with only one or rarely two

flowering nodes; calyx teeth triangular, shorter than the tube; from the Muden-Greytown-

Kranskop Districts of Natal midlands 9b. T. natalensis subsp. pseudocapitata

Bothalia 17,1 (1987)

9

5b Erect woody-stemmed subshrubs; pods narrowly oblong to lorate, l:b ±6:1; seeds transversely elongate

with the longer dimension across the width of the pod:

13a Young stems canescent; stipules linear; pseudoracemes from short peduncles, elongate and laxly

flowered; from E Swaziland 14.7. gobensis

13b Young stems sparsely to moderately pubescent; stipules narrowly triangular to lanceolate;

pseudoracemes in pseudoheads at the ends of the peduncles; from E Transvaal, Natal to

Transkei 21. 7. shiluwanensis

lb Stipules and flower bracts ovate to lanceolate, if stipules narrow then either auriculate at the base and with

reflexed margins or somewhat narrower than the bracts:

14a Prostrate to ascending herbaceous plants with many slender stems arising from a woody rootstock; usually of

open grassland and sandy places:

15a Peduncles less than 30 mm long; pseudoracemes elongated and laxly flowered; leaflets characteris-

tically obtriangular to broadly obovate, up to 20 (-26) mm long; pods shortly oblong, (8-) 12-25 mm

long with a rounded apex; seeds 2—4 (5), not transversely elongated; prostrate mat-forming herb;

stems conspicuously spreading-pubescent; from N SWA/Namibia and N Transvaal 10.7. radicans

15b Peduncles longer than 30 mm; pseudoracemes with flowers often clustered at the ends of peduncles;

leaflets variously shaped, up to 65 mm long; pods lorate, (20—) 25-60 mm long with acute apices; seeds

more than 5, transversely elongated:

16a Buds completely enclosed by large, scarious navicular or trullate bracts 4-15 (-18) mm long

(sometimes not quite masking the buds in 7. bachmannii); pseudoracemes sparsely flowered with

only 1-3 (4) flowers open at any one time:

17a Calyx densely white to yellow sericeous; leaflets abaxially canescent or sericeous; endemic to

northern Pondoland/southem Natal sandstone region 15. 7. bachmannii

17b Calyx shortly pubescent to spreading pilose; leaflets not as above:

18a Stipules maturing brown to dark brown, dying back and curling away from the stems; leaves

3-7-foliolate; leaflets obovate to narrowly obovate, 20-65 x 6-30 mm; OR (which includes

var. diffusa) 5-9-foliolate with leaflets narrower, elliptic, 8-20 x 3-6 mm; widespread in the

eastern half of the FSA area 11.7. macropoda sensu lato

18b Stipules maturing conspicuously red or brown, closely appressed to the stems; leaves 7-11

(— 13)-foliolate; leaflets mostly narrowly oblong-elliptic or oblanceolate, (12-) 18-36 (—4-2) x

3-7 (-12) mm; from N Zululand and Swaziland Lowveld 12. 7. brummittii

16b Buds visible and only partially masked by trullate, ovate to narrowly lanceolate bracts (1) 2-5 (-8) mm

long; pseudoracemes compactly 2-8 (-lO)-flowered:

19a Leaves with petiole and leaf rhachis together longer than 25 mm (except occasionally in depauperate

upland forms from the Kokstad-Harding region of E Griqualand); leaves never sessile; from

Transkei to the Natal midlands 13a. 7. albissima subsp. albissima

19b Leaves with petiole and leaf rhachis together shorter than 25 mm; leaves often sessile and with very

reduced rhachides; from W Zululand and N Natal 13b. 7. albissima subsp. zuluensis

14b Erect or bushy subshrubs 0, 5-3,0 m high, with one to a few stems, woody at least at their bases; usually of

scrub or forest margins, but where these vegetation types are evidently disturbed then plants untidy and

scrambling:

20a Calyx shortly appressed white to yellowish sericeous; leaflets abaxially grey subsericeous to sericeous:

21a Leaflets very narrowly oblanceolate or, if broader than 40 mm then longer than 26 mm; bracts 1-10

(-12) mm long, about as long as wide, broadly ovate with rounded to acute apices, moderately to

densely white sericeous; endemic from the sandstone outcrop region of N Pondoland/S Natal

15. 7. bachmannii

21b Leaflets mostly shorter than 26 mm and wider than 40 mm; bracts (8-) 10-32 mm long, navicular or

narrowly ovate, longer than wide with narrowly acute, more often acuminate apices; from the coastal

belt of SW Cape to Natal south of Durban, inland to the Albany and Amatola regions of the

ECape 16. 7. grandiflora

20b Calyx white to rufous pubescent, spreading pilose or villous; leaflets abaxially glabrous to canescent:

22a Bracts exceeding and completely enclosing young buds and also often the undeveloped apices of the

pseudoracemes:

23a Stipules broadly ovate and conspicuously cordate at the base:

24a Compact, woody shrubs, largest stipules longer than 13 mm, characteristically chartaceous,

maturing to purple or maroon, crowded and overlapping on new growth; from the Lydenburg

and Barberton Districts of E Transvaal and Swaziland 19.7. cordata

24b Lax, short-lived perennials; largest stipules never longer than 13 mm, scarious, maturing reddish

to brown; from the NE, E and SE Transvaal and N Natal, W Zululand to the Natal

midlands 18b. 7. glomeruliflora subsp. meisneri

23b Stipules ovate, not cordate at the base, or narrowly ovate to lanceolate and auriculate at the base,

often with re flexed margins:

25a Bracts — (8-) 10—32 mm long, navicular or narrowly ovate, longer than wide with narrowly acute

or more often, acuminate apices:

26a Stipules broadly ovate, closely appressed to the stems; bracts sparsely to moderately puberulent;

from the coastal belt of SW Cape to Natal south of Durban, inland to the Albany and Amatola

regions of E Cape 16.7. grandiflora

26b Stipules narrowly lanceolate to subulate, often with reflexed margins and spreading

from the stem; bracts moderately to densely spreading pilose or villous; from the Eshowe to

Inanda Districts of Natal 17. 7. inandensis

25b Bracts 4-10 (-12) mm long, trullate, broadly ovate or orbicular, ± as long as wide with rounded,

broadly acute or apiculate apices:

10

Bothalia 17,1 (1987)

27a Calyx teeth narrowly subulate, 4—7 mm long, significantly longer than the tube; from

the Barberton, Nelspruit and Komatipoort Districts of E Transvaal and Weza in

EGriqualand 20. T. subulata

27b Calyx teeth triangular, lanceolate or subulate, 1,0-3, 5 mm long, shorter than to slightly

exceeding the tube;

*28a Young inflorescences in dense glomerules at the ends of the peduncles; bracts flesh-

coloured, minutely puberulent with a ciliate margin or short and densely white pubescent;

calyx teeth triangular or appearing shortly subulate when dry, shorter than the tube except

for the lowest tooth:

29a Stems densely appressed pubescent, becoming glabrous only with age; bracts and calyces

moderately to densely white pubescent; from the coastal belt of Natal from Umzinto to N

Zululand and inland to the Boston District of the midlands in the south and to the

Hlabisa District in the north 18a. T. glomeruliflora subsp. glomeruliflora

29b Stems glabrous or only sparsely pubescent when young; bracts flesh-coloured, minutely-

puberulent with a ciliate margin; calyces sparsely to densely white to rufous pubescent or

pilose; from the NE, E and SE Transvaal and N Natal, W Zululand to the Natal midlands

18b. T. glomeruliflora subsp. meisneri

t28b Young inflorescences in dense pseudoheadsat the ends of the peduncles or pseudoracemes

up to 100 mm long; if appearing glomerule-like then bracts white to yellowish spreading

pilose; calyx teeth lanceolate-subulate, ± equal to the tube:

30a Youngest leaves densely spreading yellowish sericeous; bracts ovate-acuminate, (6) 7-12

mm long, densely white to yellowish pilose; from the Eshowe to Inanda Districts of Natal

17. T. inandensis

30b Youngest leaves densely appressed canescent; bracts ovate and apiculate, 4-7 mm long,

sparsely to moderately white pilose; from the Shilovane, Pilgrims Rest and Lydenburg

Districts of E Transvaal to N Natal, W Zululand and the Natal midlands from the Grey-

town through Pietermaritzburg to Harding Districts and the Transkei 21.7’. shiluwanensis

22b Bracts never completely exceeding or enclosing young buds and only partially obscuring them in the

pseudoracemes:

31a Calyx teeth broadly triangular, with acute apices, much shorter than the tube; bracts 1-3 mm long,

obovate or ovate with truncate or rounded to apiculate apices, shortly and densely white pubescent;

from the Boston and Durban Districts of Natal 18a. T. glomeruliflora subsp. glomeruliflora

31b Calyx teeth narrowly triangular to lanceolate or subulate with attenuate apices, ± equal to or exceed-

ing the tube:

32a Calyx teeth narrowly subulate, 4-7 mm long, significantly longer than the tube; from the Bar-

berton, Nelspruit and Komatipoort Districts of E Transvaal and Weza in E Griqualand .... 20. 7. subulata

32b Calyx teeth lanceolate or subulate, l,0-3,5(-4,0) mm long, more or less equal to the tube:

33a Youngest leaves densely appressed canescent; bracts 1-6 mm long, minutely puberulent to

sparsely pubescent; calyx lobes often maturing bristle-like and re flexed apically; from the

Shilovane, Pilgrims Rest and Lydenburg Districts of the E Transvaal to N Natal, W Zululand

and the Natal midlands from the Greytown through Pietermaritzburg to Harding Districts and

the Transkei 21.7'. shiluwanensis

33b Youngest leaves densely spreading yellowish sericeous; bracts (6-) 7-12 mm long, densely

white to yellowish pilose; calyx lobes not as above; from the Eshowe to Inanda Districts

of Natal 17. T. inandensis

SYNOPSIS OF SPECIES

1 . Tephrosia retusa Burn Davy, Flowering plants

and ferns of the Transvaal 2: xxxi + 378 (1932). Type:

Transvaal, Barberton, juxta Kaap River, Bolus 7724

(K!,BOL).

Distribution: E Transvaal and Swaziland.

Vouchers: Bolus 11821 (K, BOL); Elan-Puttick 254

(PRE); Liebenberg 3317 (K, PRE); Pott 5326 (PRE);

Rogers 24827 (K, PRE).

2. Tephrosia cephalantha Welw. ex Bak. var. de-

cumbens Welw. ex Bak. emend. Brummitt in Boletim da

* The term glomerule, used to describe certain juvenile inflorescences

in the key, refers to the spherical or globose body located at the ends of

peduncles which is formed by the compact arrangement of broadly

ovate to orbicular bracts with rounded, apiculate or broadly acute

apices.

t The term pseudoheads is used when inflorescences are congested at

the ends of the peduncles but either the bracts are too small to com-

pletely enclose the young buds or the apices project out from the inflo-

rescence because they are sharply acute to acuminate.

Sociedade Broteriana, ser. 2, 41: 384 (1968). Type: An-

gola, Hufla, Ferrao da Sola, Welwitsch 2090 (K!).

Distribution: SWA/Namibia and Botswana.

Vouchers: De Winter 2801 (K, PRE); De Winter &

Marais 4539 (K, PRE); Holub s.n. (K); Merxmiiller &

Giess 1864 (K); Smith 2311 (K).

3. Tephrosia aequalata Bak. subsp. australis

Brummitt in Boletim da Sociedade Broteriana, s6r. 2, 41:

358 (1968). Type: Zimbabwe, Melsetter Distr., Williams

146 (K, holo.!; PRE!, SRGH).

Distribution: NE Transvaal.

Vouchers: Codd 3040 (K, PRE); Hanekom 2283 (K,

PRE); Hemm 182 (K, PRE); Scheepers 341 (K, PRE);

Taylor 646 (K, PRE).

Note

This species was confused by H. M. L. Forbes: 990

(1948) with T. zombensis Bak.

Bothalia 17,1 (1987)

11

4. Tephrosia longipes Meisn. in Hooker’s London

Journal of Botany 2: 87 (1843). Type: Natal, in gramino-

sis ad latera montium Tafelberge, Port Natal, Krauss 20

(K, holo.!).

4a. subsp. longipes.

Brummitt in Boletim da Sociedade Broteriana, ser 2

41:309(1968).

Tephrosia lurida Sond.: 30 (1850). Type: Transvaal, Moorivier,

Magaliesberg, Croccxlile River, Zeyher456 (K, isosyn. !).

Tephrosia angustissima Engl.: 29 (1888). Type: Cape, pr. Kuru-

man, in summo montium ‘Ga Mhana’ , Mar loth 1086 (B, BOL GRA

PRE!, SAM).

Distribution: widespread in S Botswana, Transvaal,

OFS, Natal and N Cape.

Vouchers: Brenan & Vahrmeijer 14255 (K); Codd

2661 (K, PRE); Germishuizen 133 (K, PRE); Rogers

18129 (K, PRE); Wood 1202 (K).

Notes

(1) The typical subsp. is maintained because of the

occurrence of subsp. swynnertonii (Bak. f.) Brummitt:

315 (1968) in eastern Zimbabwe.

(2) A great deal of confusion has arisen in the past

concerning the T. longipes-T. lurida complex. Forbes

(1948) and Brummitt (1968) treated the two as separate

species while Gillett (1958) placed T. lurida as a variety

of T. longipes. Present research has shown that tradi-

tional characters used to separate them are based on eco-

logical variables which cannot be reliably upheld taxono-

mically.

(3) A number of varieties have been described in this

complex, i.e. T. longipes var. uncinata Harv.: 209

(1862). Type: Transvaal, Pretoria Distr., in nemorosis

ad Aapies R., Zeyher 455 (BM, GRA, K!, SAM); T.

longipes var. icosisperma Brummitt: 313 (1968), e.g.

Moss & Rogers 523 (K) from Komatipoort, Transvaal;

T. lurida var. lissocarpa Brummitt: 320 (1968). Type:

Transvaal, Soutpansberg, ± 9 miles E of Louis Tri-

chardt, Farm Rustfontein, Schlieben 7339 (K, holo.!)

and T. lurida var. drummondii Brummitt: 321 (1968),

e.g. Liebenberg 3313 (PRE) from Farm Schagen, Nel-

spruit, Transvaal. More field work is necessary to estab-

lish the validity of these varieties in the Flora area and

they are not considered in this preliminary treatment.

(4) This complex together with the following species,

i.e. T. reptans Bak., T. caerulea Bak. f., T. euprepes

Brummitt and T. euchroa Verdoom forms a clearly

recognizable assemblage that is widespread in tropical

Africa, and with the exception of T. longipes Meisn.

only occurs in the northern extremities of the Flora area.

5. Tephrosia euchroa Verdoorn in Bothalia 3: 239

(1937). Type: Transvaal, Soutpansberg Distr., rocky

NW slopes of Soutpansberg, Obermeyer, Schweickerdt

& Verdoorn 73 (PRE, holo. !; K!).

Distribution: N Transvaal.

Vouchers: Acocks 16762 (K, PRE); Gertenbach 4914

(PRE); Pienaar 287 (K, PRE); Smuts & Gillett 4080 (K,

PRE); Van der Schijff 1824 (PRE).

6. Tephrosia euprepes Brummitt in Boletim da

Sociedade Broteriana, ser. 2, 41: 292 (1968). Type:

Zimbabwe, western Div., Nyamandhlovu Distr., Bon-

golo, West 2694 (SRGH, holo.!; K!).

Distribution: N Botswana.

Vouchers: Allen 299BCD (K); Lugard 152, 175 (K);

Rand 19, 20 (BM).

7. Tephrosia caerulea Bak. /., Leguminosae of

Tropical Africa 1: 197 (1926). Type: Zambia, Maza-

buka, Woods 51 (BM, holo.!).

7a. subsp. caerulea.

Distribution: N Botswana.

Vouchers: De Winter 4406 (K, PRE); Smith 2211 (K).

7b. subsp. otaviensis (Dinter) Schreiber & Brum-

mitt in Boletim da Sociedade Broteriana, s6r. 2, 41: 292

(1968).

Tephrosia otaviensis Dinter in Feddes Repertorium 30: 204 (1932).

Type: SWA/Namibia, Hereroland, Klein Otavi, Dinter 5747 (NH,

iso.!; K, photo.!).

Tephrosia longipes sensu Schreiber: 298 (1957).

Distribution: SWA/Namibia and N Botswana.

Vouchers: De Winter & Marais 4832 (K); Dinter

7570 (K); Muller & Biegel 2305 (K).

8. Tephrosia reptans Bak. in Oliver, Flora of

Tropical Africa 2: 121 (1871). Types: Tanzania, Tabora

Distr., nr Tabora, Speke & Grant s.n. (K, lecto. !) and

Mozambique, Shupanga, Kirk s.n.

8a. var. reptans.

Brummitt: 284 (1968).

Distribution: NE Transvaal.

Vouchers: Smalberger 29 (PRE); Stephen 303 (PRE).

Note

Lectotypification by Brummitt: 283 (1968).

9. Tephrosia natalensis H. M. L. Forbes in Botha-

lia 4: 988 (1948). Type: Natal, Howick, Wood 8341

(NH, holo.!; K!).

9a. subsp. natalensis.

Tephrosia apiculata H. M. L. Forbes: 989 (1948). Type: Natal,

Paulpietersburg Distr. between Paulpietersburg and Sunkelsdrift,

Acocks 11749 (NH, holo.!; K, photo.!).

Distribution: S Transvaal, Swaziland, N Natal and

Natal midlands.

Vouchers: Arnold 252 (K, PRE); Compton 29168 (K,

PRE); Devenish 1722 (K, PRE); Schrire 1289, 2254

(NH).

9b. subsp. pseudocapitata (H. M. L. Forbes) B. D.

Schrire, comb, et stat. nov. Tephrosia pseudocapitata

H. M. L. Forbes in Bothalia 4: 988 (1948). Type: Natal,

Greytown Distr., Wylie s.n. (NH21787) (NH, holo.!;

K!).

Distribution: Natal midlands.

Vouchers: Buthelezi 332 (NH); Stirton 5203 (K,

PRE); Wilms 1940 (K); Wylie s.n. (NH27992) (K, NH).

12

Bothalia 17,1 (1987)

Note

The T. natalensis group may be an isolated ally of the

complex comprising species numbered four to eight.

10. Tephrosia radicans Welw. ex Bak. in Oliver,

Flora of Tropical Africa 2: 121 (1871). Type: Angola,

Huilla, in pratis humidis ad formicar monticul de

Varzeas e Catumba, Welwitsch 2082 (BM, holo. !; K!).

Distribution: SWA/Namibia and N Transvaal.

Vouchers: De Winter & Marais 4992 (K, PRE); Gal-

pin 8959 (K, PRE); Rehmann 5529 (K); Schlechter 4679

(K); Van Vuuren 1411 (K, PRE).

11. Tephrosia macropoda (E. Mey.) Harv. in

Harv. & Sond., Flora Capensis 2: 210 (1862). Syntypes:

E Cape, inter Kachu et Zandplaat, Drege s.n. (B);

Transkei, inter Gekau et Basche, Drege s.n. (K!); inter

Umtata et Umgazana, Drege s.n. (K!). See note (1)

below.

Apodynomene macropoda E. Mey.: 112 (1836). Apodynomene

macropoda E. Mey. var. latifolia E. Mey.: 1 12 (1836).

Apodynomene macropoda E. Mey. var. angustifolia E. Mey.: 112

(1836). Type: Transkei, in collibus graminosis prope Umsikaba, Drege

s.n. (?).

Apodynomene aemula E. Mey.: 113 (1836). Tephrosia aemula (E.

Mey.) Harv. & Sond.: 210 (1862), pro parte. Type: ECape, in grami-

nosis inter Zandplaat et Komba, Drege s.n. (B, holo.).

Tephrosia triphylla Harms in 0. Kuntze: 57 (1898). Type: Natal,

Krantzkloof, Kuntze s.n. (NY, holo.-K, photo.!).

Tephrosia armitageana Chiov.: 421 (1908). Type: Natal, Port Natal,

Armitage 547 (? FI).

Tephrosia spathacea sensu H. M. L. Forbes: 995 (1948), non

Hutch. & Burtt Davy: xxxi (1932).

Distribution: widespread in the eastern half of the

Flora area.

Vouchers: Compton 27355 (K, PRE); Huntley 4 72 (K,

NU); Liebenberg 7990 (K, PRE); Mogg 5812 (K, PRE);

Rogers 18020 (K, PRE).

1 la. var. diffusa (E. Mey.) B. D. Schrire, comb, et

stat. nov.

Apodynomene diffusa E. Mey.: 113 (1836). Tephrosia diffusa (E.

Mey.) Harv. in Harv. & Sond.: 210 (1862). Type: Transkei, in rupes-

tris ad fluvium quendam minorem inter Umsamvubu et Umsikaba,

Drege s.n. (K!).

Apodynomene aemula E. Mey.: 113 (1836). Tephrosia aemula (E.

Mey.) Harv. in Harv. & Sond.: 210 ( 1862) pro parte. Type: E Cape, in

graminosis inter Zandplaat et Komga, Drege s.n. (B, holo ).

Distribution: ecotypes within the range of the species.

Vouchers: Acocks 21940 (PRE); Ross 1886 (K, NH);

Strey 6620 (K, PRE); Venter 7708 (PRE); Wood 886

(K).

Notes

(1) H. M. L. Forbes examined material in B before the

Second World War for her revision (1948). She desig-

nated a specimen, Drege (Herb. Meyer 5471) as a type

thus choosing one of the syntypes in Meyer’s protologue

as the lectotype. Since this material was destroyed and it

is not clear which syntype she chose, a new lectotype can

be designated from any of the original syntypes.

(2) The T. macropoda complex is a widespread group

of trailing to prostrate open grassland herbs and over the

major part of its range, i.e. Transvaal and much of Natal,

Transkei and the eastern Cape, it is easily recognizable

as one species, T. macropoda. On the periphery of this

range, however, there is considerable divergence from

the typical form:

(a) Along the eastern coastal strip plants are particularly

robust but are clearly recognizable as T. macropoda.

(b) In the following regions plants may be much reduced

in size:

(i) the higher altitude foothills and slopes of the Dra-

kensberg in Natal to Transkei;

(ii) the southerly end of the species range in the E

Cape;

(iii) the coastal hinterland of Transkei and Natal;

(iv) parts of the E Transvaal and Swaziland.

(3) One distinct form (referred to here as var. diffusa)

occurs in these areas, and is associated with edaphically

dry granite or sandstone outcrops. Plants are procum-

bent, diffusely branching and often form dense mats.

Typical T. macropoda may be found in the surrounding

grassland and the two may appear to be quite separate

taxa.

(4) T. macropoda is, however, a variable species and

smaller-leaved forms do occur and will intergrade with

the typical form, particularly at the extremes of its range

and with increasing altitude. These are sometimes diffi-

cult to distinguish from the diffuse plant in the herbar-

ium. The latter is clearly an ecotype and the name is

retained as a variety of T. macropoda but doubtful plants

should be retained in T. macropoda sensu lato.

12. Tephrosia brummittii £. D. Schrire, sp. nov.

T. macropodae affinis, sed stipulis late ovatis, maturitate

conspicue rubris vel brunneis, arete appressis ad caules,

foliis 7—1 1(— 13)-foliolatis, foliolis anguste oblongo-

ellipticis ad oblanceolatis differt.

TYPE. — Natal, 2832 (Mtubatuba): Hlabisa Distr., St

Lucia Estuary Game Park, W of Vidal Rd (-AD), Pooley

1918e (NU, holo.; K, PRE, iso.).

Perennial spreading suffrutex up to 600 mm high, aris-

ing from a woody rootstock. Stems one to many, herba-

ceous except at the base, prostrate to ascending, glabrous

or sparsely pubescent to ascending pilose, branching

near the base. Leaves 7-11 (-13)- foliolate, (30-)

70-120(-140) mm long; stipules (6— )7— 17 x 3-11 mm,

broadly ovate, base often cordate, apex acuminate,

maturing conspicuously red or brown, closely appressed

to the stems; petiole (15-)25-55(-70) mm long, canali-

culate adaxially; leaflets ( 1 2—) 1 8— 35(— 42) x 3— 7(— 12)

mm, narrowly oblong-elliptic or oblanceolate, base

cuneate to rounded, apex emarginate, obtuse or acute,

mucronate, upper surface green, glabrous or minutely

puberulent, lower surface moderately appressed to

spreading greyish pubescent. Pseudoracemes terminal

and leaf-opposed, with 1— 2(— 3) flowers open at any one

time. Bracts 5-15 x 4-10 mm, broadly ovate-acumi-

nate, navicular, completely enclosing young buds, sca-

rious, chestnut-brown, minutely bifid at the apex. Flow-

ers 11-21 mm long. Calyx tube to 4 mm and teeth to 6

mm long, teeth lanceolate at the base narrowing to a long

Bothalia 17,1 (1987)

13

attenuate apex, longer than the tube, sparsely to modera-

tely pubescent with long spreading hairs on the teeth.

Standard 11-21 mm long, to 20 mm wide, broadly obo-

vate to orbicular, narrowing to a claw 2-3 mm long at

the base, apex rounded or emarginate, dorsally densely

pubescent, ventrally glabrous, scattered punctate-glan-

dular as are all the petals. Wings 10-18 x 7 mm includ-

ing a claw 2-3 mm long, cultrate with an auricle at the

base of the lamina above the claw, sculpturing present,

upper basal, up to 20 transcostal lamellae. Keel petals up

to 18 x 7 mm including a claw 2-3 mm long. Staminal

sheath 10-14 mm long, the stamens then free and alter-

nating in length, 5 and 4 filaments to 4 and 3 mm long

respectively, vexillary stamen to 17 mm long, free.

Ovary to 16 mm long. Style to 8 mm long, curving erect

from the ovary, flattened dorsiventrally and densely pu-

bescent on both surfaces. Stigma capitate. Fruits imma-

ture, glabrous. Seeds 10-16, transversely arranged.

SWAZILAND. — 2631 (Mbabane): Umbuluzi Falls near stream

(-AD), Compton 25094 (NBG, PRE).

NATAL. — 2732 (Ubombo): Lebombo Mountains, Gwalaweni For-

est (-AC), Brenan 14272 (K, PRE); Umpangazi Lake (-DA), Strey

5097 (NH, PRE). 2831 (Nkandla): Mtunzini Distr., 15 miles NW of

Port Dumford (-DD), Codd & Dyer 2826 (K, PRE). 2832 (Mtuba-

tuba): Hlabisa Distr., St Lucia Estuary Game Park, W of Vidal Rd

(-AD), Pooley 1918e (K, NU, PRE).

This species has often been confused with T. macro-

poda var. macropoda, from which it differs by having

stipules maturing conspicuously red or brown and re-

maining closely appressed to the stems, and leaves 7-1 1

(— 13)-foliolate with leaflets narrowly oblong-elliptic to

oblanceolate. T. macropoda var. macropoda is distin-

guished by having stipules maturing dark brown and

curling away from the stems, and leaves 3-7-foliolate

with leaflets obovate to narrowly obovate or elliptic.

T. brummitti occurs sympatrically with T. macropoda

and, although a few problem plants occur in overlapping

areas, e.g. Huntley 194 (PRE) from Mtunzini, Natal,

they are not readily identifiable as intermediates.

Two specimens from southern Mozambique, cited by

Brummitt: 372 (1968) as T. incarnata, also belong to

this species. These are: 1, Sabie, Lebombo Mountains,

Meponduine, Pedro & Pedrogao 735 (LMJ); 2,

Namaacha, near the Canada Dry factory, Barbosa &

Lemos 7534 (COI, LISC, LMJ). Both localities are near

the border with Swaziland in the Lebombo Mountains.

The plants grow at altitudes from 0-450 m in the

coastal and low veld vegetation types of N Zululand, S

Mozambique and E Swaziland. These prostrate herbs

occur in open grassland from coastal dunes to stream-

banks and forest margins along the Lebombo Mountains.

This species is named in honour of Dr R. K. Brummitt

of the Royal Botanic Gardens, Kew, who has contrib-

uted much to the knowledge of the genus Tephrosia from

the Flora Zambesiaca area.

13. Tephrosia albissima H.M.L. Forbes in

Bothalia 4; 995 (1948). Type: Natal, Greytown, Wylie

s.n. (NH21590) (NH, holo.!; K!, PRE!).

13a. subsp. albissima.

Tephrosia galpinii H. M. L. Forbes: 996 (1948). Type: Natal, Grey-

town, on railway bank on town commonage, Galpin 14755 (PRE,

holo.!; NH!).

Distribution: Natal midlands to Transkei.

Vouchers: Schrire 298, 312, 831 (NH); Stirton 5598

(K, PRE); Wylie s.n. (NH3715f) (NH).

13b. subsp. zuluensis (H. M. L. Forbes) B. D.

Schrire, comb, et stat. nov. Tephrosia zuluensis

H. M. L. Forbes in Bothalia 4: 997 (1948). Type: Natal,

Nkandhla, Wylie s.n. (NH9399) (NH, holo. !).

Tephrosia unifolia H. M. L. Forbes in Bothalia 4: 996 (1948). Type:

Natal, W Zululand, Baker s.n. (NH19781 ) ( NH, holo.!).

Distribution: W Zululand and N Natal.

Vouchers: Codd 6987 (K, NH); Grobbelaar 801

(PRE); Ngwenya 31 (NH); Schrire 2216 (NH); Turner 1

(NH).

Notes

(1) T. albissima is another variable complex including

a number of names published simultaneously by

H. M. L. Forbes (1948). A geographically distinct segre-

gate, T. zuluensis is given subspecific status and this

includes those forms from western Zululand and nor-

thern Natal which have almost sessile leaves and very

reduced densely pilose leaf rhachides. This creates prob-

lems in the choice of key characters because the typical

subspecies becomes very much reduced in size at the

opposite end of its range in the higher altitude areas of

Kokstad and Harding in eastern Griqualand (following a

similar trend to T. macropoda sensu lato.).

(2) Another entity, T. unifolia H. M. L. Forbes is

placed in synonomy under subsp. zuluensis but further

field study is needed to be certain of its position. The

best example is J. Turner 1 (NH) from the Babanango

Distr. in western Zululand. The leaves are distinctly pe-

tiolate with leaflets narrowly oblong-elliptic to oblanceo-

late, both longer and narrower than typical subsp. zu-

luensis. The collections Brown & Shapiro 73 (K) from

Itala Nature Reserve and Ross 1741 (PRE) from near

Goedgegun in the Louwsberg, need further investigation

as they appear to be somewhat intermediate between T.

albissima and T. macropoda sensu lato.

14. Tephrosia gobensis Brummitt in Boletim da

Sociedade Broteriana, ser. 2, 41: 368 (1968). Type:

Mozambique, Goba Distr., De Sousa 132 (LISC, holo.;

PRE!).

Distribution: E Swaziland.

Vouchers: Compton 31915 (PRE) Culverwell 463 (K,

PRE).

15. Tephrosia bachmannii Harms in Botanische

Jahrbiicher 26: 286 (1899). Type: Transkei, Pondoland,

Bachmann 616 (B, holo.).

Distribution: N Transkei and S Natal.

Vouchers: Crawford 390 (K, PRE); Strey 9591 (K,

my, Wood 3021 ,3101 (K).

Notes

This species is very variable, with three distinct forms:

(a) procumbent grassland suffrutices e.g. Shackleton

367 (KEI);

(b) low woody bushes with linear-oblanceolate leaf-

lets, from exposed rocky areas e.g. Acocks 13375 (PRE);

14

Bothalia 17,1 (1987)

(c) woody subshrubs up to 2 m high with broader,

obovate leaflets conspicuously white sericeous abax-

ially, e.g. Sidey4185 (PRE).

Further investigation is needed to assess this variation.

16. Tephrosia grandiflora (Ait.) Pers., Synopsis

Plantarum 2: 329 (1807). Types: Cape, Hort. Kew 1777,

seed from C. B. S., Masson s.n. (BM, syn.!); Africa

australis, Masson s.n. (BM, syn.!).

Galega grandiflora Ait.: 70 (1789).

Distribution: SW Cape to S Natal.

Vouchers: Bruce 453 (K, PRE); Marais 1195 (K,

PRE); Stirton 6297; 6320 (K, PRE); Strey 6897 (K, NH,

PRE).

Notes

The largest and most difficult complex, which is allied

to both T. albissima and T. macropoda sensu lato, com-

prises those erect or bushy subshrubs related to T. gran-

diflora (species 14 to 21). T. grandiflora has tradition-

ally been confused with many of the species in this group

but presents no problems itself. The isolated T. gobensis

from the Lebombo region seems most closely allied to it

as does the endemic T. bachmannii from the sandstone

outcrop areas of northern Pondoland/southem Natal.

17. Tephrosia inandensis H. M. L. Forbes in

Bothalia 4: 276 (1948). Type: Natal, Inanda Distr.,

Wood 289 (NH, holo.!; K!).

Distribution: Eshowe to Inanda Districts of Natal.

Vouchers: Forbes 108 (NH); J. P. Forbes s.n. (NH);

Lawn 537 (NH) Rogers 24018 (PRE); Stirton 5341

(NH).

Note

T. inandensis is closely allied to T. glomeruliflora

(no. 18), and Schrire 330 (NH) from the Natal coastal

area adjacent to Eshowe is a hybrid between them.

18. Tephrosia glomeruliflora Meisn. in Hooker’s

London Journal of Botany 2: 86 (1843). Type: Natal,

Port Natal, Gueinzius s.n. (K, holo. !).

18a. subsp. glomeruliflora.

Distribution: Natal coastal belt and adjacent midlands.

Vouchers: Baijnath 145 (K, NU); Grobbelaar 1814

(K, PRE); Nichols 840 (NH); Pole Evans 3647 (K,

PRE); Wood 11400 (NH).

18b. subsp. meisneri (Hutch. & Burtt Davy) B. D.

Schrire, comb, et stat. nov. Tephrosia meisneri Hutch.

& Burtt Davy: xxxi & 377 (1932). Type: Transvaal,

Sanderson s.n. (K, holo.!).

Tephrosia incarnata Brummitt: 370 (1968). Type: Zimbabwe,

Umtali Distr., Himalayas, Engwa, Wild 4450 (SRGH, holo.; K ! ,

PRE!).

Distribution: E half of Transvaal, N Natal to Natal

midlands. It also extends northwards into Zimbabwe.

Vouchers: Brenan 14116 (K, PRE); Germishuizen

2852 (PRE); Killian 14 (PRE); Nichols 508 (NH); Pole

Evans s.n. (PRE); Strey 7968 (NH).

Notes

(1) The glomerulose inflorescences and geographical

distribution of the two taxa T. glomeruliflora and

T. meisneri suggest one species with two distinct subspe-

cies, and Strey 8371 (K, NH) from Ismont near Pieter-

maritzburg and Coleman 973 (NH) from Byrne in the

Natal midlands indicate a possible transition from one to

the other.

(2) The typical subspecies shows considerable varia-

tion in bract size. Some individuals have minute bracts

not at all obscuring the young buds in the inflorescence

(as in the type of the species). The bulk, though, have

bracts enclosing the young buds completely. There is

little evidence of intermediates between them and on

further investigation the latter may warrant varietal

status.

(3) A few plants of subsp. meisneri are depauperate

e.g. Meeuse 9797 (K) from the Soutpansberg and Galpin

14451 (K) from Pilgrim’s Rest. These require further

investigation as mentioned under the next species.

19. Tephrosia cordata Hutch. & Burtt Davy in

Burtt Davy, Flowering plants and ferns of the Transvaal

2: xxxi (1932). Type: Swaziland, Mbabane, Burtt Davy

2886 (K, holo.!; BM!, PRE!).

Distribution: E Transvaal and Swaziland.

Vouchers: Codd 8198 (K, PRE); Compton 23302,

29985 (K); Elan-Puttick 320 (K, PRE); Stirton 1768 (K,

PRE).

Note

T. cordata requires further study in the field, particu-

larly as regards variation in stipule size and possible

links with T. glomeruliflora subsp. meisneri, e.g. Galpin

13134 (K) from Dullstroom and Pole Evans 4245 (K)

from Hendriksdal (both in the E Transvaal).

20. Tephrosia subulata Hutch. & Burtt Davy in

Burtt Davy, Flowering plants and ferns of the Transvaal

2: xxxi & 377 (1932). Type: Transvaal, Barberton

Distr., Highland Creek, Galpin 843 (K, holo.!; NH!,

PRE!, SAM).

Distribution: E Transvaal and Weza District in E

Griqualand.

Vouchers: Balsinhas 3153 (K, PRE); Stirton 10388

(NH); Thorncroft 36 (PRE); Van der Merwe 247 (K,

PRE); Viljoen 133 (K, PRE).

Note

T. subulata occurs sympatrically with T. cordata in

the E Transvaal, and Kluge 2347 (PRE) from Kaapsche-

hoop is an aberrant specimen that may represent a hybrid

between them.

21. Tephrosia shiluwanensis Schinz in Vierteljah-

resschrift der Naturforschenden Gesellschaft in Zurich

52: 423 (1907). Type: Transvaal, Soutpansberg Distr.,

Shiluwane, Junod 2355 (Z, holo.).

Tephrosia spaihacea Hutch. & Bunt Davy in Burtt Davy: xxxi

(1932). Type: Transvaal, Lydenburg Distr., Wilms 341 (K, holo.!).

Tephrosia medleyi H. M. L. Forbes: 968 (1948). Type: Natal,

Kranskop Distr. , nr Greytown, Wood 5984 (NH, holo. !).

Bothalia 17,1 (1987)

15

Tephrosia wyliei H. M. L. Forbes: 991 (1948). Type: Natal,

Elandskop Distr., Sevenfontein nr Boston, Wylie s.n. (NH7996) (NH,

holo. !).

Distribution: E Transvaal, Natal to Transkei.

Vouchers: Codd 6961 (PRE); Collins 3266 (PRE);

Germishuizen 95 (K, PRE); Junod 1106 (K, PRE);

Kluge 2253 (K, PRE); Tyson 1434 (K).

Notes

(1) 7. shiluwanensis is a variable species. At the west-

ern end of its range, i.e. from the Pietermaritzburg to

Harding Districts, plants tend to be less robust with

bracts generally more conspicuous and ovate-apiculate.

This group matches the type of T. wyliei but for the most

part they cannot reliably be separated from T. shiluwa-

nensis.

(2) Fakude 68 (PRE) is a hybrid with T. glomeruli-

flora subsp. glomeruliflora from the Hlabisa District of

Natal.

(3) An aberrant specimen, Evans 345 (NH) from the

Umlaas District between Pietermaritzburg and Durban

may be a hybrid with T. inandensis and requires further

investigation.

(4) Two additional species have been recorded as in-

troductions to southern Africa but there is no evidence

that they have become naturalized:

(a) T. Candida (Roxb.) DC.; Roxburgh s.n., Cape of

Good Hope ex Botanic Garden, Calcutta (K);

(b) T. vogelii Hook, f.; Strey 9009, Ntsubane,

Transkei (NH, PRE).

ACKNOWLEDGEMENTS

My sincere thanks to the following for much valuable

advice and constructive criticism: Mr C. H. Stirton, Dr

R. M. Polhill, Mr G. P. Lewis and Dr R. K. Brummitt.

REFERENCES

AITON, W. 1789. Horius kewensis 3: 70. London.

BAKER, E. G. 1926. The Leguminosae of tropical Africa 1: 197.

Gent.

BAKER, J. G. 1871. Papilionaceae. In D. Oliver, Flora of tropical

Africa 2: 121. Reeve, London.

BOSMAN, M. T. M. & DE HAAS, A. J. P. 1983. The genus Tephro-

sia in Malesia. Blumea 28,2: 421-487.

BRUMMITT, R. K. 1968. New and little known species from the

Flora Zambesiaca area. xx. Tephrosia. Boletim da Sociedade Bro-

teriana, ser. 2,41: 219-393.

BRUMMITT, R. K. 1980. Reconsideration of the genera Ptycholo-

bium, Caulocarpus, Lupinophyllum and Requienia in relation to

Tephrosia. Kew Bulletin 35: 459-473.

BURTT DAVY, J. 1932. A manual of the flowering plants and fems of

the Transvaal with Swaziland, South Africa 2: xxxi-xxxii,

375-379. Longmans, London.

CHIOVENDA, E. 1908. Papilionaceae in R. Pirotta, Flora della colo-

nia Eritrea. Annuario del R. Istituto botanico di Roma 8,3: 421.

DINTER, K. 1932. Diagnosen neuer sudwestafrikanischer Pflanzen.

Feddes Repertorium 30:204.

ENGLER, A. 1888. Plantae Marlothianae . Botanische Jahrbiicher fur

Systematik. Pflanzengeschichte und Pflanzengeographie 10:29.

FORBES, H. M. L. 1948. A revision of the South African species of

the genus Tephrosia Pers. Bothalia 4: 95 1-1001 .

GEESINK, R. 1981. Tephrosieae. In R. M. Polhill & P. H. Raven,

Advances in legume systematics 1: 245-260. Royal Botanic Gar-

dens, Kew.

GEESINK, R. 1984. Scala Millettiearum. Leiden Botanical Series 8:

1 18-121. Brill, Leiden.

GILLETT, J. B. 1958. Notes on Tephrosia in tropical Africa. Kew

Bulletin 13: 125.

HARMS, H. 1899. Leguminosae Africanae II. Botanische Jahrbiicher

fur Systematik. Pflanzengeschichte und Pflanzengeographie 26:

286. '

HARVEY, W. H. 1862. Leguminosae. In W. H. Harvey & O. W.

Sonder, Flora capensis 2: 203-211. Hodges & Smith, Dublin.

HOOKER, W. J. 1843. The Flora of South Africa. The London Jour-

nal of Botany 2: 86-87.

KUNTZE, C. E. O. 1898. Revisio generum plantarum 3: 57. Leipzig.

MEYER, E. H. F. 1836. Commentariorum de plantis Africae austra-

lioris: 1 1 1-1 13. Leipzig.

PERSOON, C. H. 1807. Synopsis plantarum 2,2: 329. Paris.

SCHINZ, H. 1907. Mitteilungen aus dem botanischen Museum der

Universitat Zurich (xxxvi). Vierteljahresschrift der Naturfor-

schenden Gesellschaft in Zurich 52: 423.

SCHREIBER, A. 1957. Beitrage zur Kenntms der Leguminosen Sud-

westafrikas. Mitteilungen der Botanischen Staatssammlung

Miinchen 19: 298.

SONDER, O. W. 1850. Beitrage zur Flora von Sudafrika. Linnaea 23:

30-31.

VERDOORN, I. C. 1937. An enumeration of plants collected in the

northern Transvaal. Bothalia 3: 239.

Bothalia 17,1: 17-23(1987)

A new species and a new combination of Asc/epias(Asclepiadaceae) in

southern Africa

A. NICHOLAS*

Keywords: Asclepias, Asclepiadaceae, new combination, new species, southern Africa, taxonomy

ABSTRACT

A new species is described: Asclepias gordon-grayae A. Nicholas, sp. nov. and a new combination is made: Asclepias

compressidens (N.E. Br.) A. Nicholas, comb. nov.

U1TTREKSEL

’n Nuwe spesie word beskryf: Asclepias gordon-grayae A. Nicholas, sp. nov. en ’n nuwe kombinasie word gemaak:

Asclepias compressidens (N.E.Br.) A. Nicholas, comb. nov.

1. Asclepias gordon-grayae A. Nicholas, sp. nov.

A. pateni N.E. Br. et A. praemorsae Schltr. affinis; dif-

fert a A. praemorsa appendicibus antherae brevibus et a

A. pateni margine supero coronae prope extremum dis-

tale fisso.

Herba perennis. Caudex profundus, lignosus. Caulis

unicus, erectus, gracilis, pubescens. Folia expansa,

anguste lanceolata ad linearia, 34-128 x 2,25— (—35)

mm. Inflorescentia umbellata, ad nodos et terminalis,

semi-pendens. Flores rosei ad ebumei. Corolla expansa

usque ascendens, lobis ovatis, basi connatis. Corona

cucullata-saccata, appendicibus proximalibus dentatis,

subfalcatis, extremo distali coronae brevissimo et ob-

tuso, sinus coronae saccati, carina coronae rotundata.

Appendices antherae ovatae usque ellipticae, 1,0-1, 4 x

0,5—1 ,0 mm. Apex styli truncatus. Pollinia hemitrullata

usque hemipyriformes, 0,68-0,92 x 0,40-0,52 mm.

Fructus fusiformis, apice rostrato.

TYPE.— 2832 (Mtubatuba): (-AB) Natal, Zululand,

St Lucia, Eastern Shores State Forest, Simbonvini vlei,

20 m (65 ft) alt., March 1982, Nicholas 1285 (PRE,

holo.; CPF, K, NH, MO).

Perennial herb with woody, semi-swollen, deep-

seated underground organs. Stem single, erect, 240-750

mm high. Leaves spreading, narrowly lanceolate to lin-

ear, occasionally upper leaves falcate, 34—128 x 2-25

(-35) mm, apex acute to acuminate, base minutely

auriculate to minutely hastate; petiole 0,6-1, 3 (-2,0)

mm long. Inflorescence umbelliform, semipendulous,

axillary and terminal, 1-4 per plant, 4-6 (-9)-flowered;

peduncles 7-14 (-24) mm long. Flowers 6-14 x

4, 5-8,0 mm. pink or pale pink to cream; pedicel 7-20

mm long. Sepals lanceolate, occasionally triangular or

narrowly ovate, 2, 2-3, 4 X (0,7—) 1 ,4 — 2 , 1 mm. Corolla

spreading to ascending; petals connate at base, ovate,

apex acute, 6, 4-7, 7 x 3, 2-4, 9 mm, white, cream or

pink in centre and near apex, margins white to pale pink,

abaxial surface glabrous, margins minutely scabrous.

Corona lobes fused to staminal tube 0, 3-0,5 mm above

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

corolla, saccate or cyathiform, 1,3-2, 6 x 2, 3-3, 3 mm,

white to cream with pink keel. Upper proximal ends

extended into 2 long, dentate, subfalcate appendages

1,5-2, 8 x 0,4-0, 8 mm, level with style apex, 2 tooth-

or wing-like flaps present below appendages on proximal

margin. Upper margin oblique to style apex, proximal

end usually higher than distal end, which is sometimes

raised into a short, blunt projection deeply cleft at apex,

cleft running a short distance along rounded keel. Stami-

nal column 2-3 mm high; alar fissure 1, 2-2,0 mm long;

anther wings curvirostrate, 1, 2-2,0 x 0,5-0, 9 mm;

anther appendages ovate to elliptic, 1,0-1 ,4 x 0, 5-1,0

mm, membranous, white, decumbent on style apex.

Style apex truncate, 1, 5-3,0 mm wide. Pollinia hemi-

trulloid to hemipyriform, 0,68-0,92 x 0,5-0,52 mm;

translator arms 0,2-0,32 mm long; corpusculum

0,28-0,4 x 0,5-0,52 mm. Fruits fusiform, ±65 x ± 10

mm, puberulous. Figure 1.

NATAL. — 2831 (Nkandla): Ngoye Forest Reserve (-DC), Gordon-

Gray 6191 (NU), Hilliard 2699 (NH, NU), Huntley 624 (NU), Nicho-

las 1074 (NU), Stewart 2155 (NU), Stirton 466 (PRE), Strey 6106

(NH, NU), Wood 5678 (NH), 10823 (BOL, SAM). 2832 (Mtubatuba):

St Lucia (-AB), Cawood 141 (CPF), Nicholas 1285 (CPF, K, MO,

NH, PRE); St Lucia (-AD), Crundall s.n. (PRE 51584), Pooley 1819

(NU). 2930 (Stanger): Gingindlovu (-BA), Wood 10828 (SAM).

Asclepias gordon-grayae ** is endemic to coastal

Zululand where it grows in boggy situations. This spec-

ies has been named in honour of Professor K. D. Gor-

don-Gray who has collected extensively (particularly in

Natal) and who for many years lectured in taxonomy at

the University of Natal, Pietermaritzburg. Professor

Gordon-Gray has through her many publications and her

teaching of taxonomy made a positive contribution to the

knowledge of the southern African flora. A. gordon-

grayae has in the past been confused with A. patens

N.E.Br. with which it has many affinities. Vegetatively

** In Nicholas (1982) the manuscript name A. gracilicaulis was used

(in reference to the long, thin, wavy stem of this species). This name

was never validly published and has been dropped here in favour of the

name A. gordon-grayae.

18

Bothalia 17,1 (1987)

FIGURE 1 . Asclenias gordon-grayae. 1 , flowering plant, x 0,Z5, Edwards s.n. (BOL); 2, node showing bifariously arranged hairs and leaf

bases, x 4, Gordon-Gray 6191 (NU); 3, flower, x 7, Stewart 2155 (NU); 4, corona lobe, side view, X 3, Stewart 2155 (NU); 5,

polhnanum, X 130, Stewart 2155 (NU).

Bothalia 17,1 (1987)

19

tell these two species apart. However, there are marked

differences in flower colour, corona lobe size and shape,

anther appendage size and shape and pollinia width (Fig-

ure 2; Table 1). There are also differences in distribution

(Figure 3) and in habitat preference. The number, degree

and consistency of these and other differences are diag-

nostically important and help separate A. gordon-grayae

from its two closest relatives: A. patens and A. prae-

morsa Schltr. (Figures 2, 4 & 5; Table 1). A. gordon-

grayae may be found growing at altitudes between 30

and 900 metres and flowers between September and

April, with a peak period in December.

KEY TO ASCLEPIAS GORDON-GRAYAE AND ITS CLOSEST

ALLIES

la Anther appendages 1 ,7-2,5 X 1 ,0-1 ,6 mm A. praemorsa

lb Anther appendages 0,5-1 ,4 x 0,5-1 ,0 mm:

2a Upper coronal edge entire; southern Natal and Transkei . .

A. patens

2b Upper coronal edge cleft near the distal end; Zululand

only A. gordon-grayae

FIGURE 2. — A comparison of the flowers of: la, Asclepias gordon-grayae, x 5, Stewart 2155 (NU); lb , A. patens x 9,5, Moss 470 (J); lc,

A. praemorsa X 8, Strey 6902 (NH). A comparison of the corona lobes of: 2a, A. gordon-grayae, x 4, Strey 6106 (NU); 2b, A.

patens x 10, Gordon-Gray 961 (NU); 2c, A. praemorsa x 14, Wood 1162 (NH).

FIGURE 3. — Map showing comparative distribution of Asclepias

gordon-grayae, • and A. patens,

2. Asclepias compressidens (N.E.Br.) A. Nicho-

las, comb, et stat. nov.

Asclepias navicularis (E. Mey.) Schltr. var. compressidens N.E.Br.:

683 (1908). Type: Cape, 3225 (Somerset East): Cradock (-BA), Coo-

per 1284 (K, lecto. ! ; PRE!).

Perennial herb with woody, swollen, deep-seated un-

derground organ. Stems 1-6 per plant, decumbent.

Leaves spreading to ascending, linear, occasionally nar-

rowly lanceolate or falcate, 15-85 x (l,5-)2,0-4,5 mm,

apex acuminate, base petiolate, occasionally trullate,

margins slightly revolute; petiole 1, 5-3,0 mm long. In-

florescences umbelliform, terminal, solitary, erect, 1-3

per plant, 4-9-flowered; peduncles 27-35 mm long.

Flowers ±1 X 1,1-1 ,4 mm, pink to light grey. Sepals

ascending, lobes lanceolate, 6, 6-6, 8 x 1, 2-2,0 mm.

Corolla ascending, petals lanceolate to ovate, connate at

base, 10,5-11,8 x ±5 mm, margins slightly revolute,

occasionally sinuate, central vein prominent, colour in-

side lilac-green, outside purple-green, abaxial surface

glabrous, adaxial surface canescent. Corona lobes aris-

ing from base of staminal column, erect, cucullate, sac-

cate, 3, 8-4, 2 x 1 ,7-2,0 mm, upper proximal ends form-

ing two obtuse almost triangular shoulders level with

middle of anther- wings, proximal margins concave to

20

Bothalia 17,1 (1987)

Anther-appendage length (mm)

FIGURE 4. — Histographic comparison of anther appendage length in Asclepias gordon-grayae, G; A. patens, P; and A, praemorsa, R.

lobe base and folded to meet, upper margin entire, curv-

ing to form a deltoid, obtuse distal lobe end. Keel dip-

ping near upper distal end, convex and narrow at lobe

base. Sinus a narrow central channel, 3, 8-4, 2 mm long,

with tongue-like appendage. Staminal column 3-4 mm

long; alar fissure 1,7-1 ,8 mm long; anther-wings

1,7-1, 8 x ± 0,7 mm, concave near apex with small

notch on margin; anther appendages tongue-like, ovate,

1,0-1, 3 x 1,1-1 ,2 mm, decumbent. Style apex trun-

cated to slightly conical, 2, 3-2, 8 mm wide. Pollinia

clavate to lacrimiform, 1,5-1 ,6 x ±0,56 mm; translator

arms 0,48-0,56 mm long; corpusculum 0,16-0,24 mm

wide. Figure 6.

NATAL. — 3126 (Queenstown): Queenstown (-DD), Acocks 17936

(PRE). 3225 (Somerset East): Cradock (-BA), Cooper 1284 (K, PRE).

3326 (Grahamstown): Alexandria (-CB), Acocks 17872 (PRE); South-

well (-DA), Bayliss 4631 (PRE). 3327 (Peddie): near East London

(-BB), Wood 1995 (K).

Although closely allied to Asclepias navicularis, A.

compressidens can be distinguished from it by a number

of important characters (Figure 6; Table 2). Also A.

navicularis has a distribution centred on the coastal re-

gion of the eastern Cape, whereas A. compressidens may

also be found further inland (Figure 7) and at higher

altitudes. The specific epithet refers to the compressed

tongue-like appendage that lies within the corona lobe

sinus. A. compressidens and A. disparilis N.E. Br. pos-

sess similar corona lobes, but in A. disparilis the distal

corona lobe appendage is slightly elongated and there is

a transverse shelf-like flap of tissue within the coronal

sinus just above the middle point of the lobe; the leaves

are also shorter and broader (Table 3). A. compressidens

is found at an altitude of between 30 to 140 metres and

flowers from November to February. It is found growing

in grassland and is said to be rare.

Key

Pollinia length

4 0,3-0.35

4 0,36-0,4

1 0,41-0,45

| 0,46 - 0,52

Corona lobe width

? 1,5 - 2,2

f 2,3 - 3,0

f 3,1 - 3,7

• 3,8 - 4,5

Alar fissure length Leaf width

• 0,5 -0,7

•- 0,71-1,1

1,2 -1,5

•— 1,6 - 2,0

-•1-8

-•9-17

-• 18-26

—•27-35

o A. patens

■ A. gordon-grayae

• A. praemorsa

(All measurements are in mm)

FIGURE 5. — Scatter diagram: a comparison of a number of characteristics in Asclepias gordon-grayae, A. patens and A. praemorsa.

Bothalia 17,1 (1987)

21

FIGURE 6. — Asclepias compressidens. 1, flowering plant, X 0,6, Acocks 17936 (PRE); 2b, flower, x 6,5, Rogers 12762a (BOL); 3,

pollinarium, x 120, Rogers 1276a (BOL). A comparison of the flowers of: 2a, A. navicularis X 8, Comins 1355 (PRE); 2b, A.

compressidens, x 6,5, Rogers 1276a (BOL).

22

Bothalia 17,1 (1987)

TABLE 1. — A comparison of the differences separating Asclepias gordon-grayae, A. patens and A. praemorsa

TABLE 2. — The main diagnostic differences between Asclepias compressidens and A. navicularis

Character

A. compressidens

A. navicularis

Corona lobe shape

Level of corona lobe in relation to

the style apex

Proximal corona lobe end in relation

to the distal end

Corona lobe sinus

Anther-wing (outer edge)

Notch on anther-wing margin

Pollinia

Shorter than the style apex

Lower than the distal end of the lobe

Tongue-like flap inside

Squared

Near the top

Claviform or lacrimiform

Level with the style apex

Level with the distal end of the lobe

Transverse flap-like shelf inside

Round

Near the base

± Rectangular

TABLE 3. — The main diagnostic differences found between Asclepias compressidens and A. disparilis

Character

A. compressidens

A. disparilis

Leaf length

Leaf width

Distal appendage of corona lobe

Corona lobe sinus

15-18 mm

2-4,5 mm

Not produced

Tongue-like projection

12-37 mm

3—9 mm

Slightly produced

Transverse shelf-like flap

Bothalia 17,1 (1987)

23

KEY TO ASCLEPI AS COMPRESSIDENS AND ITS CLOSEST ALLIES

la Corona lobe sinus without transverse flap of tissue near apex, compressed tongue-like appendage present

A. compressidens

lb Corona lobe sinus with transverse flap of tissue near apex, no compressed tongue-like appendage present:

2a Upper distal corona lobe edge level with style apex; leaf width at lamina base 1-5 mm A. navicularis

2b Upper distal corona lobe edge level with middle of anther-wing (not level with style apex) leaf width at

lamina base 3-9 mm a. disparilis

FIGURE 7. — Distribution of Asclepias compressidens.

ACKNOWLEDGEMENTS

I would like to thank the Directors and staff of the

following herbaria: BOL, CPF, GRA, J, K, NBG, NH,

NU, P, PRE, RUH, SAM and TCD for the loan of her-

barium specimens; the Director and staff of the Depart-

ment of Environment Affairs and the Head and staff of

the Department of Botany, University of Natal, Pieter-

maritzburg who made facilities available to me for the

study; Mr M. Lambert, Department of Classical Civili-

zation, University of Natal, Pietermaritzburg for help

with the Latin description of Asclepias gor don- gray ae.

REFERENCES

BROWN, N. E. 1908. Asclepiadeae. In W. T. Thiselton-Dyer, Flora

capensis 4,1: 518-1036. Reeve, London.

NICHOLAS, A. 1982. Taxonomic studies in Asclepias ( Asclepiada -

ceae) with particular reference to the narrow-leaved species in

southern Africa. M.Sc. thesis, University of Natal, Pietermaritz-

burg.

Bothalia 17,1:25-28(1987)

Two new brown subcrustose Parmelia species from southern Africa

(lichenized Ascomycetes)

F. BRUSSE*

Keywords: new species, lichenized Ascomycetes, Parmelia, southern Africa, subcrustose, taxonomy

ABSTRACT

Two new brown subcrustose species of Parmelia (Lichenes) are described from southern Africa on rock. They are P.

barda Brusse and P. princeps Brusse. The affinities of the two species are discussed.

UITTREKSEL

Twee nuwe bruin halfkorsagtige Parmelia- spesies (Lichenes) is op klippe uit suidelike Afrika beskryf. Die nuwe

spesies is P. barda Brusse en P princeps Brusse. Die verwantskappe van die twee spesies word bespreek.

Parmelia barda Brusse, sp. nov.

Thallus subcrustosus, saxicola, usque ad 50 mm

diametro. Lobi elongati, 0,2-0, 7 mm lati, usque ad 1,5

mm longi, 75-100 /xm crassi. Thallus superne brunneus,

nitidus, isidiis sorediisque nullis. Cortex superior 7-17

jum crassus, anticlinate prosoplectenchymatus, epicor-

tice poroso (Figure 2). Stratum gonidiale 10-35 /xm

crassum, fasciculatum, algis Trebouxiae, 5-22 /xm dia-

metro. Medulla alba, 20-80 /xm crassa. Cortex inferior

hyalinus, paraplectenchymatus, 9-13 /xm crassus. Thal-

lus inf erne pallidus. Rhizinae non bene evolutae. Apo-

thecia sessilia, numerosa, usque ad 0,8 mm diametro.

Hypothecium hyalinum, 15-30 /xm crassum. Subhyme-

nium 10-15 /xm crassum. Hymenium 44-55 /xm altum,

J+ caeruleum. Asci clavati, poricidales, cum tholis J+

caeruleis (Figure 3). Ascosporae octonae, hyalinae, sim-

plices, ellipsoideae, 7-10 x 4, 0-4, 5 /xm. Pycnidia glob-

osa, hyalina, circa 120 /xm diametro. Pycnidiosporae

hyalinae, aciculares, 7-10 x 0,8 /xm. Thallus atranori-

num (±) et materiam ignotam continens.

TYPE. — Cape, 3320 (Montagu): 4 km SW of Mon-

tagu, Kogmans Kloof near the old British Fort of 1899,

on top of an E-W ridge, on Table Mountain Sandstone

outcrops, alt. 200-250 m (-CC), Brusse 3718,

1981.05.12 (PRE, holo.). Figure 1.

Thallus subcrustose, saxicolous, up to 50 mm diam.

Lobes elongate, 0,2-0, 7 mm broad, up to 1,5 mm long,

75-100 /xm thick. Upper surface brown, non-isidiate

and non-sorediate. Upper cortex 7-17 /xm thick, anticli-

FIGURE 1. — Parmelia barda, habit.

Brusse 3718, holotype. Scale in

mm.

* Botanical Research Institute, Department of Agriculture and Water

Supply , Private Bag X 1 0 1 , Pretoria 000 1 .

26

Bothalia 17,1 (1987)

FIGURE 2. — Parmelia barda, SEM

of upper surface. Brusse 3718,

holotype. Bar = 6,3 /rm.

nally prosoplectenchymatous, epicortex pored (Figure

2). Algal layer clustered, clusters mainly subcortical, but

also scattered in the medulla, 10-35 /xm thick, algae

Trebouxia, 5-22 /xm diam. Medulla white, 20-80 /xm

thick. Lower cortex hyaline, paraplectenchymatous,

9-13 /xm thick. Lower surface pale. Rhizines not well

developed. Apothecia sessile, numerous, up to 0,8 mm

across. Hypothecium hyaline, 15-30 /xm thick. Subhy-

menium 10—15 /xm thick. Hymenium 45-55 /xm high,

J+ blue. Asci clavate, 8-spored, poricidal, tholus J+

blue (Figure 3). Ascospores hyaline, monolocular (but

equatorial ‘plasma-bridges’ common), ellipsoid, 7—10 x

4, 0-4, 5 /xm. Pycnidia hyaline, globose, about 120 /xm

diam. Pycnidiospores hyaline needles, 7-10 x 0,8 /xm.

Chemistry: Atranorin (±) and an unidentified substance

present (Table 1: bard-1).

TABLE 1. — TLC data for the two unidentified lichen substances.

using norstictic acid and atranorin as standards (Culberson 1972)

The rather narrow ascospores of this species are like

those of the genus Protoparmelia Choisy. Protoparmelia

differs from this species in many other respects even

though it is also brown. The paraphyses are not capitate

and capped, and the tholus iodine reaction differs signifi-

cantly from Protoparmelia. At present the degree of tho-

lus reaction variation is rather marked in Parmelia, but it

is not yet clear how much variation there is within one

species. If the iodine reaction pattern within the tholus

proves to be consistent, one could use these patterns for

generic segregation of the genus Parmelia, particularly if

these correlate with the presently used thalline and chem-

ical characters. The thallus anatomy of Protoparmelia

badia, the type of Protoparmelia (Hafellner 1984) is also

divergent from that of this new species. It was

therefore thought best to place this species in the genus

Parmelia. In fact the cortex reacts green with concen-

Bothalia 17,1 (1987)

27

FIGURE 5. — Parmelia princeps,

SEM of upper surface. Brusse

3074, holotype Bar = 5,9 pm.

FIGURE 4. — Parmelia princeps,

habit. Brusse 3074, holotype.

Scale in mm.

trated nitric acid, and can be considered a small subcrus-

tose member of the Neofuscae (Esslinger 1977), despite

the additional presence of atranorin. P. bar da was tested

twice for chemical contents, the second run revealed no

atranorin. The type specimen is growing on a rock with

two micro- aspects and this may explain this difference.

More exposed positions usually support specimens with

a greater quantity of a particular lichen substance.

Although the substance in the medulla was not identi-

fied further, the spot characteristics indicated that it may

be an orcinol para-depside. The Rf data for ‘bard-1’ are

similar to those given for loxodellic acid (Esslinger

1977, p. 19), but this could not be confirmed by direct

comparison, for lack of a lichen at PRE containing loxo-

dellic acid, or a confirmatory test such as a micro-crystal

test (no published data). Nevertheless all the known sub-

crustose species of the section Neofusca contain depsi-

dones, with the exception of the New Zealand endemic,

P. minuta Essl. which contains gyrophoric acid

(Esslinger 1977).

At present this species is known only from the type

specimen from Kogmans Kloof near Montagu.

Parmelia princeps Brusse, sp. nov.

Thallus subcrustosus, saxicola, usque ad 25 mm

diametro. Lobi elongati, 0, 1-0,3 lati, usque ad 1 mm

longi, 80-95 pm crassi. Thallus superne atro-brunneus,

nitidus, isidiis sorediisque destitutus. Cortex superior

7-10 /im crassus, epicortice poroso (Figure 5). Stratum

gonidiale 15-35 pm crassum, algis Trebouxiae, 5-18

pm diametro. Medulla alba, 20-45 pm crassa. Cortex

inferior paraplectenchymatus, circa 15 pm crassus,

brunneus. Thallus inf erne piceus. Rhizinae non bene

28

Bothalia 17,1 (1987)

evolutae. Apothecia sessilia, rara, usque ad 0,5 mm

diametro. Hypothecium hyalinum, 30-33 /zm crassum,

paraplectenchymatum, cellulis 3, 5-7, 5 /zm diametro.

Subhymenium 30-45 /zm crassum. Hymenium hyalinum,

50-60 /zm altum, J+ caeruleum. Asci clavati, porici-

dales, cum tholis J+ caeruleis (Figure 6). Ascosporae

hyalinae, simplices, ellipsoideae, 9,5-12,5 x 5, 0-6,0

/zm. Pycnidia non visa. Thallus acidum squamaticum

continens.

TYPE. — Cape, 3218 (Clanwilliam), 3 km W of Oly-

venboskraal, Witelskloof, on large Table Mountain

Sandstone boulder-outcrop, alt. 450-500 m (-BD), F.

Brusse 3074, 1981.05.02 (PRE, holo.). Figure 4.

Thallus subcrustose, saxicolous, up to 25 mm across.

Lobes elongate, 0, 1-0,3 mm broad, up to 1 mm long,

80-95 /zm thick. Upper surface dark brown, glossy,

non-isidiate and non-sorediate. Upper cortex 7—10 /zm

thick, epicortex pored (Figure 5). Algal layer 15-35 /zm

thick, algae Trebouxia, 5-18 /zm diam. Medulla white,

20-45 /zm thick. Lower cortex paraplectenchymatous,

around 15 /zm thick, brown. Lower surface black. Rhi-

zines not well developed. Apothecia sessile, rare, up to

0,5 mm across. Hypothecium hyaline, 30-33 /zm thick,

paraplectenchymatous, cells 3, 5-7, 5 /zm diam. Sybhy-

menium 30-45 /zm thick. Hymenium hyaline, 50-60 /zm

high, J+ blue. Asci clavate, poricidal, tholus J+ blue

(Figure 6). Ascospores hyaline, monolocular, ellipsoid,

9,5-12,5 x 5, 0-6,0 /zm. Pycnidia not seen. Chemistry:

cortical substances absent, squamatic acid and an un-

identified substance (Table 1: princ.-l) in the medulla.

Parmelia princeps is another brown lichen that is not

assignable to Protoparmelia Choisy, presently used for

the Lecanora badia group, due to important structural

differences (see the discussion of the previous species for

these). The cortex of this species does not react with

concentrated nitric acid (nor with 2 molar potassium hy-

droxide solution), but could probably still be assigned to

the Neofuscae, as some members of this group are nega-

tive with concentrated nitric acid (Esslinger 1977). The

only other subcrustose species containing squamatic acid

is Parmelia squamatica Brusse (1980, 1986) but this is

yellow-green coloured, with usnic acid in the upper cor-

tex.

Parmelia princeps is presently known only from the

type locality south-west of Clanwilliam.

ACKNOWLEDGEMENTS

The author is grateful to Mrs S. M. Perold for assis-

tance with the Scanning Electron Micrographs, Mrs A.

J. Romanowski for the photographs and Mrs E. L. Bun-

ton for the typing.

REFERENCES

BRUSSE, F. A. 1980. A taxonomic and geographic study of the genus

Xanthoparmelia in the Karoo. M.Sc. thesis. University of the

Witwatersrand, Johannesburg.

FIGURE 6. — Parmelia princeps, ascus and

paraphysis. Brusse 3074, holotype. Bar

= 10 pm.

BRUSSE, F. A. 1986. Four new effigurate-crustose species of Parme-

lia (Lichenes, Parmeliaceae) from southern Africa. Mycotaxon

27: 237-245.

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

identification of lichen products by a standardized thin-layer chro-

matographic method. Journal of Chromatography 72: 113-125.

ESSLINGER, T. L. 1977. A chemosystematic revision of the brown

Parmeliae. Journal of the Hattori Botanical Laboratory. 42:

1-211.

HAFELLNER, J. 1984. Studien in Richtung einer naturlicheren

Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae.

Beihefte zur Nova Hedwigia 79: 24 1-37 1 .

Bothalia 17,1: 29-43(1987)

Notes on African plants

VARIOUS AUTHORS

ASCLEPIADACEAE

NOTES ON ASCLEPIAS D1PLOGLOSSA , A . COGNAT A AND A. FLAV A

A.jlava N.E.Br. and A. cognata N.E.Br. have in the

past been erroneously considered conspecific with A.

diploglossa (Turcz.) Druce. Although allied, these three

taxa are morphologically distinct.

South African herbaria have, for some years, treated

A.jlava N. E. Br. and A. cognata N.E.Br. as synonyms

of the older A. diploglossa (Turcz.) Druce. This is due to

the misinterpretation of a letter sent from Kew to the

Botanical Research Institute in 1963. Marais, who was

then the South African Liaison Officer at Kew, mentions

in this letter that ‘Mr Bullock regards both A. flava

N.E.Br. and A. cognata N.E.Br. as conspecific with A.

diploglossa' . He does, however, state earlier in the same

letter ‘As far as I know A. flava N.E.Br. has not been

sunk under A. diploglossa’ . This was, and still is the

case as Bullock did not publish his opinions. Despite

Marais’s warning that these reductions to synonomy

were not validly published, Bullock’s suggestions were

unfortunately followed and to all intents and purposes A

flava and A. cognata became synonyms under A. diplog-

lossa. This situation has recently been rectified in Gibbs

Russell et al. (1984) and further details are given here.

Although vegetatively similar, close examination

shows all three species to be distinct taxonomic entities

with very different floral morphologies (Figures 1 & 2;

Table 1).

KEY TO ASCLEPIAS DIPLOGLOSSA AND ITS ALLIES

la Proximal corona lobe appendages projecting over style apex A. cognata

lb Proximal corona lobe appendages not projecting over style apex:

2a Corona lobes with arm-like proximal appendages crossing over each other and reflexing back into coronal

sinus, appendages are below style apex and level with anther-wings A. flava

2b Corona lobes without true proximal appendages, proximal ends of corona forming dentate to obtuse

(but protruding) shoulders level with top of style apex A. diploglossa

Representative specimens

Asclepias diploglossa (Turcz.) Druce

NATAL. — 2730 (Vryheid): Wakkerstroom (-AD), Devenish 759

(NH, PRE). 2828 (Bethlehem): Royal Natal Park (-DB), Acocks &

Hafstrom 1 1205 (PRE). 2829 (Harrismith): Van Reenen (-AD), Wood

11205 (PRE). 2929 (Underberg): Giants’ Castle (-AB), Symons 188

(PRE); Highmoor State Forest (-BC), Killick & Vahrmeijer 3583

(PRE); Sam Pass (-CB), Lugillies 109 (NU); Bushman’s Nek (-CC),

Killick & Vahrmeijer 3969 (PRE); Garden Castle Nature Reserve

(-CD), Hilliard & Burn 7866 (NU); Mpendle (-DB), Moll 1481

(PRE); Glengariff (-DD), Rennie 488 (NU). 2930 (Pietermaritzburg):

Greytown (-BA), Wylie s.n. (NH 21644); Inanda (-DB), Groom s.n.

(NH4106).

TRANSKEL— 3028 (Matatiele): Ramatseliso’s Gate (-BB), Board-

man 186 (PRE).

CAPE. — 3227 (Stutterheim): Hogsback (-CA), Compton 1421

(NBG); Dohne Hill (— CB), Sim 1237 (SAM). 3326 (Grahamstown):

near Grahamstown (-BC), Glass 1503 (SAM).

Asclepias flava N.E.Br.

NATAL. — 2929 (Underberg): Underberg (-CD), Dyer 3744 (NH);

Sunset (-DA), Rennie 275 (NU); Mpendle (-DB), Huntley 625 (NH);

Nkonzo State Forest (-DD), Nicholas & Norris 1159 (CPF, NH,

PRE). 2930 (Pietermaritzburg): Howick (-AC). Haygarth s.n. (SAM

8378)\ Blinkwater Range (-AD), Nicholas 1019 (NU); Zwaartkop

(-CB), Wood 11219 (SAM); Byrne (-CC), Stewart 2023 (NU); Dargle

(-DB), Nicholas & George 1040 (CPF, K, MO, NH, NU). 3030 (Port

Shepstone): Ixopo (-AA), Shirley s.n. (NU).

TRANSKEL— 3029 (Kokstad): Mt Cume (-AD), Tyson 1686

(BOL, PRE); Malowe (-BD), Tyson 1086 (BOL, SAM); Vaal Bank

(-CB), Haygarth s.n. (NH 4230)', Ngeli Forest (-DA), Coleman 813

(PRE); Fort Donald (-DC), Tyson 1660 (SAM). 3128 (Umtata): Bazija

(— CB), Baur 556 (SAM).

CAPE.— 3326 (Grahamstown): Grahamstown (-BC), Glass 1503

(NBG).

Asclepias cognata N.E.Br.

NATAL. — 2929 (Underberg): Fort Nottingham (-BD), Hilliard &

Burn 9056 (NU); Bamboo Mountain (-CB), Grice s.n. (NU); Garden

Castle Nature Reserve (-CD), Hilliard & Burn 13767 (NU); Mpendle

(-DB), Hilliard & Burn 13856 (NU).

TRANSKEL — 3029 (Kokstad): Ensikeni (-BA), Haygarth s.n. (NH

15735); Mount Insizwa (-CD), Schlechter 6496 (BOL, NH); Ngeli

Mountain (-DA), Hilliard & Burn 5769 (NH, NU).

CONCLUSIONS

A. flava and A. cognata have never been validly re-

duced to synonomy under A. diploglossa and are, as

outlined by Brown (1908), distinct species allied to, but

separable from A. diploglossa.

ACKNOWLEDGEMENTS

I would like to thank the Director and staff of the

following herbaria: BOL, CPF, GRA, J, K, NBG, NH,

NU, P, PRE, RUH, SAM and TCD for the loan of her-

barium specimens; the Head and staff of the Department

30

Bothalia 17,1 (1987)

FIGURE l.—- Comparison of flower (1) and corona lobe (2) of Asclepias species. Structure in: la, A. cognata, x 10, Hilliard & Burn 9056 (NU); lb,

A. diploglossa, x 20, Boardman A 1 1 (PRE); lc, A. flava, x 19, Coleman 813 (PRE); 2a, A. cognata, x 17, Hilliard & Burn 9056 (NU); 2b, A.

diploglossa, x 28, Boardman All (PRE); 2c, A. flava, x 38, Coleman 813 (PRE).

Bothalia 17,1 (1987)

31

TABLE 1. — A comparison of Asclepias cognata, A. diploglossa and A. flava. All measures in mm

FIGURE 2. A comparison of corona lobe variation. I, Asclepias cognata: a, Hilliard & Burtt 5769 (NH); b, Haygarih s.n. (NH 13661 ); c .Hilliard

& Burtt 13452 (NU); d, Hilliard & Burtt 13767 (NU); e, Haygarth s.n. (NH 15735); f, Hilliard & Burtt 13856 (NU). 2, A. diploglossa: a, Grice

s.n. (NU); b, Wylie s.n. (NH 21644); c, Devenish 759 (PRE); d, Rennie 488 (NU);e, Wood 11205 (PRE); f, Barker 1421 (NBG). 3, A. flava: a,

Hilliard & Burtt 5839 (NU); b, Baur556 (SAM); c, Glass 1503 (NBG); d, Tyson 1660 (SAM); e, Tyson 1686 (BOL); f. Wood 11219 (SAM).

32

Bothalia 17,1 (1987)

of Botany, University of Natal, Pietermaritzburg; and

the Director and staff of the Department of Environment

Affairs.

REFERENCES

BROWN, N. E. 1908. Asclepiadeae. In W. T. Thiselton-Dyer, Flora

capensis 4,1: 687-688. Reeve, London.

GIBBS RUSSELL, G. E., GERMISHUIZEN, G., HERMAN, P.,

OLIVIER, P., PEROLD, S. M., REID, C., RETIEF, E.,

SMOOK, L., VAN ROOY, J., WELMAN, W. G. & GON-

SALVES, P. 1984. List of species of southern African plants.

Memoirs of the Botanical Survey of South Africa No. 48.

NICHOLAS, A. 1982. Taxonomic studies in Asclepias L. ( Asclepiada -

ceae) with particular reference to the narrow-leaved species in

southern Africa. M.Sc. thesis, University of Natal, Pietermaritz-

burg.

A. NICHOLAS

ERYSIPHACEAE

TWO NEW OIDIUM SPECIES FROM THE TRANSVAAL

In January 1985 an Oidium species was observed on

leaves of Dombeya rotundifolia (Hochst.) Planch, in the

garden of the Botanical Research Institute, Brummeria,

Pretoria. The fungus was observed until leaf drop but no

teleomorph was produced. The only previous report of

an Oidium sp. occurring on a Dombeya sp. is from Natal

in the Camperdown area of Pietermaritzburg, viz. on D.

cymosa Harv. (Doidge 1950). However, this fungus was

not described and, although the fungus on both Dombeya

species is likely to be the same, this cannot be stated for

certain. As no fresh material of the Natal fungus could be

obtained it is absolutely necessary to have a detailed

description of the Transvaal fungus available for future

comparison. The fungus from D. rotundifolia is there-

fore, described here in detail.

A few months later at the same venue an Oidium

species was observed on Lannea discolor (Sond.) Engl,

at the end of its growing season. Quite a number of

powdery mildew fungi have been observed on Anacar-

diaceae, the plant family to which Lannea spp. belong.

Apart from two Phyllactinia spp., one Uncinula sp., a

Sphaerotheca sp., two Erysiphe spp. and one Macro-

sphaera sp. which have been recorded (Hirata 1966),

two Oidium spp. have been described, viz. 0. anacardii

Noack (1898) and 0. mangiferae Berthet (1914).

Identity of our Oidium species with Sphaerotheca

macularis (Wallr. ex Fries) Magnus and Erysiphe cicho-

racearum DC. can be excluded immediately on account

of the solitary production of conidia in the Lannea fun-

gus, while Phyllactinia spp. do not have an Oidium

stage. Moreover, no imperfect stage has apparently been

observed for Uncinula verniciferae P. Henn. (Salmon

1905). Identity with Microsphaera alni (Wallr.) Winter,

Erysiphe communis (Wallr.) Link and Oidium mangife-

rae must also be excluded on account of the flexuous or

bent foot cells of the conidiophores. This leaves 0. ana-

cardii which has conidia similar in size and shape to the

Lannea fungus. However, its description by Noack

(1898) does not mention whether the foot cells of the

conidiophores are straight or flexuous. In addition the

presence or absence and shape of appressoria and fibro-

sin bodies is also not recorded. This and the apparent

absence of Oidium anacardii from the African continent,

as well as the fact that the Lannea fungus occurs on a

previously unrecorded host of powdery mildew, makes it

desirable to describe this fungus as a new species.

Oidium dombeyae Gorter & Eicker, sp. nov.

Mycelium superficiale, albidum, effusum. Hyphae

hyalinae, subflexuosae, aliquando geniculatae. Cellulae

hypharum, 35-45 x 4-5 /xm. Appressoria non lobata

vel moderate lobata. Conidiophora cylindracea (70—)

90—1 15(— 130) x 7,5-10 /am, plerumque 2-septata. Cel-

lulae basales rectae, comparate longae, (50— )65— 90

(-110) x 7,5-10 pm; cellulae sequentes breviores,

(7,5— )12,5(— 17,5) x 7,5-10 gm. Conidia ovoidea, soli-

taria, (22,5-)27,5-30(-35) x (15-)17,5(-20) pm. Cor-

puscula fibrosina conspicue desunt. Ratio longitudinis

/latitudinis conidiorum circa 1,6. Tubi germinationis,

apicales vel subapicales, plerumque perbreves, interdum

autem longiores, 12,5-125 x 2, 5-5,0 pm, saepe in

appressorium non lobatum vel modice lobatum (7,5-15

x 5, 0-7, 5 pm) terminantes.

Habitat in foliis vivis Dombeyae rotundifoliae

(Hochst.) Planch., Brummeria, Pretoria. Ianuarius 1985,

PREM No. 47848.

Mycelium superficial, white, thinly spread. Hyphae

hyaline, slightly flexuous, sometimes geniculate. Hy-

phal cells, 35—45 x 4-5 pm. Appressoria unlobed or

moderately lobed. Conidiophores cylindrical, (70-)

90—1 15(— 1 30) x 7,5-10 pm, usually 2-septate. Foot

cells straight, comparatively long, (50— )65— 90(— 1 10) x

7,5-10 pm, followed by one or more shorter cells

(7,5— )12,5(— 17,5) x 7,5-10 pm. Conidia singly pro-

duced, ovoid, (22 ,5-)27 ,5-30(-35) x (15-)17*5(-20)

pm; length/width ratio ± 1,6. Germ tubes near end of

conidia, apical or subapical, usually very short but some-

times longer, 12,5-125 x 5, 0-7, 5 /am, often ending in

an unlobed or moderately lobed appressorium (7,5-15 x

5,0-7, 5 pm).

On leaves of Dombeya rotundifolia (Hochst.) Planch.,

Pretoria Botanical Garden, Brummeria. January 1985,

PREM No. 47848.

Oidium lanneae Gorter & Eicker, sp. nov.

Mycelium amphigenum, griseo-albidum, effusum vel

densum. Hyphae plus minus vae rectae, aliquando

flexuosae vel geniculatae, rectangulariter ramificatae

saepe prope septum. Cellulae hypharum (37,5-)50-60

(-70) x 5, 0-6, 2 pm. Appressoria moderate lobata vel

multilobata, aliquando binatim opposita. Conidiophora

numerosa, brevia, (l-)3(-4)-cellularia (30-)50-75

(-110) x (7,5— )8,7— 1 1 ,2 pm, saepe ab basi ad apicem

leviter dilatescentia. Cellulae basales geniculatae,

flexuosae vel curvatae, raro rectae, (25— )35— 55(— 90) x

7,5-10 pm. Conidia solitaria, ellipsoidea vel oblonga,

utrinque obtuse rotundata, (25— )30— 35(^40) x

( 1 3,7—) 1 5— 1 7 ,5(— 20) pm. Ratio longitudinis/latitudinis

conidiorum circa 2. Corpuscula fibrosina conspicue

desunt. Tubi germinationis prope apicem orientes, inter-

dum apicales, circa recti, 50-75 x 3^J pm, saepe in

appressorium non lobatum vel modice lobatum 5, 0-7, 5

pm latum terminantes. Aliquando tubi perbreves rema-

nentes ac in appressorium multilobatum terminantes.

Bothalia 17,1 (1987)

33

Habitat in foliis vivis Lanneae discoloris (Sond.)

Engl., Brummeria, Pretoria. Maius 1985, PREM Nos.

47877, 47974.

Mycelium amphigenous, greyish white, effuse to

dense. Hyphae more or less straight, occasionally

flexuous or geniculate, branching at right angles often

near a septum. Hyphal cells (37,5-)50-60(-70) x 5-6,2

pm. Appressoria moderately lobed or multilobed, some-

times opposite in pairs. Conidiophores numerous, short,

( 1— )3(— 4)-celled , (30-)50-75(-l 10) x (7,5-)8,7-l 1 ,2

pm, often slightly widening from base to top. Foot cells

geniculate, flexuous or bent, seldom straight, (25— )35

— 55(— 90) x 7,5-10 pm. Conidia singly produced, ellip-

soid to oblong, obtusely rounded at both ends,

(25— )30— 3 5 (—40) x (13,7)15-17,5(-20) pm\ length/

width ratio ± 2. No well developed fibrosin bodies are

present. Germ tubes originate near end of conidia,

usually apically, nearly straight, 50-75 x 3-4 pm, often

ending in an unlobed or moderately lobed appressorium,

5, 0-7, 5 pm wide; tubes sometimes remain very short,

ending in a multilobed appressorium.

On leaves of Lannea discolor (Sond.) Engl., Pretoria

Botanical Garden, Brummeria. May 1985. PREM Nos.

47877,47974.

Although referring and describing imperfect forms of

ectophytic powdery mildews to the form genus Oidium

does not find favour with some mycologists it is a

necessary procedure in tropical and subtropical regions

of the world where the teleomorph of many mildew fungi

is seldom produced (Doidge 1915; Hansford 1946) and

where powdery mildews in most cases can only be iden-

tified on the basis of anamorphic characters (Clare 1964;

Boesewinkel 1977; Hammett 1977).

ACKNOWLEDGEMENTS

Financial support from the Department of Agriculture

and Water Supply and the University of Pretoria is grate-

fully acknowledged.

REFERENCES

BERTHET, J. A. 1914. Molestia da mangueira. Boletim de agricul-

tura. Sao Paulo 15: 818-819.

BOESEWINKEL, H. J. 1977. Identification of Erysiphaceae by coni-

dial characters. Revue de mycologie 41: 493-507.

CLARE, B. G. 1964. Erysiphaceae of south-eastern Queensland.

Papers from the Department of Botany, University of Queensland

4, 10-11: 111-114.

DOIDGE, ETHEL M. 1915. Some notes on the South African Erysi-

phaceae. Transactions of the Royal Society of South Africa 5:

237-245.

DOIDGE, ETHEL M. 1950. The South African fungi and lichens to

the end of 1945. Bothalia 5: 1-1094.

HAMMETT, K. R. W. 1977. Taxonomy of Erysiphaceae in New

Zealand. New Zealand Journal of Botany 15: 687-711.

HANSFORD, C. G. 1946. The foliicolous ascomycetes, their parasites

and associated fungi. Mycological Papers 15: 1-240.

HIRATA, K. 1966. Host range and geographical distribution of the

powdery mildews. Mimeographed edn. Faculty of Agriculture,

Niigata University, Niigata.

NOACK, F. 1898. Congumelos parasitas das plantas di pomar, horta e

jardim. Boletim do Instituto agronomico do Estado de Sao Paulo

9: 75-88.

SALMON, E.S. 1905. The Erysiphaceae of Japan, H. Annales mycolo-

gici 3: 241-256.

G. J. M. A. GORTER* AND A. EICKER*

* Botany Department, University of Pretoria, Pretoria 0002.

FABACEAE

A NEW SPECIES OF INDIGOFERA FROM NATAL AND TRANSKEI

Indigofera rubroglandulosa Germishuizen, sp.

nov., I. hilaris Eckl. & Zeyh. similis sed dense tecta

pilis glandulosis, etiam lobis calycis longioribus, foliolis

latioribus atque caudice globoso tuberoso differt.

TYPE.— Natal, 3030 (Port Shepstone): Margate

(-CD), Strey 10990 (PRE, holo.!; NH, iso. !). Figure 3.

Small erect to semi-erect herb up to 0,3 m tall arising

from a woody base; rootstock tuberous, globose. Stems

reddish brown, longitudinally ridged, with scattered

biramous hairs densely interspersed with red glandular

hairs or occasionally with only few scattered biramous

hairs. Stipules free, adnate to base of petiole, up to 7 mm

long, glandular on outside, glabrous inside. Leaves regu-

lar, paripinnate, petiolate. Leaflets 3-7 , obovate to nar-

rowly elliptic to oblanceolate, (7—) 1 2,5—17 (—20) x

(3,5— )5— 9(— 1 1) mm, stiffly mucronate at apex, obtuse at

base, strigillose on both surfaces with biramous hairs,

red glandular-hairy on lower surface only and along mar-

gins. Petiole up to 4 mm long. Inflorescence axillary,

racemose, 4-10-flowered, up to 40 mm long, including a

10-20 mm long peduncle, covered with both biramous

and red glandular hairs. Flowers deep pink to red, 8-9

mm long, each subtended by a lanceolate caducous bract

2-3 mm long. Calyx: tube 2 mm long, lobes 3,5- 6 mm

long, glabrous inside, outside with both biramous and

glandular hairs. Standard 7-9 x 5-7,5 mm, broadly

ovate, narrowing to base, mucronate at apex, strigose

outside, glabrous inside. Wing petals 8x2 mm. Keel

blades 8-9 x 2-2,5 mm, prominently pocketed; pockets

more or less triangular, up to 2 mm long. Style 7 mm

long, upcurved for about 2 mm, white-strigose for two-

thirds of its length; stigma capitate. Stamens 5-7 mm

long; 9 fused into a staminal sheath and the 1 vexillar

stamen free; anthers uniform, 1 mm long, strongly api-

culate; apiculum up to 0,5 mm long. Fruit a cylindrical

pod, 20-25 mm long, up to 3 mm in diameter, reddish

brown, white-strigose interspersed with red glandular

hairs. Seeds 1 ,5 mm long and wide.

34

Bothalia 17,1 (1987)

NATM5E HERBARIUM DURBAN

FIGURE 3. — Holotype of Indigofera

rubroglandulosa Germishuizen

(Strey 10990).

FIGURE 4. — Distribution of Indigofera rubroglandulosa.

Location 5 (-CD), Strey 8729 ; Uvongo (-CD), Stirton 8057. 3130

(Port Edward): Beacon Hill (-AA), Strey 65 10.

TRANSKEI. — 3129 (Port St Johns): Mkambati Game Reserve

(-BD), Shackelton 89\ 5 km from Mkambati on road to Mtontsasa

(-BD), Van Wyk 1531\ Mkambati Leper Institute, along road from

hospital to office (-BD), Marais 1170\ 3 km from Mkambati hospital

(-BD), Venter 829.

I. rubroglandulosa is found in Natal and Transkei

(Figure 4) in grassland recently burnt. The first record of

the species in the PRE Herbarium was collected by

H. W. Bellmarley s.n. (PRE 58841), during July 1927

from Melmoth. It flowers in late winter to early summer.

All the material cited above was incorrectly identified as

I. hilaris or as Indigofera species. Although the distribu-

tion areas of I. hilaris and I. rubroglandulosa overlap,

the latter can be distinguished from the former by its

longer calyx lobes, wider leaflets and the red glandular

hairs. With the aid of a scatter diagram (Figure 5) using

calyx length along one axis and width of leaflets along

the other axis, /. hilaris and I. rubroglandulosa are read-

ily separated.

NATAL. — 2831 (Nkandla): 6 km south of Mtonjaneni on road to

Melmoth (-AD), Codd 1803\ Melmoth (-CB), Bellmarley s.n. 2930

(Pietermaritzburg): Camperdown (-DA), Moll 1886. 3030 (Port

Shepstone): Oribi Gorge, ‘Fairacres’ (-CB), Van Wyk 5127\ Umtam-

vuna Nature Reserve (-CC), Abbott 223 ; Izingolweni-Port Edward

road (-CC), Ward 185\ Margate (-CD), Strey 10990 ; Uvongo Bantu

ACKNOWLEDGEMENTS

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

diagnosis into Latin.

G. GERMISHUIZEN

Bothalia 17,1 (1987)

35

A I r ub ro g I and u los a

A

▲ I hilaris

A

A

A

6-

A

A

A

A

A

A

A

FIGURE 5. — Scatter diagram of

calyx length and leaflet width of

2 3 o 5 6 7 e mm Indigofera rubroglandulosa and

CALYX LENGTH /. kUdriS.

LICHINACEAE

A NEW SPECIES OF GONOHYMENIA FROM ETOSHA PAN LIMESTONE

Gonohymenia etoshica Brusse, sp. nov.

Thallus peltatus, calcicola, usque ad 3 mm diametro,

madefactus 240-320 pm crassus, homoeomerus. Pagina

superior carbo-atra, hebetata, undulata vel cerebrifor-

mis, ambitu breve lobata. Pagina inferior carbo-atra,

hebetata, laevis vel grosse et radiatim plicata. Apothecia

immersa vel substipitata, lecanorina, thallinocarpa,

usque ad 0,7 mm diametro. Excipulum thallinum in late-

ribus 70-150 pm crassum. Excipulum hyalinum, J-,

8-12 pm crassum, periclinate plectenchymatum, in late-

ribus destitutum. Hymenium hyalinum, in dimidio

superiori cum algis inspersum, 70 pm altum, J+ caeru-

leum. Hypothecium tenue, fere nullum. Paraphyses gra-

ciles, simplices vel leviter ramosae (in regionibus alga-

rum bene ramosae), luminibus 0, 8-1,0 pm crassis, sep-

tatae, gelatinosae, gelatina J+ caerulea. Asci obclavati,

8-, 12- vel 16-spori, parietibus omnibus tenuibus, J— ,

25-40 X 9-14 pm. Ascosporae hyalinae, ellipsoideae,

6-7 x 3 ,5 — 4,5 pm, simplices. Pycnidia globosa, hya-

lina, circa 50 pm diametro. Spermatia bacillaria,

2,8-3 ,6 x 0, 8-1,0 pm.

TYPE. — 1916 (Gobaub): Etosha Pan, 28 km from

Halali to Okaukuejo, Salvadora, on littoral limestone

outcrops — gentle N slope (-AB), Brusse 4155,

1984.03. 19 (PRE, holo.; LD, iso.). Figure 6.

Thallus peltate, on limestone, up to 3 mm diam.,

homoeomerous, 240-320 pm thick when wet (Figure 7).

Upper surface charcoal, matt, undulate to cerebriform-

wrinkled, simple to shortly lobate at margins. Lower

surface charcoal, matt, smooth to coarsely radiate pli-

cate. Apothecia (Figure 8) immersed to substipitate,

lecanorine, thallinocarpous, up to 0,7 mm diam. Thai-

line exciple 70-150 pm thick on sides. Exciple hyaline,

J-, 8—12 pm thick, dense periclinal plectenchyma,

absent from margins (Figure 8). Hymenium hyaline,

inspersed with algae in upper half, 70/xm high, J+ blue.

Hypothecium thin, almost absent. Paraphyses slender,

simple or lightly branched (well branched in algal areas),

lumens 0,8—1 ,0 pm thick, septate, gelatinized, gel J+

blue. Asci obclavate, 8-, 12- or 16-spored, walls thin,

J— (Figure 9), 25-40 X 9-14 pm. Asco spores hyaline,

ellipsoid, 6-7 x 3,5— 4,5 pm, simple. Pycnidia hyaline,

globose, about 50 /am diam. Spermatia hyaline rods,

2, 8-3, 6 x 0, 8-1,0 pm.

This new species is most similar to the European

species, Gonohymenia nummularia (Nyl.) Henss., but

has smaller and fewer ascospores per ascus.

The number of algae in the upper half of the hyme-

nium seems to vary considerably. Sometimes algae are

quite scarce, e.g. in the specimen cited below, but the

type is rich in hymenial algae. The asci of this species

are obclavate with acute apices and with thin walls (no

tholus development). However the hymenial gel is blue

in Lugol’s iodine solution. The asci containing 12 or 16

spores are larger than those containing only 8 asco-

spores, so that the ascospores from 8-spored asci are of

the same overall size as those from 16-spored asci. It is

36

Bothalia 17,1 (1987)

FIGURE 6. — Gonohymenia etoshica

Brusse, habit. Brusse 4155, ho-

lotype. Scale in mm.

V % 88 **>

5* ' »

7i* '

* S»

«» V/

**

• * '

if •*" , . ...

- - X

• W >:.8H fji' . «■■

*•■.' ■ » <-% iff* . A.-& '

%*

. #

; a*/

«# *»a

4 «* %»

*» * ■

v&

**>» ■

>4,

.7 ? ■"

'0k

*

FIGURE 7. — Gonohymenia etoshica

Brusse, edge of lobe section in

lactophenol cotton-blue, show-

ing undifferentiated hyphae an-

ticlinal to the surface, and algal

cells concentrated near the sur-

face. Brusse 4155, holotype.

Bar = 100 /urn.

FIGURE 8. — Gonohymenia etoshica

Brusse, part of an immersed

apothecium in section in lacto-

phenol cotton-blue. Brusse

4155, holotype. Bar =100 /urn.

Bothalia 17,1 (1987)

37

FIGURE 9. — Gonohymenia etoshica Brusse, ascus

and paraphyses. Brusse 4155, holotype. Bar =

10 pm.

interesting that the exciple terminates before reaching the

flanks of the hymenium (Figure 8), and probably allows

for intrusion of alga-bearing thalline tissue into the

hymenium, during development.

Several new combinations in this genus were made by

Henssen (1980), where good photographs of some

species are given. Poelt (1969) treated the European

species, some under Thyrea, in key form.

At present this species is known from two localities on

the edge of Etosha Pan, on limestone.

SWA/NAMIBIA. — 1816 (Namutoni): Etosha Pan, 8 km N of

Namutoni, on Pan’s Edge road, on limestone on W rise (-DD), F .

Brusse 4177, 1984.03.20 (PRE, LD).

ACKNOWLEDGEMENTS

The author is grateful to game rangers Karen and Tre-

vor Nott for company during a stay at Etosha Pan, and to

Otavi Tsumeb for armed protection against dangerous

wild animals.

REFERENCES

HENSSEN, A. 1980. Problematik der Gattungsbegrenzung bei den

Lichinaceen. Berichte der Deutschen Botanischen Gesellschaft 92:

483-506.

POELT, J. 1969. Bestimmungsschliissel europaischer Flechten.

Cramer, Lehre.

F. BRUSSE

A NEW SPECIES OF THYREA FROM OTAVI DOLOMITE (DAMARA SYSTEM)

Thyrea otaviana Brusse, sp. nov.

Thallus subfruticosus, e haptero radiatim lobatus, in

sicco usque ad 18 mm diametro, dolomiticola. Lobi

subascendentes, lineares, simplices vel ramosi, usque ad

10 mm longi et 0,4-1 ,5 (-2,5) mm lati, madefacti

400-600 pm crassi. Pagina superior carbo-atra, hebe-

tata vel granulate-isidiosa, laevis. Cortex 5-15 pm cras-

sus, paraplectenchymatus, cellulis 3, 5^4, 5 pm dia-

metro. Stratum gonidiale 20-50 pm crassum; algae

cyanescentes (Chroococcales aut Pleurocapsales).

FIGURE 10. — Thyrea otaviana

Brusse, habit. Brusse 4163, ho-

lotype. Scale in mm.

38

Bothalia 17,1 (1987)

Medulla hyalina, 390-560 pm crassa; hyphae in reticu-

lum regulare dispositae, 1,4-2, 5 pm crassae. Pagina

inferior carbo-atra, hebetata, laevis, in sicco canalicu-

lata. Apothecia laminalia, sessilia, lecanorina, in sicco

usque ad 0,5 mm diametro. Pagina hymeniorum badia,

nitida. Excipulum thallinum in lateribus 80-150 pm

crassum. Excipulum hyalinum, periclinate prosoplecten-

chymatum, 15-20 pm crassum, J— . Hypothecium hyali-

num, 40-50 pm crassum, J+ caeruleum, paraplecten-

chymatum, cellulis 2, 5—4,0 pm diametro. Hymenium

hyalinum, 70-100 pm altum, J+ caeruleum. Para-

physes ramosae et anastomosae, septatae, luminibus

1,5-1 ,7 pm crassis, gelatinosae, gelatina J+ caerulea.

Asci clavati, cum parietibus tenuibus, J— . Ascosporae

hyalinae, simplices, ovales, octonae, 11-17 x 8, 0-9, 5

pm. Pycnidia non visa.

TYPE. — 1916 (Gobaub): Etosha Pan, Halali Hill, on

S faces of dolomite outcrops on a SE slope (-AB),

Brusse 4163, 1984.03.19 (PRE, holo.; BM, COLO, LD,

MB, iso.). Figure 10.

Thallus subfruticose, radiately lobate from a central

holdfast, up to 18 mm across when dry, on dolomite.

Lobes subascending, linear, simple to branched, up to 10

mm long and 0,4-1 ,5 (-2,5) mm wide when dry,

400-600 pm thick when wet. Upper surface charcoal,

matt to granular-isidiate, smooth. Cortex 5-15 pm thick,

small-celled paraplectenchymatous, cells 3,5^4,5 /t,m

diam. (Figure 11). Algal layer 20-50 pm thick; algae

blue-green (Chroococcales or Pleurocapsales). Medulla

FIGURE 11. — Thyrea otaviana Brusse, thin section of cortex in lacto-

phenol cotton-blue. Brusse 4163, holotype. Bar = 100 /jl m.

FIGURE 12. — Thyrea otaviana Brusse, thickish section in lactophenol cotton-blue showing reticulated medulla. Brusse 4163, holotype. Bar

= 100 gm.

Bothalia 17,1 (1987)

39

FIGURE 13. — Thyrea otaviana

Brusse, section of apothecium

in lactophenol cotton-blue. Ar-

row indicates a sectioned isi-

dium. Brusse 4163, holotype.

Bar = 100 /im.

hyaline, 390-560 pm thick; hyphae reticulately arranged

(Figure 12), 1,4-2, 5 gm thick. Lower surface charcoal,

matt, smooth, canaliculate when dry. Apothecia (Figure

13) laminal, sessile, lecanorine, up to 0,5 mm (dry).

Hymenial surface brown, nitid. Thalline exciple 80-150

pm thick on sides. Exciple hyaline, periclinal prosoplec-

tenchyma, 15-20 pm thick, J-. Hypothecium hyaline,

40-50 pm thick, J+ blue, small-celled paraplecten-

chyma, cells 2, 5^,0 pm diam. Hymenium hyaline,

70-100 pm high, J+ blue. Paraphyses branched and

anastomosed, septate, lumens 1,5-1, 7 pm thick, gelati-

nized, gel J+ blue. Asci 8-spored, clavate, walls thin,

J- (Figure 14). Ascospores hyaline, monolocular, oval,

11-17 x 8, 0-9, 5 pm. Pycnidia not seen.

Two new species of this genus have recently been

described from Africa (Henssen 1986; Henssen, Biidel &

Wessels 1985), but these, like the other species of Thy-

rea (Henssen 1980; Poelt 1969; Zahlbruckner 1926),

lack a cortex, the outer surface terminating in relatively

undifferentiated hyphae. Thyrea otaviana displays a

weakly developed cortex with the hyphae becoming

swollen and conglutinate near the surface, so that it is

anticlinally paraplectenchymatous (Figure 11). The new

species is therefore the most highly evolved species of

this genus known to date (at least as far as thalline mor-

phology is concerned). The medullary hyphae are fas-

cicled or drawn together at regular intervals to give a

reticulate pattern (Figure 12) as in T. rotundata Biidel et

al. (Henssen, Biidel & Wessels 1985).

The ascospores of this new species are also larger than

any known lobate (as opposed to peltate) Thyrea, reach-

ing 17 gm long in a small sample of eight ascospores

measured.

The new species is presently known only from the

type locality, Halali Hill, Etosha Pan on dolomite on a

south-east facing slope, in the partial shade of Moringa

ovalifolia trees. Halali hill is composed of dolomite of

the Otavi group of the Damara geological sequence, and

the specific name alludes to this fact. The lichen is pro-

bably more widespread on dolomite in the northern part

of South West Africa/Namibia.

ACKNOWLEDGEMENTS

The author is grateful to game rangers Karen and

Trevor Nott for company during a visit to Etosha Pan and

to game ranger/tracker Otavi Tsumeb for armed protec-

tion against large carnivorous and other dangerous ani-

mals.

FIGURE 14. — Thyrea otaviana Brusse,

ascus and paraphyses. Brusse 4163, ho-

lotype. Bar = 10/j.m.

40

Bothalia 17,1 (1987)

REFERENCES

HENSSEN, A. 1980. Problematik der Gattungsbegrenzung bei den

Lichinaceen. Berichte der Deutschen Botanischen Gesellschaft

92: 483-506.

HENSSEN, A. 1986. Thyrea polyglossa and Thyrea divergens sp.

nov. My cotaxon 25: 493-503.

HENSSEN, A., BUDEL, B. & WESSELS, D. 1985. New or interest-

ing members of the Lichinaceae from southern Africa. 1. Species

from northern and eastern Transvaal. Mycotaxon 22: 169-195.

POELT, J. 1969. Bestimmungsschliissel europaischer Flechten.

Cramer, Lehre.

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

Prantl, Die Natiirlichen Pflanzenfamilien, edn 2, Vol. 8: 61-270.

Engelmann, Leipzig.

F. BRUSSE

POLYGONACEAE

A NEW SPECIES OF OXYGONUM FROM NATAL

Oxygonum robustum Germishuizen, sp. nov., O.

dregeano Meisn. affinis, sed ocrea setis longis rigidis,

habitu robusto decumbenti, foliisque nunquam pinnati-

sectis differt.

TYPE. — Natal, 2732 (Ubombo): east of Vazi Swamp,

Manzengwenya (-BB), Moll 4722 (PRE, holo.; NH,

iso.). Figure 15.

Robust, spreading, decumbent herb much branched

from the base; arising from a long thick vertical woody

rootstock. Old stems reddish brown, sparsely pilose;

young stems glabrous, longitudinally ridged, brown-

punctate. Ocrea membranous, reddish brown, 8-10 mm

long, fringed with rigid bristles 8-10 mm long, pilose,

brown-punctate, adnate to the base of the leaves. Leaves

simple, sessile, alternately arranged, glabrous, brown-

punctate, lamina obovate to elliptic to spathulate, (37-)

45-75 (-94) x (8-) 10-21 (-23) mm, acuminate, mar-

natal

NATAL!

-27 32 BA Ubombo

E.J. Moll

NATAISE HERBARIUM DURBAN

aftsl.; HtglO KfitJll

Oxygoniu

Edge of email vlei just e*8t of Vazi

Swamp, Manzengwenya. Graael nd.

White- flowered herb, fairly common.

FIGURE 15. — Holotype of Oxygo-

num robustum Germishuizen.

Bothalia 17,1 (1987)

41

gins entire or occasionally sharply toothed. Inflorescence

a long lax thyrse with fascicles of up to 4 flowers in the

axils of the bracts. Flowers bisexual. Perianth 5-lobed,

up to 7 mm long, glabrous, white. Stamens 8, inserted

on the inner perianth; anthers reddish brown, 1 mm long;

filaments 4 mm long. Styles 3, connate at the base; stig-

mas capitate. Fruit a 3-angled nut.

NATAL. — 2632 (Bella Vista): Maputa to Kosi Nature Reserve

(-DD), Edwards 2586. 2732 (Ubombo): 16 km north of Lake Sibayi

(—BA), Vahrmeyer & Toelken 259', Muzi Swamp ± 2 km from Phelen-

daba turnoff on road to Mbazwana (-BA), Germishuizen 3546' Vazi

Swamp (-BB), Stephen, Van Graan & Schwabe 1177', east of Vazi

Swamp, Manzengwenya (-BB), Moll 4722; Mbazwana (-BC),

Gerstner 704; Manzengwenya Forest Station (-BD), Vorster 2586;

Bangazi Lake (-DA), Strey 4991. 2832 (Mtubatuba): 25 km north of

Mtubatuba (-AB), Codd 2022; near Fames Island Camp (-AB), Ward

1493; Dukuduku U176 (-AC), Strey 5522; St Lucia, Eastern Shore

State Forest (-AB), Nicholas 1624; Nseleni Nature Reserve (-CA),

Bourquin & Ndlovu 17; 80 km from Arboretum turnoff, on old main

road to Empangeni (-CC) Ngwenya 240; Richards Bay (-CC), Lawn

1725 & 1726; Rump s.n. (NH 20120).

0. robustum is endemic to Natal, occurring in a

restricted area (Figure 16) of coastal grassland and

coastal dune forest. The first record of the species in the

Natal Herbarium was collected by W. G. Rump s.n. (NH

20120) during July 1929 at Richards Bay. All the mate-

rial cited above was incorrectly identified as O. dregea-

num. Although these two taxa are sympatric, 0. robus-

tum can be distinguished from 0. dregeanum by its

robust and decumbent habit, by the ocrea fringe having

FIGURE 16. — Distribution of Oxygonum robustum Germishuizen.

long rigid bristles and by the leaves never being pinnati-

sect.

The specific epithet robustum refers to the robust habit

of this species.

G. GERMISHUIZEN

TELOSCHIST ACEAE

A NEW SPECIES OF CALOPLACA FROM SOUTHERN AFRICA

INTRODUCTION

The genus Caloplaca Th. Fr. is one of the largest

lichen genera known. The total number of species has

been estimated at almost 500 (Santesson 1970). Al-

though many of the published species names will be

reduced to synonyms in future revisions, there are still

unknown species to be discovered in this fascinating

genus. The genus is also extremely variable and incorpo-

rates many different structural forms, which, if measured

by today’s taxonomic concepts in, for example, the Leci-

deaceae or Parmeliaceae, would have been recognized at

generic level. However, too many intermediate forms

are present to justify a more diversified generic taxo-

nomy in the Teloschistaceae (see Hafellner & Poelt

1979; Poelt & Hafellner 1980; Poelt & Pelleter 1984;

Wunder 1974).

In addition many species show extremely broad mor-

phological variations, which normally also correlate with

rather extensive geographical ranges. In taxonomic

terms this variation is usually extremely hard to define.

Yet, many characteristic species with geographically

limited distributions are also present in the genus. The

new species described here belongs to this latter cate-

gory. I recognized the unique characteristics of the

species for the first time in undetermined material from

Lesotho. On a field trip in southern Africa with Mr Frank

Brusse at the beginning of 1986, we collected the new

species in several different localities along the Drakens-

berg escarpment. The species is named in honour of my

older colleague Dr Ove Almbom, who collected it as far

back as 1953.

Caloplaca almbornii Karnefelt, sp. nov.

Thallus areolis dispersis sanguineis 0,2-3 ,0 mm

diametro compositus. Apothecia immersa, postea parum

prominentia, lecideina; discus aurantiacus. Ascosporae

polaribiloculares, 9,5-13,5 x 3, 8-4, 7 /xm, septum ± 2

/z m crassum. Thallus K+ purpureus, parietinum adest.

TYPE. — Lesotho, 2929 (Underberg): Mokhotlong,

mountain behind Pack Horse Inn (-AC), 1963, Kofler

(LD, holo.).

Thallus composed of scattered, more or less circular,

usually convex areoles, 0, 2-1,0 mm across, or when

flatter up to 3,0 mm across, then secondarily cracked

into several small areoles; usually bright reddish or occa-

sionally more orange pigmented. Cortical layer ± 40-70

/xm thick, covered by a ± 10 /xm thick pigmented layer,

paraplectenchymatous, composed of anticlinal cells,

occasionally penetrating the clustered algae in up to 150

/xm deep strands. Photobiont clustered, occasionally

nearing the surface, single cells green and spherical, ±

10 /xm in diameter. Medullary layer rather dense,

130-350 /xm deep (Figure 17F). Apothecia often pre-

sent, immersed, becoming slightly raised with maturity,

42

Bothalia 17,1 (1987)

FIGURE 17. — Caloplaca almbornii. A, specimen with very scattered areoles, Kdrnefelt 8626-7, bar = 10 mm; B & C, close up of thallus,

Kofler s.n., bar = 1 mm; D, cross section of areole showing immersed apothecium, Kdrnefelt 8625-20, bar = 100 /xm.; E, cross section

of areole with conidiomata, Kdrnefelt 8626-8, bar = 100 /xm.; F, cross section of areole, Kdrnefelt 8625-20, bar = 100 /xm.

often one per areole (Figure 17C & D), to several

together on the broader cracked areoles; disc very small,

0, 1-0,7 mm across, covered by an orange pigment.

Excipulum occasionally seen as a thin pale orange ring

around the disc, 10-20 /xm thick. Hymenium ± 65-95

/xm thick, covered by a pigmented, granular, epihyme-

nial layer up to 35 /xm thick. Asci 45-55 x 8-10 /xm.

Ascospores ellipsoidal, polaribilocular, 9,5-13,5 x

3, 8-5, 7 /xm; septum ± 2 /xm thick. Hypothecium form-

ing a relatively flat dome below the excipulum, 35-55

(-150) /xm deep, hyaline. Conidiomata scattered,

1 10-170 /xm large, pyriform (Figure 17E); conidia very

small, slightly bifusiform, ± 3,5 x 0,5 /xm. Reactions

K+ blood red, C— , KC— and P— . Caloplaca almbornii

contains parietin only (TLC).

Caloplaca almbornii is characterized particularly by

the remarkable structure of the thallus composed of scat-

tered convex areoles, forming small red-spotted patches

(Figure 17A, B & C). Apart from thallus colour and

structure of the ascomata, C. almbornii is reminiscent of

Aspicilia contorta (Hoffm.) Krempelh. which occurs on

various calcareous rocks in Europe. The areoles are often

embedded in other blackish, crustose, sterile lichens. It

is therefore not possible to demonstrate whether the

areoles are connected by anastomosing hyphae, or by a

thin hypothallus. Flatter and up to 3 mm broad areoles

can occasionally be found. The colour of the areoles is

sometimes more orange than bright reddish.

Caloplaca almbornii appears to be isolated in the

genus and is not clearly related to any other species. It

Bothalia 17,1 (1987)

43

shows some affinities with the C. cinnabarina group,

which is a very variable species complex occurring

mainly at lower altitudes in southern Africa. C. almbor-

nii and the C. cinnabarina group have similar anatomical

characters but differ considerably in the structure of the

thallus. The bright reddish colour is most reminiscent of

the colour of C. haematodes (Massal.) Zahlbr. C. alm-

bomii also differs chemically by containing only parie-

tin, whereas most other species examined contain more

than one anthraquinone.

Caloplaca almbornii is endemic to southern Africa

and is known only from mountainous regions, mainly at

rather high altitudes. It has been found locally abundant

at 2 000-3 000 m in the Drakensberg in Lesotho, the

Transkei and Natal. Furthermore, it has been found in

the Orange Free State close to the Lesotho border, in the

easternmost part of the Cape Province and in the Matopo

Hills in SW Zimbabwe (Figure 18). The species could

probably occur in the eastern mountainous region of the

Transvaal, located between the northernmost outpost in

Zimbabwe and the Drakensberg centre in the south. I

have, however, not been able to find it during several

excursions, specially undertaken to this otherwise liche-

nologically very rich eastern extension of the Drakens-

berg. The eastern Transvaal mountains are presumably

too low in altitude.

FIGURE 18. — Distribution of Caloplaca almbornii.

In the Mont-aux-Sources region in the northern part of

the Drakensberg, I found C. almbornii locally abundant

on sloping dark basaltic rocks with dripping water. In the

southern part of the Drakensberg the species was also

found locally in the cave sandstone belt, growing on flat

surfaces with stagnant water, or on sloping surfaces with

dripping water. Most of these habitats are easily recog-

nized by the blackish colour caused by a variety of crus-

tose lichens and possibly also by Cyanobacteria. C. alm-

bornii is easily discovered here, where it contrasts as

patches of bright red spots against the dark rock surface.

ZIMBABWE. — 2028 (Bulawayo): Matopo Hills, Pomowe (-AC),

1963, Kofler (LD).

O.F.S.— 2828 (Bethlehem): Clarens (-CB), 1943, Plank 1880

(PRE).

NATAL. — 2828 (Bethlehem): Royal Natal National Park, mountain

path to Mont-aux-Sources from entrance gate at the end of mountain

road from Phuthaditjhaba (-DB), 1986, Kdrnefelt 8626-8, 8626-10,

8626-20, 8626-22, 2826-46 (LD); Little Tugela area (-DB), 1943,

Esterhuysen 8875 (BOL). 2829 (Harrismith): Cathedral Peak area,

Organ Pipes (-CC), 1953, Almborn 9037, 9058 (LD).

QWA-QWA. — 2828 (Bethlehem): Mont-aux-Sources area, moun-

tain path from entrance gate to Royal Natal National Park at the end of

mountain road from Phuthaditjhaba (-DB), 1986, Kdrnefelt 8625-8,

8625-12, 8625-14, 8625-15, 8625-20-8625-23 (LD).

LESOTHO. — 2927 (Maseru): Machache Mtn (-AD), 1929-1930,

Hewitt (TRH, 4 collections); Popanyane (-BB), 1963, Kofler 3523

(LD). 2828 (Bethlehem): Leribe, Oxbow (-CC), 1953, Kofler (LD, 2

collections); Mapoteng, 30 miles E of Leribe (-CC), 1963, Kofler

310195 (LD, 2 collections). 2929 (Underberg): Mokhotlong, behind

Pack Horse Inn (-AC), Kofler (LD, 2 collections).

TRANSKEI. — 3029 (Matatiele): Qachas Nek, close to the Lesotho

border, cave sandstone formations (-BB), 1986, Kdrnefelt 8630-

44-8630-51 (LD).

CAPE. — 3027 (Lady Grey): Naude’s Nek, on R396 to Rhodes from

Elands Height (-DB), 1986, Kdrnefelt 8632-3-8632-6 (LD); mountain

6 miles from Naude’s Nek, on the way to Ben McDhuy (-DB), 1963,

Kofler 311166. 3127 (Lady Frere): Barkly Pass on R58 between Barkly

Pass and Elliot (-BB), 1986, Kdrnefelt 8633-1-8633-3 (LD).

ACKNOWLEDGEMENTS

I would like to thank Dr B. de Winter, Director of the

Botanical Research Institute, for his support in many

ways and Mr Frank Brusse, who accompanied me during

the field work along the Drakensberg escarpment.

REFERENCES

HAFELLNER, J. & POELT, J. 1979. Die Arten der Gattung Calo-

placa mit plurilocularen Sporen (Meroplacis, Triophthalmidium,

Xanthocarpia) . Journal of the Hattori Botanical Laboratory 46:

1—4 1 .

POELT, J. & HAFELLNER, J. 1980. Apatoplaca — genus novum

Teloschistacearum (Lichenes). Mitteilungen der Botanischen

Staatssammlung Munchen 16: 503-528.

POELT, J. & PELLETER, U. 1984. Zwergstrauchige Arten der Flech-

tengattung Caloplaca. Plant Systematics and Evolution. 148:

51-88.

SANTESSON, J. 1970. Antraquinones in Caloplaca. Phytochemistry

9: 2140-2166.

WUNDER, H. 1974. Schwarzfriichtige, saxicole Sippen der Gattung

Caloplaca (Lichenes, Teloschistaceae) in Mitteleuropa, mit dem

Mittelmeergebiet und Vorderasien. Bibliotheca Lichenologica 3:

1-186. Lehre.

I. KARNEFELT*

* Department of Systematic Botany, University of Lund, Ostra

Vallgatan 18, 223 61 Lund, Sweden.

Bothalia 17,1:45-49(1987)

Cetraria (Parmeliaceae) and some related genera on the African

continent

i. kArnefelt*

Keywords: Bryoria, Cetraria, new records, Parmeliaceae, Platismatia, southern Africa, taxonomy

ABSTRACT

Four species are reported new for the lichen flora of southern Africa: Bryoria fuscescens (Gyeln.) Brodo &

D. Hawksw., B. motykae (D. Hawksw.) Brodo & D. Hawksw., Cetraria chlorophylla (Humb.) Vain, and Platismatia

glauca (L.) Culb. & Culb. (Parmeliaceae), found at high elevations in Natal and south-west Cape. B. fuscescens, B.

motykae and P . glauca were previously known from a few localities, also at high elevations, in east Africa. The genus

Cetraria s. str. was previously unknown on the African continent.

UITTREKSEL

Vier spesies word nuut aangeteken vir die ligeen-flora van suidelike Afrika: Bryoria fuscescens (Gyeln.) Brodo & D.

Hawksw., B. motykae (D. Hawksw.) Brodo & D. Hawksw., Cetraria chlorophylla (Humb.) Vain, en Platismatia glauca

(L.) Culb. & Culb. (Parmeliaceae), wat hoog bo seespieel in Natal en suidwes-Kaap aangetref word. B . fuscescens, B.

motykae en P . glauca was voorheen bekend vanaf 'n paar plekke, ook hoog bo seespieel, in oos- Afrika. Die genus Cetraria

s. str. was voorheen onbekend op die vasteland van Afrika.

INTRODUCTION

During my work on the genera Coelocaulon and

Cornicularia (Kamefelt 1986), I came across a few note-

worthy specimens in the Bolus Herbarium, University of

Cape Town, collected by Ms Elsie Esterhuysen. Apart

from the material of Coelocaulon epiphorellum, which

was then new to Africa, Ms Esterhuysen’ s collections

also included four other species new to southern Africa:

Bryoria fuscescens (Gyeln.) Brodo & D. Hawksw., B.

motykae (D. Hawksw.) Brodo & D. Hawksw., Cetraria

chlorophylla (Humb.) Vain, and Platismatia glauca (L.)

Culb. & Culb. (Parmeliaceae). With the discovery of

Cetraria chlorophylla in southern Africa the genus

Cetraria s. str. is also reported as being new to the Afri-

can continent.

MATERIAL AND METHODS

The material from southern Africa was investigated

chemically for secondary compounds and was compared

anatomically with material from the northern hemisphere

in LD, and with material from the east African moun-

tains in UPS.

Cetraria chlorophylla (Humb.) Vain, in Acta So-

cietatis pro Fauna et Flora Fennica 13: 7 (1896).

Lichen chlorophyllus Humb.: 20 (1793). Type: In cortice Pyni syl-

vestris beym Vorwerk Hals copiose (B, Willdenow).

Thallus subfruticose or foliose, ± 10-20 mm across;

lobes irregularly branched, ±0,5 mm broad at tips, ±

2-3 mm at base, weakly channelled to flat, with scat-

tered marginal soralia especially towards the lobe tips,

and scattered marginal cilia; upper surface brown to pale

brown, glossy, rather smooth; lower surface paler brown

to whitish, wrinkled or sulcate, with rather frequent pale

* Department of Systematic Botany, University of Lund, Ostra

Vallgatan 18, 223 61 Lund, Sweden.

rhizinae; ascomata and conidiomata not observed. Cor-

tex paraplectenchymatous, ± 25-30 p m thick, covered

by a thin ± 5 pm thick pigmented epicortical layer,

composed of 1-2 layers of densely packed isodiametric

cells with rather distinctive lumina; photobiont green and

spherical forming a more or less continuous layer;

medulla composed of loose hyphae; lower cortical layer

similar to the upper layer (Figure 1C). Reactions K— ,

C-, KC- and P— . Cetraria chlorophylla contains pro-

tolichesterinic acid.

Cetraria chlorophylla is characterized by the rather

small, subfruticose to foliose thallus, brown to usually

more greenish brown in the northern hemisphere, and

glossy lobes with scattered marginal soralia and cilia

(Figure 1A). It is a locally common species both on

conifers and on deciduous trees in north-western and

central Europe (see Wirth 1980). In western North

America it is also one of the most common foliose spec-

ies (Hale 1979; Thomson 1984). In continental parts of

Eurasia and North America, however, C. chlorophylla is

very rare or lacking, and relatively mild and oceanic

climatic regions provide conditions favourable for the

growth of this species. C. chlorophylla has also been

reported from Japan (Sato 1965) and from Tierra del

Fuego (Rasanen 1932) and Australia (Filson 1983) in the

southern hemisphere. The species was, however, not

treated by Galloway (1985) in the new lichen flora of

New Zealand.

The material from southern Africa was collected at

750-900 m alt. on shelves on the NE side of Table

Mountain in the SW Cape (Figures 2 & 3). This is not

surprising, as Table Mountain, with its mild and humid

winters and relatively high rainfall, offers favourable

conditions for a diversified lichen vegetation. It is one of

the richest lichen localities in southern Africa and is the

type locality of numerous lichen species.

It is premature to speculate on the origin and dispersal

of C. chlorophylla in the southern hemisphere and on the

46

Bothalia 17,1 (1987)

FIGURE 1. — A, Cetraria chlorophylla, Esterhuysen 22911 , bar = 10 mm; B, Platismatia glauca, Esterhuysen 10261, bar = 10 mm; C,

C. chlorophylla, cross section of thallus, interference contrast micrograph, Esterhuysen 10261, bar 10 /xm; D, P. glauca, cross section of

thallus, light micrograph, Esterhuysen 10261 , bar = 10 /xm; E, Bryoria fuscescens, Esterhuysen 10261 , bar = 10 mm; F, B. motykae,

Esterhuysen 21356, bar = 10 mm.

only record yet known from southern Africa. I am not

inclined to believe, however, that the species has been

introduced in modem times, as it occurs at relatively

high elevations in a natural habitat (see Kamefelt 1986).

Presumably the species could occur in similar regions in

southern Africa with favourable climatic conditions,

such as the Drakensberg escarpment as well as at high

altitudes in the east African mountains.

CAPE. — 3318 (Cape Town): Table Mountain, cliffs, NE side, on

shelves, 2500-3000 ff (750-900 m] (-CD), 1954, Esterhuysen 22911

(BOL).

Platismatia glauca (L.) Culb. & Culb. in Contri-

butions from the United States National Herbarium 34:

530(1968).

Lichen glaucus L.: 1 148 (1753). Type: Europe, Linnean Herbarium

Sheet 1273-139 (LINN, lecto.).

Thallus foliose, loosely attached, 20-50 mm across;

lobes 5-10 mm broad; upper surface yellowish, with

dark brownish areas, reticulately wrinkled, with scat-

tered marginal soralia; lower surface dark brown, reticu-

lately wrinkled to pitted (Figure IB); ascomata and coni-

Bothalia 17,1 (1987)

47

diomata not observed. Upper cortex prosoplectenchyma-

tous, 20-30 pm thick, composed of strongly pachyder-

matous cells with indistinct cell lumina; photobiont

green, spherical, concentrated near the upper cortex; me-

dulla loose, 50-100 (Lim thick; lower cortex similar to the

upper cortex, heavily pigmented (Figure ID). Reactions

K-, C— , KC- and P-. Platismatia glauca contains

caperatic acid and atranorin.

This well known species from the northern hemis-

phere has been described in detail by Culberson & Cul-

berson (1968). Platismatia glauca is locally common on

conifers and deciduous trees and occasionally also on

rocks in western Europe and North America. It is ex-

tremely common especially in the western American

states, but is found more rarely in the eastern provinces

(Hale 1979; Thomson 1984). Like C. chlorophylla, P.

glauca is apparently also favoured by a rather mild

oceanic climate.

In the southern hemisphere P. glauca is also known

from the south-westernmost part of South America (Cul-

berson & Culberson 1968) and from Australia (Filson

1983), but it is so far unknown from New Zealand (Gal-

loway 1985).

On the African continent P. glauca was previously

known from Kenya, Tanzania and Uganda (Figure 3),

where it occurs at rather high altitudes from 3 200-3 500

m (Krog & Swinscow 1975). The two new records from

southern Africa were found at ± 2 400 m in the Cathe-

dral Peak area in the Natal Drakensberg and at ± 1 800

m in the Sneeuberg in the Sederberg Mountains in the

western Cape (Figure 2). The scattered montane African

records may indicate that the localities represent the rem-

nants of a much wider pre-Pleistocene distribution.

FIGURE 2.— Distribution in southern Africa of: Cetraria chloro-

phylla, ■ ; Platismatia glauca, •; Bryoria fuscescens, O; B.

motykae, A .

NATAL. — 2829 (Harrismith): Cathedral Peak area, tree trunk, 8000

ft [2 500 m], (-CC), 1944, Esterhuysen 10261 (BOL).

CAPE.— 3218 (Clanwilliam): Sederberg, Sneeuberg, shaded ledge

on cliffs, south side, 6000 ft (1 800 m] (-DA), 1952, Esterhuysen

20044 ( BOL).

Bryoria fuscescens (Gyeln.) Brodo & D. Hawksw.

in Opera Botanica 42: 83 (1977).

FIGURE 3. — Distribution in Africa of: Cetraria chlorophylla, ■; Pla-

tismatia glauca, *.

Alectoria fuscescens Gyeln.: 55 (1932). Type: Finland, Tavastia

austr. , Hollola, ad truncos Pini locis apricioribus in silva, J. P. Norrlin,

1882, Nyl. & Norrl., Lich. Fenn. Exs. no. 466 (BP, 33.947, lecto.).

Thallus subpendent, 30-60 mm across; main lobes

with rather abundant short lateral branches, up to 0,3

mm wide, pale brown to blackish; with abundant fissural

soralia, from rather small up to 0,5 mm wide; scattered

small fissural pseudocyphellae present; ascomata and

conidiomata not observed (Figure IE). Anatomical char-

acters similar to most other species of Bryoria, the corti-

cal layer composed of periclinal hyphae. Reactions of

medulla and particularly the soralia, K— , C— , KC— and

P+. B. fuscescens contains fumarprotocetraric acid,

clearly indicated by TLC of the examined material.

Within the genus Bryoria, B. fuscescens is one of the

most difficult species to circumscribe. It is widely dis-

tributed within the boreal zone in the northern hemis-

phere, where it also occurs in many different habitats,

frequently on trees, but also on the ground and on rocks.

The known morphological and chemical variation is

fairly wide and may be partly the result of different res-

ponses to the local diversified habitat selection pressures

(see Brodo & Hawksworth 1977; Bystrek 1963; Hawks-

worth 1972; Hawksworth 1973).

The material from southern Africa was collected at ±

1 680 m altitude on twigs at Horseshoe Peak, in the Hex

River Mountains in the western Cape (Figure 2). This is

the first known record of a member of the genus Bryoria

in southern Africa. The earlier records published by

Doidge (1950) as Alectoria chalybeiformis f. terrestris

and A. jubata have been re-examined and are apparently

filamentous Cyanobacteria.

Apart from southern Africa, B. fuscescens is also

known from several localities in the ericaceous zone and

48

Bothalia 17,1 (1987)

in the alpine zone from 3 200 to 4 900 m altitude, in

Ethiopia, Kenya, Tanzania and Uganda (Krog &

Swinscow 1975). Figure 4.

CAPE. — (3319 Worcester): Hex River Mountains, Horseshoe Peak,

on ledges on south side, on twigs, 5500 ft [1 680 m] (-DD), 1953,

Esterhuysen 22189 (BOL).

FIGURE 4. — Distribution in Africa of Bryoria fuscescens.

Bryoria motykae (D. Hawksw.) Brodo & D.

Hawksw. in Opera Botanica42: 155 (1977).

Alectoria motykae D. Hawksw. 124: 124 (1971). Type: Kenya, Mt

Kenya, Teleki Valley, alpine regions, on vertical surfaces of boulders

in the upper parts of the valley, alt. 4 200 m, 1948, Hedberg 1720b

(UPS, holo.).

Thallus prostrate, 10-30 mm long; main lobes with

arcuately curved lateral branches, dark brown or partly

blackish, up to 0,2 mm wide, with scattered fissural

pseudocyphellae, up to 0,5 mm broad; ascomata and

conidiomata not observed. Anatomical characters simi-

lar to most other species of Bryoria, the cortical layer

composed of periclinal hyphae. Reactions of medulla

and pseudocyphellae, K-, C-, KC- and P+. Bryoria

motykae contains fumarprotocetraric acid according to

Hawksworth (1971). However, TLC gave only weak

spots in the examined material, which included the type

specimen kept at UPS, and the P reaction was feint as

well.

Bryoria motykae is characterized by the dark brown or

partly blackish main lobes with arcuately curved lateral

branches, broad fissural soralia and scattered fissural

pseudocyphellae. The material from southern Africa,

however, differs slightly from the type material from

Mount Kenya ( Hedberg 1720b, UPS) in the absence of

soralia, but corresponds in the remaining characters with

the type (Figure IF). According to Hawksworth (1971)

pseudocyphellae are lacking, contrary to my observa-

tions on re-examination of the type material. Krog &

Swinscow (1975) had apparently also observed pseudo-

cyphellae, because they mention that the fissural soralia

developed from minute pseudocyphellae.

Among the possibly related species, Hawksworth

(1971) pointed out the resemblance to B.furcellata (Fr.)

Brodo & D. Hawksw. (as Alectoria nidulifera Norrl. in

Nyl.) in some characters. B.furcellata is, however, dis-

tinguished by the more spinulate lobes and the rather

abundant isidiate soredia. B. motykae is also reminiscent

of B. simplicior (Vain.) Brodo & D. Hawksw., in the

structure of the soredia (Hawksworth 1971). B. simpli-

cior is, however, a normally corticolous species, charac-

terized by a typical caespitose habit and very distinctive

soredia. In addition B. simplicior also lacks secondary

compounds.

B. bicolor (Ehrh.) Brodo & D. Hawksw. is somewhat

reminiscent of B. motykae in the similar fissural pseudo-

cyphellae, but this species is distinguished by the rather

short lateral spinules and differently coloured main

lobes. Furthermore, Krog & Swinscow (1975) included

B. ruwenzoriensis (D. Hawksw.) Brodo & D. Hawksw.,

recorded from east Africa, in B. bicolor.

In my opinion B. motykae is presumably associated

with the B. fuscescens aggregate, which also occurs in

the east African mountains. We have to await further

studies on this difficult species complex before the status

of B. motykae can be determined with certainty.

The material from southern Africa was collected at ±

1 300 m on sandy hollowed rock surfaces on the Groot

Drakenstein Mountain near Paarl in the western Cape

(Figure 2).

Apart from southern Africa, B. motykae is also known

from Mt Kenya, where it occurs at ± 3 500 m (Figure

5). The record published by Krog & Swinscow (1975)

FIGURE 5. — Distribution in Africa of Bryoria motykae.

Bothalia 17,1 (1987)

49

from Mt Kilimanjaro in Tanzania, proved after re-exami-

nation to belong to the B. fuscescens group.

CAPE. — 3319 (Worcester): Groot Drakenstein Peak, amongst other

lichens on sandy hollowed rock surfaces, 4500 ft [1 370m], (-CA),

1953, Esterhuysen 21356 (BOL).

REFERENCES

BRODO, I. M. & HAWKSWORTH, D. L. 1977. Alectoria and allied

genera in North America. Opera Botanica No. 42.

BYSTREK, J. 1963. De variabilitate Alectoriae fuscescentis Gyeln.

Annales Universitatis Mariae Curie-Sklodowska C 18: 411-418.

CULBERSON, W. L. & CULBERSON, C. F. 1968. The lichen

genera Cetrelia and Platismatia (Parmeliaceae). Contributions

from the United States National Herbarium 34: 449-558.

FTLSON, R. B. 1983. Checklist of Australian lichens. National

Herbarium of Victoria, Department of Conservation, Forests &

Lands.

GALLOWAY, D. J. 1985. Flora of New Zealand. Wellington.

GYELNIK, V. 1932. A/ec/on'a-Studien. Nyt Magazin for Natur-

videnskaberne 70: 35-62.

HALE, M. E. 1979. How to know the lichens. Dubuque, Iowa.

HAWKSWORTH, D. L. 1971. Regional studies in Alectoria (Lich-

enes). I. The central and southern African species. Botaniska

Notiser 124: 122-128.

HAWKSWORTH, D. L. 1972. Regional studies in Alectoria (Lich-

enes). II. The British species. The Lichenologist 5: 181-261 .

HAWKSWORTH, D. L. 1973. Ecological factors and species delimi-

tation in the lichens. In V. H. Hey wood. Taxonomy and ecology.

31-69. London & New York.

HUMBOLDT, F. W. H. A. 1793. Florae fribergensis specimen : 20.

Berlin.

KARNEFELT, I. 1986. The genera Bryocaulon. Coelocaulon and

Cornicularia and formerly associated taxa. Opera Botanica No.

86.

KROG, H. & SWINSCOW, T. D. V. 1975. Parmeliaceae, with the

exclusion of Parmelia and Usnea, in east Africa. Norwegian

Journal of Botany 22: 115-123.

LINNAEUS, C. 1753 . Species plantarum: 1 148. Stockholm.

RASANEN, V. 1932. Zur Kenntnis der Flechtenflora Feurland’s sowie

der Prov. de Magallanes, Prov. de Chiloee und Prov. de Nuble in

Chile. Annales botanici Societatis zoologicae-botanicae Fennicae

Vanamo 2.

SATO, M. 1965. Catalogus Lichenum Japonicorum, edn. 2. Miscella-

nea Bryologica et Lichenologica .

THOMSON, J. W. 1984. American arctic lichens. I. The macroli-

chens. New York.

VAINIO, E. 1896. Lichenes in Sibiria Meridionali collecti. Acta So-

cietas Pro Fauna et Flora Fennica 13: 1-20.

WIRTH, V. 1980. Flechtenflora. Stuttgart.

Bothalia 17,1:51-65 (1987)

Species groups in the genus Ehrharta (Poaceae) in southern Africa

G. E. GIBBS RUSSELL* AND R. P. ELLIS*

Keywords: anatomy, Ehrharta, morphology, Poaceae, southern Africa, taxonomy

ABSTRACT

Ehrharta Thunb. is a genus of Gondwanaland distribution with its centre of diversity in the winter rainfall Fynbos

Biome of southern Africa. In recent subfamily treatments Ehrharta has proved difficult to place satisfactorily, and during

the past five years it has been moved between Bambusoideae and Arundinoideae. However, most previous systematic

studies using cryptic characters have covered only four taxa out of about 35. The present study includes all African taxa, and

demarcates seven species groups on the basis of both spike let morphology and leaf blade anatomy. Parallelism and/or

convergence in vegetative macromorphology within and between the species groups is widespread, and is similar, in some

cases, to adaptations found in other plant families in the Fynbos Biome. However, these macromorphological trends are not

reflected in the leaf anatomy. Leaf anatomy is generally consistent with the spike let morphology. Some anatomical

differences between the species groups in Ehrharta appear to be as great as differences between taxa of much higher ranks

elsewhere in the Poaceae. This wide range of variability may be related to an early divergence of Ehrharteae from other

grasses, as suggested by the Gondwanaland distribution, and may explain the difficulty of placing this fascinating yet

baffling genus in a subfamily.

UITTREKSEL

Ehrharta is ’n genus van Gondwanaland-verspreiding met sy middelpunt van verskeidenheid in die winterreenval-Fyn-

bosbioom van suidelike Afrika. In onlangse subfamilie-behandelings is dit moeilik gevind om Ehrharta bevredigend te

plaas, en gedurende die afgelope vyf jaar is dit oor en weer in Bambusoideae en Arundinoideae geplaas. Die meeste vorige

sistematiese ondersoeke waarin verskuilde kenmerke gebruik is, het slegs vier taksons uit ongeveer 35 gedek. Die onderha-

wige ondersoek sluit al die Afrika-taksons in, en baken sewe spesiegroepe af op grond van die morfologie van die

blompakkie en die anatomie van die blaarskyf. Parallelisme en/of konvergensie in vegetatiewe makromorfologie binne en

tussen spesiegroepe is wydverspreid en is in sommige gevalle soortgelyk aan aanpassings wat by ander plantfamilies in die

Fynbosbioom aangetref word. Hierdie makromorfologie se neigings word nie in die blaaranatomie weerspieel nie. Blaarana-

tomie is oor die algemeen in ooreenstemming met die morfologie van die blompakkie. Sommige anatomiese verskille tussen

die spesiegroepe in Ehrharta blyk net so groot te wees soos verskille tussen taksons van 'n veel hoer rang elders in die

Poaceae. Hierdie wye verskeidenheid mag verband hou met ’n vroee divergensie van Ehrharteae vanaf ander grasse, soos

aangedui deur die Gondwanaland-verspreiding, en mag ’n verklaring bied waarom hierdie bekorende maar raaiselagtige

genus moeilik in 'n subfamilie geplaas kan word.

CONTENTS

Introduction 51

Materials and methods 52

Historical background 53

The classification of Ehrharta within the

Poaceae 53

The southern African species of Ehrharta 53

Taxonomic characters 54

Morphological characters 54

Anatomical characters 58

Outline of the species groups in Ehrharta 59

Setacea group 59

Capensis group 59

Longifolia subgroup 60

Capensis subgroup 61

Erecta group 61

Calycina group 62

Villosa group 62

Ramosa group 63

Dura group 63

* Botanical Research Institute, Department of Agriculture and Water

Supply, Private Bag X 101 , Pretoria 0001 .

Conclusions 64

Acknowledgements 64

References 64

INTRODUCTION

In the most recent treatment (Willemse 1982), the

tribe Ehrharteae Nevski (1937) consists of only a single

genus, Ehrharta Thunb. (1779) which includes the gen-

era Microlaena R. Br., Petriella Zotov and Tetrarhena

R. Br. Its distribution area in southern Africa, south-

western, southern and eastern Australia, Tasmania, New

Zealand and Malesia falls within the cool-temperate

Gondwanaland region (Figure 1). The centre of diversity

for the genus is the winter rainfall Cape Floristic Region

(Goldblatt 1978) or Fynbos Biome (Rutherford & West-

fall 1986) of southern Africa, with about 35 taxa, the

other 10 taxa being distributed unevenly elsewhere in the

range of the genus. In southern Africa, Ehrharta can be

divided into seven species groups demarcated on the

basis of both spikelet morphology and leaf blade anat-

omy.

An understanding of the variation within Ehrharta in

southern Africa is necessary, not only to improve the

classification of the local taxa, but also to show the

relationships between the bulk of the genus in southern

Africa and its representatives in other Gondwanaland

52

Bothalia 17,1 (1987)

FIGURE 1. — Worldwide distribution of Ehrharta, showing the number of species and infraspecific taxa reported for various areas. The

number of southern African species naturalized in India and Australia are indicated with an asterisk.

regions. Ehrharta may therefore serve as an additional

plant group to indicate the floristic history of the Gond-

wanaland continents, and is the first representative of the

Poaceae with this potential to be studied with this aim.

Additional information on Ehrharta is required

because its position within the Poaceae is not yet settled.

Since the time of Bentham & Hooker (1883), the group

has been shuffled between the Phalarideae and the Ory-

zeae of the Pooideae (Festucoideae), the Ehrharteae and

Arundineae of the Arundinoideae, the Ehrharteae of the

Oryzoideae, and the Ehrharteae and the Oryzanae of the

Bambusoideae. Modem studies based on cryptic charac-

ters such as leaf anatomy, chromosomes and embryo

types have enabled most grass tribes to be clearly placed

in one of about five subfamilies, but there is still no

general agreement regarding the affinities of the Ehrhar-

teae. Prat (1960) and Clifford & Watson (1977) place the

tribe in an unclassified residue that does not fit neatly

into any of the modem subfamilial concepts of the Poa-

ceae. Most recently, Ehrharteae has been removed from

the Arundinoideae (Renvoize 1981) to the Bambusoi-

deae (Renvoize 1985; Watson etal. 1985).

The instability of the higher classification of the

Ehrharteae may be partly a result of the high degree of

variation in its anatomical characters. Differences

between species and species groups in the southern Afri-

can species appear to be as great as differences between

taxa of much higher rank elsewhere in the Poaceae.

Overseas workers who have examined only a few taxa

may have erroneously concluded that they are represen-

tative of the entire genus or tribe. This wide range of

variability may be a result of the early divergence of the

Ehrharteae from other grasses, as suggested by the

ancient Gondwanaland distribution pattern.

The representatives of Ehrharta in southern Africa

may help to provide a better understanding both of the

history of the flowering plants and of the early evolution

of the Poaceae. They will therefore be treated in some

detail. We propose to set out the species groups infor-

mally in this introductory paper, and thereafter to publish

a parallel series of papers covering the taxonomy and

anatomy of each species group. A concluding paper will

present a formal generic and infrageneric classification

and will develop hypotheses about the phylogeny of the

species groups, and their phytogeographical signifi-

cance.

MATERIALS AND METHODS

For the morphological investigations, herbarium

specimens were examined from BOL, JF, K, NBG,

PRE, SAM and STE. The PRE material includes the

anatomical voucher specimens of Ellis, described below.

Field observations were made for about 30 of the taxa.

Spikelets were dissected without special preparation, and

were observed with a dissecting microscope.

For the anatomical investigations, specimens of

Ehrharta plants were collected in the field throughout

South Africa during the period 1974 to 1984. A total of

160 specimens were collected representing most of the

taxa recognized in this genus. This sample was designed

to incorporate considerable morphological and geo-

graphical variation in many taxa. Herbarium voucher

specimens were prepared for identification by the Natio-

nal Herbarium (PRE) after segments of leaf blade mate-

rial had been removed and fixed in FAA (Johansen

1940).

Transverse sections, 10 /zm thick, were prepared after

desilicification in 30% hydrofluoric acid (Breakwell

Bothalia 17,1 (1987)

53

1914), dehydration using the method of Feder & O’Brien

(1968) and infiltration and embedding in Tissue Prep

(Fisher Scientific). The sections were stained in safranin

and fast green (Johansen 1940). The manual scraping

method of Metcalfe (1960) was used to prepare scrapes

of the abaxial epidermis. These were either double-

stained in methylene blue and ruthenium red or only in

safranin. The anatomical detail was recorded photo-

graphically using a Reicherdt Univar microscope and

Ilford Pan F film (ASA 50).

The material fixed in FAA was also used for ultra-

structural studies with a scanning electron microscope.

The leaf blade segments were dehydrated in 2,2-

dimethoxypropane (Merck) for 2 hours and then placed

in 100% acetone for 5 minutes following the method

described by Neumann, Rushing & Mueller (1982). The

material was critical point dried with liquid C02 at 85

atmospheres at 40° C after which it was mounted on

aluminium stubs with double sided tape. It was then

glow discharge coated with a thin (± 400 A°) layer of

metallic gold in a sputter coater. Preparations were either

stored in a dessicator with silica gel or observed directly

with an ISI SX-25 scanning electron microscope ope-

rated at 25 Kv accelerating voltage. Cuticular structure

was photographed with a 6 x 7 cm camera on Ilford

FP4 120 film at varying magnification.

HISTORICAL BACKGROUND

The classification o/Ehrharta within the Poaceae

The placement of Ehrharta in the grass family has

varied at both the tribal and subfamilial level (Table 1).

Most older systems, based only on spike let morphology,

place Ehrharta in Phalarideae because of the two sterile

florets below the single fertile floret. Modem treatments

based on a number of cryptic characters place the genus

in a tribe of its own, Ehrharteae. Although first distin-

guished as an entity by Link (1827), his authorship of the

name cannot be accepted for nomenclatural reasons

(Willemse 1982). Thus, Nevski (1937) is the recognized

author for Ehrharteae, which was also described in error

as new by Tateoka (1957). Nees (1841), in the earliest

treatment of southern African grasses that adopted a

suprageneric classification, placed Ehrharta in the

Oryzeae, although Trinius (1839), working in co-opera-

tion with Nees, had earlier placed it in the Phalarideae,

where it remained until the modem system of Stebbins &

Crampton (1961), who placed the tribe back into the

Oryzoideae.

Although there is now agreement on tribal composi-

tion, the question of subfamily classification is not yet

settled. The most recent treatments include Ehrharteae in

Bambusoideae (Renvoize 1985; Watson etal. 1985), but

both discuss the difficulty of including Ehrharteae in the

subfamily as they constitute it. Renvoize (1985) states

that there is no clear correlation between anatomical and

other characters, such as embryo type and lodicules, and

Watson et al. (1985) indicate that, although they decided

to treat the group as bambusoid, other numerical

analyses either place it as arundinoid or lose it alto-

gether. Soderstrom & Ellis (in prep.), who classify bam-

busoid genera and allies, place the Ehrharteae outside the

Bambusoideae, closer to Arundinoideae.

This confused situation may be explained partly

because, as Table 1 indicates, the modem subfamily

classifications are based on a very small set of data

derived from only a few species. In this century, only

Tateoka (1963) has examined more than three taxa, and

only four taxa in all have been examined either for chro-

mosomes, embryos or leaf blade anatomy. The detailed

study of the southern African taxa of Ehrharta presented

in this and following papers should provide the breadth

of information needed to indicate the natural affinities of

this fascinating but baffling genus.

The southern African species o/ Ehrharta

The study of African Ehrharta falls into three phases,

which correspond roughly to the 18th, 19th and 20th

centuries. The earliest phase was characterized by the

description of species in four different genera. L. C. M.

Richard (1779) described Trochera striata, and a few

months later Thunberg (1779) described Ehrharta

capensis. These names were based on different types but

represent the same taxon. Soon after, Linnaeus (1791)

described two more species, but placed them in the

genus Aira. Lamarck (1786), J. E. Smith (1789, 1790)

and J. F. Gmelin (1791) each described species in Thun-

berg’s genus Ehrharta. However, Thunberg himself

(1794) accounted for his specimens collected from the

Cape in Melica, so that his Prodromus plantarum capen-

sium treatment comprised two genuine Melica species

and four species of Ehrharta.

The second phase, one of consolidation, began in the

next century with Swartz’s (1802) thorough and beauti-

fully illustrated study of the genus, in which he recog-

nized nine species, each with synonyms. He described

no new taxa, but moved Thunberg ’s ‘Melicas’ to

Ehrharta. He was aware of, but did not take up, Tro-

chera as the name for the genus. He also did not always

follow priority of publication in applying the specific

epithets, although his synonymy within each species is

otherwise sound. Palisot de Beauvois (1812) alone took

up the name Trochera, to which he attributed two

species.

Schrader (1821) covered 17 species, including all

those in Swartz’s (1802) treatment as well as eight new

species that were based on the specimens of Hesse. He

was the first to divide the genus into sections, distin-

guishing two entities on the character of bulbous versus

fibrous ‘roots’. Although published two years later,

Thunberg’s (1823) Flora capensis treatment was merely

a summary of Swartz (1802), and did not include

Schrader’s species. During this period of consolidation,

a few new species were described by Kunth (1829), by

Schultes (1830) and by Steudel (1853).

Most ambitious of the nineteenth-century studies of

Ehrharta were those of Nees ab Esenbeck (1832, 1839,

1841). His Florae Africae Australioris (1841) recog-

nized 25 species, of which nine were described by him,

using collections of Drege and Ecklon. Nees also put

forward a multitude of varieties (nine in E. calycina

alone), which, however were not validly published be-

cause he did not consistently apply Articles 24-27 of the

ICBN (Linder 1985). He adopted Schrader’s basic divi-

sion of the genus by ‘root’ type, and in addition sub-

divided the fibrous-rooted species according to size and

hairiness of the sterile lemmas. These subdivisions are

54

Bothalia 17,1 (1987)

TABLE 1. — Subfamily and tribal classifications of Ehrharta by various authors

still recognized, although a number of the species have

been re-aligned.

The last of the comprehensive nineteenth-century

accounts of Ehrharta was that of Steudel (1853). He

compiled the species known at that time, 32 in all for

southern Africa. Of greatest significance is his basic

division of these species into two groups, not by the

possession of a bulbous base, as previously, but by the

separation of E. setacea and E. rupestris into a group

(‘Racemosae’) distinct from the rest of the species (‘Pan-

iculatae’). It is unfortunate that he did not elaborate on

the characters on which he based his decision, because

this fundamental division in Ehrharta is supported by the

present study. After 1855, no further work was done in

the genus except that Kuntze (1891), rediscovering the

priority of the genus name Trochera, transferred all the

then-accepted taxa to Trochera.

The third phase in the taxonomic study of Ehrharta

has been dominated by Stapfs (1900) Flora capensis

treatment, which is the most recent critical study of the

southern African taxa. He also recognized 25 species, of

which six were newly described by him, and he placed a

number of previously recognized species into synonymy.

Chippindall (1955) adopted Stapfs concepts, only mak-

ing changes in the keys and stressing close relationships

between some of Stapfs taxa. She included a new

species of C. E. Hubbard (1933), but omitted that of

Mez (1921). Since 1955, new taxa have been described

by Launert (1961) and Gibbs Russell (1984a, 1984b).

Stapf did not follow Kuntze (1891) in taking up the

name Trochera, and soon after the conservation of

generic names was permitted, Ehrharta of Thunberg was

conserved over Trochera of L. C. M. Richard, which

was rejected in ICBN (Voss etal. 1983).

TAXONOMIC CHARACTERS

The taxa described by previous workers can be

grouped into seven easily recognizable groups on the

basis of morphological and anatomical characters as

shown in Table 2. A brief discussion of those characters

found to be useful in this regard follows.

Morphological characters

1 . Macromorphological characters

For a genus of moderate size, Ehrharta has a wide

range of macromorphological characters, a selection of

which are shown in Table 2. They are reasonably well

known, from the detailed descriptions of Stapf (1900)

and the briefer but more image-creating descriptions of

Chippindall (1955). However, unless the species are

placed in groups based on spikelet morphology and leaf

blade anatomy, the diversity of vegetative characters is

confusing. With these groupings, it can be seen that

Bothalia 17,1 (1987)

55-56

TABLE 2. — Morphological and anatomical characters used to differentiate species groups in Ehrharta. Each group is referred to by the name of a widespread species that exhibits the characters of the group

3. MESOPHYLL:

Arm cells (A)

Compact isodiametric (B)

Compact angular (C)

Diffuse irregular (I)

4. EPIDERMAL ZONATION:

Differentiated (D)

Undifferentiated (U)

5. INTERCOSTAL LONG CELLS:

Hexagonal length (JU)

Rectangular (R)

Sinuous walls (S)

Straight walls (W)

6. STOMATA:

Absent (A)

Wax bodies present (P), absent (X)

Epidermal flanges (E)

7. SILICA BODIES:

Paired (P), Short rows (S)

Rounded (R) Dumbell (D)

8. PRICKLES:

Hook-like (H), Macrohair-like (M)

9. MICROHAIRS:

Distal cell truncated (X), not taper-

ing (Y), tapering (Z)

10. EPICUTICULAR WAX:

Present (P), Absent (A)

(D)

U

(D)

U

u u u u u u

<10 <10 <10 <10 <10 <10 <10

s s s s s s s

H H H H H H H

X

p

X

p

X

A

u

<10

s

D

D

10-20 10-20

S S

* 10-20 10-20

* c c

(W) (W)

H H

M M

* Z

* p

>20 >20 >20

WWW

M M

S

D

M

W

S

D

H,M

D D

>20 10-20 >20 >20 >20 >20 >20 >20

W W W W

(S)

w

w w

h,m H.M H.M H,M h m

(B)

c

10-20

R

S

(B) B

C

D D

H H

~~U ” ~ 3 h r. trimvtata 4 E setacea subsp. setacea; 5, E. setacea subsp. uniflora ; 6, E. setacea

_ , „ oro ioHiratpd hv number as follows: 1, E. rupestris subsp. rupestris; 2, E. rupestris subsp. dodii; 3, E. rupestris suosp. in h/irhinndis' 15 £ loneiflora; 16, E. erecta Var. erecta; 17, E. erecta var.

The species and infraspecific ta^a m each group y £ ottonis- 10, E. capensis var. capensis; 11, E. capensis var. intermedia', 12, E. bulbosa; 13, E_eb“me?'. j5 £ pus ilia' 26, E. melicoides; 27, E. gigantea; 28, E. villosa var. villosa;

subsp .xabnr, 7 , E. setacea subsp. dtshcha, 8 \ E lonetfolm. . £ 22, E. brevifolia vai. cuspidata; 23. B. brevifdu, var brevi folia, 24, E catycma, IS.h.pusMc.

“ J nE — “af rSU 32, E vbspica.a; 33, * var. flUfomus; 34, * — 35, £ ^

* anatomical preparations not available.

1

E

E

A

P

R

T

Pi

M

Si

(

Ja

Cl

Cl

(

Re

Re

Wa

stil

bet

act

co

SOI

di\

po:

sep

(‘F

icu

the

thi:

pre

the

prit

the

1

has

trea

sou

whi

nun

Chi

ing

bet\

spet

Bothalia 17,1 (1987)

57

there are a few parallel trends that occur in several

species groups, and that a number of differences in struc-

ture are associated with each trend. For example,

suffrutescence occurs in four of the species groups (Seta-

cea, Capensis, Gigantea and Ramosa), and involves, in

each group, long branched rhizomes, much branched

culms that serve as the main photosynthetic organ, and

reduction of leaf blades.

a. Habit in the majority of Ehrharta species is peren-

nial, but several are annual. The annual habit seems

related to unpredictable moisture availability. All the

annual species, except one, occur in the drier north-

western part of the range of the genus, centred on the

Succulent Karoo, from about Clanwilliam north to

southern SWA/Namibia. These species fall only in the

Erecta and Calycina groups, and each has closely related

perennial species in the more mesic central part of the

distribution of the genus. The other annual species, E.

longiflora, grows in seasonally wet places, and it has,

for an Ehrharta, a wide distribution, occurring not only

in arid areas to the northwest, but as far east as Port

Alfred. It is the only species in which the spikelet

morphology is not consistent with the leaf blade

anatomy.

b. Rhizomes occur in all perennial species, but the

form of the rhizome varies widely, and is correlated with

the substrate. Species that grow between rocks have

tough, branched, naked rhizomes that are hardly diffe-

rent from the lower part of the culm. These rhizomes can

creep for several meters in deep cracks, so that the plant

forms a dense linear cushion. Species that occur in sand

have long branching rhizomes that run 0,3 m or more

below the soil surface and send up culms at the tips, so

that a single plant can appear to be a loose colony several

meters across. These rhizomes may be naked or densely

clothed by hairy cataphylls. Species that grow in clay or

loam soils usually have short knotted rhizomes that do

not spread far away from the base of the individual plant,

although in a few species unbranched rhizomes a few

centimeters long and with papery cataphylls may occur.

c. Culm characteristics are the most important in

determining the overall appearance of a grass plant, and

in Ehrharta the culms are very diverse. In different taxa

the culms vary from solitary to numerous, from herba-

ceous to wiry or woody, from erect to procumbent, from

unbranched to simply or verticillately branched. Length

varies between taxa from less than 100 mm to over 1 ,5 m

in flowering individuals.

Taxa in the Capensis groups have the lowest (some-

times the two lowest) intemodes modified into an

extremely hard cylindrical or globose bulb-like structure.

The occurrence of this ‘bulb’ may be associated with the

periodic fires in the fynbos. ‘Bulbous’ species are much

in evidence for a year or two after a fire, but apparently

do not occur in long-unbumed situations. However,

immediately following the next bum, flowering plants

may be found with many old dead ‘bulbs’ hidden under-

ground, indicating that the plant is several years old,

having survived the fireless period in this form.

Stolons are known only in a local limestone variant of

the widespread species E. calycina.

d. Leaves, as in other grasses, are composed of

sheath, ligule and blade. The sheaths are split to the base

and usually cylindrical, or rarely keeled. In some species

the upper sheaths may be longer than the intemodes, so

the leaves are closely imbricate. In suffrutescent species

with reduced leaf blades the sheaths may be persistent

around the culm, or be held outward at an angle, or they

may be lost. The basal sheaths may be persistent,

densely clothing the base of the culm, or they may slip

away from the culm with age, leaving it exposed. A few

taxa may be distinguished by a characteristic colour of

the basal sheaths.

The ligules are short, usually less than 3 mm long, and

take the form of a ciliate rim in all species groups except

the Erecta and Calycina groups, in which most taxa have

glabrous ligules.

The leaf blades vary from broad and flat, often with

one or both margins with a heavy undulate marginal

vein, to folded, rolled, setaceous or absent. Length

varies between taxa, from less than 100 mm to over 1 m.

Loss or reduction of leaf blades is correlated with suffru-

tescence. In these taxa, the phenology of leaf blade

development should be studied in the field, because it is

possible that striking differences in appearance may be

due to the age of a plant rather than to genetic dif-

ferences.

e. Inflorescences are most commonly paniculate, but

in some taxa may be reduced to a raceme with few spike-

lets, and there is often a tendency for the narrower inflo-

rescences to be secund. Generally the inflorescence is

exserted far above the highest leaf sheath, but in a few

taxa (especially annuals) it is closely subtended or even

enveloped below by an inflated leaf sheath. In some taxa

with racemose inflorescences the main axis curves sinu-

ously around the appressed spikelets.

2. Spikelet morphology

The spikelets are usually laterally compressed,

although in some taxa the sides are rounded. At matu-

rity, the spikelet is shed as a unit above the glumes,

which are persistent on the pedicel. A spikelet consists of

a pair of glumes, two empty sterile lemmas, and, at the

tip of the rachis, a fertile floret composed of lemma,

palea and bisexual flower. Each species group has a

characteristic range of spikelet sizes.

a. Glumes are subequal, and may be shorter than,

equal to or longer than the whole spikelet, and relative

glume length is a useful character for distinguishing

between species in a group. The glumes are more papery

in texture than the lemmas, and are unomamented.

b. Sterile lemmas are the single most striking spikelet

feature of Ehrharta. Their curious sculpturing in some

species is unknown elsewhere in the grass family, and

renders a detached spikelet immediately recognizable.

Shape, relative size and. ornamentation of the sterile lem-

mas are the most useful characters in separating the

species groups. In three of the species groups, Erecta,

Calycina and Dura, the tips of the sterile lemmas can

have long awns, and in the Capensis and Villosa groups

the bases are shortly stipitate. In most Ehrhartas the ster-

ile lemmas are of similar size and the smaller fertile

lemma differs from them, but in the Setacea group the

first sterile lemma is short and glume-like and the fertile

lemma is similar to the second sterile lemma.

58

Bothalia 17,1 (1987)

The lemma surfaces may be dull or shining, scaberu-

lous or smooth. The sides and margins may be glabrous

or have hairs of various lengths, and the bases may be

glabrous or bearded. The bases of the second sterile

lemma and the fertile lemma come together in a hinge-

like joint that resembles an earlobe, and which may have

a membranous appendage. All these characteristics are

used in separating the species groups. Differences in

transverse and longitudinal ribs and veins distinguish

species within the groups.

c. Fertile florets have a lemma that is smaller and

more laterally compressed than the sterile lemmas, and is

unomamented. The palea is sickle-shaped or straight, is

much smaller then the lemma and is usually hidden

within it. The fertile florets of most species are similar,

and are not useful for distinguishing species or groups.

d. Flower structure is remarkable in Ehrharta because

of the presence of six stamens in most species, although

some of the taxa with small spikelets may have five,

four, three or one stamens. The two lodicules are relati-

vely large and flat, and are usually ovate or 2-lobed.

e. Cary op sis information for comparison is lacking

because of the small number of spikelets that are found

to have mature fruits. It appears that the spikelet is shed

as soon as the fruit is mature. The lack of this informa-

tion, as well as data about embryos, is a serious defect in

our knowledge of the genus.

Anatomical characters

A suite of leaf blade anatomical characters separate

and define the various species groups in Ehrharta. These

cryptic features are manifested in the leaf blade as seen

in transverse section, in the abaxial epidermis in surface

view and ultrastructurally (Table 2). Some of these

characters distinguishing the species groups recognized

in Ehrharta are generally considered to be features of

high taxonomic value, important at the subfamily level

in the Poaceae. In Ehrharta, however, they separate taxa

below the generic level. Examples are differences in

mesophyll structure and microhair shape.

The 10 characters which were scored for all the

southern African Ehrharta taxa are diagramatically

represented in Table 4. This suite of anatomical charac-

ters in combination serves to define and diagnose each of

the seven species groups as well as two subgroups within

one of the groups. The variation encountered in most of

the characters in most of the groups will be fully

described and discussed in subsequent papers. The

character states used to define each of the groups here

are, therefore, somewhat generalized and may vary wi-

thin certain limits and intergrade between certain groups.

In the cases where variation was observed, definite lines

of development could be traced and the variation could

usually be interpreted on this basis.

1 . Midrib or keel

The vascular structure of the keel or midrib is an

important character in grass taxonomy at the subfamily

level. Complex vasculature of the keel would definitely

support bambusoid affinities as suggested by many

authors. This type of keel does not occur in any of the

species groups from South Africa but the presence of

only a median vascular bundle or the presence of a defi-

nite keel with additional parenchyma tissue differs

between the species groups recognized, and only varies

within the Calycina group.

2. Ribs and furrows

Adaxial ribs and furrows, particularly the massive ribs

of the Longifolia group, distinguish certain groups. The

raised, inflated abaxial epidermal cells in the mid-inter-

costal zones of the Calycina group are diagnostic and of

considerable phylogenetic interest because this charac-

teristic is shared with the Phalarideae with which the

Ehrharteae has been linked by earlier authors. These

epidermal cells are also evident in the epidermis as seen

in surface view.

3. Mesophyll

The chlorenchyma cells of Ehrharta are surprisingly

variable in structure and arrangement. In the Setacea

group arm cells definitely are present in some taxa. This

is a bambusoid characteristic but is also known in some

taxa without bambusoid or oryzoid affinities (Watson et

al. 1985). In some groups the chlorenchyma cells are

compact and angular with minute air spaces but, in other

groups this tissue is very diffuse, of irregular, rounded

cells with air spaces clearly visible. In the Villosa group

the abaxial layer of chlorenchyma is very regular and

almost palisade-like.

4. Epidermal zonation

Some groups are distinguished by costal and intercos-

tal zones on the abaxial surface. Zonation is not evident

in the Setacea and Longifolia groups and this is asso-

ciated with an absence of abaxial stomata as observed in

these two groups. This lack of zonation appears to be

associated with the ecological conditions of the Moun-

tain Fynbos with very low soil nutrition and also occurs

in many of the danthonoid grasses from the same habi-

tats. Species from the Lowland Fynbos, on the other

hand, have clear epidermal zonation and abaxial sto-

mata. These characters tend to vary in groups which

have a wide ecological tolerance and occur in both these

two major habitat types.

5. Intercostal long cells

Hexagonal or inflated but rather short long cells with

sinuous walls distinguish some groups, whereas others

have elongate, fusiform, hexagonal intercostal long cells

with straight walls. In the Calycina group with this latter

type of long cell, the mid-intercostal cells are much

longer than the lateral ones and these cells are also the

raised, inflated cells as seen in transverse section. In

other groups the intercostal long cells are rectangular

with sinuous walls.

6. Abaxial stomata

The stomata of the various species groups of Ehrharta

are also of taxonomic interest. They may be absent on

the abaxial surface, as in the Longifolia group and in

many taxa of the Setacea group. In the Capensis group

the guard cells are clearly visible but the subsidiary cells

are covered with wax platelets, as seen with the scanning

electron microscope. In the Erecta group the stomata are

often obscured by large, solid wax plugs, whereas, in the

Calycina group cubical wax granules often overlie the

stomata. In the Villosa group the stomata are sunken and

overarched by four characteristic papilla-like flanges

from the adjoining long cells. In the Ramosa group the

Bothalia 17,1 (1987)

59

stomata have no associated wax deposits whereas in the

Dura group the pores are obscured by dense accumula-

tions of wax platelets. Stomatal structure, and associated

epicuticular wax, therefore, serves to distinguish all but

two of the species groups in Ehrharta and provides use-

ful characters at the species level. The low dome-shaped

subsidiary cells, which are present throughout Ehrharta,

are typical of, but not exclusive to, the Arundinoideae

(Renvoize 1981).

7. Costal silica bodies

The silica bodies of Ehrharta are somewhat variable

and rather difficult to use as diagnostic characters.

Single, rounded, or paired bodies characterize some

groups whereas in others this type of body is more dumb-

bell-shaped. In the Erecta and Calycina groups the costal

silica bodies are in short chains and of the dumbbell type

but this shape can only be determined by varying the

focus. Silica body shape and arrangement is often a

character differentiating taxa at higher taxonomic levels

than that of genus.

8. Prickles

The costal macrohair-like type of prickle hair is only

found in the Erecta and Calycina groups. This type ol

hair is common in the pooid and bambusoid grasses but

is much rarer in the arundinoids. In the other species

groups of Ehrharta, only shortly barbed prickles occur,

except in the Longifolia subgroup of the Capensis group

where prickle hairs are absent.

9. Microhairs

Microhairs in Ehrharta are very small and difficult to

observe with the light microscope. However, they are

clearly visible with the SEM and clear differences

between the species groups are visible. The hairs may

have short, truncated distal cells or this cell may taper to

an acute apex. The length of the hairs also differs

between the species groups. The Longifolia subgroup is

the only grouping that does not possess abaxial micro-

hairs. Microhair shape is a very important character

separating the subfamilies of the Poaceae and the degree

of variation observed in Ehrharta is most unusual.

10. Epicuticular wax

The occurrence and nature of the epicuticular wax

deposits also appears to be a useful taxonomic character

in Ehrharta, serving to distinguish between the species

groups. The wax may be absent or present either as fine

rods or as heavier platelets. The significance of this wax

as a feature of higher taxonomic application is unknown.

The suite of leaf anatomical characters used to

distinguish the species groups in Ehrharta, therefore,

includes a wide spectrum of attributes and, in combina-

tion, these features serve to assign any given specimen to

a particular group. However, it is not easy to identify to

the species level using leaf anatomical criteria because

the species are not so distinct anatomically. Neverthe-

less, the recognition and definition of the seven groups

of species in this very difficult genus will definitely help

in providing new insights into the phylogenetic relation-

ships of the genus, until such time as sufficient embryo

and chromosome data are available.

OUTLINE OF SPECIES GROUPS

Each of the species groups in Ehrharta can be

described on the basis of a set of morphological and

anatomical characters as summarized in Tables 3 & 4

respectively. A brief synopsis of characters and diagno-

sis is then given for the constituent taxa of each group.

SETACEA GROUP

Included taxa (Gibbs Russell 1984)

E. rupestris Nees ex Trin.

subsp. rupestris

subsp. tricostata (Stapf) Gibbs Russell

subsp. dodii (Stapf) Gibbs Russell

E. setacea Nees

subsp. setacea

subsp. scabra (Stapf) Gibbs Russell

subsp. uniflora (Burch, ex Stapf) Gibbs Russell

subsp. disticha Gibbs Russell

Morphological description

1. Perennial.

2. Culms herbaceous or suffrutescent.

3. No parts bulbous.

4. Leaf ligules ciliate.

5. Leaf blades expanded in herbaceous taxa, either setaceous or

folded in suffrutescent taxa.

6. Spikelets fewer than 20.

7. Spikelets 4—8 mm long.

8. Glumes one third to almost equalling lemma length.

9. First sterile lemma reduced and glume-like, with veins.

10. Second sterile lemma base not stipitate, lacking appendages.

11. Sterile lemma surface rough.

12. Sterile lemma sides glabrous.

13. Sterile lemma bases not bearded.

14. Sterile lemma tips awnless, not mucronate.

Anatomical description

1 . No keel developed; only median vascular bundle present.

2. Adaxial ribs and furrows present; ribs rounded and well developed

(except in E. setacea subsp. disticha and subsp. uniflora).

3. Mesophyll compact, of small rounded or isodiametric cells;

tendency for arm cell-like invaginations in all taxa; E. setacea

subsp. scabra with typical bambusoid-like arm cells.

4. Costal and intercostal zones not differentiated (except in those

species with abaxial stomata — E. setacea subsp. disticha and

subsp. uniflora ).

5. Intercostal long cells inflated to hexagonal; short, less than 20 pm

long (usually less than 10 /am long); with sinuous walls.

6. Abaxial stomata absent (except in E. setacea subsp. disticha and

subsp. uniflora and then without wax platelets).

7. Costal silica bodies single or paired or absent; rounded.

8. Costal and intercostal prickles present; either hooks or asperites

with very short barbs; absent in E. setacea subsp. disticha and

subsp. uniflora.

9. Microhairs with short, truncated distal cell; longer in E. setacea

subsp. disticha and subsp. uniflora.

10. Epicuticular wax either absent or as fine rods.

Distinguished by:

First sterile lemma reduced, glume-like. Spikelets fewer than 20.

Arm cells present.

CAPENSIS GROUP

Included taxa (Gibbs Russell 1984a; Smook & Gibbs Russell 1985)

E. longifolia Schrad.

E. oltonis Kunth ex Nees

60

Bothalia 17,1 (1987)

TABLE 3. — Morphological characters diagnostic for the species groups in Ehrharta

E. barbinodis Nees ex Trin.

E. capertsw Thu nb.

E. bulbosa J. E. Sm.

E. eburnea Gibbs Russell

Morphological description

1. Perennial.

2. Culms herbaceous (except E. barbinodis where they are suffru-

tescent).

3. Culm base bulbous at lowest intemode (except E. barbinodis ) .

4. Leaf ligulesciliate.

5. Leaf blades expanded or rolled (but very short in E. barbinodis).

6. Spikelets many (but less than 35 in E. eburnea).

7. Spikelets 7-13 mm long.

8. Glumes 1 4-/4 lemma length.

9. First sterile lemma not reduced, lacking veins.

10. Second sterile lemma base stipitate, lacking appendages.

11. Sterile lemma surface rough.

12. Sterile lemma sides glabrous.

13. Sterile lemma bases bearded.

14. Sterile lemma tips awnless, mucronate.

Distinguished by:

Lowest culm node bulbous (except E. barbinodis). Second sterile

lemma stipitate. Spikelets large, 7-13 mm long, hairy at margins.

Anatomical description

Anatomically this group cannot be defined satisfactorily as the taxa

included vary greatly in anatomy. Nevertheless, two distinct and ana-

tomically homogeneous subgroups can be distingushed and these will

be dealt with separately.

LONGIFOLIA SUBGROUP

Included taxa (Smook & Gibbs Russell 1985)

E. longifolia Schrad.

E. ottonis Kunth. ex Nees

Anatomical description

1 Keels absent; median vascular bundle undifferentiated from lateral

first order vascular bundles.

2. Massive adaxial ribs and deep, cleft-like furrows with interlocking

prickles.

3. Mesophyll compact, of large angular cells; in U-shaped groups

occupying the sides and bases of the furrows.

Bothalia 17,1 (1987)

61

TABLE 4. — Anatomical characters diagnostic for the species groups in Ehrharta

4. Costal and intercostal zones only slightly differentiated.

5. Intercostal long cells inflated and short; less than 10 gm long; with

sinuous walls.

6. Abaxial stomata absent.

7. Costal silica bodies paired or single; rounded.

8. Costal prickles absent.

9. Abaxial microhairs absent.

10. Epicuticular wax absent.

Distinguished by:

Setaceous leaves without arm cells.

CAPENSIS SUBGROUP

Included taxa (Gibbs Russell 1984a; Smook & Gibbs Russell 1985)

E. barbinodis Nees ex Trin.

E. capensis Thunb.

E. bulbosa J. E. Sm.

£. eburnea Gibbs Russell

Anatomical description

1. Keel present comprising 1 or 3 vascular bundles (E. capensis) or

absent (E. barbinodis)-, margin with conspicuous sclerenchyma

cap.

2. Adaxial ribs and furrows absent.

3. Mesophyll semi-radiate; of compact but large, angular cells.

4. Costal and intercostal zones well differentiated.

5. Intercostal long cells elongate hexagonal, fusiform; between

10-20 gm long; with slightly sinuous walls.

6. Abaxial stomata present; subsidiary cells covered with wax plate-

lets but pore visible.

7. Costal silica bodies single or paired; irregularly dumbbell-shaped.

8. Costal prickles with short barbs and intercostal prickles elongated,

macrohair-like with bulbous bases.

9. Microhairs elongate, with tapering distal cell.

10. Epicuticular wax deposits as fine rods.

Distinguished by:

Presence of midrib and hexagonal long cells less than 20 gm long;

stomata with wax platelets.

ERECTA GROUP

Included taxa (Smook & Gibbs Russell 1985)

E. e recta Lam.

var. erecta

var. natalensis Stapf

var. abyssinica (Hochst. ) Pilg.

£. triandra Nees ex Trin.

£. longiflora J. E. Sm. (but spikelets similar to Capensis group)

Morphological description

1. Perennial or annual.

2. Culms herbaceous.

3. No parts bulbous.

4. Leaf ligules glabrous.

5. Leaf blades expanded.

6. Spikelets many.

62

Bothalia 17,1 (1987)

7. Spikelets 3, 0-7, 5 mm long.

8. Glumes lA-2A lemma length.

9. First sterile lemma not reduced, veined.

10. Second sterile lemma base not stipitate, lacking appendages.

11. Sterile lemma surface rough (but smooth in some specimens of

E. erecta subsp. erecta).

12. Sterile lemma sides glabrous.

13. Sterile lemma bases bearded (except in E. erecta subsp. erecta and

E. triandra).

14. Sterile lemma tips not mucronate.

Anatomical description

1. Keel present; comprises one vascular bundle with ground

parenchyma.

2. Very slight adaxial ribs and wide, shallow furrows present; may be

absent.

3. Mesophyll rather diffuse of somewhat irregular, rounded cells; air

spaces clearly visible.

4. Costal and intercostal zones clearly differentiated.

5. Mid-intercostal long cells hexagonal, fusiform in shape; elongate,

more than 30 gm long; straight- walled.

6. Abaxial stomata present; subsidiary cells always with wax depos-

its, often in the form of a solid plug blocking the stomatal aper-

ture.

7. Costal silica bodies in short chains; usually dumbbell-shaped with

the central part only visibly by varying focus.

8. Costal and intercostal prickles present; all taxa with the macrohair-

like type of prickle; asperites sometimes occur.

9. Microhairs elongate with markedly tapering distal cell.

10. Epicuticular wax always present.

Distinguished by:

Spikelets small, and first sterile lemma well developed, and glumes

short, and sterile lemmas constricted at base, lacking appendages, and

lemma sides glabrous. Raised abaxial epidermal long cells absent and

stomata with wax plugs.

CALYCINA GROUP

Included taxa (Smook & Gibbs Russell 1985)

E. brevifolia Schrad.

var. brevifolia

var. cuspidata Nees

E. calycina J. E. Sm.

E. delicatula (Nees) Stapf

E. longigluma C. E. Hubb.

E. melicoides Thunb.

E. pusilla Nees ex Trin.

Morphogical description

1. Perennial or annual.

2. Culms herbaceous.

3. No parts bulbous.

4. Leaf ligules glabrous or ciliate.

5. Leaf blades expanded.

6. Spikelets many.

7. Spikelets 3, 2-8, 5 mm long.

8. Glumes two thirds longer than lemma length.

9. First sterile lemma not reduced, veined.

10. Second sterile lemma base not stipitate, with ear-like appendages.

11. Sterile lemma surface smooth.

12. Sterile lemma sides glabrous or hairy.

13. Sterile lemma bases hairy (but not conspicuously bearded) in hairy

taxa, but glabrous taxa not bearded.

14. Sterile lemma tips mucronate or rounded.

Anatomical description

1. Keel absent, except in some specimens of E. calycina and

E. brevifolia var. cuspidata.

2. Slight adaxial ribs and furrows; may be well developed in

E. calycina and E. melicoides and may be absent in E. calycina

and E. brevifolia var. cuspidata; raised, inflated abaxial epidermal

cells always present in mid-intercostal zones.

3. Mesophyll generally compact of irregular (usually quite large),

straight-walled cells; air spaces small; chlorenchyma cells often

striated and refractive, possibly silicified.

4. Costal and intercostal zones clearly differentiated.

5. Mid-intercostal long cells elongated, fusiform (more than 25 /am

long); raised; with straight walls; shorter in E. longigluma; these

cells often stain in E. calycina and E. melicoides.

6. Abaxial stomata always present; pore aperture often obscured by

cubical wax granules.

7. Costal silica bodies in short chains; variably dumbbell-shaped of

the type which varies in shape with differing focus.

8. Costal and intercostal prickles present; sometimes with short barbs

only (E. pusilla) but otherwise all taxa with macrohair type of

prickle.

9. Microhairs finger-like with tapering distal cell.

10. Epicuticular wax always developed.

Distinguished by:

Spikelets small, and glumes long, and first sterile lemma well

developed, and second sterile lemma base not stipitate, with ear-like

appendages. Raised mid-intercostal long cells and cubical wax

granules associated with stomata.

VILLOSA GROUP

Included taxa (Smook & Gibbs Russell 1985)

E. gigantea Thunb.

E. villosa Schult. f.

var. maxima Stapf

var. villosa

Morphological description

1. Perennial.

2. Suffrutescent.

3. Rhizome intemodes sub-bulbous fE. gigantea).

4. Leaf ligules ciliate.

5. Leaf blades reduced, rolled.

6. Spikelets many.

7. Spikelets 10-18 mm long.

8. Glumes one half to longer than lemma length.

9. First sterile lemma not reduced, veined.

10. Second sterile lemma base stipitate, lacking appendages.

1 1 . Sterile lemma surface smooth.

Bothalia 17,1 (1987)

63

12. Sterile lemma sides profusely hairy.

13. Sterile lemma bases conspicuously bearded.

14. Sterile lemma tips mucronate.

Anatomical description

1 . Keel absent; median vascular bundle identical to lateral first order

bundles.

2. Adaxial ribs and furrows present; well developed; rounded ribs of

medium size.

3. Mesophyll semi-radiate of large, angular cells; abaxial layer of

chlorenchyma regular and somewhat palisade-like in arrangement.

4. Costal and intercostal zones differentiated.

5. Intercostal long cells rectangular in shape; sinuous walls; some-

times mid-intercostal long cells tend to be hexagonal.

6. Abaxial stomata present; sunken and overarched by four papilla-

like flanges from the adjacent epidermal cells.

7. Costal silica bodies absent or irregular in occurrence; single or

paired; round in shape.

8. Prickles absent (E. villosa) or costal and intercostal prickles

present (E. gigantea); hairs not macrohair-like.

9. Microhairs rather short but with tapering distal cell.

10. Epicuticular wax present but variable.

Distinguished by:

Culms over 1 m long, suffrutescent. Leaf blades reduced, rolled.

Spikelets very large, lemmas profusely hairy and conspicuously

bearded, stipitate, mucronate. Stomata with four epidermal flanges.

RAMOSA GROUP

Included taxa (Smook & Gibbs Russell 1985)

E. ramosa (Thunb. ) Thunb.

E. rehmannii Stapf

var. rehmannii

var. filiformis Stapf

E. subspicata Stapf

Morphological description

1. Perennial.

2. Herbaceous or suffrutescent.

3. Upper intemodes sometimes form bulbous galls in E. ramosa and

E. rehmannii var. filiformis.

4. Leaf ligules ciliate.

5. Leaf blades expanded to rolled in herbaceous taxa, absent or very

reduced in suffrutescent E. ramosa.

6. Spikelets few.

7. Spikelets 5, 5-8, 8 mm long.

8. Glumes two thirds longer than lemma length.

9. First sterile lemma not reduced, veined.

10. Second sterile lemma not stipitate.

1 1 . Sterile lemma surface rough.

12. Sterile lemma sides glabrous (few short hairs in E. rehmannii var.

rehmannii).

13. Sterile lemma bases bearded.

14. Sterile lemma tips truncate, not mucronate.

Anatomical description

1 . No keel; median vascular bundle only.

2. Ribs and furrows either absent or very slight ribs only present.

3. Mesophyll compact of small to medium, angular, isodiametric

chlorenchyma cells.

4. Costal and intercostal zones differentiated.

5. Intercostal long cells rectangular but tending to hexagonal shape in

mid-intercostal files in a few specimens; walls always sinuous.

6. Abaxial stomata present; no associated wax deposits; with distinct

stomatal rims.

7. Costal silica bodies single or paired; irregular in shape being short

dumbbell-shaped to rounded.

8. Costal and intercostal prickles present; with short to long barbs but

not of the macrohair-like type.

9. Microhairs with distal cell not tapering.

10. Epicuticular wax absent.

Distinguished by:

Spikelets small, and sterile lemmas well developed, with tips

rounded, sides rough and glabrous, bases glabrous and with basal

appendages. Stomata with distinct rims and no wax deposits.

DURA GROUP

Included taxa (Smook & Gibbs Russell 1985)

£. dura Nees ex Trin.

E. microlaena Nees ex Trin.

Morphological description

1. Perennial.

2. Herbaceous.

3. No parts bulbous.

4. Leaf ligules glabrous.

5. Leaf blades well developed (setaceous inf. dura).

6. Spikelets many.

7. Spikelets 9-17 mm long.

8. Glumes W-'/i lemma length.

9. First sterile lemma not reduced, veined.

10. Second sterile lemma base not stipitate, lacking appendages.

1 1 . Sterile lemma surface rough.

12. Sterile lemma sides glabrous.

13. Sterile lemma bases bearded.

14. Sterile lemma tips long awned.

Anatomical description

1 . No keel or midrib present.

2. Well developed adaxial ribs and furrows; ribs flat-topped and

furrows cleft-like.

3. Mesophyll compact, of large angular cells; airspaces not visible.

4. Costal and intercostal zones differentiated.

5 . Intercostal long cells rectangular with slightly sinuous walls; tanni-

niferous cells.

6. Abaxial stomata present; pore obscured by dense accumulation of

wax platelets.

7. Costal silica bodies single or paired; irregularly dumbbell-shaped.

8. Costal prickles only; with very short barbs.

64

Bothalia 17,1 (1987)

9. Microhairs very variable; with sharply tapering point or distal cell

blunt; distal cell apparently not dehiscent.

10. Epicuticular wax absent except in association with the stomata.

Distinguished by:

Spike lets very large, lemmas glabrous. Plants perennial and spike-

lets awned. Tanniniferous cells present and stomatal pores obscured by

wax platelets.

CONCLUSIONS

A better understanding of the taxonomy and systema-

tics of Ehrharta is needed because its morphological and

anatomical variation and its geographic distribution indi-

cate that the genus may provide important clues towards

understanding the natural relationships of the Poaceae.

The southern African species fall into seven groups

based on both spikelet morphology and vegetative ana-

tomy, while the vegetative macromorphology exhibits

parallelism and/or convergence between the groups. The

series of papers to follow will relate the details of mor-

phological and anatomical structure of each species

group to the interpretation of relationships within

Ehrharta, as it is presently circumscribed and within the

rest of the grass family.

ACKNOWLEDGEMENTS

We thank L. Smook for support in field and herba-

rium; W. Roux and H. Ebertsohn for technical assis-

tance; A. J. Romanowski for photography; and S. M.

Perold for electron microscopy. We are especially grate-

ful to J. F. Veldkamp and S. A. Renvoize for meticulous

reviewing of the manuscript.

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.

Bothalia 17,1:67-73(1987)

Taxonomy of the genus Ehrharta (Poaceae) in southern Africa: the

Setacea group

G. E. GIBBS RUSSELL*

Keywords: Fynbos, Capensis, Ehrharta, Poaceae, taxonomy

ABSTRACT

The Setacea species group in the genus Ehrharta Thunb. is differentiated morphologically by the short first sterile

lemma and by inflorescences of fewer than 20 spikelets. The Setacea group is composed of two species, each with

subspecies linked by intraspecific intermediates: E. rupestris Nees ex Trin. subsp. rupestris ; subsp. tricostata (Stapf) Gibbs

Russell; subsp. dodii (Stapf) Gibbs Russell and E. setacea Nees subsp. setacea ; subsp. scabra (Stapf) Gibbs Russell; subsp.

uniflora (Burch, ex Stapf) Gibbs Russell; subsp. disticha Gibbs Russell. All taxa are endemic to the Fynbos vegetation of

the south-western Cape Province, with distribution centred in the Caledon degree grid (3419). Parallel trends for plant size

and habit, leaf blade width and position, and spikelet size are demonstrated in both species, with similar plant types

occurring in similar geographical areas.

UITTREKSEL

Die Setacea-spesiegroep in die genus Ehrharta Thunb. word morfologies onderskei deur die kort eerste steriele lemma

en deur bloeiwyses van minder as 20 blompakkies. Die Setacea-groep bestaan uit twee spesies, elk met subspesies wat deur

intraspesifieke oorgangsvorme by mekaar aansluit: E. rupestris Nees ex Trin. subsp. rupestris ; subsp. tricostata (Stapf)

Gibbs Russell; subsp. dodii (Stapf) Gibbs Russell en E. setacea Nees subsp. setacea ; subsp. scabra (Stapf) Gibbs Russell;

subsp. uniflora (Burch, ex Stapf) Gibbs Russell; subsp. disticha Gibbs Russell. Alle taksons is endemies in die Fynbos-

plantegroei van die suidwestehke Kaap-provinsie, met die verspreiding in die Caledon-gradevierkant (3419) gesentreer.

Parallelle neigings in plantgrootte en -groeiwyse, breedte en posisie van die blaarskyf, en grootte van die blompakkies word

in albei spesies aangetoon, met soortgelyke planttipes wat in soortgelyke geografiese gebiede voorkom.

INTRODUCTION

The genus Ehrharta in Africa comprises 35 species

and infraspecific taxa, and has been divided into seven

species groups by correlating many characters of spikelet

morphology and leaf blade anatomy, as outlined by

Gibbs Russell & Ellis (1987). The leaf blade anatomy in

Ehrharta shows a wide range of variation. Characters

that normally separate groups at subfamily level else-

where in Poaceae are here used to distinguish species

groups. A detailed treatment of variation within each

species and species group is therefore called for. Ellis

(1987) demonstrates the variation and discusses the sig-

nificance of leaf anatomy in the Setacea species group.

This paper presents the taxonomy of the species in the

Setacea group, and places the morphology of the taxa in

context with the anatomy, and with the geographical

distribution. The other species groups will be similarly

treated in parallel papers in the future.

The seven species groups in Ehrharta may be classi-

fied into two major categories, those with the first and

second sterile lemmas similar to each other and different

from the fertile lemma (28 species and infraspecific taxa,

in six species groups), and those with the first sterile

lemma reduced and glume-like, and the fertile lemma

similar to the second sterile lemma (seven species and

infraspecific taxa in one species group, the Setacea

group, Figure 1). In addition, only in the Setacea group

is the number of spikelets always fewer than 20 per inflo-

rescence. Ellis (1987) has distinguished the Setacea

group anatomically by the presence of arm cells in the

* Botanical Research Institute, Department of Agriculture and Water

Supply, Private BagXIOl, Pretoria 0001.

mesophyll (a character normally of significance at sub-

family level in grass classification), the shape of the

microhairs, the silica bodies and the intercostal long

cells. Detailed morphological and anatomical descrip-

tions of the Setacea group are given by Gibbs Russell &

Ellis (1987).

FIGURE 1. — Spikelet of E. rupestris subsp. tricostata: a, whole

spikelet; b, glumes; c, first sterile lemma; d, fertile lemma; e,

second sterile lemma (Esterhuysen 26708) (x6).

Within what is now regarded as the Setacea group,

Nees (1841) recognized two species, and the fundamen-

tal difference of their spikelet plan from that of the rest of

the genus was demonstrated by Steudel (1853). Stapf

(1900) added three new species and a variety, but Gibbs

Russell (1984) placed all of Stapf s taxa, plus an additio-

nal one, as subspecies within Nees’s original two

species. These decisions, based on macromorphology

and spikelet morphology, are generally supported by

68

Bothalia 17,1 (1987)

anatomical studies (Ellis 1987). In most cases, anatomi-

cal characters have allowed better interpretation of con-

fusing morphological states; in other cases, morphologi-

cal and anatomical evidence do not agree. Some of the

variation in the Setacea group, where morphological and

anatomical characters are contradictory, may be a result

of convergent adaptations for tough, long-lived leaves

often encountered in plants of the low-nutrient Fynbos

soils.

In Africa, Ehrharta is a genus characteristic of the

winter rainfall areas, and has its centre of diversity in

Fynbos, but a number of species also occur in Succulent

Karoo, Karoo, Savanna and Forest. Six of the species

groups have taxa in more than one vegetation type. The

Setacea group is unique in being restricted to Fynbos,

and occurs only from the Cape Peninsula north to

Worcester and east to Humansdorp.

KEY TO SPECIES IN THE SETACEA GROUP

Glumes about 'A as long as spikelet 1 E. rupestris

Glumes more than 2A (very rarely only XA) as long as

spikelet 2. E. setacea

1. Ehrharta rupestris Nees ex Trin. in M6moires

de l’Acad6mie imp6riale des Sciences de St-P6tersbourg,

s£r. 6,5: 25 (1839); Nees: 227 (1841); Stapf: 668 (1900);

Chippind.: 37 (1955). Type: Cape Province, Caledon

Distr., Gnadenthal, Drige (K, isosyn.!; SAM, isosyn.!;

PRE, photo.!, fragment!).

Trochera rupestris (Nees) Kuntze: 795 (1891).

Erect or trailing tufted perennial with long creeping

rhizomes, of widely varying size and habit, from delicate

herbaceous plants 100 mm tall to robust suffrutescent

plants 450 mm tall. Culms erect or decumbent, branched

and creeping at base, leafless below. Leaves with blades

erect or spreading, inrolled, folded or flat; ligule a

membrane fringed with hairs; sheaths overlapping.

Inflorescence an erect raceme overtopping leaves. Spike -

lets 1-9, erect, often distichous. Glumes about 'A length

of entire spikelet, lower ones with tip truncate or

rounded, upper ones acute. Florets with sterile lemmas

dissimilar, the first sterile lemma a thin scale with 3-5

raised nerves, XA-'A the length of second, often appear-

ing to be a third glume; the second sterile lemma hard

and thickened, with 7 minutely tubercled nerves, tip

canoe-shaped; fertile lemma similar to second sterile

lemma, but slightly shorter, broader and with more acute

tip. Stamens 6.

(a) subsp. rupestris. Gibbs Russell: 150 (1984).

Plants erect, robust, mat- or cushion-forming, to 300

mm tall, with strong woody rhizomes. Culms suffrutes-

cent, stiffly erect above. Leaves conspicuously disti-

chous, with blades folded, usually held outward at about

45°, or sometimes erect, 20-30 x 2-4 mm, tips hooded;

sheaths strongly overlapping, basal sheaths bladeless.

Inflorescence of 4-8 spikelets, 15-35 mm long, overtop-

ping leaves by at least 3 times its own length. Spikelets

4,5-6 mm long, to 2,5 mm across above glumes, oblong

to almost square in outline.

The folded leaf blades with hooded tips and the very

broad spikelets separate this subspecies from all other

taxa in the Setacea group. In spite of these morphologi-

cal differences, anatomical characters support the combi-

nation of subsp. rupestris and subsp. tricostata under the

same species (Ellis 1987).

The distribution of subsp. rupestris is shown in Figure

2. It is never common. It is most abundant and appears in

its typical form in the Riviersonderend Mountains of the

Caledon area. From there it extends northward and

eastward, over the Langeberg at the Clock Peaks, and

out along the dry Klein and Groot Swartberg, where the

plants are smaller. In both the Caledon area and in the

Klein Swartberg, where subsp. rupestris is sympatric

with subsp. tricostata, intermediates in leaf and spikelet

characters occur between the two subspecies.

The subspecies grows on mountain slopes among

rocks at altitudes of 910 to 1 970 m. Flowering occurs

from October to January.

A particularly robust specimen otherwise typical of

the subspecies (Van Breda 4436) was collected on

coastal sand in the Vanrhynsdorp region, well outside

the usual distribution of the subspecies, and in a different

habitat. Nevertheless, the anatomical characters closely

resemble those of other examples of subsp. rupestris

(Ellis 1987). There is some doubt about the correctness

of the label of this specimen and its locality has not been

plotted on Figure 2. A parallel example is known from

E. calycina, in a different species group, where an

extremely robust plant over 2 m tall was collected on

coastal sand near Lambert’s Bay (Gibbs Russell 5615).

Vouchers: Boucher 4201 (PRE, STE); Esterhuysen 21044 (BOL,

PRE); Pocock S72 (PRE); Thomson 2255 (PRE); Van Breda 4436

(PRE).

(b) subsp. tricostata (Stapf) Gibbs Russell in

Bothalia: 150(1984).

Ehrharta tricostata Stapf: 669 (1900); Chippind.: 35 (1955). Type:

Cape Province, French Hoek, 2 400 feet, Schlechter 9292 (K, holo.!;

PRE, iso.!).

KEY TO SUBSPECIES

la Plants delicate, herbaceous, less than 250 mm tall; inflorescence of 1-4 spikelets, barely overtopping leaves;

spikelets 4,5-5 mm long lc. E. rupestris subsp. dodii

lb Plants ± robust, herbaceous to suffrutescent, 200-450 mm tall; inflorescences of 4-9 spikelets, considerably

overtopping leaves; spikelets 4, 5-6, 3 mm long:

2a Leaf blades inrolled and appearing setaceous, erect, not distichous, or sometimes flat and ± spreading,

tips not hooded; spikelets (above glumes) to 2 mm across, linear to oblong

lb. E. rupestris subsp. tricostata

2b Leaf blades folded, somewhat thickened, distichous, usually at 45° to culm, tips hooded; spikelets (above

glumes) to 2,5 mm across, oblong to nearly square la. E. rupestris subsp. rupestris

Bothalia 17,1 (1987)

69

18° 20° 22° 24°

FIGURE 2. — Distribution of E. rupestris subsp. rupestris.

Plants erect, tufted, rarely trailing, 200-450 mm tall,

with creeping rhizomes. Culms suffrutescent (rarely her-

baceous). Leaves with baldes usually inrolled and

appearing setaceous, often reduced, erect, or sometimes

flat and spreading, not distichous, to 100 x 2,5 mm, tips

tapering, acute; sheaths not strongly overlapping, basal

leaves with blades. Inflorescence of 4-9 spikelets, 18-40

mm long, usually considerably overtopping leaves.

Spikelets 4, 6-6, 3 mm long, to 2 mm across above

glumes, oblong to linear.

In its size, erect habit, and relative abundance, this

subspecies is similar to E. setacea subsp. setacea but it

may be distinguished by the leaves at the base of the

culm. E. rupestris subsp. tricostata has blade-bearing

leaves nearly to the culm base, whereas E. setacea

subsp. setacea has scale-like, overlapping, bladeless

sheaths on the lower part of the culms.

The distribution of subsp. tricostata is shown in

Fugure 3. It is the most common and the most wide-

spread of all seven taxa in the Setacea group, extending

from the mountains of the Peninsula north to the Hex

River Mountains, and eastward to the Klein Swartberg

and along the Outeniqua and Tsitsikama coastal ranges

nearly to Humansdorp. The subspecies is not recorded

from the Langeberg. Plants from the Peninsula and Cale-

don are the most robust and erect, with reduced seta-

ceous leaf blades; plants from the easternmost end of the

range are small and fine, although with suffrutescent

culm bases; plants from the Hex River Mountains merge

into subsp. dodii and are decumbent, herbaceous and

have flat leaf blades. All these forms grade gradually

into each other. In addition, intermediates link this

subspecies with the other two, which might be

considered extreme forms. It is clear on anatomical

grounds (Ellis 1987) that the smaller form of subsp. tri-

costata from the eastern part of the range is distinct from

the small subsp. dodii, even though they overlap in size.

The subspecies grows in wet places on mountain

slopes and at the base of cliffs, at altitudes of 300-2 030

m. Flowering extends from October to February, with

most plants flowering in November and December.

Vouchers: Ellis 2547 (PRE); Esterhuysen 28594 (BOL, PRE, STE);

32813 (BOL, PRE, STE); 33057 (BOL, PRE, STE); Fourcade 3132

(BOL, PRE, STE).

18° 20° 22° 24°

FIGURE 3. — Distribution of E. rupestris subsp. tricostata.

(c) subsp. dodii (Stapf) Gibbs Russell in Bothalia:

150(1984).

Ehrharta dodii Stapf: 670 (1900); Chippind.: 35 (1955). Type: Cape

Province, rocks on Constantia Berg, Wolley Dod 1961 (K, holo. ! ;

BOL, iso.!; PRE, photo.!, fragment!).

Differs from subsp. tricostata mainly in size, habit

and in number of spikelets. Plants are less than 250 mm

tall, trailing or rarely erect, and rhizomes, culms, and

leaves are herbaceous and delicate, with the culms freely

branched and the leaf blades inrolled and held erect.

Inflorescence of 1-4 spikelets, 5-10 mm long, barely

overtopping leaves. Spikelets 4,5-5 mm long, to 2 mm

across above glumes.

This subspecies is difficult to distinguish except on

glume length from the two small, delicate subspecies of

E. setacea. In general, E. rupestris subsp. dodii is trail-

ing or erect, with erect rolled leaf blades, whereas E.

setacea subsp. uniflora is also trailing, but usually has

flat leaf blades, and E. setacea subsp. disticha is erect

with spreading, rolled or folded leaf blades. However, in

each taxon the habit is variable to some extent.

The distribution of subsp. dodii is shown in Figure 4.

It has the most restricted range of any subspecies of

E. rupestris, occurring only from Constantia Berg on the

Peninsua around to Kogelberg, and northward through

Hottentots Holland and Franschhoek to the Hex River

Mountains. It is connected in the eastern part of its range

to subsp. tricostata by intermediates of slightly larger

stature, upright habit and with more numerous spikelets.

These occur within the range of typical subsp. dodii as

well as to the east as far as Montagu. The lax Hex River

Mountain form of subsp. tricostata should perhaps be

counted among these intermediates.

The subspecies grows in wet places on mountainsides

among rocks and at the bases of cliffs, at altitudes of

660-1 660 m. Flowering occurs from November to early

January.

Vouchers: Boucher 2039 (PRE, STE); Ellis 2286 (PRE); Esterhuy-

sen 33065 (BOL, PRE); 33084 (BOL, JF, PRE); 34169 (BOL, PRE).'

70

Bothalia 17,1 (1987)

FIGURE 4. — Distribution of E. rupestris subsp. dodii.

2. Ehrharta setacea Nees, Florae Africae Austra-

lioris: 228 (1841). Stapf: 668 (1900); Chippind.: 37

(1955). Type: Cape Province, in monte tabulari, alt.

3 000 ft, Drige (PRE, isosyn. , fragment!).

Erect or trailing tufted perennials with long creeping

rhizomes, sometimes stoloniferous, of widely varying

size and habit, from delicate herbaceous plants 100 mm

tall to robust suffrutescent plants 600 mm tall. Culms

decumbent or prostrate, suffrutescent or herbaceous,

often leafless below. Leaves with blades erect, recurved

or spreading, inrolled, folded or flat; ligule a membrane

fringed with hairs; sheaths overlapping. Inflorescence an

erect raceme barely to considerably overtopping leaves.

Spikelets 1-15, at first erect but spreading at anthesis.

Glumes A (very rarely only A) as long as to longer than

lemmas, both acute. Florets as in E. rupestris.

(a) subsp. setacea. Gibbs Russell: 150 (1984).

Plant erect, 250-400 mm tall. Culms suffrutescent,

branched near base, bare of leaves below, sometimes

with much shortened inflated side branches. Leaves with

blades hard, smooth, usually tightly inrolled and appear-

ing setaceous, straight and erect or curved outward and

spreading from the middle, 50-80 (110) x 4 mm;

sheaths overlapping at upper intemodes, basal sheaths

without leaves, yellowish, truncate, slightly spreading.

Inflorescence of 5-15 spikelets, 25-60 mm long, usually

overtopping leaves by at least its own length. Spikelets

5, 5-6, 8 mm long, oblong; glumes % as long as, to

longer than spikelet, not gaping at maturity.

This subspecies is most similar in habit and size to E.

rupestris subsp. tricostata, but that taxon has blade-bear-

ing sheaths at the lowest culm nodes, while E. setacea

subsp. setacea has conspicuous pale, bladeless, square-

topped sheaths at the lower nodes of the culm. Leaf

anatomical features (Ellis 1987) suggest that the two taxa

might be united. However, each is linked by interme-

diates to different but parallel lines of morphological

variation. The anatomical similarity, which extends even

to species in unrelated genera, is an example of conver-

gence in leaf characters common in Fynbos. It is note-

worthy that the two taxa with this typical ‘Fynbos ana-

tomy’ are the most widespread subspecies in the Setacea

group.

The distribution of subsp. setacea is shown in Figure

5. It is the western element of E. setacea, and is the most

widespread of the four subspecies, occurring from the

Cape Peninsula north to Baines Kloof and east only as

far as the Hottentots Holland and Kogelberg Mountains.

In the Klein Rivier Mountains and at Betty’s Bay in the

Caledon District it is linked to subsp. uniflora and subsp.

disticha by intermediates of smaller size and semi-herba-

ceous habit.

The subspecies grows on mountainsides in damp,

peaty or marshy places and in seepage areas, either in

shale or in sand derived from Table Mountain

Sandstone, at altitudes of 660-1 515 m. Flowering

occurs from September to December and sporadically

until April.

Vouchers: Esterhuysen 3574 (BOL, NBG, PRE); 26501 (PRE);

28669 (BOL, PRE, STE); 33245 (BOL, PRE); Wolley Dod 3334

(BOL, PRE).

FIGURE 5. — Distribution of E. setacea subsp. setacea.

KEY TO SUBSPECIES

la Plants 250-400 mm tall, erect; inflorescences of 5-17 spikelets; glumes appressed to florets at maturity:

2a Leaf blades smooth, tightly rolled, appearing setaceous, rigid, erect, straight or curved slightly outward from

middle; spikelets 5, 5-6, 8 mm long 2a. E. setacea subsp. setacea

2b Leaf blades scabrous, flat and broad (to 6 mm across) at base, rolled near tip, bent 45° outward from culm at

sheath; spikelets (6,5) 7-8 mm long 2b. E. setacea subsp. scabra

lb Plants shorter than 250 mm, sprawling or erect; inflorescences of 1-4 spikelets; glumes gaping widely (more

than 45°) at maturity:

3a Plants trailing or sprawling, culms herbaceous, lowest nodes bearing leaves with blades; spikelets 4,5-6 mm

long; glumes usually slightly longer than lemmas 2c. E. setacea subsp. uniflora

3b Plants erect, cushion-forming, culms suffrutescent at base, lowest nodes usually leafless; spikelets 4-5 mm

long; glumes usually slightly shorter than lemmas 2d. E. setacea subsp. disticha

Bothalia 17,1 (1987)

71

(b) subsp. scabra (Stapf) Gibbs Russell in Bothalia:

151 (1984).

Ehrharta setacea Nees var. scabra Stapf: 669 (1900). Type: Cape,

in a mountain peak near Swellendam, Swellendam Div. , Burchell 7312

(K, holo.!; PRE, photo.!).

Plant erect, 250-600 mm tall, stoloniferous. Culms

suffrutescent, branched near base, sometimes with much

shortened side branches. Leaves with blades scabrous,

flat at base, rolled near tip, held 45° from culm at sheath,

to 30-1 10x6 mm; sheaths overlapping. Inflorescence

of 5-17 spikelets, 30-55 mm long, overtopping leaves

by at least twice its own length. Spikelets (6,5) 7-8 mm

long, oblong; glumes Vt-/ as long as lemmas, not gap-

ing at maturity.

This subspecies is easily distinguished from all other

African Ehrharta species by the scabrous leaf blades,

with a texture resembling that of a Melica or a Leersia.

Anatomically, subsp. scabra and subsp. setacea differ

considerably (Ellis 1987), raising the question whether

they should be treated as separate species. Despite their

anatomical differences and the absence of intermediates

between the two allopatric entities, they were kept as

subspecies because of the confused situation in the Cale-

don area, where intermediates occur between subsp.

setacea and subsp. scabra to both subsp. uniflora and

subsp. disticha.

The distribution of subsp. scabra is shown in Figure

6. It occurs only along the Langeberg from the Clock

Peaks above Swellendam to Garcia’s Pass, and thus

forms the eastern element of E. setacea. In the Caledon

area, it is linked through a number of intermediates to

subsp. disticha and subsp. uniflora.

The subspecies grows mostly in disturbed places on

mountainsides, such as path sides and burned clearings,

and also among rocks in seepage areas, at altitudes of

350—1 212 m. Flowering occurs from October to January

and sporadically to March.

Vouchers: Crook 2268 (PRE); Du Toit 2040 (PRE, STE); Haynes

868 (JF, PRE, STE); Taylor 4236 (PRE, STE); 7613 (PRE, STE).

FIGURE 6. — Distribution of E. setacea subsp. scabra.

(c) subsp. uniflora (Burch, ex Stapf) Gibbs Russell

in Bothalia: 151 (1984).

Ehrharta uniflora Burch, ex Stapf: 670 (1900); Chippind.: 37

(1955). Type: Cape, Cape Flats near Rondebosch, Burchell 182 (K,

holo.!; PRE, fragment!).

Plants sprawling or trailing, delicate but forming

dense masses. Culms fine, herbaceous, freely branched,

lowest nodes bearing leaves with blades. Leaves not dis-

tichous, blades soft, flat, straight, 50-80 x 2 mm;

sheaths barely or not overlapping. Inflorescence of 1-4

spikelets, 5-14 mm long, barely overtopping leaves.

Spikelets 4, 5-6, 5 mm long; glumes usually slightly

longer than lemmas but sometimes slightly shorter, gap-

ing more than 45° at maturity.

This subspecies can be distinguished from the other

two small taxa in the Setacea group by its long glumes

(E. rupestris subsp. dodii has short glumes) or trailing

habit (E. setacea subsp. disticha is erect).

The distribution of subsp. uniflora is shown in Figure

7. It is known only around False Bay, from the Cape

Peninsula to Pringle Bay. In the Caledon District inter-

mediates occur to subsp. setacea and subsp. scabra.

The subspecies grows in seepage areas, marshy places

and along watercourses, as well as at forest margins. It is

the only taxon in the Setacea group to occur at low alti-

tudes, from 10-500 m. Flowering occurs from Septem-

ber to December, and occasionally to March.

Vouchers: Adamson 1331 (BOL, PRE); Cleghorn 3168 (PRE, STE);

Esterhuysen 34002 (BOL, PRE); 34039 (BOL, NBG, PRE).

FIGURE 7. — Distribution of E. setacea subsp. uniflora.

(d) subsp. disticha Gibbs Russell in Bothalia: 151

(1984). Type: Cape, Caledon District, Maanschynkop,

Rocklands Park, Esterhuysen 31735 (PRE, holo. !; BOL,

iso.!).

Plant erect, cushion-forming, to 250 mm tall, differ-

ing from subsp. scabra mainly in size, number of spike-

lets and position of glumes. Culms branched near base,

suffrutescent, lowest nodes usually leafless. Leaves dis-

tichous, blades hard, rolled, recurved or erect, to 30 mm

long. Inflorescences of 1 or 2 spikelets, 5-10 mm long,

barely overtopping leaves. Spikelets 4-5 mm long;

glumes slightly shorter than lemmas, gaping more than

45° at maturity.

This small subspecies is separated from short-glumed

E. rupestris subsp. dodii by its long glumes, and from

lax E. setacea subsp. uniflora by its erect habit and

generally smaller spikelets with glumes slightly shorter

than the spikelet.

72

Bothalia 17,1 (1987)

The distribution of subsp. disticha is shown in Figure

8. It is known only from the Babylon’s Tower and Klein

River Mountains of the Caledon area. A number of inter-

mediate specimens show that in this area it intergrades

with subsp. setacea and subsp. scabra.

The subspecies grows in dry rocky places on mountain

slopes, at altitudes of 580-1 225 m. All other taxa in the

Setacea group grow in damp or wet places. The striking

habit difference between subsp. disticha and subsp. uni-

flora may result from adaptation of the former to water

stress in a dry habitat. Flowering occurs in October and

November.

Vouchers: Esterhuysen 31735 (BOL, PRE); 32319 (BOL, PRE);

33647 (BOL, PRE); 34755 (BOL, PRE); Williams 3086 (NBG, PRE).

FIGURE 8. — Distribution of E. setacea subsp. disticha.

CONCLUSION

The Setacea group is composed of very closely related

taxa, as shown by both morphological and anatomical

characters. These taxa are distinguished from other

Ehrharta species by the spikelet plan, with a short first

sterile lemma, and by a number of anatomical features.

These distinctive characteristics indicate that the Setacea

group must be carefully reviewed when generic limits in

the Ehrharteae are examined. The Setacea group has a

more restricted distribution than any other group in the

genus. All the subspecies except E. setacea subsp. sca-

bra occur in the Caledon degree grid (3419), and exam-

ples of all the intermediates also occur in this degree

grid. Thus, the Setacea group is concentrated in the

south-western phytogeographical centre of Weimarck

(1941) and the smaller south-western centre of Oliver et

al. (1983). Outside this pivotal area, each taxon has a

distinctive distribution. Only one subspecies occurs

throughout the range of the group, E. rupestris subsp.

tricostata. The other subspecies show vicariant distribu-

tion, limited to particular mountain ranges.

Although they are very similar, the seven infraspecific

taxa recognized here are each distinguished by macro-

morphological, spikelet and anatomical differences. The

occurrence of intermediates links them into two groups

with parallel morphological trends, as shown in Table 1.

These two groups are treated as separate species, with

the linked entities which they comprise treated as sub-

species. In each species, the most widespread and com-

monly collected subspecies is a robust plant with hard

setaceous leaves: E. rupestris subsp. tricostata and

E. setacea subsp. setacea. Each species has a robust

subspecies of more restricted eastern distribution differ-

entiated from the widespread one by broader spreading

leaf blades and larger spikelets: E. rupestris subsp.

rupestris and E. setacea subsp. scabra. Each species has

little delicate subspecies with reduced inflorescences and

small spikelets in the Peninsula and Caledon area:

E. rupestris subsp. dodii, E. setacea subsp. uniflora and

E. setacea subsp. disticha. Each of these reduced taxa

has a distinctive habitat. E. rupestris subsp. dodii grows

in wet places at high altitudes, E. setacea subsp. uniflora

grows in wet places at low altitudes, and E. setacea

subsp. disticha grows in dry places at high altitudes.

Thus, the factors in the Caledon area that favour small

size are apparently independent of moisture regime and

altitude.

AKNOWLEDGEMENTS

I thank: the Directors of herbaria that have lent speci-

mens, Bolus Herbarium (BOL), University of Cape

Town; Moss Herbarium (J), University of the Witwaters-

rand, Johannesburg; Wicht Herbarium (JF), Jonkershoek

Forestry Research Centre; The Herbarium, Royal Bo-

tanic Gardens, Kew (K); National Botanic Garden,

Kirstenbosch (NBG & SAM); Botanical Research Unit,

Stellenbosch (STE). Also R. P. Ellis and W. D. Clayton

for profitable discussions, G. Condy for the spikelet

drawing, and especially W. Roux for technical assis-

tance and preparation of diagrams.

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.

ELLIS, R. P. 1987. Leaf anatomy of the genus Ehrharta (Poaceae): the

Setacea group. Bothalia 17: 75-89.

TABLE 1. — Parallel trends in subspecies for the two species of the Setacea group. Arrows indicate the existence of specimens inter-

mediate between subspecies

Adaptation and distribution Plant large, robust; Plant small; Plant large, robust;

leaves setaceous, erect; leaves delicate: leaves broad, spreading;

widespread distribution 'False Bay’ distribution 'Eastern' distribution

4 >

E. rupestris subsp. tricostata < » subsp. dodii subsp. rupestris

E. setacea

subsp. setacea <-

-> subsp. uni flora f-

subsp. disticha <r

-> subsp. scabra

Bothalia 17,1 (1987)

73

GIBBS RUSSELL, G. E. 1984. Notes on species of Ehrharta with a

short first sterile lemma (Poaceae). Bothalia 15: 149-151 .

GIBBS RUSSELL, G. E. & ELLIS, R. P. 1987. Species groups in the

genus Ehrharta (Poaceae) of southern Africa. Bothalia 17: 5 1-65.

KUNTZE, C. E. O. 1891. Revisio generum plantarum . Felix, Leipzig.

NEES AB ESENBECK, C. G. 1841. Florae Africae Australioris .

Prausnitzianus, Cracow.

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.

STAPF, O. 1900. Gramineae. In W. T. Thiselton-Dyer, Flora capen-

sis 8: 310-750.

STEUDEL, E. G. 1853. Synopsis plantarum graminearum. Metzler,

Stuttgart.

TRINIUS, C. B. 1839. Phalarideae. Memoires de I'Acadimie impi-

riale des Sciences de St-Petersbourg, s6r. 6, 5: 12-26.

WEIMARCK, H. 1941. Phytogeographical groups, centres and inter-

vals within the Cape flora. Lunds Univertsitets Arsskrift 37, 5:

1-143.

SPECIMENS EXAMINED

Adamson 596 (2a) BOL; 616 (2a) PRE; 660 (2a) BOL; 1331 (2c) PRE,

BOL. Andreae 1283 ( lb) PRE.

Bews 15169 (2a) BOL. Bolus 7978 (lb/2a) BOL; 33866 (2a) PRE.

Bond 668 (lb) NBG. Boucher 2039 ( lc) K, PRE, STE; 2069 ( lb) PRE,

STE; 2642 (2a) STE; 2645 (lc) K, STE; 4201 (la) PRE, STE. Burchell

182 (2c) K, PRE; 7312 (2b) K, PRE; 7419 (la) K.

Cleghorn 3168 (2c) PRE, STE; 3174 (2a/2c) K, PRE, STE. Compton

15486 (2a) NBG; 17505 (2a) NBG. Crook 2268 (2b) K, PRE.

Drege 1659 (2a) PRE; 1661 (la) K, PRE; 19358 (la) SAM. Du Toil

2040 (2b) PRE, STE.

Ellis 1678 (2b) PRE; 1679 (2b) PRE; 2273 (2a) PRE; 2286 (lc) PRE;

2287 (lc) PRE; 2294 (2a) PRE; 2544 (2b) PRE; 2547 (lb) PRE; 4652

(2b) PRE; 4653 (2b) PRE; 4669 (2b/2c) PRE; 4670 (2b/j2c) PRE; 4679

(2a) PRE; 4685 (la) PRE; 4689 (2b) PRE; 4690 (lb) PRE; 4700 (lb)

PRE. Esterhuysen 1541 (lb) BOL; 3574 (2a) BOL, NBG, PRE; 6770

(lb) BOL, NBG, PRE, SAM; 9498 (2a) BOL; 10876 (lb) BOL; 10974

(lc) BOL; 21044 (la) BOL, PRE; 22354 (2a) BOL, PRE; 22676 (lc)

BOL, PRE; 24081 BOL, PRE; 26501 (2a) PRE; 26708 (la/lb) BOL,

PRE; 27368 (lb) BOL, PRE; 27558 (2b/2c) BOL, K, PRE; 28067 (2a)

BOL, PRE; 28109 (lb) BOL, K, NBG, PRE; 2«267 (lb) PRE; 28594

(lb) BOL, K, PRE, STE; 28619 (lb/lc) BOL, K, PRE; 28651a (lc)

BOL, K; 28669 (2a) BOL, K, PRE, STE; 28683 (2b/2c) BOL, K;

28712 (lb) BOL, K, PRE, STE; 28777 (2b) BOL, K, PRE; 29936

(2b/2c) BOL, K, PRE, STE; 30045a (2c) BOL; 31424 (2b/2c) BOL,

K; 31735 (2d) BOL, PRE; 32319 (2d) BOL, K, PRE; 32553 (2a) BOL;

32727 (lc) BOL, K, PRE; 32792 (lb) BOL, K, PRE; 32813 (lb) BOL,

K, PRE, STE; 33040 (2c) BOL, PRE; 33047 (lb) BOL, K, PRE;

33056 (2c) BOL, PRE, STE; 33057 (lb) BOL, K, PRE, STE; 33065

(lc) BOL, K, PRE; 33084 (lc) BOL, JF, K, PRE; 33150 (lb) BOL;

33245 (2a) BOL, PRE; 33299 (lb/lc) BOL; 33511 (lc) BOL, PRE;

33520 (lb) BOL, K; 33523 (lb) BOL, PRE; 33647 (2d) BOL, K, PRE;

33662 (2a/2c) BOL, PRE; 33724 (2c) BOL, PRE; 33733 (lc) BOL, K;

33762 (lb) BOL, K; 34002 (2c) BOL, PRE; 34039 (2c) BOL, NBG,

PRE; 34122 (lb) BOL, K, PRE; 34168 (lb/lc) BOL, PRE; 34169 (lc)

BOL, K, PRE; 34173 (lb) BOL, K, PRE; 34183 (lb) BOL, K; 34497

(lc) BOL, K; 34505 (lb) K, PRE; 34557 (lb) BOL, K; 34755 (2d)

BOL, K, PRE; 34778 (2a) BOL; 34834 (lb) BOL, PRE; 35300 (lb)

BOL, K, PRE; 35364 (la) BOL, K; 35539a (2d) BOL, K; 35571 (lb)

BOL; 35620 (lb) BOL; 35621 (la) BOL; 35733 (lb) BOL; 35812 (lc)

BOL; 35822 (2a) BOL; 35S35 (lb) BOL.

Forsyth 355 (2a) JF. Fourcade 3132 (lb) BOL, K, PRE, STE.

Galpin 164 (2a/2c) PRE, STE. Gibbs Russell 5634 (2b) BOL, K,

NBG, PRE; 5635 (2b) BOL, K, PRE; 5652 (2b/2c) BOL, K, NBG,

PRE; 5671 (2a) BOL, K, NBG, PRE; 5671a (lc) PRE; 5675 (lc) PRE;

5676 (la) BOL, K, NBG, PRE; 5678 (la) PRE; 5682 (2b) BOL, K,

NBG, PRE; 5696 (lb) BOL, K, NBG, PRE.

Haynes 868 (2b) JF, PRE, STE.

Kerfoot 5709 (lb) JF. Kruger 276 (lb) JF; 521 (2a) JF, PRE; 651 (lc)

JF; 1099 ( lb) JF, STE; 1515 ( lb) JF, STE; 1599 ( lc) JF.

Pappe53 (2a) PRE. PocockS72( la) PRE.

Schlechter 1825 (2b) BOL, K; 9292 (lb) BOL, K, PRE; 9869 (la/lb)

BOL, K, PRE. Stokoe 2537 (lb/lc) PRE; 7810 (2a) BOL; 8640 (la)

BOL; 22547 (lb) BOL; 67667 (2a) SAM; 67675 (la) SAM.

Taylor 3222 (2b) PRE, STE; 4236 (2b) PRE, STE; 7613 (2b) K, PRE,

STE; 7668 (2c) PRE. Thompson 1597 (lb) PRE, STE; 2255 (la) PRE.

Thorne 43440 (2b) SAM; 44556 (la) SAM; 45847 (la) SAM. Van

Breda 4436 (la) PRE. Van der Merwe 25-52 (2a) PRE; 933 (2a) PRE,

STE; 934 (2a) PRE, STE. Van Rensburg 215 (2a) PRE.

Williams 2887 (2d) NBG; 2963 (lb) K, NBG, PRE; 3015 ( lb) K, NBG

PRE; 3086 (2d) K, NBG, PRE; 3150 (2b/2d) K. Wolley Dod 1961 (lc)

BOL, K, PRE; 3334 (2a) BOL, K, PRE.

Zeyher 19359 (2a) SAM.

.

Bothalia 17,1:75-89(1987)

Leaf anatomy of the genus Ehrharta (Poaceae) in southern Africa:

the Setacea group

R. P. ELLIS*

Keywords: arm cells, Capensis floral kingdom, Ehrharta, E. rupestris, E. setacea, leaf anatomy, Poaceae, southern Africa, systematics

ABSTRACT

The leaf blade anatomy of the taxa of the Setacea group of species of the genus Ehrharta is described and illustrated.

This group includes E. rupestris Nees ex Trin. subsp. rupestris, subsp. tricostata (Stapf) Gibbs Russell and subsp. dodii

(Stapf) Gibbs Russell, as well as E. setacea Nees subsp. setacea, subsp. scabra (Stapf) Gibbs Russell, subsp. uniflora

(Burch, ex Stapf) Gibbs Russell and subsp. disticha Gibbs Russell. All these taxa share a very characteristic leaf anatomy

with inrolled or infolded leaves without keels and have adaxial ribs with interlocking prickles. The chlorenchyma is dense

and compact with inwardly projecting invaginations visible in all taxa except £. setacea subsp. setacea. In E. setacea

subsp. scabra typical arm cells are present. Abaxial costal and intercostal zones are not differentiated and stomata are

absent. The long cells are hexagonal or inflated with sinuous walls. Silica bodies are single or paired and rounded in shape.

Small hook-like prickles with short barbs are common. Microhairs with a short, truncated distal cell occur. This leaf

anatomical structure differs considerably from that of the other species groups recognized in African Ehrharta and the

Setacea group appears to be more distinct from the other groups than they are from each other.

UITTREKSEL

Die blaarskyfanatomie van die taksons van die Setacea-spesiegroep van die genus Ehrharta word beskryf en ge'illus-

treer. Hierdie groep sluit E. rupestris Nees ex Trin. subsp. rupestris, subsp. tricostata (Stapf) Gibbs Russell en subsp. dodii

(Stapf) Gibbs Russell asook£. setacea Nees subsp. setacea, subsp. scabra (Stapf) Gibbs Russell, subsp. uniflora (Burch,

ex Stapf) Gibbs Russell en subsp. disticha Gibbs Russell in. A1 hierdie taksons het ’n baie kenmerkende blaaranatomie

gemeen. Die blare is ingerol of ingevou, sonder ’n kiel en het adaksiale ribbe met ineensluitende dorings. Die chlorenchiem

is dig en kompak met instulpings van die selwande sigbaar in alle taksons behalwe £. setacea subsp. setacea. In £. setacea

subsp. scabra is tipiese vertakte selle teenwoordig. Die abaksiale kostale en interkostale sones is nie onderskeibaar nie en

huidmondjies is afwesig. Die langselle is seshoekkig of opgeblaas en met gekart£lde wande. Silikaliggaampies is enkel of

gepaard en het ’n ronde vorm terwyl die stekelhare met kort weerhake toegerus is. Mikrohare met ’n kort afgeknotte distale

sel kom voor. Hierdie anatomiese struktuur verskil aansienlik van die van al die ander spesiegroepe in Ehrharta in Afrika en

die Setacea-groep blyk ’n afsonderlike groep te wees wat meer van die ander groepe verskil as wat hulle van mekaar verskil.

CONTENTS

Introduction 75

Leaf anatomy of the taxa of the Setacea group

E. rupestris

subsp. rupestris 76

subsp. tricostata 78

subsp. dodii 80

E. setacea

subsp. setacea 81

subsp. scabra 83

subsp. uniflora 85

subsp. disticha 86

Discussion 86

Conclusions 89

Acknowledgements 89

References 89

INTRODUCTION

The African species of Ehrharta fall into two natural

groups: those with the first sterile lemma similar to the

second, and the fertile lemma differing from both; and

those with the first sterile lemma reduced and scale-like,

* Botanical Research Institute, Department of Agriculture and Water

Supply , Private Bag X 1 0 1 , Pretoria 000 1 .

and the fertile lemma similar to the second sterile lemma

(Gibbs Russell 1984). About 20 species belong to the

first group, but the second group, to which the Setacea

group belongs, contains only two species: E. rupestris

Nees ex Trin. and E. setacea Nees, together with seven

subspecific taxa. These two species, therefore, differ

considerably in spikelet structure from the other species

groups in the genus and, morphologically, the Setacea

group appears to be rather isolated from the rest of the

genus. This paper will examine whether the leaf anatomy

of the representatives of this group confirms this obser-

vation, and will discuss the affinities of this group by

reference to the anatomical evidence.

Chippindall (1955) recognized five species with a

short sterile lemma but Gibbs Russell (1984, 1987) has

combined these into only two species, each with several

subspecies. Within each species there are parallel trends

for reduced size and spikelet number, for rolled, folded

or flat, erect or spreading leaf blades and for suffrutes-

cence. This apparently reticulate pattern of relationships,

which generally applies to the genus Ehrharta, results in

parallel or convergent trends in vegetative parts occur-

ring in different taxa recognized on the basis of spikelet

differences. It is of interest to determine whether the leaf

anatomical variation parallels that exhibited by the spike-

lets and whether it substantiates the morphological clas-

sification based on relative glume length. This paper will

copiously illustrate, describe briefly and evalute the leaf

anatomy of each of the seven subspecies of the two spe-

cies of this group with reference to the systematic posi-

tion of these taxa and of the group as a whole relative to

the genus and the higher classification of the Poaceae.

76

Bothalia 17,1 (1987)

E. rupestris NeesexTrin. subsp. rupestris

Transverse section

The median vascular bundle is indistinguishable struc-

turally from the lateral first order bundles (Figure 1.1),

as is the case throughout this group. Smaller third order

bundles occur on either side of the median bundle and

near the margin (Figure 1.1; 1.3) and, perhaps, second

order bundles should be distinguishea in this taxon.

However, if these irregular ana ill-defined additional

bundles are excluded then only two smaller bundles sep-

arate consecutive first order bundles, as is the case in all

other taxa in this group. The presence of these additional

bundles is not considered to be taxonomically significant

and probably only represents individual variation in the

specimen studied.

Rounded adaxial ribs and narrow furrows are present

(Figure 1.2; 1.3), the furrows having interlocking adax-

ial prickles (Figure 2.1). Similar pnckles also occur on

subsp. tricostata but are absent on subsp. dodii.

The chlorenchyma tissue is compact, consisting of

small round cells with dense chloroplasts (Figure 1.2).

These cells do not have straight, isodiametric sides but

do have slight, but clearly discernible wall invaginations

(Figure 1.2; 1.3). According to the generally accepted

definition of arm cells, these must be considered as such

and, in fact, they do closely resemble the arm cells of the

Oryzeae and some of the other bambusoid grasses. These

invaginations are better developed in subsp. rupestris

than in either subsp. tricostata or subsp. dodii and a

definite reduction series is evident in these three subspe-

cies.

Abaxial epidermis

The epidermis conforms very closely to the type diag-

nostic for this group, with the costal and intercostal

zones being structurally undifferentiated and with the

long cells being very short and inflated to hexagonal with

sinuous walls (Figure 1.4; 1.5). Abaxial stomata were

not detected with the light microscope, neither in

FIGURE 1. — Leaf anatomy of Ehrharta rupestris subsp. rupestris. 1-5, Ellis 4685. 1-3, leaf transverse sections; 1, infolded leaf blade with

median vascular bundle (mr) only, scale = 20 /am; 2, compact mesophyll with chlorenchyma cells with slight wall invaginations, adaxial

stomata (s) only and abaxial hooks (p) with short barbs, scale = 10 /am; 3, interference contrast showing long barbs of adaxial prickles (p)

and short barbs of abaxial hooks (p), scale = 10 /xm. 4—5, abaxial epidermis: 4, costal and intercostal zones of similar cells, note

microhair (m), scale = 10 /xm; 5, costal and intercostal long cells separated by hooks, prickles (p) or silica bodies, scale = 10 /xm. 6,

abaxial epidermis of Van Breda 4436 showing costal and intercostal hooks (p) and microhairs (m), scale = 10 /xm.

Bothalia 17,1 (1987)

77

FIGURE 2.— Epidermal ultrastructure of Ehrharta rupestris subsp. rupestris. 1-4, Ellis 4685. 1, adaxial epidermis showing ribs and furrows

with interlocking prickles, x 60. 2-4, abaxial epidermis: 2, epidermal zonation not evident and prickle hairs are widespread, x 60; 3,

detail of prickles, stomata and papilla-like silica bodies, x 340; 4, microhair and silica bodies, x 1000.

transverse sections (Figure 1.2; 1.3) nor in epidermal

preparations (Figure 1.4— 1.6). However, the SEM re-

vealed a few isolated stomata deeply sunken between the

inflated long cells and obscured by hooks (Figure 2.3).

Silica bodies are structurally identical throughout the

epidermis and appear as single, rounded bodies located

between the long cells (Figures 1.4; 1.5). Under the

SEM these are clearly seen to be raised, rounded struc-

tures covered with epicuticular wax (Figure 2.3; 2.4). As

such they are indistinguishable from papillae and do not

resemble the usual silica bodies with their concave faces

devoid of wax platelets. Their refractive nature is visible

with the light microscope (Figure 1.5) and they can also

be seen to be raised (Figure 1.6). They are undoubtedly

silica bodies but they are highly unusual and their exact

nature and structure remains somewhat uncertain. Com-

parable structures have not been described elsewhere in

the literature and they deserve further study. Similar

structures also occur in E. setacea and they appear to be

a unique characteristic of this group of Ehrharta species.

The entire abaxial epidermis is densely covered in

shortly barbed hooks occupying the same locations be-

tween the long cells as the silica bodies (Figure 1.6).

These hooks can occupy up to 50% of the available loca-

lities (Figure 1.6) but may be fewer in number (Figure

1.4; 1.5). The hooks with their reduced barbs may be

difficult to distinguish from the silica bodies with the

light microscope but are definitely distinct structures as

seen under the SEM (Figure 2.2; 2.3).

The microhairs of subsp. rupestris are also of interest

in that they are very short, with a blunt distal cell which

does not taper to a point (Figure 2.3; 2.4). They are

common throughout the epidermis (Figure 1.4; 1.6) and

are an important distinguishing feature between this

group and all the other groups of Ehrharta.

Specimens examined

CAPE. — 3118 (Van Rhvnsdorp): Vredendal Distr., Koekenap

(-CB), Van Breda 4436. 3319 (Worcester): Riviersonderend Mts,

Jonas Kop (-DC), Ellis 4685.

Discussion

A small and inadequate sample was studied anatomi-

cally and, consequently, little information on variation in

this subspecies can be deduced. However, the specimen

from the Van Rhynsdorp District in Succulent Karoo

comes from an atypical habitat and is particularly robust

(Gibbs Russell 1984) and yet closely resembles the spec-

imen from the Mountain Fynbos in leaf anatomy. It,

therefore appears as if the leaf anatomy of this taxon is

stable and consistent despite differing environmental

conditions.

E. rupestris subsp. rupestris is obviously very closely

related to subsp. tricostata and the two can hardly be

separated anatomically and, with a larger study sample,

can be expected to grade into one another. The cross-

sectional anatomy tends to support the diagnostic differ-

ence of folded (Figure 1.1) versus rolled (Figure 3.1;

3.4) leaf blades (Gibbs Russell 1984, 1987) even though

subsp. tricostata is not rolled in the same way as subsp.

78

Bothalia 17,1 (1987)

FIGURE 3. — Transectional leaf anatomy of Ehrharta rupeslris subsp. tricostata. 1-3, Ellis 4700: 1 , inrolled leaf outline with median vascular

bundle (mr), scale = 20 /xm; 2, compact mesophyll with cells densely packed with chloroplasts, scale = 10 /xm; 3, interference contrast

showing very slight inward projections of walls of chlorenchyma cells and long barbs of adaxial prickles (p), scale = 5 gm. 4-6, Ellis

4690 : 4, infolded leaf outline, scale = 20 /xm; 5, arrangement of first and third order vascular bundles, scale = 10 /xm; 6, detail of

chlorenchyma with some cells showing slight indentations and adaxially located stomata (s) only, scale = 5 gm.

dodii (Figure 6. 1; 6.3). This morphological difference is,

therefore, not very convincing when it is analysed struc-

turally. Further anatomical studies are required to con-

firm whether these two taxa are anatomically distinct; all

indications are that only infraspecific separation is justi-

fied. This study, therefore, supports the decision of

Gibbs Russell (1984, 1987) to reclassify E. tricostata as

a subspecies of E. rupestris. It must also be noted that

these two subspecies show greater similarities with each

other than either does to subsp. dodii.

E. rupestris Nees ex Trin. subsp. tricostata (Stapf)

Gibbs Russell

Transverse section

No keels or midribs are present (Figure 3.1; 3.4) and

successive first order vascular bundles are separated by

two third order bundles (Figure 3.2; 3.5). The outline of

the lamina is very similar to that of subsp. rupestris and

these two taxa differ only in subsp. rupestris having

considerably wider leaf blades which are infolded but

with the arms of the lamina curved outwards. In subsp.

tricostata the leaf blades are narrower and setaceous with

no angle being present at the median bundle (Figure 3.1;

3.4) and these leaves can be considered as being inrolled

although the margins do not overlap as in subsp. dodii.

This distinction between subsp. tricostata and subsp.

rupestris is only minor and intermediates can be ex-

pected when a larger sample is studied. Both these

subspecies also possess a ribbed adaxial surface with

interlocking prickles overlying the furrows (Figure 3.3).

The mesophyll is compact, of small regular cells. The

cells are rounded and very tightly packed (Figure 3.3)

and closer inspection reveals very slight inward projec-

tions on only a few cell walls (Figure 3.3; 3.6). These

cells resemble arm cells but cannot be considered to be

typical and may result from reduction of the type of

invagination described in subsp. rupestris.

Abaxial epidermis

The epidermal structure is virtually identical to that of

subsp. rupestris as illustrated in Figure 4. 1-4.4. The

prickle hooks are not always very common and may lack

barbs (Figure 4.4) but the solitary, round silica bodies

are clearly visible (Figure 4.2). Ultrastructurally these

bodies are also seen to be raised and papilla-like (Figure

5.3) and not concave as typical silica bodies usually are.

The microhairs have a short, blunt distal cell (Figure 5.4)

Bothalia 17, 1 ( 1987)

79

FIGURE 4. — Abaxial epidermis of Ehrharta rupestris subsp. tricostata. 1-2, Ellis 4700: 1, costal and intercostal Files of similar composition,

scale = 10 fim; 2, interference contrast showing silica cells or hooks between all long cells, scale = 10 gm. 3—4, Ellis 4690: costal and

intercostal long cells diamond shaped, scale = 10 /am; 4, intercostal long cells with thickened, pitted walls, note microhair (m), scale =

10 /xm.

FIGURE 5. — Abaxial epidermis of Ehrharta rupestris subsp. tricostata. 1, Ellis 4690 showing absence of costal and intercostal zonation, x

60. 2-4, Ellis 4700: 2, prickles common and short long cells separated by raised silica bodies, x 200; 3, prickles with short barbs,

microhairs and long cells, x 300; 4, detail of microhair and cuticular striations on long cells, x 1000.

80

Bothalia 17,1 (1987)

and are unusual in this regard in the genus. The epicuti-

cular wax layer in subsp. tricostata is very thick with

distinct horizontal striations (Figure 5.3; 5.4) and not of

fine platelets as in subsp. rupestris (Figure 2.3; 2.4).

Specimens examined

CAPE. — 3321 (Ladismith): Riversdale Distr. , Langeberg, Garcia’s

Pass (-CC), Ellis 4690. 3322 (Oudtshoom): Outeniqua Mts, Robin-

son’s Pass (-CC), Ellis 2547, 4700.

Discussion

Anatomically subsp. tricostata is very similar indeed

to subsp. rupestris and these two taxa cannot be satisfac-

torily distinguished on anatomical criteria. This is sup-

portive of their separation at only the subspecies level

(Gibbs Russell 1984) and the leaf anatomy confirms the

extremely close relationship of these two taxa. The re-

cognition of two separate species is not corroborated by

the anatomical evidence.

E. rupestrisNees ex Trin. subsp. dodii (Stapf) Gibbs

Russell

Transverse section

No keel is present (Figure 6.1; 6.3) and the adaxial

ribs and furrows are not well developed and interlocking

prickles are absent (Figure 6.2). Of particular interest is

the convolute inrolling of the leaf blade (Figure 6.1;

6.3). This type of inrolling does not occur in either

subsp. rupestris or subsp. tricostata and appears to be a

significant anatomical difference between these two

subspecies and subsp. dodii.

The mesophyll tissue is compact, of small rounded

and regular cells (Figure 6.2). These cells cannot be

regarded as being typical arm cells as they possess only

very slight invaginations.

Abaxial epidermis

The costal and intercostal zones are structurally undif-

ferentiated although their relative positions can be deter-

mined due to the staining of the sclerenchyma girders

underlying the costal zones (Figure 6.4). The long cells

are considerably longer and less inflated than in either of

the other two subspecies of E. rupestris. All three

subspecies do have sinuous long cell walls and stomata

are either very rare (subsp. rupestris ) or absent from the

abaxial surface. The silica bodies are also rounded but

they are closely associated with a cork cell. Although

these structures were not examined ultrastructurally,

there can be little doubt that they are typical graminoid

silica bodies and not the unusual type described in subsp.

tricostata and subsp. rupestris. Prickles with prominent

oval bases and very short barbs are common (Figure

6.4). Small microhairs, typical of those of the Setacea

group, are also present (Figure 6.4).

Specimens examined

CAPE. — 3418 (Simonstown): Hottentot Hollands Mts, Sir Lowry’s

Pass (-BB), Ellis 2286, 2287; Kogelberg State Forest, Platberg (-BD),

Boucher 2039.

Discussion

In its leaf anatomy E. rupestris subsp. dodii differs

more from subsp. rupestris and subsp. tricostata than

FIGURE 6. — Leaf anatomy of Ehrharta rupestris subsp. dodii. 1-3, leaf in transverse section. 1-2, Ellis 2286: 1 , inrolled outline with simple

median vascular bundle (mr), scale = 10 gm; 2, compact mesophyll of rounded cells with slight wall indentations, scale = 5 p m. 3, Ellis

2287, showing inrolled outline, few adaxial prickles and simple median bundle (mr), scale = 10 /am. 4, abaxial epidermis, Ellis 2287,

note absence of stomata, microhairs (m), costal prickles (p) and intercostal hooks, scale = 5 pm.

Bothalia 17,1 (1987)

81

these two subspecies differ from one another. The find-

ings of this study, based on only a very small sample,

appear to indicate that subsp. dodii is sufficiently distinct

from E. rupestris to warrant separate specific status.

Additional anatomical material may reveal anatomical

intermediates with subsp. tricostata but this appears

rather unlikely in view of the entirely different nature of

the involution of the blades of these two taxa. The con-

volute type of inrolling in subsp. dodii may indicate a

different type of bud vernation to that of subsp. trico-

stata where inrolling is of the involute type. If this

proves to be true then the case for separation at the spec-

ies level will be strengthened.

Although subsp. dodii differs somewhat from subsp.

rupestris and subsp. tricostata in leaf anatomy, their

anatomy is still typical of the Setacea group and different

from that of all the others of Ehrharta (Gibbs Russell &

Ellis 1987). Subsp. dodii, therefore, unquestionably be-

longs to this very distinct group.

The less acicular type of leaf transection and the

elongation of the epidermal long cells seems to be paral-

lelled in E. setacea in subsp. disticha and subsp. uni-

flora. This trend may be related to the milder climate of

the Cape Peninsula and Caledon District to which these

three taxa are restricted. These three subspecies do not

occur at higher altitudes in the mountain ranges further

inland as do E. rupestris subsp. rupestris and subsp.

tricostata and E. setacea subsp. setacea and subsp. sca-

bra. This trend, associated with a more mesophytic type

of anatomy and elongated epidermal cells with costal and

intercostal zones and with abaxial stomata, may repre-

sent the convergent trends so evident in this group.

Alternatively this pattern may reflect the reticulate nature

of relationships which also characterizes this group. It is

therefore an academic decision whether subsp. dodii

should be recognized as a separate species or not. What-

ever basis is used to define species in this group, other

characters will not necessarily follow the same pattern.

The decision is also dependant on a similar decision

being made for subsp. disticha and subsp. uniflora in E.

setacea.

E. setacea Nees subsp. setacea

Transverse section

This taxon has the leaf outline which typifies this

group: the blades are setaceous, without midribs or keels

(Figure 7.1; 7.3) and with well developed adaxial ribs

and furrows (Figure 7.2; 7.4). Subsp. setacea is notable

in having acicular leaf blades of the permanently in-

folded type with the abaxial epidermal cells being very

large and regular in size as seen in transection (Figure

7.2; 7.4).

The chlorenchyma is exceptionally compact and con-

sists of straight-walled, isodiametric cells (Figure 7.2;

7.4). The chloroplasts are dense, peripherally arranged

and with a central vacuole. No invaginations are visible.

This chlorenchyma is identical in appearance to the type

so common in the Danthonieae in genera such as Merx-

muellera, particularly M. disticha (Schrad.) Nees (Ellis

1980), Pseudopentameris (Ellis 1985a) and some species

of Pentameris (Ellis 1985b, c).

Abaxial epidermis

Costal and intercostal zones are not differentiated

(Figure 7.5; 7.7), the long cells are short and fusiform

(Figure 7.6; 7.8) and separated by a single, raised silica

body (Figure 12.1). Abaxial stomata are absent as are

hooks and prickles. Microhairs are infrequent (Figure

7.8) or absent (Figure 7.6) but when present are short

with rounded distal cells (Figure 12.2).

Specimens examined

CAPE. — 3418 (Simonstown): Hottentots Holland Mts, Sugarloaf

Peak (-BB), Ellis 2273', Sir Lowry’s Pass (-BB), Ellis 2294; Landrost-

kop, Somerset Sneeuwkop (-BB), Ellis 4679.

Discussion

The leaf anatomy conforms closely to that typical of

the Setacea group, and subsp. setacea resembles E. ru-

pestris subsp. rupestris and subsp. tricostata very clo-

sely in leaf anatomy. It differs mainly in the absence of

abaxial prickles. Other slight differences, such as the

chlorenchyma cell invaginations and the epidermal cells

in tranverse section which are regular in size, due to the

absence of prickle hair bases, are only differences of

degree. Anatomically, therefore, subsp. setacea clearly

belongs to the Setacea group. Whether it should be sepa-

rated from E. rupestris is debatable according to the

anatomical evidence and E. setacea subsp. setacea has

more in common with E. rupestris subsp. tricostata than

it has with any of the other subspecies of E. setacea. The

leaf anatomy appears to indicate that these two taxa are

conspecific and the relative glume length, which is used

to separate E. setacea from E. rupestris (Gibbs Russell

1984), results in an apparently artificial anatomical

grouping. This observation is particularly relevant when

the anatomy of subsp. setacea is compared with that of

subsp. scabra, as these taxa exhibit significant anatomi-

cal differences which would justify at least specific sepa-

ration in other grass groups.

Once again the classification of taxa within the Seta-

cea group presents problems with contradictory anatomi-

cal and morphological evidence. A possible explanation

for this situation lies in different rates of evolution in

different organs combined with a reticulate evolutionary

pattern. Present knowledge indicates that, whatever

characters are used to distinguish taxa in this group,

heterogeneity of other character sets will result. This

makes a generally acceptable classification almost im-

possible to achieve.

The anatomical resemblance between subsp. setacea

(and E. rupestris subsp. tricostata as well) and many of

the winter rainfall danthonoid grasses deserves further

mention. Both in section and epidermis subsp. setacea

closely resembles species of Pseudopentameris (Ellis

1985a), and Pentameris (Ellis 1985b, c) in particular.

These similarities may only be coincidental but it is more

probable that they result from convergent evolution.

Convergent and parallel patterns appear to be common in

Cape fynbos grass species. Most species of the mountain

regions, where nutrients are severely limiting, have

evolved a structurally complex, long-lived leaf, in con-

trast to the lowland species in which the leaves are decid-

uous (or the plants annual) with a much simpler internal

structure, with thinner cell walls, less sclerenchyma and

larger air spaces. This general pattern cuts across phylo-

genetic lines at the generic level and does not appear to

be taxonomically significant. What is of importance,

however, is that here taxa from different subfamilies

have responded identically to similar environmental con-

ditions.

82

Bothalia 17,1 (1987)

FIGURE 7. — Leaf anatomy of Ehrharta setacea subsp. setacea. 1-2, Ellis 4679: 1 , leaf outline showing inrolling and undifferentiated midrib,

scale = 20 /xm; 2, anatomical detail with compact chlorenchyma and adaxial stomata (s), scale = 5 /cm. 3—4, Ellis 22 73: 3, inrolled

outline, scale = 20 /xm; 4, isodiametric chlorenchyma cells without wall invaginations, scale = 5 /xm. 5-6, Ellis 2294, abaxial

epidermis: 5, costal and intercostal zone cell structure similar, scale = 10 /xm; 6, hexagonal long cells separated by cork-silica cell pairs,

scale = 5 /xm. 7, Ellis 2273, costal and intercostal zones not clearly differentiated, scale = 10 /xm. 8, Ellis 4679, note small, indistinct

microhairs (m), scale = 10 /xm.

Bothalia 17,1 (1987)

83

E. setacea Nees subsp. scabra (Stapf) Gibbs Russell

Transverse section

This subspecies has a wide, expanded type of lamina

(Figure 8.1; 8.3) which is not of the setaceous type of

subsp. setacea. In all specimens, even those in which the

blade is almost flat, there are indications of convolute

infolding (Figures 8.1; 8.3). This is reminiscent of the

condition in E. rupestris subsp. dodii. Slight adaxial ribs

and wide, open furrows (Figures 8.2; 8.4; 8.6) resemble

the subsp. dodii condition more than subsp. setacea.

The single most significant anatomical feature of

subsp. scabra is the possession of definite arm cells in

the mesophyll (Figures 8.2; 8.4; 8.5; 8.6). These are

typical arm cells with distinct invaginations of the cell

walls visible in all chlorenchyma cells (Figures 8.2; 8.4)

and the chloroplast pattern is also indented, indicating

the presence of these inward projections of the cell walls

(Figures 8.5; 8.6). Typical arm cells are, however,

sometimes lacking in subsp. scabra ( Ellis 4653). Arm

cells do not occur in subsp. setacea but indications of

their existence are evident in all other taxa of the Setacea

group.

Abaxial epidermis

The structure of the epidermis is very similar to that of

E. rupestris. Absence of zonation, short fusiform long

cells, no stomata, rounded silica bodies and hook-like

prickles (Figure 9. 1-9.4) occur in subsp. scabra and

characterize the taxa of the Setacea group. Ultrastructu-

rally this similarity is confirmed by the raised papilla-

like silica bodies (Figure 12.3) and short microhairs with

rounded distal cells (Figure 12.4). This epidermal struc-

ture differs somewhat from that of subsp. setacea and

subsp. uniflora but is virtually identical to that of E.

rupestris subsp. rupestris and subsp. tricostata.

Specimens examined

CAPE. — 3320 (Montagu): Barrydale Distr. , Tradouw’s Pass (-DC),

Ellis 1678, 1679, 4652, 4653 (varies anatomically). 3321 (Ladismith):

Riversdale Distr., Langeberge, Garcia’s Pass (-CC), Ellis 2544, 4689.

Discussion

Arm cells are considered to be a very important cha-

racter in the systematics of the Poaceae and the presence

of these cells in subsp. scabra may be taxonomically

meaningful. Arm cells are a diagnostic characteristic of

FIGURE 8. — Cross-sectional leaf anatomy of Ehrharta setacea subsp. scabra. 1-2, Ellis 4689 : 1 , broad, expanded outline, scale = 20 /xm; 2,

anatomical detail clearly showing arm cells (a), scale = 5 /xm. 3-4, Ellis 2544: 3, inrolled but broad leaf with median vascular bundle

(mr) only, scale = 20 /xm; 4, arm cells (a) with distinct cell wall invaginations, scale = 5 /xm. 5, Ellis 1679, leaf margin showing arm

cells and abaxial prickles (p), scale = 10 /xm. 6, Ellis 1678, arm cells (a) of chlorenchyma densely filled with chloroplasts, scale = 10

84

Bothalia 17,1 (1987)

FIGURE 9. — Abaxial epidermal structure of Ehrharta setacea subsp. scabra. 1-2, Ellis 4689 : 1 , cell arrangement, scale = 10 /im; 2, costal

and intercostal zones only slightly differentiated, scale = 10 /zm. 3-4, Ellis 4652: 3, cell pattern consistent throughout abaxial epidermis,

scale = 10 gm; 4, microhairs (m) and shortly barbed prickles (p), scale = 5 gm.

the Bambusoideae (including Oryzeae) and, as such, are

a feature of high taxonomic value. However, arm cells

are known sporadically from non-bambusoid groups.

Examples are Phragmites and Thysanolaena of the

Arundinoideae and Sclerodactylon and the Triodeae of

the Chloridoideae (Watson et al. 1985). This occurrence

of arm cells in these apparently unrelated groups is im-

possible to explain but indicates that the presence of arm

cells in the Setacea group of Ehrharta does not necessar-

ily imply bambusoid relationships for the genus. How-

ever, their presence can undoubtedly be used to substan-

tiate the arguments of those workers (Renvoize 1986) in

favour of bambusoid affinities for Ehrharta. However,

the occurrence of fully developed arm cells in only one

taxon of the genus Ehrharta urges caution in attaching

too much importance to their presence.

A certain degree of anatomical variation was observed

in this subspecies. This variation applies particularly to

the presence of stomatal bands in the abaxial epidermis

(Figure 10.3) with an associated development of clear

structural differentiation between the costal and intercos-

tal zones. The specimen displaying this type of structure

(Ellis 4653) was collected growing together with Ellis

4652, which has the anatomy typical of subsp. scabra,

whereas Ellis 4653 is a much more robust plant with an

FIGURE 10. — Anatomical variation in Ehrharta setacea subsp. scabra, Ellis 4653. 1-2, leaf sections: 1 , margin of open, expanded leaf, scale

= 20 n- m; 2, mesophyll not compact and not comprised of distinct arm cells, note abaxial stomata (s), scale = 10 /xm. 3, abaxial

epidermis showing stomatal bands (s), microhairs (m) and clear cellular differentiation between costal and intercostal zones, scale = 5

Bothalia 17,1 (1987)

85

extremely dense underground rhizome system. The ana-

tomy of this specimen is illustrated in Figure 10 and it

can be seen to differ from typical subsp. scabra in the

presence of abaxial stomata, the epidermal zonation and

by having elongated intercostal long cells (Figure 10.3)

and by the absence of typical arm cells, with the invagi-

nations being poorly developed (Figure 10.2). The mi-

crohairs are also more elongated (Figure 12.6) and there

are fewer prickles (Figure 12.5). The anatomy of this

specimen is, therefore, intermediate between that of

subsp. scabra and subsp. uniflora which has abaxial

stomata, no prickles and elongated, tapering microhairs

(Figure 12.7; 12.8). A few isolated stomata were also

observed on the epidermis of Ellis 4689, a rather robust

specimen with wide leaves (Figure 8.1). However, in

this specimen, the epidermal zones are not structurally

differentiated (Figure 9.1; 9.2) and the arm cells are

clearly developed (Figure 8.2). It is possible that a whole

range of intermediates will be found between E. setacea

subsp. scabra and subsp. uniflora and disticha. Unfortu-

nately the latter two subspecies are poorly represented in

this study and it was not possible to determine the exact

pattern of variation between these taxa.

E. setacea Nees subsp. uniflora (Burch, ex Stapf)

Gibbs Russell

Transverse section

This subspecies has a narrow but flat and expanded

leaf outline (Figure 11.1) with poorly developed ribs and

furrows (Figure 11.3). This is not the setaceous type of

leaf outline as in subsp. setacea, and abaxial stomata are

clearly visible even in transection (Figure 11.3; 11.4).

There is also no indication of convolute inrolling as in

subsp. scabra.

The mesophyll of subsp. uniflora is also not as com-

pact as in all the other taxa of the Setacea group (Figure

11.2; 1 1 .4), a difference which appears to be associated

with the development of an air-space system linked to

the abaxial stomata. The mesophyll cells themselves

cannot be considered to be classical arm cells (Figure

11.2; 11 .4) although very slight invaginations are visible

in some of the chlorenchyma cell walls.

Abaxial epidermis

The presence of a single File of stomata on either side

of each costal zone (Figure 11.5; 11.6) distinguishes the

FIGURE 11. — Leaf anatomy of Ehrharta setacea subsp. uniflora. 1-2, Ellis 4670, transections: 1, open, expanded outline with median

vascular bundle (mr) only, scale = 20 /xm; 2, anatomical detail showing chlorenchyma which is not composed of typical arm cells, scale

= 5 /xm. 3-4, Ellis 4669, transections: 3, abaxial stomata (s) common, scale = 10 /xm; 4, interference contrast showing only slight wall

indentations and abaxial stomatal aperture (s), scale = 5 /xm. 5, abaxial epidermis, Ellis 4670, with single File of stomata (s) on either

side of each costal zone, scale = 10 /xm. 6, abaxial epidermis, Ellis 4669, showing stomata (s) and clearly differentiated costal zones,

scale = 10 /xm.

86

Bothalia 17,1 (1987)

epidermis of this subspecies from all the other taxa of the

Setacea group (except subsp. disticha). There is also a

definite tendency for the costal zones to be structurally

different from the intercostal zones (Figure 11.6) with

the silica bodies being different. The intercostal long

cells are not separated by raised silica bodies as in the

other subspecies of E. setacea (Figure 12.1; 12.3; 12.5;

12.7). In this respect subsp. uniflora differs from all

other members of the Setacea group. Other differences

are the tapering microhairs (Figure 12.8) and the pre-

sence of epicuticular wax (Figure 12.7).

Specimens examined

CAPE. — 3418 (Simonstown): Cape Peninsula, Chapman’s Peak

(-AB), Esterhuysen 34039. 3419 (Caledon): Olifantsberg, Hermanus

(-AD), Ellis 4669, 4670, Esterhuysen 3 1039.

Discussion

The leaf anatomy of subsp. uniflora appears to be

intermediate between that of the Setacea group and some

of the other groups of the genus Ehrharta. This conten-

tion is supported by the epidermal zonation with abaxial

stomata, the presence of epicuticular wax, the absence of

raised silica bodies, the tapering microhairs and the less

compact mesophyll tissue. These features are all charac-

teristic of the Setacea group and subsp. uniflora differs

from the other taxa in this group in all these attributes

which tend toward the condition found in several of the

other infrageneric groupings in Ehrharta.

The leaf anatomy of subsp. uniflora is very similar

indeed to that of Ellis 4653, an atypical subsp. scabra

specimen (Figure 10). Ellis 4653 is a much more robust

plant with wider leaf blades than any subsp. uniflora

specimen. This is not reflected in the leaf anatomy, how-

ever, and leaf thickness and epidermal cell size are com-

parable. Only the width of the lamina is different. It

appears that the loss of the setaceous, acicular type of

leaf results in a very similar anatomy, irrespective of

which subspecies this occurs in. The development of

abaxial stomata appears to be the most significant change

and it is associated with several other anatomical features

such as the zonation of the epidermis and the presence of

an extensive air-space system accompanied by less com-

pact mesophyll. The loss of the Setacea type of silica

body and the possession of tapering microhairs is not

explained by the development of abaxial stomata, how-

ever, and they appear to be additional differences not

linked to the presence of these stomata.

Subsp. uniflora grows at very low altitudes, from

10-500m (Gibbs Russell 1984) and this is a possible

reason for the development of these abaxial stomata.

Temperatures are less extreme at these lower altitudes

with an equable maritime climate and the nutritional sta-

tus of the soil is considerably better than the leached soils

at higher altitudes. More material of subsp. uniflora is

required to gain a better understanding of the inter-

relationships of these taxa.

E. setacea Nees subsp. disticha Gibbs Russell

Transverse section

No fresh material of this taxon was available for ana-

tomical study and, consequently, the transectional ana-

tomy was not examined.

Abaxial epidermis

The epidermal structure is very similar to that de-

scribed for subsp. uniflora, with stomata! files and

clearly differentiated costal and intercostal zones (Fig-

ures 13.1; 13.2). Intercostal hooks are absent and the

intercostal silica bodies appear to be of the raised type

typical of the Setacea group but they are associated with

cork cells (Figure 13.2). Microhairs are common and

appear to be longer than the type usually found in this

group (Figure 13.2).

Specimens examined

CAPE. — 3419 (Caledon): Caledon Distr. , Vogelgat (-AD), Wil-

liams 3086.

Discussion

The epidermal anatomy of subsp. disticha is virtually

identical to that of subsp. uniflora and it is predicted that

the leaf anatomy of these two taxa will also be very

similar. This supports their classification as subspecies

of the same species (Gibbs Russell 1984). Freshly fixed

material for anatomical study is required to confirm the

prediction.

DISCUSSION

The leaf anatomy of several species of Ehrharta has

been studied by various authors (Metcalfe 1960; Jacques

Felix 1962; Renvoize 1985) but none of these studies,

except those of Tateoka (1963) dealt with any represen-

tatives of the Setacea group. Tateoka (1963) included

four species of this group but did not notice any arm

cells. Consequently, the unique anatomy of this most

interesting group has received very little attention. If the

presence of these arm cells had been reported earlier, it is

certain that it would have had a profound influence on

the classification of the genus in the Poaceae. On the

evidence of spikelet morphology and embryo anatomy

(Reeder 1957, 1962) Ehrharta has been placed in the

Oryzeae (Stebbins & Crampton 1961). However, the

leaf blade anatomy of the representatives studied to date

showed no affinity with the Bambusoideae (Tateoka

1963). The genus has therefore been retained in the

Arundinoideae, with which it conforms in leaf anatomy

(Renvoize 1981). Awareness of the arm cells in the Seta-

cea group would undoubtedly have strengthened the ar-

gument for retention of the genus in the Bambusoideae.

For these reasons it is regrettable that the detailed and

comprehensive study of the genus by Engelbrecht (1956)

was never published. In this thesis, 26 species of Ehr-

harta were studied in detail and their structure inter-

preted with considerable insight. Unfortunately, this

work has remained virtually unknown and has not been

considered in the taxonomy of the genus to date.

Engelbrecht (1956) observed wall invaginations in all

members of the Setacea group except E. setacea subsp.

scabra and subsp. disticha which were not examined by

him. In addition, he reported the presence of arm cell-

like invaginations in E. rehmannii, E. ramosa, E.

subspicata, E. gigantea and E. villosa. These latter ob-

servations are not substantiated by the present study

(Gibbs Russell 1987) and will be discussed fully in a

later paper. Engelbrecht (1956) used the possession of

these arm cells to divide the genus into two distinct

groups of species. The group with arm cells was further

subdivided into those without dumbbell-shaped silica

bodies (the Setacea group) and those with dumbbell-

shaped silica bodies (the Villosa and Ramosa groups).

His circumscription of the Setacea group is, therefore,

identical to the conclusions of the present study and it is

Bothalia 17,1 (1987)

87

FIGURE 12. Scanning electron micrographs of the abaxial epidermis of Ehrharta selacea. 1-2, E. setacea subsp. setacea, Ellis 4679: 1 , all

intercostal long cells interspaced by single, raised and rounded silica body, X 200; 2, microhair with distal cell collapsed, x 1000. 3-6,

E. setacea subsp. scabra. 3-4, Ellis 4652: 3, prickles and asperites separating individual long cells, x 60; 4, microhair with short,

inflated distal cell, x 1000. 5-6, Ellis 4653: 5, epidermis with few prickles, long cells separated by silica bodies and stomatal files

present, x 200; 6, microhair with slightly elongated distal cell, x 1000. 7-8, E. setacea subsp. uniflora, Ellis 4670: 7, clear zonation

with stomatal files and mid-intercostal long cells slightly inflated, x 100; 8, microhair with tapering distal cell, x 1000.

88

Bothalia 17,1 (1987)

FIGURE 13. — Abaxial epidermis of Ehrharta setacea subsp. disticha, Williams 3086. 1, clearly differentiated costal and intercostal zones,

scale = 10 jizm; 2, intercostal zones with microhairs (m) and stomata, scale = 10 fim.

truly unfortunate that these valuable observations have

had to wait over 30 years before being confirmed and

published.

Tateoka (1963), on the other hand, also examined the

leaf anatomy of several representatives of the Setacea

group (E. dodii, E. rupestris, E. setacea and E. trico-

stata ) but reported that ‘the mesophyll cells do not seem

to have an arm (a projection of cell membranes), as far as

the present author can observe’. The fact that no Ehr-

harta species were seen to possess arm cells or complex

midrib vasculature was used by Tateoka (1963) to con-

vincingly argue the case for the phylogenetic separation

of the Ehrharteae and Oryzeae into different tribes and

subfamilies. If he had been aware of the arm cells in the

Setacea group, this reasoning would not have been

nearly so sound and would probably never have been

propounded.

The present paper proves beyond doubt, with substan-

tiating photographic evidence, that arm cells, compara-

ble to those in most Oryzeae, occur in some of the taxa

of the Setacea group, E. setacea subsp. scabra in parti-

cular. These invaginations, together with chlorenchyma

cell size, appear to become progressively reduced in

most of the other taxa and are difficult to observe, even

in freshly fixed, fully hydrated material. In reconstituted

dried material the invaginations are apparently easier to

see (Engelbrecht 1956) but may be overlooked as being

artefacts due to dehydration and subsequent wall shrink-

age.

The Setacea group of species is obviously a closely

related unit with very uniform leaf anatomy. The degree

of modification of the arm cells which appears to have

occurred in the different taxa of the group is, conse-

quently, most unexpected. Two entirely different types

of modification appear to have taken place which deviate

from the E. setacea subsp. scabra condition. E. setacea

subsp. uniflora from lower altitudes with more equable

climates and possibly higher levels of soil nutrients

seems to have developed a diffuse mesophyll with an

extensive air space system associated with the possession

of abaxial stomata. No invaginations are evident in the

chlorenchyma cell walls of taxa with this type of meso-

phyll and all intermediate stages can be distinguished.

Taxa from the opposite type of habitat, from high alti-

tudes with extremes of temperature and a low level of

soil nutrients, on the other hand, have a very different

type of leaf anatomy which appears to be derived from

the basic subsp. scabra anatomy. This type of anatomy

is probably linked to the perennial nature of these leaves

and is characterized by the mesophyll cells becoming

increasingly reduced in size until they are small, straight-

walled and isodiametric with a corresponding almost to-

tal reduction in the air space system. This anatomical

type is always associated with a setaceous type of leaf

and with the absence of stomata. A clear reduction series

is evident in E. rupestris and these two very different

mesophyll structures can possibly be interpreted as dif-

ferent derivations of the E. setacea subsp. scabra type of

anatomy. It may be relevant to this argument that E.

setacea subsp. scabra itself is limited to the east of the

range of most of the taxa of the Setacea group where

summer rainfall becomes a significant factor. It can be

postulated that the loss of the arm cell invaginations may

be an adaptation to winter rainfall.

These two different anatomical types in the Setacea

group mirror the situation prevailing in many of the Dan-

thonieae of the Cape Fynbos region. Taxa from high

altitudes, such as Merxmuellera, P seudopentameris and

Pentameris (Ellis 1980, 1985a, b,c), resemble the latter

type, whereas species from the lowland Fynbos areas,

such as some species of Plagiochloa and Lasiochloa ,

have a more diffuse type of mesophyll. It appears, there-

fore, that habitat has been more significant than phyloge-

netic relationships in the evolution of these anatomical

features of the leaf blades in many Cape Fynbos grass

taxa.

The fact that the typical arm cell type in the mesophyll

of E. setacea subsp. scabra appears to be modified so

easily suggests that caution must be exercised in attach-

ing too much taxonomic significance to the presence of

this single character. Watson et al. (1985) discuss the

occurrence of arm cells in the family. They rightly con-

sider them to be a feature of high taxonomic value but

point out that they can occur sporadically in distant,

unrelated groups. No satisfactory explanation can be

given for this type of distribution. The presence of arm

cells in the Setacea group of Ehrharta does not neces-

sarily imply bambusoid affinities, particularly as they are

not accompanied by fusoids and complex midrib vascu-

lature. Rather, arundinoid links are suggested by the type

of mesophyll seen in E. setacea subsp. setacea (Tateoka

1963) and perhaps the arundinoid type of leaf anatomy

could have been derived in a similar manner to that de-

scribed in the Setacea group of Ehrharta from a prim-

itive bambusoid-like ancestor possessing arm cells. The

Bothalia 17,1 (1987)

89

Setacea group, therefore, can be viewed as a primitive

arundinoid type exhibiting some links to the peripheral

groups of the bambusoids.

CONCLUSIONS

In both leaf anatomy and spikelet morphology the Se-

tacea group differs more from all the other groups of

Ehrharta than any of these differ from each other. Apart

from arm cells, many other anatomical characters sepa-

rate the Setacea group from the other groups (Gibbs Rus-

sell & Ellis 1987). Examples are microhair and silica

body shape. Furthermore no anatomical intermediates

were found linking this group to any of the other equiva-

lent groups of Ehrharta. These other groups, on the

other hand, are sometimes linked by intermediate speci-

mens or taxa. This suggests that the Setacea group may

deserve separate subgeneric or generic status but consid-

eration of the generic and infrageneric classification of

Ehrharta will be dealt with in the last of this series of

papers.

ACKNOWLEDGEMENTS

The author is grateful to Mrs H. Ebertsohn for techni-

cal 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. CNA, Johannesburg.

ELLIS, R. P. 1980. Leaf anatomy of the South African Danthonieae

(Poaceae). U. Merxmuellera disticha. Bothalia 13: 185-189.

ELLIS, R. P. 1985a. Leaf anatomy of the South African Danthonieae

(Poaceae). X. Pseudopentameris. Bothalia 15: 561-566.

ELLIS, R. P. 1985b. Leaf anatomy of the South African Danthonieae

(Poaceae). XI. Pentameris longiglumis and Pentameris sp. nov.

Bothalia 15: 567-571.

ELLIS, R. P. 1985c. Leaf anatomy of the South African Danthonieae

(Poaceae). XIII. Pentameris macrocalycina and P. obtusifolia.

Bothalia 15: 579-585.

ENGELBRECHT, A. H. P. 1956. ' n Morfologiese studie van die

genus Ehrharta Thunb. M.Sc. thesis, University of Pretoria.

GIBBS RUSSELL, G. E. 1984. Notes on species of Ehrharta with a

short first sterile lemma. Bothalia 15: 149-151.

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 the

genus Ehrharta (Poaceae) in southern Africa. Bothalia 17: 51-65.

JACQUES-FELIX, H. 1962. Les graminees d'Afrique Tropicale. 1.

Generalites, classification, description des genres. Institut de Re-

cherches Agronomiques Tropicales et des Cultures Vivrieres,

Paris.

METCALFE, C. R. 1960 . Anatomy of the Monocotyledons . I. Grami-

neae. Clarendon Press, Oxford.

REEDER, J. R. 1957. The embryo in grass systematics. American

Journal of Botany 44: 756-768.

REEDER, J. R. 1962. The Bambusoid embryo: a reappraisal. Ameri-

can Journal of Botany 49: 639-64 1 .

RENVOIZE, S. A. 1981. The subfamily Arundinoideae and its posi-

tion in relation to a general classification of the Gramineae. Kew

Bulletin 36: 85-102.

RENVOIZE, S. A. 1985. A survey of leaf-blade anatomy in grasses.

V. The bamboo allies. Kew Bulletin 40: 509-535.

RENVOIZE, S. A. 1986. A survey of leaf-blade anatomy in grasses.

Vm. Arundinoideae. Kew Bulletin, in press.

STEBBINS, G. L. & CRAMPTON, B. 1961 . A suggested revision of

the grass genera of North America. Recent Advances in Botany 1:

133-145. University of Toronto Press.

TATEOKA, T. 1963. Notes on some grasses. XIII. Relationship be-

tween Oryzeae and Ehrharteae, with special reference to leaf ana-

tomy and histology. The Botanical Gazette 124: 264—270.

WATSON, L., CLIFFORD, H. T. & DALLWITZ, M. J. 1985. The

classification of the Poaceae: subfamilies and supertribes. Austra-

lian Journal of Botany 33: 433-484.

Bothalia 17,1:91-95(1987)

Six cultivars of Solanum macrocarpon (Solanaceae) in Ghana

Z. R. BUKENYA* and J. B. HALL**

Keywords: crop evolution, egg plant, Ghana, new cultivars, Solanum dasyphyllum, Solanum macrocarpon complex

ABSTRACT

The Solanum macrocarpon complex (the cultivated egg plant) has been studied in Ghana using morphological and

experimental methods. Six cultivars belonging to the S. macrocarpon complex have been recognized and described. The

cultivars are S. macrocarpon ‘Gboma’, S. macrocarpon ‘Mankessim’, S. macrocarpon ‘Akwaseho’, S. macrocarpon

‘Kade’, S. macrocarpon ‘Sarpeiman’ and S. macrocarpon ‘Bui’. The very spiny, hairy plant traditionally called

S. dasyphyllum is regarded as the wild ancestor from which the cultivars have been derived through a process of crop

evolution. The variation within S. macrocarpon complex is attributable to genotypic differences and environmental factors.

UITTREKSEL

Die Solanum macrocarpon- kompleks (die gekweekte eierplant) is in Ghana bestudeer deur gebruik te maak van

morfologiese en eksperimentele metodes. Ses kultivars wat aan die S. macrocarpon- kompleks behoort, is erken en beskryf.

Die kultivars is S. macrocarpon ‘Gboma’, S. macrocarpon ‘Mankessim’, S. macrocarpon ‘Akwaseho’, S. macrocarpon

‘Kade’, 5. macrocarpon ‘Sarpeiman’ en S. macrocarpon ‘Bui’. Die baie doringrige, harige plant wat tradisioneel

S. dasyphyllum genoem word, word as die wilde voorouer waaruit die kultivars deur ’n proses van gewas-evolusie ontstaan

het, beskou. Die variasie binne die S. macrocarpon- kompleks kan toegeskryf word aan genotipiese verskille en

omgewingsfaktore.

INTRODUCTION

Solanum macrocarpon L. is an important native Afri-

can vegetable, especially in west Africa where both the

leaves and fruits are eaten. In east Africa, particularly

Uganda, it is however a minor crop and it is mainly the

leaves that are eaten (Bukenya 1984). The crop is also

cultivated in South America and Asia. It was introduced

to Europe from India, described and illustrated by Miller

(1759) and named by Linnaeus (1771).

Despite its importance as a vegetable and its wide-

spread occurrence, little has been done until now to re-

cognize and describe any of the cultivars.

METHODS AND MATERIALS OF MORPHOLOGICAL STUDY

Materials belonging to the S. macrocarpon complex

were collected from different localities throughout

southern Ghana, and also (mainly seeds) obtained from

elsewhere in Africa (Togo, Nigeria and Uganda). In sou-

thern Ghana, farms where S. macrocarpon is grown,

were visited. Herbarium specimens were made. Ripe

fruits, from which seeds were later extracted, were col-

lected from farms and in some cases from markets.

Occasionally, when mature fruits were not available,

transplants were collected and later replanted in an ex-

perimental garden. Photographs of flowering and fruit-

ing plants were taken.

Each individual (transplant) or each fruit was given a

different accession number. In the laboratory, seeds

from different accessions were extracted and spread on

paper to dry at room temperature. After drying a mini-

* Botany Department, Makerere University, P.O. Box 7062, Kam-

pala, Uganda.

** Formerly: Botany Department, Makerere University, P.O. Box

7062, Kampala, Uganda. (Deceased).

mum of 20 seeds from each accession (fruit), they were

sown in compost soil in seed pots. Pots were housed in a

greenhouse at a mean daily maximum temperature of

34,8 °C and mean daily minimum temperature of

22,1 °C. Pots were watered every two to three days.

When seedlings were at the three-leaved stage (about

two weeks after germination), they were picked out of

the seed pots and transferred to seed boxes kept outside

in sunshine. When the plants were about a month old

they were removed from the seed boxes and planted in

the experimental garden.

About ten plants of each accession were planted. The

plots were constantly weeded and plants watered when

necessary. The major pests of the plants (caterpillars)

were controlled by spraying with DDT (Five grammes of

powder in about four litres of water).

The morphological characteristics of the plants were

observed in their developmental process. When plants

had fully grown, measurements of the characters of

stem, leaves, flowers and fruits were taken.

The following accessions were studied:

5. macrocarpon ‘Gboma’

Hall & Bukenya S32 & S35 (Ghana, Pamproase, nr Achimota); Hall &

Bukenya S34 (Ghana, Sarpeiman, nr Achimota); Hall & Bukenya S39

(Ghana, Avenor, nr Accra); Hall S54 (Togo, Lome); Hall S55 (Togo,

Kleve); Hall S56 (Nigeria, Lagos Campus); Hall & Bukenya S58,

(Ghana, Accra market); Hall 47172 (Ghana, Medina, nr Accra).

S. macrocarpon ‘Mankessim’

Hall & Bukenya 47124 (Ghana, Mankessim); Hall 47170 & 47171

(Ghana, Nkwanta village).

S. macrocarpon ‘Akwaseho’

Hall & Bukenya S21 (Ghana, Akwaseho, Kwahu).

92

Bothalia 17,1 (1987)

S. macrocarpon ‘Kade’

Hall 47169 (Ghana, Kade).

S. macrocarpon ‘Sarpeiman’

Hall & Bukenya S45 & S47 (Ghana, Sarpeiman).

S. macrocarpon ‘Bui’

Hall & Swaine 46212 (Ghana, Bui N. P.); Sobey 45080 (Ghana, Mole

Game Reserve).

S. dasyphyllum var. dasyphyllum (the putative wild an-

cestor)

Anoff s.n. (Ghana, Nsawam); Morton 7484 (Ghana, nr Gambaga);

Katende UG. 2 (Uganda, Kampala).

The above specimens were deposited in University of

Ghana Herbarium.

RESULTS

A summary of morphological measurements is given

in Tables 1-4.

TABLE 1.— Summary of means of leaf characters. Each figu/e represents a mean of no less than 20 measurements

TABLE 3. — Summary of means of inflorescence characters. Each figure represents a mean of no less than 20 measurements

TABLE 4. — Summary of means of characters of hermaphrodite flowers (mm). Each figure represents a mean of no less than 20

measurements

Bothalia 17,1 (1987)

93

DISCUSSION AND CONCLUSIONS

The results of morphological study show that there is

tremendous variation within the S. macrocarpon com-

plex. Some of the accessions within the complex have

big, deeply lobed leaves while others have small and less

lobed leaves. Also flowers vary in their size and depth of

corolla lobing. Whereas the flower calyx length of many

accessions was 10-22 mm, one accession had a calyx

length of about 60 mm; whereas the ratio between lamina

length and petiole length of many accessions was 8-10, a

ratio of 25-46 was obtained in one accession.

Apart from the above morphological variation, con-

sidered to be due mainly to genotypic differences, en-

vironmental factors were also seen to play a significant

part in the phenotypic variation in the S. macrocarpon

complex. One such factor was noted to be cultivation

methods. For example, at Avenor near Accra in Ghana,

5. macrocarpon was seen to be grown in crowded rows

and the leaves were removed constantly for sale when

the plants were still young. Figure 1 shows a typical

planting of S. macrocarpon on a farm at Avenor. The

plants here had a height of less than 0,3 m, and they had

small fruits (30-35 mm diam.) and small leaves (170 x

90 mm). When seeds collected from a farm at Avenor

were grown in our experimental farm with proper spac-

ing of plants and no trimming, the plants attained a

height of about 0,7 m with a fruit diameter of 40-60 mm

and a mean leaf size of 255 x 164 mm.

FIGURE 1 . — A typical planting of Solarium macrocarpon on a farm at

Avenor near Accra, July 1979.

It was Bitter (1923) who first realized the tremendous

variation within the S. macrocarpon complex and he

described several infraspecific taxa. He also noted that

the traditional separation of S. macrocarpon from S.

dasyphyllum on the basis of the latter being spiny and

hairy and the former unarmed and glabrous, was not

satisfactory. He was for example able to describe, ‘a

spineless variety of 5. dasyphyllum and a spiny subspe-

cies of 5. macrocarpon’ .

We noted that there was a continuous range of varia-

tion in hairiness and spininess from very hairy and spiny

accessions to glabrous and unarmed accessions. It also

appears that spininess is a heterozygous character. This

was demonstrated by the fact that when seeds from a

single fruit (Katende LJG.l — S. macrocarpon) were

sown, about 60 % of the plants that grew were spiny and

40 % not spiny. Some plants of accession Hall 47172 (S.

macrocarpon) from Medina near Accra, were seen to

bear spines, though S. macrocarpon, according to most

authors, is non-spiny.

The type specimen of S. macrocarpon is Thonning

117 from Guinea [Ghana]. We have studied the IDC

microfiche photographs of two duplicates on IDC micro-

fiche No. 2203-101: HI. 3-6. The specimen illustrated

in EH. 3 is glabrous, leaves oblong and subentire and that

in III. 5 is glabrous, leaves oblong. Some of the leaves

on this specimen are subentire while others are more or

less deeply double-lobed with major lobes bearing minor

lobes. The type specimens for S. dasyphyllum, Thonning

144, came from ‘Guinea’ (i.e. near Accra, Ghana (Hep-

per 1976). The IDC microfiche photographs (No.

2203-102: I. 1-4) show that the plants have oblong,

deeply double-lobed leaves. They are very hairy and the

stem, midribs and primary lateral nerves bear robust

spines.

Bitter (1923) and recent other workers like Heine

(1963) realized that the two species are very close.

Bukenya (1980 in unpublished M.Sc. thesis) con-

cluded that the species traditionally called S. macrocar-

pon and 5. dasyphyllum were one and proposed to sink

the latter under S. macrocarpon.

In this paper we have treated the S. macrocarpon com-

plex as a combination of the former two species, i.e.

S. macrocarpon L. and 5. dasyphyllum Schum. &

Thonn.

Our study has enabled us to identify seven groups

belonging to the 5. macrocarpon complex. Since this

complex is basically a cultivated crop, it is not satisfac-

tory to give formal taxonomic ranks to these groups. We

have therefore decided to give cultivar names to all these

groups, except one which is wild. The wild one matches

Bitter’s S. dasyphyllum var. dasyphyllum in its dense

spines and hairs and double lobing of the leaves. We

believe that this wild plant is the ancestor of the cultivars

and that the cultivars have arisen after a long process of

cultivation and selection for non-spininess and non-hairi-

ness. The name cultivar is used in this context in the

same sense as used in the International Code of Nomen-

clature for Cultivated Plants of 1969.

The cultivar names are either the Ghanaian native

names for the plants or names of localities where the

cultivars were collected in Ghana. An attempt was made

to compare the present cultivars with the taxa that Bitter

(1923) described in Series Macrocarpon.

The following are the named cultivars with their de-

scriptions and illustrations, where possible.

1 . S. macrocarpon ‘Gboma’ (Figure 2)

Plant of low growth, not spiny. Flowers and calyx

small; calyx 15-18 x 4—6 mm. Leaves petiolate, more or

less glabrous, occasionally with a few scattered simple

hairs, very shallowly lobed, middle lobe 20-40 mm

long. Stem glabrous. Fruit medium-sized, 30-40

x 40-50 mm, green or white when young, turning

purplish later; yellow when ripe. Locality: Ghana, Ave-

nor near Accra, 5°33'N, 0°13'W, Hall & Bukenya S39.

94

Bothalia 17,1 (1987)

This cultivar is comparable to Bitter’s S. macrocarpon

var. macrocarpon (var. culvum). Type: Guinea [Ghana],

Thonning 117.

FIGURE 2. — Solatium macrocarpon ‘Gboma’, growing at Legon,

Ghana (Mar. 1980), showing rotate corolla.

2. S. macrocarpon ‘Mankessim’ (Figure 3)

Plants fairly tall, spiny or unarmed. Calyx fairly large,

20-25 x 6 mm. Leaves more or less subsessile, fairly

deeply lobed, middle lobe 40-60 mm long, puberulous

with simple, sessile stellate and stalked stellate hairs.

Stem with same pubescence as leaves. Fruit fairly large,

40-50 x 50-60 mm, white when young, yellow when

ripe.

Locality : Ghana, Mankessim, 5°16'N, 1°0TW, Hall &

Bukenya 47 124 .

This cultivar is comparable to Bitter’s S. macrocarpon

var. primovestitum except that ‘Mankessim’ may bear

spines which are lacking in var. primovestitum and the

former has more and persistent hairs. The locality of

Bitter’s S. macrocarpon var. primovestitum is on the

coast of Mozambique.

FIGURE 3. — Solanum macrocarpon ‘Mankessim’, growing on a farm

at Mankessim, Ghana (July 1979).

3. S. macrocarpon ‘Akwaseho’ (Figure 4)

Plants similar to ‘Mankessim’, spiny or not spiny, but

leaves more deeply lobed, middle lobe up to 70 mm long

and fruits larger: about 50 x 90 mm. Leaves subsessile

with some simple and sessile stellate hairs. Calyx very

big, up to 60 x 30 mm. Fruits ivory-coloured when

young, turning yellow when ripe.

Locality: Ghana, Akwaseho, 6°37'N, 0°49'W, Hall &

Bukenya S21 .

This cultivar is comparable to Bitter’s S. macrocarpon

var. macrocarpon forma megistocalyx Bitter, in the form

of the big calyx. The calyx of our cultivar is even bigger

than in forma megistocalyx. The major difference is that

‘Akwaseho’ has some sessile stellate hairs and spines

may be present on leaves while forma megistocalyx lacks

hairs and spines. The locality of forma megistocalyx was

Congo (Lower Zaire) Region, Kisantu, Gillet s.n.

FIGURE 4. — Solanum macrocarpon ‘Akwaseho’, growing in an ex-

perimental plot at Legon, Ghana (Nov. 1979), showing big fruits,

big fruiting calyx and deeply lobed leaves.

4.5. macrocarpon ‘Kade’ (Figure 5)

Leaves very narrow and deeply lobed, middle lobe up

to 80 mm long; subsessile with simple, sessile stellate or

stalked stellate hairs. Calyx fairly long but narrow, up to

30 x 60 mm. Fruit ovoid or spherical, 30-70 x 40 mm,

green or green with patches when young, turning yellow

when ripe.

Locality: Ghana, Abodon, 6°26'N, 1°32'W, Hall 47169.

This cultivar does not match any of Bitter’s taxa. It is

unique in that its leaves are so narrow and deeply lobed

and it may have an oblong fruit which is unusual in 5.

macrocarpon.

FIGURE 5. — Solanum macrocarpon ‘Kade’, growing on a farm at

Abodon, Ghana (July 1979).

5. 5. macrocarpon ‘Sarpeiman’ (Figure 6)

Plants similar to ‘Gboma’ but almost always spiny and

fairly hairy with simple, branched, sessile stellate and

Bothalia 17,1 (1987)

95

stalked stellate hairs on leaves and stem. Leaves petio-

late. Fruit green when young, turning yellow when ripe.

Locality. Ghana, Sarpeiman, 5°43'N, 0°18'W, Hall &

Bukenya S45.

This cultivar is comparable to Bitter’s S. dasyphyllum

var. semiglabrum (C. W. Wright) Bitter, except that in

the former the spine length on the midrib ranges from

0-13 mm whereas in the latter the range is 6-10 mm.

The type locality of semiglabrum is Nupe-Benue Pro-

vince in northern Nigeria.

FIGURE 6. — Solanum macrocarpon ‘Sar-

peiman’, growing in an experimental

plot at Legon, Ghana (Nov. 1979)

showing spines.

6. S. macrocarpon ‘Bui’

Plants spiny. Leaves subsessile and deeply lobed,

middle lobe 60-70 mm long; with sparse, simple,

branched, sessile and stalked stellate hairs. Stem with

similar pubescence as leaves but denser. Fruit green

when young, turning yellow when ripe.

Locality: Ghana, Bui, 8°16'N, 2°16'W, Hall & Swaine

46212.

This cultivar is comparable to Bitter’s S. dasyphyllum

var. dasyphyllum except that the former is less hairy than

the latter, probably because the former is occasionally

cultivated. Bitter’s locality of var. dasyphyllum is Gui-

nea [Ghana], Thonning 144.

7. Putative wild ancestor (Figure 7)

Plants very hairy and very spiny. Leaves subsessile,

often doubly lobed; middle lobe up to 60 mm long. Ca-

FIGURE 7. — Putative wild ancestor ( Solanum dasyphyllum var. dasy-

phyllum) from Uganda, growing in cultivation at Legon, Ghana

(Mar. 1980), showing mature, deeply lobed leaf with numerous

spines and hairs.

lyx about 10 x 5 mm. Leaves and stem with simple,

branched, sessile or stalked stellate hairs. Fruit green

when young, turning light yellow when ripe.

Locality: Uganda, near Kampala, Katende UG.2\

Ghana, Nsawam, Anoffs.n.

This is comparable to Bitter’s S. dasyphyllum var.

dasyphyllum from Guinea [Ghana] Thonning 144.

ACKNOWLEDGEMENTS

We wish to extend our gratitude to the Uganda

Government which sponsored a two year study of Sola-

num in Ghana. Thanks go to University of Ghana, Legon

for providing the facilities for the study.

Our thanks also go to all individuals who provided us

with specimens and any form of assistance during this

study.

REFERENCES

BITTER, G. 1923. Solana africana 4. Feddes Repertorium Beihefte 16:

1-320.

BUKENYA, Z. R. 1984. Solanum macrocarpon: an under-utilized but

potential vegetable in Uganda. Uganda Journal 41:9-15.

HEINE, H. 1963. Solanaceae. In J. Hutchinson & J. M. Dalziel, Flora

of west tropical Africa, 2nd edn, rev. R. W. J. Keay & F. N.

Hepper, 2: 325-335. Crown Agents, London.

HEPPER, F. W. 1976. The west African herbaria of Isert & Thonning:

a taxonomic revision of the index to the IDC microfiche. Ben-

tham-Moxon Trust, Kew.

LINNAEUS, C. 1771. Mantissa plantarum altera. Stockholm.

MILLER, P. 1759. Gardener's dictionary. 7th edn, 2 vols.

WRIGHT, C. H. 1906. Solanaceae. In W. T. Thiselton-Dyer, Flora of

tropical Africa 4: 207-261. Reeve, London.

Bothalia 17,1:97-104(1987)

of Fusarium (Fungi: Hyphomycetes) in South Africa,

W. F. O. MARASAS*, SANDRA C. LAMPRECHT**, P. S. VAN WYKt and R. Y. ANELICH*

Bibliography

1945-1985

Keywords: bibliography, Fusarium, Fungi, Hyphomycetes, South Africa

ABSTRACT

A taxonomic study of the genus Fusarium in South Africa was published by Doidge in 1938. A record of all the fungi,

including Fusarium, known to occur in South Africa, and of the literature concerning them until the end of 1945, was

compiled by Doidge in 1950. The present bibliography attempts to provide an index to all the literature on Fusarium in

South Africa published after 1945 until the end of 1985. An alphabetical list of Fusarium species is provided and the hosts

and/or substrates from which each species has been recorded are listed alphabetically together with references to the

literature cited. A separate alphabetical list of hosts and substrates together with the Fusarium species recorded on each is

also included.

UITTREKSEL

’n Taksonomiese studie van die genus Fusarium in Suid-Afrika is in 1938 deur Doidge gepubliseer. In 1950 het

Doidge al die fungi, waaronder Fusarium, wat in Suid-Afrika bekend is en die toepaslike literatuur tot aan die einde van

1945, aangeteken. Die huidige bibliografie poog om ’n indeks te voorsien tot al die literatuur oor Fusarium in Suid-Afrika

wat na 1945 tot aan die einde van 1985 gepubliseer is. ’n Alfabetiese lys van Fusarium- spesies word voorsien en die gashere

en/of substrate waarop elke spesie aangeteken is, word in ’n alfabetiese lys aangegee tesame met verwy sings na die betrokke

literatuur. ’n Aparte alfabetiese lys van gashere en substrate tesame met die Fusarium- spesies wat op elk aangeteken is,

word ook ingesluit.

INTRODUCTION

A major taxonomic study of the genus Fusarium in

South Africa was published by Doidge (1938). In this

publication a detailed account was given of the occur-

rence of 26 species, 31 varieties, and 13 formae of Fusa-

rium on a large number of host plants, as well as insects

and other fungi. Doidge (1950) compiled a record of all

the fungi and lichens known to occur in South Africa and

of the literature concerning them, until the end of 1945.

This monumental compilation included some 29 species,

25 varieties, and 11 formae of Fusarium. These two

publications are the basis of all the existing knowledge

about the occurrence of the genus Fusarium in South

Africa. Several subsequently published lists of plant

diseases in South Africa (Doidge et al. 1953; Gorter

1961, 1973, 1977, 1981) have added some information

about the occurrence of Fusarium spp. as plant patho-

gens in South Africa, but did not include the literature

dealing with the occurrence of Fusarium spp. as sapro-

phytes or mycotoxin producers in South Africa.

The aim of the present publication is to compile an

index to all the available literature on Fusarium in South

Africa published after 1945, and thus not included in

Doidge (1950), to the end of 1985. In addition to phyto-

* National Research Institute for Nutritional Diseases, South African

Medical Research Council, P.O. Box 70, Tygerberg 7505.

** Plant Protection Research Institute, Private Bag X5017, Stellen-

bosch 7600.

t Department of Plant Pathology, University of the Orange Free

State, P.O. Box 339, Bloemfontein 9300.

t Plant Protection Research Institute, Private Bag X134, Pretoria

0001.

pathological aspects, the literature surveyed included

mycological, mycotoxicological, and ecological studies.

A few references to papers published prior to 1945 but

not cited by Doidge (1950) are included. It is hoped that

this bibliography, together with Doidge (1938) and

Doidge (1950), will be as complete a guide as possible to

the literature on Fusarium in South Africa to the end of

1985.

The material in this bibliography is arranged in three

parts: References; Fusarium species; Hosts and sub-

strates.

REFERENCES

Complete references to all the literature cited are given. The

geographical area referred to as ‘South Africa’ in this bibliography was

considered to include the Republic of South Africa, South W est Africa/

Namibia, and the Republics of Bophuthatswana, Ciskei, Transkei and

Venda. Unpublished reports, articles in popular journals, abstracts, and

theses are excluded from the literature surveyed.

AUCOCK, H. W., MARASAS, W. F. O., MEYER, C. J. &

CHALMERS, P. 1980. Field outbreaks of hyperoestrogenism

(vulvo-vaginitis) in pigs consuming maize infected by Fusarium

graminearum and contaminated with zearalenone. Journal of the

South African Veterinary Association 5 1 : 163-166.

BOOTH, C. 1971. The genus Fusarium. Commonwealth Mycological

Institute, Kew, England, pp. 237.

BREW'ER, P. J. 1962. Potato stem-end rot caused by Fusarium spp.

South African Journal of Agricultural Science 5: 47 5 — 48 1 .

BRYSON, R. W. 1982. Kikuyu poisoning and the army worm. Jour-

nal of the South African Veterinary Association 53: 161-165.

BURGESS, L. W., NELSON, P. E. & TOUSSOUN, T. A. 1982.

Characterization, geographic distribution and ecology of Fusa-

rium crookwellense sp. nov. Transactions of the British Myco-

logical Society 79: 497-505.

BURGESS, L. W., NELSON, P. E. , TOUSSOUN. T. A. & MARA-

SAS, W. F. O. 1985. Fusarium scirpi: emended description and

notes on geographic distribution. Mycologia 77: 212-218.

98

Bothalia 17,1 (1987)

COMMONWEALTH MYCOLOGICAL INSTITUTE (CMI). 1977a.

Distribution maps of plant diseases. Fusarium oxysporum /. sp.

cubense. Map No. 31. Edn 4. Commonwealth Mycological Insti-

tute, Kew, England.

COMMONWEALTH MYCOLOGICAL INSTITUTE (CMI). 1977b.

Distribution maps of plant diseases. Gibberella fujikuroi. Map

No. 102. Edn 5. Commonwealth Mycological Institute, Kew,

England.

COMMONWEALTH MYCOLOGICAL INSTITUTE (CMI). 1978.

Distribution maps of plant diseases. Gibberella fujikuroi var.

subglutinans. Map No. 191. Edn 3. Commonwealth Mycological

Institute, Kew, England.

COMMONWEALTH MYCOLOGICAL INSTITUTE (CMI). 1984a.

Distribution maps of plant diseases. Fusarium culmorum. Map

No. 440. Edn 2. Commonwealth Mycological Institute, Kew,

England.

COMMONWEALTH MYCOLOGICAL INSTITUTE (CMI). 1984b.

Distribution maps of plant diseases. Fusarium oxysporum /. sp.

cucumerinum. Map No. 558. Edn 1. Commonwealth Mycological

Institute, Kew, England.

COETZER, J. A. W., KELLERMAN, T. S. & NAUDE, T. W. 1985.

Neurotoxicoses of livestock caused by plants and fungi in southern

Africa. Republic of South Africa Department of Agriculture and

Water Supply Technical Communication No. 199: 1-38.

COMBRINK, J. C., KOTZE, J. M., WEHNER, F. C. & GROBBE-

LAAR, C. J. 1985. Fungi associated with core rot of Starking

apples in South Africa. Phytophylactica 17: 81-83.

COMBRINK, N. J. J. & PRINSLOO, K. P. 1975. Houvermoe van

aartappels soos beinvloed deur bemestingstowwe. Crop Produc-

tion 4: 41-45.

COMBRINK, N. J. J., PRINSLOO, K. P. & JANDRELL, A. C.

1975. Die invloed van kalsium, fosfaat en boor op die houvermoe

en kwaliteitsbepalende knoleienskappe by aartappels. Agro-

plantae 7: 81-84.

CROOKES, C. A. & RUKENBERG, F. H. J. 1985a. Isolation of

fungi associated with blossom malformation of mangoes. South

African Mango Grower' s Association Research Report 5: 10-14.

CROOKES, C. A. & RUKENBERG, F. H. J. 1985b. A literature

review of the distribution, symptomology, cause and control of

mango blossom malformation. South African Mango Grower's

Association Research Report 5: 15-24.

DARVAS, J. M. 1978. Stem-end rot and other post-harvest diseases.

South African Avocado Grower’s Association Research Report 2:

49-50.

DARVAS, J. M. 1979. Ecology of avocado root pathogens. South

African Avocado Grower’s Association Research Report 3:

31-32.

DARVAS, J. M. & KOTZfi, J. M. 1979. Stem-end rot and other post-

harvest diseases. South African Avocado Grower’s Association

Research Report 3: 41—43.

DARVAS, J. M. & KOTZ£, J. M. 1981. Post-harvest diseases of

avocados. South African Avocado Grower's Association Year

Book 4: 63-66.

DARVAS, J. M., SCOTT, D. B. & KOTZE, J. M. 1978. Fungi

associated with damping-off in coniferous seedlings in South Afri-

can nurseries. South African Forestry Journal 104: 15-19.

DEACON, J. W. 1984a. Panama wilt in bananas. Farming in South

Africa, Bananas. Pamphlet No. H. 1/1984: 1-4. Department of

Agriculture and Water Supply, Pretoria.

DEACON, J. W. 1984b. Panama disease of bananas in South Africa.

Horticultural Science 1 : 29-3 1 .

DEACON, J. W., HERBERT, J. A. & DAMES, J. 1985. False

panama disorder of bananas. Subtropical. 15-18.

DIECKMAN, F., MANICOM, B. Q. & COETZEE, K. 1982. An

attempt to control blossom malformation of mangoes with che-

mical sprays. Subtropica 3: 15-16.

DOIDGE, E. M. 1938. Some South African Fusaria. Bothalia 3:

331-483.

DOIDGE, E. M. 1950. The South African fungi and lichens to the end

of 1945. Bothalia 5: 1-1094.

DOIDGE, E. M., BOTTOMLEY, A. M., VAN DER PLANK, J. E. &

PAUER, G. D. 1953. A. revised list of plant diseases in South

Africa. Union of South Africa Department of Agriculture Science

Bulletin No. 346: 1-122.

DU TOIT, J. J. 1968. Root rot of young maize plants — the causal

fungi. South African Journal of Agricultural Science 1 1: 595-604.

DU TOIT, J. J. 1969. Root rot of young maize plants and certain

mycological factors pertaining to the soil. Phytophylactica 1:

51-56.

DU TOIT, J. J. & INGAMELLS, C. J. 1972. Stubby root of chicory

induced by Fusarium oxysporum. Phytophylactica 4: 101-104.

DUTTON, M. F. & WESTLAKE, K. 1985. Occurrence of myco-

toxins in cereals and animal feedstuffs in Natal, South Africa.

Journal of the Association of Official Analytical Chemists 68:

839-842.

EDMONSTONE-S AMMONS, C. P. 1957. Pineapples. Black spot:

symptoms and control. Farming in South Africa 32, 11: 22-25.

EDMONSTONE-S AMMONS, C. 1958. Some aspects of black spot in

pineapple. South African Journal of Agricultural Science 1:

111-120.

EICKER, A. 1969. Microfungi from surface soil of forest communities

in Zululand. Transactions of the British Mycological Society 53:

381-392.

EICKER, A. 1970a. Ecological observations on soil fungi. South Afri-

can Journal of Science 66: 327-334.

EICKER, A. 1970b. Vertical distribution of fungi in Zululand soils.

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EICKER, A. 1974. The mycoflora of an alkaline soil of the open-

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EICKER, A. 1975. A survey of the antimycotic and antibacterial activ-

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Botany 41: 187-198.

EICKER, A. 1976. Non-parasitic mycoflora of the phylloplane and

litter of Panicum coloratum. Transactions of the British Myco-

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ENGELBRECHT, M. C., SMIT, W. A. & KNOX-DAVIES, P. S.

1983. Damping-off of rooibos tea, Aspalathus linearis. Phytophy-

lactica 15: 121-124.

FERREIRA, J. F. & KNOX-DAVIES, P. S. 1984. Occurrence and

control of Fusarium oxysporum on sweet melon seed. Phytophy-

lactica 16: 67-69.

FERREIRA, J. F. & KNOX-DAVIES, P. S. 1985. Germination of

microconidia of Fusarium oxysporum f. sp. melonis in melon seed

exudates. Phytophylactica 17: 51-52.

GELDERBLOM, W. C. A., THIEL, P. G., VAN DER MERWE,

K. J., MARASAS, W. F. O. & SPIES, H. S. C. 1983. A

mutagen produced by Fusarium moniliforme. Toxicon 21:

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GELDERBLOM, W. C. A., MARASAS, W. F. O., STEYN, P. S.,

THIEL, P. G., VAN DER MERWE, K. J., VAN ROOYEN,

P. H., VLEGGAAR, R. & WESSELS, P. L. 1984a. Structure

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GELDERBLOM, W. C. A., THIEL, P. G., MARASAS, W. F. O. &

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GERLACH, W. 1977. Fusarium lunulosporum spec. nov. von Grape-

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GERLACH, W. & NIRENBERG, H. 1982. The genus Fusarium— a

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Bothalia 17,1 (1987)

101

FUSARIUM SPECIES

The Fusarium species are arranged alphabetically and the hosts and/or substrates from which each species has been recorded are listed

alphabetically under that species together with references to the literature cited. The system of taxonomy and nomenclature of Fusarium used is

that of Nelson el al. (1983). In cases where the names of Fusarium species used in the literature cited cannot be accommodated according to this

system, e.g. F. roseum, the name is indexed as used in the relevant publication(s) with a note that the identity is uncertain. In cases where the

names used in the literature cited are considered to be synonyms of the names accepted by Nelson et al. ( 1983), these names are cited as synonyms

under each species and are also cross-referenced in the alphabetical list, e g. F. verticillioides (Sacc.) Nirenberg is considered to be a synonym

under F . moniliforme Sheldon, but is also included in the alphabetical list under F. verticillioides with a note to ‘see F. moniliforme' . In no case

did the present authors attempt to confirm the identity of any of the Fusarium species or the hosts referred to in the literature cited. The names of

the host plants are given as in the original reference.

acuminatum Ell. & Ev.

Conifers: Darvas etal. (1978)

Eleusine sp.: Kellerman et al. (1980)

Medicago saliva'. Marasas & Bredell (1973); Thompson (1985)

Medicago spp. : Lamprecht et al. (1984)

Panicum coloratum : Kellerman etal. (1980)

Pastures: Kellerman etal. (1980)

Pennisetum typhoides: Rabie etal. (1982)

Sorghum cajfrorum : Rabie etal. (1982)

Tribulus terrestris: Kellerman etal. (1980)

Triticum aestivum: Kellerman et al. (1980)

aquaeductuum (Radik. & Rabenh.) Lagerh.

= aquaeductuum (Radik. & Rabenh.) Lagerh. var. medium

Wollenw.

Conifers: Darvas etal. (1978)

aquaeductuum var. medium (See aquaeductuum )

avenaceum (Fr.) Sacc.

Hordeum vulgare: Rabie etal. (1982)

Medicago spp.: Lamprecht etal. (1984)

Panicum coloratum: Eicker(1976)

Pastures: Kellerman et al. (1980)

Soil: Eicker(1969)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985)

Triticum aestivum: Putterill (1954)

chlamydosporum Wollenw. & Reinking

= chlamydosporum Wollenw. & Reinking vai.fuscum Gerlach

= fusarioides (Frag. & Cif.) Booth

Aspalathus linearis: Engelbrecht etal. (1983)

Cynodon lemfuensis: Gerlach & Nirenberg (1982)

Pastures: Kellerman etal. (1980)

Pennisetum typhoides: Kriek & Marasas (1983); Marasas et al.

(1984a); Rabie etal. (1975, 1982)

Phaseolus vulgaris: Marasas et al. (1984a); Rabie et al. (1978)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985);

Rabie etal. (1978, 1982)

Tribulus terrestris: Kellerman et al. (1980)

Zea mays: Kriek & Marasas (1983); Marasas (1982); Marasas et al.

(1981)

chlamydosporum vai.fuscum (See chlamydosporum)

ciliatum Lk.

Epichloe zahlbruckneriana: Gerlach & Nirenberg (1982)

concolor Reinking (Identity uncertain)

Pennisetum typhoides: Gerlach & Nirenberg (1982); Rabie et al.

(1982)

Pyrus malus: Combrink et al. (1985)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985)

crookwellense Burgess, Nelson & Toussoun

Panicum laevifolium: Burgess et al. (1982)

Triticum aestivum: Burgess et al. (1982)

culmorum (W. G. Smith) Sacc.

Conifers: Darvas etal. (1978)

Eleusine sp.: Kellerman et al. ( 1980)

Panicum coloratum: Kellerman et al. (1980)

Pastures: Kellerman et al. (1980)

Persea americana: Darvas & Kotze (1981)

Soil: Eicker (1969, 1970b); Smith et al. ( 1984)

Triticum aestivum: CMI (1984a); Jooste (1965); Kellerman et al.

(1980); Maas & Kotze (1981); Scott (1980); Smith et al. ( 1984)

decemcellulare Brick

Persea americana: Darvas (1978); Darvas & Kotze(1979, 1981)

dlamini Marasas, Nelson & Toussoun

Soil: Marasas etal. (1985a)

equiseti (Corda) Sacc.

Arachis hypogaea: Martin etal. (1971)

Conifers: Darvas etal. (1978)

Cucumis melo: Reuveni (1985)

Lycopersicum esculentum: Visser (1980)

Medicago sativa: Thompson (1985)

Medicago spp.: Lamprecht etal. (1984)

Pastures: Kellerman etal. (1980)

Pennisetum typhoides: Rabie etal. (1982)

Soil: Eicker ( 1974, 1975); Papendorf (1976); Smith et al. (1984)

Sorghum caffrorum: Martin et al. (1971); Rabie & Liibben (1984);

Rabie & Thiel (1985); Rabie etal. (1982)

Triticum aestivum : Maas & Kotze (1985); Smith et al. (1984)

Zea mays: Kriek & Marasas (1983); Marasas (1982); Marasas et al.

(1981)

equiseti var. longipes (See longipes)

fusarioides (See chlamydosporum)

graminearum Schwabe

Medicago spp. Lamprecht et al. (1984)

Panicum coloratum: Eicker ( 1976)

Soil: Eicker (1976)

Triticum aestivum: Maas & Kotze (1981); Oelofsen (1950); Scott

(1980)

Zea mays: Aucock et al. ( 1980); Gelderblom et al. (1984b); Kriek &

Marasas (1983); Kriiger (1970a, 1970b, 1970c); Kriiger &

Jooste (1967); Kriiger et al. (1965); Marasas (1982); Marasas et

al. (1977, 1979a, 1979b, 1981, 1984a); Steyn (1933); Thiel et

al. (1982); Watt & Breyer-Brandwijk ( 1962)

heterosporum Nees

Trifolium subterraneum: Mills (1984)

heterosporum Nees var. congoense (Wollenw.) Wollenw. (Identity

uncertain)

Grasses infected with ergot or smut: Gerlach & Nirenberg ( 1982)

lateritium Nees

= lateritium Nees var. longum Wollenw.

= stilboides Wollenw.

Citrus spp.: Booth (1971); Gerlach & Nirenberg (1982); Gordon

(1960)

Coffea spp.: Gerlach & Nirenberg ( 1982)

lateritium var. longum (See lateritium)

longipes Wollenw. & Reinking

= equiseti (Corda) Sacc. var. longipes (Wollenw. & Reinking)

Joffe

Eleusine sp.: Kellerman etal. (1980)

Panicum coloratum: Kellerman et al. (1980)

Triticum aestivum: Kellerman et al. (1980)

lunulosporum Gerlach

Citrus paradisi: Gerlach ( 1977); Gerlach & Nirenberg (1982)

lycopersici (See oxysporum f. sp. lycopersici)

merismoides Corda

Sorghum caffrorum: Rabie & Liibben ( 1984); Rabie & Thiel (1985)

moniliforme Sheldon

= verticillioides (Sacc.) Nirenberg

Ananas comosus: Edmonstone-Sammons (1957, 1958)

Arachis hypogaea: Van Warmelo (1967)

Cereals: Scott (1965); Van Rensburg et al. (1971)

102

Bothalia 17,1 (1987)

Conifers: Darvas etal. (1978)

Feeds: Van Warmelo (1967)

Hay: Van Warmelo (1967)

Legumes: Scott (1965); Van Rensburg etal. (1971)

Medicago sativa : Van Warmelo (1967)

Musa spp: Swarts (1978)

Oryza sativa: Gorter (1977)

Panicum coloratum : Eicker (1976)

Pastures: Kellerman etal. (1980)

Pennisetum typhoides : Marasas et al. (1984a); Rabie et at. (1982);

Steyn etal. (1978)

Per sea americana: Darvas (1979)

Phaseolus vulgaris: Van der Walt & Steyn (1943)

Saccharum officinarum: Booth (1971); CMI (1977b)

Soil: Eicker ( 1969); Papendorf( 1976)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985);

Rabie etal. (1982)

Tribulus terrestris: Kellerman etal. (1980)

Voandzeia subterranea: Martin et al. ( 197 1 )

Zea mays: Aucock etal. (1980); CMI (1977b); Coetzer et al. (1985);

Du Toit (1968); Gelderblom et al. (1983, 1984a, 1984b,

1984c); Kellerman et al. (1972); Kriek & Marasas (1983);

Kriek et al. (1981a, 1981b); Kruger (1970a, 1970b, 1970c,

1970d); Kruger & Jooste (1967); Kruger et al. (1965); Marasas

etal. (1976, 1977, 1979a, 1979b, 1981, 1984a, 1984b, 1985b);

Pienaar (1977); Pienaar et al. (1981); Russell & Berjak (1983);

Russell et al. (1983); Steyn (1933, 1950); Steyn et al. (1979);

Thiel etal. (1982); Van Warmelo (1967)

moniliforme var. subglutinans (See subglutinans)

oxysporum Schlecht. emend. Snyd. & Hans.

= oxysporum Schlecht. var. redolens (Wollenw.) Gordon

Arachis hypogaea: Rabie et al. (1982)

Aspalathus linearis: Engelbrecht et al. (1983)

Chrysanthemum cinerariaefolium: Ingamells (1974)

Cichorium intybus: Du Toit & Ingamells (1972)

Citrus spp.: Gordon (1960)

Conifers: Darvas etal. (1978)

Cucumis melo: Ferreira & Knox-Davis (1984, 1985); Reuveni

(1985)

Hordeum vulgare: Rabie etal. (1982)

Man: Young & Meyers (1979)

Mangifera indica: Rijkenberg & Crookes (1985)

Medicago sativa: Thompson (1985)

Medicago spp.: Lamprecht etal. (1984)

Musa spp.: Deacon etal. (1985)

Narcissus pseudonarcissus: Gordon ( 1 960); Wager (1972)

Panicum coloratum: Eicker (1976)

Persea americana: Darvas (1979)

Soil: Darvas (1979); Eicker (1969, 1970a, 1970b, 1974, 1975);

Papendorf (1976); Rijkenberg & Crookes (1985)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985);

Rabie etal. (1982)

Trifolium subterraneum: Mills (1984)

Vitis vinifera: Le Roux etal. (1973)

Zea mays: Du Toit (1968); Kruger (1970a, 1970b, 1970c, 1970d);

Kruger & Jooste (1967)

oxysporum Schlecht. f. sp. cepae (Hanz.) Snyd. & Hans.

Allium cepa: Holz & Knox-Davies (1974, 1976, 1985a, 1985b,

1985c)

Oxalis pes-caprae: Holz & Knox-Davies (1976)

Oxalis zeekoevleyensis: Holz & Knox-Davies (1976)

oxysporum Schlecht. f. sp. cubense (E. F. Smith) Snyd. & Hans.

Musa spp.: CMI (1977a); Deacon (1984a, 1984b); Deacon et al.

(1985); Swarts (1978)

oxysporum Schlecht. f. sp. cucumerinum Owen

Cucumis sativus: Booth (1971); CMI (1984b)

oxysporum Schlecht. f. sp. lycopersici (Sacc.) Snyd. & Hans.

= lycopersici Bruschi

Lycopersicum esculentum: Hofmeyr et al. (1948); Holz (1976); Nel

(1949); Visser(1980)

oxysporum Schlecht. f. sp. melonis Snyd. & Hans.

Cucumis melo: Ferreira & Knox-Davies (1984, 1985)

oxysporum Schlecht. f. sp. niveum (E. F. Smith) Snyd. & Hans.

Citrullus vulgaris: Booth (1971)

oxysporum Schlecht. f. sp. phormii Wager

Phormium tenax: Wager (1947)

oxysporum Schlecht. f. sp. vasinfectum (Atk.) Snyd. & Hans.

Gossypium spp.: Booth (1971)

oxysporum var. redolens (See oxysporum)

poae (Peck) Wollenw.

Saccharum officinarum: Booth (1971)

Zea mays: Kriek & Marasas (1983); Marasas (1982); Marasas et al.

(1981)

roseum Link (Identity uncertain)

Cereals: Scott (1965); Van Rensburg etal. (1971)

Legumes: Scott (1965); Van Rensburg etal. (1971)

Soil: Eicker (1975)

sacchari var. subglutinans (See subglutinans)

sambucinum Fuckel

Humulus lupulus: Gorter (1977)

Persea americana: Darvas (1978); Darvas & Kotze (1979)

Soil: Eicker (1974, 1975)

scirpi Lambotte & Fautr.

Medicago spp.: Burgess etal. (1985); Lamprecht et al. (1984)

Soil: Burgess etal. (1985)

Sorghum caffrorum: Rabie & Liibben (1984)

Zea mays: (Kriiger ( 1970a)

scirpi Lambotte & Fautr. var. compactum Wollenw. (Identity uncer-

tain)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985)

semitectum Berk. & Rav.

Conifers: Darvas etal. (1978)

Musa sp.: Swarts (1978)

Pennisetum typhoides: Rabie etal. (1982)

Soil: Eicker (1969)

Sorghum caffrorum: Martin etal. (1971); Rabie & Liibben (1984)

solani (Mart.) Appel & Wollenw. emend. Snyd. & Hans.

Chrysanthemum cinerariaefolium: Ingamells (1974)

Citrus limonia: Graham etal. (1985)

Crotalaria juncea: Hean (1947)

Cucumis melo: Reuveni (1985)

Hyacinthus sp.: Gordon (1960); Wager (1972)

Insects: Gordon (1960)

Medicago sativa: Thompson (1985)

Persea americana: Darvas (1978); Darvas & Kotze (1979)

Soil: Eicker (1974, 1975); Papendorf (1976)

Sorghum caffrorum: Rabie & Liibben (1984); Rabie & Thiel (1985)

Trifolium subterraneum: Mills (1985)

Vitis vinifera: Le Roux etal. (1973)

Zea mays: Kriiger (1970a)

sp. (See Fusarium species undetermined at end of this list)

sporotrichioides Sherb.

Soil: Papendorf (1976)

Voandzeia subterranea: Kriek & Marasas (1983); Martin et al.

(1971)

stilboides (See lateritium)

subglutinans (Wollenw. & Reinking) Nelson, Toussoun & Marasas

= moniliforme Sheldon var. subglutinans Wollenw. & Reinking

= sacchari (Butler) Gams var. subglutinans (Wollenw. & Rein-

king) Nirenberg

Air: Crookes & Rijkenberg (1985a)

Ananas comosus: Gordon (1960)

Conifers: Darvas etal. (1978)

Crotalaria juncea: Hean (1947)

Mangifera indica: Crookes & Rijkenberg (1985a, 1985b); Dieckman

etal. (1982); Kotz6 (1985)

Morus japonica: Crookes & Rijkenberg (1985a)

Saccharum officinarum: Booth (1971)

Soil: Crookes & Rijkenberg (1985a); Rijkenberg & Crookes (1985)

Sorghum caffrorum: CMI (1978); Rabie & Liibben (1984); Rabie &

Thiel (1985); Rabie etal. (1982)

Syzygium cordatum: Crookes & Rijkenberg (1985a)

Zea mays: Aucock et al. (1980); CMI (1978); Gelderblom et al.

(1984b); Kriek & Marasas (1983); Kriek et al. (1977); Kriiger

et al. (1965); Marasas (1982); Marasas et al. (1977, 1979a,

1979b, 1981, 1984a); Steyn (1933); Steyn et al. (1978); Thiel

(1978); Thiel et al. (1982); Watt & Breyer-Brandwijk (1962)

verticillioides (See moniliforme )

Bothalia 17,1 (1987)

103

Fusarium species undetermined

Ananas comosus: Malan (1954)

Arachis hypogaea: Van Warmelo (1967)

Cereals: Dutton & Westlake (1985)

Conifers: Darvas etal. (1978)

Crotalaria juncea : Hean (1947)

Eleusine sp.: Kellerman etal. (1980)

Feeds: Dutton & Westlake (1985); Van Warmelo (1967)

Glycine max : Dutton & Westlake (1985)

Hay: Van Warmelo (1967)

Mangifera indica: Crookes & Rijkenberg (1985a)

Medicago sativa: Thompson (1985); Van Warmelo (1967)

Musa spp.: Deacon etal. (1985)

HOSTS AND

Panicum coloratum : Eicker (1976); Kellerman et al. (1980)

Pastures: Kellerman etal. (1980)

Pennisetum clandestinum: Bryson (1982); Newsholme etal. (1983)

Pennisetum typhoides : Rabie et al. (1975)

Soil: Crookes & Rijkenberg (1985a); Eicker (1969, 1970b, 1974,

1975); Van Wyk & Baard (1970)

Solanum tuberosum: Combrink & Prinsloo (1975); Combrink et al.

(1975); Visser&Kotzd (1979)

Tribulus terrestris: Kellerman et al. (1980)

Triticum aestivum: Kellerman etal. (1980)

Zea mays: Du Toit (1969); Dutton & Westlake (1985); Kruger

(1970a, 1970b); Kruger & Jooste (1967); Van Warmelo (1967)

The hosts and substrates are listed alphabetically and the Fusarium species recorded on each are listed alphabetically. References to the

literature cited can be found under the relevant Fusarium species. The names of the host plants are given as in the original reference.

Air

F. subglutinans

F . oxysporum f. sp. melonis

F. solani

Allium cepa L.

F. oxysporum f. sp. cepae

Cucumis sativus L.

F. oxysporum f. sp. cucumerinum

Ananas comosus (L.) Merr.

F. moniliforme

F. subglutinans

F. sp.

Arachis hypogaea L.

F. equiseti

F. moniliforme

F. oxysporum

F. sp.

Aspalathus linearis (Burm. f.) R. Dahlg.

F. chlamydosporum

F. oxysporum

Cereals

F. moniliforme

F. roseum

F. sp.

Chrysanthemum cinerariaefoiium (Trev.) Bocc.

F. oxysporum

F. solani

Cichorium intybus L.

F. oxysporum

Citruilus vulgaris Schrad.

F. oxysporum f. sp. niveum

Citrus limonia Osbeck

F . solani

Citrus paradisi Macf.

F. lunulosporum

Citrus spp.

F . latent ium

F. oxysporum

Cynodon lemfuensis Vanderyst

F. chlamydosporum

Eleusine spp.

F . acuminatum

F. culmorum

F. longipes

F. sp.

Epichloe zahlbruckneriana P. Henn.

F . ciliatum

Feeds

F . moniliforme

F. sp.

Glycine max (L.) Merr.

F. sp.

Gossypium spp.

F. oxysporum f. sp. vasinfectum

Grasses infected with Ergot or Smut

F. heterosporum var. congoense

Hay

F. moniliforme

F. sp.

Hordeum vulgare L.

F. avenaceum

F. oxysporum

Humulus lupulus L.

F. sambucinum

Hyacinthus sp.

F. solani

Coffea spp. Insects

F. lateritium F ■ solani

Conifers

F. acuminatum

F. aquaeductuum

F . culmorum

F. equiseti

F. moniliforme

F. subglutinans

F. oxysporum

F. semitectum

F. sp.

Crotalaria juncea L.

F. solani

F. subglutinans

F. sp.

Legumes

F. moniliforme

F. roseum

Lycopersicum esculentum Mill.

F. equiseti

F. oxysporum f. sp. ly coper sici

Man

F. oxysporum

Mangifera indica L.

F. oxysporum

F. subglutinans

F. sp.

Cucumis melo L.

F. equiseti

F. oxysporum

Medicago sativa L.

F. acuminatum

F. equiseti

104

Bothalia 17,1 (1987)

F. moniliforme

F. oxysporum

F. solani

F. sp.

Medicago spp.

F. acuminatum

F. avenaceum

F. equiseti

F. graminearum

F. oxysporum

F. scirpi

Morus japonica L.

F. subglutinans

Musa spp.

F. moniliforme

F. oxysporum

F. oxysporum f. sp. cubense

F. semitectum

F. sp.

Narcissus pseudonarcissus L.

F. oxysporum

Oryza sativa L.

F . moniliforme

Oxalis pescaprae L.

F . oxysporum f. sp. cepae

Oxalis zeekoevleyensis R. Knuth

F. oxysporum f. sp. cepae

Panicum coloratura L.

F. acuminatum

F. avenaceum

F. culmorum

F . graminearum

F. longipes

F. moniliforme

F. oxysporum

F. sp.

Panicum laevifolium Hack.

F. crookwellense

Pastures

F. acuminatum

F. avenaceum

F. chlamydosporum

F. culmorum

F. equiseti

F. moniliforme

F. sp.

Saccharum officinarum L.

F . moniliforme

F. poae

F. subglutinans

Soil

F. avenaceum

F. culmorum

F. dlamini

F. equiseti

F. graminearum

F. moniliforme

F. oxysporum

F. roseum

F. sambucinum

F. scirpi

F. semitectum

F . solani

F. sporotrichioides

F. subglutinans

F. sp.

Solanum tuberosum L.

F. sp.

Sorghum caffrorum Beauv.

F. acuminatum

F. avenaceum

F. chlamydosporum

F. concolor

F. equiseti

F. merismoides

F. moniliforme

F. oxysporum

F. sambucinum

F. scirpi

F. scirpi var. compactum

F. semitectum

F. solani

F. subglutinans

Syzygium cordatum Hochst.

F. subglutinans

Tribulus terrestris L.

F. acuminatum

F. chlamydosporum

F. moniliforme

F. sp.

Trifolium subterraneum L.

F. heterosporum

F. oxysporum

F. solani

Pennisetum clandestinum Hochst. ex Chiov.

F. sp.

Pennisetum typhoides (Burm.) Stapf & Hubbard

F. acuminatum

F. chlamydosporum

F. concolor

F. equiseti

F. moniliforme

F. semitectum

F. sp.

Persea americana Mill.

F. culmorum

F. decemcellulare

F. moniliforme

F. oxysporum

F. sambucinum

F. solani

Phaseolus vulgaris L.

F. chlamydosporum

F. moniliforme

Phormium tenax Forst.

F. oxysporum f. sp. phormii

Pyrus malus L.

F. concolor

F. lateritium

Triticum aestivum L.

F. acuminatum

F . avenaceum

F . crookwellense

F. culmorum

F. equiseti

F. graminearum

F. longipes

F. sp.

Vitis vinifera L.

F. oxysporum

F. solani

Voandzeia subterranea (L.) Thomas

F. moniliforme

F. sporotrichioides

Zea mays L.

F. chlamydosporum

F. equiseti

F. graminearum

F. moniliforme

F. oxysporum

F. poae

F. scirpi

F. solani

F. subglutinans

F. sp.

Bothalia 17,1: 105-119(1987)

The genus R ubus (Rosaceae) in South Africa. IV. Natural hybridiza-

tion

SPIES, J. J.*, STIRTON, C. H.** and DU PLESSIS, H.*

Keywords: chromosomes, hybridization, polyploidy, reproduction, Rosaceae, Rubus, South Africa, speciation

ABSTRACT

The genus Rubus L. is represented in southern Africa by the subgenera Eubatus Focke and Idaeobatus Focke. A

combination of morphological data, data on the reproductive systems of some collections and meiotic chromosome beha-

viour indicates that a hybrid swarm in the eastern Transvaal was formed subsequent to the hybridization between

R. cuneifolius Pursh. taxon B (subgenus Eubatus) and R. longepedicellatus (C. E. Gust.) C. H. Stirton (subgenus Idaeoba-

tus). Other examples of intra- and intersubgeneric hybridization were found during this study of the South African material.

These instances, with examples found in the literature, indicate that the subgeneric subdivisions of Rubus are artificial.

Three different methods were used to analyse the meiotic chromosome configurations. The genome relationship

system of Alonso & Kimber (1981) and Kimber & Alonso (1981) and the modification of the binomial system of Jackson &

Casey (1980) by Spies (1984) proved to be the most sensitive for distinguishing between alio-, segmental alio- and

autoploids.

U1TTREKSEL

Die genus Rubus L. word in suidelike Afrika verteenwoordig deur die subgenera Eubatus Focke en Idaeobatus Focke.

’n Kombinasie van morfologiese data, data rakende die voortplantingsisteem van sommige eksemplare en meiotiese

chromosoomgedrag het aangetoon dat ’n basterkompleks in die oostelike Transvaal gevorm is na die verbastering van R.

cuneifolius Bailey takson B (subgenus Eubatus) en R. longepedicellatus (C. E. Gust.) C. H. Stirton (subgenus Idaeobatus).

Ander voorbeelde van intra- en intersubgeneriese verbastering is tydens hierdie studie in Suid-Afrika gevind en in samehang

met verdere voorbeelde in die literatuur toon dit aan dat die onderverdeling van die genus Rubus in subgenera kunsmatig is.

Drie verskillende metodes is gebruik om die meiotiese chromosoomgedrag van die plante te vergelyk. Die genoom-

verwantskapsisteem van Alonso & Kimber ( 198 1 ) en Kimber & Alonso (1981) en die modifikasies op die binomiale sisteem

van Jackson & Casey (1980) deur Spies (1984) toon aan dat hierdie twee metodes die sensitiefste is om tussen alio-,

segmentele alio- en outoplo'iede plante te onderskei.

CONTENTS

Introduction 105

Materials and methods 105

Results 106

Morphology 106

Reproductive system 107

Chromosome behaviour 110

Discussion Ill

Morphology Ill

Reproductive system 114

Chromosome behaviour 114

Hybridization 116

T axonomic implications of hybridization 117

Conclusions 118

References 118

INTRODUCTION

The genus Rubus is somewhat enigmatic in South

Africa. It forms part of our indigenous flora but natura-

lized species also occur. Most taxa are considered weedy

and yet they are included in a breeding programme to

* Botanical Research Institute, Department of Agriculture and Water

Supply, Private Bag X101 , Pretoria 0001 , South Africa.

** The Herbarium, Royal Botanical Gardens, Kew, Richmond,

Surrey, TW9 3AE, United Kingdom.

improve their agricultural production. The genus also

contains agamic species as well as sexual species. In

short, it is a taxonomist’s nightmare.

The genus Rubus comprises 12 subgenera of which

two are represented in South Africa: Eubatus Focke and

Idaeobatus Focke. In South Africa the subgenus Euba-

tus, or true brambles or blackberries, includes only

exotics, whereas the subgenus Idaeobatus, or rasp-

berries, contains a few exotics and a number of indige-

nous species (Spies & Du Plessis 1985).

It has been proposed (Stirton 1981a & b; Spies & Du

Plessis 1985) that the problems with Rubus taxonomy in

South Africa are caused by the occurrence of apomixis,

hybridization among indigenous species and between in-

digenous and exotic species, the variation produced by a

breeding program with subsequent escape from cultiva-

tion and inadequately collected herbarium material.

Each paper in this series has dealt with a different

aspect of the cytogenetics of Rubus in South Africa. The

aim of this paper is to determine whether natural hybridi-

zation occurs in the South African Rubus complex and

whether this hybridization, if it does occur, is restricted

to intrasubgeneric taxa.

MATERIALS AND METHODS

The following specimens were collected in the veld,

transplanted in the Pretoria National Botanical Garden

106

Bothalia 17,1 (1987)

and subsequently examined for this study [All the her-

barium specimens are housed in the Pretoria National

Herbarium (PRE)]:

R. cuneifolius Pursh taxon A*

NATAL. — 2929 (Underberg): 14 km from Swartberg to Underberg

(-CD), Stirton 8154. 2930 (Pietermaritzburg): 3 km from Midmar Dam

to Lions River (-CB), Henderson & Gaum 93; 5 km from Pieter-

maritzburg to Mooi River (-CB), Liengme s.n.; Highlands Farm

(-CD), Beard 720. 3029 (Kokstad): 40 km from Underberg to Swart-

berg (-BA), Stirton 8157; 1 1 km from Harding to Weza (-DB), Stirton

8102.

R. cuneifolius Pursh taxon B*

TRANSVAAL. — 2329 (Pietersburg): 3 km from Haenertsburg to

Boyne (-CC), Stirton 8033 . 2330 (Tzaneen): Modderfontein (-CC),

Stirton 8013. 2430 (Pilgrim’s Rest): 1 km from Graskop to Sabie

(-DD), Stirton 9800, 9859, 9861, 9868, Henderson & Gaum 18. 2530

(Lydenburg): 5 km from Lydenburg to Sabie (-AB), Henderson &

Gaum 37; Dullstroom (-AC), Stirton 7255. 2628 (Johannesburg):

Heidelbergkloof (-CA), Bredenkamp 123.

R. longepedicellatus (C. E. Gust.) C. H. Stirton

TRANSVAAL. — 2329 (Pietersburg): 10 km from Tzaneen to Hae-

nertsburg (-CC), Stirton 5755; near Pietersburg (-CD), McCullum 13.

2330 (Tzaneen): Pietersburg District (-CC), McCullum 887. 2430 (Pil-

grim’s Rest): Pilgrim’s Rest (-DB), Killick & Strey 2420; Mariepskop

(-DB), Van der Schijff4562; Bourke’s Luck (-DB), Viljoen 27; 1 km

from Graskop to Sabie (-DD), Henderson & Gaum 22, Stirton 9862.

2530 (Lydenburg): 5 km from Lydenburg to Sabie (-AB), Henderson

& Gaum 36; Brooklands (-BA), Henderson cfc Gaum 14; Nelspruit

(-BD), Mogg (PRE 55710). 2531 (Komatipoort): Kruger National Park

(-AB), Van der Schijff 1228 .

NATAL. — 3029 (Kokstad): Ngeli Forest (-DA), Stirton 8135.

R. x proteus sp. ined.

TRANSVAAL. — 2329 (Pietersburg): 10 km from Tzaneen to Hae-

nertsburg (-CC), Stirton 5756, 5783. 2430 (Pilgrim’s Rest): Spekboom

River, Burgersfort (-CB), Henderson 319; Bourke’s Luck (-DB),

Henderson & Gaum 27 , 28, 29, 31, 32; Mac-Mac Waterfalls (-DD),

Henderson & Gaum 20; 6 km from Pilgrim’s Rest to Lydenburg

(-DD), Henderson & Gaum 33; 1 km from Graskop to Sabie (-DD),

Stirton 9797, 9798, 9799, 9801, 9855, 9860, 9862, 9863, 9864, 9865,

9866, 9867, 9869. 2530 (Lydenburg): 3 km from Brooklands to

Hendriksdal (-BA), Henderson & Gaum 72; 33 km from Nelspruit to

Sabie (-BD), Henderson & Gaum 11.

R. rigidus X R. cuneifolius taxon A

NATAL. — 2929 (Underberg): 25 km from Himeville to

Boesmansnek (-DC), Henderson & Gaum 50, 51 .

This cytotaxonomic study concentrated upon a possi-

ble hybrid swarm in the area between Graskop and Sabie

in the eastern Transvaal Lowveld (2430DD) (Stirton

1984). The cytogenetical methods and results were re-

ported by Spies & Du Plessis (1985 & 1986) and Spies,

Du Plessis & Liebenberg (1985). These investigations

included meiotic analyses of aceto-carmine anther

squashes and embryo sac studies.

In order to compare morphological characters of the