Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 23,2

Oct./Okt. 1 993

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

TEGNIESE PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag XI 01,

Pretoria 0001, Republic of South Africa. A catalogue 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 National Botanical Institute, Pretoria, 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 published annually.

A booklet of the contents to Vols 1-20 is available.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,

Pretoria 000 1 , Republiek van Suid-Afrika. 'n Katalogus van alle beskik-

bare publikasies kan aangevra word.

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

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

lyfblad van die Nasionale Botaniese Instituut, Pretoria, is gewy aan die

bevordering van die wetenskap van plantkunde. Die hoofgebiede wat

gedek word, is taksonomie, ekologie, anatomie en sitologie. Twee dele

van die tydskrif en ’n indeks van die inhoud, outeurs en onderwerpe

verskyn jaarliks.

'n Inhoudsopgawe tot volumes 1-20 is beskikbaar.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

The memoirs are individual treatises usually of an ecological nature, but

sometimes dealing with taxonomy or economic botany.

THE FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with accompanying

text. The plates are prepared mainly by the artists at the National

Botanical 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.

Twenty plates are published annually.

An index to Vols 1 — 49 is available.

FLORA OF SOUTHERN AFRICA (FSA)

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

Ciskei, Transkei. Lesotho, Swaziland, Bophuthatswana, Namibia,

Botswana and Venda. The FSA contains descriptions of families, genera,

species, infraspecific taxa, keys to genera and species, synonymy, liter-

ature and limited specimen citations, as well as taxonomic and ecological

notes.

Contributions to the FSA will also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

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 populations. Now

available:

’n Reeks van losstaande omvattende verhandelings oor vemaamlik eko-

logiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.

Die skilderye word meestal deur die kunstenaars van die Nasionale

Botaniese Instituut voorberei. Talle bekende botaniese kunstenaars het

tot die reeks bygedra, soos Cythna Letty (meer as 700 plate), Kathleen

Lansdell, Stella Gower, Betty Connell, Peter Bally en Fay Anderson. Die

Redakteur verwelkom lewende plante van algemene belang of ekono-

miese waarde vir afbeelding.

Twintig plate word jaarliks gepubliseer.

'n Indeks tot volumes 1-49 is beskikbaar.

’n Taksonomiese verhandeling oor the flora van die Republiek van

Suid-Afrika, Ciskei, Transkei. Lesotho, Swaziland, Bophuthatswana,

Namibie, Botswana en Venda. Die FSA bevat beskrywings van families,

genusse, spesies, infraspesifteke taksons, sleutels tot genusse en spesies,

sinonimie, literatuur. verwysings na enkele eksemplare. asook beknopte

taksonomiese en ekologiese aantekeninge.

Bydraes tot die FSA sal ook in Bothalia verskyn.

’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.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium). by/deur J.M. & H.M. Anderson.

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

A series devoted to the publication of monographs and major works on

southern African flora. Published: Vol. 14—19 (earlier volumes published

as Supplementary volumes to the Journal of South African Botany)

'n Reeks gewy aan die publikasie van monografiee en belangrike werke

oor flora van suidelike Afrika. Gepubliseer: vol. 14—19 (vroeere volumes

gepubliseer as Supplementary volumes van die Journal of South African

Botany).

BOTH ALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 23,2

Scientific EditorAVetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur: B.A. Momberg

N A T I O N A L

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

OctyOkt. 1993

Editorial Board/Redaksieraad

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

CONTENTS-INHOUD

Volume 23,2

1. Synopsis of the genus Brachylaena (Asteraceae) in southern Africa. S.S. CILLIERS 175

2. Nomenclatural changes and additions to the genus Ophioglossum in Africa (Ophioglossaceae:

Pteridophyta). J. E. BURROWS and T. J. EDWARDS ..." 185

3. Systematic studies in the genus Mohria (Anemiaceae: Pteridophyta). IV. Comparative gametophyte

morphology in Mohria and Anemia. J.P ROUX 191

4. Synopsis of the genus Disparago (Asteraceae). M. KOEKEMOER 197

5. Studies in the Marchantiales (Hepaticae) from southern Africa. 2. The genus Athalamia and A. spathysii\

the genus Oxymitra and O. cristata. S.M. PEROLD 207

6. Studies in the Marchantiales (Hepaticae) from southern Africa. 3. The genus Targionia and T. hypophylla

with notes on T. lorbeeriana and Cyathodium foetidissimum (Targioniaceae). S.M. PEROLD . . . 215

7. The hepatic, Jensenia spinosa (= Pallavicinia stephanii: Pallaviciniaceae), in southern Africa. S.M.

PEROLD 223

8. Notes on African plants:

Asclepiadaceae. Validation of the combination Aspidonepsis reenensis (Asclepiadaceae): the type

species of the subgenus Unguilobium. A. NICHOLAS and D.J. GOYDER 236

Curcurbitaceae. A new species of Kedrostis from the western Cape. P. BRUYNS 233

Sapotaceae. A ‘ Bequaertiodendron by any other name? C.L. BREDENKAMP and G.F. SMITH . 235

Vitaceae. A new species of Rhoicissus from the Transvaal. E. RETIEF 231

9. The vegetation of the southern Langeberg, Cape Province. 3. The plant communities of the Bergfontein,

Rooiwaterspruit and Phesantefontein areas. D.J. MCDONALD 239

Errata in Bothalia 23,1: 157: The vegetation of the southern Langeberg, Cape Province. 2. The plant

communities of the Marloth Nature Reserve, by D.J. McDonald 263

10. The efficient use of small plots in a fynbos phytosociological study in the northern Cederberg: a quick way

to collect plant-environmental data. P.J. MUSTART, E.J. MOLL and H.C. TAYLOR 265

11. The vegetation ecology of municipal Durban, Natal. Floristic classification. D.C. ROBERTS 271

12. National Botanical Institute, South Africa: list of staff and publications 327

13. Guide for authors to Bothalia 339

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2323unse_0

Bothalia 23,2: 175-184(1993)

Synopsis of the genus Brachylaena (Asteraceae) in southern Africa

S.S. CILLIERS*

Keywords: Asteraceae, Brachylaena, southern Africa, synopsis, taxonomy

ABSTRACT

A synopsis of the southern African representatives of the genus Brachylaena R. Br. (Asteraceae) is presented, in which seven

species and two subspecies are recognized. B. uniflora Harv. is placed in synonymy with B. discolor subsp. transvaalensis (Phill.

& Schweick.) J. Paiva and the division of subsp. discolor in two varieties is not accepted. Descriptions, synonymy, voucher

specimens, distribution maps, line drawings and keys to the species and subspecies are given.

UITTREKSEL

’n Sinopsis van die Suider-Afrikaanse verteenwoordigers van die genus Brachylaena R. Br. (Asteraceae) word aangebied

waarin sewe spesies en twee subspesies erken word. B. uniflora Harv. is in sinonimie met B. discolor subsp. transvaalensis

(Phill. & Schweick.) J. Paiva geplaas en die verdeling van subsp. discolor in twee varieteite. word nie aanvaar nie. Beskrywings,

sinonimie, verwysingseksemplare, verspreidingskaarte. lyndiagramme en sleutels tot die spesies en subspesies word voorsien.

INTRODUCTION

The latest taxonomic revision of all southern African

species of Brachylaena was published by Phillips &

Schweickerdt (1937). They recognized nine species in

South Africa, including a new species, B. transvaalensis

Phill. & Schweick., as well as B. discolor DC. and B.

uniflora Harv.

According to Hilliard & Burtt (1971) it is not always

possible to distinguish between B. discolor, B. uniflora

and B. transvaalensis on the basis of the characters de-

scribed by Phillips & Schweickerdt (1937). These three

species became known as the B. discolor-uniflora com-

plex, which was regarded as the only remaining taxo-

nomic problem in the genus.

In an effort to solve that problem, Paiva (1972) reduced

B. transvaalensis to subspecific rank, under B. discolor.

He also distinguished two varieties (var. mossambicensis

and var. discolor) in B. discolor subsp. discolor on the

basis of reproductive characteristics. Unfortunately he did

not study B. uniflora because it does not occur in his study

area, namely that of the Flora zambesiaca.

Hilliard (1977) did not agree with Paiva (1972) and

speculated that B. discolor and B. uniflora represented

merely the extremes in a wide range of head sizes within

the B. discolor-uniflora complex. According to Hilliard

(1977) the variation of the characteristics used by Paiva

(1972) as diagnostic, is continuous. Hilliard also men-

tioned that the variation can be interpreted in two ways:

a single species showing clinal variation in the number of

flowers in a head loosely linked to an ecological cline, or

two species which are hybridizing.

^Department of Plant and Soil Sciences, Potchefstroom University for

CHE, Potchefstroom 2520.

MS. received: 1991-06-19.

Gibbs Russell et al. (1987) followed the delimitation

proposed by Paiva (1972), and recognized eight species

and four infraspecific taxa in southern Africa, including

B. discolor DC. subsp. discolor with the varieties discolor

and mossambicensis J. Paiva; subsp. transvaalensis (Phill.

& Schweick.) J. Paiva; as well as B. uniflora Harv.

The views presented here are supported by a detailed

study of the vegetative and reproductive morphology and

the anatomy of leaves and stems of the southern African

representatives of Brachylaena (Cilliers 1990). Material

was gathered during extensive field studies and loaned

from all major South African and some overseas herbaria.

A brief review is given of the research on the mor-

phology of the heads of the B. discolor-uniflora complex.

The variation of certain quantitative characters was pre-

sented in the form of dice diagrams (Radford et al. 1974)

as shown in Figure 1.

The diagrams in Figures 2 & 3 show the following:

1, on the basis of four parameters (Figures 2A-C; 3A)

the four entities clearly fall into two groups with two

members each: a, the varieties of B. discolor subsp. dis-

color, and b, B. discolor subsp. transvaalensis and B. uni-

flora',

2, the two varieties of B. discolor subsp. discolor cannot

be separated on the basis of any of the eight parameters;

3, on the basis of six of the parameters shown, B. discolor

subsp. transvaalensis and B. uniflora cannot be separated;

the number of flowers both on male and female heads,

however, indicate a discontinuity between the two entities;

4, the parameters of the male heads shown are taxonom-

ically more useful than those of the female heads, which

tend to show continuous variation; the length of the female

involucre, however, clearly supports the recognition of the

two groups mentioned under 1.

176

Bothalia 23,2(1993)

FIGURE 1. — Dice diagram to show variation in certain quantitative

characters: where: x = mean, s.e.t = standard error, c = stan-

dard deviation, and r = range (Radford et al. 1974).

On this evidence, backed by other findings by Cilliers

(1990), I propose to sink B. uniflora under B. discolor

subsp. transvaalensis and not to recognize varieties under

B. discolor subsp. discolor.

TAXONOMIC TREATMENT

Brachylaena R. Br. in Transactions of the Linnean

Society of London 12: 115 (1817); DC.: 430 (1836);

Harv.: 116 (1865); Benth. & Hook.: 228 (1873); Engl. &

Prantl: 174 (1890); Phill. & Schweick.: 206 (1937); Paiva:

368 (1972); Hilliard: 105, 106 (1977). Type species: B.

neriifolia (L.) R. Br.

Oligocarpha Cass.: 22 (1817).

Trees or shrubs, dioecious. Leaves alternate, simple,

petiolate or subsessile, entire, toothed or crenate, some-

times 3-lobed at apex, mostly coriaceous, often white- or

rusty-tomentose abaxially. Heads in open or dense axillary

or terminal racemes or panicles, rarely solitary; unisexual,

rarely with fertile hermaphrodite flowers; 1 — 30(— 50) flow-

ers per head; involucre oblong, ovoid or campanulate,

bracts in 3-7 rows, free, dry, inner ones progressively

longer. Receptacle epaleate, honeycombed. Male flowers:

corolla tubular, 3-5-lobed; anthers tailed at base, exserted;

style filiform, sometimes thickened above, simple or bifid,

branches very short, flat, acute or obtuse; ovary usually

abortive, pubescent; pappus poorly developed, of scabrid

bristles, uniseriate. Female flowers: corolla tubular, 5-

lobed; staminodes occasionally present; style filiform,

bifid, branches very short, flat, acute or obtuse. Achenes

4—5-angled, pubescent or subglabrous, glandular.

The genus consists of twelve species, confined to Af-

rica and the Mascarene Islands. Seven species are found

in southern Africa.

KEY TO SPECIES

la Leaves glabrous abaxially. if tomentose then rusty brown; male flowers with an abortive ovary (0. 7-1.9 mm long);

female flowers with staminodes (0. 6-2.0 mm long):

2a Leaf base decurrent into petiole, petiole often up to 8 mm (rarely up to 10 mm) long, not clearly distinct from leaf

base; leaf lamina 5-10 (rarely 4) times as long as broad 1.6. neriifolia

2b Leaf base acute, petiole mostly longer than 1 0 mm, clearly distinct from leaf base; leaf lamina 1 -4 times as long

as broad 2. 6. glabra

lb Leaves white-tomentose abaxially, never glabrous; ovary in male flowers, if present, shorter than 0.6 mm;

staminodes in female flowers, if present, shorter than 0.6 mm:

3a Heads in dense axillary racemes or solitary; usually less than six heads per raceme; leaf apex usually mucronate;

anthers usually shorter than 1.9 mm:

4a Involucral bracts glabrous to white-tomentulose, in 5-8 series; leaves mostly oblong; petiole up to 3 mm, rarely

up to 5 mm long; heads sometimes solitary 3.6. ilicifolia

4b Involucral bracts white-tomentose, never glabrous, in 3 or 4, rarely 5 series; leaves mostly obovate to

oblanceolate, never oblong; petiole usually longer than 5 mm; heads never solitary 4. 6. huillensis

3b Heads in open axillary or terminal racemes or panicles; usually more than eight heads per raceme or panicle; leaf

apex rarely mucronate; anthers up to 3.7 mm long:

5a Leaf apex 3-lobed or 3-toothed (leaves from coppice growth sometimes without lobes or teeth); involucral bracts

in up to 5 series in male heads, in up to 6 series in female heads 5. 6. elliptica

5b Leaf apex not 3-lobed or 3-toothed; involucral bracts in male heads in 4—10 series, in female heads in 5-10

series:

6a Leaf base obtuse to rounded; leaves mostly elliptic; outer bark dark brown to black 6. 6. rotundata

6b Leaf base decurrent into a petiole; leaves mostly oblanceolate to narrowly obovate; outer bark light grey or

light brown 7. 6. discolor

1 . Brachylaena neriifolia (L.) R. Br. in Transactions

of the Linnean Society of London 12: 115 (1817); Less.:

208 (1832); DC.: 430 (1836); Drege: 169 (1843); Sch.

Bip.: 671 (1844); Harv.: 116(1865); Bolus & Wolley-Dod

14: 277 (1904); Sim: 246 (1907); Juel: 381 (1918); Sim:

44 (1921); Phill. & Schweick.: 207 (1937). Type: Aethio-

pia |South Africa], LINN 992.2, lecto.-PRE, photo.!

Baccharis neriifolia L.: 860 (1753); Willd.: 1904 (1804); Pers.: 423

(1807); Spreng.: 462 (1826); Steud.: 178 (1841). Oligocarpha neriifolia

(L.) Cass.: 22 (1817). Conyza neriifolia (L.) L’Herit. ex Steud.: 414

(1841).

Tarchonanthus lanceolatus Thunb.: 145 (1800); Willd.: 1793 (1804);

Pers.: 405 (1807); Thunb.: 638 (1823); Spreng.: 456(1826). Type: Cape,

Thunberg s.n. (UPS, holo.-PRE (18917), photo.!).

Multistemmed shrubs or small trees (2— )4— 6(— 8) m

high; bark dark grey to brown, longitudinally fissured;

young branches with rusty brown down, glabrescent.

Leaves petiolate, oblanceolate to narrowly oblanceolate,

sometimes very narrowly elliptic or lorate, (20— )45— 110

(-180) x (5—) 1 0— 20(— 25) mm; coriaceous, glabrous adaxi-

ally, glabrous or rusty brown-tomentose abaxially; mar-

gins entire or serrated on coppice shoots, base decurrent

in a short petiole, rarely acute; petiole (3-)5-8 (-10) mm

long. Heads many, in open axillary or terminal racemes

or panicles; peduncles of male heads 1 .5— 3.0(— 5.0) mm,

and of female heads 1. 8-3.0 mm long. Involucral bracts

in 4-7 series, 2.5-3. 0(-5.0) x 2.0— 3.5(— 5.0) mm; individ-

ual bracts very narrowly to very broadly ovate, or nar-

rowly to widely elliptic, rusty brown. Flowers 8-13 in

Bothalia 23,2(1993)

177

D r

M -

T -

U -

^ I 2 3456 789 10

MALE HEADS: NUMBER OF SERIES OF

INVOLUCRAL BRACTS

INVOLUCRAL BRACTS

D

M

T

U

12 3 4 5

C MALE HEADS: PEDUNCLE LENGTH (mm)

D

M

T

u

-^3-

c

10 20 30

MALE HEADS: NUMBER OF FLOWERS

40

D

M

J[

U

12 3 4 5 6

D FEMALE HEADS: PEDUNCLE LENGTH (mm)

D

M

T

U

10

20

30

FEMALE HEADS: NUMBER OF FLOWERS

FIGURE 2. — Dice diagrams of all studied material of the B. discolor-

uniflora complex. Variation in involucral length (mm): A,

male, and B, female heads; variation in peduncle length (mm)

of C, male and D, female heads. D = B. discolor subsp. dis-

color var. discolor. ; M = B. discolor subsp. discolor var.

mossambicensis; T = B. discolor subsp. transvaalensis ; and U

= B. uniflora.

FIGURE 3. — Dice diagrams of all studied material of the B.

discolor-uniflora complex. Variation in number of se-

ries of involucral bracts: A, male, and B. female

heads; variation in number of flowers of C, male and

D, female heads. D = B. discolor subsp. discolor var.

discolor. M = B. discolor subsp. discolor var.

mossambicensis; T = B. discolor subsp. transvaalen-

sis; and U = B. uniflora.

178

Bothalia 23,2 (1993)

FIGURE 4. — Brachylaena neriifolia (L.) R. Br. A, branch with leaves

and female heads, Cilliers 69: B, male head, Cilliers 73: C, fe-

male head, Cilliers 65. Scale bars: 10 mm.

male and 7-11 in female heads. Pappus 3-5 mm long.

Ovary in male flowers 0.7-1 .1 mm long, pubescent, rarely

absent, sometimes fertile. Staminodes in female flowers,

0.6-1 .0 mm. Achene oblanceolate to narrowly elliptic, 1-2

x 0.5-1. 0 mm, pubescent glandular. Flowering time: De-

cember to February. Figure 4.

Leaves from coppice growth of B. neriifolia may some-

times be confused with leaves of B. glabra. The species,

however, can usually be distinguished by the features

given in the species key.

Distribution and habitat: B. neriifolia occurs in the

southern Cape, extending from Giftberg'and Piketberg in

the north to the Storms River in the east (Figure 5). It is

common along streambanks on mountains and in forests.

Vouchers: Barker 2955 (NBG); Cilliers 72 (PUC); Orchard 543 (STE);

Paterson 1267 (GRA ); Jacot Guillarmod 8700 (GRA).

2. Brachylaena glabra (Lf) Druce in Report of

the Botanical Society and Exchange Club of the British

Isles 1916: 611 (1917); Phill. & Schweick.: 209 (1937);

Hilliard: 106 (1977). Type: Cape, Thunberg s.n. (UPS,

holo.-PRE (microfiche 18914!).

Tarchonanthus glaberL.i.: 360 ( 1781 ); Thunb.: 638(1823).

T. dentatus Thunb.: 145 (1800); Willd.: 1793 (1804); Pers.: 405 (1807);

Thunb.: 638 (1823); Less.: 208 (1832). B. dentata (Thunb.) Harv.: 116

(1865); Sim: 246 (1907); Juel: 381 (1918); Sim: 44 (1921); Bews: 215

(1921); Henkel: 72 (1934); non DC. (1836). Type: Cape, Thunberg s.n.

(UPS, holo.-PRE (18915), photo.!).

B. grandifolia DC.: 430 (1836); Drege: 136(1843). Type: Zuurbergen,

Drege 2122 (G-DC, holo.!; K!).

Single-stemmed trees to multistemmed shrubs (2-)

5-20 m high; bark light grey to brown, not fissured; young

branches with rusty brown down, glabrescent. Leaves pet-

iolate, narrowly obovate to oblanceolate, sometimes nar-

rowly elliptic, (30— )90— 1 30(— 225) x (12-) 20-60(-90)

mm, coriaceous, glabrous adaxially, glabrous or rusty

brown-tomentose abaxially, margins entire or serrated in

upper half and on coppice shoots, base acute; petiole (8-)

10— 15(— 27) mm long. Heads many, in open axillary ra-

cemes or terminal panicles or racemes; peduncles of male

heads (2.5-)4.0-6.0 mm, and of female heads 1-3 mm

long. Involucral bracts in 4-7 series, 2.3-5.0(-7.0) x 0.6-

4.0(— 6.0) mm; individual bracts very narrowly to very

broadly ovate, rusty brown. Flowers 13-24 in male heads,

3— 10(— 22) in female heads. Pappus 3-6 mm long. Ovary

in male flowers 1.3-1. 9 mm long, oblong, rarely absent,

sometimes fertile. Staminodes in female flowers 0.8-2.0

mm long. Achene narrowly obovate to oblanceolate, 1 .3-

3.8 x 0.7-1. 4 mm, pubescent glandular. Flowering time:

March and April. Figure 6.

Distribution and habitat: B. glabra occurs mainly

along the eastern Cape coast (Figure 5), in open spaces

of high forest. At its northern limit in Natal (southeastern

parts) (Figure 5) it forms a small tree or a shrub, mainly

on south-facing sandstone outcrops and escarpment edges

near the coast.

Vouchers: Abbott 1192 (NH); Cilliers 63. 133 (PUC); Fourcade 537

(BOL.GRA, PRE), 4499(BOL); Ward 197 (NU); West267(BOL, GRA.

PEU).

3. Brachylaena ilicifolia (Lam.) Phill. & Schweick.

in Bothalia 3: 212 (1937); Hilliard: 108 (1977). Type:

Cape, Sonnerat s.n. (K, iso.!).

Baccharis ilicifolia Lam.: 345 (1783); Pers.: 423 (1807); Steud.: 177

(1841).

FIGURE 5. — Known geographic distribution of Brachylaena neriifolia.

•; and B. glabra. ▲.

Bothalia 23,2 (1993)

179

FIGURE 6. — Brachylaena glabra (L.f.) Druce. A, branch with leaves

and female heads, Thode A860', B, male head, Cilliers 133\ C,

female head, Cilliers 63. Scale bars: 10 mm.

Tarchonanthus racemosus Thunb.: 145 (1800); Thunb.: 638 (1823);

Less.: 208 (1832). Brachylaena racemosa (Thunb.) DC.: 430 (1836);

Harv.: 1 16 (1865); Sim: 246 ( 1907); Wood: 170 ( 1908); Juel: 381 ( 1918);

Sim: 44 (1921); Bews: 215 (1921). Type: Cape. Thunberg s.n. (UPS.

holo.-PRE (microfiche 18919!).

B. elliptica auct., non (Thunb.) DC.: 430 (1836) quoad descr. et spec.

Multistemmed shrubs or small trees ( 1— )3 — 4(— 6) m

high; bark dark grey to brown, fissured; young branches

with white down, glabrescent. Leaves short-petiolate to

subsessile, oblong to narrowly oblong, sometimes nar-

rowly elliptic to elliptic or lanceolate to narrowly ovate,

( 10— )25— 45(— 75) x (4— )10— 15(— 30) mm, chartaceous, gla-

brous adaxially, white-tomentose and glandular abaxially,

apex acute or obtuse to rounded, usually mucronate, mar-

gin entire or dentate, base acute to obtuse, sometimes

rounded; petiole ( 0.5—) 1 .0— 3.0(— 5.0) mm long. Heads few,

in dense axillary racemes, sometimes solitary; peduncles

of male heads 0.5-1. 0(-2.0) mm and of female heads 0.8-

1 ,5(— 2.0) mm long. Involucral bracts in 5-8 series, 2.5-

5.0C-9.0) x 1 .5— 3.0( — 4.0) mm; individual bracts narrowly

to very widely ovate, sometimes narrowly to widely el-

liptic, glabrous to white tomentulose. Flowers 9-1 1 (-18)

in male and 8-12 in female heads. Pappus 2-5 mm long.

Ovary in male flowers, if present, up to 0.6 mm long.

Staminodes in female flowers, if present, up to 0.2 mm

long. Achene oblanceolate or narrowly elliptic, 2. 1-3.6 x

0.5-1. 3 mm, pubescent glandular. Flowering time'. August

and September. Figure 7.

Distribution and habitat'. B. ilicifolia occurs in scrub

in the drier parts of northeastern Transvaal, northern and

central Natal, and in the eastern Cape (Figure 8). It grows

in dry river valleys and on dry hillslopes.

Vouchers: Bayliss 5052 (GRA); Cilliers 53, 147 (PUC); Dahlstrand

1995( GRA, PRE); Feely 62 (NH. PRE, UN); Van Wyk5246tPRE. PRU).

4. Brachylaena huillensis O. Hoffm. in Botanische

Jahrbiicher 32: 149 (1902); Phill. & Schweick.: 212

(1937); Wild; 124 (1969); Paiva: 369 (1972); Hilliard: 107

(1977). Type: Angola, Huilla, Antunes 121 (K, iso.!).

FIGURE 7. — Brachylaena ilicifolia (Lam.) Phill. & Schweick. A.

branch with leaves and male heads, Bayliss 5052', B. male

head. Bayliss 5052\ C, female head. Cilliers 53. Scale bars: 10

mm.

180

Bothalia 23,2 (1993)

FIGURE 8. — Known geographic distribution of Brachvlaena ilicifolia ;

•; and B. huillensis, ▲, within southern Africa.

Tarchonanthus camphoratus auct. non L.: Hiem: 554 ( 1898); Gossw. &

Mendon^a: 121-123 ( 1939), Welwitsch 6745 (could not be found).

B. hutchinsii Hutch.: 126 (1910); Brenan & Greenway: 149 (1949);

Eggeling: 95 (1951); Dale & Greenway: 155 (1961); Cufod.: 1091

(1966). Type: Nairobi, Hutchins s.n. (K, holo. !); Kenya, Nairobi,

Battiscombe 27 & 54 (K, para.!).

Single-stemmed medium to large trees (5-)6-10(-15)

m high; bark light grey to brown, deeply fissured; young

branches with white down. Leaves petiolate, narrowly ob-

ovate to oblanceolate, sometimes narrowly elliptic to ellip-

tic, (26— )45— 75(— 1 25) x (8—) 1 6— 30(— 50) mm, coriaceous,

glabrous adaxially, glandular and white-tomentose abaxi-

ally, apex acute, usually mucronate, margin entire, some-

times sinuate in front part or irregularly dentate, base

acute, sometimes decurrent; petiole (3— )5— 8 (-12) mm

long. Heads few, in dense axillary racemes; peduncles of

male heads 0.5— 1 .0(— 2.0 ) mm and of female heads 0.5- 1.0

mm long. Involucral bracts in 3-5 series, 2-4(-6) x 1.5-

3.0(— 5.0) mm, individual bracts ovate to very widely

ovate, sometimes narrowly to widely elliptic, white-to-

mentose. Flowers 6-9(-10) in male and 5 in female heads.

Pappus ( 1 .5— )3.0— 4.0(— 6.0) mm long. Ovary in male

flowers, if present, up to 0.3 mm long. Staminodes in fe-

male flowers, if present, up to 0.4 mm long. Achene ob-

lanceolate or narrowly elliptic, 3.5— 3.7(— 4.0) x 1.0-1. 3

mm, pubescent glandular. Flowering time ; July. Figure 9.

Distribution and habitat : within southern Africa B.

huillensis occurs in the northern parts of Transvaal, in

Zululand and Natal (Figure 8). It is confined to short for-

est, woodland and bush, usually on sandy soils.

Vouchers: Cilliers 79, 100, 104 (PUC); Lang s.nl (BOL, NBG); Moll &

Strey 3755 (NH); Pooley 883 (UN); Ward 7119 (NH).

5. Brachylaena elliptica (Thunb.) DC. in Pro-

dromus systematis naturalis regni vegetabilis 5; 430

(1836), quoad basionym excl. spec, et descr.; Harv.: 116

(1865); Sim: 92 (1907); Wood: 169 (1908); Juel: 381

(1918); Sim: 44 (1921); Bews: 215 (1921); Henkel: 72

(1934); Phill. & Schweick.: 216 (1937). Type: Cape,

Thunberg s.n. (UPS, holo.-PRE (microfiche 18916!).

Tarchonanthus ellipticus Thunb.: 145 (1800); Willd.: 1793 (1804);

Pers.: 405 (1807); Thunb.: 638 (1823); Spreng.: 456 (1826) excl. syn.;

Less.: 208(1832).

B. dentate DC.: 430 (1836), non (Thunb.) Harv.; Type: Zuurbergen,

Dr'ege 3659 (G-DC, lecto.!, here designated); Albany, Burchell 3400

(G-DC!); Kafffaria, Ecklon 791. 793 (G-DC!).

Multistemmed shrubs or small trees (2— )4— 6(— 8) m

high; bark dark grey to brown, fissured; young branches

with white down, glabrescent. Leaves petiolate, widely to

narrowly obovate or narrowly oblanceolate, sometimes

narrowly elliptic to oblong, (21— )40— 70(— 76) x (6—) 1 5—30

(-62) mm, chartaceous, glabrous adaxially, glandular and

white-tomentose abaxially, apex acute or obtuse, some-

times mucronate, usually 3-lobed, margins entire or den-

tate, base acute, rarely obtuse; petiole (l-)3^f(-6) mm

long. Heads many, in open axillary or terminal racemes

FIGURE 9. — Brachylaena huillensis O. Hoffm. A, branch with leaves

and male heads, Cilliers 104\ B, male head, Cilliers 104 ; C, fe-

male head. Ward 7119. Scale bars: 10 mm.

Bothalia 23,2(1993)

181

FIGURE 10. — Brachylaena elliptica (Thunb.) DC. A, branch with

leaves and male heads, Bayliss 8747', B, male head, Bayliss

8747', C, female head, Cilliers 38. Scale bars: 10 mm.

or panicles; peduncles of male heads 0.5-1. 0 mm and of

female heads 0.5-1. 5 mm long. Involucral bracts in 3-6

series, 1 .5— 3.0(— 5 .0) x 0.5— 1 .5 (— 2.0 ) mm; individual

bracts narrowly to very widely ovate, sometimes lanceo-

late, white tomentulose. Flowers (4 — )7— 1 1(— 15) in male

and 4-10 in female heads. Pappus 1.5-2.5(-4.0) mm long.

Ovaries in male flowers, if present, up to 0.4 mm long.

Staminodes in female flowers, if present, up to 0.8 mm

long. Achene narrowly obovate to narrowly oblanceolate,

2. 1-3.2 x 0.5-1. 0 mm, pubescent glandular. Flowering

time : April, May and June. Figure 10.

The leaves of B. elliptica are very variable in their

shape and often conspicuously 3-lobed or 3-toothed at the

apex; they sometimes resemble those of B. glabra, but

are shorter, not as wide and white-tomentose (never gla-

brous) abaxially.

Distribution and habitat : B. elliptica is widespread in

the eastern Cape from Uitenhage eastwards through Trans-

kei to Natal and Zululand (Figure 11). It can be found in

coastal and river bush, valley bushveld, in scrub or on

grassy slopes, often on rocky ridges.

Vouchers: Cilliers 27, 38, 50 (PUC); MacOwan 244 (NH); Olivier 2141

(PEU); Osborne 232 (GRA); Strey 6699 (NH, PRE).

6. Brachylaena rotundata S. Moore in Journal of

Botany: 131 (1903); Burtt Davy & Pott-Leend.: 168

(1912); Eyles: 508 (1916); Sim: 44 (1921); Phill. &

Schweick.: 218 (1937); Wild: 123 (1969); Paiva: 372-375

(1972). Type: Transvaal, north escarpment of Witwaters-

rand series, Rand 738 (BM, holo. ! ).

B. rhodesiana S. Moore 37: 448 (1906); S. Moore: 108 (1911 ); Eyles:

508 (1916). Type: Rhodesia, Matopo hills, Gibbs 72 (BM, holo.!; K,

iso.!).

B. discolor auct. non DC.: Munro: 90 ( 1908); Eyles: 508 (1916).

Single-stemmed large shrubs or medium-large trees

(3— )5— 8(— 1 5 ) m high; bark dark brown to black, deeply

fissured; young branches white-tomentose, glabrescent.

Leaves petiolate, narrowly to widely elliptic, sometimes

narrowly obovate or oblong, (25— )50— 100(— 1 60) x (15-)

30-50(-90) mm, coriaceous, glabrous to white-tomentose

adaxially, white-tomentose abaxially, glandular, apex

acute to obtuse, sometimes rounded, margins entire or cre-

nate smoothly rounded, sometimes dentate in coppice

shoots, base obtuse to rounded; petiole ( 1— )3— 7(— 10) mm

long. Heads many, in open axillary or terminal racemes

or panicles, peduncles of male heads 1 .0—1 .5(— 2.0) mm

and of female heads 0.5-3 .0 mm long. Involucral bracts

in 4-10 series, 3— 6(— 8) x 2— 3(— 5) mm; individual bracts

ovate to very widely ovate, sometimes lanceolate, white-

tomentulose. Flowers 1 3— 28(— 3 1 ) in male and 11-27 in

female heads. Pappus 3.0-6.5 mm long. Ovaries absent

in male flowers. Staminodes absent in female flowers.

Achene narrowly elliptic, 1.6-3. 8 x 0.5- 1.0 mm, pubes-

cent glandular. Flowering time : August and September.

Figure 12.

Distribution and habitat : within southern Africa B.

rotundata occurs in the northeastern, southern and western

parts of Transvaal and occasionally on the Free State side

of the Vaal River (Figure 11). It is common on rocky

ridges and hills and the edges of dry kloof bush.

Vouchers: Adlam & Wood 5633 (NH); Botha & Ubbink 1556 (PUC,

PRE); Cilliers 149, 151 (PUC); Leendertz 235 (GRA, KMG. NH); The-

ron 1557 (PRE, PRU).

FIGURE 1 1 . — Known geographic distribution of Brachylaena elliptica,

•; and B. rotundata, ▲, within southern Africa.

182

Bothalia 23,2 (1993)

FIGURE 12. — Brachylaena rotundata S. Moore. A, branch with leaves

and female heads, Cilliers 149: B, male head, Cilliers 151\ C,

female head, Cilliers 150. Scale bars: 10 mm.

7. Brachylaena discolor DC. Prodromus systematis

naturalis regni vegetabilis 5: 430 (1836); Drege: 155, 157

(1843); Harv.: 117 (1865); Wood & Evans: 23, 24 (1898);

Sim: 247 (1907); Wood: 169 (1908); Sim: 44 (1921);

Bews: 215 (1921); Henkel: 72 (1934); Phill. & Schweick.:

219 (1937); Macnae & Kalk: 154 (1958); Wild: 123

(1969); Hilliard & Burtt: 3 (1971); Paiva: 369 (1972);

Hilliard: 109 (1977). Type: Cape, Uitenhage District, Bur-

chell 3751 (G-DC, lecto.!, here designated); Uitenhage,

Ecklon 1422 (G-DC!); between the Umzimkulu and Um-

tentu Rivers, Drege 5043 (G-DC!); Mozambique, Delagoa

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

Multistemmed shrubs or small to large trees (2-)6-10

(-20) m high; bark light grey to brown, fissured; young

branches with white down, glabrescent. Leaves petiolate,

oblanceolate to narrowly obovate, sometimes elliptic to

narrowly elliptic; (50— )70— 100(— 1 90) x (15-) 25-45(-60)

mm, glabrous adaxially, white-tomentose abaxially, glan-

dular, apex acute, sometimes obtuse, rarely rounded, mar-

gin entire, sinuated or serrated over whole margin or in

upper half and in coppice shoots, base decurrent into a

petiole; petiole (3— )8— 10(— 12) mm long. Heads many, in

open axillary or terminal racemes or panicles, peduncles

of male heads 0.5-3.0(-5.0) mm and of female heads 0.4-

4.0 mm long. Involucral bracts in 4—10 series, ( 1 .5— )3.0—

1 0.0(— 1 5.0) x (0.5-)2.0-4.0 (-6.0) mm; individual bracts

narrowly to very widely ovate, sometimes elliptic to nar-

rowly elliptic, white-tomentulose to tomentose. Flowers

1-45 in male and 1-26 in female heads. Pappus 2.0-8. 5

mm long. Ovary in male flowers, if present, up to 0.5

FIGURE 13. — Brachylaena discolor DC. subsp. discolor. A, branch

with leaves and female heads, Harrison 29: B, male head,

Cilliers 135: C, female head, Cilliers 109. Scale bars: 10 mm.

mm long. Staminodes in female flowers, if present, up to

0.7 mm long. Achene narrowly obovate to linear, 1.0-3. 9

x 0.4—1. 0 mm, pubescent glandular. Flowering time : July,

August and September.

B. discolor can usually be distinguished from the other

species by the leaf base which is mostly decurrent into

the petiole.

Distribution and habitat: B. discolor, one of the most

widely distributed species in the genus, occurs in northern

and eastern Transvaal, through Swaziland and the eastern

FIGURE 14. — Known geographic distribution of Brachylaena discolor

subsp. discolor within southern Africa.

Bothalia 23,2 (1993)

183

FIGURE 15. — Brachylaena discolor DC. subsp. transvaalensis (Phill.

& Schweick.) J. Paiva. A, branch with leaves and female

heads, Cilliers 134; B. male head, Cilliers 137; C, female head,

Cilliers 144. Scale bars: 10 mm.

parts of Natal and Transkei up to the mouth of the Bush-

mans River in the eastern Cape. Its habitat varies from

inland forests on valley slopes and plateaux to coastal for-

ests, scrub and dune bush.

Key to subspecies

In 10-15% of the specimens investigated, the values of

the characters used in this key overlapped.

la Involucral bracts of male heads in 7-10 (rarely 6) series;

peduncle of male heads up to 5 mm long; involucral bracts

of female heads in 6-10 series; peduncle of female heads

\-4 mm long; involucral bracts up to 12 mm long in male

heads and up to 15 mm long in female heads

7a. subsp. discolor

lb Involucral bracts of male heads in 4-6 series; peduncle of male

heads up to 1 mm long; involucral bracts of female heads

in 5-7 series; peduncle of female heads 0.4— 1.5 mm long;

involucral bracts up to 4 mm long in male heads and up to

6 mm long in female heads 7b. subsp. transvaalensis

7a. subsp. discolor

J. Paiva in Boletim da Sociedade Broteriana 46: 369

(1972).

B. natalensis Sch. Bip.: 972 (1843); Sch. Bip.: 671 (1844); Harv.: 117

(1865). Type: prope Natal-Bay [Durban], Krauss 243 (G, holo.!).

B. discolor subsp. discolor var. mossambicensis J. Paiva 46: 369 ( 1972).

Type: Nhacoongo, 90 km south of Inhambane, Gomes & Sousa 1667

(COI, holo.!).

B. rhodesiana auct. non S. Moore: S. Moore: 108 (1911) quoad specim.

Swynnerton 6515.

Multistemmed shrubs or small trees, usually not higher

than 6-8 m. Male heads with involucral bracts in 7-10

series, involucre 3—1 Of— 1 2) mm long; peduncles 1-5

mm long; 1 1-39 flowers per head. Female heads with

involucral bracts in 6-10 series, involucre 6.0— 13.5(— 1 5.0)

mm long; peduncles 1-4 mm long; 9-26 flowers per head.

Figure 13.

Distribution and habitat: B. discolor subsp. discolor is

a common coastal species (with an occasional inland oc-

currence, e.g. Hluhluwe Game Reserve) found from the

mouth of the Bushmans River in the eastern Cape to

Transkei, Natal and Zululand (Figure 14).

Vouchers: Cilliers 108. 109. 135 (PUC); Flanagan 861 (NBG);

Muirhead K17 (RUH); Schweickerdt 1384 (NH, STE); Strey 8751. 8753.

8783. 8793, 8795, 8798 (NH).

7b. subsp. transvaalensis (Phill. & Schweick.) J.

Paiva in Boletim da Sociedade Broteriana 46: 369 (1972).

B. transvaalensis Phill. & Schweick.: 214 (1937); Hilliard & Burtt: 3

(1971). Type: South Africa, Transvaal, Pietersburg. between Houtbos and

Haenertsburg, Hutchins s.n. (K, holo.!).

B. uniflora Harv.: 117 (1865); Bews: 215 (1921); Phill. & Schweick.:

211 (1937); Hilliard & Bum: 3 (1971); Hilliard: 109 (1977). Type: Natal,

on mountains, 30-60 miles from the sea, Sutherland s.n. (K, holo.!).

Single-stemmed medium-large to large trees, not

higher than 20 m. Male heads with involucral bracts in

4--6 series, involucre 1-4 mm long; peduncles 0.5-1. 2 mm

long; 1-14 flowers per head. Female heads with involu-

cral bracts in 5-7 series, involucre 3-6 mm long; pedun-

cles 0.4-1. 5 mm long; 1-11 flowers per head. Figure 15.

Distribution and habitat: within southern Africa B. dis-

color subsp. transvaalensis occurs in the northern and

eastern Transvaal and in Swaziland, as well as in the

northeastern and southern parts of Natal and the Transkei

(Figure 16). It can be found in inland and coastal forests,

but not in dune bush or forests directly facing the sea.

FIGURE 16. — Known geographic distribution of Brachylaena discolor

subsp. transvaalensis.

184

Bothalia 23,2 ( 1993)

Vouchers: Cilliers 120, 137, 144 (PUC); Compton 29073 (NBG); Keet

1130 (STE); Schrire, Van Wvk & Abbott 1782 (NH); Strey 8752, 8819,

8820 (NH).

ACKNOWLEDGEMENTS

The financial support by the Council for Scientific and

Industrial Research, Pretoria and the Potchefstroom Uni-

versity for Christian Higher Education is gratefully ac-

knowledged. Thanks are also due to Dr O.A. Leistner

(National Botanical Institute, Pretoria) and Dr G.F. Smith

(formerly Potchefstroom University for Christian Higher

Education) for their constructive comments on the text,

and the directors and staff of the cited herbaria for the

loan of specimens.

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Bothalia 23,2: 185—190(1993)

Nomenclatural changes and additions to the genus Ophioglossum in Africa

(Ophioglossaceae: Pteridophyta)

J.E. BURROWS* and T.J. EDWARDS**

Keywords: Africa, new combination, new species, Ophioglossum, Pteridophyta, taxonomy

ABSTRACT

Recent research has necessitated a review of the nomenclature of the genus Ophioglossum in Africa, This paper establishes

a new species, O. caroticaule J. E, Burrows, a new subspecies, O. vulgatum L. subsp. africanum Pocock ex J. E. Burrows, and

a change in status, O. latifolium (Prantl) J.E. Burrows (basionym: O. gomezianum var. latifolium Prantl). Lectotypification is

provided for O. gracillimum Welw. ex Hook. & Bak., O. lusoafricanum Welw. ex Prantl, O. latifolium (Prantl) J.E. Burrows, and

O. rubellum Welw. ex A. Braun. Nomenclatural problems in O. lancifolium Presl are discussed.

UITTREKSEL

Onlangse navorsing het ’n hersiening van die nomenklatuur van die genus Ophioglossum in Afrika noodsaaklik gemaak. 'n

Nuwe spesie, O. caroticaule J.E. Burrows, ’n nuwe subspesie, O. vulgatum , L. subsp. africanum Pocock ex J.E. Burrows, en 'n

verandering in status, O. latifolium (Prantl) J.E. Burrows (basioniem: O. gomezianum var. latifolium Prantl), word in hierdie

artikel beskryf. Lektotipifikasie word verskaf vir O. gracillimum Welw. ex Hook. & Bak., O. lusoafricanum Welw. ex Prantl, O.

latifolium (Prantl) J.E. Burrows, en O. rubellum Welw. ex A. Braun. Nomenklatoriese probleme in O. lancifolium word

bespreek.

INTRODUCTION

The genus Ophioglossum L. has received no taxonomic

attention in Africa since Clausen’s (1938) confusing

monograph of the genus. The simple morphological struc-

ture of the plants together with apparently considerable

intraspecific variation has caused compilers of African re-

gional floras to either follow Clausen’s nomenclature or

assume a conservative stance by maintaining broadly cir-

cumscribed species which, in the light of recent studies,

are often composed of two or more clearly distinct taxa.

The most important development since Clausen’s work

is the publication by Wagner & Wagner (1983) of their

'genus communities’ concept, based upon work on the

genus Botrychium Swartz (Ophioglossaceae) in North

America. This concept made use of the fact that Botrych-

ium (like Ophioglossum) frequently grows together in

multiple species communities and, when this occurs, one

is able to make rational comparisons between taxa occur-

ring within a community, particularly if these differences

are maintained in communities found elsewhere under dif-

ferent edaphic and climatic conditions. This concept has

provided taxonomists with a valuable additional tool with

which to distinguish between environmentally-induced

variation and differences that are genetically controlled.

During recent studies, the senior author has discovered

an unique area in northwestern Zimbabwe which supports

1 1 species of Ophioglossum. Additional genus communi-

* RO. Box 710, Lydenburg, 1120.

** UN/FRD Research Unit for Plant Growth and Development, Depart-

ment of Botany, University of Natal, P.O. Box 375, Pietermaritzburg,

3200.

MS. received: 1992-04-24.

ties have since been located in both Zimbabwe and South

Africa which have made a better delimitation and defini-

tion of the southern African species of Ophioglossum pos-

sible. However, these studies have resulted in the need to

revise the nomenclature of several of the taxa occurring

on the African subcontinent.

DISCUSSION

In 1983, the senior author collected nine species of

Ophioglossum growing within an area of about six square

kilometres in the Sengwa Wildlife Research Area of north-

western Zimbabwe. Subsequent collecting at Sengwa has

yielded a further two species, bringing the total to eleven,

a situation not recorded to date elsewhere in the world for

the genus Ophioglossum. Additional genus communities

have been discovered by the senior author on Ngoma-

kurira, northeastern Zimbabwe (6 species), Elim Hospital,

eastern Zimbabwe (4 species), Treur River, eastern Trans-

vaal (5 species) and several communities composed of

two or three species each. These localities occur on var-

ious substrates (sandstone, granite, quartzite and dolomite)

and under varying climatic conditions, and the consistent

differences displayed between members of these genus

communities have allowed for clearer definitions to be

established between taxa.

However, the specific concepts within the African

members of Ophioglossum entrenched by Tardieu-Blot

(1953, 1964) and Schelpe (1970, 1977) do not adequately

cover the number of taxa that are evident from genus com-

munity studies. This was noted by Dr Mary Pocock of

Grahamstown who spent much time studying the South

African species of Ophioglossum. She recorded her con-

cepts in a manuscript which unfortunately was never pub-

186

Bothalia 23,2 (1993)

lished. The author’s studies corroborate some of Pocock’s

species concepts and one of her taxa is published below.

Spore morphology is a valuable taxonomic tool within

the southern African species of Ophioglossum even

though it cannot be readily employed for rapid determi-

nations in herbaria.

1. The Ophioglossum lancifolium aggregate

Key to species

la Trophophore linear with a length:width ratio >10:1; tropho-

phore frequently falcate O. gracillimum

lb Trophophore narrowly lanceolate to narrowly elliptic with a

length:width ratio <10: 1 ; trophophore not falcate:

2a Old rhizomes strongly linear, with a length:width ratio >6: 1 ;

spore wall lurnina distinct, muri clearly reticulate ....

O. lusoafricanum

2b Old rhizomes short or ellipsoid, with a length:width ratio

<6: 1 ; spore wall lurnina reduced to conical pits:

3a Trophophore linear-elliptic, length: width ratio >6:1; rhi-

zome ellipsoid O. caroticaule

3b Trophophore elliptic, length:width ratio <6:1; rhizome

linear sp. aff. O. lancifolium

1.1. Ophioglossum caroticaule J.E. Burrows , sp.

nov., ab O. gracillimo sports aletis et trophophoris majori-

bus et latioribus differt; etiam O. lusoafricano similis sed

rhizomate brevi et latiore, trophophoro apiculato et sports

lurnina parva et conica ostendentibus differt.

TYPUS. — Zimbabwe: Gokwe Dist., Sengwa Wildlife

Research Area, Leguaan Vlei, 880 m, 11-2-1991, J.E. &

S.M. Burrows 5153 (K, holo.; PRE, SRGH, iso.). Figures

1; 2A, B.

Rhizome 4—14 x 2.0-3. 5 mm (ratio 2.6:1), ellipsoid,

with wiry, semi-descending roots which are rarely pro-

liferous. Stipe 18-40 mm long, ± 0.5 of its length subter-

ranean, dead stipes sometimes slightly persistent.

Trophophore usually solitary, erect, 14—39 x 2. 5-5.0 mm

(ratio 7.2:1), very narrowly elliptic, sterile trophophore

flat, fertile trophophore shallowly folded, apex sharply

apiculate, venation linear, epidermal cell walls sinuous,

stomata anomocytic. Sporophore 40-60 mm long, bearing

(6—) 1 2— 1 5(— 1 9) pairs of sporangia. Spores alete, 43—49

pm in equatorial diameter, muri thick, ± 3 pm wide, con-

tinuous and somewhat undulate, lurnina conical, 1. 5-2.3

pm wide, proximal surface concave, pitted (Figure 2A,

B).

This taxon was included under O. lancifolium s.l.

which comprises three or more species in Africa. It closely

resembles O. lusoafricanum but is clearly differentiated

from it by the spores with their conical lurnina, and the

short, ellipsoid rhizome. The apex of the trophophore is

generally sharply apiculate compared to the acute apex of

O. lusoafricanum. O. gracillimum is separated from O.

caroticaule by its trilete and smaller spores, 27-38 pm in

equatorial diameter (Figure 2D, E), and its much narrower

and slightly falcate trophophore.

Ophioglossum caroticaule appears, from the few

known collections, to be a species of shallow, sandy or

humic loams which are seasonally moist or wet, growing

among sparse, short grasses in subtropical deciduous

woodland or on open sheetrock on seasonally wet vege-

tation islands. It is a gregarious species and, in all popu-

lations so far seen in the field, grows with either O.

lusoafricanum, O. gomezianum Welw. ex A. Br., O. rubel-

lum Welw. ex A. Br. or O. costatum R. Br.

Specimens examined

(Focalities identified in terms of grid reference system —

see Edwards & Eeistner 1971.)

ETHIOPIA. — 0539: Sidamo, Bitata, 20 km from Negele on road to

Menghist, 1 450 m, 24-5-1983, Gilbert, Ensermu & Vollesen 7762 (K).

KENYA. — 0237: K4 Machakos, Nairobi-Mombasa Road, 7.4 km to-

wards Nairobi from Hunter’s Lodge, 960 m, 6-1-1972, Faden & Faden

72/18 (K, MO).

ZAMBIA. — 0831: Abercom Dist., old road to Cascalawa, 16-2-1960,

Richards 12480 (K).

ZIMBABWE. — 1828 (Gokwe): Gokwe Dist., Sengwa Wildlife Re-

search Area, Leguaan Vlei, 13-2-1983, Burrows 3016 (J.E.B. Herb.);

11-2-1991, Burrows & Burrows 5153 (K, holo., PRE, SRGH). 2027

(Plumtree): Matopo Hills, Bulalima-Mangwe, Embakwe, (-DD), 10-5-

1942, Feiertag s.n. (BM, SRGH). 2028 (Bulawayo): Matopo Hills,

Bulalima-Mangwe, Greystone, (-CA), 1-5-1939, Feiertag s.n. (BM,

SRGH).

SOUTH AFRICA

TRANSVAAL. — 2429 (Zebediela): Potgietersrus, Percy Fyfe Nature

Reserve, (-AA), 1 500 m, 22-2-1971, Huntley 2013 (PRE). 2430

(Pilgrim’s Rest): Bourke’s Luck. Goedgeloof Plantations, Treur River,

(-DB), 1 200 m, 27-12-1990, Burrows 5113 (J.E.B. Herb.).

1 .2. Ophioglossum gracillimum Welw. ex Hook. &

Bak., Synopsis filicum: 445 (1868); Burrows: 40 (1990);

Johns: 8 (1991). Type: Angola, Pungo Andongo, near

Catete, Jan. 1857, Welwitsch 36 (BM, lecto.!; K!, FISU,

iso.).

FIGURE 1 . — Ophioglossum caroticaule, Burrows 3016, Sengwa Re-

search Area, Zimbabwe.

Bothalia 23,2 (1993)

187

FIGURE 2. — Spores of Ophioglossum. O. caroticaule, J.E. & S.M. Burrows 5153: A. proximal, B. distal. O. lusoafricanum : C. J.E. & S.M. Burrows

5147 , distal; F, Burrows 4949, distal. O. gracillimum, J.E. & S.M. Burrows 5138: D. proximal, E, distal. O. sp. aff. lancifolium, J.E. & S.M.

Burrows 5158: G, proximal, FI, distal. O. vulgatum subsp. africanum, J.E. & S.M. Burrows 5122: J. proximal, K. distal. O. convexum, J.E.

& S.M. Burrows 5213: 1, L. distal. Scale bars: 20 pm.

O. lusitanicum L. var. gracillimum Welw. ex A. Br.: 177 (1868).

O. gramineum Willd., sensu Schelpe: 34 (1977).

Although the collection Welwitsch 36 is repeatedly

quoted as the type material of this species, no single ele-

ment has been selected from what are, by Welwitsch's

own admission (on the BM sheet), several different collec-

tions, albeit from the same area or locality. After his initial

gathering in January 1857, to which he allocated the num-

ber 36, Welwitsch apparently re-collected this species on

three occasions. Subsequent collections made in February

were then numbered 36a, 36b and 36c and the BM sheet

bears plants from all four gatherings. Since the January

collection (No. 36) is the first collection, and is typical of

the species, the group of seven central plants immediately

above the brown label on the sheet housed in BM, is

hereby selected as the lectotypic element for the name of

the species, with isotypic specimens in K and LISU.

The practice of placing Welwitsch's species under the

superficially similar Australo- Asian species O. gramineum

(Prantl 1883; Carruthers 1901; Clausen 1938; Tardieu-B lot

1953, 1964; Schelpe 1977) is, in my opinion, unfounded

and O. gracillimum is hereby maintained as a clearly de-

fined species confined to Africa.

188

Bothalia 23,2 (1993)

Distribution : Ghana, Tanzania, Angola, Zimbabwe,

South Africa (Transvaal).

1.3. Ophioglossum lusoafricanum Welw. ex Prantl

in Berichte der Deutschen Botanischen Gesellschaft 1:

351 (1883); Pichi Sermolli: 641 (1954); Faden: 25 (1974).

Type: Angola, Pungo Andongo, Welwitsch 34 (B, holo.

[destroyed]; BM!, neo.; K!, LISU, iso.).

Soon after the publication of Welwitsch’s epithet, the

taxon was sunk into O. gramineum Willd. by Clausen

(1938) who then, in July 1939, annotated Welwitsch’s

sheet of O. lusoafricanum in Kew as O. lancifolium Presl.

In southern Africa, Schelpe (1970) and Schelpe & An-

thony ( 1 986) have retained the epithet O. lancifolium for

all species of Ophioglossum with a narrowly elliptic to

lanceolate trophophore which is held clear of the ground

on a distinct stipe. Recent studies have shown that there

are three distinct taxa that display these general characters

within south-central Africa. O. caroticaule (described

above) appears to be rare. The most common species is

a taxon with a rather variable trophophore shape and size

but with two characters which clearly differentiate it from

what is believed to be typical O. lancifolium: spore sculp-

turing (Figure 1A & B) and rhizome shape. This common

taxon matches perfectly Welwitsch’s O. lusoafricanum , a

fact also recognized by Pichi Sermolli (1954) and Faden

(1974). What is thought to be true O. lancifolium is re-

corded from central Africa but is not recorded from south

of the 20°S line of latitude, being apparently confined to

the Zambezian phytogeographical region (White 1983).

In his type description, Prantl (1884) quotes a specimen

in Berlin as the type collection. Since all Berlin’s Ophio-

glossum material was destroyed during World War 2, a

neotype must be selected.

All the sheets seen of Welwitsch 34 in K and BM com-

prise more than one gathering. Although probably all of

the same species, in a genus with species so poorly dif-

ferentiated as Ophioglossum , it is essential that a single

gathering is selected as the type to avoid taxonomic am-

biguity. Therefore the top collection on the sheet in BM,

composed of nine plants collected in January 1857, is

hereby selected as the neotype of O. lusoafricanum.

O. lusoafricanum displays a confusing trait whereby

the first emerging trophophore from a root sucker is fre-

quently ovate to orbicular and appressed to the ground,

thereby resembling O. convexum J. E. Burrows. These

small, ovate forms may sometimes produce fertile spikes,

giving the impression that there are two topopatric species.

However the spore pattern of the two species is quite dif-

ferent (Figure 2C, F, I, L) with O. convexum being con-

sistently trilete and O. lusoafricanum being alete or trilete,

or rarely monolete. The spore sculpturing of O. convexum

is variable which may indicate hybrid activity in some

forms.

1.4. Ophioglossum lancifolium Presl , Supplemen-

tum tentaminis pteridographiae (1845). Type: Mauritius,

Du Petit-Thouars s.n. (PR, holo., P!, iso.?).

O. lancifolium is a name that has been widely applied

to any species of Ophioglossum in tropical Africa that

bears a narrow trophophore well clear of the ground

(Schelpe 1970; Tardieu-Blot 1953; Jacobsen 1983). Al-

though plants which roughly correspond to Thouars’

plants have been collected in central Africa, the status of

O. lancifolium cannot be verified on the African mainland

until good topotypic material is located from Mauritius.

The extant specimens in Paris collected by Du Petit-

Thouars from Mauritius are extremely poor and inade-

quate for a sound comparison. Specimens from mainland

Africa share the same trophophore shape, the same rhi-

zome form and (possibly) the same spore ornamentation

(Figure 2G, H). However it is vital that live plants are

seen and collected from Mauritius before the African

taxon can be equated with the type of O. lancifolium.

Unfortunately Lorence, in his survey of the pteridophytes

of Mauritius (1978), records against O. lancifolium : ‘not

seen recently’.

The distribution of the African taxon currently attrib-

utable to O. lancifolium: Angola, Namibia, Botswana,

Zambia, Zimbabwe.

2. Ophioglossum latifolium (Prantl) J.E. Burrows,

stat. nov.

O. gomezianum Welw. ex A. Br. var. latifolium Prantl in Jahrbuch des

Koniglichen Botanischen Gartens und des Botanischen Museums zu

Berlin 3: 307-333 (1884). Type: Angola, Pungo Andongo, near Catete,

Feb. 1857, Welwitsch 32 (K, lecto.!, the right-hand specimen here desig-

nated; BM!, LISU, iso.).

O. latifolium most closely resembles O. rubellum

Welw. ex A. Br. (with which it is most frequently con-

fused) but differs in its much larger size, its more slender

rhizome with wider-spreading proliferous roots, the lack

of orange pigmentation in the pedicel and trophophore,

and its completely different spore ornamentation.

Up to now we have thought that this taxon could be

a hybrid since three spore samples examined have all had

malformed or undeveloped spores. This fact is also noted

by Prantl in his original description where he states

‘sporae degenerae’. However, apparently normal spores

have been found in some collections which, together with

this species’ wide distribution, suggests that it is probably

a stabilised hybrid with a very low percentage of viable

spores produced. This is borne out by the populations seen

being extremely localised and of obvious clonal origin.

Nonetheless, there is no reason not to consider it a good

species.

There seems to be no valid reason to assume that O.

latifolium is allied to O. gomezianum since it shares very

few characters with the latter species. Most noticeably, it

lacks the suborbicular or ellipsoid rhizome that is typical

of O. gomezianum , whereas its consistently broader

trophophores which are held almost flat on the ground,

clearly separate it from O. gomezianum, and the taxon is

therefore elevated to species level.

Although no holotype was designated by Prantl in his

description, Schelpe (1977) illegitimately quotes the Kew

collection as the holotype. The Kew sheet comprises two

collections and the right-hand collection on the Kew sheet

is hereby selected as the lectotype of the name O. latifol-

Bothalia 23,2 (1993)

189

ium. Ballard’s determination of that collection as O. cos-

tatum is incorrect.

Distribution : Sierra Leone, Nigeria, Kenya, Zaire, An-

gola, Zambia, Zimbabwe.

3. Ophioglossum vulgatum L. subsp. africanum Po-

cock ex J.E. Burrows, subsp. nov., ab O. vulgato subsp.

vulgato sporis glabrioribus reticulo murorum tenuiore

aequabiliore luminisque fovearum conicis ± 1 .0-1.5 |im

latis diminutis differt.

TYPUS.— 2929 (Underberg): South Africa, Natal,

Bergville Dist., Cathedral Peak Forest Research Station,

Catchment 9, (-AB), 1 905 m, 9-1-1951, Killick 1267

(NU, holo.; PRE, iso.).

The epithet vulgatum has been generally applied to

southern African species of Ophioglossum that are mor-

phologically very similar to the north temperate O. vul-

gatum L. During a recent study of the genus in Africa,

numerous SEM photographs have been made of spores of

the southern African taxon, which has revealed a striking

difference in spore morphology between the northern and

southern hemisphere taxa. The spore morphology of O.

vulgatum has been well documented in Europe (Paul

1987; Ferrarini et al. 1986) and Asia (Kurita 1981;

Nakamura & Shibasaki 1959; Pant & Misra 1975). All

these studies reveal spore ornamentation characterised by

a very irregular and coarse reticulum of muri rising up to

shallow points resulting in a distinctly jagged profile.

In contrast, all the spores of the southern African taxon

examined have a relatively uniform and fine reticulum

which, in profile, is smooth (Figure 2J, K), lacking the

jagged reticulum of the northern hemisphere taxon. Spore

size, however, does not differ greatly [southern Africa:

(27-)33^15(-60) jim; Europe/Asia: (28-)34-40(-47) pm]

although the southern African material tends to be slightly

larger.

Pocock, in her unpublished manuscript on southern Af-

rican Ophioglossum , describes the southern African ma-

terial of O. vulgatum as a subspecies of the typical

material, based, as she puts it, largely upon geographical

considerations’. This, in itself, is not sufficient grounds

for taxonomic distinction; however, the additional infor-

mation now available on spore morphology supports this

separation, and the southern African material (none of the

specimens examined for spore pattern have shown any

resemblance to the northern hemisphere subspecies) is

now regarded as a separate subspecies. Spores of O. vul-

gatum from East Africa have not yet been examined.

Selected specimens examined

Note: specimens of which spores have been examined

by SEM are marked with an asterisk *.

ZIMBABWE. — 1731 (Harare): Domboshawa, (-CA), 26-3-1983,

Burrows 3037 (SRGH, J.E.B. Herb.). 1828 (Gokwe): Gokwe Dist.,

Sengwa Wildlife Research Area. Froggy Pond, (-AA), 10-2-1991. Bur-

rows & Burrows 5144* (K. MICH. NU, PRE, SRGH. TI); Leguaan Vlei,

(-AA), 11-2-1991, Burrows & Burrows 5155* (J.E.B. Herb., TI). 1832

(Mutare): Nyanga National Park, (-BD), 16-1-1951, Chase 3758* (BM,

BOL, SRGH). 1932 (Melsetter): Chimanimani Mts, Bundi Plain, (-DD),

26-10-1959, Goodier & Phipps 280 (BM, PRE, SRGH).

TRANSVAAL. — 2230 (Messina): Funduzi, (-CD), 21-1-1931,

Bremekamp & Schweickerdt 321 (PRE). 2328 (Baltimore): Blouberg,

(-BB), 11-1-1955, Codd & Dyer 8979 (PRE). 2430 (Pilgrim’s Rest):

Graskop, The Pinnacle, (-DD), 2-1-1961, Braithwaite 203* ( BOL ). 2530

(Lydenburg): 0.25 km N of Sabie, (-BB), 30-12-1990, Burrows & Bur-

rows 5122* (K, MICH, PRE, TI); Kaapsehoop village, (-DB), 19-12-

1990, Burrows & Burrows 5093* (K, MICH, PRE, TI).

NATAL. — 2730 (Vryheid): Utrecht, (-AD), 3-11-1985, Devenish

1884 (PRE). 2929 (Underberg): Fort Nottingham Commonage, (-BD),

30-10-1987, Goldblatt & Manning 85 1 0 (MO, PRE). 2930 (Pietermaritz-

burg): New Hanover/Mapumulo, Kingcliffe, (-BD), 5-1-1965, Moll 1519

(PRE). 3029 (Kokstad): Alfred Dist., Weza, southern slopes of Ingeli

Mtn, (-DA), 2-1-1966, Strey 6395 (BR, K, NU, PRE).

4. Ophioglossum rubellum Welw. ex A. Br. in

Kuhn, Filices africanae: 179 (1868); Hook. & Bak.: 445

(1868); Prantl: 321 (1884); Clausen: 152 (1938); Tardieu-

Blot: 23 (1953); Faden: 25 (1974); Schelpe: 33 (1977);

Johns: 10 (1991). Type: Angola, Pungo Andongo, near

Catete, Welwitsch 33 (K, lecto.!; BM!, LISU, iso.).

As with most of Welwitsch’s collections of Ophio-

glossum, his No. 33, the type of O. rubellum , is composed

of a number of collections gathered at different localities

in the Pungo Andongo District during January and Feb-

ruary 1857. The sheet in BM and one of the two sheets

in K both consist of more than one gathering. Clausen, in

1946, annotated the one sheet in K as ‘Type’, although

he never published a lectotypification. There are, however,

two (apparently) different collections on the Kew sheet.

In order to avoid confusion, the group of four plants

mounted above Welwitsch’s printed label at the bottom of

the sheet at Kew (already annotated as the type by

Clausen) is hereby selected as the lectotypic element for

the name of the species.

Distribution : Guinea (?), Sierra Leone, Ghana, Nigeria,

Central African Republic, Ethiopia, Uganda, Kenya, Tan-

zania, Gabon, Angola, Zambia, Zimbabwe, South Africa

(Transvaal).

ACKNOWLEDGEMENTS

Sincere thanks are extended to the directors and cura-

tors of the following herbaria for allowing me to make

use of their collections: BM, K, NU, P, PRE, SRGH.

WIND; Mr Ian Coulson, Officer-in-Charge of Sengwa

Wildlife Research Institute and the staff of the Zim-

babwean Dept of National Parks for allowing me to study

and collect plants on their property; the staff of the E.M.

unit of the University of Natal for help with the SEM

photographs of spores; and Mr M. Lambert for the Latin

translations.

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du Cameroun 3. Museum National d’Histoire Naturelle, Paris.

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Bothalia 23,2: 191-196(1993)

Systematic studies in the genus Mohria (Anemiaceae: Pteridophyta). IV.

Comparative gametophyte morphology in Mohria and Anemia

J.P. ROUX*

Keywords: Anemia , Anemiaceae, gametophyte, Mohria , morphology, Pteridophyta

ABSTRACT

The sporophyte morphology in Mohria and Anemia (Anemiaceae) is dissimilar. However, similarities in their anatomy,

trichomes, spores and chromosome numbers show clearly that these genera are related. The contribution of the gametophyte to

the classification and phylogeny of the Pteridophyta is largely neglected. The gametophyte of Anemiaceae is primitive in many

features when compared with that of other leptosporangiate fems. The prothallus of Mohria is considered more advanced than

that of Anemia in the germination pattern of the spores, the presence of mycorrhiza in the cushion, the permanently lateral

position of the meristematic region and the simpler trichome types.

UITTREKSEL

Die sporofiet van Mohria en Anemia (Anemiaceae) verskil morfologies van mekaar. Ooreenkomste in hul anatomie.

trigome, spore en chromosoomgetalle toon egter duidelik dat hierdie genusse verwant is. Die gametofiet van Anemiaceae is in

baie opsigte primitief wanneer dit met die van ander leptosporangiate varings vergelyk word. Die protallus van Mohria word as

meer gevorderd beskou as die van Anemia in die ontkiemingspatroon van die spore, die teenwoordigheid van mikorisa in die

protalluskussing, die meristematiese streek wat permanent in ’n laterale posisie bly en die meer eenvoudige trigoomtipes.

INTRODUCTION

Mohria Swartz, together with Anemia Swartz, Schizaea

J. Smith, Actinostachys Wall, and Lygodium Swartz are

commonly placed in a single family, the Schizaeaceae

(Engler & Prantl 1898-1902; Bower 1923; Christensen

1938; Copeland 1947; Tryon & Tryon 1982; Tryon &

Lugardon 1990), because of their exindusiate, mono-

sporangiate sori and characteristic apical ring of annulus

cells. Eggert & Delevoryas (1967) have shown, however,

that ‘schizaeaceous’ sporangia evolved independently in

coenopteridid derivatives that are quite unrelated to the

schizaeoid fems. Senftenbergia Corda, formerly included

in the Schizaeales (Reed 1947) because of its apical an-

nulus, has been noted to occur on zygopteridid foliage

(Mickel 1974). This sporangium type therefore appears to

be polyphyletic in origin. The subdivision of the schi-

zaeoid assemblage into distinct families, Anemiaceae

(. Anemia and Mohria ), Lygodiaceae ( Lygodium ) and

Schizaeaceae ( Schizaea and Actinostachys ) has recently

gained wider acceptance (Nayar 1970; Bierhorst 1971;

Love et al. 1977; Clifford & Constantine 1980; De la Sota

& Morbelli 1987).

Although Anemia and Mohria are dissimilar in mor-

phology they are evidently related in view of similarities

in their anatomy (Prantl 1881; Bower 1918; Roux et al.

1992), trichomes (Mickel 1962; Roux 1992a), spores

(Mickel 1962; Hill 1977, 1979; Dettman & Clifford 1991;

Roux 1992b) and chromosome numbers (n = 38, 76).

Bower (1923) discouraged the use of gametophyte

morphology in pteridophyte classification. However, stu-

dies in that field were continued and results have proven

* National Botanical Institute, Compton Herbarium, Private Bag X7,

Claremont 7735.

MS. received: 1992-08-03.

useful as taxonomic and phylogenetic tools. The purpose

of the present study is to summarize previously published

work, and to extend the information on the anatomy and

morphology of the prothallus. All the information relevant

to the prothallus of Mohria is synthesized and analysed.

MATERIAL AND METHODS

Anatomical studies were carried out on prothalli col-

lected in the wild. Wax embedding was done using stan-

dard techniques (Johansen 1940). Serial sections 8-10 pm

thick were taken with a rotary microtome and stained with

fast green and safranin. Photography was done with a

Zeiss ‘Axoscop’ fitted with a M35W camera. Ilford PanF

film was used throughout.

SEM studies were done on spores cultivated on clay

pots. The prothalli were critical-point dried using CO2 as

a transitional fluid. Specimens were affixed to aluminium

stubs using glue and sputter coated with Au/Pd and were

viewed in a Cambridge S200 SEM at 5 kV.

Specimens examined

Mohria caffrorum (L.) Desv., 3318 (Cape Town): Stellenbosch, (-DD),

Roux 2363.

M. marginalis (Sav.) J.P. Roux, 2829 (Harrismith): Cathedral Peak,

(-CC), Roux 2297.

M. vestita Bak., 2430 (Pilgrim's Rest): Graskop-Sabie rd, (-DD), Roux

2236. 2630 (Carolina): Swaziland, (-DB), Roux 2261 .

Voucher specimens are all deposited in the Compton

Herbarium (NBG).

192

Bothalia 23,2 (1993)

RESULTS AND DISCUSSION

Spore germination

Atkinson (1960, 1962), based her observations on

spore germination in Mohria and Anemia on whole

mounts and found the pattern in both these genera to be

identical. Atkinson (1960, 1962) recorded the first division

of the spore protoplast to be in an equatorial plane, form-

ing an apical cell at the proximal pole, this being the pole

bearing the triradiate scar, and a distal cell at the distal

pole. With the second division the apical cell divides at

a right angle to the first to form a larger green prothallial

cell and a smaller, almost colourless, rhizoidal cell (Figure

1). Nayar & Kaur (1968, 1971) categorised this germina-

tion pattern as polar and more specifically as the Anemia-

type.

Observations made by Raghavan & Huckaby (1980)

on sectioned material are in conflict with those of Atkin-

son (1960, 1962). They convincingly showed that in Moh-

ria the first division is in an equatorial plane resulting in

a larger distal cell at the distal pole and a smaller rhizoidal

cell at the proximal pole. With the second division, how-

ever, it is the distal cell that divides to form a larger distal

cell and a smaller protonemal cell (Figure 1 ). This pattern

corresponds with the Osmunda- type of polar germination

as defined by Nayar & Kaur (1968, 1971).

Raghavan & Huckaby (1980) found Mohria and Ane-

mia to be dissimilar. In Anemia , as in Mohria , the first

division is in an equatorial plane giving rise to a larger

distal cell at the distal pole. However, at the proximal pole

a smaller protonemal cell is formed. With the second cel-

lular division the distal cell divides to form a larger distal

cell and a smaller rhizoidal cell (Figure 1 ). This pattern

of spore germination does not fit into the scheme of Nayar

& Kaur (1971).

The distinct modes of gennination in Mohria and Ane-

mia probably indicate different trends of specialization

which is also evident in the sporophyte morphology. The

origin of the rhizoid and protonemal cell in Mohria by a

route similar to that seen in genera considered advanced,

confirm its specialized status.

Prothallial development

Prothallial development in Mohria (Bauke 1878; At-

kinson 1960) and Anemia (Momose 1949; Twiss 1910;

Kaur 1961; Atkinson 1962) is known in some detail and

shows no differences which may be of any significance.

Following the formation of the prothallial cell it continues

to grow and divide by transverse divisions producing a

filament five to six cells long. Longitudinal divisions are

soon evident in cells behind the apex and these continue

both in an anterior and posterior direction. Vertical, lon-

gitudinal and oblique divisions take place and the position

of the localized dividing area in relation to the rest of the

expanding but less rapidly growing gametophytes deter-

mines the shape of the early cellular plate. The margin of

the prothalli may be variously lobed. I often found uni-

cellular trichomes to be associated with the sinuses. A true

apical initial is evidently never formed. A large wing is

eventually formed with the more rapidly dividing meri-

stem in a lateral position which, in Mohria , is maintained

throughout the life of the plant. Circumstances contribu-

ting to the lateral position of the meristem can be ascribed

to the anticlinal divisions which are more numerous than

periclinal divisions. Also no wing is formed on the prox-

imal side of the meristem but only hairs and gametangia.

Divisions in a third plane give rise to a cushion directly

behind the meristem. The gametophyte grows at first more

or less parallel to the substrate but as the cushion contin-

ues to develop the thallus assumes an oblique position

supported by long rhizoids. In Mohria the prothallus even-

tually becomes upright.

Mature prothallus

In the mature prothallus of Mohria the lateral meristem

remains hidden by the continued development of the vo-

luminous spiralling wing (Figure 2A). The older basal

parts of the wing eventually turn brown as the thallus

grows upwards. Marginal areas of all but the last formed

parts of the wing are irregular in outline. The wings are

one cell layer thick except close to the cushion where it

may be two cells thick. As growth continues the massive

cushion grows upwards and appears as a column. Small

flaps of sterile tissue often appear on the cushion. These

flaps of tissue may become winglike and aid in photosyn-

MOHRIA (Raghavan & Huckaby 1980)

^^||||^

MOHRIA & ANEMIA (Atkinson 1960, 19621 /|TTT|\

4|||^

ANEMIA (Raghavan & Huckaby 19801

FIGURE 1 . — Spore germination pat-

terns in Mohria and Anemia as

observed by Atkinson (1960,

1962) and Raghavan & Huck-

aby (1980). Dark-coloured =

spore or distal cell; vertical

lines = protonemal cell; dots =

rhizoidal cell.

Bothalia 23,2 (1993)

193

FIGURE 2. — Prothallus andprothallial structures in Mohria vestita, Roi£x 2261. A, mature prothallus showing spirally formed wing; B. prothallus

cushion showing interspersed antheridia, archegonia and trichomes; C, three-celled hair on prothallus wing; D. unicellular hair on

prothallus wing; E, two-celled hair on prothallus wing; F, mature antheridium on abaxial surface of prothallus cushion; G. archegonium

on abaxial surface of prothallus cushion; H, cushion of M. vestita, Roux 2236 , prothallus showing endofungal hyphae (H). Scale bars:

A, 2 mm; B. 500 pm; C, E, 50 pm; D, F, 20 pm; G. 100 pm; H, x 1000.

194

Bothalia 23,2 (1993)

thesis. Antheridia, although mostly confined to the poste-

rior regions of the thalli may occur interspersed with ar-

chegonia (Figure 2B & G) which are formed later than

the antheridia. As the columnar cushion increases in

height, archegonia and hairs continue to be fomied in

great numbers. Abundant rhizoids are formed on the cush-

ion. Older rhizoids are brown, long and stiff and continue

to be produced along the columnar cushion. Mature thalli

of Mohria are chlorophyllous and mycorrhizal (Figure

2H) with the mycorrhiza restricted to the ventral side of

the cushion. The endophytic fungus evidently plays only

a minor role in the nutrition of the thallus, as in cultivated

material, where the fungus is absent, the thalli show no

adverse effects.

In Anemia, cells on the proximal side of the meristem

grow outwards until a small wing is formed. As this wing

becomes larger, the meristem is carried into a vertical po-

sition and the only indication of an earlier lateral meristem

is the inequality of the wings. This change in position of

the meristem is accompanied by the development of a

thick cushion bulging towards the ventral side, and by

uplifting and recurving of the wings over the dorsal sur-

face of the cushion (Atkinson 1962). Unlike Mohria the

near vertical position of the prothallus is not maintained.

In Anemia the thalli are elongate-cordate in form, more

or less prostrate in position, green at the anterior end and

dying off behind. Rhizoids and archegonia are produced

in great numbers on the ventral surface of the cushion.

Nayar & Kaur (1971) suggested the more primitive

thallus to be dorsiventral with a massive median midrib

and heavy wings several cells thick near the midrib but

progressively becoming one cell layer thick towards the

margins. In this respect the prothalli of Mohria and Ane-

mia can be considered primitive. A step in advancement

in Mohria and Anemia, however, is the elimination of the

apical initial and the development of a multicellular mer-

istematic region. Nayar & Kaur (1971) furthermore sug-

gested that among those genera with a thalloid-cordate

type gametophyte, the most primitive is the symmetrical

type. In Anemia the prothallus is temporarily asymmetrical

but later becomes symmetrical. The gametophyte of Moh-

ria remains permanently asymmetrical and is therefore

considered more advanced.

Prothallial trichomes

The thallus of Mohria bears small one- to three-celled

hairs (Figure 2C) on the wing and cushion surfaces among

the antheridia and archegonia. Atkinson (1960) found

most hairs in older thalli to occur mainly on the surface

of the cushion. My observations, however, showed that

the cushion is not significantly more hairy than the wings.

Unicellular trichomes, up to 60 pm long, similar to the

naviculate trichomes found on the fronds of the sporo-

phyte (Roux 1992a), are the most common type in Mohria

and occur on the wings as well as on the cushion (Figure

2D). Two-celled hairs common on young thalli, have pre-

viously been described by Bauke (1878) and Stokey

(1960). The outermost colourless cell is at least twice as

long as the basal cell. A chlorophyllous basal cell very

frequently bears two colourless cells (Figure 2E). This was

also reported by Bauke (1878) and Atkinson (1960). These

cells are usually of unequal size. Atkinson (1960) also

reported branched hairs and hairs up to three cells long

from the wings.

One- to four-celled hairs also occur on mature prothalli

in Anemia. At first, the hairs are marginal, developing

from a cell adjacent to the meristem. As in Mohria, the

basal cell of two-celled hairs is chlorophyllous and the

outer cell colourless. Marginal multicellular hairs have

been reported in some Anemia {A. adiantifolia , A. aurita)

species (Atkinson 1962).

Prothallial trichomes have been considered of little

value in taxonomic and phylogenetic studies (Stokey

1951, 1960; Atkinson & Stokey 1964) because similar tri-

chome types occur in apparently unrelated groups of ferns,

a situation that may be ascribed to parallel evolution. Nev-

ertheless Nayar & Kaur (1971) suggest that the restricted

distribution of hairy prothalli among the various phyletic

groups may be of value in comparative studies. Naked

prothalli appear to be the more primitive condition among

the homosporous ferns. Unicellular hairs are more com-

mon in advanced families such as Polypodiaceae,

Davalliaceae, Lomariopsidaceae and Grammitidaceae and

are usually secretory (Nayar & Kaur 1971). The prothallus

of Schizaeaceae is either subterranean or terrestrial and is

devoid of any trichomes. In Lygodium the prothallus has

been described as naked (Bauke 1878; Twiss 1910) but a

few clavate trichomes have been reported for L. flexuosum

(L.) Swartz ( Nayar & Kaur 1971, contra Mahabale &

Kulkami 1949). I have observed clavate trichomes in a

marginal position in cultured prothalli of L. japonicum

(Thunb.) Swartz. In the Anemiaceae, however, non-secre-

tory, often multicellular trichomes occur.

Gametangia

Antheridium

Antheridia are fonned on the margins and ventral sur-

faces of the thallus in the region of the meristem. They

are produced in great numbers on the cushion and may

extend to the wing. Antheridia in Mohria and Anemia are

similar in ontogeny and morphology. Atkinson (1960,

1962) described the antheridium as developing from a su-

perficial initial. A thin disc-shaped cell is cut off from the

antheridial initial to form a proximal ring cell and a distal

terminal cell. This is followed by the formation of a dome-

shaped wall which divides the terminal cell into an outer

wall or ring cell and an inner primary spermatogenous

cell. A division of the ring cell gives rise to the cover cell

of the antheridium. The antheridium structure is thus typ-

ical of the leptosporangiate ferns (Figure 2F). Successive

division of the spermatogenous cell gives rise to a small

number of sperm. In Mohria and Anemia each spermato-

zoid is contained within a cell wall at the time of release

(Atkinson 1960; Nester 1985). At dehiscence the cover

cell is shed explosively and the spermatocytes emerge one

by one through a pore.

Archegonium

My observations in the ontogeny of the archegonium

in Mohria conform with those of Atkinson (1960). Arche-

gonia are borne on the abaxial side of the cushion only.

In mature archegonia I found the archegonial neck, which

Bothalia 23,2 (1993)

195

FIGURE 3. — Section through archegonium of Mohria marginalis, Roux

2297, x 400, E, egg cell; NCC, neck canal cell; VCC, ventral

canal cell.

consists of four cells and up to eight tiers high (Figure

2G), to curve in the direction of the substrate (Figure 2B),

a system which would be advantageous to the fertilization

process. Prior to the opening of the archegonium the distal

part of the neck becomes bulbous and the neck canal cell

contorts to form a globular mass containing two to four

nuclei (Figure 3) confirming Atkinson’s (1960) observa-

tions. The venter is well embedded in the cushion and is

surrounded by a layer of small cells containing a dense

protoplasm. Ontogenetically and structurally the archego-

nia in Mohria and Anemia are similar.

Nayar & Kaur (1971) described an advanced archego-

nium as having a neck consisting of three to four tiers of

cells high, which curve away from the apex and possess

an undivided neck canal cell. Archegonia in Anemiaceae

thus conform largely with the primitive type.

Embryogenesis

The division of the zygote has not been observed by

me or by Atkinson (1960, 1962). De la Sota & Morbelli

(1987), however, claim it to be of the ‘leptosporangiate’

type, in which the first division is longitudinal or parallel

to the main axis of the archegonium.

The young embryo of Mohria is protected by a well-

developed calyptra (Figure 4A). Initially only the foot and

the stem can be identified. The foot is well embedded in

the ground tissue of the prothallus cushion. The young

embryos I examined showed the frond to differentiate be-

fore the root. The stem soon developed an apical initial

with three cutting faces. The primary root developed en-

dogenously and contained a large apical initial and a well-

defined rootcap even before breaking through the cortical

tissue of the stem (Figure 4B).

Like Atkinson (1960) I also found prothalli attached to

young sporophytes containing up to five fronds. Sections

through these prothalli show that the cell walls separating

the foot from the prothallus tissue thicken and form an

abscission layer. Mycorrhiza were also observed in tissue

FIGURE 4. — Transverse sections of embryo in Mohria "ertita, Roux

2236. A, young embryo showing calyptra (C), indicated by

arrows, x 68; B, first root before breaking through prothallus

tissue showing root cap and large apical initial, x 270; C, prothal-

lus of a developing young plant with first root, frond and shoot

apex, x 68. A, apical initial; C, calyptra; F, frond; R, root; RA.

root apex; RC. root cap.

196

Bothalia 23,2 ( 1993)

of the young sporophyte, especially the stem and roots,

as well as in the tissues of the gametophyte.

Vascular tissue in the root, stem and frond is formed

at an early stage (Figure 4C). Initially the tracheids show

spirally arranged secondary thickenings but later pitting

is of the scalariform or reticulate scalariform type.

CONCLUSIONS

Classification and phylogeny of the Pteridophyta is

largely based on the dominant sporophyte generation. The

small, usually overlooked, gametophyte generation or pro-

thallus may, however, also contribute to the understanding

of the evolutionary processes and phylogeny of the ferns.

Bower (1923) and Holttum ( 1949), cautious of the effects

of external influences during the development of the pro-

thallus, realised the meagre but important contribution it

can make with a view to classification.

Stokey (1951) and Nayar & Kaur (1971) suggested

possible evolutionary trends in the structure of the pro-

thallus and gametangia. Features that are considered ad-

vanced in the prothallus of the leptosporangiate ferns are:

a small, short-lived, autotrophic, symmetric, cordate thal-

lus with a centrally situated meristematic region and

poorly developed midrib. Trichomes are present but are

simple, unicellular and secretory. The antheridium is a

three-cellular structure, consisting of a basal, ring and cap

cell. The cap cell dehisces in its entirety and the anther-

idium has a small sperm output. The archegonia are small,

form later than the antheridia, and are situated closer to

the meristematic region. The mature archegoniunr has a

neck that curves away from the meristem and consists of

up to four tiers of cells. The neck canal cell is undivided.

Considering these changes, the prothallus of Ane-

miaceae is in many respects phylogenetically primitive, a

feature which is also expressed in many morphological

features of the sporophyte. On grounds of the sporophyte,

Mohria is considered phylogenetically more advanced

than Anemia (Bower 1923; Mickel 1962). The perma-

nently laterally placed meristematic region and the ab-

sence of multicellular hairs from the prothallus are

supportive of such an assessment.

It is thus evident that the prothallus can make an im-

portant contribution to an understanding of the phylogeny

of the Pteridophyta at the family as well as at the generic

level.

ACKNOWLEDGEMENTS

I wish to express my sincerest thanks to Dr J. Manning

for valuable suggestions.

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Pteridophyta). III. Comparative sporangium and spore morphol-

ogy. Bothalia 22: 199-204.

ROUX. JR, VAN DER WALT. .I.J.A. & VAN DER MERWE, R.B. 1992.

Systematic studies in the genus Mohria (Pteridophyta: An-

emiaceae). I. Comparative morphology and anatomy of the rhi-

zome and frond. South African Journal of Botany 58: 83-89.

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Bothalia 23,2: 197-206(1993)

Synopsis of the genus Disparago (Asteraceae)

M. KOEKEMOER*

Keywords: Asteraceae, Disparago , new sections, synopsis, taxonomy

ABSTRACT

The genus Disparago comprises nine very distinct species and is here divided into four new, well-defined sections:

Monticapra Koekemoer, comprising three recently described species, Disparago gongylodes Koekemoer, D. barbata

Koekemoer, D. pilosa Koekemoer; section Laevicarpa Koekemoer (D. kolbei) and section Uniflos Koekemoer (D. tortilis )

which are both monotypic, and section Disparago which has four species (D. ericoides. D. anomala, D. kraussii, D. laxifolia).

The previous revision (Levyns 1936a) included seven species of which six are retained. A key to the sections and the species and

a diagrammatic key are given as well as diagnostic descriptions, synonyms, typification and notes on the distribution and habitat

of each species.

UITTREKSEL

Die genus Disparago bestaan uit nege baie duidelike spesies en word hier verdeel in vier nuwe, goed omskryfde seksies:

Monticapra Koekemoer, met drie onlangs beskryfde spesies, Disparago gongylodes Koekemoer, D. barbata Koekemoer, D.

pilosa Koekemoer; seksie Laevicarpa Koekemoer ( D . kolbei ) en seksie Uniflos Koekemoer (D. tortilis) wat albei monotipies

is en seksie Disparago met vier spesies (D. ericoides, D. anomala, D. kraussii, D. laxifolia). Die vorige hersiening (Levyns

1936a) het sewe spesies ingesluit waarvan ses behou word, 'n Sleutel tot die seksies, die spesies en 'n diagrammatiese sleutel

word gegee sowel as diagnostiese beskrywings, sinonieme, tipifikasie en notas oor die verspreiding en habitat van elke spesie.

INTRODUCTION

The genus Disparago is confined to South Africa and

occurs in the fynbos of the southwestern Cape. It now

comprises nine species: six retained from the previous re-

vision (Levyns 1936a) and three recently described

(Koekemoer 1991). The genus is remarkable in many

ways — the biogeography, morphology, evolutionary de-

velopment and speciation (Koekemoer in prep, a, b).

There are many unanswered questions about Disparago

and the genus holds challenging clues to a better under-

standing of the fynbos flora.

Disparago Gaertn.. Fructibus et seminibus plan-

tarum 2: 463, t. 173 (1791); DC.: 257 (1837); Cass.: 348

(1819); Less.: 362 (1832); Benth. 323 (1873); Harv.: 277

(1894); Schltr.: 203 (1900); Levyns: 95 (1936a); Dyer:

682 (1975). Type species: Disparago ericoides (Berg.)

Gaertn. (= Stoebe ericoides Berg.).

Wigandia Neck.: 95 (1790) non Kunth; Less.: 362

(1832).

Seriphia et Stoebes spp. auctt.

The name Disparago is listed as conserved, although

conservation against Wigandia was superfluous (Rickett

& Stafleu 1960).

Shrublets or shrubs, usually profusely, sympodially

branched, larger plants dense, upright or cushion-like and

spreading; new shoots developing from below terminal

* National Botanical Institute, Private Bag X101, Pretoria 0001, South

Africa.

MS. received: 1992-02-27.

inflorescence of previous season. Leaves alternate, small,

filiform, oblong or obovate; adaxial surface villous, hairs

unicellular: margins entire, involute; apex mucronate,

apiculate or obtuse. Synflorescences terminal. Heads sol-

itary, in small clusters or in very dense globose, sub-

globose or cylindrical aggregations; 1 -flowered or up to

10-flowered; individual heads homogamous or heteroga-

mous, composed of ray and/or tubular flowers. Involucre

in 1-4 series, chaffy, lacking a distinct stereome, broadly

or narrowly cymbiform; apices acute or acuminate; mar-

gins entire, sometimes undulate or with clavate out-

growths. Ray flowers 3-lobed, female or sterile, white or

pink, ray 1. 5-4.0 x 1. 2-3.0 mm; cypselas fertile or sterile;

surface glabrous, puberulous or villous. Tubular flowers

5-lobed, lobes erect or spreading star-like, male or her-

maphrodite, white or pink; cypselas fertile or sterile, sur-

face glabrous, puberulous or villous. Pappus present or

absent, caducous or persistent, 3— 1 5(— 20), plumose or

barbed bristles, free or connate in a ring or tube at base.

Distribution : the main area of distribution is the fynbos

of the southwestern Cape, a winter rainfall area. One spe-

cies, D. tortilis , is more widely distributed and spreads

along the south and east coast to Umtamvuna in Natal

where it occurs in relict fynbos areas. In the western Cape

the genus does not occur further north than the Great Win-

terhoek mountains near Porterville. The inland boundaries

of its distribution are along the mountains bordering the

karoo or the transition between fynbos and other biomes.

DISCUSSION

The original generic description by Gaertner (1791 ) in-

cluded only species now accommodated in the section

Disparago. The generic description given above was

198

Bothalia 23,2 (1993)

therefore expanded to include all the other species subse-

quently found.

Disparago can be distinguished from its closest rela-

tives, Stoebe and Elytropappus , by the presence of ray

flowers in some/all the heads of the synflorescence. Al-

though the monotypic genus Bryomorphe also has ray

flowers, it is distinguished by a moss-like habit. The leaf

morphology of Disparago is very similar to that of Stoebe

and it is therefore very difficult to identify sterile material.

Most species of Elytropappus can be distinguished by the

presence of glands on the leaves. Generic and infrageneric

relationships will be discussed in a separate paper

(Koekemoer in prep. a).

SPECIES TREATMENT

The genus Disparago can be divided into four natural

groups which are here treated as sections. Section

Monticapra: D. gongylodes, D. barbata, D. pilosa. Sec-

tion Laevicarpa: D. kolbei. Section Uniflos: D. tortilis.

Section Disparago: D. anomala, D. ericoides, D. kraussii,

D. laxifolia.

Key to sections

la Ray flowers sterile or reduced section Disparago

lb Ray flowers female:

2a Heads 1-flowered section Uniflos

2b Heads 5-8-flowered:

3a Cypselas of both ray and tubular flowers fully developed;

tubular flowers hermaphrodite section Laevicarpa

3b Cypselas of ray flowers fully developed, those of tubular

flowers reduced; tubular flowers male . section Monticapra

In the two monotypic sections ( Uniflos and Laevi-

carpa), the cypselas of both the ray and tubular flowers

are fertile. The main differences between the sections are

in the cypsela surface, the number of flowers per head

and the arrangement of heads. Species in the section

Monticapra have inflorescences similar to those of

Bryomorphe and further investigation is needed to deter-

mine whether Bryomorphe, a monotypic genus, should be

given generic status.

Key to species (see also diagrammatic key in Table 1)

la Ray flowers female, cypsela fully developed:

2a Heads 1-flowered 5. D. tortilis

2b Heads 5-8(-10)-flowered:

3a Cypselas fully developed in ray and in tubular flowers . . .

4. D. kolbei

3b Cypselas fully developed in ray flowers, sterile in tubular

flowers:

4a Leaves obovate 1 . D. gongylodes

4b Leaves linear or lanceolate:

5a Pappus plumose, connate at base; leaves spreading;

involucre tomentose 3. D. pilosa

5b Pappus barbed, loose; leaves appressed; involucre

smooth 2. D. barbata

lb Ray flowers sterile, cypselas reduced:

6a Cypsela surface woolly 7. D. ericoides

6b Cypsela surface puberulous:

7a Pappus absent 6. D. anomala

7b Pappus present:

8a Leaf tips obtuse; leaves more than half their length apart,

mostly spreading; west of 20 degrees longitude;

flowering in spring 9. D. laxifolia

8b Leaf tips mucronate; leaves less than half their length

apart, mostly appressed at least near inflorescence;

east of 20 degrees longitude; flowering in autumn .

8. D. kraussii

TABLE 1.— Diagrammatic key to the species of Disparago

Bothalia 23,2(1993)

199

Sectio 1. Monticapra Koekemoer, sect, nov., capi-

tula heterogama, 5-8-flora; flores radii feminei; flores

disci fungentes masculini; flores radii albi, flores disci

vinacei; cypselae florum radii fertiles, florum disci steriles.

Ad altitudines altas limitata. Typus sectionis: D. gon-

gylodes Koekemoer.

Capitula heterogamous, 5-8-flowered; ray flowers fe-

male; tubular flowers functionally male; ray flowers

white, tubular flowers wine-red; cypselas fertile in ray

flowers, sterile in tubular flowers. Confined to high alti-

tudes.

The name Monticapra was chosen in honour of Miss

E. Esterhuysen (with all due respect), who like a ‘moun-

tain goat’ explored remote mountains to discover the three

species in this section.

1. Disparago gongylodes Koekemoer in Bothalia

21: 158 (1991). Type: Cape, Great Winterhoek Wilderness

Area, Esterhuysen 35788 (BOL, holo.!; NBG!, K!, PRE

(2x)!, S!, STE!).

Shrublets (50—) 100-350 mm tall, profusely branched

mainly from base. Leaves obovate, 2.5-3.5(-5.0) x 1-2

mm, involute at margins only, adaxial surface villous,

apex obtuse; pedicel twisted near base. Synflorescence of

5-12 heads in a loose arrangement. Involucre of about 12

bracts in three series, outermost series foliaceous, inner

series broadly cymbiform, narrowing to the innermost se-

ries; dark brown, often with reddish tint, margins often

undulate in upper half, apex acuminate and often reflexed.

Heads composed of two ray flowers and five tubular flow-

ers. Ray flowers white, female, rays 3-4 x 1. 5-2.5 mm;

cypselas fertile, surface woolly. Tubular flowers pink,

male; cypselas sterile, surface puberulous. Pappus present

in ray and tubular flowers, plumose, of 2 or 3 caducous

hairs, not connate, capillaries in pappus tips inflated and

these tips obtuse and loosely coalescent, tips of capillaries

acute in lower part.

Distribution: this species is only known from one area:

the top of the Great Winterhoek Mountain near Porterville

(Figure 1 ) where it grows very abundantly on coarse Table

Mountain Sandstone, on flat areas where small Restio spe-

cies are dominant.

Specimens examined

CAPE. — 3319 (Porterville) Great Winterhoek Reserve, (-AA), Es-

terhuysen 34840 (PRE, BOL), 35788 (BOL, K, NBG, STE); Twentyfour

Rivers Mountains, (-AA), Esterhuysen 21894 (BOL, PRE); Great

Winterhoek Mountains, Farm Berghof, (-AA), Koekemoer 315, 440

(PRE); Great Winterhoek Reserve, 2 km from Zuurvlakte entrance, (-

AA), Koekemoer 439 (PRE).

2. Disparago barbata Koekemoer in Bothalia 21:

158 (1991). Type: Cape, Matroosberg, SE slopes, Ester-

huysen 28615 (BOL 48345, holo.!; BOL 48346!, PRE!,

S!).

Shrublets 200-400 mm tall, erect or trailing. Leaves

filiform, 4-9 x 0.5-0. 8 mm, spirally twisted, acute,

spreading. Synflorescence of numerous heads crowded in

ovoid secondary heads 10-15 mm in diameter, dark brown

and globular. Heads with three ray flowers and three tu-

bular flowers. Involucre of about 12 bracts in 3 series,

4.5-6.0 x 0.6-1. 3 mm, narrowly cymbiform, abaxial sur-

face sparsely villoUs, margins entire, apex acuminate,

often reflexed, usually tri-coloured: light brown near base,

dark brown towards apex and tinted red in centre. Ray

flowers female, ray 2.0-2. 5 x ± 1.5 mm, white; cypselas

fertile, puberulous, dorsiventrally flattened. Tubular flow-

ers functionally male, purple-red; cypselas sterile, puberu-

lous. Pappus present in ray and tubular flowers, of 2 or

3 bristles, barbed, caducous, not connate.

Distribution: this species is only known from the

Matroosberg near De Dooms in the southwestern Cape

(Figure 1).

Specimens examined

CAPE. — 3319 (Worcester): Matroosberg SE slopes, (-BC), Es-

terhuysen 28615 (BOL, PRE, S); Matroosberg E slopes above De Dooms,

(-BC), Esterhuysen 28119 (BOL).

3. Disparago pilosa Koekemoer in Bothalia 21: 159

(1991). Type: Cape, Roodeberg, near Matroosberg, N

slopes, Esterhuysen 29721 (BOL 48349, holo.!; BOL

48350!, NBG!, PRE!, S!).

Shrublets, up to 250 mm tall, profusely branched from

base, branches woolly or sparsely villous, appearing 4-an-

gled because of leaf arrangement. Leaves lanceolate, 2-6

x 0.5-0.9 mm, appressed or slightly spreading, not spirally

twisted. Synflorescence with secondary heads composed

of a few heads, loosely grouped. Heads with 3 ray flowers

and 5 tubular flowers. Involucre 11-13, in 2 or 3 series,

outermost broadly cymbiform, decreasing in width to the

inner series which can be filiform, margins undulate,

sometimes with membranous edge, apex acuminate,

sometimes reflexed, adaxial surface villous in lower half,

bicolorous, yellow-brown in lower half, black-brown in

upper half with reddish tint near apex. Ray flowers female,

white, 2. 5-3.5 x 1. 5-2.0 mm; cypselas fertile, surface

puberulous. Tubular flowers functionally male, reddish

pink, often with glands on surface of lobes; cypselas ster-

ile. Pappus present in ray and tubular flowers, plumose,

of about 10 hairs, connate in a ring at base.

Distribution: this species has only been collected on

Matroosberg, Roodeberg, Roelofsberg and at the foot of

Bokkeveld Sneeukop in the Worcester District (Figure 1).

It is reported to occur in sand in rocky areas.

Specimens examined

CAPE. — 3219 (Wuppertal): Ceres Dist., northern base of

Roelofsberg, (-CC), Esterhuysen 34675 (BOL, S); Waboomsrivier, Cold

Bokkeveld, (-CC), Hanekom 624 (PRE). 3319 (Worcester) Ceres Dist.,

Rosendal & Bo-Rosendal, at foot of Bokkeveld Sneeukop, (-AB), Es-

terhuysen 33928 (BOL, S); Ceres Dist., Roodeberg near Matroosberg, N

slopes, (-BC), Esterhuysen 29721 (BOL, NBG, PRE, S); Matroosberg

northern plateau, (-BC), Esterhuysen 27704 (BOL); Matroosberg near

the top along pipeline, (-BC), Koekemoer 361 (PRE).

Sectio 2. Laevicarpa Koekemoer , sect, nov., capit-

ula heterogama, 5-8-flora; flores radii feminei; flores disci

hermaphroditi; flores et radii et disci: rosei; cypselae

florum et radii et disci fertiles. Ad montes regionem

200

Bothalia 23,2 (1993)

FIGURE 1. — Geographical distribution of D. gongylodes , •; D. barbata,

O; and D. pilosa, ▼.

‘Karoo’ cognitam conterminos limitata. Typus sectionis:

D. kolbei (H. Bol.) Hutch.

Capitula heterogamous, 5-8-flowered; ray flowers fe-

male; tubular flowers hermaphrodite; ray as well as tubu-

lar flowers pink; cypselas fertile in ray as well as tubular

flowers. Confined to mountains bordering the Karoo.

The name Lcieviccirpa refers to the smooth surface of

the cypselas in D. kolbei.

4. Disparago kolbei (H. Bol.) Hutch.-. 511 (1932).

Amphiglossa kolbei Bolus: 394 (1909). Type: Cape, Oudtshoorn Dist.,

Swartberg Pass, Kolbe 1477 (BOL, holo!).

D. rosea Hutch.: 511 (1932); Levyns: 99 (1936a). Type: Cape, Uni-

ondale Dist., hills near Avontuur, Fourcade 1663 (BOL 48241, lecto.!,

here designated; GRA!, PRE!, K).

The name D. kolbei was given to commemorate F.C.

Kolbe, the collector of the type specimen and friend of

H. Bolus.

Shrublets usually 100-300 mm tall, but old plants up

to 700 mm tall and about 1.5 m in diameter. Leaves fili-

form to lanceolate, 2-4 x 0.3-0. 5 mm, shorter leaves

straight and appressed, longer leaves spirally twisted and

spreading, often woolly. Synflorescence of 1— 3(— 5) heads.

Heads with 2 or 3 ray flowers and 2 or 3 tubular flowers.

Involucre of about 10 bracts in 3 series, outermost series

ovate, about 1/3 as long as inner series of broadly cymbi-

form, light brown bracts. Ray flowers female, pink; cyp-

selas fertile, surface glabrous, apical annulus usually well

developed. Tubular flowers hermaphrodite, pink; cypselas

identical to those of ray flowers. Pappus present in all

flowers, of 8-12 plumose hairs, basally connate for about

Vs of their length.

Distribution: this species is confined to the mountain

ranges bordering the Karoo, stretching from Antoniesberg

in the east to the Witteberge in the west (Figure 2).

Specimens examined

CAPE. — 3320 (Ladismith): Witteberg, (-BC), Compton 2689 (BOL),

3335 (STE), 12201 (NBG); Esterhuysen s.n. (NBG); Goldblatt 3794 (K,

PRE, S); Matjiesfontein, Witteberg near radio mast, (-BC), Van Zyl 3561

(PRE, STE); Anysberg, (-DA), Esterhuysen 17325 (BOL), 25986 (BOL,

PRE); Koekemoer 305 (PRE); Marshall 19 (SAAS), 163 (STE); Van Wvk

1032. 1048 (PRE, STE); Van Zyl 3394 (PRE, STE); Vlok 451 (PRE, STE).

3321 (Ladismith): Elandskloof between Vleiland and Seweweekspoort,

(-AD), Moffett & Steensma 3870 (STE). 3322 (Oudtshoorn): Swartberg

Pass, (-AC), Bond 1556 (NBG); Esterhuysen 19683, 28823 (BOL);

Hafstrom & Acocks 2338 (PRE, S); Koekemoer 299, 332 (PRE); Kolbe

1477 (BOL); Levyns 6655 (BOL); Markotter 9946 (STE); Schlechter s.n.

(BOL); Stokoe 9070 (BOL), 58107 (SAM), 65910 (PRE, SAM), 64077

(SAM); Taylor 6974 (PRE, STE); Wall 277 (S); Swartberg, Hat-

tinghskloof, (-AC), Thompson 1319 (PRE, STE); Swartberg, EVKOM-

road on ridge, (-AC), Pienaar 38 (SAAS, STE). 3323 (Willowmore):

Antoniesberg, N slopes, (-AD), Esterhuysen 24944 (BOL); Uniondale,

hills near Avontuur, (-CA), Fourcade 1663 (BOL, GRA, PRE); Union-

dale, hill N of Avontuur, (-CA), Fourcade 4611 (BOL, PRE, STE);

Avontuur, hill NE of Avontuur, about 5 m from beacon, (-CA), Koeke-

moer 486 (PRE); Hoopsberg, accessed from Farm Hoogste Drift, lower

slopes of western ridge, (-CB), Koekemoer 480 (PRE).

Sectio 3. Uniflos Koekemoer , sect, nov., capitula

homogama, uniflora; flores radii feminei; flores disci

hermaphroditi; flores radii et disci: rosei vel albi; cypselae

florurn et radii et disci fertiles. Sectio vulgaris. Typus

sectionis: D. tortilis (DC.) Sch. Bip.

Capitula homogamous, 1-flowered; ray flowers female;

tubular flowers hermaphrodite; colour of ray and tubular

flowers similar: either pink or white; cypselas fertile in

ray as well as tubular flowers. A widespread section.

The name Uniflos refers to the one-flowered heads of

the species in this section.

5. Disparago tortilis (DC.) Sch. Bip. in Jahres-

bericht der Pollichia: 25(181) (1861). Type: C.B.S., Uiten-

hage, Ecklon 1576 (G-DC, lecto., here designated;

microfiche PRE!).

Stoebe tortilis DC. 260 (1837).

D. ericoides auct. non Berg, nec Gaertn.: 463 (1791); Cass.: 348

(1825); DC.: 257 (1837); Harv.: 278 (1894); Levyns: 100 (1936a); R.A.

Dyer: t. 1102(1951).

The name D. tortilis has not been used extensively in

the history of the genus. This taxon was widely known

under a misapplied name, D. ericoides. Although the type

specimen of D. tortilis is very poor, there is no doubt

about its identity or about the identity of D. ericoides.

Shrubs up to 1 m tall, profusely branched, erect or

bushy. Leaves filiform, 4— 8(-10) x 0.5-0.8 mm, spirally

twisted with 1-1 1/2 twists, usually spreading, sometimes

appressed, margins tightly involute, apex mucronate. Syn-

florescence usually with numerous, sometimes only about

5 heads, in usually globose aggregations, or if tubular then

often interrupted at base. Heads 1-flowered, with either

tubular or ray flower. Involucre of 6-8 bracts, in 2 series,

narrowly cymbiform, in outermost series often chaffy in

lower half and foliaceous in upper half. Ray flowers fe-

male, usually pink (various shades), occasionally white,

2. 7-3.3 x 2.5-3.0 mm; cypselas fertile, surface puberu-

lous, apical annulus often well developed. Tubular flowers

hermaphrodite, usually pink, or white, glands sometimes

Bothalia 23,2 (1993)

201

FIGURE 2. — Geographical distribution of D. kolbei. ■; D. kraussii, •; and D. laxifolia, O.

present on outside of lobes; cypselas fertile and identical

to those of ray flowers. Pappus usually of 5 plumose hairs,

not connate, occasionally up to 20 hairs in robust forms

and then slightly connate at base.

Distribution of this species is disjunct: it is found from

Jonkershoek and Simon’s Bay along the south and east

coasts (not more than 150 km inland) to the Dweza State

Forest in Transkei and re-appears around Port Edward and

Port Shepstone, where it occurs in what appear to be relict

fynbos areas (Figure 3). D. tortilis is very well adapted

to the grassy fynbos of the eastern Cape and is a pioneer

in disturbed areas.

Specimens examined

NATAL. — 3030 (Port Shepstone): Murchison, (-AA), Medley Wood

3078 (NH); Oribi Gorge, (-CA), Me Clean 578 (PRE. NH); Van Wyk

5415 (PRE); top of Horseman's Point, (-CB). Glen 473 (STE); Oribi

Nature Reserve, footpath between Lukonka Point and Umfezi Point,

(-CB), Abbott 1879 (NH); Fourman's Hill, (-CB), Koekemoer 264

(PRE); Umtamvuna Nature Reserve, Eon Crown, (-CC); Abbott 1929

(NH); Koekemoer 261 (PRE); Uvongo River tributary, Wichman's Farm,

(-CD), Nicholson 1704 (PRE); Port Shepstone Sea Park, (-CD), Strey

8116 (PRE); Umzumbi, (-DA), Medley Wood 12857 (PRE). 3130 (Port

Edward): Port Edward, (-AA), Acocks 10910 (PRE).

CAPE — 3227 (Stutterheim): Dohne Research Station, (-CB), Perks

s.n. (GRA A1370); Mount Coke, (-CD), Sim 1418 (GRA, NBG); East

London, Bonza Bay, (-DD), Acocks 9552 (PRE). 3228 (Butterworth):

Dwesa, (-BD), Abbott 2633 (NH); East London, Kei Mouth, (-CB),

Flanagan 213 (BOL, GRA, NBG, SAM); Stutterheim, Kentani, (-CB),

Pegler 260 (BOL, NBG); East London, Gonubi Springs, (-CC), Acocks

10986 (PRE); Kentani River mouth. (-CC), Galpin 5789 (PRE); Gonubi

Springs, (-CC), Compton 17043 (NBG); Levyns 10826 (BOL). 3318

(Cape Town): Jonkershoek, (-DD), Kruger KR339 (PRE). 3319 (Worces-

ter): Jonaskop, (-DC), Boucher 5012 (PRE). 3320 (Montagu): Witteberg,

(-BC), Humbert 9804 (PRE); Marloth Nature Reserve, (-CD), Taylor

7686 (PRE). 3321 (Ladismith): Garcia’s Pass, (-CC), Leipoldt 16049

(BOL); Fouriesberg, (-DD), Koekemoer 475 (PRE). 3322 (Oudtshoom):

Swartberg. (-AC), Pienaar 41 (STE); Spitskop, (-AD), Koekemoer 33 1

(PRE); Meiringspoort, (-BC), Esterhuysen 24855 (PRE); Swartberg east,

Magas, (-BC), Marshall 253 (PRE); Klein Moeras River, (-CC),

Koekemoer 473 (PRE); Robinson Pass, (-CC), Koekemoer 476 (PRE);

Waboomskraal, (-CD), Lewis 68721 (BOL); Kamanassie Mountains,

Laudina, (-DB), Esterhuysen 18337 (BOL, PRE); Mannetjies Mountain,

(-DB), Esterhuysen 6404 (BOL); Taylor 1469 (SAAS). 3323

(Willowmore): Slypsteen Mountain, (-AC), Esterhuysen 6305 (BOL,

PRE), 6316 (PRE); Antonies Mountain, (-AD), Esterhuysen 24965

(BOL); Potjies River Height, (-CA), Hugo 1473 (PRE. STE); Thompson

964, 1335 (PRE. STE); Hill NE of Avontuur. (-CA); Fourcade 4607

(BOL); Koekemoer 485 (PRE); Langkloof, near Ongelegen siding on

road to Kareedouw, (-CB), Immelman 324 (PRE, STE); Hoopsberg,

(-CB); Esterhuysen 6579 (BOL); Koekemoer 479 (PRE); Kouga Moun-

tains, Farm Hoeree, (-DB), Oelofsen 43 (PRE); Tuschen Bij, Tsitsikama,

(-DD), Fourcade 487 (STE); 7 miles east of Joubertina, (-DD), Ward

5697 (PRE); The Hoek, valley at foot of Outeniqua Mountains, (-DD),

Esterhuysen 10606 (BOL); Joubertina, (-DD), Esterhuysen s.n.

(BOL48263). 3324 (Steytlerville): Baviaanskloof Mountains, between

Smitkraal and Wilgerhof, (-CB), Thompson 1945 (PRE. STE); between

Patensie and Willowmore, (-CB), Koekemoer 281 (PRE); Kareedouw

Pass, (-CC/CD), Fourcade S.D. (STE); Assegaai Bosch road to Karee-

douw. (-CD), Britten 1247 (PRE); Assegaaibosch, (-CD), Breyer 23313

(PRE); Levyns 5589 (BOL); Kareedouw Pass, (-CD), Gillett 1499 (STE);

Koekemoer 324 (PRE); Farm Goedehoop, (-DA), Stirton 9593 (PRE).

3325 (Port Elizabeth): Rietberg, above Kirkwood, (-AD), Nordenstam

282 (S): Somerset Oos, Kommadagga, (-BB), Bayliss BS-1589 (PRE);

Zuurberg. above Slagboom, (-BC), J.R. & B.R. 452 (BOL, GRA);

Zuurberg National Park. Brandrug, (-BC), Koekemoer 64, 169 (PRE);

Addo Elephant Park, Zuurkop, (-BD), Archibald 3867 (PRE); Fer-

niebrae, (-BD), Koekemoer 63 (PRE); Zuurberg Inn, (-BD), Koekemoer

170 (PRE); Addo Elephant Park, Zuurkop, (-BD), Barnard 575 (PRE);

Zuurberg Pass, (-BD), Nordenstam 355, 367 (S), Groendal Wilderness

Catchment area, (-CA), Scharf 1172, 1455, 1608 (PRE); Port Elizabeth,

1 mile west of Rocklands, (-CA), Wisura 2858 (NBG); Port Elizabeth,

(-CB), Cruden 441 (GRA); Couga Mtns, (-CB), Taylor 899 (NBG); Van

Stadens Reserve, (-CC), Dahlstrand 1977, 2538 (STE), 3209. 3220

(PRE); Mkhotyana 20 (GRA); 18 km from Port Elizabeth on road to Van

Stadens Pass, (-CC), Koekemoer 61 (PRE); Witteklip, (-CC) .Rodin 1011

(P, PRE); near Port Elizabeth, (-DC), Tyson 2179 (PRE, SAM, STE);

flats near Van Stadens River Mtn, (-CC), Ecklon & Zeyher s.n. (GRA);

Blue Mtn, Loerie, (-CC), Dix 7 (GRA); Van Stadens Mtn. (-CC), Ecklon

c£ Zeyher 51 (5.2) (SAM ), 166 (4.2) (S); Zeyher 712 (NBG); Bethelsdorp.

(-CD). Bolus s.n. (NBG. PRE); Van Stadens River, (-CD), Koekemoer

323 (PRE); Port Elizabeth, Springs Nature Reserve. (-CD), Olivier 2517

(GRA. PRE); Ten Top Hill, Groendal, (-CD), Olivier 3193 (PRE);

Uitenhage, (-CD), Pappe s.n. (GRA); near ‘Thees’, Van Stadens River,

(-CD), Scott Elliot 295 (NBG); Theescomb. (-CD), Bolus s.n. (BOL);

Van Stadens Reserve, (-CD), Pienaar 57 (GRA); 14 km NW of

Rocklands on Elands River rd, (-CD), Snijman 325 (NBG PRE); flats

between Krakakamma and Van Stadens Mtn, (-CD), Ecklon & Zeyher

99 (5.2) (S); Uitenhage, (-CD), Pappe s.n. (GRA); Addo National Park,

Zuurkop, (-DA), Archibald 3867 (BOL); Botha 5677 (GRA); 2 miles

SW of Addo Drift, (-DB ), Fries, Norlindh & Weimarck 761, 1175 (BOL);

Alexandria, 6.5 miles N of Nanaga shop, (-DB), Acocks & Story 2795

(PRE); near Port Elizabeth golf course, (-DC), Britten 1789 (GRA); 8

miles from Port Elizabeth to Grahamstown, (-DC), Comins 751 (PRE);

Markman Industrial Area. Port Elizabeth, (-DC), Dahlstrand 2835

202

Bothalia 23,2 (1993)

(GRA, PRE. STE); Coega, (-DC), Horn D.H.S. (PRE); 26 km from Port

Elizabeth on N2 to Grahamstown, (-DC), Koekemoer 278, 279 (PRE);

24 km from Port Elizabeth, (-DC), Koekemoer 280 (PRE); Parsons Vlei,

(-DC), Long 36 (GRA); Redhouse, (-DC), Long 1062 (GRA); Koega,

Farm Sonop, (-DC), Olivier 708 (NBG), 1582 (PRE); Redhouse, (-DC),

Paterson 478 (GRA); between Port Elizabeth and Grahamstown, (-DC),

Story 2821 (PRE); Port Elizabeth, Kabega Park, (-DC), Olivier 440

(GRA); Port Elizabeth golf course, (-DC), Walters P(l) (NBG); Port

Elizabeth, (-DC), West 167 (GRA); Walmer, (-DC), Ecklon s.n.

(BOL48256, STE); Paterson 1174 (BOL); Port Elizabeth, (-DC),

Drege 149 (PRE); Algoa Bay, (-DD), Zeyher 2921 (P, S, SAM, STE).

3326 (Grahamstown): Alicedale, (-AC), Bavliss BR-404 (PRE);

Hofman’s Bosch, (-AC), Britten 1016, 1137 (GRA); Alexandria, Hillary,

(-AC), Burtt Davy 14245 (BOL); Tsitsikama National Park, (-AC),

Relief 396 (PRE); Highlands road, Farm Atherstone, (-AD), Boucher

2484 (PRE, STE); Highlands road, near railway crossing, (-AD), Brink

213 (GRA, PRE, STE); Alicedale road near Grahamstown, (-AD), Brink

378 (GRA, PRE); Howieson’s Poort, (-AD), Batten s.n. (NBG); Comins

1701 (GRA, NBG, STE), 1707 (GRA); Rennie 327 (BOL); Grahams-

town, (-AD), Guthrie 3310 (NBG); Coldspring, (-AD), Jacot

Guillarmod 8720 (PRE), 9569 (GRA, PRE); Highlands station, (-AD),

Burtt Davy 7951 (BOL, PRE); near Coldstream station, (-AD), Killick

776 (PRE); between Grahamstown and Alexandria, 1 km from Salem,

(-AD), Koekemoer 174 (PRE); Atherstone, (-AD), Koekemoer 277

(PRE); Grahamstown, Frasers Reserve, (-AD), Shumane P28 (GRA);

Salem, (-AD), Zeyher 291 (S, SAM); Albany, (-BC), Erens PRE28359

(PRE); 5 km from Grahamstown on road to Port Alfred, (-BC),

Germishuizen 1511 (PRE); on R67, 13 km from Grahamstown near

Belmont Valley, (-BC), Koekemoer 272 (PRE); Grahamstown near Sett-

lers Monument, (-BC), Koekemoer 273 (PRE); Gunfire Hill, (-BC),

Richardson 21, 27 (GRA); Grahamstown, (-BC), Britten 1939-11

(NBG); Dyer 153 (GRA); MacOwan 1002 (BOL); Marloth 6130 (PRE);

Rogers 27267 (GRA), 27367 (BOL, STE)); Theron 496 (PRE); Tyson

16837 (BOL); Signal Hill (-BC), Levyns 3765 (BOL); Grahamstown

hills, (-BC), Tyson 16837 (SAM); Port Elizabeth, Vaal Vlei Estate,

(-BC), Mogg 4702 (PRE); Bathurst, 8.7 miles WNW of Kleinemonde,

(-BD), Acocks 21820 (BOL, PRE); Round Hill, (-BD), Bolus 4110

(PRE); Bathurst, Farm Hopewell, (-BD), Compton 19839 (NBG); Gra-

hamstown, (-CA), Britten 6559 (PRE); Alexandria Dist., (-CB), De Vos

c-18446 (STE); Alexandria, near station, (-CB), Galpin 10634 (PRE);

Alexandria, Nauaga, (-CB), Bond 1241 (NBG); Alexandria, NE of

Nananga Post Office, (-CB), Acocks 12795 (PRE); Alexandria, (-CB),

Archibald 4381 (GRA); Grahamstown, Barville Park near Kasouga,

(-DA), Bayliss 8582 (GRA); 8 km from Port Alfred at Hayes siding,

(-DB), Relief 299 (PRE); Kowie West, (-DB), Tyson s.n. (BOL13299,

PRE); betw. Grahamstown and Coegha, (-DC), Sidey 1678 (PRE). 3327

(Grahamstown); near Bell, Dist. Peddie, (-AB), Galpin 7688 (GRA);

Grahamstown, near war monument at Kaffirdrift Police Sta., (-AC),

Burrows 2776 (GRA); East London, above Kwani River, (-BA), Comins

1519 (PRE); East London, The Springs, (-BA), Rattray 1370 (BOL).

3418 (Cape Town): Noordhoek Peak, (-AB), Gillett 3568 (PRE); above

Chapmans Peak Drive, (-AB), Whellan 1786 (PRE); Palmiet River

Mouth, (-BD), Boucher 1045 (PRE). 3419 (Caledon): Lebanon Reserve,

Grabouw, (-AA), Verdoucq 87 (STE); Hermanus, Femkloof, (-AD),

Orchard 321 (PRE); Zwarteberg and Klein Rivers Mountain, (-AD),

Ecklon s.n. (S); 2 miles W of Papiesvlei Post Office, (-BC), Acocks 22312

(PRE); Paardeberg Mountain, (-BC), Stokoe 64076 (PRE); The Kelders

(-CB), Van derMerwe 2033 (PRE). 3420 (Swellendam): Bontebok Park,

(-AB), Liebenberg 6751 (PRE, STE), 7179 (PRE); De Hoop, Oulande,

(-AD), Eellingham 874 (PRE); Van Wyk 2179 (PRE); Potberg, (-BC),

Acocks 23011 (PRE); Bredasdorp, Die Poort, (-CA), Compton 22609

(STE). 3421 (Riversdale): Langeberg, above Herbertsdale, (-BB), Muir

1259 (BOL); Skulpiesbaai, (-BC), Van Schalkwyk 84 (PRE). 3422

(Mossel Bay): Goukamma Nature Reserve, (-BB), Heinecken 217

(PRE). 3423 (Knysna): The Heads, (-BB), Phillips 1050 (SAAS);

Tsitsikama Coastal Park, (-BB), Retief 396 (PRE); Plettenberg Bay,

(-AB ), Rogers 22690 (PRE); Ratels Bosch, Tsitsikama, (-BB ), Fourcade

336 (BOL, GRA). 3424 (Humansdorp): E of Witelsbos, (-AA), Hutch-

inson 1418 (K); Humansdorp, (-BB), Burtt Davy 11984 (PRE); Levyns

3788 (BOL); Rogers 2946 (PRE); Cape St Francis, (-BB), Thompson

896 (PRE, STE); Kareedouw Pass, (-BB), Gillett 1373 (STE), 2023 (K);

Humansdorp, Majo, (-BB), Ecklon & Zeyher 51 (4.2) (SAM); Port

Elizabeth, Winterhoek Mtn, (-DC), Fries, Norlindh & Weimarck 1082

(BOL, S); Redhouse, (-DC), Fries, Norlindh & Weimarck 545 (BOL);

Port Elizabeth, (-DC), Fries, Norlindh & Weimarck21 (BOL). 3425 (Port

Elizabeth): Cape Recief, (-BA), Ecklon & Zeyher 113 (93.7) (S).

Sectio 4. Disparago

Capitula heterogamous, 2-flowered; ray flowers sterile;

tubular flowers hermaphrodite, colour of ray and tubular

flowers similar: either white or pink; cypselas reduced in

ray flowers, fertile in tubular flowers. Confined to coastal

flats.

18° 20° 22° 24° 26° 28° 30°

28°

30°

32°

34°

FIGURE 3. — Geographical distribution of D. tortilis.

Bothalia 23,2 (1993)

203

6. Disparago anomala Schltr. ex Levyns in Journal

of South African Botany 2: 101 (1936a). Type: Cape,

Worcester, Papiesvlei, Schlechter 10443 (PRE, lecto.!,

here designated; G(2x)!; GRA!; PRE(4x)!, S!).

D. anomala Schltr, in sched.

D. anomala Schltr. ex Bolus & Wolley-Dod: 280 (1904), nom. nud.

The specific name refers to the absence of a pappus.

Shrublets very variable, 80-150 or 300-800 mm tall,

profusely and densely branched and cushion-like or large

open shrubs. Leaves filiform, 3— 7(— 10) x 0.4-0.6 mm, spi-

rally twisted with 1-11/2 twists, mucronate, mucro unob-

trusive or to 0.8 mm long, margins fully involute.

Synflorescence : heads in globose or subglobose aggrega-

tions. Heads 2-flowered or sometimes 1 -flowered, usually

1 ray flower and 1 tubular flower. Involucre in 2 series,

inner series of 3 or 4 bracts, chaffy, 3-5 x 0.4— 0.7 mm,

usually bicoloured, outer series often only partly chaffy.

Ray flowers sterile, 1. 7-2.0 x 1.3-1. 6 mm, usually white

or sometimes pink; cypselas reduced, often not distinct

from corolla. Tubular flowers hermaphrodite, ?lobes ap-

parently not opening in star-like manner; cypselas pentag-

onal, apical annulus usually well developed, surface

puberulous, hairs falcate. Pappus absent.

Distribution : D. anomala occurs on coastal flats in the

southwestern Cape. It is very common in the Peninsula

and spreads northwards to Blackheath and eastwards to

the Breede River (Figure 4). It very often grows on

calcrete ridges or calcareous soils.

Specimens examined

CAPE. — 3318 (Cape Town): Blackheath, (-DC), Boucher 5083

(STE): Raitt 44 (STE, PRE). 3418 (Simonstown): Peninsula. Witsand,

(-AB), Penfold 228 (NBG); Boon Mountain, (-AB), Pillans 4919 (BOL);

Salter 4277 (K); Klipfontein road, E of Mowbray, (-AB). Salter 280-15

(BOL); Buffels Bay, (-AB), Salter 280-15b (BOL); Rheboks Dam, Good

Hope Nature Reserve, (-AB), Taylor 6732 (PRE); Sirkelsvlei, (-AD),

Compton 19371 (NBG, STE); Bonteberg, (-AD), Compton 10627

(NBG); Buffels Bay, (-AD), Compton 13000 (NBG); Brightwater, (—

AD), Compton 14571 (NBG); slope above Smitswinkel Bay, (-AD),

Esterhuysen 32819 (BOL); Galpin 12785 (PRE); Gillen 1508 (K); Cape

Point, (-AD), Gillett 3474 (STE); Good Hope Nature Reserve. (-AD),

Koekemoer 354 (PRE); Smitswinkel Bay, (-AD), Levyns 4935 (BOL);

Salter 4281 (PRE, K); Buffels Bay, (-AD), Salter 2933, s.n. (BOL); Good

Hope Nature Reserve, near Gifkommetjie turnoff, (-AD), Taylor 6717a

(STE); Smitswinkel Vlei, (-AD), Wolley-Dod 766 (K); Cape Rats, (-

BA), Adamson 4256 (BOL); Cape Flats near beacon FO-ST2, (-BA),

Taylor 3 150 (PRE. STE); near Faure, (-BB), Galpin 12565 (PRE); W of

Faure, (-BB), Salter 4326 (PRE, BOL, K); Die Punt, Pringle Bay, (-BD),

Boucher 1461 (STE, PRE); Caledon, Hanglip, (-BD), Esterhuysen 33494

(BOL); Betty’s Bay, (-BD), Levyns 10072 (BOL); Pringle Bay, (-BD),

Levyns 10847 (BOL); Rooiels, (-BD), Parker 4180 (BOL, NBG). 3419

(Worcester): Houwhoek, (-AA), Bolus s.n. (PRE); Guthrie 3601 (NBG);

Schlechter 7412 (BOL, G. P); S of Caledon, (-AB), Salter 9034 (SAM);

Hawston, (-AC), Leighton 410 (BOL); Onrust River, (-AC), Schlechter

1986 (PRE), 10393 (BOL. G. GRA. PRE, S); border between Wortelgat

and Walker Bay State Forest, (-AD), Bosenberg & Rutherford 226 (STE);

near Shaw’s Mtn, (-AD), Salter 9034 (BOL); Papiesvlei, (-AD/BC),

Schlechter 10443 (G. GRA, PRE); Genadendal, (-BA), Gillett 899

(STE); Flouhoogte, E of Stanford, (-BC), Hugo 1544 (PRE, STE); Die

Skeiding, between Bredasdorp and Elim, (-BC/BD), Fellingham 249

(PRE. STE); Bredasdorp Mountain, (-BD/DB), Galpin 10479 (G, PRE);

Gans Bay, (-CB), Hall 4610 (NBG, PRE); Die Kelders, (-CB),

Koekemoer 347 (PRE); Hagelkraal, (-DA), Barker 5276 (NBG); Comp-

ton 19035 (STE); Hugo 1591 (STE), 1721 (PRE. STE); Lewis 63089

(SAM); Viljoenshof between Elim and Die Dam, (-DA), Koekemoer 348

(PRE); Van Wyk 799 (STE); Groot Hagelkraal, near Pearly Beach. (-DA),

Van Wyk 1146 (PRE, STE); Elim, (-DB). Barker 5288 (NBG); Schlechter

7679 (BOL, G, GRA); Die Poort, between Bredasdorp and Elim, (-DB),

Bolus 20538 (BOL); between Bredasdorp and Elim, (-DB), Gillett 1444

(STE); flats between Bredasdorp and Elim, (-DB). Levyns 4871 (PRE,

BOL); Bredasdorp Forest Reserve, (-DC), Hugo 1643 (PRE, STE); 2 km

from Elim, (-DC), Koekemoer 346 (PRE). 3420 (Bredasdorp): Bontebok

Park, (-AB), Compton 21929, 22630 (NBG); De Hoop, Potberg, (-BC),

Burgers 1759 (PRE, STE), 1815 (PRE); Potberg, Farm Albertsdale. (-

BC), Compton 19531 (NBG); Potberg, (-BC), Koekemoer 182, 183

(PRE); Levyns 8180, 11554 (BOL); Lewis 63082 (SAM); Thompson 3766

(PRE, STE); Bredasdorp, Die Poort, (-CA), Compton 22609 (NBG);

Bredasdorp, (-CA), Levyns 3523 (BOL); between Bredasdorp and Struis

Bay, (-CA), Levyns 3534a (BOL); Bredasdorp, (-CA), Levyns 11547

(BOL); Bredasdorp, Heuningberg Nature Reserve, (-CA), Koekemoer

184 (PRE); Bredasdorp, 1 km from Struis Bay to Elim, (-CC),

Koekemoer 343 (PRE).

7. Disparago ericoides (Berg.) Gaertn., Fructibus

et seminibus plantarum 2: 463 (1791); Fess.: 363 (1832).

Type: ‘e Cap. b. Spei’, Thunberg s.n. (SBT; photocopy

and fragment in PRE!).

Stoebe ericoides Berg.: 339 (1767); Thunb.: 169: (1800);Thunb.: 726

(1823); Spreng.: 442 (1826). Seriphium ericoides (Berg.) Pers.: 500

(1807).

Stoebe torta Spreng.: 442 (1826). Type not seen.

D. lasiocarpa Cass.: 42 (1825); DC.: 258 (1837); Harv.: 278 (1894);

Levyns: 102 (1936a), synon. nov. Type not seen.

Wigandia disparaginoides Less.: 362 (1832). Type not seen.

D. seriphioides DC.: 257 (1837). Type: Burchell 705, not seen.

D. hoffmanniana Schltr.: 203 (1900). Type: in regio austro-oc-

cidentalis, in dictione Caledon, alt. c. 1500 ped., 21 December 1896,

Schlechter 9801 (PRE, lecto.! here designated: BOL!, GRA!).

Because of the nature and uniqueness of the cypselas

of this species it is very easy to identify, even from a

description. A fragment of the type specimen in the

Bergius Herbarium and a drawing by Mr Ears Kers con-

firmed that this name was misapplied to D. tortilis for

many years.

Small compact or larger open shrubs, 100-300(-800)

mm tall, profusely branched. Leaves filiform, 2-8 x 0.3-

0.6 mm, spirally twisted with 1—1 twists, usually

spreading but often appressed especially near inflores-

cence, mucronate, margins involute with margins touch-

ing. Synflorescences variable in robustness, number of

heads and degree of aggregation. Heads with 1 ray and

1 tubular flower; flowers strongly honey-scented. Involu-

cre of 4—6 bracts, in 2 series; bracts in inner series nar-

rowly cymbiform, acute, yellowish with reddish tinge near

apex; outer series protected by 4—6 spathulate, foliaceous

bracts. Ray flowers sterile, ray 2.5-3. 5(-4.0) x 1. 5-2.0

mm, in shades of pink, or white; cypselas reduced. Tubu-

lar flowers hermaphrodite; cypselas woolly, apical annu-

lus a loose arrangement of hairs. Pappus of 15-20

plumose hairs, connate at base.

Distribution : from Cape Point northwards to Malmes-

bury, De Dooms and Montagu and eastwards to Potberg.

It was also collected at Ystervark Point and Plettenberg

Bay. The distribution is disjunct with gaps between Hei-

delberg and Riversdale and between Mossel Bay and

Knysna (Figure 5).

204

Bothalia 23,2 (1993)

FIGURE 4. — Geographical distribution of D. anomala.

Specimens examined

CAPE. — 3318 (Cape Town): Springfontein, (-CB), Axelson 376

(NBG); Bokke River, (-CB), Hugo 974 (PRE, STE); near Melkbosch,

(-CB), Wasserlau 775 (NBG); Kenilworth, (-CD), Bolus 3297 (PRE);

Flanagan 2448 (PRE); top of Table Mountain, (-CD), Schonberg 4888

(PRE); S of Koeberg, (-CD), Pillans 6793 (BOL), s.n. (GRA); Farm

Burgers Post, near Pella, (-DA), Boucher & Shepherd 4923 (PRE, STE);

Koeberg, (-DA), Goldblatt 5313 (PRE); between Cape Town and

Malmesbury, (-DA), Montgomery 382 (STE); Kraaifontein, (-DC), Low

794 (STE); Stellenbosch, (-DD). Drege s.n. (PRE-12826). 3319 (Worces-

ter): Waterval, Tulbagh, (-AC), Drege s.n. (GRA); De Dooms, Groot

Kloof, (-BC), Stokoe 63085 (PRE); Matroosberg, N slope, (-BC), Es-

terhuysen 18713 (BOL); between Worcester and Rawsonville, (-CB),

Esterhuysen 19677 (PRE, NBG); near Edelweiss garage, Breede River,

(-CB), Walters 2394 (SAAS); Franshoek, Zachariashoek Catchment,

(-CC), Kruger 887 (STE); La Motte Forest Station, Zachariashoek, (-

CC), Vtviers 104 (STE); Franshoek Pass, (-CC), Compton 12966 (NBG);

Franshoek Pass, near Catspad memorial needle, (-CC), Koekemoer 352

(PRE); Audensberg, (-CD), Esterhuysen 28185 (BOL, PRE); Blaaskloof,

W slope of Keeromsberg, (-DA), Esterhuysen 27583 (?BOL); Jonaskop,

(-DC), Boucher 4244 (STE). 3320 (Montagu): Pypsteelfontein,

Waboomsberg, (-CA), Moffett & Steensma 4118 (STE); between

Swellendam and Barrydale, (-CC), Marsh 1135 (PRE, STE). 3418 (Cape

Town): near Muizenberg, (-AB), Bolus 3297 (GRA); Peninsula, near

Kommetjie, (-AB), Galpin 12783 (PRE); Noordhoek Mountain, (-AB),

Gillett 3568 (STE); Simonstown, top of Redhill near Pinehaven, (-AB),

Koekemoer 353 (PRE); S slopes of Chapmans Peak, (-AB). Pillans 3027

(PRE); Simonstown, Farm Bergvliet, (-AB), Purchell 90475 (BOL);

Good Hope Nature Reserve, Olifantbos road near Klein Rondevlei, (-

AB), Taylor 7632 (PRE, STE); Good Hope Nature Reserve, (-AB ), Rodin

3304 (PRE); Kommetjie, (-AB), Penfold 237 (BOL); Smitswinkel. (-

AD), Galpin 12782 (PRE); Red Hill, '(-AD), Taylor 5686 (PRE, STE);

near Platboom Street, Southern hills. Plot 33, (-AD), Taylor 6652 (PRE;

STE); Helderberg, (-BB), Galpin 12308 (PRE); Palmiet River Mouth,

(-BB), Levyns 3337 (BOL); Steenbras Valley, (-BB), Stokoe 2568 (PRE);

Cape Flats, (-BC), Ecklon 1841 (S); near Palmiet River Mouth, (-BD),

Boucher 1045 (STE); Betty’s Bay, (-BD). Boucher 1446 (PRE, STE);

Kogelberg, (-BD), Boucher 1780 (PRE, STE); Hangklip, (-BD), Comp-

ton 6071 (NBG); Rooiels, (-BD), Koekemoer 351 (PRE); Palmiet River

Valley, (-BD), Stokoe 8718 (PRE); Palmiet River, J-BD), Stokoe 58132

(PRE. BOL); Kogelberg Reserve, (-BD), Van Wilgen 108 (PRE, STE).

3419 (Caledon): Viljoen’s Pass, (-AA), Rodin 3099 (PRE); top of

Viljoen’s Pass, (-AA), Stokoe 65913 (PRE); Caledon (-AB), Bolus 9172

(PRE); Hermanus, (-AC), Compton 23223 (NBG); Kleinmond, (-AC),

De Vos 541 (STE); Onrust River Mountain, (-AC), Esterhuysen 4916

(NBG, PRE); Botrivier, Kleinmond, (-AC), Taylor 3785 (NBG); Tulbach

waterfall, (-AC), Ecklon 51 ( 1.11 ) (SAM-38472); Hermanus golf course,

(-AC), Walters 55 (PRE); Onrust River, (-AD), Compton 4916 (NBG);

Hermanus, (-AD), De Beer 23224 (PRE); Babylons Tower, (-AD), Es-

terhuysen 1941 (BOL); Hermanus, flats near Klein River, (-AD), Galpin

12764 (PRE); Shaw’s Mountain Pass, (-AD), Hafstrom & Acocks 2251

(PRE); Femkloof Nature Reserve, (-AD), Orchard 537 (PRE, STE); 1

km from Caledon on Shaw’s Pass, (-AD), Stirton 11103 (STE); Klein

River Mountain, (-AD), Stokoe 68737 (BOL, PRE); Genadendal, near

mission station, (-BA), Le\yns 4847 (PRE); Genadendal Mountains,

(-BA), Schlechter 9801 (BOL, GRA, PRE); Zondagskloof, (-BC),

Compton 10228 (NBG); Hartebees River, (-BC), Elbrecht 22151 (PRE);

Bredasdorp, (-BD), Barker 7259 (GRA, NBG); Paardenberg, (-BD),

Jordaan c!8458 (STE); The Kelders, (-CB), Van derMerwe 2033 (STE);

Danger Point, (-CB), Leighton 1560 (PRE); Danger Point, (-CB), Comp-

ton 10262 (NBG); Baardscheerdersbosch, (-DA), Compton 20454

(NBG); Elitn, (-DA), Drake 9643\ Schlechter 3515. 9643 (PRE);

Blaaskloof. Keeromsberg, (-DA), Esterhuysen 27583 (PRE); 1 km from

Viljoenshof between Elim and Die Dam, (-DA), Koekemoer 349 (PRE);

Ratel River, (-DA), Lewis 5250 (NBG); Hagel Kraal. (-DA), Lewis

63088 (BOL); between Pearly Beach and Viljoenshof, (-DA), Simpson

213 (STE); Uintjieskuil, (-DB), Van Breda & Admiraal 2353 (PRE); 3

km from Die Dam to Elim, (-DC), Koekemoer 344 (PRE). 3420

( Riversdale ): Bontebok National Park, (-AB), Grobler 592 (PRE, STE);

Koekemoer 308, 339 (PRE); Liebenberg 7970 (PRE); De Hoop, (-AD),

Van der Merwe 1362 (PRE, STE); Potberg, above Diepkloof, (-BC),

Burgers 1640b (STE); Potberg, (-BC), Taylor 4337 (PRE); Bredasdorp.

(-CA), Hafstrom & Acocks 2250 (PRE). 3421 (Riversdale): Still Bay,

(-AD), Muir 439 (PRE); Ystervarkpunt, Kampong entrance, N block,

(-BC). Willems 109 (STE).

8. Disparago kraussii Sch. Bip. in Flora 2: 693

(1844); Harv.: 278 (1894); Levyns: 100 (1936a). Type:

Cape Province, near Tsitsikama, Krauss s.n. (P-CO, spec-

imen on the far left, lecto.!, here designated).

The type, Krauss s.n., is mounted with two other speci-

mens on the same sheet in the Cosson Herbarium. The

specimen of Drege in the centre is D. laxifolia , whereas

the two others are of D. kraussii. The specimen on the

left is chosen as lectotype because the original label of

Krauss is mounted with it. All three specimens are pro-

vided with notes in Schultz Bipontinus’s handwriting.

Small shrublets, often profusely branched and dwarfed,

1 5 0—5 00 ( —800 ) mm tall. Leaves linear, 4—8 x 0.3-0.5 mm,

spirally twisted, appressed or slightly spreading, margins

involute and almost touching, mucronate at apex. Syn-

florescence globular or ovate aggregation of heads. Heads

with 1 ray and 1 tubular flower. Involucre of about 6

bracts in 1 or 2 series; narrowly cymbiform, smooth, yel-

lowish. Ray flowers sterile, ray 1. 8-2.2 x 0.7-1. 5 mm,

usually white or sometimes pink; cypselas reduced. Tu-

bular flowers hermaphrodite; cypselas puberulous, hairs

falcate, more prominent on ribs, apical annulus well de-

veloped. Pappus present in tubular flowers, of (3— )5(— 8)

plumose hairs, not coalescent; occasionally poorly devel-

FIGURE 5. — Geographical distribution of D. ericoides.

Bothalia 23,2 (1993)

205

oped and then only 1 or 2 hairs per flower and lacking

in some heads.

In Taylor 7720 a pappus, identical to that in the tubular

flowers, was present in the sterile ray flowers, whereas in

Compton 23519 , reduced stamens were present in the ster-

ile ray flowers.

Distribution : D. kraussii can be found along the south-

ern coast of the Cape from De Hoop Nature Reserve to

Still Bay (Figure 2).

Specimens examined

CAPE. — 3420 (Riversdale): De Hoop, (-AD), Barker 8743 (NBG);

Burgers 162 (PRE, STE); Van der Merwe 1181 (PRE); Die Poort,

Bredasdorp, (-AD), Compton 22609 (STE); De Hoop game camp, (—

AD), Van der Merwe 1112 (PRE); De Hoop, calcrete ridges, (-AD),

Barker 8693 (NBG); Heidelberg, Witsand, (-BB), Lexyns 10729 (BOL);

De Hoop, Farm Hamerkop, (-BC), Van Wyk 1605, 2311, 2315 (PRE);

Whitesands, (-BD), Esterhuysen 16963 (BOL); De Hoop, Witwater, (—

BD), Van Wyk 2293 (PRE); Farm Meulvlei, (-CA). Bosenberg & Ruther-

ford 272 (STE); Albertinia, (-CA), Compton 23519 (NBG); Uyshoek, 7

km N of Amiston, (-CA), Hugo 843 (PRE, STE); Bredasdorp, (-CA), Dix

41927 (SAM); Bredasdorp, Skipskop, (-CB), Acocks 22258 (PRE);

Moerasfontein, between Bredasdorp and Skipskop, (-CB). Hugo 848

(PRE, STE); De Hoop, between Buffelsfontein and Ryspunt, (-CB), Van

Wyk 2217 (PRE); flats between Bredasdorp and Struis Bay, (-CC), Levyns

3534 (BOL). 3421 (Riversdale): Plattebosch, (-AA), Muir 438(5217)

(PRE); between Riversdale and Albertinia, (-AB), Compton 23327

(NBG); Puntjie along Riversdale coast, (-AB), Esterhuysen 16976

(BOL); Still Bay, Panorama circle, (-AD), Bohnen 3823 (STE), 5122

(PRE, STE); Still Bay Reserve, (-AD), Fellingham 464 (PRE. STE);

above bridge at Still Bay strand, (-AD), Nordenstam 402 (S);

Schoemanshoek between Albertinia and The Fisheries, (-BA), Boucher

3717 (PRE, STE); Albertinia Downs, (-BA), Muir 1975 (PRE); near

Albertinia, (-BA), Muir 1978 (PRE); Cloetes Pass, Herbertsdale, (-BB),

Zinn 54598 (PRE); 'Cancase leegte', S of Albertinia, (-BC), Oliver 5722

(STE); Gouriqua, Ystervarkpunt, (-BC), Willemse 110 (STE);

Ystervarkpunt, Aulax Hill, (-BD), Willemse 162 (STE). 3422 (Mossel

Bay): Cloete's Pass, (-AA), Lewis 5406 (NBG); hill 2.5 km W of Mossel

Bay near reservoir, (-AA), Vlok 1423 (STE); George, Gwaing River,

(-AB), O’Callagan, Fellingham & Van Wyk 226 (STE); George, Chris-

tina Bay, (-BA), Schlechter 2449 (BOL, G, K, PRE, S, STE); Knysna,

Buffels Bay, (-BB), Keet 960 (PRE); Sedgefteld, (-BB), O'Callagan 576

(STE). 3423 (Knysna): Lake Pleasant Hotel, (-AA). Acocks 21221

(PRE); Groenvlei, (-AA), Levyns 10310 (BOL); Robberg Nature Re-

serve, highest point, (-AB ), Taylor 7720 (STE); Robberg Nature Reserve,

top of central dune, (-AB), Taylor 7720a (STE); Plettenberg Bay,

Robberg, (-AB), Vdjoen 62 (PRE).

9. Disparago laxifolia DC., Prodromus systematis

naturalis regni vegetabilis 6: 257 (1837); Harv.: 278

(1894); Levyns: 101 (1936a). Type: ‘ad Cap. Bonae-Spei

in distr. Caledon ad Zwartberg et Kleinrivier’s-berge',

Ecklon s.n. (G-DC in herb. Dunant, holotype; photo and

microfiche in PRE!).

Small trailing shrublets, 100-300 mm tall, branches

usually lax and far apart. Leaves linear, 3— 8(— 10) x 0.6-

2.5 mm, widely spaced, spirally twisted, usually with Vl-l

twist, spreading, older leaves reflexed; margins slightly

involute, apices blunt. Synflorescence globular aggrega-

tion of heads. Heads with 1 ray and 1 tubular flower (a

single case with 2 ray flowers was observed). Involucre

of about 6 bracts in 2 series, inner series chaffy, narrowly

cymbiform, outer series foliaceous, spathulate. Ray flow-

ers sterile, ray 1.5-2.0 x 1.0-1. 5 mm, white; cypselas re-

duced. Tubular flowers hermaphrodite; cypsela surface

puberulous, apical annulus a well-developed fringe of

hairs. Pappus present in tubular flowers, of 5 plumose

hairs, not connate.

Distribution : D. laxifolia has a very local and scattered

distribution that stretches from Simonstown to the

Houwhoek Mountains, Grabouw, Caledon and Hermanus.

It is very common in the Femkloof Nature Reserve, near

Hermanus (Figure 2).

Specimens examined

CAPE. — 3418 (Simonstown): Redhill, (-AB), Levyns 5903 (BOL);

Salter 5703 (BOL, K); Peninsula, Sirkelsvlei, (-AD), Esterhuysen 7793

(BOL); Sir Lowry's Pass (-BB), Drege s.n. (P); Schlechter 595 (PRE),

7223 (BOL, GRA. S); Steenbras area, (-BB), Levyns 8753 (BOL); Sir

Lowry's Pass, Palmiet River, (-BB), Penther 1415 (S): Cape Point Gap,

Klawer Valley, (-BB), Salter 6426 (BOL, K); near Somersfontein, (-BD),

Boucher 720 (PRE, STE); Betty’s Bay, (-BD), Esterhuysen 34768

(BOL). 3419 (Caledon): Grabouw, Palmiet River, (-AA), Bolus 5077

(BOL); Bond 1531 (NBG); Esterhuysen 35273 (BOL); Levyns 5358,

7778 (BOL); Stokoe 63084 (SAM); Palmiet River Valley, Platteberg,

(-AA), Stokoe 65911 (SAM); Houwhoek, (-AA), Esterhuysen 33688

(BOL); Galpin 4152 (GRA. PRE); Levyns 11404 (BOL); Schlechter 5447

(BOL, G, GRA, K, PRE, S), 5506 (GRA, PRE); Good Hope Reserve.

(-AB), Wright s.n. (K); Palmietberg, E of bridge, (-AC), De Vos 1431

(PRE. STE); Hermanus, Vogelgat Reserve, (-AC), Esterhuysen 35540

(BOL); Hermanus, Femkloof Nature Reserve, (-AC). Koekemoer 192

(PRE); Palmiet River mouth, (-AC), Koekemoer 439 (PRE); Femkloof

Nature Reserve, (-AD). Orcltard 321 (STE); Vogelgat, Sea Saddle, (—

AD), Wdliams 2909 (NBG): Caledon, Highlands, (-DB ), Compton 12259

(NBG); Highlands Estate, Elgin, (-DB), Lewis 63083 (SAM).

Excluded species

Disparago gomphrenoides Sch. Bip.: 180 (1861). Type:

C.B.S., Zeyher 46 (P!) = Stoebe sp.

ACKNOWLEDGEMENTS

I wish to thank the following: all the curators of herba-

ria who sent specimens on loan; the NBI, my employers

for creating opportunities for research and for financial

support; my colleagues, for advice and moral support; A.

Romanowski, for the developing and printing of photo-

graphs; Dr S.M. Perold for assistance with the SEM; li-

brary staff for handling numerous requests. Dr O.A.

Leistner for translating the diagnoses; E. du Plessis for

proofreading the manuscript; Prof. B-E. van Wyk, for su-

pervising my M.Sc. thesis on Disparago.

REFERENCES

BENTHAM, G. 1873. Compositae. In G. Bentham & J.D. Hooker,

Genera plantarum 2: 163-533. Reeve, London.

BERGIUS, PJ. 1767. Descriptiones plantarum ex capite bonae spec.

399-440. Salvius, Stockholm.

BOLUS, H. 1909. Contributions to the African Flora. Transactions of

the South African Philosophical Society 18: 378^-00.

BOLUS, H. & WOLLEY-DOD, A. 1 904. A list of flowering plants and

ferns of the Cape Peninsula, with notes on some of the critical

species. Transactions of the South African Philosophical Soci-

ety 14: 207-280.

CASSINI, A.H.G. 1819. Dictionaire des sciences naturelles 13: 348.

349.

CASSINI, A.H.G. 1 825. Dictionaire des sciences naturelles 34: 42, 43.

DE CANDOLLE, A.P. 1837. Prodromus systematis naturalis regni

vegetabilis 6: 257, 258. Treuttel & Wiirtz, Paris.

DYER, R.A. 1951. Disparago ericoides. The Flowering Plants of

Africa 28: t. 1102. Government Printer, Pretoria.

DYER, R.A. 1975. The genera of southern African flowering plants.

Government Printer, Pretoria.

GAERTNER. J. 1791 . Fructibus et seminibus plantarum 2: 463, t. 173.

Schramm, Tubingen.

206

Bothalia 23,2 (1993)

HARVEY, W.H. 1894. Compositae. In W.H. Harvey & O.W. Sonder,

Flora capensis 3: 277 , 278. Hodges, Smith, Dublin.

HUTCHINSON, J. 1932. Notes on the flora of southern Africa: III.

Miscellaneous new species. Kew Bulletin 10: 465-512.

KOEKEMOER, M. 1991. New species in the genus Disparago.

Bothalia 21: 158-161 .

KOEKEMOER, M. in prep. a. Generic and infrageneric relationships

in the genus Disparago (Asteraceae). South African Journal of

Botany.

KOEKEMOER, M. in prep. b. Speciation and biogeography in the

genus Disparago (Asteraceae) — with reference to the distribu-

tion of related genera. South African Journal of Botany.

LESSING, C. F. 1832. Synopsis generum compositarum'. 362, 363.

Black, Young & Young, London.

LEVYNS, M.R. 1936a. A revision of Disparago Gaertn. Journal of

South African Botany 2: 95-103.

LEVYNS, M.R. 1936b. Notes on some members of Compositae.

Transactions of the Royal Society of Southern Africa 23: 91-95.

NECKER, N.J. DE 1790. Elementa botanica. Societatem Typo-

graphicam, Neuwied.

PERSOON, C.H. 1807. Synopsis plantarum : 500. Treuttel & Wiirtz,

Tubingen.

RICKETT, H.W. & STAFLEU, F.A. 1960. Nomina generica con-

servanda et rejicienda spermatophytorum 6. Taxon 9: 153-161.

SCHLECHTER, R. 1900. Plantae Schlechterianae novae vel minus

cognitae describuntur. II. Botanische Jahrbiicher 27: 203, 204.

SCHULTZ BIPONTINUS, C.H. 1844. Enumeratio Compositarum.

Flora 2: 692, 693.

SCHULTZ BIPONTINUS, C.H. 1861. Cassiniaceae uniflorae. Jahres-

bericht der Pollichia 18/19: 24(180), 25(181).

SPRENGEL, K.P.J. 1826. Systema vegetabilium 3: 442.

THUNBERG, C.P. 1800. Prodromus plantarum capensium: 169.

Edman, Uppsala.

THUNBERG, C.P. 1823. Flora capensis'. 726. Stuttgart.

Bothalia 23,2: 207-214(1993)

Studies in the Marchantiales (Hepaticae) from southern Africa. 2. The genus

Athalamia and A. spathysii ; the genus Oxymitra and O. cristata

S.M. PEROLD*

Keywords: Athalamia, A. spathysii , Cleveaceae, Hepaticae, Marchantiales, Oxymitra, O. cristata, Oxymitraceae, southern Africa, taxonomy

ABSTRACT

The genera Atltalamia (Cleveaceae) and Oxymitra (Oxymitraceae), each respectively represented in southern Africa by a

single species, A. spathysii (Lindenb.) Hattori and O. cristata Garside ex Perold, are discussed.

UITTREKSEL

Die genusse Athalamia (Cleveaceae) en Oxymitra (Oxymitraceae), wat in Suider-Afrika elk respektiewelik verteenwoordig

word deur 'n enkele spesie, A. spathysii (Lindenb.) Hattori en O. cristata Garside ex Perold, word bespreek.

ATHALAMIA

Athalamia Falconer in Annals and Magazine of

Natural History, Ser. 2,1: 375 (1848); ibid.: 397 (1851);

Shimizu & Hattori: 52 (1954); S. Amell: 57 ( 1963). Type

species: Athalamia pinguis Falconer.

Clevea Lindb. 9: 289 (1868); Steph.: 769 (1898);

Schiffn.: 29 (1893); Howe: 36 (1899); K. Mull.: 368

(1951-1958); Hassel de Menendez: 133 (1963). Type spe-

cies: Clevea hyalina (Sommerf.) Lindb.

Spathysia Nees ex Trevis.: 439 (1877). Type species:

Spathysia lindenbergii Trevis. nom. illeg.

Gollaniella Steph.: 74 (1905). Type species: Golla-

niella pusilla Steph.

Thallus medium-sized, medianly concave, bright green,

in crowded patches; on soil in rocky clefts or under over-

hangs. Branches simple or once pseudodichotomously fur-

cate; thickened over midrib, thinning toward undulate,

attenuate margins; apex hardly notched, dorsally not

grooved. Dorsal epidermis hyaline, cell walls sometimes

thickened at comers. Air pores simple, small, slightly ele-

vated, stellate or hardly so, surrounded by a ring of cells,

the radial walls generally thickened, leading below into

individual empty air chambers, these in one or several

layers, bounding walls chlorophyllose, rarely with smaller

cells containing a single, large oil body; storage tissue

with rounded cells fairly closely packed together. Rhizoids

both smooth and pegged. Scales hyaline or purple red, in

2 or more forwardly directed ventral rows, extending be-

yond thallus margins or not, obtusely triangular, append-

age long-acuminate.

Monoicous. Antheridia embedded in slightly raised

central cushion, or all along midline, opening into pro-

jecting, conical papillae. Archegoniophores dorsal, single

^National Botanical Institute, Private Bag X101, Pretoria 0001.

MS. received: 1992-06-03.

to several along midline, on short unfurrowed stalk, fili-

form scales at its top, involucres 1 — 3, basally connate,

each with single, obliquely erect capsule, wall dehiscing

by irregular valves, cells with thickening bands. Spores

rounded distally, somewhat flattened proximally, lacking

triradiate mark, densely covered with conical spines. El-

aters long, tapering, bispiral.

Athalamia spathysii (Lindenb.) Hattori in Shimizu

& Hattori in Journal of the Hattori Botanical Laboratory

12: 54 (1954); S. Amell: 57 (1963); Vanden Berghen: 168

(1965); Volk: 230 (1979). Type: Greece, Corfu, leg.

Spathys (W, holo.!; STR, iso.).

Marchantia spathysii Lindenb.: 104 (1829); Bisch.: 1018 (1835).

Dumortiera spathysii (Lindenb.) Nees: 171 (1838); Gott. et al.\ 546

(1846). Clevea spathysii (Lindenb.) K. Mull.: 75 (1940); K. Miill.: 374

(1951-1958).

Plagiochasma rousselianum Mont.: 334 (1838). Clevea rousseliana

(Mont.) Leitgeb in Steph.: 771 (1898). Type: Algeria. 'Boudjareah’,

Roussel.

Thallus medium-sized, slightly concave along middle,

oblong to broadly ovate or obovate (Figure 1A), bright

green, sometimes purple along margins, polygonal out-

lines of subdorsal air chambers clearly visible from above,

i.e. reticulate dorsally, central areolae at apex small, en-

larging toward margins and then in parallel, radiating

rows; air pores tiny, slightly raised, singly placed over

each polygonal area, wet; thallus margins clasped together,

revealing deep purple, transversely wrinkled underside of

wings, dry; in rather crowded patches, simple or once

pseudodichotomously furcate. Branches (3.5-) 5.0-12.0 x

2.5-1. 5 mm, 825-1075 pm thick over midrib, laterally

thinning out into attenuate wings; apex slightly notched

and dorsally not grooved, with tips of 2 or 3 purple-red

ventral scales recurving over edge; margins acute, thin,

scalloped, undulate, often somewhat erect, older parts

dead, ventrally purple; flanks sloping obliquely; ventral

face medianly keeled, green, midrib with row of purple-

red scales on either side (Figure 1C).

208

Bothalia 23,2(1993)

FIGURE 1. — Athalamia spathysii. Thalli. A, dorsal view of thallus with stalked archegoniophore and several young sessile ones; inset with

archegoniophore and 2 dehiscing sporangia; B, dorsal view of thallus with rows of antheridia; C, ventral view of thallus; D, transveise section

of thallus; E, F, dorsal air pores, with thickened radial walls; G, transverse section of air pore; H, ventral scale; I, transverse section of stalk;

J, scale from top of stalk; K capsule wall with cells containing annular thickenings; L, transverse section of capsule wall. A, I-L, Volk 00589\

B Volk 61 24\ C-H, Volk 00904. Scale bars: A-C, 2 mm; D, 1 mm; E-G, K, L = 50 pm; H, J = 200 pm; I = 100 pm. Illustration by A. Pienaar.

Bothalia 23,2(1993)

209

Dorsal epidermal cells unistratose, hyaline, polygonal

or isodiametric, (32.5-)57.5-62.5 x 30.0-42.5 pm, thin-

walled, but frequently thickened at comers, in transverse

section 40 pm thick; air pores very slightly raised (Figure

1G), simple, small, 7.5-12.5 pm wide, 65-150 pm distant

from each other and usually bounded by innermost hyaline

circle (Figure IF), the remains of a collapsed cell ring,

otherwise surrounded by a ring of 4 — 6(— 7) bluntly trian-

gular cells, 15.0 x 22.5-32.0 pm, the radial walls mostly

thickened and pores thus stellate (Figure IE, IF), partly

overlying adjacent dorsal cells. Assimilation tissue 350-

500 pm thick with air chambers empty, in one layer or

sometimes in several layers, 50-95 pm wide, vertical or

sloping forward, oblique laterally (Figure ID), unistratose

walls consisting of chlorophyllose cells, roundish or oval,

40.0-62.5 x 45.0 pm; storage tissue ± 600 pm thick, cells

fairly densely packed together, rounded, up to 62.5 pm

wide; rhizoids mostly smooth, 25-35 pm wide. Scales

purple-red, arranged in 2 forwardly directed ventral rows,

one on either side of midrib, not or somewhat extending

beyond margins of thallus, recurved at apex, obtusely tri-

angular (Figure 1H), margins entire or sometimes slightly

dentate on oblique side of base, not constricted where

joined with acuminate, apically pointed and tapering ap-

pendage, total length (including 750 pm appendage) ±

1300 pm, width at base up to 1000 pm, cells (4-)5- or

6-sided, 90-137 x 40-50 pm; oil bodies absent.

Monoicous, but occasionally only antheridia or arche-

gonia found. Antheridia , when present with archegonia in

a group, proximal to archegoniophore, embedded in cen-

tral, slightly raised, ill-defined cushion lacking scales, oth-

erwise in rows along entire middle of thallus (Figure IB),

sunken, saccate, 500 x 230 pm, narrowed at neck, opening

into raised, conical papillae projecting 200 pm above sur-

face and 300 pm wide at base. Archegoniophores single

or several, linearly and acropetally arranged on dorsal

face, when young sessile, rounded, and basally surrounded

by inconspicuous, filiform scales, air pores and air cham-

bers lacking; stalk cylindrical, pellucid, eventually 1 .6-7.0

mm long, round to ovate in cross section (Figure II). di-

ameter 250 pm, without rhizoidal furrow; scales eventu-

ally carried upwards and then only present at top of stalk,

where joined to archegoniophore, hyaline or pink, basally

750 x 500 pm, with 4 or 5 filiform, apical strands (Figure

1 J), up to 750 pm long, cells 75 x 48 pm; archegoniophore

white to greenish white, central disc absent, with 1-3 in-

volucres, connate at their bases and attached to top of

stalk, bilabiate by vertical cleft, each with single capsule,

brown, globular, 1250 pm in diameter, borne obliquely

erect on 850 x 600 pm seta with bulbous foot, dehiscing

by several irregular valves, wall unistratose (Figure 1L),

cells spindle-shaped, up to 75 x 30 pm, with annular or

spiral thickening bands (Figure IK). Spores (60— )65— 75(—

78) pm in diameter, globular, with distal face convex (Fig-

ure 2A, B), no triradiate mark on somewhat flatter

proximal face, golden brown, semitransparent, thickly

covered with dense conical spines (Figure 2A-D), 5 pm

high and 15 pm wide at base, sprinkled with fine granules

(Figure 2E). Elaters yellow-brown, slightly tapering to-

ward ends, 275-350 pm long, 7.5-10.5 pm wide in mid-

dle, bispiral (Figure 2F), very occasionally branched,

sometimes bent. Chromosome number: n = 9 (Bomefeld

1987); as Clevea rousseliana: n = 9 (Heitz 1927).

DISCUSSION

Athalamia spathysii is rarely collected in southern Afri-

ca, the gatherings by Volk ( 1979) being restricted to Nami-

bia (Figure 3). It grows in periodically dry areas, on soil

overlying slate, granite or sandstone, in rocky crevices or

under overhangs, where runoff may be concentrated and

some protection against radiation is afforded; sometimes

it occurs together with Plagiochasma spp., Targionia

FIGURE 2. — Athalamia spathysii. SEM micrographs of spores. A, B. distal face; C, side view of distal face; D, side view of both faces; E, spinous

processes much enlarged; F, elater. A-F, Volk 00589. A-D, x 580; F. x 1000.

210

Bothalia 23,2 (1993)

FIGURE 3. — Distribution of Athalamia spathysii, •; and Oxymitra

cristata , ♦ in southern Africa.

hypophylla etc., but is distinguished from them by the

reticulate dorsal surface, small and slightly raised, mostly

stellate air pores, empty air chambers, scales with acumi-

nate appendages, cells of the capsule wall with spiral

thickenings and by bluntly spinous spores.

It has also been reported from further north in Africa,

i.e. fromTanzania (= Tanganyika) by Vanden Berghen

(1965). Athalamia spathysii is furthermore known from

Israel, North Africa, the Canary Islands and the Mediter-

ranean countries (Muller 1951-1958), the type specimen

having been collected at Corfu. Frey & Kiirschner (1988)

report it from Yemen and Oman in the Arabian Peninsula,

and state that its distribution includes Turkey and Jordan.

Athalamia is classified in the family Cleveaceae Cav-

ers, together with the genera Peltolepis Lindb. and Saute-

ria Nees, but neither of the latter two occur in southern

Africa.

There are at least 10 species world-wide in the genus

Athalamia, with A. spathysii the only representative in

southern Africa, A. pulcherrima (Steph.) Hatt. being from

Ethiopia and distinguished by larger, hyaline ventral scales

and cells with much thickened radial walls surrounding

the dorsal air pores. Another species recorded from Africa,

in Algeria, is Clevea (Athalamia) trabutiana Steph., but

it is regarded by Grolle (1976) as conspecific with A.

hyalina (Sommerf.) Hatt. In A. spathysii the radial walls

of the pore cells are only somewhat thickened, a possible

response to growing in sunlight, as was found in cultiva-

tion studies by Muller (1951-1958). Shimizu & Hattori

(1954), however, reported for this genus, that they had

observed pores with both thickened radial walls (stellate)

or with thin walls (not stellate) on the same plant!

According to Muller (1939), Arnell (1963), Vanden

Berghen (1968) and Volk (1979) oil bodies are absent in

the genus Athalamia. I also found them to be absent in

A. spathysii , but fresh material was not available for study.

Shimizu & Hattori (1954), however, reported oil cells in

Athalamia glauco-virens as well as in A. nana (Shimizu

& Hattori 1955). Schuster (1992) states that sparse, scat-

tered oil cells may be present or absent.

Asexual reproduction is said to be absent. Stephani

(1895) suspected that adventitious growth in the form of

ventral shoots from the costa was partly the reason for

the ‘dichten verfilzten niedrigen Rasen’ in Clevea pul-

cherrima Steph. As in Plagiochasma spp., the apical part

of the thallus continues to grow and increase in length,

beyond the archegoniophore after the latter has differen-

tiated.

Kashyap (1915) described a mycorrhizal region in the

midrib of A. pinguis, the hyphae having penetrated the

thallus via the rhizoids. Sterile specimens of A. spathysii

have been confused with Asterella muscicola (Arnell

1957), because of the stellate dorsal pores, but the latter

does not grow in Namibia (Volk 1979), its ventral scales

have scattered oil bodies, the stalk has a rhizoidal furrow

and the capsule wall lacks thickenings.

Specimens examined

NAMIBIA. — 2217 (Windhoek): WIN 63 (Neudamm), schattige

Glimmerschieferfelsen am Rivier, (-AD), Volk 00952 (BOL; PRE); un-

terhalb Briicke, Glimmerschiefer, (-AD), Volk 86-877 (Herb. Volk); 1

mile west of Windhoek, (-CA), Schelpe 4763 (BOL); WIN 62,

Hatsamas/Dordabis, Kalkfels, (-DC), Volk 00589 p.p. (Herb. Volk). 2218

(Gobabis): WIN 70, schattiges Talchen an den Gobabisstrasse,

Glimmerschiefer, (-??), Volk 00904 (BOL; PRE). 2316 (Nauchas): WIN

39 (Mahonda), schattige Uberhange, Glimmerschiefer, (-BA), Volk 6124

(PRE).

OXYMITRA

Oxymitra Bisch. ex Lindenb., Synopsis hepati-

carum europaearum: 124 (1829); Gott., Lindenb. & Nees:

597 (1846); Frye & Clark: 40 (1937); K. Mull.: 410

(1951-1958); Hassel de Menendez: 200 (1963); Arnell:

11 (1963). Lectotype species: O. incrassata (Brotero)

Sergio & Sim-Sim.

Pycnoscenus Lindb.: 606 (1863), nom. illeg.

Tessellina Dumort.: 164 (1874), non Dumort.: 78

(1822) orth. var.: Tesselina Dumort. mut., Schiffn.: 15

(1893). Type species: Tessellina coriandrina (Spreng.)

Dumort.

Thallus medium-sized, deeply and sharply grooved

dorsally, pale green to deep green, in gregarious or ro-

sette-like stands; rare, on soil, sometimes derived from

ironstone. Branches broadly ovate or oblong, pseudo-

dichotomously furcate; thickened over midrib, thinning to-

ward margins; apex emarginate. Dorsal epidermis hyaline,

persistent. Air pores tiny, simple, radial walls thickened,

stellate, leading below into tall, narrow, subvertical, empty

air chambers, bounded by unistratose, chlorophyllose cell

walls. Storage tissue with rounded or angular cells, closely

packed together; rhizoids numerous, some smooth, others

pegged. Scales large and conspicuous, arising ventrally

and projecting far above thallus margins, triangular, dark

red to almost black or hyaline.

Monoicous or dioicous. Antheridia inedianly grouped

behind archegonia or mixed with them when monoicous,

embedded, with hyaline necks projecting, at their bases

surrounded by capillary hairs, vestigial or prominent. Ar-

chegonia several in a row along midline, flask-shaped,

protected in a continuous, crest-like, fleshy involucre, or

Bothalia 23,2(1993)

211

individually enclosed in conical or pyramidal, chambered

involucres with scales around the base. Sporangia without

stalk or foot, globose, capsule wall hyaline, unistratose,

delicate. Spores triangular-globular, anisopolar, black,

opaque; distal face with several large areolae, containing

a central nodule or not; proximal face with triradiate mark

distinct or not, irregularly reticulate or smooth. Elaters

absent, but some sterile cells reputed to be present.

Oxymitra cristata Gar side ex Perold , sp. nov.

Type: Cape, near Bredby Mine (25 miles S of Kuruman),

Schelpe 5900 (BOL, holo.).

Frondes caespitosae confertae, virides; squamae

ventrales atro-violaceae, oblique triangulares, marginem

frondis bene superantes. Antheridia in pulvillos medianos

lineares immersa. Archegonia ante antheridia posita, in

serie unica lineari disposita atque in involucra connata im-

mersa, cristam medianam prominentiam crenatam for-

mantia, bracteae desunt. Sporae 100-115 Jim diametro:

exterior (distalis) facies irregulariter reticulata, interior

(proximalis) facies laeves.

O. incrassata, species altera generis, archegoniis in in-

volucris conicis vel pyramidalibus singulatim inclusis,

bracteis multis concomitatis ab O. cristata differt.

Thallus smallish to medium-sized, apically emarginate,

dorsally deeply grooved along entire midline (Figure 4A,

C), except where interrupted by gametangia, broadly ovate

or oblong, bright green to deep green, dark red along mar-

gins, sometimes bronze-brown proximally, forming a

broad, deeply coloured ‘V’ over base, reticulate from

above, with outlines of subdorsal air chambers clearly vis-

ible, air pores tiny, hardly raised (Figure 4G), numerous

but singly placed above each polygonal air chamber, wet;

thallus margins incurved or raised and tightly clasped to-

gether, flanks covered by large, triangular, shiny dark red

scales, dry; in crowded, overlying patches, once to several

times pseudodichotomously furcate. Branches (3— )5 — 10(—

12) x 2.5^4.8 mm, 1 100-2000 |im thick over midrib; mar-

gins rather obtuse to acute, scalloped or undulating, much

exceeded by projecting, triangular scales; flanks deep pur-

ple, steeply rising (Figure 4C), to sloping obliquely up-

ward and outward (Figure 4D); ventral face rounded to

flattish, green (Figure 4B).

Dorsal epidermal cells unistratose, hyaline, from above

polygonal, (22.5-)37.5-45.0(-52.0) x 27.5-37.5(-47.5)

pm, in transverse section brick-shaped, 25-35 pm thick.

Air pores stellate (Figure 4H), small, simple, slightly

raised (Figure 4G), 5-10 pm wide, 4- or 5-(6)-sided (Fig-

ure 4H), 75-112 pm distant from each other, bounded by

4 or 5, rarely 6, small, bluntly wedge-shaped cells, 12.5-

15.0 x ± 25 pm across base, radial walls slightly thick-

ened, partly overlying, or occasionally adjoining larger,

polygonal cells. Assimilation tissue 400-600 pm thick

with air chambers empty (Figure 4F), in one layer, 32-65

(-100) pm wide, bounding walls unistratose, cells 37^17

x 25-35 pm; storage tissue composed of rounded or an-

gular cells, ± 50 pm wide. Rhizoids numerous, 17-25 pm

wide, some pegged, others smooth. Scales arising ven-

trally and projecting up to 375 pm above thallus margins,

dark red to almost black, shiny, basal part of apical scales

hyaline, densely imbricate, obliquely triangular (Figure

41), 1250-1375 pm long, 1075-1200 pm wide across

base, margins entire, sometimes crenate or shortly toothed

toward base of forwardly directed side, cells in body of

scale (4-)5- or 6-sided, ± 100 x 25 pm, smaller at rounded

margin.

Monoicous. Antheridia in a row along midline, proxi-

mal to archegonia (Figure 4A, E), immersed, necks hya-

line, cylindrical, 75 pm wide, protruding 210 pm above

dorsal surface. Archegonia in a median row, in flask-

shaped cavities, partly sunken into thallus and as a group

enclosed above by a 600 pm high, irregular, crest-like,

fleshy ridge (Figure 4D), tinged reddish on top around

openings of archegonial necks, these long and turning pur-

ple with age. Sporangia ventrally partly sunken into thal-

lus, dorsally bulging on either side of central crest, capsule

± 750 pm wide, wall hyaline, thin and delicate. Spores

( 108— )1 10—1 15(— 1 25) pm in diameter, triangular-globular,

polar, black, opaque, wing ± 5 pm wide, entire, anisopolar,

with ornamentation different on two faces: distal face

(Figure 5A-C) with 6 or 7 large central areolae, up to 30

pm wide, smaller toward margin, generally with a nodule

and thin radiating ridges in the middle of each, areolar

walls thick; proximal face with triradiate mark distinct

(Figure 5D, E), 3 facets with incomplete areolae, walls

irregular (Figure 5E, F), sometimes rather indistinct (Fig-

ure 5D). Chromosome number n = 18 (Baudoin 1976).

DISCUSSION

Grolle (pers. comm.) has pointed out to me that O.

cristata Garside was invalidly published as it did not com-

ply with Art. 37.1 of ICBN because it was published after

1 January 1958 without indication of the holotype.

Garside’s paper was published in April 1958 and two type

specimens were cited: Schelpe 5900 and Duthie 5531. but

no holotype was designated. Oxymitra cristata Garside ex

Perold is therefore accordingly newly described here, to-

gether with a Latin diagnosis; Schelpe 5900 is designated

as the holotype.

Oxymitra cristata is endemic to southern Africa, and

very rarely collected. Garside (1958) remarked that ‘the

detailed geographical distribution is not yet completely

known’ and this is still true today, only a few more col-

lections having been made during the intervening 35

years. It is known from Namibia, Orange Free State and

northern and eastern Cape (Figure 3). It has been found

mainly on substrates derived from ironstone, and it occa-

sionally grows in association with Plagiochasnra spp. and

Riccia spp. Oxymitra cristata is distinguished by its large,

obliquely triangular, shiny, dark red scales and by the row

of archegonia enclosed in an irregular, but continuous,

crest-like, fleshy ridge of tissue. Garside (1958) regarded

the ridges as composed of fused involucres. In the only

other species in the genus, O. incrassata (Brotero) Sergio

& Sim-Sim, better known as O. paleacea Bisch. ex

Lindenb. (Sergio & Sim-Sim 1989), each archegonium is

individually enclosed in a conical or pyramidal involucre,

containing air chambers and air pores. These involucres

are arranged in two rows along the midline and are ac-

companied by numerous bracts. Oxymitra incrassata also

differs from O. cristata by its hyaline ventral scales, by

its spore ornamentation, with the areolae on the distal face

lacking a central nodule and by the smooth facets on the

212

Bothalia 23,2 (1993)

FIGURE 4. — Oxymitra cristata. Thalli. A, dorsal view of thallus with crest-like involucre and, proximally, row of antheridia; B, ventral view of

thallus; C, transverse section of thallus through dorsal groove; D, transverse section of thallus through involucre with archegonium; E,

transverse section through antheridia, dorsal cells and air chambers; G, transverse section through much enlarged air pore; H, air pore and

surrounding cells seen from above; I, ventral scale. A, B, D, Volk 00906 ; C, F— 1, Volk 81/050', E, Volk 00957. Scale bars: A-E, 2 mm; F-H,

50 pm; 1, 500 pm. Illustration by A. Pienaar.

Bothalia 23,2(1993)

213

FIGURE 5. — Oxymitra cristata. SEM micrographs of spores. A, B, distal face; C, side view of distal face; D. E, proximal face; F, side view of

proximal face with areolar walls more pronounced. A, D, Volk 00906 ; B, C, E, F, Volk 5050. A, B, D, E, x 320; C, F, x 345.

proximal face, by the chromosome number n = 9 and by

its distribution: it is known from Europe, North Africa,

the Canary Islands, North and South America.

Bischler (1988) records the presence of tubers in Oxy-

mitra, but these have not been observed in the present

investigation of O. cristata. The two species of Oxymitra

are placed in the family Oxymitraceae K. Mull, ex Grolle.

The sporangia, partially embedded in the thallus tissue,

the loss of a foot and seta of the capsule, the absence of

elaters, spore release only after destmction of surrounding

tissue and the sporeling type (Baudoin 1976), all indicate

the relationship of Oxymitraceae to Ricciaceae, and they

have been classified together in the suborder Ricciineae

(Schuster 1984), although Schier (1974), on the basis of

biochemical studies, had segregated the Oxymitraceae into

their own suborder, the Oxymitrineae.

Oil bodies have not been observed in O. cristata , but

Muller (1939) reports their presence (after Gyorffy) in the

storage tissue of O. incrassata during spring and autumn.

The generic description has had to be emended somewhat

to accommodate the gametangial differences in the two

species, the first set of particulars in each case referring

to O. cristata.

SPECIMENS EXAMINED

NAMIBIA. — 2216 (Otjimbingwe): OM 37 Otjua, (-AA), Volk 00957

(herb. Volk). 2217 (Windhoek): Voigtland, feuchte Kliifte, (-CB), Volk

5050 (herb. Volk). 2218 (Gobabis); WIN 70, schattiges Talchen,

Glimmerschiefer (+ Plagiochasma sp. & Athalamia spathysii), (-??), Volk

00906 (PRE, herb. Volk).

O.F.S. — 2826 (Brandfort): Glen Agricultural College, on stony kop-

pie, (-CD), Zietsman 943 (PRE). 2926 (Bloemfontein): Bloemfontein,

Botanical Garden, not cultivated part, on hill, (-AA), Volk 81/050 (PRE);

Bloemfontein, next to Botanical Garden, shallow soil over rock, (-AA),

Volk 81/291 (PRE); Bloemfontein, near Hillandale, (-AA), Gemmell

( =Duthie 5501 ) (BOL); Bloemfontein. Eagle’s Nest, on ironstone kopje,

in shelter of grass and boulders, (-AA) Gemmel & Lutjeharms ( =Duthie

5460 ) (BOL).

CAPE. — 2723 (Kuruman): near Bredby Mine, 25 miles S of Kuru-

man, (-CA), Schelpe 5900 (BOL) (holotype). 2822 (Glen Lyon):

Olifantshoek area, on farm belonging to Mr Rossouw, on stony koppie,

with Riccia spp. and Plagiochasma sp. (-BA), Koekemoer 1024 (PRE);

Griqualand West, Groenwater Valley, Hay, east of Postmasberg (sic)

(Postmasburg], (-DD), Cooke s.n. ( =Duthie 5531) (BOL). 3027 (Lady

Grey): Herschel, mudbanks of Sterkspruit, with Riccia sp., (-CB), Hep-

bum CH 1043 (PRE).

ACKNOWLEDGEMENTS

I wish to thank the following: Dr R. Grolle, Jena, for

critically reading the manuscript and for his information

regarding the earlier publication date of Athalamia by Fal-

coner as well as for drawing my attention to the invalidity

of Oxymitra incrassata Garside, which I had missed in

Geissler & Bischler (1989); Prof. T. Poes, Eger, for also

refereeing the manuscript and Prof, (emer.) Dr O.H. Volk,

Wurzburg University, for the loan of some of his speci-

mens. My thanks also to the curator of BOL for the loan

of specimens; and to Ms M. Koekemoer, NBI, for col-

lecting specimens; to the artist, Ms A. Pienaar; the typist,

Mrs J. Mulvenna, and the photographer, Mrs A.

Romanowski, for their valued contributions.

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FRYE, T.C. & CLARK, L. 1937. Hepaticae of North America. University

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GARSIDE, S. 1958. Studies in South African Ricciaceae. III. A new

species of Oxymitra. The Journal of South African Botany 24:

83-87.

GEISSLER, P. & BISCHLER, H. (Eds) 1989. Naiadea to Pycnoscenus.

Hepaticarum 11: 353.

GOTTSCHE, C.M., LINDENBERG, J.B.G. & NEES AB ESENBECK

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printed 1967, Cramer, Lehre.

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ter Gebiete. Feddes Repertorium 87: 171-279.

GROLLE, R. 1983. Nomina generica hepaticarum: references, types and

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Bothalia 23,2: 215-221 (1993)

Studies in the M archantiales (Hepaticae) from southern Africa. 3. The genus

Targionia and T. hypophylla with notes on T. lorbeeriana and Cyathodium

foetidissimum (Targioniaceae)

S.M. PEROLD*

Keywords: Cyathodium , Hepaticae, Marchantiales, southern Africa, Targionia , Targioniaceae, T. hypophylla , T. lorbeeriana, Targionioideae

ABSTRACT

A taxonomic account of the genus, Targionia, and its only representative in southern Africa, the almost cosmopolitan T.

hypophylla , is given here. Much more southern African and other material, than was available to the previous authors, Sim

(1926) and Amell (1963) has been examined.

Material collected at Knysnaby Duthie was identified by Arnell (1963) as Cyathodium foetidissimum, but is shown to have

been misidentified and is actually a specimen of Riccia rubricollis. The presence of the genus, Cyathodium in southern Africa,

has not been confirmed.

U1TTREKSEL

’n Verslag oor die genus Targionia en sy enigste verteenwoordiger in Suider-Affika, die byna kosmopolitiese T. hypophylla,

word hier gegee. Baie meer materiaal is ondersoek as wat vir die vorige outeurs, Sim (1926) en Amell (1963), beskikbaar was.

Materiaal wat Duthie by Knysna versamel het, is deur Amell ( 1 963) gei'dentifiseer as Cyathodium foetidissimum. Dit is egter

verkeerdelik benaam en die materiaal is "n voorbeeld van Riccia rubricollis. Die teenwoordigheid van die genus Cyathodium in

Suider-Afrika is nie bevestig nie.

Targionia (Micheli) L., Species plantarum 2: 1136

(1753); Gott. et al. : 574 (1846); Schiffn.: 26 (1893);

Steph.: 763 (1898); Macvicar: 33 (1926); Sim: 16 (1926);

K. Mull.: 325 (1951-1958); Hassel de Menendez: 68

(1963); Arnell: 46 (1963). Type species: Targionia

hypophylla L.

Thallus , medium-sized, fleshy, lobes linear, sometimes

widening toward apex or cordate, somewhat leathery, light

green to darker green, often with a waxy bloom, in

crowded patches or extensive sheets; on soil in sheltered

rocky clefts or a pioneer on disturbed soil.

Branches often simple, sometimes repeatedly furcate

or only apically branched, or with ventral innovations;

thickened over midrib, rather abruptly thinning into wings;

apex notched, tips of scale appendages curving backwards

over it; not grooved. Dorsal epidermis hyaline, cell walls

thickened, especially at comers; air pores simple, conspic-

uous, slightly raised, with 3 concentric rings of cells, their

walls thin, leading below into shallow air chambers con-

taining chlorophyllose filaments; oil cells present; storage

tissue 3-5 times thicker than assimilation tissue above;

ventrally purple-black; some rhizoids smooth, others

pegged. Scales in one row on each side of midrib, dark

purple, large, obliquely triangular with broadly ‘awf-

shaped appendage.

Autoicous or dioicous. Antheridia mostly embedded in

swollen disciform apex of small male branch, arising ven-

trally at side of midrib and emerging laterally. Archegonia

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

MS. received: 1992-10-23.

formed at apex of thallus, and subsequently capsule dis-

placed to just below apex, enclosed in bivalved involucre

which opens at central narrow vertical fissure; capsule

wall cells with annular thickening bands. Spores convex

distally, ornamented with a double network, the larger net-

work forming strongly raised, rounded ridges which en-

close polygonal areas, all covered by a reticulum of

smaller, fine ridges. Elaters bispiral, sometimes branched.

Targionia hypophylla L., Species plantarum: 1136

(1753); Steph.: 764 (1898); Macvicar: 33 (1926); Sim: 16

(1926); K. Mull.: 326 ( 1951-1958); Hassel de Menendez:

69 (1963); Amell: 46 (1963); Volk: 241 (1979); Piippo:

274 (1991). Types: ‘Italia, Hispania, Constantinopoli’-t- ci-

tation (syn.); (OXF, syn.) Dill.: 532. Lichen no. 9, tab. 78,

fig. 9. (1741); (H-SOL, isosyn.) [according to Isoviita

(1970) and quoted by Grolle (1976)].

T. michelii Corda: 649 (1829). Type: Italy, leg. Sieber.

T. mexicana Lehm. & Lindenb. in Lehm.: 27 (1832). Type: Mexico,

leg. Schiede.

T. capensis Hiibener: 17 (1834).

T. hypophylla var. capensis (Hiib.) Krauss: 135 (1846). Type: Cape,

‘in promont. bonae Spei-, leg. Zeyher.

T. bifurca Nees & Mont, in Mont.: 113 (1838); Nees: 315 (1838).

Type: Chile, ‘prope Quillota", leg. Brotero.

T. convoluta Lindenb. & Gott. in Gott. et al:. 576 (1846). Type:

Mexico, ad Chinantla, leg. Liebman.

Thallus medium-sized, somewhat leathery, linear to lig-

ulate, apically often slightly broader, dark green, some-

times with bluish tint, marginally purple and entire to

somewhat crenate or scalloped, flat above, not grooved.

216

Bothalia 23,2 (1993)

FIGURE 1 — Targionia hypophylla , A, dorsal aspect of thallus with tip of involucre protruding at apex; B, thallus with short ventrally innovating

branch bearing terminal disc with antheridia; C, ventral aspect of thallus with pouch-like involucre at apex and 2 rows of ventral scales; D

tran verse section of thallus; E, air pore from above; F, transverse section of air pore, dorsal cells and chlorophyllose filaments; G, scale with

margins ± intact; H, scale with fimbriate appendage; I, cells of capsule wall with thickening bands. A, C, D. F, I, Koekemoer477a\ B, E, G,

Garside 6674\ H, Schelpe 4947. A, B, C, x 1 1 ; D, x 22; E, x 300; F, x 275; G, H, x 55; I, x 250.

Bothalia 23,2(1993)

217

polygonal outlines of subdorsal air chambers clearly vis-

ible from above, each with large and well-spaced air pore,

wet (Figure 1A); flanks ventrally deep purple and fre-

quently incurved, branches becoming almost tubular, or

sides clasped together, mostly covering dorsal epidermis

with its conspicuous white-encircled air pores, distal ven-

tral scales exposed, leading branch often becoming erect

and elevating ventral sporophyte, dry; in crowded patches,

simple to repeatedly furcate or with ventral or apical in-

novations. Branches 5-10 mm long between furcations,

total length up to 30 mm, 2. 7-3. 5 mm wide, 550 pm thick

over midrib, laterally thinning out into wings; apex

slightly notched with tips of scales reflexed over edge in

sterile plants or, when female receptacle present, entire

and lacking apical scales; margins acute, thin, ventrally

purple; flanks sloping obliquely outward and upward (Fig-

ure ID); ventral face medianly keeled, mostly purple, on

either side of midrib with row of imbricate, apically di-

rected, purple to reddish ± triangular scales (Figure 1C).

Dorsal epidermal cells unistratose, hyaline, from above

rounded to oval, 22.5-37.5 x 17.5-25.0 pm, trigones con-

spicuous, in transverse section cells ± 45 pm high, rec-

tangular, lateral walls thickened; marginal cells mostly

purple, from above rectangular, trapezoid or triangular,

(25-)30— 45(-50) x 20-32 pm, second row purple as well

or sometimes hyaline, also variable in shape and size,

22.5-50.0 x 15.0-17.5 pm, an inner third or fourth row

of cells frequently also present, all lacking trigones; mar-

ginal cells at apex of thallus ( 1 7— )22— 35 x ( 12— )25— 37

pm; air pores somewhat raised, simple, 52.5-67.5 x 35.0-

40.0 pm. oval or rounded (Figure IE), 250-375 pm distant

from each other, bounded by 3 concentric rings of cells:

innermost annular row of ± 6 thin-walled, curved, hyaline

cells, ± 15 x 25 pm, the remains of a collapsed cell ring,

next row of 8-11 sausage-shaped cells, ± 22 x 30 pm,

slightly thicker-walled, partly overlying outermost row of

14 also sausage-shaped cells, ± 27 x 35 pm; assimilation

tissue with air chambers in one layer, ± 80 pm thick, con-

taining simple or branched. 2- or 3-celled chlorophyllose

filaments, cells (20.0— )22. 5-34.0 x 25.0 pm, those below

air pores larger (Figure IF), 47.0 x 37.5 pm, with numer-

ous chloroplasts, and except at air pores, in close prox-

imity to dorsal cells; storage tissue ± 425 pm thick, cells

closely packed, angular, variable in size, 17.5-50.0 pm

wide, occasional cells with oil bodies, ± 27.5 pm wide;

rhizoids mainly between scales, some smooth, others

pegged. ± 15 pm wide. Scales in 2 longitudinal rows, one

on either side of costa, reddish to dark purple, obliquely

triangular, imbricate, not or hardly reaching thallus mar-

gins, 875-1750 pm long and up to 1325 pm wide at base,

if receptacle present, those covering it proximally larger,

2250 pm in length, basally up to 2000 pm wide, apically

continuing into single, forwardly directed, long-acuminate

appendage (Figure 1G), ± 375 pm long; cells in body of

scale 4-6-sided, ± 80.0 x 17.5 pm. in appendage 32.5-

62.5 x 22.5-27.5 pm, below appendage, polygonal, (20—

)37-60 x 25-45 pm, scattered oil cells present, margin

entire or not, sometimes with single-celled papillae, 30.0

x 17.5 pm or with fimbria (Figure 1H) ± 75 x 25 pm or

longer, especially at appendage of apical scales.

Autoicous or dioicous. Androecia borne on short, ven-

trally innovating branches, protruding at sides of thallus,

on terminal disc, 1.3-1. 8 mm wide, 750 pm thick and

encircled above by low, frilly membrane; antheridia seve-

ral, with conical protruberances, 150 pm high, each with

opening leading into antheridial cavity below. Gynoecia

terminal, enclosed in large, ± 3125 x 3600 pm, shiny

black, mussel-like, bivalved involucre, displaced below

apex of thallus, wall 4-layered, cells in outer layer thick-

walled on outside, 4-7-sided. 62.5-150.0 x 32.5-45.0 pm,

in inner layers thin-walled, elongated, ± 100 x 40 pm,

interspersed with numerous oil cells; to release spores and

elaters, involucre opening along narrow central vertical

fissure fringed with several irregular, cellular protuber-

ances; pseudoperianth lacking; calyptra hyaline, delicate,

cells irregular in shape and size, 45.0-55.0 x 27.5 pm;

capsule subsessile, spherical, wall yellowish brown, cells

± rectangular to mostly spindle-shaped, ± 75 x 35 pm,

with thickening bands (Figure II) ± 5 pm wide. Spores

(47 — )62— 77(— 95) pm in diameter, light brown to brown

or dark reddish brown, anisopolar, 2 faces different in

shape and ornamentation, distal face convex, rounded,

with double network (Figure 2A, D-F), the larger network

consisting of raised, rounded ridges, enclosing 16 or 17

polygonal areas, 12.5-17.5 pm wide, all covered by su-

perimposed, fine reticulum, the areolae of which mostly

smaller on sides and crests of primary ridges and larger

within enclosures; proximal face only slightly rounded to

flattish or even somewhat hollowed, generally with very

irregularly contorted, closely drawn together, reticulated

ridges separated by narrow fissures (Figure 2B); at junc-

ture of proximal and distal faces, when seen from inner

face, encircling row of reticulated primary ridges on distal

face projecting beyond rim of smaller proximal face, ap-

pear to form an undulating wing, ± 10 pm wide. Elaters

yellowish brown, not tapering but sometimes branched,

doubly spiral (Figure 2C). up to 290 x 12.5 pm. Chro-

mosome number : n = 9 (Bomefeld 1987).

DISCUSSION

In the family Targioniaceae, two subfamilies, Tar-

gionioideae and Cyathodioideae Grolle, are recognized,

each with only a single genus. They are characterized by

gynoecia that become ventrally displaced at the apex of

the thallus and capsules which are enclosed in a bivalved.

mussel-shaped involucre.

In Targionia species the thalli are compact and some-

what leathery, with low air chambers containing chloro-

phyllose filaments and a thick layer of storage tissue

below; in Cyathodium species the thalli are delicate, with

tall air chambers lacking chlorophyllose filaments and

with much reduced storage tissue.

To date, no monographic studies have been done on

the genus Targionia , although as many as 26 names have

been recorded under it (Geissler & Bischler 1990). Piippo

(1991), however, recently speculated that most of these

names will undoubtedly prove to be synonyms of T.

hypophylla. There are at least two other Targionia species

with well-defined specific limits, namely T. stellaris (K.

Mull.) Hassel de Menendez (1963) from Argentina and

a new species from India reported by Udar & Gupta

(1983).

Targionia hypophylla , the only representative of the

genus, is a widespread, almost cosmopolitan species [for

total range see Piippo (1991)], occurring especially in tem-

perate and seasonally dry areas where it grows as a xero-

218

Bothalia 23,2 (1993)

FIGURE 2. — Targionia hypophylla. SEM micrographs of spores and an elater. A, D-F, distal face of spore; B, proximal face of spore; C, elater.

A-C, S.M. Perold 2365\ D, Koekemoer 433\ E, Garside 6573 (Round House); F, Schelpe 4947. A, x 500; B, x 530; C, x 490; D, x 445; E,

x 560; F, x 460.

phyte on soil in rock crevices, on soil over rock outcrops

or under rock overhangs, or as a pioneer on disturbed

earth banks, like road cuttings. Sometimes other liverworts

such as Riccia spp., Plagiochasma spp., or Mannia spp.

are also present.

Targionia hypophylla is easily recognised, when fertile,

by the ventrally displaced capsule contained in a shiny

black pouch at the apex, by which further lengthening of

the branches is arrested. Sterile plants are distinguished

by their dark green, somewhat leathery appearance, the

conspicuous white-encircled air pores and reddish to pur-

ple-black triangular ventral scales with a single append-

age. Kashyap (1914) regards it as a very variable species,

and so does Schuster (1992), who thinks that incipient

speciation is under way; on the other hand Muller (1951-

1958) observes that, in spite of its wide distribution, plants

display little variation and the species is probably very

old. It is somewhat similar to Mannia spp., but these differ

from it by their less conspicuous air pores and by capsules

borne aloft on a longish stalk. Plagiochasma spp. have

smaller air pores and more pronounced spicules on the

dorsal epidermal cells, with the capsules also raised on a

stalk.

T. hypophylla was previously described from southern

Africa by Sim (1926) and Arnell ( 1963). It has been quite

rarely collected in Namibia (from whence Volk (1979)

recorded 14 collections), central Transvaal, Natal, Orange

Free State, Lesotho, Transkei, central, southern and

eastern Cape, but frequently in the winter rainfall areas

of the north western and southwestern Cape (Figure 4).

Further north in Africa it has also been recorded from

Ghana by Jones & Harrington ( 1983), Tanzania (Bizot el

al. 1978), Kenya and again Tanzania (Bizot et al. 1985),

Malawi (S.M. Perold 2665 PRE), Zambia (Vanden

Berghen 1972), Zimbabwe (Sim 1926; Best 1990) and the

adjacent islands of Madagascar (Arnell 1963) and

Reunion (Bischler 1990), although in the latter report, it

is listed as T. hypophylla and in brackets, ‘or T. lor-

beeriana ’. Frey & Kiirschner (1988) report T. hypophylla

from the Arabian Peninsula (together with T. lorbeeriana)

as well as a possible new morphotype, I hypophylla spp.

linealis (spp. nova?). T. elongata is also known from Af-

rica (Ethiopia), and Scott & Pike (1988) have published

SEM micrographs of its spores, which are very distinct

and clearly different from those of T. hypophylla , in that

there are only 5 or 6 large, ± smooth areas on the distal

face, as opposed to the usual 16 or 17 reticulated ones in

T. hypophylla. On a recent visit to Malawi, some speci-

mens of Targionia were collected, one of which, S.M. Per-

old 2653, had up to 30 reticulated areas (Figure 3F),

7.5-10.0 pm wide, on the distal face; the spore diameters

were 65-72 pm. It is not certain, however, whether we

are dealing with a different species or not.

Targionia hypophylla L. var . fhnbriata K. Muller is not

treated as a distinct variety in this study of southern Af-

rican specimens as was done by Arnell (1963), because

the presence of fimbria along the ventral scale margins is

very variable, even in thalli from the same population.

Apical scales are also more frequently fimbriate than older

ventral scales on the same branch. Schuster (1992) regards

the intrapopulational variation in this feature as so great

that no taxonomic segregation of the two extremes seems

possible to him.

Statistical and other studies by several authors (Sergio

& Queiroz Lopes 1972; Zamora et al. 1990; Jovet-Ast

Bothalia 23,2 (1993)

219

FIGURE 3. — Targionia spp. SEM micrographs of spores. A-E, T. lorbeeriana : A-D, distal face; E. proximal face. F, distal face of T. sp. A, Sergio

& Sim-Sim Lev. 70: B. Cros et al. 1 2- VII- 1 984; C, Sergio 148161: D, E, S. Arnell s.n. (Tenerife); F. S.M. Perold 2653 (Malawi). A, x 550;

B. x 490; C, x 635; D. E, x 580; F, x 620.

1956; Jovet-Ast & Zigliara 1967 and Zigliara 1970) have

confirmed the existence of an allied species, T. lorbeeriana

(Muller 1940) of which I have studied the type specimen

(leg. Huber-Tharandl, (5) collected in Sicily). Arnell

(1963) and Magill & Schelpe (1979) record the presence

of T. lorbeeriana in southern Africa, but this could not be

confirmed in the present investigation although (Arnell

(1963) even records T. lorbeeriana var. fimbriata from

here). The specimens which Arnell placed under T.

lorbeeriana , Garside 6573 [spore diameter 57.5-62.5 pm

(Figure 2E)] and Schelpe 4947 [spore diameter 75-85 pm

(Figure 2F)] are undoubtedly T. hypophylla, judging by

their spore ornamentation and cell measurements of the

thalli and scales. The correct naming of old dried herbar-

ium material is, however, difficult, since the colour of the

thalli fades, as does the distinctive odour of T. lorbeeriana.

Spore ornamentation (see later) seems to be a more reli-

able and certainly more permanent distinguishing charac-

ter. Grolle ( 1983) regards reports of the occurrence of T.

lorbeeriana in the East African Chyulu Mountains (Bizot

et al. 1978), as well as in the Cape (and India), as needing

verification. Targionia lorbeeriana is thought to be a "trip-

loid race' of T. hypophylla, as it has 27 chromosomes. It

is said to be characterized by a strong smell of acid pear

drops when fresh; by the cordate shape and light green

colour of the thallus; by larger, oval air pores and by dif-

ferent cell dimensions [for the latter see Zamora et al.

(1990)]. Schier (1974) reports slight differences in the

flavonoids of the two species. Differences in the spore

ornamentation of the two species were demonstrated by

Jovet-Ast & Zigliara ( 1967). Scott & Pike (1988) consider

the spore sculpturing as the clearest and least ambiguous

way of separating the two species: in their Australian ma-

terial they consistently found that T. hypophylla has reg-

ular areolae, whereas in T. lorbeeriana the areolae are

irregular in arrangement.

Spore samples of 20 different specimens of T. lorbeeri-

ana (on loan from European herbaria) were presently stud-

ied with the SEM and compared with more than 40 spore

samples of local T. hypophylla. In T. lorbeeriana spores

there are fewer ridges on the proximal face (Figure 3E)

and on the distal face, the fine reticulum, particularly that

covering the inside of the polygonal areas of the coarse

reticulum, has thicker walls (Figure 3 A-D); the proximal

spore face in T. hypophylla has numerous contorted ridges

and the walls on the distal face are thin and the areolae

oval or irregular in shape and size, the fine mesh present-

ing an altogether 'looser' appearance (Figure 2A, D-F).

Of the southern African material examined, spore orna-

mentation of Magill 6606. Van Rooy 2973, 3142a seems

to be intermediate between T. hypophylla and T. lorbeeri-

ana. Judging by my findings, the illustrations by Scott &

220

Bothalia 23,2 (1993)

Pike (1988: plate 1.1), appear to be of T. lorbeeriana and

1.2 of T. hypophylla , in other words, the reverse of what

they are stated to be. The T. hypophylla spores illustrated

by Taylor et al. ( 1974) are closely similar to those in my

study.

Cyathodium foetidissimum Schiffner

Arnell (1963) observed that a specimen in the National

Herbarium, Pretoria, collected by A.V. Duthie at Belve-

dere, Knysna, probably belongs to this species, i.e.

Cyathodium foetidissimum. This specimen has been ex-

amined and its spores photographed. It is undoubtedly a

species of Riccia and the ornamentation of the spores

proves it to be R. rubricollis Garside & Duthie ex Perold

(Perold 1991). On the small specimen packet inside the

larger envelope, in Sim’s handwriting is the note ‘Miss

Duthie’s (No. 23) new Riccia , Belvedere, Aug. 1928’ and

below it, in ArneH’s writing and signed by him is the

annotation ‘Cyathodium sp.'. One wonders why Arnell

had ignored Sim’s identification, as the latter must have

received the specimen from Duthie herself. In mitigation,

it may be added though, that the material had been pressed

and it is only a small sample, perhaps even a little atypical,

but the internal sporangia and spores immediately place

it correctly.

The other two species of Cyathodium , C. africanum

and C. aureonitens which Arnell (1963) thought would

possibly be found in southern Africa, have so far not been

collected here and it is doubtful if they ever will be, since

they are conspicuous and luminous green, and it is un-

likely that they could have been overlooked. The presence

of the genus, Cyathodium in southern Africa has therefore

not been confirmed although it is very widely distributed

in the rest of Africa (Jones 1952).

SPECIMENS EXAMINED

NAMIBIA. — 2217 (Windhoek): Dassieskuppe, Ostseite, an schat-

tigen Felsen, in Moosgesellschaft, (-CB), Volk 11357 (M, PRE).

TRANSVAAL. — 2428 (Nylstroom): Waterberg Game Park along

Melk River, shady riverbank, (-AB), Leistner 3558 (PRE). 2527 (Rusten-

burg): Swartkop Picnic Resort, near Randburg, on steep earth bank above

stream, (-DD), S.M. Perold 249 (PRE). 2528 (Pretoria): Pretoria, along

Apies River, (-CA), Scott 23 (PRE). 2529 (Witbank): Witbank. C.R.

Swart Nat. Res., at subsidiary stream entering Olifants River from west,

on streambed, clay soil, (-CD), Glen 1475 (PRE).

NATAL.- — 2828 (Bethlehem): Drakensberg, Royal Natal Nat. Park,

Broome Hill, in forest, (-DB), Cholnoky 182 (BOL); Drakensberg, lower

western slopes of Sentinel along footpath to chain ladder, alpine

heath/grassland with igneous rock outcrops, (-DB), Magill 6606 (PRE).

2929 (Underberg): Sani Pass Hotel, opposite bank of Mkhomazana River,

at waterfall, on soil, (-CB), S.M. Perold 2509 (PRE); halfway up Sani

Pass, vertical rock wall, with earth pockets, on soil, (-CB), S.M. Perold

2517 ( PRE).

O.F.S. — 2926 (Bloemfontein): Bloemfontein, Eagle’s Nest, (-AA),

Potts CH 1 143 (PRE); Bloemfontein, (-AA), T.R. Sim CH 1137 (PRE);

Farm Lastpoort on the Ruigtespruit, between Reddersburg and Helvetia,

grassland with trees and shrubs, on soil, (-DC), Van Rooy 2345 (PRE).

LESOTHO. — 2828 (Bethlehem): 5 km from New Oxbow Lodge to

Mokhotlong, waterfall over basalt cliff in tributary of Fanana River, near

Maiuti Club Ski Chalet, alpine heath-grassland, S aspect, on soil in

sheltered rock crevice, (-DC), Van Rooy 2961, 2973 (PRE); 6 km from

New Oxbow Lodge to Mokhotlong, alpine heath-grassland, on soil over

basalt, (-DC), Van Rooy 3058 (PRE); 4 km from New Oxbow Lodge to

Butha Buthe, alpine heath-grassland, on soil bank at flattish basalt out-

crops, NW aspect, (-DC), Van Rooy 3142a (PRE). 2929 (Underberg):

Kotisephola Pass, between Sani Top and Mokhotlong, alpine heath-grass-

land, basalt outcrops at top of pass, rock overhang, (-CA), Van Rooy 3399

(PRE); Sani River banks, ± 7 km from Sani Top, along road to Mokhot-

long, alpine heath-grassland, on soil in rock crevice, (-CA), Van Rooy

346 T. Sani Top, mountain slopes, W of Border Post, alpine heath-grass-

land, on soil, (-CB), Van Rooy 3554 (PRE).

TRANSKEI. — 3127 (Lady Frere): near top of Cala Pass, moist verti-

cal bank in road cutting, (-BC), Glen 1718 (PRE).

CAPE. — 2824 (Kimberley): Kimberley, (-DB), Welman CH 1141

(PRE). 2917 (Springbok): in valley leading from Ookiep to 'Ropeway’,

damp corners under ledges, (-DB), Giffen CH 1136 (PRE). 3018

(Kamiesberg): Inkruip, northern Khamiesberg, south slopes, damp

ground in lee of large granite boulders, (-AA), Oliver 7220 (PRE); Sors

Sors area, NE of Kamieskroon, (-AA), Oliver 9206 (PRE): 19 km NE of

Kamieskroon, 5 km after turnoff on road to Rooifontein, (-AA), S.M.

Perold 1473 (PRE); 2 km beyond Willem Stone Bridge, Pedroskloof, on

road to Rooifontein, on soil under rock, (-AA), S.M. Perold 1496 (PRE);

3 km along Rooifontein road, after turnoff from Kamieskroon-Leliefont-

ein road, on soil beneath rock overhang, (-AA), S.M. Perold 2138 (PRE);

5.2 km along Rooifontein road, after turnoff from Kamieskroon-

Leliefontein road, (-AA), S.M. Perold 2169 (PRE); southern Khamies-

berg, Klippoort, SE of Hoedberg, granite hill, slope facing south, sandy

hollows, (-CB). Oliver 9557 (PRE). 3023 (Britstown): Britstown, Farm

Jakkalskuile, (-AD ),Viljoen CH 4523 (PRE). 3026(Aliwal North); Albert

Dist., ± 12 km SE of Bethulie, Cliftonvale Farm, foot of rocky cliffs,

seepage zone, (-CA), H.H. Burrows 2522 (PRE). 3027 (Lady Grey): Dist.

Herschel, Sterkspruit, (-CA), Hepburn CH 1138. CH 1139 (PRE);

Witteberg Mts, Joubert’s Pass, 8 km east of Lady Grey, western aspect ,

alpine heath-grassland, on soil, (-CB), Van Rooy 2700 (PRE). 3118

(Vanrhynsdorp): Gifberg. SE of Van Rhynsdorp, at top of mountain,

underneath rock, drier area away from stream, (-DC), S.M. Perold 2754

(PRE). 3119 (Calvinia): Van Rhyns Pass, at top of plateau, on soil, (-AC),

S.M. Perold 2186 (PRE); Hantamsberge, top of mountain near FM tower,

mountain rhenosterveld, rock and soil pockets, (-BD), Koekemoer 822

(PRE); south side of Hantams Mountain, near FM tower, on soil between

rocks, (-BD), S.M. Perold 1824 (PRE); Farm Daantjie-se-kraal, 37 km

along road between Soetwater and Clanwilliam. 8 km before Botterkloof

Pass, (-CB/CD), S.M. Perold 1871 (PRE); Vondelingsfontein, northern

Roggeveld, damp ground in lee of shrubs, (-DD), Oliver 8911 (PRE);

Vondelingsfontein Farm, 10 km from Calvinia on road to Middelpos,

along small streamlets at roadside, (-DD), S.M. Perold 1839 (PRE). 3120

(Williston): northern Roggeveld. Knegtsbank, kloof north of farm, south

slopes with dense rhenosterbos scrub, (-CC), Oliver 8920 (PRE). 3218

(Clanwilliam): middle slopes of Pakhuis Pass, on Clanwilliam side, (-

BB), Koekemoer 433 (PRE); Aggensbachsberge, ground forest, (-BB),

Stirton 9280 (PRE); 20 km N of Citrusdal, past Hexriver Farm, on rocky

outcrops at roadside, above Olifantsrivier, (-BD), S.M. Perold 530 (PRE);

5 km along road to Algeria, after turnoff from Cedarberg road, under

damp overhang near road, (-BD), S.M. Perold 2351 (PRE). 3219

(Wuppertal): 3 km before turnoff to BiedouwAVupperthal, on R364,

between Soetwater and Clanwilliam. at streamlet, on sand over sand-

stone, (-AA), S.M. Perold 1883 (PRE); 4 km south of Algeria Forest

Station, (-AC), S.M. Perold 2365 (PRE); Ceres, Kaggakamma Nat. Res.,

near beacon 3 in streambed, at foot of boulders, (-DA), Koekemoer 768,

769, 770 (PRE). 3220 (Sutherland): 24 km NW from Sutherland

(Roggeveld Mtns), occasional in S aspect rock crevices, (-AB) Schelpe

4952 (BOL); 2 mis N from Sutherland, occasional on S aspect rock edges

under bushes, (-BC), Schelpe 4947 { BOL); Verlaten Kloof, 18 mis S from

Sutherland, locally frequent under asparagus bushes on S aspect, shale

slope, (-DA). Schelpe 4942 (BOL); Sutherland, near top of Komsberg

Pass, in rhenosterveld, on sandy-loamy soil, (-DB), Vlok 2667 (PRE);

Smoushoogte, Klein Roggeveld, damp southern slope, with short

rhenosterbos scrub, (-DC), Oliver 8969 (PRE). 3318 (Cape Town): Pen-

insula, Constantia slopes, (-CD), S. Amell 379 (BOL); Kirstenbosch,

(-CD), S. Arnell 565 (BOL); Cave Peak (-CD), S. Arnell 615 (BOL);

Lion's Head, Round House, (-CD), S. Arnell 1176 (PRE); behind Round

House, Camps Bay, on granite soil, (-CD), Garside 6495 (BOL); Round

House, Camps Bay, on soil, (-CD), Garside 6573 (BOL); Camps Bay,

below Round House, (-CD), Garside 6674 (BOL); Table Mountain,

Slongoli, (-CD), T.R. Sim CH 1149 (PRE); Devil’s Peak, (-CD), T.R. Sim

CH 1142 (PRE); Cape Town, (-CD), H.A. Wager CH 1133 (PRE);

Stellenbosch Flats, (-DD), Duthie CH 1148 (PRE); Stellenbosch, (-DD),

Duthie CH 1147 (PRE); Jonkershoek road, Stellenbosch, earth bank by

side of road, (-DD). Garside H60 (BOL); Jonkershoek, Stellenbosch,

(-DD), Garside 9 (PRE). 3319 (Worcester): Gydo Pass, on soil, (-AB),

Stirton 9160 (PRE); Hex River Pass road, occasional on shaded earthbank

in dry stream bank, (-BD), Schelpe 4918 (BOL); Hex River Pass, near

summit, gulley, on damp ground, (-BD), Stirton 9482 (PRE); Bainskloof,

Bothalia 23,2 (1993)

221

NE of Wellington, vertical rock wall at roadside, on soil, (-CA), S.M.

Perold 2785 (PRE); Worcester Karroo Garden, amongst karoid shrubs,

(-CB). Koekemoer 364 (PRE); Farm Leipzig, east of Worcester, at foot of

Rabiesberg, on soil on rock face, (-DA), Morley 289 (PRE); Sandhills,

north of Worcester, near railway line, on soil on rock outcrop, (-DA), S.M.

Perold 579 (PRE). 3320 (Montagu): Montagu, Bath Kloof, (-CC), S.

Amell 749. 758 (BOL). 3322 (Oudtshoom): Prince Albert, at northern

base of Swartberg Pass, in arid mountain fynbos, on loamy sandy soil,

(-AC), Vlok 2660 (PRE); Meirings Poort, North of De Rust, near

Oudtshoom, on earth bank above road, (-BC), S.M. Perold 898 (PRE);

near Hoekplaas, on gravel road from De Rust to Uniondale, on rocky

slope under karoo bushes, (-DB), Koekemoer 477a (PRE). 3325 (Port

Elizabeth): Hells Gate, Uitenhage, (-CD), T.R. Sim CH 1140 (PRE). 3419

(Caledon): Greyton Kloof; on soil on rock wall next to footpath, (-BA),

S.M. Perold 607. 1170 (PRE); Betty’s Bay, (-BD), S. Amell 695 (BOL).

ACKNOWLEDGEMENTS

My sincere gratitude to Drs C. Sergio, LISU, and A.J.E.

Smith, Bangor, for refereeing this article and for their

helpful suggestions. I also wish to thank the following for

the loan of specimens; Dr C. Sergio, LISU, the curators

of BOL and S; Prof. Dr O.H. Volk, Wurzburg, and Dr R.

Liibenau-Nestle, Kempten, Germany; my thanks to my

colleagues at NBI, Dr H.L. Glen, Miss M. Koekemoer and

Mr J. van Rooy, for collecting specimens; also to the artist,

Ms G. Condy, the typist Mrs J. Mulvenna and the pho-

tographer Mrs A. Romanowski, for their valued contribu-

tions. Miss S.K. Marner, OXL, is thanked for kindly

sending photographs of Targionia hypophylla from

Dillenius's herbarium.

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*» -*■, ■

Bothalia 23,2: 223-229 (1993)

The hepatic, Jensenia spinosa (= PaUavicinia stephanii: PaUaviciniaceae), in

southern Africa

S.M. PEROLD*

Keywords: hepatic, Jensenia spinosa , Metzgeriales, PaUaviciniaceae, PaUavicinia stephanii, Pallavicinioideae, southern Africa

ABSTRACT

Recently, specimens of Jensenia spinosa (= PaUavicinia stephanii) were collected at the type locality ‘Spitskop, bei

Lydenburg', in the Transvaal. The type, which previously was the only specimen from southern Africa, is held at G. A detailed

description of J. spinosa, illustrated with photographs of the thalli and spores is presented here, since earlier descriptions are

sketchy and lack illustrations.

UITTREKSEL

Onlangs is eksemplare van Jensenia spinosa (= PaUavicinia stephanii) by die tipelokaliteit, "Spitskop, bei Lydenburg' in

Transvaal versamel. Die tipe, wat voorheen die enigste eksemplaar vanaf Suider-Afrika was, word by G gehou. 'n Beskrywing

van J. spinosa, geYllustreer met foto’s van die tallusse en spore word gegee, aangesien vroeere beskrywings onvolledig en sonder

illustrasies is.

INTRODUCTION

The subfamily Pallavicinioideae (Migula) Grolle of the

family PaUaviciniaceae Migula, order Metzgeriales, com-

prises the genera PaUavicinia and Jensenia (see below).

Jensenia Lindb. (1868) is the older name for the latter of

the two genera and had to replace Makednothallus Ver-

doom (1932) (Grolle 1964), which was based on Mittenia

Gott. (1864). This name, however, was taken up already

by Mittenia Lindb. (1863), a genus of mosses. Jensenia

was treated by Schiffner (1893) and later authors such as

Schuster & Inoue (1975); Grolle (1984); Grolle & Piippo

(1984) and Schuster (1992), not as a genus, but as a sub-

genus of PaUavicinia. It corresponds to Stephani’s (1900)

PaUavicinia sectio Dendroideae. Differences between

Jensenia and PaUavicinia are shown in Table 1.

Grolle & Piippo (1986) list several of the above dif-

ferences to demonstrate that Jensenia is a natural group

which they prefer to treat at the generic level. I have fol-

lowed them in this study even though Schuster (1992)

adopts a broadened concept of PaUavicinia to include

Jensenia because of the following: in PaUavicinia levieri

Schiffn. the androecia have scales scattered over the costal

surface in several poorly demarcated rows, i.e. more like

those in Jensenia (or Mittenia, as Schuster prefers to call

it, since Mittenia has priority at subgeneric rank). Further-

more, in the Japanese PaUavicinia longispina Steph.,

Schuster (1992) often found a basal stipe that was creeping

and rhizoidal, contrasted to an ascending (but not erect)

frond, that was 1 or 2 (very rarely 3) times dichotomous,

i.e. also as in Jensenia , as opposed to PaUavicinia which

is supposedly always prostrate and without a creeping rhi-

zome. The differences between PaUavicinia and Jensenia

are therefore not clear-cut in the above two characters,

but in the remaining ones (see Table 1), the differences

are deemed to be sufficiently marked to treat Jensenia as

* National Botanical Institute, Private Bag X 101. Pretoria 0001 .

MS. received: 1992-11-20.

a separate genus, rather than as a subgenus of PaUavicinia.

Six species (Grolle 1964) or seven (Engel 1990), are clas-

sified in Jensenia, which belongs to a southern and prob-

ably Gondwanaland element. Jensenia spinosa ( =

PaUavicinia stephanii ; = Makednothallus stephanii ) is the

only African representative. Initially Grolle (1965) had not

recognized the combination Makednothallus stephanii

(Jack) Schust. (Schuster 1963) and Jones (1990) made the

combination Jensenia stephanii (Jack) Jones. The type

specimen of PaUavicinia stephanii was collected by

Wilms at Spitskop, ‘bei Lydenburg', Transvaal, in 1888.

This collection is held at G. No specimens of J. spinosa

were housed at PRE, except for the recent acquisition of

a De Sloover collection from Rwanda, through the kind-

ness of Dr Riclef Grolle. Amell (1963) admits that he did

not see examples of this taxon himself, but based his de-

scription on that of Stephani. Two recent attempts were

made to collect this species again at a locality called

Spitskop, situated south of Lydenburg, but were unsuc-

cessful. Only upon coming across a copy of Wilms's

( 1898) ‘Ein botanischer Ausflug ins Boerenland’ did it be-

come clear that the Spitskop to which he referred, lay

about 50 km east of Lydenburg and 9 km south of the

present town of Sabie, which was only proclaimed later.

Fresh material of this locally scarce species was recently

collected within sight of this Spitskop, on the protected

vertical banks of a small streamlet, close to the water sur-

face. Jensenia spinosa and its spores are herewith de-

scribed in detail and illustrated with photographs, as

earlier descriptions are sketchy and not illustrated.

Jensenia spinosa (Lindenb. & Gott.) Grolle in

Grolle & Piippo in Acta Botanica Fennica 133: 65 (1986).

Symphyogyna spinosa Lindenb. & Gott. in Gott. et al .: 786 (1847).

PaUavicinia spinosa (Lindenb. & Gott.) Grolle: 268 (1979). Type: Masca-

renes. Ins. Mauritius, leg. Bory (W, lecto.).

S. serrata Mitten in Melliss: 572 (1875). Type: St Helena, leg. Melliss

(NY, holo.)

224

Bothalia 23,2 (1993)

TABLE 1. — Comparison of characters of the genera Jensenia and Pallavicinia

Pallavicinia stephanii Jack in Steph.: 129 (1892). Pallavicinius

stephanii Jack in Steph.: 324 (1900). Makednothallus stephanii (Jack)

Schust.: 292 (1963). Jensenia stephanii (Jack) Jones: 15 (1990). Type:

Transvaal, Spitskop bei Lydenburg, leg. Wilms G 007994 , lecto., selected

here; Wilms G 007992, para. [Synonymy mostly fide Grolle ( 1979)].

Terricolous, on damp soil; thallose, medium-sized,

green; shoots erect, dendroid, in crowded tufts, expanded

into flabellate aerial fronds; once dichotomously branched

(Figure 3C), to several (2 — 4) times (Figure 1A). Terminal

branches generally 6 or 7(8), very rarely 11 in older

plants, up to 10 mm long and tapering distally; proximal

branches 2-5 x 1.0- 1.7 mm and 165 pm thick over costa,

apices blunt to rounded, entire or slightly notched, bearing

2-cel led slime papillae (Figure 2C); central conducting

strand visible from above (Figure 1A, B) and bifurcating

some distance below separation of branches; margins of

wings with small remote teeth in sterile plants, but with

prominent spines in male and some female plants, lacking

slime papillae, plane to undulate, bilaterally rather grad-

ually expanded. Stipe ascending, wingless and un-

branched, purplish red, basally bearing some rhizoids, ±

10 x 500 pm, with single strand below, double strand

above (Figure 1H). Rhizome horizontally creeping,

branched, dark brown, embedded in the soil, in section

(Figure ID) cortical cells with outer walls slightly thick-

ened, medullary cells thin-walled, larger, ± 45 pm wide,

central strand cells thick-walled, 7.5 pm wide. Rhizoids

mostly ventral, smooth, brownish, slightly thick-walled, ±

12.5 pm wide. Wings with forwardly directed marginal

teeth, sometimes curved and projecting at angles of 20°-

90°, low and blunt (Figure IF), or up to (3)4 or 5 cells

long, basally 2 to 4 cells wide, tapering to pointed uni-

seriate apices (Figure 2H); marginal cells from above rec-

tangular or polygonal, 30-52 x 17-25 pm (Figure IJ),

lamina! cells hexagonal, ± 70 x 30 pm (Figure IK), epi-

dermal cells above midrib rectangular, 35^47 x 20-22 pm

(Figure I L), all containing up to 9 spindle- or rod-shaped

011 bodies, finely granulose (Figure II), ± 10 pm long, as

well as numerous rounded chloroplasts, 2.5 pm wide; in

section wings unistratose along margins for 6-8 cell rows,

then 2, soon 3 cell layers deep, grading into thickened

costa (Figure 1C, E), parenchymal cells enclosed by chlo-

rophyllose epidermis above and below, colourless and

empty, rounded, up to 45 x 45 pm. Costa with central

conducting strand up to 50 x 50 pm, consisting of a group

of ±11 thick- walled and elongated, brown, sclerenchymal

cells, ± 5 pm wide, surrounded by parenchymal cells (Fig-

ure 1C, E, G).

Dioicous. Androecia above and lateral to costa, cover-

ing dorsal central parts of segments of once or twice di-

chotomously branched, backwardly arched frond (Figure

2A, B, D), margins of segments undulate, prominently

toothed (Figure 2H); antheridia numerous, not in distinct

rows but crowded together, up to 4 alongside each other,

globose (Figure 2D, E), 225 x 200 pm, yellowing with

age, in axils of and covered by apically directed, dentate

to laciniate scales (Figure 2F, G), the latter imbricate, fre-

quently connate at the bases, turning orange, laciniae up

to 400 pm long, apically uniseriate, 3 or 4(6) cell rows

wide basally, cells up to 75 x 22 pm. Gynoecia single,

dorsally situated immediately distal to a bifurcation in

lower part of frond, which when young is somewhat fun-

nel-shaped with connivent wings (Figure 3A). Involucre

cup-like, surrounding a group of archegonia (Figure 3E),

basally 4 cell layers thick in transverse section (Figure

3G), otherwise 2- or 3-layered (Figure 3F), 1300 pm high,

including laciniae 375 pm long projecting upwards from

the mouth, 2750 pm wide when opened up and spread

out. Pseudoperianth developing inside involucre after fer-

tilization, becoming cylindrical (Figure 3D) and eventu-

ally ± 4500 pm long, 3 or 4 cell layers thick, irregularly

laciniate at mouth, laciniae up to 330 pm long, uniseriate

above, 3 or 4 arising from each basal part, up to 500 x

300 pm, separated from each other by slits, sometimes

with 2-celled slime papillae at the base, outer cells hex-

agonal, ± 70 x 37 pm above, elongating to + 200 x 37

pm below. Calyptra initially enclosing capsule and seta

(Figure 3D, K), inserted into mature pseudoperianth and

basally fused with it, 2825 x 750 pm, creamish white, at

apex old archegonial neck projecting and at sides several

unfertilized archegonia (Figure 3K) as well as 4-celled

slime papillae, 67 x 15 pm, mostly 2 cell layers thick

(Figure 3L), outer cells smaller, 60-75 x 30 pm, irregu-

larly shaped, with sinuous walls. Capsule cylindrical,

1050 x 750 pm, brown, containing a mass of spores and

elaters without elaterophore, wall bistratose (Figure 31),

with outer cells up to 75 x 20 pm, lacking band-like thick-

enings (Figure 3H), eventually perforating top of calyptra

and carried aloft as seta elongates, dehiscing by 2 slits.

Bothalia 23,2 (1993)

225

FIGURE 1. — Jensenia spinosa: LM photographs of thalli. A, terminal branching; B, single branch with toothed margin; C, ts. of branch showing

costa with central strand and lateral wings; D, ts. of rhizome; E, ts. of branch; F, small, remote teeth at margin; G, ts. of costa and central

strand; H. ts. of apical part of stipe with double conducting strand, note also 2-celled slime-papilla; I, cells containing oil bodies and

chloroplasts; J, cells along margin of branch; K, cells in lamina; L, dorsal cells over costa. A-C, Wilms 007994 ; D-L, Perold & Koekemoer

2923. A, x 7.5; B. x 25; C. x 125; D. H, x 50; F, x 100; G, J-L, x 250; I, x 500.

226

Bothalia 23,2 (1993)

at base as in G); H, highly toothed margin of male thallus. A, B, x 1 8, C, x 250, D, x 25, E, > ’ ’ ' '

Bothalia 23,2(1993)

227

FIGURE 3. — Jensenia spinosa: LM photographs. A, female thalli with young gynoecia; B, female thallus with older gynoecia; C. female branch

with single gynoecium: D. pseudoperianth opened to show capsule sheathed in calyptra; E. involucre; F. ts. of involucre: G, ts. of base of

involucre, 4 cell layers thick; H, capsule wall cells; I, ts. of capsule wall: J, ts. of young seta; K. calyptra wall with unfertilized archegonium

still attached; L, ts. of calyptra wall. A-G, Perold & Koekemoer 2923\ H-L. De Sloover 13459. A, C, x 10: B, x 12.5; E, F. x 25: G-I, x

250; J. x 50; K, L, x 125.'

228

Bothalia 23,2 (1993)

FIGURE 4. — Jensenia spinosa : SEM of spores and elater. A-D, distal face of spore; E. proximal face of spore; F, elater. A. Perold & Koekemoer

2923 ; B-F. De Sloover 13459. A, x 1255: B, x 1295; C, E, x 1215; D. x 1 193; F, x 375.

with the valves coherent. Seta before elongation and when

still enclosed by calyptra, ± 875 pm long, diameter in

section 450 jam, cortical cells 27.5-37.5 x 32.5-37.5 jam,

in ± 43 cell rows, medullary cells, 30.0-52.5 jam wide,

angular, ± nine across (Figure 3J); foot 625 pm long, nar-

rowing to a pointed base. Spores light brown, globular to

oval, 25-32 pm in diameter, ornamentation cristate with

numerous short and long, branched and unbranched, bent

or curved ridges (Figure 4A-E), in between with short

rods and dots, 2.5 pm high, expanded and flattened on

top: proximal face, besides being ornamented as described

above, also with a discrete, round area, ± 10 pm wide,

lacking ridges and with only a few granules visible (Figure

4E). Elaters light brown, tapering slightly at ends, up to

290 x 7.5 pm in the centre, 2-spiral (Figure 4F).

Besides the type locality in southern Africa (Figure 5),

./. spinosa is known elsewhere in Africa from Malawi,

Tanzania, Rwanda and Zaire, as well as from the islands

Mauritius, Reunion and St Helena (Grolle 1979), gener-

ally from high elevations. Vana et al. (1979) reported it

as Pallavicinia stephanii from Rwanda and Reunion. It is

therefore quite widespread, but scarce.

According to Grolle (1979) ./. spinosa is very similar

to the neotropical species, ./. erythropus (Gott.) Grolle;

Schuster (1992) concludes that in the description and

illustrations of J. erythropus by Hiissel de Menendez

(1961 ), the scales are scattered over the dilated sectors of

the frond segments. The only other dendroid member of

the family Pallaviciniaceae in Africa, is Symphyogyna

podophylla (Thunb.) Mont. & Nees (Perold 1993) which,

in the absence of female material, is quite difficult to dis-

tinguish from J. spinosa. Grolle (1979) gives a key to

distinguish them (on vegetative characters), in which he

draws attention to the smaller, as measured by him, 21-27

x 24—35(42) pm, inframarginal cells of the thallus wings

with the cuticle finely punctate to striate in J. spinosa ,

whereas in S. podophylla he found these cells to be larger

at 50-60(65) x 50-75 pm and the cuticle smooth. Van

der Gronde (1980), who studied the genus Jensenia in

Colombia, states that female material of Jensenia is neces-

sary to distinguish it with certainty from a dendroid Sym-

phyogyna, which has a scale-like involucre and lacks a

pseudoperianth. In Jensenia the involucre is cup-like and.

after fertilization, a long pseudoperianth develops; the lat-

ter is joined with the calyptra at the base. There are thus

three structures surrounding the sporophyte in Jensenia'.

the involucre, the pseudoperianth and the calyptra.

SPECIMENS EXAMINED

RWANDA. — Parc des Volcans, 3 650 m, Vysoke sur pan rocheux

(lave), vertical moussu, De Sloover 13459 (ex Herb. Bryologicum R.

Grolle).

FIGURE 5. — Known distribution of Jensenia spinosa in southern

Africa.

Bothalia 23,2(1993)

229

TRANSVAAL. — 2530 (Lydenburg): Spitskop, bei Lydenburg, (-

BB), Wilms (G 007992, G 007994)', 9 km S of Sabie, 0.5 km N of Spitskop

Store, on road R537 from Sabie to White River, at small streamlet on left

side of road, on vertical earth bank, (-BB). Perold & Koekemoer 2923

(PRE).

ACKNOWLEDGEMENTS

I wish to express my sincere gratitude to Drs R. Grolle

and E.O. Campbell for refereeing this article and for their

helpful suggestions; also to Dr P. Geissler, curator at G

for the loan of the type specimen and to Ms M. Koeke-

moer, curator of PRE, for accompanying me and helping

me on field trips as well as for actually finding the locality

where Jensenia spinosa grows. My thanks also to Mrs A.

Romanowski, photographer, for developing and printing

the photographs (taken by the author), and to the typist.

Mrs J. Mulvenna, for her valued contribution.

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Bothalia 23,2: 231-237(1993)

Notes on African plants

VARIOUS AUTHORS

VITACEAE

A NEW SPECIES OF RHOICISSUS FROM THE TRANSVAAL

Rhoicissus laetans Retief sp. nov., R. microphyllae

(Turcz.) Gilg & Brandt similis sed foliis ellipticis vel ob-

ovatis, glaucoviridibus, glabris, in sicco valde discolori-

bus, sine domatiis differt; R. microphyllae folia ovata vel

elliptica sunt, pilis rufis vestita, cum domatiis fasciculis

trichomatorum in axillis venarum principalium lateralium.

2 mm long, yellowish green. Stamens with filaments 1

mm long, dorsifixed, opposite petals, bending over gynoe-

cium. Disc entire, with ovary immersed in it. Style simple,

entire, 2 mm long. Stigma not broader than style. Berries

12 mm in diameter. Seed 1— 3(— 4); testa smooth, dark

brown, usually with a prominent, longitudinal furrow.

Type. — Transvaal, 2430 (Pilgrim's Rest): along Drie

Rondawels-Uitkyk road, (-DB), Herman 737 (PRE,

holo.). Figure 1.

A shrub, up to 1.5 m high, sometimes scrambling. Ten-

drils absent. Leaves simple, petiolate; lamina elliptic to

obovate, 18-50 x 10-25 mm, glabrous, net-veining prom-

inent on undersurface, margin entire, base obtuse, apex

obtuse to slightly acute, very shortly mucronate; stipules

absent; petiole up to 5 mm long. Inflorescence a leaf-

opposed cyme, tendril-like. Calyx entire. Petals 4, ovate.

TRANSVAAL. — 2430 (Pilgrim’s Rest): Blydepoort Nature Reserve,

Lowveld look-out, (-BD), Van Wyk 10233 (PRU); Farm Steenveld

229KT, kranzes overlooking Ohrigstad River, (-DA), Fourie 1316 (PRE);

along Drie Rondawels-Uitkyk road, (-DB), Herman 737 (PRE); 18.3

radial miles from Pilgrim’s Rest, banks of Blyde River, (-DB), Mogg &

Davidson 33547 (J, PRE); Blydepoort Nature Reserve, F.H. Odendaal

Rest Camp, (-DB), Van Wyk 5200 (PRE, PRU).

Rhoicissus laetans is endemic to the northeastern

Transvaal Escarpment, where it occurs in a small area

(Figure 2). The species is found in mountain grassland

with stunted shrub vegetation (North-eastern Mountain

Sourveld) or on steep, densely wooded kloof slopes with

mixed bushveld vegetation. Occasionally it occurs in riv-

erine forest. The species grows on soils derived from

quartzite and sandstone. It flowers from at least November

(probably earlier, judged by the occurrence of fruits on

specimens) till February.

Classification of the new species in the genus Rhoi-

cissus is supported by the shape of its flower buds, struc-

ture of the floral disc and inflorescence morphology. In

Rhoicissus the buds are globose in outline (Figure 3).

Flower buds of Cissus are typically conical to ovoid with

the petals cucullate at the apex, whereas those of Cy-

phostemma are cylindric or flask-shaped, constricted in

the middle and inflated at the apex (Figure 3). Flower

buds of Ampelocissus and Cayratia are also globose in

FIGURE 1 . — Holotype of Rhoicissus laetans, Herman 737 (PRE).

FIGURE 2. — Distribution of Rhoicissus laetans.

232

Bothalia 23,2 ( 1993)

FIGURE 3. — Characteristic flower bud shape in three genera of

the Vitaceae: A, Rhoicissus: B. Cissits ; C, Cyphostemma.

outline, but Ampelocissus differs from Rhoicissus in hav-

ing an inflorescence with tendrils and a floral disc with

vertical furrows. Cayratia has a thin disc and axillary

cymes, whereas in Rhoicissus the disc is annular, entire

and thick and the inflorescence is a leaf-opposed cyme.

The species of Rhoicissus usually have 5 or 6 petals

per flower. Although only 4 petals are present in R. lae-

tans and R. microphylla (as is the case in Cissus, Cypho-

stemma and Cayratia ), the prominent globose flower buds

place the species in Rhoicissus.

Rhoicissus laetans is distinguished from all the other

members of the genus by its shrubby habit and simple,

glabrous, glaucous green leaves. In southern Africa R.

tomentosa (Lam.) Wild & Drummond and R. microphylla

(Turcz.) Gilg & Brandt also have simple leaves. The

leaves of R. tomentosa are broadly transversely elliptic to

reniform and the species is a high-climbing liane. R.

microphylla , a small shrub, has ovate leaves with the un-

dersurface of the lamina covered with reddish brown

hairs. The other southern African species, namely R.

digitata (L.f.) Gilg & Brandt, R. revoilii Planch., R.

rhomboidea (E. Mey. ex Harv.) Planch., R. sessilifolia

Retief and R. tridentata (L.f.) Wild & Drummond all have

3-foliolate leaves.

FIGURE 4. — SEM micrograph of leaf undersurface: A, Rhoicissus

tuft domatia in axils of principal lateral veins, x 16.

For nearly twenty years it was thought that specimens

of R. laetans belonged to R. microphylla , a species oc-

curring only in the eastern Cape. The two taxa differ not

only in their widely disjunct distribution but R.

microphylla has ovate to elliptic leaves, with reddish hairs

and hair-tuft domatia on the undersurface (Figure 4B). R.

laetans, on the other hand, has elliptic to obovate leaves

which are glaucous green and glabrous when fresh (Figure

4A), and prominently discolorous when dried.

The geographical range of R. laetans falls within the

Wolkberg Centre of endemism, and more specifically the

Blyde Subcentre (Matthews et al. 1993; listed as

Rhoicissus sp. nov.). Other woody species endemic/near-

endemic to the Blyde Subcentre include Euclea dewinteri

Retief, Protea laetans L.E. Davidson and Combretum

petrophilum Retief. Rhoicissus laetans and its sister spe-

cies, R. microphylla, which occurs in the eastern Cape,

mirror a disjunct distribution pattern shown by a number

of other Blyde Subcentre endemics, e.g., Streptocarpus

meyeri B.L. Burtt, Cyrtanthus huttonii Baker and Hae-

manthus paucifolius Snijman & Van Wyk (Hilliard &

Burtt 1971; Dyer 1972; Reid & Dyer 1984; Snijman &

Van Wyk 1993). The repeated pattern shown by these

vicarious taxa should have value for tracing the history

and development of the southern African flora.

The specific epithet laetans refers to the area from

which the species has mainly been collected: Blydepoort

Nature Reserve, with 'bly’ meaning ‘joyful’.

ACKNOWLEDGEMENTS

The author wishes to thank Drs O.A. Leistner and H.F.

Glen of the National Botanical Institute, Pretoria, and

Prof. A.E. van Wyk of the University of Pretoria, for their

assistance in the preparation of this manuscript.

REFERENCES

DYER, R.A. 1972. Cyrtanthus huttonii. The Flowering Plants of Africa

42: t. 1660.

laetans. x 35; B, R. microphylla. note scattered trichomes and hair-

Bothalia 23,2(1993)

233

HILLIARD. O.M. & BURTT. B.L. 1971. Streptocarpus: an African plant

study. University of Natal Press, Pietermaritzburg.

MATTHEWS, W.S., VAN WYK, A.E. & BREDENKAMP. G.J. [1992]

1993. Endemic flora of the northeastern Transvaal Escarpment,

South Africa. Biological Conservation 63: 83-94.

REID, C. & DYER, R.A. 1984. A review of tlte southern African

species of Cyrtanthus. American Plant Life Society, La Jolla,

California.

SNIJMAN, D.A. & VAN WYK, A.E. 1993. A new species of

Haemanthus (Amaryllidaceae) from the eastern Transvaal Es-

carpment, South Africa. South African Journal of Botany 59:

247-250.

E. RETIEF

MS. received: 1992-10-19.

CUCURBITACEAE

A NEW SPECIES OF KEDROST1S FROM THE WESTERN CAPE

Kedrostis psammophila Bmyns , sp. nov., a ceteris

speciebus capreolis destitutis, caulibus horizontalibus sub-

terraneis repentibus, receptaculo longissimo in flore

femineo cum ovario subterraneo differt. Type: South Af-

rican Cape. Nortier, Bruyns 4569 (BOL, holo.; PRE, K,

M, iso.).

Plant with carrot-shaped to cylindrical tuber up to 200

x 100 mm or more, usually at least 150 mm beneath soil

surface and pale grey-brown with white flesh, with several

smaller subsidiary tubers produced elsewhere along stems,

very strongly foetid, monoecious. Stems soft, creeping

horizontally to 1 m or more and extensively branched be-

neath soil surface, cylindrical, white; portions above

ground usually not more than 150 mm long, clustered,

horizontally spreading, bearing leaves along whole length,

grey-green, hairy. Leaves palmately 3- to 5-lobed often to

at least halfway but sometimes ± entire, 6-20 x 10-25

mm, segments irregularly dentate, almost all of equal size,

grey-green, hairy on both sides; petiole 8-15 mm long,

grooved above, setose. Tendrils and stipules none. Flow-

ers unisexual, male and female quite different and borne

separately, inflorescences arising from underground stems.

Male flowers on peduncle ± 80.0 x 1. 5-2.0 mm of which

usually 50 mm protruding from soil bearing 25 or more

flowers, green and pilose above ground, white and gla-

brous below; bracts ± 4 mm long, often with few small

teeth on margin, lanceolate, spreading; pedicel 1 2-22 mm

long, slightly over 1 mm thick; calyx green, pilose outside,

receptacle ± 4 mm long; sepals 1. 5-2.0 x± 1.5 mm, acute;

corolla lobes spreading with recurved obtuse tips, 5-7 x

3 mm at base, outside green and pilose, inside green with

darker longitudinal stripes, fine hairs, and yellow multi-

cellular short clavate papillae towards edge and apex; an-

thers 3, white, erect. ± 3 mm long, very hairy inside

around base, one small with 1 locule, two larger with 2

locules each. Female flowers on subterranean peduncle

0-20 x 1.0-1 .5 mm. bearing 1 flower and 1-2 small bracts

at apex, white, glabrous; pedicel curving downwards, ± 2

mm thick, white with few scattered hairs; calyx pale yel-

low, sparsely and finely pilose, smooth; receptacle ini-

tially horizontal then erect, cylindrical and solid, 35-70 x

2-3 mm, widening to 5-6 mm in last 5 mm, mostly sub-

terranean; sepals to 2 mm long, lanceolate; corolla lobes

10-12 x 3-4 mm, broadest just above middle, slightly

narrower at base, with rounded obtuse apex, ascending-

spreading. pale greenish and smooth outside, bright yellow

and papillate within; staminodes 0-2 per lobe (5 per

flower), pilose, erect, ± 2 mm long, inserted near base of

petals; ovary ± ovoid, horizontal, 5-7 x 4—5 mm, pale

yellow, finely papillate; style 5-7 x 1. 5-2.0 mm with

broad cauliflower-like obscurely trifid apex, fused to re-

ceptacle about 4 mm below sepals. Fruit (only 1 seen)

± spherical, 22 mm diam., finely pubescent, green where

exposed, white beneath soil. Seeds 8, white, flattened

ovoid. 10x7 mm (not yet ripe). Flowering April to June.

Figure 5.

Specimens examined

CAPE. — 2917 (Springbok): Wildeperdehoek Pass, (-DC), Bruyns

5171 (BOL); 5 km S Pass, (-DC), Bruyns 5174 (BOL). 3017 (Hon-

deklipbaai): Riethuis, (-AB), Bruyns 4593 (BOL); 7 km S Taaibosduin,

(-AD), Bruyns 4592 (BOL): NE Soebatsfontein, (-BA), Bruyns 5184

(BOL): Swartvlei. (-BD). Bruyns 5354 (BOL); Sandkraal, (-DA), H -

Taylor 1378 (BOL); Nariep, (-DC), no material deposited. 3018

(Kamiesberg): Stofkraal, (-CB), Bmyns 4712 (BOL): Kliprand, (-DA),

Bruyns 5268 (BOL). 3118 (Vanrhynsdorp): Swartbooisvlei, (-AC),

Bayer 6229 (BOL); Draaihoek. (-CB), H, -Taylor 1146 (BOL). 3218

(Clanwilliam): Nortier, (-AB). Bayer (BOL); Bmyns 4569 (BOL. PRE.

K, M); Grootdrif, (-AD). Bruyns 4738 (BOL); Redelinghuys, (-BC),

Bruyns 4737 (BOL).

The Cucurbitaceae are a relatively small family in

southern Africa with eight genera and 72 species (Meeuse

1962; Gibbs Russell et al. 1987; De Wet et al. 1991). The

family is mainly found in those parts of the subcontinent

receiving summer rainfall with only about five species (of

the genera Melothria, Kedrostis and Cucumis) native to

the western Cape. Most of them are creepers or climbers,

often with a large tuberous rootstock and characteristic

cucumber- or pumpkin-like fruit.

According to the key in Meeuse (1962), the present

new species belongs to Kedrostis Medik. and is the ninth

species known in southern Africa.

K. psammophila has less of a climbing or scandent

habit than any other southern African species. The aerial

stems are short and usually prostrate. They lack tendrils,

which are found on all other southern African species of

Kedrostis. Beneath the surface the stems spread exten-

sively and are repeatedly branched so that the largest, cen-

tral tuber can be very difficult to locate. Subsidiary tubers

are sometimes found at the nodes at intervals along these

underground stems and it is from these underground stems

too that the inflorescences arise.

According to Meeuse ( 1962: 24), in Kedrostis , the male

and female flowers are often borne on separate inflores-

cences with the males usually clustered and the females

either clustered or solitary. This always appears to be the

234

Bothalia 23,2 (1993)

FIGURE 5. — Kedrostis psammophila. A, part of plant showing male inflorescence with many small flowers and female inflorescence

with single flower longer and larger than male; dotted line on left indicates ground level. B, C, male flower; D, dissection of

male flower; E, dissection of top of female flower; F, dissection of inflated base of female flower showing ovules; G,

staminode from female flower showing hairs; H, hairs on edge of leaf; I, papillae on corolla surface. Scale bars: A, 10 mm;

B-F, 3 mm (at B); G, I mm; H, I, 0.5 mm.

case in K. psammophila. He found that in general the

female and male calyces and corollas are identical, but

this is not true of the new species. Here the males are

borne on a ± dense pedunculate inflorescence protruding

from the soil and they are small and green. The bright

yellow female corolla is about twice the size of the male.

It has an extremely long solid receptacle with the ovary

at its base close to the stem and well below the ground

and the corolla situated just above the surface of the soil.

This leads to the remarkable situation of the fruit devel-

oping beneath the soil and makes K. psammophila, along

with Cucumis humifructus Stent, only the second known

geocarpic cucurbit in southern Africa (Meeuse 1962: 62).

Female (lowers were found to be much rarer that male

but a few were observed on the same plants as males.

The plants are therefore generally assumed to be monoe-

cious.

I have been unable to locate any herbarium records

made prior to 1986 in any of the western Cape herbaria

and it is remarkable that this widespread and common

species has not been noticed earlier. The species was

brought to my attention by M.B. Bayer, who noticed the

unique flowering habit in plants growing wild near the

Agricultural Research Station at Nortier. It has since

turned out to be widespread in the fine reddish sand found

over most of western Namaqualand from Redelinghuys

in the south to around Port Nolloth (Figure 6). It occurs

on this sand eastwards to the foothills of the Kha-

miesberge and is also found in patches of relatively soft

Bothalia 23,2(1993)

235

FIGURE 6. — Distribution of Kedrostis psammophila.

gneissic sand (much more gravelly than the coastal sands)

higher up on the southern side of the Khamiesberge as

far east as Kliprand.

Finding fruit of this species proved difficult and 1 have

only seen one (. Bruyns 5174). This had been partially ex-

posed by erosion of the soil above it and was probably

not quite fully developed. It contained eight relatively

large seeds (some dissected ovaries contained up to 10

ovules) and was extremely smelly when opened. In fact,

when damaged all parts of the plant give off the unpleas-

ant foetid odour characteristic of Kedrostis. Meeuse men-

tions that C. humifructus is dispersed by antbears. I have,

though, noticed no particular concentrations of K.

psammophila around the groundsquirrel warrens which

are common in the sandier parts of Namaqualand and its

seeds may be dispersed by moles.

REFERENCES

DE WET, B.C., ARCHER, R„ FISH, L.. GERMISHUIZEN, G„ HER-

MAN, P.P.. JORDAAN, M„ PEROLD, S.M., REID, C„ VAN

ROOY, J., WELMAN, W.G. & GLEN, H.F. 1991. New taxa, new

records and name changes for southern African plants. Bothalia

21:211.

GIBBS RUSSELL, G.E., WELMAN, W.G.. RETIEF, E„ IMMELMAN,

K.L., GERMISHUIZEN, G.. PIENAAR, B.J., VAN WYK. M. &

NICHOLAS, A. 1987. List of species of southern African plants.

Memoirs of the Botanical Sur\’ey of South Africa No. 56: 201-203.

MEEUSE, A.D.J 1962. The Cucurbitaceae of southern Africa. Bothalia

8: 1-111.

P. BRUYNS*

* Bolus Herbarium, Univ. of Cape Town, Private Bag, Rondebosch 7700.

MS. received: 1992-09-07.

SAPOTACEAE

A ■BEQUAERTIODENDRON' BY ANY OTHER NAME?

The taxonomy and nomenclature of the genus Be-

quaertiodendron De Wild, have caused considerable con-

troversy over the last thirty years. To complicate matters,

two recent revisions, one of Bequaertiodendron (Liben

1989) and the other of the genera of the Sapotaceae (Pen-

nington 1991), differ markedly from each other.

Chrysophyllum magalismontanum was described by

Sonder (1850) and based on Zeyher 1849 , a specimen

from the Magaliesberg. This species as well as C.

natalense Sond. and C. glomemliferum Hutch. & Dalz.

have fruit characters which separate them from the rest

of the genus. The need to recognise this group as a sep-

arate entity prompted various researchers to describe new

genera and sections in which members of the group could

be accommodated. Thus, Chrysophyllum L. sect. Zey-

herella Pierre ex Engl., Pachystela Pierre ex Engl. sect.

Zeyherella (Engl.) Lecomte, Tisserantiodoxa Aubrev. &

Pellegr., Zeyherella (Engl.) Aubrev. & Pellegr., Boivinella

Aubrev. & Pellegr. and Neoboivinella Aubrev. & Pellegr.

originated.

The genus Bequaertiodendron was established by De

Wildeman (1919) and was based on Bequaert 2483 from

the Belgian Congo. Chrysophyllum magalismontanum

Sond. was considered to be congeneric with the type spe-

cies of the genus, namely Bequaertiodendron congolense

De Wild. This meant that the concept of Bequaertioden-

dron had to be broadened, and this was done by Heine

& Hemsley in 1960, hence Bequaertiodendron De Wild,

emend. Heine & J.H. Hemsley.

The genus Englerophytum was described by Krause in

1914 and typified by Englerophytum stelechantha Krause.

It comprises five to ten poorly defined species in tropical

Africa. Bequaertiodendron was reduced to synonymy

under Englerophytum by Aubreville in 1960. However,

Heine & Hemsley (1960, 1968) did not recognise this

genus for the following reasons: the type material was

thought to have been destroyed, the diagnosis and

illustrations did not fully agree, and the genus was de-

scribed without fruits.

Liben (1989), in a revision of the West and Central

African species, concluded that the material placed in

Bequaertiodendron magalismontanum by Heine &

Hemsley (1960) could be divided into eight different spe-

cies. On the basis of macromorphological floral and seed

characters, these species were grouped into three genera,

namely Englerophytum Krause, Wildemaniodoxa Aubrev.

& Pellegr., and Zeyherella (Pierre ex Engl.) Aubrev. &

Pellegr. Liben (1989) reduced Bequaertiodendron maga-

lismontanum to synonymy under Zeyherella magalismon-

tana (Sond.) Aubrev. & Pellegr.

Steyn (1990), in an attempt to retain Bequaer-

tiodendron magalismontanum, used macromorphological

characters of female and hermaphroditic flowers from

gynodioecious populations to evaluate Liben' s concept of

B. magalismontanum sensu lato. According to Steyn

( 1990), characters which distinguish the southern African

B. magalismontanum from the tropical African material

are the presence of a stylar compitum, an ovarial nectary.

236

Bothalia 23,2 (1993)

introrse apiculate stamens, or staminodes derived from

such stamens, and a tendency in female plants to increase

to ten the number of corolla lobes, of epipetalous

staminodes and of locules. It is also emphasised that the

staminal structure of southern African plants and En-

glerophytum ( sensu Liben 1989) differ considerably. In

the latter genus short filaments are united in a staminal

tube that adheres tightly to and completely covers the pis-

til and bears nonapiculate anthers.

Pennington (1991) reduced Bequaertiodendron to syn-

onymy under Englerophytum, because of the rediscovery

of authentic material of the type species of E. ste-

lechanthum (Aubreville 1971). He pointed out that closely

related species groups in Sapotaceae are often highly vari-

able in the number of their floral parts. An examination

of the Englerophytum complex throughout its range

showed that there was no clear distinction between free

stamens and those fused into a tube, but rather a range of

conditions from free, through slightly fused to completely

fused. As a whole, the complex is characterised by a ten-

dency towards fusion.

Concerning the genus Bequaertiodendron in southern

Africa, it is clear that Englerophytum is the earlier name

and Pennington’s treatment of the genus should be fol-

lowed. Nevertheless, on the basis of characters pointed

out by Steyn (1990), it is recommended that more research

should be undertaken at species level within Engle-

rophytum. The NBI agrees with Pennington’s views and

accepts the following name changes:

Englerophytum magalismontanum (Sond.) Pen-

nington, The genera of Sapotaceae: 252 (1991).

Chrysophyllum magalismontanum Sond.: 72 (1850). Pachystela

magalismontana (Sond.) Lecomte: 189 (1919). Zeyherella magalismont-

ana (Sond.) Aubrev. & Pellegr.: 37 (1958). Pouieria magalismontana

(Sond.) Meeuse: 335 (1960). Bequaertiodendron magalismontanum

(Sond.) Heine & Hemsley: 307 (1960).

Englerophytum natalense (Sond.) Pennington,

The genera of Sapotaceae: 252 (1991).

Chrysophyllum natalense Sond.: 72 (1850). Boivinella natalensis

(Sond.) Pierre ex Aubrev. & Pellegr.: 37 (1958). Neoboivinella natalensis

(Sond.) Aubrev. & Pellegr.: 23 (1959). Pouteria natalensis (Sond.)

Meeuse: 339 (1960). Bequaertiodendron natalense (Sond.) Heine &

Hemsley: 308 (1960).

ACKNOWLEDGEMENTS

Dr Donald J.B. Killick is thanked for constructive criti-

cism.

REFERENCES

AUBREVILLE. A. 1960. Notes sur les Sapotacees de l'Afrique

equatorial e. Notulae Systematicae 16: 253.

AUBREVILLE. A. 1971. Essais de geophyletique des Sapotacees 2.

Adansonia 11: 425.

AUBREVILLE, A. & PELLEGRIN. F. 1958. Rehabilitation de deux

genres de Sapotacees. Bulletin Societe botanique de France 105:

35-37.

AUBREVILLE, A. & PELLEGRIN, F. 1959. Rectification au sujet de

Sapotacees africaines. Bulletin Societe botanique de France 106:

22, 23.

DE WILDEMAN, E. 1919. Sur quelques especes congolaises de la

famille des Sapotacees. Revue Zoologique Africaine 7, Sup-

plement Botanique: 21, 22.

HEINE. H. & FIEMSLEY, J.H. 1960. Notes on African Sapotaceae II.

The genus Bequaertiodendron De Wild. Kew Bulletin 14: 304.

HEINE. H. & HEMSLEY. J.H. 1968. Sapotaceae. In E. Milne-Redhead

& R.M. Polhill, Flora of tropica! East Africa: 18-24. Whiteffiars

Press, London.

KRAUSE, K. 1914. Englerophytum , eine neue afrikanische Gattung der

Sapotaceen. Botanische Jahrbiicher fur Systematik, Pflan-

zengeschichte und Pflanzengeographie 50: 343.

LECOMTE. H. 1919. Quelques Sapotacees africaines. Bulletin du Mu-

seum d'histoire naturelle 25: 189-193.

LIBEN, L. 1989. La veritable identite des genres et especes confondus

sous le nom de Bequaertiodendron magalismontanum (Sond.)

Heine & Hemsley (Sapotaceae) en Afrique centrale et oc-

cidentale. Bulletin du Jardin botanique national de Belgique 59:

151-169.

MEEUSE, A.D.J. I960. Notes on the Sapotaceae of southern Africa.

Bothalia 7: 339.

PENNINGTON. T.D. 1991. The genera of Sapotaceae. Royal Botanic

Gardens. Kew, Surrey.

SONDER. W. 1850. Beitriige zur Flora von Siidafrika. Linnaea 23: 72.

STEYN. E.M.A. 1990. Macromorphological floral characters in

Bequaetliodendrott magalismontanum: possible taxonomic sig-

nificance. South African Journal of Botany 56: 440-442.

C. L. BREDENKAMP* and G.F. SMITH*

*National Botanical Institute, Private Bag X 101 . Pretoria 0001 .

MS. received: 1993-10-05.

ASCLEPIADACEAE

VALIDATION OF THE COMBINATION ASPIDONEPSIS REENENSIS (ASCLEPIADACEAE):

THE TYPE SPECIES OF THE SUBGENUS UNGUILOBIUM

In our paper describing the new African genus As-

pidonepsis (Nicholas & Goyder 1992), the basionym of

A. reenensis was inadvertently omitted, so invalidating the

proposed combination. As this invalid combination is the

type of the proposed subgenus Unguilobium this too be-

comes invalid. The present note is intended to correct

these omissions.

Unguilobium A. Nicholas & D.J. Goyder ex A.

Nicholas & D.J. Goyder in Bothalia 22: 31, 32 (1992),

subgen. nov.

Folia ascendentia, margine manifeste revoluta. //;-

florescentia 4-11 -flora. Corolla reflexa; pagina abaxialis

pubescentia. Coronae lobi ad columnam staminalem circa

1 mm super insertionem corollae conjuncti, cucullati; ap-

pendix proximalis ad apicem deltato-falcata et apicem

styli aequans vel superans impendensque; extremum dis-

tale coronae appendice parva omatum (A. reenensis) vel

appendice carente (A. shebae ); sinus profundus rimi-

formis.

TYPUS. — Aspidonepsis reenensis (N.E. Br.) A. Nich-

olas & D.J. Goyder vide infra.

Bothalia 23,2 (1993)

237

Aspidonepsis reenensis (N.E. Br.) A. Nicholas &

D.J. Goyder , comb. nov. Type: South Africa, Natal, Van

Reenen, Wood 8635 (K!, holo.; GRA!, NH!. PRE!, SAM!,

iso.)

Asclepias reenensis N.E. Br. in W.T. Thiselton-Dyer, Flora capensis

4,1: 1131 (1909).

ACKNOWLEDGEMENTS

We are most grateful to both Colin Walker of the Open

University, Milton Keynes, U.K. and Piet Vorster of the

Botany Department, Stellenbosch University, South Africa

for drawing the invalid publication of Aspidonepsis

reenensis to our attention.

REFERENCE

NICHOLAS. A. & GOYDER, D.J. 1992. Aspidonepsis (Asclepiadaceae),

a new southern African genus. Bothalia 22: 23-35.

NICHOLAS A* & GOYDER, D.J.**

* Botany Department, University of Durban-Westville, Private Bag

X54001, Durban 4000, Natal, South Africa.

** The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey,

TW9 3AB, England.

Bothalia 23,2: 239-263 ( 1993)

The vegetation of the southern Langeberg, Cape Province. 3. The plant

communities of the Bergfontein, Rooiwaterspruit and Phesantefontein areas

D.J. MCDONALD*

Keywords: classification, forest, fynbos, Langeberg, phytosociology

ABSTRACT

The fynbos shrubland communities of the southeastern Langeberg, Cape Province were analysed after sampling 97 quadrats

in the Bergfontein, Rooiwaterspruit (Garcia’s Forest Reserve) and Phesantefontein areas. Data were initially analysed using

TWINSPAN and the resulting classification refined using Braun-Blanquet (BB) procedures. One Afromontane forest commu-

nity and 16 fynbos communities are recognized and described. A hierarchical classification of the fynbos communities is

proposed.

UITTREKSEL

Die fynbos-struikgemeenskappe van die suidoostelike Langeberge, Kaapprovinsie, is geanaliseer. ’n Totaal van 97

monsterpersele in die Bergfontein-. Rooiwaterspruit- (Garcia Staatsbos) en Phesantefontein-gebiede is ondersoek. Data is

aanvanklik deur Twinspan geanaliseer en die resultaat d.m.v. Braun-Blanquet prosedures (BB) verfyn. Een Afro-montane

woudgemeenskap en 16 fynbosgemeenskappe word erken en beskryf. 'n Hierargiese klassifikasie van die fynbosgemeenskappe

word voorgestel.

CONTENTS

Introduction 239

Study area

Location 240

Physiography and geology 241

Soils 241

Climate 242

Methods 243

Vegetation 243

Afromontane Forest 244

Fynbos 244

1 . Erica hispidula Shrublands 244

1 . 1 Erica hispidula-Brunia alopecuroides Shrub-

lands 244

1 .2 Erica hispiduIa-Restio inconspicuus Shrub-

lands 249

1.2.1 Restio inconspicuus-Chondropetalum mu-

cronatum Shrublands 249

1 .2.2 Restio inconspicuus-Selago serrata Shrub-

lands 251

1 .2.3 Restio inconspicuus-Erica melanthera

Shrublands 252

1 .3 Erica hispidula-Tetraria bromoides

Shrublands 252

1 .3. 1 Tetraria bromoides-Berzelia galpinii

Shrublands 253

1.3.2 Tetraria bromoides-Ischyrolepis hystrix

Shrublands 253

1 .3.2.1 Ischyrolepis hystrix-Phylica rubra

Shrublands 253

1.3. 2. 2 Ischyrolepis hystrix-Phylica pinea

Shrublands 254

* Conservation Biology Research Unit, National Botanical Institute, Pri-

vate Bag X7, Claremont 7735.

MS. received: 1992-1 1-02.

1.3.3 Tetraria bromoides-Hypodiscus striatus

Shrublands 254

1.3.4 Tetraria bromoides-Protea coronata Shrub-

lands 255

1.4 Erica hispuhda-Protea nitida Shrublands .. 256

2. Erica versicolor-Agathosma ovata Shrub-

lands 257

3. Cullumia aculeata var. aculeata Shrublands . 257

3.1 Cullumia aculeata-Eeucadendron eucalyp-

tifolium Shrublands 258

3.1.1 Leucadendron eucalyptifolium-Protea ne-

riifolia Shrublands 258

3. 1 . 1 . 1 Protea neriifolia—Merxmuellera decora

Shrublands 258

3. 1 . 1 .2 Protea neriifolia-Erica articularis Shrub-

lands 258

3.1 .2 Leucadendron eucalyptifolium-Elegia fda-

cea Shrublands 259

3.2 Cullumia aculeata-Protea lorifolia Shrub-

lands 259

3.2.1 Protea lorifolia-Ficinia laciniata Shrub-

lands 260

3.2.2 Protea lorifolia-Leucospermum calligerum

Shrublands 260

Discussion and conclusions 261

Acknowledgements 262

References 262

Errata: The vegetation of the southern Langeberg,

Cape Province. 2 263

INTRODUCTION

A survey of the vegetation of the southern Langeberg

was undertaken to describe the plant communities of this

previously poorly documented mountain range. The de-

scriptions of plant communities presented here are based

on samples taken on the third of three sample transects

240

Bothalia 23.2 (1993)

FIGURE 1 . — Map of the mountains of the Fynbos Biome showing the position of the Langeberg and the location of the Bergfontein-Rooiwaterspruit-

Phesantefontein areas. B, Barrydale; C, Cape Town; G. George; H, Heidelberg; M, Mossel Bay; P, Port Elizabeth; R. Riversdale; S.

Swellendam and W, Worcester.

straddling the Langeberg at intervals between Swellendam

and the Gouritz River. Each transect is treated separately

(McDonald 1993a & b) but whereas the Marloth Nature

Reserve (MNR) and Boosmansbos Wilderness Area

(BWA) transects were more or less continuous from south

to north, the third transect is a composite of samples from

the Bergfontein area and the Rooiwaterspruit-Phe-

santefontein area (BRP) (Figure 1). This presented certain

problems with respect to data analysis and interpretation

which are discussed below.

STUDY AREA

Location

Numerous extensive wildfires have occurred on the

Langeberg between Garcia's Pass and the Gouritz River

in the past ten years (C. Martens pers. comm.). The fynbos

vegetation on this part of the Langeberg is therefore al-

most all in a juvenile phase (< 1 0 years) (Kruger 1979).

Little choice was left in finding mature fynbos along a

continuous transect for sampling vegetation on the third

transect over the southern Langeberg. A transect which

satisfied most logistic and sampling criteria was selected

in the Bergfontein area, 33° 58' S, 21° 33' E. This area

lies northeast of Riversdale, approximately midway be-

tween Garcia’s Pass and the Gouritz River and is in the

most easterly zone of the Langeberg. One negative feature

was that the vegetation at Bergfontein was seven years

old. However, despite the structural immaturity of the

fynbos communities, the vegetation was in the ‘maturing’

successional phase (Kruger 1979) with the non-ephemeral

species well established. It was therefore accepted that the

Hol istic composition would reflect the ‘mature’ phase ad-

equately and sampling could proceed.

The boundaries of the study area at Bergfontein fol-

lowed the boundary between State Forest land and private

agricultural land. The northern extreme of the designated

transect extended onto private land where the vegetation

was not disturbed by agriculture.

The Bergfontein transect followed the bridle path

(Muir 1929) and sampling was largely confined to southl-

and southwest-facing slopes east of Witelsrivier. Four

sample plots were located west of Witelsrivier opposite

the western end of Koksposberg to accommodate the

Erica hispidula-Protea nitida Shrublands not found in the

remainder of the study area.

Shortly after 74 releves had been recorded on the

southern slopes of the Bergfontein transect, a wildfire de-

stroyed all the vegetation on the transect. Therefore alter-

native sites with vegetation and habitat as similar as

possible to the vegetation on the northern slopes and high-

altitude south-facing slopes had to be found. Since no suit-

able sites were available east of Garcia’s Pass, alternative

sites were chosen west of Garcia’s Pass above Rooi-

waterspruit (high-altitude south-facing slopes) and above

Phesantefontein (north-facing slopes) 33° 57' S, 21° 9' E.

At Rooiwaterspruit, which falls within the Garcia’s

Forest Reserve, four releves (301-304) were recorded in

vegetation on the Cedarberg Formation shaleband east of

Stinkhoutbos. One plot (297) was located on the western

side of the watershed between the headwaters of

Rooiwaterspruit and Korinte River and three plots (298-

300) were located on the steep, high-altitude south-facing

slopes of the unnamed peak immediately west of Sleeping

Bothalia 23,2(1993)

241

Beauty (near trigonometric beacon 48). This latter peak

is referred to as Korinteberg.

Above Phesantefontein, eight plots (286-293) were lo-

cated in seven-year-old veld on the mid-altitude north-fac-

ing slopes east of Aasvoelkrans (Garcia's Forest Reserve)

and a further six (282-285, 295 & 296) in mature veld

(>20 years) on the lower northern foothills of the

Langeberg (Phesantefontein Farm).

Physiography and geology

Bergfontein

The south slopes of the Langeberg above Bergfontein

are not as steep as the south slopes of the range further

west. At the forefront of the mountain is Koksposberg, a

prominent, isolated, low ridge-like hill or ‘koppie’. This

hill and others similar to it, to the west and east, ranging

in altitude from 457-727 m and situated between the

Kaffirkuils and Gouritz Rivers, are the remnants of the

Peninsula Formation sandstone which is much more

prominent further west. South of Koksposberg is a

silcrete-capped mesa which supports fynbos. This area

was excluded from the study since it fell outside the south-

ern boundary of the transect and has been disturbed by

agricultural practices. Immediately north of Koksposberg

is the Cedarberg Formation shaleband which separates the

Peninsula Formation sandstone from the Nardouw Sub-

group sandstone. There is no deeply incised intennontane

valley following the shaleband as there is in other parts

of the Langeberg e.g. in BWA (McDonald 1993a). North

of the shaleband is another 'koppie', of higher altitude

than Koksposberg consisting of Nardouw Subgroup sand-

stone. Further north is Witelsberg which is a massive block

of Nardouw Subgroup sandstone with steep south and

north sides but with an expansive 'flat' top with shallow

gradient from the west to the peak at 1 166 m at the

eastern end.

Below the steep north slopes of Witelsberg is a plateau

extending from west of Witelsberg to Waboomsrivier.

North of the plateau is a dissected sandstone ridge which

gives way to arid sandstone slopes. They in turn make

contact with the Bokkeveld Group shales of the Little

Karoo.

Rooiwaterspruit and Phesantefontein

In the Garcia’s Forest Reserve west of Garcia’s Pass,

the Peninsula Formation sandstone forms massive peaks

on the southern side of the mountain range. Sleeping

Beauty (1 343 m) is the most prominent peak with

Korinteberg ( 1 330 m) to the west and lower peaks further

westwards towards Gysmanshoek. Behind the peaks lie

two incised valleys where streams have eroded the shales

of the Cedarberg Formation. North of the shaleband lies

the Nardouw Subgroup sandstone with Aasvoelkrans west

of Stinkhoutbos reaching 1 341 m and another prominent

peak east of Stinkhoutbos with a height of 1 330 m, re-

ferred to here as Phesanteberg.

Below the southern slopes of the mountain the Penin-

sula Formation sandstones make contact with silcrete-

capped plateau-like mesas which in turn are dissected by

numerous watercourses. The high northern slopes of Nar-

douw Subgroup sandstone grade at a moderate angle to

the undulating foothills at Phesantefontein. The foothills

consist of sandstone overburden and the remains of a once

extensive silcrete-capped landscape. Remnant mesas are

encountered extending well into the Little Karoo (Figure

2).

Soils

The soils of the Bergfontein and Rooiwaterspruit-

Phesantefontein areas of the Langeberg are the same as

those of the MNR and BWA (McDonald 1993a & b) at

the level of 'form' (SCWG 1991). The climate of the three

areas is similar (see below) as is the geology. Topograph-

ical variation and changes in parent material therefore ac-

count for most differences between the soils encountered

on the BRP 'transect' and those of BWA and MNR. A

description of the soil forms found on the southern

Langeberg is given in McDonald (1993a); a summary of

soil forms found on the BRP ‘transect’ follows:

Champagne Form soils are found at sites where there

is a a deep (>200 mm) accumulation of organic material.

On the BRP transect such sites are found mainly at high

altitude on cool, moist south-facing slopes e.g. on Ko-

rinteberg. These soils also occur at lower elevations often

in depressions where plant remains collect.

The catena of 'non-organic’ soils derived from sand-

stone parent material in the study area include Houwhoek,

Cartref, Glenrosa and Mispah Forms. Houwhoek and Car-

tref Form soils are closely related. The characteristics they

share are an orthic A-horizon and presence of an E-hori-

zon. They differ in that Houwhoek Form has a podzolised

B-horizon overlying saprolite, whereas Cartref Form has

a lithocutanic B-horizon. Glenrosa Form is similar to Car-

tref but lacks the eluviated E-horizon of the former.

Mispah Form in turn is similar to Glenrosa Form, but here

the orthic A-horizon overlies hard rock as opposed to the

lithocutanic B-horizon of Glenrosa Form.

FIGURE 2. — Silcrete mesas on the north flank of the Langeberg, ex-

tending into the Little Karoo, at Phesantefontein.

242

Bothalia 23,2 (1993)

Clovelly Form soils which have an orthic A-horizon

over a yellow-brown apedal B-horizon are found in two

situations derived from different parent materials: 1, on

the Cedarberg Formation shale both at Bergfontein and at

Rooiwaterspruit, behind Korinteberg; and 2, where there

is accumulation of sand derived from Nardouw Subgroup

sandstones, on dry north-facing slopes where leaching is

limited.

Oakleaf Form soils which have limited extent in BWA

(McDonald 1993a) but which were not identified in MNR

occur on the lower south slopes at Bergfontein, below

Koksposberg and on the west side of Witelsrivier. These

soils which result from mixing of shale and sandstone

have an orthic A-horizon and a reddish brown neocutanic

B-horizon.

Climate

Characteristics of the climate of the BPR ‘transect1 are

similar to those described by McDonald (1993a & b) for

BWA and MNR. The southern Langeberg falls within a

uniform bioclimatic zone, transitional between the winter

rainfall region in the west and the year-round rainfall re-

gion in the east. The major climatic events which affect

the weather of the southern Cape coast and the coastal

mountains are the passage of cold fronts advancing from

west to east, coastal lows and cutoff lows (Fuggle 1981;

Van Heerden & Flurry 1987). A Walter-Lieth climate di-

agram for Riversdale (Figure 3A) gives an approximation

of the climate of the lower south slopes of the Langeberg

in the Riversdale-Gouritz River District.

Winds

The direction of surface winds affecting the southern

Langeberg is dependent on season with summer winds

blowing mainly onshore and winter winds mainly off-

shore. Calm periods occur approximately one-third of the

time in both summer and winter (Schulze 1965).

Extreme fire-hazard conditions prevail when hot, dry

berg winds occur in the southern Cape, mainly in winter

(Van Wilgen 1984). They arise from air subsiding in re-

sponse to pressure gradients between an established anti-

cyclone and an advancing depression (Fuggle 1981). The

lire which swept through the Bergfontein area in May

1991 occurred in such conditions.

Precipitation

Precipitation on the Langeberg is from rain, mist and

snow. Snowfalls occur once or twice each winter mainly

on the high peaks and ridges and do not persist. Mist is

a year-round phenomenon, usually occurring at elevations

from 1 000 m upwards. Rainfall occurs throughout the

year with peaks in autumn (April) and spring (October).

It may occur from unstable prefrontal conditions but is

most often associated with postfrontal events. Eastward-

moving cold fronts are usually followed by a ‘ridging-in’

by the South Atlantic anticyclone behind the fronts. Air

movement over the warm Agulhas Current results in on-

shore advection of cool moist air (Cowling 1984; Pres-

ton-Whyte & Tyson 1988). Orographic rain occurs when

this moist air strikes the coastal mountains.

RIVERSDALE (122 m)

FIGURE 3. — A, climate diagram for Riversdale which approximates the

climate of the lower south slopes of the Langeberg in the

Riversdale-Gouritz River District; B. monthly rainfall ( 1 984—

1991) at Waboomsrivier on the north side of Witelsberg, showing

peaks in April (autumn) and October (spring).

The high peaks in the MNR at Swellendam and in

BWA receive an estimated 1 200-1 400 mm rainfall per

annum. Proceeding eastwards, Kanetberg receives 500-

600 mm, Aasvoelkrans 600-700 mm, Korinteberg 700-

800 mm and Sleeping Beauty 800-900 mm rainfall p.a.

East of Garcia’s Pass the peaks receive from 800-900 mm

rainfall but the Bergfontein area (including Witelsberg

south slopes) is much drier, receiving 600-700 mm rain-

fall p.a. (Dent et al. 1987). This gradient is opposite to

that reflected in the mean annual rainfall for Heidelberg

(378 mm) and Riversdale (426 mm) (Rebelo et al. 1991).

The difference is attributed to the effect of orographic

rainfall on the Langeberg; the more westerly higher alti-

tude peaks receive more rainfall than the somewhat lower

eastern part of the Langeberg close to the Gouritz River.

The north slopes of Witelsberg are in a rain shadow

and are consequently much drier than the southern slopes.

The mean annual precipitation for a seven-year period

(1984—1990) at Waboomsrivier at the northern base of

Witelsberg was 496 mm (P.E. Wadman pers. comm.); sea-

sonal distribution of the rainfall is as shown in Figure 3B.

Rooiwaterspruit and Korinteberg receive a mean annual

precipitation of 700-800 mm, whereas the upper north

slopes above Phesantefontein (east of Aasvoelkrans) re-

ceive 500-600 mm and the lower north slopes 300-500

mm depending on locality (Dent et al. 1987).

Temperature

No temperature data are available for the study area;

a situation commonplace in the Cape mountains where

there are few weather stations (Bond 1981; Fuggle & Ash-

Bothalia 23,2 ( 1993)

243

ton 1979; Fuggle 1981). Extrapolation of temperature data

from lowland' weather stations to montane situations has

been done (Van Wilgen 1984) but this does not reflect the

true montane temperature regime in most cases. The clos-

est temperature recording station to the study area is at

Riversdale. Temperatures recorded here may approximate

those experienced on the lower south slopes of the

Langeberg. Therefore, if the environmental lapse rate of

0.6°C/100 m (Cowling 1984) is used to predict montane

temperatures, the high altitude south-facing slopes of

Witelsberg at 1 1 50 m are predicted to have temperatures

6.26°C lower than Riversdale at 106 m.

Solar radiation

Incoming radiation may be measured directly (Morris

1981), which is cumbersome in mountainous terrain, or

derived from sunshine duration. No data are published for

sunshine duration on the Langeberg. Bond (1981) used

Swift’s (1976) algorithm to calculate potential radiation

for a range of slopes and aspects for the 33° 30' S latitude

which is roughly equivalent to the latitude for the southern

Langeberg. Bond (1981) found that potential radiation on

the Swartberg and Outeniqua Mountains is relatively sim-

ilar on all aspects and slopes in summer, with marked

differences between north and south aspects, particularly

on steep slopes, in winter. This holds for the Langeberg

as well.

METHODS

Ninety-seven 5 x 10 m quadrats (McDonald 1983,

1988, 1993a & b; Campbell 1985; Boucher 1987) were

sampled on a 'composite transect’ over the Langeberg in

the Bergfontein. Rooiwaterspruit and Phesantefontein

areas. No stratification of the study area was undertaken

since no suitable aerial photography was available. Plots

were therefore subjectively placed at sites representative

of major landscape features and vegetation communities.

Floristic, structural and environmental data were col-

lected from each sample plot. Permanently recognizable

species were recorded using the Braun-Blanquet cover-

abundance scale (Mueller-Dombois & Ellenberg 1974;

Werger 1974; Westhoff & Van der Maarel 1973). The mid-

point of the BB values given as percentage cover is as

follows: 5 = 87.5%; 4 = 62.5%; 3 = 37.5%; 2 = 15.0%;

1 = 2.5%; + = 0. 1% ; R = value ignored. Ephemeral geo-

phytes and annuals were noted in each releve but were

not used in analyses and descriptions of communities. A

border zone of 1 .5 m from the perimeter of each plot was

searched for any species not found in the marked plot.

Species occurring outside the plot are represented by ‘O’

in the phytosociological tables. Vegetation structure was

measured by estimating the projected canopy cover and

height of the respective strata. Environmental variables re-

corded include altitude, aspect, slope, geology, soil form,

drainage and estimated soil depth. Mean annual rainfall

was estimated from isohyet maps prepared by Dent et al.

(1987).

Samples were taken only in fynbos shrubland commu-

nities. Stinkhoutbos in the Rooiwaterspruit area is a well-

developed stand of Afromontane Forest but was not

formally sampled (see below).

Two-way Indicator Species Analysis — TWINSPAN

(Hill 1979a) was used to obtain an initial tabular clas-

sification of the data. This was followed by successive

refinement of the phytosociological tables following the

Braun-Blanquet method (Mueller-Dombois & Ellenberg

1974; Werger 1974) using the PCTables programs (Bou-

cher pers. comm.). Detrended Correspondence Analysis —

DECORANA Hill 1979b) was used to assess the re-

lationship of the Erica versicolor-Agathosma ovata

Shrublands to the other shrublands in the Bergfontein area

(see below).

The plant communities are described in the order of

the proposed hierarchical classification (see above). No

syntaxonomic rank is assigned to any given community.

Structural description follows the a priori system of

Campbell et al. (1981).

The ‘relationships’ between communities described in

this study and those described in other, previous studies

of Mountain Fynbos were determined on the basis of flor-

istic (mainly) and structural similarity. The similarities

were determined from published descriptions and phyto-

sociological tables. No rigid system was applied and the

relationships serve merely as a guide for future synthesis

of fynbos communities.

VEGETATION

The primary objective of a study such as this is to

characterize the plant communities of a given area so that

they may be repeatedly identified where they occur in the

landscape. The early descriptions of the vegetation of the

southeastern Langeberg by Muir (1929) in his treatment

of the vegetation of the Riversdale area are highly infor-

mative but, apart from a broad classification, do not pro-

vide clear delineations of the fynbos communities. The

fynbos vegetation sampled on the composite BPR

‘transect’ is classified into 16 communities: 10 at Berg-

fontein, two at Rooiwaterspruit and the remaining four at

Phesantefontein. One Afromontane Forest community is

recognized. The classification of the plant communities of

the BPR transect is not complete since the scale of the

study dictated that not all communities could be exhaus-

tively sampled in the limited study area. Emphasis is on

the sclerophyllous fynbos plant communities, since com-

munities of this type make up the major part of the vege-

tation in the study area. Afromontane Forest communities

are extremely limited in extent.

The vegetation of the south slopes of the Bergfontein

area is mainly wet to mesic proteoid fynbos with a few

isolated patches of trees in protected places. These trees

such as Cunonia capensis are representative elements of

Afromontane Forest. A well-developed stand of this forest

type is located on an east-facing cliff of Perdeberg oppo-

site Witelsberg. This forest is difficult to reach and fell

outside the study area so it was not sampled. On the north-

ern extreme of the Bergfontein transect, i.e. on the lower

north slopes of Witelsberg, dry proteoid fynbos is encoun-

tered.

At Rooiwaterspruit. the south slopes also support wet

proteoid and ericaceous fynbos with one well-developed

patch of Afromontane Forest, Stinkhoutbos, in a moist

244

Bothalia 23,2 (1993)

kloof or ravine. The north-facing slopes above Phe-

santefontein have mesic proteoid to dry asteraceous

fynbos communities.

During preparation of the phytosociological tables it

was initially doubted whether releves taken in mature or

senescent vegetation (sensu Kruger 1979) would be sat-

isfactorily accommodated, for reasons such as lack of dif-

ferential species and overriding dominance of tall shrubs.

It was interesting to note, however, that these releves were

appropriately placed in the tables and that they gave in-

sights into the nature of the respective communities when

they reach the mature and senescent phases (see 1.3.1 and

1.3.2. 1 below).

Afromontane Forest

The description of Afromontane Forest is confined to

the community found at Stinkhoutbos (Figure 4). No sam-

ples were taken in this forest which covers approximately

2 ha. A list of species confirms that it may be classified

as the Cunonia capensis-Platylophus trifoliatus Sub-

association (McKenzie 1978), also found in BWA (Mc-

Donald 1993a). As the colloquial name of the forest stand

suggests, the stinkwood Ocotea bullata is common and

co-dominant with Cunonia capensis and Platylophus tri-

foliatus in the canopy. Virgilia oroboides forms large trees

mainly on the forest margins. Plectranthus fruticosus is

the dominant understorey shrub.

Fynbos

1 . Erica hispidula Shrublands

The role of Erica hispidula in linking the fynbos com-

munities of the mesic to wet slopes of the southwestern

and southern Cape mountains has been indicated by Mc-

Donald (1993a & b). The south slopes of the Langeberg

at Bergfontein and Rooiwaterspruit are no exception.

Erica hispidula is found in all the fynbos communities at

Bergfontein except in the Tetraria bromoides-Phylica

FIGURE 4. — Stinkhoutbos, a patch of Afromontane Forest in a secluded

kloof at Rooiwaterspruit.

pinea Shrublands and with little occurrence in the wet

Erica hispidula-Brunia alopecuroides Shrubland sampled

on Korinteberg. The Erica hispidula-Restio inconspicuus

Shrublands include the very wet high altitude shrublands

on the south-facing slopes of the peaks, as well as shrub-

lands forming part of the mosaic of communities on the

mid and lower south-facing slopes of the Bergfontein area.

1 . 1 Erica hispidula-Brunia alopecuroides Shrublands

Differential species: Brunia alopecuroides, Erica al-

bens, E. mucronata, E. regerminans, E. transparens, Hip-

pia integrifolia, Helichrysum capense, Indigofera con-

cava, Lobelia pubescens var. rotundifolia, Restio fragilis.

Dominant species: Anthochortus crinalis, Brunia alo-

pecuroides, Spatalla parilis, Platycaulos compressus.

Structural formation: Low Closed Restioid Shrubland.

Relationships: Brunia alopecuroides-Restio bifidus

Community (Kruger 1974); Subcommunity E2 of the

Erica-Penaea Community (Glyphis et al. 1978); Restio-

Hypolaena Subcommunity (H & I) (Laidler et al. (1978);

Ericoid-Restioid Zone Fynbos (Taylor 1978); Low Nar-

row-sclerophyllous Heathland ( Kruger 1979); Simocheilus

carneus-Restio anceps Community (Bond 1981); Wet

Mountain Fynbos (Moll et al. 1984); Ruitersberg Wet Eri-

caceous Fynbos (Campbell 1985); Erica hispidula-Spa-

talla nubicola Shrublands and Restio inconspicuus-

Anthochortus crinalis Shrublands (McDonald 1993a): An-

thochortus crinalis-Erica curviflora Shrublands (McDon-

ald 1993b).

Owing to the fire in the Bergfontein area during this

survey, sampling of the vegetation of the high altitude

south-facing slopes of Witelsberg was not possible. As a

substitute, four samples (297-300) were taken on similar

slopes on Korinteberg. The community represented is

equivalent to that found on Witelsberg prior to the fire

(D.J. McDonald pers. obs.).

Moist air moving onshore from the southern Cape

coast, together with mist and stratus cloud result in high

orographic precipitation on the high altitude (above 1 000

m) south-facing slopes. The high precipitation, low tem-

perature and reduced insolation result in low rates of or-

ganic matter decay and consequent accumulation of deep

layers of peat-like material. This deep acid peat (pH 3.0

in 1 mol/1 CaCb) with underlying Peninsula Formation

sandstone, or Champagne Form soil, supports the Erica

hispidula-Brunia alopecuroides Shrublands which are

typical of these habitats (McDonald 1993a & b).

The Erica hispidula-Brunia alopecuroides Shrublands

on Korinteberg (Figure 5) are well differentiated tloristi-

cally (Table 1 ) with dense stands of mid-high Brunia alo-

pecuroides shrubs dominating the upper stratum. Spatalla

nubicola (Proteaceae), a narrow endemic, which is found

in the equivalent shrubland community in the BWA (Mc-

Donald 1993a) is replaced by the closely allied S. parilis

on Korinteberg. Anthochortus crinalis (Restionaceae), a

dense mat-res, tio, dominates the understorey.

A specimen of Erica dodii was found in plot 298 on

Korinteberg, which is the first record of this species on

Bothalia 23,2(1993)

245

TABLE 1. Phytosociological table of the Erica hispidula Shrublands on the south slopes of the Bergfontein area, southern Langeberg

246

Bothalia 23,2 (1993)

TABLE 1. — Phytosociological table of the Erica hispidula Shrublands on the south slopes of the Bergfontein area,

southern Langeberg (continued)

Releve number 1 2223*2222222*2222222222*222222222222*2222222222*3333*222222*22222222222*222222*2222 1

19990*5566777*3335555777*223334445577*1144444566*0000*233335*11122222466*111224*6666 i

j 7890*8909346*1271267012*890890563457*2313489534*1234*734560*78902346212*456157*5678 j

Bothalia 23,2(1993)

247

TABLE 1. Phytosociological table of the Erica hispidula Shrublands on the south slopes of the Bergfontein area,

southern Langeberg (continued)

Re 1 eve number

2223*2222222*2222222222*222222222222*2222222222*3333*222222*22222222222*222222*2222

9990*5566777*3335555777*223334445577*1144444566*0000*233335*11122222466*111224*6666

7890*8909346*1271267012*890890563457*2313489534*1234*734560*78902346212*456157*56781

248

Bothalia 23,2 (1993)

TABLE 1. Phytosociological table of the Erica hispidula Shrublands on the south slopes of the Bergfontein area,

southern Langeberg (continued)

Bothalia 23,2(1993)

249

TABLE 1. — Phytosociological table of the Erica hispidula Shrublands on the south slopes of the Bergfontein area,

southern Langeberg (continued)

the Langeberg mountain range. A small population of the

rare Langeberg endemic species Empleurum fragrans

(Rutaceae) was also located close to plot 298 but not in

the Erica hispidula-Brunia alopeciiroides Shrubland; this

represents a range extension of some 30 km east of its

previously recorded range (Williams 1984). Eeucadendron

radiation (Proteaceae), also a Langeberg endemic, is

found on rocky promontories amongst the more uniform

Erica hispidula-Brunia alopecuroides Shrubland.

FIGURE 5. — The Erica hispidula-Brunia alopecuroides Shrublands on

the high altitude south-facing slopes of Korinteberg.

1 .2 Erica hispidula-Restio inconspicuus Shrublands

This community, similar to the shrublands of the same

name in BWA, comprises most of the shrublands where

Restio inconspicuus is present. R. inconspicuus is not

found in the Erica hispidula-Bmnia alopecuroides Shrub-

lands and is sparingly present in the Tetraria bromoides-

Berzelia galpinii Shrublands (Table 2). The Erica his-

pidula-Restio inconspicuus Shrublands described here

were sampled on the southern mid- to lower slopes of

the Bergfontein area and are characterized by presence of

Agapanthus africanus, Berzelia galpinii, Nevillea sp. nov.

(Restionaceae) (newly discovered in the Bergfontein area)

amongst other species (Table 2).

Penaea cneorum subsp. ovata, Leucadendron spissi-

folium and Drosera aliceae , all species which prefer moist

slopes, occur in the Erica hispidula-Brunia alopecuroides

Shrublands and the Erica hispidula-Restio inconspicuus

Shrublands.

1.2.1 Restio inconspicuus-Chondropetalum mucrona-

tum Shrublands

Differential species: Chondropetalum mucronatum,

Chironia jasminoides.

Dominant species: Blaeria coccinea, Platycaulos com-

pressus, Restio inconspicuus.

Structural formation: Closed Graminoid Shrubland.

Relationships: Chondropetalum-Restio Tussock Marsh

(Boucher 1978); ‘Slope-type’ Restioid Marsh (Taylor

1978); Erica mollis Fynbos Community (Glyphis et al.

1978); Restio-Hypolaena Subcommunity (Laidler et al.

1978); Restiad Herblands (in part) (Kruger 1979); Sneeu-

250

Bothalia 23,2 (1993)

TABLE 2.— A phytosociological table of the Cullumia aculeata var. aculeata Shrublands on the north slopes of Witelsberg (Bergfontein) and Phesantefontein, southern

Langeberg

Bothalia 23,2(1993)

251

TABLE 2, — A phytosociological table of the Cullumia aculeata var. aculeala Shrublands on the north slopes of Witelsberg (Bergfontein) and Phesantefontein , southern

kop Azonal Restioid Fynbos (Campbell 1985); Restio in-

conspicuus-Anthochortus crinalis Shrublands (McDonald

1993a); Erica hispidula-Anthochortus crinalis Shrublands

(McDonald 1993b).

Chondropetalum mucronatum is widely distributed

from the eastern Langeberg to Bainskloof and the Cape

Peninsula in marshy places (Linder 1985). In the Cape

Hangklip area Boucher (1978) records it as occurring on

mountain slopes at sites with impeded drainage irrespec-

tive of aspect or altitude. This is similarly true on the

Langeberg, however, at BWA and MNR (McDonald

1993a & b), stands of C. mucronatum are scattered and

simply form part of the vegetation mosaic on south-facing

wet slopes. At Bergfontein the distribution of C.

mucronatum is not so scattered and the species differen-

tiates a distinct community on localized seepages with a

'peaty coarse sand’ substratum as it does in the Cape

Hangklip mountains (Boucher 1978). Chironia jasmi-

noides is also characteristic of marshy places from the

southwestern Cape to the Riversdale District (Bond &

Goldblatt 1984) and at Bergfontein. is almost at the eastern

limit of its range.

At Bergfontein the Restio inconspicuus-Chondro-

petalum mucronatum Shrublands (Figure 6) occur within

an altitudinal range from 560 m to 850 m on moderate

slopes (14°-30°) with southeast- to southwest-facing as-

pects.

This community has an upper stratum dominated by

Chondropetalum mucronatum (up to 1.5 m) which

emerges above a low closed stratum (<0.5 m) where

grasses, restios and ericas are present in more or less equal

proportions. Blaeria coccinea (Ericaceae) is the dominant

shrub with Platycaulos compressus and Restio incon-

spicuus (Restionaceae) and Pentameris macrocalycina and

Pentaschistis malouinensis (Poaceae) comprising the

major part of the graminoid component. The Nevillea sp.

nov. recorded here is closely allied to the N. obtusissima

(continued)

recorded by Boucher (1978) in the Chondropetalum-

Restio Tussock Marsh of the Cape Hangklip area.

1.2.2 Restio inconspicuus-Selago serrata Shrublands

Differential species: Othonna quinquedentata, Selago

serrata, Syncarpha vestita.

Dominant species: Elegia juncea, Leucadendron spis-

sifolium, Pentameris macrocalycina, Pentaschistis malou-

inensis, Restio inconspicuus, Staberoha cernua.

Structural formation: Low Open to Closed Graminoid

Shrubland with sparse emergent shrubs.

Relationships: Tetraria thermalis-Hypodiscus aristatus

Community (Kruger 1974); Leptocarpus membranaceus

( Calopsis membranacea)-Hypodiscus aristatus Commu-

FIGURE 6. — The Restio inconspicuus-Chondropetalum mucronatum

Shrublands on southeast- to southwest-facing slopes at Bergfon-

tein.

252

Bothalia 23,2 ( 1993)

nity (McKenzie et al. 1977); Subcommunity C of the Pen-

aea-Erica Fynbos Community ( Glyphis et al. 1978); Low

Ericoid Open Heath or Open graminoid-heath (Kruger

1979); Erica viridescens-Hypodiscus aristatus Commu-

nity (Bond 1981); Nuweberg Mesic Ericaceous Fynbos

(Campbell 1985); Hvpodiscus aristatus-Berzelia interme-

dia Shrublands (McDonald 1993a).

This community (Figure 7) is found on east- to south-

west-facing rocky sites where surface rock was estimated

at 32% on average. Slope inclination ranges from 17°-37°

and the sandy Mispah Form soils (lithosols) are well

drained. In general this shrubland has a single low stratum

(<1 m) with Psoralea pinnata occasionally emergent to 2

m.

This shrubland community is poorly differentiated. It

has only three character species, Othonna quinquedentata,

Selago serrata and Syncarpha vestita which are poorly

represented. It lacks the moisture-loving species common

to communities 1.1 and 1.2.1 but has a strong graminoid

component, a characteristic which it shares with the Restio

inconspicuus-Chondropetalum mucronatum Shrubland.

Apart from the dominant species, Calopsis membvanacea

and Hvpodiscus aristatus (Restionaceae) are well repre-

sented, whereas Berzelia galpinii is not prominent. The

low cover-abundance of Berzelia galpinii is ascribed to

the shallow, rocky, well-drained nature of the soil.

1.2.3 Restio inconspicuus-Erica melanthera Shrublands

Differential species; none.

Dominant species: Berzelia galpinii, Elegia asperi-

flora, Erica cubica, Pentaschistis malouinensis, Staberoha

cemua, Tetraria flexuosa.

Structural formation: Low Closed Graminoid Shrub-

land.

FIGURE 7. — The Restio inconspicuus-Selago serrata Shrublands

found on rocky sites with east- to southwest-facing aspects above

Bergfontein.

FIGURE 8. — The Restio inconspicuus-Erica melanthera Shrublands on

grey shallow sandy lithosols at Bergfontein.

Relationships: as for 1.2.2 above but Berzelia interme-

dia-Erica melanthera Shrublands (McDonald 1993a) and

Leucadendron eucalyptifolium-Erica melanthera Shrub-

lands (McDonald 1993b).

This community (Figure 8) is found on a variety of

aspects from northeast through south to southwest. The

soils are grey sandy shallow lithosols, seldom exceeding

0.3 m and mostly <0.2 m deep. Amounts of exposed rock

range from very low to 98% with most sample sites hav-

ing 85% rock cover. Despite this, total vegetation cover

is high at 94%.

The absence of differential species in this community

is apparently related to the drainage regime. Species such

as Berzelia galpinii, Erica cubica and Elegia asperiflora

which have a strong preference for sites with impeded

drainage, although occurring in other communities, play

a dominant role here. Erica melanthera which favours

shallow sandy soils is also mainly found in these

shrublands, but this species is generally much less com-

mon than in the Hvpodiscus aristatus-Berzelia intermedia

Shrublands of BWA (McDonald 1993a).

The low stature of the community is ascribed to the

shallowness of the soil and high percentage rock cover.

It could be speculated, however, that some stands would

reach mid-high to tall stature in older vegetation.

1.3 Erica hispidula-Tetraria bromoides Shrublands

These shrublands include five distinct communities that

have T. bromoides as the common denominator. T.

bromoides (Cyperaceae) is a robust leafy sedge that is

known for its preference for heavier soils i.e. soils with

a high clay fraction, derived from shale or sandstone

(Boucher 1978; McDonald 1993a & b). Of note is the

occurrence of Penaea mucronata in these shrublands. This

species shows a distinctly different habitat preference

(lower, drier slopes) to Penaea cneorum subsp. ovata

Bothalia 23,2(1993)

253

which is found in communities of the Erica hispidula-

Restio inconspicuus Shrublands on wetter, higher slopes.

l. 3.1 Tetraria bromoides-Berzelia galpinii Shrublands

Differential species: none.

Dominant species: Erica hispidula, Leucadendron eit-

calyptifolium, Penaea mucronata, Tetraria flexuosa.

Structural formation: Low Closed Graminoid Shrub-

land with Mid-high Emergent Shrubs in some stands.

Relationships: Berzelia-Leucadendron Moist Tall Fyn-

bos (Boucher 1978); Mixed-sclerophyllous Scrub (Kruger

1979); Mangold Wet Proteoid Fynbos (Campbell 1985).

This community (Figure 9) is found on south and

southwest-facing slopes at altitudes ranging from 350-550

m. Soils are derived from sandstone of either the Peninsula

Formation or Nardouw Subgroup and are generally 0.5

m deep, with soils at some sites as shallow as 0. 1 m. The

average slope of sampled stands is 21.3° (8°-32°). Rock-

iness is generally low, with an average of 12%, but with

some sites having as much as 90% surface rock. Amount

of litter depended on the age of the stand, with the old

vegetation having an estimated 75% litter cover below the

shrub canopy.

Two of the stands sampled (releves 212 & 213) of the

community were located in vegetation estimated to be 16

years or possibly older. The remainder of the samples were

in six-year-old vegetation. The two mature stands gave a

clear indication of how the Tetraria bromoides-Berzelia

galpinii Shrublands would appear over a much wider area

when mature. In the mature state they would be classified

structurally as Tall Closed Proteoid Shrublands with a

Closed Ericoid Shrubland Understorey, in contrast with

the structural formation given above.

The Tetraria bromoides-Berzelia galpinii Shrublands

are transitional between the Erica hispidula-Restio in-

FIGURE 9. — The Tetraria bromoides-Berzelia galpinii Shrublands

found on southerly slopes with sandy soils at altitudes from

350-550 m.

conspicuus Shrublands on shallow sandy soils and the

Erica hispidula-Tetraria bromoides Shrublands on soils

with a high clay fraction derived either from shale or an

admixture of sandstone and shale. Since the Tetraria

bromoides-Berzelia galpinii Shrublands are ecotonal their

floristic composition is complex, displaying elements of

both the Erica hispidula Shrubland types given above

(Table 1).

1.3.2 Tetraria bromoides-Ischyrolepis hystrix Shrublands

The two communities found on the Cedarberg Forma-

tion shaleband, the Tetraria bromoides-Phylica rubra and

Tetraria bromoides-Knowltonia capensis Shrublands have

many species in common but also a number of differences

(discussed below). However, the absence of many species

from these communities which are otherwise generally

found in the Erica hispidula-Tetraria bromoides Shrub-

lands sets these shrublands apart. These distinctions are

ascribed to soil-related rather than climate-related factors.

1.3. 2.1 Ischyrolepis hystrix- Phylica rubra Shrublands

Differential species; Cyclopia sessiliflora, Indigofera

sarmentosa, Phylica rubra.

Dominant species: Ischyrolepis hystrix, Leucadendron

eucalyptifolium, Phylica rubra. Protea aurea, P neriifolia,

Tetraria bromoides.

Structural formation: Tall Proteoid Shrubland with

Mid-high. Mid-dense Shrub Understorey and Low Mid-

dense Graminoid Field Layer.

Relationships: Broad-sclerophyllous Scrub (Kruger

1979); Protea aurea-Pteridium aquilinum Community

(Bond 1981); Outeniqua Wet Proteoid Fynbos (Campbell

1985); Restio inconspicuus-Protea aurea Shrublands (Mc-

Donald 1993a); Cliff ortia serpyllifolia-Leucadendron eu-

calyptifolium Shrublands and Widdringtonia nodiflora-

Tetraria bromoides Shrublands (McDonald 1993b).

The Ischyrolepis hystrix-Phylica rubra Shrublands

(Figure 10) were sampled on the Cedarberg Formation

shaleband. east of Stinkhoutbos, behind Korinteberg. The

mean altitude of the sample sites (releves 301-304), which

were located relatively close to each other in mature pro-

teoid fynbos, is 824.5 m. Aspect ranged from southwest

to west on shallow (5°) to moderate (22°) slopes. The

yellow-brown sandy clay loam soils are classified as

Clovelly Form, reaching a mean depth of 0.45 m. Surface

rock cover is low, not exceeding 10% and the soils are

moderately to well drained. Litter cover is high (60-80%)

and the shrub canopy cover is closed (100%).

The age of the Ischyrolepis hystrix-Phylica rubra

Shrublands, estimated at 15 years, could be a factor in-

fluencing the difference in species composition between

this community and the closely allied but younger

Ischyrolepis hystrix-Phylica pinea Shrublands found at

Bergfontein (see below). Alternatively it may be postu-

lated that geographical separation of the two areas where

the shrublands were sampled may account for the differ-

ences between them (gamma diversity). These two

shrublands may also be interpreted as variants of a broader

shaleband community dominated by Cannomois virgata.

Erica grata, Ischyrolepis hystrix, Phylica pinea and Protea

254

Bothalia 23,2 (1993)

FIGURE 10. — The Ischyrolepis hystrix-Phylica rubra Shrublands

found on the Cedarberg Formation shaleband east of Stink-

houtbos. Note the tall Protea aurea shrubs behind the figure.

aurea, since P. aurea and Tetraria bromoides dominated

shaleband shrublands have been recorded in other parts

of the Langeberg (McDonald 1993a & b). As much as

the differential species separate the above two communi-

ties from each other, absence of numerous species, the

most notable of which are Gnidia galpinii, Lanaria lanata,

Penaea mucronata and Struthiola garciana , reinforce the

distinction between the two shrublands (Table 2).

1.3. 2. 2 Ischyrolepis hystrix-Phylica pinea Shrublands

Differential species: Carpacoce spermacocea, Erica

coccinea, Knowltonia capensis, Pelargonium cordifolium,

Serruria fasciflora.

Dominant species: Cannomois virgata, Ischyrolepis

hystrix, Leucadendron eucalyptifolium , Phylica pinea.

Structural formation: Tall Mid-dense to Closed

Restioid/Proteoid Shrubland with Sparse Mid-high Shrub

Understorey and a Low Mid-dense to Closed Graminoid

Shrubland field layer.

Relationships: as for 1.3.2. 1 above.

Apart from releve 227, samples representing the

Ischyrolepis hystrix-Phylica pinea Shrublands (Figure 1 1 )

were found at altitudes from 30CM-00 m on southeast- to

southwest-facing slopes of the Cedarberg Formation

shaleband, where it traverses the Bergfontein area. Releve

227 was sampled on a shale lens on a moderate north-

west-facing slope above Rooiwaterspruit, where the soil

is a 0.3 m deep, yellow-brown loamy Clovelly Form soil.

The soils at Bergfontein were not classified except that

they were noted to have a grey-brown orthic A-horizon.

At Rooiwaterspruit, the vegetation in releve 227 was

dominated by Leucadendron eucalyptifolium in the tall

(1. 5-3.0 m) mid-dense canopy. Non-ericoid shrubs and

graminoids combined to form a closed understorey, 1.5

m high. The community represented by releve 227 is

somewhat different to the Bergfontein Community since

many of the typical shaleband-associated species are ab-

sent (Table 2). However, it is similar enough to the

Ischyrolepis hystrix-Knowltonia capensis Shrublands and

different enough from the Ischyrolepis hystrix-Phylica

rubra Shrublands for it to be included in the former. The

community at Bergfontein was dominated by the tall ( 1 .2-

5.0 m) Cannomois virgata in the canopy, with a mid-high,

mid-dense stratum dominated by Phylica pinea and Pro-

tea aurea and a low, closed stratum dominated by res-

tioids.

The Ischyrolepis hystrix-Phylica pinea Shrubland is

the only community from which Erica hispidula is totally

absent. No explanation for this can be advanced since this

species occurs in similar communities in the MNR and

BWA (McDonald 1993a & b). A combination of wetness

and shale-derived soil may account for the absence of this

species.

1.3.3 Tetraria bromoides-Hypodiscus striatus Shrublands

Differential species: Anomalanthus scoparius, Diosma

tenella, Heteropogon contortus, Hypodiscus striatus , Peu-

cedanum ferulaceum, Polygala sp.

Dominant species: Anomalanthus scoparius, Erica

hispidula, Hypodiscus striatus, Lanaria lanata, Leucaden-

dron eucalyptifolium, L. salignum, Tetraria bromoides.

Structural formation: Low Mid-dense to Closed

Graminoid Shrubland with Mid-high to Tall, Sparse to

Mid-dense Proteoid Shrubland Overstorey.

Relationships: Mixed-sclerophyllous Scrub (Kruger

1979); Protea neriifolia-Leucadendron eucalyptifolium-

Erica triceps Community (Bond 1981); Robinson Mesic

Proteoid Fynbos (Campbell 1985); Hypodiscus aristatus-

Leucadendron eucalyptifolium Shrublands (McDonald

1993a); Leucadendron eucalyptifolium-Erica melanthera

Shrublands (McDonald 1993b).

J

FIGURE 1 1 . — The Ischyrolepis hystrix-Phylica pinea on the Cedarberg

Formation shaleband in the Bergfontein area. Note the tall restio,

Cannomois virgata, characteristic of this community.

Bothalia 23,2(1993)

255

These shrublands (Figure 12) were sampled on the

north and northwest slopes of Koksposberg and on the

west-facing mid-slopes of Witelsberg, overlooking

Witelsrivier (Bergfontein area). The community occurs at

altitudes from 300-550 m on sites with shallow ( 0.2 m)

well-drained sandy loam soils, where the slopes are mod-

erate to steep ( 16°-36°) and where rock cover varies from

almost nil to 90%.

Presence of Tetraria bromoides points to soils with a

higher fertility, however, soil-vegetation relationships of

this community are not clear and require further investi-

gation. The community is clearly defined and character-

ized by differential species (Table 1) which have a

preference for stony or rocky well-drained sites. All the

differential species are also species of low stature which

are found in the understorey of mature stands of the Tetra-

ria bromoides-Hypodiscus striatus Shrublands. Ano-

malanthus scoparius (Ericaceae) grows as a prostrate

dwarf shrub covering stony soil but not over rocks (E.G.H.

Oliver pers. comm.) and the erect Hypodiscus striatus

(Restionaceae) is found as tussocks in shallow pockets of

soil. Heteropogon contortus is a subtropical C4 grass

which Bond (1981 ) recorded as differential for his Protea

nitida Community (Waboomveld) on fertile soils. At

Bergfontein, H. contortus is faithful to the Tetraria bro-

moides-Hypodiscus aristatus Shrublands and not found at

all in the Erica hispidula-Protea nitida Shrublands de-

scribed below. Diosma tenella (Rutaceae) is a shrublet

which Williams (1982) described as ‘rather rare' but hav-

ing a wide tolerance for different soil types, from shales

to sandy gravels and silcrete.

From Table 1 it is seen that there are numerous species

which, although not differential or dominant, are strongly

represented in this community: Acmadenia trigona,

Leucadendron salignum, Lanaria lanata, Penaea mucro-

nata, Staberoha cemua, Tetraria flexuosa and Rhodocoma

fruticosa.

One or two strata were found in these shrublands at

the time of sampling. One stratum was found where the

shrubs were co-dominant with the graminoid component

up to a height of 1 m. Two strata occurred where the

shrubs, mainly proteoids such as Leucadendron eu-

calyptifolium and Protea neriifolia , exceeded 1 m, forming

a sparse to mid-dense overstorey. The immaturity of the

vegetation gave a false impression of the potential struc-

ture of the community, which is predicted to become a

tall closed proteoid shrubland with a low closed ericoid

or restioid shrubland understorey when mature (as in

releves 261 & 262).

1.3.4 Tetraria bromoides-Protea coronata Shrublands

Differential species: Alepidea capensis , Erica cf. zwart-

bergense, Protea coronata.

Dominant species: Erica hispidula, Lanaria lanata,

Leucadendron eucalyptifolium, Tetraria bromoides.

Structural formation: Mid-high Closed Proteoid Shrub-

land with Low Closed Graminoid Shrubland Understorey

or a Low Closed Graminoid Shrubland with Sparse emer-

gent Proteoid shrubs.

Relationships: Broad-sclerophyllous Scrub (Kruger

1979) ; Outeniqua Wet Proteoid Fynbos (Campbell 1985);

Wet Proteoid Fynbos (Rebelo et al. 1991).

This community (Figure 13) occurs on the lower south-

east-, southwest- and west-facing slopes of Koksposberg

in the Bergfontein area. At plot 221 the soil is derived

from Cedarberg Formation shale, whereas at the remainder

of the plots (214—216 & 247) the soils are apparently de-

rived from accumulated material at the footslopes. The

soils are well-drained shallow (0. 2-0.3 m) fine-textured

grey-brown to dark brown sandy loams, with negligible

surface rock cover.

The distribution of Protea coronata is strongly related

to edaphic factors with this species favouring heavy soils

with a high clay fraction (Rourke 1980). At Bergfontein,

Protea coronata occurs on sandy loam soils but, contrary

to Rourke’s description where Protea coronata is said to

form ‘densely massed stands’, the species occurs as scat-

tered individuals. Frequent fires may have excluded P.

coronata , a reseeding proteoid, at three of the five sample

sites. Alternatively these sites may be marginally more

mesic and P. coronata, which prefers wetter sites ( Rourke

1980) , may have been outcompeted by Protea neriifolia

and Leucadendron eucalyptifolium (Table 2). Even though

the Tetraria bromoides-Protea coronata Shrubland is

poorly characterized by the differential species Alepidea

capensis. Erica cf. zwartbergense and P. coronata, it is

substantiated by absence of numerous species found in

the Tetraria bromoides-Hypodiscus striatus Shrublands on

the one hand and the Tetraria bromoides-Protea nitida

Shrublands on the other. It also has many species in com-

mon with the latter community, in particular' the ubiquitous

‘waboomveld’ species Montinia caryophyllacea and Rhus

rosmarinifolia. Helichrysum cymosum is not a true differ-

ential species of the Tetraria bromoides-Protea coronata

Community as stated by Rebelo et al. (1991) since this

species is also well represented in the more mesic Erica

FIGURE 12. — The Tetraria bromoides-Hypodiscus striatus Shrublands

on the west-facing slopes of Koksposberg with the south slopes

of Witelsberg visible behind.

256

Bothalia 23,2 (1993)

FIGURE 1 3. — The Tetraria bromoides-Protea coronata Shrublands on

the lower slopes of Koksposberg, in the Bergfontein area.

hispidula-Protea nitida Shrublands which were not sam-

pled or described by these authors.

Two strata were found in these shrublands. Graminoids

are marginally more abundant than woody shrubs in the

low stratum, which was the dominant stratum when sam-

pled. In releve 247, the low stratum had been disturbed

by grazing animals. The upper stratum was dominated by

mid-high proteoid shrubs, Leucadendron eucalyptifolium

and Protea neriifolia. Following Campbell's (1985) sys-

tem, the Tetraria bromoides-Protea coronata Shrublands

would be placed in the Mesic Proteoid Subseries, how-

ever, based on lloristic composition and relationships, the

community is placed unequivocally in Wet Proteoid

Fynbos.

Aloe gracilis Haw. var. decumbens Reynolds was

found in this community and the Erica hispidula-Protea

nitida Shrublands. Reynolds (1950) gives the distribution

of this endemic variety of Aloe gracilis as from near

Garcia’s Pass westwards to Kleinberg at altitudes 272-364

m. Records of A. gracilis var. decumbens at Bergfontein

represents a range extension for this species on the

Langeberg, 30 km east of Garcia's Pass. Future searches

could reveal that it may occur further east, perhaps beyond

the Gouritz River.

1 .4 Erica hispidula-Protea nitida Shrublands

Differential species: Anthospermum aethiopicum, Ca-

lopsis filiformis, Clutia laxa, Eragrostis capensis, Erica

glandulosa, E. peltata, Pelargonium candicans, Protas-

paragus rubicundus. Protea nitida, Themeda triandra.

Dominant species: Lanaria lanata, Leucadendron sa-

lignum, Protea nitida, Restio triticeus, Themeda triandra.

Structural formation: Closed Graminoid Shrubland

with Mid-high Sparse to Open Proteoid Overstorey.

Relationships: ‘Waboomveld’ (Taylor 1963, 1978; Tay-

lor & Van der Meulen 1981); Protect arborea (P. nitida )

Pseudo-Savannah (Taylor 1969); Protea arborea-Rhus

angustifolia Community (Werger et at. 1972); Protea-

Tetraria Dry Short Fynbos (Boucher 1978); Restio gau-

dichaudianus ( Ischyrolepis gauclichaudiana )-Lobostemon

glaucophyllus Community (McKenzie etal. 1977); Protea

arborea (P nitida) Tall Broad-sclerophyllous Shrubland

or Open Shrubland with Heathland (Kruger 1979); Protea

nitida Community (Waboomveld) [Outeniqua Mountains]

& Protea nitida-Protea repens Community [Swartberg]

(Bond 1981); Protea nitida Woodland ('Waboomveld')

(Taylor 1984); Rooiberg Talus Asteraceous Fynbos

(Campbell 1985); Ischyrolepis gaudichaudiana-Myrsine

africana High Closed Shrubland (McDonald 1983, 1988).

Protea nitida is most often found on colluvial soils on

debris or talus slopes but may also be found on fine-tex-

tured soils on lower mountain slopes (Taylor 1978; Kruger

1979; R.M. Cowling pers. comm.). These soils generally

have a higher nutrient status than that of leached sandstone

soils, due to colluvial mixing of soil derived from various

parent materials such as sandstone, granite or shale. Fyn-

bos with P nitida as a prominent, if not dominant shrub

or tree, is found throughout the Fynbos Biome and ac-

cording to Taylor (1978) and Kruger (1979) characterizes

a distinct formation, 'waboomveld'. The term 'waboom-

veld’ encompasses more than one community where P.

nitida is present, but throughout the distribution range of

this species from the northwestern to the southern Cape

there is a remarkably constant group of associated species

including Anthospermum aethiopicum, Ischyrolepis gaudi-

chaudiana, Montinia caryophyllacea, Rhus rosmarinifolia

and Themeda triandra. Campbell (1985) explicitly stated

that P. nitida is diagnostic and must be present for Moun-

tain Fynbos to be classified as Talus Asteraceous Fynbos

but he also indicated that P. nitida occurs in other com-

munities as well.

At Bergfontein these shrublands (Figure 14) have ei-

ther two or three strata. P. nitida forms a sparse canopy

up to 2 m high. Where present the second stratum from

0.5-1. 2 m is dominated by Leucadendron salignum and

the low stratum 0.0-0.5 m by Themeda triandra. The

community was restricted to well-drained east-facing

slopes above Witelsrivier, directly opposite the west-fac-

ing slopes of Koksposberg where the closely related Tetra-

ria bromoides-Protea coronata Shrublands occur. The

altitude at which these shrublands are found ranges from

300-350 m with the slopes averaging 19°. Annual precip-

itation is estimated at 600-700 mm (Dent et al. 1987).

The soils with pH 4.9 (n = 3) are a shallow (0.2 m) well-

drained mix of sandstone debris from the Peninsula For-

mation sandstone and Cedarberg Formation shale. They

are classified as Oakleaf Form soils. Rock cover ranges

from 10-40%.

Apparently the Erica hispidula-Protea nitida Shrub-

lands at Bergfontein are not burnt often, so the composi-

tion of the understorey is not attributed to regular fires or

other disturbances. The grassy understorey is rather attrib-

uted to soil factors since the grasses in the community are

C4 grasses which show preference for more fertile soils

(Bond 1981; Cowling 1983a & b; Linder 1989). This

community qualifies for inclusion in Campbell’s (1985)

Rooiberg Talus Asteraceous Fynbos on the basis of pres-

ence of Protect nitida and 20 % grass cover. However,

Bothalia 23,2 (1993)

257

the lack of asteraceous ‘elytropappoid' species and the

high grass cover (70%) indicate closer affinities to Grassy

Fynbos ( sensu Cowling & Holmes 1992).

A stand of tall P. nitida with an understorey composed

almost entirely of grasses was located on deep sandy soils

on the plateau below the upper north slopes of Witelsberg.

This community was not sampled but was observed to be

quite different in structure and composition from the Erica

hispidula-Protea nitida Shrublands. Further investigation,

not possible in this study due to the fire, would be required

to clarify the relationships.

2. Erica versicolor-Agathosma ovata Shrublands

Differential species: Adromischus triflorus, Agathosma

ovata, Centella virgata var. congesta, Crassula atropur-

purea var. atropurpurea, C. muscosa, C. rupestris, Cras-

sula sp. (unidentified). Erica tenuis, Ficinia elongata,

Lampranthus deltoides, Lobelia pubescens var. pubescens,

Senecio aizoides.

Dominant species: Agathosma ovata, Ehrharta ramosa,

Erica melanthera, E. versicolor, Phylica puipurea var.

floccosa.

Structural formation: Sparse to Open Low Graminoid

Shrtibland.

Relationships: unclear.

The estimated annual precipitation for Koksposberg is

500-600 mm (Dent et al. 1977). However, the north-fac-

ing slopes are in a rain shadow as well as being equatori-

ally oriented. They are therefore hot and dry and probably

receive much less rainfall than the above estimate. A sim-

ilar situation also exists on the upper north-facing slopes

of Witelsberg. These dry. rocky slopes (Figure 15) with

high incoming radiation host the Erica versicolor-

Agathosma ovata Shrublands. The soils are lithosols

(Mispah Form), are shallow and acid (pH 3.4 in 1 mol/l

CaCb) and consist merely of accumulated sand and or-

ganic material in shallow depressions amongst the rocks.

FIGURE 14. — The Erica hispidula-Protea nitida Shrublands on the

east-facing slopes, west of Bergfontein, above Witelsrivier.

FIGURE 15. — The Erica versicolor-Agathosma ovata Shrublands

found on the rocky north slopes of Koksposberg and Witelsberg.

The parent rock of Koksposberg is Peninsula Formation

sandstone and rock cover is 90% in the four releves (208—

211) representing this community. On Koksposberg the

community is found at altitudes from 550-606 m on

slopes varying from almost Hat (7°) on the ridge to mod-

erately steep (18°) on the north face. On Witelsberg, con-

sisting of Nardouw Subgroup sandstone, the community

is found from 788 m to 1 090 m.

Two strata are present in the Erica versicolor-Aga-

thosma ovata Shrublands. The upper stratum is from 0.5-

1.2 m and is dominated by the shrubby Erica versicolor.

The lower stratum where shrubs and graminoids are co-

dominant is from 0-0.5 m. Dominant species in this stra-

tum are Ehrharta ramosa and Erica melanthera. However,

the prevalence of succulent species in the genera Ad-

romischus, Aloe, Crassula, Lampranthus and Senecio

which prefer dry, rocky sites gives this community its

character and sets it apart from all other communities de-

scribed for the Bergfontein area (releves not included in

Table 1 or Table 2). A. ovata also favours a dry, rocky

habitat as does Oldenburgia paradoxa which grows in

rock crevices forming large, compact, rounded masses of

short shoots.

A detrended correspondence analysis (DCA) (Hill

1979b) of the Bergfontein data showed a clear distinction

between the Erica versicolor-Agathosma ovata Shrubland

and the other shrublands. DCA Axis I represented a mois-

ture gradient and the Erica versicolor-Agathosma ovata

Shrublands are towards the drier end of the gradient. DCA

Axis II apparently represented a soil depth/rockiness gra-

dient, showing that the community inhabits the rockiest

sites with shallowest soils in the study area.

3. Cullumia aculeata var. aculeata Shrublands

The Cullumia aculeata Shrublands include the non-

Erica hispidula shrublands represented by Table 2. All

these shrublands are on the northern slopes of the

258

Bothalia 23,2 (1993)

Langeberg range adjacent to the Little Karoo. Most of the

samples were taken in the Phesantefontein area; the ex-

ceptions are releves 278-281 which were recorded on the

north slopes of Witelsberg before the May 1 99 1 fire.

3.1 Cullwnia aculeata-Leucadendron eucalyptifolium

Shrublands

This community is found on the mid- to upper slopes

immediately east of Aasvoelkrans. They are mesic pro-

teoid shrublands and are separated into three distinct com-

munities with Leucadendron eucalyptifolium as the

common factor.

3.1.1 Leucadendron eucalyptifolium-Protea neriifolia

Shrublands

Two communities are found in the Leucadendron eu-

calyptifolium-Protea neriifolia Shrublands. P neriifolia

links these shrublands, setting them apart from the

Leucadendron eucalyptifoliutn-Elegia filacea Shrublands

where P. neriifolia is absent. These shrublands have affi-

nity to the Tetraria bromoides-Hypodiscus striatus Shrub-

lands found on the southern slopes at Bergfontein with

Anomalanthus scoparius and Hypodiscus striatus , differ-

ential species of the latter community, being found here

as well.

3. 1 . 1 . 1 Protea neriifolia-Merxmuellera decora Shrub-

lands

Differential species: Ficinia trichodes, Ischyrolepis sp.,

Merxmuellera decora.

Dominant species: Hypodiscus argenteus, Lanaria la-

nata, Merxmuellera decora.

Structural formation: Mid-high Closed Graminoid

Shrubland.

Relationships: Proteoid Zone Fynbos (Taylor 1978);

Broad sclerophyllous- Scrub or Open Scrub (Kruger

1979); Sanddrift Mesic Proteoid Fynbos (Campbell 1985);

affinities with Tetraria bromoides-Hypodiscus striatus

Shrublands and Tetraria bromoides-Protea coronata

Shrublands described above.

This community (Figure 16) is represented by five

releves (286-290) at altitudes from 630-850 m on the

northeast- to northwest-facing slopes east of Aasvoelkrans.

These slopes are moderate, with shallow ( 0.2 m), yel-

low-brown sandy loam soil, classified as Glenrosa Form.

The parent material of the soil is taken to be Nardouw

Subgroup sandstone, but the heavy nature of the soil

(releves 286-289) suggests a high clay fraction possibly

derived from a clay lens or local exposure of the

Cedarberg Formation shale. This requires verification but

the vegetation reflects a soil with a relatively higher nu-

trient status (see below). Surface rock cover is low to

moderate (3^40%) except in releve 290 which is different

from the other samples. It was located on a rocky sand-

stone outcrop with 95% rock cover. The soil is of the

Mispah Form, but the Holistic composition of the releve

places it in the Protea neriifolia-Merxmuellera decora

Shrublands.

At the time of sampling, the vegetation was seven years

old. Depending on the site, the community had one or

two strata,' with height not exceeding 1 .2 m. The shrubland

was thus classified as above, a Mid-high Closed Gra-

minoid Shrubland, but it is predicted that with time the

community would mature to a Tall Closed Proteoid Shrub-

land with a Graminoid Understorey.

The Protea neriifolia-Merxmuellera decora Shrub-

lands are poorly differentiated from the closely allied Pro-

tea neriifolia-Erica articularis Shrublands based on the

differential species of the first community. Merxmuellera

decora presents problems with identification in mature

vegetation since it usually flowers only in the immediate

post-fire phase; Ischyrolepis sp. (Table 2) was not identi-

fied and Ficinia trichodes was poorly represented. Ab-

sence of the differential species of the latter community

from the Protea neriifolia-Merxmuellera decora Shrub-

lands provides a sounder basis for the separation of the

two communities.

3. 1 .1.2 Protea neriifolia-Erica articularis Shrublands

Differential species: Aristea racemosa, Cassytha cilio-

lata, Coelidium cymbifolium, Erica articularis, Euryops

abrotanifolius, Ischyrolepis sieberi, Zygophyllum fulvum.

Dominant species: Ceratocaryum decipiens, Elegia

galpinii, Leucadendron salignum, Staberoha cernua.

Structural formation: Low Closed Herbland or Low

Mid-dense to Closed Graminoid Shrubland.

Relationships: Proteoid Zone Fynbos (Taylor 1978);

Broad-sclerophyllous Scrub or Open Scrub (Kruger

1979); Elegia galpinii-Metalasia pulcherrima Commu-

nity (Outeniqua Mountains) and Protect repens-Protea

lorifolia-Hypodiscus striatus Community (Swartberg)

(Bond 1981); Mesic Mountain Fynbos (Moll et al. 1984);

Thomas Dry Proteoid Fynbos (Campbell 1985). Leu-

cadendron eucalyptifolium-Staberoha cernua Shrublands

(McDonald 1993a).

FIGURE 1 6. — The Protect neriifolia-Merxmuellera decora Shrublands

found on the northeast- to northwest-facing slopes east of

Aasvoelkrans, above Phesantefontein.

Bothalia 23,2(1993)

259

This community (Figure 17) occurs on the same north-

facing slopes as the Protea neriifolia-Merxmuellera

decora Shrublands but at lower altitudes (590-630 m).

The shallow (0.2— 0.3 m), light grey-brown soils were clas-

sified as Glenrosa Form (releves 291-293) and Mispah

Form (releve 294). All the releves were situated on shal-

low slopes (9°-14°), between rock outcrops, where rock

cover was very low except in releve 294 where it was

estimated at 20%.

The differential species of the Protea neriifolia-Erica

articularis Shrublands are not constant throughout the

community, nor do they have high cover-abundance. Of

these species, Erica articularis and Ischyrolepis sieberi

have highest cover-abundance whereas the remaining spe-

cies are rare (Table 2). The very rare legume, Coelidium

cymbifolium , previously known from only two collections,

Muir’s type collection and a collection of Middlemost

(Granby 1980), was found in releves 293 and 294 and

appears to be narrowly endemic to this community. Protea

aspera , a prostrate rhizomatous shrublet. was also re-

corded in this community and the Leucadendron eu-

calyptifolium-Elegia filacea Shrublands (see below and

Table 2). This species is best known from Onrust to

Bredasdorp but has also been recorded from an isolated

population in the Langkloof, east of Garcia's Pass (Rourke

1980). Records of Protea aspera at Phesantefontein rep-

resent an extension of the range of the Langkloof popu-

lation 5 km west of Garcia's Pass.

The Protea neriifolia-Erica articularis Shrublands are

dominated by restios such as Ceratocaryum decipiens,

Elegia galpinii, Mastersiellci purpurea, Restio filifonnis,

Thamnochortus ellipticus and Staberoha cernua, all spe-

cies preferring ntesic to dry habitats. T. ellipticus was orig-

inally collected by Muir in the Phesantefontein area and

is only known from this, the type locality (Linder 1985).

3.1.2 Leucadendron eucalyptifolium-Elegia filacea

Shrublands

Differential species: Calopsis filifonnis, Elegia filacea,

Hypodiscus laevigatas.

Dominant species: Elegia filacea, Protea repens.

Structural formation: Low Closed Graminoid (Res-

tioid/Cyperoid) Shrubland with Tall Open Proteoid Over-

storey.

Relationships: as for 3. 1.1. 2 above and Leucadendron

eucalyptifolium-Protea lorifolia Shrublands (McDonald

1993a).

Two releves, 295 and 296, represent this community

(Figure 18). It is found below the Leucadendron euca-

lyptifolium-Erica articularis Shrublands, and is floristi-

cally transitional between them and the C. aculeata-

Protea lorifolia Shrublands. Both releves were at 530 m

on slopes with a shallow gradient. The soil which is 0.25-

0.3 m deep is a light grey sandy loam with quartz pebbles

and gravel on the surface. Rock cover is nil and the soils

are classified as Glenrosa Form.

Elegia filacea is most often found on accumulated sand

and the sandy nature of the soil accounts for its presence

here. In this community it dominates the understorey in

FIGURE 17. — The Protea neriifolia-Erica articularis Shrublands on

the lower north-facing slopes above Phesantefontein.

which grasses are absent and sedges few, mainly of the

genus Ficinia. Low shrubs play a subordinate role in the

understorey composition, but the presence of Protea as-

pera., which also favours sandy soil, should be noted. The

upper (tall) stratum is composed of proteoid shrubs with

Protect repens dominant.

The rare, endemic Erica rhodantha Guth. & Bol. was

found in this community. This species is apparently re-

stricted to the fynbos shrublands of the lower north slopes

of the Langeberg in the region of Garcia's Pass. Few col-

lections of this species have been made (E.G.H. Oliver

pers. comm.).

The Leucadendron eucalyptifolium-Elegia filacea

Shrublands are the same as the Leucadendron euca-

lyptifolium-Protea lorifolia Shrublands described from

BWA (McDonald 1993a), but for the purposes of this

paper are treated as slightly different. The difference is

based on the presence of Calopsis filifonnis, Hypodiscus

laevigatas. Protea aspera and Thamnochortus ellipticus

at Phesantefontein and not at BWA.

3.2 Cullumia aculeata-Protea lorifolia Shrublands

This community, found at the driest extreme of the

south-north climatic gradient of the Langeberg, is div ided

into two: the Protea lorifolia-Ficinia laciniata Shrublands

(which fall within the Leucadendron eucalyptifolium-Pro-

tea lorifolia Shrublands of the BWA) and the Protect

lorifolia-Leucospermum calligerum Shrublands (which

are equivalent to the Passerina obtusifolia-Leucospermum

calligerum Shrublands at BWA). The Cannomois pan’if-

lora-Passerina obtusifolia Shrublands found at BWA

(McDonald 1993a) were not identified as a separate com-

munity at Phesantefontein but as part of the Passerina

obtusifolia-Leucospennum calligerum Shrublands.

260

Bothalia 23,2 (1993)

FIGURE 18. — The Leucadendron eucalyptifolium-Elegia filacea

Shrublands.

3.2.1 Protect lorifolia-Ficinia laciniata Shrublands

Differential species: Cannomois parviflora, Dodonaea

angustifolia , Ficinia laciniata , Gnidia francisci, Linum

gracile, Othonna sp., Polygala refracta, Siphocodon cf.

spartioides, Syncarpha milleflora, iWesium subnudum.

Dominant species: Leucadendron salignum, Penta-

schistis colorata, Hypodiscus aristatus, Tetraria ustulata.

Structural formation: Low Mid-dense Sedgeland or

Graminoid Shrubland with Mid-high Sparse Proteoid

Shrubs in some stands.

Relationships: Proteoid Zone Fynbos (Taylor 1978);

Broad-sclerophyllous Scrub or Open Scrub (Kruger

1979); Elegia galpinii-Metalasia pulcherrima Commu-

nity (Outeniqua Mountains) and Protea repens-Protea

lorifolia-Hypodiscus striatus Community (Swartberg)

(Bond 1981); Mesic Mountain Fynbos (Moll et al. 1984);

Thomas Dry Proteoid Fynbos (Campbell 1985); Leu-

cadendron eucalyptifolium-Protea lorifolia Shrublands

(McDonald 1993a).

This community was not found or sampled at Phe-

santefontein. The four releves (278-281) representing the

Protect lorifolia-Ficinia laciniata Shrublands were situ-

ated on the moderately steep, rocky, north-facing slopes

of Witelsberg on the Bergfontein transect at altitudes from

545-730 m. The soils are shallow (0.1-0.15 m) sandy

Mispah Form lithosols and surface exposure of rock was

estimated to be 91% on average.

This community is well differentiated from the other

shrublands represented in Table 2. This could be ascribed

either to geographical separation of these releves from the

Phesantefontein area or to the habitat where this shrubland

occurs not being present or not sampled at Phesantefon-

tein. Topographically, in rockiness and in climatic regime,

the north-facing slopes of Witelsberg closely resemble the

north-facing slopes of Deception Ridge in BWA (McDon-

ald 1993a). At Phesantefontein, however, moderately

steep, rocky, well-drained slopes of the same type are ab-

sent. The "habitat explanation’ for the absence of this com-

munity at Phesantefontein therefore seems the most

plausible.

Leucadendron salignum , Hypodiscus aristatus, Pen-

taschistis colorata and Tetraria ustulata dominate the

community at this early stage of its development (6 years

old), forming a low, mid-dense graminoid shrubland. As

these shrublands mature it is predicted that the proteoid

shrubs, Protea lorifolia and Protea repens will form a

mid-high to tall, open to mid-dense overstorey.

3.2.2 Protea lorifolia-Leucospermum calligerunt

Shrublands

Differential species: Adrontischus triflorus, Anthosper-

mum spathulatum, Calopsis rigida, C. marlothii, Cliffortia

pitlchella, Clutia laxa, Cytnbopogon marginatus, Hennan-

nia odorata, Heterolepis sp. nov., Lightfootia rigida,

Machairophyllum cookii, Metalasia massonii, Montinia

caryophyllacea, Passerina obtusifolia, Pentaschistis ma-

louinensis, Phylica purpurea, Thcunnochortus karooica.

Dominant species: Elegia galpinii, Leucosperntum

calligerunt, Thoracospernta galpinii, Willdenowia bolusii.

Structural formation: Low to Mid-high Mid-dense

Graminoid Shrubland with Sparse emergent Proteoid

Shrubs in some stands.

Relationships: Arid Fynbos (Taylor 1978; Kruger

1979); Phylica axillaris-Felicia filifolia Community (Out-

eniqua Mountains) and Passerina obtusifolia-Felicici fili-

folia-Pentaschistis eriostoma Community (Swartberg)

(Bond 1981); Dry Mountain Fynbos (Moll et al. 1984);

Sebrafontein Dry Asteraceous Fynbos (Campbell 1985);

Dry Proteoid Fynbos (Rebelo et al. 1991); Passerina ob-

tusifolia-Leucospermum calligerunt Shrublands (McDon-

ald 1993a); Leucadendron eucalyptifolium-Hypodiscus

argenteus Shrublands in part (McDonald 1993b).

This community (Figure 19) is found on the low, arid

northern foothills of Phesantefontein at altitudes from

450-520 m. The slope varies from almost level (5°) to

14° and the soils are mostly shallow ( <0. 1 m), well-

drained, rocky (75-98%) Mispah Form soils. The excep-

tion is the Glenrosa Form soil of releve 285 which is 0.25

m deep with an estimated surface rock cover of only 3%!

Releve 283 was situated on a rocky outcrop and was the

only site where Passerina obtusifolia was recorded. The

silcrete-capped mesas at Phesantefontein present a similar

substratum and habitat to that found in releve 283 and

therefore, although these mesas were not sampled, it is

predicted that they will support shrublands of the Protea

lorifolia-Leucospermum calligerunt type (Phesantefon-

tein) or the Cannomois parviflora-Passerina obtusifolia

type (Witbooisrivier; McDonald 1993a).

These shrublands are well defined with numerous dif-

ferential species. Heterolepis sp. nov. (as yet undescribed),

the succulent Machairophyllum cookii (Mesembryan-

themaceae) and Thamnochortus karooica are endemic to

this community. Species with single occurrences have

been retained in Table 2 specifically for this community,

for the sake of completeness. They are, however, not truly

differential.

Bothalia 23,2(1993)

261

Leucospennum saxatile which is endemic to the Arid

Fynbos east of Garcia’s Pass (Rourke 1972) was noted in

this community on the northern footslopes of Witelsberg

prior to the May 1991 fire. Serruria balanocephala

Rourke ined. found in this community at Witbooisrivier

(McDonald 1993a) has not been found at Phesantefontein

nor further east. Leucospennum embescens recorded from

Springfontein immediately west and at Garcia’s Pass di-

rectly east of Phesantefontein was also not encountered at

the latter locality and is apparently also not found much

further east than Garcia’s Pass.

Owing to the extensive fires on the Langeberg east of

Garcia’s Pass in the recent past, much of the vegetation

on the lower northern slopes of the range is immature and

thus not suitable for sampling. However, it can be confi-

dently stated that the community named the Passerina

obtusifolia-Leucospennum calligerum Shrublands (Mc-

Donald 1993a) and the Protect lorifolia-Leucospermum

calligerum Shrublands (this paper) extend along the base

of the Langeberg from Barrydale to the Gouritz River,

fringing the Little Karoo.

DISCUSSION AND CONCLUSIONS

The hierarchical classification and description of the

fynbos communities of the Bergfontein and Rooi-

waterspruit-Phesantefontein areas satisfies the primary ob-

jective of the study. It also provides a framework which,

together with differential species for each community, is

a starting point for describing the communities over a

wider area on the southeastern Langeberg. Further sam-

pling in other areas would serve to test this classification

and would most likely identify additional communities.

From the perspective of management of the fynbos of

the Langeberg mountain catchments, this classification is

perhaps too detailed and, as may be argued, too floristi-

cally orientated. However, it is important to be able to

FIGURE 19. — The Protea lorifolia-Leucospermum calligerum Shrub-

lands found on the flow, arid foothills of the Phesantefontein

area.

classify plant communities at all stages of the succession.

This has raised much debate (Cowling & Holmes 1992)

and it has been generally concluded that when applying

structural classifications it is only possible to satisfactorily

classify mature fynbos plant communities (Campbell

1985). Finding stands of mature fynbos (> 10 years old)

is becoming increasingly difficult (Campbell 1985; this

study) so applying structural classifications becomes

equally difficult. Despite the complexity of the vegetation

in this study, the successful hierarchical classification of

the mainly immature fynbos, and the determination of

character species for communities highlights a major ad-

vantage in using the floristic classification approach. If

suitably interpreted, by using the character species for dif-

ferent communities, the complex taxonomy of the fynbos

need not be the reputed daunting obstacle for managers

of fynbos vegetation (Bond 1981).

In general, the communities described for the Bergfont-

ein and Rooiwaterspruit-Phesantefontein areas are similar

to those described by McDonald (1993a & b) for the

Boosmansbos Wilderness Area and the Marloth Nature

Reserve. Degree of difference or conversely, similarity of

the fynbos communities of the southern Langeberg, which

is beyond the scope of this paper, can now be assessed.

A proposed synthesis is now possible which is aimed at

being generally applicable in the Langeberg fynbos

shrublands, and hopefully beneficial to the conservation

and management of these shrublands.

Apart from the purely descriptive reference to environ-

mental conditions pertaining to each community, little in-

dication is given of environmental variables as the causal

agents of species distribution and species composition of

communities. This forms a separate aspect to the study of

the Langeberg vegetation and will be reported separately

(McDonald unpublished data).

The Erica versicolor-Agathosma ovata Shrubland

found on Koksposberg does not fit well within the hier-

archy of communities. This was evident in both the phy-

tosociological tables and the ordination of the data. Many

of the fynbos species found generally on the south slopes

of the Bergfontein area are found in this community but

the succulent differential species indicate an azonal xeric

community. A clear definition of this community and its

relationships to other fynbos vegetation will only be pos-

sible given additional samples. It is predicted to occur on

the sandstone outliers of the Langeberg east of Garcia's

Pass as described and possibly on the Aasvogelberg near

Albertinia.

Garcia’s Pass has historically given ready access to the

Muiskraal area at its northern entrance. It has thus been

an area much visited by botanists since the time of

Thunberg and Masson in the late 1700’s. Many collections

of Langeberg north slope endemic species have Muiskraal

as the type locality, notably many of Muir's collections.

Many of these species are considered rare. Some of these

species such as Coelidium cymbifolium and Erica rho-

dantha have been found in this survey at Phesantefontein

which lies just west of Muiskraal. The intensity of sam-

pling in this study has, however, been too limited to cover

the area adequately. It is proposed therefore that the areas

on the north slopes of the Langeberg west and east of

262

Bothalia 23,2(1993)

Garcia’s Pass require thorough botanical investigation. If

these areas have high numbers of endemic plant species

as is indicated by this survey, these north slopes of the

Langeberg deserve a higher conservation status than is

currently the case.

ACKNOWLEDGEMENTS

The hospitality of the Oosthuizen family of the Farm

Bergfontein is gratefully acknowledged. Mr C. Martens,

Forester at Grootvadersbosch, gave invaluable support; he

and Messrs C. Ruiters and J. Solomon, also of Groot-

vadersbosch, who assisted with strenuous fieldwork, are

thanked sincerely. The advice and encouragement of Prof.

R.M. Cowling and Dr C. Boucher at times when it was

most needed is much appreciated. My wife and family

and my colleagues at the Stellenbosch Herbarium, Na-

tional Botanical Institute, are thanked for their support

during this project. Two anonymous referees are also

thanked for their suggested improvements to the manu-

script.

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Errata in Bothalia 23,1: 157

The vegetation of the southern Langeberg, Cape Province. 2. The plant communities of the

Marloth Nature Reserve, by D.J. McDonald.

Replace 3rd paragraph, left column with:

The climate of the MNR is typical of the southern

Langeberg since it falls within the transition zone be-

tween winter and year-round rainfall areas. The mean

annual rainfall for the peaks is estimated to be >1 400

mm (Dent et al. 1987). However, the climate on the

south side of the range is distinctly different from that

on the north side. The south slopes of the southern

Langeberg experience the highest rainfall in late sum-

mer to autumn (March & April) and late winter to

spring (August, October & November). The driest pe-

riods are early summer (December) and early winter

(June), Figure 5A. A more marked bimodality is evi-

dent in the rainfall pattern on the north slopes which

are in a summer rainshadow. Here peaks in rainfall

occur in autumn (April & May) and late winter (Au-

gust). The driest period extends from spring to late

summer (October-March), Figure 5B. The mean sum-

mer maximum and mean winter minimum temperatures

for Swellendam are 29.4°C (January) and 6.6°C (July)

respectively. On the opposite side of the mountain at

Weltevrede the equivalent temperatures are 30.0°C

(February) and 2.9°C (July).

.

Bothalia 23,2: 265-269 ( 1993)

The efficient use of small plots in a fynbos phytosociological study in the

northern Cederberg: a quick way to collect plant-environmental data

P.J. MUSTART*, E.J. MOLL* and H.C. TAYLOR**

Keywords: Cederberg, phytosociological survey, small plot sizes

ABSTRACT

A phytosociological survey of selected plant communities in the northern Cederberg was made using small (4-16 m2) plot

sizes. A satisfactory phytosociological table was obtained, and plant-environmental relationships were inferred from it. The use

of small plot sizes enabled ecological information about plant communities to be quickly, easily and efficiently obtained. This

method could be of considerable use for establishing and monitoring vegetation patterns.

UITTREKSEL

’n Fitososiologiese opname van geselekteerde plantgemeenskappe in die noordelike Cederberg is gemaak, en klein (4—16

nr) perseelgroottes is gebruik. 'n Bevredigende fitososiologiese tabel is verkry, en plant-omgewingsverhoudings is daaruit

afgelei. Die gebruik van klein perseelgroottes het daartoe gelei dat ekologiese inligting oor plantgemeenskappe vinnig, maklik

en doeltreffend verkry kon word. Hierdie metode sou heel nuttig vir die bepaling en monitering van plantegroeipatrone gebruik

kon word.

INTRODUCTION

The Zurich-Montpellier, or Braun-Blanquet. approach

has been widely used to survey and classify vegetation

(Werger 1974). The aim of such studies is to describe the

communities of the chosen area and to correlate them with

environmental factors. In South Africa these vegetation

studies have been carried out at scales ranging from re-

gional, comprising several thousands of hectares ( Boucher

1978; Cowling 1984; Taylor 1984), to smaller areas of a

few hundred hectares and less (Werger et al. 1972; Cowl-

ing et al. 1976; Campbell & Moll 1977; McKenzie et al.

1977; Glyphis et al. 1978; Laidler et al. 1978; Campbell

et al. 1980; Van Wilgen & Kruger 1985). In these studies

plot size has ranged from 4 to 200 m": choice of plot size

has largely depended on floristic richness and structure of

the communities sampled, and usually ignores the small-

scale pattern. Relatively small plots (e.g. 25 m~) have been

suggested as suitable for reflecting less habitat hetero-

geneity than larger plot sizes, in turn leading to the un-

masking of subtle variations within the observed com-

munities (Van Wilgen & Kruger 1985).

Large plots possibly covering heterogeneous habitats

and hence reflecting transitional communities, are time-

consuming for data collection. Small plots obviate both

these problems. The aim of this study was to perform a

phytosociological survey in the northern Cederberg using

plots of small size (4-16 m~) in order to see if a satisfac-

tory Braun-Blanquet table could be obtained, and if plant-

environmental patterns could be inferred.

♦Department of Botany, University of Cape Town, Private Bag,

Rondebosch 7700. Cape Town.

**9 Dorries Drive, Simonstown 7995.

MS. received: 1992-08-17.

STUDY AREA

The area surveyed was in the Pakhuis Pass area of the

northern Cederberg at altitudes ranging from 600 to 1 000

m. The communities sampled lay between 18° 59' E to

19° 04' E, and 32° 08' S to 32° 1 V S, covering an area

of approximately 300 hectares. The vegetation is described

as Mesic Mountain Lynbos (Moll et al. 1984). The geol-

ogy of the area is quartzitic sandstone of the Peninsula

Lormation of the Table Mountain Group. The area falls

within the 60% winter rainfall region. The annual rainfall

is 480 mm of which 72% falls between April and August

(winter). Mean monthly maximum temperatures range

from 20°C (winter) to 30°C (summer). Mean minimum

temperatures drop to 5°C in midwinter.

METHODS

The area was surveyed using aerial photographs in

order to broadly identify communities. Braun-Blanquet

techniques as described by Werger (1974) were used.

Since plant relationships with the environment, and with

other plants, will be related to both the scale of environ-

mental variation and to plant size, releve size was chosen

in relation to vegetation structure (height) as follows: 2 x

2 m (<1 m in height); 2 x 4 m (1-2 m in height); and 4

x 4 m (>2 m in height). The following data were collected

in January 1986: floristic lists and cover abundance for

each of five to six releves per community (22 plots in

total); species not in the releve, but occurring in the sur-

rounding area in a 1-2 m border round the plot recorded

as (+). Environmental data, as set out in Table 1, were

recorded for each releve. The computer program

TWINSPAN (two-way indicator species analysis, Hill

1979) was applied to the raw data, producing a tabular

matrix approximating Braun-Blanquet table work. The

matrix was further sorted by hand.

266

Bothalia 23,2 (1993)

TABLE 1. — Phytosociological table of northern Cederberg communities

Subcommunity 2b

Cannomois parviflora

Anlhospermum aethiopicum

Telraria ustulata

Ficinia deusla

Metalasia densa

Cymbopogon marginatus

Chasmanlhe sp.

Telraria cuspidata

Cluna alaternoides

Elytropappus sp.

Hypodiscus neesii

Pentaschistis curvifolia

Strulhiola ciliata

Selago sp.

Campanulaceae

Subcommunity 2a

Ischyrolepis gaudichaudiana

Merxmuellera arundinacea

2

(+)

1

2

Community 2

Ischyrolepis sieberi

Sloebe plumosa

Ficinia nigrescens

Clifforlia ruscifolia 1 2

Lobostemon glaucophyllus

Subcommunity 3b

Indigofera frutescens

Pentaschistis sp.

Udbeckia quinqueloha

Rhus rimosa

+

+ 1

+

(+)

+ (+)

+ +

(+) (+)

(+) (+)

BB cover values: (+) = not in quadrat, but occurring in surrounding area in a I to 2 m border round plot; + = <1% of quadrat area; 1 = 1-5%;

2 = 5-25%; 3 = 25-50%; 4 = 50-75%; 5 = 75-100%.

Rock cover (superscripts are BB cover values): n = none; b = bedrock; III = rock size (ht), >0.5 m; II = 0.25-0.5 m; I = 0.05-0.25 m.

Soil depth: 0 = skeletal, <0.05 m; I = shallow, 0.05-0.15 m; 01 = mixed skeletal and shallow; 2 = medium, 0.15-1.00 m; 3 = deep, >1.0 m.

Soil moisture: d = dry, well drained; df = dry, flat, hard surface; dr = dry, well drained with runoff.

Slope: 0 = level, 0-3°; 1 = gentle, 4-8°; 2 = moderate, 9-16°; 3 = steep, 17-26°.

Bothalia 23,2 ( 1993)

267

TABLE 1.— Phytosociological table of northern Cederberg communities (continued)

Widespread species

Eriocephalus africanus (+) 2 (+)

Ehrhana ramosa + 2 + ++ I121++ + +

BB cover values: (+) = not in quadrat, but occurring in surrounding area in a 1 to 2 m border round plot; + = <1% of quadrat area; 1 = 1—5%;

2 = 5-25%; 3 = 25-50%; 4 = 50-75%; 5 = 75-100%.

Rock cover (superscripts are BB cover values): n = none; b = bedrock; III = rock size (ht), >0.5 m; II = 0.25-0.5 m; I = 0.05-0.25 m.

Soil depth: 0 = skeletal, <0.05 m; 1 = shallow, 0.05-0.15 m; 01 = mixed skeletal and shallow; 2 = medium, 0.15-1.00 m; 3 = deep, >1.0 m.

Soil moisture: d = dry, well drained; df = dry, flat, hard surface; dr = dry, well drained with runoff.

Slope: 0 = level, 0-3°; 1 = gentle, 4-8°; 2 = moderate, 9-16°; 3 = steep, 17-26°.

Species not included in the above and found in one releve only (species name is followed by releve number, cover abundance value): Agathosma sp. 13, +;

A. puhigera/esterhuyseniae 6, 1; Arctotis laeris 1, (+); A. sp. 2, 2; Aspalathus spinosissima 8, 1; A. tridentata/quinquefolia 2, (+); Arislea singularis 20, +;

Babiana sp. 8, 1; Cassine peragua 19, (+); Chrysocoma tenuifolia 22, (+); Crassula atropurpurea 9, +; Cullumia bisulca 8, 2; Dilairis ixioides 17, (+); Diosma

acmaeophylla 9, (+); Diospyros austro-africana 19, 1; Dodonaea viscosa 22, +; Erica cf. articularis 7, +; Eriocephalus sp. 5, 1; Euclea linearis 22, 1; E.

undulata 22, +; Ficinia cedarbergensis 15, (+); F. compar 5, 1; Gnidia sp. 3, I; Helichrysum rutilans 3, +; Hypodiscus argenteus lb, 1; Kiggelaria africana

19, (+); Knowhonia capensis 22, +; Leucadendron concavum 1, +; Lobelia sp. 14, +; Macrosrylis squarrosa 7, 2; Manulea sp. 3, (+); Maytenus helerophylla

22, 1; Murallia sp. 16, +; Olhonna amplexifolia 22, (+); Passerina glomeraia 8, 1; Pelargonium cf. crithmoides 8, +; P. scabrum 12, I; Pharnaceum sp. 2,

+ ; Podocarpus elongatus 19, 2; Prismatocarpus sp. 1, +; Proiasparagus sp. 18, +; Protea acaulos 2, (+); Restio sp. (H.T. 00714) 6, 3; Rhus dissecta 22, (+);

R. scylophylla 19, +; R. tomentosa 18, 1; Scabiosa sp. 3, 1; Secamone alpinii 18, (+); Selago sp. 13, +; Serruria aitonii 2, (+); Stoebe sp. 4, +.

RESULTS AND DISCUSSION

There are three major floristic communities (Table 1).

1. Thamnochortus plcitypteris-Willdenowia arescens

Community

This community is an open herbland occurring in

releves 1 to 6, and differential species are Thamnochortus

plcitypteris and Willdenowia arescens. This graminoid

guild is further represented by high cover of Ischyrolepis

monanthos in all releves except number 6 (which in gene-

ral has few species); as well as Ficinia dunensis in three

releves; and Ischyrolepis cf. curviramis, Pentaschistis

viscidula , Tetraria nigrovaginata and Ficinia bidbosa in

two releves each. Macrostylis decipiens and Rafnia diffusa

are low sprawling bushes (± 300 mm high), together form-

ing an understorey to the restioid stratum in three releves.

Metalasia agathosmoides occurs in four releves, and there

is an occasional low presence of further asteraceous spe-

cies, Phaenocoma sp., Helichrysum sp„ Elytropappus sp.

and Ursinia sp., in some of the releves.

This community occurs on flat, sandy plains that are

fully exposed to sun and wind. The soil is well drained

268

Bothalia 23,2 (1993)

and deep (>1 m in four releves, and 0.15-1.00 m in two

releves), and consists of a coarse-textured pale grey sand.

There is no rock cover. These factors, together with the

hot, dry summers and low annual rainfall create generally

very dry conditions.

The widespread and generally high cover of Res-

tionaceae species substantiates the hypothesis that these

shallow-rooted species are suited to survive in areas ex-

posed to severe summer drought due to their ability to

respond to summer moisture flushes with very rapid pho-

tosynthesis (Van der Heyden & Lewis 1989). The pres-

ence of both graminoids and Asteraceae concurs with

Campbell's (1986) finding that these taxa predominate at

the dry end of fynbos gradients. The occasional presence

of deeper-rooted proteoids ( Leucadendron concavum, Pro-

tect acaulos and Semina aitonii), each occurring in only

one releve, indicates the infrequent availability of suffi-

cient water at deeper levels.

2. Ischyrolepis sieberi-Ficinia nigrescens Community

This mid-dense graminoid shrubland has Ischyrolepis

sieberi and Ficinia nigrescens as differential species.

There is also a high cover of the shrubs Stoebe plumosa

and Cliffortia ruscifolia, in many of the releves. This com-

munity is spread over two different landscape forms:

releves 7 to 1 1 occur on level areas with much exposed

bedrock, and releves 12 to 17 occur on well-drained rocky

slopes (gentle to steep). Both environments have soils

varying in depth from skeletal to shallow, and consisting

of dark brown, mixed fine and coarse sand particles. Soil

of the bedrock environment type was dry and well drained

when examined in mid-summer, but during winter rains

the underlying bedrock could cause water accumulation.

Soils on the rocky slopes would be well drained during

the rainy season.

Subcommunity 2a

This subcommunity of five releves (7-11) on level bed-

rock has Ischyrolepis gaudichaudiana as sole differential

species. Merxmuellera arundinacea occurs in two releves.

Subcommunity 2b

This subcommunity occurs in the six releves (12-17)

of the rocky slopes environment, and has Cannomois

parviflora and Anthospermum aethiopicum as differential

species. Other graminoid species such as Tetraria ustulata,

T. cuspidata and Ficinia deusta commonly occur as do

the shrubby components Metalasia muricata and Stru-

thiola ciliata. Within sight of each releve was a mature

Protea nitida and/or P. laurifolia forming a sparse wood-

land overstorey. The aspect of these slopes is mostly east

with the one northwest-facing releve (No. 14) being near

to a seepage area. This, together with the high rock cover,

would make it a less dry and less wind-exposed environ-

ment than the bedrock one, and could account for the

absence of most of this subcommunity’s species in the

level bedrock environment (releves 7-11). Furthermore,

the rockiness in combination with slope-related drainage

patterns would result in a variety of moisture-related

microhabitats. This could also be the cause of the greater

numbers of species found on the rocky slopes than on the

level bedrock. Greater species richness on rocky slopes

than on level areas in arid areas has been attributed to

these factors (Barbour & Diaz 1973; Olsvig-Whittaker et

al. 1983).

3. Olea europaea subsp. africana-Maytenus oleoides

Community

The differential species of this closed woodland are

Olea europaea subsp. africana and Maytenus oleoides.

These, as well as Heeria argentea and Euclea natalensis,

are common to most of the releves of subcommunities 3a

and 3b. This community occurs adjacent to large boulders

(4 m and higher) which provide shelter from wind, sun

and fire, as well as extra moisture due to shade and water

runoff. Soils are of medium depth and the slope level to

gentle. This has allowed the development of a closed

woodland of trees and shrubs. Cassine peragua , a forest

tree (in one releve. No. 19), and Protasparagus scandens,

common in forest undergrowth, indicate the tendency of

succession to low forest. Substantial forest development

in this community is undoubtedly limited by the low rain-

fall of the area, and the fact that the prevalence of fire on

the adjacent plains severely limits the forest area.

Subcommunity 3a

The differential species are Myrsine africana and

Chironia baccifera in this subcommunity which occurs in

three releves. These, together with Diospyros glabra and

Euclea acutifolia form an understorey to the closed wood-

land canopy. The ground layer is poorly developed and

consists of sparsely occurring Chironia baccifera. These

species are mostly excluded from the drier subcommunity

3b.

Subcommunity 3b

Indigofera frutescens and Pentaschistis sp. are differ-

ential species. This subcommunity represents a drier gra-

dient of Community 3 since the releves in which it occurs

(20 and 21) have more sunny aspects (SW and NW) than

the other releves which face S or SE. This could account

for the absence of most of these species in the former

subcommunity, and vice versa. These two subcommuni-

ties appear to reflect moisture-related microhabitat differ-

ences within the woodland community.

CONCLUSIONS

A satisfactory phytosociological table was produced

using small plot sizes. We obtained a neat phytosociolog-

ical table, with well-defined communities. This in itself

indicates the existence of small-scale pattern in fynbos.

The survey was quickly (five days field work) and easily

achieved. The efficiency of this method would allow com-

paratively easy monitoring and quick establishment of

plant-environmental relationships.

ACKNOWLEDGEMENTS

We thank R Masson for assistance in using computer

programs, and Prof. R.M. Cowling for commenting on

the manuscript.

Bothalia 23,2(1993)

269

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bajada and moisture gradients in the United States and Argentina.

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BOUCHER, C. 1978. Cape Hangklip area II. The vegetation. Bothalia 12:

455^497.

CAMPBELL, B.M. 1986. Montane plant communities of the Fynbos

Biome. Vegetatio 66: 3-16.

CAMPBELL, B„ GUBB, A. & MOLL, E. 1980. The vegetation of the

Edith Stephens Cape Flats Flora Reserve. Journal of South Afri-

can Botany 46: 435^144.

CAMPBELL, B.M. & MOLL, E.J. 1977. The forest communities of

Table Mountain. South Africa. Vegetatio 34: 105-1 15.

COWLING, R.M. 1984. A syntaxonomic and synecological study in the

Humansdorp region of the Fynbos Biome. Bothalia 15: 175-227.

COWLING, R.M., MOLL, E.J. & CAMPBELL, B.M. 1976. The ecolog-

ical status of the understorey communities of pine forests on Table

Mountain. South African Forestry Journal 99: 13-24.

GLYPHIS, J., MOLL, E. & CAMPBELL. B. 1978. Phytosociological

studies on Table Mountain, South Africa. 1. The Back Table.

Journal of South African Botany 44: 281-289.

HILL, M.O. 1979. TWINSPAN — a FORTRAN programme for arranging

multivariate data in ordered two-way tables by classification of

individuals and attributes. Section of Ecology and Systematics,

Cornell University, Ithaca, New York.

LAIDLER, D„ MOLL, E.J., CAMPBELL, B.M. & GLYPHIS, J. 1978.

Phytosociological studies on Table Mountain, South Africa. 2.

The Front Table. Journal of South African Botany 44: 291—

295.

MCKENZIE, B„ MOLL, E.J. & CAMPBELL, B.M. 1977. A phytosocio-

logical study of Orange Kloof, Table Mountain, South Africa.

Vegetatio 34: 41-53.

MOLL, E.J., CAMPBELL, B.M., COWLING, R.M., BOSSL L„ JAR-

MAN, M.L. & BOUCHER, C. 1984. A description of major

vegetation categories in and adjacent to the Fynbos Biome. South

African National Scientific Programmes Report No. 83.

OLSVIG-WHITTAKER, L„ SHACKAK, M. & YAIR. A. 1983. Vegeta-

tion patterns related to environmental factors in a Negev Desert

watershed. Vegetatio 54: 133-145.

TAYLOR, H.C. 1984a. A vegetation survey of the Cape of Good Hope

Nature Reserve. 1. The use of Association Analysis and Braun-

Blanquet methods. Bothalia 15: 245-258.

TAYLOR, H.C. 1984b. A vegetation survey of the Cape of Good Hope

Nature Reserve. 2. Descriptive account. Bothalia 15: 259-

291.

VAN DER HEYDEN, F. & LEWIS, O.A.M. 1989. Seasonal variation in

photosynthetic capacity with respect to plant water status of five

species of the mediterranean climate region of South Africa. South

African Journal of Botany 55: 509-515.

VAN WILGEN, B.W. & KRUGER, F.J. 1985. The physiography and

vegetation communities of the Zachariashoek catchments; south-

western Cape Province. South African Journal of Botany 51:

379-399.

WERGER, M. 1974. On concepts and techniques applied in the Ziirich-

Montpellier method of vegetation survey. Bothalia 1 1: 309-323.

WERGER, M.J.A., KRUGER, F.J. & TAYLOR, H.C. 1972. A phytoso-

ciological study of the Cape Fynbos and other vegetation at

Jonkershoek, Stellenbosch. Bothalia 10: 599-614.

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Bothalia 23,2:271-326(1993)

The vegetation ecology of municipal Durban, Natal. Floristic classification

D.C. ROBERTS*

Keywords: floristic classification, municipal Durban, phytosociology. PHYTOTAB, urban open space

ABSTRACT

The vegetation of municipal Durban is classified according to the Braun-Blanquet method, using the PHYTOTAB program

package. Five major vegetation types with 18 communities are described, as are the major environmental factors influencing

vegetation composition and structure.

UITTREKSEL

Die plantegroei van Durban se munisipale gebied is volgens die Braun-Blanquet-metode met behulp van die PHYTOTAB-

programpakket geklassifiseer. Vyf hoof plantegroeitipes met 1 8 gemeenskappe word beskryf, asook die bepalende om-

gewingsfaktore wat plantegroeisamestelling en -struktuur be'i'nvloed.

CONTENTS

Introduction 271

Study area 271

Methods 273

Results 274

Description of the plant communities 274

Mangrove Forest 274

Community 1 : Avicennia marina-Bruguiera gym-

norrhiza Mangrove Short Forest 274

Coastal Forest Complex 276

Community 2: Transitional Eugenia capensis-

May terms procumbens High Closed Shrubland 276

Community 3: Mimusops caffra-Allophylus nata-

lensis Low/Short Thicket 278

Community 4: Manilkara discolor-Tricalysia lan-

ceolata Short Thicket 280

Community 5: Protorhus longifolia (sapling)-FVy-

chotria capensis Short Thicket 282

Valley Bushveld Complex 283

Community 6: Rhus pentheri-Cussonia spicata

Low Thicket 283

Community 7: Dovyalis rhamnoides-Hippobro-

mus pauciflorus Low Thicket 285

Shrubland Complex 286

Community 8: Transitional 286

Community 9: Acacia gerrardii-Rhus chirinden-

sis Tall Closed Shrubland//Low/Short Thicket

Mosaic 287

Community 10: Transitional 289

Grassland Complex 291

Community 1 1 : Bothriochloa insculpta-Hypar-

rhenia hirta Short Closed Grassland 291

Community 12: Phragmites australis-Cyperus im-

mensus Short Closed Grassland//Tall/High

Closed Shrubland//Tall Closed Reedbed Mosaic 292

Community 13: Senecio madagascariensis-Pani-

cum maximum Tall Closed Shrubland//Short

Closed Grassland Mosaic 293

* Department of Geographical and Environmental Sciences, University of

Natal, King George Vth Ave, Durban 4001.

MS. received: 1992-04-03.

Community 14: Phyllanthus burchellii-Digitaria

eriantha Short Closed Grassland//Tall

Sparse/Closed Shrubland Mosaic 295

Community 15: Sutera kraussianci-Aristida junci-

fonnis Low/Short Closed Grassland 298

Community 16: Hypoxis gerrardii-Alloteropsis

semialata Low Closed Grassland 299

Community 17: Helichrysum aureum-Themeda

triandra Short Closed Grassland 300

Community 18: Diantluis zeyheri-Eragrostis

curvula Short/Low Closed Grassland 302

Discussion and conclusions 303

Acknowledgements 304

References 304

Appendix 304

INTRODUCTION

In order to facilitate ecologically effective open space

planning and management within municipal Durban, a

comprehensive survey of all the remaining vegetated areas

within the city was undertaken. This excluded landscaped

or formally managed areas (e.g. parks, gardens and agri-

cultural lands). The study included classification of the

vegetation in terms of both floristics and structure, as well

as correlation of the vegetation classification with a range

of environmental parameters. This latter facilitated a more

accurate interpretation of the current ecological status and

conservation value of the City’s urban open space re-

sources.

STUDY AREA

The study area incorporates two local authority areas:

namely municipal Durban and Yellowwood Park. Al-

though Yellowwood Park is administered by an indepen-

dent health committee, it is completely encircled by

municipal Durban, and is included within the boundaries

of the study area to ensure physical continuity (Figure 1).

At the commencement of the study in 1983, municipal

Durban and Yellowwood Park together covered an area

of approximately 300.13 km“ (P. Johnson pers. comm.);

8.5 km" of which was covered by open water in Durban

272

Bothalia 23,2 (1993)

FIGURE 1 . — Location of the study

area.

Harbour. In 1985/1986, the Durban municipal boundaries

were altered as a result of extensions made in the Isipingo

and Phoenix areas. Because of the difficulty of altering

the boundaries of the study area midway through the re-

search, and the fact that these changes would not signif-

icantly affect or compromise the outcome of the study,

the original municipal boundaries were retained.

Accordingly, the Umlaas River marks the southern

limit of the study area as far as the Umlaas canal, from

whence the boundary follows the route of the previously

uncanalised river to the coast at Isipingo. To the west and

north the study area is flanked by the districts of Klaarwa-

ter, Pinetown, Shallcross, Umhlatuzana, Queensburgh,

Westville, Chesterville, Clermont, KwaDabeka, Kwa-

Mashu, Duffs Road, Inanda, Mount Edgecombe, Glen

Anil and Umhlanga. This area is bounded by latitudes 29°

40' 20" and 29° 55' 55" south, longitude 30° 50' 03" east

and the Indian Ocean (Figure 1).

Low rolling hills characterise most of the study area,

with the terrain varying in altitude from sea level along

the eastern boundary to over 350 m above sea level in

Chatsworth, a distance of approximately 15 km. A major

topographic feature of the area is the steep-sided sand

ridge known as the Bluff Ridge. This ridge stands over

80 m (maximum height 108 m) in altitude and diverges

from the normal line of the coast at an angle of about 14

Bothalia 23,2 (1993)

273

degrees (Preston- Whyte 1980). Inland and parallel to the

Bluff Ridge is a second, smaller ridge with a maximum

height of 91 m above sea level, which is often regarded

as part of the Bluff Ridge (Preston-Whyte 1980; King

1982). For the purposes of this study this ridge was re-

garded as a distinct land form and referred to as the Went-

worth Ridge (Alexander 1987). Both the Bluff and the

Wentworth Ridge trend in a southwest to northeast direc-

tion.

West of the Bluff and Wentworth Ridges, a third ridge,

known as the Berea Ridge, parallels the coast and rises

to about 130 m (maximum height 151m above sea level)

in altitude. Natal Bay, about which the city of Durban has

grown, lies between these two sets of ridges and is en-

closed by a curved spit which extends from the Umgeni

River estuary to the Point, and never exceeds 6 m in al-

titude (King & Maud 1964). The floor of the corridor

between the ridges is flat and low-lying and consists

largely of alluvial deposits from rivers draining into the

bay. These rivers have entrenched deep, youthful valleys

through the Berea Ridge but the coast of the Bluff Ridge

is unbroken in the bay area (Preston-Whyte 1980).

West of the Berea Ridge, away from the foreshore and

central city area, the terrain rises rapidly and becomes

progressively more rugged towards the interior (King &

Maud 1964). Here a low-lying hill and valley region

slopes up to interfluvial crests separated by the entrenched

east-west valleys (Preston-Whyte 1980) of the Umlaas,

Umhlatuzana, Umbilo and Umgeni Rivers. The valley

sides of these major rivers are steep, and where they cross

Natal Group Sandstone, gorges have been formed. These

large rivers have greatly influenced the topography of the

study area (Cawood 1980).

The geological units found within the study area are

considerably diverse and are discussed in detail by King

& Maud (1964). Archaean granite-gneiss underlies the

whole of the study area and is unconformably overlain

by: the arenaceous Natal Group Sandstones; the glacial

Dwyka Series and the argillaceous and arenaceous

Pietermaritzburg Shale and Vryheid Formations, both of

the Karoo sequence. There are several minor intrusions

(dykes and sills) of Karoo Dolerite scattered throughout

these formations. All other geological formations within

the Durban area have accumulated during the Quaternary

Era. These are: the Bluff Sandstones: the Berea Red Sand

and boulderbed; and sundry unconsolidated sand, grit and

clay associated with the harbour and city area; with alluvia

along the major river courses (King & Maud 1964).

The range of soils found in the Durban area is linked

not only to the nature of the parent geological material

from which they are derived, but also to the topography

of the land. The soils represented include: Arcadia

Rydalvale, Cartref Cartref, Dundee Dundee. Femwood

Fernwood, Glenrosa Williamson, Hutton Clansthal, Kat-

spruit Katspruit, Kroonstad Avoca. Kroonstad Mkambati.

Longlands Waldene, Milkwood Milkwood, Mispah

Mispah, Rensburg Phoenix, Shortlands Shortlands and

Swartland Swartland (Mac Vicar et al. 1977).

According to Acocks (1988) the study area contains

elements of both Coastal Forest and Thomveld (Veld Type

1) and Valley Bushveld (Veld Type 23), but both have

been badly disturbed by the process of urbanisation. The

climate in Durban is generally warm and humid with a

wet summer season, but with some form of precipitation

in all seasons (Cawood 1980). The main form of precip-

itation is rain, with an average of between 1 000 mm and

1 250 mm per annum (Preston-Whyte 1980). According

to Koppen’s classification (Schulze 1947), Durban falls

within the Cfwa unit. The mean annual temperature is

20.5°C with an annual range of 8.3°C (Preston-Whyte

1980).

METHODS

The Braun-Blanquet method of sampling and synthesis

followed in the study is described by Westhoff & Van der

Maarel ( 1973), Mueller-Dombois & Ellenberg ( 1974) and

Werger ( 1974).

The study area was stratified into physiognomic-phys-

iographic units using 1:1 750 aerial photography. Sam-

pling intensity was determined using the relationship

between mapping scale, vegetation structure and floristic

community described by Rutherford & Westfall (1986).

Sample sites were located by means of random co-ordi-

nates within each physiognomic-physiographic unit iden-

tified. A total of 345 sample sites, each of 1 1 3 m“ (as

determined by Rutherford & Westfall's (1986) proposed

relationship between mapping scale and smallest mappa-

ble unit area) were sited in homogeneous stands repre-

senting the different physiognomic-physiographic units.

At each sample site all plant species were listed to-

gether with their Domin-Krajina cover abundance values

and growth forms. The total canopy cover and height

range of each stratum of vegetation was also recorded for

the purpose of physiognomic-structural classification (Ed-

wards 1983). The environmental parameters recorded

were altitude, aspect, slope, geological formation and li-

thology, soil texture, soil depth, soil form and series, geo-

morphology and land use.

The vegetation was classified according to the Braun-

Blanquet method using the PHYTOTAB program package

(Westfall et al. 1982; Westfall & De Wet 1988). The main

environmental factors influencing the communities were

derived from gradients determined by an ordination of flo-

ristic data by detrended correspondence analysis (DCA)

(Hill & Gauch 1980) using the DECORANA program

(Hill 1979). Species diversity for each community was

expressed as a function of average species number per

releve/area of the releve. Furthermore, because of the sig-

nificant impact of alien invasive species on the plant

ecolgy of municipal Durban, figures for percentage occur-

rence and average cover of the two predominant species:

Chromolaena odorata * and Lantana camara* are record-

ed for all community types. (Note: * denotes alien spe-

cies).

Community structure is illustrated by means of layer

diagrams (Ito 1979) whereby vegetation height classes are

grouped into three strata, namely a herb/grass stratum (0-

0.5 m), a shrub stratum (0.5-5.0 m) and a tree stratum

(>5.0 m), and determining the highest mean percentage

cover in each stratum. The following symbols, determined

274

Bothalia 23,2 (1993)

by the highest mean cover in a stratum, are used to classify

each layer diagram type.

Layer diagram type

L-type

rL-type

D-type

C-type

I-type

Cover of strata

herb>shrub>tree

herb<shrub<tree

herb<shrub>tree

herb>shrub<tree

herb=shrub=tree

RESULTS

DESCRIPTION OF THE PLANT COMMUNITIES

In the community descriptions woody and herbaceous

species are both listed in order of constancy followed by

mean percentage cover, with the respective values indi-

cated next to each species. The general species character-

istics of each community are omitted from the community

descriptions because they are directly apparent from the

Appendix (p. 304). Where species occurrences have spe-

cial significance, however, these are noted.

Classification of the vegetation, according to the

Braun-Blanquet method, revealed 18 plant communities

within five major vegetation types. Communities are

named according to the preliminary rules and recommen-

dations for a standardised South African syntaxonomic no-

menclature system proposed by the South African

Syntaxonomic Nomenclature Committee (Scheepers et al.

unpubl.). Each name comprises a diagnostic species fol-

lowed by a dominant species and a physiognomic struc-

tural term (Edwards 1983). Because of the highly

disturbed nature of most of the remnant plant communities

found within the Durban area, it is not possible to assign

a single physiognomic structural term to many of the

groupings described. A long history of disturbance, vary-

ing spatially, temporally and in intensity, has produced a

mosaic of structural types within each distinct floristic

grouping. In order to accommodate this variation and give

a truer representation of the nature of the plant commu-

nities involved, each community is described as a physi-

ognomic mosaic where appropriate. Edwards' (1983)

structural classification makes provision for this through

the use of / 'where straddling of height classes occurs’

and // ‘where mosaics of classes are found’.

The phytosociological classification of communities is

shown in the synoptic Appendix (p. 304). Diagnostic spe-

cies are listed in groups 1—15, and non-diagnostic and

infrequent species in groups 46-52**. The taxa on the left

of the matrix, are grouped into simplified lifeform classes

to facilitate 'veld condition’ assessment (Westfall et al.

1983); the life form classes are based on Dyer's (1976)

description of the genera of South African flowering

plants.

MANGROVE FOREST

Community 1: Avicennia marina-Bruguiera gymnor-

rhiza Mangrove Short Forest (Figure 3)

This short mangrove forest (Edwards 1983) is found

along the Natal coast in bays, lagoons and at river mouths

where suitable mudflats occur (0-5 m). It is represented

by releves 11, 13, 14, 201 and 202, with 2-5 species per

releve. This short forest community has a rL structure (Ito

1979; Figure 2A) with the greatest average cover of 5 1.6%

in the upper height class of higher than 5 m. The two

lower height strata are comprised almost exclusively of

the saplings and seedlings of the three dominant tree spe-

cies.

Habitat

The soils are of the Femwood Form, Femwood Series

and are derived from grey sands of the Berea System and

recent alluvial deposits. The terrain is generally flat with

a slight easterly (Beachwood Nature Reserve) or westerly

(Bayhead mangroves) aspect.

Floristics

The community is diagnosed by the Avicennia marina

species group (Appendix .1), and has no species in com-

mon with any other species group listed in the Appendix.

The character species for this group consist exclusively

of three halophytic mangrove tree species: Avicennia ma-

rina, Bruguiera gymnorrhiza and Rhizophora mucronata.

The species diversity per unit area for this community is

therefore low, with an average of 0.03 species per m",

and an average of three species per releve.

In cases where various life forms, such as trees and

shrubs have adopted a scandent growth form, these are

referred to as climbers. A distinction is also made within

species in terms of developmental stage; with seedlings,

saplings and fully grown forms of each species being

clearly differentiated from one another. In instances where

plants could not be identified beyond the generic level,

the epithet sp. is appended (e.g. Ipomoea sp.). It should

be noted that such a name may not necessarily refer to a

consistent species concept, but may include various spe-

cies within the genus. The structural classification of each

community together with appropriate layer diagram type

is shown in Figure 2.

** complete phytosociological tables for both the diagnostic species and

the non-diagnostic species are available from the author or Dr R.H.

Westfall, Agricultural Research Council, Roodeplaat Grassland Institute,

Private Bag X05, Lynn East 0039, Pretoria.

Trees and shrubs

% Mean

Occurrence cover

Chromolaena odorata* (shrub) 0.0% 0.0%

Lantana camara* (shrub) 0.0% 0.0%

Conspicuous woody species with more than 5% mean

cover and occurring in more than 20% of the releves rep-

resenting this community are:

Avicennia marina ( tree) 100.0% 45.8%

Bruguiera gymnorrhiza (tree) 100.0% 38.6%

Rhizophora mucronata (tree) 60.0% 20.4%

Herbs

The only herb species present in more than 20% of

the releves representing the community is:

Bothalia 23,2(1993)

275

Arthrocnemum perenne (undershrub) 20.0% 0.002%

(non-diagnostic)

The presence of this genus as the only representative

of the herb stratum (other than seedlings of the three dom-

inant tree species) indicates the saline, mesic conditions

experienced in this community.

Saplings and seedlings

Saplings and seedlings occurring in more than 20% of

the releves representing the community are:

General

Community 1 is floristically unrelated to any of the

other vegetation types identified within the study area, and

thus shares no common species groups with any of the

other communities. It is equivalent to Acocks’s (1988)

Mangrove Forest, Moll’s (1976) Mangrove Swamp and

allied communities and Ward’s (1980) Mature Mangrove

Communities.

Because of its floristic isolation, and the fact that this

community depends entirely on within-community dyna-

mics for long-term survival, the mangroves are considered

to have a high conservation status. This ranking is further

re-inforced by the restricted occurrence of this community

within the municipal Durban area. Only two sites remain:

the first and largest at Beachwood Nature Reserve, and

the second in a limited area on the mudflats of Durban

Bay to the west of Salisbury Island. A small, threatened

community occurs at the Isipingo Lagoon, just outside of

Durban. If these remaining sites are lost, there is little or

no potential for re-establishment of similar communities

elsewhere in the municipal area due to loss of suitable

habitat.

The ecological autonomy of this vegetation type may

be attributed to the specific habitat requirements of the

community (Hutchings & Saenger 1987). Mangroves

5m <

0,5 < 5m

Om < 0,5m

0 25 50 75 100

Community 5 type rL

I

5m <

0,5 < 5m

Om < 0,5m

0 25 50 75 100

Community 9 Type D

0 25 50 75 100

Community 14 Type D

5m <

0,5 < 5m

Om < 0,5m

0 25 50 75 100

Community 17 Type D

0 25 50 75 100

Community 3 Type D

0 25 50 75 100

Community 1 1 Type D

0 25 50 75 100

Community 18 Type D

0 25 50 75 100

Community 4 Type rL

0 25 50 75 100

Community 16 Type L

FIGURE 2. — Average community structure of the vegetation of muncipal Durban, showing height classes, strata, mean percentage cover and layer

diagram types (see Methods).

276

Bothalia 23,2 (1993)

FIGURE 3. — Community 1: Avicen-

nia marina-Bruguiera gym-

norrhiza Mangrove Short For-

est. Note continuous canopy of

Avicennia marina and large

number of establishing Avi-

cennia marina propagules in

foreground. Location: Bay-

head Mangroves.

occur only on shores where the vigour of the surf is bro-

ken by sand bars or coral reefs or islands (White 1983).

This niche is effectively unavailable to plant species other

than die specifically adapted halophytes which occupy it.

As a result the mangrove community has the lowest spe-

cies diversity of all the communities described in this

study (Figure 4). The extreme environmental conditions

have also prevented invasion by Chromolaena odorata*

and Lantana camara* (Figures 5 & 6), the two most im-

portant invasive alien species in Natal (Macdonald & Jar-

man 1985).

Habitat

The soils are of the Femwood Form, Fernwood Series,

and Hutton Form, Clansthal Series. The latter occur in the

two atypical releves found within this community, 205

and 219, and should not be regarded as characteristic. The

terrain varies from flat to a slope of 26.6°. Aspect is also

variable, with all points of an eight-point rose equally rep-

resented except for north, northeast and southwest.

FI ori sties

The rL structure illustrated in the structure diagram

(Figure 2 A) is a product of these environmental restraints,

as the limit placed on the number of species capable of

surviving in these areas means that the community is

dominated exclusively by the three mangrove tree species,

Avicennia marina, Bruguiera gymnorrhiza, Rhizophora

mucronata. This produces a community structure cha-

racterised by a single layer of trees with little or no un-

derstorey. The cover within the 0.5-5.0 m category is

made up of saplings and young individuals of the three

tree species. Seedlings and/or saplings for all three domi-

nant tree species were recorded only within the releves

sampled at Bayhead in the harbour.

COASTAL FOREST COMPLEX

Community 2: Transitional Eugenia capensis-Maytenus

procumbens High Closed Shrubland (Figure 7)

This high closed shrubland//low thicket community

mosaic (Edwards 1983) is found between 5 and 75 m

above sea level. It is represented by seven releves (Ap-

pendix) with 4—10 species per releve. This community has

a D structure (Ito 1979; Figure 2B) with the greatest av-

erage cover of 75.3% in the 0.5-5.0 m height class.

This community is differentiated by the absence of

character species. The species diversity per unit area is

low and averages 0.06 species per nT for the community,

with an average of 6.6 species per releve.

Trees and shrubs

Percentage occurrence and average cover of Chromo-

laena odorata* and Lantana camara* are:

Average number of species per m2

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

Community number

FIGURE 4. — Species diversity for Communities 1-18.

Bothalia 23,2(1993)

277

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

Community number

Percentage of releves

Community number

FIGURE 5. — Average percentage cover of Chromolaena odorata* and

Lantana camara* in Communities 1-18.

Chromolaena odorata* (shrub) 42.9% 23.9%

Lantana camara* (shrub) 28.6% 14.6%

Conspicuous woody species with more than 5% mean

cover and occurring in more than 20% of the releves rep-

resenting this community are:

Ficus burtt-davyi (shrub) 28.6% 23.9%

Rhus nebulosa (shrub) 28.6% 14.6%

Albizia adianthifolia (tree) 28.6% 8.1%

Herbs

There are no herb species occurring in more than 20%

of the releves representing the community.

Climbers

The only climber present in more than 20% of the

releves representing the community is:

Rhoicissus digitata ( shrub) 28.6% 28.6%

FIGURE 6. — Percentage of releves in Communities 1-18 in which

Chromolaena odorata* and Lantana camara* were recorded.

General

The majority of the releves constituting this community

occur in the frontal section of the secondary dunes (Tinley

1985) and are floristically equivalent to Tinley’s (1985)

Scrub-Thicket Community, Cawood's (1980) Dune Scrub

and Ward's (1980) Closed Dune Scrub. Acocks (1988)

does not recognise this as a separate community and de-

scribes this formation as part of the Dune Forest unit.

According to the classification presented in this study,

also included within this community are two releves (205

and 219) comprised predominantly of dense stands of the

two invasive species Chromolaena odorata * and Lantana

camara *, situated well inland of the secondary dune area.

Although these releves are floristically different from

those located within the dune area, they are similar in

terms of cover, structure, density and species richness. For

the purposes of community description, the frontal dune

scrub/thicket community (Moll 1976; Tinley 1985) repre-

FIGURE 7. — Community 2: Tran-

sitional Eugenia capensis -

Maytenus procumbens High

Closed Shrubland. Note wind-

clipped, compact, even canopy

of closed frontal dune commu-

nities (in the middle and back-

ground) containing Eugenia

capensis. Maytenus procum-

bens and Rhus nebulosa. Dune

pioneer Scaevola thunbergii

(of the Strand community) in

foreground. Location: Vir-

ginia Beach.

278

Bothalia 23,2 (1993)

sented by Community 2 (excluding the atypical releves)

could be more accurately described as Eugenia capensis-

Maytenus procumbens High Closed Shrubland rather than

as a transitional group as indicated in the Appendix.

Community 2 is related to Community 3 through the

shared presence of the Eugenia capensis species group

(Appendix .3), and to Communities 3 and 5 through the

shared presence of the Ficus burtt-davyi species group

(Appendix .9), this indicates a common indigenous shrub

element throughout all three communities. Its association

with the other vegetation complexes is limited to gener-

alist species groups (both indigenous and alien) that are

ubiquitous throughout the study area, and is the result of

the presence of the two atypical releves already noted.

The strong floristic relationship between Communities 2,

3 & 5 is the first indication that the community spectrum

should be treated as the fundamental unit for conservation.

This implies that it is insufficient merely to preserve a

single area of each community type without preserving

ecological/physical continuity with viable examples of the

associated types.

As with the mangroves, the species diversity in the

frontal dune area is restricted by the severe environmental

conditions to which the community is exposed (Ward

1980). The diversity of the frontal dune communities is,

however, higher than in the mangroves (Figure 4). The

presence of Chromolaena odorata * and Lantana camara*

in this community (Figures 5 & 6) is due only to the

inclusion of the two atypical releves, although Cawood

(1980) and Ward (1980) both note that Lantana camara*

is an important invasive species in areas of disturbance

in the frontal dune community. In this connnunity a D

structure predominates (Figure 2B), and is the result of

the ‘salt-spray factor' which produces a clipped-hedge ap-

pearance (Ward 1980). The predominant structural stratum

in this community therefore falls within the 0.5-5.0 m

category. The existence of a cover value in the greater

than 5 m category, is the result of releves sampled further

inland, here ‘as the scrub-thicket matures it provides in-

creasing protection and humus, with patches of uneven

canopy in lee sites and hollows’ (Tinley 1985).

Community 3: Mimusops caffra-Allophylus natalensis

Low/Short Thicket (Figure 8)

This low/short thicket community (Edwards 1983) is

found between 5 and 142 m above sea level. It is repre-

sented by 23 releves (Appendix) with 7-26 species per

releve, and has a D structure (Ito 1979; Figure 2C) with

the greatest average cover of 68.9% in the 0.5-5 .0 m

height class.

Habitat

The soils are mainly of the Femwood Form, Femwood

Series (8 releves) and Hutton Form, Clansthal Series (13

releves). However, the two releves located within

Stainbank Nature Reserve have soils of the Cartref Fonn,

Cartref Series and Kroonstad Form, Mkambati Series. The

terrain varies from flat to a slope of 45°. The aspect of

the land in each of the releves is also variable, with all

points of an eight-point rose represented except northeast.

The southeasterly aspect is the most common, and found

in six out of the 23 releves.

Floristics

This community is differentiated by the Mimusops

caffra-Allophylus natalensis species group (Appendix .2).

The species diversity averages 0.15 species per rn with

an average of 14.1 species per releve. A notable feature

of this community is that 47.1% of the diagnostic species

for this community are either seedlings or saplings.

Trees and shrubs

Percentage occurrence and average cover of Chro-

molaena odorata* and Lantana camara* are:

FIGURE 8. — Community 3: Mimu-

sops caffra-Allophylus nata-

lensis Low/Short Thicket.

Note the lack of large trees or

forest patches. Predominant is

a thicket of Brachylaena dis-

color, Chrysanthemoides mo-

nilifera, and Strelitzia nicho-

lai , interspersed with repre-

sentatives of Allophylus na-

talensis and Mimusops caffra.

Chromolaena odorata* and

Lantana camara* in fore-

ground indicate previous dis-

turbance. Location: southeast-

ern Bluff slopes.

Bothalia 23,2(1993)

279

Saplings and seedlings

The only sapling occurring in more than 20% of the

releves representing the community is:

Deinbollia oblongifolia sapling (tree) 39.1% 0.04%

General

This community can be equated with Acocks’s ( 1988)

and Moll's (1976) Dune Forest, Tinley’s (1985)

Thicket/Forest Community and Ward's (1980) Mimusops

cajfra Woodland. Within the study area no significant

patches of forest remain due to previous logging of the

larger trees for building poles and fuel wood (Ward 1980;

Director Parks, Beaches and Recreation Department

1989). To a large extent the community is dominated by

heliophytic thicket elements such as Mimusops and

Brachylaena with a significant creeper component (Moll

1976; Tinley 1985). The dense tangle of creepers, climb-

ers, scandents and woody lianas which entwine the canopy

is one of the most striking features of this community.

These creepers are most abundant on the margins of

thicket/forest, in openings where the canopy is light and

especially in secondary habitats.

Moll (1976) (also Ward 1980) notes that ‘much of the

high density of these lianas can be attributed to recent

disturbance, particularly where Flagellaiia is present.' The

predominance of Brachylaena discolor over large areas of

this community can also be attributed to previous damage

and disturbance (Moll 1976). Based on his observations

at Isipingo during the period 1949-1960, Ward (1980)

concluded that Allophylus natalensis and Brachylaena dis-

color, both characteristic species in this community, are

not tolerant of deep shade, and unless there is disturbance,

do not occur in the Dune Forest proper. The fact that

47.1% of the diagnostic species for this community are

either seedlings or saplings indicates that regeneration is

a significant aspect of this community’s dynamics, and

could also be related to prior disturbance.

Communities 2 and 3 are clearly related through the

common presence of the Eugenia capensis species group

(Appendix .3) and the Ficus burtt-davyi species group

(Appendix .9). The latter also shows floristic associations

with Community 5. The association between Communities

2 & 3 indicated by the Eugenia capensis species group

(Appendix .3), is dependent on the fact that these two

groups form the dune component of the Coastal Forest

Complex. Together they represent a gradation from the

high closed shrubland of the foredunes through to the

low/short thicket of the backdunes.

The affinities of the Ficus burtt-davyi species group

(Appendix .9) indicate that the dune components of the