ISSN 0006 8241 = Bothalia

Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Vol. 22,2 Oct./Okt. 1992

PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE, PRETORIA

PUBLIKASIES VAN DIE NASIONALE BOTANIESE INSTITUUT, PRETORIA

Obtainable from the National Botanical Institute, Private Bag X101,

Pretoria 0001, Republic of South Africa. A catalogue of all available

publications will be issued on request.

BQTHALIA

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.

Verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101,

Pretoria 0001, 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.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The memoirs are individual treatises usually of an ecological nature, ’n Reeks van losstaande omvattende verhandelings oor vemaamlik

but sometimes dealing with taxonomy or economic botany. ekologiese, maar soms ook taksonomiese of plantekonomiese onderwerpe.

THE FLOWERING PLANTS OF AFRICA / DIE BLOMPLANTE VAN AFRIKA

This serial presents colour plates of African plants with 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 TOO 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.

From Vol. 50, one part of twenty plates is published annually. A volume

consists of two parts. The publication is available in English and Afrikaans.

Hierdie reeks bied kleurplate van Afrikaanse plante met bygaande teks.

Die skilderye word meestal deur die kunstenaars van die Nasionale

Botaniese Instituut voorberei. Thlle 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

ekonomiese waarde vir afbeelding.

Vanaf Vol. 50 word een deel, bestaande uit twintig plate, jaarliks

gepubliseer. ’n Volume bestaan uit twee dele. Die publikasie is beskikbaar

in Afrikaans en Engels.

FLORA OF SOUTHERN AFRICA / FLORA VAN SUIDELIKE AFRIKA

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

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

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

species, infraspecific taxa, keys to genera and species, synonymy, literature

and limited specimen citations, as well as taxonomic and ecological

notes.

’n Thksonomiese verhandeling oor die flora van die Republiek van Suid-

Afrika, Ciskei, Transkei, Lesotho, Swaziland, Bophuthatswana, Namibie,

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

spesies, inffaspesifieke taksons, sleutels tot genusse enspesies, sinonimie,

literatuur, verwysings na enkele eksemplare, asook beknopte taksonomiese

en ekologiese aantekeninge.

PALAEOFLORA OF SOUTHERN AFRICA / PALAEOFLORA VAN SUIDELIKE AFRIKA

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

southern Africa. Much of the information is presented in the form

of tables and photographic plates depicting fossil populations. Now

available:

’n Palaeoflora met ’n uitleg veigelykbaar met di6 van die Flora van suide-

like 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.

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

Digitized by the Internet Archive

in 2016

https://archive.org/details/bothaliavolume2222unse_0

BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 22,2

Scientific Editor/ Wetenskaplike Redakteur: O.A. Leistner

Technical Editor/Tegniese Redakteur B.A. Momberg

NATIONAL

N S T I T U T E

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

1992

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

Editorial Committee

Redaksiekomitee

O.A. Leistner

B.A. Momberg

M.C. Rutherford

R.A. DYER NBI RESEARCH PRIZE

This prize is awarded to NBI staff members for the most

meritorious contribution(s) (actions or publications)

that have promoted the research aims of the NBI during

the previous financial year or over an extended period.

During a ceremony on 22 May of this year, commemo-

rating the founding of the Pretoria National Botanical

Garden by Dr R.A. Dyer, this Prize was awarded to

Mrs S.M. Perold.

Mrs Perold is internationally known for her research

on mosses, more specifically the liverworts. She started

her work on this group of plants some 13 years ago and

since 1984, when she began publishing her results, 35

of her publications have been produced, mostly in this

journal.

We congratulate you, Sarie Perold, and wish you many

more productive and fulfilling years.

CONTENTS — INHOUD

Volume 22,2

I . New combinations and resurrected names in Microcharis and Indigastrum (Fabaceae— Papilionoideae).

B. D. SCHRIRE 165

2 . Studies in the Justicia and Siphonoglossa (Acanthaceae) species of southern Africa: final conclusions.

K.L. IMMELMAN 171

3 . Cololejeunea cardiocarpa, an epiphyllous liverwort in southern Africa (Lejeuneaceae). S.M. PEROLD 177

4 . Notes on African plants:

Bryophyta. New and interesting records of mosses in the Flora of southern Africa area: 3. Miscel-

laneous acrocarpous taxa. J. VAN ROOY and S.M. PEROLD 195

Bryophyta. Notes on the moss flora of Zimbabwe. J. VAN ROOY 196

Fabaceae. A new species of Coelidium (Liparieae). A.L. SCHUTTE and B-E. VAN WYK ... 189

Fabaceae. Lessertia sneeuwbergensis, a new species from the Middelburg District of the central

Cape Province. G. GERMISHUIZEN 189

Fabaceae. Cyclopia squamosa (Podalyrieae), a new species from the southwestern Cape Province.

A.L. SCHUTTE 190

Geraniaceae. Typification of Pelargonium section Polyactium. P. VORSTER and G.L. MAGGS 195

Gesneriaceae. Notes on the genus Streptocarpus . T.J. EDWARDS, C. KUNHARDT and S. VENTER 192

Liliaceae/Asphodelaceae. Lectotypification of Apicra jacobseniana (Alooideae). G.F. SMITH 196

Pyxinaceae. A new species in the lichen genus Heterodermia, from coastal Namaqualand. F. BRUSSE 183

Ricciaceae. Lectotypification of Riccia crystallina. S.M. PEROLD 185

Stilbaceae. Stilbe verticillata , the correct name for the species previously known as Stilbe mucronata.

J.P. ROURKE 192

5. Systematic studies in the genus Mohria (Anemiaceae: Pteridophyta). III. Comparative sporangium

and spore morphology. J.P. ROUX 199

6. Richness, composition and relationships of the floras of selected forests in southern Africa. C.J.

GELDENHUYS 205

7. An analysis of the orchid flora of Mt Mulanje, Malawi. H. KURZWEIL 235

8. Die fitososiologie van die Bankenveld in the Grootvlei-omgewing, Suid-Transvaal. W.J. MYBURGH,

P.J.J. BREYTENBACH, E.K. THERON en G.J. BREDENKAMP 245

9. A phytosociological study of Signal Hill, Cape Town, utilizing both perennial and ephemeral species.

C. JOUBERT and E.J. MOLL 255

10. The recovery and dynamics of submerged aquatic macrophyte vegetation in the Wilderness lakes,

southern Cape. P.J. WEISSER, A.K. WHITFIELD and C.M. HALL 283

I I . Flora and vegetation of the Mbonambi Beach Arcuate Scar on the Zululand dune barrier, Natal,

South Africa. P.J. WEISSER, E.C.A. SMITH, A.P. BACKER and S. VAN EEDEN 289

1 2 . Obituaries: Amy Frances Gordon Jacot Guillarmod, nee Hean (1911-1992). M.J. WELLS and E. BRINK.

Lucy Kathleen Armitage Chippindall (1913—1992). N.P. BARKER 295

13. National Botanical Institute, South Africa: list of staff and publications 301

1 4 . Book review 313

■ 4 '■

Bothalia 22,2: 165-170 (1992)

New combinations and resurrected names in Microcharis and

Indigastrum (Fabaceae-P^pilionoideae)

B.D. SCHRIRE*

Keywords: Indigastrum, Indigofera, Microcharis, new combinations, resurrected names, taxonomy

ABSTRACT

As a result of cladistic studies in the tribe Indigofereae in Africa and Madagascar (Schrire 1991) evidence was accumulated

to justify resurrecting the genera Microcharis Benth. and Indigastrum Jaub. & Spach, which were previously considered

part of Indigofera L. All described species in each genus are listed alphabetically. Forty-seven new combinations, six changes

in rank and four resurrected names are published.

UITTREKSEL

Na aanleiding van kladistiese studies van die tribus Indigofereae in Afrika en Madagaskar (Schrire 1991), is getuienis

versamel ter regverdiging van die herinstelling van die genusse Microcharis Benth. en Indigastrum Jaub. & Spach, wat voorheen

as deel van Indigofera L. beskou is. Alle spesies wat in elke genus beskryf is, word alfabeties gelys. Sewe-en-veertig nuwe

kombinasies, ses veranderinge in rang en vier heringestelde name word gepubliseer.

MICROCHARIS Benth.

Microcharis Benth. in Benth. & Hook, f., Genera

plantarum 1: 501 (1865a); Benth.: 297 (1865b); Bak.: 132

(1871); Taub.: 277 (1894); Engl.: 213 (1895); Hutch. &

Dalziel: 388 (1927); Bak. f.: 256 (1929). Lectotype

species: Microcharis tenella Benth., selected by J.B.

Gillett: 127 (1958).

Indigofera subgenus Microcharis (Benth.) J.B. Gillett:

127 (1958c); J.B. Gillett: 321 (1971).

Indigofera subgenus Indigastrum sensu Cronquist: 124

p.p. excl. sp. 64.

Indigofera sensu Hutch.: 400 (1964); Polhill: 291 (1981);

De Kort & Thijsse: 104 (1984); Lock: 289 (1989); Thu-

lin: 119 (1990).

1. Microcharis ammophila (Thulin) Schrire, comb.

nov.

Indigofera ammophila Thulin in Nordic Journal of Botany 2: 47 (1982).

Type: Kenya, Mandera Dist., 12 km S of El Wak on Wajir road, 11-5-1978,

Gilbert & Thulin 1672 (UPS, holo.; BR, C, EA, K!, KRA, MO, WAG).

2. Microcharis angolensis Bak. in Flora of tropical

Africa 2: 132 (1871); Hiern: 230 (1896). Type: Angola,

Pungo Andongo, banks of river Cuanza, near Calunda-

Quisonde, fl. 1-1857, Welwitsch 2004 (LISU, lecto.; BM!,

K!). Selected by Torre (1962), although incorrectly desig-

nated a holotype.

Indigofera richardsiae J.B. Gillett: 583 (1956); J.B. Gillett: 131 (1958c);

Torre: 137 (1962); J.B. Gillett: 323 (1971). Type: Zambia, Mbala Dist.,

Pans road, Richards 1268 (K, holo.!). J.B. Gillett published this name

because of Indigofera angolensis D. Dietr.

* Botany Department, University of Natal, Pietermaritzburg, South

Africa. Present address: Royal Botanic Gardens, Kew, Richmond, Surrey,

TW9 3AB, UK.

MS. received: 1991-08-27.

3. Microcharis annua (Milne-Redh.) Schrire, comb.

nov.

Indigofera annua Milne-Redh. in Kew Bulletin 1936: 470 (1936);

J.B. Gillett: 133 (1958c). Type: Zimbabwe, Hwange, 4-1932, Levy 36

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

4. Microcharis aphylla (Viguier) Schrire, Du Puy &

Labat, comb, et stat. nov.

Indigofera phyllogramme Viguier var. aphylla Viguier in Notulae

Systematicae 13: 369 (1949) p.p., lecto. inch; J.B. Gillett: 132 (1958c).

Type: Madagascar, Perrier 13845 (P, lecto.!; K!, MO, P!). Lectotype

selected here.

5. Microcharis asparagoides (Taub.) Schrire, comb.

nov.

Indigofera asparagoides Taub. in Engl., Die Pflanzenwelt Ost-Afrikas

und der Nachbargebiete. C: 210 (1895); Bak. f. : 125 (1926); Cronq.:

172 (1954); J.B. Gillett: 176 (1958b); J.B. Gillett: 135 (1958c); J.B. Gillett:

326, t.47 (1971). Types: Tanzania, Bukoba, Stuhlmann 3989, 4101 (B,

syn. f); Ihangiro, Stuhlmann 3353 (B, syn. t).

6. Microcharis brevistaminea (J.B. Gillett) Schrire,

comb. nov.

Indigofera brexistaminea J.B. Gillett in Kew Bulletin, Additional Series

1: 130 (1958c). Type: Zaire, NE of Lake Moero, Niembe River, 27-5-1908,

Kassner 3009 (K, holo.!; BM).

7. Microcharis buchneri (Taub.) Schrire, comb. nov.

Indigofera buchneri Taub. in Botanische Jahrbucher 23: 180 (1896);

Bak. f.: 105 (1926); J.B. Gillett: 131 (1958c); Torre: 138 (1962). Type:

Angola, Malanje, Buchner 644 (B, holo. f; BM, sketch!).

8. Microcharis butayei (De Wild.) Schrire, comb.

nov.

Indigofera butayei De Wild, in Annales du Musee du Congo Beige,

Botanique Serie 5, 1: 132 (1904); J.B. Gillett: 135 (1958c); J.B. Gillett:

325 (1971); Thulin: 100 (1983); Thulin: 138 (1990). Type: Zaire, Lower

Congo, N'Lemfu, Butaye, coll. Gillet 1208 (BR, lecto.!). Lectotype

selected here.

166

Bothalia 22,2 (1992)

I. medicaginea sensu Bak. f.: 120 (1926); Cronq.: 171 (1954) p.p.,

non Welw. ex Bak.

9. Microcharis cana (Thulin) Schrire, comb. nov.

Indigofera cana Thulin in Nordic Journal of Botany 2: 48 (1982);

Thulin: 99 (1983); Thulin: 138 (1990). Type: Ethiopia, Wello Region,

below Bati on Assab road, 18-9-1962, Mooney 9620 ( ETH, holo. ; UPS).

10. Microcharis contorta (7.5. Gillett) Schrire, comb.

nov.

Indigofera contorta J.B. Gillett in Kew Bulletin, Additional Series 1:

133 (1958c). Type: South Yemen, Riyan Airport, Guichard KG/HAD/349

(BM, holo.!).

11. Microcharis cufodontii (Chiov.) Schrire, comb.

nov.

Indigofera cufodontii Chiov. in Cufodontis, Missione biologica nel paese

dei Borana: 71, t.12 (1939); J.B. Gillett: 134 (1958c); J.B. Gillett: 324

(1971); Thulin: 100 (1983); Thulin: 138 (1990). Types: Ethiopia, Sidamo

Region, Yavello, Cufodontis 473, 613 (FI, syn.).

12. Microcharis disjuncta (7.5. Gillett ) Schrire,

comb. nov.

var. disjuncta

Indigofera disjuncta J.B. Gillett var. disjuncta in Kew Bulletin 1955:

584 (1956); J.B. Gillett: 543 (1958a); J.B. Gillett: 132 (1958c); A. Schreib.:

55 (1970). I. acutifolia Schinz: 163 (1888) non Schlechtd. (1838). Type:

Namibia, Kuisib, 24-4-1885, H. Schinz 245 (Z, holo.!).

I. arenaria sensu Bak.: 79 (1871); Bak. f.: 192 (1903); Bak. f. : 108

(1926), non A. Rich.: 183 (1847) nec E. Mey. : 107 (1836).

I. semhaensis Vierh.: 362 (1907) p.p. quoad typum.

I. welwitschii Bak. var. simplicifolia Bak. f.: 253 (1932). Type:

Namibia, Karibib, Dinter 2523 (B, holo. t); Dinter 6971 (BM, neo.!;

K!). Neotypified by J.B. Gillett: 585 (1956).

var. failax (7.5. Gillett) Schrire, comb. nov.

Indigofera disjuncta J.B. Gillett var. failax J.B. Gillett in Kew Bulletin

1955: 585 (1956); J.B. Gillett: 132 (1958c). Type: Socotra, Bayley Balfour

674 (K, holo.!).

I. leptocarpa Hochst. & Steud. ex Balf. f.: 510 (1882) p.p., non Eckl.

& Zeyh.: 236 (1836).

13. Microcharis ephemera (J.B. Gillett) Schrire,

comb, et stat. nov. Type; Tanzania, 6.5 km from Mbeya

on Tukuyu Rd, 1 680 m, fl. 12-5-1956, Milne-Redhead &

Taylor 10197 (K, holo.!; EA, B, BR, LISC).

Indigofera asparagoides Taub. subsp. ephemera J.B. Gillett in Kew

Bulletin 13: 176 (1958b); J.B. Gillett: 328 (1971).

14. Microcharis galpinii N.E.Br. in Kew Bulletin

1897: 258 (1897); N.E. Br.: 375 (1932). Type: South Africa,

Transvaal, nr Barberton, summit of Saddleback Mtn,

± 1 500 m, 8-3-1891, Galpin 1315 (K, holo.!; GRA!,

PRE!).

Microcharis pseudo-indigofera Merxm.: 22 (1951). Indigofera pseudo-

indigofera (Merxm.) J.B. Gillett: 132 (1958c). Type: Zimbabwe,

Marondera Dist., Dehn 627 (M, holo.; SRGH).

Indigofera graminea Schltr. , nom. nud.

15. Microcharis garissaensis (7.5. Gillett) Schrire,

comb. nov.

Indigofera garissaensis J.B. Gillett in Kew Bulletin 24: 502 (1970);

J.B. Gillett: 324 (1971). Type: Kenya, near Tana River, 16 km SE of

Garissa, 180 m, 1-1961, Lucas 18 (K, holo.!; EA).

16. Microcharis gyrata ( Thulin ) Schrire, comb.

nov.

Indigofera gyrata Thulin in Nordic Journal of Botany 2: 45, 46 (1982);

Thulin: 138 (1990). Type: Kenya, Mandera Dist., 4 km on the Mandera-

Ramu road, 3-5-1978, Gilbert & Thulin 1372 (UPS, holo.; BR, C, EA,

K!, KRA, MO, WAG).

17. Microcharis karinensis (Thulin) Schrire, comb.

nov.

Indigofera karinensis Thulin in Nordic Journal of Botany 8: 469 (1989).

Type: Somalia, Bari Region, 15 km on the El Gal-Qandala road,

23-11-1986, Thulin & Warfa 6003 (UPS, holo.; K!, MOG).

18. Microcharis kucharii (Thulin) Schrire, comb.

nov.

Indigofera kucharii Thulin in Nordic Journal of Botany 8: 471 (1989).

Type: Somalia, Hiran Region, Bulo Burti Dist., 12-12-1987, Kuchar 17609

(UPS, holo.; K!, MOG).

19. Microcharis latifolia Benth. in Transactions of

the Linnean Society 25: 298, t. 33 B (1865b); Taub. in

Engl.: 213 (1895); Bak. f.: 256 (1926). Type: Mozambique,

Luabo, mouth of Zambezi, Kirk s.n. (K, holo.!).

Indigofera lobata J.B. Gillett: 131 (1958c); J.B. Gillett: 322 (1971).

J.B. Gillett published this nom. nov. because of Indigofera latifolia

Micheli.

20. Microcharis longicalyx (7.5. Gillett) Schrire,

comb. nov.

Indigofera longicalyx J.B. Gillett in Kew Bulletin 1955: 583 (1956);

J.B. Gillett: 543 (1958a); J.B. Gillett: 130 (1958c). Type: Nigeria, Jos

(or Bauchi) plateau, without exact locality, plains, 8-1930, Lely P.643

(K, holo.!).

I. lateritia Chev. : 174 (1920) non Willd.: 1233 (1802) nec Bertol.:

5 (1832).

Microcharis tenella sensu Bak. f.: 256 p.p.; Hutch. & Dalz.: 388 (1927)

p.p., non Benth.

21. Microcharis medicaginea (Welw. ex Bak.)

Schrire, comb. nov.

Indigofera medicaginea Welw. ex Bak. in Oliver, Flora of tropical Africa

2 : 86 (1871); Hiern: 210 (1896); Harms: 578 (1915); sensu Bak. f.: 120

(1926) and sensu Cronq.: 171 (1954) p.p., quoad typum solum; J.B. Gillett:

135 (1958c); Torre: 139 (1962) non A. Schreib.: 293 (1957). Type: Angola,

Pungo Andongo, by the stream of Catete, 5-1857, Welwitsch 2026 (BM,

lecto.!; K!, LISU). Type selected by Torre (1962).

22. Microcharis microcharoides (Taub.) Schrire,

comb. nov.

Indigofera microcharoides Taub. in Engl., Die Pflanzenwelt Ost-Afrikas

und der Nachbargebiete C: 209 (1895); Bak. f. : 106 (1926); J.B. Gillett:

131 (1958c); J.B. Gillett: 323 (1971). Type: Tanzania or Kenya, ‘? Masai

steppe’, Fischer 288 (B, holo. t; BM, sketch!); Tanzania, Pare Dist.,

Haarer 1355 (K, neo.!; EA). J.B. Gillett (1958c) neotypified this species

but the neotype was not cited in his revision for the Flora of tropical

East Africa (1971).

var. microcharoides

J.B. Gillett in Kew Bulletin 24: 502 (1970); J.B. Gillett:

324 (1971).

Bothalia 22,2 (1992)

167

Rhynchotropis curtisiae Johnst.: 23 (1925); Bak. f.: 167 (1926). Type:

Kenya, Mau range SW of Kijabe, Curtis 814 (GH, holo.).

var. latestipulata (J.B. Gillett) Schrire, comb. nov.

Indigofera microcharoides Taub. var. latestipulata J.B. Gillett in Kew

Bulletin 24: 502 (1970); J.B. Gillett: 324 (1971). Type: Kenya, between

Umba and Mwena Rivers on Lungalunga-Msambweni road, 100 m,

14-8-1953, Drummond & Hemsley 3784 (K, holo.; EA).

23. Microcharis phyllogramme (Viguier) Schrire,

Du Puy & Labat, comb. nov.

Indigofera phyllogramme Viguier in Notulae Systematicae 13: 369

(1949); J.B. Gillett: 132 (1958c). Type: Madagascar, Perrier 4876 (P, lec-

to. ! ; K!, P!). Lectotype selected here.

24. Microcharis praetermissa (Bak. f.) Schrire ,

comb. nov.

Indigofera praetermissa Bak. f.. The Leguminosae of tropical Africa:

121 (1926); J.B. Gillett: 134 (1958c). Type: Zaire, Lake Mweru, Kassner

2809 (K, holo.!).

I. welwitschii var. remotiflora sensu Cronq.: 171 (1954) p.p. non Bak.

25. Microcharis remotiflora (Taub. ex Bak. f)

Schrire, comb. nov.

Indigofera remotiflora Taub. ex Bak. f. , The Leguminosae of tropical

Africa 1: 121 (1926) p.p. excl. var. angolensis Bak. f. , loc. cit. Type:

Cameroon, Staudt 486 (B, holo. t); Zenker 486 (BM, neo.!). Neotypified

by J.B. Gillett (1958c). The identity of var. angolensis Bak. f. is uncertain

but it is not conspecific with I. remotiflora.

I. welwitschii Bak. var. remotiflora (Taub. ex Bak. f.) Cronq.: 172

(1954); J.B. Gillett: 543 (1958a); J.B. Gillett: 134 (1958c); Torre: 139

(1962); Binns: 82 (1968); J.B. Gillett: 326 (1971).

26. Microcharis sessilis (Thulin) Schrire, comb. nov.

Indigofera sessilis Thulin in Nordic Journal of Botany 8: 475 (1989).

Type: Somalia, Bari Region, 8 km NE of Gorgore, 26-11-1985, Thulin

& Warfa 5561 (UPS, holo.; E, FT, K!, MOG).

27. Microcharis spathulata (J.B. Gillett) Schrire,

comb. nov.

Indigofera spathulata J.B. Gillett in Kew Bulletin, Additional Series

1: 129 (1958c); J.B. Gillett: 322 (1971). Type: Zambia, Mweru-Luapula

Dist., near Mbereshi, Walter 7 (K, holo.!).

28. Microcharis stipulosa (Chiov.) Schrire, comb.

nov.

Indigofera stipulosa Chiov. in Cufodontis, Missione biologica nel paese

dei Borana 75: t.13 (1939); J.B. Gillett: 132 (1958c); Thulin: 99 (1983);

Thulin: 137 (1990). Type: Ethiopia, Sidamo Region, Neghelli, Cufodontis

228 (FT, holo.).

29. Microcharis tenella Benth. in Transactions of the

Linnean Society 25 : 297, t. 33A (1865b); Bak.: 132 (1871);

Bak. f.: 256 (1929) p.p.. Type: Nigeria, Jeba on the

Kworra, Barter s.n. (K, lecto.!). Lectotype selected by J.B.

Gillett: 584 (1956).

Indigofera hutchinsoniana J.B. Gillett: 584 (1956); J.B. Gillett: 543

(1958a); J.B. Gillett: 132 (1958c). J.B. Gillett published this nom. nov.

because of Indigofera tenella Vahl ex DC.

I. welwitschii sensu Hutch. & Dalz.: 393 (1927) non Bak.

30. Microcharis tenuirostris (Thulin) Schrire, comb.

Indigofera tenuirostris Thulin in Nordic Journal of Botany 8: 476 (1989).

Type: Somalia, Nugal Region, 11 km from Garoe along road to Halin,

8-2-1984, Warfa 1000 (UPS, holo.).

31. Microcharis tisserantii (Pellegr.) Schrire, comb.

nov.

Indigofera asparagoides Taub. var. tisserantii Pellegr. in Bulletin Societe

de Botanique de France 72: 538 (1925). Type: Chad [Oubangui], Waka

Region, Bayedou rocks, Tisserant 1650 (P, holo.!).

I. tisserantii (Pellegr.) Pellegr.: 6 (1947); Cronq.: 173 (1954); J.B.

Gillett: 136 (1958c).

32. Microcharis tritoides (Bak.) Schrire, comb. nov.

Indigofera tritoides Bak. in Kew Bulletin 1895: 214 (1895); Bak. f. :

108 (1926). Type: Somalia, Golis Range, near Dara-as, Miss E. Cole

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

subsp. tritoides

Indigofera tritoides Bak. var. mtoides

J. B. Gillett in Kew Bulletin, Additional Series 1: 133 (1958c); Thulin:

100 (1983); Thulin: 138 (1990).

I. leptocarpa Hochst. & Steud. ex Balf. f.: 510 (1882) p.p., non Eckl.

& Zeyh.: 236 (1836). Type: Arabia, 1837, Schimper 771 (K, isosyn.!).

/. tenuisiliqua Schweinf. : 241 (1896); Bak. f.: 109 (1926). Type: Sudan,

1868, Schweinfurth 297 (K, isosyn.!).

I. spinosa Forssk. var. spiniflora sensu Schweinf.: 237 (1896) p.p.

subsp. obbiadensis (Chiov.) Schrire, comb, et stat.

nov.

I. sesquijuga Chiov. var. obbiadensis Chiov.: 139 (1929). Type: Somalia,

Obbia, in dunes, Puccioni & Stefanini 387 (439) (FI, holo.).

Indigofera tritoides Bak. var. obbiadensis (Chiov.) J.B. Gillett in Kew

Bulletin, Additional Series 1: 133 (1958c).

33. Microcharis wajirensis (J.B. Gillett) Schrire,

comb. nov.

Indigofera wajirensis J.B. Gillett in Kew Bulletin, Additional Series

1: 135 (1958c); J.B. Gillett: 325 (1971). Type: Kenya, Wajir Region,

27-5-1952, J.B. Gillett 13368 (K, holo.!; EA).

34. Microcharis welwitschii (Bak.) Schrire, comb.

nov.

Indigofera welwitschii Bak. in Oliver, Flora of tropical Africa 2: 84

(1871); Hiern: 209 (1896); Harms: 577 (1915); Bak. f.: 122 (1926); Bak.

f.: 103 (1928); Cronq.: 171 (1954), pro maj. parte; J.B. Gillett: 543 (1958a);

J.B. Gillett: 134 (1958c); Torre: 138 (1962); J.B. Gillett: 326 (1971); Thulin:

100 (1983); Thulin: 138 (1990). Anila welwitschii (Bak.) Kuntze: 940

(1891). Type: Angola, Pungo Andongo, Pedras de Cabondo, fl. & fr.

4-1857, Welwitsch 2066 (BM, lecto.!; COL K, LISU, P). Lectotype select-

ed here.

I. variabilis De Wild.: 134 (1904) non N.E. Br. (1903) nom. illegit.

Type: Zaire, Lower Congo, Gillet 734 (BR, holo.).

INDIGASTRUM Jaub. & Spach

Indigastrum Jaub. & Spach, Illustrationes Plantarum

Orientalium 5: 101, t. 492 (1857). Type species: Indigas-

trum deflexum (Hochst. ex A. Rich.) Jaub. & Spach

(= Indigastrum parviflorum [Heyne ex Wight & Arn.]

Schrire).

Indigofera subgenus Euindigofera Benth. in Benth. &

Hook.f.: 494 (1865).

nov.

168

Indigofera ‘group’ Indigastrum (Jaub. & Spach) Bak.

f.: 161 (1926); Cronq.: 124 (1954) p.p. excl. spp. 65-68.

Indigofera subgenus Indigastrum (Jaub. & Spach) J.B.

Gillett: 123 (1958c); Torre: 136 (1962); J.B. Gillett: 320

(1971).

Indigofera sensu Hutch.: 400 (1964); Polhill: 291 (1981);

De Kort & Thijsse: 104 (1984); Lock: 289 (1989); Thu-

lin: 119 (1990).

1. Indigastrum argyraeum (Eckl. & Zeyh.) Schrire ,

comb. nov.

Indigofera argyraea Eckl. & Zeyh., Enumeratio plantarum: 239 (1836);

Harv.: 201 (1862); A. Schreib. : 50 (1970). Am la argyraea (Eckl. & Zeyh.)

Kuntze: 50 (1898). Type: Cape, in locis saxosis (altit. Ill) laterum montium

prope Shiloh ad flumen Klipplaat R. turn ad Swart Kei R., Jan. -Mar.,

Ecklon & Zeyher 1595 (S, lecto.!). Lectotype selected here.

I. collina Eckl. & Zeyh.: 239 (1836). Type: Cape, in petrosis (altit.

II) collium continuorum inter flumina Koonap & Kat R., fr. Jul., Ecklon

& Zeyher 1596 (S, lecto.!). Lectotype selected here.

I. burchellii E. Mey. var. paucifolia E. Mey. : 106 (1836) non DC. Type:

Cape, prope Zwartbulletjie in collibus, alt. 2500' (II, d), 6-3-1827, Drege

6689 (P, lecto.!). Lectotype selected here.

I. burchellii E. Mey. var. multifolia E. Mey.: 106 (1836) non DC. Type:

Cape, in rupestribus siccis ad Groot Vis R., alt. 300' (V, a), 20-6-1832,

Drege 3802 (P, lecto.!). Lectotype selected here.

I. medicaginea sensu A. Schreib.: 293 (1957) p.p., non Welw. ex Bak.

/. pseudoaltemans Dinter, nom. nud.

2. Indigastrum argyroides (E. Mey. ) Schrire, comb.

nov.

Indigofera argyroides E. Mey., Commentariorum de plantis Africae

australioris: 106 (1836); Harv.: 191 (1862); J.B. Gillett: 126 (1958c);

A. Schreib.: 51 (1970). Type: Cape, ad ripas limosas fluvii Gariep, 200'

(III, B), 16-9-1830, Drege 3313 (P, lecto.!). Lectotype selected here.

/. saxicola Engl.: 28 (1888) non F. Muell. (1864). I. engleri Bak. f.:

194 (1903); Bak. f. : 105 (1926). Type: Namibia, lower Swakop River,

250 m, Marloth 1209 (SAM, lecto.!).

I. medicaginea sensu A. Schreib.: 293 (1957) p.p., non Welw. ex Bak.

3. Indigastrum burkeanum (Benth. ex Harv.)

Schrire, comb. nov.

Indigofera burkeana Benth. ex Harv. in Harv. & Sond., Flora capensis

2: 197 (1862); N.E. Br.: 373 (1932); Dyer: t. 939 (1944); J.B. Gillett:

125 (1958c) p.p., excl. specimens from Cape, Botswana and Namibia.

Type: Transvaal, Magaliesberg, Dec., Zeyher 476 (K, holo.!; S, TCD).

4. Indigastrum candidissimum (Dinter) Schrire,

comb. nov.

Indigofera candidissima Dinter in Feddes Repertorium 18: 428 (1922);

Dinter: 198 (1932); A. Schreib.: 52 (1970). Type: Namibia, Kegelberg,

4-4-1911, Dinter 2107 (B, t).

5. Indigastrum costatum (Guill. & Perr.) Schrire,

comb. nov.

Indigofera costata Guill. & Perr., Florae Senegambia tentamen: 187

(1832); Bak.: 95 (1871); Bak.f. : 163 (1926). Type: Senegal, near mouth

of Senegal, Ghiandoum, Perrottet s.n. (P, holo.!; BM).

subsp. costatum

Indigofera costata Guill. & Perr. subsp. costata

J.B. Gillett in Kew Bulletin, Additional Series 1: 125 (1958c); J.B.

Gillett: 543 (1958a); J.B. Gillett: 320 (1971); Thulin: 99 (1983); Thulin:

137 (1990).

Bothalia 22,2 (1992)

subsp. theuschii (O. Hoffm.) Schrire, comb. nov.

Indigofera costata Guill. & Perr. subsp. theuschii (O. Hoffm.) J.B.

Gillett in Kew Bulletin, Additional Series 1: 125 (1958c); Torre: 136

(1962). /. theuschii O. Hoffm.: 126(1880-82); Bak. f.: 162 (1926); Bak.

f. : 105 (1928) p.p.; A. Schreib.: 294 (1957). Type: Angola, Pungo

Andongo, Theusch & Mechow 128 (B, t; BM, photo.!)

I. affinis Harv.: 184 (1862) non De Wild.: 144 (1923). Type: Botswana,

Lake Ngami, J. McCabe s.n. (K, holo.!).

/. heptaphylla Hiern: 209 (1896); Eyles: 373 (1916). Type: Angola,

Pungo Andongo, near de Quibinda, Quitage, Welwitsch 4143 (BM, lecto.!;

LISU). Selected by Torre: 136 (1962).

I. goniodes Hochst. ex Bak. var. damarensis Bak. f.: 163 (1926). Type:

Namibia, Een s.n. (BM, holo.!).

/. burkeana sensu J.B. Gillett: 125 (1958c) p.p. , excl. specimens from

Transvaal; A. Schreib.: 52 (1970), non Benth. ex Harv.

subsp. goniodes (Hochst. ex Bak.) Schrire, comb. nov.

Indigofera costata Guill. & Perr. subsp. goniodes J.B. Gillett in Kew

Bulletin, Additional Series 1: 125 (1958c); J.B. Gillett: 320 (1971); Thulin:

99 (1983); Thulin: 137 (1990). I. goniodes Hochst. ex Bak.: 85 (1871).

Type: Ethiopia, Tigray region, Dscha Dscha, 1854, Schimper 2242 (K,

holo.!; P!).

Indigastrum macrostachyum Jaub. & Spach: t. 493 (1857). Type:

Ethiopia, Tigray region, Dscha Dscha, 1853, Schimper 1305 (P, holo.!).

subsp. macrum (E. Mey.) Schrire, comb. nov.

Indigofera costata Guill. & Perr. subsp. macra (E. Mey.) J.B. Gillett

in Kew Bulletin, Additional Series 1: 125 (1958c); A. Schreib.: 53 (1970).

I. macra E. Mey.: 105 (1836); Harv.: 197 (1862); N.E. Br.: 373 (1932).

Anila macra (E. Mey.) Kuntze: 939 (1891). Type: Cape, ad ripas fluvii

Bashee, 500' (V, b), 26-1-1832, Drege 5419 (P, lecto.!). Lectotype selected

here.

I. macra E. Mey. var. aequalis N.E. Br. : 44 (1926). Type: Transvaal,

Zoutpansberg, Messina, Moss & Rogers 131 (K, holo.!).

I. goniodes sensu Eyles: 373 (1916) non Hochst. ex Bak.

I. goniodes Hochst. ex Bak. var. rhodesica Bak. f. : 76 (1920). Types:

Zimbabwe, Bulawayo, F.A. Rogers 13666 (BM, syn.!), Gardner 62 (K,

syn.!).

6. Indigastrum fastigiatum (E. Mey. ) Schrire, comb.

nov.

Indigofera fastigiata E. Mey., Commentariorum de plantis Africae

australioris: 102 (1836); Harv.: 183 (1862); N.E. Br. : T73 (1932); J.B.

Gillett: 124 (1958c). Type: Cape, inter Umzimvubu et Umsikaba, 1200'

(V, b), 2-1832, Drege 5433 (P, lecto.!). Lectotype selected here.

/. fastigiata E. Mey. var. angustata Harv.: 183 (1862). Type: Cape,

Swartkops River, Zeyher 2442 (S, lecto.!). Lectotype selected here.

I. rostrata Conrath: 223 (1908) non H. BoL: 23 (1896). Type: Trans-

vaal, Modderfontein, 1898, P. Conrath 1196 (K, holo.!).

7. Indigastrum guerranum (Torre) Schrire, comb.

nov.

Indigofera guerrana Torre in Memorias da Junta investigates do

Ultramar, ser. 2, 19: 61, t. 39 (1960); A. Schreib.: 57 (1970). Type: Angola,

Mossamedes, entre o apeadeiro do C.F. Dois Iramaos e Virei, 9-5-1957,

B. Teixeira 2249 (LISC, holo.!; COI, LUA).

8. Indigastrum parviflorum (Heyne ex Wight &

Arn.) Schrire, comb. nov.

Indigofera parviflora Heyne ex Wight & Arn., Prodromus florae

peninsulae Indiae orientalis; 201 (1834); Harv.: 198 (1862); Bak.: 83 (1871);

Fries: 78 (1914); Bak. f.: 162 (1926); N.E.Br.: 367 (1932); J.B. Gillett:

126 (1958c); Cronq.: 170 (1954); A. Schreib.: 60 (1970); J.B. Gillett: 321

(1971); Thulin: 99 (1983); Thulin: 137 (1990). Anila parviflora (Heyne

ex Wight & Arn.) Kuntze: 939 (1871). Type: India, in herbarium Wallich

no. 5457, Heyne s.n. (K, holo.!).

Bothalia 22,2 (1992)

169

subsp. parviflorum

var. parviflorum

Indigofera parviflora Heyne ex Wight & Am. var. parviflora in J.B.

Gillett in Kew Bulletin 1955: 582 (1956); J.B. Gillett: 126 (1958c).

/. deflexa Hochst. ex A. Rich.: 178 (1847). Indigastrum deflexum

(Hochst. ex A. Rich.) Jaub. & Spach: t. 492 (1857). Type: Ethiopia,

Tacaze River near Dschelad-scheranne, fl. 14-9-1840, Schimper 1467 (P,

lecto.!; G, K!, S! , UPS). Lectotype selected here.

Indigofera barcensis Chiov. : 36 (1932). Type: Eritrea, De Benedictis

200 (FT, holo.).

var. crispidulum (J.B. Gillett) Schrire, comb. nov.

Indigofera parviflc. a Heyne ex Wight & Am. var. crispidula J.B. Gillett

in Kew Bulletin, Additional Series 1: 126 (1958c). Type: Zimbabwe, Mato-

bo Dist., ± 1 350 m, fl. 11-3-1947, West 2252 (K, holo.!; SRGH).

subsp. occidentalis (J.B. Gillett) Schrire, comb, et

stat. nov.

Indigofera parviflora Heyne ex Wight & Arn. var. occidentalis J.B.

Gillett in Kew Bulletin 1955: 582 (1956); J.B. Gillett: 543 (1958a); J.B.

Gillett: 178 (1958c); Torre: 137 (1962). Type: Senegal, Leprieur s.n. (P,

holo.; BM!).

REFERENCES

BAKER, E.G. 1903. The Indigoferas of tropical Africa. Journal of Botany

41: 185-194, 234-245, 260-267, 323-334.

BAKER, E.G. 1920. Leguminosae. Journal of Botany 58 : 74 —76.

BAKER, E.G. 1926. The Leguminosae of tropical Africa 1: 95—166. Gent.

BAKER, E.G. 1928. Leguminosae. Journal of Botany 66, Suppl. 1:

98-159.

BAKER, E.G. 1929. The Leguminosae of tropical Africa 2: 256. Gent.

BAKER, E.G. 1932. New African species of Leguminosae. Journal of

Botany 70 : 251-255.

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

2: 65—132. Reeve, London.

BAKER, J.G. 1895. Indigofera tritoides. Kew Bulletin 1895: 214.

BALFOUR, I.B. 1882. Indigofera leptocarpa. Proceedings of the Royal

Society of Edinburgh 11: 510.

BENTHAM, G. 1865a. Leguminosae. In G. Bentham & J.D. Hooker,

Genera plantarum 1: 494, 501. Reeve, London.

BENTHAM, G. 1865b. Description of some new genera and species

of tropical Leguminosae. Transactions of the Linnean Society,

London 25 : 297- 320.

BERTOLONI, A. 1832. Dissertatio de quibusdam novis plantarum specie-

bus et de Bysso antiquorum: 5. Bononia.

BINNS, B. 1968. A first checklist of the herbaceous flora of Malawi:

81, 82. Government Printer, Zomba.

BOLUS, H. 1896. Contributions to the flora of South Africa. Journal

of Botany 34: 16-25.

BROWN, N.E. 1897. Microcharis galpinii. Kew Bulletin 1897: 258.

BROWN, N.E. 1926. Indigofera. In J. Burtt Davy, A manual of flowering

plants and ferns of the Transvaal with Swaziland 1: 44.

BROWN, N.E. 1932. Indigofera. In J. Burtt Davy, A manual of flowering

plants and ferns of the Transvaal with Swaziland 2 : 360 — 373.

Longmans, London.

CHEVALIER, A.J.B. 1920. Exploration botanique de l Afrique occiden-

tal franchise 1: 174. Paris.

CHIOVENDA, E. 1929. Flora of Somalia 1: 139. Rome.

CHIOVENDA, E. 1932. Indigofera barcensis. Atti della Societa dei

naturalisti di Modena 63: 36.

CHIOVENDA, E. 1939. In Cufodontis, Missione biologica nel paese

dei Borana: 71, t. 12.

CONRATH, P. 1908. Indigofera rostrata. Kew Bulletin 1908: 223.

CRONQUIST, A. 1954. Papilionaceae. V. Galegeae. Flora du Congo

Beige et du Ruanda-Urundi 5 : 72—175.

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

(Leguminosae-Papilionoideae) in southeast Asia. Blumea 30:

89-151.

DE WILDEMAN, E. 1904. Indigofera butayei. Annales du Musee du

Congo Beige, Botanique Ser. 5,1: 134.

DE WILDEMAN, E. 1923. Indigofera affinis. In E. Baker & E. de

Wildeman, Sur quelques Indigofera du Congo beige. Bulletin du

Jardin Botanique de I’Etat a Bruxelles 8: 143-159.

DINTER, K. 1922. Index der aus Deutsch-Sudwestafrika bis zum Jahre

1917 bekannt gewordenen Pflanzenarten XII. Feddes Repertorium

18: 423-444.

DINTER, K. 1932. Diagnosen neuer siidwestafrikanischer Pflanzen.

Feddes Repertorium 30: 180—205.

DYER, R. A. 1944. Indigofera burkeana. The Flowering Plants of Africa

24: t. 939.

ECKLON, C.F. & ZEYHER, C. 1836 (Jan.). Enumeratio plantarum

africae australis extratropicae 2: 232—245. Hamburg.

ENGLER, H.G.A. 1888. Plantae Marlothianae. Botanische Jahrbiicher

10,1: 1-50.

ENGLER, H.G.A. 1895. Die Pflanzenwelt Ost-Afrikas und der Nach-

bargebiete. Theil C: 209—211. Berlin.

EYLES, F. 1916. A record of plants collected in southern Rhodesia.

Transactions of the Royal Society of South Africa 5 : 372 — 375.

FRIES, R.E. 1914. Botanische Untersuchungen; wissenschaftliche

Ergebnisse der Schwedischen Kongo-Rhodesia-Expedition: 78.

Stockholm.

GILLETT, J.B. 1956. Indigofera. New species, varieties and names from

West tropical Africa. Kew Bulletin 1955 : 573 —585.

GILLETT, J.B. 1958a. Indigofera. In R.W.J. Keay, Flora of west tropical

Africa, edn 2, 1: 533—543.

GILLETT, J.B. 1958b. Indigofera asparagoides. Kew Bulletin 13: 176.

GILLETT, J.B. 1958c. Indigofera (Microcharis) in tropical Africa with

the related genera Cyamopsis and Rhynchotropis. Kew Bulletin,

Additional Series 1: 1—166.

GILLETT, J.B. 1970. Additions to our knowledge of Indigofera L. in

East tropical Africa. Kew Bulletin 24, 3: 465—506.

GILLETT, J.B. 1971. Indigofereae. In E. Milne-Redhead & R.M. Polhill,

Flora of tropical East Africa Leguminosae 3, Papilionoideae:

321—328. Crown Agents.

GUILLEMIN, A., PERROTTET, S. & RICHARD, A. 1832. Florae

Senegambia tentamen: 172—189.

HARMS, H. 1915. Indigofera. In N.G.A. Engler, Die Pflanzenwelt Afrikas

3,1: 577, 578.

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

Flora capensis 2: 163—203. Hodges, Smith, Dublin.

HIERN, W.P. 1896. Catalogue of the African plants collected by Dr

Friedrich Welwitsch in 1853-1861 , 1: 206—218. London.

HOFFMANN, O. 1880—1882. Indigofera theuschii. Linnaea 43: 126.

HUTCHINSON, J. & DALZIEL, J.M. 1927. Flora of west tropical Africa

1, edn 1. Crown Agents, London.

HUTCHINSON, J. 1964. Papilionaceae. In Genera of flowering plants

Vol. 1: 400. Oxford.

JAUBERT, H.-F. & SPACH, E. 1857. Ulustrationes plantarum orientalium

5: 101 t. 481, 482 and t. 492, 493 (1857). Paris.

JOHNSTON, I.M. 1925. Further new plants collected by Mrs. Richard

C. Curtis in tropical Africa. Contributions from the Gray

Herbarium of Harvard University, N.S. 75: 23.

KUNTZE, C.E.O. 1891—1898. Revisio generum plantarum 1: 159, 2:

938-940, 3,2: 50-52. Leipzig.

LOCK, J.M. 1989. Legumes of Africa: a checklist: 289—332. Royal

Botanic Gardens, Kew, London.

MERXMULLER, H. 1951. Microcharis pseudo-indigofera. Proceedings

and Transactions of the Rhodesia Scientific Association 43: 22.

MEYER, E.H.F. 1836 (Feb.). Commentariorum de plantis Africae

australioris: 92—108. Leipzig.

MILNE-REDHEAD, E.W.B.H. 1936. Tropical African plants. Kew

Bulletin 1936: 470.

PELLEGRIN, F. 1925. Indigofera asparagoides Taub. var. tisserantii.

Bulletin Societe botanique de France 72 : 53 8.

PELLEGRIN, F. 1947. Indigofera tisserantii. Bulletin Societe de bota-

nique de France 94: 6.

POLHILL, R.M. 1981. Indigofereae. In R.M. Polhill & PH. Raven,

Advances in legume systematics 1: 289—291. Royal Botanic

Gardens, Kew.

RICHARD, A. 1847. Tentamen florae Abyssinicae 1: 178—185. Paris.

SCHINZ, H. 1888. Beitrage zur Kenntnis der Flora von Deutsch-Siidwest-

Afrika und der angrenzenden Gebiete II. Abhandlungen des

Botanischen Vereins der Provinz Brandenburg 30: 138—186.

SCHREIBER, A. 1957. Beitrage zur Kenntnis der Leguminosen Stid-

westafrikas. Mitteilungen der Botanischen Staatssammlung

Munchen 16: 283-299.

SCHREIBER, A. 1970. Fabaceae. In H. Merxmiiller, Prodromus einer

Flora von Siidwestafrika 60: 38—63. Cramer, Lehre.

SCHRIRE, B.D. 1991. Systematic studies in African Indigofereae

( Leguminosae-Papilionoideae ) Ph.D. thesis. University of Natal,

South Africa.

SCHWEINFURTH, G.A. 1896. Indigofera. Bulletin de I'Herbier Boissier

4, app. 2: 237, 241.

170

Bothalia 22,2 (1992)

TAUBERT, P. 1894. Leguminosae. In A. Engler & K. Prantl, Die

natiirlichen Pflanzenfamilien edn 1, 3,3: 184—384. Berlin.

TAUBERT, P. 1896. Leguminosae Africanae 1. Botanische Jahrbiicher

23: 172-196.

THULIN, M.L. 1982. New and noteworthy species of lndigofera

(Leguminosae) from NE Africa. Nordic Journal of Botany 2:

41-50.

THULIN, M.L. 1983. Leguminosae of Ethiopia. Opera Botanica 68:

82-100. Copenhagen.

THULIN, M.L. 1989. lndigofera tenuirostris. Nordic Journal of Botany

8: 469.

THULIN, M.L. 1990. In O. Hedberg & Edwards, Flora of Ethiopia 3:

119-138.

TORRE, A.R. DA 1960. Taxa angolensia nova vel minus cognita 1.

Memdrias da Junta de Investigates do Ultramar, ser. 2, 19:

50-62.

TORRE, A.R. DA 1962. Papilionoideae. Conspectus florae angolensis

3: 84-140. Junta de Investigates do Ultramar, Lisbon.

VIERHAPPER, F. 1907. lndigofera semhaensis. Denkschriften der

Akademie der Wissenschaften 71: 362.

VIGUIER, R. 1949. Leguminosae madagascariensis novae. Notulae

systematicae 13: 333—369.

WIGHT, R. & ARNOTT, G.A.Q. 1834. Prodromus florae peninsulae

Indiae orientalis: 201. London.

WILLDENOW, C.L. 1802. Caroli a Linne species plantarum 3:

1220-1239. Berlin.

Bothalia 22,2: 171-175 (1992)

Studies in the Justicia and Siphonoglossa (Acanthaceae) species of

southern Africa: final conclusions

K.L. IMMELMAN*

Keywords: Acanthaceae, Justicia, Siphonoglossa, South Africa, taxonomy

ABSTRACT

On the basis of seed surface micromorphology, pollen and inflorescence type, as well as characters of the capsule, seven

sections can be distinguished in the southern African species of Justicia. The relationship of these to the sections recognised

by Graham is discussed. A key to the sections and a synopsis of their main distinguishing characters are presented. One

new section, sect. Minima Immelman, is described. A short note on the generic position of the Siphonoglossa species of

the region, following on from a previous paper, is also given.

UITTREKSEL

Op grand van saadoppervlak-mikromorfologie, stuifmeel en bloeiwysetipe, sowel as kenmerke van die kapsule, word sewe

seksies onder die Suider-Afrikaanse Justicia- spesies onderskei. Die verwantskap van die seksies met die van Graham word

bespreek. ’n Sleutel tot seksies en ’n oorsig van hul belangrikste onderskeidende kenmerke word gegee. Een nuwe seksie,

seksie Minima Immelman, word beskryf. ’n Kort nota oor die generiese posisie van die Siphonoglossa- spesies van die streek,

wat volg op 'n vorige artikel, word ook gegee.

INTRODUCTION

In the preceding four papers on the southern African

species of Justicia and Siphonoglossa, the various states

of indumentum, pollen and seed surface were described.

Their possible value both in identifying species and

delimiting sections within Justicia, was discussed. One

of the points made throughout was that none of these

organs could be used in isolation in the delimiting of

sections.

The species studied were chosen on the basis of their

geographical distribution rather than as being (putatively)

representative of the whole genus. As was stated by

Graham (1988), small sample size created difficulties in

the delimitation both of sections and of the genus Justicia.

She studied 58 species in detail, but concluded that ‘It is

over-optimistic to hope that all the remaining known

species of Justicia will fit neatly into the sectional

classification proposed here .... However, it is hoped that

this study will provide the basis for future work.’ The

present study of the southern African representatives of

Justicia and the closely related genus Siphonoglossa used

Graham’s work as a starting point for discussion, since

it gives a worldwide perspective on the genus. Graham’s

sectional treatment will be discussed in the light of results

obtained from the detailed study of the ± 30 southern

African taxa traditionally included in Justicia and Siphono-

glossa.

Some confusion surrounds the designation of a type

species in Justicia. A summary of the problems involved

was given by Steam (1971), and the problem is also dealt

with by Graham (1988). She considered the type species

to be J. hyssopifolia L., as did Hedren (1989), and this

decision is followed here.

* Genetic Resources Section, Grasslands Research Centre, Private Bag

X05, East Lynne, Pretoria 0039.

MS. received: 1992-01-23.

METHODS

All southern African taxa tradionally included in Justicia

and Siphonoglossa were studied, and sections elucidated.

The area termed ‘southern Africa’ is that defined by the

Flora of southern Africa, i.e. the area south of the Cunene

and Limpopo Rivers, including Botswana.

Dried material was examined from the following

herbaria: BOL, GRA, KMG, KNP, NBG, NH, NU, SAM,

STE and WIND. Living material was collected on trips

to the northern Transvaal, northern Natal and Zululand,

the eastern Cape and the north-central part of Namibia.

Some of this material was grown in the nursery of the

National Botanical Institute, Pretoria.

Seed testa surface, indumentum and pollen were

examined under the Scanning Electron Microscope.

Inflorescence structure was also examined. Various charac-

ters of leaves, bracts, calyx, corolla, capsules and habit

were taken into consideration in the conclusions reached.

Nomenclature of the sections was taken from Graham

(1988), and her sectional synonomy and type species are

not repeated here.

RESULTS AND DISCUSSION

In this series of studies, striking correlations between

inflorescence form, seed testa morphology and pollen were

found in the genus. Largely on this basis, the species

studied could be grouped into seven sections, one of them

new.

The primitive inflorescence is considered to probably

be a thyrse which has been reduced in various ways. Where

it has been reduced to sessile cymes/single flowers, these

are in some cases re-aggregated to form a more or less

dense, terminal synflorescence.

172

Bothalia 22,2 (1992)

The pollen in the genus is three- or two-colporate or

two-porate, with entire or areolate margocolpi. The sexine

is reticulate to lophate in all species, with a pseudocolpus

(colpoid streak) on either side of the colpus (Immelman

1989a). The area between the colpus and the pseudocolpus

(the margocolpus, see Immelman 1989a) is either entire

or divided into circular areolae. In the southern African

taxa, as well as in seven tropical African species examined

later, the inflorescence and pollen types are closely

correlated and valuable in dividing the genus into sections.

Seed surface, described simply as ‘rough’ in earlier

accounts of the genus, is one of the characters that is used

to distinguish Justicia from the related genus Monechma

Hochst. (Munday 1980). It is, however, very varied in

Justicia , and under the SEM reveals a number of useful

characters, a fact also noted by Graham (1988). In a

number of cases seed surface also correlates with the

sections, which lends weight to their naturalness. It is

considered unlikely that a number of characters, taken

from three different organs, viz. pollen, inflorescence and

seeds, would have evolved in parallel.

A number of other characters were considered, and

some were found to reinforce these divisions, especially

capsule texture. Capsule shape is mainly useful at generic

level. Munday (1980) considered a clavate capsule charac-

teristic of Monechma , as it was constant in the genus, being

correlated with the two-seeded condition. Some Justicia

species (also two-seeded) also have this type of capsule,

but in most cases it can be used to distinguish the

two genera. (It is possible that Monechma should be

considered congeneric with Justicia, as no one character

or combination of characters could be found to completely

separate the two genera, but this will not be discussed

here.)

The capsule of Justicia is usually cylindrical-clavate with

an apiculate apex and a short or long stipe. The stipe is

laterally flattened. The fertile portion contains four seeds,

sometimes reduced to two or one, each subtended by a

hook-shaped retinaculum. The texture of the capsule varies

from woody and thick-walled (sects. Raphidospora,

Justicia ) or papery and thin-walled (sects. Tyloglossa,

Minima), most species of sect. Hamieria and sect.

Ansellia. Intermediate between these two extremes are the

capsules of, for example, sect. Betonica and some species

of sect. Hamieria.

As stated in the introduction, the sections were proposed

on the basis of a study of southern African material only.

However, Justicia is probably a tropical genus, with

southern Africa being at the southern end of its distribu-

tion range in Africa and it was therefore considered

important to assess whether the apparent pattern seen in

the southern African species also existed north of the

Cunene and Limpopo Rivers. As a test, therefore, seven

tropical species (listed in Immelman 1989a) were selected

from a number of the sections, their probable section being

first assessed by examining the type of inflorescence

present. The pollen of each was then viewed under the

SEM and it was found that in every case it was of the pollen

type expected for the section in which they had provision-

ally been placed (Immelman 1989a). It was not, unfor-

tunately, possible to examine seeds of the species.

KEY TO SECTIONS IN SOUTHERN AFRICAN SPECIES

OF JUSTICIA

la Pollen three-colporate; inflorescence of pedunculate lax

axillary cymes, or sessile condensed cymes aggregated into

a terminal inflorescence:

2a Pollen with entire margocolpi:

3a Seeds one per capsule, smooth; inflorescence a simple

axillary cyme; flowers with a pouched throat

II sect. Justicia

3b Seeds 2—4 per capsule, rough; inflorescence a terminal or

axillary simple cyme or a compound axillary cyme; flowers

without a pouched throat:

4a Inflorescence terminal, an aggregate of sessile cymes, each

usually reduced to a single flower; seed testa with shortly

sinuate-dentate pattern Ill sect. Betonica

4b Inflorescence axillary, a lax branching raceme-like com-

pound cyme; seed testa either with long barbed scales

or reticulate with each cell outlined by minute papillae

I sect. Raphidospora

2b Pollen areolate:

5a Calyx four-lobed; inflorescence a terminal elongated ‘spike’;

seeds without crystals in testa; flowers small and white

V sect. Minima

5b Calyx five-lobed; inflorescence a terminal aggregate of

sessile cymes; seeds with crystals in testa; flowers medium

to large, yellow or blue IV sect. Tyloglossa

lb Pollen two-colporate or two-porate; inflorescence of axillary

lax or condensed cymes:

6a Pollen with entire margocolpi; shrubs, sometimes woody;

cymes often reduced to a single axillary flower; bracts and

leaves often reduced II sect. Justicia

6b Pollen areolate; plants herbaceous or shrubby; inflorescence

rarely reduced to a single flower:

7a Pollen porate, small (22-36 pm long); inflorescence an

axillary elongate ‘spike’; flowers white, small; herbaceous;

seeds irregularly rugose at high magnifications

VII sect. Ansellia

7b Pollen colporate, medium-sized (34—60 pm long); inflores-

cence of axillary sessile cymes; flowers and habit various;

seeds usually papillate at high magnifications, sometimes

reticulate VI sect. Hamieria

I. Sect. Raphidospora (Nees) T. Anders, (sect. Ill

in Graham: 587). Species included: J. glabra Koenig ex

Roxb. , J. campylostemon (Nees) T. Anders.

Inflorescence of axillary laxly-branching thyrses. Phero-

phylls and prophylls reduced, subulate. Pollen 3-colporate,

with entire margocolpi. Capsule relatively large, hard and

woody, with a long stipe. Seeds four, reticulate or with

long barbed scales.

The inflorescence and pollen are considered in most

respects to be the most primitive in southern Africa. The

seeds of the two southern African species, however, differ

greatly and, in/, glabra at least, may be considered highly

advanced, as they are covered with long barbed scales

(Immelman 1990a).

All except one species of Justicia seen have anthers with

the lower locule spurred, J. campylostemon being an

occasional exception. A few specimens were seen where

the anther locules were not spurred, but only mucronate.

This is the regular condition in Siphonoglossa. However,

as this is variable within J. campylostemon, and its other

characters are typical of Justicia, it was not considered

a species of Siphonoglossa.

From the three-colporate pollen with entire margo-

colpi and the laxly branching inflorescence of sect.

Raphidospora, a number of lines of specialization can be

traced.

Bothalia 22,2 (1992)

173

n. Sect. Justicia (sect. VI in Graham: 595). Species

included: J. bolusii C.B. Cl., J. orchioides L.f., J. cuneata

Vahl, J. thymifolia (Nees) C.B. Cl., J. guerkeana Schinz

and J. platysepala (S. Moore) P.G. Mey.

Inflorescence of stout-peduncled axillary elongated

thyrses or nearly sessile or flowers solitary, pedicelled or

sessile. Pherophylls and prophylls reduced and subulate

or large and lanceolate to broadly obovate. Flowers hooded

or upper line straight, with or without pouched throat.

Pollen two- or three-colporate, with entire margocolpi.

Capsule relatively large, hard and woody, with a long stipe.

Seeds one (two), smooth or rough. Shrubby, in J. bolusii

with suckers from stems below ground.

J. bolusii is a rare eastern Cape endemic, and in flowers,

seeds and pollen, differs from other species of the section

in the southern African region. The pollen type is the same

as sect. Raphidospora. J. bolusii and J. campylostemon

were formerly placed together in sect. Gendarussa by

Clarke (1901) (= sect. Raphidospora), but / bolusii differs

from both /. campylostemon and /. glabra in the single

smooth seed, the pouched corolla throat and the un-

branched inflorescence.

/. bolusii is probably most similar to some East African

species of Justicia, e.g. /. cordata (Nees) T. Anders., a

common species in East Africa. J. bolusii and /. cordata

are very similar in appearance, both have a pouched throat

in the corolla, a clavate capsule and a similar inflorescence,

in addition to the unusual smooth seeds. Graham (1988)

states that the pollen of /. cordata is 2-colporate, but under

the SEM, it was seen to be 3-colporate as in /. bolusii

(Immelman 1989a). It is reasonable to surmise that the

section to which /. bolusii and /. cordata belong was once

more widely distributed in Africa, J. bolusii being a relict

of this distribution.

The pouched corolla throat of J. bolusii and /. cordata

is like that described for Monechma (Munday 1980) and

so are the 1- or 2-seeded capsules and smooth-surfaced

seeds. It was therefore considered whether J. bolusii should

not be better placed in that genus than in Justicia, but the

inflorescence and pollen are not like those of Monechma.

Monechma usually has sessile cymes or single sessile

or pedicellate flowers and has two-colporate areolate

pollen, whereas /. bolusii has inflorescences as described

above and 3-colporate pollen with entire margocolpi. The

one or two smooth seeds and pouched corolla throat

are considered to be a parallel adaptation in Monechma,

rather than indicating relationship between it and J.

bolusii.

Sect. Justicia sensu Graham (1988) also includes /.

orchioides L.f. and J. platysepala (S. Moore) P.G. Mey.

This has been accepted here with some hesitation, as it

is felt that differences exist. However, the whole of sect.

Justicia is in need of revision, which would have to include

more tropical African species. Graham states that J.

orchioides and J. platysepala have ‘type 4’ pollen, i.e.

2-porate with two raised bands (entire margocolpi), which

applies to these species but not to J. bolusii or J. cordata,

which are both 3-colporate. These two groups do not

resemble each other in inflorescence or seed testa either.

It is possible to place them in separate sections.

Within the second group (/. orchioides, J. thymifolia,

J. platysepala and J. guerkeana ) there is a tendency to

increased woodiness and to a reduction in the size of

the bracts and the number of flowers in the cyme, the

three trends progressing together. J. guerkeana and

J. platysepala have numerous flowers in the cyme, broad

bracts with green centres and white membranous margins,

and rather soft shrubby stems, whereas J. thymifolia is

woody, has small bracts and still has multiflowered

cymes. J. orchioides and /. cuneata invariably have small

bracts, solitary axillary flowers and are hard-stemmed

(in J. orchioides sometimes spiny) shrublets. It is not

suggested that any of these is the direct ancestor of any

other species.

III. Sect. Betonica (Nees) T. Anders, (sect. II in

Graham: 585). Species included: J. betonica L., J. montis-

salinarum A. Meeuse.

Inflorescence a terminal spike-like aggregate of axillary

cymes reduced to 1(3) flowers each. Pherophylls and

prophylls reduced and subulate or large and green-veined.

Pollen 3-colporate, with entire margocolpi. Capsule

relatively large, semi-woody, with a long stipe. Seeds 4,

rough.

The common and widespread J. betonica (South Africa

to India) and the narrowly endemic J. montis-salinarum

(Soutpansberg of the Transvaal) have pollen similar to that

of sect. Raphidospora, being 3-colporate with entire

margocolpi, but the cymes have been reduced to 1 (rarely

3) axillary flower(s) each. These flowers have been secon-

darily aggregated to form a dense, terminal ‘spike’. In J.

betonica there are large, colourful, imbricate pherophylls

and prophylls, wherease in J. montis-salinarum there

apears to have been even further reduction, with both

pherophylls and prophylls being small and subulate. In

both species, the seed testa is drawn into sinuate, dentate

structures, which in J. betonica have a micro-structure of

colliculate individual cells and in /. montis-salinarum are

finely striate. A sinuate or dentate testa was not seen in

any other species in the genus. Although the two species

placed in this section are very different superficially, their

similar pollen and inflorescence structure, and their dis-

tinctive seed testa, indicate they should be placed together.

IV. Sect. Tyloglossa (Hochst.) Lindau (sect. IV in

Graham: 390). Species included: J. petiolaris (Nees) T.

Anders, (with three subspecies), J. flava (Vahl) Vahl and

J. kirkiana T. Anders.

Inflorescence terminal, a dense or lax aggregate of

sessile axillary cymes. Pherophylls and prophylls reduced,

oblanceolate to obovate. Pollen 3-colporate, areolate.

Capsule of medium size, texture papery, stipe short. Seeds

4, reticulate, with crystals in testa.

This is a well-defined section, one of the major unifying

characters of the section being the seeds, which under the

SEM are seen to have one to many cubic or rectangular

crystals in each cell of the testa. This character is unique

to the section. The pollen is 3-colporate and areolate.

The inflorescence in the section is basically composed

of sessile cymes, as in the preceding section, but these

are aggregated into a terminal compound ‘spike’ with

.174

Bothalia 22,2 (1992)

numerous flowers at each node. The pherophylls and

prophylls are somewhat reduced, usually spatulate and

often with very long-stalked glandular hairs (Immelman

1990b). Though the inflorescence type of J. petiolaris

subsp. incerta and subsp. bowiei tends towards that of the

sect. Hamieria, being more open than in subsp. petiolaris ,

the pollen and seeds are quite different from those of sect.

Hamieria.

V. Sect. Minima Immelman, sect. nov. Type species:

J. minima A. Meeuse.

Inflorescentia terminalis elongata simplex spiciformis

e cymis flore solitario redactis composita. Pherophylla

prophyllaque redacta subulata. Calyx quadrilobatus. Pol-

linis grana tricolporata areolata. Semina quatuor, testa

reticulata.

Inflorescence a terminal, elongate, unbranched spike-

like aggregate of cymes reduced to a single flower each.

Pherophylls and prophylls reduced and subulate. Calyx

four-lobed. Pollen 3-colporate, areolate. Seeds 4, testa

reticulate.

J. minima is a rare endemic from the Waterberg of the

Transvaal. It is not here considered closely related to any

of the southern African species, and its affinities probably

lie north of the South African border. Its pollen is similar

to, though much smaller than that of sect. Tyloglossa

(above), but its inflorescence is a terminal, unbranched,

lax ‘spike’ (actually a reduced series of cymes forming a

terminal florescence) similar to that of sect. Ansellia. Both

pherophylls and prophylls are greatly reduced, and the

calyx is four-lobed. The reticulate micro-pattern of the

seed surface did not aid in placing the seed surface, as

the reticulate pattern is scattered in the genus, e.g. J.

campylostemon and J. capensis Thunb. , and therefore was

not considered significant. This species does not key out

in Graham’s key to sections, the nearest being sect.

Rostellaria subsect. Anisostachya (if the bracts are ignored)

or subsect. Ansellia (if the number of calyx lobes, the

pollen and the details of the seed are ignored).

VI. Sect. Hamieria (Solms-Laub.) Benth. (sect. V

in Graham: 591). Taxa included: J. capensis Thunb.,

J. protracta (Nees) T. Anders, (with two subspecies),

J. heterocarpa subsp. dinteri (S. Moore) Hedren, J. parvi-

bracteata Immelman and J. odora (Forssk.) Vahl. A very

complete synonymy for the section is given in Hedren:

62 (1989).

Inflorescence of scattered sessile axillary cymes, or these

reduced to a single flower. Pherophylls and prophylls

foliose, bracteoles reduced and subulate or absent. Pollen

2-colporate, areolate. Capsule of medium size, texture

papery, stipe short or, in one species, hard and woody with

long stipe (J. cuneata ) with J. odora and J. capensis

intermediate. Seeds 4, or only one in the dimorphic

capsules of J. heterocarpa subsp. dinteri and (rarely) J.

protracta subsp. protracta. Seed surface varied, usually

papillate or reticulate.

Both pollen and inflorescence in this section are

specialized with respect to the condition seen in sect.

Raphidospora, the pollen being 2-colporate and areolate

and the inflorescence reduced to pedunculate or sessile

axillary cymes with up to five flowers, scattered in the leaf

axils. The seeds of four of the five species in the section

have a micro-pattern of a single papilla in the centre of

each cell of the testa, but in J. capensis the seed

testa is reticulate. I would agree with Hedren (1989) in

considering J. capensis and J. odora as belonging in sect.

Hamieria, while being in some respects very different

from the majority of species in the section.

A feature of interest in this section is the dimorphic

capsules that are present in two members, regularly in

J. heterocarpa subsp. dinteri and encountered once in

J. protracta subsp. protracta. These are also occasionally

seen in Siphonoglossa leptantha (Nees) Immelman subsp.

leptantha. Specimens of J. protracta and S. leptantha

showing this condition are preserved at PRE and NU

( Balkwill 217 (NU), Balkwill 649 (NU, PRE), Immelman

s.n. (630291 in PRE)). As well as the normal four-seeded

capsule, a smaller, one-seeded, non-opening (indehiscent)

capsule is produced, with four irregularly serrate wings.

The seed does not differ from those in the normal capsules

except in being larger (Immelman 1990a). This was also

seen by Hedren (1989) in a number of tropical African

taxa in sect. Hamieria. The possession in common of a

specialised type of capsule is evidence of a close relation-

ship between Justicia sect. Hamieria and Siphonoglossa.

The seed of Siphonoglossa, like that of most of the

species of sect. Hamieria, is papillate, the pollen is simi-

lar, and its inflorescence is also of many-flowered scattered

sessile axillary cymes. In Hedren (1989), the J. striolata

species group of sect. Hamieria corresponds to Siphono-

glossa as recognised by me (non Oerst, Immelman 1989b),

and he has placed Siphonoglossa sensu Moore non Oerst.

in the synonymy of sect. Hamieria. He found it to separate

rather widely from the rest of the section on MDS

ordination, but nevertheless retained it in Justicia sect.

Hamieria. Though I gave it as my opinion (Immelman

1989b) that there are reasons for not combining the two

genera, if Siphonoglossa sensu Moore is placed in Justicia,

I feel it might best be as a separate section close to sect.

Hamieria.

Note on the genus Siphonoglossa : when I published my

paper on Siphonoglossa and Aulojusticia (Immelman

1989b), I was not aware of the existence of a series of

papers by Henrickson & Hilsenbeck (1979) and Hilsenbeck

(1990a, 1990b) on the South American representatives of

Siphonoglossa. Hilsenbeck’s opinion is that there are three,

or possibly four diverse taxa included under the name

Siphonoglossa, which are held together mainly by having

in common a long narrow corolla tube (Hilsenbeck 1990b).

After comparing the South African species with his

descriptions of the South American taxa, I would agree

that they do not belong together in a common genus,

separate from Justicia. Rather the South African (and other

African) taxa should be placed either in a section/sub-

section of Justicia (the latter being the course followed

by Hedren), or in a genus with a name other than

Siphonoglossa, the type species of Siphonoglossa being

a South American species. Aulojusticia, which already

exists, would be a suitable name. Before either of these

is done, however, it would be necessary to compare

the South African species with the South American

species and the tropical African species placed in Justicia

Bothalia 22,2 (1992)

175

sect. Hamieria species group J. striolata by Hedren,

looking especially at pollen, inflorescence structure and

seed testa.

VII. Sect. Ansellia (C.B. Cl.) Ensermu. Species

included: J. anselliana (Nees) T. Anders., J. crassiradix

Burkill & C.B. Cl. and / anagalloides (Nees) T. Anders.

Sect. Rostellaria T. Anders, subsect. Ansellia (C.B. Cl.)

V.A.W. Graham (sect. Vllb in Graham: 598).

Inflorescence axillary, an elongated, slender-peduncled

spike-like aggregate of cymes reduced to a single flower

each. Pherophylls and prophylls greatly reduced, subulate.

Pollen 2-porate or shortly 2-colporate, areolate. Capsule

of medium size, texture papery, stipe short. Seeds 4,

rough. Herbaceous, may be annual or creeping and rooting

at the nodes.

This well-defined section has inflorescence and flowers

resembling those of the sect. Minima, but the inflores-

cences are axillary not terminal, the calyx is five- not

four-lobed and the pollen is quite different. The two upper

(adaxial) lobes of the calyx are noticeably longer than the

three lower (abaxial) lobes. The pollen, which is the

smallest in the genus in southern Africa, is areolate and

either 2-porate or shortly 2-colporate. In /. crassiradix

there were short colpi present and it is considered probable

that the porate condition seen in /. anagalloides and /.

anselliana is ‘ derived from the colporate by gradual

reduction of the colpus length.

ACKNOWLEDGEMENTS

This work was done as part of a Ph.D. thesis in the

Department of Botany, University of Natal, Pietermaritz-

burg. I would like to thank my supervisor. Dr F. Getliffe

Norris, for her help and advice in the writing up of my

thesis, and the curators of all herbaria that loaned

specimens.

REFERENCES

CLARKE, C.B. 1901. Acanthaceae. In W.T. Thiselton Dyer, Flora

capensis 5,1: 1—92. Reeve, London.

ENSERMU, K. 1990. Justicia sect. Ansellia (Acanthaceae). Symbolae

botanicae upsaliensis 29: 2.

GRAHAM, A.W. 1988. Delimitation and infra-generic classification of

Justicia (Acanthaceae). Kew Bulletin 43,4: 551—624.

HEDREN, B.C.M. 1989. Justicia sect. Hamieria (Acanthaceae) in

tropical Africa. Acta Universitatis Upsaliensis Symbolae Botanicae

Upsaliensis XXIX, 1: 1-141. Almqvist & Wiksell International,

Stockholm, New York.

HENRICKSON, J. & HILSENBECK, R.A. 1979. New taxa and com-

binations in Siphonoglossa (Acanthaceae). Brittonia 31: 373 — 378.

HILSENBECK, R.A. 1990a. Pollen morphology and the systematics

of Siphonoglossa sensu lato (Acanthaceae). American Journal

of Botany 77: 27—40.

HILSENBECK, R.A. 1990b. Systematics of Justicia sect. Pentaloba

(Acanthaceae). Plant Systematics and Evolution 169: 219—235.

IMMELMAN, K.L. 1989a. Studies in the southern African species of

Justicia and Siphonoglossa (Acanthaceae): palynology. Bothalia

19: 151-156.

IMMELMAN, K.L. 1989b. Siphonoglossa and Aulojusticia in southern

Africa (Acanthaceae). Bothalia 19: 209, 210.

IMMELMAN, K.L. 1990a. Studies in the southern African species of

Justicia and Siphonoglossa (Acanthaceae): seeds. Bothalia 20:

49-59.

IMMELMAN, K.L. 1990b. Studies in the southern African species of

Justicia and Siphonoglossa (Acanthaceae): indumentum. Bothalia

20: 61-66.

MUNDAY, J. 1980. The genus Monechma Hochst. ( Acanthaceae tribe

Justiciae) in southern Africa. M.Sc. thesis. University of the

Witwatersrand, Johannesburg.

STEARN, W.T. 1971. Taxonomic and nomenclatural notes on Jamaican

gamopetalous plants. Journal of the Arnold Arboretum 52:

614-647.

Bothalia 22,2: 177-182 (1992)

Cololejeunea cardiocarpa , an epiphyllous liverwort in southern Africa

(Lejeuneaceae)

S.M. PEROLD*

Keywords: Amell, Cololejeunea cardiocarpa, epiphyllous liverwort, Lejeuneaceae, Leptocolea cristata var. lanciloba, Platycolea, southern Africa,

taxonomy

ABSTRACT

Cololejeunea cardiocarpa was misidentified by Arnell (1963) and described by him as a new variety, Leptocolea cristata

var. lanciloba. Gradstein et al. (1983) drew attention to this error. Fresh material has now been collected and the species

is described here and illustrated with photographs.

UITTREKSEL

Amell (1963) het Cololejeunea cardiocarpa verkeerd geidentifiseer en as ’n nuwe varieteit, Leptocolea cristata var. lanciloba

beskryf. Gradstein et al. (1983) het die aandag op hierdie fout gevestig. Vars materiaal is nou versamel en die spesie word

hier beskryf en met foto’s gel llustreer.

INTRODUCTION

Amell (1963) cited Alice Pegler PRE-CH 916 , as the type

of the name of his new variety, Leptocolea cristata var.

lanciloba S. Arnell, a leafy liverwort epiphyllous on

Encephalartos villosus Lem., from Kentani Dist., eastern

Cape Province. Recently, however, Gradstein et al. (1983)

recognized that Arnell’s description and illustrations

clearly referred to Cololejeunea cardiocarpa (Montagne)

Schuster.

This report is given for the following reasons: 1, the

above information may easily be overlooked; 2, the

distribution of this species in southern Africa, as reported

by Gradstein et al. (1983) in Plate 1/3, is indicated by a

dot in the southwestern Cape, instead of in the eastern Cape

(Transkei) as it should be, and 3, fresh material has re-

cently come to hand.

In the highly specialized family, Lejeuneaceae Cavers,

to which Cololejeunea cardiocarpa belongs, the subfamily

Cololejeuneoideae Herz. ex Grolle, is said to represent

a pinnacle in the evolution of the leafy hepatics (Schuster

1980). Two tribes are recognized: Displasiolejeuneae

Schuster and Cololejeuneae, the latter with two genera,

Cololejeunea (Spruce) Schiffn. and Aphanolejeunea

Evans, but the characters separating them are not at all

clear-cut. They are generally distinguished by the

following: the absence of underleaves, the small size of

the plants, the cells being more or less uniform, the stems

with one row of ventral cortical cells, the leaves with a

narrowly ovate or lanceolate shape, and the rhizoids in

a small group, one group per lateral leaf. The species of

Cololejeunea have vegetative lateral branching of, what

is termed, the Lejeunea- type, i.e. a branch does not

originate from a cortical cell and is thecal or ‘collared’;

there are numerous discoid gemmae with smooth outlines,

mostly originating from intramarginal leaf cells and with

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

MS. received: 1991-10-24.

their plane parallel to the lobe face; the leaves have inflated

lobules with a hyaline papilla usually more or less proximal

to the apical tooth and the perianth is distinctly beaked

with five carinae.

Schuster (1980) states that currently as many as about

200 species have been described under Cololejeunea sensu

lato. Of the three subgenera in the genus Cololejeunea

treated by him (Schuster 1980), namely Protocolea

Schuster, Platycolea Schuster and Cololejeunea, those

assigned to Protocolea and Platycolea never have tubercles

or connoidal processes on the cells of the leaf lobes,

lobules, bracts and perianths, as are present in Colole-

jeunea. Species in subgenus Platycolea, with its subgeneric

type, Cololejeunea cardiocarpa (selected by Schuster), are

characterized by laterally connate, finger-like hyaline cells

at the lobe apices and compressed perianths with low an-

tical and postical keels. Platycolea includes at least 31 spe-

cies which, however, display a wide diversity of characters.

Accordingly, E.W. Jones (pers. comm.) argues that there

is much to be said for keeping those species with ‘hyaline

border cells to the leaves’ united in the genus

Pedinolejeunea Chen & Wu (or in the subgenus Pedinole-

jeunea Benedix ex Mizut.), rather than splitting them up

by the shape of the lobules into Platycolea and Pedinole-

jeunea, as was done by Schuster. Clearly, there still is

much that is controversial about the matter.

Tixier (1985) places C. cardiocarpa in section Platycolea

Schuster s. em., subsection Fimbriatae Tixier; lately, he

(Tixier unpublished) has attempted to divide it into several

species because of its extreme variability (E.W. Jones pers.

comm.).

Cololejeunea cardiocarpa (Montagne) Stephani in

Hedwigia 29: 91 (1890).

Cololejeunea (Platycolea) cardiocarpa (Montagne) Schuster: 54 (1959);

Schuster: 173 (1963); Schuster: 1278 (1980); Vanden Berghen: 226 (1977);

Vanden Berghen: 448 (1978).

Lejeunia cardiocarpa Montagne: 446, 447 (1838—1842). Type: Cuba.

Leptocolea cardiocarpa (Montagne) Evans: 268 (1911); Schuster: 144

(1955); Jones: 200 (1957).

Bothalia 22,2 (1992)

3URE 1 -Cololejeunea cardiocarpa , vegetative phase. A ventra! view ol °f lobl S muchetUarged and, toward centre,

antical margin of lobe; D, apex of lobe, with several hyabne cells’^h^" Bruise 5W A x 35' B X 70; C D, x 175; E, x 350.

showing 2 subrectangular hyaline cells at the.r base. A-C, Brusse 5869, D, E, Brusse 5721. A, X b,

Bothalia 22,2 (1992)

179

Lejeunia (subg. Colo-Lejeunea sect. Leptocolea ) cardiocarpa

(Montagne) Spruce: 300 (1884-1885).

Lejeunia jooriana Austin: 20 (1875). Lejeunea (Colo-Lejeunea) jooriana

Stephani: 171 (1892). Cololejeunea jooriana (Stephani) Evans: 173 (1902).

Leptocolea jooriana (Stephani) Evans: 270 (1911); Stephani: 844 (1915).

Type: Louisiana.

Physocolea savesiana Stephani: 904 (1912—1917).

Synonymy mostly after Schuster: 1278 (1980) and partly after Tixier:

46 (1985).

Epiphyllous or corticolous on mostly smooth bark, in

small patches, closely appressed to substrate, flat, thin,

freely branching, yellow-green to green, small to medium-

sized. Stems up to ± 6 mm long, slender, 50-55 /zm wide

and together with the leaves up to 1500 gm wide at

maturity (Figure 1A), lacking underleaves, but bearing

small fascicles of rhizoids near each leaf; on cross section

with 1 medullary and 5 rows of cortical cells (more if

section taken close to origin of a branch). Leaves flat and

appressed both wet and dry, ovate to ovate-triangular,

obtusely rounded apically, widest below midline (Figure

IB), spreading at an angle of 55°— 75°, rather imbricate,

dorsal base usually extending across and slightly beyond

stem. Lobes (600— )850— 1000 x 300—550 pm, narrowed

at base, antical (Figure 1C) and postical margins smooth,

nearly straight, apex fairly obtuse to rounded, generally

bordered with a row of 4—16 hyaline cells, elongated and

rather finger-like (Figure ID), 40—65 X 15—20 pm,

sometimes at their base small, subrectangular cells,

12.5-25.0 x 20 pm, also hyaline (Figure IE); chlorophyl-

lose cells at apex ± 17.5 x 12.5 ^m, medially 25—30 x

15.0-22.5 /im and at base + 17.5 x 12.5 pm, all of one

type, thin-walled, with small trigones, containing oil

bodies, usually 3—7 per cell, fusiform in shape; antical

margin rarely with a row of ± quadrate, hyaline cells.

Lobules inflated, ovate to broadly ellipsoidal, constricted

at base (Figure 2F), extending b3— 2/5 the distance from

stem to apex of lobe, 300—325 x 250 /zm, cells ± 20.0

x 12.5 )im, keel convex, nearly continuous with postical

margin of lobe, lobule apex with obtuse, slightly jutting

out, 1-celled tooth, 20 x \5 pm\ about midway between

apical tooth and keel, a more prominent 3- or 4-celled

proximal tooth, with an inconspicuous, concealed hyaline

papilla at its base (Figure 2B). Stylus 2-celled, 45.0—52.5

x 10 pm, basal cell often larger than apical cell (Figure

2C). Gemmae numerous, discoid, broadly oval with

smooth outline, unspecialized and lacking adhesive cells,

usually on lower lobe surface and mostly originating from

intramarginal cells, ± 60—70 (tm in diameter, composed

of ± 29 cells of which ± 17 around margin (Figure 3D).

Auto- or paroicous. Androecia situated below perianth

or on a short lateral branch, bracts mostly unmodified but

distinguishable from vegetative leaves by the absence of

apical hyaline cells, in several pairs; antheridia 1 or 2 per

bract (Figure 2D), flask-shaped, 150 x 90 pm, with

hyaline neck ± 100 pm long. Perichaetial bracts with lobes

somewhat smaller than those of vegetative leaves, 450 x

200 pm, widely spreading, apex obtuse with a few hyaline

cells, lobule 200 x 180 pm. Perianth terminal or some-

times on a lateral branch, sessile, usually with a single

innovation, rarely 2, 560 x 440 pm, more or less

compressed, outline obovate, with 2 sharp lateral keels

and 2 smaller ventral ones (Figures 2E; 3C), apex with

short beak (Figure 3B). Sporangium initially enclosed in

a delicate calyptra, containing numerous spores. Spores

irregularly ovate, green, 25—45 x 20 pm, consisting of

several cells (Figure 2H). Elaters attached to inner wall

of valves of sporangium (Figures 2F; 3A), distributed 4,

4, 3, 3 per valve, 175 x 25 pm, free apices blunt (Figure

2G). This description was based on southern African

collections.

Arnell (1963) could not have been familiar with the

above species, hence he described the Pegler specimen as

a new variety, Leptocolea cristata var. lanciloba. Accord-

ing to Jones (pers. comm.) Cololejeunea cardiocarpa ap-

pears to differ from Leptocolea cristata (Stephani) E.W.

Jones by being a smaller plant with a more delicate tex-

ture; its leaves are ovate (not oval or obovate as in L. crista-

ta and are broadest proximally, tapering to a narrow apex

(but they are variable); the lobule is short with the free

margin inflexed and not visible in situ, as it is in L. cristata

and its cell walls are thin and lack trigones, the cells be-

ing transparent.

Cololejeunea cardiocarpa is a widespread pantropical

and oceanic subcosmopolitan (Poes 1978) species, known

from North, Central and South America, Africa, Mada-

gascar and New Caledonia. In Africa it has been recorded

from Sierra Leone (Jones & Harrington 1983), Cameroon

(Tixier 1975), Zaire, Kenya, Uganda, Tanzania (Bizot &

Poes 1979; Bizot & Poes 1982; Gradstein etal. 1983; Jones

1957; Poes 1985) and from Zimbabwe (Vanden Berghen

1978). In the checklist by Magill & Schelpe (1979)

Cololejeunea cardiocarpa is reported as occurring in

southern Africa; there are few records from the sub-

continent but new collections, such as the two from

Swaziland cited under specimens examined, gradually

increase the known distribution area in southern Africa.

It is, apparently, able to tolerate considerable desiccation

and, though usually epiphyllous, it becomes corticolous

under dry conditions, but probably is more difficult to find

on bark. Jones (1957) reports it as growing in mostly

montane localities in the tropics, whereas Schuster (1980)

regards it as confined to the Coastal Plain in the United

States. Poes (1978) reported it from lowland forest,

submontane rain forest and montane rain forest, with the

highest frequency per leaf in the submontane rain forest.

Gradstein et al. (1983) found it occurring from lowland

to lower montane forests under humid to subxeric

conditions. Two of the southern African collections are

from a coastal forest (Figure 4).

Schuster (1980) states that the finger-like, hyaline cells

of the leaf apices are absolutely diagnostic. These are,

however, not always well developed and may sometimes

even be lacking, when it may be more difficult to place

a specimen correctly. Schuster (1980) also remarks that

the stylus in this species is merely a slime papilla, but

I found it to be 2-celled in the specimens I examined. It

probably is not of diagnostic significance.

SPECIMENS EXAMINED

SWAZILAND. — 2631 (Mbabane): Malolotja Nature Reserve,

evergreen riverine forest, on leaflets of a cycad, alt. 930 m, (— AA),

Prendergast s.n. ; small tree-lined gulley in Highland Grassland, on small,

thin dentate leaves of bush tree, alt. 1 390 m, (— AA), Prendergast s.n.

(both specimens fide Dr E.W. Jones (pers. comm.) and held at the

herbarium of the Malolotja Nature Reserve, Mbabane, Swaziland).

180

Bothalia 22,2 (1992)

FIGURE 2. — Cololejeunea cardiocarpa. A-C, vegetative phase: A, lobule in situ ; B, lobule with apical and proximal teeth much enlarged,

C, stylus. D-H, sexual phase: D, 2 antheridia in lobule; E, perianth with perichaetial leaves at base; F, open perianth with

sporangium; G, 2 elaters partially shown; H, spore. A-F, H, Brusse 5869; G, Brusse 5721. A, X 175; B, C, G, X 350; D, X 87, E,

x 70; F, H, x 700.

Bothalia 22,2 (1992)

181

FIGURE 3. — Cololejeunea cardiocar-

pa. A— C, sexual phase: A,

opened sporangium, showing

attached elaters; B, tip of

perianth from above; C, cross

section taken ± midway

through perianth. D, gemma.

A, Brusse 572T, B— D, Brusse

5869. A-C, x 87; D, x 350.

TRANSKEI.— 3228 (Butterworth) : Kentani Dist., growing on

Encephalartos villosus Lem. leaves, (—AD), Pegler PRE-CH 916 (PRE);

Dwesa Nat. Res., ‘4 km from campsite on road to mBashe, 10—50 m

beyond nGoma River Bridge, near road, growing on Buxus natalensis

(Oliv.) Hutch, leaves, in understorey vegetation of coastal forest, (— BD),

Brusse 5721 (PRE); Dwesa Nat. Res., about 5 km from campsite on

road to mPume gate, growing on Encephalartos villosus Lem. leaves,

in understorey vegetation of coastal forest, (— BD), Brusse 5869 (PRE).

ACKNOWLEDGEMENTS

I wish to express my gratitude to Dr E.W. Jones, Oxford,

and to Prof. T. Poes, Hungary, for critically reading the

manuscript, and for their constructive suggestions, as well

as for all manner of other help; also to Mr Franklin Brusse,

lichenologist at NBI, for collecting fresh specimens of

Cololejeunea cardiocarpa. Mrs A. Romanowski, photo-

grapher, is thanked for developing and printing the

photographs and Mrs J. Mulvenna, typist, is also sincerely

thanked for her valued contribution.

FIGURE 4. — Distribution of Cololejeunea cardiocarpa in southern

Africa.

REFERENCES

ARNELL, S. 1963. Hepaticae of South Africa. Swedish Natural Science

Council, Stockholm.

AUSTIN, C.F. 1875. New Hepaticae. Bulletin of the Torrey Botanical

Club 6: 17-21.

BIZOT, M. & P6CS, T. 1979. East African bryophytes HI. Acta Botanica

Academiae Scientiarum Hungaricae 25: 223—261.

BIZOT, M. & P6CS, T. 1982. East African bryophytes V. Acta Acade-

miae Scientiarum Hungaricae 28: 15—64.

EVANS, A.W. 1902. The Lejeuneae of the United States and Canada.

Memoirs of the Torrey Botanical Club 8: 113—183.

EVANS, A.W. 1911. Hepaticae of Puerto Rico. 10. Cololejeunea,

Leptocolea and Aphanolejeunea. Bulletin of the Torrey Botanical

Club 38: 251-286.

GRADSTEIN, S.R, P6CS, T. & vA^A, J. 1983. Disjunct Hepaticae

in tropical America and Africa. Acta Botanica Hungarica 29:

127-171.

JONES, E.W. 1953. African Hepatics II. Leptocolea with hyaline-

margined leaves. Transactions of the British Bryological Society

2: 144-157.

JONES, E.W. 1957. African Hepatics XII. Some new or little-known

Lejeuneaceae. Transactions of the British Bryological Society 3:

191-207.

JONES, E.W. & HARRINGTON, A.J. 1983. The hepatics of Sierra Leone

and Ghana. Bulletin of the British Museum (Natural History),

Botany 11: 215-289.

MAGILL, R.E. & SCHELPE, E.A. 1979. The bryophytes of southern

Africa. Memoirs of the Botanical Survey of South Africa No. 43.

MONTAGNE, C. 1838—1842. In Ramon de la Sagra, Histoire physique,

politique naturelle de V He de Cuba, Botanique 9: 427—492.

POCS, T. 1978. Epiphyllous communities and their distribution in East

Africa. Bryophytorum Bibliotheca 13: 681—713.

POCS, T. 1985. East African bryophytes. VII. Acta Botanica Hungarica

31: 113-133.

SCHUSTER, R.M. 1955. North American Lejeuneaceae. 0. Paradoxae:

genera Aphandejeunea and Leptocolea. Journal of the Elisha

Mitchell Scientific Society 71: 126—148.

SCHUSTER, R.M. 1959. Epiphyllous Hepaticae in the southern

Appalachians. The Bryologist 62: 52—55.

SCHUSTER, R.M. 1963. An annotated synopsis of the genera and sub-

genera of Lejeuneaceae 1. Nova Hedwigia, Beihefte 9: 1—203.

SCHUSTER, R.M. 1980. The Hepaticae and Anthocerotae of North

America. Vol. 4. Columbia University Press, New York.

SPRUCE, R. 1884—1885. Hepaticae amazonicae et andinae. Transac-

tions of the Proceedings of the Botanical Society, Edinburg 15:

1-590.

182

Bothalia 22,2 (1992)

STEPHANI, F. 1890. Die Gattung Lejeunea in Herbarium Lindenberg.

Hedwigia 29: 68—99.

STEPHANI, F. 1892. The North American Lejeuneae. Botanical Gazette

17: 170-173.

STEPHANI, F. 1912—1917. Species hepaticarum 5: 1-1044.

TIXIER, P. 1975. Contribution it l’6tude de l’h^paticologie africaine. 1.

R6coltes en bordure du Golfe de Guinee (Cameroun et Gabon).

Annales de la Faculte des Sciences, Yaounde 20: 3—10.

TIXIER, P. 1985. Contribution a la connaissance des Cololejeunoideae.

Bryophytorum Bibliotheca 27. Cramer, Vaduz.

VANDEN BERGHEN, C. 1977. H6patiques epiphylles recoltees par J.L.

De Sloover au Kivu (Zaire) et au Burundi. Bulletin du Jardin

botanique national de Belgique 47: 199—246.

VANDEN BERGHEN, C. 1978. Hepatiques epiphylles recoltees en

Rhodesie. Revue bryologique et lichenologique 44: 443—452.

Bothalia 22,2: 183-197 (1992)

Notes on African plants

VARIOUS AUTHORS

PYXINACEAE

A NEW SPECIES IN THE LICHEN GENUS HETERODERMIA , FROM COASTAL NAMAQUALAND

Heterodermia namaquana Brusse, sp. nov. , thallo

ut in H. erinacea (Ach.) W. A. Weber, sed subtus

sorediato.

Thallus foliosus vel subfruticosus, ramunculicola vel

corticola, usque ad 60 mm longus, laxe adnatus. Lobi sub-

lineares, ascendentes, 1.0— 3.5 mm lati, 70-200 /zm crassi,

sympodiales; margines perciliati, ciliis simplicibus, albis,

vel interdum apicem versus denigratis, 2—7 mm longis,

basin versus 85-110 /xm crassis. Thallus supeme cinereus

vel albidus, opacus vel interdum pruinosus, sparse ciliatus,

undulatus vel glebosus. Cortex superior hyalinus, peri-

clinate et longistrorsum prosoplectenchymatus, 25—130

crassus. Stratum gonidiale sinuatum vel confornica-

tum, in crassitudine pervarians; algis Trebowciis, 7—19

/zm diam. Medulla albida, demum in sorediis omnino

dissoluta, ultimo evanescens. Cortex inferior deficiens.

Thallus infemq albidus, nervatus, sparse ciliatus, ad

apicem versus viridi-sorediatus; areae sorediorum diffu-

sae, virides, saepe leviter convexae; soredia 15—40 nm

diam. Apothecia infrequentia, substipitata, laminalia,

usque ad 4 mm diam.; discus carbo-nigrescens vel

cinereus; margines ciliati, cinerei vel albidi. Cortex

hyalinus, anticlinate prosoplectenchymatus, 45-80 gm

crassus. Stratum gonidiale ut in thallo. Medulla albida,

laxa. Hypothecium hyalinum, 30—50 ^trn crassum, J-,

cyanophilum, granulis inspersum. Epihymenium badium,

granulare. Hymenium hyalinum, 55-70 gm altum, J+

caeruleum; paraphyses septatae, capitatae; asci clavati,

cum tholis, J+ caeruleis vel pallide caeruleis, typi Leca-

norae. Ascosporae octonae, fuscae, 2(l)-loculares, ad

septum leviter constrictae, primum in aqua typi Pachy-

sporariae, dein in solutione aquosa hydroxidi kalii typi

Physciae, 14.5—22.0 x 6.5— 8.5 gm. Pycnidia non visa.

Thallus atranorinum et zeorinum continens.

TYPUS. — Cape, 2917 (Springbok): (—AC), Namaqua-

land coastal plain, 1.6 km from the first Kleinzee turnoff

on the main Port Nolloth-Steinkopf tarmac road, to

Kleinzee. Port Nolloth Allotment Area. Common lichen

growing on twigs of various shrubs, in gently undulating

terrain. Succulent shrubland with Stoeberia one of the

dominant bushes. Alt. 190 m. F. Brusse 5930, 9-9-1991

(PRE, holo. ; B, BM, COLO, CTES, E, LD, S, TNS, UC,

UPS, US, iso.). Figurae 1, 2 & 3.

Thallus as in Heterodermia erinacea (Ach.) W. A.

Weber, except lower surface sorediate.

Thallus foliose or subfruticose, on twigs or corticolous,

up to 60 mm long, loosely adnate. Lobes sublinear,

ascending, 1.0— 3.5 mm broad, 70—200 gm thick, sym-

podial; margins abundantly ciliate, cilia simple, white, to

sometimes blackened towards the tips, 2-7 mm long,

85-110 gm thick near base. Upper surface ash-grey to

whitish, opaque or sometimes pruinose (frosted), sparsely

ciliate, undulate to lumpy (lumps corresponding to pockets

on the lower surface). Upper cortex hyaline, periclinally

and longitudinally prosoplectenchymatous, 25—130 gm

thick. Algal layer vaulted, very variable in thickness; algae

Trebouxia, 7—19 gm diam. Medulla whitish, at length

completely degenerating into soredia, finally vestigial.

Lower cortex lacking. Lower surface whitish, veined,

sparsely ciliate (the cilia originating from the lower side

of the upper cortex), green sorediate at the lobe tips;

sorediate areas diffuse, green, often mildly convex

(corresponding to a concavity in the upper surface of the

FIGURE 1. — Heterodermia nama-

quana Brusse, habit photo-

graph. E Brusse 5930, holo-

type. Scale in mm.

184

Bothalia 22,2 (1992)

FIGURE 2.—Heterodermia nama-

quana Brusse, in habitat, on

shrub, Stoeberia beetzii (Din-

ter) Dinter & Schwant. The

shrub is about 1.5 m tall. F.

Brusse 5930, type locality, with

Fifteen Miles Mountains in the

distance.

lobe tip; lobe tip subinvolute); soredia 15-40 pm diam.

Apothecia infrequent, substipitate, laminal, up to 4 mm

across; disc charcoal to grey (when heavily pruinose);

margins ciliate, ash-grey to whitish. Cortex hyaline,

anticlinally prosoplectenchymatous, 45-80 pm thick.

Algal layer as in thallus, absent under hymenium. Medulla

whitish, lax. Hypothecium hyaline, 30—50 pm thick,

J-, cyanophilic (stains deep blue in lactophenol Cotton

Blue), granular inspersed. Epihymenium reddish brown,

granular. Hymenium hyaline, 55—70 pm high, J+ blue;

paraphyses septate, capitate; asci clavate, eight-spored,

tholus J+ blue or pale blue, Lecanora-type. Ascospores

greyish brown, 2(l)-locular, slightly constricted at septum,

Pachysporaria-type in water, changing to Physcia-type in

an aqueous solution of potassium hydroxide, 14.5—22.0

x 6.5— 8.5 pm. Pycnidia not seen. Chemistry. Atranorin

and zeorin present (as well as traces of leucotylin and 6a,

16/3-di-O-acetylleucotylin).

This new species was previously thought to be Hetero-

dermia erinacea (Ach.) W.A. Weber (Brusse 1988), but

material of the American species has since been seen

(Sipman, Lichenotheca Latinoamericana no. 23). The

most conspicuous difference is that H. erinacea is non-

sorediate (Kurokawa 1962), whereas H. namaquana is

sorediate. The soredia in this species are formed at the

expense of the medullary tissue, so that older lobes have

only vestiges of the medulla left. Consequently, older lobes

consist solely of the upper cortex, with very little else.

This lichen is also ciliate on all surfaces, which includes

the lower surface, which is sparsely ciliate. Interestingly,

the cilia on this surface arise from the lower side of the

upper cortex, and pass through the medulla and soralia,

when these are present. Like H. erinacea, this new species

has a lower surface with a veined appearance. In H. nama-

quana, this is due to vein-like thickenings in the lower side

of the- upper cortex. The lobes are mildly pocketed in

places which gives the upper side a lumpy appearance.

The sorediate lobe tips are also often subinvolute, giving

the diffusely sorediate areas of the lower surface a convex

appearance.

The ascospores of these two species are of similar size

and ascospore-type in an aqueous solution of potassium

hydroxide (the recommended observation medium

(Mayrhofer & Poelt 1979) for ascospore-types of fresh

material), but differ in water. Freshly prepared sections

FIGURE 3. — Heterodermia namaquana Brusse, growing on twigs of

Stoeberia beetzii (Dinter) Dinter & Schwant. F. Brusse 5930, type

locality.

Bothalia 22,2 (1992)

185

of the apothecia show ascospores of the Physcia-type in

H. erinacea (same as in KOH), but of Pachysporaria-type

in H. namaquana. The changeover from Pachysporaria-

type to Physcia-type in H. namaquana can be observed

by allowing a solution of potassium hydroxide to pass under

the cover slip of freshly prepared apothecial sections in

water. No such changes can be observed for

H. erinacea. This represents a fundamental difference

between the ascospores of these two species, and for this

reason, they do not represent a true species pair.

The tholus of both species is fairly pale blue in Lugol’s

iodine solution, and most of the blue colour of hymenial

sections in this solution is due to the hymenial gel. The

tholus illustrated by Honegger (1978, 1980) for Physcia

stellaris, is fairly representative of Heterodermia nama-

quana as well.

In the sterile state this new species may be confused

with Heterodermia comosa (Eschw.) Follm. & Redon, a

similar-looking sorediate lichen (Swinscow & Krog 1988).

However, this species is unrelated, because of its larger

(30—35 x 13-16 fim), Polyblastidium-type ascospores

(Kurokawa 1962).

At present this new species is known from a 250 km

stretch of coastal succulent shrubland, ranging from the

Klinghardt Mountains in southwestern Namibia to the Port

Nolloth area of the northwestern Cape Province.

REFERENCES

BRUSSE, F.A. 1988. Five new species of Parmelia (Parmeliaceae,

lichenized Ascomycetes) from southern Africa, with new com-

binations and notes, and new lichen records. Mycotaxon 31:

533-555.

HONEGGER, R. 1978. The ascus apex in lichenized Fungi I. The

Lecanora-, Peltigera- and Teloschistes-types. The Lichenologist

10: 47-67.

HONEGGER, R. 1980. The ascus apex in lichenized Fungi II. The

Rhizocarpon- type. The Lichenologist 12: 157—172.

KUROKAWA, S. 1962. A monograph of the genus Anaptychia. Beihefte

zur Nova Hedwigia 6: 1—115.

MAYRHOFER, H. & POELT, J. 1979. Die saxicolen Arten der Flechten-

gattung Rinodina in Europa. Bibliotheca Lichenologica 12: 16.

SWINSCOW, T.D.V. & KROG, H. 1988. Macrolichens of East Africa :

91. British Museum (Natural History), London.

F. BRUSSE*

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

MS. received: 1991-11-08.

RICCIACEAE

LECTOTYPIFICATION OF RICCI A CRYSTALLINA

While working on a taxonomic revision of the liverwort

family Ricciaceae for the Flora of southern Africa

Cryptogam series, it was brought to my attention that no

lectotype had been designated for Riccia crystallina L. The

1988 loan to PRE from Helsinki (Herb. S.O. Lindberg,

H-SOL), of Lichen no. 12 ( Riccia crystallina ), a Dillenian

specimen presumably examined by Linnaeus, was accom-

panied by a loan form bearing the following message from

Dr Pekka Isoviita: ‘The type of Riccia crystallina, as now

used, should be Micheli’s illustration (with typotype at

FI?)’. Mme Jovet-Ast, in her authoritative work on the

Mediterranean species of Riccia (Jovet-Ast 1986), had,

however, cited as the type specimen of R. crystallina : ‘tres

petit specimen, Herbier Dillenius, Isoviita (1970) (H-SOL)’.

Puzzled by this discrepancy, I wrote to both Dr Isoviita

and Mme Jovet-Ast, enquiring about the typification.

Previously, Jovet-Ast (1964, 1966) had thoroughly in-

vestigated the history and taxonomy of R. crystallina L.

emend. Raddi and of R. cavernosa Hoffm. emend. Raddi.

This was necessitated by the confusion, which had reigned

up to that time concerning these two species, although

Micheli (1729) had clearly distinguished between them

(nos. 1 and 5 in his Nova plantarum genera ):

1. Riccia minor, latifolia, pinguis, aspergine chrystallina

perfusa Tab. 57. fig. 3. Hepatica palustris, lobis cristatis

Bot. Paris, num. 5 Tab. XIX. fig. 2. Brumali, atque Vemali

tempore in urbanis horris ubique, et ita copiose crescit,

ut ab hac hortulani, et floricultores abhorreant.

5. Riccia minima, pinguis, aspergine chrystallina perfusa

Tab. 57. fig. 7. In sylva Coenobii Patrum Franciscanorum,

vulgo la Doccia supra memorata.’ (Micheli’s No. 1 Riccia

minor, latifolia... represents the current R.r crystallina, his

No. 5 Riccia minima pinguis... the current R. caverno-

sa). Linnaeus (1753) however, cited ‘Riccia minima et

minor pinguis aspergine crystallina perfusa. Mich. gen.

107. t. 57. f. 7.3, thus combining the two under R.

crystallina.

As Linnaeus’s (1753) Species plantarum is the starting

point of the nomenclature of the Hepaticae, his treatment

superseded that of Micheli, regardless of the fact that he,

Linnaeus, was not very knowledgeable about the liverworts

and had even grouped them together with the Algae. In

feet, Linnaeus’s concepts of the hepatics were mainly based

on Dillenius’s (1741) Historia muscorum, to which he also

referred in the protologue of Riccia crystallina, as well

as to Vaillant’s (1723, 1727) Botanicon parisiense and to

his own works, Flora suecica (Linnaeus 1745a) plus ‘Oland

& Gotland journey’ (Linnaeus 1745b).

In his later emendation Raddi (1818) distinguished

between R. crystallina and R. cavernosa, for which he

respectively cited Micheli’s drawings Tab. 57. fig. 3 and

fig. 7, but Raddi’s implicit treatment was not a typifica-

tion, not even under the ‘residue method’: he did not

mention the Dillenius element of R. crystallina, also cited

by Linnaeus (1753), anywhere in his paper, (as Isoviita

recently wrote to Jarvis — see below), nor did Raddi

directly indicate that Micheli’s Tab. 57 fig. 3 was the type

(ICBN 1988 Art. 8.3). The confusion thus persisted, even

into this century: in southern Africa, Duthie & Garside

(1937, 1939) recognized R. crystallina as R. plana Tayl.

and R. cavernosa as R. rautanenii Steph. Sim (1926) men-

tions R. crystallina, but cites no collections which could

be checked. According to Duthie & Garside (1937), Sim’s

drawings of R. crystallina ‘A, of part of thallus, x 7’ is

that of R. cupulifera AN. Duthie and of ‘D, four spores

before separation, x 50’ is that of R. curtisii (Aust.) James,

thus illustrating that Sim’s concept of the species was not

clear. Amell (1963) described both R. chrystallina (sic)

and R. plana, although he (Amell 1953) had remarked

that their relationship had previously been pointed out

186

Bothalia 22,2 (1992)

..... .

JA x

/ £ l Cw — ^ £ ^r^yv ;

fa ( *

^ 4. l7i v- //^3 %&dri ^*9 iW'

Herbarium

Universitatis Fiorentinae

FOTOTECA

y, Wj}^ J'>*

***#v

’*M

- P\UCUI^

fc S i

*>xoy*' £Kt>n%t*.

N.° 6' ? 8

Data <£. |o- l-^i

frfiVf

j&u-U %Uts6c* x

*Zx£/i2iir!tjri

<*> ±*Vi m 4*RSl> 17109.

Fy-z

G

B

FIGURE 4. — A, photograph of herbarium sheet (with enlargements of all the relevant specimen photographs and labels) of Micheli’s Riccia species

0:1. n: 1 = Ordo : l.no.:l and 0:2. n:l = Ordo : 2.no.:l from which were illustrated Tab. 57. fig. 1 (I on photo of label) and fig. 3, (3

on photo of label) respectively; B, xerox copy of Micheli’s Tab. 57. fig. 3G.

Bothalia 22,2 (1992)

187

by Garside. Amell’s (1963) drawings of the spores of R.

crystallina are those of R. cavernosa, and R. cavernosa

was described by him (Amell 1963) as R. rautanenii. Jovet-

Ast (1964, 1966) placed R. plana in synonymy under R.

crystallina and R. rautanenii under R. cavernosa.

As observed by Garside in 1957 and quoted by Jovet-

Ast (1964), there is no type specimen of R. crystallina in

the Herbarium of the Linnean Society in London, nor had

Jovet-Ast designated a lectotype. Nevertheless, ever since

her publications (Jovet-Ast 1964, 1966), it had been

accepted practice to follow her example by citing the

Micheli element as R. crystallina L., rather than the

Dillenian element, which actually represents R. cavernosa

Hoffm. (Koponen et al. 1977). My examination by SEM

of the spores of the latter specimen certainly confirms this

observation, although the spores are not very well

preserved. Isoviita’s (1970) suggestion that the Dillenian

specimen should be selected as lectotype, also failed to

‘preserve current usage’ (ICBN 1988, Recommendation

7B.5) (see filed at PRE, copy of Dr Isoviita’s letter, dated

19 February 1990, and addressed to Dr Jarvis at The

Natural History Museum, London; Dr Isoviita kindly

forwarded this letter to me).

In explaining the reason for her citation of the small

Dillenian specimen as the type of R. crystallina, Jovet-

Ast (pers. comm.) wrote that she had thought that Isoviita

would have been aware of her earlier work (Jovet-Ast

1964, 1966) ahd I presume, perhaps she may also have

assumed that he was in possession of additional informa-

tion, even though his concept of R. crystallina sensu

stricto was shown by Koponen et al. (1977) to have been

erroneous.

In her correspondence with me (letter dated 28 January

1990), Mme Jovet-Ast very graciously outlined several

options open to one in selecting a lectotype for R. crys-

tallina. Among these, she suggested that one could choose

Micheli’s figure of R. minor (Tab. 57. fig. 3) (although

no spores were illustrated), or else, one could select a

neotype from among the specimens collected and named

by Raddi as R. crystallina. It proved to be unnecessary

to resort to the latter option as Dr Chiara Nepi, curator

at FI, kindly sent micrograph negatives of three of

Micheli’s Riccia specimens (two of which are shown on

Figure 4) assuring me that they were of ‘the only ( Riccia )

specimens that may have some references with the de-

scriptions in Nova plantarum genera'. I ascertained that

the labels numbered I Riccia 0:1. n:l and 3 Riccia 0:2. n:l

correspond to Micheli’s descriptions on pp. 106 and

107 (Micheli 1729) of Riccia Ordo l.numero 1 and

of Ordo 2.numero 1 respectively. Each of the labels

also bears some words that can be linked with the descrip-

tions, namely on I (specimen unfortunately in fragments)

is written ‘fructu globoso’ and in the description it

reads ‘fructu subrotundo’ and on 3 is written ‘cristallina

aspersa’, with the description reading ‘aspergine chrystal-

lina perfusa Tab. 57. fig. 3’. Below the latter label is an

unnumbered one of ‘ Riccia major' (the ink is some-

what smeared) that clearly pertains to Riccia 0:1. n:l, since

it is described in the text on p. 106 as Riccia major.

The third specimen, which is not shown on Figure 4, is

labelled ‘2 Riccia 0: (figure illegible as scored through). n:

2’ and ‘fructu pyramidato’ as in Micheli’s description on

page 107.

Unfortunately, the rosette, which clearly must have

served as the model for Micheli’s drawing of Tab. 57. fig.

3, has become detached from its original position (notice

smudge on Figure 4A) and has come to lie between the

two specimens, but there is no doubt about its identity nor

of the remaining two pieces on the herbarium sheet.

Micheli’s drawing Tab. 57. fig. 3 (Micheli 1729)

(reproduced in Figure 4B from a xerox copy of the

original) is therefore selected as the lectotype of R. crys-

tallina, with the typotype, Riccia 0:2. n:l, held at FI.

ACKNOWLEDGEMENTS

I wish to thank Mme Jovet-Ast most sincerely for her

kindness and generosity in sharing her experience of the

above problem with me; also Dr Isoviita for the loan of

the Dillenian specimen held at H-SOL and for forwarding

to me a copy of his letter to Dr Jarvis. I had several

discussions about this matter with my colleagues at NBI,

Pretoria, and wish tp thank Dr H.F. Glen, Dr D.J.B.

Killick, Dr O.A. Leistner and Mr J. van Rooy for their

advice; also Mr Ashley Nicholas, South African Liaison

Officer at Kew at the time, for all his help in obtaining

literature. I particularly wish to thank the curator of FI,

Dr Chiara Nepi, for a copy of the relevant pages of

Micheli’s work and for the micrograph negatives; also the

curator of PI for the loan of Raddi’s specimens.

REFERENCES

ARNELL, S. 1953. Notes on South African Hepaticae. Extrait de la

Revue bryologique et lichenologique T. XXII fasc. 1—2.

ARNELL, S. 1963. Hepaticae of South Africa. Swedish Natural Science

Council, Stockholm.

DILLENIUS, J.J. 1741. Historia muscorum. Oxford.

DUTHIE, A.V. & GARSIDE, S. 1937. Studies in South African Riccia-

ceae 1. Transactions of the Royal Society of South Africa 24:

93-133.

DUTHIE, A.V. & GARSIDE, S. 1939. Studies in South African

Ricciaceae II. Transactions of the Royal Society of South Africa

27: 17-28.

ISOVIITA, R 1970. Dillenius’s Historia muscorum as the basis of hepatic

nomenclature, and S.O. Lindberg’s collection of Dillenian bryo-

phytes. Acta Botanica Fennica 89: 1—28.

JOVET-AST, S. 1964. Riccia crystallina L. emend. Raddi et Riccia

cavernosa Hoffm. emend. Raddi (Note preliminaire) . Revue

bryologique et lichenotogique 33: 459—483.

JOVET-AST, S. 1966. Riccia crystallina L. emend. Raddi et Riccia

cavernosa Hoffm. emend. Raddi. 13. Revue bryologique et

lichenologique 34: 82—90.

JOVET-AST, S. 1986. La Riccia de la Region Mediterraneenne.

Cryptogamie, bryologique et lichenologique 7: 283—431.

KOPONEN, T., ISOVIITA, P. & LAMMES, T. 1977. The bryophytes

of Finland: an annotated checklist. Flora Fennica 6: 1—77.

LINNAEUS, C. 1745a. Flora suecica. Stockholm.

LINNAEUS, C. 1745b. Olandska och Gothlandska Resa. Stockholm &

Upsala.

LINNAEUS, C. 1753. Species plantarum. Bradbury & Evans.

MICHELI, P.A. 1729. Nova plantarum genera. Florentiae.

RADDI, G. 1818. Novarum vel rariorum ex Cryptogamia Stirpium in

agro Florentino collectarum Decades duae. Opusculi Scientifici

di Bologna 2: 349—361.

SIM, T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1—475. Cape Town.

VAILLANT, S. 1723. Botanicon Parisiense Prodromis 5.

VAILLANT. S. 1727. Botanicon Parisiense.

S.M.PEROLD

MS. received: 1991-06-10.

188

Bothalia 22,2 (1992)

FABACEAE

A NEW SPECIES OF COEUD1UM (LIPARIEAE)

Of the twenty known species of the genus Coelidium

Vog. ex Walp., seventeen have white, pink or purple

flowers and only three are yellow-flowered (Granby 1980,

1987). A new species with ivory-yellow flowers was

discovered recently.

Coelidium vlokii Schutte & Van Wyk, sp. nov., C.

obtusilobo Granby valde affinis sed habitu valde maiore

robustiore, floribus maioribus cremeis (in C. obtusilobo

violaceis ), apice carinae (rostro) purpureo atque lobis

calycis acutis (non obtusis), glabris (in C. obtusilobo

sericeis ) differt.

TYPE. — Cape Province, 3323 (Uniondale): Uniondale

commonage, S aspect of Fortkoppie, (— CA), 19-09-1990,

Schutte 665 (PRE, holo.; BOL, JRAU, K, NBG, iso.).

Virgate multistemmed shrublet up to 1.0 x 0.8 m,

sprouting from a woody rootstock. Branches slender,

densely leafy; sericeous when young, glabrescent. Leaves

alternate, sessile, simple, erect, linear to narrowly oblong

with strongly involute margins, non-twisting, (2— )3(— 7)

mm long, up to 1 mm wide; apex acute; densely sericeous

adaxially, glabrescent abaxially. Stipules minute. Inflores-

cences bifloral, decussate, axillary, aggregated in terminal

clusters of up to 8 flowers. Bracts lanceolate, 1.5— 2.0 x

1.0 mm, glabrescent; margins thinly silky. Bracteoles

absent. Flowers sessile, 7—9 mm long, ivory-yellow with

purple keel tip; glabrous. Calyx 5 mm long, greenish-

yellow, tubular with prominent basal hypanthium; upper

two lobes fused higher up, glabrous; lateral and lower

sinuses of equal depth; lobes triangular, acute, margins

ciliate. Standard 7.0— 8.5 X 5.0 mm; lamina widely

obovate; apex emarginate; base somewhat indented; claw

± 2 mm long. Wing petals slightly longer than keel, 7

x 2 mm; lamina oblong to narrowly oblong, with distal

part imbricate, pocketed; sculpturing upper central, 2 rows

of intercostal lunae; claw ± 2 mm long. Keel petals 6.5

mm long, lamina half oblong-elliptic, auriculate, distinctly

pocketed; apex obtuse; claw ± 2.5 mm long. Stamens 6.5

mm long; monadelphous, 4 short and 6 long filaments

fused for ± */3 the length; anthers varying in size, alter-

nately dorsifixed and sub-basifixed. Pistil ± 7 mm long,

sessile; style slender, slightly upcurved, glabrous; ovary

uni-ovulate, densely sericeous. Young pods 8—9 X 4 mm,

ovate, sericeous. Figure 5.

Judged by morphological similarities, the closest relative

of C. vlokii is probably C. obtusilobum Granby, a species

known only from the type specimen collected on the

Kamiesberg in Namaqualand. C. vlokii differs from the

latter in the larger and more robust habit, the larger cream-

yellow flowers with only the tip of the keel purple (flowers

uniformly violet in C. obtusilobum ), the glabrous calyx

(sericeous in C. obtusilobum) and the acute calyx lobes

(distinctly obtuse in C. obtusilobum).

C. vlokii is known only from the type locality at Union-

dale in the southern Cape Province (Figure 6), where it

FIGURE 5. — Coelidium vlokii. A, flowering twig with young leaves, showing sessile flowers; B, mature leaves; C, leaf in transverse section,

showing involute margins; D, flower in lateral view; E, bract; F, calyx opened out with upper lobes to the left; G, standard petal; H, wing

petal; I, keel petal; J, stamens; K, anthers; L, pistil; M, young pod in lateral view (note that a short pedicel develops in the fruiting stage).

Bothalia 22,2 (1992)

189

FIGURE 6. — The known geographical distribution of Coelidium vlokii.

is locally common in Elytropappus-Eriocephalus veld on

clayish soil.

This distinct new species is named after Mr J.H.J. Vlok

(Cape Department of Nature and Environmental Conser-

vation), who has collected several rare and unknown

legumes in recent years.

CAPE. — 3323 (Uniondale): Uniondale commonage, S aspect of

Fortkoppie, (-CA), 19-09-1990, Schulte 665 (PRE, holo.; BOL, JRAU,

K, NBG, iso.), 19-09-1990, Schulte 661-664 (JRAU), 21-09-1991, Schulte

729 (JRAU); 1 km S of Uniondale, along road in poort, (— CA), 01-11-1990,

Van Tonder 79 (JRAU).

REFERENCES

GRANBY, R. 1980. A revision of the genus Coelidium (Liparieae-

Fabaceae). Opera Botanica 54: 1—47.

GRANBY, R. 1987. Coelidium flavum, a new species of Fabaceae—

Liparieae from the Cape Province. Nordic Journal of Botany 7:

51, 52.

A.L. SCHUTTE* and B-E. VAN WYK*

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

Johannesburg 2000.

MS. received: 1991-11-22.

FABACEAE

LESSERT1A SNEE UWBERGENSIS, A NEW SPECIES FROM THE MIDDELBURG DISTRICT OF THE CENTRAL CAPE PROVINCE

Lessertia sneeuwbergensis Germishuizen, sp. nov.,

L. cryptanthae Dinter affinis, sed longitudine calycis

florumque, magnitudine et forme fructus, floribus ex

axillis foliorum summorum ortis, atque seminis multis in

quoqe fructu differt.

TYPE. — 3124 (Hanover): Middelburg District, Gordon-

ville, Sneeuwberge, (—DA), 1952-10-28, Acocks 16536

(PRE, holo.).

Small, erect annual herb 30—200 mm high. Stems much

branched from base, yellow-brown, strigose, with scat-

tered white hairs. Taproot slender, simple or branched,

up to 200 mm long, up to 3 mm thick at base. Stipules

paired, obliquely ovate, acute, up to 2.0 mm long, white,

strigose along margins. Leaves variable; leaflets usually

solitary, terminal, rarely up to 4, in short-stalked, oppo-

site to subopposite pairs, linear, 2.0-12.0 (—14.0) x

1.0— 1.5 mm, acute or obtuse at apex, obtuse at base,

strigose, with white hairs on both surfaces; petioles up

to 65 mm long, strigose, with scattered white hairs.

Inflorescence few-flowered, pedunculate axillary raceme,

bracteate. Bracts broadly ovate, acute, strigose, brown hairs

intermixed with white hairs, especially at apex and along

margins; bracteoles present, small, up to 0.5 mm long.

Flowers small, flower colour not known. Calyx

campanulate, 3-4 mm long, subequal, 5-toothed, densely

brown, strigose, with a few white hairs intermixed. Vexil-

lum broadly obovate, shallowly bilobed, up to 9 x 6 mm,

glabrous. Wing petals narrowly elliptic, shorter than keel,

up to 8 x 2 mm, with little sculpturing. Keel petals oblong

obtuse, up to 9 x 4 mm; auricle and pocket absent.

Staminal sheath up to 8 mm long, persistent in fruit;

anthers uniform. Style hooked, with conspicuous tuft of

white hairs behind stigma, persistent in fruit. Pods inflated,

broadly oblong ovoid, 14—24 x 9—12 mm, yellow-brown,

sometimes mottled with purple, glabrous, conspicuously

veined. Seeds broadly reniform or ‘Lftshaped, yellow-

brown, slightly wrinkled, up to ± 17 per pod. Figure 7.

CAPE.— 3124 (Hanover): Oppermanskraal, Sneeuwberge, (—BA),

1952-10-28, Acocks 16516 (PRE); Gordonville, Sneeuwberge, (—DA),

FIGURE 7. — Holotype of Lessertia sneeuwbergensis, Acocks 16536.

190

Bothalia 22,2 (1992)

FIGURE 8. — Distribution of Lessertia sneeuwbergensis.

1952-10-28, Acocks 16536 (PRE); Lootsberg Pass, (—DC), 1947-03-28,

Acocks 13547 (PRE).

Lessertia sneeuwbergensis is endemic to the Sneeuw-

berge and Lootsberg of the Middelburg District of the

central Cape Province (Figure 8). Found mainly in Karroid

Merx-muellera Mountain Veld of upper southern slopes.

Fruiting takes place between October and March.

The specific epithet sneeuwbergensis is derived from

the locality in which this new Lessertia species was

collected. Acocks collected the type species on the

southern slopes of the Sneeuwberg Mountain Range.

In 1975, when R. Dahlgren was studying material for

a revision of the genus Wiborgia, he thought the specimen

of Acocks 13547 belonged to the genus Lebeckia because

of its unusual leaves. However, it belongs to the genus

Lessertia because of its membranous inflated fruits and

the presence of a tuft of hairs around the stigma.

Lessertia sneeuwbergensis and L. cryptantha closely

resemble one another, but are separated by a number of

morphological differences (see Table 1) and by their

distribution. L. cryptantha occurs in the Klinghardts

Mountains in central Namibia and L. sneeuwbergensis

occurs in the Sneeuwberg Mountain range and Lootsberg

in the central Cape Province.

TABLE 1. — Comparison of morphological characters in Lessertia

sneeuwbergensis and L. cryptantha

ACKNOWLEDGEMENTS

My thanks go to the Director and staff (particularly

Dr H.F. Glen for translating the diagnosis into Latin and

Mrs A.J. Romanowski for the photographing of the type)

of the National Botanical Institute.

G. GERMISHUIZEN

MS. received: 1991-10-18.

FABACEAE

CYCLOPIA SQUAMOSA (PODALYRIEAE), A NEW SPECIES FROM THE SOUTHWESTERN CAPE PROVINCE

The genus Cyclopia Vent, is currently under revision.

A study of the morphological variation within the genus

indicated the existence of an undescribed species, quite

unlike any of the 26 taxa recognized by Kies in her treat-

ment of the genus in 1951.

Cyclopia squamosa A.L. Schutte, sp. nov., C. sessi-

liflorae Eckl. & Zeyh. affinis sed petiolis prominentibus

persistentibus, internodiis perbrevibus, minus quam 1 mm

longis (petioli inconspicui et intemodia 3 mm vel longiora

in C. sessiliflora et speciebus omnibus aliis), foliolis linear-

ibus marginibus valde revolutis (foliolis oblongo-ellipticis

marginibus tantum subrevolutis in C. sessiliflora) , calyce

bracteisque maioribus (valde minoribus in C. sessi-

liflora) atque pedicellis longioribus (valde brevioribus in

C. sessiliflora) differt.

TYPE. — Cape Province, 3319 (Worcester): Paarl Dis-

trict, Wemmershoek Peak, on cliff and at base of cliff,

on steep southern slope, (— CC), 25-10-1981, Esterhuysen

35695 (BOL, holo.; C, K, S, iso.).

Woody shrubs. Branches rigid, internodes short, less

than 1 mm long, covered with prominent partially over-

lapping persistent petioles, appearing scale-like, + gla-

brous; young twigs densely leafy, sparsely villous. Leaves

alternate, trifoliolate, (8 — )14 — 17( — 22) mm long, sparsely

villous, glabrescent. Leaflets linear, (6 — )12 — 15 (—20) mm

long, less than 1 mm wide, margins strongly revolute,

caducous; apex acute, mucronulate; base distinctly

pulvinate. Petioles flattened, up to 3 X 2 mm, persistent.

Stipules less than 1 mm long, fused with petiole. Inflores-

cences axillary, single-flowered. Bracts paired, subequal,

Bothalia 22,2 (1992)

191

FIGURE 9. — Cyclopia squamosa. A, flowering branchlet showing the almost hidden flowers, conspicuous persistent petioles and short internodes;

B, mature leaf, abaxial view; C, leaflet in transverse section (enlarged) to show strongly revolute margins; D, flower in lateral view, showing

paired bracts; E, calyx opened out with upper lobes to the left; F, standard petal; G, wing petal; H, keel petal; I, anthers; J, pistil; K,

young pod in lateral view. Scales in mm.

lanceolate, 6 mm long, keeled, outer surface glabrous,

inner surface pubescent, margins thinly silky. Bracteoles

absent. Flowers yellow, 15—17 mm long, almost hidden

between leaves, glabrous. Pedicels 6—7 mm long, gla-

brous. Calyx intrusive at base, 8—9 mm long, glabrous;

upper two lobes fused higher up, carinal lobe longer than

the other four, keeled; lobes triangular-acute to lanceolate-

acuminate, margins ciliate. Standard 15.0—15.5 x 11—12

mm; lamina obovate to suborbicular; apex emarginate;

base cuneate; claw + 3 mm long. Wing petals slightly

longer than keel, 15.5-17.0 x 4. 5 -5.0 mm; lamina oblong

to narrowly oblong, with distinct pocket ± 6 mm long;

claw + 3.5— 4.0 mm long. Keel petals 14.5—16.5 x

3.5-4. 5 mm, lamina oblong-elliptic, with distinct pocket

± 6 mm long; apex rostrate; claw 3.5 -5.0 mm long.

Stamens 10, filaments free; anthers distinctly dimorphic,

5x5, alternately short dorsifixed and long basifixed. Pistil

sessile, 14—15 mm long; ovules 3—7; ovary glabrous; style

slender, curved inwards at the apex, glabrous. Young pods

13 x 6 mm, ovate, glabrous. Figure 9.

This species is related to C. sessiliflora Eckl. & Zeyh.

but differs in the presence of prominent petioles and very

short intemodes of less than 1 mm long (petioles incon-

spicuous and intemodes 3 mm or longer in C. sessiliflora

and elsewhere in the genus), the linear leaflets with

strongly revolute margins (leaflets oblong-elliptic with

margins only slightly revolute in C. sessiliflora), the larger

calyx and bracts, and longer pedicels (calyx 5 mm or

shorter, bracts up to 2 mm long and pedicels 2—3 mm

FIGURE 10. — The known geographical distribution of Cyclopia squa-

mosa, *.

192

Bothalia 22,2 (1992)

long in C. sessiliflora). In the latter the standard is

orbicular and very shortly clawed, versus obovate to sub-

orbicular with a longer claw in C. squamosa.

C. squamosa is a rare legume known only from the

type locality on the Wemmershoek Mountains in the south-

western Cape Province (Figure 10). The conspicuous

persistent petioles and very short intemodes, giving

the branches a scaly appearance, are useful diag-

nostic characters for this species, hence the specific

epithet.

CAPE. —3319 (Worcester): Paarl District, Wemmershoek Peak, on cliff

and at base of cliff, on steep southern slope, (-CC), 25-10-1981,

Esterhuysen 35695 (BOL, holo.; C, K, S, iso.), 04-04-1982, Esterhuysen

35764 (BOL, C).

ACKNOWLEDGEMENTS

I am grateful to Dr H.F. Glen (National Botanical

Institute, Pretoria) for the Latin translation and to Prof.

B-E. van Wyk for commenting on the manuscript. The

directors and staff of the mentioned herbaria are also

thanked for the loan of specimens.

REFERENCES

KIES, P. 1951. Revision of the genus Cyclopia and notes on some other

sources of bush tea. Bothalia 6: 161—176.

A.L. SCHUTTE*

* Department of Botany, Rand Afrikaans University, P.O. Box 524,

Auckland Park, Johannesburg 2006.

MS. received: 1992-03-18

STILBACEAE

STILBE VERTICILLATA , THE CORRECT NAME FOR THE SPECIES PREVIOUSLY KNOWN AS STILBE MUCRONATA

During the course of revising the Cape endemic family

Stilbaceae, it has come to my attention that the name of

the species currentiy known as Stilbe mucronata N.E. Br.

(Pearson 1901; Gibbs Russell et al. 1987), is antedated by

an earlier name. Moldenke’s combination, Stilbe verticil-

lata (Ecklon & Zeyher) Moldenke made in 1948 is the

correct name for this species but has been overlooked until

now. Stilbe chorisepala Suesseng. , currently upheld as a

distinct species (Gibbs Russell et al. 1987), is here reduced

to synonymy. The full synonymy is as follows:

Stilbe verticillata (Eckl. & Zeyh.) Moldenke in

Phytologia 2 : 474 (1948).

Trichocephalus verticillatus Eckl. & Zeyh.: 131 (1835). Type: In

lateralibus montium prope Palmietrivier, supra Grietjiesgat, June, Alt.

4, Ecklon & Zeyher 1003 (SAM, iso.!).

Stilbe mucronata N.E. Br.: t. 2526 (1897); Pearson: 184 (1901). Type:

In declivibus montium Houwhoek, April 1895, 1400 ped.. Bolus 8409

(K, lecto.! here designated).

Stilbe chorisepala Suesseng.: 56 (1950). Type: Franschhoek Pass,

17-11-1946, S. Rehm (M, holo.!).

Stilbe mucronata N.E. Br. var. cuspidata H.H.W. Pearson: 184 (1901).

Stilbe verticillata (Eckl. & Zeyh.) Moldenke var. cuspidata (H.H.W.

Pearson) Moldenke: 474 (1948). Type: Zwartberg, Caledon, 3000 ft,

Dec., Bolus s.n. (K, holo.!).

REFERENCES

BROWN, N.E. 1897. Stilbe mucronata. In Hooker’s leones Plantarum

6: t. 2526.

ECKLON, C.F. & ZEYHER, K.L.P. 1835. Enumeratio plantarum africae

australis: 131. Hamburg.

GIBBS RUSSELL, G.E. etal. 1981. List of species of southern African

plants. Edn 2, Part 2. Memoirs of the Botanical Survey of South

Africa No. 56: 168.

MOLDENKE, H.N. 1948. Notes on new and noteworthy plants. V.

Phytologia 2 : 474.

PEARSON, H.H.W. 1901. Verbenaceae. In W.T. Thiselton-Dyer, Flora

capensis 5: 184. Lovell Reeve, London.

SUESSENGUTH, K. 1950. Stilbe chorisepala. Mitteilungen der

Botanischen Staatssammlung, Munchen Heft 2: 56.

J.P. ROURKE*

* National Botanical Institute, Kirstenbosch, Private Bag XI, Claremont

7735, South Africa.

MS. received: 1992-05-19.

GESNERIACEAE

NOTES ON THE GENUS STREPTOCARPUS

Within the genus Streptocarpus the rosulate species com-

prising the rexii aggregate sensu Hilliard & Burtt (1971)

include S. rexii (Hook.) Lindl., S. primulifolius Gandoger,

S. cyaneus S. Moore and S. parviflorus Hook. f. A new

species recently discovered in the eastern Transvaal is

included within the aggregate. In addition 5. caeruleus

subsp. longiflorus is raised to specific status.

Streptocarpus fasciatus T. Edwards & C. Kunhardt,

sp. nov. S. cyaneo affinis sed corolla leviter dorsiventraliter

compressa et sine taenia flava in fauce.

TYPE. — South Africa, eastern Transvaal, 25 km from

Nelspruit on the Komatipoort Rd, C. Kunhardt & J. Kluge

s.n. (NU, holo.; E, PRE, iso.).

Robust rosulate perennial. Leaves oblong, reaching 290

x 100 mm, lamina slightly decurrent, pilose, margins

crenate; petiole up to 20 mm. Inflorescence up to

12 -flowered; peduncle to 300 mm tall, pilose. Sepals

narrowly deltoid, 6.5 x 1.5 mm, pilose, tips red-brown.

Corolla 70—76 mm long; tube 50 mm long, lower half

linear, 5 mm broad, upper half expanding and slightly

dorsiventrally compressed, 20 x 11 mm at the throat,

pilose externally and inside along corolla roof; limb

bilabiate, upper lobes 12 X 14—15 mm, rounded, with a

medial violet streak in each; lower lip 22-24 mm long,

violet-streaked, lobes 15—16 X 15 mm (Figure 11).

Stamens arising from waist of corolla; filaments 9—10 mm

long, white, glabrous basally but with stalked glandular

trichomes near connective; anthers 3 x 1 mm, pale

mauve, connectives deep violet; posterior staminode

Bothalia 22,2 (1992)

193

FIGURE 11. — S. fasciatus, flowering plant, X 0.4.

minute, within a dorsal channel, lateral staminodes 2 mm

long. Pollen prolate, 20—23 x 10—12 fin i, tricolporate,

poles apocolpoid, exine minutely scabrate. Ovary ± 24

mm long, with dense appressed filiform trichomes; style

12 mm long, dorsiventrally compressed, with stalked

glandular trichomes; stigma bifid, white, peripherally

papillate. Fruit 100—120 x 2 mm.

In S. cyaneus the occurrence of a yellow bar in the throat

is remarkably constant despite variation in the ground

colour and dimensions of the corolla. This yellow bar is

entirely lacking in S. fasciatus which is at the eastern limit

of S. cyaneus. Nearby populations of S. cyaneus have short

corolla tubes with a pink ground colour and 2—4 flowers

per inflorescence.

S. cyaneus is a notoriously difficult species to define.

This is due partly to the inclusion of diverse forms within

the species and possibly due to introgression with

S. parviflorus in the north. Hilliard & Burtt (1971) recog-

nise a degree of geographic patterning in the variation

but state that there is seldom correlation of several charac-

ters over a wide area. As a consequence they reduced

S. polackii B.L. Burtt and S. junodii Beauv. to synonymy.

S. fasciatus does not conform to these previously recog-

nised entities.

The specific epithet fasciatus refers to the well-defined

longitudinal stripes which mark the corolla lobes. Plants

are very floriferous with individual peduncles often

supporting 12 flowers, a feature shared with S. parviflorus.

Corolla size approaches the upper limits found in S.

cyaneus whereas the slight dorsiventral compression of the

corolla is reminiscent of the situation found in S. gardenii

Hook.

S. fasciatus has only been recorded from an enclave of

woodland in the Crocodile River Gorge. The specimen

illustrated was found growing in association with S. pole-

evansii Verdoom under the protection of granite boulders.

Flowers are produced from late summer into autumn while

in S. cyaneus the main flowering period is spring to sum-

mer. The discovery of the species is recounted by Kunhardt

(1991).

Streptocarpus caeruleus Hilliard & Burtt in Strep-

tocarpus, an African plant study: 387 (1971). Type. — N.

Transvaal, Blaauwberg, cult. R.B.G. Edinburgh (e semi-

nibus R. Story 6512), C. 3824 (E, holo.; NU! iso.).

TRANSVAAL. — 2329 (Pietersburg): Blouberg, middle buttress,

(— AA), Stirton, Edwards & Venter 12646 (NU); Soutpansberg, Farm

Lejuma, (— AB), Hilliard 4760 (NU).

For discussion see S. longiflorus.

Streptocarpus longiflorus (Hilliard & Burtt) T.

Edwards, stat. nov. Type. — N. Transvaal, Blaauwberg,

cult, in R.B.G. Edinburgh (e seminibus Burtt 2918 ex hort.

C. Thompson) C. 4895 (E, holo.; NU! iso.).

Streptocarpus caeruleus Hilliard & Burtt subsp. longiflorus Hilliard

& Burtt: 388 (1971).

Rosulate perennial. Leaves 5-10, oblong, reaching 300

x 70 mm, lamina slightly decurrent, pilose, margins

crenate. Inflorescence 5— 15(— 20)-flowered; peduncle to

200 mm tall, pilose. Sepals narrowly deltoid, 3.5— 4.0 x

1.5 mm, pilose, red-brown. Corolla 38-47 mm long; tube

26—32 mm long, lower half linear, 5 mm broad, upper

half expanding, 15—18 mm at the throat, pilose externally

and inside along corolla roof; limb bilabiate, upper lobes

8-10 x 10—12 mm, rounded; lower lip 15-18 mm long,

lobes 8—10 x 10-12 mm. Stamens arising from waist of

corolla; filaments 9—13 mm long, white, glabrous basally

but with stalked glandular trichomes near connective;

anthers 2x1 mm, pale mauve, connectives deep violet;

posterior staminode minute, within a dorsal channel,

lateral staminodes 2 mm long. Pollen prolate, 24—29 x

10—12 pm, tricolporate, poles apocolpoid, exine minutely

scabrate. Ovary ± 20 mm long, with dense appressed

filiform trichomes; style 18 mm long, dorsiventrally com-

pressed, with stalked glandular trichomes; stigma bifid,

white, peripherally papillate. Fruit 90-110 x 3 mm,

pendulous when mature.

TRANSVAAL. — 2329 (Pietersburg): Bloubeig, upper slopes, Edwards,

Stirton & Venter 979 (NU).

Plants of S. caeruleus are weakly rosulate, seldom com-

prising more than four leaves, the corolla is short ((18mm)

and mauve with two yellow spots flanking the lower medial

lobe (Figure 12B), the fruits are held erect and seldom

exceed 50 mm in length. In contrast, plants of S. longi-

florus have rosettes of up to 10 leaves, their corollas are

38—47 mm long and have a y-shaped yellow bar in the

throat (Figure 12A), and the pendulous capsules are

90—100 mm long.

Burtt (1962: 42) suggested that the long and short-

flowered plants of ‘this species’ may represent floral dimor-

phism. This idea was rejected (Hilliard & Burtt 1971) due

to the intermediate flowers produced by FI hybrids. To

accommodate this genetic distinction Hilliard & Burtt ( l.c .)

placed the taxa together under S. caeruleus but

distinguished two subspecies S. caeruleus subsp. caeruleus

and S. caeruleus subsp. longiflorus. On a recent trip to

the Blaauwberg (Blouberg) mixed populations of these taxa

were frequently encountered but no hybrids were seen.

Different breeding systems were apparent in cultivated

specimens. The type subspecies is predominately autoga-

mous and good seed set occurs in die absence of pollina-

tors. There is little spatial separation between the anthers

194

Bothalia 22,2 (1992)

FIGURE 12. — Dissections of the

corollas of: A, S. longiflorus',

B, S. caeruleus.

and the stigma. Often the two fused fertile anthers split

apart in the bud, thus achieving pollination before anthesis.

In S. longiflorus the stigma extends about 3 mm above

the anthers and seed is seldom set on cultivated plants.

Such breeding barriers are rare in Streptocarpus. Thus by

virtue of their distinctive morphology, distributional

overlap, intermediate FI hybrids and the occurrence of

breeding barriers between these taxa, subsp. longiflorus

is raised to specific status.

S. longiflorus is endemic to the Blaauwberg. Plants are

found in more exposed habitats than those occupied by

S. caeruleus, frequently well above the tree line in the

shade of rocks. Bees were seen visiting the flowers, but

whether they are effective pollinators was not ascertained.

In most aspects 5. longiflorus resembles S. cyaneus, which

we consider to be its closest ally. These species are

allopatric. The nearest populations of S. cyaneus on the

Soutpansberg are distinguished by the simple yellow bar

on the corolla floor.

ACKNOWLEDGEMENTS

Sincerest thanks are extended to the late Chris Kunhardt

who collected the type of S. fasciatus and to his son

Martin who provided the photographs used in this paper.

The Natal University Research Fund is thanked for its

financial support and the Natal University Electron Micro-

scope Unit is thanked for their assistance in viewing

pollen. Mr D. Pike is acknowledged for his assistance with

the Latin diagnosis. The referees are thanked for their

comments.

REFERENCES

BURTT, B.L. 1962. Studies in the Gesneriaceae of the Old World, miscel-

laneous transfers and new species. Notes of the Royal Botanic

Gardens Edinburgh 24: 41-49.

HILLIARD, O.M. & BURTT, B.L. 1971. Streptocarpus, an African plant

study. Natal University Press, Pietermaritzburg.

KUNHARDT, M. 1991. A special Streptocarpus. The Gloxinian 41: 7—9.

T.J. EDWARDS* C. KUNHARDT and S. VENTER**

* F.R.D. Unit for Plant Growth and Development, Department of

Botany, University of Natal, P.O. Box 375, Pietermaritzburg 3200.

** Botany Department, University of the North, P/Bag 1106, Sovenga 0727.

MS. received: 1992-02-13.

Bothalia 22,2 (1992)

195

GERANIACEAE

TYPIFICATION OF PELARGONIUM SECTION POLYACT1UM

The name Polyactium DC. originated in sectional rank

(De Candolle 1824: 655) and was raised to generic rank

by Ecklon & Zeyher (1835: 65). Harvey (1860: 272) again

reduced it to sectional rank, a status which it retains to

the present.

Farr et al. (1979: 1386) stated that a type had not been

designated for the name Polyactium. This is incorrect, as

Knuth (1912: 317) indicated Pelargonium lobatum (Burm.

f.) L’Hdrit. as the type in his key to the section of the ge-

nus. We consider this to be an arbitrary designation, con-

trary to Art. 8.1(b) of the I.C.B.N. (Greuter et al. 1988),

as P. lobatum did not form part of the protologue of

Polyactium. The protologue consists of a single element,

Pelargonium multiradiatum Wendl., which therefore has

the status of type species.

In accordance with Art. 10.2 of the I.C.B.N. (Greuter

et al. 1988), we therefore propose that Pelargonium

multiradiatum be accepted as the type species of Pelargo-

nium section Polyactium DC.

ACKNOWLEDGEMENT

Our research on the taxonomy of the genus Pelargo-

nium is financed by the University of Stellenbosch, and

the South African Foundation for Research Development.

REFERENCES

DE CANDOLLE, A.R 1824. Prodromus systematis naturalis regni

vegetabilis ... 1. Treuttel & Wiirtz, Paris.

ECKLON, C.F. & ZEYHER, K.L. 1835. Enumeratio plantarum africae

australis extratropicae ... 1. Perth & Besser, Hamburg.

FARR, E.R., LEUSSINK J.A. & STAFLEU, F. 1979. Index nominum

genericorum (plantarum) 3. Regnum vegetabile 102.

GREUTER, W., BURDET, H.M., CHALONER, W.G., DEMOULIN,

V., GROLLE, R., HAWKSWORTH, D.L., NICOLSON, D.H.,

SILVA, P.C., STAFLEU, F.A., VOSS, E.G. & McNEILL, J. 1988.

International code of botanical nomenclature adopted by the Four-

teenth International Botanical Congress, Berlin, July— August

1987. Regnum vegetabile 118.

HARVEY, W.H. 1860. Pelargonium. In W.H. Harvey & O. Sonder, Flora

capensis 1: 259—308. Hodges Smith, Dublin.

KNUTH, R. 1912. Pelargonium. In A. Engler, Das Pflanzenreich 4, 129,

53: 316-545.

P. VORSTER* and G.L. MAGGS*+

* Botany Department, University of Stellenbosch, Private Bag X5018,

Stellenbosch 7599.

+ Present address: Government Herbarium, Private Bag 13184, Windhoek,

Namibia.

MS received: 1992-04-24.

BRYOPHYTA

NEW AND INTERESTING RECORDS OF MOSSES IN THE FLORA OF SOUTHERN AFRICA AREA:

3. MISCELLANEOUS ACROCARPOUS TAXA

Additional new records (see Van Rooy & Perold 1990,

1992) identified from geographical regions referred to in

the first two fascicles of the moss Flora of southern Africa

(Magill 1981, 1987) are reported here.

FISSIDENTACEAE

Fissidens fasciculatus Homsch. (Magill 1981: 67)

Eastern Transvaal (2430 DD: Rehmarm 593c; Stirton 7036) and Natal

(locality not precise: H hger sub Sim 7759, Wager PRE-CH 258, PRE-

CH 11976).

DICRANACEAE

Campylopus

introflexus (Hedw.) Brid. (Magill 1981: 150)

Northern Cape (2822 BA: Magill 6417, 6419).

pilifer Brid. (Magill 1981: 152)

Namibia (2217 CA: Gibbs Russell & Smook 5458).

Leucobryum acutifolium (Mitt.) Card. (Magill 1981: 156)

Transkei (3130 AA: Abbott 4185; Smook 6159. 3130 AC: Van Rooy

1776, 1778, 1787. 3129 BC: Van Rooy 1867, 1888, 1892).

CALYMPERACEAE

Octoblepharum albidum Hedw. (Magill 1981: 163)

Northern Transvaal (2329 AB: Glen 2198 ) and Transkei (3130 AA:

Smook 6169).

POTTIACEAE

Phascum leptophyllum C. Mull. (Magill 1981: 204)

Zululand (2832 CA: Glen 2746).

Tortula fragilis Zay/. (=T.schmidii (C.Mull.) Broth.) (Magill 1981: 215)

Namibia (1918 AD: Volk 81/133 ) and central Transvaal (2427 BD:

Glen 2599)).

Hyophila baginsensis C. Mull. (Magill 1981: 230)

Namibia (2017 AC: Schelpe 4813, 4815, 4816).

Didymodon xanthocarpus (C. Mall.) Magill (Magill 1981: 235)

Southern Transvaal (2628 AD: Wager sub Sim 7833; Wager PRE-

CH 34) and Natal (precise locality not known: H tiger PRE-CH 11857).

Barbula acutata C. Mall. (Magill 1981: 239)

Natal (2828 DB: Magill 6706, 6719).

Oxystegus cylindricus (Brid.) Hilp. (Magill 1981: 259)

Orange Free State (2828 DB: Magill 6596, 6628).

GIGASPERMACEAE

Oedipodiella australis (Wager & Dix.) Dix. (Magill 1987: 303)

Lesotho (2928 BD: Van Rooy 3161).

BARTRAMIACEAE

Bartramia hampeana C. Mall. (Magill 1987: 417)

Northern Transvaal (2329 AB: Glen 2199).

REFERENCES

MAGILL, R.E. 1981. Bryophyta. In O.A. Leistner, Flora of southern

Africa. Part 1, Mosses, Fascicle 1, Sphagnaceae— Grimmiaceae.

National Botanical Institute, Republic of South Africa.

MAGILL, R.E. 1987. Bryophyta. In O.A. Leistner, Flora of south-

ern Africa. Part 1 Mosses, Fascicle 2, Gigaspermaceae-Bartrami-

aceae. National Botanical Institute, Republic of South Africa.

VAN ROOY, J. & PEROLD, S.M. 1990. New and interesting records

of mosses in the Flora of southern Africa area: 1. Sphagnaceae -

Grimmiaceae. Bothalia 20: 211—213.

VAN ROOY, J. & PEROLD, S.M. 1992. New and interesting records

of mosses in the Flora of southern Africa area: 2. Gigasper-

maceae-Bartramiaceae. Bothalia 22 : 37.

J. VAN ROOY* and S.M. PEROLD*

MS. received: 1992-07-07.

196

Bothalia 22,2 (1992)

BRYOPHYTA

NOTES ON THE MOSS FLORA OF ZIMBABWE

Macrocoma (Homsch. ex C. Miill.) Grout

In recently published checklists by Kis (1985) and Best

(1990), three species of Macrocoma are reported from

Zimbabwe. The reports of M. lycopodioides (Schwaegr.)

Vitt and M. pulchella (Homsch.) Vitt are based on T.R.

Sim’s treatments of southern African bryophytes (Sim

1926, 1932). However, Vitt (1980a, 1980b) revised the

genus Macrocoma and together with R.E. Magill (Magill

& Vitt 1981) studied the phytogeography and ecology of

the African members of this genus. Vitt (1980a) noted that

Sim (1926) did not fully understand the variability

expressed by the southern African species. Magill & Vitt

(1981) found that M. lycopodioides occurs only south of

FIGURE 13. — Distribution of Tayloria isleana.

the Limpopo River and M. pulchella is restricted to the

southwestern Cape Province of South Africa. These two

species should therefore be excluded from the moss flora

of Zimbabwe. Only M. tenue (Hook. & Grev.) Vitt subsp.

tenue, the most widely distributed of the African taxa, is

known from Zimbabwe (Magill & Vitt 1981).

Tayloria Hook.

A Vahrmeijer collection from Castle Beacon, Mount

Vumba, Zimbabwe ( Vahrmeijer PRE-CH13514) , was recent-

ly identified at PRE as Tayloria isleana (Besch.) Broth.

It is a new record for Zimbabwe. The occurrence of T.

isleana in Zimbabwe is not unexpected as the species was

previously known from East Africa (Kis 1985), Natal in

South Africa (Magill 1987), Madagascar (Crosby et al.

1983) and the Mascarene Islands (Figure 13). Tayloria

orthodonta (P. Beauv.) Wijk & Marg. is also known from

Zimbabwe and occurs in tropical and subtropical Africa

as far south as the northern Transvaal in South Africa and

the Mascarene Islands.

REFERENCES

BEST, E.B. 1990. The Bryophyta of Zimbabwe — an annotated check-

list. Kirkia 13: 293-318.

CROSBY, M.R., SCHULTZE-MOTEL, U. & SCHULTZE-MOTEL,

W. 1983. Katalog der Laubmoose von Madagaskar und den

umliegenden Inseln. Willdenowia 13: 187—255.

KIS, G. 1985. Mosses of southeast tropical Africa. Institute of Ecology

and Botany of the Hungarian Academy of Sciences, Vacratot,

Hungary.

MAGILL, R.E. 1987. Bryophyta. In O.A. Leistner, Flora of southern

Africa. Part 1, Mosses, Fascicle 2, Gigaspermaceae-Bartrami -

aceae. Botanical Research Institute, South Africa.

MAGILL, R.E. & VITT, D.H. 1981. The phytogeography and ecology

of Macrocoma (Orthotrichaceae, Musci) in Africa. Bothalia 13:

463-466.

SIM, T.R. 1926. The Bryophyta of South Africa. Transactions of the

Royal Society of South Africa 15: 1—475.

SIM, T.R. 1932. South African Bryophyta. Further notes. Transactions

of the Royal Society of South Africa 20: 15—31.

VITT, D.H. 1980a. The nomenclature and taxonomy of Macrocoma

lycopodioides (Schwaegr.) Vitt. Journal of Bryology 11: 219—229.

VITT, D.H. 1980b. The genus Macrocoma I. Typification of names and

taxonomy of the species. Bryologist 83: 405—436.

J. VAN ROOY*

MS. received: 1992-07-07.

LILIACEAE/ASPHODELACEAE

LECTOTYPIFICATION OF APICRA JACOBSENIANA (ALOOIDEAE)

An uncommon, but very distinctive, alooid plant

(Asphodelaceae: Alooideae) from the southwestern Cape

Province, South Africa, has been placed in several

different genera, namely Apicra Haw. non Willd., Poell-

nitzia Uitewaal, Haworthia Duv. and Aloe L. Contempo-

rary consensus is that it is properly placed in the genus

Poellnitzia (Smith 1991), typified by P. rubriflora (L. Bol.)

Uitewaal (1940). One year before Poellnitzia was estab-

lished for the aberrant Apicra rubriflora L. Bol. (1920),

Von Poellnitz (1939) described a second species of Apicra,

namely A. jacobseniana, which he regarded as closely

related to A. rubriflora. The species, which was named

in honour of Dr Hermann J. H. Jacobsen, German

horticulturist and curator of the Kiel Botanical Garden,

should, however, not be confused with Pfaworthia

jacobseniana V. Pbelln. (1937), which is currently included

in the synonymy of H. glauca var. herrei (V. Poelln.)

Bayer.

Current indications are that the name Apicra jacob-

seniana (which was never transferred to Poellnitzia) should

be included in the synonymy of Poellnitzia rubriflora

Bothalia 22,2 (1992)

197

FIGURE 14. — Specimen of Poellnitzia rubriflora,

collected 5 km W of Bonnievale, southwestern

Cape Province, which agrees in all respects with

the description of Apicra jacobseniana.

(Smith 1991; Figure 14). It has also been omitted from the

synoptic work of Gibbs Russell et al. (1985). In this paper

the affinity of Apicra jacobseniana is established and the

name is lectotypified.

Material on which Von Poellnitz (1939) based the name

Apicra jacobseniana was introduced to Europe in the

1930’s by Wilhelm Triebner. He gave the locality as

‘Worcester District’. The genus Poellnitzia, under which

A. jacobseniana is currently classified, is endemic to the

Robertson-McGregor-Bonnievale region of the south-

western Cape (Smith & Van Wyk 1991; Wood 1991).

Triebner was notorious for providing insufficient and

unreliable details of his collecting localities and those of

his correspondents (Bayer 1982), and he probably took

‘Worcester District’ to include the geographical distribu-

tion range of Poellnitzia. Mrs F. N. Morris of Oudtshoom,

an avid succulent plant collector and regular correspon-

dent of Triebner (Codd & Gunn 1985), is accredited as

having been the first to collect Apicra jacobseniana in the

vicinity of Worcester.

The type collection of A. jacobseniana ( Triebner 34)

was probably sent directly to Kiel where it was cultivated

in the Botanical Garden. However, no herbarium specimen

of the material could be traced. If duplicates of the type

collection were kept at Von Poellnitz’ home, they were des-

troyed when the place was bombed early in 1945. The rem-

nants of Von Poellnitz’ material were transferred from

Oberlodla to Berlin-Dahlem by the late Prof. Dr E.

Werdermann (Werdermann 1949; Obermeyer 1964), but

no material of Apicra jacobseniana could be found in B

(Dr Th. Raus pers. comm.).

Von Poellnitz did not cite any additional specimens when

he established A. jacobseniana, but he accompanied the

description with an illustration of a sterile specimen. This

illustration is here chosen as the lectotype:

TYPE. — J.K.L.A. von Poellnitz, Apicra jacobseniana

v. P., Kakteenkunde (1939): 95, unnumbered plate, lecto.

here designated (icono.).

ACKNOWLEDGEMENTS

I am grateful to Dr Th. Raus, curator of Liliaceae at

B, for providing useful documentation.

REFERENCES

BAYER, M.B. 1982. The new Haworthia handbook. National Botanic

Gardens of South Africa, Kirstenbosch, Cape Town.

BOLUS, H.M.L. 1920. Novitates Africanae: Apicra rubriflora L. Bolus

(Liliaceae -Aloineae). Annals of the Bolus Herbarium 3: 1—14.

CODD, L.E. & GUNN, M.D. 1985. Additional biographical notes on

plant collectors in southern Africa. Bothalia 15: 631—654.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK, L.

1985. List of species of southern African plants, edn 2, part 1.

Cryptogams, gymnosperms, monocotyledons. Memoirs of the

Botanical Survey of South Africa No. 51.

GUNN, M.D. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

OBERMEYER, A. A. 1964. The South African species of Anthericum,

Chlorophytum and Trachyandra. Addenda et Corrigenda. Bothalia

8: 147, 148.

SMITH, G.F. 1991. Contributions to the systematics of selected genera

of the Alooideae (Asphodelaceae) . Ph.D. (Botany) dissertation.

University of Pretoria.

SMITH, G.F. & VAN WYK, B-E. 1991. Generic relationships in the

Alooideae (Asphodelaceae). Taxon 40: 557—581.

UITEWAAL, A.J.A. 1940. Een nieuw geslacht der Aloineae. Succulenta

22: 61-64.

VON POELLNITZ, J.K.L.A. 1937. Six new species of Haworthia Duval.

Desert Plant Life 9: 102, 103.

VON POELLNITZ, J.K.L.A. 1939. Apicra jacobseniana v.P. Kakteen-

kunde (1939): 95.

WERDERMANN, E. 1949. Aufbau und Schicksal der Dahlemer Kakteen-

sammlungen. Sukkulentenkunde 3: 324—338.

WOOD, J. 1991. Threats facing the Robertson karoo. Veld & Flora 77:

16, 17.

G.F. SMITH*

* Department of Plant and Soil Sciences, Potchefstroom University for

Christian Higher Education, Potchefstroom 2520.

MS. received: 1992-03-25.

Bothalia 22,2: 199-204 (1992)

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

HI. Comparative sporangium and spore morphology

J.P. ROUX*

Keywords: Mohria , morphology, sporangium, spores

ABSTRACT

The genera Mohria and Anemia (Anemiaceae: Pteridophyta) can be separated on both their sporangia and spores. In Mohria

the capsule is globose with an apical annulus but in Anemia it is ovate-globose to cylindrical with a subapical annulus. The

spores of both genera are radially symmetrical, tetrahedral and trilete with near parallel muri. The exinal sculpture in Mohria

is cicatricose and in Anemia it can be canaliculate or cicatricose. In both genera the mural sets anastomose to form common

muri that extend from near the distal pole to the equatorial radial region. The muri in Mohria are hollow and differ from

those in Anemia which are solid or microporate. Supramural sculpturing in Mohria and Anemia is perinous. Spores of the

other schizaealean ferns show no or little taxonomic affinities with Mohria and Anemia. In Actinostachys and Schizaea the

spores are monolete and in Lygodium trilete but the exinal sculpture is smooth.

UITTREKSEL

Mohria en Anemia (Anemiaceae: Pteridophyta) kan onderskei word op grond van hul sporangiums en spore. By Mohria

is die sporangiumkapsule bolrond met ’n apikale annulus, maar by Anemia is dit eiervormig-bolrond tot silindries met ’n

subapikale annulus. Die spore van albei genusse is radiaal simmetries, tetrahedraal en trileet met feitlik parallelle muri.

By Mohria is die eksien gelitteken, maar by Anemia kan dit gelitteken of gekanaliseerd wees. Die muri-stelle by albei genusse

verenig om gemeenskaplike muri te vorm wat vanaf die distale pool tot by die ekwatoriaal radiale gebied strek. By Mohria

is die muri hoi teenoor did van Anemia wat solied of mikroporaat is. Supramurale skulptuur in Mohria en Anemia is pennies

van aard. Spore van die ander Schizaea-aglige varings toon geen of min verwantskappe met did van Mohria en Anemia.

By Actinostachys en Schizaea is die spore monolities en by Lygodium is hulle trileties maar die eksien is glad.

INTRODUCTION

Mohria Swartz, together with Anemia Swartz, Schizaea

J. Sm., 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) because of

their sporangium morphology. The subdivision of this

assemblage into distinct families, the Anemiaceae ( Anemia

and Mohria), Lygodiaceae ( Lygodium ) and Schizaeaceae

{Actinostachys and Schizaea ) has recently gained wider

acceptance (Nayar 1970; Bierhorst 1971; bove et al. 1977;

De la Sota & Morbelli 1987).

Although Anemia and Mohria are dissimilar in mor-

phology they are evidently related in view of similarities

in their plant anatomy (Prantl 1881; Bower 1918; Roux et

al. 1992), trichomes (Mickel 1962; Roux et al. 1992),

spores (trilete, tetrahedral, radially symmetrical with a

cicatricose or canaliculate exinal sculpture) and chromo-

some numbers (n = 38, 76). Anemia is considered the out-

group for Mohria.

The spore morphology of Anemia has been well

documented (Mickel 1962; Hill 1977, 1979; Dettmann &

Clifford 1991). Only recently, however, has their mor-

phology been appraised in a systematic context (De la Sota

& Morbelli 1987; Dettmann & Clifford 1991). Since an

intensive study of Mohria has been lacking for some time,

the spore morphology of merely two or three species has

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

Claremont 7735.

MS. received: 1992-03-18.

been appraised in these studies. In a critical review of

the genus, however, I found it to comprise seven species.

The spore morphology of all the species recognised is

investigated and their phylogenetic relationships evalua-

ted.

MATERIAL AND METHODS

For SEM study untreated sporangia and spores were af-

fixed to aluminium stubs using glue and sputter-coated

with Au/Pd. Specimens were viewed in a Cambridge S200

SEM at 5 or 10 kV. Wax embedding was done using

standard techniques. Sections 8—10 pirn thick were serially

sectioned with a rotary microtome and stained with

safranin and fast green. Photography was done with a Zeiss

Axoskop’ fitted with a M35W camera. Ilford Pan F film

was used for all photography.

Specimens examined

M. caffrorum (L.) Desv., 3218 DC Clanwilliam, Roux 2008 (NBG);

3318 CD Cape Town, Roux 1260 (NBG).

M. lepigera (Bak.) Bak., 1832 DD Zimbabwe, Taylor 3279 (NBG);

1932 CD Zimbabwe, Phipps 281 (BOL); Madagascar, De la Bdthie

7894 (P).

M. marginalis (Sav.) J.P. Roux, Reunion, Commerson s.n. (P); 2155

BC Reunion, De Marne s.n. (NBG); 2828 DB Bethlehem, Roux 907

(NBG); 2829 CA Harrismith, Roux 1524 (NBG).

M. nudiuscula J.P. Roux, 2828 CB Bethlehem, Roux 947 (NBG).

M. rigida J.P. Roux, 2828 DB Bethlehem, Roux 1910 (NBG), Steiner

s.n.

M. saxatilis J.P. Roux, 3219 AA Wuppertal, Roux 2012 (NBG).

M. vestita Bak., 2530 BA Lydenburg, Steiner s.n. (NBG).

200

Bothalia 22,2 (1992)

RESULTS AND DISCUSSION

Sporangial morphology

The sporangium morphology of the schizaealean ferns

has been discussed by Goebel (1905). Prantl (1881) showed

that sporangia of Mohria are derived from a single

marginal protodermal cell, the position of which, as a

result of continual adaxial growth, eventually becomes

superficial. This has been confirmed by Bower (1918).

Mohria, along with all the other schizaealean ferns, is

generally considered as belonging to the leptosporangia-

tes because of their single-layered capsule wall.

Goebel (1905) defined leptosporangia as originating from

a single cell and having a one-layered capsule wall.

Eusporangia, on the other hand, develop from several cells

and have, at least in the primordium, a many-layered

capsule wall. In Mohria, neighbouring cells contribute to

the formation of the massive stalk. Therefore, in gross

morphology the sporangium of Mohria is intermediate

between the two types.

During the early development of the sporangium a

tapetum cannot be distinguished. Later, however, it

separates from the sporogenous tissue as a bicellular layer

(Figure 1A & B). Cells of the outer tapetal layer are

rectangular in transverse section and periclinal. The inner

tapetal layer consists of much larger cells (Figure IB).

Relatively early in the development of the sporangium the

tapetum disintegrates leaving only the proximal part of the

outer tapetal layer intact.

In Mohria I found that the distal portion of the lamina

curves around the sporangia in a protective manner. It does

not differ morphologically or anatomically from the lamina

tissue, and the sporangia are therefore exindusiate. Based

on the grounds of outgroup comparison this feature is

considered plesiomorphic. Anemia is also exindusiate.

Sporangia in the schizaealean ferns are not borne in sori.

In Mohria sporangia occur individually near each vein

ending, or one or two sporangia may be borne at the

terminal vein dichotomy. Sporangia in Mohria thus have

a proper vascularization. In Anemia the vascularization

is rudimentary or vestigial.

Sporangia in Mohria are large, up to 670 /on in diameter,

globose and are borne on a short, massive stalk. The

annuli, which are situated distally, consist of a single row

of radially arranged elongated cells with densely lignified

walls. These indurated cells may number up to 22. The

stomium is formed by a few smaller cells with less ligni-

fied walls which interrupt the ring of annulus cells. The

sporangia dehisce vertically with the slit always facing

away from the lamina margin.

The number of rhomboidal apical plate cells varies. In

M. saxatilis the cells may number three but in the other

taxa there may be as many as six. Bierhorst (1971) considers

a low number of apical plate cells as derived. Spore output

per sporangium, even from the same frond, is extremely

inconsistent. Bower (1923) gave the estimated number of

spores per sporangium as 128 but reported counts of 101

and 107. The number, however, appears to be more

variable, as I have made counts ranging from 57 to ± 196.

Bower (1923) and Wagner (1974) associated a large spore

output with the primitive state. In Anemia the sporangia

are also short-stalked or sessile but with an ovate-globose

to cylindrical capsule. Annuli are subapically positioned.

Schizaealean ferns are held together largely by the

morphology of the sporangium with its group of radially

arranged annulus cells which may be terminal or lateral.

Senftenbergia, a Carboniferous genus with a similar

sporangium structure, has long been suggested as an

ancient element of the schizaealean ferns (Radforth 1939).

More recently, however, Mickel (1974) suggested that it

is a member of the coenopteridalean ferns. If this is true,

the ‘schizaeoid’ sporangium type must therefore have

evolved more than once and thus cannot be used as a

character diagnostic of the Schizaeales. The ‘schizaeoid’

ferns therefore appear to be polyphyletic.

FIGURE 1. — Transverse sections of a sporangium of Mohria caffrorum. A, tapetal development and division of the sporogenous tissue,

X 400; B, mature tapetum and sporocytes before meiosis, Roux 2002 (NBG). S, sporocytes; T, tapetum; ST, sporogenous tissue.

Bothalia 22,2 (1992)

201

FIGURE 2. — Scanning electron micrographs of spore surfaces of Mohria species. A & B, M. caffrorum, Roux 2008 (NBG); C & D,

M. lepigera, Taylor 3279 (NBG); E & F, M. marginalis, De Marne s.n. (NBG); G & H, M. nudiuscula, Roux 947 (NBG). Scale

bar = 5 pm.

mmm

202

Bothalia 22,2 (1992)

FIGURE 3. — Scanning electron micrographs of spore surfaces of Mohria species. A & B, M. rigida, Roux 1910 (NBG); C & D, M. saxatilis,

Roux 2012 (NBG); E & F, M. vestita, Steiner s.n. (NBG). Scale bar = 5 /im.

Spore morphology

A general overview of spore ornamentation in

schizaealean ferns is provided by De la Sota & Morbelli

(1987), whereas Dettmann & Clifford (1991) examined

those of Anemia, Mohria and Ceratopteris Brongn.

Mohria spores are tetrahedral, trilete and radially

symmetrical with a triangular to subtriangular amb 70—20

pm in diameter. They possess a series of conspicuous

parallel ridges or muri which are as broad or broader than

the lumina or grooves. The muri are relatively straight and

usually run parallel to the equatorial plane. In M. lepigera

(Bak.) Bak. and M. marginalis (Sav.) J.P. Roux, however,

they may often be somewhat spirally arranged. Muri of

neighbouring series anastomose in the disto-equatorial

radial regions which often coalesce with the laesura arms.

The laesura, like the proximal surface, is slightly elevated

whereas the distal surface is broadly convex. Muri are

formed by a thick cicatricose exine and are hollow

(Erdtman 1957; Dettmann & Clifford 1991).

The supramural sculpture in Mohria, which is of peri-

nous derivation, ranges from almost smooth in M. vestita

Bak. (Figure 3E & F), reticulate-verruculate in M.

caffrorum (Figure 2 A & B) to colliculate in M. marginalis

(Figure 2E & F) and M. lepigera (Figure 2C & D),

verrucate in M. nudiuscula J.P. Roux (Figure 2G & H)

and finely granulate in M. rigida J.P. Roux (Figure 3A

& B). I have been unable to determine polarity of the fine

sculpture.

Spore diameter ranges from 70—120 pm with no

distinction between taxa. Slight variations also occur

Bothalia 22,2 (1992)

203

within a species. Spore diameter in the genus, however,

averages 87.77 /xm (n = 85, SD = 9.16 /xm) with the largest

percentage (31.76%) felling within the range 81—90 /xm

(Figure 4). Wagner (1974) suggested that spores exceeding

70 /xm are derived.

Spore ornamentation provides evidence of the close

relationship between Mohria and Anemia, especially

subgenus Coptophyllum, as was suggested by Mickel

(1962). De la Sota & Morbelli (1987), identified three spore

types in Mohria based on the structure of the muri. My

observations, however, showed only two types: spores with

closely spaced muri and narrow lumina as in M. lepigera

and M. marginalis (Figure 2C, D, E & F) and spores with

broad muri and lumina as in M. caffrorum (L.) Desv. , M.

nudiuscula, M. rigida and M. vestita (Figures 2A, B, G

& H; 3A, B, E & F). M. saxatilis J.P. Roux belongs to

the latter type but here the muri are somewhat fossulate

(Figure 3C & D).

CONCLUSIONS

Mohria and Anemia can be separated on features of both

the sporangia and spores. Sporangium ontogeny in Mohria

is not of the true leptosporangiate type but is rather inter-

mediate between the developmental patterns of the

leptosporangium and the eusporangium. The annulus of

Anemia is subapical, whereas in Mohria it is distal and

composed of a row of indurated trapezoid cells which

intergrade proximally with a few small isodiametric apical

plate cells. In Mohria the sporangium capsule is globose,

whereas that of Anemia is ovate-globose to cylindrical.

Mickel (1962) considered the globose sporangium of

Mohria to be derived.

(Figure 2G & H). The perine of Mohria and Anemia

subgenus Coptophyllum is thicker and more ornate than

that found in the two other subgenera of Anemia (Dettmann

& Clifford 1991).

Spores of the other extant schizaealean ferns show little

or no morphological similarities to those of Mohria and

Anemia. In Actinostachys and Schizaea the spores are

monolete, whereas those of Lygodium are trilete with a

smooth exinal sculpture.

Mohria and Anemia exhibit numerous plesiomorphic

features with reference to the sporangium and spores.

The sporangia are large, exindusiate, non-soriate (mono-

sporangiate), sessile or short-stalked and have neither a

true eusporangiate or leptosporangiate development.

Annuli consist of a ring of apically or subapically

positioned end cells. The spore output per sporangium is

large (>64 spores per sporangium), and the spores are

trilete and large ( > 70 /xm) in diameter. These features are

significant in explaining evolutionary pathways within the

pteridophytes and also indicate the phylogenetic position

of the schizaealean ferns.

ACKNOWLEDGEMENTS

This study formed part of an M.Sc. degree under the

mentorship of Prof. J.J. A. van der Walt of the University

of Stellenbosch. I wish to express my thanks to him for

his encouragement and valuable discussions throughout

the study and to Dr J. Manning for his valuable comments

on an earlier draft of this paper.

REFERENCES

Mohria and Anemia have radially symmetrical, tetra-

hedral, trilete spores with the exine sculptured to form

near-parallel muri overlain by a thin perine. The mural

sets in both genera anastomose with each other to form

three common muri that extend from near the distal pole

to the equatorial region. In Mohria these muri often

coalesce with the laesura. The cicatricose muri of Mohria

are hollow. In Anemia the exinal sculpture may be

canaliculate or cicatricose but the muri are solid or

microporate. Hollow muri may therefore, on the principle

of outgroup comparison, be considered derived. Only

M. nudiuscula has spores with a supramural sculpture

FIGURE 4. — Range of spore size in Mohria.

BIERHORST, D.W. 1971. Morphology of vascular plants. Macmillan,

New York.

BOWER, F.O. 1918. Studies in the phytogeny of the Filicales. VII. The

Pteroideae. Annals of Botany 32: 1—68.

BOWER, F.O. 1923. The ferns ( Filicales ). Vol. I. University Press,

Cambridge.

CHRISTENSEN, C. 1938. Filicinae. In F. Verdoom, Manual of

Pteridology. The Hague, Nijhoff.

COPELAND, E. B. 1947. Genera filicum. Waltham, Massachusetts.

DE LA SOTA, E.R. & MORBELLI, M.A. 1987. Schizaeales. Phyto-

morphology 37: 365—393.

DETTMANN, m’e. & H.T. CLIFFORD. 1991. Spore morphology of

Anemia, Mohria , and Ceratopteris (Filicales). American Journal

of Botany 78 : 303 — 325.

ENGLER, A. & PRANTL, K. 1898-1902. Die naturlichen Pflanzen-

familien 1,4. Engelmann, Leipzig.

ERDTMAN, G. 1957. Pollen and spore morphology/plant taxonomy.

Almquist & Wiksell, Stockholm.

GOEBEL, K. 1905. Organography of plants. (Translation by Balfour,

I.B.) Clarendon Press, Oxford.

HILL, S.R. 1977. Spore morphology of Anemia subgenus Coptophyl-

lum. American Fern Journal 67: 11—17.

HILL, S.R. 1979. Spore morphology of Anemia subgenus Anemia.

American Fern Journal 69: 71—79.

LOVE, A.D., LOVE, D. & PICHI SERMOLLI, R.E.G. 1977. Cyto-

taxonomical atlas of the Pteridophyta. Cramer, Vaduz.

MICKEL, J.T. 1962. A monographic study of the genus Anemia , subgenus

Coptophyllum. Iowa State Journal of Science 36: 349—482.

MICKEL, J.T. 1974. Phyletic lines in the modem ferns. Annals of the

Missouri Botanic Garden 61: 474—482.

NAYAR, B.K. 1970. A phylogenetic classification of the homosporous

ferns. Taxon 19: 229—236.

PRANTL, K. 1881. Untersuchungen zur Morphologie der Gefasskryp-

togamen. Die Schizaeaceen. Engelmann, Leipzig.

RADFORTH, N.W. 1939. Further contributions to our knowledge of the

fossil Schizaeaceae; genus Senftenbergia. Transaction of the Royal

Society, Edinburgh 59 : 745 —759.

204

Bothalia 22,2 (1992)

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

Systematic studies in the genus Mohria (Pteridophyta: Ane-

miaceae). I. Comparative morphology and anatomy of the rhizome

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

TRYON, R.M. & TRYON, A.F. 1982. Ferns and allied plants with special

reference to tropical America. Springer-Verlag, New York.

WAGNER, W.H. Jr 1974. Structure of spores in relation to fern phylogeny.

Annals of the Missouri Botanic Garden 61: 332—353.

Bothalia 22,2: 205-233 1992)

Richness, composition and relationships of the floras of selected forests

in southern Africa

C.J. GELDENHUYS*

Keywords: biogeography, environmental gradients, flora, forest, fragmentation, growth form, species-area curves, species lists

ABSTRACT

Species lists of 14 widely separated forests representing particular geographic regions in southern Africa were used to

study the size and composition of the individual floras, the similarities between them, and possible determinants of the observed

patterns. The forests contain 1 438 species which belong to 155 families and 661 genera. The growth form spectra show

specific patterns amongst the individual forests such as an abundance of ferns in montane forests, and of woody plants and

vines in coastal forests. The richness of a forest flora increases with increasing altitudinal range within the forest. Significant

linear species-area relationships exist for both woody and herbaceous species, but explain only 30% and 38% of the variation

respectively in the size of the floras. In a multiple regression model the number of dispersal corridors, the proximity to

other forests and mean altitude explained 81% of the variation in the number of woody species. The number of landscape

types and of dispersal corridors explained 75 % of the variation in number of herbaceous species. Several other factors contribute

to the disproportionately large floras of relatively small forests such as at Umtamvuna, Sabie and Richards Bay. A high

proportion of unique taxa are present (30% woody and 42% herbaceous species). The shared taxa show definite trends of

the southward attenuation of species and the presence of elements of the Afromontane and Indian Ocean Coastal Regions.

In conclusion, it is suggested that the southern Cape forests have been isolated from forests along the escarpment and mountains

to the east since at least the Pliocene due to the Sundays River valley which stretches from the coast to the escarpment

in the arid interior.

UITTREKSEL

Soortlyste van 14 geisoleerde woude wat spesifieke geografiese streke in suidelike Afrika verteenwoordig, is gebruik om

die grootte en samestelling van individuele floras, die ooreenkomste tussen hulle, en moontlike bepalende faktore van die

waaigenome patrone te bestudeer. Die woude bevat 1 438 spesies wat tot 155 families en 661 genera behoort. Die verskeidenheid

groeivorms toon spesifieke patrone by individuele woude, soos ’n oorvloed van varings in bergwoude, en van houtagtige

soorte en rankers in kuswoude. Die rykdom van ’n woudflora neem toe met toenemende wydte van die grense in hoogte

bo seespieel binne die woud. Betekenisvolle liniere spesies-area-verhoudings bestaan vir beide houtagtige en kruidagtige

soorte, maar verklaar slegs 30% en 38% onderskeidelik van die variasie in die grootte van die floras. In ’n meervoudige

regressie-model verklaar die aantal migrasieroetes, die nabyheid aan ander woude en gemiddelde hoogte bo seespieel 81%

van die variasie in die aantal houtagtige soorte. Die aantal landskaptipes en migrasieroetes verklaar 75% van die variasie

in aantal kruidagtige soorte. Verskeie ander faktore dra by tot die buitengewoon groot floras van relatief klein woude soos

by Umtamvuna, Sabie en Richardsbaai. ’n Hoe persentasie unieke taksons kom voor (30% houtagtige en 42% kruidagtige

soorte). Die gedeelde soorte toon definitiewe neigings tot die suidwaartse vermindering van soorte en die teenwoordigheid

van soorte van die Afromontaanse streke en die kusgebiede van die Indiese Oseaan. Ten slotte word voorgestel dat die Suid-

Kaapse woude ten minste sedert die Plioseen van die woude langs die eskarp en berge na die ooste geisoleer is as gevolg

van die Sondagsriviervallei wat strek vanaf die kus tot by die eskarp in die droe binneland.

INTRODUCTION

Many forest species have a wide distribution in southern

Africa (Palgrave 1977; Von Breitenbach 1986) and charac-

terize two main floristic regions (White 1978, 1983; Moll

& White 1978). Forests of the Afromontane Region occur

along the Drakensberg escarpment, the Natal and eastern

Cape midlands and the southern and southwestern Cape

mountains and coastal plateaux. Tongaland-Pondoland

forests of the Indian Ocean Coastal Region occur along

the coastal dunes and lowlands. The distribution of many

other species overlaps the two regions. Transitional forests

in the drier lowlands and river valleys between the two

regions such as Kafffarian Subtropical Transitional Thicket

in the eastern Cape (Cowling 1984; Everard 1987) and

similar types in Natal (Edwards 1967) contain species of

both regions. The strong southern attenuation of species

has been attributed to the subtropical temperate transition

* CSIR Division of Forest Science and Technology, P.O. Box 395,

Pretoria 0001.

MS received: 1991-05-26.

(Scheepers 1978; Tinley 1985; Cawe 1986) and the increas-

ing fragmentation of forests due to climatic deterioration

(Geldenhuys 1989). The few widely separated, large forests

are interspersed with many smaller forests (Anon. 1987).

The aims of this study were twofold; firstly, to deter-

mine the floristic richness of widely separated forests

which represent the different geographic regions in

southern Africa and for which comprehensive checklists

exist, and the floristic relationships between them. There

is a need for this because recent studies of southern African

flora have excluded the forest flora because of the small

size of the forest biome and the difficulties posed by the

techniques used to study relationships between floras

(Gibbs Russell 1985, 1987). Furthermore, the studies of

White (1978, 1983) and Moll & White (1978) focused on

tree species only; secondly, to determine the most likely

of several possible sources for the variation in size,

composition and interrelationships of the floras. Based on

biogeographic principles, the following factors are con-

sidered (Brown & Gibson 1983): the size and spatial

separation of the individual forests; the role of dispersal

206

Bothalia 22,2 (1992)

corridors and barriers; the climatic gradient from tropical

northeast to temperate southwest and from the mountains

to the coast; habitat diversity within a forest, including

climatic and edaphic gradients and disturbance regimes;

speciation centres, the development of wider tolerance

ranges through different ecotypes of a species, and the

increase in smaller and more herbaceous growth forms

with a more confined distribution due to increased stress.

STUDY AREA AND METHODS

Fourteen forests or forest complexes (several smaller

forests in close proximity in the same geographical region)

were selected because relatively detailed floristic in-

formation was available for them and because they

represented different geographic areas of the forest biome

in southern Africa (Figure 1). The forests varied greatly

in extent, altitudinal range, geographic location, degree

of isolation, geology, landscape types, surrounding vege-

tation types, and rainfall and temperature regimes (Tables

1 & 2). Values for size of the Transvaal and Natal forests

were obtained from Cooper (1985), and for forests in

Transkei, Ciskei and the Cape Province from relevant

floristic sources.

Various published and unpublished species lists (Taylor

1955; Killick 1963; Moll 1969, 1978, 1980; Van der Schijff

& Schoonraad 1971; Venter 1972; Campbell & Moll 1977;

McKenzie et al. 1977; McKenzie 1978; Scheepers 1978;

Weisser 1980; Weisser & Drews 1980; Nicholson 1982;

Abbott 1985; Deall 1985; Burns 1986; Cawe 1986; Phil-

lipson 1987; Lubke & Strong 1988; Geldenhuys 1989; C.J.

Geldenhuys unpubl. data) were used to compile a list of

species for each forest (see Appendix). Each species was

classified as canopy tree, subcanopy tree, woody shrub,

soft shrub, liane (woody climber), vine (herbaceous

climber), fern (terrestrial) with erect or creeping rhizome,

epiphyte, geophyte, graminoid or forb using the system

of Geldenhuys et al. (1988). Only presence or absence of

a species was indicated for each forest.

Woody and herbaceous plants were separated for the

different analyses because the two categories show con-

trasting patterns along the climatic gradients from

mountain to coast (Geldenhuys & MacDevette 1989).

The effect of forest size on species richness was in-

vestigated by means of the species-area relationship

S = cAz, where S is the number of species, A is area and

c and z are constants. These were fitted by means of a

linear log. log regression. The relationship between the

logarithm of the number of woody or herbaceous species

in a forest and several environmental variables was

determined by means of the stepwise forward selection

procedure of multiple regression analysis (STSC 1986;

Kleinbaum & Kupper 1978). The following independent

variables were included: log forest size (ha); log mean al-

titude (m); log altitudinal range (m); distance from the

tropical source as measured along the forest zone from

arbitrary points, i.e. the Zimbabwe border for the moun-

tain forests, and the Mozambique border for coastal

FIGURE 1.— Distribution of the forests in southern Africa. International political boundaries are not indicated in order not to clutter the forest

pattern. The location of the study areas is indicated as follows: Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast;

Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc, Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape

Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

Bothalia 22,2 (1992)

207

TABLE 1. — Environmental data for forests included in this study

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

forests; the proximity to other large forests (1 for close

to several large forests; 2 for close to several small forests

but distant from large forests; 3 for very isolated from most

forests); the number of geological types (quartzite, sand-

stone, mudstone, limestone, dolerite, dolemite, shale,

schist, conglomerate, granite and dune sand); the number

of landscape types (mountain slope, mountain plateau,

escarpment, valley, gorge, estuary and dune); the number

of plant dispersal corridors present (mountain range,

escarpment, river, and coastal dune system); and the

number of different structural vegetation types surrounding

the forest (fynbos, renosterveld, grassland, thomveld,

woodland and thicket).

Information for the last four variables was obtained from

descriptions of the study areas of the relevant floristic

sources.

The index of similarity of Czekanowski (IsC) (as used

by Rogers & Moll 1975), expressed as percentage, was

208

Bothalia 22,2 (1992)

used to compare similarity between forests, where

IsC = 200w/(a+b), a and b are the numbers of species

present in each forest, and w is the number of species com-

mon to both forests.

RESULTS

Size and composition of total forest flora

Number of taxa

Table 3 lists the number of families, genera and species,

as well as the species/family and species/genus ratios for

the vascular plants in each forest and for the total forest

flora. The list of species (Appendix) represents 1 438

species, i.e. the bulk of species occurring in the southern

African forests.

Twenty-six families each contain 1% (14) or more of the

taxa (species, subspecies and varieties) of the total forest

flora. These families are (number of species in brackets):

Acanthaceae (45), Adiantaceae (21), Anacar-

diaceae (29), Apocynaceae (19), Asclepiadaceae (31),

Aspleniaceae (24), Asteraceae (81), Capparaceae (14),

Celastraceae (40), Convolvulaceae (15), Crassulaceae (20),

Cyperaceae (35), Ebenaceae (19), Euphorbiaceae (67),

Fabaceae (79), Flacourtiaceae (21), Lamiaceae (33), Lilia-

ceae (42), Malvaceae (15), Moraceae (14), Oleaceae (17),

Orchidaceae (53), Poaceae (57), Rubiaceae (66), Scro-

phulariaceae (19) and Vitaceae (14). These same families

also represent 17 % of all families present and include 55 %

of the genera and 62% of all forest species. Fifty-four

percent of families have four or fewer species. Sixty-five

families are represented by a single genus and 37 by a

single species.

Only 15 genera contain 10 or more species. Of these,

only Streptocarpus (12) (Gesneriaceae) does not belong

to one of the largest families. The other genera are

TABEL 2. — Rainfall and temperature data for forests included in this

study. Data were obtained from the respective study reports or

from published and unpublished sources for nearby stations

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London

coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape

Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern Cape;

Trm, Transkei mountains; Umt, Umtamvuna Gorge.

TABEL 3. — Number of families, genera and species, and species/

family and species/genus ratios for the different forests and the

total forest flora

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London

coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape

Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

Asplenium (23), Crassula (18), Cyperus (11), Diospyros

(12), Ficus (12), Helichrysum (10), Isoglossa (10), Mayte-

nus (14), Pavetta (13), Plectranthus (18), Protasparagus

(10), Rhus (21), Senecio (19) and Vemonia (10). Sixty-one

percent of the genera are represented by a single species.

Growth forms

The growth form spectra varied significantly between

the different forests (Table 4; Chi-square value = 593.7,

df = 143, P< 0.001). Values with a particularly high

Chi-square value for a particular cell are indicated in the

table. None of the forests contain canopy trees, soft shrubs

or geophytes in disproportionate numbers. The forests

which contain species of a particular growth form in excess

of the expected number are Maputaland (subcanopy trees

and graminoids), Umtamvuna (woody shrubs), Hawaan

(lianes), Transkei mountains and Cape Peninsula (erect

ferns), Mariepskop (epiphytes) and the southern Cape

(forbs). Growth forms in numbers less than the expected

number occur in the southern Cape (subcanopy trees

and lianes), Transkei mountains (vines, graminoids and

forbs), Umtamvuna (graminoids), Richards Bay (erect

ferns), Maputaland (all ferns and forbs) and northeastern

Transvaal (woody shrubs).

Woody species constitute approximately 50% of the total

flora in all forests but this percentage varies greatly be-

tween individual forests (Table 4). In general, coastal

forests have a percentage of woody species in excess of

60%, whereas for montane forests the percentage varies

between 39% and 53%. However, the Transkei mountain

forests have a percentage of 68% and the Richards Bay

coastal forests a percentage of 57 % .

The geographical ranges of species are significantly

related to their growth form (Chi-square value based on

absolute frequencies = 246.7, df = 99, P< 0.001). Cell

Bothalia 22,2 (1992)

209

TABLE 4. — Number of species by growth forms for the different forests. The signs following some numbers indicate that the number is much

higher (+) or lower (— ) than the expected number under assumption of independence (Chi-square analysis)

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

** Woody species include trees, shrubs and lianes.

values which have made a large contribution to the

significant Chi-square value are indicated in Table 5. Trees,

lianes and ferns are well represented: 37 % of canopy trees,

26% of subcanopy trees, 27% of lianes, 31% of erect ferns

and 24% of creeping ferns occur in more than five of the

forests. Fifteen per cent or less of the other growth forms

occur in more than five forests. No species occurs in all

forests but the species which occur in more than 10 forests

(75%) are Apodytes dimidiata, Calodendrum capense,

Canthium inerme, Celtis africana, Clausena anisata,

Cussonia spicata, Dietes iridioides, Ekebergia capensis,

Galopina circaeoides, Grewia occidentalis, Halleria

lucida, Ilex mitis, Maytenus heterophylla, Maytenus

undata, Olea capensis subsp. macrocarpa, Oplismenus

hirtellus, Pittosporum viridiflorum, Protasparagus

setaceus, Psychotria capensis, Psydrax obovata, Rapanea

melanophloeos, Rhoicissus tridentata, Scutia myrtina,

Secamone alpinii and Zanthoxylum capense.

Regression analyses

Size and species richness of the different forests vary

greatly (Tables 1 & 3). The number of species of both woo-

dy and herbaceous plants shows a significant log

TABLE 5. — The frequency of occurrence of species of different growth forms in 14 widely separated forests of southern Africa

* 1, canopy trees; 2, subcanopy trees; 3, woody shrubs; 4, soft shrubs; 5, lianes; 6, vines; 7, erect ferns; 8, creeping ferns; 9, epiphytes;

10, geophytes; 11, graminoids; 12, forbs.

210

Bothalia 22,2 (1992)

TABLE 6. —Constants and significance of the linear log-log models of

the species-area relationships for the forests

species-log area relationship (Table 6). However, this

relationship explains only 30% and 38% respectively of

the variation in the size of the floras. In both models a

number of forests lie outside the 95 % confidence intervals.

The Umtamvuna, Richards Bay and Sabie forests have

many more plants of both categories, whereas the

Peninsula, Grootvadersbosch and Cathedral Peak forests

have far fewer woody species, and the Transkei mountain,

East London coast, Hawaan and Maputaland coast forests

have much fewer herbaceous species than the number

predicted by the linear log species-log area regression

model.

LOG NUMBER WOODY SPECIES OBSERVED

• Observed values 95% Intervals — Predicted values

LOG NUMBER HERBACEOUS SPECIES OBSERVED

LOG NUMBER HERBACEOUS SPECIES PREDICTED

• Observed values 95% Intervals — Predicted Values

FIGURE 2. — Observed and predicted values, and 95% confidence

intervals in relation to predicted values for the number of woody

and herbaceous plants in a forest. The coefficients for the multi-

ple regression equations are presented in Table 7. The study areas

are indicated as follows: Ama, Amatole Mountains; Cap,

Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch;

Haw, Hawaan; Mak, Mariepskop; Mlc, Maputaland coast; Net,

northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib,

Richards Bay; Sab, Sabie transect; Soc, southern Cape; Trm,

Transkei mountains; Umt, Umtamvuna Gorge.

In the multiple regression analysis for woody plants,

proximity to other forests, the number of dispersal

corridors and mean altitude explained 81.6% of the varia-

tion in the observed values (Table 7). The use of fewer

or more variables in the model caused a reduction in the

coefficient of determination (R2). The number of land-

scape types and dispersal corridors explained 75.1% of the

observed variation in the number of herbaceous plants

(Table 7). Data for the Transkei mountain forests were

excluded from this analysis because Cawe (1986)

undersampled herbaceous plants other than ferns. All

observed values, except those for herbaceous species in

the Transkei mountain forests, fall within the 95%

confidence intervals around the values predicted by the

multiple regression model (Figure 2).

Shared and unique taxa and percentage similarity

Shared species

Shared taxa show at least three distinct patterns (Table

8). Firstly, forests share many more of their species with

forests to their north and east than they share with forests

to their south and west. This indicates an erosion of species

from the two tropical source areas, i.e. the Transvaal and

Maputaland forests, to the southwestern Cape forests.

Secondly, forests share many more species with their

nearest neighbours than with forests further away. Note

that the forests to the south share more species with the

Mariepskop forest than with either the Sabie or the north-

eastern Transvaal forests. Thirdly, the Affomontane forests,

i.e. including forests from the southern to western Cape,

share relatively fewer species with the forests of the coastal

areas. The Umtamvuna and Transkei mountain forests,

however, share relatively many species with both the

coastal and montane areas.

Unique species

A large proportion of the species are unique to individual

forests: 33% of the woody and 42% of the herbaceous spe-

cies (Table 9). Canopy trees and ferns have the lowest

proportions of unique species, whereas these proportions

are >40% for the shrubs, geophytes and forbs (Table 5).

Umtamvuna (20%), southern Cape (16%), Richards Bay

(13%), Maputaland coast (13%) and the Sabie transect

(12%) together contributed 74% of the unique species, and

were the most important contributors to the unique species

of each growth form. The Mariepskop and northeastern

Transvaal escarpment forests contain relatively many

unique soft shrubs and epiphytes.

Percentage similarity

The mean percentage similarity between any two forests

is 34.4% for woody plants and 23.7% for herbaceous

plants (Table 10). The individual forests differ widely in

the number of forests and in the particular forests with

which they share a similarity higher than the mean for the

particular plant group.

DISCUSSION

Before the results are discussed, it is necessary to note

that some components of the flora, in particular some

Bothalia 22,2 (1992)

211

TABLE 7. — Analysis of variance for the significant regression variables in the order in which they were fitted, and estimates of the regression coefficients

TABLE 8. — The percentage shared taxa for the 14 forests. The upper triangle gives the values for the woody plants and the lower triangle the

values for the herbaceous plants. In each cell of two values in the triangles, the upper value indicates the percentage of the species of

the forest of that row which is shared with the forest of that column. The bottom value of the cell shows the reverse relationship

Forest*

Pen Gvb Soc Ama Elc Trm Umt Haw Rib Mlc Cap Sab Mak Net

% species shared between forests

Woody species

Herbaceous species

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

212

Bothalia 22,2 (1992)

TABLE 9.— The number of unique species over growth forms, and the unique species as a percentage of all plants, for each forest

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast. Net, northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

+ 1, canopy trees; 2, subcanopy trees; 3, woody shrubs; 4, soft shrubs; 5, lianes; 6, vines; 7, erect ferns; 8, creeping ferns; 9, epiphytes;

10, geophytes; 11, graminoids; 12, forbs.

herbaceous growth forms, may have been undersampled.

This is understandable in studies of forests because atten-

tion is invariably focused on the trees and conspicuous

understorey plants. Firstly, this occurred in the Transkei

mountain forest data where Cawe (1986) was concerned

with the timber resource potential but also sampled the

conspicuous fern understorey as a possible indicator of

site productivity. This undersampling of herbaceous

species in Transkei was considered in the regression

analyses for herbaceous species, and explains the devia-

tions of the Transkei herbaceous data from the observed

general trends (see Tables 4, 8, 10). Secondly, several

species, especially herbaceous plants, tend to exhibit a

disjunct distribution, being absent in a given forest but

present in neighbouring ones. This may reflect inadequate

collection of inconspicuous and rare plants. The recently

published species list for the Amatole forests (Phillipson

1987) lacked several species which by that time had been

collected from the eastern parts of the forests (C.J.

Geldenhuys unpubl. data). By contrast, the lists may also

reflect the inclusion of species which are usually associated

with other biomes (such as grasses, shrubs and pioneer

trees) but which are contained in particular development

stages of some forests. Finally, the lists may merely reflect

the true distribution pattern of some species. More detailed

studies may clarify this and the Appendix is included to

assist in this clarification.

Flora size and relationships

The evergreen forests in southern Africa cover only

0.08% of the area and contain only 7.1% of the indigenous

vascular species, and thus have a relatively rich 0.58

species/km2, making it the second richest biome per unit

area in southern Africa. The overall ratio for southern

Africa with over 20 227 indigenous vascular taxa is 0.0079

TABLE 10.— Percentage Czekanowski similarity of the woody and herbaceous components of the floras of the 14 forests. The upper triangle gives

the values for the woody plants (trees, shrubs and lianes) and the lower triangle the values for the herbaceous plants

Forest*

* Ama, Amatole Mountains; Cap, Cathedral Peak; Elc, East London coast; Gvb, Grootvadersbosch; Haw, Hawaan; Mak, Mariepskop; Mlc,

Maputaland coast; Net, northeastern Transvaal escarpment; Pen, Cape Peninsula; Rib, Richards Bay; Sab, Sabie transect; Soc, southern

Cape; Trm, Transkei mountains; Umt, Umtamvuna Gorge.

Bothalia 22,2 (1992)

213

plant species/km2 (Gibbs Russell 1985). Fynbos has 1.36

species/km2 with a total of 7 316 species, and grassland

has 0.25 species/km2 with 3 788 species (Gibbs Russell

1987).

Sixteen of the largest families of the forest flora are

included amongst the 38 largest flowering plant families

listed by Gibbs Russell (1985). The other large forest

families are Adiantaceae, Apocynaceae, Aspleniaceae,

Capparaceae, Celastraceae, Ebenaceae, Flacourtiaceae,

Moraceae, Oleaceae and Vitaceae. Of these latter families

only Capparaceae is indicated by Gibbs Russell (1987) as

a characteristic family of any other biome, i.e. the desert

biome. However, most families listed by Gibbs Russell

(1987) as characteristic of other biomes occur in the forest

list. Notable absences are two large families listed by Gibbs

Russell (1985), namely Restionaceae and Chenopodiaceae,

which respectively partly distinguish fynbos and desert

(Gibbs Russell 1987).

Gibbs Russell (1985) suggested that families with a

species/genus ratio more than twice the overall ratio of

9.6 for southern African seed plants have diversified

extensively within southern Africa. The species/genus ratio

for the total forest flora is only 2.2 with the ratio for the

individual forests ranging between 1.3 and 1.6 (Table 3).

Forty-nine of the families have a species/genus ratio greater

than 2.2 (Appendix). The families with a species/genus

ratio of more than 4.4, i.e. twice the overall mean, are

Aspleniaceae (12.0), Crassulaceae (6.7), Dioscoreaceae

(6.0), Ebenaceae (9.5), Gesneriaceae (12.0), Lycopodiaceae

(6.0), Moraceae (4.7), Ochnaceae (7.0), Folygalaceae (9.0),

Solanaceae (5.0) and Thelypteridaceae (4.5). The high

ratio of these families can be attributed to a single genus

with many species. They are mostly forest understorey or

subcanopy plants.

Size of individual forest floras

Species-area relationships

Forest size determines the richness of the flora but only

in simple linear regression and explains 30% to 38% of

the observed variation in species richness. It explains the

rich southern Cape forest flora despite its extreme southern

location at the western end of the larger forests of southern

Africa (see Anon. 1987). However, size does not explain

the rich floras of the small Umtamvuna, Richards Bay and

Sabie forests. In the multiple regression analyses, forest

size was an insignificant variable, whereas variables which

explain dispersal patterns and habitat diversity (proximity

to other forests, the number of dispersal corridors and

landscape types, and mean altitude) explained 75 % to 82 %

of the variation in species richness.

Number of dispersal corridors

The number of dispersal corridors meeting in a particu-

lar forest is one of the strongest variables determining the

number of woody plants in a forest (Table 7). A dispersal

corridor provides environments which are similar to the

two source areas at either end of it, or it is a broad band

of similar habitat (Brown & Gibson 1983). Mountain

chains (Transkei and Amatole Mountains), escarpments

(Natal and Transvaal Drakensberg), river valleys (Tugela

River, Edwards 1967) and coastal dune systems (Zululand

and eastern Cape) link forests into larger complexes and

link forest complexes on either side of dry, open valleys

and lowlands (see Anon. 1987). The most prominent dry

zone stretches from the Transvaal lowveld to the eastern

Cape between the southwest-northeast mountain chains and

escarpment, and the Indian Ocean coast (Zucchini &

Adamson 1984).

Each type of corridor provides a different set of environ-

mental conditions and provides for a specific direction of

dispersal for the plants.

The I\igela River basin is a good example of a corridor

which allows coastal and montane species to mix along

the rivers and escarpments, at a distance from both

sources, for example in the Qudeni, Nkandhla and Ngoye

forests on the eastern margin of the Tugela River basin

(Edwards 1967; Anon. 1987). This explains in part the high

degree of similarity between the small Sabie and Richards

Bay forests and the higher degree of similarity of the Trans-

vaal escarpment forests to the Richards Bay forest rather

than to the other two Natal north coast forests (Table 10).

But coastal and montane forest species cannot establish

themselves in the area between the rivers due to unfavour-

able climatic conditions (drought and frosts) and the

frequent occurrence of fires.

The corridor provided by the Drakensberg escarpment

explains the high similarity amongst the Transvaal forests,

and between these and those occurring on the Transkei

and Amatole Mountains (Table 10). The Transvaal escarp-

ment provides sites with very uniform climate over several

degrees of latitude, and which protect the forest against

the frequent grassland fires such as the Wonderwoud near

Tzaneen. This escarpment is also part of the chain of

mountains which extend more or less uninterrupted as far

south as the Amatole forests.

Mountain ranges and dune systems provide for large

habitat diversity through climatic (altitudinal range and

different slopes and exposures), edaphic and disturbance

gradients (Van der Schijff & Schoonraad 1971; Scheepers

1978; Deall 1985; Bums 1986; Geldenhuys 1989). The

diversity of habitats allows species to migrate within the

system during conditions of environmental change

(Scheepers 1978). Mountain ranges also allow forests to

persist within larger areas of totally different, extreme

climatic and disturbance regimes such as the Karoo and

Fynbos (Anon. 1987; Geldenhuys 1985, 1989).

Both the number of corridor types present in a forest

and the proximity of the forest to other forests contribute

significantly to the number of woody species in that forest

(Table 7). This concept is demonstrated in the rich woody

flora of the small Umtamvuna gorge forest. It exists in

a central position between the coastal and mountain forests

of the eastern Cape, Transkei and Natal. It is linked to

those different types of forests by different types of

corridors which allow an interchange of species between

forests along the coast, and on mountain ranges and

the Drakensberg escarpment. This is shown by the

high similarity between the Umtamvuna and the other

mentioned forests. The gorge is relatively deep, and there-

fore protected from fires, but at the same time it is unob-

structed, which allows coastal elements to migrate inland

and mountain elements to migrate towards the coast.

214

Proximity between forests

The greater floristic similarity between forests of the

larger complexes which occur in relatively close proximity

is attributed to the similarity of their environments.

Examples are the close affinity between the Transvaal

escarpment forests, between the Natal coastal forests and

between the Amatole, Transkei and Umtamvuna forests

(Table 10). The probability of successful establishment

after chance events of long-distance dispersal (Brown &

Gibson 1983) is increased if the forests in close proximity

share similar environments. By contrast, the Natal coastal

forests share much fewer species with the distant Drakens-

berg escarpment forests which is presumably due to great

climatic and edaphic dissimilarity.

The smaller similarity between relatively isolated forests

is attributed to the effective abiotic and biotic barriers to

dispersal of propagules between them, and the lack of

effective dispersal corridors. Firstly, the climate in the

valleys and lowlands between adjacent forest complexes

(Muir 1929; Edwards 1967; Cowling 1984; Everard 1987),

the more extreme fire regimes of adjacent woodlands,

grasslands and fynbos (Granger 1984; Edwards 1984), and

the exposed mountain peaks and ridges (Killick 1963;

Geldenhuys 1989) are barriers to the successful dispersal

of forest biota. Van Daalen (1981) noted the inability

of forest species to establish in fynbos. Secondly, the

Peninsula, Grootvadersbosch, southern Cape, Hawaan and

Cathedral Peak forests occur isolated from most other

forests and are linked with them by few and ineffective

corridors.

The Peninsula, Grootvadersbosch and Cathedral Peak

forests have high similarities only with their nearest neigh-

bours, and share mostly the widespread species.

The Peninsula forests are presently very isolated from

the main western Cape mountain ranges. However, their

species richness is higher than that of the forests of those

mountains (for example McKenzie 1978). They share

several species with forests along the coast to the east

(Masson & McKenzie 1989) which makes a coastal

corridor very likely.

Grootvadersbosch is very isolated from other forests,

even the southern Cape forests. The links between Groot-

vadersbosch and the coast are poor and cross relatively

dry country (Muir 1929).

Cathedral Peak forests are isolated from the rest of the

Drakensberg escarpment forests. They have very poor links

with the Natal midlands and coastal forests. They are

surrounded by extensive grasslands which bum frequently

(Edwards 1984; Tainton & Mentis 1984; Everard 1986).

Hawaan forest shares several species with smaller forests

in the vicinity such as Steinbank and Krantzkloof (coastal

scarp), and Karkloof (Natal midlands) although it is most

similar to the Hlogwene dune forest (Rogers & Moll 1975;

Moll 1978).

The southern Cape forest is large, covers several land-

scape types and is linked with the forests to the east mainly

through the discontinuous mountain ranges and along

the coast. The rivers provide only local links with the

Bothalia 22,2 (1992)

inland mountains which have very small, isolated forests

(Geldenhuys 1989).

Altidude

Mean altitude improved the coefficient of determination

of the number of woody plants in the multiple regression

model, but was an insignificant variable in linear regres-

sion (Table 7). Its negative coefficient emphasizes the

higher richness of coastal forests compared to the mountain

forests. This was also shown by Geldenhuys & MacDevette

(1989) for both the southern Cape and Natal. I attribute

its insignificance in linear regression to the wide altitudinal

range of many forests along the eastern escarpment and

mountains.

Number of landscape types

The number of landscape types in a forest is the most

significant variable determining the number of herbaceous

species (Table 7). Different landscape types provide

different combinations of slopes, aspects, soil depths, soil

nutrient and moisture status, and different disturbance

regimes (Scheepers 1978; Deall 1985; Geldenhuys 1989).

Each landscape type carries a subset of unique species with

narrower habitat tolerances. Geldenhuys & MacDevette

(1989) have shown that different herbaceous growth forms

show different habitat preferences along gradients from

the coast to the mountain, both in Natal and the southern

Cape. This is particularly evident in the southern Cape,

Amatole, and Transvaal escarpment forests which include

the largest number of landscape types (Table 1) and which

have many species in most of the herbaceous growth forms

(Table 4).

Habitat requirements and distribution of species

Physiological tolerances of species to climatic conditions

are reflected in the growth form spectra of different forests

(Table 4) and the distribution ranges of species of different

growth forms. This would also contribute to the observed

variation in the richness of the floras of different forests.

The southern African forest environment is characterized

by relatively steep climatic gradients (Killick 1963; Venter

1972; Scheepers 1978; Campbell & Moll 1977; McKenzie

1978; Deall 1985; Bums 1986). Mountains are cool to cold

and the coastal areas warm to hot. The northeastern parts

are subtropical-tropical with summer rain, and the south-

western parts almost cool temperate with winter rain. The

mountains and coast receive high rainfall with relatively

dry areas in-between.

Growth form spectra indicate that cooler mountain

forests have a larger proportion of herbaceous plants

whereas the warm, humid coastal forests have a larger

proportion of woody plants (Table 4; Geldenhuys &

MacDevette 1989). Coastal forests are particularly rich in

trees, woody shrubs, lianes and vines. Mountain forests

are particularly rich in ferns, which are far less common

in the coastal forests, and are deficient in climbers except

in the lower-lying (drier and/or warmer) parts. Fern and

bryophyte epiphytes are generally associated with moun-

tain forests and mistbelts (Poes 1982) and epiphytic orchids

with tropical lowlands (Harrison 1972). Mountain forests

generally contain many epiphytes (e.g. Scheepers 1978).

Bothalia 22,2 (1992)

215

Notable exceptions are the Peninsula, Grootvadersbosch,

Transkei (where they were not collected) and Cathedral

Peak forests. In the southern Cape epiphytes are abundant

and represented by numerous species. This feature is most

pronounced in the large, less frequently disturbed forests

(by fire) of the coastal platform and river valleys, rather

than in the small, more frequently disturbed mountain

forests (Geldenhuys & MacDevette 1989). More frequent

disturbance by fire could explain, in part, the lack of

epiphytes in the smaller mountain forests of this study.

Protection from fire could explain the high species richness

in the larger montane forests of the Transvaal escarpment

and in the well-protected but small Umtamvuna forests.

Many species drop out along the tropical-temperate

gradient (Table 8). This southward attenuation of species

was noted in several studies (Phillips 1931; McKenzie 1978;

Moll & White 1978; Scheepers 1978; Tinley 1985; Cawe

1986; MacDevette 1987; Geldenhuys 1989). The high

number of unique trees and shrubs of the Maputaland dune

forests has been related to the deterioration of the tropical

climate to the south (Table 9; Moll & White 1978; Tinley

1985). Further south, I have related the sharp decline in

numbers of species from the southern Cape to the Cape

Peninsula to the increasing aridity, fire frequency and forest

fragmentation since the Pliocene (Geldenhuys 1989).

Steep gradients imply that the widespread species have

wide habitat tolerances, and that the restricted species have

narrower tolerances. Tree species have much wider ranges

than shrubs, and ferns have much wider ranges than the

other herbaceous growth forms (Table 5). However, only

7 % of all species occur in eight or more forests, and no

species occur in all forests.

Interaction with adjacent vegetation types

The climatic and disturbance regimes and structure of

surrounding vegetation types will determine the interaction

of the forest with those vegetation types. This interaction

can increase the number of species in the forest in several

ways:

Forest margin in close contact with disturbance regimes

of adjacent vegetation types

Small forests have a large ratio of forest margin to forest

area. As such they may contain proportionately more

species which are usually associated with adjacent

vegetation types but which appear in forest communities

during the successional stages. The Richards Bay and

Sabie forest communities in particular contained shrub,

graminoid and forb species which were common in

communities other than forest. These forests occurred in

complex mosaics with other vegetation communities

(Venter 1972; Deall 1985). This partly explains the high

species richness of these two forests in relation to their

small size (Figure 2). The inclusion of many ecotonal

species in the forest floras of Sabie, Richards Bay and the

southern Cape could also explain the high number of

unique species of several different growth forms of these

forests (Table 8). In contrast, Hawaan forest is well

protected and mature but surrounded by cultivated land

(Moll 1969; Cooper 1985). It therefore lacks an ecotone

and this could, at least in part, explain its low number

of herbaceous species. Everard (1986) also pointed to the

negative effect on species richness of a forest if the forest

ecotone is frequently destroyed by fire.

Vegetation types with structure and disturbance regimes

somewhat similar to forest

Subtropical transitional thicket in the eastern Cape

(Cowling 1984; Everard 1987), similar types in Natal

(Edwards 1967) and moist savanna (Huntley 1984) of the

Transvaal (Van der Schijff & Schoonraad 1971; Scheepers

1978; Deall 1985) and Natal (Edwards 1967) share various

proportions of forest taxa. As such they provide corridors

for the dispersal of forest species across the barriers

(Edwards 1967; Moll & White 1978; Cowling 1984;

Everard 1987). Current land use practices, such as

intensive agriculture in the eastern Cape and Natal, remove

this corridor and may intensify the isolation of the forests.

However, plantation forestry and the associated reduction

of fire and amelioration of the microclimate provide

corridors for plant species migration (Geldenhuys et al.

1986; Knight et al. 1987).

Environmental change

The present patterns of composition and interrelation-

ships of the different forests suggest that their high degree

of similarity may have been established before major

fragmentation of the forests occurred. For example, the

southern Cape forests are relatively similar to the Amatole,

Transkei and Transvaal forests. Yet they are linked with

the forests to the east by broken mountain ranges which

are separated by relatively dry wide open valleys and

extensive lowlands. One particularly prominent gap in

forest distribution is formed by the Sundays River valley

east of Port Elizabeth (Figure 1). It stretches in a north-

westerly direction towards remnants of the escarpment of

the African Surface in the vicinity of Graaff-Reinet in the

arid interior. East of this valley a massive uplift occurred

during late Pliocene (+2.5 million years ago) along the

Ciskei-Swaziland axis, whereas west of the valley the uplift

was of lesser magnitude. This resulted in significant

rejuvenation along the major inland drainage lines which

are evident in the high accumulation rates of sediment at

the mouths of major rivers along the southeastern coast

(Partridge & Maud 1987).

I suggest that the forests in the southern Cape became

isolated from the forests along the escarpment to the east

of the Sundays River valley by the late Pliocene. The maps

of Partridge & Maud (1987) suggest that the Sundays River

was already extensive by the Miocene but indications

are that aridity increased rapidly towards the Pliocene-

Pleistocene (Deacon 1983). The relatively dry Suurberg

forests immediately to the east of the Sundays River valley

are the closest forests to the southern Cape (Geldenhuys

1985). The only connections between the southern Cape

forests and those to the east would have been along the

coast and by means of the subtropical transitional thicket.

The increasing aridity which followed the Pliocene

(Deacon 1983) increasingly fragmented the forests. Forests

were probably most limited during the last cold, dry

Glacial Maximum of 18 000 years ago (Deacon et al. 1983;

Scholtz 1986). Acocks (1988) and White (1983) attributed

216

Bothalia 22,2 (1992)

the relic nature of the forests within the grassland and

fynbos biomes to the destructive activities of man during

the relatively recent 100 to 300 years. However, Feely

(1980, 1986) indicated that most of the present southern

African grassland existed throughout the Holocene and

was not induced by recent forest clearing. Forests still

persist today in areas where Iron Age formers in Transkei

settled in high density for at least the last 1 400 years. I

have indicated that fires associated with hot, desiccating

winds have confined forests to shadow areas of fire-bearing

winds (Geldenhuys 1989) whereas others (Story 1952;

McKenzie 1978; Scheepers 1978; Deacon etal. 1983) have

also commented on the role of fire.

During this long period of forest fragmentation, forests

and forest biota survived in areas which we now consider

as dispersal corridors. I suggest that the forest species

responded in different ways to the increasing pressures of

drought and fire. Some species survived in the specific

landscape types because of better availability of moisture

and protection against fires. Outside of these sites many

species were eliminated due to pressures from droughts

and fires. Species with wider climatic tolerances persisted

with a wide distribution range and with the adoption of

a range of sizes and shapes. The pressures of drought and

fire caused many other species to evolve into smaller

growth forms. This view is supported by two findings of

this study. Firstly, forests in closer proximity share more

species than forests further apart. Disperse may play a

role, but I suggest that this role is of lesser significance.

Secondly, most of the large families, and many of the other

important families and genera are shared between the forest

and the other vegetation types. Their species/fomily ra-

tios are small in the forest compared to the large ratios

outside the forest. They have few but widespread species

in the forest, and many but relatively localized species in

the surrounding vegetation types. Species with the taller,

longer-living growth forms occur in the forest, whereas

the smaller and often herbaceous growth forms occur in

the vegetation types which are exposed to more extreme

environmental conditions.

CONCLUSION

I suggest that fragmentation of the forests and an increase

in vegetation types which are tolerant of frequent fires

and/or droughts had a profound effect on the speciation

of the southern African flora. Most of the large plant

families, and many of the other important families, are

shared between the forest and the other vegetation types.

This sharing suggests that forest might have been the

original gene source for the speciation of many of the

families and genera. Examples are the Anacardiaceae,

especially Rhus, Asteraceae, Liliaceae, Orchidaceae,

Proteaceae and Rosaceae. This effect of increasing aridity

and disturbance on the radiation of species beyond forests

should be considered in studies of the phylogenies of many

of the groups.

I have indicated that a variety of factors contributed to

the variation in the size of the floras of individual forests.

Forests where several positive factors operate have rich

floras compared to the poorer floras of forests with fewer

positive factors (Table 1). However, the significant vari-

ables do not explain the large number of both woody and

herbaceous plants of the Umtamvuna forest, except

perhaps the number of corridors. The Umtamvuna forest

forms part of the southern Natal/Fondoland quartzite sand-

stone complex which is known to have a remarkably high

number of endemic woody species (Van Wyk 1981). This

whole complex requires a detailed study to determine the

composition and distribution of different plant communi-

ties, and the distribution of the rare and endemic species.

This would allow a more objective explanation of its high

number of species in the relatively confined area.

The fragmentation had been aggravated by current land

use practices, such as clearing for agriculture, forestry and

subsistence utilization, and veld burning practices for

grazing and improved water runoff in catchments (Phillips

1963; Feely 1980, 1986; Cooper 1985) and the develop-

ment of coastal resorts and townships. I suggest that more

localized studies should be conducted to determine the

effect of these land use practices on the survival of species

in different regions.

The suggestion of the isolation of the southern Cape

forests from those to the east already by the Pliocene

implies long isolation and stability of the forest species.

Several well-defined ecotypes may exist in many of the

taxa. Collection of seed of those species for planting in

other parts of their range may have serious implications

for the conservation of the ecotypes within those species.

ACKNOWLEDGEMENTS

This study formed part of the Conservation Forestry

Research Programme of the former South African Forestry

Research Institute (presently the CSIR Division of Forest

Science and Technology) of the Department of Environ-

ment Affairs, Forestry Branch. The study was done as part

of a Ph.D. thesis under the supervision of Prof. E.J. Moll,

Department of Botany, University of Cape Town.

REFERENCES

ABBOTT, A. 1985. Flora of the Umtamvuna Nature Reserve. Unpublished

interim list of plants, Clearwater, Port Edward.

ACOCKS, J.P.H. 1988. Veld types of South Africa. 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

ANON. 1987. Map of South African indigenous evergreen forest. Forest

Biome Project, Ecosystem Programmes, FRD: CSIR, Pretoria.

BROWN, J.H. & GIBSON, A.C. 1983. Biogeography. Mosby, St. Louis.

BURNS, M.E.R. 1986. A synecological study of the East London coast

dune forests. M.Sc. thesis, Rhodes University, Grahamstown.

CAMPBELL, B.M. & MOLL, E.J. 1977. The forest communities of

Table Mountain, South Africa. Vegetatio 34: 105—115.

CAWE, S.G. 1986. A quantitative and qualitative survey of the inland

forests of Transkei. M.Sc. thesis. University of Transkei, Umtata.

COOPER, K.H. 1985. The conservation status of indigenous forests in

Transvaal, Natal and O.F.S., South Africa. Wildlife Society of

Southern Africa, Durban.

COWLING, R.M. 1984. A syntaxonomic and synecological study in the

Humansdorp region of the Fynbos Biome. Bothalia 15: 175—227.

DEACON, H.J. 1983. The comparative evolution of Mediterranean-type

ecosystems: a southern perspective. In F.J. Kruger, D.T. Mitchell

& J.U.M. Jarvis, Mediterranean-type ecosystems— the role of

nutrients: 3-40. Ecological Studies 43. Springer-Verlag, Berlin.

DEACON, H.J., HENDY, Q.B. & LAMBRECHTS, J.J.N. (eds) 1983.

Fynbos palaeoecology: a preliminary synthesis. South African

National Scientific Programmes Report No. 74, FRD: CSIR,

Pretoria.

DEALL, G.B. 1985. A plant ecological study of the eastern Transvaal

escarpment in the Sabie area. M.Sc. thesis, University of Pretoria,

Pretoria.

Bothalia 22,2 (1992)

225

Rhynchosia caribaea (Jacq.) DC.

Rhynchosia hirta (Andr. ) Meikle

& Verde.

Rhynchosia komatiensis Harms

Rhynchosia monophylla Schltr.

Rhynchosia ovata Wood & Evans

Rhynchosia stenodon Bak. f.

Rhynchosia thorncroftii (Bak. f.)

Bum Davy

Rhynchosia totta (Thunb.) DC.

Schotia brachypetala Sond.

Schotia afra (L.) Thunb. var. afra

Schotia latifolia Jacq.

Sophora inhambanensis Klotzsch

Sophora tomentosa L.

Sphenostylis marginata E. Mey.

Tephrosia pondoensis (Codd)

Shrire

Tephrosia shiluwanensis Schinz

Teramnus labialis (L. f.) Spreng.

Virgilia divaricata Adamson

Virgilia oroboides (Berg.) Salter

subsp. ferruginea B.-E. van Wyk

subsp. oroboides

Umtiza listeriana Sim

Zornia capensis Pers.

FLACOURTIACEAE 11, 21, 1.9

Aphloia theiformis (Vahl) Benn.

Casearia gladiiformis Mast.

Dovyalis caffra (Hook. f. &

Harv. ) Hook. f.

Dovyalis longispina (Harv.) Warb.

Dovyalis lucida Sim

Dovyalis rhamnoides (Burch, ex

DC.) Harv.

Dovyalis rotundifolia (Thunb.)

Thunb. & Harv.

Dovyalis sp.

Dovyalis zeyheri (Sond.) Warb.

Gerrardina foliosa Oliv.

Homalium dentatum (Harv.)

Warb.

Homalium rufescens Benth.

Kiggelaria africana L.

Pseudoscolopia polyantha Gilg

Rawsonia lucida Harv. & Sond.

Scolopia mundii (Eckl. & Zeyh.)

Warb.

Scolopia oreophila (Sleum.)

Killick

Scolopia zeyheri (Nees) Harv.

Trimeria grandifolia (Hochst.)

Warb.

Trimeria trinervis Harv.

Xylotheca kraussiana Hochst.

FLAGELLARIACEAE 1, 1, 1.0

Flagellaria guineensis Schumach.

GENTIANACEAE 2, 3, 1.5

Chironia laxa Gilg

Chironia peglerae Pram

Neurotheca schlechteri Gilg ex

Bak. & N.E. Br.

GERANIACEAE 3, 6, 2.0

Geranium omithopodon Eckl. &

Zeyh.

Monsonia natalensis Knuth

Pelargonium alchemilloides (L.)

L'Herit.

Pelargonium cordifolium (Cav.)

Curtis

Pelargonium papilionaceum (L.)

L’Herit.

Pelargonium zonale (L.) L'Herit. 4

FOREST

GF Fre P G S A E T U H R MC S MN

evomlrmailaaae

nbcacmtwbcpbkt

6 10 00111110101111

6 1 00000000000100

3 1 00000000000100

6 1 00000000000100

3 1 00000000100000

6 1 00000000100000

3 1 00000000000100

62 00000000110000

1 3 00000100100010

1 2 00101000000000

1 3 00111000000000

3 1 00000000010000

3 3 00000010100100

3 1 00000010000000

3 2 00000010000100

6 1 00000000000001

1 1 00100000000000

12 11 000000000000

2 1 00001000000000

12 1 00000000100000

GESNERIACEAE 1, 12, 12.0

Streptocarpus confusus Hilliard

Streptocarpus cyaneus S. Moore

Streptocarpus gardenii Hook.

Streptocarpus haygarthii N.E. Br.

ex C.B. Cl.

Streptocarpus micranthus C.B. Cl.

Streptocarpus parviflorus Hook. f.

Streptocarpus polyanthus Hook.

Streptocarpus porphyrostachys

Hilliard

Streptocarpus primulifolius Gand.

Streptocarpus pusillus Harv. ex

C.B. Cl.

Streptocarpus rexii (Hook.) Lindl.

Streptocarpus wilmsii Engl.

GLEICHENIACEAE 2, 3, 1.5

Dicranopteris linearis (Burn. f. )

Underw.

Gleichenia polypodioides (L.)

J.E. Sm.

Gleichenia umbraculifera (Kunze)

T. Moore

FOREST

GF Fre P G S A E T U H RMC S MN

evoml rmai laaae

nbcacmtwbcpbkt

2 00110000000000

12 1 00000000000010

12 2 00000000000110

12 1 00000000001000

12 1 00000010000000

12 1 00000000000010

12 2 00000000000011

12 2 00000010000100

12 1 00000010000000

12 1 00000000001000

12 4 00111100000000

12 1 00000000000010

8 1 00000000000100

86 11110010000010

82 00000100001000

3 1 00000000000001

3 1 00000010000000

2 3 00001000010010

23 00000001110000

2 6 00110110100100

3 8 00101111110010

2 3 00111000000000

3 1 00000000010000

2 5 00010100001101

2 1 00000010000000

1 3 00000010000011

3 1 00000010000000

1 10 11110 1 1 1 0 0 0 111

3 1 00 0J3 0010000000

37 00000110110111

1 10 11110 110 11110 0

2 1 00000100000000

1 8 00111110110001

2 10 00111110101111

23 00011100000000

3 4 00000101110000

65 00001011110000

12 1 00000010000000

12 1 0000000000 1000

12 1 00000000010000

GREYIACEAE 1, 1, 1.0

Greyia radlkoferi Szyszyl.

HALORAGACEAE 1, 1, 1.0

Laurembergia repens Berg.

HAMAMELIDACEAE 1, 3, 3.0

Trichocladus crinitus (Thunb.)

Pers.

Trichocladus ellipticus Eckl. &

Zeyh. ex Walp.

Trichocladus grandiflorus Oliv.

HYMENOPHYLLACEAE 2, 6, 3.0

Hymenophyllum capense Schrad.

Hymenophyllum marlothii Brause

Hymenophyllum peltatum (Poir. )

Desv.

Hymenophyllum polyanthos

Swartz

Hymenophyllum tunbridgense (L.)

J.E. Sm.

Trichomanes pyxidiferum L.

HYPOXIDACEAE 1, 3, 3.0

Hypoxis membranacea Bak.

Hypoxis rooperi S. Moore

Hypoxis sp.

ICACINACEAE 3, 6, 2.0

Apodytes dimidiata E. Mey. ex

Am.

Apodytes sp. nov.

Cassinopsis ilicifolia (Hochst.)

Kuntze

Cassinopsis tinifolia Harv.

Pyrenacantha grandiflora Baill.

F’yrenacantha scandens Planch,

ex Harv.

2 1 00000000000100

12 4 00110010000001

3 3 01100010000000

3 6 0011 1 1 1 000000 1

3 3 00000010000110

93 10100000000010

9 1 00100000000000

9 1 00000000000010

94 11100000000010

97 00110010001111

10 1 00000010000000

10 1 00000000100000

10 1 00100000000000

1 13 11111110 111111

3 1 00000010000000

3 6 01110100001100

3 1 00000010000000

5 2 00000000000011

64 01100000110000

12 3 00010010100000

3 1 00000010000000

42 00010000100000

42 00110000000000

4 1 00100000000000

ILLECEBRACEAE 1, 3, 3.0

Silene bellidioides Sond.

Silene burchellii Otth

Silene undulata Ait.

IRIDACEAE 7, 8, 1.1

Anomatheca laxa (Thunb.)

Goldbl.

Aristea ecklonii Bak.

12 1 00100000000000

12 1 00000000100000

12 2 00110000000000

10 2 00000000100010

10 4 00010010000011

226

Bothalia 22,2 (1992)

FOREST

GF Fre PGSAETUHRMCSMN

evomlrmailaaae

i be acmtwbc pbkt

FOREST

227

Bothalia 22,2 (1992)

FOREST

GF Fre PGSAETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

Cyphia sylvatica Eckl.

FOREST

GF Fre PGSAETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

228

Bothalia 22,2 (1992)

FOREST

PGSAETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

111111 10000000

00110000000000

00000000100000

00000000100001

00 100000000000

00 1 100000000 1 0

0 1 10 1000000000

00 1 100000000 1 0

00000000000100

00100000100000

00010010000100

000000000001 1 1

00100000000001

00000010000000

00100010100001

0000000000001 1

00000100001 100

00 1 1000000000 1

00010000000000

001000001 01000

00010000000000

00000000100 100

00100000000010

00010100000000

10000000000000

00010010001000

00100000000000

00010100001000

00000000000001

00000000001001

00100000000000

00100010000000

00000001 1 10000

00 100000000000

00000100000001

00100001000010

00010001100000

00010000000000

00000010000001

00000000000001

00000000010000

00000000100000

00000000000001

000000000001 0 1

01110100001111

00010000100000

000001000000 1 0

00000000000001

00 1 10000000000

00000000000001

000 1 0 10000000 1

Bothalia 22,2 (1992)

229

Stenoglottis fimbriata Lindl.

Tridactyle bicaudata (Lindl.)

Schltr.

Tridactyle tricuspis (H. Bol.)

Schltr.

Tridactyle tridentata (Harv. )

Schltr.

OSMUNDACEAE 2, 2, 1.0

Osmunda regalis L.

Todea barbara (L.) T. Moore

OXALIDAC E AE 1, 4, 4.0

Oxalis incamata L.

Oxalis purpurea L.

Oxalis semiloba Sond.

Oxalis stellata Eckl. & Zeyh.

var. gracilior Salter

PASSIFLORACEAE 1, 3, 3.0

Adenia digitata (Harv.) Engl.

Adenia gummifera (Harv. ) Harms

Adenia hastata (Harv.) Schinz

PEDALIAC E AE 1, 1, 1.0

Cera to theca triloba (Bemh.)

Hook. f.

PERIPLOCAC E AE 4, 5, 1.3

Cryptolepis capensis Schltr.

Cryptolepis oblongifolia ( Meisn .)

Schltr.

Mondia whitei (Hook. f. ) Skeels

Petopentia natalensis (Schltr.)

Bullock

Tacazzea apiculata Oliv.

PHYTOLACCACEAE 1, 2, 2.0

Phytolacca americana L.

Phytolacca octandra L.

PIPERACEAE 2, 4, 2.0

Peperomia blanda (Jacq.) H.B.K.

Peperomia retusa (L. f ) A. Dietr.

Peperomia tetraphylla (G. Forst.)

Hook. & Am.

Piper capense L. f.

PITTOSPORACEAE 1, 1, 1.0

Pittosporum viridiflorum Sims

PLUMBAGINACEAE 2, 3, 1.5

Limonium scabrum (Thunb .)

Kuntze

Plumbago auriculata Lam.

Plumbago zeylanica L.

POACEAE 34, 57, 1.7

Agrostis lachnantha Nees

Aristida junciformis Trin. &

Rupr.

Brachiaria chusqueoides (Hack.)

Clayton

Brachypodium flexum Nees

Cymbopogon validus ( Stapf) Stapf

ex Bum Davy

Dactyloctenium australe Steud.

Dactyloctenium geminatum Hack.

Digitaria diversinervis (Nees)

Stapf

Digitaria eriantha Steud.

Digitaria natalensis Stent

Ehrharta calycina J.E. Sm.

Ehrharta capensis Thunb.

Ehrharta erecta Lam.

var. erecta

var. natalensis Stapf

Ehrharta rehmannii Stapf

FOREST

PGS AETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

00100000000000

00001000000000

000000001 10000

00000000 1 1 0000

00000000100100

00000000 1 10000

00100000000000

00000000100100

00000000 1 10000

00000000 1 10 100

00000000000100

00 1 000000000 1 1

0000000000 1000

01111110101111

000 1 1000 1 1 1000

00111011110110

00010010001 1 10

0000000 1 1 10000

00010001110110

00010000000000

00100000000000

00000000000001

00000000 1 10000

00000000100000

00000000000100

0001001000001 1

000000000000 1 1

00000000100000

00000000 1 1 1 10 1

00000000 1 1 1 1 1 1

00000001000000

00 100000000000

00000000010000

00000000 1 10000

00101000100000

11101 10000 1000

00 1 1 1000000000

00001000100000

000000001 10000

00000000000100

000000001 10000

01111100101011

00000100001000

1 1 1 10 1 1000 1 1 1 1

0000000000 1000

00000010000000

00100000000000

00000000100100

101 1000000 1000

00010010 100000

230

Bothalia 22,2 (1992)

Polygala serpentaria Eckl. &

Zeyh.

Polygala sp.

Polygala virgata Thunb.

POLYGONACEAE 3, 3, 1.0

Oxygonum dregeanum Meisn.

Polygonum salicifolium Willd.

Rumex sagittatus Thunb.

POLYPODIACEAE 6, U, 1.8

Loxogramme lanceolata (Swartz)

Presl

Microgramma lycopodioides (L.)

Cope l .

Microsorium ensiforme (Thunb.)

Schelpe

Microsorium punctatum (L.)

Cope l .

Microsorium scolopendrium

(Burnt, f ) Copel.

Pleopeltis excavata (Bory ex

Willd.) Sledge

Pleopeltis macrocarpa (Bory ex

Willd.) Kaulf.

Pleopeltis schraderi (Mett.)

Tardieu-Blot

Pleo podium simianum Schelpe &

N.C. Anthony

Polypodium polypodioides (L.)

Hitchc.

Polypodium vulgare L.

PORTULACACEAE 1, 1, 1.0

Portulacaria afra Jacq.

PRIMULACEAE 1, 1, 1.0

Samolus valerandi L.

PROTEACEAE 2, 3, 1.5

Brabejum stellatifolium L.

Faurea macnaughtonii Phill.

Faurea speciosa (Welw. ) Welw.

PSILOTACEAE 1, 1, 1.0

Psilotum nudum (L.) Beauv.

PTAEROXYLACEAE 1, 1, 1.0

Ptaeroxylon obliquum (Thunb.)

Radik.

RANUNCULACEAE 4, 8, 2.0

Clematis brachiata Thunb.

Knowltonia cordata H. Rasm.

Knowltonia bracteata Harv. ex

Zahlbr.

Knowltonia filia (L. f. ) Dur.

& Schinz subsp. scaposa

H. Rasm.

Knowltonia vesicatoria (L. f.)

Sims

subsp. grossa H. Rasm.

subsp. humilis H. Rasm.

Ranunculus multifidus Forssk.

Thalictrum rhynchocarpum Dill.

& Rich.

RHAMNACEAE 6, 8, 1,3

Berchemia discolor (Klotzsch)

Hemsl.

Berchemia zeyheri (Sond.)

Grubov

Helinus integrifolius (Lam.)

Kuntze

Phylica buxifolia L.

Phylica paniculata Willd.

FOREST

OF Fre PGS AETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

3 1 00000010000000

12 2 000 1 0 100000000

3 5 00010000101 1 10

12 1 00000000 100000

12 5 001 10010100100

6 8 0 1 1 100 10001 1 1 1

9 1 000000000000 1 0

9 1 000000 10000000

9201 100000000000

9 3 0000001 0110000

9 2 000000 10110000

9 2 000000000000 1 1

9 10 11111010001111

96 001101 10001 0 1 0

9 2 001 10000000000

9 9 00011110101111

8 3 00010100001000

3 1 00000010000000

12 2 0000 1000100000

2 1 10000000000000

1 3 00100010000010

2 2 000000000001 10

8 2 00000010100000

2 6 00011110010010

5 8 00110010101111

12 2 00 1 10000000000

12 1 00000 100000000

12 1 00 100000000000

12 3 1 1 100000000000

12 1 00 100000000000

12 7 011100 10101001

12 6 0001010000 1111

2 1 000000000 10000

2 2 00000000000 1 1 0

3 6 00011010110010

3 1 10000000000000

3 2 000000 10000100

Rhamnus prinoides L'Herit.

Scutia myrtina (Burn. f. ) Kurz

Ziziphus mucronata Willd.

RHIZOPHORACEAE 2, 5, 2.5

Bruguiera gymnorrhiza (L.) Lam.

Cassipourea flanaganii (Schinz)

Alston

Cassipourea gerrardii (Schinz)

Alston

Cassipourea gummiflua Tul.

Cassipourea malosana (Bak.)

Alston

ROSACEAE 5, 9, 1.8

Alchemilla capensis Thunb.

Alchemilla rehmannii Engl.

Cliffortia linearifolia Eckl. &

Zeyh.

Cliffortia serpyllifolia Cham. &

Sclechtd.

Leucosidea sericea Eckl. & Zeyh.

Prunus africana (Hook, f) Kalkm.

Rubus pinnatus Willd.

Rubus rigidus J.E. Sm.

Rubus rosifolius J.E. Sm.

RUBIACEAE 31, 66, 2.1

Alberta magna E. Mey.

Anthospermum herbaceum L. f.

Breonadia salicina (Vahl) Hepper

& Wood

Burchellia bubalina (L. f. ) Sims

Canthium ciliatum (Klotzsch)

Kuntze

Canthium gueinzii Sond.

Canthium inerme (L. f.) Kuntze

Canthium mundianum Cham. &

Schlechtd.

Canthium pauciflorum (Klotzsch)

Kuntze

Canthium setiflorum Hiem

Canthium spinosum (Klotzsch)

Kuntze

Canthium sp. nov.

Canthium suberosum Codd

Catunaregam spinosa (Thunb.)

Tirvengadum

Cephalanthus natalensis L.

Coddia rudis (E. Mey. ex Harv. )

Verde.

Coffea racemosa Lour.

Conostomium natalense (Hochst.)

Brem.

Galium mucroniferum Sond.

Galium thunbergianum Eckl. &

Zeyh.

Galopina circaeoides Thunb.

Gardenia thunbergia L. f.

Hyperacanthus amoenus (Sims)

Bridson

Kraussia floribunda Harv.

Lagynias lasiantha (Sond.)

Bullock

Mitriostigma axillare Hochst.

Otiophora cupheoides N.E. Br.

Oxyanthus speciosus DC.

subsp. gerrardii (Sond.) Bridson

Pachy stigma cymosum Robyns

Pachystigma macrocalyx (Sond.)

Robyns

Pavetta barbertonensis Brem.

F’avetta bowkeri Harv.

Pavetta capensis (Houtt.) Brem.

subsp. komghensis (Brem.)

Kok

Bothalia 22,2 (1992)

231

Pavetta cooperi Harv. & Sond.

Pavetta galpinii Brem.

Pavetta gerstneri Brem.

Pavetta gracilifolia Brem.

Pavetta inandensis Brem.

Pavetta kotzei Brem.

Pavetta lanceolata Eckl.

Pavetta natalensis Sond.

Pavetta revoluta Hochst.

Pavetta sp.

Pentanisia prunelloides (Eckl. &

Zeyh.) Walp.

Pentas micrantha Bak.

Plectroniella armata (K. Schum.)

Robyns

Psychotria capensis (Eckl.) Vatke

Psychotria zombamontana

(Kuntze) Petit

Psydrax livida (Hiem) Bridson

Psydrax locuples (K. Schum.)

Bridson

Psydrax obovata (Eckl. & Zeyh.)

Bridson subsp. obovata

Rothmannia capensis Thunb.

Rothmannia globosa (Hochst.)

Keay

Rubia cordifolia L.

Rubia petiolaris DC.

Tarenna junodii (Schinz) Brem.

Tarenna littoralis (Hiem) Bridson

Tarenna pavettoides (Harv. ) Sim

Tarenna supra-axillaris (Hemsl.)

Brem.

Tricalysia capensis (Meisn.) Sim

Tricalysia lanceolata (Sond.) Burn

Davy

Tricalysia sonderiana Hiem

Vangueria cyanescens Robyns

Vangueria esculenta S. Moore

Vangueria iniausta Burch.

Vangueria randii S. Moore subsp.

chartacea (Robyns) Verde.

RUTACEAE 7, 10, 1.4

Calodendrum capense (L. f.)

Thunb.

Clausena anisata (Willd.) Hook. f.

ex Benth.

Oricia bachmannii (Engl.)

Verdoom

Teclea gerrardii Verdoom

Teclea natalensis (Sond.) Engl.

Toddalia asiatica (L.) Lam.

Vepris undulata (Thunb.) Verdoom

& C.A. Sm.

Zanthoxylum davyi (Verdoom)

Waterm.

Zanthoxylum capense (Thunb.)

Harv.

Zanthoxylum thomcroftii

(Verdoom) Waterm.

SALVADORACEAE 1, 1, 1.0

Azima tetracantha Lam.

SANTALACEAE 3, 3, 1.0

Colpoon compressum Berg.

Osyndicarpos schimperianus

(Hochst. ex A. Rich.) A. DC.

Rhoiacarpos capensis (Harv.) A.

DC.

SAPINDACEAE 8, 12, 1.5

Allophylus decipiens (Sond.)

Radik.

FOREST

GF Fre PGSAETUHRMCSMN

evomlrmailaaae

nbcacmtwbcpbkt

3 3 00000 1 0000 1 1 00

3 2 00000010000100

3 1 00000000010000

3 1 00000010000000

3 1 00010000000000

2 8 000 1 1 1 10 1 100 1 1

3 1 00000010000000

3 4 000010011 10000

3 1 00000000010000

12 1 000000000000 10

12 1 0000000000000 1

1 1 00000000010000

3 11 00111111110111

2 2 00000000000 1 1 0

2 2 00000000000 1 0 1

2 1 00000000000100

1 11 01111111110101

2 8 0 1 1 10 1 10000 1 1 1

2 8 00 10 1 1 1 1 1 10 100

12 2 000000 10 100000

12 3 00 1 100000000 1 0

3 1 00000000010000

3 2 00000000 1 10000

3 3 00000 1 1 0 100000

3 1 00000000010000

2 6 00000 1 100 10 1 1 1

2 6 0000 1110010110

2 3 0000000 1 1 1 0000

2 1 000000000000 1 0

2 1 00000100000000

2 1 00000000 1 10000

2 3 0000000 1 1 1 0000

1 u o n liiionioii

2 12 0011 1 1 1 1 1 1 1 1 1 1

3 2 00000 1 10000000

2 3 0000000 11 1 0000

29 00011111110011

5 2 00000000000101

1 10 0 1 1 1 1 1 10 1 10 0 1 1

1 7 00110110000111

2 11 0 0 111111110 111

2 1 00000000000100

3 3 00 1 0 1 0 1 0000000

3 7 01111010011000

5 4 000 100 10000 1 0 1

3 1 00001000000000

2 4 00 111 100000000

Allophylus dregeanus (Sond.) De

Winter

Allophylus melanocarpus (Sond.)

Radik.

Allophylus natalensis (Sond.) De

Winter

Allophylus transvaalensis Bunt

Davy

Atalaya natalensis R.A. Dyer

Blighia unijugata Bak.

Deinbollia oblongifolia (E. Mey.

ex Am.) Radik.

Dodonaea angustifolia L. f.

Hippobromus pauciflorus (L. f.)

Radik.

Pancovia golungensis (Hiem)

Exell & Mendonca

Pappea capensis Eckl. & Zeyh.

SAPOTACEAE 7, 13, 1.9

Bequaertiodendron magalismonta-

num (Sond.) Heine & J.H.

Hemsl.

Bequaertiodendron natalense

(Sond.) Heine <4 J.H. Hemsl.

Chrysophyllum viridifolium J.M.

Wood & Franks

Inhambanella henriquesii (Engl.

& Wjrb.) Dubard

Manilkara concolor (Harv. ex

C.H. Wr. ) Gerstner

Manilkara discolor (Sond.) J.H.

Hemsl.

Manilkara nicholsonii Van Wyk

Mimusops caffra E. Mey. ex A.

DC.

Mimusops obovata Sond.

Mimusops zeyheri Sond.

Sideroxylon inerme L.

Vitellariopsis dispar (N.E. Br.)

Aubrev.

Vitellariopsis marginata (N.E. Br.)

Aubrev.

SCHIZAEACEAE 1, 1, 1.0

Mohria caffrorum (L.) Desv.

SELAGINACEAE 1, 1, 1.0

Tetraselago natalensis (Rolfe)

Junell

FOREST

GF Fre PGSAETUHRMCSMN

evomlrmailaaae

nbc acmtwbc pbkt

2 3 00000110100000

2 7 000000 1 1 1 1 1 1 1 0

2 5 0000 1 1 0 11 10000

2 3 00000000000 1 1 1

2 1 00000010000000

1 1 00000000010000

3 5 0000 1 0 1 1 1 10000

3 3 00100010010000

27 00111110100010

3 2 00000000 1 1 0000

2 3 00001000010010

2 1 00000000000100

2 4 00000010110010

2 3 000000 1 10 10000

1 1 00000000010000

1 2 000000001 10000

1 2 00000000 1 10000

2 1 00000010000000

1 4 0000 100 1 1 10000

1 7 00011011110010

1 3 00000000000 1 1 1

1 9 011111111 10000

2 1 00000000010000

2 2 00000010010000

7 7 00 1 10 10000 1 1 1 1

12 1 00000000000 100

SCROPHULARIACEAE 10, 19, 1.9

232

Bothalia 22,2 (1992)

SELAGINELLACEAE 1, 3, 3.0

Selaginella dregei (Presl) Hieron.

Selaginella kraussiana (Kunze) A.

Br. ex Kuhn

Selaginella mittenii Bak.

SIMAROUBACEAE 1, 1, 1.0

Kirkia acuminata Oliv.

SOLANACEAE 2, 10, 5.0

Solanum aculeastrum Dun.

Solanum aculeatissimum Jacq.

Solanum americanum Mill.

Solanum didymanthum Dun.

Solanum geniculatum E. Me y.

Solanum giganteum Jacq.

Solanum hermannii Dun.

Solanum retroflexum Dun.

Solanum terminale Forssk.

Withania somnifera (L.) Dun.

STERCULIACEAE 3, 6, 2.0

Cola greenwayi Brenan

Cola natalensis Oliv.

Dombeya pulchra N.E. Br.

Dombeya rotundifolia (Hochst.)

Planch.

Dombeya tiliacea (Endl.) Planch.

Sterculia mutex Hemsl.

STRELITZIACEAE 1, 4, 4.0

Strelitzia alba (L. f) Skeels

Strelitzia caudata R.A. Dyer

Strelitzia nicolai Regel <6 Koem.

Strelitzia reginae Ait.

THELYPTERIDACEAE 2, 9, 4.5

Macrothelypteris torresiana

(Gaud.) Ching

Thelypteris bergiana (Schlechtd.)

Ching

Thelypteris confluens (Thunb .)

Morton

Thelypteris dentata (Forssk.) E.

St. John

Thelypteris gueinziana (Mett.)

Schelpe

Thelypteris interrupta (Willd.) K.

Iwats.

Thelypteris knysnaensis N.C.

Anthony & Schelpe

Thelypteris madagascariensis

(Fee) Schelpe

Thelypteris pozoi (Lagasca)

Morton

THYMELAEACEAE 6, 11, 1.8

Dais cotinifolia L.

Englerodaphne pilosa Burtt Davy

Englerodaphne ovalifolia (Meisn.)

Phill.

Gnidia denudata Lindl.

Gnidia polyantha Gilg

Gnidia pulchella Meisn.

Gnidia woodii C.H. Wr.

Passerina falcifolia C.H. Wr.

Passerina rigida Wikstr.

Peddiea africana Harv.

Struthiola pondoensis Gilg ex

C.H. Wr.

TILIACEAE 3, 8, 2.7

Grewia caffra Meisn.

Grewia lasiocarpa E. Mey. ex

Harv.

Grewia occidentals L.

FOREST

GF Fre PGS AETUHRMCSMN

e v o m I rmai laaae

nbc acmtwbc pbkt

8 3 000000000001 1 1

8 6 001 10 1000001 1 1

8 2 0000000000001 1

1 1 00000000000100

4 3 00101000000001

12 4 00 1 100000000 1 1

4 1 00001000000000

4 1 00000010000000

5 2 00100000000010

4 6 0 1 1 1000000 10 1 1

4 1 00100000000000

4 2 00010010000000

5 2 00000010000001

3 3 000 1 0000 1 000 1 0

2 1 00000000010000

1 3 0000001 1010000

3 1 00000000000100

1 2 00000000100100

2 2 00001010000000

2 2 000000000001 10

2 1 00100000000000

2 2 00000000000 1 0 1

2 4 00001 0101 10000

12 1 00001000000000

7 1 00000010000000

7 6 0 1 1 10000000 1 1 1

7 4 00 1 10000001001

7 3 00000100100010

7 5 00 100 1 10000 1 1 0

7 2 00100000000100

7 1 00100000000000

7 3 00000 1 00000 1 0 1

7 6 00 1 10 1000001 1 1

2 5 00000 1 1000 10 1 1

3 2 00010100000000

3 1 00000010000000

3201 100000000000

3 1 00000000000010

3 2 00010010000000

3 1 00000010000000

3 1 00100000000000

3 3 0000 1000 1 1 0000

2 8 00000 111110111

3 1 00000010000000

5 3 0000000 1 1 1 0000

3 3 0000 1 1 1 0000000

3 13 11111111110 111

Sparrmannia africana L. f.

Sparrmannia ricinocarpa (Eckl. &

Zeyh.) Kuntze

Triumfetta annua L.

Triumfetta pilosa Roth

Triumfetta rhomboidea Jacq.

TRIM ENIAC E AE 1, 1, 1.0

Xymalos monospora (Harv.)

Baill.

TYPHACEAE 1, 1, 1.0

Typha capensis (Rohrb.) N.E. Br.

ULMACEAE 3, 4, 1.3

Celtis africana Burm. f.

Celtis durandii Engl.

Chaetacme aristata Planch

Trema oriental is (L.) Blume

VELLOZIACEAE 2, 2, 1.0

Talbotia elegans Balf.

Xerophyta retinervis Bak.

VERBENACEAE 5, 7, 1.4

Avicennia marina (Forssk.) Vierh.

Clerodendron glabrum E. Mey.

Clerodendron myricoides

(Hochst.) Vatke

Clerodendron suffruticosum

Guerke

Lantana meamsii Moldenke

Lippia javanica (Burm. f. ) Spreng.

Priva meyeri Jaub. <& Spach

VIOLACEAE 1, 3, 3.0

Rinorea angustifolia (Thouars)

Baill.

Rinorea domatiosa Van Wyk

Rinorea ilicifolia (Welw. ex Oliv.)

Kuntze

VISCACEAE 1, 4, 4.0

Viscum nervosum Hochst. ex A.

Rich.

Viscum obovatum Harv.

Viscum obscurum Thunb.

Viscum rotundifolium L. f.

VITACEAE 4, 14, 3.5

Cayratia gracilis (Guill. & Perr.)

Suesseng.

Cissus fragilis E. Mey. ex Kunth

Cissus quadrangularis L.

Cyphostemma anatomicum (C.A.

Sm.) Wild & Drum.

Cyphostemma cirrhosum (Thunb.)

Descoings ex Wild & Drum.

FOREST

GF Fre PGS AETUHRMCSMN

evomlrmai laaae

nbcacmtwbcpbkt

4 2 01 100000000000

4 3 00010000001001

12 1 00000000000010

3 3 00000010100100

3 2 00000000100100

1 6 000 1 0 1 10000 1 1 1

111 00000000100100

1 13 11111101111111

1 2 00000010100000

3 7 000 1 1 1 1 1 1 10000

2 6 000000101 10 1 1 1

10 1 00000000001000

10 1 00000000000100

2 1 00000000100000

29 00110011110111

2 2 00000001 000 100

2 1 00000000000100

3 1 00000000000100

3 3 000000 1 0 1 00 100

3 2 00010000010000

2 2 00000010000001

2 1 00000010000000

3 2 00000000 1 10000

3 1 00000000000010

3 1 00000000100000

3 5 001 10 1 1 00000 1 0

3 1 00100000000000

6 1 00000000000001

5 2 000000 1 0 100000

5 1 00000000010000

5 3 00000000000 1 1 1

6 6 00011001 1 10001

URTICACEAE 9, 12, 1.3

Bothalia 22,2 (1992)

233

Cyphostemma hypoleucum

(Harv. ) Descoings ex Wild &

Drum.

Cyphostemma sp. nov.

Cyphostemma woodii (Gilg &

Brandt ) Descoings

Rhoicissus digitata (L. f.) Gilg &

Brandt

Rhoicissus revoilii Planch.

Rhoicissus rhomboidea (E. Mey.

ex Harv.) Planch.

Rhoicissus sp.

Rhoicissus tomentosa (Lam.) Wild

& Drum.

FOREST

GF Fre P G S A E T U H RMC S MN

evomlrmailaaae

nbcacmtwbcpbkt

6200 100010000000

5 1 00000010000000

5 1 00000000000100

5 7 011110101 10000

5 6 00010100010011

5 8 000001 11110111

5 2 000000001 10000

5 10 01101111100111

Rhoicissus tridentata (L. f.) Wild

& Drum.

VITTARIACEAE 1, 1, 1.0

Vittaria isoetifolia Bory

ZAMIACEAE 1, 2, 2.0

Encephalartos altensteinii Lehm.

Encephalartos villosus Lem.

ZYGOPHYLLACEAE 1, 2, 2.0

Zygophyllum morgsana L.

Zygophyllum uitenhagense Sond.

FOREST

GF Fre P G S A E T U H RMC S MN

evomlrmailaaae

nbcacmtwbcpbkt

512 01111110111111

9 3 00 1 000 1 00000 1 0

3 1 00001000000000

3 2 00001 0 10000000

4 1 00 100000000000

12 1 0000 1000000000

Bothalia 22,2: 235-243 (1992)

An analysis of the orchid flora of Mt Mulanje, Malawi

H. KURZWEIL*

Keywords: altitudinal distribution, extralimital distribution, floristic composition, Mt Mulanje, Malawi, Orchidaceae

ABSTRACT

The composition of the orchid flora of Mt Mulanje, Malawi, is analysed. The altitudinal distribution of the orchids, the

distribution of the terrestrial and epiphytic species and the extralimital distribution of the species is assessed for both genera

and subfamilies (after the systematic concept of Dressier 1981). The altitudinal distribution of species endemic to Malawi

is also assessed. The terrestrial species show a significant increase with altitude whereas the epiphytic species are more

dominant at the lower levels. A similar increase is also found in the species endemic to Malawi. An analysis of the distributions

shared with other African countries reveals that most species also occur in Zambia, Zimbabwe and East Africa, whereas

significantly fewer species are shared with Angola, southern Africa, Mocambique, Zaire and West Africa. Most species

shared with tropical African countries are found on the lower slopes of Mt Mulanje.

UITTREKSEL

Die samestelling van die orgidee-flora van Mt Mulanje, Malawi, word ondersoek. Die verspreiding van die orgidee bo

seevlak, die verspreiding van die terrestriele en epifitiese spesies, en die verspreiding van die spesies buite Mt Mulanje

word vir genusse sowel as subfamilies (volgens die sisteem van Dressier 1981) bepaal. Die verspreiding bo seevlak van die

soorte endemies aan Mt Mulanje word ook ondersoek. Die aantal terrestriele spesies neem aansienlik toe met hoogte bo

seevlak, terwyl epifitiese spesies meer dominant laer af is. Spesies endemies aan Malawi toon ’n soortgelyke toename. ’n

Analise van die verspreidings toon dat die meeste soorte ook in Zambie, Zimbabwe en Oos-Afrika voorkom. Minder spesies

word egter met Angola, suidelike Afrika, Mosambiek, Zaire en Wes-Afrika gedeel. Die meeste spesies wat ook in tropiese

Afrika voorkom, word teen die onderste hange van Mt Mulanje aangetref.

INTRODUCTION

The Mulanje massif (Figure 1) is an isolated mountain

block in the southeastern corner of Malawi. The massif

is comparatively small, covering only about 640 km2

(Eastwood 1988; measured at the 800 m contour). From

the surrounding plains at around 600—700 m the steep

slopes rise abruptly to plateaux at 1 800—2 000 m, which

are divided by ridges and peaks of up to 3 000 m. The

slopes and the plateaux offer a wide variety of different

habitats, ranging from lowland rain forests (Figure 2A)

and savanna woodland (Figure 2B) to montane grassland

(Figure 2C) and seepage areas overlying bare rock (Figure

2D). Almost the entire mountain massif is now a Forest

Reserve, but very few of the lowland forests at the base

of the mountain and on the lower slopes are left following

the planting of tea and coffee.

The botanical exploration of Mt Mulanje began in 1891

(Chapman 1962) when Alexander Whyte made the first

extensive collections. The earliest extensive botanical

survey of the mountain was that of Chapman (1962) on

the vegetation and the plant communities of Mt Mulanje,

in which also climatological, geological and soil data were

accumulated. Particularly the forest flora has since then

attracted a great deal of attention, and has been the subject

of several exhaustive publications (most notably Dowsett-

Lemaire 1988).

The orchid flora of Mt Mulanje is one of the richest

in Malawi and is especially rich in terrestrial species.

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

Claremont 7735.

MS. received: 1992-04-21.

Although the orchid flora of the whole of Malawi has been

written up already (La Croix et al. 1991; epiphytic orchids

only: La Croix et al. 1983; Morris 1970), neither a local

orchid flora of Mt Mulanje nor a complete checklist are

available to date. However, some detailed information on

the distribution of orchids on the massif was given in the

orchid flora of Malawi (La Croix et al. 1991) as well as

by Chapman (1962), Dowsett-Lemaire (1988) and in some

taxonomic papers.

As Mt Mulanje is relatively isolated and may be a

halfway station between the eastern highlands of Zimbabwe

and the large upland areas of Nyika and southern Tanza-

nia, an analysis of the composition of the orchid flora of

Mt Mulanje is of interest. Consequently, special attention

is paid to the altitudinal distribution of orchids, to the dis-

tribution of terrestrial and epiphytic species, to the distri-

bution of the species in other African countries, and to

the distribution of species endemic to Malawi. It may be

somewhat premature for an analysis of this kind as the

data base may be incomplete, but it is unlikely that fur-

ther records will affect the main results of the study. The

mountain massif has been divided into four altitudinal

zones:

1, Mulanje base; this area comprises lowland rain forests

in the adjacent parts of the surrounding plain as well as

the foothills; it is now largely confined to forest patches

in cultivated land; approximately up to 900 m;

2, slopes and gorges; savanna woodland and mid-altitude

to afro montane forests; approximately 900-1 900 m;

3, plateaux; grassland and afromontane forest; approxi-

mately 1 700—2 300 m;

4, peaks and ridges above the plateaux.

236

Bothalia 22,2 (1992)

FIGURE 1. —Map of the Mt Mulanje

Massif. Major settlements, • ;

peaks, ▲ ; mountain huts, ■ ;

R, rivers. Dotted lines indicate

footpaths. Tea Estates and

other localities: 1, Lujeri

Estate; 2, Likhubula Forest

Station; 3, Chikatali Estate; 4,

Esperanza Estate; 5, Pwazi

Estate; 6, Mlanje Boma; 7,

Limbuli Estate; 8, Mimosa

Estate; 9, Boma Cottage; 10,

Chisambo.

RESULTS AND DISCUSSION

The orchid flora

A total of 173 orchid species in 45 genera have been

recorded on Mt Mulanje (Tables 1, 2). A recent enumera-

tion of orchids has also been provided for the Nyika Plateau

in northern Malawi (Williamson 1979), and some com-

parisons between Mulanje and Nyika are made below.

However, such comparisons are somewhat problematic as

low-lying areas were not included in Williamson’s (1979)

study.

Almost all of the southern African terrestrial orchid

genera are represented on Mt Mulanje. Most of them

reveal affinities to the orchid floras of South Africa and

south central Africa where the largest concentration of

species as well as most of the related species are found

(Disa, Brownleea, Satyrium, Disperis, Schizochilus,

Neobolusia, Stenoglottis ). Disa, Disperis and Satyrium are

rather poor in species numbers on Mt Mulanje when com-

pared to Nyika, which is probably partly due to the

considerably larger grassland area on Nyika. As far as

Disa is concerned, the watershed between the rivers Zaire

and Zambesi which includes the Nyika Plateau is generally

rich in Disinae and was considered an important centre

of the subtribe (Linder 1983). As in most other African

countries the large pantropical genera Habenaria and

Eulophia are well represented on Mt Mulanje. Eulophia

(20 species) is the largest terrestrial orchid genus, followed

by Habenaria (14 species) and Satyrium (12 species; Tables

1, 3). This is in contrast to the Nyika Plateau where

Habenaria as well as Disa and Satyrium outnumber

Eulophia (Williamson 1979). However, the abundance of

low-lying areas on Mt Mulanje partly accounts for the

predominance of Eulophia. Only six species of the largely

Madagascan genus Cynorkis occur on Mt Mulanje. Several

other terrestrial orchid genera occur with a single or few

species, including the palaeotropical genus Nervilia, the

primarily Asian genus Calanthe, the cosmopolitan genera

Liparis and Malaxis, and the southern African elements

Herschelianthe and Monadenia. It is interesting that one

largely terrestrial species belongs to the primarily epiphytic

genus Polystachya.

It is interesting to note that three terrestrial orchid

genera, Bonatea, Centrostigma and Pterygodium, occur-

ring in southern and East Africa do not occur on Mt

Mulanje. Pterygodium, although well represented in South

Africa and also occurring in Tanzania, has not yet been

collected in Malawi or central Africa. Corycium, closely

related to Pterygodium and with a similar distribution (well

represented in South Africa, also known in Tanzania;

P. Cribb pers. comm.) has only recently been discovered

in a rather restricted area on Mt Mulanje (Kurzweil,

unpublished). Possibly the lack of Pterygodium in central

Bothalia 22,2 (1992)

237

Africa is similarly the result of under-collecting of certain

areas. Further terrestrial genera which occur in Malawi

but have not yet been found on Mt Mulanje are Zeuxine,

Epipactis, Epipogium, and Phaius.

Because of the occurrence of lowland forests at the base

of the mountain and on the lower slopes, an abundance

of epiphytes have been recorded in the past, although some

species have been recorded before the expansion of the

tea and coffee plantations and may since have disappeared.

The epiphytic orchids mainly belong to the genera

Polystachya, Bulbophyllum, Aerangis, Angraecum, Cyr-

torchis, Diaphananthe, Tridactyle and their allies, although

epiphytic orchids are also known in some primarily

terrestrial genera (Disa, Brownie ea, Stenoglottis). Liparis

is represented by terrestrial as well as epiphytic species.

The latest epiphytic genera are the predominantly African

genus Polystachya (20 species) and the large pantropical

FIGURE 2. — Various habitats on Mt Mulanje. A, lowland forest; B, savanna woodland; C, plateau grassland, Satyrium trinerve in the foreground;

D, seepage area on bare rock, here colonized by Satyrium rhynchantoides.

238

Bothalia 22,2 (1992)

TABLE 1. —The genera represented on Mt Mulanje, their systematic po-

sition, worldwide number of species as well as number of species

recorded on the mountain massif, and their overall distribution

(classification after Dressier 1981, number of species after La

Croix el al. 1991).

c = cosmopolitan; pantr = pantropical; palaeotr = palaeotropical;

Afr = primarily African; Mad = primarily Madagascan; As = primarily

Asian

genus Bulbophyllum (15 species; Tables 1, 3). Several ol

the species have their southern-most and western-most

limits of distribution in Malawi (see also Dowsett-Lemaire

1989). Comparatively few of the African epiphytic genera

are not found on the mountain (e.g. Chamaeangis,

Solenangis, Ancistrorrhynchus) . The epiphytic genera

12 3 4

ZONE

■iSPIR H ORCH CZlEPID IH VAND

FIGURE 3. — Distribution of subfamilies in the zones 1—4.

belonging to the tribe Vandeae clearly outnumber all other

epiphytic genera, which, however, may partly also be the

result of extensive taxonomic splitting in this tribe.

The systematic position of the genera found on Mt

Mulanje is shown in Table 1. The classification follows

Dressier (1981) throughout. All of the orchid subfamilies

are represented except the non-African Cypripedioideae

and Apostasioideae. However, the subfamily Spiranthoi-

deae is very poorly represented (Table 4), and is confined

to the base of the mountain (Figure 3). The three remaining

subfamilies Orchidoideae, Epidendroideae and Vandoideae

are rich in species, and all their African tribes and

subtribes except Vanilleae and Epipogieae (both Epiden-

droideae) are represented. The percentage of Orchidoideae

increases significantly with altitude, and all of the orchids

found in zone 4 belong to this subfamily (Figure 3).

Evidently the two subfamilies Epidendroideae and Vandoi-

deae are less dominant at higher levels than they are in

zone 1.

Terrestrial and epiphytic species

Whereas the overall percentage of terrestrial and

epiphytic species on the mountain is almost equal (51%

and 49%, respectively), significant altitudinal differences

exist (Table 2 and Figure 4). Only 30% of the species grow

terrestrially at the Mulanje base, but the terrestrial species

account for 44% in zone 2 and for 61% in zone 3. The

orchids in zone 4 are exclusively terrestrial. Evidently, the

percentage of epiphytic orchids decreases with altitude.

Extralimital distribution of the species

The Africa-wide distributions of the species recorded

on Mt Mulanje were arranged after zones as well as after

10° [ MM

12 3 4

ZONE

M TERRESTRIAL EPIPHYTIC

FIGURE 4. —Percentage of terrestrial and epiphytic orchids in zones 1—4.

Bothalia 22,2 (1992)

239

TABLE 2. — The number of orchid species recorded on Mt Mulanje, the number of terrestrial and epiphytic species and their altitudinal distribution.

The altitudinal distribution and the distribution of terrestrial and epiphytic species is also summarized for the subfamilies and for the whole

orchid family. An asterisk indicates taxa with species which may be epiphytic or terrestrial

subfamilies and are shown in Table 5. The figures for the

zone 4 and those for the subfamily Spiranthoideae are

evidently statistically less reliable than the others as

the number counted is rather low (n=10 and n=2,

respectively).

Within Malawi, the highest number of species is shared

with the remaining parts of the Southern Region (particu-

larly the Zomba Plateau) where 80 % of all Mulanje species

occur. Sixty eight percent of orchids are shared with the

Northern Region. The comparatively high species number

in the Northern Region is mainly due to the occurrence

of two orchid-rich areas, the Viphya Plateau and the Nyika

Plateau. Only 44% of the species from Mulanje have also

been recorded from the Central Region. Sixty-six to 77 %

TABLE 3. — The ten largest genera occurring on Mt Mulanje. The number

of species follows La Croix et al. (1991)

240

Bothalia 22,2 (1992)

TABLE 4. —The division of orchid genera and species on Mt Mulanje

according to subfamilies. The classification follows Dressier

(1981). Numbers of genera and species after La Croix et al. (1991)

of the orchids found on Mt Mulanje also occur in Zambia,

East Africa and Zimbabwe, while considerably fewer

species are shared between Mt Mulanje and Angola,

southern Africa, Mozambique, Zaire and West Africa.

The figure for Madagascar is, as could be expected, the

lowest.

Most orchids shared with tropical, continental African

countries occur in zone 1, and their number successively

decreases with altitude on Mt Mulanje (Table 5). The sharp

rise in zone 4 in the orchids shared with Madagascar is

noteworthy although it may partly also be the result of a

low overall number of species (n=10). This is to a lesser

extent also found in the orchids shared with West Africa.

It is also interesting that the tropically widespread spe-

cies are most common in zone 1, but they are missing in

zone 4.

The Spiranthoideae are represented by widespread

species only (Table 5). Rather surprisingly, the shared dis-

tributions are similar in the two subfamilies Orchidoideae

and Vandoideae. The Epidendroideae stand out due to the

high percentage of species shared with Madagascar, East

and West Africa, and their low percentage of species

shared with southern Africa. This is obviously the result

of the distribution of Bulbophyllum to which the majority

of the Epidendroideae on Mt Mulanje belong. The Epiden-

droideae are also significantly more frequently tropically

widespread in their distribution than Orchidoideae and

Vandoideae.

MALAWI ENDEMICS M S MALAWI ENDEMICS

□ MT MULANJE ENDEMICS

FIGURE 5. —Percentage of endemic species in zones 1—4.

Endemic species

A list of endemic species is given in Table 6. Four

species are endemic to Mt Mulanje, three of them being

terrestrial. A further six species recorded on Mt Mulanje

are endemic to the South Region, and two of the species

are endemic to Malawi. It is interesting to note that the

percentage of endemic species increases with altitude

(Figure 5). This is, however, due to the increase of the

S Malawi and the Mt Mulanje endemics, whereas the

overall Malawi endemics decrease with altitude.

The number of endemics on Mt Mulanje is considerably

lower than on the Nyika plateau where 18 ‘forms of

life’ (probably varieties, subspecies and species) are

thought to be endemic including six species (Williamson

1979).

ACKNOWLEDGEMENTS

I am most grateful to Dr H.R Linder for critically

reading the manuscript and making valuable comments

on the text. I also wish to thank Mr J.R Roux for technical

assistance and for translating the abstract, and Mrs Jeanette

TABLE 5. —Percentages of the orchids shared with other African countries, arranged after zones 1—4 as well as after subfamilies (classification

after Dressier 1981). Records in tropical Africa after La Croix et al. (1991); in southern Africa from Stewart et al. (1982)

S = Malawi, Southern Region excluding Mt Mulanje; C = Malawi, Central Region; N = Malawi, Northern Region; Zam = Zambia;

Zim = Zimbabwe; Ang = Angola; SA = southern Africa; Moc = Mozambique; EA = East Africa; Mad = Madagascar; Zai = Zaire;

WA = West Africa; Trap = percentage of species which are widespread in tropical Africa.

Bothalia 22,2 (1992)

241

TABLE 6. —Species endemic to Malawi recorded on Mt Mulanje

Loedolff for the photographic work. The use of litera-

ture held in the Bolus Herbarium library is gratefully

acknowledged.

REFERENCES

CHAPMAN, J.D. 1962. The vegetation of the Mlanje Mountains,

Nyasaland. Government Printer, Zomba.

CRIBB, P.J. 1978. New orchids from South Central Africa. Kew Bulletin

32: 137-187.

CRIBB, P.J. 1979. A revision of Stolzia (Orchidaceae). Kew Bulletin 33:

79-89.

CRIBB, P.J. & STEWART, J. 1985. Additions to the orchid flora of tropical

Africa. Kew Bulletin 40: 399—419.

DOWSETT-LEMAIRE, F. 1988. The forest vegetation of Mt Mulanje

(Malawi): a floristic and chorological study along an altitudinal

gradient (650—1 950m). Bulletin du Jardin Botanique National

de Belgique 58 : 77—107.

DOWSETT-LEMAIRE, F. 1989. The flora and phytogeography of the

evergreen forests of Malawi I: Afromontane and mid-altitude

forests. Bulletin du Jardin Botanique National de Belgique 59:

3-131.

DRESSLER, R.L. 1981. The orchids — natural history and classifica-

tion. Harvard University Press, Cambridge, Massachusetts and

London.

EASTWOOD, F. 1988. Guide to the Mulanje Massif Lorton Commu-

nications, Johannesburg.

LA CROIX, I.F., LA CROIX, E.A.S., LA CROIX, T.M., HUSTON,

J.A. & JOHNSTON-STEWART, N.G.B. 1983. Malawi orchids.

Vol. 1. Epiphytic orchids. National Fauna Preservation Society

of Malawi.

LA CROIX, I.F., LA CROIX, E.A.S. & LA CROIX, T.M. 1991. Orchids

of Malawi. Balkema, Rotterdam, Brookfield.

LINDER, H.P. 1980. An annotated revision of the genus Schizochilus

Sond. (Orchidaceae). Journal of South African Botany 46:

379-434.

LINDER, H.P. 1983. The historical phytogeography of the Disinae

(Orchidaceae). Bothalia 14: 565 —570.

LINDER, H.P. 1985. Notes on the orchids of southern tropical Africa

I: Brownleea and Herschelia. Kew Bulletin 40: 125—129.

MORRIS, B. 1970. The epiphytic orchids of Malawi. The Society of

Malawi.

STEWART, J. 1980. A revision of the African species of Aerangis

(Orchidaceae). Kew Bulletin 34: 239—319.

STEWART, J., LINDER, H P., SCHELPE, E.A. & HALL, A.V. 1982.

Wild orchids of southern Africa. Macmillan South Africa,

Johannesburg.

STEWART, J. & LA CROIX, I.F. 1987. Notes on the orchids of southern

tropical Africa HI. Aerangis. Kew Bulletin 42: 215—219.

SUMMERHAYES, V.S. 1962. African orchids: XXVIH. Kew Bulletin

16: 253-314.

WILLIAMSON, G. 1979. The orchid flora of the Nyika Plateau. Jour-

nal of South African Botany 45: 459—467.

WOOD, J.J. 1982. A new species of Jumellea (Orchidaceae) from tropi-

cal Africa. Kew Bulletin 37: 77—79.

APPENDIX: A PROVISIONAL CHECKLIST

The species recorded on Mt Mulanje are listed in alphabetical order. The nomenclature follows La Croix et. al. (1991) throughout. Most of the

geographical names are indicated in the map provided (Figure 1). Habitats: e, epilithic or epiphytic; t, terrestrial; gr, grassland; If, lowland and

mid-altitude forest; ma, marsh or bog; mf, montane or submontane forest; pi, pine plantation; ro, on or among rocks; sa, savanna woodland;

se, seepage area. Altitudinal zones: 1, Mulanje base; 2, slopes; 3, plateaux; 4, peaks and ridges. Locality and literature references: 1, La Croix

et al. (1983); 2, La Croix et al. (1991); 3, Morris (1970); 4, Dowsett-Lemaire (1988); 5, Herbarium in Zomba (MAL); 6, Stewart (1980); 7, Chap-

man (1962); 8, Stewart & La Croix (1987); 9, Cribb (1978); 10, Linder (1985); 11, Cribb & Stewart (1985); 12, Linder (1980); 13, Wood (1982);

14, Cribb (1979); 15, Summerhayes (1962); 16, pers. obs. (collecting numbers are given in case of new records only).

Acampe praemorsa (Roxb.) Blatter & McCann , e. If, sa; zone 1, Limbuli

stream and Chitakali Estate, 3; Ruo River, 1, 2; Esperanza Estate,

4; Mimosa Estate and Likhubula Valley, 1

Aerangis

alcicomis (Reichb. f.) Garay, e. If; zones 1 & 2, Phalombe Gorge, 1,

3, 6

appendiculata (De Wild.) Schltr., e. If, sa; zone 2, Crater, 3, 6

distincta Stewart & La Croix, e. If; zones 1 & 2, Phalombe Gorge,

8; Mimosa Estate, 2; zone 3, Chambe Plateau, 2

kotschyana (Reichb. f.) Schltr., e. If, sa; zone 1, Phalombe Plain, 1,

3; zone 2, Little Ruo Basin, 6

mystacidii (Reichb. f.) Schltr., e. If, sa; zone 1, 1, 2, 3, 4, 6; zone 2,

Crater, 4; Likhubula Forest Station, 2, 3, 6; Lichenya River,

2, 3

splendida Stewart & La Croix, e, If; zones 1 & 2, Ruo Gorge, 2, 4,

8; Limbuli stream, 8

verdickii (De Wild.) Schltr., e, sa; zone 1, Limbuli Estate, 2

Angraecopsis

amaniensis Summerh., e. If, sa; zone 2, Likhubula Valley, 3

malawiensis Cribb, e, If, mf; zone 2, Little Ruo Plateau, 1, 2; Crater,

2, 4

parviflora (Thouars) Summerh. ', e. If; zone 1 , 1, 3, 4; zone 2, Crater,

2, 3, 4; Ruo Gorge, 1, 2, 3, 4; Likhubula Valley, 3; Chisongeli

and Lichenya, 4

Angraecum

aff. angustipetalum Rendle, e; locality unknown, 2, 3

chamaeanthus Schltr., e, mf; zones 2 & 3, 2, 3, 4

conchiferum Lindl., e, mf; zone 3, Lichenya and Sombani Plateaux,

2, 3

cultriforme Summerh., e. If; species identity doubtful (La Croix et al.

1991); zone 1, 1, 2, 3

sacciferum Lindl., e, mf; zone 2, Crater, Likhubula Valley, 3; Chisonge-

li, 4; zone 3, 1, 2, 3, 4

stella-africae Cribb, e, sa; zone 2, Ruo Gorge, 2

Ansellia africana Lindl., e. If; zones 1 & 2, Lower Ruo River,

1

Bolusiella

iridifolia (Rolfe) Schltr. subsp. picea Cribb, e, mf; zone 3, 1, 2,

4

maudae (Rolfe) Schltr., e, If, sa; zone 1, 1

Brachycorythis pleistophylla Reichb. f. subsp. pleistophylla, t, gr, sa;

zones 2 & 3, Lichenya Path and Chambe Plateau, 2

Brownleea

maculata Cribb, e/t, mf; zone 2, Ruo Gorge, 2; zone 3, 1, 2, 9

mulanjiensis Linder, t, gr; zone 3, Madzeka Plateau, 2, 10; Chinzama

Plateau, 2; zone 4, Sapitwa peak, 2, 10

parviflora Harv. ex Lindl., t, gr; zones 3 & 4, Lichenya Plateau and

Thuchila Path, 2; Chinzama Plateau, 16

242

Bothalia 22,2 (1992)

Bulbophyllum

elliotii Rolfe, e, sa; zones 1 & 2, 1, 2, 3

encephalodes Summerh., e, If, sa; zone 1, Pwazi River, 3; Lujeri Estate,

2; zone 2, Ruo Gorge, 1, 3

fuscum Lindl. var. melinostachyum (Schltr.) J.J. Vermeulen, e. If, sa;

zones 1 & 2, Likhubula Valley and Fort Lister, 2

gravidum Lindl., e, If, sa; zone 2, Ruo Gorge, 1, 2, 3, 4

humblotii Rolfe, e, If; zone 1, 1, 2, 3

intertextum Lindl., e, If; zone 2, Ruo Gorge, 1, 2, 3, 4; Crater, 3, 4;

Chisongeli, 4

josephii (Kuntze) Summerh., e, If, sa; zone 2, SE slopes, 2; foothills,

3; Ruo Gorge, 1, 3, 4; Chisongeli, 4

longiflorum Thouars, e. If; zone 1, 1, 2, 3; zone 2, Mlanje Boma, 3

maximum (Lindl.) Reichb. /., e, sa; zone 1, 1, 2, 3, 4; zone 2, Ruo

Gorge, 4

oreonastes Reichb. f, e, If, sa; zone 2, 1, 3; probably misidentified

and is in fact B. fuscum Lindl. (La Croix et al. 1991)

rugosibulbum Summerh., e, sa; zone 2, 1, 2

sandersonii (Oliv.) Reichb. f, e, If, mf, sa; zone 2, Ruo Gorge, 3, 4;

Chisongeli, 4; Likhubula River, 2, 3; Limbuli stream and

Sombani Path, 3; zone 3, Chambe Plateau, 3; Chisongeli and

Lichenya Plateau, 4

scaberulum (Rolfe) H. Bot., e, If, sa; zone 2, 1, 2

stolzii Schltr., e, mf; zone 3, 2, 4

unifoliatum De Wild. var. infracarinatum (Williamson) J.J. Vermeulen,

e. If; zones 1 & 2, Ruo Gorge, 1, 2, 3, 4

Calanthe sylvatica (Thouars) Lindl., t, If; zone 2, Lichenya Path, 2

Calyptrochilum christyanum (Reichb. f.) Summerh., e. If, sa; zone 1,

1, 2, 3, 4; zone 2, Likhubula Valley, 2, 3; Ruo Gorge, 2, 3, 4

Corycium dracomontanum Parkman & Schelpe, t, gr; zone 3, Sombani

Plateau, 16 ( Kurzweil 1492, NBG!)

Corymborkis corymbis Thouars, t, If; zone 1, Lichenya Forest, 2, 7;

Ruo Estate, 2; zone 2, Crater, 2

Cynorkis

anacamptoides Kraenzl., t, gr, ma, se; zone 2, Lichenya Path and

Likulezi Path, 2; zone 3, Chinzama Plateau, 2; Chambe Plateau

and Ruo Gorge, 3

brevicalcar Cribb, t, se; zones 3 & 4, 2, 11, 16

buchananii Rolfe, t, se, pi; zone 3, Lichenya and Madzeka Plateaux,

2; Chambe Plateau, 5

hanningtonii Rolfe, t, If, mf, pi, sa; zone 2, Boma Path, 2, 3; Likhubula

Valley and Limbuli stream, 3; Sombani Path, 16

kassneriana Kraenzl., t, mf, pi; zones 2 & 3, Lichenya Path, 2, 16;

Chambe Plateau, 2, 3; Sombani Path, 16

kirkii Rolfe, t. If, gr, se; zone 2, Likhubula Valley, 3; Sombani Path,

16; Lichenya Path, 16, 5; zone 3, Thuchila Plateau, 16; Chambe

Plateau, 5

Cyrtorchis

arcuata (Lindl.) Schltr.

subsp. variabilis Summerh., e, If, sa; zone 1, 1, 2, 3

subsp. whytei (Rolfe) Summerh., e. If, sa; zones 1 & 2, Likhubula

Valley, 2, 3; Crater, Pwere stream, Limbuli stream, Pwazi River

and Chapaluka stream, 3

crassifolia Schltr., e, sa; zone 2, 1, 2, 3

praetermissa Summerh., e, sa; zones 1 & 2, 1, 2, 3, 4

ringens (Reichb. f.) Summerh., e. If, mf, sa; zone 1, Lujeri Estate, 3;

zone 2, Ruo Gorge, 2, 3, 4; Likhubula River, Crater and

Sombani Path, 3; Chisongeli, 4

Diaphananthe

fragrantissima (Reichb. f.) Schltr., e, If, sa; zone 1, 1, 2, 3, 4; zone

2, Crater, 4

rutila (Reichb. f.) Summerh., e, If; zone 1, Lower Ruo River, 1, 2;

Lichenya forest, 2; Chisambo, 4

stolzii Schltr., e, If; zone 2, Crater, Limbuli stream and Lujeri Estate,

3; Ruo Gorge, 1, 2, 3, 4; Chisongeli, 2, 4

subsimplex Summerh., e, mf; zone 3, Lichenya Plateau, 1, 2, 4

xanthopollinia (Reichb. f.) Summerh., e. If, mf, sa; zone 1, Lower Ruo

River, 1, 2

Didymoplexis africana Summerh., t, If; zone 1, Lujeri Estate, 2

Disa

aconitoides Sond. subsp. concinna (N.E.Br.) Linder, t, gr, ma, sa; zone

3, Chambe Plateau, 2, 5; Thuchila Plateau, 16

erubescens Rendle subsp. erubescens, t, gr, ma; zone 3, 2, 16

fragrans Schltr., t, se; zone 3, Lichenya Plateau, 2

ochrostachya Reichb. /., t, gr, ma; zone 4, Sapitwa peak, 2

perplexa Linder, t, ma; zone 3, Lichenya, Chinzama, Sombani and

Madzeka Plateaux, 2, 16

saxicola Schltr. e/t, ro, se; zone 2, Sombani Path, 16; zone 3, Lichenya

and Madzeka Plateaux, 2; zone 4, Sapitwa and Chilemba peaks,

5

Disperis

anthoceros Reichb. /., t, mf, sa; zone 2, Limbuli stream, 3

dicerochila Summerh., t, mf, pi; zone 3, Chambe, Thuchila and Madzeka

Plateaux, 2

leuconeura Schltr., t. If; zones 1 & 2, Likhubula Valley, Esperanza

Estate, 2

Eggelingia clavata Summerh., e, If; zone 2, Ruo Gorge, 1, 2, 4

Eulophia

callichroma Reichb. f, t, sa; zone 2, Likhubula Valley, 2

clavicomis Lindl. var. nutans (Sond.) A.V. Hall, t, gr, ma; zones 2 &

3, 2

cucullata (Afzel. ex Swartz) Steud., t, gr, ma, sa; zones 1—3, 2

eylesii Summerh., t, sa; zone 2, Phalombe Gorge, Likulezi Path and

Likhubula Valley, 2

fridericii (Reichb. f.) Hall, t, sa; zone 2, Likhubula Valley, 2

horsfellii (Bateman) Summerh., t, If; zone 1, Lujeri Estate, 2

livingstoniana (Reichb. f.) Summerh., t, gr, ma, sa; zones 1 & 2, 2

longisepala Rendle, t, sa; zone 2, Likhubula Valley, 2

macrantha Rolfe, t, under bamboo; zone 1, Limbuli and Ruo Estates, 2

monticola Rolfe, t, gr; zone 3, Lichenya Plateau, 2; Chambe Plateau, 5

norlindhii Summerh., t, sa; zones 1 & 2, Lichenya Path and Phalombe

Gorge, 2

nyasae Rendle, t, sa; zone 2, 2

odontoglossa Reichb. /., t, gr, sa; zone 2, Lichenya, Sombani and

Chapaluka Paths, 2

rara Schltr., t, gr; zone 2, Chambe, Chapaluka and Lichenya Paths, 2

rolfeana Kraenzl., t, sa; zone 2, Likhubula Valley, 2

seleensis (De Wild.) Butzin, t, sa; zone 2, Likhubula Valley, 2; Sombani

Path, 5

streptopetala R. Br. ex Lindl., t, If, mf, sa; zones 2 & 3, 2, 16

subsaprophytica Schltr., t, sa; zone 2, Likhubula Valley, 2

venulosa Reichb. /., t, sa; zones 1 & 2, Likhubula Valley, 2

zeyheri Hook. /., t, gr; zone 2, Lichenya Path, 2

Habenaria *.

amoena Summerh., t, sa; zone 2, Likhubula Valley, 2

comuta Lindl., t, gr, sa; zones 2 & 3, Sambani Path, 2

disparilis Summerh., t, sa; zone 2, 2

humilior Reichb. /., t, gr; zone 3, Thuchila Plateau, 2

macrandra Lindl., t, If; zone 2, Ruo Gorge, Crater, 2

macrostele Summerh., t, gr, se; zone 2, Lichenya and Sombani Paths,

2; zone 3, Thuchila and Chambe Plateaux, 5; Chinzama,

Lichenya, Sombani and Madzeka Plateaux, 16

malacophylla Reichb. /., t, If; zone 2, Ruo Gorge, 5

myodes Summerh., t, sa; zone 2, 2, 15

nyikana Reichb. f. subsp. nyikana, t, sa; zone 2, Likhubula Valley, 2;

zone 3, Madzeka Plateau, 5

petitiana (A. Rich.) Dur. & Schinz, t. If, mf; zones 2 & 3, Lichenya

Path, 2; zone 3, Thuchila Plateau, 5

praestans Rendle, t, gr, sa; zones 2 & 3, Lichenya Path, 2; Sombani

Path, 16

tentaculigera Reichb. /., t, sa; zone 2, Sombani Path, 16 (Kurzweil 1413,

NBG!)

trilobulata Schltr., t. If, sa; zone 1, Likhubula Valley, 2

welwitschii Reichb. f, t, gr; zone 3, 2

Herschelianthe baurii (H. Bol.) Rauschert, t, gr; zone 3, 2; zone 4, 7

Holothrix

johnstonii Rolfe, t, se; zone 2, Likhubula Valley and Lichenya Path,

5; zone 3, 2, 3; zone 4, Path to Sapitwa, 3

longiflora Rolfe, t, gr, sa, se; zone 2, Lichenya Path, 2; Sombani Path,

16; zone 3, 2, 3, 7

orthoceras (De Wild.) Reichb. /., t, If, mf; zone 2, Lichenya Path, 2;

Likhubula Valley, 2, 3; zone 3, Chambe Plateau, 3

Jumellea

filicornoides (De Wild.) Schltr., e. If, sa; zone 2, Crater, 2, 3;

Chisongeli, 2; Likhubula Valley, 3; zone 3, 1

usambarensis Wood, e, mf; zone 2, Chisongeli, 4; slopes below

Lichenya, 13; zone 3, 1, 2, 3, 4

Liparis

bowkeri Harv., t/e. If, mf, sa; zone 1, 1; zone 2, Ruo Gorge, 1, 2, 3;

Lichenya Path and Likhubula Valley, 2; Sombani Path and slope

above Mulanje Village, 16; zone 3, 1, 2, 3

caespitosa (Thouars) Lindl., e, If, mf; zone 2, 1, 2

nervosa (Thunb.) Lindl., t, gr, sa; zones 1 & 2, Lichenya Path, 2

Malaxis weberbaueriana (Kraenzl.) Summerh., t. If, mf; zone 2, Ruo

Gorge, 2

Microcoelia

exilis Lindl., e, If; zones 1 & 2, 1, 2

moreauae L. Jonsson, e, If; zone 1, 1, 2

Monadenia brevicornis Lindl., t, gr; zone 3, Lichenya and Madzeka

Plateaux, 2; Chinzama and Sombani Plateaux, 16; Thuchila

Bothalia 22,2 (1992)

243

Plateau, 5; zone 4, Thuchila-Sombani divide, 16; Chilemba and

Sapitwa peaks, 5

Mystacidium tanganyikense Summerh., e, mf; zone 2, Crater, 3;

Chisongeli, 4; zone 3, 1, 2, 3, 4

Neobolusia stolzii Schltr., t, gr, se; zone 3, Lichenya Path, 2; Chambe,

Sombani and Chinzama Plateaux, 16

Nervilia

adolphii Schltr . , t, sa; zone 2, Lichenya Path, 2

pectinata Cribb, t, If, mf; zones 2 & 3, Ruo gorge, 2

kotschyi (Reichb. f.) Schltr., t, sa; zone 2, Likhubula Valley, 2

shirensis (Rolfe) Schltr., t, sa; zone 2, Likhubula Valley, 2

Oeceoclades maculata (Lindl.) Lindl., t. If; zones 1 & 2, Likhubula Valley,

2

Platycoryne

mediocris Summerh., t, gr, se; zones 1 & 2, Likhubula Valley, 2

pervillei Reichb. /., t, gr; zone 1, Lujeri Estate, 2; Limbuli Estate, 5;

zone 2, Thuchila and Lichenya Paths, 5

Platylepis glandulosa (Lindl.) Reichb. f, t. If; zone 1, Lujeri Estate,

2

Polystachya

adansoniae Reichb. /., e, sa; zone 2, 1, 2

albescens Ridley subsp. imbricata (Rolfe) Summerh., e, If, sa; zone 1,

1, 3; zone 2, Crater, 2, 4; Ruo Gorge, 1, 2, 3, 4; Chisongeli,

4

brassii Summerh., e, sa; zones 2 & 3, 1, 2, 3

campyloglossa Rolfe, e, mf; zone 2, Ruo Gorge, 1; Lichenya Path,

2

cultriformis (Thouars) Spreng., e. If, mf; zone 2, Ruo Gorge, 1, 2, 4

fusiformis (Thouars ) Lindl. , e. If, mf, sa; zone 1, 1; zone 2, Ruo Goige,

2, 3, 4; Likhubula Valley, 3; Chisongeli, 4

greatrexii Summerh., e, If, sa; zone 1, Limbuli Estate, 2; zone 2, SE

slopes, 1, 2; Crater, 4

heckmanniana Kraenzl., e, mf; zone 2, SE slopes, 2; Ruo Gorge, 1,

2, 4; Chisongeli, 4; zone 3, 2; Lichenya Plateau, 4

johnstonii Rolfe, e on Xerophyta sp. ; zone 3, 1, 2, 3

lawrenceana Kraenzl., e, ro, sa; zone 2, Thuchila Path, 2

minima Rendle, e, sa; zone 2, SE slopes, 2

modesta Reichb. f, e, sa; zones 1 & 2, 1, 2

purpureobracteata Cribb & I. la Croix, e, If, mf; zone 2, Chisongeli

and Ruo Gorge, 2

simplex Rendle, e, If, sa; zone 2, Ruo Gorge, 1, 2, 4

songaniensis Williamson, e/t, gr, se; zone 2, Boma Path, 2; zone 3,

Lichenya Plateau, 2

stuhlmannii Kraenzl ., e. If, sa; zone 2, Crater, 2

tessellata Lind/. , e. If, ro, sa; zone 1, 1; Limbuli stream, Pwazi River,

Chitakali and Mimosa Estates, Pwere stream, 3; Esperanza

Estate and Chisambo, 4; zone 2, Likhubula Valley, 1, 2, 3; Crater

and Ruo Gorge, 1, 2, 3, 4; Sombani Path, 2

transvaalensis Schltr., e, mf, ro; zone 3, 1, 2, 3, 4

villosa Rolfe, e. If, mf, sa; zones 1 & 2, 1, 2; zohe 2, Likhubula Val-

ley, 3; zone 3, Chambe Plateau, 3

zambesiaca Rolfe, e, If, ro, sa; zone 2, Ruo Gorge and SE slopes, 2;

Chisongeli, 4; Boma Path, 3; zone 3, 1, 2, 3, 4

Rangaeris muscicola (Reichb. f) Summerh., e. If, sa; zone 2, Crater,

2, 3, 4; Ruo Gorge, 1, 2, 3, 4; Likhubula Valley, 3; Chisongeli, 4

Roeperocharis wentzeliana Kraenzl., t, gr; zone 3, Chinzama, Madzeka

and Lichenya Plateaux, 2; Sombani Plateau, 5

Satyrium

aberrans Summerh., t, gr; zone 3, Madzeka Plateau, 2

amblyosaccos Schltr., t, gr; zone 3, Lichenya Plateau, 2

anomalum Schltr., t, sa; zone 2, Lichenya Path, 2; Sombani Path,

16

breve Rolfe, t, ma, se; zone 3, Lichenya Plateau, 2

chlorocorys Rolfe, t, gr; zone 3, Lichenya Plateau, 2; Chinzama,

Madzeka and Sombani Plateaux, 16

ecalcaratum Schltr., t, ro; zone 4, 2

macrophyllum Lindl., t, gr, sa, se; zone 3, Sombani, Thuchila and

Chambe Plateaux, 2

neglectum Schltr., t, gr, ma; zone 2, Lichenya Path, 2; zone 3, Thuchila

Plateau, 2; Sombani Plateau, 16

oliganthum Schltr., t, gr; zone 3, Lichenya, Chambe and Sombani

Plateaux, 2

rhynchantoides Schltr., t, gr, se; zone 3, 3, 16

shirense Rolfe, t, gr, se; zone 3, Lichenya, Thuchila, Sombani and

Madzeka Plateaux, 2

trinerve Lindl., t, gr, ma; zone 3, Lichenya Plateau, 2; Thuchila, Madze-

ka and Sombani Plateaux, 16

Schizochilus sulphureus Schltr., t, gr, se; zones 3 & 4, 2, 12, 16

Schwartzkopffia lastii (Rolfe) Schltr., t, sa; zone 2, Likhubula Valley,

2, 16; Lichenya Path, 2

Stenoglottis fimbriata Lindl., e/t, If, mf, sa; zone 2, Crater, 3, 4;

Chisambo, Ruo Gorge and Chisongeli, 4; Likhubula Valley, 5;

Sombani Path, 16; zone 3, Lichenya Plateau, 4; Thuchila Plateau,

5

Stolzia

compacta Cribb subsp. purpurata Cribb, e, If, mf; zone 2, Ruo Gorge,

2; Chisongeli, 4; zone 3, 1, 2, 4, 14

repens (Rolfe) Summerh., e. If, mf, sa; zone 3, 1, 2, 3, 4

Tridactyle

anthomaniaca (Reichb. f.) Summerh., e, sa; zone 1, 1, 2

bicaudata (Lindl.) Schltr., e, If, sa; zone 1, 1, 2, 3, 4; zone 2, Crater

and Ruo Gorge, 3, 4; Chisongeli, 4; Limbuli stream and Chambe

Path, 3

inaequilonga (De Wild.) Schltr., e, If, mf, sa; zone 2, Ruo Gorge, 2,

3; Chisongeli, 4; zone 3, 1, 2, 3, 4

tricuspis (H. Bol.) Schltr., e, mf, sa; zone 3, 2, 3

tridactylites (Rolfe) Schltr., e, If; zone 1, 1, 2, 3, 4; zone 2, Ruo Gorge,

2, 3, 4; Likhubula Valley, 2, 3; Chisongeli, 4

tridentata (Hare.) Schltr., e, sa; zone 1, 1; zone 2, Likhubula Valley,

2, 3; Limbuli Estate and Sombani Path, 3

verrucosa Cribb, e, ro; zone 2, Chisongeli, 2, 4

Ypsilopus erectus (Cribb) Cribb & Stewart, e, mf, sa; zone 2, Lichenya

Path, 2; zone 3, 1, 2, 3, 4

The following species apparently do not occur:

Aerangis brachycarpa (A. Rich.) Dur. & Schinz; zone 2, Ruo Gorge,

1; zone 3, Chambe Plateau, 1. The specimens are considered to

be either A. distincta or A. splendida (Stewart & La Croix 1987;

La Croix et al. 1991).

Bulbophyllum oxypterum (Lindl.) Reichb. /.; 3. The plant belongs to

B. sandersonii (La Croix et al. 1983: 53).

'

-

■

.

Bothalia 22,2: 245-254 (1992)

Die fitososiologie van die Bankenveld in die Grootvlei-omgewing,

Suid-Transvaal

W.J. MYBURGH* P.J.J. BREYTENBACH* G.K. THERON** en GJ. BREDENKAMP**

Sleutelwoorde: Bankenveld, Braun-Blanquet, Grootvlei-area, fitososiologie

UITTREKSEL

Die plantegroei van die Bankenveld in die Grootvlei-omgewing is fitososiologies ondersoek. ’n Totaal van 50 re 1 eves is

numeries (TWINSPAN) geklassifiseer deur van Braun-Blanquet-prosedures gebruik te maak. Daar is ses plantegroei-eenhede,

insluitend twee variante, onderskei. Die resultate van ’n DECORANA-ordening stem grootliks ooreen met die plantegroei-

eenhede, met geassosieerde omgewingsgradiente van die Braun-Blanquet-opname.

ABSTRACT

The phytosociology of the Bankenveld in the Grootvlei area is presented. The results of a numerical classification (TWIN-

SPAN) of 50 relevds were defined by Braun-Blanquet procedures. The classification revealed six vegetation units, with two

variations. The results of a DECORANA ordination corroborate largely the vegetation units with associated environmental

gradients of the Braun-Blanquet survey.

INLEIDING

Die noordoostelike gedeelte van die Hoeveldstreek,

meer spesifiek die Heidelberg-substreek, beslaan ’n totale

oppervlakte van 2 433 000 ha waarvan 1 237 600 ha onder

bewerking is (Landbou-ontwikkelingsprogram 1986). Die

oorblywende natuurlike weiveld is beperk tot die rante en

koppies met vlak litosols en die laagliggende gebiede,

dreineringsbane en vleie met periodies waterversadigde

vertiese kleie.

Die Bankenveld (Veldtipe 61: Acocks 1988) word beskou

as ’n skyngrasveld en vorm deel van die Grasveldbioom.

Die plantegroei van die Grasveldbioom is fisionomies

monolities en word gekenmerk deur ’n sterk dominansie

van hemikriptofiete van die Poaceae. Die kruinbedekking

is vogafhanklik en toon ’n afname in gemiddelde jaarlikse

reenval (Rutherford & Westfall 1986). Die Bankenveld is

’n gespesialiseerde nis waar borne, struike, kruide en

grasse assosieer (Rutherford & Westfall 1986).

In die ‘Witskrif oor Landboubeleid’ word verwys na

die kommerwekkende agteruitgang van natuurlike weiveld

(Nasionale Weidingstrategie 1985). Dit is geen uit-

sondering in die Hoeveldstreek nie. Daar is ’n merkbare

verswakking in veldtoestand van oos na wes vanwee

oorbeweiding deur skape en beeste.

Die bree plantegroei-indeling van Acocks (1988) is

onvoldoende by streek- en substreekbeplanning (Deall et

al. 1989). Die doel van die studie is om die floristiese

samestelling, struktuur en omgewingsinteraksies van die

Bankenveld in die Grootvlei-distrik te bestudeer, die

plantegroei en omgewingsverhoudings te verstaan en te

organiseer en om die plantegroei in homogene eenhede

of plantgemeenskappe te onderskei. Dit is nodig om

* Huidige adres: Roodeplaat Weidingsinstituut, Privaatsak X05, Lynn

East 0039.

** Departement Plantkunde, Universiteit van Pretoria, Pretoria 0002.

MS. ontvang: 1991-09-06.

plantegroei-eenhede te identifiseer en klassifiseer sodat

die natuurlike weidings optimaal ontwikkel, bestuur en

benut kan word sonder ’n verdere verswakking van weiveld

(Nasionale Weidingstrategie 1985).

Die studiegebied dien as ’n skakel tussen die oostelike

en westelike dele van die Grasveld-bioom (Breytenbach

et al. in druk). Die data wat tydens die studie ingewin is,

sal gebruik word om die ekstrapoleringsmoontlikhede na

die omliggende gebiede te ondersoek.

DIE STUDIEGEBIED

Die studiegebied is gelee in die omgewing van Groot-

vlei, suidelike Transvaal (Figuur 1) en vorm deel van die

grasveldbioom. Die studiegebied is die Bankenveld gelee

tussen 26°45' en 27°00' suiderbreedte en 28°30' en 28°45'

oosterlengte en varieer in hoogte bo seespieel vanaf 1 500

m tot 1 827 m by Korporaalskop in die noorde. Die

Bankenveld word omring deur die Cymbopogon—Themeda-

veld (Veldtipe 48), Themeda-\e\d (Veldtipe 52), Gemengde

Themeda-ve\d na Cymbopogon-Themeda-ve\d oorgang

(Veldtipe 53) en Bankenveld na Ihemeda-ve\d oorgang

(Veldtipe 55) (Acocks 1988). ’n Volledige beskrywing van

die fisiese omgewing van die gebied word in Breytenbach

et al. (in voorb.) weergegee.

Volgens die Bree Terreinpatrone-kaart van Suidelike

Afrika (Kruger 1983) kom daar twee terreinmorfologiese

klasse, naamlik 3 en 18, in die studiegebied voor. Klas 3

word beskryf as effens golwende vlaktes met ’n lae relief

van 0—130 m. Klas 18 word beskryf as heuwels en

laaglande met ’n hoe relief van 130—450 m (Kruger 1983)

en sluit grootliks die Bankenveld (Acocks 1988) in.

In teenstelling met klas 3 waar 80% van die oppervlak

hellings met minder as 5° het, besit slegs 20—50% van

die oppervlak van klas 18 hellings van minder as 5°.

Studies het getoon dat daar ’n noue assosiasie bestaan

tussen die geologie en plantegroei van die Bankenveld

246

Bothalia 22,2 (1992)

(Bredenkamp 1975; Bredenkamp & Theron 1978, 1980).

Die studiegebied word gekenmerk deur die Opeen-

volging Karoo, die Supergroep Witwatersrand en die

Supergroep Ventersdorp [South African Committee for

Stratigraphy (SACS) 1980].

Die studiegebied word verdeel in die Ba-, Bb-, Ea- en

Ib-landtipes (Van der Bank et al. 1978). Gronde van die

landtipes is klipperig en word oor die algemeen nie geploeg

nie. Hierdie landtipes word meestal as natuurlike weid-

ing vir beeste gebruik.

Die geologie van die Ea-landtipe bestaan oorwegend uit

die Opeenvolging Karoo met sandsteen, skalie en doleriet-

dagsome (Landtipe opname personeel 1984). Die hoerlig-

gende gronde (Terreineenhede 1 & 3, Figuur 2) sluit die

Mayo-, Milkwood-, Shortlands-, Swartland-en Avalongrond-

vorms (Macvicar et al. 1977) in. Die laerliggende gebiede

(Terreineenhede 4 & 5, Figuur 2) word gekenmerk deur die

margalitiese waterversadigde gronde van die Willowbrook-,

Sterkspruit-, Estcourt-, Rensburg- en Arcadiavorms.

Landtipes Ba en Bb word gedefmieer as die plintiese

katena en sluit onder andere die Hutton-, Avalon-,

Longlands-, Shortlands- en Mispahgrondvorms (Terrein-

eenhede 1 & 3, Figuur 2) in. Gronde van die voethellings

en valleivloere (Terreineenhede 4 & 5, Figuur 2) sluit die

Bainsvlei-, Estcourt-, Willowbrook-, Westleigh-, Bonheim-

en Rensburgvorms in (Landtipe opname personeel 1984).

Supergroep Ventersdorp met ondesitiese tot diasitiese

lawas, tuf, chert, agglomeraat en kwartsiet is kenmerkend

van die Ba-landtipe.

Die Bb-landtipe bestaan oorwegend uit sandsteen,

skalies en grintsteen van die Opeenvolging Karoo. Die

Ib-landtipe beslaan die grootste gedeelte van die Banken-

veld in die studiegebied en word aangetref waar bloot-

gestelde rots, afkomstig van die Ventersdorpsisteem,

60—80% van die oppervlak beslaan (Landtipe opname

personeel 1984). Die vlak, klipperige gronde van die hoer-

liggende gebiede (Terreineenhede 1, 2 & 3, Figuur 2) is

gewoonlik van die Shortlands-, Glenrosa- en Mispahgrond-

vorms (Landtipe opname personeel 1984).

Die laaglande (Terreineenhede 4 & 5, Figuur 2) word

gewoonlik deur die Bonheim-, Valsrivier-, Arcadia- en

Rensburggrondvorms gedomineer (Landtipe opname per-

soneel 1984).

METODES

Die floristiese samestelling wat by elke monsterperseel

waargeneem is, is op die Ziirich-Montpellier

fitososiologiese benadering (Braun-Blanquet 1932; Werger

1974) gebaseer. Die verspreiding van monsterpersele is

gebaseer op landtipes en geologie.

Die aantal monsterpersele wat in elke homogene eenheid

uitgeplaas is, is proporsioneel op grond van eenheidgrootte

Bothalia 22,2 (1992)

247

bepaal. Daar is ’n totaal van 50 releves elk met ’n opper-

vlakte van 200 m2 uitgeplaas.

In elke monsterperseel is al die identifiseerbare spesies

aangeteken en versamel. Daar is ook ’n bedekkingswaarde

toegeken volgens die Braun-Blanquet bedekkingsgetal-

sterkteskaal (Werger 1974). Name van die taksons stem

ooreen met die van Gibbs Russell et al. (1985, 1987) en

is saamgevat in ’n floristiese analise en spesielys (Myburgh

et al. in voorb.).

Habitatsdata wat versamel is, sluit in hoogte bo see-

spieel, geologie, klipbedekking, topografiese posisie,

helling en aspek. Die grondklassifikasie is verkry van die

1:50000 2628 DC Grootvlei-bodemkaart (Van der Bank

et al. 1978).

Die politetiese verdelingstegniek, TWINSPAN (Hill

1979a & b), is gebruik om die data in tabelvorm op te

stel. Braun-Blanquet-prosedures (Braun-Blanquet 1932)

is gebruik om die klassifikasie verder te verfyn. Die

resultate word weergegee in die fitososiologiese tabel

(Tabel 1).

RESULTATE

Klassifikasie

grasveld met koppies, riwwe en dagsome waar struike en

borne enkel of in bosgroepe voorkom. Die grasse Eragros-

tis curvula, Themeda triandra, Heteropogon contortus en

Elionurus muticus is dominant. Dominante kruide sluit

in Commelina africana en Crabbea acaulis (Spesiegroep

M, Tabel 1). Die plantegroei varieer met verandering in

habitat. Geologie, klipbedekking, hoogte bo seespieel en

aspek dra by tot die herkenning van plantgemeenskappe.

Die aantal spesies aangeteken in elke releve wissel van 21

tot 49, met ’n benaderde gemiddeld van 33 spesies per

releve.

Daar word twee hoofhabitattipes aangetref, naamlik die

hoogliggende klipperige gebiede en die laagliggende nie-

klipperige gebiede. Die hoogliggende klipperige gebiede

word gekenmerk deur struikveld, terwyl die laagliggende

nie-klipperige gebiede oorwegend uit grasveld bestaan.

Twee struikgemeenskappe en twee grasveldgemeenskappe

kan onderskei word; die eerste struikgemeenskap het twee

variante.

1. Diospyros lycioides—Euclea cmpa-struikveld.

1.1 Euclea crispa—Aloe davy ana-variant.

1.2 Euclea crispa—Rhus pyroides-v ariant.

2. Diospyros lycioides—Diheteropogon amplectens-

struikveld.

Daar kan algemeen na die Bankenveld in die gebied 3' Themeda "dandra-Cymbopogon plurinodis-gTWdd.

verwys word as ’n Themeda triandra— Eragrostis curvula- 4. Heteropogon contortus— Loudetia simplex- grasveld.

FIGUUR 2. — ' Terreinvormsketse van die Ea-, Ba-, Bb- en Ib-landtipes aangetref in die studiegebied (Scheepers, Smit & Ludick 1984). Terreineen-

heid: 1, kruin; 2, vryhang; 3, middelhang; 4, voethang; 5, valleivloer.

248

Bothalia 22,2 (1992)

TABEL 1. — ’n Plantsosiologiese tabel van die Bankenveld in die Grootvlei-omgewing

Gemeenskap- 1 234

nommer 1 . 1

1 .2

BBCCCBBABBBBAABCCCBC BBCAAAAABA AAAAAAAAAA ABA

12222222222223223322 2322221232 1111131132 231

NONWNNNSSSSSWSSSSWSS SSSWOOSSSS NWNNNNNNNN WSS

W W WWW WOW WO W WWWW WW

ABDEABCACCAABCCCCACE OBAAAAAAAA AAAAAAAAAA AAA

43333311333334333333 3313333133 3333331341 333

RRVRRWRRVRRRRRRRRRR RRVJJJRJRP JJJPPRRPPP RRR

KKDKKDDTKDKKKKKKKKKK KKDDDDTDTV DDDWGGVW TTT

BCDCBCEDCCBBCBCDDOCO BBCBCBACAB AAAAABAAAA AAC

ABCDADDDDDADDDCADAAC CDODDDODDD OOODDODDOD ODD

33033113011131311130 0113311113 1111113211 133

23333344333434333333 3354432444 2222245225 422

23224343434443433433 3332232343 2222222323 322

52682147363253673965 6339972834 6158474088 169

Re I eves

0000000 00000000000000000000 0000000000 0000000000 000

2222345 31333231113343444444 4212221234 0000001000 111

6789080 93568041582397356401 2391244517 3245670189 267

SPES I EGROEP A

Di ospyros /ycioides

Cymbopogon excavatus

Rhus rigida

Pellaea calomelanos

Hyparrheni a hirta

Chei I anthes hirta

Indigot'era obscura

Maytenus heterophyl / a

Rhyne he lytrum re pens

Argyro / ob i um ve / ut i num

Aloe transvaa I ens i s

/ pomoea crass i pes

P / ectranthus madagascar i ens i s

Chaetacanthus costatus

12122

+ 11 +

+ Rim

i i i++++

+ +i i

++ ++

R

1 1222+

1 11212

+R

222++1+

++ 1

1

+ + R

222122221132231112+

+ 2 1 1++1212+ 12

++R 111+ RR2-1 + ++

R+R+ ++++ ++R1+ +2

11 +22++22+2 1

+1 ++++1+++++1+ +2

+R + RR++ + R+ R+

R+ 12 R+1+1 1R 1 4

+++ + 1+ + + + 1

++ R ++++++ ++

++113 R 1 +

1++ R 1 +11+1

1+ ++1 ++ +1 +

+R ++ ++

+1 +11112

1+ + 1 2++

R++RR+ 1 +

RR +

2 12+

R+ + +

R+ +++ +R

+ 1 +

++ ++ +

++ +++ +

1 +R

+ +R R

+ 1 +

+ + +

+ 1-

1

+++

SPES I EGROEP C

Aloe davyana

I pomoea obscura

Ka / anchoe thyrs i f / ora

I 3 32++++ I

| 1 1 ++++ 1 ■

I R+ +R I

Bothalia 22,2 (1992)

249

TABEL 1. — ’n Plantsosiologiese tabel van die Bankenveld in die Grootvlei-omgewing

Gemeenskap-

nommer

Hoogte bo seespieel

Geomorfo I ogie

Aspek

He I I i ng

Terre i nvorm

Geo I og i e

K I i pbedekk i ng

Eros i e

Bewe i d i ng

B lootste I I i ng

Aantal spes i es/re I eve

1 2 3 4

1.1 1.2

BBBBBBB BBCCC8BABBBBAABCCCBC

2332222 12222222222223223322

NNONNNN NONWNNNSSSSSWSSSSWSS

00 WO 0 W W WWW W 0

DOCBOCC ABDEABCACCAABCCCCACE

3333333 43333311333334333333

JJJRRRR RRVRRWRRVRRRRRRRRRR

DDDKKKK KKDKKDDTKDKKKKKKKKKK

DOCODDC BCDCBCEDCCBBCBCDDDCO

AAAAAAA ABCDADDDDDADDDCADAAC

0003313 33033113011131311130

3333333 23333344333434333333

3343332 23224343434443433433

3003296 52682147363253673965

BBCAAAAABA AAAAAAAAAA ABA

2322221232 1111131132 231

SSSWOOSSSS NWNNNNNNNN WSS

W WO W WWWW WW

OBAAAAAAAA AAAAAAAAAA AAA

3313333133 3333331341 333

RRVJJJRJRP JJJPPRRPPP RRR

KKDDDDTDTV DDDWGGVW TTT

BBCBCBACAB AAAAABAAAA AAC

CDDDDDDODD 0DD0D0DDD0 ODD

0113311113 1111113211 133

3354432444 2222245225 422

3332232343 2222222323 322

6339972834 6158474088 169

Re I eves

0000000 00000000000000000000 0000000000 0000000000 000

2222345 31333231113343444444 4212221234 0000001000 111

6789080 93568041582397356401 2391244517 3245670189 267

SPES I EGROEP E

250

Bothalia 22,2 (1992)

TABEL 1. — ’n Plantsosiologiese tabel van die Bankenveld in die Grootvlei-omgewing

Gemeenskap- 1 2 3 a

SPESIEGROEP N

Bothalia 22,2 (1992)

251

TABEL 1. — ’n Plantsosiologiese tabel van die Bankenveld in die Grootvlei-omgewing

SPES I EGROEP N

252

Bothalia 22,2 (1992)

1. Diospyros lycioides—Euclea crispa - stru ikveld

Die gemeenskap word aangetref teen steil hange met ’n

helling van ongeveer 15° en die grondoppervlak word met

46% klip bedek. Die basaltiese lawas van die Supergroep

Ventersdorp vorm dagsome met groot rotsblokke. Die

plantgemeenskap is gelee tussen 1 580 en 1 800 m bo

seespieel en toon ’n wye toleransie ten opsigte van glooiing

met ’n voorkoms wat strek van noord na suid.

Die borne en struike Canthium gilfillanii, Celtis afri-

cana, Cussonia paniculata, Ehretia rigida, Euclea crispa,

Rhoicissus tridentata en kruide Lippia rehmannii en

Teucrium trifidum (Spesiegroep B, Tabel 1) is diagnostics

vir die gemeenskap.

Die boomstratum het ’n gemiddelde hoogte van 3 m en

’n gemiddelde kroonbedekking van 8 % . Die opvallendste

boomspesies is Maytenus heterophylla, Canthium gil-

fillanii, Celtis africana en Cussonia paniculata. Die

struikstratum toon ’n gemiddelde kroonbedekking van 18%

by ’n hoogteklas van 2 tot 3 m. Die struikstratum word

gekenmerk deur die teenwoordigheid van Diospyros

lycioides, Rhus rigida, Rhus pyroides en Diospyros

whyteana. Die xerofitiese varings Pellaea calomelanos en

Cheilanthes hirta asook grasspesies soos Eragrostis

curvula, Heteropogon contortus en Themeda triandra is

kenmerkend van die kruidstratum. Die plantgemeenskap

besit gemiddeld 36 spesies per releve.

1.1. Euclea crispa— Aloe davyana-w ariant

Die variant, wat tussen 1 600 tot 1 700 m bo seespieel

gelee is, word teen steil (11° — 20°), oorwegend noordelike

tot noordoostelike glooiings aangetref. Die klipbedekking

(46—60%) is oorwegend basaltiese lawas van die Venters-

dorpsisteem.

Diagnostiese spesies vir die variant is onder andere die

sukkulent Aloe davyana, Ipomoea obscura, Kalanchoe

thyrsiflora, Pavonia transvaalensis en Hypoxis argentea

(Spesiegroep C, Tabel 1). Die variant besit ’n gemiddeld

van 33 spesies per releve.

1.2. Euclea crispa— Rhus pyroides-v ariant

Die variant het ’n wyer toleransie ten opsigte van aspek

en hoogte bo seespieel as die vorige en word op 1 580 tot

1 800 m bo seespieel teen noordelike tot suidelike glooi-

ings aangetref. Die klipbedekking (46— 60%) is oorwegend

basaltiese lawas van die Ventersdorpsisteem. Rhus

pyroides, Diospyros whyteana, Rhus discolor, Myrsine

africana, Felicia filifolia en Cheilanthes eckloniana

(Spesiegroep D, Tabel 1) is diagnostics vir die variant wat

’n gemiddeld van 37 spesies per releve het.

2. Diospyros lycioides— Diheteropogon amplectens-

struikveld

Die plantgemeenskap word hoogliggend op die suid-,

wes- en oosfrontglooiings van bulte, maar op ’n meer

gelyke (0°— 10°) tot effens golwende topografie as die

vorige plantgemeenskap aangetref. Die bogrondse klipbe-

dekking van 30% is afkomstig van die Ventersdorpsisteem.

Diagnostiese spesies van die gemeenskap is onder

andere die grasse Diheteropogon amplectens, Digitaria

tricholaenoides en die kruide Berkheya onopordifolia en

Gomphrena celosioides (Spesiegroep E, Tabel 1). Daar

is gemiddeld 33 spesies per relev6.

Die boomstratum ontbreek, maar daar is wel enkele

individue van Maytenus heterophylla buite die relev^s

aangeteken. Die opvallende struike Diospyros lycioides en

Rhus rigida dra by tot die 12% gemiddelde kroonbedek-

king van die struikstratum. Die opvallendste dikotiele

kruide is Berkheya setifera en Crabbea acaulis. Promi-

nente grasse is Tristachya leucothrix, Trachypogon

spicatus, Themeda triandra, Heteropogon contortus en

Elionurus muticus. Die kruidstratum het ’n gemiddelde

kroonbedekking van 57%.

3. Themeda triandra— Cymbopogon plurinodis-grasveld

Die plantgemeenskap word op die laagliggende, effens

golwende, nie-klipperige habitatte met hellings van 0°— 5°

aangetref. Die gemiddelde klipbedekking van 5% is

afkomstig van die Opeenvolging Karoo.

Die diagnostiese spesies is Cymbopogon plurinodis,

Vemonia natalensis, Sonchus nanus en Tephrosia longipes

(Spesiegroep G, Tabel 1). Die dikotiele kruide Stoebe

vulgaris, Tagetes minuta, Bidens pilosa en die dwerg-

struikie Ziziphus zeyheriana asook grasse soos Aristida

congesta en Aristida mollissima dui op eig versteurde veld.

Die gemeenskap het gemiddeld 27 spesies per releve.

Die kruidstratum word gekenmerk deur die dikotiele

kruide Helichrysum rugulosum, Hermannia depressa,

Solanum panduriforme, Conyza podocephala en Schkuhria

pinnata. Algemene grasse is: Cymbopogon plurinodis,

Cynodon dactylon, Eragrostis curvula en Elionurus

muticus. Die kruidstratum toon ’n gemiddelde kroon-

bedekking van 63 % .

4. Heteropogon contortus— Loudetia simplex- grasveld

Die plantgemeeskap word net soos die Themeda

triandra— Cymbopogon plurinodis- grasveld op die laer-

liggende, nie- klipperige habitatte met hellings van 0°— 5°

aangetref. Die gemiddelde klipbedekking van 15% is

oorwegend kwartsiete afkomstig van die Witwatersrand-

sisteem.

Die spesies Euryops transvaalensis, Gazania krebsiana,

Loudetia simplex en Diheteropogon filifolius (Spesiegroep

H, Tabel 1) is diagnostics vir die gemeenskap.

Die kruidstratum word gekenmerk deur die teenwoor-

digheid van Solanum panduriforme, Commelina africana,

Helichrysum cerastioides asook grasspesies soos Loudetia

simplex, Tristachya leucothrix, Eragrostis racemosa en

Aristida junciformis met ’n gemiddelde kroonbedekking

van 52%. Die kruide Stoebe vulgaris, Euryops trans-

vaalensis, Felicia fascicularis en grasse Cynodon dactylon

en Aristida junciformis dui op versteuring en oorbeweiding

in die gemeenskap.

Bothalia 22,2 (1992)

253

400

AS 1

300

200.-

100

*23

•12

*31

•9

•1

;io *3

•42 4*V&-6-8*5

22 24" »2

! »4

•15

»40

•11

*32

*30

STEIL HELLINGS

*27

•43

•45

*50

*46 *26 #29

37

•48

AFNAME

* 2

w -r

m ±

o o

m

39

MINDER STEIL HELLINGS

100

200

300

400

500

AS 2

FIGUUR 3. — Die multidimensionele verspreiding van releves langs die eerste en tweede asse van ’n DECORANA-ordening van die Bankenveld

in die Grootvlei-distrik.

ORDENING

Die ordeningsdiagram (Figuur 3) onderskei vier plant-

gemeenskappe en twee variante op grond van diskon-

tinuTteite in die habitat. Die Diospyros lycioides—Euclea

crapa-struikveld word hoogliggend op klipperige habitatte

aangetref en word in die ordeningsdiagram (Figuur 3) deur

eenhede A en B verteenwoordig. Die Euclea crispa—Aloe

davyana-vahani (Eenheid A) kom voor teen steil noorde-

like tot noord-oostelike glooiings in teenstelling met die

Euclea crispa—Rhus pyroides-v ariant (Eenheid B, Figuur

3) wat op minder steil glooiings voorkom.

Die Diospyros lycioides—Diheteropogon amplectens-

struikveld en die Themeda triandra—Cymbopogon

plurinodis- grasveld word onderskeidelik deur Eenhede

C en D (Figuur 3) verteenwoordig. Die Heteropogon

contortus—Loudetia simplex- grasveld kom voor in laaglig-

gende habitatte by intermediere hellings en klipbedekking

en word deur Eenheid E in die ordeningsdiagram (Figuur

3) verteenwoordig.

GEVOLGTREKKING

Die plantgemeenskappe toon duidelike voorkeure ten

opsigte van spesifieke omgewingstoestande. Die resultate

verkry tydens ordening dui daarop dat hoogte bo seespieel,

klipbedekking, aspek en helling die grootste bydrae tot die

onderskeiding van die plantgemeenskappe lewer.

BEDANKINGS

Departement Landbou-Ontwikkeling word bedank vir

fmansiele ondersteuning; mnr H. Bezuidenhout vir hulp

tydens die verwerking van data, en die personeel van die

Nasionale Herbarium, Pretoria, vir die identifikasie van

eksemplare.

LITERATUUR

ACOCKS, J.P.H. 1988. Veld types of South Africa, 3de uitg. Memoirs

van die Botaniese Opname van Suid-Afrika No. 57.

BRAUN-BLANQUET, J. 1932. Plant sociology. McGraw Hill, New

York.

BREDENKAMP, G.J. 1975. 'n Plantsosiologiese studie van die Suiker-

bosrandnatuurreservaat . M.Sc.-verhandeling, Universiteit van

Pretoria.

BREDENKAMP, G.J. & THERON, G.K. 1978. A synecological account

of the Suikerbosrand Nature Reserve. 1. The phytosociology of

the Witwatersrand geological system. Bothalia 12: 513—529.

BREDENKAMP, G.J. & THERON, G.K. 1980. A synecological account

of the Suikerbosrand Nature Reserve. 2. The phytosociology of

the Ventersdorp geological system. Bothalia 13: 199—216.

BREYTENBACH, P.J.J., MYBURGH, W.J., THERON, G.K. &

BREDENKAMP, G.J. in voorb. The phytosociology of the

Villiers— Grootvlei area. South Africa. 1. Physical environment

and the plant communities of the Bb Land Type. South African

Journal of Botany.

DEALL, G.B., SCHEEPERS, J.C., & SCHUTZ, C.J. 1989. The vege-

tation ecology of the Eastern Transvaal Escarpment in the Sabie

area. 1. Physical environment. Bothalia 19: 53—67.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK. L.

1985. List of species of southern African plants. Edn 2, Part 1.

Memoirs van die Botaniese Opname van Suid-Afrika No. 51.

254

Bothalia 22,2 (1992)

GIBBS RUSSELL, G.E., WELMAN, W.G., RET1EF, E., IMMEL-

MAN, K.L. GERMISHUIZEN, G., PIENAAR, B.J., VAN

WYK, M. & NICHOLAS, A. 1987. List of species of southern

African plants. Edn. 2, Part 2. Memoirs van die Botaniese

Opname van Suid-Afrika No. 56.

HILL, M.O. 1979a. DECORANA: A FORTRAN program for detrended

correspondence analysis and reciprocal averaging. Cornell

University New York, Ithaca.

HILL, M.O. 1979b. TWINSPAN: A FORTRAN program for arranging

multivariate data in an ordered two-way table by classification

of the individuals and attributes. Cornell University, New York,

Ithaca.

KRUGER, G.P. 1983. Terreinmorfologiese kaart van Suidelike Afrika.

Navorsingsinstituut vir Grand en Besproeiing. Departement Land-

bou, Pretoria.

LANDBOU-ONTWIKKELINGSPROGRAM 1986. Departement van

Landbou en Watervoorsiening, Hoeveldstreek, Fotchefstroom.

LANDTIPE OPNAME PERSONEEL 1984. Landtipes van die kaarte

2628 Oos Rand, 2630 Mbabane. Memoirs on the Agricultural

Natural Resources of South Africa No. 5.

MACVICAR, C.N., DE VILLIERS, J.M., LOXTON, R.F., VERSTER,

E., LAMBRECHTS, J.J.N., MERRYWEATHER, F.R., LE

ROUX, J., VAN ROOYEN, T.H. & HARMSE, H.J. VON M.

1977. Soil classification: a binomial system for South Africa.

Department of Agricultural Technical Services, Pretoria.

MYBURGH, W.J., BREYTENBACH, P.J.J., THERON G.K. &

BREDENKAMP, G.J. in voorb. The phytosociology of the

Villiers-Grootvlei area. South Africa. 5. Floristic analysis and

checklist. South African Journal of Botany.

NASIONALE WEIDINGSTRATEGIE 1985. Departement van Land-

bou en Watervoorsiening, Pretoria.

RUTHERFORD, M.C. & WESTFALL, R.H. 1986. Biomes of southern

Africa — an objective categorization. Memoirs van die Botaniese

Opname van Suid-Afrika No. 54.

SCHEEPERS, J.J., SMIT, J.A., LUDICK, B.P. 1984. 'n Evaluasie van

die landboupotensiaal van die Hoeveldstreek in terme van

droelandgewasverbouing en veeproduksie. Hoeveldstreek,

Departement van Landbou.

SOUTH AFRICAN COMMITTEE FOR STATIGRAPHY (SACS) 1980.

Stratigraphy of South Africa. Part 1 (Comp. l.e. Kent). Litho-

stratigraphy of the Republic of South Africa, South West Africa/

Namibia, and the Republics of Bophuthatswana, Transkei and

Venda. Handbook of the Geological Survey of South Africa 8.

VAN DER BANK, W.J., VERSTER, E., ROBERTS, V.E. & MAC-

VICAR, C.N. 1978. Soil survey of Grootvlei. Technical

Communication No. 145. Dept. Agricultural Technical Services,

Pretoria.

WERGER, M.J.A. 1974. On concepts and techniques applied in the

Zurich-Montpellier method of vegetation survey. Bothalia 11:

309-323.

WESTFALL, R.H., VAN STADEN, J.M. & PANAGOS, M.D. 1987.

Predictive species-area relations and determination of subsam-

ple size for vegetation sampling in the Transvaal Waterberg. South

African Journal of Botany 53: 44—48.

Bothalia 22,2: 255-282 (1992)

A phytosociological study of Signal Hill, Cape Town, utilizing both

perennial and ephemeral species

C. JOUBERT* and E.J. MOLL**

Keywords: ephemeral, perennial, phytosociology, West Coast Renosterveld

ABSTRACT

A phytosociological study based on the collection of vegetation and environmental data from 53 randomly stratified sample

plots on Signal Hill, Cape Town, was carried out over an area of 124 ha. The survey extended over 12 months to ensure

the inclusion of as many plant species as possible, and a list of the vascular plant species was compiled. A total of 81 families,

255 genera and 460 species was identified. The phytosociological method revealed that only one major plant community

occurs in the study area and two subcommunities, with a total of five variants correlated mostly with aspect and historic

land use, were identified. The perennially and seasonally identifiable species were analysed separately to determine their

relative contribution to the phytosociological classification. The two data sets gave similar classifications. A vegetation map

as well as a soil map was compiled.

UITTREKSEL

’n Fitososiologiese studie, gegrond op die versameling van plantegroei- en omgewingsdata by 53 ewekansige gestratifiseerde

monsterpersele oor ’n gebied van 124 ha op Seinheuwel, Kaapstad, is gedoen. Die opname is oor 12 maande uitgevoer om

te verseker dat soveel plantspesies as moontlik ingesluit word, waama ’n lys van die vaatplante opgestel is. Altesaam 81

families, 255 genusse en 460 spesies is geidentifiseer. Die fitososiologiese metode het aan die lig gebring dat slegs een

hoofplantgemeenskap in die studiegebied voorkom, en twee subgemeenskappe met altesaam vyf variante wat meestal ten

opsigte van aspek en historiese grondgebruik korreleer, is geidentifiseer. Die meerjarige en efemere spesies is afsonderlik

ontleed om elke groep se bydrae tot die plantegroeitipes te bepaal, en die twee stelle data het soortgelyke klassifikasies opgelewer.

’n Plantegroeikaart asook ’n grondkaart is opgestel.

INTRODUCTION

Since the earliest times the flora of the Cape has

fascinated travellers, visitors and scientists (Raven-Hart

1967, 1971). The vegetation of Signal Hill has been used

for grazing and fuel supplies since prehistoric times, and

after European settlement some areas were cultivated and

afforested (Joubert 1991). In 1964 the area was proclaimed

a nature reserve (Ashton 1985). The natural vegetation has

been protected since then.

The vegetation is broadly classified as West Coast

Renosterveld (Moll & Bossi 1984) and falls within the

Fynbos Biome (Kruger 1978). The area is a unique West

Coast Renosterveld site as it is the only area on Malmes-

bury shale influenced by coastal fog. No plant community

study has previously been made of the area except for a

post-fire study (Michell 1922) of the vegetation along the

eastern slopes.

Werger (1974) states that ‘In floristically rich areas ...

communities can be clearly characterised floristically on

the bases of floristic lists in which only permanently recog-

nisable species are entered’. He further states that in arid

regions perennials are generally better indicators of

specific habitat factors because annuals are a relatively

unimportant component. Le Roux (no date) found in

Namaqualand that species composition and cover of

ephemerals vary during the growth period and from year

to year as new vegetation associations are formed annually.

* Present address: Cape Technikon, P.O. Box 652, Cape Town, 8000.

** Botany Department, University of Cape Town, Private Bag, Ronde-

bosch 7700.

MS. received: 1992-03-25.

Annuals have, however, been found to be very useful in

some studies of arid vegetation (Werger 1974). Thus one

aim of the present study was to evaluate whether

ephemerals were important for plant community studies.

Therefore both perennially and seasonally identifiable

species were recorded and the data were analysed to

determine their relative contribution to the phytosocio-

logical classification. This form of comparison has not

previously been published.

STUDY AREA

Locality

Signal Hill (latitude 33° 54' and longitude 18° 21') is

a 2.5 km long ridge above Cape Town and is joined to

the rest of the Table Mountain Nature Reserve by Lion’s

Head (Figure 1). The reserve was proclaimed in 1964 and

is managed by the Cape Town City Council (CCC) (Ashton

1985).

The study area (124 ha) is situated on the northwest,

north and southeast-facing slopes of Signal Hill, ranging

between 120 m and 350 m above sea level. Several valleys

and dry ravines of more or less equal depth dissect the

hillside. The slope varies between 17° and 39°.

Geology

The rocks comprising Signal Hill form part of the Tyger-

berg Formation of the Malmesbury Group which consists

mainly of irregular alternations of grey to green phyllitic

shale, siltstone and medium to fine-grained greywacke

256

Bothalia 22,2 (1992)

which is generally more massively bedded than the pelitic

rocks. Ripple cross-lamination as well as micro-layering

is present in the steeply dipping grey wacke beds and ripple

marks, slumping and sole structures also occur on the

bedding planes. A few thin layers of lava, pyroclastics,

quartzite, grit as well as conglomerate are present in the

Formation (Theron 1984).

The rocks of the Tygerberg Formation are to a large

extent covered by superficial sediments and are frequently

deeply weathered to red-brown or yellow clay and loamy

soil. The degree and depth of weathering change consider-

ably over relatively short distances (Theron 1984).

Climate

According to the Koppen classification the study area

experiences a typical Mediterranean climate; i.e. the Csb

type (Schulze & McGee 1978). There is no weather station

in the study area, though a rain gauge existed on the

summit from 1882 until 1950. Data of various kinds are

available from stations surrounding the area, and these

indicate that local topography plays an important role in

influencing the mesoclimates of the three respective slopes.

Rain is brought in winter by northwesterly winds when

a cold front approaches the land (Schulze 1972). In late

winter or spring an influx of cold air may occur, causing

stormy weather and gales. As the cold front moves across

the coast, showers occur after the passage of the front and

clear up rapidly from the west (Schulze 1972).

Hot, dry, gusty berg winds prevail mainly during sum-

mer. Maximum temperatures recorded in the Fynbos

Biome appear to be associated with these winds. With the

onset of the wind, temperatures rise sharply and humidity

Bothalia 22,2 (1992)

257

drops markedly (Fuggle 1981), creating a fire hazard in

the area. The prevailing winds in the dry summer are

strong southerly to southeasterly winds (Fuggle 1981).

Signal Hill is about 350 m high and forms a barrier against

the prevailing winds. When the southeaster blows, the

northwestern aspect is completely protected, but the

northern aspect less so. Likewise, when the cold north-

wester blows from across the sea, the southeastern aspect

is protected, while the northern aspect is once again

exposed to the wind. The latter aspect is, therefore, more

often exposed to wind.

Signal Hill receives a mean of 463.5 mm rain annually.

Figure 2 illustrates the data collected during the period

1882 to 1950 (Weather Bureau 1986). Data for the study

area and five stations from surrounding areas were

collected over a period of about 70 years and appear in

Table 1. Topography plays a major role as illustrated by

the data collected on different sides of Signal Hill. In all

cases the highest precipitation, i.e. 60% of the total, is

experienced from May to August with a peak during June,

while January and February are the driest months. The

mean number of rainy days for Signal Hill is 86. July has

the highest number of rainy days i.e. 12, and during

December to February there are on average only three

rainy days per month.

No temperature data for the study area are available,

though data from stations in the vicinity show very clearly

that a moderate temperature is experienced with no

extremes on a daily or a seasonal basis. The hottest month

is February and the coldest month is July.

Fog forms when warm air is blown over the cold Atlantic

sea on northwesterly to westerly air drifts (Fuggle 1981),

thus the western slopes receive most fog, with a peak

during April and May according to data from the Cape

Town Harbour.

Flora — historical records

It is impossible to visualise the vegetation of the study

area before European settlement and the extent of the

disturbances which followed, as the early descriptions only

give a sketchy impression. Sparrman (1785) noted that such

reports are not always reliable, as these men had spent

months at sea, with the result that they tended to over-

praise the Cape at the sight of greenery after such a long

time at sea (Skead 1980).

City Hospital Temp — 1 — Signal Hill Rainfall

FIGURE 2.— Walter-Lieth climate diagram for Signal Hill and City

Hospital, Cape Town.

TABLE 1. — Annual rainfall for Signal Hill, Cape Town and five

surrounding stations

One of the first descriptions of the study area was made

by Van Riebeeck who, on 27 April 1652, ‘went along the

downs behind the rump of the Lion Mountain where we

found between the mountain and the downs the most

beautiful land for sowing and for grazing cattle that one

can desire . . . Crossing the Lion Mountain on the seaward

side of the head, found the slopes on the other side dry

and stony ...’ (Skead 1980). A seaman who visited the

seaward slopes during 1685 found the area ‘not at all

Rocky, but cover’d over with Grass’ (Raven-Hart 1971). It

appears that grass was already very common at this time

as another seaman wrote in 1702 that the ‘. . . Lion’s Rump

... is grown over with luxuriant grass and a few trees . . .’

(Raven-Hart 1971). A photograph dated ± 1910 (Cape

Archives: E 8144) shows grassland along the northern

slopes, while another dated 1899 (Cape Archives: Dr. J

80) shows areas densely populated by low scrub, possibly

Elytropappus rhinocerotis (renosterbush) .

Between 1657 and 1665, some 80 Khoikhoi were living

in a kraal on the eastern slopes of Signal Hill. During

Kolbe’s visit between 1707 and 1713 two large Khoikhoi

kraals existed at the foot of the eastern slopes of the Lion’s

Hill (Fagan 1989). It is possible that their fires maintained

the grassland on the slopes, and that they started this firing

in an attempt to promote firewood and grazing for their

herds, or to stimulate the growth of geophytes (Deacon

1983).

Michell (1922) compiled a detailed description of her

post-fire study site on the eastern slopes (1919—1921).

Unfortunately no study was made prior to the fire, with

the result that the relative importance of renosterbush and

Rhus lucida at this time is not known. She states that the

vegetation is ‘sclerophyllous bush, the characteristic

vegetation of the southwestern region’, though the area is

‘deficient in several typical southwestern families’. All the

species encountered during this period are listed by her,

many with notes on their numbers and localities. Of

particular interest is the following: The vegetation of the

valleys ‘show certain marked differences’ from that of the

open slopes. Michell states that the vegetation in the valleys

was not badly burnt owing to the somewhat sheltered

position of the watercourse in each valley, and that

the valley bottoms were ‘covered with Acacia karroo ’.

Adamson & Salter (1950) also note the presence of this

species on Signal Hill. Today only isolated individuals

occur. The slopes were dominated by R. lucida after the

fire, which ‘dotted’ the landscape at ‘frequent intervals’.

The north-facing slopes had a ‘more open type of vegeta-

tion’ than the south-facing slopes. The latter had a

conspicuously different plant population during the winter

months, though the contrast was less during summer.

258

Bothalia 22,2 (1992)

Clutia pulchella was common at the foot of one of the

valleys (similar to relev6 28 of the present study). Michell

(1922) also mentions the presence of Noltea africana.

Hyparrhenia hirta was common and mentioned several

times. Some 20 other grass species are also mentioned.

Protea repens, P. nitida and Leucadendron argenteum

are reported as being scarce. A number of annuals was

associated with cattle manure.

Michell (1922) concludes by stating that the fire favoured

the renosterbush which was ‘far in advance of any

others’ and, with the exception of the northern slopes

of the valleys, evenly distributed all over the area. The

vegetation type established after the fire is termed

‘Rhenosterveld’ and considered an ‘artificial one’. Mem-

bers of the Proteaceae, Rutaceae and Ericaceae were ‘only

occasionally seen’. She states that ‘especially in the case

of the Proteaceae, bush fires have been largely instrumen-

tal in eradicating large numbers of species from these

slopes’.

A few other descriptions from the early part of this

century give an idea of what the vegetation was like at

the time. According to these descriptions Proteaceae were

common on the slopes of Signal Hill. Liickhoff (1951)

states: ‘Most of the original Signal Hill flora has been

destroyed. The older generation still speaks of fields of

proteas that once grew on the hill. Today only a few plants

survive. For the rest we find pines, gums, taaibos and

abundant grass — the latter always reliable evidence of

repeated burning’. It appears that Protea repens was once

extremely common (Jackson 1977) on the lower slopes of

the study area, whereas large tracks of P. lepidocarpoden-

dron were described by Marloth during the early part of

this century (Liickhoff 1951). ‘Almost the whole of Signal

Hill used to be covered with Proteaceae, mainly P. repens,

P lepidocarpodendron, P. nitida, Leucospermum conocar-

podendron and Leucadendron argenteum' (Liickhoff 1951).

Today only isolated individuals of some of these species

have survived. Adamson (1929) noted that renosterbush

is ‘well developed on the slopes of Signal Hill’ and that

the community is ‘relatively pure and slow changing’ on

the western slopes. The area carried no appreciable forests

(Liickhoff 1951).

METHODS

Soil map

A soil map was compiled for Signal Hill. Soil profiles

were studied throughout the area. Soil samples were taken

at four different localities and analysed by the Faculty of

Agriculture at the University of Stellenbosch. The pH was

measured in KC1. Resistance is expressed in ohms, using

a standard USDA soil cup. The localities and soil profiles

are marked A, B, C and D (Figure 3). The results of the

soil analyses and the exposition as regards the symbols

in Tables 3 & 4, are given in Table 2, as well as dominant

soil families, a brief description of the soils and a

topographical description of the sample plots.

Phytosociological study

The field work for the vegetation survey was started in

December 1988 and completed in December 1989. Releves

were compiled from 53 stratified random plots (Werger

FIGURE 3. — Soil map of Signal Hill, Cape Town. See Table 2 for expo-

sition of symbols. Soil profiles for four localities in the study

area (marked A, B, C and D on the soil map). A, Glenrosa soil

form: non-bleached A horizon (0.0— 0.3 m) on a non-hard, non-

wet, and non-calcareous lithocutanic B horizon (0.3 -0.9 m) on

saprolite (0.9— 1.3 m); A horizon, pH = 4.7, R = 2730; B horizon,

pH = 4.2, R = 2960. B, Oakleaf soil form: non-bleached A

horizon (0.0— 0.3 m) on a non-red, non-luvic B horizon on saline

saprolite (shale); A horizon, pH = 5.0, R = 1360; Saprolite, pH

= 5.1, R = 168. C, Oakleaf (to Inhoek) soil form: non-bleached

(weak to moderate structured, dark coloured) A horizon (0.0— 0.5

m) on a reddish coloured non-luvic neocutine B horizon (0.50—1.3

+ m); A horizon, pH = 5.2, R = 2420; B horizon, pH = 4.4,

R = 2630. D, Hutton (to Inanda or Sweetwater) soil form: meso-

trophic (dark coloured, probably humic or humic phase A

horizon) non-luvic red apedal B horizon (A + B horizon 0.9 m

deep) on well-drained weathered saprolite (shale); A horizon,

pH = 4.8, R = 2980; B horizon, pH = 4.5, R = 4450.

1974). These were permanently marked with a steel

dropper, 1.3 m high. A 700 mm long white PVC conduit

tubing with a diameter of 20 mm was fixed onto the

dropper (Van Blerk 1990), rendering the plots easily

detectable from a distance, especially in mid-high,

mid-dense shrubland (Campbell etal. 1981). A numbered

metal tag was fixed to the top of the dropper to ensure

the location of the plots even after a veld fire.

Stratification was done on a topographical basis. Six

such units were distinguished within the study area. The

number of plots was determined on an area basis within

the three major units, i.e. the three aspects with open slopes

and ravines (Figure 4). Cultivated and built-up areas were

excluded from the survey. Randomly stratified sample plots

were sited in stands of vegetation which appeared floristi-

cally, structurally and environmentally as homogeneous

as possible (Campbell & Moll 1977). The sample intensity

Bothalia 22,2 (1992)

259

TABLE 2.— Exposition of symbols in Table 3 and on Figure 3 with brief description of soils

and topography of Signal Hill, Cape Town

in topsoil; rocky outcrops are

rare.

260

Bothalia 22,2 (1992)

TABLE 2.— Exposition of symbols in Table 3 and on Figure 3 with brief description of soils

and topography of Signal Hill, Cape Town

Phyto- Map Dominant Description of soils Topography Releve number

socio- symbol soil

logi- families

cal

table

symbol

SHALLOW TO MODERATELY DEEP LITH0S0LS WITH/WITHOUT AN E HORIZON

Bothalia 22,2 (1992)

261

TABLE 3.— Phytosociological table of the perennial species on Signal Hill, Cape Town

262

Bothalia 22,2 (1992)

TABLE 3. — -Phytosociological table of the perennial species on Signal Hill, Cape Town

Variant

1.1.1

Total no. spp. .655

.235

No. perennials

No. ephemera Is

Relev6 number

223

985

322

350

000

74

1.1.2 . 1.1.3

655565 . 7777666566648364645544544

279228 . 3375748045563 1 85086306886

332333 . 44444342333242323232222 1 2

259573.3140562984969178152124693

323122. 3333222223223 1312223 1 2322

7020755 . 0235286161 604017934282293

0000030.3112132111133142132222231

268913 . 4760922280 13050645 1734593

1.2.1

5543322334645

0021586793124

3221222222424

0639822469580

2211000111111

0492764334644

4353342544544

3826729165017

1.2.2

45455

48139

24333

80214

11022

68925

42454

98834

Species Group C

Aspalathus cymbiformis

Conroe l ina africana

Orbea variegata

Cynanchun zeyheri

Berkheya car l ini folia

Anthospermum galioides

Zygophyllum sessilifolium

Crassula nudicaulis

Species Group D

Gnidia inconspicua

Erica baccans

Diosma hirsuta

Knowltonia capensis

Euryops abrotanifolius

Species Group E

Zantedeschia aethiopica

Noltea africana

Pentaschistis aspera

Ehrharta erecta

Species Group F

Cliffortia polygonifolia

Rhus tomentosa

Olea europaea

Chironia baccifera

Stoebe cinerea

Myrsine africana

Cheilanthes capensis

Kiggelaria africana

Selago corymbosa

Rhus angustifolia

Protasparagus rubicundus

Lobostemon fruticosus

Putterlickia pyracantha

* Pittosporum undulatum

* Centranthus rober

Euphorbia arceuthobioides

Species Group G

Chrysanthemoides monilifera

Hibiscus aethiopicus

Felicia fruticosa

Ischyrolepis capensis

Species Group H

Ehrharta calycina

Li nun thesioides

++

0

0++

+

+ + 1

+ 1

+ +

0 +0

++ + 0

+ +

0+ +

1 1

00

0B1+++AAA1+++

10 +1111A11A

010 + +

+0+ +010+

+AA0 1 1B

1+111 1

++ + +

0 01

+ +

0+

+ 0

A1

+00

0

+ 0

100

0+

++

0 0

+ +++

1

+1+

+0+++

011 00+0 ++0 10 1

+0+ + ++ + +

00 + + + +++

1 0 1 1++11 + 0

+++++ + 0++ + + +++++++

0 0

+ 0 0

+ 00 + +

0+ + 000

+++++

B 0

0+

1+0+

B11 + 1

01++1

1 +00

A 00

11101

B1BA4

0++A0

0+1

433

+0+

1A

00

+ +1

Bothalia 22,2 (1992)

263

TABLE 3. — Phytosociological table of the perennial species on Signal Hill, Cape Town

264

Bothalia 22,2 (1992)

TABLE 3.— Phytosociological table of the perennial species on Signal Hill, Cape Town

Species of single occurrence not included in the phytosociological table: species

(relev6:cover); Acacia karroo (35:0); Adenandra uniflora (50:+); Aspalathus hispida (9:+);

Blaeria ericoides (50:0); Cliffortia hirta (47:1); Clutia alaternoides (47:+); Clutia pulchella

(28:3); Crassula subulata (40:0); Dodonaea angustifolia (32:0); Ehrharta melicoides (38:+);

Erepsia bracteata (14:+); Erica mauritanica (40:0); Erica hispidula (47:+); Erica plukenetii

(47:+); Ficinia bergiana (44:+); Ficinia indica (9:+); Ficinia nigrescens (11:+); Helichrysum

nudifoliun (47:+); Heteropogon contortus (9:0); Hymenolepis parviflora (48:0); Lampranthus

mul t i seriatus (1:+); Lessertia capensis (50:+); Metalasia muricata (50:+); Myrsiphyllum

undulatum (2:0); Pennisetum setaceum * (5:0); Plantago lanceolata * (38:+); Podalyria sericea

(17:+); Protasparagus africanus (48:+); Rubus cf. cuneifolius * (47:+); Ruschia pulchella

(18:0); Selago adpressa (31:+); Selago serrata (42:0); Senecio rigidus (48:0); Senecio

subcanescens (48:0); Solanum americanum (25:0); Solanum pseudocapsicum * (35:0); Stipagrostis

zeyheri (9:+); Ursinia dentata (50:+). * Introduced species

Bothalia 22,2 (1992)

265

TABLE 4. — Phytosociological table of the ephemeral*

species on Signal Hill, Cape Town

Variant 1

Number of spp

No. ephemerals

No. perennials

Re l eve number

1.1. 1.1.2 . 1.1.3

655 . 4655565 . 7777666566648364645544544

235 . 0279228 . 33757480455631 85086306886

322 .1323122.3333222223223131222312322

350 . 7020755 . 0235286161 604017934282293

223 . 2332333 . 44444342333242323232222 1 2

985 . 3259573.31405629849691 781521 24693

000.0000030.3112132111133142132222231

574.1268913.4760922280130506451734593

1.2.1

5543322334645

0021586793124

2211000111111

0492764334644

3221222222424

0639822469580

4353342544544

3826729165017

1.2.2

45455

48139

11022

68925

24333

80214

42454

98834

Species Group A

* Raphanus raphanistrun

* Stachys arvensis

Lotononis penduncularis

Species Group B

Dolichos decumbens

Ornithogalum graminifoliun

Senecio arenarius

Silene clandestina

Sutera anti r rhino ides

Ornithogalum hispidum

Oxalis lanata

Lotononis prostrata

Hemi men's montana

Pelargonium hirtum

Species Group C

. 0

. + + + + + +

.+ 0 0 +0

+ . + +

. ++ +

. + +

. + + +

1 +

. + +

0

Lachenalia fistulosa

Babiana disticha

Ornithogalum thyrsoides

Pentaschistis airoides

Trachyandra muricata

Oxalis hirta

Dimorphotheca pluvial is

Felicia bergerana

Albuca canadensis

Cenia turbinata

Crassula campestris

Bulbine alooides

Pelargonium rapaceum

Ursinia anthemoides

* Gnaphalium subfalcatun

Cyphia digitata

Manulea cheiranthus

Hebenstretia repens

* Solanum nigrum

Diascia capensis

Oxalis obtusa

* Petrorhagia prolifera

Phyllopodium cordatum

Bulbine tuberosa

Gladiolus brevifolius

Arctotheca calendula

Pelargonium proliferum

* Anti rrhi nun oronti urn

Adenogranrana glomerata

Tribolium echinatum

* Conyza canadensis

Medicago polymorph a

+++.

++.

4

+++,

+ +,

+++.

+++,

+0 .

+0.

++.

+++.

+ +.

+++++ ,

++++++,

+0+++

++ + ++

+ ++++

+++ +

+ ++ +

++ + ++

+++++++

++++++

+ +

++ +

+ + +

+

,+ + ++

, +++ +

,++++++++0++++ + + ++ +

,+ +++ 4- +++++ ++ +++ 1++

,+++++ + 0++ + +++ +++0

, + +++ ++++ + +++++++ 4

, + +++ ++++ ++++++ 4

,4-4-+++++++++ ++ + +4

,+++ +++0++++ + ++ 0+

,4-+++ + + ++++ +++ +++

.+ 0+ ++0 +++ +++0

,+++ 0+ 0 +++ +0 +

.4- ++ + ++ ++ +++++

, ++ + + + + + +++ +

.+ +++00++++ 0+0 + + c

,++ + ++ ++++ +0 0

. +++ + + + o ++ +++

. + + 0+ + +

+ ++ +

++ .

+ +.

++

+ +

0+ +

00

# In this study ephemerals are those species not perennially identifiable e.g. geophytes

(cryptophytes) and annuals (therophytes).

* Introduced species.

266

Bothalia 22,2 (1992)

J i

TABLE 4. — Phytosociological table of the ephemeral

species on Signal Hill, Cape Town

Variant 1.1.1

Number of spp

No. ephemerals

No. perennials

Re l eve number

655

235

322

350

223

985

000

574

1.1.2 . 1.1.3

4655565 . 7777666566648364645544544

0279228.3375748045563185086306886

1323122.3333222223223131222312322

7020755 . 0235286161604017934282293

2332333 . 4444434233324232323222212

3259573.3140562984969178152124693

0000030.3112132111133142132222231

1 2689 1 3 . 4760922280 130506451734593

1.2.1

5543322334645

0021586793124

2211000111111

0492764334644

3221222222424

0639822469580

4353342544544

3826729165017

1.2.2

45455

48139

11022

68925

24333

80214

42454

98834

# In this . .

(cryptophytes) and annuals (therophytes).

* Introduced species.

geophytes

Bothalia 22,2 (1992)

267

TABLE 4.-— -Phytosociolog i cal table of the ephemeral^ species on Signal Hill, Cape Town

Species of single occurrence not included in the phytosociological table: species

(relev£:cover);

Aristea africana (47:+); Brachypodiun distachyon * * (25:+); Bromus pectinatus * (2:+); Carduus

pycnocephalus * (44:+); Cerastiun capense (35:+); Didymodoxa capensis (35:+); Eriospermum

capense (30:+); Gastridium phleoides * (53:0); Geissorhiza imbricata (45:+); Heliophila

diffusa (44:+); Hesperantha falcata (39:+); Homeria flaccida (22:0); Ixia dubia (14:+); Ixia

odorata (17:+); Lophochloa cristata (20:0); Micranthus alopecuroides (43:+); Microdon

sparsiflorum (14:+); Osteospermum clandestinum (5:+); Otholobium uncinatum (45:0); Oxalis

polyphylla (40:+); Oxalis pusilla (10:+); Pelargonium auritum (10:+); Pelargonium pillansii

(47:+); Picris echioides * (23:+); Ranunculus multifidus (48:+); Rapistrum rugosum * (7:+);

Senecio abruptus (44:+); Sonchus asper * (23:+); Spiloxene capensis (38:+); Vulpia bromoides

* (25:+); Wahlenbergia capensis (32:+); Wahlenbergia obovata (20:+); Wurmbea spicata (38:+);

Zehneria scabra (26:+).

# In this study ephemerals are those species not perennially identifiable e.g. geophytes

(cryptophytes) and annuals (therophytes).

* Introduced species.

for the vegetation of the study area is about one releve per

2.4 ha. Sample plot size was 10 x 10 m on the open slopes

(McKenzie et al. 1977; Campbell & Moll 1977) and 5 x

20 m in the dry ravines and valleys (Campbell & Moll

1977), resulting in plots of 100 m2 each.

A total of 293 species was recorded for the phytosocio-

logical tables. The Braun-Blanquet method (Werger 1974)

was used to classify the vegetation of the area in an effort

to show the variation within, and the relationship between,

the floristic units. Cover values for all species and heights

for the perennial species were recorded. In addition to the

values stated by Werger, an ‘O’ was used to indicate species

found within a 10 m wide belt outside the plots (McKenzie

et al. 1977). Additional environmental data were recorded

at each plot and tabulated at the bottom of the phytosocio-

logical table of the perennially identifiable species (Table

3). The plots were visited on a regular basis for 12 months

in order to record as many species as possible and a

separate table of these seasonally identifiable species was

made (Table 4).

Two-way indicator species analysis TWINSPAN (Hill

1979) was used to derive a first approximation of the vege-

tation types. A suite of computer programs used by

Boucher & Shepherd (1988) was also used and proved most

useful in this respect. The results of this classification were

refined by the Braun-Blanquet classification technique as

described by Mueller-Dombois & Ellenberg (1974), using

a word processor package to shuffle releves and species

in order to obtain a final classification. All plots sampled

are retained in the phytosociological tables (Tables 3 & 4).

The perennial species were treated separately from the

ephemeral species to determine the contribution of each

to the vegetation. Perennial species are considered to be

those which are perennially identifiable (Werger 1974)

and ephemeral species those which are not perennially

identifiable. The latter includes both therophytes and

cryptophytes. The results from the perennial species are

listed separately from those of the ephemeral species in

order to determine the relationship between the two

groups. Species with a single occurrence were listed below

268

Bothalia 22,2 (1992)

II I II //////// /1 1 ~ Contour interval 10 m

FIGURE 4. — Contour map showing the position of sample sites.

the relevant table together with the releves in which they Campbell et al. (1981) was adopted in the description of

occurred and their cover-abundance values. the community types.

Diagnostic species listed in Tables 3 & 4 were used to A list of 81 families, 255 genera and 460 vascular plant

identify the vegetation types in the veld as well as for species was compiled. This list is based on species

mapping purposes. The system and terminology of recorded during the present study as well as those recorded

Bothalia 22,2 (1992)

269

by other botanists in the past. Old taxon names were

updated using Gibbs Russell et al. (1985, 1987), or recent

revisions, or by consulting specialists in particular taxa.

Outdated taxon names which could not be updated were

excluded from the list. Species mentioned by Adamson

& Salter (1950) as occurring on Signal Hill were checked

with the Guthrie Collection (Bolus Herbarium). Only

those species which occur as actual specimens in the

herbarium and which were collected on Signal Hill were

added to the list. Species encountered incidentally during

herbarium work, for which Signal Hill is given as the

locality, were also included.

RESULTS

Soil

The more gentle slopes of Signal Hill consist mainly

of shale horizons (Smith-Baillie etal. 1976). Oakleaf (Oa)

and Glenrosa (Gs) overlying shale rock are the dominant

soil forms in the study area. Other soil families are Mispah

(Ms), Cartref (Cf), Hutton (Hu), Inanda (la), Inhoek

(Ik), Estcourt (Es), Vilafontes (Vf), Klapmuts (Km) and

Sweetwater (Sr). See Figure 3 for the soil map and Table

2 for the exposition of the symbols on the soil map.

Along the northern open slopes shallow, gravelly

lithosols with 6-15% clay in the topsoil generally occur.

E horizons are rare. An area of stony, moderately deep,

cobbly colluvial soils with salt accumulation in the subsoils

is also present. Rocky outcrops are common in the v lley

bottoms. Shallow to moderately deep, well-drained cobbly

pedisediments with 10—20% clay in the topsoil occur in

the valley bottoms.

The northwest-facing slopes generally have shallow,

gravelly to cobbly lithosols with 10—20% clay in the

topsoil. The deepest soils occur along the lower south-

feeing slopes and in valley bottoms. In the area transitional

to the northern slopes, rocky outcrops are common as well

as in the valley bottoms. Well-drained pedisediments of

varying depth with 10—20% clay in the topsoil occur in

the valleys.

Shallow to moderately deep lithosols with/without an

E horizon occur on the open east-feeing slopes. The topsoil

is usually gravelly to cobbly with 15—25% clay in the

topsoil. Deep, moderately to highly organic-rich well-

drained soils with 20—40% clay in the topsoil occur in

the southeast-facing valleys. The organic carbon content

of the valley soils is the highest recorded in the study area,

i.e. 1-5%.

Phytosociological study

The analysis resulted in the identification of one major

plant community and two subcommunities, one of which

has three and the other two variants. Each of these

vegetation types is related to a particular set of environ-

mental conditions, of which aspect and historic land use

differences were the most significant. There appears to

be no clear correlation between soil forms and vegetation

types. These community types have been mapped in

Figure 5.

The phytosociological classification, as well as the

floristic relationship between the different subcommuni-

ties and variants, is given in Tables 3 & 4. A list of the

vascular plant species recorded appears in the Appendix.

In this survey the taxonomic entities Mohria caffrorum

and Cheilanthes contmcta were merged. Introduced woody

species such as Pinus pinaster, P radiata, Acacia cy clops,

A. saligna and Eucalyptus spp. , which occur as scattered

individuals, were excluded from Table 3.

1. Rhus lucida—Chrysocoma coma-aurea Community

The community occupies the entire study area and

consequently occurs in a variety of habitats; i.e. on open

slopes as well as in dry ravines, on different geological

formations and soil types, on various aspects and at

different altitudes. The community is characterised by the

species of Species Group K (Tables 3 & 4).

The vegetation comprises an evergreen, mid-high to low,

small to large-leaved shrubland with a grassy understorey.

FIGURE 5. — Vegetation map of Signal Hill, Cape Town. Rhus

lucida—Chrysocoma coma-aurea Community. Hyparrhenia

hirta—Rhus glauca Subcommunity: Variants 1.1.1, 1.1.2 & 1.1.3.

Cliffortia polygonifolia—Rhus tomentosa Subcommunity: Vari-

ants 1.2.1 & 1.2.2.

270

Bothalia 22,2 (1992)

Depending on local site factors, the shrub canopy closure

varies from sparse to mid-dense; similarly die grass

understorey varies from very sparse to open (Campbell

et al. 1981). There are very few trees, but the multi-

stemmed shrub, Rhus lucida, is abundant and more or less

evenly distributed.

Three strata are generally recognised, namely a canopy

of mid-high sparse to mid-dense shrubs; a sparse to open

grassy/low shrub stratum; and a very sparse seasonal herb

layer. Rhus lucida, Anthospermum spathulatum and Otho-

lobium hirtum are the most constant species present in the

canopy, having a height of 1—2 m and a cover of 10— 60% .

Viscum capense was often recorded as growing on R.

lucida. The second stratum is dominated by the woody

shrubs Chrysocoma coma-aurea, Helichrysum patulum,

Salvia africana-caerulea, Rhus laevigata, Hermannia

hyssopifolia and Senecio pterophorus, and the grass Merx-

muellera stricta with a height of 0.25— 1.0 m and a cover

of 10-25%.

The herbaceous layer with a cover of generally < 5 %

consists of the forbs Senecio hastatus and Orobanche

ramosa, and the fern Cheilanthes hastata. Geophytes are

a feature on the northern aspects in particular, and autumn

to spring ephemerals are prominent in areas of sparse vege-

tation. Characteristic ephemeral species are the forbs

Pelargonium elongatum, P. lobatum\ the geophytes Oxalis

glabra, O. purpurea, O. pes-caprae, O. bifida, Cyanella

hyacinthoides, Moraea bellendenii and the introduced

grass Briza maxima. Lichens and mosses were recorded,

but omitted from the phytosociological table. It appears

that lichens are generally less common on the east and

southeast-facing slopes than in the rest of the area.

Plant litter cover varies between 5% and 95%, and

consists mainly of leaves of R. lucida and H. patulum,

grasses and twigs. The total canopy cover of the vege-

tation varies from 35-140%, and an average of 31

perennial and 22 ephemeral species was recorded per

releve.

Very few species were found growing under the Rhus

bushes. Those which did occur include Senecio hastatus,

Stachys aethiopica, Scabiosa columbaria, Annesorrhiza

capensis, Oxalis spp., annual orchids and fern clusters

under the larger bushes. The climber, Helichrysum

patulum, with a height of about 0.7 m, often scrambles

into other shrubs such as R. lucida and R. glauca,

and when thus supported can reach a height of about 1.2

m. Other scramblers in R. lucida are Helichrysum

cymosum and Chrysocoma coma-aurea, though to a lesser

extent.

This community is divided into two subcommunities.

Subcommunity 1.1, the Hyparrhenia hirta—Rhus glauca

Subcommunity, occurs in the drier parts, and is charac-

terised by the species listed in Species Group C (Tables

3 & 4). Subcommunity 1.2, the Cliffortia polygonifolia—

Rhus tomentosa Subcommunity occurs in the wetter parts,

and is characterised by the species listed in Species Group

F (Table 3). The diagnostic ephemeral species for the latter

subcommunity are less well defined and are listed in

Species Groups D and E (Table 4).

1.1 Hyparrhenia hirta—Rhus glauca Subcommunity

This subcommunity of 35 releves is the most extensive

and occupies approximately 80 ha (64% of the study area).

It occurs mostly on the north and northwest-feeing seaward

side, though two releves, Nos 39 and 40, face east-north-

east. This is the more diverse subcommunity, probably

related to the diversity of microhabitats available because

of the occurrence of a variety of soil types, and as a result

of disturbance history including fire. The soils are

generally shallow, usually gravelly to cobbly, with the

topsoil overlying shale rock and having 6—40% clay in

the topsoil. Rocky outcrops are rare to common.

The high constancy (100%) and relatively high canopy

cover, ranging from <1% to 40%, of H. hirta and

relatively constant presence (80%) and locally high canopy

cover, up to 45%, of R. glauca, are features of this sub-

community. The rest of the differential species in this

subcommunity have a relatively low canopy cover. The

high constancy and canopy cover of H. hirta, and the low

cover of woody species is an indication of the generally

poor condition of this vegetation, attributed to frequent

fires, especially in variants 1.1.1 and 1.1.2.

Species which differentiate this subcommunity are listed

in Species Group C (Tables 3 & 4). An average of 32

perennial and 26 ephemeral species was recorded per

releve.

In this open grassy/low shrubland three strata are

evident. The mid-high sparse shrub stratum, with a canopy

cover of 10—20%, is 1.3 m tall and consists mainly of

Rhus lucida, Anthospermum spathulatum and Otholobium

hirtum. The open grassy/low shrub stratum is dominated

by the woody R. glauca (0.90—0.96 m tall; 7% cover),

and the grass H. hirta (0.86 m tall; 10—25% cover). Other

species in this stratum of 0.25-1.0 m tall and cover

of 35-40% are the low woody shrubs Protasparagus

capensis, Hermannia alnifolia, H. prismatocarpa, Aspala-

thus ericifolia, A. cordata, A. cymbiformis and Sutherlan-

dia frutescens. The sparse herbaceous component

includes, apart from H. hirta, other grass species such

as Ehrharta calycina and Tribolium hispidum.

The very sparse herb/dwarf-shrub stratum '(<0.25 m)

with a cover of <5% is represented by the woody

Hermannia althaeifolia, Helichrysum cylindriflorum, H.

asperum, Aspalathus acuminata, Pelargonium myrrhifo-

lium and Leysera gnaphalodes. A fairly common climber

in R. lucida is Pelargonium candicans. The most common

succulents are Lampranthus emarginatus, Ruschia rubri-

caulis, Orbea variegata and Cynanchum zeyheri. The forbs

include Indigofera incana, Cineraria geifolia, Euphorbia

crispa, Aizoon sarmentosum and Commelina africana\ and

the ephemerals as listed in Species Group C (Table 4).

Of these ephemerals, species such as the geophytes

Lachenalia fistulosa, Babiana disticha, Omithogalum

thyrsoides, Trachyandra muricata, Oxalis hirta, Albuca

canadensis, Bulbine alooides\ the herbs Dimorphotheca

pluvialis, Felicia bergerana, Cenia turbinata, Crassula

campestris, Pelargonium rapaceum, Ursinia anthemoides,

Gnaphalium subfalcatum\ and the grass Pentaschistis

airoides are the most common.

Bothalia 22,2 (1992)

271

This subcommunity is divided into three variants.

Species listed in Species Group C (Tables 3 & 4) indicate

the relationship among these three variants.

Variants 1.1.1 and 1.1.2

Most of this area was burnt during early 1986, less than

three years prior to the survey.

These two variants can be treated as a single unit as they

are closely related. The difference between the two variants

is characterised by the two species, Eragrostis curvula and

Erepsia anceps present in variant 1.1.1, but absent from

variant 1.1.2. The relationship between these two variants

is shown by the relative absence of species listed in

Species Group B (Tables 3 & 4). These variants are also

discerned from variant 1.1.3 by the absence of these

species.

This vegetation occurs on open slopes as well as in

dry ravines (Figure 6). These variants occur on steep

(22°-31°), predominantly convex, mainly north-facing

slopes characterised by extreme exposure to solar radia-

tion in the driest and hottest part of the study area. They

cover some 20 ha (16%) of the study area on a variety of

soils. The vegetation generally (seven of the 10 releves)

occurs on shallow lithosols, some 200 mm deep. These

are usually gravelly, with the topsoil overlying shale rock,

and with 6—15% clay in the topsoil. Rocky outcrops and

E horizons are rare. Releve 6 occurs on soils similar to

FIGURE 6.— Variant 1.1.1. Dry north-facing ravine.

those associated with variant 1.1.3. Moderately deep

(600—900 mm) stony colluvial soils with salt accumulation

in the subsoils were recorded in relev6s 3 and 8. These

are cobbly soils with 15—25% clay in the topsoil. The

substrate of these variants is generally rather rocky and

most of the rocks fall into the category 20—200 mm

diameter. Bedrock occurs in most of the releves,

irrespective of altitude whether they are in dry ravines or

on open slopes.

Between 15% and 55% of the surface is bare. Litter

cover is between 5% (releve 7, a fire having been

experienced a year previously) and 60% (releve 6, on the

open slope). Total canopy cover during summer varies

between 45 % and 65 % . During winter cover increases by

about 10% and is associated with the appearance of

ephemeral species. The height of the vegetation does not

exceed 1.5 m.

The total number of plant species recorded per releve

varies between 40 and 62, with about 32 perennial and

24 ephemeral species per releve. Species Groups I and

J (Table 3) list those perennial species which occur

throughout the study area, though generally not in these

two variants. Species listed in Species Group I do not occur

in the valleys on the southeast-facing slopes either.

Conspicuous species which are absent from these two

variants are the woody shrubs Elytropappus rhinocerotis,

Pelargonium cucullatum, Lobostemon argenteus, Passe-

rina filiformis , Cliffortia ruscifolia, Gnidia laxa, Montinia

caryophyllacea, Athanasia trifurcata, Diospyros glabra',

and the herb Scabiosa columbaria.

Species Group J (Table 4) lists the ephemeral species

which occur all over the study area except in these variants.

The most prominent of these are the forbs Anagallis

arvensis, Sebaea aurea, S. exacoides, Pelargonium colum-

binum; and the geophytes Geissorhiza aspera, Crassula

capensis and Annesorrhiza capensis. Species Group I

(Table 4) lists the ephemeral species which occur all over

the study area except in these variants and generally not

in the valleys on the southeast-facing slopes either. Such

absent species are the geophytes Urginea altissima, Oxalis

tomentosa, O. punctata and Haemanthus sanguineus; and

the forbs Pterygodium catholicum, Arctopus echinatus and

Wachendorfia paniculata.

Three strata are evident in this xeric open grassy/low

shrubland. The vegetation (Figure 7) is almost entirely

dominated by the constant occurrence of the grass

Hyparrhenia hirta (0.8 m tall) and, to a lesser extent, the

woody shrub Rhus glauca (0.9 m tall). The mid-high

sparse shrub stratum, about 1.2 m tall, with a canopy cover

which varies from absence to 18 % and an average of 10 % ,

consists mainly of Rhus lucida, Anthospermum spathula-

tum and Otholobium hirtum. The open grassy/low shrub

stratum is dominated by H. hirta and R. glauca with a

cover of 25% and 6.5% respectively. Other species in this

stratum with a cover of some 40% as well as those in the

very sparse herb/dwarf-shrub stratum are similar to those

listed in the general discussion on this subcommunity and

to a lesser extent those listed in Species Groups A and

B (Table 4).

This vegetation creates an appearance of almost pure

grassveld since woody plants are not prominent. The

272

Bothalia 22,2 (1992)

FIGURE 7.— Variant 1.1.3. Rhus

lucida dotting the slopes with

Hyparrhenia hirta and Heli-

chrysum patulum.

floristic analysis, however, revealed that the woody species

in many areas, though not conspicuous, are present.

Species such as R. lucida (1.4 m), R. glauca (0.9 m),

R. laevigata (1 m) and Anthospermum spathulatum (1 m)

are often dwarfed. Conspicuous in this grassveld are the

often tall (up to 1.45 m) A. spathulatum individuals at

higher altitudes.

It appears that frequent fires played an important role

in determining vegetation structure and floristic compo-

sition. Evidence from historical photographs indicates that

parts of this vegetation have been grassveld for at least nine

decades. The high cover of Hyparrhenia hirta seems to

be a result of these frequent fires. The frequency of fires

has, however, decreased in recent years.

There appears to be no difference floristically between

the open slopes and dry ravines. An attempt was conse-

quently made to determine whether a difference on a

structural basis existed between these two habitats. The

canopy cover and height of the woody R. lucida, R. glauca

and Otholobium hirtum were studied. It was found that

R. glauca attains the highest canopy cover value and height

in the ravines; i.e. in releves 1, 2 and 5. A mean canopy

cover of 18% and a height of 1.1 m was recorded in the

ravines, versus a mean canopy cover of 2% and a height

of 0.75 m on the open slopes. The other two species did

not show this tendency. It was also noted during field work

that O. hirtum forms extensive thickets in the valley

bottoms, though no data were collected in these areas. A

mean canopy cover of < 3 % for this species in these two

variants was recorded, though local high values were

recorded; i.e. 15% in releve 2 (a dry ravine) and 10%

in releve 8 (with moderately deep soil on the open

slope).

The cause of the similarity in species composition

between the dry ravines and open slopes may be the

extremely shallow soil in the former as bedrock attains

its highest cover in the ravines. None of these ravines is

wet for a long period, and runoff is probably rapid.

It appears that these two variants are at present in a

relatively stable subclimax stage induced abiotically by

excessive firing over a relatively long period.

Variant 1.1.3

This is the most extensive variant in the study area and

covers some 60 ha (48%) of the study area. It occurs on

the less xeric, mostly west and northwest-facing slopes.

The slope varies between 17° and 39°. Only three ravines

occur and these are situated on the north-north-west-facing

side (releves 24, 25 and 35), although open low-lying areas

occur on the west-facing side (releve 26). With an average

of 33 perennial and 26 ephemeral species recorded per

releve, it is floristically the richest of the five variants.

The vegetation occurs on a variety of soils, though

generally on shallow lithosols (E horizons are rare), which

are <0.3 m deep and usually with gravelly to cobbly

topsoil overlying shale rock with 10—20% clay in the

topsoil. Rocky outcrops are rare to common in places, and

also occur in the valley bottoms. The deepest soils occur

along the lower southwest-facing slopes and in valley

bottoms. Releves 20 and 30 occur on well-drained

pedisediments of moderate depth ( < 600 mm) which are

gravelly to non-gravelly.

The age of the vegetation also varies significantly, i.e.

between 8 and 20 years. Between 10% (releve 26) and 50%

(releves 12, 33 and 35) of the surface is bare during

summer. During winter this percentage decreases by about

10%. Species responsible for this increase in cover are

mainly Senecio hastatus, Stachys aethiopica and the

ephemeral species listed in Species Groups B, C, H, I,

J and K (Table 4). The total canopy cover for summer

varies between 35% (releve 17) and 70% (releve 19).

The relatively high canopy cover of Elytropappus

rhinocerotis in releve 33 suggests frequent fires in the past

(Michell 1922). The extreme rockiness of this releve

may be attributed to surface erosion under such conditions.

Brownlie (1982) found a positive correlation between

E. rhinocerotis and eroded areas on clay soil. Many large

(4 m2) bare patches are conspicuous in releves 20 and

32 which occur on the same southwest-facing slope.

In these releves, 30—35% of the surface is bare. The

vegetation is about 20 years old. Very large specimens

of both old and dead E. rhinocerotis individuals were

recorded.

Bothalia 22,2 (1992)

273

Litter cover varies between 30% in a high lying rocky

relev6 with several succulent species (releve 12) to about

75 — 85% in ravines at lower altitudes (releves 13, 24, 25

and 26). The highest litter values appear to be correlated

with wet areas and not with the age of the vegetation.

This variant is characterised by the prominence of

species listed in Species Group B (Tables 3 & 4), although

these species also occur to a lesser extent in variant 1.1.2

of this subcommunity.

Three strata are evident in this sparse grassy/low to mid-

high open shrubland. The mid-high sparse shrub stratum,

about 1.3 m tall and with a canopy cover of about 20%,

is dominated by R. lucida, which has a canopy cover of

11% in this variant versus 6% in the former two variants.

Other species in the mid-high shrub stratum (1-2 m

tall) are Anthospermum spathulatum, Otholobium hirtum,

Chrysanthemoides monilifera, Pelargonium cucullatum,

Cliffortia ruscifolia and Atharuxsia trifurcata.

Characteristic woody shrubs in the sparse grassy/low

sparse to open shrub stratum (0.25—1.0 m) with a canopy

cover of about 35%, are Elytropappus rhinocerotis,

Lobostemon argenteus, Eriocephalus africanus, Salvia

africana-lutea, Chrysanthemoides incana and Euclea

tomentosa. The tall H. hirta (0.85-0.90 m) with a canopy

cover which ranges from < 1— 30% , and a mean of 10% ,

is generally conspicuous, and the stunted Themeda trian-

dra (<1— 20%), Cymbopogon marginatus (<1— 6%) and

Tribolium uniolae are locally prominent.

In the sparse herb/dwarf-shrub layer (<0.25 m) with

a canopy cover of <5%, Hermannia althaeifolia,

Helichrysum cylindriflorum, Leysera gnaphalodes, Pelar-

gonium myrrhifolium and Helichrysum asperum are

prominent woody shrubs. The climbers Indigofera

psoraleoides , Cissampelos capensis, Lessertia excisa and

Tephrosia capensis occur locally. This layer generally has

a canopy cover of < 5 % in summer which can increase

to 75% in winter (releve 25). Prominent herb species in

this stratum are Senecio hastatus, Stachys aethiopica, the

fern Mohria cajfrorum and the ephemeral species listed

in Species Group B (Table 4) which occur scattered with

low canopy covers.

Low-growing succulents are found scattered in this

variant. These are often found among rocks and on rocky

outcrops at higher altitudes in particular. Soil is generally

shallow ( < 250 mm) and common species are Othonna

arborescens, Euphorbia caput-medusae, Tylecodon gran-

diflorus, Adromischus hemisphaericus , Crassula scabra,

Lampranthus emarginatus and Ruschia rubricaulis.

At higher altitudes, such as in releve 19, wind-pruned

shrubs of R. lucida and Euclea tomentosa occur. The latter

is particularly prominent locally in these rocky parts.

Isolated pine trees, mainly P pinaster, occur on the slopes.

As in the former variant, the ravines appear to be no

different floristically from the open slopes. Structural

differences occur in that both Rhus glauca and Hyparr-

henia hirta are taller in some ravines (releves 24 and 25),

though they do not have a higher canopy cover. The same

applies to H. hirta in releve 35, though not to R. glauca

(Figure 8). Species listed in Species Groups G, I and J

(Table 3), appear to be almost absent from releves 24, 25

and 35 (ravines) as well as from releves 21, 23 and 27 (open

slopes).

Conspicuous in this variant is the much drier appearance

of the north-north-west-facing slopes where H. hirta is

prominent, versus the southwest-facing slopes where

shrubs are dominant, giving the latter a green lush

appearance. A vertical firebreak on a west-facing slope

is maintained by using bush-cutters (Figure 9). The shrubs

are thus prevented from reaching any considerable height.

The vegetation in this firebreak is consequently artificially

converted to grassveld dominated by H. hirta. The occur-

rence of this grass firebreak and the high cover of H. hirta

on the north-north-west-facing slopes indicate the strong

relationship among the three variants in this subcom-

munity.

Two of the releves on the east-facing slopes appear to

belong to this variant rather than to subcommunity 1.2.

This may be due to the fact that these releves occur along

an east-north-east-facing slope. Variant 1.1.3 may be con-

sidered as transitional between the two subcommunities.

Species listed in Species Group I (Tables 3 & 4) indicate

FIGURE 8.— Variant 1.1.3. Dry ravine

with Acacia karroo, Rhus glau-

ca and Hyparrhenia hirta

(releve 35).

274

Bothalia 22,2 (1992)

the relationship between variants 1.1.3 and 1.2.1. The

woody shrubs Elytropappus rhinocerotis , Pelargonium

cucullatum, Lobostemon argenteus, Passerina filiformis,

Clijfortia ruscifolia, Gnidia laxa, Montinia caryophyl-

lacea ; the herbs Scabiosa columbaria, Pterygodium

catholicum, Arctopus echinatus, Wachendorfia paniculata ;

and the geophytes Urginea altissima, Oxalis tomentosa,

O. punctata and Haemanthus sanguineus indicate this

relationship.

1.2 Clijfortia polygonifolia—Rhus tomentosa

Subcommunity

This subcommunity of 18 releves occupies some 45 ha

(36% of the study area) and occurs on the mesic eastern

and southeastern inland slopes and valleys. The slope

varies between 17° and 34°. The soils are generally shallow

to deep, gravelly or non-gravelly with 15—40% clay in the

topsoil. The deepest soils occur in the valley bottoms.

The high constancy and canopy cover values of many

species such as Rhus lucida, Chrysocoma coma-aurea,

Helichrysum patulum, Clijfortia polygonifolia, Rhus

tomentosa, Olea europaea subsp. africana, Chironia

baccifera, Stoebe cinerea and Myrsine africana are

features of this subcommunity. Rhus lucida, with a canopy

cover of about 17%, is the most constant species (100%).

Diagnostic species listed in Species Group F (Table 3) are

the trees Kiggelaria africana and Olea europaea subsp.

africana -, the woody shrubs Clijfortia polygonifolia, Rhus

tomentosa, Chironia baccifera, Stoebe cinerea, Myrsine

africana, Selago corymbosa, Rhus angustifolia, Protaspa-

ragus rubicundus, Lobostemon fruticosus, Putterlickia

pyracantha ; and the fern Cheilanthes capensis.

This vegetation has been protected from fire for about

16 years. Prior to 1972, extensive pine plantations existed

on the southeastern slopes, but it appears that the veld

along these slopes has recovered to a large extent, except

for the area in which releves 29 and 51 are situated. This

latter area has been subjected to afforestation for a much

longer period than the rest of the area. Floristically this

part has also recovered and cannot be distinguished from

the rest of the slope, though the general height of the

FIGURE 9.— Variant 1.1.3. Grass fire-

break along west-facing slope.

vegetation is lower than that in the rest of the subcom-

munity.

Two variants are recognised in this subcommunity; i.e.

variant 1.2.1 which occurs on the open slopes and variant

1.2.2 which occurs in the valleys. The floristic differences

between the valleys and the open slopes are rather weak.

Species Groups D and E (Tables 3 & 4) list the charac-

teristic species for the open slopes and valleys respectively.

Variant 1.2.1

This is the most extensive variant in this subcommunity

and occurs on the east and southeast-facing inland slopes

except for releves 39 and 40 which appear to be more

closely related to variant 1.1.3 on the northwest-facing

slopes. The vegetation on the east-facing slopes generally

occurs on shallow to moderately deep (<350 mm)

lithosols with or without an E horizon. The topsoil is

usually gravelly, overlying shale rock with 15—25% clay

in the topsoil. The southeast-facing releves occur mostly

on moderately deep ( < 500 mm) lithosols with gravelly

to cobbly topsoil overlying shale rock with 15—25% clay

in the topsoil. Rocks are predominantly of the smaller size

(20—200 mm), with bedrock only in 3 of the 13 releves.

The total number of species per releve varies between

26 and 61, and for perennials between 19 and 45. An

average of 29 perennial and 12 ephemeral species per

releve was recorded. The number of ephemerals was

exceptionally low in some releves and varied between 4

and 24 species. Between 5% and 40% of the surface is

bare in summer. Litter cover varies between 45% and

95%, while total canopy cover varies between 50% and

80%.

The east-facing slopes (releves 36, 37, 38, 41 and 42)

are significantly disturbed and occur on a multiple fire-

break which is shifted periodically. The vegetation cover

in this firebreak is not lower than in the rest of the variant.

The species richness in these releves, as well as in releves

29 and 51, is generally lower than in the rest of the area.

In the disturbed area occupied by releves 29 and 51,

Helichrysum patulum attains its highest canopy cover in

Bothalia 22,2 (1992)

275

the study area, namely up to 25% with a mean of 16%,

giving this short veld a silver-grey colour.

Some of the former eucalypt firebreaks along these

slopes are at present reduced to the occasional tree stump

on which coppices frequently appear. Isolated pine trees,

mainly Pinus pinaster, occur on the slopes. Dead, burnt

pine tree stumps and relicts from dry-packed stone walls

are also encountered.

Evidence from historical photographs [Cape Archives:

J 765(5)] dated 1885, indicates that this vegetation has

recovered structurally to a very large extent and possibly

floristically as well. Considering that the last pine

plantations were destroyed by fire some 16 years ago, the

recovery of these slopes is indeed remarkable.

Three strata are recognised in this mid-high open

shrubland. The open canopy, 1— 2 m tall, with a cover of

40%, is formed by the small tree Olea europaea subsp.

africana and mid-high shrubs Rhus lucida, Anthospermum

spathulatum, Otholobium hirtum, Cliffortia polygonifolia,

R. tomentosa, R. angustifolia, Chrysanthemoides monili-

fera, Elytropappus rhinocerotis, Cliffortia ruscifolia, Erica

baccans and Euryops abrotanifolius (Figure 10).

The low sparse shrub stratum, (0.25—1.0 m tall) with

a canopy cover of 20%, consists of Chrysocoma coma-

aurea, Helichrysum patulum, R. laevigata, Chironia

baccifera, Stoebe cinerea, Myrsine africana, Selago

corymbosa, Lobostemon fruticosus, Felicia fruticosa,

Pelargonium cucullatum, Lobostemon argenteus, Passe-

rina filiformis, Gnidia laxa, Montinia caryophyllacea,

Helichrysum cymosum and Alhanasia trifurcata. The most

common grass species is Merxmuellera stricta with a

height up to 0.5 m and a cover of <1%.

The third stratum, the sparse herbaceous layer (<0.25

m tall), consists of the very sparse dwarf shrubs Hibiscus

aethiopicus and Gnidia inconspicua\ the forbs Knowl-

tonia capensis and Scabiosa columbaria ; ephemeral

species and ferns, and has a cover of 15%.

The total canopy cover for this variant is 60% which

increases by 10—20% during winter. Ephemeral species

listed in Species Groups D, I, J and K (Table 4) contribute

to this increase in cover. Apart from the ephemerals listed,

ferns such as Mohria caffrorum and Cheilanthes capensis,

also increase in cover during winter.

The high constancy and canopy cover values of many

species such as Cliffortia polygonifolia, Rhus tomentosa,

Olea europaea subsp. africana, Stoebe cinerea and

Myrsine africana are features of this subcommunity. Rhus

lucida is the most constant species (100%) with a canopy

cover of about 21% . Diagnostic species are listed in Species

Groups D (Tables 3 & 4) and include the perennial Gnidia

inconspicua, Erica baccans, Diosma hirsuta, Knowltonia

capensis and Euryops abrotanifolius.

Some of the species which were recorded in this variant

and nowhere else in the study area are the woody shrubs

Adenandra uniflora, Blaeria ericoides, Cliffortia hirta,

Clutia alatemoides, Diosma hirsuta, Erica hispidula, E.

mauritanica, E. plukenetii, Euryops abrotanifolius, Heli-

chrysum nudifolium, Knowltonia capensis and Selago

serrata.

This variant is characterised by the absence of the peren-

nial and ephemeral species in Species Groups A, B, C,

E and H (Tables 3 & 4).

Variant 1.2.2

This variant occurs in five narrow valleys along the

southeast-facing slope. These valleys occur on moderately

to highly organic-rich, well-drained soils which are deep

(>1.0 m), usually reddish in colour, apedal, with 20—40%

clay and 1—5% organic carbon in the topsoil. This is the

highest organic carbon content found in the study area.

Rocks are mostly of a smaller size (20—40 mm), though

larger rocks as well as bedrock were recorded. The slope

varies between 23° and 30°.

The total canopy cover varies between 50% (releve 44)

and 140% (releve 48) in summer. During summer between

5% and 15% of the surface is bare. In winter the canopy

cover increases, mainly due to the growth of the fern

Cheilanthes capensis (especially in releve 44) and an added

component in the form of ephemeral species as listed

FIGURE 10. -Variant 1.2.1. Rhus

lucida amongst a variety of

mid-high shrubs.

276

Bothalia 22,2 (1992)

in Species Groups E, H, J and K (Thble 4). The litter cover

is generally high (90%).

Four strata are recognised in this mid-high mid-dense

shrubland. The low tree layer with an open canopy cover

of 15% is represented by Noltea africana, Kiggelaria

africana and the shrub Maytenus heterophylla. The mid-

high mid-dense shrub layer (1.0-2. 0 m) has a canopy cover

of 60% and includes the shrubs Rhus lucida, R. tomentosa,

R. angustifolia, Otholobium hirtum, Cliffortia polygoni-

folia, Anthospermum spathulatum, Clutia pulchella and

the small tree Olea europaea subsp. africana. Low shrubs

and forbs occur in open places in the canopy. The low

sparse shrub layer (0.25—1.0 m), with a canopy cover of

about 10%, includes the woody Chrysocoma coma-aurea,

Helichrysum cymosum, Myrsine africana, Diospyros

glabra, Putterlickia pyracantha, Rhus laevigata and the

scrambler Helichrysum patulum. The latter varies in height

and can reach up to 1.5 m when supported by the shrubs

previously mentioned or by M. heterophylla, as is the case

in relevd 49.

Owing to the relatively dense shrub strata, the

herbaceous layer is sparse and poorly developed directly

under large shrubs such as Rhus angustifolia and Clutia

pulchella. This layer consists mainly of low-growing ferns

such as Cheilanthes capensis, C. hastata, Mohria caffro-

rum and Asplenium aethiopicum, with a cover generally

not higher than 15 % . In the case of Cheilanthes capensis

very high cover values, up to 75% (releve 44), were

recorded. Zantedeschia aethiopica was recorded in all the

valleys. Grasses are poorly represented and include low

cover values for Ehrharta calycina which was recorded

in three valleys. This species was not recorded on the open

slopes of this subcommunity. In the dry valley releve 44,

it was recorded under R. angustifolia. Other grass species

recorded are Pentaschistis aspera and Ehrharta erecta.

Shrubs in this variant generally have a higher cover than

in variant 1.2.1 and trees are more common and also have

a higher cover than in the previous variant. Species which

differentiate this variant on their characteristically higher

cover are Rhus angustifolia, R. tomentosa and Putterlickia

pyracantha. Two species which were not found anywhere

else in the study area, i.e. Clutia pulchella and Noltea

africana, have a cover of about 30% in releve 28. In re-

leves 44, 48 and 53, R. angustifolia is the most dominant

species with a cover which ranges from 30—55%, whereas

both R. tomentosa and Maytenus heterophylla have a cover

of 20% in releve 49.

The number of species recorded per releve varies from

41 to 59. An average of 33 perennial and 18 ephemeral

species per releve was recorded. This variant is charac-

terised by the presence of the perennial species in Species

Groups E, F, H, J and K and the absence of species in

Species Groups A, B, C, D, G and I (Table 3). The

presence of ephemeral species listed in Species Groups

E, H, J and K and the absence of species listed in Species

Groups A, B, C, D and I (Table 4) are characteristic of

this variant. Diagnostic species are listed in Species Group

E (Tables 3 & 4) and include the tree Noltea africana,

the forb Zantedeschia aethiopica, and the fern Asplenium

aethiopicum. Several species were recorded in this variant

only and nowhere else in the study area. The most

prominent are the woody Noltea africana and Clutia

pulchella and the fern Asplenium aethiopicum.

CONCLUSION

One community and two subcommunities with a total

of five variants were recognised. These are correlated with

differences mainly in aspect, climate and historic land use.

The ephemeral species confirm the fact that only two sub-

communities with their variants are recognised and this

corresponds with Werger’s (1974) statement that commu-

nities in floristically rich areas can be clearly characterised

floristically on the bases of floristic lists in which only

permanently recognisable species are entered. Le Roux

(no date), however, found in Namaqualand that most

ephemeral vegetation units occur dispersed among the

perennial plant- communities on the plains, with little

relationship between ephemeral vegetation units and

perennial communities, whereas releves with no ephemeral

species occur mainly on the rocky outcrops.

Variants 1.1.1 and 1.1.2, which occur on the hot, dry

northern slopes as well as in the dry ravines are dominated

by the grass Hyparrhenia hirta and stunted woody shrubs

of which Rhus lucida and R. glauca are the most

prominent. These two variants are related to the more

mesic variant 1.1.3 on the northwest-facing slopes and

ravines where H. hirta is less conspicuous and the woody

elements are more prominent. Table 5 shows an increase

in cover by R. lucida from 6% on the north-facing slopes

to 11% on the northwest-facing slopes. H. hirta shows a

decrease from 25% to 10% in these areas. The prominence

of H. hirta on the north-north-west-facing slopes in variant

1.1.3 as well as in a firebreak maintained by bush-cutting

shows the relationship among the three variants of this

subcommunity.

Rhus lucida shows an increase from 11% in variant 1.1.3

to 21% on the wetter open slopes and to about 13 % in the

valleys of subcommunity 1.2. A large number of other

mid-high shrubs was also recorded in these valleys. R.

lucida showed an increase in height from the drier to the

wetter parts. H. hirta is almost completely absent from

TABLE 5. — The mean height (H) and cover (C) of Rhus lucida, Hyparrhenia hirta, and the mid-high and low shrub strata of the five variants

distinguished on Signal Hill, Cape Town

Species/Stratum R. lucida H. hirta Mid-high Low

Variant H(m) C(%) H(m) C(%) H(m) C(%) H(m) C(%)

Bothalia 22,2 (1992)

277

subcommunity 1.2 and is replaced by the grass Merx-

muellera stricta on the open slopes, though the cover

values are much lower. Introduced grass species of

Mediterranean origin were recorded throughout the study

area, though only low cover values occur. The most

constant of these was the annual Brim maxima.

Mid-high shrubs (1—2 m) showed an increase from the

drier to the wetter parts throughout the five variants. Low

shrubs (0.25—1.0 m) also showed this tendency, though a

decrease was recorded in the wet valleys on the southeast-

facing slopes.

Variant 1.1.3 is a transitional area between the xeric

north-facing parts and wetter east and southeast-facing

parts. This is indicated by the presence of certain shrub

and forb species and the high local cover values for H.

hirta. Releves 39 and 40 occur on an east-north-east-facing

slope, which means that these slopes receive more solar

radiation than the other slopes in subcommunity 1.2. Its

relationship with variant 1.1.3 (subcommunity 1.1) shows

that the east-north-east-facing slopes of subcommunity 1.2

can with inadequate management be converted into the

more xeric variant 1.1.3.

Subcommunity 1.1 indicated that Elytropappus rhino-

cerotis can increase with an increase in fire frequency

and/or erosion (releve 33). With even higher fire

frequencies grassland may become dominant (Variants

1.1.1 and 1.1.2). Very few young E. rhinocerotis individuals

were recorded. These occurred mainly along the edge of

natural vegetation adjacent to the eucalypt firebreaks.

During winter an increase in total canopy cover is

experienced which is caused by ferns in the wetter valleys

and forbs, especially ephemeral species, along the drier

northern slopes. A mean of 26 ephemeral species per

releve was recorded for subcommunity 1.1, versus a mean

of 15 ephemeral species per releve for subcommunity 1.2.

The relative stability of the vegetation was indicated by

examination of aerial and historical photographs. There

appears to have been little permanent change in the

vegetation structure during the last century.

ACKNOWLEDGEMENTS

The study was partly funded by the South African

Nature Foundation. The Cape Town City Council granted

permission for work on Signal Hill. A special word of

thanks to the following people who assisted in various

ways: Pierre Nel (field work), Coert Smit (computer

work), Carol Wagner, staff of the Bolus Herbarium at UCT

and at the NBI at Stellenbosch (plant identification),

Dr Charlie Boucher (computer package), Dr Freddie Ellis

(soil map) and Liz Swart (editing).

REFERENCES

ADAMSON, R.S. 1929. The vegetation of the southwestern region. In

R.S. Adamson et al.. Botanical features of the southwest Cape

Province. Speciality Press, Cape Town & Wynberg.

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

Juta, Cape Town.

ASHTON, E.R. 1985. Cape Town Municipality: Table Mountain and

Silvermine Nature Reserves. In I.A.W. Macdonald, M.L. Jarman

& P. Beeston, Management of invasive alien plants in the Fynbos

Biome. South African National Scientific Programmes Report

No. 111. CS1R, Pretoria.

BOUCHER, C. & SHEPHERD, PA. 1988. Plant communities of the

Pella site. In M.L. Jarman, A description of the Fynbos Biome

Project intensive study site Pella, Ch. 4. Occasional Report 33.

CSIR, Pretoria.

BROWNLIE, S.F. 1982. The effects of recent land use on a fynbos site.

Final report. School of Environmental Studies, University of Cape

Town.

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

1. Longmans, London.

CAMPBELL, B.M., COWLING, R.M., BOND, W. & KRUGER, F.J.

1981. Structural characterization of vegetation in the Fynbos

Biome. South African National Scientific Programmes Report

No. 52. CSIR, Pretoria.

CAMPBELL, B.M. & MOLL, E.J. 1977. The forest communities of

Table Mountain, South Africa. Vegetatio 34: 105—115.

DEACON, H.J. 1983. The peopling of the Fynbos Region. In H.J.

Deacon, Q.B. Hendey & J.J.N. Lambrechts, Fynbos palaeo-

ecology: a preliminary synthesis. South African National Scientific

Programmes Report No. 75. CSIR, Pretoria.

FAGAN, G. 1989. The Company’s garden. Vol. 1. Municipality of Cape

Town. Unpublished report.

FUGGLE, R.F. 1981. Macro-climatic patterns within the Fynbos Biome.

National Programme for Environmental Sciences. Fynbos Biome

Project. CSIR, Pretoria.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK,

L. 1985. List of species of southern African plants. Edn 2,

Part 1. Memoirs of the Botanical Survey of South Africa No.

51.

GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., IMMEL-

MAN, K.L., GERMISHUIZEN, G., PIENAAR, B.J., VAN

WYK, M. & NICOLAS, A. 1987. List of species of southern

African plants. Edn 2, Part 2. Memoirs of the Botanical Survey

of South Africa No. 56.

HILL, M.O. 1979. TWINSPAN. A FORTRAN program for arranging

multivariate data in an ordered two-way table by classification

of the individuals and attributes. Ecology and Systematics, Cornell

University, Ithaca, New York.

JACKSON, W.P.U. 1977. Wild flowers of Table Mountain. Timmins, Cape

Town.

JOUBERT, C. 1991. History and description of contemporary vegeta-

tion, Signal Hill, Cape Town. M.Sc. thesis, University of Cape

Town.

KRUGER, F.J. 1978. A description of the Fynbos Biome Project. South

African National Scientific Programmes Report No. 28. CSIR,

Pretoria.

LE ROUX A. no date, ’n Fitososiologiese studie van die Hester Malan-

natuurreservaat. M.Sc. thesis. University of Pretoria.

LUCKHOFF, C.A. 1951. Table Mountain. Our national heritage after

three hundred years. Balkema, Cape Town.

McKENZIE, B., MOLl, E.J. & CAMPBELL, B.M. 1977. A phyto-

sociological study of Orange Kloof, Table Mountain, South

Africa. Vegetatio 34: 41—53.

MICHELL, M.R. 1922. Some observations on the effects of a bush fire

on the vegetation of Signal Hill. Transactions of the Royal Society

of South Africa 10: 213—232.

MOLL, E.J. & BOSSI, L. 1984. Assessment of the extent of the natural

vegetation of the Fynbos Biome of South Africa. South African

Journal of Science 80: 355—358.

MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and

methods of vegetation ecology. John Wiley, New York.

RAVEN-HART, R. 1967. Before Van Riebeeck. Callers at South Africa

from 1488 to 1652. Struik, Cape Town.

RAVEN-HART, R. 1971. Cape Good Hope 1652—1702. The first fifty

years of Dutch colonisation as seen by callers. 2 Volumes,

Balkema, Cape Town.

SCHULZE, B.R. 1972. South Africa. In J.F. Griffiths, Climates of Africa.

World survey of climatology. Vol. 10. Elsevier, Amsterdam.

SCHULZE, R.E. & McGEE, O.S. 1978. Climatic indices and classifi-

cations in relation to the biogeography of southern Africa. Junk,

The Hague.

SKEAD, C. J. 1980. Historical mammal incidence in the Cape Province.

Vol. 1. The western and northern Cape. Department of Nature

and Environmental Conservation, CPA.

SMITH-BAILLIE, A.L., RUDMAN, R.B., OOSTHUIZEN, A.B.,

ELLIS, F. & DOHSE, T.E. 1976. Soil survey of the Cape

Peninsula. Soil and Irrigation Research Institute, Department of

Agricultural Technical Services, Pretoria.

278

SPARRMAN, A. 1785. Voyage to the Cape of Good Hope. Vol. 1. London.

THERON, J.N. 1984. The geology of Cape Town and environs.

Department of Mineral and Energy Affairs, Pretoria.

THUNBERG, C.P. 1793-1795. Travels at the Cape of Good Hope

1772-1775. London.

VAN BLERK, E.F. 1990. A census ofgrysbok Raphicerus melanotis in

the Rondevlei Nature Reserve, Cape Town. Unpublished report.

Bolhalia 22,2 (1992)

Department of Biological Sciences: Nature Conservation, Cape

Technikon.

WEATHER BUREAU 1986. Climate of South Africa. Climate Statistics

up to 1984. WB 40, Pretoria.

WERGER, M.J.A. 1974. On concepts and techniques applied in the

Ziirich-Montpellier method of vegetation survey. Bolhalia 11:

309-323.

APPENDIX

List of the vascular plant species on Signal Hill, Cape Town. The following symbols are used: * introduced species; # recorded by Michell

(1922) after a fire; + sight record only; • isolated individuals; J, author’s collection number.

PTERIDOPHYTA

SCHIZAEACEAE

Mohria caffrorum (L.) Desv., J189

ADIANTACEAE

Cheilanthes Swartz

capensis (Thunb.) Swartz, J51

contracta Mett. ex Kuhn, J52

hastata (L. f.) Kunze, J53

ASPLENIACEAE

Asplenium aethiopicum (Burm. f. ) Becherer, J27

GYMNOSPERMAE

PINACEAE

Pinus L.

halepensis Mill.*+

pinaster Ait.* J254

pinea L.*+

radiata D. Don* J255

ANGIOSPERMAE— MONOCOTYLEDONAE

POACEAE

Cymbopogon marginatus (Steud.) Stapf ex Burn Davy, J77

Hyparrhenia hirta (L.) Stapf, J150

Heteropogon contortus (L.) Roem. & Schult., J145

Themeda triandra Forssk. , J320

Paspalum dilatatum Pair*, J414

Pennisetum setaceum (Forssk.) Chiov.* J246

Ehrharta Thunb.

calycina J.E. SM., J92

erecta Lam. , J93

melicoides Thunb., J94

villosa Schult. f, J435

Aira caryophyllea L* J7

Avena barbata Brot.* J31

Helictotrichon leoninum (Steud.) Schweick., Levyns s.n.

Merxmuellera stricta (Schrad.) Conert, J185

Pentaschistis Stapf

airoides (Nees) Stapf, J247

aspera (Thunb.) Stapf, J248

Gastridium phleoides (Nees & Meyen ) C.E. Hubb.*, J162

Stipagrostis zeyheri (Nees) De Winter, J312

Sporobolus africanus (Poir.) Robyns & Toumay-V

Eragrostis curvula (Schrad.) Nees, J96

Cynodon dactylon (L.) Pers. #

Cynosurus echinatus L.*, J78

Lophochloa cristata (L.) Hyl., J366

Melica racemosa Thunb., J183

Tribolium Desv.

ciliare (Stapf) Renvoize #

echinatum (Thunb.) Renvoize, J323

hispidum (Thunb.) Renvoize, J324

uniolae (L. f.) Renvoize, J325

Briza L.

maxima L.*, J39

minor L.*, J40

Festuca scabra Vahl #

Vulpia bromoides (L.) S.F. Gray* J335

Bromus L.

catharticus Vahl *#

diandrus Roth* J41

hordeaceus L. subsp. molliformis (J. Lloyd) Maire & Weiller* J43

pectinatus Thunb.*, J44

Brachypodium distachyon (L.) Beauv.*, J38

Lolium L.

multiflorum Lam. , J177

rigidum Gaudin* #

Hordeum murinum L.*, J436

CYPERACEAE

Cyperus tenellus L.f#

Ficinia Schrad.

bergiana Kunth, J364

filiformis (Lam.) Schrad., J118

indica (Lam.) Pfeiffer, J119

nigrescens (Schrad.) J. Raynal, J120, J381

ARACEAE

Zantedeschia aethiopica (L.) Spreng., J349

RESTIONACEAE

Restio multiflorus Spreng. , J437

Ischyrolepis Steud.

capensis (L.) Under, J156

triflora (Rottb.) Under, J438

COMMELINACEAE

Commelina africana L. , J66

JUNCACEAE

Juncus bufonius L* #

ANTHERICACEAE

Chlorophytum triflorum (Ait.) Kunth, Levyns s.n.

ASPARAGACEAE

Asparagus crispus Lam. , J196

Protasparagus Oberm.

africanus (Lam.) Oberm., J262

capensis (L.) Oberm., J263

rubicundus (Berg.) Oberm., J264, J372a

Myrsiphyllum undulatum (L. f.) Kunth, J197

ASPHODELACEAE/LILIACEAE

Bulbine Willd.

alooides (L. ) Willd. , J45

favosa (Thunb.) Roem. & Schult. #

tuberosa (Mill. ) Oberm. , J47

Trachyandra Kunth

chlamydophylla (Bak.) Oberm., Levyns s.n.

muricata (L. f.) Kunth, J322

COLCHICACEAE

Baeometra uniflora (Jacq.) G.J. Lewis, J34

Wurmbea spicata (Burm. f) Dur. & Schinz, J340

ERIOSPERMACEAE

Eriospermum Jacq. ex Willd.

Bothalia 22,2 (1992)

279

capense (L.) Salter, JlCf7

lancifolium Jacq. #

HYACINTHACEAE

Albuca L.

canadensis (L.) Leighton, J9

spiralis L. f, Michell s.n.

Urginea altissima (L. f.) Baker., J329

Tenicroa exuviata (Jacq.) Speta, J432

Dipcadi brevifolium (Thunb. ) Fourc. , J433

Omithogalum L.

gramini folium Thunb., J202

hispidum Homem. subsp. bergii (Schlechtd.) Oberm., J203

pilosum L. f, Michell s.n.

suaveolens Jacq. , Levyns s.n.

thyrsoides Jacq. , J204

Lachenalia Jacq. f. ex Murray

fistulosa Bak. , J163

orchioides (L.) Ait.

var. glaucina, J164a

var. orchioides, J164b

unifolia Jacq. #

HAEMODORACEAE

Wachendorfia paniculata Burm. , J336

ARYLLIDACEAE

Haemanthus sanguineus Jacq. , J130

HYPOXIDACEAE

Empodium plicatum (Thunb.) Garside, J434

Hypoxis sp. #

Spiloxene Salisb.

capensis (L.) Garside, J309

serrata (Thunb.) Garside, J417a

TECOPHILAEACEAE

Cyanella hyacinthoides L. , J76

IRIDACEAE

Romulea Maratti

flava (Lam.) De Vos var. flava, J281, J417

hirsuta (Eckl. ex Klatt) #

rosea (L.) Eckl. var. australis (Ewart) De Vos, J282

Galaxia ovata Thunb. #

Moraea Mill.

bellendenii (Sweet) N.E. Br., J192

ciliata (L. f. ) Ker-Gawl. , J193

gawleri Spreng. , J423

papilionacea (L. f. ) Ker-Gawl. #

tripetala (L. f. ) Ker-Gawl. #

vegeta L. , Levyns s.n.

Ferraria crispa Burm. , J117

Homeria Vent.

collina (Thunb.) Salisb. #

flaccida Sweet, J148

Bobartia indica L. #

Aristea africana (L. ) Hoffmg. , J19

Geissorhiza Ker-Gawl.

aspera Goldbl. , J122

imbricata (Delaroche) Ker-Gawl. var. bicolor (Thunb.) Goldbl., J123

inflexa (Delaroche) Ker-Gawl., J430

Hesperantha Ker-Gawl.

falcata (L. f. ) Ker-Gawl. , J144

spicata (Burm. f.) N.E. Br. , J431

Ixia L.

dubia Vent., J157

odorata Ker-Gawl. , J158

scillaris L. , J159

Sparaxis grandiflora (Delaroche) Ker-Gawl. subsp. fimbriata (Lam.)

Goldbl. , J308

Babiana Ker-Gawl.

disticha Ker-Gawl. , J32

stricta (Ait.) Ker-Gawl., J33

Gladiolus L.

brevifolius Jacq. var. brevifolius, J126

gracilis Jacq. #

Homoglossum priori i (N.E. Br.) N.E. Br. , J46, J408

Micranthus (Pers.) Eckl.

alopecuroides (L.) Rothm. , J187

tubulosus (Burm.) N.E. Br., J188

Lapeirousia corymbosa (L.) Ker-Gawl., J167

Watsonia marginata (L. f. ) Ker-Gawl. , J339

ORCHIDACEAE

Holothrix villosa Lindl. var. villosa, J147, J371

Satyrium Swartz

bicome (L.) Thunb. #

cdorum Sond. , J418

Disa tenuis Lindl. #

Monadenia bracteata (Swartz) Dur. <& Schinz, J190

Disperis Swartz

circumflexa (L.) Dur. & Schinz #

villosa (L. f.) Swartz #

Pterygodiuin Swartz

alatum (Thunb.) Swartz, J266

catholicum (L.) Swartz, J267

Corycium orobanchoides (Swartz) Schltr. , J68, J426

DICOTYLEDONAE

URTICACEAE

Didymodoxa capensis (Lf.) Friis & Wilmot-Dear var. integrifolia (Wedd.)

Friis & Wilmot-Dear, J84

PROTEACEAE

Protea L.

lepidocarpodendron (L.) L.», J428

repens (L.) L. •, J427

Leucadendron salignum Berg. , J170

Hakea suaveolens R. Br* #

LORANTHACEAE

Viscum capense L. f. , J334

SANTALACEAE

Colpoon compressum Berg. , J65

POLYGONACEAE

Rumex L.

acetosella L* #

cordatus Poiret, J283

Polygonum undulatum (L. ) Berg. , J411

CHENOPODIACEAE

Chenopodium ambrosioides L* #

Atriplex semibaccata R. Br. , J30

AIZOACEAE

Adenogramma glomerata (L. f. ) Druce, J5

Limeum africanum L. , Adamson 1858

Galenia pubescens (Eckl. & Zeyh.) Druce, J121, J384

Aizoon sarmentosum L. f. , J8

Tetragonia herbacea L. , J319, J379

MESEMBRYANTHEMACEAE

Delosperma asperulum (Salm-Dyck) L. Bol., Page s.n.

Drosanthemum striatum (Haw. ) Schwant. , J90

Erepsia N.E. Br.

anceps (Haw. ) Schwant. , J98

bracteata (Ait.) Schwant., J99

Lampranthus N.E. Br.

emarginatus (L.) N.E. Br., J165

multiseriatus (L. Bol. ) N. E. Br. , J166

Ruschia Schwant.

pulchella (Haw.) Schwant., J284

rubricaulis (Haw. ) L. Bol. , J285

CARYOPHYLLACEAE

Cerastium capense Sond., J50

Silene L.

burchellii Otth, J439

clandestina Jacq. , J302

ILLECEBRACEAE

Petrorhagia prolifera (L.) Ball & Heywood* J249

RANUNCULACEAE

Anemone tenuifolia (L. f. ) DC. , JU

Knowltonia capensis (L.) Huth, J161

280

Bothalia 22,2 (1992)

Ranunculus L.

multifidus Forssk. , J269

muricatus L. , J270

MENISPERMACEAE

Cissampelos capensis L. f. , J59

FUMARIACEAE

Phacocapnos c race a (Cham. & Schlechtd.) Bemh. , J251

Fumaria muralis Sond. ex Koch* #

BRASSICACEAE

Heliophila diffusa (Thunb.) DC. var. diffusa, J138

Lepidium africanum (Burm. f. ) DC. , J403

Raphanus raphanistrum L.*, J271

Rapistrum rugosum (L.) All.*, J272

DROSERACEAE

Drosera cistiflora L. , J91

CRASSULACEAE

Tyiecodon grandiflorus (Burm. f.) Toelken, J328

Crassula L.

brevifolia Harv. #

campestris (Eckl. & Zeyh.) Endl. ex Walp. subsp. campestris, J69

capensis (L.j Baill., J70

ciliata L. , J424

fascicularis Lam. , J71

muricata Thunb. #

nudicaulis L. , J72

saxifraga Harv. #

scabra L. , J73

subulata L. , J74

tetragona L. , Levyns s.n.

Adromischus hemisphaericus (L.) Lem., J6

MONT1NIACEAE

Montinia caryophyllacea Thunb. , J191

PITTOSPORACEAE

Pittosporum undulatum Vent.* J256

ROSACEAE

Rubus cf. cuneifolius Pursh.* J358

Cliffortia L.

hirta Burm. f. , J60

polygonifolia L. , J61

ruscifolia L. , J62

FABACEAE

Paraserianthus lophantha (Willd.) Nielsen subsp. lophantha* J227

Acacia Mill.

cyclops A. Cunn. ex G. Don* J1

karroo Hayne, J2

meamsii De Wild.*, J367

saligna (Labill.) Wendl.* J413

Virgilia oroboides (Berg.) Salter, J333

Podalyria Willd.

calyptrata Willd. , J258

sericea R. Br. , J259

Lotononis (DC.) Eckl. & Zeyh.

oxyptera (E. Mey.) Benth., Salter 7686

peduncularis Benth., J178

perplexa (E. Mey.) Eckl. & Zeyh., Penfold 6812

prostrata Benth. , J179

cf. umbellata Benth. , J440

Aspalathus L.

acuminata Lam. , J21

cephalotes Thunb. subsp. violacea Dahlg. , J22

chenopoda L. , J23

cordata (L.) Dahlg., J24

cymbiformis DC. , J25

ericifolia L. subsp. ericifolia, J26

hispida Thunb. subsp. hispida, J365

spinosa L. #

uniflora Dahlg. , Salter 6408

Melolobium aethiopicum (L.) Druce, JI84

Spartium junceum L.*+

Medicago polymorpha L. var. polymorpha, J182

Trifolium L.

angustifolium L. , J326

campestre Schreb.*#

glomeratum L* #

repens L* J327

Indigofera L.

cardiophylla Harv., J151

incana Thunb. , J152

psoraleoides L. , J153, J154

racemosa L. , J155

Psoralea imbricata (L. f. ) Thunb. , J442

Otholobium C.H. Stirton

decumbens (Ait. ) C.H. Stirton, J208, J385

fruticans (L.) C.H. Stirton, J441

hirtum (L.) C.H. Stirton, J209

uncinatum (Eckl. & Zeyh.) C.H. Stirton, J356

Tephrosia capensis (Jacq. ) Pers. , J318

Sutherlandia frutescens R. Br. , J316

Lessertia DC.

capensis (Berg.) Druce, J168

excisa DC. , J169

tomentosa DC. #

Vicia L.

benghalensis L * J351

sativa L.*, J443

Dolichos decumbens Thunb., J89

GERANIACEAE

Geranium L.

dissectum L* #

molle L* J124

purpureum Vill.* J125

Erodium L'Herit..

malachoides (L.) Willd. #

moschatum (L.) L'Herit. ex Ait.*, J108

Pelargonium L’Herit.

alchemilloides (L.) L’Herit., J230

auritum (L.) Willd. subsp. auritum, J231

candicans Spreng. , J232

capitatum (L.) L’Herit., J444

chamaedryfolium Jacq. , J233

columbinum Jacq. , J234

cucullatum (L.) L'Herit. subsp. tabulare Volschenk, J235

elongatum (Cav.) Salisb. , J236

grossularioides (L.) L’Herit. #

hirtum (Burm. f. ) Jacq. , J237

lobatum (Burm. f.) L'Herit., J239

longifolium (Burm. f. ) Jacq. , V.d. Walt 483

myrrhifolium (L.) L'Herit. var. myrrhifolium, J240

pillansii Salter, J241

pinnatum (L.) L’Herit., J242

proliferum (Burm. f. ) Steud. , J243

rapaceum (L.) L’Herit., J244

tabulare (L.) L’Herit., V.d. Walt 478

triste (L.) L’Herit., J245, J375

OXALIDACEAE

Oxalis L.

bifida Thunb., J212

compressa L. f. J213

flava L. , J214, J380

glabra Thunb., J215

hirta L. , J210, J216, J415

incarnata L. , J217

lanata L. f. , J225

livida Jacq. #

luteola Jacq. , J218

obtusa Jacq. , J219

pes-caprae L. , J220

polyphylla Jacq. , J221

punctata L. f, J222, J382

purpurea L. , J223

pusilla Jacq. , J224

tomentosa L. f, J226, J409

LINACEAE

Linum thesioides Bartl. , J173

ZYGOPHYLLACEAE

Zygophyllum L.

cf. fulvum L. , J445

sessilifolium L. , J343

Bothalia 22,2 (1992)

281

RUTACEAE

Adenandra uniflora (L.) Willd., J4

Diosma hirsuta L. , J86

POLYGALACEAE

Polygala L.

bracteolata L. , J260

myrtifolia L. , J261

Muraltia heisteria (L.) DC., J194

EUPHORBIACEAE

Clutia L.

alatemoides L. , J63

pulchella L. , J64

Euphorbia L.

arceuthobioides Boiss. , Jill

caput-medusae L. , JU2

crispa (Haw.) Sweet, JU3

genistoides Berg. #

ANACARDIACEAE

Schinus molle L*, J353

Rhus L.

angustifolia L. , J274

glauca Thunb. , J275

laevigata L. , J276

lucida L. , J277, J279, J402

rosmarinifolia Vahl, J278

tomentosa L. , J280

CELASTRACEAE

Maytenus heterophylla (Eckl. & Zeyh.) N.K.B. Robson, J181

Putterlickia pyracantha (L.) Szyszyl., J268

SAPINDACEAE

Dodonaea angustifolia L. f. , J88

RHAMNACEAE

Noltea africana (L.) Reichb. f, JI99

Phylica L.

imberbis Berg., J373

plumosa L. , J252

MALVACEAE

Hibiscus aethiopicus L. , J146

STERCULIACEAE

Hermannia L.

alnifolia L. , J140

althaeifolia L. , J141

cuneifolia Jacq. #

hyssopifolia L. , J142

prismatocarpa E. Mey. ex Harv. , J143

FLACOURTIACEAE

Kiggelaria africana L. , J160, J348, J350

THYMELAEACEAE

Gnidia L.

inconspicua Meisn. , J129

laxa (L. f. ) Gilg, J128

sericea L. , J446

Passerina L.

filiformis L. , J228

vulgaris Thoday, J229

MYRTACEAE

Eucalyptus

cladocalyx F. Muell.*, J461

ficifolia F. Muell.*, J458

gomphocephala DC.* J462

leucoxylon F. Muell.*, J460

tereticomis Smith* J459

ONAGRACEAE

Oenothera cf. noctuma Jacq.*, J447

APIACEAE

Centella L.

affinis (Eckl. & Zeyh.) Adamson, Levyns 2542

glabrata L. , J42, J407

Arctopus echinatus L. , J17

Torilis arvensis (Huds.) Link*, J32I

Lichtensteinia lacera Cham. & Schlechtd., J172

Annesorrhiza Cham. & Schlechtd.

capensis Cham. & Schlechtd., J12

hirsuta Eckl. & Zeyh., Levyns s.n.

Peucedanum galbanum (L.) Benth. <6 Hook, f, J250

ERICACEAE

Erica L.

baccans L. , JI01

hispidula L. , JI04

mauritanica L. , JI02

plukenetii L. , J105

strigosa Soland. , J448

Blaeria ericoides L. , J37, J406

MYRSINACEAE

Myrsine africana L. , J195

PRIMULACEAE

Anagallis arvensis L.*, J10

EBENACEAE

Euclea Murray

racemosa Murray, J109

tomentosa E. Mey. ex A. DC. , J110

Diospyros glabra (L.) De Winter, J87

OLEACEAE

Olea europaea L. subsp. africana (Mill.) P.S. Green, J200

GENTIANACEAE

Sebaea Soland. ex R. Br.

aurea (L. f. ) Roem. & Schult. , J289

exacoides (L.) Schinz, J290

Chironia baccifera L. , J54

ASCLEPIADACEAE

Microloma sp. #

Asclepias cancellata Burm. f. , J20

Cynanchum zeyheri Schltr., J75

Orbea variegata (L.) Haw., J201

BORAGINACEAE

Lobostemon Lehm.

argenteus (Berg.) Buek, J175

fruticosus (L.) Buek, J176

glaucophyllus (Jacq.) Buek, J449

Echium plantagineum L.*, J450

LAMIACEAE

Leonotis leonurus (L.) R. Br.+

Stachys L.

aethiopica L. , J310

arvensis L.*, J3U

Salvia L.

africana-caerulea L. , J286

africana-lutea L. , J287

SOLANACEAE

Lycium afrum L. , J451

Solanum L.

cf. americanum Mill. , J303

guineense L. , J452

hermannii Dun. #

nigrum L.*, J304

pseudocapsicum L.*, J305

SCROPHULARIACEAE

Diascia capensis (L.) Britten, J83

-Hemimeris montana L. f, J139

Nemesia Vent.

barbata (Thunb. ) Benth. , J198

parviflora Benth. #

Linaria spuria (L.) Mill.*, Kelmann s.n.

Antirrhinum orontium L * J16

Manulea cheiranthus (L. ) L. , J180

282

Bothalia 22,2 (1992)

Sutera Roth

antirrhinoides (L. f ) Hiem, J314

hispida (Thunb.) Druce, J315

Phyllopodium cordatum (Thunb .) Hilliard, J253

Zaluzianskya divaricata Whip. , J341

SELAGINACEAE

Hebenstretia repens Jarosz, J131

Selago L.

adpressa Choisy, J291

corymbosa L. , 729 2

fruticulosa Rolfe, J293

ramosissima Rolfe, Levyns 7090

serrata Berg. , J294

spuria L. , J453

BIGNONIACEAE

Teco maria capensis (Thunb.) Spach, J401

OROBANCHACEAE

Orobanche ramosa L.* J205

PLANTAGINACEAE

Plantago lanceolata L* J257

RUBIACEAE

Anthospermum L.

aethiopicum L. , J13

galioides Reichb. f. subsp. galioides, J14

spathulatum Spreng. subsp. spathulatum, J15

Nenax hirta (Cruse) Salter, Salter 6407

Galium sp., J347

VALERIANACEAE

Centranthus ruber (L.) DC.*, J332

DIPSACACEAE

Scabiosa columbaria L. , J288

CUCURBITACEAE

Zehneria scabra (L. f. ) Sond. subsp. scabra, J342

CAMPANULACEAE

Prismatocarpus sessilis Eckl. ex A. DC. #

Wahlenbergia Schrad. ex Roth

capensis (L.) A. DC., J337

obovata V. Brehm., J338

Microcodon cf. sparsiflorum A. DC. , J352

Cyphia Berg.

bulbosa (L.) Berg., J79

digitata (Thunb.) Willd., J80

incisa (Thunb.) Wild., Levyns 2883

phyteuma (L.) Wild., J81

volubilis (Burnt, f. ) W'illd. #

zeyheriana Presl ex Eckl. & Zeyh. , J82

Lobelia erinus L. , J174

Monopsis lutea (L.) Urb. #

ASTERACEAE

Pteronia hirsuta L. f, J454

Felicia Cass.

bergeriana (Spreng.) O. Hoffin., JU5

fruticosa (L.) Nicholson, JU6

tenella (L.) Nees #

Conyza Less.

canadensis (L.) Cronq.*, J67

scabrida DC. #

Chrysocoma coma-aurea L. , J57

Tarchonanthus camphoratus L. , J317

Gnaphalium subfalcatum Cabrera* J127

Troglophyton parvulum (Harv.) Hilliard & Bum#

Pseudognaphalium undulatum (L.) Hilliard & Bum, J265

Helichrysum Mill.

asperum (Thunb.) Hilliard & Bum

var. asperum, J132

var. glabrum Hilliard, J132a

cylindriflorum (L.) Hilliard & Bum, J133

cymosum (L. ) D. Don subsp. cymosum, J134

nudifolium (L.) Less., 7555, J405

patulum (L.) D. Don, J135

revolutum (Thunb). Less., J136

tereti folium (L.) D. Don, J137

S toe be L.

alopecuroides (Lam.) Less. #

cinerea Thunb., J313

Elytropappus rhinocerotis (L. f. ) Less. , J95

Metalasia muricata (L.) D. Don, J186

Relhania L’Herit. emend. Bremer

fruticosa (L.) Bremer, J273

genistifolia (L.) L’Herit., J372

Ley sera gnaphalodes (L.) L., J171

Eriocephalus africanus L. , J106

Athanasia L.

capitata (L.) L., Levyns s.n.

dentata (L.) L., J28

trifurcata (L.) L. , J29

Hymenolepis parviflora (L.) DC., J149

Cenia turbinata (L.) Pers., J49

Cineraria geifolia (L.) L., J58

Senecio L.

cf. abruptus Thunb. , J295

arenarius Thunb. ^ J296

burchellii DC., J297

hastatus L. , J298

pinifolius (L. ) Lam. , J299

pterophorus DC. , J300

pubigerus L. , J412

rigidus L. , J301

subcanescens (DC.) Compton, J363

Euryops abrotanifolius (L.) DC., JU4

Othonna arborescens L. , J211

Dimorphotheca pluvialis (L.) Moench, J85

Osteospermum L.

clandestinum (Less.) T. Norl., J206

spinosum L. , J207

Chrysanthemoides Toum. ex Medik.

incana (Burm. f.) T. Norl. , 755

monilifera (L.) T. Norl., J56

Ursinia Gaertn.

anthemoides (L.) Poir. subsp. anthemoides, 7550

dentata (L.) Poir., J331

Arctotheca calendula (L.) Levyns, J18, J421

Gorteria personata L. subsp. personata, J348

Gazania f>ectinata (Thunb.) Spreng., J457

Berkheya Ehrh.

armata (Vahl) Druce, J35

carlinifolia (DC.) Roessl. subsp. promontorii Roessl., J36

pinnatifida (Thunb.) Thell. Levyns s.n.

rigida (Thunb.) H. Bol. & Wolley-Dod ex Adamson & Salter, J455

Carduus pycnocephalus L.*, J48

Hypochoeris L.

glabra L* #

radicata L.*, J368

Urospermum picroides (L.) Scop, ex F.W. Schmidt *#

Picris echioides L.*, J456

Sonchus L.

asper (L.) Hill* J306

dregeanus DC. #

oleraceus L* J307

Bothalia 22,2: 283-288 (1992)

The recovery and dynamics of submerged aquatic macrophyte vegeta-

tion in the Wilderness lakes, southern Cape

P.J. WEISSER* A. K. WHITFIELD** and C.M. HALL***

Keywords: biomass, Characeae, fluctuations, macrophytes, monitoring, Najas marina, Potamogeton pectinatus, succession, vegetation dynamics,

Wilderness lakes

ABSTRACT

Between 1979 and 1981, the submerged aquatic macrophyte vegetation in the Wilderness lakes died back significantly,

and in some areas disappeared altogether. This study documents the senescent phase and describes the recovery of the plant

populations between May 1982 and May 1983. In two lakes, namely Langvlei and Eilandvlei, the plant biomass approximately

doubled between the winters of 1982 and 1983. Seasonal changes in species composition are documented and possible factors

accounting for the collapse and recovery of the plant populations are discussed.

UITTREKSEL

Gedurende 1979—1981, het die makrofitiese onderwaterplantegroei in die Wildemismere aansienlik teruggesterf en in sommige

gebiede heeltemal verdwyn. Hierdie opname beskryf die verouderende fase en die herstel van die plantbevolkings vanaf

Mei 1982 tot Mei 1983. In beide Langvlei en Eilandvlei het die plantbiomassa nagenoeg verdubbel tussen die winters van

1982 en 1983. Seisoenale veranderinge in die spesiesamestelling word beskryf en faktore wat moontlik verantwoordelik kon

wees vir die ondergaan en herstel van die plantbevolkings word bespreek.

INTRODUCTION

Several authors (Howard-Williams 1980; Jacot Guillar-

mod 1982; Weisser & Howard-Williams 1982) have studied

the aquatic macrophyte communities in the Wilderness

lakes. Their work, which was conducted prior to a major

die-back of aquatic plants throughout the lakes, provide

base-line data for describing vegetation changes arising

from a combination of seasonal and successional processes

(Miles 1979).

Between 1975 and 1979, submerged macrophytes were

an important biological component of the Wilderness

lakes, producing in excess of 750 t of dry plant mass per

annum (Howard-Williams 1980). The most important

species were Potamogeton pectinatus, Chara globularis,

Lamprothamnium papulosum and Ruppia cirrhosa. The

aquatic macrophyte population beds in this system

died back significantly between 1979 and 1981. This

study describes the senescent phase and documents

seasonal changes in species composition and biomass

which occurred during part of the recovery period

(1982-1983).

STUDY AREA

The Wilderness lakes comprise a segmented barrier

lagoon system lying parallel to the southern Cape coast.

There are three interconnected lakes (Rondevlei, Langvlei

and Eilandvlei) linked to the Wilderness Lagoon and the

Touws River by a narrow winding channel known as the

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

** J.L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown

6140.

*** Institute for Water Research, Rhodes University, Grahamstown 6140.

MS. received: 1992-05-19.

Serpentine (Figure 1). The Wilderness Lagoon is shallow,

varying in depth from 1 m in its lower reaches to 3.5 m

in die vicinity of the Serpentine entrance. The lakes are

all deeper than the lagoon, with a maximum depth of 6.5

m in Eilandvlei, 4.0 m in Langvlei and 6 f Rondevlei.

The whole lake system (91 km2) is situated on a series

of Quaternary sands known as the Wilderness-Knysna

embayment. Immediately inland of this embayment is a

200 m high Tertiary platform (foothills of the Outeniqua

Mountain range) consisting of folded sediments of the

Table Mountain Sandstone group, which forms part of the

catchment area of streams and rivers that feed the lakes

and Wilderness Lagoon. The lake system is estuarine in

nature and only periodically open to the sea at the Touws

River Mouth.

An important characteristic of the Wilderness lakes is

its reversed salinity gradient (Allanson & Whitfield 1983).

Rondevlei, which only receives fresh water from precipi-

tation and is the lake furthest from the sea, has the highest

salinity, whereas Eilandvlei, which is also influenced by

fresh water input from the Touws River has the lowest sa-

linity. Evaporation at Rondevlei (1 100—1 200 mm per an-

num) generally exceeds precipitation and this leads to

increasing salinity during dry periods. Heavy rainfall years

have the opposite effect. A series of floods in 1981 resulted

in a dilution of salt present in all three lakes. At Rondevlei

the salinity decreased from 16 g kg'1 to 10 g kg'1 between

September 1980 and September 1981, whereas over the

same period in Langvlei and Eilandvlei salinity decreased

from 10 g kg 1 to 5 g kg'1 and 9 g kg'1 to 4 g kg 1

respectively (Allanson & Whitfield 1983).

The waters throughout the system are generally well oxy-

genated, though low oxygen concentrations (< 2 mg 1 ') are

occasionally recorded in deeper areas of all three

lakes during the summer months (December— February).

284

Bothalia 22,2 (1992)

FIGURE 1. — Map of the Wilderness lakes showing localities mentioned in the text.

Surface water temperatures in the central region of the

lakes range from between 14°C in midwinter (July) to 26°C

in midsummer (January). The pH values range between

pH 7 and pH 9 despite the acidic (pH 5—6) nature of the

rivers and streams entering the system. Nutrient levels are

generally low, with total phosphorus and nitrate (No3N)

values seldom exceeding values of 100 ^tg/T1 and normal-

ly <40 fig l1. Chlorophyll a values are also generally

less than 15 fig l'1 though readings as high as 146 fig 1 1

have been recorded during phytoplankton blooms in

Langvlei (Allanson & Whitfield 1983).

MATERIALS AND METHODS

Aquatic macrophyte distributions in the Wilderness Lake

System prior to the die-back were mapped according to

a method described by Weisser & Howard-Williams (1982).

During the senescent phase (1979—1981), the degree of

aquatic macrophyte die-back in each part of the system

was visually monitored by boat at monthly intervals. The

recovery phase was more intensively monitored and

sampling commenced in May 1982. Six littoral transects

were positioned at approximately equidistant intervals

around Eilandvlei and five around Langvlei. The limits

of each transect were defined as the edge of the emergent

zone and the deepest point at which macrophyte growth

was recorded. A submerged macrophyte sampler de-

scribed by Howard-Williams & Longman (1976) was used

to sample a 0.0625 m2 area at variable intervals along the

length of each transect. Samples were returned to the

laboratory for sorting and drying. Results are expressed

as g m 2 dry weight. The total species biomass along

each transect was established by multiplying the mean bi-

omass for each plant species by the total transect length.

The above procedure was repeated in May, August and

November 1982, and in February and May 1983. In this

study, the period December— February is regarded as

summer, March— May as autumn, June— August as winter

and September-November as spring.

The standing biomasses of submerged macrophyte popu-

lations in Rondevlei and Wilderness Lagoon were too low

to sample effectively with the macrophyte collector. There-

fore only changes in species composition in these two

sections of the Wilderness lakes were recorded.

Past distribution patterns of aquatic macrophytes in the

Wilderness lakes were obtained from direct inspection of

historical aerial photographs using a Wild Aviopret Stereo-

scope (Weisser & Stadler 1983).

RESULTS

Eilandvlei

In 1975 most of Eilandvlei (Figure 2) supported a littoral

plant community dominated by Potamogeton pectinatus,

which occurred in water up to 3 m deep. At depths greater

than 3 m, no macrophytes were recorded. Zones of

Characeae and Ruppia cirrhosa occurred on the landward

side of the P. pectinatus zone (Figure 2).

Between 1976 and 1978, the P. pectinatus zone (Figure

3) receded by almost 40 m at the southeastern side of the

lake. This decline continued during 1979, so that by January

1980 no mature plant canopy was evident. A series of se-

vere floods in 1981 caused silt-laden waters (400—500 mg

dry mass 1 ') to enter Eilandvlei from the Duiwe River,

increasing water turbidity.

The recovery of submerged aquatic plant beds com-

menced during 1982 with Najas marina occupying large

areas previously colonized by P. pectinatus (Figures 2 &

4) . Changes in species composition were most apparent

in the northeastern section of Eilandvlei. In May 1982 the

mean dry biomass (six sampling transects around the lake)

of N. marina was 635 g m 2 (S.D. ± 1032), that of P

pectinatus 109 g m 2 (S.D. + 81) and Characeae 4 g m 2

(S.D. + 5). The average transect length in Eilandvlei

Bothalia 22,2 (1992)

285

FIGURE 2.— Distribution of submerged aquatic macrophytes in

Eilandvlei during 1975 (adapted from Weisser & Howard-Williams

1982). Dashed line = limit of study area; stippled areas =

Potamogeton pectinatus-, areas with vertical lines = Characeae;

oblique lines = Ruppia cirrhosa\ horizontal lines = Najas marina.

increased from 19 m in September 1982 to 27 m in March

1983.

Proportional contributions of different aquatic plant taxa

to biomass in different seasons during 1982 and 1983 are

shown in Figure 5. N. marina beds were a major compo-

nent of the aquatic plants in the winter of 1982 but during

the spring these beds started to recede and were replaced

by P. pectinatus. Characeae beds also showed signs of

recovery. By the summer of 1982/83 large amounts of

filamentous algae ( Enteromorpha sp.) started to grow both

epiphytically and epipsammically. The smothering effect

of this alga on other submerged plants resulted in their

reduced growth and disappearance from several areas of

the lake. Although this alga gradually diminished over the

rest of the study period, it did not disappear entirely.

Langvlei

In 1975 the entire bottom of Langvlei was covered by

dense Characeae beds (Figure 6) comprising Chara

globularis and Lamprothamnium papulosum. Both species

occurred throughout the lake, but the western region was

colonized almost exclusively by L. papulosum, and C.

globularis dominated the deeper central areas. These

extensive Characeae beds were still evident during April

1978, but by March 1979 had virtually disappeared. Aerial

photographs taken of Langvlei during December 1981 (Job

391, 1:10000) showed an absence of any aquatic macro-

phyte canopy or submerged Characeae beds. In 1975 the

lake bottom was clearly visible at 4 m. Thereafter, the

water transparency decreased as a result of a dinoflagel-

late bloom (Weisser & Howard-Williams 1982). The mean

Secchi disc value for the period January— December 1978

was 0.8 m (Coetzee & Palmer 1982).

During 1982, water transparency in Langvlei increased

again, resulting in a recovery of littoral submerged plant

beds but not those in deeper water (Figure 7). Five littoral

transects around the lake in May 1982 revealed a

Potamogeton pectinatus mean dry biomass of 750 g m 2

(S.D. ± 1138), Characeae 48 g m2 (S.D. ± 54) and

Najas marina 1 g m2 (S.D. + 2). A band of N. marina

was present along the northern shore of Langvlei during

the winter of 1982 but was replaced by P pectinatus in

the spring of that same year. The latter species contributed

more than 70% to the total submerged biomass in the lake

in all seasons (Figure 8).

The extensive charophyte meadows, which covered the

deeper areas of the lake prior to the vegetation die-back,

did not reappear. P pectinatus spread into these areas

during the summer of 1982/83, and by autumn 1983 this

species covered virtually the entire water surface area of

some regions of the lake. Filamentous algae were recorded

in the lake during the spring and summer of 1982/83, but

they did not proliferate as in Eilandvlei.

Rondevlei

In 1975, submerged macrophyte beds were absent in

Rondevlei, possibly due to the relatively high salinity (22

g kg ') prevailing at the time. During 1978 the submerged

FIGURE 3. — Photograph of Eiland-

vlei in March 1978 showing

bands of Potamogeton pectina-

tus (arrowed) surrounding both

Drommedaris Island and the

lake. These aquatic plant beds

had disappeared by January

1980.

286

Bothalia 22,2 (1992)

FIGURE 4. — Distribution of submerged aquatic macrophytes in Eiland-

vlei during 1982. Areas with vertical lines = Najas marina;

stippled = Potamogeton pectinatus.

vegetation of Rondevlei comprised a peripheral band of

the dominants Ruppia cirrhosa and P. pectinatus (Weisser

& Howard-Williams 1982) extending to a depth of 1.3 m.

In March 1979 the submerged macrophytes of Rondevlei

had diminished considerably compared to March 1978, and

the few scattered remaining plants did not reach the

surface. Aerial photographs taken during December 1981

(Job 391, No. 291/3) confirmed the macrophyte senescent

phase. This situation persisted until 1982 when evidence

of a recovery was recorded. In May 1982 sparse patches

of R. cirrhosa covered approximately 30% of the littoral

zone to a depth of about 1 m. By May 1983 these patches,

together with P. pectinatus, had expanded to cover approxi-

mately 90% of the littoral zone.

Winter Spring Summer Autumn Winter

FIGURE 5. — Seasonal changes in composition (pie chart) and biomass

(graph) of submerged aquatic plants in Eilandvlei during 1982

and 1983. Pie chart legend: vertical lines Potamogeton pectina-

tus; horizontal lines = Najas marina; black - Characeae; white

= Enteromorpha.

Wilderness Lagoon

In March 1978, the submerged plants in the Wilderness

Lagoon comprised Potamogeton pectinatus, Ruppia

cirrhosa, Zostera capensis and filamentous algae (Weisser

& Howard-Williams 1982). Isolated patches of P pectina-

tus were recorded growing in the Wilderness Lagoon

during 1982/83, and epipsammic Enteromorpha mats

proliferated during lagoon mouth closure. These mats

covered up to 60% of the water surface at such times, but

were dissipated and washed out to sea whenever the estuary

mouth opened. Z. capensis was not recorded from the

Wilderness Lagoon during 1982/83.

DISCUSSION

Aquatic macrophyte populations of brackish waters are

characterized by a low species diversity and vulnerability

to changes in environmental conditions, whether natural

or anthropogenically induced. Howard-Williams & Liptrot

(1980) for example suggested that if human activities cause

the decline of a macrophyte species in estuarine waters.

FIGURE 6. — Distribution of submerged aquatic macrophytes in Langvlei during 1975 (after Howard-Williams & T. Longman in Weisser & Howard-

Williams 1982). Vertical lines = Characeae; stippled = Potamogeton pectinatus.

Bothalia 22,2 (1992)

287

FIGURE 7. — Distribution of submerged aquatic macrophytes in Langvlei during 1982. Areas with vertical lines = Najas marina ; horizontal lines

= Potamogeton pectinatus', stippled = Ruppia cirrhosa.

there are generally few replacement taxa. This hypothesis

is supported by the current study which found that, during

the senescent phase, no new macrophyte species invaded

the previously occupied littoral zone. Recolonization

potential was limited to five species and contrasts to the

situation pertaining in most terrestrial environments e.g.

tropical forests.

The rapidity with which vegetation changes proceed in

brackish waters is noteworthy. In this study community

‘succession’ was most prevalent in Eilandvlei (Figure 5).

Nevertheless, in both Eilandvlei and Langvlei, seasonal

changes in mean standing crops followed a similar pattern

and were of the same order of magnitude. Mean biomass

per square metre approximately doubled between the

winter of 1982 and the winter of 1983 in both lakes.

However, the rate of expansion of plant beds towards

the centre of the lake in Langvlei was more rapid than in

Eilandvlei, a much deeper system. Average water trans-

parencies (Secchi disc) in Langvlei were 1.7 m and in

Eilandvlei 1.2 m over the period April 1982 to January

Winter Spring Summer Autumn Winter

FIGURE 8.— Seasonal changes in composition (pie charts) and biomass

(graph) of submerged aquatic plants in Langvlei during 1982 and

1983. Pie chart legend: vertical lines = Potamogeton pectinatus',

horizontal lines = Najas marina ; black = Characeae; white =

Enteromorpha.

1983, indicating better light penetration in Langvlei. In

nearby Swartvlei, it has been shown that the distribution

of Potamogeton pectinatus is closely associated with water

depth and degree of penetration of phytosynthetically active

radiation within the water column (Howard-Williams &

Allanson 1981).

Evidence from Langvlei suggests that aquatic vegeta-

tion proceeds through cycles; from an aquatic macrophyte

dominated system through a phytoplankton dominated

system and back to a macrophyte dominated system.

However, the dominant submerged plants at the beginning

of the study (Characeae) were not well represented at the

end of the study period, when P. pectinatus was dominant.

The cause(s) of most of the submerged macrophytes

dying in the Wilderness lakes has not been identified and

may well differ from one part of the system to another.

According to Weisser & Howard-Williams (1982) the high-

water level management policy, planktonic algal blooms,

development of periphyton on submerged plants and up-

rooting of plants by wave action during strong winds may

all have contributed to declines in submerged macrophytes

within the system. It is interesting to note that a similar

collapse in aquatic macrophytes was recorded from nearby

Swartvlei in 1979 (Davies 1982; Whitfield 1984), and 10

years later these Characeae and P. pectinatus beds had still

not fully recovered (A.K. Whitfield pers. obs.). In Natal,

Breen & Weisser (1986) recorded die disappearance of

previously abundant Potamogeton schweinfurthii and

P pectinatus in Lake Mzingazi, but could not identify any

definite factor(s) which caused the die-back. Breen &

Weisser (op. cit.) did, however, suggest that rapid changes

in lake water levels may have contributed to the collapse.

This investigation has shown that aquatic plant commu-

nities of the Wilderness lakes are highly variable, both on

a spatial and temporal scale, and that this dynamism

is due to a complex interplay of environmental and

288

Bothalia 22,2 (1992)

biotic factors. The study has also provided base-line in-

formation which will facilitate future monitoring of the

system. In conclusion we would like to recommend regular

vegetation mapping and aerial photographic coverage of

the study area as a valuable tool in the future management

of the Wilderness lakes.

ACKNOWLEDGEMENTS

We thank the Lake Areas Development Board, National

Parks Board and Cape Department of Nature & Environ-

mental Conservation for permission to work in areas under

their control. Technical assistance was provided by Mr

G.L. Christmas, Miss A.P. Backer and Mr K. Wagner.

Financial, logistical and infrastructural support was pro-

vided by the CSIR, Estuaries Joint Venture Programme

(FRD), Institute for Freshwater Studies, National Botan-

ical Institute and J.L.B. Smith Institute of Ichthyology. We

also thank Mr M.G. O’Callaghan and Mr D.J. McDonald

for commenting on the manuscript.

REFERENCES

ALLANSON, B.R. & WHITFIELD, A.K. 1983. The limnology of the

Touws River floodplain. South African National Scientific

Programmes Report No. 79.

BREEN, C.M. & WEISSER, P.J. 1986. Description and status of

the aquatic vegetation. In R.M. Walmsley & J.H. Grobler, An

evaluation of the impact of urban and recreational development

on the ecology and usage of Lake Mzingazi: 38—42. Foundation

for Research Development, Ecosystem Programme, Occasional

Report Series No. 6.

COETZEE, D.J. & PALMER, N.G. 1982. Algemene fisiese en chemiese

toestande in Eilandvlei, Langvlei en Rondevlei gedurende 1978.

Bontebok 2: 9—12.

DAVIES, B.R. 1982. Studies on the zoobenthos of some southern Cape

coastal lakes. Spatial and temporal changes in the benthos of

Swartvlei, South Africa, in relation to changes in the submerged

littoral macrophyte community. Journal of the Limnological

Society of Southern Africa 8: 33—45.

HOWARD-WILLLAMS, C. 1980. Aquatic macrophyte communities of

the Wilderness lakes: community structure and associated

environmental conditions. Journal of the Limnological Society

of Southern Africa 6: 85—92.

HOWARD-WILLLAMS, C. & ALLANSON, B.R. 1981. An integrated

study on littoral and pelagic primary production in a southern

African coastal lake. Archiv fUr Hydrobiologie 92: 507-534.

HOWARD-WILLLAMS, C. & LIPTROT, M.R.M. 1980. Submeiged mac-

rophyte communities in a brackish South African estuarine lake

system. Aquatic Botany 9: 101—106.

HOWARD-WILLLAMS, C. & LONGMAN, T.G. 1976. A quantitative

sampler for submerged aquatic macrophytes. Journal of the

Limnological Society of Southern Africa 2: 31—33.

JACOT GUILLARMOD, A. 1982. Checklist of the aquatic and flood-

plain vegetation of the Wilderness lakes, southern Cape. Bontebok

2: 41-51.

MILES, J. 1979. Vegetation dynamics. Outline studies in ecology.

Chapman & Hall, London.

WEISSER, P.J. & HOWARD-WILLLAMS, C. 1982. The vegetation of

the Wilderness lake system and the macrophyte encroachment

problem. Bontebok 2: 19—40.

WEISSER, P.J. & STADLER, A. 1983. Suitability of aerial photographs

for monitoring emergent and submerged macrophyte vegetation

in the Wilderness lakes, South Africa. Proceedings of the

International Symposium on Aquatic Macrophytes, Nijmegen,

Netherlands: 298—305.

WHITFIELD, A.K. 1984. The effects of prolonged aquatic macrophyte

senescence on the biology of the dominant fish species in a

southern African coastal lake. Estuarine, Coastal and Shelf

Science 18: 315—329.

Bothalia 22,2: 289-294 (1992)

Flora and vegetation of the Mbonambi Beach Arcuate Scar on the

Zululand dune barrier, Natal, South Africa

P.J. WEISSER* E.C.A. SMITH* A.P. BACKER* and S. VAN EEDEN*

Keywords: arcuate scar, dune valley, South Africa, vegetation, Zululand

Editor’s note: It is appreciated that this study is descriptive rather than quantitative but, in view of the unique vegetation of the arcuate scars and

the current spotlight on conservation in southern Zululand, it is felt that publication of this purely descriptive account is justified.

ABSTRACT

A hundred and seventy-nine species belonging to 63 families were found in 13 communities in the Mbonambi Arcuate

Scar. This is a rare type of dune valley that is described here and its vegetation is mapped. The great botanical diversity

in a small area (7 ha) can be related to the variety in ecological conditions and the juxtaposition of pristine and disturbed

areas. Because most of the adjacent area has been mined and this type of dune valley is rare, it deserves a high conservation

status rating. A Pereskia aculeata infestation was found that requires urgent attention.

UITTREKSEL

Honderd nege-en-sewentig spesies uit 63 families in 13 gemeenskappe is in ’n seldsame tipe duinvallei (arcuate scar) by

Mbonambi aangeteken. Die duinvallei word beskryf en die plantegroei word karteer. Die groot botaniese diversiteit oor ’n

klein gebied (7 ha) is die gevolg van die verskille in ekologiese toestande en die feit dat ongeskonde en versteurde gebiede

saam voorkom. Aangesien ’n groot gedeelte van die aangrensende gebied alreeds gedelf is, behoort ’n hoe bewaringstatus

aan hierdie duinvallei toegeken te word, veral omdat dit so seldsaam is. Die vervuiling van Pereskia aculeata het dringend

aandag nodig.

INTRODUCTION

During vegetation mapping work in May 1975 near the

Mbonambi Mission Station, 17 km north of Richards Bay,

a steep dune valley with densely vegetated slopes was

discovered (Figure 1). The valley bottom presented a

swamp, mainly colonized by reeds, sedges and grasses and

the slopes were covered with woody vegetation. The

absence of any apparent anthropogenic interference and

the lushness of the vegetation contrasted sharply with the

degraded, depauperate vegetation of the surroundings. This

motivated us to focus our research in this valley as our

mandate was to find areas of conservation value (Weisser

1978).

Venter (1972) and Weisser (1978) reviewed the dune

literature for the area between Richards Bay and St Lucia.

At that time no author had reported the presence of the

arcuate scars. The first reference to some of these dune

valleys and their location was published by Weisser (1978)

and the structures were termed ‘arcuate scar’ following

a suggestion by Hobday (pers. comm.). Vegetation map-

ping and fieldwork done by the senior author between Kosi

Bay and Tugela River showed that arcuate scars are rare,

only occurring in two coastal stretches, one north of

Richards Bay and one south. From the arcuate scars

visited, the Mbonambi arcuate scar was the one with the

densest vegetation. In 1983 the Mbonambi and other

arcuate scars were shown to K. Tinley. He reported on

their geomorphology (Tinley 1985). In his report he refers

to them as ‘cirques’.

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

MS. received: 1991-10-21.

The purpose of this work is to report on the discovery

and describe the Mbonambi Arcuate Scar, to make a

floristic inventory, to map the vegetation using aerial

photographs, and to describe the main plant communi-

ties.

MATERIALS AND METHODS

General reconnaissance and plant collecting were

undertaken, followed by a more detailed study of specific

areas. For the collecting of the specimens, standard

herbarium methods were employed. The facilities of the

National and Natal Herbaria of the National Botanical

Institute were available for the identification of specimens.

Plant names are mostly given in accordance with Gibbs

Russell et al. (1985, 1987).

Vegetation mapping was done from aerial photo No.

35234 (1985) supplied by the Air Survey Co. of Africa

Ltd. Photo interpretation was done through direct inspec-

tion, enlargement and transfer onto a base map using a

Bausch & Lomb ZT-4 Zoom Transfer Scope. The mapping

units were delimited according to features visible on

the aerial photo. The areas stratified were studied in

the field and the plant communities described in situ

structurally and floristically. Additional structural and

floristic information was obtained by doing a topo-

graphic profile using a measuring tape and an abney

level. The anthropogenic mapping units ‘ Casuarina

equisetifolia and ‘ Eucalyptus sp.’ were not studied.

Ground truth and field data were gathered from 1975 to

1987 (Weisser 1987).

Bothalia 22,2 (1992)

290

FIGURE 1. —Aerial view of the Mbo-

nambi Beach Arcuate Scar,

October, 1982. The central

Stenotaphrum secundatum—

Phragmites australis Reed-

swamp is clearly visible. The

seaward face is slumping owing

to undercutting by the sea.

STUDY AREA

The study area is situated on the dune barrier at

Mbonambi Beach, about 20 km northeast of Richards Bay

(28°4113" south and 32°13'20" east, Figure 2). The climate

is humid and warm to hot with a high year-round rainfall

(Schulze 1965), the mean annual temperature at the Cape

St Lucia Station being 21.5° C and the mean annual rain-

fall 1 292 mm. In this climate plant growth is luxuriant

and of a tropical nature (Aubert de la Rue et al. 1958, in

FIGURE 2. — Map showing the position of the arcuate scar (arrow).

Venter 1972). A climatic diagram can be consulted in

Weisser (1978).

RESULTS

Description of arcuate scars

Arcuate scars are valleys opening seaward (Figure 1)

caused by the erosive action of springs linked with

impermeable beds in the dune barrier. Water from rain

percolates through the dunes and accumulates on a clay

layer under the dunes. Because of the seaward dip of the

impermeable layer, the water is drained seaward causing

springs. These springs may cause slumping and back-

cutting, resulting in an arcuate valley with steep sides and

a swamp in the centre. The circular form offers protec-

tion against the strong sea winds. The high humidity and

steep valley sides are a good protection against fires and

diminish the hours of exposure to direct sunlight as well.

The semicircular form of the valley allows the existence

of north, south, east and west exposed slopes. There is

a marked soil-moisture gradient from the swampy bottom

to well-drained dune summit areas. For additional geomor-

phological information consult Tinley (1985).

Floristics

One hundred and seventy-nine species belonging to 63

families were recorded in the 7 ha area, which includes

some areas outside the scar. The most well-represented

family was Asteraceae (14 spp.), followed by Poaceae

(12 spp.), Cyperaceae (11 spp.), Rubiaceae (10 spp.),

Fabaceae (8 spp.) and Celastraceae (8 spp.). The checklist

is available on request from the National Botanical

Institute, Pretoria.

Main plant communities and their description

Fourteen mapping units with thirteen plant community

units were distinguished and their spatial distribution is

shown on the vegetation map (Figure 3) and on a profile

(Figure 4). The numbering of communities in the text,

Bothalia 22,2 (1992)

291

Beach and dune pioneers; 2, Cliff vegetation; 3, Dune Thicket

and Low Forest; 4, Dune Forest; 5, Trema orientalis Woodland;

6, Hygrophilous Forest; 7, Cyperus latifolius Marsh; 8,

Stenotaphrum secundatum-Phragmites australis Reedswamp; 9,

Secondary Grassland— Dwarf Shrubland Mosaic; 10, Acacia

karroo Woodland; 11, Secondary Dune Scrub and Forest; 12,

Casuarina equisetifolia Plantation; 13, Eucalyptus sp. Plantation:

14, Clearings.

the map and the profile is the same, e.g., areas marked

with 4 correspond to Dune Forest. The mapping units

distinguished were:

1. Beach and dune pioneers

2. Cliff vegetation

3. Dune Thicket and Low Forest

4. Dune Forest

5. Trema orientalis Woodland

6. Hygrophilous Forest

7. Cyperus latifolius Marsh

8. Stenotaphrum secundatum-Phragmites australis Reed-

swamp

9. Secondary Grassland— Dwarf Shrubland Mosaic

10. Acacia karroo Woodland

11. Secondary Dune Scrub and Forest

12. Casuarina equisetifolia Plantation

13. Eucalyptus sp. Plantation

14. Clearings

1. Beach and dune pioneers

In this mapping unit the plant cover is very low or non-

existent and, if present, concentrated at the landward zone

in the form of dune pioneer plants, including Ipomoea

pes-caprae, Sporobolus virginicus, Carpobrotus dimi-

diatus, and Gazania rigens var. uniflora. This community

has a low species diversity and is structurally simple. It

generally presents one layer of herbaceous plants, with an

average height of 0.10—0.65 m and a cover of 10-40%.

During our study this community was absent at times,

probably destroyed by high tides.

Where the streamlet flows onto the beach, a

hygrophilous community developed after 1985, with spe-

cies such as Phragmites australis, Stenotaphrum secun-

datum, Typha capensis, Hydrocotyle bonariensis and

Zantedeschia aethiopica. On drier substrates other spe-

cies such as Casuarina equisetifolia, Chrysanthemoides

monilifera, Panicum repens, Lobelia anceps, Senecio sp.

and Anthospermum littoreum were present. This area was

too small to be mapped separately and was included in

the beach mapping unit.

2. Cliff vegetation

Steep cliffs are present on the seaward side of the dune

barrier as a result of landslides caused by undercutting

by sea waves during storms and high tides. The slumped

material and debris accumulates on the base forming a

‘piedmont’. Most of this mapping unit is bare or covered

with remnants of slumped vegetation. Strelitzia nicolai is

often conspicuous. Depending on the stability of the sub-

strate and length of exposure, pioneer shrubs and herbs

establish themselves, such as Chrysanthemoides monilife-

ra, Ipomoea wightii, Carpobrotus dimidiatus, Cynanchum

obtusifolium, Anthospermum littoreum and Carissa macro-

11

FIGURE 4. — Schematic profile through the arcuate scar, October 1982, revealing the topography and the position of the communities found.

1, Beach and dune pioneers; 4, Dune Forest; 6, Hygrophilous Forest; 7, Cyperus latifolius Marsh; 8, Stenotaphrum secundatum-Phragmites

australis Reedswamp; 11, Secondary Dune Scrub and Forest.

292

Bothalia 22,2 (1992)

carpa. Some cliff areas have been planted with Casuarina

equisetifolia for erosion control (mapping unit 12).

Where cliffs are stable and the environment is shady,

Senecio sp., Samolus valerandi, Lobelia anceps, Pycreus

polystachyos and mosses establish themselves.

3. Dune Thicket and Low Forest

This mapping unit comprises the woody vegetation of

the southwest-facing side of the arcuate scar (Figure 1).

The vegetation is dense and its crown is pruned by onshore

winds. The limits with adjacent mapping units are not

always clear because transition is often progressive.

Thicket varies in height from 0.5— 5.0 m with a total

cover of 60— 100%. This community is generally formed

by a single layer of densely growing woody plants. Occa-

sionally herbs occur but then usually with low cover

values. Woody species common in this community are

Mimusops caffra, Strelitzia nicolai, Brachylaena discolor

and Chrysanthemoides monilifera. In the field layer

Microsorium scolopendrium was present, as well as

Crocosmia aurea. Climbers noted include Rhus nebulosa,

Rhoicissus digitata, Cynanchum obtusifolium and Ipomoea

wightii.

4. Dune Forest

This mapping unit is typical of most of the higher level

slopes of the arcuate scar, where there is no influence of

the water table (Figure 1). When exposed to sea winds,

the vegetation is stunted and it leads to coastal thicket.

The forest has been degraded through clearing on the out-

skirts and there it is replaced by secondary communities

such as Secondary or Grassland-Dwarf Shrubland Mosaic.

In its structure and species composition it is similar to

other Zululand dune forest on steep slopes. The height-

cover values estimated were 5—8 m /60% cover for the

canopy tress, 1-3 m /60% for the subcanopy layer, 1-3

m /10% and 0—1 m /60% for the field layer. Structurally

this forest is characterized by a few dominant trees forming

a dense canopy, a rather sparse intermediate layer and a

field layer that varies in its height and cover depending

on which species were able to establish themselves.

Canopy trees include Mimusops caffra, Apodytes dimi-

diata, Canthium inerme, Euclea natalensis, Strelitzia

nicolai, Allophylus natalensis, Dovyalis longispina, and

Psydrax obovata. In the subcanopy Tricalysia sonderiana,

Kraus sia floribunda, Psychotria capensis, Peddiea

africana and Phoenix reclinata are common and Deinbol-

lia oblongifolia, Carissa bispinosa and Pavetta revoluta

are occasionally found. In the field layer Commelina spp.,

the ferns Microsorium scolopendrium and Stenochlaena

tenuifolia, with herbs Drimiopsis maculata and Asy stasia

gangetica were recorded. Locally Strelitzia nicolai groups

become dominant. Climbers recorded were Protaspara-

gus falcatus, Pyrenacantha scandens, Smilax kraussiana

and Tylophora anomala.

5. Trema orientalis Woodland

The dominant species are Trema orientalis and Strelitzia

nicolai. In the subcanopy Canthium inerme, Psychotria

capensis, Ficus burtt-davyi and Brachylaena discolor are

present and Stenochlaena tenuifolia is dominant in the field

layer. This community was found as a patch in one area

and is a stage in the regeneration of the vegetation on a

landslide. Trema orientalis will probably soon be replaced

with species from the Dune Forest.

In the sampled area the canopy was 5-6 m high with

an estimated cover of 50%. The subcanopy was 3— 5 m

high with 25% cover and the field layer 0—1 m high

with 40% cover. Pammenter et al. (1985) reported Trema

orientalis regenerating after fire at the Mlalazi Nature

Reserve. No evidence of fire was found inside the arcuate

scar. During 1987, after this survey had been completed,

a landslide occurred in the scar. The regeneration of

vegetation on this landslide, on which Trema orientalis

was dominant, has been reported on by Weisser et al.

(1991).

6. Hygrophilous Forest

This forest occurs in patches growing between the cen-

tral area of the arcuate scar and its slopes (Figures 1 &

4). The forest presented the following height/cover charac-

teristics: trees, 3—12 m/50%; subcanopy trees and shrubs,

1—3 m/30%; tall herbs, 1—2 m/70% and field layer, 0—0.6

m/10%.

The canopy and subcanopy trees recorded are Ficus sur,

Rapanea melanophloeos, Bridelia micrantha, Syzygium

cordatum, Halleria lucida, Berchemia discolor, Maesa

lanceolata and Phoenix reclinata. In the field layer

Stenochlaena tenuifolia is often dominant. Other species

were Commelina spp, Aneilema schlechteri and

Phragmites australis. Climbers noted included Coccinia

palmata, Dioscorea sylvatica and Ipomoea conge sta.

1. Cyperus latifolius Marsh

This community occurs along the main water course of

the arcuate scar. It tends to occupy heavy, wet soil. The

clay content of these soils is high, and they have either

a high water table or surface water. Cyperus latifolius is

dominant in some areas forming monospecific stands.

Other species recorded, especially near the borders of the

mapping unit, were Stenotaphrum secundatum, Thelyp-

teris confluens and Phragmites australis. The height of C.

latifolius is about 1 m and the total cover 70—80%.

8. Stenotaphrum secundatum— Phragmites australis

Reedswamp

This unit consists of a dense grass mat with occasional

reeds and ferns (Figure 5). The height/cover estimates

obtained at the site studied were high layer, 2.0— 3.5 m

(P. australis stalks) 5% cover and grass layer, 0—0.8

m/90%.

Important species are the grass Stenotaphrum secunda-

tum, the reed Phragmites australis, Zantedeschia aethio-

pica, Stenochlaena tenuifolia and Smilax kraussiana. Also

present were Rhoicissus digitata, Rhus nebulosa, Ficus

burtt-davyi and Cyperus sp.

Bothalia 22.2 (1992)

293

FIGURE 5. —The central part of the

arcuate scar is colonized by

a dense grass cover main-

ly formed by Stenotaphrum

secundatum with scattered

Phragmites australis.

9. Secondary Grassland— Dwarf Shrubland Mosaic

At the western boundary of the arcuate scar the plant

cover is mainly secondary. The vegetation varies according

to successional stage. No cover-height estimates were taken

because the mosaic nature of this community makes such

data inadequate. Previously this area was used for grazing.

Fires were probably common before the Department of

Forestry took over the management of the area. The woo-

dy element is formed by pioneer trees and shrubs, such

as Dodonaea angustifolia, Eugenia capensis, Tricalysia

sonderiana, Phoenix reclinata, Bridelia cathartica,

Canthium inerme, Rhus nebulosa, Allophylus natalensis,

Enterospermum littorale, Diospyros villosa var. villosa,

Chrysanthemoides monilifera, Rhus nebulosa, Strychnos

spinosa, Maytenus sp., Scutia myrtina and Antidesma

venosum.

Occasional trees, e.g. Syzygium cordatum, Brachylae-

na discolor, Erythrina sp. and Carissa macrocarpa may

occur. The grasses Aristida junciformis subsp. junciformis

and Imperata cylindrica are common and the sedge

Cyperus obtusiflorus was recorded. In some areas the

shrubs Salacia kraussii, Helichrysum kraussii and Passe-

rina rigida are dominant. Climbers present include

Rhoicissus digitata, Smilax kraussiana, Senecio mikani-

oides, and Dalbergia armata.

10. Acacia karroo Woodland

This woodland represents a successional stage that has

developed from dune grassland. Some areas that were

grassland in 1937 are today covered with Acacia karroo

of about 5 m tall. The following height/cover values were

recorded: field layer, 0—0.3 m/50%; understorey, 0.3— 2.5

m/20%; tree layer, 2.5— 5.0 m/60%.

Apart from Acacia karroo the following woody species

were recorded: Brachylaena discolor, Eugenia capensis

and Rhus natalensis. Climbers present in this community

are Senecio mikanioides, Cynanchum natalensis,

Mimusops cajfra, Canthium inerme, Bridelia cathartica,

Rhus nebulosa, Phoenix reclinata, Tricalysia sonderiana,

Pavetta revoluta and Apodytes dimidiata. In the field layer

Commelina spp., cf. Brachiaria (Poaceae), Achyranthes

aspera, Laportea peduncularis , Asystasia gangetica and

Microsorium scolopendrium were noted.

11. Secondary Dune Scrub and Forest

This unit follows Communities 9 and 10 in the succes-

sion and, depending on the degree of development, it will

be a secondary scrub or forest. The physiognomy is very

variable. The values of the field layer were 0-0.75 m and

the shrub layer, 0.2— 6.0— 8.0 m. The field layer ranges

from 5—60% cover and the shrub layer from 60—90%.

The following species were encountered: Brachylaena

discolor, Eugenia capensis, Carissa macrocarpa,

Dodonaea angustifolia, Strelitzia nicolai, Rhus natalen-

sis, Rhus nebulosa, Apodytes dimidiata, Canthium inerme,

Kraussia floribunda, Scutia myrtina and Allophylus

natalensis. In the subcanopy Peddiea africana, Eugenia

capensis, Tricalysia sonderiana, Psychotria capensis and

Phoenix reclinata were present. The field layer was mainly

formed by Microsorium scolopendrium, Asystasia gange-

tica and Commelina spp. Climbers were Dioscorea

sylvatica, Dalbergia armata, Secamone alpinii, Cynan-

chum sp., Cissampelos torulosa, Pupalia cf. atropurpurea

and Rubia cordifolia.

12. Casuarina equisetifolia Plantation

Some areas were planted with C. equisetifolia to stabilize

erosion zones, mainly near the seaside. Some selfseeding

and colonization of areas by C. equisetifolia was observed.

13. Eucalyptus sp. Plantation

This community is artificial and was not studied.

14. Clearings

Vegetation clearing was done as part of the mining

exploration activities. These clearings are outside the

arcuate scar.

294

Bothalia 22,2 (1992)

ADDITIONAL OBSERVATIONS AND CONSERVATION

PRIORITIES

The structure of the arcuate scar with an impermeable

layer at its base causes a hydric gradient. There is also

a gradient in relation to exposure to sun and to saltspray-

bearing winds that increase the variety of ecological

conditions. These varied ecological conditions give rise

to the presence of a variety of communities with many

species, 179 species having being recorded on 7 ha.

Most of the vegetation in the arcuate scar is pristine,

as the steep landward slopes are not affected by the

previously frequent fires. Moreover, topography is too

steep to allow cultivation and therefore the area was not

cleared by the local inhabitants.

Arcuate scars should be conserved because 1, pristine

vegetation is rarely found in the area; 2, they are reservoirs

of species from where recolonization of dunes can take

place after mining (Camp & Weisser 1991); 3, they are

rare geomorphological features and (4) they have high

species and habitat diversities in a very small area. From

the scientific point of view we have here a model situa-

tion where the effect of parameters such as exposure, relief

and water factor can be preferentially studied. For these

reasons the Mbonambi Arcuate Scar deserves to be

conserved. The Richards Bay Minerals has therefore

agreed not to mine this area and to contribute to its

protection and conservation. One of the conservation

aspects that need urgent attention is the control of the

noxious weed Pereskia aculeata that was observed growing

in the arcuate scar.

ACKNOWLEDGEMENTS

Thanks are due to Dr J.C. Scheepers, for advice and

proofreading of the manuscript; to Messrs C. Buthelezi

and A. Ngwenya for their help during field work; to the

staff of the Herbarium of the National Botanical Institute

for plant identifications, especially Mrs M. Jordaan;

Mr A. Jeffrey from the Air Survey Co. of Africa Ltd. for

his help in obtaining aerial photographs; the Director of

Richards Bay Minerals, Mr R.D. MacPhersen, Mr J.

Goedhals and Mr P. Camp for taking protection measures

to keep the arcuate scar outside the mining area and for

aerial photos and Mrs A. Romanowski for the photo-

graphic work.

REFERENCES

CAMP, P. & WEISSER, P.J. 1991. Dune rehabilitation, flora and

plant succession after mining at Richards Bay, South Africa.

Proceedings of the Forest Biome Meeting, August 1990, Richards

Bay.

GIBBS RUSSELL, G.E. et al. 1985. List of species of southern African

plants. 2nd edn, part 1. Memoirs of the Botanical Survey of South

Africa No. 51.

GIBBS RUSSELL, G.E. etal. 1987. List of species of southern African

plants. 2nd edn, part 2. Memoirs of the Botanical Survey of South

Africa No. 56.

PAMMENTER, N.W., BERJAK, M. & MACDONALD, I.A.W. 1985.

Regeneration of a coastal dune forest following fire. South African

Journal of Botany 51: 453—459.

SCHULZE, B.R. 1965. General survey. Climate of South Africa Part

8. Weather Bureau Publications, Department of Transport,

Pretoria.

TINLEY, K.L. 1985. Coastal dunes of South Africa. South African

National Scientific Programmes Report No. 109. CSIR, Pretoria.

VENTER, H.J.T. 1972. Die plantekologie van Richardsbaai, Natal. D.Sc.

thesis. University of Pretoria.

WEISSER, P.J. 1978. A vegetation study of the Zululand Dune areas.

Conservation priorities in the dune area between Richards Bay

and Mfblozi River Mouth based on a vegetation survey. Natal Town

and Regional Planning Report Vol. 38, Pietermaritzburg.

WEISSER, P.J. 1987. Dune vegetation between Richards Bay and Mlalazi

Estuary and its conservation priorities in relation to dune min-

ing. Natal Town and Regional Planning Report Vol. 19.

WEISSER, P.J., BACKER, A.P., GLEN, H.F. & NGWENYA, A. 1991.

Recolonization of a landslide in the Kwambonambi Dune Area,

Natal, South Africa 1987—1990. Environmental Forum Report :

165-169. Foundation for Research Development. CSIR, Pretoria.

Bothalia 22,2: 295-300 (1992)

OBITUARIES

AMY FRANCES MAY GORDON JACOT GUILLARMOD, NEE HEAN (1911-1992)

During a working career that spanned 54 years Amy Jacot

Guillarmod (Figure 1) was a stalwart of botany in southern

Africa. Her collecting label duplicates, housed at GRA,

indicate 10241 collections, mainly from Lesotho, from

Port St Johns in the Transkei, from the Albany, Bathurst

and Victoria East Districts of the eastern Cape and from

George and Knysna in the southern Cape. They are housed

mainly at PRE, GRA, RUH and overseas at Kew (K) and

Missouri (MO). Her publications, of which there are

records of 198, range from the cornerstone Flora of

Lesotho which appeared in 1971, through research papers

on wetlands, ‘bogs and sponges’ and on cycads, tip

numerous popular articles showing her concern for

conservation and education. She is commemorated in the

names of the grass Merxmuellera guillarmodiae, the

diatoms Navicula jacotiae and Pinnularia guillarmodiae

and in several entomological names. She was also

honoured by numerous societies and by the Botanical

Research Institute, which dedicated Volume 50 part 1 of

The Flowering Plants of Africa to her. She particularly

appreciated its salutation ‘khotso’, meaning ‘peace be unto

you’ which acknowledged her bond with the Basotho. But

probably Amy’s most lasting contribution, and the one for

which she would most like to be remembered, was to

the knowledge of and enthusiasm for botany that she

engendered in students and colleagues alike.

Amy was bom at Hillcrest in Natal on the 23rd of May

1911. Her father was in the Royal Navy Reserve and she

spent much of her early childhood in Durban and Cape

Town. Until 1922, when she was 11 years old, she was

tutored at home. From 1923 until 1926 she attended

primary school, briefly in Durban and then in Scotland.

Her secondary schooling from 1926 to 1929 was at Durban

Girls High School where she matriculated. Amy returned

to Scotland to attend the University of St Andrews from

1930 to 1935. She first obtained her M.A. in English and

History but then established the basis for her career in

science by switching to Botany and Zoology and obtaining

her B.Sc.

On returning to South Africa in 1935, Amy taught briefly

in Durban before taking up an appointment as a plant

pathologist at the Pretoria headquarters of the Division

of Botany and Plant Pathology of the Union Department

of Agriculture. Her first five publications from this period

were on vims diseases of tobacco and of other crops — an

interest that continued long after she left the Department

in 1940. Although her early work was on viruses, she was

part of a team whose responsibilities ranged from the

taxonomy of fungi to that of higher plants, and they were

expected to give advice on any plant related subject. Thus

was nurtured a wide knowledge of things botanical

and agricultural, with personal interests as divergent as

conservation and economic botany.

Whilst in Pretoria, Amy met the entomologist ‘Chariot’

Charles Frederic Jacot Guillarmod, and they were

married in Bethlehem in the Orange Free State in 1940.

In the same year Chariot’s father died. Chariot and Amy

left their employment in science and went to help his

mother run the family store at ‘Mamathe’s in Basutoland

(now Lesotho). They were to spend the next 17 years,

a period crucial in career building, in the Mountain

Kingdom. To Amy, however, it was a golden opportunity

to collect and study the little known flora of the area. It

was by no means easy to concentrate on this task. She had

to learn both French and Sesotho. She taught at a girls

school and helped with the store. In 1949 young Francois

arrived. Scientific work and publications were reduced to

a trickle, but 1956/7 saw Amy’s return to academic

life — as lecturer and Head of the Botany Department of

the University College of Basutoland at Roma.

In 1958 the family moved to Grahamstown, Chariot to

join the staff of the Albany Museum, and Amy to lecture

in the Botany Department of Rhodes University. It was a

post that she was to hold for 15 years, until her retirement

as senior lecturer in 1973. During this period her list of

publications grew to 60, and she established her reputation

both as a teacher and as a researcher. She was rewarded in

1962/63 with the Mary E. Woolley Fellowship for overseas

study of the International Federation of University Women

and in 1967 with a D.Sc. in Botany from the University

of St Andrews for her studies on the Flora of Lesotho.

FIGURE 1. — Amy Frances May Gordon Jacot Guillarmod, n6e Hean

(1911-1992).

296

Bothalia 22,2 (1992)

Retirement, to Amy, meant new challenges: taking a

succession of new jobs and moving from the secure

comfort of her home in African Street to a 31 ha plot which

she was determined to clear of invading scrub pine, wattle

and stinkbean. Her first appointment was, as a Senior

CSIR Bursar, with the Institute for Freshwater Studies at

Rhodes University. This enabled her to continue her

interest in wetland plants, begun many years before in

Lesotho and developed through her long association

with the Limnological Society of Southern Africa. She

remained a Research Associate, retaining a working place

at Rhodes University, until the time of her death. In

addition to this position she joined the staff of the Botanical

Research Institute, in a half-day capacity as Research

Assistant in the Albany Museum Herbarium, from 1981

to 1987. Then, from 1987 to 1990, i.e. until she was 79

years old, Amy was part-time curator of the M. A. Pocock

Collection of Marine Algae at the Albany Museum.

She loved her plot ‘Faraway’, just outside Grahamstown

on the Highlands road which was later to be declared a

heritage site. When she and Chariot moved to the plot in

1974, a great number of pines grew on it and their

friends had a standing invitation for tea after work or on

weekends — but only after said friends had sawn down at

least one pine tree. She and Chariot also sawed down 2

or 3 trees daily after work before enjoying their coffee.

The system paid off and within three years the natural

vegetation started coming back. Amy has systematically

recorded every species she found on the plot from Fungi

to Asteraceae. The new Marasmius titanosporus described

by Derek Reid at Kew in 1988 was collected by Amy in

the little forest patch on ‘Faraway’. There was also a ‘magic

tree’ in the forest which produced Easter eggs, Christmas

gifts, and birthday treats for her little grandchildren whom

she adored.

Fires were always a threat and Amy and her staff

fought many over the years — not just on ‘Faraway’ but on

surrounding farms as well. They were all linked by citizen

band radio and kept a day and night vigil to spot the fires

early. Spoomet was the worst culprit and started many

a fire by ditching clinkers — Amy fought them with her

usual energy and I think the railway types involved must

have shaken in their boots whenever her name was men-

tioned!

When her husband. Chariot, died so unexpectedly and

suddenly in 1977 we all watched anxiously as she lost

centimetres in height — but that questing spirit did not let

her down. Her therapy was work. She returned to Rhodes

as an undergraduate student in Geography and at the end

of the year wiped the floor with all the bright young minds

pitted against her, coming first in class.

In searching for one word to describe Amy, we found

many: indomitable, indefatigable, intrepid, redoubtable —

all the names of battleships. Amy, the daughter of a

Commander in the Royal Navy, would have loved that, just

as she enjoyed sharing her birthday with Linnaeus, father

of taxonomic botany, for whom she gave a party every year.

The day-long Scottish breakfast party that co-incided with

Amy’s own 80th birthday was very special. It provided

a reminder, that behind Amy the teacher and researcher

was Amy the housewife — ever active and creative.

‘Active’ is also the word to describe Amy’s commitment,

both scientific and social. Her scientific societies included:

Botanical Society of South Africa (Life Member); Dendro-

logical Foundation (Publications Committee); Limnologi-

cal Society of Southern Africa (Honorary Life Member);

Linnean Society of London (Fellow); Royal Society of

South Africa; Lesotho Scientific Association (Vice

President); South African Archaeological Society (Life

Member); South African Association for the Advancement

of Science (Council Member); South African Association

of Botanists (Founder Member and Vice President); South

African Association of University Women (National

Fellowship Secretary and President of the Grahamstown

Branch); Systematic Association (Life Member).

She was also an avid supporter of local horticultural and

conservation groups, was active in church affairs, in the

Girl Guide Movement and in hockey circles, and received

the S.A. War Services Medal.

Amy was not only a producer, she was also an avid

critic. One always waited with bated breath to hear what

unforgivable flaw Amy had discovered in one’s latest work.

I have taken the liberty of slipping in one such (unbota-

nical) error of fact that Amy would undoubtedly have

found. I am sure that it would please her if you were to

do a little detective work on her behalf.

Amy died in Settlers Hospital, Grahamstown on the 7th

of May 1992. Shortly before her death she was planning

to continue working on her specimens in the Albany

Museum Herbarium.

Amy was much loved by her students who never returned

to Grahamstown without visiting her — some of them now

lecturers and professors themselves. Even students of re-

cent years knew her and their high regard showed in the

standing-room-only service on Tuesday when the little

Christ Church she attended regularly was simply packed.

Some flew to Grahamstown from Cape Town and Pieter-

maritzburg just for the service.

Botany is not the best paid of professions and it did not

enable Amy to fulfil all her ambitions, but in 1979 she

wrote to a colleague ... ‘isn’t it pleasant to be a botanist,

much nicer than any other profession?’

M.J. WELLS and E. BRINK

PUBLICATIONS OF A.F.M.G. JACOT GUILLARMOD

HEAN, A.F. 1938a. Diseases of tobacco. South African Department -1938c. The control of tobacco mosaic. Farming in South Africa, May.

of Agriculture. Bulletin 188: 45-50. -1940. Kromnek (spotted wilt virus disease) in South Africa. Farming

—1938b. Kromnek disease of flowers and vegetables (spotted wilt virus in South Africa, October.

disease). Farming in South Africa, April. —1941. Our Child the Earth. African Home Library Series, ICCLA

Bothalia 22,2 (1992)

297

No. 87. Sheldon Press, London.

—1943. Notes on maternal care in thrips. Journal of the Entomological

Society of South Africa 6: 81—83.

—1945. They threaten our food supply. South African Insect Life 1: 35-37.

—1947a. A wilt disease o/Crotalaria juncea (Sun Hemp). South African

Department of Agriculture. Science Bulletin 255.

—1947b. A South African virus disease of crucifers. South African Depart-

ment of Agriculture. Science Bulletin 254.

—1949. Yesterday and today in Basutoland. Health practices of the Basuto.

Health Horizon: 43—48. National Association for the Preven-

tion of Tuberculosis, Jan. London.

HEAN, A.F. & MOKHEHLE, C. 1947a. Some Basuto beliefs about wild

life. African Wildlife 1: 62—63.

—1947b. Some Basuto beliefs about wild life. African Wildlife 1: 68, 69.

—1947c. Some Basuto beliefs about wild life. African Wildlife 1: 80-82.

HORAK, I.G., JACOT GUILLARMOD, A., MOOLMAN, I.C. & DE

VOS, V. 1987. Parasites on domestic and wild animals in South

Africa. XXII. Ixodid ticks on domestic dogs and on wild

carnivores. Onderstepoort Journal of Veterinary Research 54:

573 -580.

JACOT GUILLARMOD, A. 71950. ‘Me, ke rata ho ba Lesupatsela. In

Sesotho for Girl Guides Association, Basutoland (now Lesotho).

—1958. Temperature variations in male cones of Encephalartos. Nature

182 : 474. London.

—1959a. Variations of temperature in male cones of Encephalartos

altensteinii. Journal of South African Botany 25: 93—99.

—1959b. The flora of Basutoland I. Ecological introduction. Basutoland

Notes and Records 1: 5—14.

—1959c. Aseroe in South Africa. South African Journal of Science 55:282.

—1960. The flora of Basutoland II . The flora of the dolerite dikes and

sills. Basutoland Notes and Records 2: 5—10.

—1961. The botany of Basutoland. Journal of the Botanical Society of

South Africa 47: 12—14.

—1962a. The botany of Basutoland: 29—34. ‘Science in General Edu-

cation’ Conference Handbook, Roma, Basutoland.

—1962b. Common wild flowers of Basutoland: 98—100. Year Book, British

Red Cross Society, Basutoland Branch, Maseru.

—1962c. The bogs and sponges of the Basutoland mountains. South

African Journal of Science 58: 179—182.

—1963a. Further observations on the bogs of the Basutoland mountains.

South African Journal of Science 58: 115—118.

—1963b. Missionaires botanistes au Bassoutoland. Musees de Geneve

38: 12, 13.

-1964a. The flora of Basutoland III. Water plants and waterside plants.

Basutoland Notes and Records 3: 7—15.

—1964b. Plant hunting in Europe. The Bluestocking 30: 32—35.

—1964c. Botany and bogs. Scottish Rock Garden Club Journal 9:

125-129.

-1964d. Wild animals forecast rain. African Wildlife 18: 256.

— 1964e. Habits of chamaeleons. African Wildlife 18: 256.

— 1964f. Interesting and rare plants in Basutoland. African Wildlife 18:

283-291.

—1965a. Notes on ‘Les plantes et l’ethnographie au Basutoland’. Annali

lateranensi 29: 397—414.

—1965b. A contribution towards the economic botany of Basutoland.

Botaniska Notiser 119: 209 —212.

—1966. Botanical exploration in Basutoland. Basutoland Notes and

Records 5: 22—31.

—1967a. Botanical exploration in Basutoland (abridged form of above

article). South African Journal of Science 63: 81—83.

—1967b. Phoenix reclinata — distribution. African Wildlife 20: 257.

-1968a. From the roof of southern Africa. African Wildlife 21: 304,

313-320.

—1968b. Selmar Schonland. Dictionary of South African Biography 1:

695, 696.

-1968c. Lesotho. In I.O. Hedberg, Conservation of vegetation in Afri-

ca south of the Sahara, edn 1. Acta Phytogeographica Suecica

54: 253-256.

-1969a. The effect of land usage on aquatic and semi-aquatic vegeta-

tion at high altitudes in southern Africa. Hydrobiologia 34: 3-13.

—1969b. Modem agricultural practice and limnology in Lesotho.

Limnological Society of southern Africa. Newsletter 13: 41—44;

—1970a. Notes on Aloe polyphylla. Basutoland Notes and Records 8:

30-36.

—1970b. Where does it come from? Veld & Flora 1: 8, 9.

—1970c. Two baboon tales. African Wildlife 24: 51, 52.

— 1970d. The Orange River headwaters. Limnological Society of Southern

Africa. Newsletter 15: 46—51.

— 1970e. Aquatic weeds in southern Africa. Probe (Science Students Coun-

cil Journal, Univ. Wits.): 47-49, 53.

-1970f. Review: Principles of dispersal in higher plants, by Van der Pijl.

Journal of South African Botany 36: 331.

—1971. The flora of Lesotho. Cramer, Lehre, Germany.

—1972a. The bogs and sponges of the Orange River catchment within

Lesotho. Civil Engineer in South Africa 14: 84, 85.

-1972b. The Caledon River in the early days of European settlement.

Civil Engineer in South Africa 14: 93, 94.

—1973a. Biography of Banks. Dictionary of South African Biography 2:

24, 25.

—1973b. Biography of Burchell. Botanical aspects. Dictionary of South

African Biography 2: 104—106.

—1973c. Biography of Schinz. Dictionary of South African Biography

2: 629, 630.

—1973d. Biography of Stoneman. Dictionary of South African Biography

2: 111, 718.

— 1973e. Biography of Trimen. Dictionary of South African Biography

2: 750, 751.

— 1973 f . Biography of Wahlberg. Dictionary of South African Biography

2: 829, 830.

— 1973g. Fish River vegetation. Veld & Flora 3: 47—49, 52.

—1974a. Plumbago and Tecomaria. Veld & Flora 4: 34—36.

—1974b. Vegetation: technical data report in environmental study of the

Swartkops River Basin, Vol. 3, Part 2. City of Port Elizabeth.

—1974c. Vegetation of the Grahamstown area. In J. Daniel, Grahams-

town and environs: 16—19.

— 1974d. Pelargoniums and geraniums. Veld & Flora 4: 43—45.

—1975a. Limnological bibliography for Africa south of the Sahara. Journal

of the Limnological Society of Southern Africa 1: 37—51.

—1975b. Review: History of British vegetation, by Pennington. South

African Geographical Journal 57: 150, 151.

—1975c. Point of no return. African Wildlife 29: 28—31.

—1976a. Use of odourless carrier, a petroleum product, in preparing

herbarium material. Taxon 25: 219—221.

—1976b. The floras of Lesotho and of the Orange Free State (Republic

of South Africa). Boissiera 24: 623, 624.

—1976c. Myriophyllum, an increasing water weed menace for South Afri-

ca. Proceedings of the South African Association for the Advance-

ment of Science , Nelspruit: 93—100.

— 1976d. Review: A treatise on limnology, Vol. in, Limnological Botany,

by G.E. Hutchinson. South African Journal of Science 72: 254,

255.

— 1976e. Letter re ‘Flora of Lesotho’. South African Archaeological So-

ciety Bulletin 31: 68, 69.

—1977a. Myriophyllum. Duplicated report to Inland Waters and Water

Research Commission.

—1977b. Myriophyllum, a powerful threat to our water supplies. Environ-

ment RSA 4: 3—5.

—1977c. Review: Aquatic weeds in South East Asia, by Varshney & Rzos-

ka. South African Journal of Science 73: 29.

— 1977d. Some waterweeds of the eastern Cape Province I. Eichhomia

crassipes. Eastern Cape Naturalist 60: 4—8.

— 1977e. Notes on various uses for gladioli in Lesotho, southern Africa.

Gladiolus Annual 1977 (Golden Jubilee year): 44, 45.

— 1977f. Myriophyllum, an increasing water weed menace for South

Africa. South African Journal of Science 73: 89, 90.

— 1977g. Biography of Bews. Dictionary of South African Biography 3:

63, 64.

— 1977h. Biography of Christol. Dictionary of South African Biography

3: 148.

— 1977i. Biography of Clarke. Dictionary of South African Biography 3:

154, 155.

— 1977j. Biography of Cooper. Dictionary of South African Biography

3: 175, 176.

—1977k. Biography of Dyke. Dictionary of South African Biography

3: 260, 261.

—19771. Biography of Ellenberger. Dictionary of South African Biography

3: 267, 268.

—1977m. Biography of Meisner. Dictionary of South African Biography

3: 596.

— 1977n. Biography of Sloley. Dictionary of South African Biography

3: 742.

— 1977o. Biography of Tyson. Dictionary of South African Biography

3 : 792 , 793.

— 1977p. Emersed water milfoil/waterduisendblaar, a new pest plant in

southern Africa. Nurserymen's Journal 13: 3—12.

— 1977q. Dr Mary Agard Pocock. The Bluestocking (Journal of the South

African Association of University Women) 31: 40.

— 1977r. Review: Making aquatic weeds useful, by Ruskin & Shipley.

South African Journal of Science 73: 157.

298

-1977s. Review: Making aquatic weeds useful, by Ruskin & Shipley.

Journal of the Limnological Society of Southern Africa 3: 8.

— 1977t. Some water weeds of the eastern Cape Province II. Myriophyl-

lum. Eastern Cape Naturalist 61: 14—17.

— 1977u. Letter to Editor, Piscator: Myriophyllum distribution. Piscator

99: 8.

— 1977v. Rhodohypoxis again! and notes on Aponogeton ranunculiflorus.

Veld & Horn 63: 21-23.

-1977w. Some water weeds of the eastern Cape Province m. Salvinia.

Eastern Cape Naturalist 62: 10—0.

—1978a. Lesotho. In P. Stegmann, Excursion Guide, SAAB/SAGP Tour,

1978 : 21-27. Bloemfontein SAAB Regional Centre.

—1978b. Some water weeds of the eastern Cape Province IV. Azolla.

Eastern Cape Naturalist 63: 16—18.

-1978c. Obituary: Dr M.A. Focock. The Bluestocking (Journal of the

South African Association of University Women): 43, 44.

-1978d. Can we come to terms with our water weeds? Proceedings of

the South African Association for the Advancement of Science :

409-416.

-1978e. Where did plant invaders come from?: the eastern Cape: 34,

35. Parrot’s Feather: 96-99. Kariba Weed: 02-05. In C.H.

Stirton, Plant Invaders — beautiful, but dangerous. Department

of Nature and Environmental Conservation of the Cape.

-1978f. Notes on distribution and biology of Aponogeton ranunculiflorus.

Kew Bulletin 32: 781-783.

— 1978g. Obituary: Mary Agard Pocock 1886—1977. Phycologia 17:

440-445.

—1979a. Oldenburgia— Rabbit’s ears. Eastern Cape Naturalist 66: 24, 25.

-1979b. Phytochemistry. Report at Stellenbosch SAAB Congress, January

1979. South African Journal of Science 75 : 428.

—1979c. Water weeds in southern Africa. Aquatic Botany 6: 377—391.

— 1979d. Myriophyllum in southern Africa. Proceedings of the third

National Weeds Conference of South Africa: 141—150. Balkema,

Cape Town & Rotterdam.

-1979e. Lesotho. In I. Hedberg, Systematic botany, plant utilisation

and biosphere conservation. Almqvist & Wiksell International,

Stockholm.

-1979f. Taxonomy and economics of water plants in Africa. Proceed-

ings of the IX Plenary Meeting of A.E.T.F.A.T., Las Palmas. 1978:

85-87.

— 1979g. Report to the Lakes Area Development Board on invasive weed

species in the area of the Wilderness lakes. Institute for Fresh-

water Studies Special Report No. 79/3.

—1980a. Exotic weeds and macchia vegetation in the eastern Cape.

Eastern Cape Naturalist 69: 21—23.

—1980b. Something from almost nothing. Eastern Cape Naturalist 24:

30, 31.

—1981a. Mistaken identity and false charges. The Naturalist 25: 11—13.

—1981b. Review: Succulent flora of southern Africa, by D. Court. The

Naturalist 25: 41, 42.

—1981c. A medicine chest from the veld: Basotho materia medica. Veld

& Mora 67: 113-115.

— 1981d. Biography of J. Hewitt. Dictionary of South African Biography

4: 232, 233.

— 1981e. Biography of Nikolaus Joseph Jacquin. Dictionary of South

African Biography 4: 256, 257.

—1982a. Review: Botanical exploration in southern Africa, by M.D. Gunn

& L.E. Codd. Eastern Province Herald, March 30.

—1982b. Porcupine and preservation. The Naturalist 25: 20—22.

—1982c. In my opinion. South African Weed Science Society Newsletter

No. 12: 1-3.

— 1982d. Checklist of the aquatic and floodplain vegetation of the Wilder-

ness lakes, southern Cape. Bontebok 2: 41-51.

— 1982e. A weed — yet beautiful and useful. The Naturalist 26: 17, 18.

— 1982f. (reed. 1983). A list of Sesotho plant names, with corresponding

scientific names. Ministry of Agriculture, Lesotho.

—1983a. A weed, yet beautiful and useful! The Rose Bulletin (Royal

National Rose Society), Autumn 1983: 22-24.

—1983b. Recovery of eastern Cape heathland after fire. Bothalia 14:

701-704.

—1983c. Cosmos, so very beautiful! The Naturalist 21: 3, 4.

—1984a. Special report — rare and endangered: 25—27. Albany Museum

Report Jan. 1981-March 31, 1983.

—1984b. Review: The ferns and fern allies of southern Africa, by W.B.G.

Jacobsen, 1983. The Naturalist 28: 40, 41.

—1984c. Coral trees (Erythrina species). The Naturalist 28: 7—9.

— 1984d. New series on local noxious weeds. Grocott’s Mail: 19. 25 Sept.

— 1984e. Noxious weeds of the eastern Cape: 1. Acacia cyclops. Grocott’s

Mail: 12. 16 Oct.

Bothalia 22,2 (1992)

-1984f. Noxious weeds of the eastern Cape: 2. Acacia longifolia.

Grocott’s Mail: 26. 26 Oct.

-1984g. Noxious weeds of the eastern Cape: 3. Acacia meamsii.

Grocott’s Mail: 6. 16 Nov.

-1984h. Noxious weeds of the eastern Cape: 4. Acacia saligna. Grocott’s

Mail: 10. 30 Nov.

— 1984i. Review: Wild flowers of Lesotho, by Marthe Schmitz, 1982.

Bothalia 15: 342, 343.

—1985a. Noxious weeds of the eastern Cape: 6. Pinus pinaster. Grocott’s

Mail: 14. 4 Jan.

-1985b. Noxious plants of the eastern Cape. 7. Albizia lophantha.

Grocott's Mail: 9. 1 March.

—1985c. Silk and spikes: 8. Hakea sericea. Grocott’s Mail: 2. 20 Sept.

— 1985d. Gums: the water thieves: 9. Eucalyptus species. Grocott’s Mail:

3. 8 October.

—1986. The vegetation of the Grahamstown area. In J.B.McI. Daniel,

W. Holleman & A. Jacot Guillarmod, Grahamstown and its

environs: 18—25. Albany Museum, Grahamstown.

-1987a. Amy remembers. Ichthos Newsletter No. 17: 16.

—1987b. That amazing woman: Mary Agard Pocock. The Elephant's

Child 10: 14-19.

—1987c. Obituary: Dr R.A. Dyer. The Elephant’s Child 10: 29.

— 1987d. Ethel Mary Doidge. Dictionary of South African Biography 5:

202, 203.

—1988a. Pot-pourri — sweetly scented mystery. The Naturalist 32: 39—41.

-1988b. Exotic plant invasion, man’s activities and indigenous vegeta-

tion. In N.M. Bruton & F.W. Gess, Towards an environmental

plan for the eastern Cape: 88, 99.

—1988c. Looking back on twenty-five years of our history. Journal of

the Limnological Society of Southern Africa 14: 4, 5.

— 1988d. Review: The Botany of the southern Natal Drakensberg, by O.M.

Hilliard & B.L. Burtt, 1987. Annals of Kirstenbosch Gardens 15;

Koedoe 31: 251-254.

— 1988e. Did you know? (Gyrrmosperms). The Elephant’s Child 5/6: 12.

—1989a. Phoenix reclinata Jacq. , the world’s most southerly palm. The

Phoenix (Magazine of the Albany Museum) 2: 7, 8.

—1989b. Fun with the Pocock collection of seaweeds. The Elephant’s

Child 12: 5, 6.

—1989c. Faraway. Conserva 4: 16, 17.

JACOT GUILLARMOD, A. & ALLANSON, B.R. 1978. Eradication

of water hyacinth. South African Journal of Science 74: 122.

JACOT GUILLARMOD, A. & CROSS, R. 1972. The problem of wax

on cycad leaves: a scanning electron microscope investigation.

Proceedings of the southern African Electron Microscopy Society

Johannesburg 1972: 34—36.

JACOT GUILLARMOD, A. & EVA, P. 1975. Limnological bibliography

for Africa south of the Sahara. Institute for Freshwater Studies,

Rhodes University, Grahamstown. Nov. 1975.

—1976a. Limnological bibliography for Africa south of the Sahara. Institute

for Freshwater Studies, Rhodes University, Grahams- town. Feb.

1976.

-1976b. Limnological bibliography for Africa south of the Sahara. Institute

for Freshwater Studies, Rhodes University, Grahams- town. May

1976.

—1976c. Limnological bibliography for Africa south of the Sahara. Institute

for Freshwater Studies, Rhodes University, Grahamstown. Aug. 1976.

— 1976d. Limnological bibliography for Africa south of the Sahara.

Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1976.

—1977a. Limnological bibliography for Africa south of the Sahara. Institute

for Freshwater Studies, Rhodes University, Grahams- town. Feb.

1977.

—1977b. Limnological bibliography for Africa south of the Sahara.

Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1977.

—1977c. Limnological bibliography for Africa south of the Sahara.

Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1977.

-1977d. Limnological bibliography for Africa south of the Sahara.

Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1977.

—1978a. Limnological bibliography for Africa south of the Sahara. Institute

for Freshwater Studies, Rhodes University, Grahams- town. Feb.

1978.

-1978b. Limnological bibliography for Africa south of the Sahara No.

11. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1978.

—1978c. Limnological bibliography for Africa south of the Sahara No.

12. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1978.

Bothalia 22,2 (1992)

299

-1978d. Limnological bibliography for Africa south of the Sahara No.

13. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1978.

—1979a. Limnological bibliography for Africa south of the Sahara No.

14. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Feb. 1979.

—1979b. Limnological bibliography for Africa south of the Sahara No.

15. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1979.

—1979c. Limnological bibliography for Africa south of the Sahara No.

16. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1979.

— 1979d. Limnological bibliography for Africa south of the Sahara No.

17. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1979.

-1980a. Limnological bibliography for Africa south of the Sahara No.

18. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Feb. 1980.

—1980b. Limnological bibliography for Africa south of the Sahara No.

19. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1980.

—1980c. Limnological bibliography for Africa south of the Sahara No.

20. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1980.

— 1980d. Limnological bibliography for Africa south of the Sahara No.

21. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1980.

—1981a. Limnological bibliography for Africa south of the Sahara No.

22. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Feb. 1981.

—1981b. Limnological bibliography for Africa south of the Sahara No.

23. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1981.

—1981c. Limnological bibliography for Africa south of the Sahara No.

24. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1981.

— 1981d. Limnological bibliography for Africa south of the Sahara No.

25. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1981.

—1982a. Limnological bibliography for Africa south of the Sahara No.

26. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Feb. 1982.

—1982b. Limnological bibliography for Africa south of the Sahara No.

27. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1982.

—1982c. Limnological bibliography for Africa south of the Sahara No.

28. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1982.

— 1982d. Limnological bibliography for Africa south of the Sahara No.

29. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1982.

—1983a. Limnological bibliography for Africa south of the Sahara No.

30. Institute for Freshwater Studies, Rhodes University, Grahams -

town. Feb. 1983.

—1983b. Supplementary bibliography to Limnological bibliography for

Africa south of the Sahara. Institute for Freshwater Studies,

Rhodes University, Grahamstown.

—1983c. Limnological bibliography for Africa south of the Sahara No.

31. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1983.

—1983d. Limnological bibliography for Africa south of the Sahara No.

32. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1983.

— 1983e. Limnological bibliography for Africa south of the Sahara No.

33. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1983.

-1984a. Limnological bibliography for Africa south of the Sahara No.

34. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Feb. 1984.

—1984b. Limnological bibliography for Africa south of the Sahara No.

35. Institute for Freshwater Studies, Rhodes University, Grahams-

town. May 1984.

—1984c. Limnological bibliography for Africa south of the Sahara No.

36. Institute for Freshwater Studies, Rhodes University, Grahams-

town. Aug. 1984.

—1985. Limnological bibliography for Africa south of the Sahara No. 38.

Institute for Freshwater Studies, Rhodes University, Grahams-

town. Nov. 1985.

JACOT GUILLARMOD, A., GETLIFFE, F. & MZAMANE, M. 1973.

On our borders (Sehlabathebe Wild Life Reserve and National

Park). Eastern Cape Naturalist 48: 9—12.

JACOT GUILLARMOD, A., JUBB, R.A. & SKEAD, C.J. 1979. Field

studies of six southern African species of Erythrina. Annals of

Missouri Botanical Garden 66: 521—527.

JACOT GUILLARMOD, A. & MARAIS, W. 1972. A new species of

Aponogeton (Aponogetonaceae). Kew Bulletin 21: 563-565.

JACOT GUILLARMOD, A. & RAUTENBACH, P. 1985. Limnological

bibliography for Africa south of the Sahara No. 37. Institute

for Freshwater Studies, Rhodes University, Grahamstown. Mar.

1985.

JESSOP, J.P. & JACOT GUILLARMOD, A. 1969. The vegetation of

the Thomas Baines Nature Reserve. Journal of South African

Botany 35: 367-392.

MOORE, E.S. & HEAN, A.F. 1938. Frenching in tobacco. Farming

in South Africa. June.

REID, D.A. & JACOT GUILLARMOD, A. 1988. Marasmius titanospo-

rus , a new species from the eastern Cape, South Africa. Trans-

actions of the British Mycological Society 91: 707-709.

RICHARDSON, G.R., LUBKE, R.A. & JACOT GUILLARMOD, A.

1984. Regeneration of grassy fynbos near Grahamstown (eastern

Cape) after fire. South African Journal of Botany 3: 153—162.

ROSE, E.F. & JACOT GUILLARMOD, A. 1974. Plants gathered as food-

stuffs by the Transkeian peoples. South African Medical Journal

48: 1688-1690.

LUCY KATHLEEN ARMITAGE CHIPPINDALL (1913-1992)

Bom in Pretoria on the 15th February 1913, Lucy

Chippindall (Figure 2) attended St Mary’s Diocesan

School, Pretoria, from 1919 to 1929. During this time she

developed her interest in biology, in particular bird-

watching and botany.

Initially employed as a shop assistant in Pretoria, she

soon moved to the Division of Botany in 1931, where she

was employed as an herbarium technician mounting grass

specimens. With the assistance of Dr Dyer, Lucy

Chippindall learned to use taxonomic keys to identify

grasses. Much of her time was subsequently spent

answering queries concerning the identity of pasture

grasses. Perceiving a need for a guide to the grasses of

southern Africa, she began planning a taxonomic treatment

which, when finally published in 1955, was to see service

for the next 35 years.

During the Second World War the demands placed on

the National Herbarium in Pretoria increased, as botanists

in central and southern Africa could not send material to

the European herbaria for identification. As a result, Lucy

identified a large number of African grass specimens and

greatly extended her knowledge of the grasses. Because

of this work, she was able to identify and describe several

new grass species (Chippindall 1946a). In addition to these

taxonomic descriptions, she also published a paper on seed

dispersal in the grasses (1946b).

In 1946, Lucy Chippindall left the Division of Botany

and spent three years at the University of the Witwaters-

rand, graduating in 1948 with a B.Sc. in botany.

After marrying Paddy Crook, she moved to Melsetter,

in what is now Zimbabwe, where Paddy was employed

300

Bothalia 22,2 (1992)

FIGURE 2.— Lucy Kathleen Armitage Chippindall (1913-1992)

as an agricultural extension officer. It was in Melsetter

that she continued her work towards a treatise on southern

African grasses. This, her major work, was the culmina-

tion of her efforts initiated in the 1940’s. The text, titled

A guide to the identification of grasses in South Africa,

was formalised at the request of the Board of Trustees of

the book The grasses and pastures of South Africa, edited

by D. Meredith.

This book has been described as a classical contribution

to the literature on pasture science in the subcontinent.

Lucy Chippindall’s contribution formed the major com-

ponent of this important reference work, incorporating

taxonomic keys to genera and species, descriptions,

illustrations by (among others) Cythna Letty and Gertrude

PUBLICATIONS OF

CHIPPINDALL, L.K.A. 1943. Rhynchelytrum repens. The Flowering

Plants of Africa 23: t. 914.

—1944. Andropogon amplectens. The Flowering Plants of Africa 24: t.

922.

—1946a. Contributions to the grass flora of Africa. I. Three new grasses

from tropical Africa and Panicum lanipes in South and South

West Africa. Blumea Supplement 3: 25—33.

—1946b. Contributions to the grass flora of Africa: II. A brief survey

:>f some lesser known dispersal mechanisms in African grasses.

Laurence, and distribution maps based on the data

gathered by John Acocks. Despite other texts that have

appeared subsequently, this work will remain in use by

farmers, researchers and conservationists for many years

to come. It has also become a valuable piece of Africana

and is sought after by book collectors.

During the 1950’s and 1960’s, the Crooks lived in

Salisbury (now Harare) and Urn tali (now Mutare). During

this time Lucy was employed in the Government

Herbarium in Salisbury. There, with the help of her

husband, she began to prepare other manuscripts on

southern African grasses. These were published as three

volumes of pamphlets, and dealt with 240 southern African

grasses (Chippindall & Crook 1976-1978). These pam-

phlets were intended to provide the layman with a means

to identify the more common grass species. Each pamphlet

deals with a single species, providing illustrations, distribu-

tion data (in Africa south of the equator) and habitat data.

In addition, common names, descriptions and notes on

the economic value are also provided.

Around 1978 the Crooks moved to Cape Town, where

Lucy continued her work on the grasses in the Bolus

Herbarium of the University of Cape Town. During this

period, her interaction with students and staff stimulated

an interest in the unusual grass flora of the fynbos, a

subject that has continued to receive the attention of the

researchers at the University of Cape Town.

During her career she collected more than 2 000

herbarium specimens, the majority of which are housed

in SRGH, PRE and BOL. In January 1989 she was

awarded the Senior Medal for Botany of the South African

Association of Botanists (SAAB). In 1990 she was made

an honorary member of the Grassland Society of southern

Africa.

She is commemorated in the species Pentaschistis

chippindalliae Linder (Linder & Ellis 1990), and the latest

work on southern African grasses (Gibbs Russell et al.

1990) is dedicated to her in recognition of her efforts in

the field of grass systematics.

Her passing on the 4th April 1992, marks the end of

an era, an era in which Lucy Chippindall provided a solid,

enduring foundation in what were then the young and

exciting fields of pasture science and grass taxonomy.

LUCY CHIPPINDALL

Blumea Supplement 3: 33—41.

—1946c. The common names of grasses in South Africa. Department

of Agriculture Pamphlet No. 265. (Botany and Plant Pathology

Series No. 7). Government Printer, Pretoria.

-1955. A guide to the identification of grasses in South Africa. In D.

Meredith, The grasses and pastures of South Africa. Part I. CNA,

Cape Town.

CHIPPINDALL, L.K.A. & CROOK, A.O. 1976-1978. 240 Grasses of

southern Africa. Collins, Salisbury.

REFERENCES

LINDER, H.P. & ELLIS, R.P. 1990. A revision of Pentaschistis.

Contributions from the Bolus Herbarium No. 12.

GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK,

L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical Survey

of South Africa No. 58.

N.P. BARKER*

* Botany Department, University of Cape Town, Private Bag, Ronde-

bosch. Cape Town 7700.

Bothalia 22,2: 301-312 (1992)

NATIONAL BOTANICAL INSTITUTE

SOUTH AFRICA

Scientific, Technical and Administrative Staff

(31st March 1992)

CHIEF DIRECTORATE

CAPE TOWN

Huntley, Prof. B.J. M.Sc. Chief Director

Comelissen, Mrs A.M. Secretary to the Board

McLean, Mrs S.E. Secretary. Nat. Higher Seer. Cert.

ADMINISTRATION DIRECTORATE

CAPE TOWN

Jordaan, A.W. M.B.A. Director: Administration

Hughes, W.S.G. Head: Finance

Jansen van Vuuren, J.G. Head: Administration (Pretoria) ■»

Van Zyl, J.M. Head: Personnel & Administration

ADMINISTRATION PRETORIA

Jansen van Vuuren, J.G. Provisioning Administration Officer

Albertse, Miss L. B.A. Assistant Provisioning

Administration Officer

Britz, Mrs E.P Telephonist

Joubert, Mrs I.J.M. Senior Provisioning Administration

Clerk

Kama, Mrs P.B. General Assistant I

Kekana, Mrs M.R. General Assistant I

Kgaditsi, W.T. Photocopy Machinist

Koehne, Mrs R.W.R. Registration Clerk

Letebele, M.M. General Assistant I

Makgobola, Mrs M.R. General Assistant I

Maphuta, Mrs M.S. General Assistant I

Nkosi, Mrs P.M. General Assistant I

Phaala, M.C. General Assistant I

Phiri, Mrs E. General Assistant I

Prinsloo, Mrs M.M. Senior Provisioning Administration

Clerk

Smuts, Ms W.E. Assistant Provisioning Administration

Officer

Thiart, Mrs S.P. Dip. (Typing). Senior Typist

Tloubatla, J.M. Courier

Venter, W.A. N2 (Technical). Supervisor: Cleaning

Services

FINANCE— CAPE TOWN

Hughes, W.S.G. Senior Accountant

Armitage, Mrs C.S. Accountant

Johnson, J.M. Salaries Clerk

Kriel, Ms C.M.M. Accounts Clerk

Neuwirth, Ms E.V. Salaries Accountant

October, A. Accounts Clerk

Petersen, R.E. Salaries Clerk

Simon, Ms P.I. Assistant Accountant

Traut, G.D. Salaries Clerk

Yeoman, Ms I.N. Senior Accounts Clerk

302

Bothalia 22,2 (1992)

PERSONNEL & ADMINISTRATION -CAPE TOWN

Van Zyl, J.M. Assistant Director

Barnard, Mrs A. Chief Telephonist

Blanche, J.G. Principal Personnel Officer

Britz, Mrs M.J. Typist

Dollie, Mrs N.J. Cleaner

Haupt, Mrs C.S. Cleaner

Mirkin, Mrs Y.A. Typist

Mulder, Mrs G.P. Typist

Nicholas, Mrs W.L. Photocopier

Schutte, Mrs L.A.J. Senior Registration Clerk

Smith, Miss L.G. Chief Typist

Solomons, Mrs. H.J. Receptionist/Typist

Staal, P.B. Senior Personnel Practitioner

Theron, R. Courier

Van Heerden, L.S. Personnel Officer

EDUCATION & INFORMATION DIRECTORATE

CAPE TOWN

Low, A.B. M.Sc. Director: Education & Information (from Aug. 1991)

Davidson, D.C. Head: Graphic Services

Leistner, O.A. Head: Publications (Pretoria)

Potgieter, Mrs E. Head: Mary Gunn Library (Pretoria)

Reynolds, Ms P.Y. Head: Kirstenbosch Library

Ross, Ms S.I. M.Ed. Head: Communication and Promotions

Wells, M.J. Head: Information (Pretoria)

GRAPHIC SERVICES -CAPE TOWN

Davidson, D.C. B.A. Hons, PRISA Final Dipl. Chief Liaison Officer

Loedolff, Mrs J. B.Sc. Industrial Technician. Photographer/Illustrator

INFORM ATION -PRETORIA

Wells, M.J. M.Sc. Assistant Director. Weeds research, botanical horticulture, fynbos utilization

and conservation.

Fourie, Mrs D.M.C. B.Sc. Scientific Information Officer

Joffe, Mrs H. B.Sc. Garden Utilization Officer

LIBRARIES

Potgieter, Mrs E. B.Bibl. Senior Librarian (Mary Gunn Library, Pretoria)

Lategan, Mrs B.F. Library Assistant

Reynolds, Ms P.Y. B.A., H.L.D.S., B.Proc. Senior Librarian (Kirstenbosch Library, Cape Town)

PUBLICATIONS -PRETORIA

Leistner, O.A. D.Sc., F.L.S. Assistant Director. Editing

Brink, Mrs S.S. Dip. (Typing). Chief Typesetter. Typesetting, lay-out, computers

Condy, Ms G.S. M.A. Botanical Artist. Colour/pen-and-ink art work, exhibitions, curation

Du Plessis, Mrs E. B.Sc. Hons, S.E.D. First Communication Officer. Editing, translating, lay-out

Momberg, Mrs B.A. B.Sc. (Entomology & Zoology). Chief Research Technician. Editing, lay-out

Mulvenna, Mrs J.M. Dip. (Typing). Senior Typist. Word processing, typesetting

Bothalia 22,2 (1992)

303

GARDENS DIRECTORATE

CAPE TOWN

Botha, D.J. D.Sc. Director: Gardens

Britz, R.M. Curator: Harold Porter NBG

Carstens, Mrs M.M. Secretary. Seer. Dip.

Chapiin, P.J. Curator: Witwatersrand NBG

Engelbrecht, B. Curator: Orange Free State NBG

Grobler, P.J. Deputy Director Special Projects

Heilgendorff, J.P. Curator: Pretoria NBG

Kluge, J.P. Curator: Lowveld NBG

Oliver, I.B. Curator: Karoo NBG

Tarr, B.B. Curator: Natal NBG

Winter, J.H.S. Curator: Kirstenbosch NBG

PLANNING, MAINTENANCE & DEVELOPMENT— CAPE TOWN

Theron, B. de W. Town Planning. Senior Landscape Architect

Linde, D.C. N.T.C. (Technician and Inspector of Works). Control Works Inspector

Manasse, S.P. Dip. (Masonry). Artisan

Petersen, R.H.W. Artisan’s Assistant

HAROLD PORTER NBG -BETTY’S BAY

Britz, R.M. Dip. (Forestry). Chief Research Technician

Abrahams, F. General Assistant I

Abrahamse, S.J. Foreman I

Dramat, N. General Assistant I

Forrester, J.A. NTC HI (Hort.). Senior

Research Technician

Harper, C.C. Driver

Jacobs, L.H. General Assistant I

Martinka, A.G. General Assistant II

Rex, G. General Assistant I

Van Coller, A.E. General Assistant I

Visser, C.J.J. General Assistant II

KAROO NBG —WORCESTER

Oliver, I.B. Chief Research Technician

Dlikilili, N.S. General Assistant EH

Kecela, S. General Assistant I

Kumeke, A.M. General Assistant H

Made, M.A. General Assistant I

Madumane, M. General Assistant II

Makubalo, F.N. General Assistant III

Makubalo, J. General Assistant n

Mcinziba, S.S. General Assistant I

Monisi, M.M. General Assistant II

Mpeke, Mrs E.N. General Assistant I

Mtetwa, A.M. General Assistant III

Ngqwazi, D.C.T. General Assistant ni

Nkoloti, P.M. General Assistant H

Perry, Miss P.L. B.Sc. (Hort.), M.Sc. Principal Research

Technician

Qumba, G.A. General Assistant I

Radebe, M.B. General Assistant EH

Sibeko, L.A. General Assistant EH

Sibozo, N.E. Driver

Simane, D.K. Foreman Gr. I

Smit, Mrs A.C. Senior Provisioning Administration Clerk

Theoha, P.P. General Assistant En

Tuswa, M.H. General Assistant I

Viljoen, D.M. N.D. (Hort.). Senior Research Technician

Yekiso, S.M. General Assistant H

KIRSTENBOSCH NBG -CAPE TOWN

Winter, J.H.S. N.D. (Hort.). Control Research Technician

CENTRAL

Adonis, J.J.M. General Assistant, Cleaning Services

Arends, Miss L.D. Assistant, Plant Recording

Basson, W. Head, Cleaning Services

Crowie, A.C. Driver

De Jonge, Miss K. Typist

De Kock, Mrs M.E. General Assistant, Cleaning Services

304

Bothalia 22,2 (1992)

CENTRAL

Ficks, T.E. Driver

Fredericks, Miss N.C.E. General Assistant

Geduldt, D.C. Engraver

Grace, T. Storeman/Sen. Provisioning Administration

Clerk

Haynie, R. General Assistant, Workshop

Jacobs, A.P. Information Officer

Jacobs, F.H. General Assistant, Stores

Jacobs, K.C. Factotum

Jansen, KJ. Driver

Labuschagne, Mrs C.E. B.Sc. (Hon). Education Officer

Marent, Mrs. H.C. Senior Provisioning Administration

Clerk, Plant Recording

McLean, N.S. General Assistant, Workshop

Newman, W. Driver, Workshop

Nicoll, Mrs. R.C. Senior Provisioning Administration

Clerk

Trautman, C.E. Artisan/Supervisor, Workshop

Williams, W.P. Foreman. Drivers

GARDEN

Adonis, A. General Assistant. Ericas

Adams, H. General Assistant. New development

Balabala, L. General Assistant. Dell

Benjamin, R.C.J. Foreman. Annuals

Boonzaaier, I. General Assistant. General garden

Booysen, J.D. General Assistant. Annuals

Bowler, J.H. Foreman. New development

Bowler, M.A. General Assistant. Annuals

Claasens, D. General Assistant. New development

Crowie, H.R. General Assistant. General garden

Crowie, R.W. Foreman. General garden

Hendricks, S. General Assistant. Proteas

Jansen, W. General Assistant. Annuals

Jenkins, A. General Assistant. New development

Johnson, J. General Assistant. Annuals

Julius, J.A. Foreman. Dell

Lewis, D.P. General Assistant. Proteas

Lewis, PS. General Assistant. Proteas

Loft, G.E. General Assistant. Proteas

Lukas, K. General Assistant. General garden

Manuel, D.R. General Assistant. Annuals

Maxwell, PE. General Assistant. New development

McKlein, P. General Assistant. Dell

Morris, J.N.M. General Assistant. Proteas

Palmer, I. General Assistant. General garden

Petersen, A. General Assistant. Cycads

Philander, N. General Assistant. New development

Pietersen, J. General Assistant. New development

Plaatjies, S.D. General Assistant. Proteas

Ruiters, M. General Assistant. New development

Sampson, D. General Assistant. Dell

Sampson, R. General Assistant. General garden

Solomons, E.A. General Assistant. Proteas

Solomons, K.J. General Assistant. Proteas

Van der Westhuizen, A.J. Foreman. Proteas

Van Rooy, K. General Assistant. Annuals

Williams, M.L.J. General Assistant. New development

ESTATE

Le Roux, PH. Chief Research Technician. Head: Estate

Abrahams, M. General Assistant. Estate & Trails

Adams, J. General Assistant. Estate & Trails

Adonis, S.J. General Assistant. Aliens

Alfreds, M. General Assistant. Aliens

Andrews, M.M. General Assistant. Estate & Trails

Arendse, B.A. General Assistant. Aliens

Baadjies, I. General Assistant. Estate & Trails

Bezuidenhout, A.K. General Assistant. General main-

tenance

Bouwers, G.G. General Assistant. Construction

Claasen, F. General Assistant. Aliens

Dollie, Y. General Assistant. Estate & Trails

Fienies, C. General Assistant. General maintenance

Geswind, A.J. General Assistant. Lawnmowers

Grootboom, C.J. General Assistant. Construction

Hendricks, M. General Assistant. Estate & Trails

Hope, C. General Assistant. Construction

Isaacs, M. General Assistant. Aliens

Jackson, P General Assistant. Lawnmowers

Jaftha, R. General Assistant. General maintenance

Jaftha, W.R. General Assistant. Construction

Kayster, G.J. Foreman. Construction

Kuscus, G.W. Foreman. General maintenance

Matthews, I.N. General Assistant. Estate & Trails

McLean, A. General Assistant. Aliens

Mitchells, G. Foreman. Estate & Trails

Petersen, J. General Assistant. Aliens

Petersen, N.H. General Assistant. Construction

Plaatjies, D. General Assistant. Aliens

Plaatjies, M.P General Assistant. Estate & Trails

Rhode, W.C. General Assistant. Estate & Trails

Sampson, J. Foreman. Aliens

Sampson, J.J. General Assistant. Lawnmowers

Snyders, S.G. General Assistant. Lawnmowers

Solomons, G. General Assistant. Aliens

Solomons, S. General Assistant. Construction

Van der Meulen, C.A. General Assistant. Construction

Van Gusling, E.J. Foreman. Lawnmowers

Willis, J.P.P General Assistant. Estate & Trails

Wyngaard, D.J. General Assistant. Estate & Trails

Bothalia 22,2 (1992)

305

NURSERY

Saunders, R.C. N.T.C. Ill (Hort.). Chief Research Technician. Head: Nursery

Adams, H. Foreman, Plant utilization

Adonis, M. General Assistant. Nurseryman

Apolis, A. General Assistant

August, C. General Assistant. Seed room

Berman, R.C. Security

Britz, R.M. N.D. (Forestry). Principal Research Tech-

nician. Head: Security

Carrol, R.R. General Assistant

Daniels, A. General Assistant. Plant utilization

Davids, M. General Assistant. Seed room

Davids, M.I. General Assistant. Senior Nurseryman

Davids, N. General Assistant. Nurseryman

Davids, M. General Assistant. Senior Nurseryman

Duncan, G.D. N.D. (Hort.). Principal Research Tech-

nician

Eksteen, M. General Assistant. Succulents

Erasmus, S. Security

Francis, J. General Assistant. Plant utilization

Goliath, Mrs L. General Assistant. Seed room

Gould, Mrs M. N.D. (Hort.). Senior Research Tech-

nician

Hendricks, B.D. General Assistant. Succulents

Hitchcock, A.N. N.D. (Hort.), N.H.D. (Hort.). Senior

Research Technician

Jacobs, C.W. General Assistant. Nurseryman

Jacobs, D.G. Foreman. Plant utilization

Jacobs, E.C. General Assistant. Seed room

Jacobs, H.C. Security

Jamieson, Mrs H.G. N.D. (Parks & Recreation). Research

Technician

January, C. Security

January, PC. General Assistant. Plant utilization

Juta, E.C. General Assistant. Plant utilization

Kettledas, P.G. General Assistant. Nurseryman

King, O. General Assistant. Nurseryman

Koma, B. General Assistant. Succulents

Kotze, F.G. N.T.C. HI (Hort.). Principal Research Tech-

nician

Lawrence, E. General Assistant

Lewin, T.B. General Assistant. Nurseryman

Manuel, I.P. Foreman. Seed room

Marthinus, E. General Assistant. Succulents

Mulder, G.R. General Assistant. Nurseryman

Notten, Miss A.L. Student (Temporary)

Pick, Miss U.M. General Assistant. Seed room

Powrie, Miss F.J. B.Sc. (Hon) N.D. (Hort.), Senior

Research Technician

Rudolph, A. Security

Sardien, T.P. General Assistant. Group Leader, succulents

Sauls, C.J. General Assistant. Nurseryman

Siljeur, H.V. General Assistant. Security

Smith, D. General Assistant

Solomons, T. Foreman. Security

Tamboer, J.S. Foreman. Nursery

Thomas, Mrs M.L. Senior Research Technician

Van der Walt, Mrs L.E. N.D. (Hort.). Research Tech-

nician

Van Jaarsveld, E.J. M.Sc. N.D. (Hort.). Chief Research

Technician

Van Rooyen, Miss S. General Assistant. Seed room

Van Schalkwyk, J. General Assistant. Succulents

Von Somnitz, Miss B.D. N.D. (Hort.). Kirstenbosch

Scholar (Temporary)

Williams, G.C. General Assistant. Security

LOWVELD NBG— NELSPRUIT

Kluge, J.P. B.Sc. Hons, T.H.O.D. Chief Research Technician

Froneman, W.C. N.D. (Nature Conservation & Manage-

ment), N.D. (Parks & Recreation Admin.), N.T.C. HI

(Hort.). Principal Research Technician

Hurter, P.J.H. B.Sc. Research Technician

Khoza, D.E. General Assistant

Khoza, F.D. General Assistant

Khumalo, N.S. General Assistant

Khumalo, S.S. General Assistant

Magagula, K.E. General Assistant

Magagula, N.R. General Assistant

Mahlahlubane, F.J. General Assistant

Makamo, Mrs J.E. Geheral Assistant

Makhubela, B.J. General Assistant

Mantseke, N.A. General Assistant

Maqungo, Miss VL B. General Assistant

Mazibuko, F.E. General Assistant

Mdhluli, M.B. General Assistant

Mdluli, M.E. General Assistant

Mdluli, S. General Assistant

Mkhatshwa, Mrs N.S. General Assistant

Mteto, E.M. General Assistant

Muswili, K.J. General Assistant

Ngomane, S. General Assistant

Ngqani, Mrs L.S. General Assistant

Ngwengoma, P.N. General Assistant

Ngwenya, PS. General Assistant

Ngwenyama, K.A. General Assistant

Ngwenyama, M.M. General Assistant

Nkosi, M.P. General Assistant

Nkosi, Mrs P.B. General Assistant

Nkosi, Mrs S.L. General Assistant

Nyathi, R.M. General Assistant

Shabangu, M.E. General Assistant

Shabangu, S.L. General Assistant

Shabangu, W.N. General Assistant

Shawe, S.A. Foreman

Sibule, B.F. General Assistant

Sibure, M.E. General Assistant

Sibure, W.F. General Assistant

Sigudla, B.M. General Assistant

Soka, M.P. General Assistant

Thabethe, S.S. General Assistant

Van der Walt, Mrs G.A. Provisioning Administration

Clerk

306

Bothalia 22,2 (1992)

NATAL NBG -PIETERMARITZBURG

Tarr, B.B. N.D. (Parks & Recreation Admin.), Advanced Dip. (Adult Education).

Chief Research Technician

Busani, M.A. Driver

Dlamini, N.S. General Assistant n. Nursery foreman

Dlungwane, R. Foreman

Gabuza, A. General Assistant II

Gates, Ms J.E. N.D. (Hort.), N.D. (Parks & Recreation

Admin.). Senior Horticulturist. Kniphofia, forest

species.

Kistner, H.A. N.D. (Hort.)

Madonda, M. General Assistant I

Mbense, A. General Assistant n. Machine operator

Mdluli, K. General Assistant I

Mkize, M. General Assistant I

Mncwabe, Ms A. General Assistant I

Mncwabe, P. General Assistant II

Mpangase, Z. General Assistant II

Mpulo, D.H. General Assistant I

Mpulo, Ms E. General Assistant I

Mthalane, A. General Assistant II

Mthembu, D. General Assistant I

Mtolo, C. General Assistant HI. Team leader

Nkabini, A. General Assistant II

Nzakwe, W. General Assistant I

Radebe, A. General Assistant I

Tshangase, M. General Assistant II

Van der Merwe, Ms M.E.H. Provisioning Administration

Clerk

Zimu, J. General Assistant II

Zimu, S. General Assistant I

Zondi, Ms B.P. General Assistant I

Zondo, Z. General Assistant II

Zuma, J. General Assistant I

Zuma, Ms K. General Assistant I

ORANGE FREE STATE NBG— BLOEMFONTEIN

Engelbrecht, B. N.D. (Hort.), N.D. (Parks & Recreation Management), N.D. (Forestry)

Chief Research Technician

Eysele, Mrs J.P. Senior Provisioning Administration Clerk

Kokela, Mrs C.L. General Assistant II . Nursery

Lekheto, M.J. General Assistant HI. Maintenance

Lekheto, T.S. General Assistant n. Bulb area

Lumley, M.J. Principal Research Technician. Nursery

Mbolekwa, G.M. General Assistant n. Grass garden

Mbolekwa, L.M. General Assistant II. Rhus, display

Mofokeng, J.M. General Assistant II. Entrance area

Mofokeng, M.S. General Assistant II. Braai area

Mohokare, J. Driver

Mohapi, Mrs M.A. General Assistant II. Cleaner

Mohapi, T.A. General Assistant III. Grower, nursery

Moima, K.H. Foreman I. Foreman garden

Moima, T.J. General Assistant I. Maintenance

Mopeli, M.J. General Assistant I. Nursery

Moticoe, Mrs M.A. General Assistant II. Nursery

Nakanyane, R.B. General Assistant II. Pond, display

area

Nakedi, M.J. General Assistant II. Estate paths

Olifant, M.D. General Assistant 13. Kiosk area

Rampai, M.A. General Assistant I. Maintenance

Sebolai, P.R.A.N. General Assistant II. Nursery, tools

maintenance

Semeyane, T.D. General Assistant II. Maintenance

Thaele, Mrs M.E. General Assistant n. Seed room,

nursery

PRETORIA NBG

Heilgendorff, J.P. N.H.D. (Hort.). Chief Research Technician

Baloi, R.F. General Assistant II

Baloyi, K.J. General Assistant III

Baloyi, S.J. Driver/Operator

Baloyi, S.M. Research Assistant

Chipi, S. Security Assistant

Chuma, S.J. Security Assistant

De Ruiter, J. General Assistant III

Dry, D.H. N.D. (Hort.) Chief Research Technician.

Technical papers on horticulture and plants

Key ter, B.A. Senior Security Officer

Klapwijk, N.A. N.D. (Hort.), N.D. (Plant Prod.), N.D.

(Diesel Fitting). First Research Technician. Proteas,

fuchsias, southern part of Pretoria garden

Lephera, J. General Assistant II

Letsoalo, H.M. General Assistant I

Mabasa, J.R. Security Assistant

Mabasa, P.P. General Assistant II

Mabunda, Z.S. General Assistant II

Machika, S.M. General Assistant II

Mahlangu, J.J. General Assistant I

Makafula, F. General Assistant I

Bothalia 22,2 (1992)

307

PRETORIA NBG

Makena, M.S. Driver/Operator

Makena, S.N. Foreman

Makena, T.J. General Assistant II

Makgopo, C.K. General Assistant II

Makhubela, D. Foreman

Makhubela, K.P. General Assistant II

Makoeng, P.T. General Assistant II

Makola, J. General Assistant I

Makola, L.M. General Assistant HI. Tractor driver

Makua, E.G. General Assistant I

Malewa, D. General Assistant II

Malobola, L. General Assistant II

Malobola, M. General Assistant m

Maluleke, M.J. Security

Mametja, A. General Assistant II

Mariri, N.J. Factotum

Marule, PM. General Assistant HI. Tractor driver

Masango, M.G. General Assistant II

Masokwameng, T.P. General Assistant II

Mathabathe, D.S. General Assistant II

Matlala, S.M. General Assistant II

Matshika, S.P. General Assistant II

Mnyangeni, L.D. General Assistant II

Mogoru, M.F. General Assistant II

Mogoru, S. General Assistant II

Mohale, F.R. Foreman

Mohale, J.N. General Assistant I

Mokawe, N.R. General Assistant I

Molefe, J.R. General Assistant EH

Molomo, S.E. General Assistant I

Mononyane, J.B. General Assistant II

Morifi, L.J. General Assistant I

Motshweni, V. General Assistant II

Msisa, S.K. General Assistant II

Mudau, R.T. General Assistant I

Muhali, B. General Assistant II

Nkambule, J. General Assistant I

Nkoane, J.M. General Assistant I

Nkwana, F.N. Driver/Operator

Noko, J.M. Research Assistant

Noku, A.Y. General Assistant HI. Tractor driver

Ramakgaphola, A.M. General Assistant I

Ramatsetse, PM. Security Assistant

Rampopana, A.M. General Assistant I

Sete, L. Foreman

Shirindi, J.R. General Assistant I

Shilubane, E. Storeman Assistant

Sithole, J. General Assistant I

Steenkamp, L.C. Foreman

Strydom, D.J.F. N.T.C. HI (Hort.), N.D. (Parks & Rec.

Management). Chief Research Technician. Cultiva-

tion of mass plants, northern part of Pretoria garden

Swartz, Ms P. M.Sc. (Botany). Senior Horticulturist

Tefu, P.R. General Assistant H

Tloubatla, J.L. General Assistant I

Tolo, P.K. General Assistant I

WITWATERSRAND NBG — WILROPARK

Chaplin, P.J. NTC Dip. (Hort.). Chief Research Technician

Behr, Miss C.M. B.Sc. Hons. Principal Research Tech-

nician. Education and information, phytosociologi-

cal classification of the Witwatersrand garden

Bonewe, N.W. General Assistant H. Machine operator

Head, Mrs S.E. Senior Provisioning Administration Clerk

Khedzi, K.P. General Assistant H. Nursery

Lxirenzo, T.C. N.D. (Hort.), Dip. (Small Bus. Man.).

Senior Research Technician

Lukhwa, N.A. General Assistant I. Garden

Luvhimbi, T.S. General Assistant I. Garden

Manyikana, T.M. General Assistant I. Garden

Mbulaheni, N.P. General Assistant II. Garden

Mulibana, N.S. General Assistant II. Machine operator

Mmola, Ms B.E. General Assistant I. Cleaner

Ncuba, M.J. General Assistant I. Garden

Ndou, M.W. General Assistant H. Machine operator

Ndwambi, N.W. General Assistant I. Garden

Nedambale, M.P. General Assistant in. Nursery

Nemalili, M.E. Driver

Nemalili, A.S. General Assistant EH. Driver

Nekhavhambe, S.P. General Assistant I. Garden

Nemavhulani, M.R. General Assistant I. Storeman

Nenungwi, M.S. General Assistant I. Nursery

Rammela, N.N. General Assistant H. Machine operator

Raphalalani, V.S. General Assistant I. Nursery

Ravhuhali, P.W. General Assistant I. Garden

Steel, Miss B.S. N.D. (Nature Conservation), Dip.

(Journalism). Research Technician

Tebeile, Ms Z.M. General Assistant H. Clerical assistance

Tshisikule, G.M. General Assistant II. Garden

RESEARCH DIRECTORATE

PRETORIA

Eloff, Prof. J.N. M.Sc. (Chemistry), D.Sc. (Plant Biochemistry). Director: Research

Meyer, Ms M.C. Dip. Gim. Man. Personal Secretary

Saayman, Mrs E.J.L. M.Sc. Hons. Scientific Liaison Officer. Cytotaxonomy

308

Bothalia 22,2 (1992)

Arnold, T.H. Programme Leader: Data Management

Brink, Mrs E. Curator: Albany Museum Herbarium (Grahamstown)

Donaldson, Dr J.S. Programme Leader: Conservation Biology (Cape Town)

Du Plessis, Mrs H. Head: Research Support Services

Oliver, E.G.H. Curator: Stellenbosch Herbarium (Stellenbosch)

Rourke, Dr J.P. Curator: Compton Herbarium (Cape Town)

Rutherford, Dr M.C. Programme Leader: Stress Ecology (Rondebosch)

Welman, Miss W.G. Acting Curator: National Herbarium

Williams, Ms R. Curator: Natal Herbarium (Durban)

ALBANY MUSEUM HERBARIUM -GRAHAMSTOWN

Brink, Mrs E. B.Sc. Scientist. General taxonomy and botanical information

particularly trees and woody plants

Booi, A.D. Herbarium Assistant. General herbarium practice, identification and information

Verwey, Mrs L.M. Provisioning Administration Clerk. General administration, information and curation

Zenzile, J.M. General Assistant

DATA MANAGEMENT— PRETORIA

Arnold, T.H. M.Sc. Assistant Director. Computer application especially in taxonomy

De Wet, Mrs B.C. B.Sc., B.A., H.D.L.S. Datametrician

Evenwel, Mrs E. Scientific Assistant

Harris, Mrs B.J. Scientific Assistant

COMPTON HERBARIUM -CAPE TOWN

Rourke, J.P. Ph.D., F.L.S. Specialist Scientist. Systematics of southern African Proteaceae, Stilbaceae

Cupido, Mrs C. General Assistant

Forster, Mrs S.E. Provisioning Administration Clerk

Holm, Mrs K. Scientific Assistant

Kurzweil, H. Ph.D. Scientist. Systematics of southern African terrestrial orchids

Paterson-Jones, Mrs D.A. (nee Snijman). M.Sc. Scientist. Systematics of Amaryllidaceae

Roux, J.P. N.T.C. (Hort.), M.Sc. Scientific Officer. Systematics of Pteridophyta

Steiner, K.E. Ph.D. Scientist. Systematics of Scrophulariaceae and evolutionary interactions between oil-secreting

flowers and oil-collecting bees

CONSERVATION BIOLOGY-CAPE TOWN

Donaldson, J.S. M.Sc. (Entomology), Ph.D. (Zoology). Assistant Director. Cycad biology,

plant/insect interactions, conservation biology

Botha, P.A. N.H.D. (Hort.). Scientific Officer. Tissue culture research

Bowler, Mrs M. General Assistant

Brown, N.A.C. Ph.D. Specialist Scientist. Seed biology research, plant growth regulators

De Lange, J.H. B.Sc. (Hort.), M.Sc. (Plant Physiology), D.Sc. (Agric.). Specialist Scientist. Ecology, tissue culture,

horticulture

Jita, Ms G.N. General Assistant

Kohly, Miss N. B.Sc. (Entomology & Microbiology). Scientific Officer. Scientific illustration

Leivers, S. B.Sc. (Microbiology, Plant Virology). Commercial tissue culture

Manning, J.C. Ph.D. Scientist. Systematics of Iridaceae and Orchidaceae, cladistics and biogeography

Nanni, Ms I. B.Sc., H.E.D. Scientific Officer. Ecology, seed biology

Parenzee, Ms H.A. Administrative Assistant

Rebelo, A.G. Ph.D. (Zoology). Scientist. Conservation biology, biogeography.

Scott, Mrs G. B.Sc. (Pharmacy), M.Sc. Scientific Officer. Plant secondary compounds, medicinal plants

Bothalia 22,2 (1992)

309

NATAL HERBARIUM -DURBAN

Williams, Ms R. B.Sc. Hons, H.D.E. Scientific Officer

Mbonambi, M.B. General Assistant. Gardener

Ngwenya, M.A. Herbarium Assistant. Identification, information

Noble, Mrs H-E. Provisioning Administration Clerk. General administration

Nzimande, S.B. General Assistant

Sikhakhane, T.B. Herbarium Assistant. General herbarium practice

NATIONAL HERBARIUM— PRETORIA

Welman, Miss W.G. M.Sc. Acting Curator. Scientist. Taxonomy of Convolvulaceae— Asteraceae

Glen, H.F. Ph.D. Assistant Curator: information. Scientist. Taxonomy of trees and succulents,

especially Aloe, also cultivated plants

Jordaan, Mrs M. B.Sc. Assistant Curator: scientific curation of dicotyledons. Scientific Officer.

Taxonomy of Casuarinaceae— Connaraceae

Koekemoer, Miss M. M.Sc. Assistant Curator: monocotyledons, cryptogams and fossils. Scien-

tist. Taxonomy of Poaceae, Asteraceae; Disparago and related genera

Weisser, P.J. Ph.D. Assistant Curator: herbarium support services. Scientist. Ecology, dune vege-

tation, vegetation dynamics and vegetation mapping

Anderson, H.M. Ph.D. Scientist. Palaeobotany, palaeogeography

Anderson, J.M. Ph.D. Specialist Scientist. Palaeobotany, palaeogeography.

Archer, R.H. M.Sc. Scientific Officer. Taxonomy of mainly Celastraceae, Euphorbiaceae

Brusse, F.A. M.Sc. Scientist. Taxonomy of the southern African lichens

Burger, Mrs S.J.C. Scientific Assistant. General herbarium practice

Cloete, Mrs M. Dip. (Typing). Typist

Dreyer, Miss L.L. M.Sc. Scientific Officer. Taxonomy of mainly Geraniaceae

Fish, Mrs L. B.Sc. Scientific Officer. Poaceae. Identifications, collecting *

Germishuizen, G. M.Sc. Scientist. Plant identifications, taxonomy of Polygonaceae, Fabaceae, Loranthaceae, Viscaceae

Glen, Mrs R.P. M.Sc. Scientific Assistant. Ferns

Herman, P.P.J. M.Sc. Scientific Officer. Identifications, Rubiaceae— Asteraceae, Flora of Transvaal

Heymann, Mrs M.Z. T.E. Dip. Scientific Assistant

Lephaka, M.G. Scientific Assistant. Parcelling and pressing

Makgakga, S.K. Scientific Assistant. Mounting and filing of herbarium specimens

Nicholas, A. M.Sc. Scientist. Taxonomy of Asclepiadaceae (incl. Periplocaceae), carnivorous plants (particularly

Droseraceae), herbarium pests

Perold, Mrs S.M. M.Sc. Scientific Officer. Taxonomy of Ricciaceae (Hepaticae)

Phahla, T.J. Scientific Assistant. Mounting and filing of herbarium specimens

Pretorius, Mrs. R. B.Sc. Hons, H.E.D. Scientific Assistant

Reid, Miss C. M.Sc. Scientist. Monocotyledons. Taxonomy of Cyperaceae

Retief, Miss E. M.Sc. Scientist. Pollen studies of Boraginaceae. Taxonomy of Boraginaceae, Verbenaceae, Lamiaceae,

Scrophulariaceae, Acanthaceae

Strohmaier, Mrs S.M. T.E. Dip. Scientific Assistant

Taussig, Miss J.A. N.D. (Hort.). Scientific Assistant

Van Rooy, J. M.Sc. Scientist. Taxonomy and biogeography of mosses

Van Wyk, Mrs C.M. M.Sc. Scientist. Melolobium, Pelargonium. Taxonomy of Rutaceae, Thymelaeaceae, Apiaceae,

Ericaceae

Veldman, Mrs J.M. Provisioning Administration Clerk

STELLENBOSCH HERBARIUM

Oliver, E.G.H. M.Sc. Curator. Scientist. Taxonomy of the Ericoideae (Ericaceae)

Beyers, Mrs J.B.P. B.Sc. Hons. Scientist. Taxonomy of the Gnidieae (Thymelaeaceae)

Davidse, Mrs E. Scientific Assistant

Fellingham, Mrs A.C. B.Sc. Scientific Officer. Taxonomy of Cliffortia (Rosaceae)

Leith. Mrs J. Provisioning Administration Clerk

310

Bothalia 22,2 (1992)

STRESS ECOLOGY-RONDEBOSCH

Rutherford, M.C. Ph.D., Dip. Datamet. Programme Leader. Stress and disturbance ecology

Bailey, Ms C.L. B.Sc. Hons. Scientific Officer. Terrestrial and aquatic ecology/ecophysiology

Bennett, N.N. Scientific Assistant

Davis, G.W. Ph.D. Scientist. Ecophysiology, resource modelling

De Witt, D.M. Scientific Assistant

Hoffman, M.T. Ph.D. Scientist. Disturbance ecology, desertification, photography

Hunter, Ms D.A. Administrative secretary

Jacobs, Q.C. General Assistant

Jagger, B.W Scientific Assistant

McDonald, D.J. M.Sc. Scientist (Stellenbosch). Vegetation ecology, pollination biology

Midgley, G.F. M.Sc. Scientist. Plant stress physiology /ecology

Musil, C.F. Ph.D. Scientist. Aquatic and terrestrial plant ecophysiology

O’Callaghan, M.G. M.Sc. Scientist (Stellenbosch). Wetlands, salt marshes, coastal vegetation

Pickett, Ms G.A. B.Sc. Hons. Scientific Officer. Vegetation dynamics

Powrie, L.W. M.Sc. Scientist. Karoo ecology, education, computer programming/operations

Wand, S.J.E. M.Sc. (Agric.). Scientific Officer. Ecophysiology

RESEARCH SUPPORT SERVICES -PRETORIA

Du Plessis, Mrs H. M.Sc. Head of Cost Centre. Assistant Scientist. Cytogenetics

Botha, Mrs A.G. Scientific Assistant. Grass leaf anatomy

Romanowski, Mrs A.J. Dip. (Photography). Industrial Technician (Photography). Scientific photography

Roux, Mrs W.J.G. Dip. (Private Secretary). Scientific Assistant. Graphic artist, biology

Steyn, Miss C.C. Scientific Assistant. Embryology

PUBLICATIONS BY THE STAFF

(1991-04-01—1992-03-31)

ARMBRUSTER, W.S. & STEINER, K.E. 1992. Pollination ecology

of four Dalechampia species (Euphorbiaceae) in northern Natal,

South Africa. American Journal of Botany 79: 306—313.

BAILEY, C. & RUTHERFORD, M.C. 1992. Impressions of the Richters-

veld. Veld & Flora 78: 21.

BRITS, G.J. & BROWN, N.A.C. 1991. Control of seed dormancy in

Leucospermum. Proceedings of the Sixth Biennial Conference of

the International Protea Association, Perth, Australia, September

1991.

BROWN, N.A.C. & DREWES, F.E. 1991. Germination of achenes of

members of the Proteaceae following pretreatment with the growth

regulator promalin (GA4 + GA7 + BA) and its components.

Proceedings of the Sixth Biennal Conference of the International

Protea Association, Perth, Australia, September 1991.

BRUSSE, F.A. 1991a. Review: Macrolichens of East Africa, by T.D.V.

Swinscow, & H. Krog. British Museusm (Natural History),

London. South African Journal of Botany 57: 179.

BRUSSE, F.A. 1991b. Two new species in the Agyriaceae (Lichenized

Ascomycotina, Lecanorales) from southern Africa. Bothalia 21:

154-156.

BRUSSE, F.A. 1991c. A new species in the lichen genus Parmelia

(Parmeliaceae, Ascomycotina), from the Blouberg, Northern

Transvaal, South Africa, with further notes on southern African

lichens. Mycotaxon 42: 163—169.

BRUSSE, F.A. & DICKINSON, C.H. 1991. A new foliicolous species

in the lichen genus Porina (Porinaceae, Pyrenulales) from

southern Africa. Mycotaxon 42: 347—350.

BRUSSE, F.A. & KARNEFELT, I. 1991. The new southern hemisphere

lichen genus Coelopogon (Lecanorales, Ascomycotina), with a

new species from southern Africa. Mycotaxon 42: 35—41.

DAVIS, G.W. 1991. Botany without flowers: technological impressions

of the struggle for survival. Veld & Flora 77: 54, 55.

DAVIS, G.W. , FLYNN, A.P. & MIDGLEY, G.F. 1992. Growth and

gas-exchange responses of Leucadendron xanthoconus (Protea-

ceae) seedlings to different nutrient and water regimes. South

African Journal of Botany 58: 56—62.

DE LANGE, J.H. & BOUCHER, C. 1991. Autecological studies on

Audouinia capitata (Bruniaceae). 1. Plant-derived smoke as a seed

germination cue. South African Journal of Botany 56: 700, 701.

DE WET, B.C., ARCHER, R., FISH, L., GERMISHUIZEN, G.,

HERMAN, 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: 191-213.

DONALDSON, J.S. 1991. Three new species .of Microgastrinae

(Hymenoptera: Braconidae) from South Africa with notes on

Glyptapanteles acraeae. Journal of the Entomological Society of

Southern Africa 54: 29—37.

DREYER, L.L. & GERMISHUIZEN, G. 1991. Guidelines for the use

of the genspec programme. Addendum 6. Ms for Standards and

procedures for scientific curation.

DUNCAN, G.D. 1991a. Clivias and their cultivation. Parks & Grounds

59: 21, 22.

DUNCAN, G.D. 1991b. Review: Bulbs. Veld & Flora 77: 63.

DUNCAN, G.D. 1991c. Plant moraeas and be peacock proud! Southern

Argus (24-10-91), Cape Town.

DUNCAN, G.D. 1991d. Spring babianas for pot or garden. Southern

Argus (31-10-91), Cape Town.

DUNCAN, G.D., ROURKE, J.P. & CONDY, G. 1991. Bulbs and bulbous

plants. 1992 Kirstenbosch Year Book. Struik Publishers, Cape

Town.

FOURIE, D.M.C. 1991. Indigenous trees/Inheemse borne. Venda special

stamp issue. Bulletin TPT49. Philatelic Services & Interspa,

Pretoria.

FOURIE, D.M.C. 1992. Acacia trees/Akasia borne. Bophuthatswana

special stamp issue. Bulletin TPT 63. Philatelic Services &

Interspa, Pretoria.

Bothalia 22,2 (1992)

311

FRONEMAN, W. 1992. Doringbosviooltjies. Veld & Flora 78 : 52 , 53.

GERMISHUIZEN, G. 1991a. Review: Flora of south-eastern Queens-

land Vol. 1, by T.D. Stanley & E.M. Ross. Bothalia 21: 115.

GERMISHUIZEN, G. 1991b. Dichilus strictus. The Flowering Plants

of Africa 51: t. 2037.

GERMISHUIZEN, G. 1991c. Caesalpinia bracteata , a new species from

the Onseepkans area of the northern Cape Province (Fabaceae).

Bothalia 21: 152.

GLEN, H.F. 1991a. The Mary Gunn Library of the National Botanical

Institute. Palm Enthusiast 8: 14—16.

GLEN; H.F. 1991b. The paper chase [notes and reviews of new litera-

ture]. Trees in South Africa 42: 48 —50.

GLEN, H.F. & HARDY, D.S. 1991a. Unusual fruit of nyala tree. Trees

in South Africa 42: 22— 27.

GLEN, H.F. & HARDY, D.S. 1991b. The type specimen of Aloe sout-

pansbergensis Verdoom (Liliaceae/Asphodelaceae). Bothalia 21:

151, 152.

GLEN, H.F. & LAVRANOS, J.J. 1991. Juniperus phoenicea. The

Flowering Plants of Africa 51: t. 2022.

GLEN, H.F., MOMBERG, B.A. & POIGIETER, E. 1991. Bothalia:

Contents to vols 1—20. Bothalia supplement: 1-32.

GLEN, H.F. & ONDERSTALL, J. 1991. Steganotaenia araliacea. The

Flowering Plants of Africa 51: t. 2031.

GLEN, H.F. & TAPSON, A. 1991. Notes on the proposed guide to exotic

trees cultivated in southern Africa. Palm Enthusiast 8: 10—13.

GLEN, H.F. , WEISSER, P.J. & Members of the Tree Society of Southern

Africa 1991. The tree vegetation of Basfontein Farm. Trees in South

Africa 42: 11—22.

GOLDBLATT, P, MANNING, J.C. & BARI, A. 1991. Sulcus and oper-

culum structure in the pollen grains of Iridaceae subfamily

Ixioideae. Annals of the Missouri Botanic Garden 78: 950—961.

HERMAN, P.P.J. 1991a. Stachytarpheta species in southern Africa

(Verbenaceae). Bothalia 21: 57.

HERMAN, P.P.J. 1991b. Plectroniella armata. The Flowering Plants of

Africa 51: t. 2029.

KOEKEMOER, M. 1991. New species in the genus Disparago (Aster-

aceae). Bothalia 21: 158—161.

KURZWEIL, H. 1991a. The unusual structure of the gynostemium of

the Orchidaceae— Coryciinae. Botanische Jahrbilcher fur Systema-

tik, Fflanzen-geschichte und Pflanzen-geographie 112: 273—

293.

KURZWEIL, H. 1991b. Zur Orchideenflora der Kapprovinz. (Teil 2).

Die Orchidee 42: 139—142.

KURZWEIL, H. 1991c. A method to illustrate the delicate flower

structures of the genus Disperis (Orchidaceae). Communications

of the Electron Microscopy Society of South Africa, Cape Town

Conference, December 1991 Vol. 21, Abstract p. 25.

KURZWEIL, H. 1992. The structure of the gynostemium in the

Orchidaceae-Coryciinae. Proceedings of the 13th World

Conference (Auckland 1990): 268-274.

KURZWEIL, H., LINDER, H.P. & CHESSELET, P. 1991. The phylo-

geny and evolution of the Pterygodium—Corycium complex

(Coryciinae, Orchidaceae). Plant Systematics and Evolution 175:

161-223.

KURZWEIL, H. & WEBER, A. 1991. Floral morphology of southern

African Orchideae. I. Orchidinae. Nordic Journal of Botany 11:

155-178.

KURZWEIL, H. & WEBER, A. 1992. Floral morphology of southern

African Orchideae. II. Habenariinae. Nordic Journal of Botany

12: 39-61.

MANNING, J.C. 1991. A new species of Rhigioglossa (Mesomyia)

Macquart 1850 (Diptera: Tabanidae) from the Natal Drakensberg.

Annals of the Natal Museum 32: 163—166.

MANNING, J.C. & GOLDBLATT, P. 1991. Systematic and phylogenetic

significance of the seed coat in the shrubby African Iridaceae

Nivenia, Klattia and Witsenia. Botanical Journal of the Linnean

Society 107: 387—404.

MANNING, J.C. & LINDER, H.P. 1992. Pollination and evolution in

Disperis (Orchidaceae), or why are there so many species? South

African Journal of Science 88: 38—49.

MIDGLEY, G.F. & HOFFMAN, M.T. 1991. Heuweltjies: nutrient

factories. Veld & Flora 77: 72 —75.

MUSIL, C.F. 1991. Seed bank dynamics in sand plain lowland fynbos.

South African Journal of Botany 57: 131-142.

MUSIL, C.F. & DE WITT, D.M. 1991. Heat stimulated germination

in two Restionaceae species. South African Journal of Botany 57:

175, 176.

OLIVER, E.G.H. 1991a. The Ericoideae— a review. Contributions from

the Bolus Herbarium 13: 158—208.

OLIVER, E.G.H. 1991b. Obituary: Hedley Brian Rycroft. Bothalia 21:

109—114.

OLIVER, E.G.H. & FELLINGHAM, A.C. 1991. A new species of

Cliffortia from the southwestern Cape (Rosaceae). Bothalia 21:

60-62.

OLIVER, E.G.H. & OLIVER, I.M. 1991. Studies in the Ericoideae

(Ericaceae). V1H. New species in Erica (section Pseuderemia)

from southern Africa. Bothalia 21: 137—142.

PEROLD, S.M. 1991a. Studies in the genus Riccia (Marchantiales) from

southern Africa. 22. R. rubricollis now validated, typified and

described. Bothalia 21: 51-54.

PEROLD, S.M. 1991b. Studies in the genus Riccia (Marchantiales) from

southern Africa. 23. R. bullosa-, typification and a full descrip-

tion. Bothalia 21: 129—135.

PEROLD, S.M. 1991c. Two new species of Riccia L. from tropical Africa:

R. somaliensis and R. erubescens. Journal of Bryology 16:

367-377.

PICKETT, G.A. & HOFFMAN, M.T. 1992. Karosion: soil erosion in

the Karoo. Veld & Flora 78: 8, 9.

POWRIE, F.J. 1991a. Who says they are boring? Grow South African

pelargoniums. Veld & Flora 77: 84-86.

POWRIE, F.J. 1991b. The plant collector and collecting, the ethics of

collecting. British Pelargonium and Geranium Society,

Geraniaceae Group News Autumn 91.

POWRIE, F.J. 1992. Some thoughts and comments on Pelargonium

tricolor, relatives and hybrids. British Pelargonium and Geranium

Society, Geraniaceae Group News Winter 91/92.

POWRIE, L.W. 1992. How alert are those Richtersveld plants? Veld &

Flora 78: 18, 19.

REBELO, A.G. 1992a. Preservation of biotic diversity. In R.M. Cowling,

The ecology of fynbos: nutrients, fire and diversity. 309—344.

Oxford University Press, Cape Town.

REBELO, A.G. 1992b. Red Data Book species in the Cape Floristic

Region: threats, priorities and target species. Transactions of the

Royal Society of South Africa 48: 55—86.

REBELO, A.G., COWLING, R.M. & HOLMES, P.M. 1992. Plant

diversity and endemism. In R.M. Cowling, The ecology of fynbos:

nutrients, fire and diversity: 62—110. Oxford University Press,

Cape Town.

REYNOLDS, Y. 1991. The NBI Library at Kirstenbosch. Veld & Flora

77: 44-46.

ROUX, J.P. 1991. A new species of Elaphoglossum (Pteridophyta:

Lomariopsidaceae) from the Tristan da Cunha Island group.

Central South Atlantic. South African Journal of Botany 57: 234,

235.

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

Systematic studies in the genus Mohria (Pteridophyta: Anemia-

ceae) I. Comparative morphology of the rhizome and frond. South

African Journal of Botany 58: 83—89.

RUTHERFORD, M.C. 1991a. Diversity of photosynthetic responses in

the mesic and arid Mediterranean-type climate regions of southern

Africa. In G. Esser & D. Overdieck, Modem ecology. Basic and

applied aspects: 133—160. Elsevier, Amsterdam.

RUTHERFORD, M.C. 1991b. Review: Germination physiology and desert

ecology, by A.M. Mayer & G. Orshan. Bothalia 21: 227, 228.

SAUNDERS, R.C. 1992. Ceraria namaquensis. Grafting as a solution

to the problem of growing succulents in non-arid areas. Veld &

Flora 78: 4.

SCOTT, G. 1991. Botanical exploration of the West Coast, and medicinal

plants of the West Coast: a circle of seasons in South Africa:

19—21, 171-174. Struik, Cape Town.

SERGIO, C. & PEROLD, S.M. 1992. A new species of Riccia L. from

the island of Madeira, Riccia atlantica, sp. nov. Journal of

Bryology 17: 127—132.

SNUMAN, D.A. 1991. Kamiesbergia, a new monotypic genus of the

Amaryllideae— Strumariinae (Amaryllidaceae) from the north-

western Cape. Bothalia 21: 125—128.

SPIES, J.J., VAN DER MERWE, E., DU PLESSIS, H. & SAAYMAN,

E.J.L. 1991. Basic chromosome numbers and polyploid levels

in some South African and Australian grasses (Poaceae). Bothalia

21: 163-170.

STEINER, K.E. 1992a. A new Diascia species (Scrophulariaceae) from

the Richtersveld, South Africa. South African Journal of Botany-

Si,: 36-38.

STEINER, K.E. 1992b. Two new Diascia species (Scrophulariaceae)

from the Little Karoo. South African Journal of Botany 58: 39—47.

STEINER, K.E. 1992c. Two new Diascia species from the Nieuwoudt-

ville area, western Cape. South African Journal of Botany 58:

202 -206.

312

Bothalia 22,2 (1992)

STEINER, K.E. 1992d. Three new species of Diascia (Scrophularia-

ceae) from the western Cape. Bothalia 22: 0—18.

STEINER, K.E. & WHITEHEAD, V.B. 1991a. Oil flowers and oil bees:

further evidence for pollinator adaptation. Evolution 45:

1493-1501.

STEINER, K.E. & WHITEHEAD, V.B. 1991b. Resin collection and the

pollination of Dalechampia capensis (Euphorbiaceae) by Pachyan-

thidium cordatum (Hymenoptera: Megachilidae) in South Africa.

Journal of the Entomological Society of Southern Africa 65:

67-72.

VAN DER WALT, J.J.A. & ROUX, J.P. 1991. Taxonomy and phylogeny

of Pelargonium section Campy lia (Geraniaceae). South African

Journal of Botany 57: 291—293.

VAN JAARSVELD, E.J. 1991a. The Blouberg botanical expedition. Veld

& Flora 78: 27-29.

VAN JAARSVELD, E.J. 1991b. Successful cultivation, knowledge of a

plant’s growing requirements necessary. Southern Argus. Thursday

September 26, 1991.

VAN JAARSVELD, E.J. 1991c. Hardy Cape Vygies with beautifiil floral

character. The Argus. Oct. 1991.

VAN JAARSVELD, E.J. 1991d. Aloes for your garden. The Argus. Oct.

1991.

VAN JAARSVELD, E.J. 1991e. Honeybells. There’s a honeybell to suit

almost every garden. Veld & Flora 77: 90, 91.

VAN JAARSVELD, E.J. 1991f. The Blouberg botanical expedition, with

specific reference to the succulent vegetation plants. Aloe 28:

78-83.

VAN JAARSVELD, E.J. 1991g. The subfamily Asphodelaceae with

specific reference to Trachyandra tortilis. Cactus and Succulent

Journal (U.S.) 63: 196-199.

VAN JAARSVELD, E.J. 1991h. Tylecodon bodleyae, a new species from

the northwestern Cape. Cactus and Succulent Journal (U.S.) 64:

57-61.

VAN JAARSVELD, E.J. 1991i. The conservation status of Gasteria

baylissiana Rauh, a rare endemic of the Zuurberg (Eastern Cape).

The Cactus and Succulent Journal of Great Britain 9: 99—

101.

VAN JAARSVELD, E.J. 1992. The Genus Gasteria, a synoptic review.

Aloe 1: 1-30.

VAN ROOY, J. 1991. The genus Rhabdoweisia in southern Africa: R.

crispata new to Africa, and R. fugax. The Bryologist 94: 409-

412.

WAND, S.J.E., CUTTING, J.G.M. & JACOBS, G. 1991a. Nectarine fruit

growth as influenced by TIBA sprays and girdling. Journal of

the South African Society of Horticultural Science 1: 43—46.

WAND, S.J.E., CUTTING, J.G.M. & JACOBS, G. 1991b. The relation-

ship between basipetal auxin transport and calcium and glucose

uptake try young peach fruits in vitro. Journal of the South African

Society of Horticultural Science 1: 47—50.

WAND, S.J.E., CUTTING, J.G.M., JACOBS, G. & THERON, K.I. 1991.

Calcium and indole-3-acetic acid contents of developing nectarine

fruits after 2,3,5 triiodo-benzoic acid sprays and girdling. Journal

of the South African Society of Horticultural Science 1: 3—7.

WEISSER, P.J., BACKER, A., GLEN, H.F. & NGWENYA, A. 1991.

Recolonization of a landslide in the Kwambonambi dune area.

Natal, South Africa 1987—1990. In D.A. Everard & G.P. von

Maltitz, Dune forest dynamics in relation to land use practices:

72—80. Environmental Forum Report, Foundation for Research

and Development, CSIR, Pretoria.

WELLS, M.J. 1991. Introduced plants of the Fynbos Biome of South

Africa. In R.H. Groves & F. di Castri, Biogeography of Medi-

terranean invasions: 115—129. Cambridge University Press,

Cambridge.

WRIGHT, M.G., VISSER, D. & DE LANGE, J.H. 1991. Autecological

studies on Audouinia capitata (Bruniaceae). 2. Insects as pollen

vectors. South African Journal of Botany 57: 260—263.

Bothalia 22,2: 313 (1992)

Book Review

LIFE STRATEGIES OF SUCCULENTS IN DESERTS. With special

reference to the Namib desert, by D.J. VON WILLERT, B.M. ELLER,

M.J.A. WERGER, E. BRINCKMANN & H.-D. IHLENFELDT. 1992.

Cambridge Studies in Ecology, Cambridge University Press, The Press

Syndicate of the University of Cambridge, The Pitt Building, Trumpington

Street, Cambridge CB2 1RP. Pp. 340. ISBN 0 521 24468 4.

A remarkable book has been produced by a team of authors, all well

known in the field of research on succulent plants and/or the arid regions

of southern Africa. The book is of particular relevance to that sub-

continent, because it is based on the solid results of 12 years field research

in the Richtersveld and the Knersvlakte in the northwestern Cape Province,

and represents a major contribution to the ecology of the flora of the

Succulent Karoo.

The book concentrates on the ‘life strategies’ of succulent plants. ‘Life

strategy’ is defined as ‘the plant’s set of integrated . . . features that are

suitable, in the context of the whole life cycle of the plant, to maximize

the chances to successfully meet the specific ecological problems set

by its habitat, to exploit the resources in its habitat and thus to maximize

its chances for survival.’

This very wide definition is filled with ample life in five subsequent

chapters.

In the first chapter it is shown that a purely morphological or anatomical

definition of succulents cannot be satisfying. An ecophysiological

definition is proposed focusing on the storage of utilizable water which

makes the plant temporarily independent of external water supply. It is

shown that the storage of water results in structural requirements, espe-

cially with respect to the static problem caused by the heavy water bur-

den and with respect to protective tissues.

The second chapter deals with the climate and vegetation of deserts

in general. The characteristics of arid climates are shown together with

peculiarities of the life cycle of desert plants. More detailed information

on the Namib desert in a very wide sense is given in the subsequent

chapter, concentrating on the very steep coast-inland gradient. The typical

diurnal and annual sequences of periods with either high air humidity

from the Benguela current or the very dry conditions caused by berg

winds are analysed in detail.

The core of the book is found in the fourth chapter, which over 176

pages introduces the reader to the ecophysiology of the succulent plants

which is subdivided into (a) energy fluxes, (b) water fluxes and (c) carbon

fluxes. All three subchapters partly have textbook character because they

all start off with a good explanatory introduction to the physical and

ecophysiological background.

In the chapter on energy fluxes it is shown that the non-visible thermal

radiation forms an important part of the total radiation budget. Several

ways for a succulent plant to avoid excessive thermal stress by

manipulating its radiation budget are demonstrated using field measure-

ments or theoretical calculations. Unfortunately it is not shown how

important the radiation budget really is for succulent plants and no figures

on the upper limits of heat stress are given. Very important are the energy

fluxes at night. It is shown that in nights with a clear sky, plant surfaces

soon cool down below the temperatures of the surrounding air.

This observation is of importance for the subsequent subchapter on

water fluxes. Numerous means of water loss and water uptake are

demonstrated using field measurements. It is perhaps the most interesting

observation that, in times of drought stress, many CAM-succulents are

able to compensate for up to 49 % of their daytime water losses at night

when the leaves take up water directly from the atmosphere. This process

is described as reverse transpiration and its driving force is a water

vapour gradient from the atmosphere to the substomatal cavities of the

leaf.

In this context the high osmolarity due to accumulation of soluble salts

in many Mesembryanthemaceae (but not in Crassulaceae!) might play

an important role, as it is involved in the translocation of internal water.

In the chapter on carbon fluxes the different modes of C02 fixation

are discussed. It is shown that, although CAM is the dominating

metabolism in succulents, the role of CAM remains doubtful. With respect

to the carbon budget, the authors propose to use the amount of water

that can be stored by the expenditure of one gram of organic matter as

a suitable quotient (succulence quotient) for the ecophysiological

characterization of a succulent plant.

In a short final chapter it is demonstrated how structural and ecophysio-

logical features can be placed in various feasible combinations that form

part of the whole life strategy.

It is not surprising nor important that in a book of such volume, one

will also find a few weak points, errors, missing literature, and misiden-

tifications. Furthermore, I feel that the ‘special reference to the Namib

desert’ (as expressed in the subtitle; in the discussion of climatic gradients

and in some measurements of the non-succulent Welwitschia mirabilis)

was not helpful, because the proper Namib with its summer rainfall

seasonality and palaeotropical flora in my view represents a completely

different biome compared to the winter rainfall Succulent Karoo and its

flora of warm-temperate origin, which is analysed in the book. Also,

it was surely not necessary to discredit the old ‘degree of succulence’

(Delf 1912) by stating that in succulents no saturating water content exists

(page 4), while on page 248 the ‘water content at full hydration’ is used

for the definition of the new ‘succulence quotient’.

In summary, this book represents an excellent analysis of the ecophysio-

logical peculiarities of succulent plants. For me this is the best existing

book on the plant ecology of the Succulent Karoo. Every botanist and

ecologist working in the region should know it, and, when planning a

trip to the Richtersveld, this would be the book one should take along.

N. JURGENS*

* Institut fur Allgemeine Botanik, Universitat Hamburg, Ohnhorststr.

18, D— 2000 Hamburg 52, Germany.

BOTHALIA

Volume 22,2

Oct./Okt. 1992

CONTENTS— INHOUD

1 . New combinations and resurrected names in Microcharis and Indigastrum (Fabaceae— Papilionoideae).

B.D. SCHRIRE 165

2 . Studies in the Justicia and Siphonoglossa (Acanthaceae) species of southern Africa: final conclusions.

K.L. IMMELMAN 171

3 . Cololejeunea cardiocarpa, an epiphyllous liverwort in southern Africa (Lejeuneaceae). S.M. PEROLD 177

4 . Notes on African plants:

Bryophyta. New and interesting records of mosses in the Flora of southern Africa area: 3. Miscel-

laneous acrocarpous taxa. J. VAN ROOY and S.M. PEROLD 195

Bryophyta. Notes on the moss flora of Zimbabwe. J. VAN ROOY 196

Fabaceae. A new species of Coelidium (Liparieae). A.L. SCHUTTE and B-E. VAN WYK .... 189

Fabaceae. Lessertia sneeuwbergensis, a new species from the Middelburg District of the central

Cape Province. G. GERMISHUIZEN 189

Fabaceae. Cyclopia squamosa (Podalyrieae), a new species from the southwestern Cape Province.

A.L. SCHUTTE 190

Geraniaceae. Typification of Pelargonium section Polyactium. P. VORSTER and G.L. MAGGS 195

Gesneriaceae. Notes on the genus Streptocarpus . T.J. EDWARDS, C. KUNHARDT and S. VENTER 192

Liliaceae/Asphodelaceae. Lectotypification of Apicra jacobseniana (Alooideae). G.F. SMITH 196

Pyxinaceae. A new species in the lichen genus Heterodermia , from coastal Namaqualand. F. BRUSSE 183

Ricciaceae. Lectotypification of Riccia crystallina. S.M. PEROLD 185

Stilbaceae. Stilbe verticillata, the correct name for the species previously known as Stilbe mucronata.

J.P. ROURKE 192

5. Systematic studies in the genus Mohria (Anemiaceae: Pteridophyta). III. Comparative sporangium

and spore morphology. J.P. ROUX 199

6. Richness, composition and relationships of the floras of selected forests in southern Africa. C.J.

GELDENHUYS 205

7. An analysis of the orchid flora of Mt Mulanje, Malawi. H. KURZWEIL 235

8. Die fitososiologie van die Bankenveld in the Grootvlei-omgewing, Suid-Transvaal. W.J. MYBURGH,

P.J.J. BREYTENBACH, E.K. THERON en G.J. BREDENKAMP 245

9. A phytosociological study of Signal Hill, Cape Town, utilizing both perennial and ephemeral species.

C. JOUBERT and E.J. MOLL 255

10. The recovery and dynamics of submerged aquatic macrophyte vegetation in the Wilderness lakes,

southern Cape. P.J. WEISSER, A.K. WHITFIELD and C.M. HALL 283

11. Flora and vegetation of the Mbonambi Beach Arcuate Scar on the Zululand dune barrier, Natal,

South Africa. P.J. WEISSER, E.C.A. SMITH, A.P. BACKER and S. VAN EEDEN 289

1 2 . Obituaries: Amy Frances Gordon Jacot Guillarmod, nee Hean (1911-1992). M.J. WELLS and E. BRINK.

Lucy Kathleen Armitage Chippindall (1913—1992). N.P BARKER 295

13. National Botanical Institute, South Africa: list of staff and publications 301

14. Book review 313

Abstracted, indexed or listed in AGRICOLA, Biological Abstracts, Current Advances in Plant Science, Current Contents, Field Crop Abstracts,

Forestry Abstracts, Herbage Abstracts, Excerpta Botanica, Revue of Plant Pathology, Revue of Medical and Veterinary Mycology and The Kew

Record of Taxonomic Literature. / Opgesom, in indeks opgeneem of gelys in AGRICOLA, Biological Abstracts, Current Advances in Plant Science,

Current Contents, Field Crop Abstracts, Forestry Abstracts, Herbage Abstracts, Excerpta Botanica, Revue of Plant Pathology, Revue of Medical

and Veterinary Mycology en The Kew Record of Taxonomic Literature.

ISSN 0006 8241

Reproduction and printing by Execuprint Printers, Adriana Crescent, Gateway Industrial Park, Verwoerdburg. Tel. (012) 66-5131. / Gepubliseer

deur/verkrygbaar van die Nasionale Botaniese Instituut, Privaatsak X101, Pretoria 0001, Suid Afrika. Kopieset: S.S. Brink (NBI). Reproduksie

en drukwerk deur Execuprint Printers, Adriana Singel, Gateway Industriele Park, Verwoerdburg. Tel. (012) 66-5131.