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Bothalia

A JOURNAL OF BOTANICAL RESEARCH

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

Obtainable from the National Botanical Institute, Private Bag XlOl, Pretoria 0001, Republic of

South Africa. A catalogue of all available publications will be issued on request.

BOTHALIA

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

the Union of South Africa. This house journal of the National Botanical Institute, Pretoria, is

devoted to the furtherance of botanical science. The main fields covered are taxonomy, ecology,

anatomy and cytology. Two parts of the journal and an index to contents, authors and subjects are

published annually.

Three booklets of the contents (a) to Vols 1-20, (b) to Vols 21-25 and (c) to Vols 26-30, are available.

STRELITZIA

A series of occasional publications on southern African flora and vegetation, replacing Memoirs of

the Botanical Survey of South Africa and Annals of Kirstenbosch Botanic Gardens.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

taxonomy or economic botany. Published; Nos 1-63 (many out of print). Discontinued after No. 63.

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

A series devoted to the publication of monographs and major works on southern African flora.

Published: Vols 14-19 (earlier volumes published as Supplementary volumes to the Journal of

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

This serial presents colour plates of African plants with accompanying text. The plates are prepared

mainly by the artists at the National Botanical Institute. Many botanical artists have contributed to

the series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian Condy, Betty Connell, Stella

Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700 plates), Claire Linder-

Smith and Ellaphie Ward-Hilhorst. The Editor is pleased to receive living plants of general interest

or of economic value for illustration.

Erom Vol. 55, twenty plates are published at irregular intervals.

An index to Vols 1-49 is available.

FLORA OF SOUTHERN AFRICA (FSA)

A taxonomic treatise on the flora of the Republic of South Africa, Lesotho, Swaziland, Namibia

and Botswana. 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.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the informa-

tion is presented in the form of tables and photographic plates depicting fossil populations. Now

available:

Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidiurn, 1983, by J.M. & H.M.

Anderson.

Molteno Formation (Trias.sic) Vol. 2. Gymnosperms (excluding Dicroidiurn), 1989, by J.M.

& H.M. Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, 1985, by J.M. &

H.M. Anderson. Obtainable from: A. A. Balkema Marketing, Box 317, Claremont 7735,

RSA.

Towards Gondwana Alive. Promoting biodiversity and stemming the Sixth Extinction, 1999,

by J.M. Anderson (ed.)

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 30,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

- - J rA.sjJtSE

IN3T1TUUT

NATIONAL

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ISSN 0006 8241

October 2000

Editorial Board

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

Acknowledgements to referees

Anderson, Prof. E.F. Desert Botanical Garden, Phoenix, USA.

Burrows, J.E. Private Nature Reserve, Lydenburg, RSA.

Campbell, Dr B.M. University of Zimbabwe, Harare, Zimbabwe.

Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.

Hilton-Taylor, C. lUCN/SSC UK Office, Cambridge, UK.

Killick, Dr D.J.B. c/o National Botanical Institute, Pretoria, RSA.

Leistner, Dr O.A. National Botanical Institute, Pretoria, RSA.

Le Maitre, D.C. CSIR, Environmentek, Stellenbosch, RSA.

Liebenberg, Prof. H. University of Pretoria, RSA.

Moll, Prof. E.J. South African Wildlife College, Hoedspruit, Northern Province, RSA.

Nelson, Dr E.C. Outwell, Wisbech, UK.

Perry, Ms PL. Bangor, Wales, UK.

Phiri, Dr P.S.M. Herbarium, University of Zambia.

Stuppy, Dr W. Royal Botanic Gardens, Kew, UK.

Thiede, Dr J. University of Cologne, Germany.

Verdcourt, Dr B. Royal Botanic Gardens, Kew, UK.

Weisser, Prof. PJ. University of Venda, Northern Province, RSA.

Westfall, Dr R.H. Range and Forage Institute, Agricultural Council, Pretoria, RSA.

Willis, C.K. National Botanical Institute, Pretoria, RSA.

Wilson, Ms K.L. National Herbarium of New South Wales, Sydney, Australia.

CONTENTS

Volume 30,2

1. Studies in the Sphaerocarpales (Hepaticae) from southern Africa. 3. The genus Riella and its local

species. S.M. PEROLD 125

2. FSA contributions 17: Casuarinaceae. C.M. WILMOT-DEAR 143

3. Three new species of Erica (Ericaceae) from Western Cape, South Africa. E.G.H. OLIVER and

I.M. OLIVER 147

4. Notes on African plants:

Convallariaceae. A new combination in Eriospermum. J.C. MANNING 157

Hepaticae and Zannichelliaceae. New records from an ephemeral pan, Blouvlei, in Western

Cape, South Africa. W.R. HARDING, S.M. PEROLD and R.P. GLEN 157

Pteridophyta. A new combination and new records for the Flora of Malawi. J.P. ROUX 155

5. Combining floristic and growth form composition in a gradient-directed vegetation survey of

Matjiesrivier Nature Reserve, Western Cape, South Africa. R.G. LECHMERE-OERTEL and

R.M. COWLING 161

6. Wetland vegetation of southern KwaZulu-Natal, South Africa. L. PERKINS, G.J. BREDENKAMP

and J.E. GRANGER 175

7. Wetland vegetation in the North-eastern Sandy Highveld, Mpumalanga, South Africa. PM. BUR-

GOYNE, G.J. BREDENKAMP and N. VAN ROOYEN 187

8. Miscellaneous notes:

Aloaceae. The conservation status of Aloe in South Africa: and updated synopsis. G.F. SMITH,

E.M.A. STEYN, J.E. VICTOR. N.R. CROUCH, J. GOLDING and C. HILTON-TAYLOR . . 206

Apocynaceae. Chromosome studies on African plants. 15. Periplocoideae. J.J. SPIES,

H.J.T. VENTER and S.M.C. VAN WYK 211

Poaceae. Chromosome studies on African plants. 14. Panicoideae. A. STRYDOM, J.J. SPIES

and S.M.C. VAN WYK 201

Picking up the pieces: Red Data Lists in southern Africa. J.S. GOLDING 213

9. Obituaries:

Otto Heinrich Volk (1903-2000). H.E. GLEN and S.M. PEROLD 215

In memory of S.W. Amell. Hepaticologist ( 1895-1970). G. EEN and S.M. PEROLD 218

Rosemary Charlotte Holcroft (1942-2000). D.J.B. KILLICK 221

Werner Rauh (1913-2000), one of the World’s most prolific authors on succulent plants.

W. BARTHLOTT and G.E. SMITH 223

10. Book reviews 225

1 1 . National Botanical Institute South Africa: administration and research staff 3 1 March 2000, publica-

tions 1 April 1999-31 March 2000. Compiler: B.A. MOMBERG 229

12. Guide for authors to 243

New species and combinations in Bothalia 30,2 (2000)

Crepidomanes mannii {Hook.) J.P Roux, comb, nov., 155

Erica humidicola E.G.H. Oliv., sp. nov., 149

Erica rimarum E.G.H. Oliv., sp. nov., 152

Erica rusticula E.G.H. Oliv., sp. nov., 147

Eriospermum flagelliforme (Baker) J.C. Manning, comb, nov., 157

IV

Bothalia 30,2: 125-142 (2000)

Studies in the Sphaerocarpales (Hepaticae) from southern Africa. 3.

The genus Riella and its local species

S.M. PEROLD*

Keywords: Hepaticae, Riella Mont, R. affinis M.Howe & Underw., R. alatospora Wigglesworth, R. capensis Cavers, Riellaceae Engl., R. echinospora

Wigglesworth, R. purpureospora Wigglesworth, Riellineae R.M.Schust., southern Africa, Sphaerocarpales Cavers, subgenus Riella,

subgenus Trabiitiella Porsild

ABSTRACT

A taxonomic account of the aquatic liverwort genus Riella Mont, its two subgenera and five local species is presented.

This comprises descriptions and illustrations of these taxa together with a distribution map and a key to the subgenera and

species. The taxa are classified in the order Sphaerocarpales Cavers, suborder Riellineae R.M.Schust. A description of the

order Sphaerocarpales and a key to the three local suborders are given in Perold (2000) in the present series.

INTRODUCTION

The genus Riella Mont, was first recorded from

southern Africa by Cavers (1903). Dried mud, containing

Crustacea, the raison d’etre for its collection, was taken

from a shallow pond near Port Elizabeth, Eastern Cape,

in 1897 by a Mr Hodgson and sent to Owens College,

Manchester (via Plymouth), where it was placed in a

small aquarium. In a few weeks a number of green

shoots had grown out of the mud. Upon fruiting, they

were recognized as hepatics and sent to Cavers at

Yorkshire College, Leeds. He identified and described

them as a new species, R. capensis.

In 1926 and 1932 Miss E.L. Stephens sent a number

of South African Riella plants together with algae and

samples of mud to Manchester. Three new species of

Riella from the Cape were isolated, one of which was

from Valkenberg Vlei, unfortunately without ripe spores.

A fourth species was from Schonken’s Salt Pan near

Brandfort in the Free State. These species were studied

and described by Wigglesworth (1937). Her descriptions

were subsequently supplemented by Proskauer (1954),

who also identified a Pocock specimen from a farm dam,

4 miles (6.4 km) from Grahamstown, as the widely dis-

tributed R. affinis, thus adding a fifth species to the

southern African records of Riella. Amell (1957, 1963)

did not record any new species, but listed two new col-

lections from Namibia, the whereabouts of which have

not been traced. Unfortunately, very few collections have

been made in the last 30^0 years.

The occurrence of these plants is stated to be rare and

sporadic, which can perhaps ‘be attributed to the fact that

minor changes in the environment can result in their dis-

appearance’ (Schuster 1992). Rapid urbanization in

southern Africa in recent years, has also led to the

destruction of natural habitats, particularly on the Cape

Flats.

* National Botanical Institute, Private Bag XI 01, 0001 Pretoria.

MS. received: 1999-12-10.

The gametophytes of Riella are delicate, short-lived

and highly susceptible to differing environmental condi-

tions, which, if not leading to their disappearance, may

cause major changes in their size and form. The spores,

however, are exceedingly resistant, surviving in the dried

state for years. With ornamentation that is regarded as

species-specific, spores are essential for identification, as

few Riella species show well-marked, distinguishing veg-

etative characters. Sometimes only the mode of branching

may be of some significance. This study was undertaken,

even in the absence of recent collections, because infor-

mation gained by means of newer techniques such as

SEM micrographs of the spores (and thalli) of southern

African species has not been published before. It also

completes this series of studies in the local Sphaero-

carpales, which were excluded in my treatment of the

Marchantiidae for the Flora of southern A frica (Perold

1999a).

MATERIAL AND METHODS

The same procedures as outlined in Perold (1999b)

were employed in the preparation of the material for

examination and photography by compound light micro-

scope and scanning electron microscope.

Throughout this treatment of the Sphaerocarpaceae

and Riellaceae I have used the term ‘stem’, although it

is usually referred to as ‘axis’ or ‘rib’. Some of the

species descriptions and illustrations of Riella provide

rather less detail than others, because of a lack of suit-

able material.

Differences and similarities between the five southern

African Riella species are presented in tabular form

(Table 1 ).

A description of the order Sphaerocarpales and a key

to the three local suborders, i.e. Monocarpineae,

Sphaerocarpineae and Riellineae, are provided in the

previous article, no. 2 (Perold 2000), in the present

series.

TABLE 1. — Comparison of southern African Riella species

126

Bothalia 30,2 (2000)

127

Suborder Riellineae R.M.Schust.: 32 (1958);

R.M.Schust.: 827 (1992).

Plants aquatic, usually entirely submerged, thallus

developing in vertical plane, bilaterally symmetrical in

plane of wing, secondarily asymmetrical. Ste?n (or axis)

slender, erect, elongate, simple or furcate, in section

ellipsoid or subround, invested with unistratose wing

along its dorsal side. Wing undulate or ruffled, thin, over-

arching stem at coiled apex. Scales unistratose, leaflike,

mostly dimorphic, the two forms not always easy to dis-

tinguish: lateral leaf scales basally attached to stem and

formed at juncture of wing and stem, at maturity on both

sides of wing and often associated with young involu-

cres; ventral leaf scales produced along ventral surface of

stem, smaller and often constricted in middle, with lam-

inar attachment to stem by single cell or row of cells.

Cells thin-walled, here and there with a single oil body.

Rhizoids generally only borne basally on stem, all

smooth-walled.

Asexual reproduction by gemmae from ventral side of

stem and similar to ventral leaf scales.

Dioicous or rarely monoicous. Antheridia individual-

ly developed along thickened, free margin of wing,

sunken in pockets, ovoid, on very short pedicel.

Archegonia single, when fertilized enclosed in flask-

shaped involucres, smooth or rarely fluted with parallel,

longitudinal ribs or lamellae, in acropetal sequence, usu-

ally to right and left of wing. Sporophyte with globose

capsule, short seta and spherical or ± uniformly wide

foot, cleistocarpous; spores released on decay of capsule

wall and involucre. Spores large, single, not permanent-

ly united in tetrads, brown or purple to red; distal face

ornamented with fine or coarse spines, tips truncate,

sometimes wider below and basally connected by mem-

branes, rarely with prominent wing; proximal face with

few to many, finer spines. Nutritive cells present, 4-

nucleate. Elaters absent.

Riellaceae Engl, in Syllabus der Pflanzenfamilien,

edn 1; 45 (1892); Schiffn.: 51 (1893) as Rielloideae;

Mull.Frib.: 314 (1951-1958); S.W.Arnell: 6 (1963);

R.M.Schust.: 836 (1992).

The Riellineae include only the family Riellaceae;

hence the details are not repeated in the subordinal

description.

Riella Mont, in Annales des Sciences Naturelles,

Ser. 3, Bot. 18: 11 (1852); Mull.Frib.: 314 ( 1951-1958);

R.M.Schust.: 836 (1992). Type: R. notarisii (Mont.)

Mont.

Duriaea Bory & Mont.: 1115 (1843), not of Merat:

432 (1829).

Maisonneuva Trevis.: 442 (1877).

Plants green, aquatic, erect in growth, 10-60 mm tall.

Stem (or axis) sparsely branched, slender, invested along

dorsal side with undulate or ruffled wing and on each

side with lateral leaf scales and ventrally with smaller,

ventral leaf scales. Rhizoids usually only at base of stem,

hyaline, smooth.

Dioicous or rarely monoicous. Antheridia in pockets

along free wing margin. Archegonia in flask-shaped

involucres, smooth or rarely with longitudinal ribs or

lamellae. Sporophyte globose. Spores not remaining in

tetrads, large, 100-120 pm diam.

Riella is unique among Hepaticae in that there is an

intermediate stage in thallus development, i.e. the juve-

nile stage, in which growth is initiated by an intercalary

meristem (Thompson 1941).

Key to two subgenera and locally occurring species

of Riella

la Involucral flasks surrounding sporophytes with 8, or some-

times more, longitudinal ‘ribs’ or lamellae; plants

monoicous. widespread; spores light brown, subglobose

subgenus Trabutiella Porsild (= section Plicatae hWoxgt)

1 . R. affinis

lb Involucral flasks surrounding sporophytes smooth, without

‘ribs’ or lamellae; plants dioicous, endemic to South

Africa; spores brown or purple to crimson, triangular or

subglobose subgenus Riella (= section Euriella Porsild

and section Laevigatae KWorgs)'.

2a Spores brown, with or without wings;

3a Spores winged 2. R. alatospora

3b Spores without wings:

4a Spores triangular; involucre ovoid, long-acuminate,

capsule occupying its lower ‘A 3. R. capensis

4b Spores subglobular; involucre ovoid, shortly acumi-

nate, capsule occupying its lower ±

4. echinospora

2b Spores purple or crimson, marginal webbing prominent . .

5. R. purpureospora

Subgenus Trabutiella Porsild in Botanisk Tids-

skrift 24: 323 (1902).

1. Riella affinis M. Howe & Underw. in Bulletin

of the Torrey Botanical Club 30: 221 (1903);

R.H.Thomps.: 110 (1940); ibid.: 845 (1941); ibid.: 275

(1942); Prosk.: 69 (1955); S.W.Arnell: 7 (1963); Magill

& Schelpe: 9 ( 1 979). Type: on bank of a reservoir, Tafira,

Grand Canary, June 1897, O.E. Cook 729 (US, Smith-

sonian Inst., presumed iso., fide Prosk.).

R. vishwanatai Pande et al.: 166 (1954). Type: India, Lake Latif Shah,

Uttar Pradesh, Misra 3590. Synonymy fide Prosk. ( 1954).

Plants erect (Figure 1 A, B) or semi-erect, wholly sub-

merged or sometimes partly exposed, delicate, up to 23

mm tall, stems simple or sparsely furcate, occasionally

with adventitious branching toward base, the latter gener-

ally attached to substrate by rhizoids. Stem slender, in

cross section (Figure 1C) slightly flattened on dorsal side,

rounded on ventral side, 155-175 pm or ± 8 cell rows

thick, 200-255 pm wide, outer cells small, ± ovoid,

12.5-17.5 X 22.5-27.5 pm, inner cells somewhat larger,

angular, 30.0-37.5 x 20.0-37.5 pm. Wing unistratose,

with margin entire, overarching stem apex and rounded

above, 2.4-3. 0 mm wide, narrowing below and undulate,

gradually disappearing toward base, becoming deeply

notched at sinuses (Figures lA, B; 2A), containing 1 or

128

Bothalia 30,2 (2000)

FIGURE 1. — Riella ajfmis, Pocock BOL20503. A, B, monoicous plants seen from side, with involucres (containing capsules) in acropetal

sequence along stem; at wing margin, antheridia (indicated by d) 1 or 2(3) in notches or more numerous in a row, side by side; C, c/s stem

and part of wing. D-G, lateral leaf scales [G, after Thompson (1941) with smaller cells containing a single oil body], H-J, ventral leaf

scales [J, after Thompson (1941 )]. K, much enlarged row of antheridia at margin of narrowed part of wing; L, sporophyte with envelop-

ing involucre and calyptra; M, surface view of seta, upper part up to 4-stratose, below unistratose, supported on bulbous foot; N, c/s calyp-

tra surrounding upper part of seta; O, detail of involucral rib (one of 8-10); P, c/s involucral wall (rather flattened) with ribs. Scale bars:

A, B, 2 mm; C, 200 pm; D-J, 250 pm; K, L, P, 500 pm; N, 50 pm; O, M, 100 pm.

more antheridial involucres; cells near stem 5- or 6-sided,

87.5-1 12.5 X 27.5-50.0 (tm, near margin smaller, mostly

4-sided, 27.5-35.0 x 25-^5 gm. Scales dimorphic: later-

al leaf scales (Figure ID-G) at juncture of wing and stem,

often associated with archegonia or young involucres, ±

regularly distributed along both sides of stem, unistratose,

linear-lanceolate or tapering apically, 540-730 x 100-150

pm, marginal cells 4- or 5-sided, 17.5-35.0 x 17.5-20.0

pm, smaller ones sometimes containing an oil body, cells

in body angular, 25-60 x 22.5-37.5 pm; ventral leaf

scales (Figure I H-J) obliquely attached to morphological

ventral side of stem, fewer and smaller than lateral leaf

scales, often constricted in middle, 350^00 x 170-190

pm, marginal cells ± rectangular in shape, 12.5-25.0 x

7.5-12.5 pm, some smaller ones with an oil body, cells in

scale body 4-6-sided, 32.5-35.0 x 20-25 pm. Rhizoids

arising from base, or along length of stem, hyaline,

smooth, 20-30 pm wide.

Asexual reproduction reportedly by gemmae, but not

seen in present investigation; said to develop periodical-

ly along stem and to be similar to ventral scales.

Monoicous and protandrous. Antheridia discharged in

specimens examined, involucres flask-shaped, ± 250 x

1 20 pm, pockets variable in number, discontinuous along

Bothalia 30,2 (2000)

129

FIGURE 2. — Riella affinis, Pocock BOL20503. SEM micrographs of plants. A, stem of plant with wing and ribbed female involucres; B, several

involucres; C, single involucre. A, x 8.3; B, C, x 25.6.

wing margin, if only 1 or 2(3), then in notches (Figure

lA, B) at irregular intervals, otherwise in a row (Figure

lA, K), with as many as 13 lying side by side, ducts

oblique, up to ± 125 pm long, opening through pores at

wing edge, width of wing here reduced (Figure IK).

Archegonia borne on dorsal surface of stem, on either

side at join with wing, in acropetal sequence. Involucres

(Figures IL; 2A-C) up to 8 or even 10, along stems,

ellipsoid-ovoid, ribbed, 1575-1700 pm long, 900-1000

pm wide across middle, including ribs, gradually con-

tracted and eventually occluded at beak, also narrowed

toward base, ribs (Figure lO) mostly 8 in number

(Figure IP), joined at apex, unistratose, undulate, almost

as long as involucre, 110-160 pm wide across central,

widest part, narrower above and below, cells 4-6-sided,

30.0-42.5 X 20-25 pm, at entire margin rectangular,

25^5 X 15.0-17.5 pm, cells in involucral wall densely

chlorophyllose, 4—6-sided, 35-50 x 30.0-37.5 pm. Stalk

very short, obliquely inserted on stem, internally occu-

pied by seta and foot, as well as surrounding 3- (or 4-)

stratose, basal calyptra wall, and externally covered by

lower part of involucral wall. Calyptra closely investing

mature capsule, hyaline, outer cells in surface view, 6- or

7-sided, 60.0-82.5 x 40-50 pm, crowned above by

archegonial neck, below closely surrounding seta and

foot; in cross section (Figure IN) cells in outer row larg-

er, 20-25 X 32.5-37.5 pm, in central row, 12.5-20.0 x 25

pm, in innermost row 12.5-15.0 x 15-20 pm. Arche-

gonial neck ± 100 x 37.5 pm, upper part purple, below

colourless, with 6 rows of cells and 4 neck canal cells.

Capsule ovoid to subglobose, 750-800 pm diam., wall

unistratose, thickenings absent, cells ± rectangular,

50.0-67.5 X 35-50 pm. Seta (Figure IM) ± 150 pm long,

dark red, upper end wider, 3- or 4-seriate, below ± 37.5

pm wide, uniseriate, soon becoming necrotic. Foot bul-

bous, multicellular, 150-170 x 140 pm, in cross section

with large, closely packed, roughly triangular, haustorial

cells, 55-70 x 27.5-35.0 pm, closely surrounded by 3- or

4- stratose base of calyptra. Spores 80-100 pm diam.,

including spines, light brown, without wing, subglobose;

FIGURE 3. — Riella affinis, Pocock BOL20503. SEM micrographs of spores. A, B, distal face; C, lateral view of part of distal face above and

proximal face below; D, E, proximal face; F, proximal face, with shorter spines, seen slightly from side, exposing larger spines of distal

face on the right. A, x 373; B, x 361; C, x 418; D, x 350; E, x 407; F, x 441 .

130

Bothalia 30.2 (2000)

distal face (Figure 3A-C) densely covered with ± 11

irregular rows of spines across diam. and ± 36 projecting

at periphery, 7.5-12.5 pm long, apices blunt, truncate,

rarely acute, basal membranes interconnecting spines

below quite faint, sometimes forming imperfect reticula-

tions; proximal face (Figure 3D-F) with triradiate mark

absent, spines small and fine, ± 2.5 pm long, not basally

interconnected. Nutritive cells no longer present in mate-

rial examined. Spore release by disintegration of capsule

wall, then of calyptra and finally of involucral wall.

DISCUSSION

Riella affinis is a widely dispersed species, originally

known from Grand Canary; then from Stanford Uni-

versity campus, California; Uttar Pradesh, India (as R.

vishwanatai); near Grahamstown, South Africa (Figure

4), and finally, from Israel (unpublished, according to

Lipkin & Proctor 1975). Schuster (1992) states that it

was also reported from Argentina by Hassel de

Menendez (1959), but this is incorrect, as R. affmis is not

mentioned in this paper, which deals with R. americana

M.Howe & Underw.

On account of its ribbed involucre, R. affinis is easily

recognised and has been placed in subgenus Trabutiella,

together with two other species from elsewhere, but it is

the only monoicous one. The two other species are R.

cossoniana Trabut (= R. paulsenii Porsild, placed in syn-

onymy by Lipkin & Proctor 1975) and R. garnimdiae

Hassel de Menendez.

Schofield (1985: fig. 15-2A, B) illustrates what he

calls Riella affmis. ‘A’ is of an ‘antheridium-producing

gametophore’ and ‘B’, ‘detail of marginal chambers with

antheridia’. He cites Wigglesworth (1937) as his source

for A and B. ‘A’ is actually an enlargement of Wiggles-

worth’s fig. 49 of a male plant of R. echinospora

Wigglesworth, whereas his ‘B’ is an exact copy of her

fig. 51 of the same species. Schofield does not cite a

source for his ‘C’, which illustrates a ‘sporophyte-bear-

ing gametophore’; the involucres are, however, without

ribs. It should be emphasized here, that Wigglesworth

FIGURE 4. — Distribution of Riella species in southern Africa. A, R.

affmis. ▲; R. alatospora, □;/?. capensis, ♦; R. echino.spora. ■;

R. purpureospora. •.

(1937) did not treat R. affinis, as it was first collected in

South Africa in 1953 by Pocock and recorded by

Proskauer (1954).

Thompson (1942) refers to ‘explosive discharge of

the antherozoids’ in both culture and in temporary

mounts, but Proskauer (1955) comments that he had

never observed explosive discharge of antheridia in

Riella. Proskauer also found wider variation in spore

size, 70-130 pm diam., than I did. He remarks that,

‘spore size is of limited diagnostic value in the order,’ an

admonition it would be well to keep in mind. Spore orna-

mentation, on the other hand, particularly as illustrated

on SEM micrographs, is crucial to correct identification.

Proskauer (1955) reports that the specimen collected

by Dr Pocock on 31 March 1953, was from ‘Farm dam,

4 miles from Grahamstown on Cradock Road ... about V4

mile down from road’. On the label of another Pocock

specimen collected on 10 May 1953 (BOL20505), this

locality (Figure 4) is given as ‘Dam on Table Hill Farm,

Cradock Road, 6 miles’. I visited the area in October

1999 and found the name of this farm now to be ‘Table

Fann, Hilton’ owned by the White family. I collected

some mud from the bed of the Palmiet River, where the

mostly dried up river runs under a bridge on the road

(R350) from Grahamstown. Subsequently the techniques

for cultivation, as described by Proctor (1972) and

Hassel de Menendez (1987), were followed, but without

success.

Specimen examined

Pocock BOL20503.

Subgenus Riella

Euriella Porsild in Botanisk Tidsskrift 24: 327 (1902).

2. Riella alatospora Wigglesworth in Journal of

the Linnean Society of London, Botany 5: 317 (1937);

Prosk.: 68 (1955); S.W.Arnell: 7 (1963); Magill &

Schelpe: 9 (1979). Type: Cape Town, original locality,

vlei at Salt River between main road and railway line,

legit E.L. Stephens, atque usu sporarum illae originis

coluit G. Wigglesworth (BOL!; MANCH!) (? type not

designated).

Plants erect, 20-35 mm tall, some with stem simple,

others once-furcate near base and bilateral, with 2 nearly

equal shoots connected below by intermediate membra-

nous portion (Figure 5A), occasionally with clusters of

branches, possibly developed from numerous adventi-

tious shoots. Stem (Figure 5D) in cross section ± ovoid or

slightly flattened dorsally and rounded ventrally, 350-

400 pm or 9 cell rows thick, 280-300 pm wide, outer

cells rectangular or isodiametric, rather smaller, 25-50 x

22.5-37.5 pm, inner cells angular, 50-60 x 42.5-50.0

pm. Wing unistratose, sometimes bistratose at join with

stem, margin slightly eroded, overarching stem apex,

1. 8-3.4 mm wide, undulate, narrowing below and soon

disappearing; cells near stem 5- or 6-sided, 62.5-95.0 x

37.5^2.5 pm, near margin smaller and mostly 4-sided,

Bothalia 30,2 (2000)

131

FIGURE 5. — Riella alatospora, E.L. Stephens BOL26400. A, male plant once furcate near base, with numerous antheridia along wing margins

(after Wigglesworth 1937); B, female plant with 3 involucres; C, wing margin with several rows of smaller cells; D, c/s stem and part of

wing; E-K, lateral leaf scales; L, M, ventral leaf scales; N, sporophyte with stalked involucre; O, mouth of involucre from above; P, c/s

foot; Q, c/s stalk; R, surface view of cells in involucral wall; S, cells in capsule wall. Scale bars: A, B, 2 mm; C, O, P, R, 50 pm; D, 200

pm; E-M, 250 pm; N, 500 pm; Q, S, 100 pm.

132

Bothalia 30.2 (2000)

FIGURE 6. — Riella alatospora, E.L. Stephens BOL26400. SEM micrographs of plants. A, several involucres along stern; B, close-up view of

mature involucre raised on stalk; C, leaf scales along stem. A, x 8.3; B, x 18.4; C, x 45.5.

25-45 X 22.5-25.0 ptm, sometimes outer 5, 6 or more

rows (Figure 5C) without chloroplasts; several smaller

cells, ± 20 X 20 pm, filled with an oil body, scattered

throughout wing. Scales dimorphic (Figure 6C): lateral

leaf scales (Figure 5E-K) often paired, regularly and

obliquely attached along stem, some bluntly triangular in

shape, others irregular, 400-720 x 410-720 pm, margin-

al cells 4- or 5-sided, 25^5 x 15-30 pm, cells in body

5- or 6-sided, 27.5-50.0 x 22.5^2.5 pm, smaller cells

scattered throughout, 17.5-22.5 x 17.5-22.5 pm, each

containing an oil body; ventral leaf scales (Figure 5L, M)

fewer and smaller than lateral leaf scales, shape variable,

160-280 X 180^00 pm, marginal cells 4- or 5-sided,

40.0^7.5 X 27.5^2.5 pm, in between with occasional

smaller, wedge-shaped cells ± 25 x 25 pm in largest

dimension, containing an oil body, cells in body angular,

42.5-67.5 X 37.5^5.0 pm.

Dioicous. Male plants same size as female plants or

often smaller. Antheridia numerous, in a single, linear

series in pockets along thickened wing margin.

Archegonia near apex of stem, on both sides of wing.

Involucres (Figures 5B, N; 6A, B) up to 5 produced in

acropetal sequence along stem, obovoid, ± smooth, up to

2125 pm long, 1450 pm wide across widest part, nar-

rowing upwards to beak, ±175 pm wide and surrounded

by as many as 13 crowded, slightly projecting cells

(Figure 50), below also contracted toward stalk, cells in

unistratose involucral wall (Figure 5R) 5- or 6-sided,

42.5-57.5 X 32.5^2.5 pm. Stalk fleshy below, oblique-

ly attached to stem, ± 350 x 350^30 pm, in cross sec-

tion (Figure 5Q) cells in outer row angular, 50-55 x

30.0^7.5 pm, inner cells ± ovoid or angular, ± 50 x

45-60 pm, foot resting on internal cells of fleshy part.

Calyptra bi- or tristratose, outer layer of cells in surface

view polygonal, 37.5-65.0 x 35.0-67.5 pm, above

archegonial neck remaining attached, ± 100 x 40 pm.

Capsule globose, 1050-1150 pm diam., cells in wall

(Figure 5S) angular, 50-80 x 30-50 pm. Seta very short,

50-60 pm long, dark red, expanded above, up to ± 165

FIGURE 7. — Rietta alatospora, E.L. Stephens BOL26400. SEM micrographs of spores. A, B. distal face; C, lateral view of part of distal face; D,

E. proximal face with prominent wing; F. lateral view of proximal face, part of wing and some spines on distal face. A, x 367; B. D, x 344;

C, x394; E, x 329; F, x4l3.

Bothalia 30,2 (2000)

133

|im wide and 4-seriate, below only 50-55 )am wide and

uniseriate. Foot not bulbous, width nearly uniform along

its length, 420^30 x 220-240 gm, in cross section sur-

rounded by calyptra (Figure 5P). Spores 105-125 gm

diam., including prominent discoid wing, ± 20 gm wide,

increasing to ± 27.5 gm wide at angles, golden brown, ±

triangular; distal face (Figure 7A-C) with ± 11 or 12

irregular rows of spines across diam., 5. 0-7. 5 gm long,

frequently dilated above, truncate, extending onto wing

and ± 32 projecting beyond margin, 5. 0-7. 5 gm between

spines, central ones linked by basal connecting mem-

branes forming indistinct reticulations; proximal face

(Figure 7D-F) somewhat raised as centrally flattened

dome, without triradiate mark, medianly with irregularly

spaced papillae and marginally sprinkled with granules

which extend onto wing, laterally surrounded by ± con-

cave wing, with radiating striations and marginally pro-

jecting spines.

DISCUSSION

Wigglesworth (1937) reports that there is a striking

difference from the other Riella species in the appear-

ance of the majority of cultured young plants of R.

alatospora because they become heart-shaped at the top,

instead of protruding at only one side as was usual in R.

purpureospora Wigglesworth. I cannot comment on this,

not having observed plants in culture.

In some of the specimens I examined, the marginal 5

or 6 rows of cells along the wing were without chloro-

plasts, which may perhaps be ascribed to the effects of

partial drying.

In Wigglesworth (1937) there is a typographical error

in the length of the plant as it is given in mm (3.5) instead

of cm. She gives the size of the spores as ± 120 pm,

whereas Proskauer’s (1955) measurements varied from

80 to 140 pm and my own from 105 to 125 pm.

Riella alatospora is easily distinguished by spores

with a prominent discoid wing. Whether it has survived

in ponds on the Cape Flats (Figure 4) is a matter of con-

jecture; at least it will not have been another victim of

‘anonymous extinction’ (Campbell 1989), thanks to the

laudable efforts of the ladies Stephenson and Wiggles-

worth.

Specimens examined

Stephens BOL26399-2640P, CC {= computer catalogue) 1627 (cul-

ture MANCH).

3. Riella capensis Cavers in Revue Bryologique 5:

81 (1903); Wigglesworth: 316 (1937); Prosk.: 68 (1955);

S.W.Amell: 7 (1963); Magill & Schelpe: 9 (1979). Type:

cultivated from mud collected at shallow pond in neigh-

bourhood of Port Elizabeth, leg. Hodgson in 1897

(MANCH 1630 = Manchester Museum, Owens College

22799, holo.!).

Plants erect, stems simple (Figure 8 A, B), or irregu-

larly pseudodichotomously branched, 10-30 mm tall,

usually with V-shaped, twin, winged shoots at base from

common stem, then repeatedly, and reputedly becoming

shrub-like with stalked, adventitious shoots, but not

observed in scanty, remaining material. Stem slender, in

cross section (Figure 8C) subround, slightly flattened

dorsally and ventrally, 200-300 pm or 8 cell rows thick,

300-350 pm wide, outer cells smaller, almost isodia-

metric, 25-35 x 27.5 pm, inner cells mostly larger,

rounded or angular, 25.0-37.5 x 30.0-42.5 pm. Wing

unistratose, but often bistratose at join with stem, over-

arching stem apex, where rounded and circinate,

1.5- 2. 8(^.0) mm wide, narrowing below, but not dis-

appearing, basally present, undulate; cells near stem 5-

or 6-sided, 62.5-87.5 x 25.0-32.5 pm, near margin

smaller, quadrate, 15.0-22.5 x 12.5-22.5 pm, scattered

throughout wing, smaller cells filled with an oil body

(Figure 8D). Scales dimorphic: lateral leaf scales crowd-

ed at apices of branches, further down sometimes

paired, but mostly distant and alternate, rounded to

obtusely triangular (Figure 8E-K), 430-750 x 360-800

pm, marginal cells ± rectangular, 22.5-37.5 x 27.5^7.5

pm, occasionally smaller and filled with an oil body,

cells in body of scale angular, 37.5-50.0 x 32.5^2.5

pm, those containing an oil body scattered about,

22.5- 25.0 X 32.5-37.5 pm; ventral scales smaller

(Figure 8L, M), few, roughly triangular or rounded,

300-380 X 310-330 pm, marginal cells rectangular

across, 20.0-32.5 x 35^0 pm, or in between, toward

base, ± 50 X 25 pm, with smaller cells containing an oil

body, cells in body of scale larger, 37.5-60.0 x

27.5- 35.0 pm, with an occasional oil cell present, ± 25.0

X 17.5 pm.

Gemmae not seen.

Dioicous. Male plants absent in material studied, but

described by Cavers (1903) as less branched (shrubby)

and robust than female plants. Involucres (Figures

8N-P; 9F), up to 5, produced in acropetal sequence

along same side of stem and only fairly rarely on the

other side as well, 1875-2675 x 1050-1350 pm, ±

smooth, ovoid-acuminate with gradual attenuation and

then long drawn-out toward beak, mouth located apical-

ly on slender, fmger-like projection, ± 200 x 100 pm,

fringed with papillae, below also narrowed toward stalk,

cells in involucral wall 5- or 6-sided, 32.5-50.0 x

27.5- 37.5 pm, toward base somewhat longer. Stalk

(Figure 80) rarely almost absent, generally 500-625 x

1 50-175 pm, obliquely attached to stem where it widens

(Figure 8N), mostly occupied by seta and foot, except

for extreme base. Calyptra persistent, tristratose.

Capsule globose, 750-825 pm diam., occupying lower

V3-V2 of involucre, wall (Figure 8Q) unistratose, yellow,

cells 4-6-sided, 55-80 x 45.0-82.5 pm. Seta 275-350

pm long, dark brown, expanded above and 4-seriate,

narrow below, uniseriate. Foot ± 225 pm long, yellow-

ish, hardly bulbous, gradually widening from ± 100 pm

above to ± 225 pm below. Spores 95-120 pm diam.,

including spines, without wing, light brown, ± triangu-

lar; distal face (Figure 9A, B) with 10-12 rows of spines

across diam. and 30-36 projecting around margin, a few

protruding from periphery of proximal face, though not

excluded in the count, mostly broadly conical, tapering

to an acute tip, but sometimes blunt, up to 10 or 12 pm

long, some basally connected by membranes not forming

distinct reticulations, between spines lightly sprinkled

134

Bothalia 30,2 (2000)

FIGURE 8. — Riella capensis, Hodgson MANCH1630. A, B, female plants with involucres mostly on same side of stem; C, c/s stem and part of

wing; D, cells in wing, few smaller ones containing an oil body; E-K, lateral leaf scales; L, M, ventral leaf scales; N, P, involucre con-

taining capsule, obliquely raised on stalk; O, involucre almost sessile; Q, cells in capsule wall. Scale bars: A, B, 2 mm; C, 200 pm; D, 100

pm; E-M, 250 pm; N, O, P, 500 pm; Q, 100 pm.

Bothalia 30,2 (2000)

135

FIGURE 9. — Riella capeiisis, Hodgson MANCH1630. SEM micrographs of spores and involucre. A, distal face; B, distal face seen slightly from

side; C, D, proximal face; E, proximal face partly from side, only two facets showing; F, apex of involucre collapsed, below obliquely

raised on stalk, paired lateral leaf scales at stem. A, x 352; B, x 367; C, x 382; D, x 394; E, x 375; F, x 25.

with some granules; proximal face (Figure 9C-E) slight-

ly raised, not flat, marginal bases of spines joined by

slightly striated webbing, from which individual, coni-

cal spines project outwards, triradiate mark faintly visi-

ble, papillae sparsely scattered over face, which is rather

roughened with tiny, fairly indistinct granules.

DISCUSSION

Cavers did not give a Latin description of his new

species, as it only became compulsory with the 1935

ICBN code (Briquet 1935); subsequently, Wigglesworth

supplied a Latin description in 1937.

Cavers (1903) described the stem as circular in cross

section, which probably would be more representative of

the species than my section of it (Figure 8C). Fie also

stated that he would describe the developmental stages of

R. capensis, but 1 have not found a reference to such an

article. He expressed the opinion that his new species

came nearest to R. helicophylla Mont, from Spain,

Algiers and Tunis.

Hassel de Menendez, in her 1959 paper, points out the

differences between R. americana and R. capensis, the

latter much branched, the male plants with up to 100 (or

more) antheridia, the female plants with up to 50 sporan-

gia on a single plant, spore diam. 80 pm and the spines 8

pm long. Sim’s (1926) record of it from Cape Town is

evidently incorrect (Wigglesworth 1937).

Proskauer (1955) assigns a Pocock specimen collect-

ed on 12 December 1952 at the Palmiet River, Table

Rock Farm, seven miles from Grahamstown on the

Cradock Road, to R. echinospora, but then refers to the

marginal spines of the spores as showing light webbing,

which, in my opinion, would place it nearer to R. capen-

sis. I have seen no such webbing in R. echinospora

spores, which are subround. His determination is there-

fore suspect.

Wigglesworth ’s (1937: fig. 7) illustration of the

involucre of R. capensis, is rather less attenuate toward

the beak, than those that 1 examined; her figure 9 of the

proximal spore face, suggests some webbing at the base

of the spines, whereas her figures 11 and 12 of 7?. echi-

nospora spores clearly rule out the possibility of any

webbing. Furthermore, she supposes that the course of

growth in R. capensis plants followed the same lines as

that of R. alatospora.

My visit to Port Elizabeth in October 1999 in an effort

to find more material of R. capensis proved unsuccessful.

It was recently brought to my attention by Dr W.R.

Harding, that Coetzer (1987) had reported R. capensis

from Rocher Pan on the west coast. This collection has

not been traced and the determination could not be veri-

fied.

Specimen examined

Hodgson MANCH1630 (22799).

4. Riella echinospora Wigglesworth in Journal of

the Linnean Society of London, Botany 5: 321 (1937);

Prosk.: 68 (1955); S.W.Arnell: 7 (1963); Magill &

Schelpe: 9 (1979). Type: Orange Free State, Brandfort,

salt pan, leg. Schonken (BOL!; MANCH!), presumed

iso., from sticker on packet held in BOL.

136

Bothalia 30,2 (2000)

FIGURE U).~Riella echinospora. Schonken BOL26029. A, male plant with row of antheridia along wing margin and numerous leaf scales along

stem; B, female plant with several involucres; C, apical part of wing and involucre raised on stalk; D, c/s stem; E— H, lateral leaf scales, 1,

J, ventral leaf scales; K-M, involucres; N, detail of apical part of involucre with mouth occluded; O, cells in lower part of involucral wall;

P, c/s involucral wall; 0, cells in capsule wall; R, c/s foot and 3-stratose calyptra. Scale bars: A, B, 2 mm; C, K-M, P, 500 pm; D, 200 pm;

E-J, 250 pm; N, O, 0, HK) pm; R, 50 pm.

Bothalia 30,2 (2000)

137

FIGURE 1 1 . — Riella echinospora, Schonken BOL26029. SEM micrographs of plants. A, B, female plants with involucres and leaf scales; C, leaf

scales along stem. A, x 19; B, x 8; C, x 44.

Plants erect (Figure lOA, B), up to 35 mm tall; those

developed from gemmae and remaining sterile, accord-

ing to Wigglesworth (1937), branching freely and larger

than fertile thalli. Stem (Figure lOD) slender, in cross

section ± ovoid, slightly flattened dorsally and ventrally,

150-225 pm or ± 6 cell rows thick, 230-250 pm wide,

cells angular to rounded, 30.0-37.5 x 25.0-42.5 pm.

Wing (Figure IOC) unistratose, 1. 6-2.1 mm wide, highly

undulate, narrowing below; cells near stem 5- or 6-sided,

85.0-137.5 X 42.5-50.0 pm, near margin smaller, 4- or

5-sided, 25.0-37.5 x 17.5-25.0 pm; at margin, wedged

between others, small cells, 12.5-17.5 x 10-15 pm, con-

taining an oil body. Scales dimorphic: lateral leaf scales

(Figures lOE-H; IIC) sometimes very numerous,

crowded in pairs along stem, often associated with

archegonia, oblong or tapering slightly toward apex,

concave, attached to stem by row of cells, near tip with

single mucilage cell, 400-510 x 160-350 pm, at margin

cells quadrate or rectangular, 25.0-32.5 x 25 pm, small

cells with oil bodies in between, ± 12.5 x 17.5 pm, inner

cells 4- or 5-sided, 30-35 x 37.5-50.0 pm, mostly larger

below, 45-55 x 30.0^2.5 pm; ventral leaf scales (Figure

101, J) very similar to lateral leaf scales, but attached to

stem by single cell.

Asexual reproduction by gemmae which are described

as constricted in the middle (Wigglesworth 1937).

Dioicous. Male plants (Figure lOA) generally smaller

than female plants. Antheridia in continuous series or

interrupted, along wing margin. Involucres 4 or 5 in

acropetal sequence along stem, ovoid, sometimes acumi-

nate (Figures lOK-M; llA, B), ± smooth, 1375-1675 x

950-1050 pm, tapering above, mouth sometimes still

occluded (Figure ION), surrounded by smallish, apically

rounded cells, 27.5-45.0 x 15.0-22.5 pm, below con-

tracted toward stalk, in cross section ± ovoid (Figure

lOP), cells in involucral wall (Figure lOO) 4-6-sided,

40-75 X 25-40 pm, larger toward base, 87.5-147.5 x

27.5^2.5 pm. Stalk ± 375 x 270 pm, obliquely attached

FIGURE 12. — Riella echinospora, Schonken BOL26029. SEM micrographs of spores. A, B, distal face; C, lateral view of part of distal face above

and proximal face below; D, E, proximal face; F, lateral view of part of proximal face above and distal face below. A, x 417; B, x 455; C,

X 53 1 ; D, X 409; E, x 394; F, x 489.

138 Bothalia 30,2 (2000)

FIGURE 1 3. — Riella purpureospora, Harding CHI 3724 (PRE). A, Furcate male plant; B, female plant with involucres near apex; C, c/s stem and

part of wing; I), dimorphic cells in wing with numerous chloroplasts, scattered smaller cells with oil body in each; E-L, lateral leaf scales;

M, N, ventral leaf scales; O, apical part of male plant with interrupted row of antheridia, proximal ones discharged; P, archegonium; Q, R,

involucres; S, capsule wall; T, surface view of seta and foot; U, e/s calyptra surrounding seta. Scale bars: A, B, 2 mm; C, 200 pm; D, S,

T, 100 pm; E-N, 250 pm; O, 500 pm; P, U, 50 pm; Q, R, 500 pm.

Bothalia 30,2 (2000)

139

FIGURE 14. — Riella purpureospora. Harding CHI 3724 (PRE). SEM micrographs of plants. A, B, apical part of male plant with antheridia near mar-

gin and leaf scales along stem; C, pseudodichotomous branching in male plant; D, female plant with 3 involucres and leaf scales along stem;

E, more enlarged lateral view of involucre; F, small part of stem with leaf scales. A, x 9.6; B, x 15.7; C, x 8.8; D, x 8.4; E, x 18.4; F, x 34.8.

to Stem, occupied by seta and foot. Calyptra mostly bis-

tratose. Capsule occupying lower ± V4 of involucre, sub-

globose, 775-850 pm diam., wall pale yellow, cells

irregular in shape and size (Figure lOQ), 30-90 x

40.0-52.5 pm. Seta ± 100 pm long, dark red, not expand-

ed above. Foot ± 250 pm long, width ± uniform through-

out its length, in cross section (Figure lOR), surrounded

by calyptra and up to 300 pm wide. Spores 87.5-97.5 pm

diam., including spines, without wing, light brown, sub-

globose; distal face (Figure 12A-C, F) fairly densely

covered with 13-16 irregular rows of spines across

diam., 50 or more projecting around periphery, but rather

difficult to count as several rows involved due to round-

ness of spore, mostly slender and acute, occasionally api-

cally truncate and slightly swollen or dilated at tips, up to

10 pm long, not basally connected, ± 5 pm between

spines; proximal face (Figure 12D-F) raised, often

indented in the centre, covered with ± 19 rows of small-

er and finer spines across, up to 4 or 5 pm long, in

between with some papillae, triradiate mark absent.

DISCUSSION

It is possible that Riella echinospora is more wide-

spread than just the Brandfort area, as Arnell (1957,

1963) also recorded it from a Volk collection in Namibia

at Flaribes, Marienthal, ‘In seichtem Wasser auf feinem

Sand, haufig’. Proskauer (1955) also reported that ‘a

sporeling with attached spore probably belonging to this

species was isolated during class work at Berkeley from

a culture prepared from soil gathered by Dr Pocock on

the Cape Flats’. Regarding his reference to the specimen

from ‘the Palmiet River, Table Rock Farm’, I have

already commented on its spores in my discussion of R.

capensis.

In October 1 999 on my way through the Free State to

the Eastern Cape, it was noticed that there were numer-

ous pans visible from the highway. It would be worth

investigating them for the presence of Riella plants.

Specimens examined

Schonken BOL26029', CC (= computer catalogue) 1632-1635',

1637-1640', 1642-1649 (cultures MANCH).

5. Riella purpureospora Wigglesworth in Journal

of the Linnean Society of London, Botany 5: 312 (1937);

Prosk.: 66 (1955); S.W.Amell: 7 (1963); Magill &

Schelpe: 9 (1979). Type: prope Cape Town, legit E.L.

Stephens, atque usu sporarum illae originis coluit G.

Wigglesworth (BOL!, MANCH!) (? type not designated).

Plants erect, 20-60 mm tall, stems sparsely to fre-

quently furcate, some branches again furcate, occasion-

ally with several pseudodichotomies close together, the

daughter stems growing new wings; stalked adventi-

tious shoots formed anywhere along parent stem

(Figures 13A, B; 14C). Stem in cross section (Figure

13C) subround, 330-350 pm or 10 cell rows thick,

350-380 pm wide, outer cells isodiametric, 25.0-37.5 x

20.0-32.5 pm, inner cells angular, mostly larger, 30-60

X 30^0 pm. Wing unistratose, but bistratose at join

with stem, overarching stem apex, 1.3-2.75 pm wide,

undulate, narrowing below and then disappearing alto-

gether, leaving basal part of stem wingless; cells near

stem 5- or 6-sided, 50.0-87.5 x 30-50 pm, near margin

smaller, 4- or 5-sided, 20.0-32.5 x 15-20 pm; scattered

throughout wing, numerous small cells filled with an

oil body (Figure 13D). Scales dimorphic (Figure 14F):

lateral leaf scales (Figure 13E-L) at irregular intervals.

140

Bothalia 30.2 (2000)

FIGURE 15. — Riella piirpureospora, Garside 6656 (BOL). SEM micrographs of spores. A, B, distal face; C, part of distal face; D, proximal face,

showing webbing at base of spines; E, one facet of proximal face; F, lateral view of proximal face, part of wing and some spines on dis-

tal face. A, X 340; B, x 455; C, x 382; D, x 344; E, x 524; F, x 554.

obliquely or vertically attached on both sides of stem,

often in pairs, oblong. Ungulate or tapering apically,

570-925 X 310-800 pm, unistratose, marginal cells

mostly 4-sided, 15-30 x 20.0-37.5 pm, with smaller

cells, ± 20.0 X 17.5 pm, containing an oil body, wedged

in between, cells in body of scale 4-6-sided, 25.0-57.5

X 20.0-52.5 pm, scattered about in between, smaller

cells ± 25 X 25 pm, each with an oil body; ventral leaf

scales (Figure 13M, N) fewer and rather smaller, irreg-

ularly shaped or bluntly triangular, 270-450 x 320-480

pm, marginal cells rectangular or ± isodiametric,

25.0-37.5 X 17.5-32.5 pm, with smaller cells contain-

ing an oil body wedged in between, cells in scale body

4-6-sided, 25^5 x 25-30 pm, and in between an occa-

sional smaller cell with an oil body.

Dioicous. Male plants somewhat smaller than female

plants. Anthehdia numerous, in a single, linear series

(Figure 14A, B), in pockets of up to 23, in acroscopic

sequence along wing margin, but sometimes interrupted

(Figure 130), ovoid, 350 x 250-270 pm, discharging

through individual ducts, mostly sloping toward and

opening by pores at edge of wing; cells in wing covering

antheridia rather larger than those at periphery of

antheridia. Archegonia (Figure 13P) at maturity with 4

cover cells at apex of neck, these swollen and separating

from each other, leaving neck open for entrance of

antherozoids. Involucres (Figures 13Q, R; 14D, E) up to

7 produced in acropetal sequence along stem, pyriform, ±

smooth, 2375-2575 pm long, ± 1600 pm wide across

widest part, abruptly narrowing to beak, ± 200 pm wide

and surrounded by ± 12 cells in an irregularly protruding

row, below also contracted toward stalk, cells in involu-

cral wall 5- or 6-sided, 40-60 x 32. 5-^5. 0 pm. Stalk ±

375 X 250 pm, obliquely attached to stem, upper ± 125

pm occupied by basal part of foot. Calyptra multi stratose.

in cross section up to 4 layers of cells surrounding inner

haustorial cells of foot (Figure 13U). Capsule globose, up

to 1200 pm diam. at maturity, wall red or mauve, unis-

tratose, cells (Figure 13S) 4- or 5-sided, 42.5-70.0 x

30.0-47.5 pm. Seta (Figure 13T) ±210 pm long, dark

red, upper ± 100 pm expanded, funnel-shaped, narrow

below, only ± 35 pm wide, uniseriate. Foot (Figure 13T)

± 260 pm long, gradually expanding from narrow upper

part to ± 200 pm wide below. Spores 82.5-117.5 pm

diam., including spines, without wing, purple or red, ± tri-

angular; distal face (Figure 15A-C), with ± 12 rows of

spines across diam. and 25-30 projecting around margin,

mostly stout and truncate, rarely acute, 7.5-10.0 pm long,

5-10 pm between spines, basally connected by mem-

branes forming irregular reticulations; proximal face

(Figure 15D-F) raised, not flat, marginally with basally

webbed spines, sometimes webbing very prominent,

appearing almost wing-like, 7.5-12.5 pm wide, with

spines projecting outwards from it, triradiate mark occa-

sionally nearly complete, but mostly only present toward

angles, rest of face irregularly dotted with up to 15 low

spines per facet.

DISCUSSION

So far, R. piirpureospora is the only species to have

been collected in recent times, viz. by Dr W.R. Harding.

It is quite a robust plant and is probably the easiest species

to identify because of its purple or red spores and capsule

wall, as well as the pronounced webbing between the

bases of the marginal spines on the proximal spore face.

The reference to the separation of the four cover cells

of the archegonial neck in my description is from

Thompson ( 1 942) who referred to R. affinis, but it is

equally applicable to other species, i.e. R. piirpureospora.

Bothalia 30,2 (2000)

141

The Harding specimen was collected at Blouvlei, Cape

Town vicinity (Figure 4), in an ephemeral pan which con-

tains water between April and September, together with

Pseudalthenia aschersoniana and Bolboschoenus mar-

itimus. The pH of the water was 9.6, the alkalinity (as

CaCOs) 246 mg per litre and the salinity (as Na) 3813 mg

per litre (W.R. Harding pers. comm.).

Specimens examined

Garside 6656 (BOL); Harding CHI3724 (PRE); Stephens CC (=

computer catalogue) 1613-1626, 1631, 1636 (cultures MANCH);

Walgate 999 (BOL).

Riella sp. — from Valkenberg vlei, Wigglesworth, in Journal of the

Linnean Society of London, Botany 5: 324 (1937); Prosk.: 68 (1955).

Cape Town, Valkenberg Vlei, legit E.L. Stephens BOL2603P.

This species has not been treated in this study, as there

are no ripe spores in the original collection and it has not

been collected again. As mentioned in the introduction,

the spore ornamentation is essential for correct identifi-

cation of Riella species.

ECOLOGY

The five Riella species known from southern Africa,

are from widely scattered localities, ranging from the

summer rainfall area of central Free State (R. echinospo-

ra), to the winter rainfall areas of the Cape Flats (R.

alatospora and R. piirpnreospora), in Western Cape, and

extending to parts of Eastern Cape (R. affinis and R.

capensis), which receive sparse summer and winter

rains. The vegetation types in these localities, according

to Low & Rebelo (1996), are the following: central Free

State: Dry Sandy Highveld Grassland; Western Cape,

Cape Flats: Sandplain Fynbos; Eastern Cape, Port

Elizabeth area: Mesic Succulent Thicket; north of

Grahamstown: Eastern Mixed Nama Karoo.

Riella species grow in temporary or permanent pools,

vleis or intermittent streams, containing fresh or brackish

water. Hassel de Menendez (1987) on the other hand,

found that Argentinian Riella species did not grow in

temporary dry ponds, but rather in lakes, some of which

are artificial. Riella thalli cannot, however, withstand

desiccation, even for a short while.

It is thought that Riella spores may be transported by

wind or by birds (Hassel de Menendez 1987). Schuster

(1992) is of the opinion that it is unlikely, although theo-

retically possible, that thalli (and spores) may be dissem-

inated on the feet of wading birds from one site to the

next. Apparently spores can pass through their gut

unharmed and may be transported in this way over dis-

tances limited to under 80-100 km.

ACKNOWLEDGEMENTS

My sincere thanks to the curators of BOL and

MANCH for the loan of specimens; also to Dr W.R.

Harding for collecting and sending a specimen of R. pur-

pureospora, as well as a copy of Coetzer’s paper; to Dr

G.G. Hassel de Menendez for kindly sending reprints of

her papers on Riella species and to Mr F. White, Table

Farm, for his assistance. My thanks to Ms G. Condy for

the drawings, Mrs A. Romanowsky for developing and

printing many photographs and to Ms D. Maree for typ-

ing the manuscript.

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COETZER, A.H. 1987. Succession in zooplankton and hydrophytes of

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(Hepaticae) la segunda especie del genero hallado en

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HASSEL de menendez, G.G. 1987. Progress with knowledge of

the submerged genus Riella (Hepaticae) in Argentina. Symposia

Biologica Hungarica 35: 335-342.

HOWE, M.A. & UNDERWOOD, L.M. 1903. The genus Riella.

Bulletin of the Torrey Botanical Club 30: 214-224.

LIPKIN, Y. & PROCTOR. V.W. 1975. Notes on the subgenus

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1^68.

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Bothalia 30,2: 143-146 (2000)

FSA contributions 17: Casuarinaceae

CM. WILMOT-DEAR*

Genera 3(or 4) with about 90 species native to

Australia, S.E. Asia and Polynesia; many widely cul-

tivated. One species, behaving as if indigenous in

Madagascar and the East coast of tropical Africa is

thought to be self-sown from sea-borne Pacific fruits,

since infructescences can survive long periods of salt

water immersion; this same species has more recently be-

come naturalized in southern Africa. The family is easily

distinguished by the ‘Equisetiim-Vike' green deciduous

branchlets unlike those of any other tree.

1855000 CASU ARINA

Casuarina L., Amoenitates academicae seu disser-

tationes variae physicae 4: 143 ( 1759); Adans.: 481, 543

(1763); G.Forst. & J.R.Forst.: 105, t. 52 (1776); Friis:

499 (1980); Wilmot-Dear: 1 (1985); Wilmot-Dear &

M.G. Gilbert: 262 (1989); K.L.Wilson & L.A.S.Johnson:

100 (1989); Wilmot-Dear: 116 (1991). Type species: C.

eqiiisetifolia L.

Allocasuarina L.A.S.Johnson: 73 (1982); K.L.Wilson

& L.A.S.Johnson: 110 (1989) (see note below).

Trees, dioecious, rarely monoecious. Branches of two

kinds: main persistent woody branches bearing decidu-

ous, little-branched, green, thin ± flexible branchlets.

Leaves on both types of branch reduced to whorls of tri-

angular scales united at base, midribs decurrent to lower

node giving ribbed or grooved appearance to intemode;

leaf whorls alternating at consecutive nodes; on persis-

tent branches leaves becoming separated as stem thick-

ens. Inflorescences with closely-spaced, alternating

whorls of bracts similar to scale leaves. Male inflores-

cences terminal on deciduous branches (rarely also axil-

lary on woody stems), spicate, cylindrical but tapering to

sterile basal region; flowers sessile and solitary in axils

of bracts, enclosed by pair of lateral membranous-scari-

ous bracteoles; perianth segments 1 or 2 (anterior and

posterior), membranous, concave, enclosing single sta-

men and falling as stamen develops; mature anther

exserted. Female inflorescences axillary towards apex of

woody branches, short-stalked or ± sessile, globose or

ovoid; bracteoles as in male; perianth 0; ovary 1 -locular

with short terminal style; stigmas 2, long, slender, well

exserted at maturity. Infructescence cone-like and woody

due to enlargement and thickening of accrescent bracts

and bracteoles, the latter much the larger (often with dor-

sal protuberance) and forming a pair of ‘valves’ enclos-

ing fruit in a ‘cell’. Fruit a seed-like samara, compressed,

dark brown-black and shiny or pale grey-fawn and rather

dull, bearing large ± translucent wing with single longi-

tudinal nerve excurrent at apex. Seed ovoid, somewhat

flattened laterally, narrowly acute at apex, embedded in

spongy air-filled tissue.

A genus of ± 75 species, distribution as for family;

many widely cultivated as ornamentals and (especially in

the past) for timber; one, much-used for soil stabilization

in coastal area, now naturalized in parts of southern Africa.

Fifty-nine species, comprising all those with dorsally

thickened fruit valves and dark shiny seeds and including

all the shrubs, are now separated into a new genus,

Allocasuarina L.A.S.Johnson ( 1982); for the sake of con-

sistency with other African floras a broad concept of

Casuarina is retained here. Keys and very full descriptions

of both genera are given in Wilson & Johnson ( 1989).

Key to cultivated species and hybrids

la Ribs on both deciduous and persistent branches 4; scale leaves broadly triangular, free part 0.3-0. 6 mm long and

wide, adpressed; stigma yellowish; infrutescence cells 5 per whorl, separated by 7-10 mm of muricately-

pattemed surface formed from dorsal thickening of valves:

2a Deciduous branchlets 5-6 mm diam., strongly 4-angled C. decussata

2b Deciduous branchlets 3-4(5) mm diam.. almost terete C. toridosa

lb Ribs on deciduous branchlets (6)7-many; scale leaves on persistent branches narrowly triangular, at least 1 mm

long, ± reflexed; stigma red; infructescence cells (6)7-9 per whorl, dorsal thickening where present rarely as

above:

3a Infructescence cells 6 or 7 per whorl, separated by 4—6 mm of muricate or irregularly rugose surface formed from

dorsal thickening of valves; ribs on deciduous branchlets 6 C. fraseriana

3b Infructescence cells (6)7-9 per whorl, separated by 4 mm or less, dorsal thickening where present never as

above; ribs on deciduous branchlets (6)7-15:

4a Deciduous branchlets with 14 or 15 ribs; scale leaves adpressed, free for 0.5-0. 7 mm; samaras brownish, not

shiny C. glauca

4b Deciduous branchlets with (6)7-10 ribs; scale leaves and samaras various:

5a Deciduous branchlets 0.8-1 mm diam.; leaf tips free for 1.0-1. 3 mm; samaras dark brown, shiny . ... C. verticillata

5b Deciduous branchlets 0.4— 0.6(-0.9) mm diam.; leaf tips free for 0.3-0. 7 mm; samaras various:

6a Infmctescence valves with large, triangular, transverse, dorsal ridge ± 1 mm high; samaras rich red-brown,

shiny; male inflorescence 0.5 mm or less diam., whorls hardly overlapping, axis stalk often visible

between; scale leaves and male bracts ± uniformly pale C. littoralis

* Mrs C.M. Thomas, The Herbarium. Royal Botanic Gardens. Kew. Richmond, Surrey TW9 3AB, England.

144

Bothalia 30,2 (2000)

FIGURE \ —Casuahna equisetifolia. A, tlowering branch, x 0.8; B, portion of deciduous branchlet showing whorl of scale leaves, x 9.6; C,

male inflorescence, x 3.6; D, whorl of male flowers, x 12; E, male flower showing bract, bracteoles and perianth (anther removed), x 24;

F, male bract, x 24; G, male bracteole, x 24; H, male perianth segment, x 24; I, portion of woody branch with female inflorescence, x 3.6;

J, female flower, x 24; K, infructescence, x 2.4; L, samara, x 6. A-H, Ward 1331\ I-K, Stapleton 9398\ L, H. Haig 2364. (Reproduced

with the permission of the Director, Royal Botanic Gardens, Kew. Artist: Eleanor Catherine).

Bothalia 30,2 (2000)

145

6b Infructescence valves not dorsally thickened nor transversely ridged; samaras pale, not shiny; male inflores-

cence whorls usually crowded and overlapping; scale leaves various;

7a Scale leaves uniformly pale; intemodes with prominent angled ribs; infructescence valve apices free for

1. 5-3.0 mm, backs longitudinally wrinkled or ridged C. equisetifolia

7b Scale leaves with distinct transverse brown band; intemodes with inconspicuous rounded ribs; in-

fmctescence valves various:

8a Deciduous branchlets long, ( 1 10-)330-540 mm, and robust, 0.6-0. 9 mm diam., ribs 9; infmctescence

valve apices free if at all up to 1 mm, backs thus not easily visible C. jimghuhniana

8b Deciduous branchlets usually comparatively short, 90-200(-250) mm, and thin, 0.4— 0.6(-0.8) mm

diam., ribs various; infructescence valve apices free for 2-3 mm, backs easily visible:

9a Deciduous branchlets 0.4— 0.6 mm diam. with 7-9 ribs; infructescence valve backs with several

irregular longitudinal wrinkles C. cimninghamiana

9b Deciduous branchlets 0. 7-0.8 mm diam. with 10-13 ribs; infructescence valve backs with single

longitudinal ridge C. cunninghamiana x lobesa

Casuarina equisetifolia L., Amoenitates academ-

icae seu dissertationes variae physicae 4: 143 (1759);

Engl.; 159 (1895); C.H. Wright: 315 (1917); Battis-

combe: 68 (1926); Battiscombe: 83 (1936); Brenan; 122

(1949); Williams: 182, photo opp. p. 135 (1949);

Cufod.: 2 (1953); Dale & Greenway: 130, t. 26, photo

26 (1961); Hutch.: 142 (1967); J.H.Ross: 147 (1972);

R. A. Dyer: 29 (1975); Wilmot-Dear: 5 (1985); Wilmot-

Dear & M.G. Gilbert: 262 (1987); Wilmot-Dear: 120

( 1991). Type: Rumph., Herb. Amb. 3(4), t. 57 ( 1743).

Tree 7-25 m tall, monoecious; bark grey-brown.

Deciduous branchlets 0.5-0. 7 mm diam., ribs 7 or 8,

angled, prominent. Scale leaves (6)7 or 8 per whorl,

greenish or straw-coloured; on persistent branches free

to 2-3 mm, thickly chartaceous, much reflexed, pubes-

cent; on deciduous branchlets (0.4-)0.5-0,7 mm, thin-

ly chartaceous, adpressed, glabrous, margin ciliate.

Male inflorescences 10-30(-40) x 1. 2-2.0 mm (ex-

cluding anthers), whorls 15-25; bracts adpressed,

1.1-1. 8 X 0.4-0. 5 mm, pubescent outside; bracteoles

ovate, 0. 7-1.0 x 0.3 mm, acute, erose-dentate-ciliate.

Perianth segments 2, up to 0.7 x 0.4 mm, rounded.

Filaments exserted 1.5 mm; anthers 0. 8-1.0 mm, long,

brownish. Female inflorescences 3-5 mm long; stalk

3-10 mm long; bracts as male. Stigmas exserted 3-4

mm, red. Infructescence shortly cylindrical-subglo-

bose, apex flattened, 8-17(-25) x 10-16 mm; whorls

(6-)8 or 9(-12); valves 7 or 8 per whorl, 1.5-3. 2 mm

wide (but smaller and fewer towards apex), ± obovate,

acute to mucronate, apical 1. 5-3.0 mm free, gap

between pairs 0. 5-1.0 mm, valve backs with 2(3) lon-

gitudinal ridges. Samaras pale brown, dull, 5-7 mm

long, to 1 mm thick; wing 3. 5-4. 5 x 2-3 mm (those

from small valves smaller). Seeds slightly over 'A

length of whole fruit. Figure 1.

Naturalized on coast of southeastern KwaZulu-Natal

in sandy areas and on seashore; also planted in

Mpumalanga and Eastern Cape to stabilize coastal dunes

and as an ornamental street tree. Very ancient introduc-

tion (fruits ?sea-bome, self sown; see note on p. 143

above) in coastal East Africa and Madagascar; indige-

nous in Malaysia, Australasia and Polynesia; cultivated

widely throughout the world in tropical and warm tem-

perate regions. Figure 2.

The nodular roots fix nitrogen.

Vouchers: Ross 2306 (NH); Van der Meulen 1682 (PRE); Ward 1331

(NU).

FIGURE 2. — Distribution of Casuarina equisetifolia: naturalized. •;

cultivated. A.

Ornamental species and hybrids of cultivated origin

C. cunninghamiana Miq.

Deciduous branchlets 0.4-0. 6 mm diam., ribs 7-9,

inconspicuous. Scale leaves free for 0.3-0.5(-0.6) mm,

0. 1-0.2 mm wide with distinct dark red-brown band.

Male inflorescences 1.0-1. 5 mm diam.; bracts adpressed.

Infructescence s ovoid, 8-12 x 7-10 mm; valves with

several irregular longitudinal fine wrinkles. Samaras

pale brown, dull. Cultivated in Pretoria and Johan-

nesburg in Gauteng, Cape Town in Western Cape and

Grahamstown in Eastern Cape.

A hybrid between C. cunninghamiana and C. equi-

setifolia, which is not in the key, with prominent ribs but

dark-banded scale leaves, has been cultivated in Namibia

(Ombangua, Bethanie), Johannesburg, and in and near

Cape Town.

A hybrid of C. cunninghamiana, possibly with C.

obesa Miq. or C. glauca Miq. (see 9b of key), with

deciduous branchlets 0.7-0. 8 mm. diam., ribs 10-13

very inconspicuous, scale leaves as in C. cunninghami-

ana, male inflorescences ± 2 mm diam. with adpressed

bracts, and infructescences up to 18 x 12 mm with

valve backs bearing single well-defined off-central

dorsal ridge, has been cultivated in Cape Town,

Knysna and Johannesburg; it is also widespread in

Ethiopia.

146

Bothalia 30,2 (2000)

C decussata Benth. = Allocasuarina decussata (Benth.)

L.A.S.Johnson

Tree very similar to C. tomlosa (see below) but decidu-

ous branchlets 5-6 mm diam., strongly angled with very

prominent ribs. Cultivated in Cape Town.

C. fraseriana Miq. = Allocasuarina fraseriana (Miq.)

L.A.S.Johnson

Deciduous branchlets 0.8- 1.0 mm diam.; ribs 6, con-

spicuous. Scale leaves ± reflexed, free for 0.6-0.8 mm,

0.2-0. 3 mm wide, straw-coloured. Male inflorescences ±

1 mm diam., whorls little-overlapping. Infructescences

cylindrical, 20-30 x 15-25 mm, cells separated widely

by dorsal protuberances as in C. tomlosa. Cultivated in

Cape Town.

C. glauca Sieber ex Spreng.

Deciduous branchlets 0.7-0. 9 mm diam.; ribs 14 or

15, inconspicuous. Scale leaves adpressed, free for

0.5-0. 7 mm, ± 0.15 mm wide, banded as in C. cunning-

hamiana. Male inflorescences ± 2 mm diam., bracts ±

reflexed, free for 1. 6-2.0 mm, banded as leaves.

Infructescences cylindrical or depressed-ovoid, 10-15 x

10-15 mm, valve backs, where visible, with several fine

longitudinal ridges. Samaras pale brownish, dull. Culti-

vated near Pretoria and in parts of Cape (Simonstown,

Port Elizabeth, Queenstown).

C. jungliuhniana Miq. = C. montana Miq.

Deciduous branchlets 0. 6-0.9 mm diam., ribs 9.

Scale leaves free for 0.5-0. 6 mm, 0.2-0.25 mm wide

with distinct dark red-brown band. Male inflorescences

1.3-1. 5 mm diam.; bracts adpressed. Infructescences

spherical to elongate- or depressed-ovoid, valve backs

hardly visible, often with 1 or 2 longitudinal ridges.

Samaras light brown, dull. Cultivated in Cape Town.

C. littoralis Salisb. = C. suherosa Otto & Dietr. = Allo-

casuarina littoralis (Salisb.) L.A.S.Johnson

Deciduous branchlets 0.5 mm diam.; ribs 8 often

prominent. Scale leaves free for ± 0.7 mm, 0.2 mm wide,

straw-coloured (in dry state), apex sometimes indistinct-

ly darkened. Male inflorescences up to 0.5 mm diam.,

axis often visible between widely spaced whorls.

Infructescences long-cylindrical, valve backs with large

transverse ridge-like protuberances. Samaras rich red-

brown, shiny. Cultivated in Northern Province,

Johannesburg in Gauteng, Pietermaritzburg in KwaZulu-

Natal, and near Cape Town in Western Cape.

C. torulosa Aiton = Allocasuarina tomlosa (Aiton)

L.A.S.Johnson

Deciduous branchlets resembling permanent ones,

0.3-0.4(-0.5) mm diam., almost terete since ribs not

well-marked; ribs 4. Scale leaves broadly triangular, free

part 0. 3-0.4 mm long and wide. Male inflorescences up

to 0.6 mm diam., axis often visible between well-spaced

whorls. Infructescences cylindrical or depressed-ovoid,

15-22 X 15-18 mm; cells separated very widely by large

dorsal protuberances forming a regularly and deeply

muricate-pattemed surface. Samaras very dark brown,

shiny. Cultivated in Empangeni (eastern KwaZulu-

Natal), parts of Eastern Cape and near Cape Town.

C. verticillata Lam. = C. quadrivalvis Labill. = C. stricta

Aiton = Allocasuarina verticillata (Lam.) L.A.S. John-

son

Deciduous branchlets, 0. 8-1.0 mm diam., ribs 9-10,

inconspicuous. Scale leaves somewhat reflexed, free for

1.0-1. 3 mm , ± 0.2 mm wide, rather indistinctly darken-

ing towards apex. Male inflorescences 2-3 mm diam.,

whorls often little-overlapping. Infructescences ovoid-

elongate, 20-35 X ± 20 mm; valve backs much thick-

ened, sometimes with small, ± triangular dorsal thicken-

ing near base, usually irregularly longitudinally wrin-

kled. Samaras as in C. torulosa. Cultivated near Johan-

nesburg, Cape Town, Port Elizabeth, Grahamstown, and

Pietermaritzburg .

REFERENCES

ADANSON, M. 1763. Families des plantes. Part 1. Vincent. Paris.

BATTISCOMBE, E. 1926. Trees and shrubs of Kenya Colony: 68. The

Government Printer, Nairobi.

BATTISCOMBE, E. 1936. Trees and shrubs of Kenya Colony, edn 2:

83. The Government Printer, Nairobi.

BRENAN, J.P.M. 1949. Check-lists of the forest trees and shrubs of the

British Empire, Tanganyika territory No. 5, 2: 122. Imperial

Eorestry Institute, Oxford.

CUFODONTIS, G. 1953. Casuarinaceae. Enumeratio plantarum

aethiopiae spermatophyta. Bulletin du Jardin Botanique de TE-

tat, Bruxelles, Suppl. Vol. 23: 2.

DALE, I.R. & GREENWAY, P.J, 1961 , Casuarinaceae. Kenya trees and

shrubs: 130, fig. 26, photo 26. Buchanan’s Kenya Estates in

association with Hatchards, London.

DYER, R.A. 1975. The genera of southern African flowering plants 1:

29. Department of Agricultural Technical Services, Pretoria.

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

bargebiete, Theil C: 159. Reimer. Berlin.

FORSTER, J.G.A. & FORSTER, J.R. 1776. Characteres genericum

plantarum: 105. White, Cadell & Elmsly, London.

FRIIS, I. 1980. The authority and date of publication of the genus

Casuarina and its type species. Taxon 29: 499.

HUTCHINSON, J. 1967. Casuarinaceae. The genera of flowering

plants. Vol, 2: 142. Oxford University Press, Oxford.

JOHNSON, L.A.S. 1982. Notes on Casuarina II. Journal of the

Adelaide Botanic Gardens 6: 13-S7.

LINNAEUS, C. 1759. Amoenitates academicae .sen dissertationes

variae physicae 4: 143. Laurentius Salvius, Stockholm.

ROSS, J.H. 1972. Flora of Natal. Memoirs of the Botanical Survey of

South Africa No. 39: 147.

RUMPHIUS, G.E. 1743. Herbarium Amboinense 3,4: t. 57. Amster-

dam.

WILLIAMS, R.O. 1949. Useful and ornamental plants in Zanzibar and

Pemba: 1 82. Zanzibar.

WILMOT-DEAR., C M. 1985. Casuarinaceae. Flora of tropical East

Africa: 1-8.

WILMOT-DEAR., C M. 1991. Casuarinaceae. Elora zambesiaca 9,6:

116-120.

WILMOT-DEAR, C.M. & GILBERT, M.G. 1989. Casuarinaceae. Elora

of Ethiopia 3: 262.

WILSON, K.L. & JOHNSON, L.A.S. 1989. Casuarinaceae. Elora of

Australia 3: 100-202. Canberra.

Bothalia 30,2: 147-153 (2000)

Three new species of Erica (Ericaceae) from Western Cape, South

Africa

E.G.H. OLIVER* and I.M. OLIVER*

Keywords: Erica, taxonomy, new species. South Africa, Western Cape

ABSTRACT

Three new species of Erica L. from the mountains of Western Cape, South Africa, are described. E. rusticula

E.G.H.Oliv. with an indehiscent fruit, is confined to sandy places in the eastern Cold Bokkeveld, E. humidicola E.G.H.Oliv.

is a highly localized endemic in seepages in the Kogelberg Biosphere Reserve and E. rimarum E.G.H.Oliv. is restricted to

rock faces at high altitudes in the Hex River Mountains.

Erica rusticula E.G.H.Oliv., sp. nov. in grege

specierum olim in generibus minoribus fructibus inde-

hiscentibus positorum, Ericae bokkeveldiae E.G.H.Oliv.

affinis, sed ab ea antheris quatuor, calcaribus minimis,

filamentis latis, ovario villoso sine nectariis, tubo calycis

non crasso, foliis adaxialiter pubescentibus differ!.

TYPE. — Western Cape, 3219 (Wuppertal); northern

Cold Bokkeveld, Kleinveld 100, main valley NE of

Sneeukop, 1 240 m, (-CD), 6 May 2000, Oliver 11517

(NBG, holo.; BM, K, MO, NY, PRE, S).

Shrub compact rounded, 100-150(-300) x 100-300

mm, erect to semi-spreading, single-stemmed reseeder.

Branches', numerous main branches of short growth

10-15 mm long, terminating in an inflorescence or con-

tinuing vegetative growth, each with 2-5 short, recurved

secondary branchlets, 2-10 mm long, not from each

node, intemodes very short, ± 1 mm or less, all covered

with dense, very short, white reflexed hairs. Leaves 3-

nate, imbricate, ± 1. 6-2.0 x 1.0 mm, adpressed, elliptic,

subacute, adaxially shortly hairy and flattened, abaxially

glabrous and rounded, margins shortly ciliate when

young, sulcus narrow and closed at base; petiole

adpressed, ± 0.4 mm long, glabrous, shortly ciliate.

Inflorescence', flowers 3(6)-nate in 1(2) whorls at ends of

main and secondary branches, nodding, umbel-like when

2-whorled; pedicel 0.3 mm long, glabrous, pinkish red;

bract partially recaulescent and approximate to calyx, ±

1.0 X 0.5 mm, elliptic to oblong, subacute, pale pink to

pink and sometimes green-tipped, glabrous, margins

shortly ciliate and with a few subsessile, dark red non-

sticky glands towards apex, sulcus small, narrow; bracte-

oles 2 approximate and adpressed to calyx, otherwise

like the bract. Calyx 4-lobed; ± 1 .5 x 2.2 mm, shortly and

broadly funnel-shaped, hard and wax-like, pink; tube ± 1

mm long; lobes adpressed to corolla, ± 0.8 x 1.0 mm,

broadly elliptic, subacute, sulcus 0.2-0. 3 mm long, nar-

row. Corolla 4-lobed, 2.5 x 2. 5-3. 5 mm, shortly and

widely funnel-shaped, glabrous, pink, with thin semi-

transparent broad tube 1 mm long; lobes semi-spreading,

± 1.5 X 2.0 mm, broadly ovate, rounded or subacute.

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

7735 Claremont, Cape Town.

MS. received: 2000-07-03.

margins entire, with sessile, dry, dark-red glands and

short reflexed hairs. Stamens 4, subexserted, free; fila-

ment broad ± 2.5 x 0.5 mm, oblong, straight to slightly

curved apically, glabrous, white; anthers bilobed, ± 0.8 x

0.7 mm. obovate in outline in adaxial view, dorsally

attached near base, minutely appendiculate, spurs very

small, ± 0. 1-0.3 mm long; thecae ± 0.8 x 0.35 mm, ovate

in lateral view, erect, with a few papillae, otherwise

glabrous, almost black; pollen in monads. Ovary 2-locu-

lar, ± 0.7 X 0.6 mm, ovoid, complanate, obtuse, upper

three quarters covered with dense straight long hairs,

lower quarter glabrous, septum very thin and fragile,

nectaries absent; ovules 1 per locule, pendulous from a

subapical placenta; style exserted, ± 5 mm long, filiform,

curved, glabrous, reddish pink; stigma simple-truncate to

slightly widened. Fruit indehiscent, ± 1.2 x 2.0 mm,

ovoid, pericarp very thin and papery with exocarp thin,

mesocarp composed only of numerous crystals and

endocarp thin and brown. Seed one per fruit, ± 1.2 x 1.8

mm, ovoid, testa thin, very shallowly reticulate, yellow-

brown; cells 25-50 x 75-100 pm, irregularly elongate

with slightly thickened jigsaw-like anticlinal walls,

numerous small pits in inner periclinal walls. Figure 1.

This new species was discovered just as the revision

of the 84 species of indehiscent-fruited Erica was in

press (Oliver 2000).

The species is at first sight very similar to Erica

bokkeveldia E.G.H.Oliv. in the structure of the shrublet

and of the flower — the pink, open flowers with dark

exserted stamens and very long style, the petaloid pink

bract, bracteoles and calyx and the 2-locular ovary with

a single ovule in each locule. A close examination of the

flowers reveals that it differs from that species in having

only four stamens (not eight), minute spurs on the

anthers (not long, broad and serrated spurs), oblong,

broad filaments (not linear and thin), an ovary covered

all over with long thin hairs (not shortly hairy at the apex

only) and lacking any nectaries, and the corolla tube thin,

delicate and semitransparent (not hard and thickened). E.

bokkeveldia was included in the former genus Eremia as

Eremia calycina Compton (Oliver 1976).

There are also slight differences between the two

species in other organs — the leaves in E. rusticula are

148

Bothalia 30,2 (2000)

FIGURE 1. — Erica msticula. A. flowering branch; B, flowering branchiet; C, stem; D, leaf, abaxial view (left), adaxial view (right); E, flower;

F, bract; G, bracteole; H, calyx; I, J, stamen, front, side and back views; K, gynoecium; L, ovary cut open longitudinally; M, fruit; N, testa.

All drawn from type, Oliver 11517.

pubescent adaxially (not glabrous), narrower sepals

which are broadly elliptic and subacute (not very broad-

ly orbicular and broadly obtuse). In the seeds the testa is

slightly thicker, being yellowish with slightly thickened,

more jigsawed anticlinal walls and numerous small pits

in the periclinal walls (not transparent with undulate non-

thickened anticlinal walls and large-pitted periclinal

walls in E. bokkeveklia).

The texture of the corolla tube is significantly differ-

ent in the two species. In E. bokkeveldia the hard tube

envelops the delicate indehiscent fruit and remains

around it as a protection when the whole flower is shed

onto the ground. The fruit is extremely delicate with a

very thin, parchment-like pericarp and almost non-exis-

tent, transparent testa. In E. rusticula this tube is not hard

and therefore does not provide the same protection to the

fruit which is, however, a bit tougher with a slightly

thicker pericarp and with a slightly thickened seed testa.

Erica bokkeveldia is a somewhat isolated species in

the genus and is hypothesized to be allied to E. cetrata

E.G.H.Oliv. (Oliver 2000). We would suggest that E. rus-

ticula also belongs in this alliance. The position of these

species within the genus Erica, which has now been con-

siderably enlarged with the inclusion of all the indehis-

cent-fruited genera (Oliver 2000), is unresolved due to

the lack of any clear indications of a ‘natural’ infra-

generic division of the genus. All the indehiscent-fruited

species are retained at the end of the genus as an interim

measure. E. rusticula is therefore placed with E.

bokkeveldia as species number 45a (Oliver 2000).

The above three species are known from the Ceres

District with E. cetrata being more widespread to the

base of the Hex River Mountains and eastwards to the

Bonteberg near Touws River. The other two are Cold

Bokkeveld endemics. E. rusticula is confined to the

rocky ridges between Bokkeveld Sneeukop and Bloukop

in the northeastern areas, whereas E. bokkeveldia occurs

at lower altitudes just further south and to the west — they

do not co-occur. Both inhabit sandy or sandy-gravelly

flat areas where the other shrubs are also very low.

Erica rusticula has been found in five populations on

the Farm Klein veld 100, hence the epithet — rusticu-

lus = of the countryside [diminutive form], ‘Klein-veld’.

This farm covers the areas east of the Skurweberg Range

from Bokkeveld Sneeukop to Bloukop (Figure 2). In the

higher altitude populations the associated vegetation is

currently mature, rather old and undisturbed, whereas at

the type locality there is a fair amount of wind erosion

and farming disturbance on the pure quartzitic sandy

flats. In the latter area the plants are extremely old with

very gnarled woody stems. Unfortunately, due to the

extremely dry summer of 1999/2000 most of the popula-

Bothalia 30,2 (2000)

149

FIGURE 2. — Known distribution of Erica rusticula, •; and E. humidi-

cola, D.

tions were not flowering well except for the population

at the northeastern base of Sneeukop, hence its selection

as the type. The species was noted among Erica materi-

al collected for our interest by a team from the Protea

Atlas Project for which we are always very grateful.

Apart from variation in the size of the plants and their

flowering condition between the populations, no varia-

tion in morphological characters was noted.

The lack of nectaries in E. rusticula is an unusual fea-

ture since there is no accompanying enlargement of the

stigma that is usually associated with this condition. The

nectaries are clearly present in E. bokkeveldia and bees

have been seen visiting a population of the species in

Hartebeeskloof. No pollinating insects were noted visit-

ing the plants of E. rusticula and there was no indication

of pollen being shed when the plants were manually dis-

turbed. The pollination syndrome needs to be investigat-

ed in the light of the differences between the two species.

Elowering time: April and May. This is a very dry period

of the year in the area. However, in good years the nor-

mal early autumnal rains around the Easter period could

be very beneficial and trigger the flowering of the

species. This flowering time is in contrast to the

September/October flowering period for E. bokkeveldia.

Paratype material

WESTERN CAPE. — 3219 (Wuppertal): northern Cold Bokkeveld,

Bloukop area. (-CB), 30-04-2000. P. Holmes in NBG345123 (NBG);

ridge between Bloukop and Patryskop, N end SSW of Bloukop, 1 380

m, (-CB), 6-05-2000, Oliver 11508 (BOL, K. NBG, NY); ibid., neck at

S end ENE of Beacon 128. 1 440 m, (-CD), 5-05-2000, Oliver 11496

(BOL, NBG, P, PRE); ibid., S end ENE of Beacon 128, 1 540 m,

(-CD), Oliver 11506 (NBG).

Erica humidicola E.G.H.Oliv., sp. nov. (§Evanthe),

Ericae fervidae L. Bolus et E. pillansii Bolus maxime sim-

ilis, sed ab eis corolla breviore campanulata rosea (non

longa tubuloso-campanulata vel tubulosa rubra) differ!.

TYPE. — Western Cape, 3418 (Simonstown): Kogelberg

Reserve, SpinnekopsneskJoof, west-facing lower slopes

below Dwarsrivierberg, 180 m, (-BD), 22 September 1999,

Oliver 11353 (NBG, holo.; BM, K, MO, NY, PRE, S).

Shrub erect. 0. 5-1.0 m tall, bushy in open areas or

long and lanky in thicker, older vegetation, single-

stemmed reseeder. Branches: a few main erect ones,

100-300 mm long, mostly with continuous apical

growth, numerous secondary branches, 10-20 mm long

at every node, few, very short. mm long tertiary

branchlets, intemodes on main branches 8-10 mm long,

all branches with short spreading hairs. Leaves 4-nate, ±

3.5 X 0.6 mm subspreading, adaxially flattened, abaxial-

ly rounded, hirsute all over, sulcus narrow and closed at

base; petiole 0.6 mm long, adpressed, glabrous, ciliate.

Inflorescence: flowers 1-3 at ends of short lateral sec-

ondary and tertiary branchlets crowded along main

branches into loose spike-like synflorescence; pedicel 2

mm long, pubescent, red; bract partially recaulescent

about quarter to third way up pedicel, ± 0.6 x 0.3 mm,

ovate-triangular, subacute, puberulous, ciliate, with

small subapical sulcus, pink; bracteoles 2, positioned

slightly higher than bract, slightly longer than bract, oth-

erwise similar. Calyx 4-partite, segments ± 1 .5 x 1 .2 mm,

broadly ovate-elliptic, acute, adpressed to corolla, short-

ly and narrowly sulcate up to quarter their length, finely

puberulous, shortly ciliate, reddish pink. Corolla 4-

lobed, ± 4.5 x 3.0 mm, broadly campanulate, very finely

puberulous, dark pink; lobes ± 2.0 x 1.5 mm, broadly

rounded, entire, subspreading to spreading, devoid of

hairs towards margins abaxially. Stamens 8, included or

just manifest, free; filaments ± 2.5 mm long, linear-

oblong, straight or with slight S-bend, glabrous, white;

anthers bifid, dorsally attached near base, erect, sub-

quadrate in adaxial view, appendiculate, spurs ± 0.5 mm

long, pendulous, slightly decurrent along apex of fila-

ment, thick, entire, sparsely papillate, golden; thecae

dark brown, slightly parted but erect, ± 0.8 x 0.4 mm,

oblong-elliptic with slight nose in lateral outline, obtuse,

papillate-strigose on adaxial margins; pore about half

length of theca; pollen in tetrads. Ovary 4-locular, ± 1.3

X 1.1 mm, broadly obovoid, emarginate, with small basal

nectaries, hirsute; ovules 4 or 5 per locule, spreading

from placenta centrally placed on axis; style exserted, 3

mm long, cylindrical, glabrous; stigma capitellate. Eruit

a dehiscent capsule, ± 4.5 x 4.0 mm, valves free almost

to base but not spreading much, septa about 80% on

valves and 20% on columella. Seeds ± 0.5 x 0.4 mm,

subspheroid-ellipsoid, shallowly reticulate, golden brown;

cells subequally subquadrate, ± 45 x 55 |im. with slight-

ly undulate anticlinal walls and small pores in inner per-

iclinal walls. Figure 3.

This new species falls within a large group of water-

loving species with 4-nate leaves such as the large tubu-

lar-flowered Erica macowanii Cufino and the common

small-flowered E. parxhflora L. They all have in com-

mon, leaves with a thin cuticle, large epidermis and few

sclereids (< 5), nearly all nodes on the main branches

bearing secondary branchlets, terminating in 1^-flow-

ered inflorescences, the main branches continuing with

150

Bothalia 30,2 (2000)

FIGURE 3. — Erica humidicola. A, flowering branch; B, stem and whorl of leaves; C, flower; D, flower cut open laterally showing androecium

and gynoecium; E, bract; F, bracteole; G, sepal; H, anther, front, side and back views; 1, gynoecium; J, ovary cut longitudinally; K, cap-

sule with one valve removed; L, seed; M, testa cells. All drawn from type, Oliver 11353.

vegetative growth; corolla usually hairy and often finely

so as in E. feminanun E.G.H.Oliv., E. fervida and E. pil-

lansii. Most species in the alliance have no anther

appendages — the few exceptions being E. pannflora L.,

E. velitaris Salisb., the E. fervida/pillansii complex and

this new species, and most with no hairs on the ovary.

The closest relatives of E. humidicola appear to be the

showy scarlet-llowered E. fervida and E. pillansii, which

form a complex and are both also confined to the

Kogelberg Biosphere Reserve. It can, however, be distin-

guished by the pink colour of its flowers which have a

shorter open campanulate corolla (not long, tubular-cam-

panulate to tubular) and appendiculate anthers which

have papillate hairs on them (not smooth muticous

anthers). The species is placed here in ^Evanthe with

these two species despite the very short corolla tube, the

others being in ^Ephehus (E. parviflora) and ^Orophanes

{E. velitaris). This clearly points to the unsatisfactory

subgeneric system mentioned above under E. rusticula.

The species is very restricted in its distribution being

known thus far from only two small marshy/seepage

areas in Spinnekopsneskloof below Dwarsrivierberg

(Figure 2). In both populations the species grows in

stands of much taller, erect shrubs of species of

Leucadendron (Proteaceae) and Psoralea (Fabaceae). In

the northern population it occurs as the only heath, where-

as the southern population is dominated by taller plants of

the yellow-flowered E. campanularis Andr. In both there

are few plants, perhaps only several dozen in each. Two

other species which inhabit damp/wet places were noted

nearby, the small pink-flowered E. intervallaris at the

edges of the seeps and the tall, orange tubular-flowered E.

curviflora alongside the stream near the northern popula-

tion. Elowering time: September-October.

Bothalia 30,2 (2000)

151

A single plant of undoubted hybrid origin between E.

humidicola and E. campanularis was recorded in the

southern population by Mrs Amida Johns of the

Kogelberg Biosphere Reserve who accompanied us on

the investigation of the new species. The hybrid had

paler pink flowers with whitish tips and the narrower

campanulate corolla shape of E. campanularis.

The name is derived from the habitat preference —

hiimidus = damp, wet, incola or -cola - dweller.

The species was brought as a small branchlet by Ion

Williams in 1964 and again in 1969 by former colleague.

Charlie Boucher, during his survey of the Kogelberg

Reserve. It is the second species recorded as endemic to the

Spinnekopsneskloof, the other being E. vallis-araneanim

E.G.H.Oliv. which occurs at I 000 m at the head of the kloof

on steep south-facing slopes and which is extremely rare.

Paratype material

WESTERN CAPE. — 3418 (Simonstown): Kogelberg Reserve,

Spinnekopsnes, (-BD), 8-10-1964, Williams 534 (NBG); ibid., end of

Spinnekopsneskloof road, 1000 ft [± 300 m], 23-10-1969. Boucher 801

(NBG); ibid., 190 m, (-BD), Johns 38 (reference collection at Kogelberg

Reserve and at Harold Porter Botanical Garden, Betty’s Bay).

FIGURE 4. — Erica rimarum. A, flowering branch; B, stem with two whorls of leaves; C, flowering branchlet; D, leaf, abaxial view; E, flower; F,

bract/bracteole; G, sepal; H, corolla; I, flower cut open laterally showing androecium and gynoecium; J, stamen, side, back and front views;

K, gynoecium; L, ovary cut open laterally; M, fruit; N, capsule with one valve removed; O, seed; P, testa cells. A-L drawn from type,

Esterhuysen 14852', M-P from Esterhuysen 7796.

152

Bothalia 30,2 (2000)

Hybrid material {E. campanutaris x E. humidicola) — 3418

(Simonstown); Kogelberg Reserve, Spinnekopsneskloof, 200 m,

(-BD), 22-09-1999, Johns sub Oliver 11354 (NBG).

Erica rimarum E.G.H.Oliv., sp. nov. (%Eurystoma),

Ericae brevicaulis primo adspectu similis, sed E. navi-

gatoris maxime affinis et foliis 2-natis, corolla marroni-

na cyathiformi minori (non albida urceolata), antheris

inclusis (non manifestis) calcaribus in thecis (non in fil-

amento) dignoscenda.

TYPE, — Western Cape, 3319 (Worcester): Hex River

Mtns, crevices in rock face above stream on vlakte (flats)

on north side of Milner Peak, 3 000^ 000 ft [900-1 200

m], (-AD), 14 December 1948, Esterlmysen 14852

(BOL, holo.; NBG, PRE).

Shrub compact, gnarled and twiggy, 40-100 mm tall,

single-stemmed, stem very old and woody. Branches:

many main branches ± 10 mm long, erect or recurved,

mostly terminating in an inflorescence, occasional sec-

ondary branchlets up to 5 mm long, all glabrous. Leaves

2-nate, imbricate, ± 3.0 x 0.7 mm, erect, lanceolate-

oblong, adaxially concave, abaxially rounded, with acute

margins, glabrous, with a few non-sticky short-stalked

glands on basal margins and petiole, apex yellow-cuspi-

date, sulcus very narrow and closed at base; petiole

adpressed, + 0.5 mm long, glabrous. Inflorescence: flow-

ers 2-nate in one whorl at ends of main branches; pedicel

± 2.0 mm long, curved at base, glabrous, reddish; bract

partially recaulescent, about one third up pedicel, ± 2.0 x

0.6 mm, lanceolate, acute, glabrous, maroon, margin in

lower half with sessile or very short-stalked, non-sticky,

dark red glands, sulcus narrow, about one third its length;

bracteoles 2, attached about two thirds up pedicel, simi-

lar to bract. Ca/yx 4-partite, maroon; segments adpressed

to corolla, imbricate, ± 2.5 x 1.1 mm, ovate, acute,

glabrous, margins with small, sessile or short- stalked,

non-sticky, dark red glands, mainly in lower half, sulcus

narrow, about one third length of segment. Corolla 4-

lobed, ± 3.0 x 2.5 mm, cyathiform, maroon, thick in tex-

ture, glabrous; lobes 1.5 x 1.0 mm, rounded, entire, erect.

Stamens 8, included, free; filament ± 1.1 x 0.2 mm,

broadened towards base, with small apical sigmoid bend

below anther, glabrous, white; anthers ovate in adaxial

view, bilobed, dorsally attached, appendiculate, spurs

pendulous, ± 0.5 x 0.1 mm [as long as theca], serrate and

finely ciliate; thecae ± 0.5 x 0.4 mm, broadly elliptic-

rhombic in lateral view, dark brown, glabrous; pollen in

tetrads. Ovary 4-locular, ± 0.9 x 1 .0 mm, broadly ovoid,

emarginate, with well-developed nectaries around base,

glabrous; ovules ± 15 per locule, spreading from placen-

ta, central on axis; style included, 1.1. mm long, broadly

cylindical, tapering towards base; stigma capitate. Fruit

a dehiscent capsule, valves splitting about 70% with

septa on valves only and free from columella. Seed ± 0.5

X 0.3 mm, unequally ellipsoid with flat and rounded

sides, light brownish orange, shallowly reticulate; testa

thin, cells elongate, ± 100 x 30 pm, irregularly elliptic,

with slightly raised, jigsawed anticlinal walls and dense-

ly pitted, inner periclinal wall. Figure 4.

This new species is most similar to E. brevicaulis

Guthrie & Bolus and the recently described E. naviga-

toris E.G.H.Oliv. (Oliver & Oliver 1998) both of which

occur in the same mountain range. In growth and habitat

FIGURE 5. — Known distribution of Erica rimarum.

and flower colour it is most similar to the former which

has hairy margins to the leaf, a hairy ovary and short

anther appendages, which are lateral on the filament.

Despite the erect, much larger size of the plants and

flowers of E. navigatoris, the new species is probably

more closely related to it — it is like a miniature version

of that species. They share glabrous branches, cuspidate

leaves, non-sticky, slightly stalked red glands on the

edges of the bract, bracteoles and sepals which are all

similar in shape and position and the anthers with broad

irregularly serrate and ciliate appendages. E. rimarum

differs, however, in a number of characters — 2-nate

leaves (not 3-nate), corolla cyathiform and maroon (not

urceolate and white), anthers included (not manifest),

stigma included (not exserted) and anther appendages on

the anther theca (not along the apex of the filament).

Erica rimarum forms small, gnarled, woody shrub-

lets, growing on south-facing ledges and clefts on large

rocks or cliff-faces, hence the epithet, rima = a cleft or

fissure. The species is widespread on the peaks in the

southwestern part of the Hex River Mountain range from

Milner Peak to Waaihoek, where it grows at high alti-

tudes (Figure 5). Miss Esterhuysen showed the first

author the plants growing on the northern slopes west of

Milner Peak and at the time we discussed its probable

identity as E. brevicaulis. Many collections have been

made by her and also by Thomas Stokoe, both intrepid

collectors of high-altitude species, often growing in inac-

cessible rocky places in these rugged mountains. The last

collection of the species was that by Esterhuysen and the

first author in 1960. The record from Stettynsberg is

interesting, since no collections have been made in any

of the intervening mountains of the Du Toitskloof com-

plex. Flowering time: October-December.

The species possesses large nectaries around the base

of the ovary, which feature should indicate that the small

flowers are pollinated by insects.

Bothalia 30,2 (2000)

153

Paratype material

WESTERN CAPE. — 3319 (Worcester): Buffelshoek Twins, 5 500

ft [1 680 m], (-AD), 25-12-1942, Esterhuysen 8390 (BOL);

Buffelshoek Peak, 4 000 ft [1 220 m], (-AD),' 26-04-1942, Ester-

huysen 7796 (BOL); ibid., 6 500 ft [1 980 m], Esterhuysen 14849

(BOL, PRE); ibid., 2-01-1955, Esterhuysen 24046 (BOL, K); Castle

Rocks, 4 000 ft [1 220 m], (-AD), 5-12-1948, Esterhuysen 14708

(BOL, PRE); Waaihoek Mtn, 5 500 ft [1 680 m], (-AD), 15-12-1942,

Esterhuysen 8353 (BOL, NBG, NY); ibid., 30-05-1942. Stokoe 8558

(BOL); Milner Peak, 6 000 ft [1 820 m], (-AD), 27-12-1942, Ester-

huysen 8478 (BOL, NBG, PRE); ibid., 11-11-1960, Esterhuysen

28590 (BOL); ibid., N rocky slope and flats, 4 900 ft [1 490 m],

(-AD). 11-11-1960, Oliver 1009 (NBGl; Michell's Pass. (-AD). 12-

1929. Stokoe 2043 (BOL, PRE); Skilderberg [mountain unknown].

5 000-6 000 ft [1 520-1 830 m], (?AD), 12-1931. Stokoe 2642 (BOL);

Brandwacht, 6 000 ft, [1 830 m], (-CB), 26-11-1944, Esterhuysen

10986 ihOL); Stettynsberg, 3 000-4 000 ft [900-1 200 m], (-CD), 16-

12-1944, Esterhuysen 11447 (BOL),

REFERENCES

OLIVER. E.G.H. 1976. Studies in the Ericoideae. 1. The genera Eremia

and Eremiella. Bothalia 12: 29-48.

OLIVER, E.G.H. 2000. Systematics of Ericeae (Ericaceae-Ericoi-

deae): species with indehiscent and partially dehiscent fruits.

Contributions from the Bolus Herbarium 19: 1^83.

OLIVER, E.G.H. & OLIVER. I.M. 1998. Three new species of Erica

(Ericaceae) from South Africa. Novon 8: 261 -21 A.

'4.

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r

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Bothalia 30,2: 155-159 (2000)

Notes on African plants

VARIOUS AUTHORS

PTERIDOPHYTA

A NEW COMBINATION AND NEW RECORDS FOR THE FLORA OF MALAWI

This note updates the checklist of the Pteridophyta of

Malawi (Burrows & Burrows 1993) by recording some

recent collections.

HYMENOPHYLLACEAE

Crepidomanes mannii {Hook.) J.RRoux, comb.

nov.

Trichomanes mannii Hook, in Hook. & Baker. Synopsis filicum: 75

(1867).

Illustration: Komas: t. 7a, b (1994).

The classical bigeneric system of dividing the

Hymenophyllaceae into Hymenophylliim and Trichoma-

nes is not natural. The system proposed by Iwatsuki

( 1984) is followed here, thus necessitating the new com-

bination.

Crepidomanes mannii is a small and easily over-

looked fern that is widely distributed in the tropical parts

of continental Africa and Madagascar. Although wide-

spread in tropical Africa the species was first recorded

for the Flora zambesiaca area (Zambia) by Komas

(1976). The plants mostly grow on wet rocks or on the

lower parts of tree trunks in moist, shady forests.

MALAWI. — Zomba Plateau. Mandala Falls. Rou.x 2870 (NBG).

VITTARIACEAE

Antrophyum mannianum Hook., A second cen-

tury of ferns: t. 73 (1861).

Illustration: Schelpe: t. 30 ( 1970).

This rare fern occurs in East and West tropical Africa.

Until now, the species has only been known from one

collection made at Namuli in the Zambezia Province of

Mozambique (Schelpe 1970). This location could not be

found but it may be a typographical error for Nalume, a

river originating in the Gurue Mountains in the north-

western comer of Zambezia Province. Plants are mostly

found near streams on moist, deeply shaded rocks.

MALAWI. — Mt Mulanje, Ruo Gorge, Roux 2887 (MAL, NBG).

ASPLENIACEAE

Asplenium gemmascens Alston in Boletim da

Sociedade Broteriana, Ser. 2, 30: 10 (1956). Eigure 1.

Asplenium torrei Schelpe: 209 (1967).

Alston (1956) described A. gemmascens as differing

from A. hemitomnm Hieron. in the proliferous bud at the

apex of the rachis, the short-creeping rhizome, the herba-

ceous pinnae, and the conspicuous veins. In describing A.

torrei, based on a single collection made on the slopes of

Mt Nhandore in the Serra da Gorongosa, Mozambique,

Schelpe (1967) (erroneously) stated that this and A.

blastophorum Hieron. are the only proliferous African

species belonging to the A. aethiopicum complex. All these

species belong to Asplenium section Sphenopteris Mett.

I have studied the isotype of A. torrei in the herbarium

of the Universidade Eduardo Mondlana (EMU) in Maputo,

Mozambique, and found it to be identical to my collection

made on Mt Mulanje. The material fits both the descrip-

tions of A. gemmascens and A. torrei. As there are now dis-

tinctive characters to separate the taxa, A. torrei is placed in

synonymy. This supports the view of Pichi Sermolli ( 1985)

who also found the taxa to be conspecific.

Asplenium gemmascens is a widespread but not very

common species occurring in East and West tropical

Africa. Plants form small clonal clusters on moist, deeply

shaded rocks, or terrestrially, in evergreen forests.

MALAWI. — Mt Mulanje, Ruo Gorge. Roux 2901 (NBG).

ACKNOWLEDGEMENTS

I wish to express my thanks to the Southern African

Botanical Diversity Network (SABONET) Steering

Committee for financial support to visit Zimbabwe,

Malawi, Zambia and Mozambique. My gratitude also

goes to Mr Edwin Kathumba who so competently acted

as guide and assisted whilst visiting Malawi.

REFERENCES

ALSTON, A.H.G. 1956. New African ferns. Boletim da Sociedade

Broteriana, Ser. 2, 30: 5-27.

BURROWS, J.E. & BURROWS, S.M. 1993. An annotated checklist of

the pteridophytes of Malawi. Kirkia 14: 78-99.

HOOKER. W.J. 1861. A second century of ferns', t. 73. Pamplin,

London.

HOOKER. W.J. 1867. In W.J. Hooker & J.G. Baker, Synopsis filicum.

Hardwicke, London.

IWATSUKI. K. 1984. Studies in the systematics of filmy ferns. VII. A

scheme of classification based chiefly on the Asiatic species.

Acta Phytotaxonomica Geobotanica 35: 265-279.

KORNAS. J. 1976. Notes on African Hymenophyllaceae. 1. Tricho-

manes mannii Hook, new to the Flora zambesiaca area.

Bulletin du Jardin Botanique National de Belgique 46:

387-392.

156

Bothalia 30,2 (2000)

FIGURE 1. — Aspleniwri gemmascem, Roux 2901 (NBG). A, habit; B, abaxial surface of pinna; C, rhizome palea; D. section of C showing cellu-

lar structure; E & G, stipe paleae; F. section of E showing cellular structure; H, palea from abaxial surface of pinna. Scale bars: A, 20 mm;

B. 10 mm; C, E, G, 0.5 mm; D, F, FI, 200 pm. Drawn by J.P. Roux.

KORNAS, J. 1994. Filmy ferns (Hymenophylaceae) of Central Africa

(Zaire, Rwanda, Burundi). 2. Trichomanes (excl. subgen. Micro-

gonium). Fragmenta Florislica el Geohotariica 39: 33-75.

PICHI SERMOLLI, R.E.G. 1985. A contribution to the knowledge of

the Pteridophyta of Rwanda. Burundi, and Kivu (Zaire). Bulle-

tin du Jardln Bntanique National de Belgique 55: 123-206.

SCHELPE, E.A.C.L.E. 1967. New taxa of Pteridophyta from southeast

tropical Africa. Bolelin da Sociedade Broieriana, Ser. 2, 41:

203-217.

SCHELPE, E.A.C.L.E. 1970. Pteridophyta. In A.W. Exell & E. Launert,

Flora zambesiaca. Crown Agents, London.

J.P. ROUX*

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

7735 Claremont, Cape Town.

MS. received: 1999-05-18.

Bothalia 30.2 (2000)

157

CONVALLARIACEAE

A NEW COMBINATION IN ERIOSPERMUM

In 1 872 J.G. Baker described the species Anthericiim

flagellifortne from material at Kew that had been collect-

ed by Joseph Burke in South Africa on his trip through

the country in 1841-1842. The collection comprises

rather withered inflorescences, without leaves or root-

stock (Baker noted at the time that the leaves were per-

haps developed at a different time to the flowers). Baker

later transferred the species to Schizobasis, obviously

impressed by the persistent inflorescence axis which,

like that of Schizobasis, remains green and photosynthet-

ic for some time after the flowers have abscised. He dis-

tinguished it from the other species of Schizobasis which

he recognized by the erect, unbranched inflorescence.

Burke’s companion on the trip, C.F. Zeyher, also collect-

ed specimens of this taxon in the Magaliesberg, and the

sheet deposited in the South African Museum collection

(SAM 22787) bears the much more prescient determina-

tion " Eriospermttm ' in Zeyher ’s hand. Examination of

the Burke and Zeyher material reveals that it lacks the

spurred bracts and marcesant flowers which are charac-

teristic of Schizobasis and its other urgineoid allies. The

collections in fact accord in all respects with Eriosper-

mum abyssimcum Baker, probably the most common and

widespread species in the genus (Perry 1994). This

species was described by Baker in 1 876 in the same pub-

lication in which he transferred Anthericum flagellifonne

to Schizobasis . The name Anthericiim flagellifonne is

now the earliest available name for the species currently

known as Eriospermum abyssinicwn and is accordingly

removed from the synonymy of Schizobasis vohibilis.

where it was placed by Jessop (1977). The name

Eriospermum abyssimcum Baker becomes a taxonomic

synonym of Eriospermum flagelliforme. A full synono-

my of E. abyssimcum is given in Perry (1994).

Eriospermum flagelliforme (Baker) J.C. Manning,

comb. nov.

Anthericum flagelliforme Baker in Journal of Botany I: 140 (1872).

Schizobasis flagelliformis (Baker) Baker: 261 (1876). Type: South

Africa, Apies River (Aapages River), Burke s.n. (K!, holo.).

Eriospermum abyssimcum Baker: 263 (1876). Type: Ethiopia,

Gallabat, Gendua, Schweiiifurth 26 (K, lecto.; G), syn. nov.

REFERENCES

BAKER, J.G. 1872. Revision of the nomenclature and arrangement of

the Cape species ot Anthericum. Journal of Botany 1 : 1 35-141 .

BAKER. J.G. 1876. Revision of the genera and species of Anthericeae

and Eriospermeae. Journal of the Linnean Society 15: 253-363.

JESSOP, J.P. 1977. Studies in the bulbous Liliaceae in Africa: 7. The

taxonomy of Drimia and certain allied genera. Journal of South

African Botany 43: 265-319.

PERRY, PL. 1994. A revision of the genus Eriospermum (Eriosperm-

aceae). Contributions from the Bolus Herbarium 17: 1-320.

J.C. MANNING*

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

7735 Claremont, Cape Town.

MS. received: 2000-06-28.

HEPATICAE AND ZANNICHELLIACEAE

NEW RECORDS FROM AN EPHEMERAL PAN, BLOUVLEl, IN WESTERN CAPE, SOUTH AFRICA

Seasonal vleis and ephemeral pans were once common

in the vicinity of Cape Town, but have now all but disap-

peared as a consequence of urban development, freshen-

ing of previously saline systems or eutrophication.

Blouvlei, probably the last ephemeral pan within the

Cape Metropolitan Area, was incorporated into a business

park. Century City, during 1997 (Lochner & Rossouw

1997). As part of the Environmental Management Plan

for this development, an eight-hectare portion of the orig-

inal vlei environment was rehabilitated (alien removal)

and conserved. Blouvlei has been classified by the

Botanical Society as one of 37 Core Conservation Areas

on the Cape Flats (Maze & Rebelo 1999).

Associated water quality’

Surface water quality in the ephemeral pans on this

site is typically heavily humic-stained and saline, reach-

ing salinities in excess of 40 parts per thousand during

the last weeks in which the pans contain water.

Composition of the major ions is typically Na > Mg > Ca

> K, and Cl > SO4. At the time of the collections report-

ed on here, i.e. August 1998, concentrations of the major

ions were (in mg per liter; corresponding values for July

1999 in parentheses): K = 94 (55); Na = 3813 (2908); Ca

= 232 (174); Mg = 509 (400); Sulphate =1217 (1142)

and chloride = 7010 (4863). The pH of the water was 9.6

(8.5), and the alkalinity (as CuCOb) was 246 mg per liter.

HEPATICAE

Riella purpureospora

Riella purpureospora Wigglesworth (Sphaerocar-

pales) is an annual, salt-tolerant aquatic liverwort, first

described from the Cape Peninsula (Wigglesworth 1937;

Proskauer 1955). It was last collected in 1954 by Mr S.

Garside who worked on bryophytes at the Bolus Her-

barium, Cape Town. No collections were recorded dur-

ing the subsequent 44 years until the senior author found

158

Bothalia 30,2 (2000)

it growing in a restored ephemeral pan system at

Blouvlei, Goodwood, Cape Town. R. purpiireospora was

discovered for the first time during 1998, in a shallow,

exposed area of the pans containing no other hydro-

phytes. Although the fomiation of involucres was not

noted during 1998, prolific production of these structures

occurred for an extended period during 1999.

A short description of the species follows, but a

detailed account may be found in Perold (2000). ARiella

species was also recorded in Rocher Pan, West Coast,

during 1981/2 (Coetzer 1987); it had been identified as

R. capensis, but this has not been confinned.

Riella species are unique in being the only truly

aquatic liverworts. They are characterized by entirely

submerged, erect, up to 60 mm long stems, invested

along one side with a thin, 3^ mm wide, undulating

wing, which overarches the apex of the stem, giving it a

somewhat circinate appearance. The spore-containing

capsules are enveloped by large, bottle-shaped involu-

cres, borne along the stem, often on the same side as the

wing, the youngest being near the apex. The antheridia

are mostly borne on separate plants in a row of pockets

along the thickened free margin of the wing. The spores

are released upon decay of the involucre and capsule

wall; they are large and the ornamentation, as well as

sometimes the colour, are regarded as species-specific

and essential for correct identification. R. purpiireospora

has purple spores, hence its specific epithet; it is thus

easily recognized, since the other four southern African

Riella species have brown spores.

ZANNICHELLIACEAE

Pseudalthenia aschersoniana

Pseudalthenia aschersoniana (Graebn.) Hartog

(Zannichellioideae) is a member of the eurysaline group

of the Zannichelliaceae. This ‘vlei grass’ is an endemic

component of the ephemeral estuarine environments of

the Cape Peninsula (Den Hartog 1980, 1981). Earliest

herbarium records report P. aschersoniana from the

southwestern Cape Peninsula (Kommetjie) in 1897, from

Rietvlei (Milnerton) in 1930, and again from both

Rietvlei and Kommetjie during the early to mid-1960s.

Routine monitoring of the ephemeral pans, usually

wet between April and September, during the late winter

and spring of 1997 and 1998, revealed Bolboschoenus

maritimus and P. aschersoniana as the dominant aquatic

plant species, with B. maritimus dominant in the deeper

(0.3-0. 5 m) water, and P. aschersoniana in the shallow

(< 0.2 m) areas.

The dominant plants in both areas during the dry sea-

son is a species of Sarcocornia. Collections of Pseudal-

thenia aschersoniana were made on 3 1 August 1 998, and

submitted to the National Botanical Institute for identifi-

cation.

Subsequent to the pans drying out during September

1998, healthy specimens of P. aschersoniana were dis-

covered during early 1 999, growing in ± 0.3 m of water

on the top of gabion baskets forming the sides of an arti-

ficial canal system on the property, and constituting a

boundary of the ephemeral pan area.

These plants are regarded as pond weeds (Reinecke

1964) and are annuals with slender, sympodially

branched rhizomes. The nodes give rise to upright stems,

up to 400 mm long, the length apparently governed by the

depth attained by the seasonal body of water in which

they are growing. The leaves are alternate and linear,

20-75 X 1-2 mm, gradually narrowing to the rounded

apex, and expanded below to form a 5 mm long sheath

which is auriculate above. The floral parts are extremely

reduced, male and female flowers are borne together in

the axils. The male flower is a single stamen, at first

enveloped in a thin membranous spathe, the filament

lengthens rapidly at anthesis, up to 25 mm long and the 4

locules form a rectangular, 8-sporangiate anther that

dehisces longitudinally. The female flowers are 1 -carpel-

late and surrounded by a vaginate spathe and an inner,

flask-shaped ‘perianth’, the style lengthens rapidly during

anthesis, the ephemeral stigma is large and obliquely fun-

nel-shaped. The fruits are asymmetrically graniform to

somewhat bean-shaped drupelets, ± 4 mm long and

stalked at maturity. The stalks of fruits borne higher up on

the erect shoots are only 1 or 2 mm long, whereas they are

20-60 mm long on those formed lower down, as well as

on the rhizomes and are positively geotropic (Obermeyer

1966), burying the fruits in the mud, thus affording them

greater protection during the dry season. Germination

starts soon after the first rains begin to fill the ponds.

DISCUSSION

The rare and sporadic occurrence of species of Riella

and Pseudalthenia may be attributed to loss or alteration

of habitat, particularly those changes brought about by

urbanization. During August/September 1999 R. pur-

pureospora was absent in other Western Cape vlei local-

ities where it was previously collected, or likely to have

been present, namely Zandvlei (Westlake), Wildevoel-

vlei, Papkuilsvlei, Bloubergsvlei and Valkenberg Vlei. It

should be noted that the level of disturbance of these

vleis is considerably higher than in Blouvlei, and that in

many cases the ‘original’ vlei environment is no longer

discernible. The reappearance of R. purpiireospora and

P. aschersoniana at Blouvlei provides a valuable indica-

tion of the resilience of ephemeral pan floral communi-

ties to rehabilitation, and of the unique value of this con-

servation area within a high-density commercial develop-

ment.

ACKNOWLEDGEMENTS

The authors wish to thank Dr O.A. Leistner for kind-

ly reading the manuscript and for his helpful suggestions.

REFERENCES

COETZER, A.H. 1987. Succession in zooplankton and hydrophytes of

a seasonal water on the west coast of South Africa, Hydro-

hiologia 148: 193-210.

DEN HARTOG, C 1980. Pseudalthenia antedates VIeisia. A nomen-

clature note. Aquatic Botany 9: 95.

/

Bothalia 30,2 (2000)

159

DEN HARTOG, C. 1981. Aquatic plant communities of poikilosaline

waters. Hvdrobiologia 81: 15-22.

LOCHNER, R & ROSSOUW, N. 1997. The development of an envi-

ronmental management plan for incorporating a wetland into a

large mixed use development: the Century’ City example. Paper

submitted to the 1997 conference of the International

Association for Impact Assessment (lAIA).

MAZE, K.E. & REBELO, A.G. 1999. Core flora conservation areas on

the Cape Flats. FCC Report 99/1. Flora Conservation Commit-

tee, Botanical Society of South Africa.

OBERMEYER, A. A. 1 966. Zannichelliaceae. Flora of southern Afiica

1:77-81.

PEROLD, S.M. 2000. Studies in the Sphaerocarpales (Flepaticae) from

southern Africa. 3. The genus Riella and its local species.

Bothalia 30: 125-142.

PROSKAUER, J. 1955. The Sphaerocarpales of South Africa. Journal

of South Afi'ican Botany 21 : 63-75.

REINECKE, P. 1 964. A contribution to the morphology of Zannichellia

aschersoniana Graebn. Journal of South African Botany 30:

93-101.

WIGGLESWORTH, G. 1937. South African species of Riella, includ-

ing an account of the developmental stages of three of the

species. Journal of the Linnean Society of London. Botany 5:

309-332.

W.R. HARDING*, S.M. PEROLD** and R.P. GLEN**

* DH Environmental Consulting cc, P.O. Box 5429, 7135 Helderberg,

South Africa

** National Botanical Institute, Private Bag XI 01, 0001 Pretoria.

MS. received: 2000-06-14.

Bothalia 30,2: 161-173 (2000)

Combining floristic and growth form composition in a gradient-

directed vegetation survey of Matjiesrivier Nature Reserve, Western

Cape, South Africa

R.G. LECHMERE-OERTEL* and R.M. COWLING**

Keywords: biomes, Cederberg, floristic, fynbos, gradsects. growth forms, Matjiesrivier Nature Reserve, Succulent Karoo, survey, Western Cape

ABSTRACT

The floristically complex vegetation of Matjiesrivier Nature Reserve (MNR), which spans the ecotone between the

Fynbos and Succulent Karoo Biomes in the eastern Cederberg Mountains, Western Cape, was surveyed using a gradient-

directed transect (gradsect). The gradsect was aligned with a topo-climatic aridity gradient across MNR. The vegetation was

classified using TWINSPAN, based on a combination of floristic and growth form characteristics, and an understanding of

the main ecological gradients controlling vegetation distribution. The final classification described seven robust and eco-

logically meaningful communities that represented a trade-off between statistical rigour and practicality for management.

The seven communities were mapped using a geographical information system (GIS).

CONTENTS

Abstract 161

Introduction 161

Study area 162

Methods 162

Results 163

Floristic classification 163

Growth form classification 163

Community descriptions 163

1. Fynbos Communities 163

1.1. Asteraceous Fynbos Matrix (AFM) 163

1.2. Restioid Sandy Fynbos (RSF) 167

2. Transitional Communities 168

2.1. Dwarf Bedrock Shrubland (DBS) 168

3. Non-Fynbos Communities 168

3.1. Kloof Thicket (KT) 168

4. Succulent Karoo Communities 169

4.1. Shale Succulent Karoo (SSK) 169

4.2. Succulent Karoo Matrix (SKM) 170

4.3. Sandy Succulent Karoo (SaSK) 170

Discussion 171

Mountain Fynbos 171

Succulent Karoo 171

Conclusion 172

Community-environment generalizations 172

References 172

Appendix 173

INTRODUCTION

The vegetation in the Fynbos-Succulent Karoo eco-

tone, Western Cape is structurally and floristically com-

plex, as it comprises elements from three distinct and

typically speciose vegetation types (Table 1): Mountain

Fynbos (Rebelo 1996a), Central Mountain Renosterveld

* Department of Zoology. University of Port Elizabeth, P.O, Box 1 600,

6000 Port Elizabeth.

** Institute for Plant Conservation, Department of Botany, University

of Cape Town, Private Bag, 7701 Rondebosch, Cape Town.

MS. received: 1998-06-02.

(Rebelo 1996b) and Lowland Succulent Karoo (Hoffman

1996). Although the Fynbos Biome has received consid-

erable attention in terms of vegetation classification, lit-

tle work has been done in the arid fynbos and Renos-

terveld of the northwest region of the Fynbos Biome

(Cowling et al. 1997) or in the Fynbos-Succulent Karoo

ecotone.

This ecotonal vegetation is a mosaic of arid Fynbos,

Renosterveld and Succulent Karoo separated out along

gradients of moisture, soil conditions and disturbance

regime. Fynbos is typically found on fire-prone mesic

sites with sandstone-derived soils, Renosterveld on

mesic sites with shale-derived soils and Succulent Karoo

on fire-free xeric sites, irrespective of soil type. Fire is

the driving disturbance in fynbos, imposing an element

of stochasticity on the establishment of species assem-

blages (Cowling et al. 1997). Herbivory and drought are

the major disturbances that influence species assem-

blages in Succulent Karoo (Milton et al. 1997).

TABLE 1. — The major structural differences between the three vege-

tation types converging at Matjiesrivier Nature Reserve (adapt-

ed from Low & Rebelo 1996)

162

Bothalia 30.2 (2000)

The juxtaposition of three major vegetation types in a

single reserve raises problems for the management and

conservation of the flora at MNR. Each vegetation type,

particularly succulent karoo and fynbos, needs to be man-

aged separately, as they respond differently to certain

management actions such as fire and have different graz-

ing capacities (Van Wilgen et al. 1992; Milton & Dean

1996). There was thus a need for the vegetation in MNR

to be classified into management units, which would

allow the application of different management strategies

to different vegetation communities. To be meaningful for

managers, who may not necessarily have botanical train-

ing, the classification needed to be relatively simple and

based on easily identifiable structural characters, higher

taxa and dominant genera or species.

The aim of this survey was to classify the vegetation

of MNR into ecologically meaningful units (communi-

ties) that were easily identifiable using simple growth

form and floristic characters. These vegetation commu-

nities of MNR were mapped within a GIS framework.

STUDY AREA

Matjiesrivier Nature Reserve (32° 25' S, 19° 17' E) lies

to the east of the Cederberg Mountains, Western Cape.

MNR experiences a Mediterranean-type climate with a

strong topo-climatic gradient (± 100-300 mm annual

rainfall) superimposed over an abrupt transition from

sandy, infertile soils (over Table Mountain Group

Sandstone) to clayey, fertile soils (over Bokkeveld Group

Shale) (Figure 1 ). MNR, prior to being purchased for con-

servation in 1991, was a commercial stock farm for

approximately 150 years, with a little arable land in the

Matjiesrivier Valley. There are no farm records detailing

stocking strategies or fire history. The vegetation on

MNR reflects the main environmental gradients of mois-

ture and soil with a gradual transition from Mountain

Fynbos to Succulent Karoo from west to east, and a sharp

boundary over the transition from Table Mountain

Sandstone to Bokkeveld Shale. Patches of Renosterveld

are found on mesic shale-derived soils.

METHODS

The abundance and percentage cover of all perennial

plant species with a cover of > 5% was estimated in 125,

10 X 10 m sites during the winter of 1996. The cover and

abundance data for each species were combined into nine

categories using the Domin scale (Causton 1988). Species

occurring in fewer than three sites were removed from the

data analyses. Every species was classified into a unique

growth form: forb, sedge, grass, restioid, geophyte, leaf-

succulent shrub, stem-succulent shrub, evergreen shrub

and deciduous shrub. The restioids and shrubs were fur-

ther divided into height classes: dwarf (< 0.25 m), low

(0.25-1.0 m), medium (1-2 m) and tall (> 2 m). Each

growth form represented a functional type with known

differences in eco-physiological behaviour (Cowling et

al. 1994). The cover data for all species that fell into the

same growth form were totaled within each site. Several

climatic, topographic and edaphic variables were mea-

sured or estimated for each site. The mean and standard

deviation of several environmental variables were calcu-

lated for all the sites comprising a community.

The sites were positioned through MNR using an

approach similar to that employed by gradsect sampling

(Gillison & Brewer 1985; Austin & Heyligers 1991).

Gradsect sampling is the deliberate positioning of tran-

sects along the potentially most important environmental

gradient in the area. Assuming that plant species or com-

munities are separated along an environmental gradient,

positioning sites within a gradsect ensures that the great-

est range of vegetation is included in the sampling

(Austin & Heyligers 1991).

A conspicuous environmental gradient at MNR was

an eastward increase in aridity that corresponded to an

altitudinal gradient from ± 1 200 m in the west to ± 500

m in the east. Interpolated rainfall data from the

Computing Centre for Water Research (CCWR 1996)

minute-by-minute rainfall database, supported this ob-

servation. The gradsect used to survey the vegetation at

MNR was aligned from west to east and corresponded to

the main road access. The sampling effort was mostly

confined to areas within about one kilometre from the

road. The gradsect was further stratified using geology

(Sandstone or Shale) and land type (bedrock sheet, sandy

plain, gravel pan or other).

The data were analysed using the polythetic classifica-

tion computer program TWINSPAN (Hill 1979). The

default settings were used except for the floristic classifi-

cation where the five pseudospecies cutoff levels were set

FIGURE 1 , — The west-east aridity

gradient superimposed over

the geological transition from

sandstone to shale.

Bothalia 30,2 (2000)

163

at 0, 3, 5, 7, 9 to account for the Domin scale. We initially

classified the vegetation using both the floristic and

growth form data sets separately and compared the two

classifications. Ultimately, the vegetation was classified

into communities or management units based on an intu-

itive combination of the growth form and floristic classifi-

cations and field notes made during the survey. In the final

classification, we used a combination of obvious growth

form characteristics combined with the dominant families,

genera or species. Although not strictly objective, this

method provided a workable and robust set of communi-

ties, which were a satisfactory compromise between eco-

logical meaning and manageability. A map of the vegeta-

tion at MNR, based on the communities derived from the

classification was prepared using a combination of GIS

coverages, aerial photographs and field notes.

RESULTS

Floristic classification

Seven major communities were derived from the

floristic classification (Figure 2). The differential species

from TWINSPAN were not always used as the diagnos-

tic species in the final classification. The first division of

sites was into fynbos and thicket and succulent karoo. At

the second division the succulent karoo sites were divid-

ed into two communities based on underlying geology:

one located exclusively on shale-derived soils (Shale

Succulent Karoo) and the other on sandstone-derived

soils. The fynbos sites did not split into meaningful com-

munities at the second division.

At the third level, the succulent karoo sites on shale

were divided into two forms (1 and 2), with no obvious-

ly apparent ecological significance. The succulent karoo

sites on sandstone were divided into two communities

based on land type. Sandy Succulent Karoo was found

on deep sand plains whereas Succulent Karoo Matrix

was found on the rocky slopes. The fynbos sites were

divided into four communities based on land type. Dwarf

Bedrock Shrubland was found on bedrock sheets,

Restioid Sandy Fynbos on deep sand plains. Kloof

Thicket around large outcrops of rock and Asteraceous

Fynbos Matrix on the rocky slopes and cliff tops. Dwarf

Bedrock Shrubland, which was not derived in the growth

form classification, was included as an important com-

ponent of the vegetation based on field observations. The

communities and their ecological significance are dis-

cussed in the community descriptions. A list of all the

species names used in the classification, with their

authors is given in the appendix.

Growth form classification

The growth form classification (Figure 3) was funda-

mentally similar to the floristic classification, and all the

communities except Dwarf Bedrock Shrubland were

resolved. The differential growth forms at each division

of the classification gave some indication of the domi-

nant growth forms characteristic of each community. The

initial division of sites was into fynbos and non-fynbos

or transitional. At the second division, a small group of

sites, which were congruent with the Restioid Sandy

Fynbos in the floristic classification, split off from the

fynbos sites. The remaining fynbos sites made up the

Asteraceous Fynbos Matrix from the floristic classifica-

tion. The non-fynbos sites were split into Succulent

Karoo and a Fynbos-Succulent Karoo transitional group.

At the third division, the Asteraceous Fynbos Matrix

sites were divided into two forms not similar to the two

forms in the floristic classification. The group of

Succulent Karoo sites remained largely intact except for

four sites that split off for no apparent ecological reason.

The transitional sites were split into two groups. The

sites in the first group were congruent with those of the

Kloof Thicket Community in the floristic classification.

The sites in the second group were a combination of

Shale Succulent Karoo and Asteraceous Fynbos Matrix

from the floristic classification and probably represented

a transitional zone between these two communities.

At the fourth level of division, further resolution was

only obtained by splitting the Succulent Karoo sites into

two groups. The one group had sites congruent with

Sandy Succulent Karoo whereas the other had sites con-

gruent with Succulent Karoo Matrix in the floristic clas-

sification.

COMMUNITY DESCRIPTIONS

The final classification of the vegetation was made

from an intuitive combination of the growth form and

floristic classifications. Seven major communities were

recognised and are described below according to their

growth form and floristic composition. These communi-

ties were thought to be ecologically significant and were

used as units in the vegetation map (Figure 4).

1. Fynbos Communities

Fynbos at MNR is characterised by a few distinct

growth forms, families and species. The dominant

growth forms are low to medium woody shrubs with lep-

tophyllous and sclerophyllous ericoid leaves, dwarf to

tail restioids and dwarf to low sedges. One species that

repeatedly came up as an indicator species is Diosma

acmaeophylla (Rutaceae), which is a medium to tall

evergreen, ericoid shrub. Fynbos communities are

restricted to the western half of MNR where rainfall and

altitude are relatively high and the soils derived from

sandstone. Fynbos communities made up a mosaic in this

half of MNR, depending on the land type.

1.1. Asteraceous Fynbos Matrix (AFM)

Related communities: Campbell’s (1985) Dry Astera-

ceous Fynbos.

AFM is a heterogeneous community and is more a

‘dumping’ category than a distinct community. Plant

cover in the sites is ± 40%. A mesic and a xeric form are

defined in the floristic classification (Figure 2). The

mesic form is the ‘true’ fynbos and the xeric form is tran-

sitional from fynbos into succulent karoo.

164

Bothalia 30,2 (2000)

FIGURE 2. — The hierarchical classification of the vegetation at Matjiesrivier Nature Reserve based on floristic characters.

Bothalia 30.2 (2000)

165

FIGURE 3. — The hierarchical classification of the vegetation at Matjiesrivier Nature Reserve based on growth form characters.

166

Bothalia 30,2 (2000)

FIGURE 4. — A map of the vegetation of Matjiesrivier Nature Reserve, showing the distribution of the communities.

Bothalia 30,2 (2000)

167

Floristic composition

All the typical elements of arid mountain fynbos

(members of the Asteraceae, Proteaceae, Restionaceae

and Poaceae) are present in AFM. Shrubs from the

Asteraceae are the most abundant of all the families in

the fynbos. Diosma acmaeophylla is the only indicator

species for AFM. However, a number of genera and

species are relatively common throughout AFM and are

locally abundant in the fine-scale species associations.

The mesic and xeric forms of AFM are separated on the

basis of dominant genera. The mesic form has a greater

proportion of more ‘typical’ fynbos taxa including:

Diosma acmaeophylla, Phylica odorata, Leucadendron

pubescens, Aspalathus spp., Agathosma spp., Stoebe

fusca, and Felicia scabrida. Geophytes and members of

the Restionaceae (particularly of the genus Cannomois)

and Cyperaceae (particularly Ficinia dunensis) dominat-

ed the understorey of the mesic form.

The xeric form is characterised by a greater propor-

tion of woody taxa from the Asteraceae including: Erio-

cephalus africanus, E. ericoides, Pteronia incana, Euryops

wageneri, E. othonnoides, Dolichothrix ericoides and

Elytropappus rhinocerotis, and several species of Othonna.

Members of the Mesembryanthemaceae and Crassula-

ceae dominate the field layer. The xeric form of AFM

shows affinities to Central Mountain Renosterveld

(Rebelo 1996b). However, the relatively high cover of

restioids means that this type is classified as a form of

fynbos (Campbell 1988).

Structure

Many of the sites classified as AFM in the floristic

classification are classified as transitional with Shale

Succulent Karoo in the growth form classification

(Figure 2). Low to medium evergreen shrubs, restioids

and low leaf-succulent shrubs are the most abundant

growth forms in AFM. The vegetation canopy is 0.5-1. 5

m tall with a few taller (> 2 m) evergreen shrubs in par-

ticularly mesic sites. In terms of leaf size and form, the

leaves of the characteristic shrubs are typically sclero-

phyllous, leptophyllous and either ericoid or elytropap-

poid. Low (0.25-1.0 m) sedges and low to medium

(0.25-2.0 m) restioids occur in dense clumps which are

often locally dominant throughout AFM on small sandy

patches. Low (< 0.1 m) leaf-succulent shrubs make up a

significant proportion of the field layer of AFM, espe-

cially where the soil is shallow.

Distribution and habitat

Asteraceous Fynbos Matrix is widely distributed

through the western half of MNR (Figure 4). The eastern

extreme of xeric AFM (and fynbos in general) is approx-

imately in the middle of MNR (longitude: 19° 25' E; alti-

tude: 800 m; estimated annual rainfall: 250 mm). The

boundary between xeric AFM and Succulent Karoo

Matrix is characterized by a gradual loss of fynbos ele-

ments and a corresponding increase in succulent karoo

elements (mostly non-succulent asteraceous shrubs). An

increase in aridity, associated with a decrease in altitude,

accompanied this transition.

AFM occurs in all topographic positions from flat

plateaus and plains to steep slopes, wherever the soil is

derived from sandstone or quartzite and where there is a

rocky substrate (either as talus or as partially exposed

bedrock). Soil conditions are invariably sandy, with a

very low clay content and electrical conductivity

(Lechmere-Oertel 1998). Isolated patches of AFM are

scattered away from the centres of AFM throughout the

western half of MNR, in the Fynbos-Succulent Karoo

Transition zone (Figure 4). These AFM patches occur at

locally mesic sites within the transitional zone. Such

sites include many of the high elevation ridges that are

more mesic, south-facing slopes and sites where there is

precipitation runoff into the soil due to the presence of

large rock sheets.

1.2. Restioid Sandy Fynbos (RSF)

Related communities: Campbell’s (1985) Dry Restioid

Fynbos; Taylor’s (1996) Community 18 Wildenowia

arescens-Thamnochortus platypteris.

RSF is generally low to medium in height (< 2 m)

with a canopy layer dominated by either restioids or pro-

teoids. The understorey is either bare or dominated by

dwarf (< 0.25 m) graminoids (restioids, grasses and

sedges) and leaf-succulent woody shrubs from the

Mesembryanthemaceae and Crassulaceae. The percent-

age cover and average height of plants in RSF appear to

depend on moisture availability. In mesic areas there is

greater cover (up to 70%) of tall restioids and proteoids,

whereas the more xeric areas have lower cover (< 40%)

of shorter restioids.

Eloristic composition

Members of the family Restionaceae (restioids) have

the greatest cover in RSF. The most conspicuous and

locally dominant genera in the Restionaceae are typical-

ly Willdenowia, Restio, Thamnochortus, Hypodiscus,

Ischyrolepis and Elegia. Large isolated individuals (up to

2 X 2 m) from the genus Willdenowia (mostly Willde-

nowia incurvata) are scattered through RSF. Some of the

patches of RSF are also characterised by the abundance

(> 10%) of Leucadendron brunioides var. brunioides

(Proteaceae) or other Leucadendron species.

Depending on the apparent moisture availability,

there is considerable variation in the understorey of RSF.

Grass taxa, particularly the genera Pentaschistis and

Ehrharta, are more apparent in the understorey of the

xeric areas. There is also an increase in the proportion of

mesembs and crassulas in the understorey with increas-

ing aridity. In mesic areas, sedges, notably of the genera

Eicinia (mostly Eicinia dunensis) and Tetraria replace

the grasses and shrubs in the understorey. Geophytes and

annual forbs are also quite common.

There is a distinct form of RSF that has a considerably

shorter canopy than the other patches of RSF, and is

dominated by Ischyrolepis unispicatus (> 40% cover)

and Elytropappus rhinocerotis (Asteraceae). This form

of RSF only occurs where there is a high cover of small

(< 0.05 m) pebbles scattered on or near the surface of the

168

Bothalia 30,2 (2000)

sand. These pebbles were possibly derived from the ero-

sion of an ancient Tertiary surface (Taljaard 1949; Peter

Holmes pers. comm.).

Structure

In terms of structure, RSF is quite uniform, with the

biggest variation being attributable to differences in

height of the canopy. The dominant growth forms in RSF

are medium to tall restioids and medium to short sedges

and grasses. In a few patches there are very tall proteoid

shrubs; otherwise there are few woody shrubs typical of

AFM.

Distribution and habitat

Restioid Sandy Fynbos is found exclusively on patch-

es of deep sand throughout the western half of MNR. The

orange or white sand is probably of aeolian origin. There

are several large areas of deep sands in the western half

of MNR and all of these are covered by RSF (Figure 4).

The patches of RSF form islands within the fynbos

matrix and have many elements in common with AFM.

The sand patches are often interspersed with large sand-

stone outcrops and bedrock sheets, which harbour Kloof

Thicket and Dwarf Bedrock Shrubland Communities.

2. Transitional Communities

2.1. Dwarf Bedrock Shrubland (DBS)

Related communities'. Taylor’s (1996) Communities 5, 6

& 7.

Dwarf Bedrock Shrubland is represented by only two

sites in the floristic classification and is not defined as a

community in the growth form classification. DBS is

included as a distinct community because, based on field

observations, it appears to be an important component of

the vegetation that may harbour rare or endemic species.

Typically, there is a very low cover of vegetation on the

sheetrock, as there are only a limited number of cracks

and erosion hollows where enough soil can collect to

support woody shrubs. The limited soil volume available

to most of the shrubs has resulted in stunted growth.

Individuals of species which are typically medium to tall

in more suitable habitats, remain at < 0.5 m, with only

their thick and twisted stems testifying to their age.

Floristic composition

In mesic areas, DBS is characterised by stunted eri-

coid, sclerophyllous members of the Ericaceae,

Proteaceae and Rhamnaceae. Three species: Erica maxi-

miliani, Protea glabra and Phylica buxifolia, are partic-

ularly conspicuous on bedrock sheets throughout the

western half of MNR. Large individuals of P. glabra (up

to 5 m high), with roots penetrating deep into cracks in

the bedrock, are quite common up to the extreme eastern

boundary of the fynbos/succulent karoo transition. The

dominant species in DBS (based on abundance, as there

was never a high cover of vegetation) depend on the sur-

rounding vegetation.

Towards the arid east of MNR, taxa common in suc-

culent karoo increase relative to the fynbos taxa. Even

within the fynbos matrix there are small populations of

members of the Mesembryanthemaceae (Ruschia,

Lampranthus, Conophytum) and Crassulaceae {Crassu-

la, Tylecodon) that are able to survive the harsh, xeric

conditions of the shallow soil.

Structure

DBS is structurally diverse and combines typical fyn-

bos and succulent karoo growth forms. Ericoid, sclero-

phyllous and dwarf leaf-succulent shrubs are found

together, which is why DBS is considered as transitional

and does not arise as a distinct community in the growth

form classification.

Distribution and habitat

As the name suggests, DBS is confined to the surfaces

of the large sheetrock outcrops that occur throughout

MNR (Figure 4). The sheetrock is exposed sandstone or

quartzite bedding planes that are typically flat. There are

few occurrences of exposed sheets of horizontal shale

strata at MNR for comparison between geological

groups. The surfaces of the bedrock sheets are typically

eroded into channels, crevices and depressions where

enough soil and organic matter collects to provide micro-

sites for the growth of woody plants. The depth of the

crack (and thus soil) appears to dictate the type and struc-

ture of plants that can grow there.

The rocky habitat presents two extremes of water

availability to plants that survive on them. During the dry

summer season, the exposed sheetrock is exceptionally

xeric as there is little soil to act as a reservoir for mois-

ture. During the wet season, the depressions in a

sheetrock have an over-abundance of water due to the

very high runoff and may become waterlogged. Plants

are unable, in most cases, to penetrate subsurface water

unless the rock crack in which they are growing is deep

enough (as is the case for P. glabra). Thus the plants liv-

ing in rock cracks must be able to survive extended peri-

ods of extreme drought during summer, very wet condi-

tions during winter, and rapid changes between a state of

moist soil and dry soil.

3. Non-Fynbos Communities

3.1. Kloof Thicket (KT)

Related communities: Campbell’s (1985) Forest and

Thicket; Taylor’s (1996) Community 4, Olea europaea

subsp. africana-Myrsine africana on sand flats and

mesic screes.

Kloof Thicket is confined to locally mesic habitats

between large rock outcrops. The vegetation canopy is

tall to very tall (> 2 m) and cover is high compared to the

surrounding fynbos matrix. The floristic composition of

KT is derived from a combination of typical subtropical

thicket shrubs and a number of typical fynbos shrubs.

The restioid and proteoid growth forms are mostly absent

from KT.

Bothalia 30,2 (2000)

169

Floristic composition

Kloof Thicket appears in the floristic classification as

a subset of fynbos, with Rhus undulata as a differential

species. Other thicket species, which we observed as

common in KT (but are not apparent in the floristic clas-

sification), included Maytenus oleioides and Dodonaea

angustifolia. Depending on the location of the rock out-

crop, there are several other woody shrub species more

typical of fynbos or the transition from fynbos into suc-

culent karoo such as Diosma acmaeophylla, Phylica bux-

ifolia, Pteronia spp., Eriocephaliis ericoides and

Helichrysum spp.

Structure

Kloof Thicket is structurally easy to identify. It is

characterised by tall (> 2 m) woody shrubs with large

(mesophyllous) evergreen leaves, and by a lack of the

restioid and proteoid growth forms. Forbs, ferns and

deciduous dwarf shrubs are characteristic of the under-

storey.

Distribution and habitat

Kloof Thicket is typically found between the large

sandstone outcrops that are dotted through the western

half of MNR. These thicket clumps are restricted to the

higher rainfall areas of MNR and become less frequent

with increasing aridity. KT occurs on patches of shallow

aeolian sand trapped between the large outcrops. There is

probably considerable runoff from the outcrops during

precipitation and thus the soil around the edge of the out-

crops receives more effective precipitation than the sur-

rounding vegetation.

The protected nature of the KT sites, which are

hemmed in by large outcrops of sandstone, suggests that

they are fire refuges. Fire is believed to be a factor that

maintains the boundary between fynbos and thicket

(Cowling & Holmes 1992) and it is possible that infre-

quent fires restrict KT into fire refugia.

4. Succulent Karoo Communities

Succulent Karoo sites are separated from fynbos sites

on the basis of a number of floristic and growth form

characteristics. The main Succulent Karoo families at

MNR include the Mesembryanthemaceae, Asteraceae

and Crassulaceae. The abundance of leaf- and stem-suc-

culent and deciduous shrubs, and the absence of ever-

green leptophyllous shrubs with ericoid leaves, are prob-

ably the most important growth form features.

4.1. Shale Succulent Karoo (SSK)

Related communities'. Campbell’s ( 1985) Succulent Shrub-

land.

SSK is exclusively confined to a band of exposed

shale strata from the Bokkeveld Group. Deciduous

shrubs from the Asteraceae, stem-succulent species from

Euphorbia, and leaf- and stem-succulent members of the

Mesembryanthemaceae and Crassulaceae dominate SSK.

The vegetation in SSK is relatively sparse, with less than

30% cover and with a canopy height of about one metre.

The understorey is also typically sparse, comprising

dwarf leaf- and stem-succulent shrubs and many annual

ferns, forbs and geophytes.

Eloristic classification

Two differential species, Tylocodon wallichii (Cras-

sulaceae) and Pteronia incana (Asteraceae) are charac-

teristic of SSK in the floristic classification. Other gen-

era from the Asteraceae that are particularly abundant in

SSK included Euryops, Pteronia and Eriocephalus.

Pteronia divaricata, Galenia africana, Eriocephalus

africanus and E. ericoides are most abundant. Large

individuals of Euphorbia mauritanica and Tylecodon

paniculatus are dotted throughout the community.

Although these individuals do not account for a large

proportion of the cover, they are a conspicuous and diag-

nostic feature of SSK. Species diversity in SSK is rela-

tively low compared to the other succulent karoo com-

munities.

Two formations, SSK-1 and SSK-2, are derived in the

floristic classification based on the presence of Euryops

othonnoides and Pteronia divaricata in SSK-1 and

Galenia africana and Eriocephalus africanus in SSK-2.

The separation of SSK sites into these formations did not

make ecological sense, therefore, SSK was mapped as a

single management unit.

‘Heuweltjies’ or mima-like earth mounds (Knight et

al. 1989; Moore & Picker 1991) are quite common

throughout SSK. These mounds are about 0. 1-0.5 m

high and ± 2-10 m diam. ‘Heuweltjies’ differ from the

surroundings in terms of soil nutrient, organic matter and

moisture status (Milton et al. 1997) and those at MNR

support ‘islands’ of vegetation that are noticeably differ-

ent from the surrounding matrix vegetation. We only

sampled three sites on these ‘heuweltjies’. In both the

floristic and growth form classifications these sites are

assigned to SSK-2, with two characteristic species:

Pteronia divaricata and Tylecodon paniculatus.

Structure

There is no distinct group of sites in the growth form

classification that corresponds to the SSK sites in the

floristic classification. There is considerable overlap

between AFM and SSK and thus, in the growth form

classification, SSK appears to be part of a transitional

community from fynbos into succulent karoo. This is not

surprising considering that most of the growth forms

found along the shale band are also found in the fyn-

bos/succulent karoo transition. It is only the charismatic

and conspicuous growth forms such as the large stem-

and leaf-succulents [Tylecodon paniculata and Euphor-

bia mauritanica) which cause the vegetation on the shale

band to look different to the surrounding matrix.

Distribution and habitat

SSK is found exclusively on the bands of shale and

sandstone from the Bokkeveld Group that are exposed

170

Bothalia 30,2 (2000)

along their bedding planes near the homestead at MNR

(Figure 4). These alternating layers of shale and sand-

stone give rise to a series of parallel north-south orien-

tated ridges of sandstone and valleys of shale. The shale

bedrock is exposed in many places and there is often an

overburden of sandstone debris on the slopes from the

sandstone ridges. The soil on these slopes is dark brown

and has a relatively high proportion of silt and clay, giv-

ing rise to a fine-textured loam compared to the sur-

rounding sandstone-derived sands. It is these fine-tex-

tured soils which are thought to increase the effect of

summer drought on the vegetation, resulting in the

occurrence of more succulent karoo taxa compared to the

surrounding matrix vegetation (Lechmere-Oertel 1998).

The succulent karoo vegetation growing on gravel

patches is also classified as SSK. There is another

localised occurrence of shale in MNR, on a south-facing

slope in the middle of the Matjiesrivier Gorge. There is

extensive folding of the rock strata at this point, and the

river has exposed an anticline of Bokkeveld Group Shale

beneath the Witteberg Group Sandstone. The vegetation

on this shale is dominated by dwarf shrubs (mostly

Pteronia spp.), leaf-succulent mesembs and the stem-

succulent Euphorbia liamata. There is a very low total

cover; probably because the surface is unstable with

almost no soil (steep slope and gravelly shale soil which

must have had a very high rate of erosion).

The proximity of the Bokkeveld Group ridges and

valleys to the homestead and perennial water (the

Matjiesrivier) suggests that the SSK community would

have been under considerable grazing pressure during

the ± 200 years MNR was a stock farm. Thus the current

vegetation and community description is unlikely to

reflect what the vegetation would be like in ungrazed

conditions.

4.2. Succulent Karoo Matrix (SKM)

Related communities: a xeric version of Campbell’s (1985)

Succulent Shrubland that occurs on quartzitic soils.

The most distinguishing feature of SKM is the domi-

nance of leaf-succulent members of the Mesembry-

anthemaceae and Crassulaceae, both in terms of relative

cover in the vegetation and species diversity. The vege-

tation in SKM is generally less than one metre high and

total vegetation cover relatively low (< 30%). As in

AFM, there are two forms within SKM, which reflect the

transition from fynbos into succulent karoo. One form

occurs in the arid part of the transitional zone and has a

combination of typical succulent karoo and fynbos taxa

and growth forms. The other form, which occurs in the

extreme arid part of MNR is ‘proper’ succulent karoo

with very few fynbos elements. These forms comprise an

intricate matrix, depending on local environmental con-

ditions that inlluence water availability, and are not map-

pable as separate units.

Florist ic composition

Numerous members of the Mesembryanthemaceae and

Crassulaceae dominate SKM. There is high species rich-

ness in these two families in SKM, which is expected,

considering that the succulent karoo is the centre of diver-

sity for these families (Milton et al. 1997). In the floristic

classification, two Ruschia species, the grass genus Pen-

taschistis and the shrub Gnidia deserticola (Thymelae-

aceae) are differential taxa for SKM (Figure 2).

In the transitional form of SKM, shrub genera from

the Asteraceae, particularly Pteronia, Eriocephalus and

Elytropappus, are most common. There are also a num-

ber of fynbos taxa present wherever there is enough

moisture, such as on south-facing slopes. A number of

other conspicuous species such as Euphorbia hamata, E.

mauritanica, Tylecodon paniculatus and Hoplophyllum

spinosa are distributed sparsely through the arid part of

the transitional formation.

The ‘true’ SKM form is almost totally dominated by

members of the Mesembryanthemaceae and Crassula-

ceae and there are few shrubs from other families

(Gnidia deserticola is a notable exception as a locally

dominant species). Two very common genera from the

Mesembryanthemaceae found in SKM are Ruschia and

Lampranthus .

Structure

Dwarf to low leaf-succulent woody shrubs dominate

the vegetation in SKM, increasing relative to the non-

succulent shrubs along the aridity gradient. In the ‘true’

SKM form there are few typical fynbos growth forms

such as evergreen ericoid shrubs. Medium-height leaf-

succulent shrubs, geophytes, grasses and deciduous

dwarf shrubs are differential growth forms for this for-

mation. In the ‘transitional’ form, low to medium ever-

green shrubs, growth forms more typical of the fynbos

communities, are differential. The transitional forms

reflect a change from evergreen to deciduous and an

increase in leaf succulence.

Distribution and habitat

SKM occurs in the eastern half of MNR (Figure 4).

There is a gradual transition from Fynbos into Succulent

Karoo marked by an increase in the appearance of typi-

cal karroid elements with increasing aridity and a corre-

sponding loss of fynbos elements. SKM occurs in the

fynbos part of the transitional zone wherever chere is

localised aridity. Examples of this include the steep

slopes of the Matjiesrivier Gorge and its tributaries in the

western half of MNR. The ‘true’ SKM form is found in

the eastern extreme of MNR, near the Doring River.

4.3. Sandy Succulent Karoo (SaSK)

Related communities: none.

SaSK is the karroid equivalent of Restioid Sandy

Fynbos on arid sites. SaSK generally has a sparse cover

of vegetation with large spaces between the individual

plants. The canopy height is generally less than 0.5 m

and there are few understorey species. The vegetation is

quite patchy with relatively dense monospecific popula-

tions of several species occurring repeatedly through the

Bothalia 30.2 (2000)

171

landscape. In total, SaSK comprises a relatively small

area of MNR, and is confined largely to a single valley

floor in the eastern part (Figure 3).

Floristic composition

SaSK is characterised by locally dense populations of

Ruschia spp., Euphorbia deccusata and Zygophyllum

retrofractum. Perennial grasses such as Stipagrostis

namaquensis are also common except near the old kraal

sites, which are dotted through the landscape. Very infre-

quent patches of Willdenowia incurx’ata occur in locally

mesic areas of deep sand.

Structure

SaSk is characterised by localised patches of dwarf

leaf-succulent or fleshy-leaved shrubs and stem-succu-

lent. aphyllous plants. Grasses are common, and form

relatively dense stands except around the old kraal sites.

In the more mesic sites of SaSK. localized patches of low

to medium restioids occur.

Distribution and habitat

Sandy Succulent Karoo is found exclusively on deep

sands within the karoo matrix in the eastern part of MNR

(Figure 4), where the yellow soil is generally deeper than

1.2 m and there is a very low surface cover of rocks. The

sandy soil appears to be well drained and is probably of

aeolian origin, similar to the patches in the fynbos

matrix.

DISCUSSION

Mountain Fynbos

Campbell (1985) classified the mountain vegetation

of the Fynbos Biome, including the Cederberg Moun-

tains that lie just to the west of MNR. He used a combi-

nation of structural characteristics and higher taxa (fam-

ilies) to derive four major vegetation categories: three

non-fynbos and one fynbos. Within each category, he

described a number of communities. We only discuss

those communities that are relevant to the vegetation at

MNR.

Forest and Thicket is characterized by a high cover of

large shrubs or trees with leaves other than leptophylls.

In MNR, this community is usually found within the fyn-

bos matrix at sites that receive runoff from rocky out-

crops and appear to be protected from fire. Karroid and

Renoster shrubland occur over a large part of MNR

where the annual rainfall is too low to support fynbos, or

where soil conditions result in a pronounced summer

aridity (Cowling & Holmes 1992; Cowling et al. 1997;

Lechmere-Oertel & Cowling 1999). Dry Asteraceous

Fynbos is found on the rocky slopes and Dry Restioid

Fynbos on the sand plains. Dry Asteraceous Fynbos is a

xeric form of asteraceous fynbos (Table 1 ) characterized

by evergreen ericoid shrubs, mainly members of the fam-

ily Asteraceae, and by a low occurrence of restioids. Dry

Restioid Fynbos is characterized by the dominance of

restioids and other graminoids, and the relatively low

occurrence of ericoid shrubs.

Using floristic characters, Taylor (1996) classified the

vegetation of the northern Cederberg Mountains into 26

communities. The only non-fynbos community identi-

fied was thicket, and no succulent karoo or renoster

shrubland was encountered. Considering that this classi-

fication covers a centre of diversity for mountain fynbos,

it is not surprising that Taylor (1996) derived so many

communities. However, many of his communities cannot

be treated as management units as they form part of a

complex mosaic that would not be practical to resolve

and map for management purposes.

The classification of vegetation in other regions of the

Fynbos Biome has received attention from a number of

authors. However, because these surveys were carried

out at different scales and due to the high turnover in

species between regions (gamma diversity) (Bond 1981),

we have not referred to these studies in any detail.

Cowling & Holmes (1992) and Cowling et al. (1997)

provide good reviews of these phytosociological studies.

Succulent Karoo

Little work has been done on classifying succulent

karoo vegetation, partly because of taxonomic problems

in the dominant family, the Mesembryanthemaceae

(Hilton-Taylor 1987). Milton (1978) classified the vege-

tation at Andriesgrond near Clanwilliam, which is on the

western side of the Cederberg (60 km northwest of

MNR). Ten communities, including succulent and bro-

ken karoo, were classified, based on floristic and envi-

ronmental characteristics. However, none of these com-

munities were found at MNR, although there were defi-

nitely karroid elements common to both areas. Lane

(1978) briefly surveyed the Tanqua/Doom Karoo using

structural characters and derived four communities sepa-

rated along a moisture gradient. Snijman & Perry (1987)

described the flora of the Nieuwoudtville Wild Flower

Reserve that lies on the ecotone between Mountain

Fynbos and karroid shrublands. Their survey, however,

did not produce any communities that were apparent at

MNR, probably because the Nieuwoudtville Reserve lies

on dolerite, which gives rise to soil very different from

shale- or sandstone-derived soils.

Cowling & Holmes (1992) and Taylor (1996) provide

good summaries of the comparative benefits of a floris-

tic and structural approach to vegetation surveys.

Floristic-based methods, which require relatively

detailed knowledge of the flora, are more appropriate for

long-term management including the conservation of

rare and endemic species. They are also more applicable

to smaller areas. Structure-based methods are more use-

ful for experimental and autecological studies, which

require a detailed record of short-term changes in the

vegetation (e.g. establishing how structural traits vary

along resource and nutrient gradients). Structure-based

methods, being quicker to use than floristic-based meth-

ods, are also more appropriate for surveying extensive

areas of floristically complex vegetation such as fynbos

and succulent karoo, especially considering time and

financial constraints.

172

Bothalia 30,2 (2000)

The main aim of vegetation classification is to simpli-

fy the vegetation into robust units that are recognizable

and repeatable through the landscape. Mapping the nat-

ural resources benefits nature conservation because the

landscape is divided into homogenous units that can be

used as a basis for management planning (Pressy &

Bedward 1991). The vegetation at MNR is difficult to

classify phytosociologically because it comprises a com-

plex mosaic of communities at a fine scale, which would

be impossible to map or treat as management units.

Therefore it was necessary to design a classification sys-

tem for management purposes that overlooked the fine-

scale complexity and defined more pragmatic communi-

ties that were both mappable and meaningful for man-

agement (Pressy & Bedward 1991). It is, however, still

important to recognise the fine-scale mosaic of commu-

nities particularly in terms of conserving species diversi-

ty (Campbell 1985).

CONCLUSION

Community-environment generalizations

The fynbos communities are confined to the west of

MNR, which is generally higher in altitude, and thus

cooler, and receives more precipitation. Asteraceous

Fynbos Matrix, which occurs on all the rocky talus

slopes and areas of partially exposed bedrock, covers

most of this mesic area. Within the fynbos matrix, the

other fynbos communities are separated according to

land type. The deep sandy soils on the flat plains support

RSF. Fire-protected and mesic outcrops of sandstone

support Kloof Thicket with its tall shrubs and trees that

typically have mesophyllous evergreen leaves. The

rocky outcrops and exposed sheetrock support a combi-

nation of stunted fynbos shrubs and leaf-succulent

shrubs with Succulent Karoo affinity that are confined to

growing wherever enough soil has collected.

Moving eastward along the aridity gradient, there is a

gradual transition from fynbos into succulent karoo. This

transition is characterized by a slow turnover of growth

forms and species from leptophyllous and sclerophyllous

fynbos shrubs to leaf-succulent dwarf shrubs, and by the

loss of the restioid growth form. The transitional zone

between fynbos and succulent karoo is indistinct. There

are many areas where fynbos and succulent karoo ele-

ments extend deep into the other, in response to localised

moisture gradients caused by geological or topographic

features such as the Matjiesrivier Gorge. The steep north-

facing slopes of the Matjiesrivier Gorge, which represent

a relatively arid environment, give rise to a corridor of

succulent karoo deep into the fynbos side of the transi-

tional zone.

By combining the differential species from both the

growth form and floristic classifications with personal

notes made during the field trips, we believe that the

communities derived during this survey are ecologically

meaningful and will have use in management planning.

The communities were easily identifiable by a few key

species, simple growth form groups and knowledge of

the different land types. Thus the communities fulfil the

desired characteristics of being easily identifiable, eco-

logically meaningful and useful for mapping and man-

agement.

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CAMPBELL, B.M. 1988. Plant form in the mountains of the Cape,

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Pretoria.

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Vegetation of South Africa, Lesotho and Swaziland. Department

of Environmental Affairs and Tourism, Pretoria.

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of Stellenbosch Publications.

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National Botanical Institute, Pretoria.

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nutrients, fire and diversity. Oxford University Press, Cape

Town.

APPENDIX

List of species used in the classification, with their authors and families. Nomenclature from PRECIS (Arnold & De

Wet 1993) and authors of plant names from Brummitt & Powell (1992).

'.I

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Bothalia 30,2: 175-185 (2000)

Wetland vegetation of southern KwaZulu-Natal, South Africa

L. PERKINS*, G.J. BREDENKAMP* and J.E. GRANGER**

Keywords: Braun-Blanquet, classification. Grassland Biome, KwaZulu-Natal, South Africa, vegetation, wetlands

ABSTRACT

Vegetation data from southern KwaZulu-Natal were analysed, TWINSPAN classification separated the entire data set

(547 releves) into five subsets. One subset, representing the vegetation of the wetlands of southern KwaZulu-Natal, was further

classified by Braun-Blanquet procedures. Eight plant communities and fourteen subcommunities were identified and

described. The results can be integrated with existing phytosociological classifications of the Grassland Biome.

INTRODUCTION

The Grassland Biome Project was launched in

response to growing concerns about the state of South

African grasslands (Mentis & Huntley 1982). These

grasslands are under threat from afforestation, agricul-

ture, urbanization, invasive plants, overgrazing and plant

harvesting (Cowling & Hilton-Taylor 1994). A better

understanding and knowledge of grassland structure and

functioning is required so as to permit efficient land use

planning, utilization, conservation and management

(Mentis & Huntley 1982). To this end, a considerable

body of studies covering different parts of the Grassland

Biome have been produced (Van Wyk & Bredenkamp

1986; Bezuidenhout 1988; Turner 1989; Kooij 1990;

Matthews et al. 1991; Fuls et al. 1993; Smit et <7/. 1993;

Eckhardt et al. 1993; 1996a, b, c).

KwaZulu-Natal grasslands south of Estcourt (Figure

1 ) were identified as one area where insufficient phyto-

sociological data existed. This area was therefore select-

ed for a syntaxonomical and synecological study to fur-

ther the aims of the Grassland Biome Project. Previous

ecological work in southern KwaZulu-Natal has been

carried out by Bews (1917), Pentz (1949), West (1951),

Acocks (1953), Killick (1963), Edwards (1967), Moll

(1968), Granger (1976) and Granger & Schulze (1977).

However, most of the above studies concentrated on

broad vegetation types at a reconnaissance level, and are

not compatible with the present more detailed investiga-

tion using releve data and aiming at a hierarchical classi-

fication of vegetation at the plant community level.

Furthermore, no specific attempt has been made to cor-

relate the vegetation between Estcourt and the Eastern

Cape with Land Types (Smit et al. 1993) or any other

environmental stratification system.

The wetlands of southern KwaZulu-Natal are one of

the most important and neglected ecosystems in the

region (Begg 1986), and were investigated as one of the

main vegetation groups in the present study.

* Department of Botany, University of Pretoria, 0002 Pretoria.

** Department of Botany, University of Natal, Private Bag XOl, 3200

Scottsville, South Africa.

MS. received: 1999-05-26.

The present study, in line with the Grassland Biome

Project, aims in part to contribute to the updating of

FIGURE 1. — Location of study area within the Grassland Biome, and

distribution of Acocks' Veld Types within and adjacent to study

area: 44, Highland Sourveld; 45, Mist Belt ‘Ngongoni Veld; 56,

Highland Sourveld to Cymhopogon-Themeda Veld Transition;

58, Themeda-Festuca Alpine Veld; 65, Southern Tall Grassveld

(After Acocks 1988).

176

Bothalia 30.2 (2000)

Acocks’s Veld Type maps and to provide more detailed

phytosociological data within each Veld Type (Mentis &

Huntley 1982).

STUDY AREA

The study area is situated in southern KwaZulu-Natal,

between Estcourt and the KwaZulu-Natal-Eastern Cape

border. The northern boundary is 29° 00' latitude, the

southern boundary the Eastern Cape (former Transkei)

border, the western boundary is the Themeda-Festuca

Alpine Veld (Veld Type 58) of Acocks ( 1988) of the High

Drakensberg, and the eastern boundary is the irregular

transition area between grassland and other veld types

(mainly Valley Bushveld, Acocks 1988) (Figure 1). The

enclave of the Eastern Cape (formerly Transkei) within

KwaZulu-Natal was included in the study area. The

whole study area covers ± 14 400 km^ and includes large

areas under conservation management, large areas of

commercial farm land, much afforestation, and parts of

the former Homelands of KwaZulu and Transkei where

subsistence agriculture is chiefly practised.

Begg (1986) accepted the definition of a wetland by

the United States Fish and Wildlife Service: ‘land where

an excess of water is the dominant factor determining the

nature of soil development and the types of plant and ani-

mal communities living at the soil surface. It spans a

continuum of environments where terrestrial and aquatic

systems intergrade’.

Wetlands, defined as such, are found throughout this

area, from the high altitudes of the Drakensberg spurs

and foothills (1 800 m) to the relatively low incised river

valleys (900 m). Many wetlands have, however, been

drained for agricultural and other purposes. The alluvial

soils bordering certain wetlands are particularly fertile

(Phillips 1973) and agricultural practices in such areas

have altered many wetlands.

METHODS

Releves were compiled in 547 stratified random sample

plots. Stratification of the landscape was based on

Turner’s physiographic regions of Natal in Phillips

(1973) and Phillips’ bioclimatic subregions (Figures 2 &

3). Allocation of plots within the various regions was

done on a pro rata and area basis (Smit et al. 1993).

Representative numbers of plots were included in the fol-

lowing terrain units: crest, scarp, midslope, footslope and

valley floor/river bed (Land Type Survey Staff 1984).

Two hundred and fifteen sample plots were thus placed

in wetlands, typically on footslopes or valley bottoms.

Plot sizes were 25 m^(Eckhardt 1993).

The total floristic composition, and cover-abundance

values for each species, was recorded in each sample plot

using the Braun-Blanquet cover-abundance scale (Mueller-

Dombois & Ellenberg 1974). Environmental data recorded

included geology, topography, rockiness, soil form, gra-

dient, aspect and altitude (Matthews et al. 1991 ).

Data were incorporated into a data base using the

TURBOVEG programme (Hennekens 1996a). Two-Way

EIGURE 2. — Physiographic regions within and adjacent to study area

(stippled), after Turner in Phillips (1973): 1, Lesotho Plateau &

High Drakensberg E.scarpment; 2, Spurs & Foothills of the

High Drakensberg (the ‘Little Berg’); 13, Imphendle Block; 14,

Natal Midlands; 15, Howick Benchland; 16, Bulwer Block;

16a, Swartberg Block; 16b, Kokstad Block; 16c, Matatiele

Block; 16d. East Griqualand Uplands; 17, Winterton-Estcourt-

Muden Plain; 27, Underberg-Himeville Plain; 27a, East

Griqualand Plains; 28, Greytown-Pietermaritzburg-Richmond

Benchland; 29. Kranskop Divide; 32, Ixopo-Highflats Bench-

land; 33, Harding Benchland; 33a, Umzimkulu Benchland; 43,

Incised River Valleys.

Indicator Species Analysis (TWINSPAN: Hill 1979) was

applied to the entire data set in order to identify the major

plant communities. This initial classification was then

refined, using the programme MEGATAB (Hennekens

FIGURE 3. — Bioclimatic subregions within and adjacent to study area,

after Phillips (1973): 3, Mistbelt; 4c. Highland Montane (Open

Grassland to Wooded Savanna to Forest, Sub-Montane and

Highland); 4d, Montane Protea Savanna; 4e, Highland

Montane (Open Grassland to Wooded Savanna); 6. Open

Grassland to Wooded Savanna (Moister Faciation); 8, Open

Grassland to Wooded Savanna (Drier Faciation).

Bothalia 30.2 (2000)

TABLE 1 . — Phytosociological table of the wetlands of southern KwaZulu-Natal

177

Plant community number

Releve number In table

Unique releve nr.

(Turboveg database, Pretoria)

SPECIES GROUP A

Leucosidea sericea

Gunnera perpensa

Salix mucronata subsp. woodii

Rhus dentata

Cymbopogon validus

Cliffortla nitidula

Ischaemum fasciculatum

Rubus ludwigii

SPECIES GROUP B

Rhamnus prinoides

Euclea crispa

Ilex mitis

Geranium pulchrum

Hallerla luckja

Olinia emarginata

Buddleja salviifolia

Diospyros austro-africana

SPECIES GROUP C

Myrsine africana

Cephalarla natalensis

Pteridium aquilinum

Passerjna montana

Erica anomala

Schizostylis coccinea

Cliffortia paucistaminina

Passerina fUitormis

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Erica evansii

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Hyparrhenia tamba

Setaria sphacelata

SPECIES GROUP L

Miscanthus capensis

Acacia meamsil

SPECIES GROUP M

Elionurus muticus

Helichrysum argyrophyllum

Senecio retrorsus

SPECIES GROUP N

Themeda triandra

Tristachya leucothrix

Helichrysum pilosellum

Harpochloa falx

Oxalis depressa

Eragrostis racemosa

Mohocymbium ceresitforme

Trachypogon spicatus

Vemonia natalensis

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Bothalia 30,2 (2000)

179

1996b) with Braun-Blanquet procedures (Westhoff &

Van der Maarel 1987; Behr & Bredenkamp 1988). The

results of this classification for the Incised River Valleys

and Upland Savanna are presented in a phytosociological

table (Table 1).

RESULTS

The 215 releves from wetlands of all parts of the study

area. i.e. the Drakensberg foothills and Little ‘Berg’, mid-

lands, mistbelt and incised river valleys, were grouped

together by the TWINSPAN classification and after

removing the outlier releves, are presented in Table 1 .

These wetlands in general can be classified as Anmdi-

nella nepalensis-Paspalum dilatatiim (species group S,

Table 1) wetlands. Some of the wetlands are relatively

undisturbed, especially those in the Natal Drakensberg

Park, whereas others in agricultural areas show signs of

trampling and overgrazing. Deliberate or accidental dry-

ing up of wetlands for agricultural purposes has reduced

or eliminated many wetlands from southern KwaZulu-

Natal (Begg 1986). Franklin Vlei (30° 15' S, 29° 25' E),

for example, the third largest wetland in KwaZulu-Natal

after the Pongola Floodplain and Mkuze Swamp, is the

largest wetland in the present study, but has been reduced

by agricultural activities to less than half of its former

area (Begg 1986). These agricultural activities continue

unabated. Other wetlands have been incorporated into

exotic timber plantations, or drained for grazing land, but

the overall reduction of wetland habitat is impossible to

quantify (Begg 1986).

Encroachment of exotic weeds such as Verbena

bonariensis and Rubus cuneifoliiis or indigenous species

such as the grass Sporoboliis africanus and the forb

Helichrysum argyrophyllitm are indications of distur-

bance. A number of the wetlands and stream margins are

dominated by woody species: in protected areas these

are often Leucosidea sericea. Biiddleja salviifoUa and

Rhus species, whereas in disturbed areas Acacia mearn-

sii prevails. Even within protected areas Acacia mearn-

sii, A. dealbata. Verbena bonariensis and Rubus

cuneifoliiis are present in small quantities and the Park

authorities maintain ongoing control measures against

these invaders.

Other wetlands, particularly on moist footslopes

between a midslope and valley bottom, have no woody

species and instead support a variety of grasses and

forbs. Grazing and trampling by livestock is a common

problem on these footslopes. except in nature reserves

and other well-managed areas.

The terrain unit classified as valley bottom (Land

Type Survey Staff 1984) may contain a flowing stream

or a still pan (vlei) or marsh. A feature of the vleis and

marshes is the low species diversity in individual marsh-

es, but a high species turnover from one marsh to the

next. This phenomenon was also noted by Fuls et al.

(1993) and Eckhardt (1993). A large number of relative-

ly small communities and subcommunities is therefore

apparent in Table 1. Eight communities and 14 subcom-

munities were identified. These are:

1. Miscanthus capensis-Leucosidea sericea-Cliffortia

nitidida Scrub Community of stream margins

1.1. Miscanthus capensis-Leucosidea sericea Disturbed

Scrub Subcommunity

1.2. Miscanthus capensis-Leucosidea sericea-Euclea

crispa Scrub Subcommunity

1.3. Miscanthus capensis-Leucosidea sericea-Myrsine

africana Streambank Subcommunity

1 .4. Miscanthus capensis-Passerina montana-Erica

anoinala Fynbos Subcommunity

1.5. Carex austro-africana-Oenothera indecora Disturb-

ed Streambank Subcommunity

1.6. Miscanthus capensis-Artemisia afra Streambank

Subcommunity

2. Typha capensis-Paspalum urx’illei Pan Community

2.1. Typha capensis-Mariscus congestus Undisturbed

Pan Subcommunity

2.2. Typha capensis-Solanum mauritianum Disturbed

Pan Subcommunity

3. Hyparrhenia rufa-Conyza albida Disturbed Flood-

plain Community

3.1. Conyza albida-Sporobolus africana Moderately

Disturbed Floodplain Subcommunity

3.2. Conyza albida-Hyparrhenia tamba Severely Dis-

turbed Floodplain Subcommunity

4. Elionurus muticus-Helichrysum argyrophyllum Over-

grazed Floodplain Community

4.1. Elionurus muticus-Senecio retrorsus Wet Over-

grazed Floodplain Subcommunity

4.2. Elionurus muticus-Thenieda triandra Drier Over-

grazed Floodplain Subcommunity

5. Themeda triandra-Helichrysum aureonitens Grassy

Floodplain Community

5.1. Themeda triandra-Monopsis decipiens Wet Grassy

Floodplain Subcommunity

5.2. Themeda triandra-Tristachya leucothrix Drier

Grassy Floodplain Subcommunity

6. Hyparrhenia hirta-Eragrostis plana Grassy Wetland

Community

7. Rubus cuneifolius-Verbena bonariensis Disturbed

Wetland Community

8. J uncus effusus-Polygonum hystriculum Wetland

Community

PLANT COMMUNITY DESCRIPTIONS

In the following descriptions, the species groups al-

ways refer to Table 1 .

1. Miscanthus capensis-Leucosidea sericea-Cliffortia

nitidida Scrub Community

Diagnostic species: the woody Leucosidea sericea,

Clijfortia nitidida, Salix miicronata subsp. woodii, Rubus

ludwigii and Rhus dentata, the grasses Cymbopogon

validus and Ischaemum fasciculatum and the forb

Giinnera perpensa (species group A).

Dominant species: the shrub/small tree Leucosidea

sericea (species group A) and the grasses Miscanthus

180

Bothalia 30,2 (2000)

capensis (species group L) and Anmdinella nepalensis

(species group S).

Common species: the shrubs Cliffortia nitidula, Salix

mucronata subsp. woodii and Rhus dentata (species

group A) and the large-leaved forb Gimnera perpensa

(species group A).

This community is restricted to the western part of the

study area, generally above 1 500 m and is found on the

banks of steep gullies and in protected places near water

where there is generally low grazing pressure, and where

the occurrence of fire is infrequent. The soils are clayey

and drainage is often slow. A visually conspicuous fea-

ture of these gullies is the prominence of the exotic

woody encroacher Acacia mearnsii (species group L).

This encroachment is considered as a serious threat to the

indigenous wetland vegetation of KwaZulu-Natal.

1.1. Miscanthus capensis-Leiicosidea sericea Disturbed

Scrub Subcommunity

Diagnostic species: the woody Leucosidea sericea,

Salix mucronata subsp. woodii and Rhus dentata, the

grass Cymbopogon validus and the forb Gimnera per-

pensa (species group A).

Dominant species: the small tree Leucosidea sericea

(species group A) and the tall grasses Miscanthus capen-

sis (species group L) and Arundinella nepalensis.

Common species: the forb Senecio inornatus (species

group S).

This subcommunity tends to form long thin patches of

woody vegetation alongside streams and gullies and

rarely exceeds a width of three metres. It is found in gul-

lies and streambank verges in the Little ‘Berg’ but shows

signs of disturbance, either in the form of woodcutting or

utilization by livestock. The relative paucity of species is

a result of this utilization. The soils are loamy clay and

slow-draining.

1.2. Miscanthus capensis-Leucosidea sericea-Euclea

crispa Scrub Subcommunity

Diagnostic species: the shrubby Halleria lucida,

Buddleja salviifolia, Rhamnus prinoides, Euclea crispa.

Ilex mitis^ Diospyros austro-africana and Olinia emar-

ginata and the forb Geranium pulchrum (species group

B), with the absence of species group C.

Dominant species: the small tree Leucosidea sericea

(species group A) and the grass Arundinella nepalensis

(species group S).

Common species: the woody Rhus dentata and Salix

mucronata subsp. woodii (species group B), the grasses

Paspalum dilatatum and Paspalum urvillei (species

group S), the forbs Gunnera perpensa (species group A),

Polygonum hystriculum and Senecio inornatus (species

group S) and the sedges Mariscus conge stus and J uncus

ejfusus (species group S).

This subcommunity is found on streambanks and in

gullies in the Little ‘Berg’ above about 1 600 m, in shel-

tered spots where there is little disturbance, which

accounts for the greater species richness compared with

the Miscanthus capensis-Leucosidea sericea Subcom-

munity. The soils tend to be moist and fairly clayey. The

herbaceous cover varies from 80-90 % and the arboreal

cover, from 55-65 %. Many of the plants in species

group B are shade-loving. Killick (1990) refers to this

subcommunity as the Leucosidea sericea Scrub.

1.3. Miscanthus capensis-Leucosidea sericea-Myrsine

africana Streambank Subcommunity

Diagnostic species: the shrubs Myrsine africana and

Passerina montana and the fern Pteridium aquilinum

(species group C).

Dominant species: the grasses Miscanthus capensis

(species group L) and Arundinella nepalensis (species

group S).

Common species: Leucosidea sericea (species group

A), Rhamnus prinoides and Buddleja salviifolia (species

group B).

This subcommunity is most common in fire-protected

areas along streams, in gullies and on rocky outcrops.

The herbaceous layer of this community is sparse, possi-

bly due to the shade created by the woody canopy.

1.4. Miscanthus capensis-Passerina montana-Erica

anomala Fynbos Subcommunity

Diagnostic species: the woody Cliffortia lanceolata

and Erica evansii and the herbaceous Phygelius

aequalis, Anthospermum monticola, Cycnium racemo-

sum and Valeriana capensis (species group D).

Dominant species: a scrubby component of Leucosidea

sericea and the grass Miscanthus capensis (species

group L).

This subcommunity is located at fairly high altitudes

in the Little ‘Berg’ above 1 800 m. It corresponds rough-

ly with the Cliffortia linearifolia scrub described by

Killick (1963), although different Cliffortia species were

found in the present study. Streambanks and gullies with

moist clayey soil are the habitat of this subcommunity.

Erica anomala and Erica evansii together with the

Passerina species form the heathland element in this

subcommunity. The Ericaceous belt or ‘Sub-alpine

Fynbos’ (Killick 1963) is typically found in more exten-

sive patches higher up the Drakensberg range, above

1 900 m (Killick 1963; Hilliard & Burtt 1987). The pre-

sent study included sample plots in the vicinity of 1 800

m, and in some of these the heathland elements were

recorded. The ‘Sub-alpine Fynbos’ was described as the

natural climax community of the sub-alpine belt by

Killick (1963).

1 .5. Carex austro-africana-Oenothera indecora Disturb-

ed Streambank Subcommunity

Diagnostic species: the sedge Carex austro-africana,

the weedy Oenothera indecora. Ranunculus multifidus,

Ciclospermum leptophyllum, the grass Agrostis lachnan-

tlia and the fern Cheilanthes quadripinnata (species

group E).

Dominant species: the sedge Carex austro-africana

(species group E) and locally, the reed Phragmites aus-

tralis (species group S).

Bothalia 30,2(2000)

181

Common species: the weedy Cirsium vulgare and

Pseudognaphalium luteo-album (species group S) and

the sedge Jiincus ejfusus (species group S).

Certain streambanks and marshes are disturbed by

overgrazing and trampling from domestic livestock,

causing soil erosion and consequent prominence of

weedy species. The presence of the exotic, though natu-

ralized tree Salix babylonica (species group E) is a con-

spicuous feature of this landscape.

1.6. Miscanthus capensis-Artemisia afra Streambank

Subcommunity

Diagnostic species: Artemisia afra, Acalypha punctata

(species group F).

Dominant species: the woody Leucosidea sericea

(species group A) and the grasses Miscanthus capensis

(species group L) and Hsparrhenia hirta (species group

Q).

Common species: the forbs Gunnera perpensa and

Senecio isatideus (species group A), and the grass

Cymbopogon validus (species group A).

The banks of flowing streams are the site of this sub-

community, which is not extensive but shows little of the

signs of disturbance of the Carex aiistro-africana-

Oenothera indecora Subcommunity. The soils are sandy,

which accounts for the dominance of Hyparrhenia hirta.

2. Typha capensis-Paspalum utxhllei Pan Community

Diagnostic species: Typha capensis (species group G).

Dominant species: Typha capensis (species group G),

the grass Paspalum urx’illei and the sedge Mariscus con-

gestiis (species group S).

Common species: the exotic wattle Acacia meamsii

(species group L), the forbs Pseudognaphalium luteo-

album and Polygonum hystriculum, the grass Arundi-

nella nepalensis and the sedge J uncus ejfusus (species

group S).

The many small pans found throughout the study

area commonly have Typha capensis which appears to

favour the still water, growing in the shallows. The

alpha species diversity of these pans is quite low, with

few species recorded per sample plot. However the

Beta diversity, or the diversity between one pan and the

next, can be quite high. Thus a nearby pan can have a

considerably different species composition, with only

Typha capensis being common to them both. This phe-

nomenon is reflected in the table, where species group

G consists only of Typha capensis. Few woody species

are found in or near these pans, although the exotic wattle

Acacia meamsii was recorded several times at disturbed

pans.

2.1. Typha capensis-Mariscus congestus Undisturbed

Pan Subcommunity

Diagnostic species: Typha capensis (species group G).

Dominant species: Typha capensis (species group G).

Common species: the forb Polygonum hystriculum, the

sedges Mariscus congestus and Juncus ejfusus and the

grasses Paspalum dilatatum, P. urx’illei and Arundinella

nepalensis (species group S).

A number of the pans were in relatively good condi-

tion, in that signs of trampling and overgrazing were

minimal or absent. The soils in this subcommunity are

clayey and drainage is slow.

2.2. Typha capensis-Solanum mauritianum Disturbed

Pan Subcommunity

Diagnostic species: the exotic bugweed Solanum mauri-

tianum and the forb Senecio harx’eianus (species group H).

Dominant species: Typha capensis (species group G)

and the reed Phragmites australis (species group S).

Common species: the grass Paspalum urxnllei, and the

sedge Mariscus congestus (species group S).

Some of these pans show signs of disturbance, in the

form of trampling and overgrazing, by domestic live-

stock. Moist clayey soils with slow drainage are charac-

teristic of these pans.

3. Hyparrhenia ruja-Conyza albida Disturbed Flood-

plain Community

Diagnostic species: the grasses Hyparrhenia ruja and

Sporobolus ajricanus and the weed Conyza albida (spe-

cies group I).

Dominant species: the grasses Hyparrhenia ruja and

Eragrostis plana (species group S). Common species:

the introduced weed Verbena bonariensis (species group

S), the exotic tree Acacia meamsii (species group F), the

sedge Mariscus congestus and the grasses Paspalum

urx’illei and P. dilatatum (species group S), and Mis-

canthus capensis (species group F).

This community is one of the most disturbed in terms

of grazing and trampling pressure, hence the high occur-

rence of exotic plants and weeds. Most of the sites of this

community are close to standing or running water, and

subjected to trampling pressure when animals approach

to drink. It is also extensively grazed by the animals at

this time, causing prominence of Sporobolus ajricanus

(species group I), which is associated with disturbed

areas (Gibbs Russell et al. 1991). The community is also

more accessible than the others described, generally

occurring on flatter terrain to the east of the Drakensberg

foothills. A large proportion of Midlands wetlands, habi-

tat for rare bird and amphibian species (Grimsdell &

Raw 1984), are disturbed in this way.

3.1. Conyza albida-Sporobolus africana Moderately

Disturbed Floodplain Subcommunity

Diagnostic species: the weed Conyza albida and the

grasses Hyparrhenia ruja and Sporobolus ajricanus

(species group I), with the absence of species group J.

Dominant species: the grasses Hyparrhenia ruja

(species group I), Paspalum urvillei and Phagmites aus-

tralis (species group S).

182

Bothalia 30,2 (2000)

Common species: the ruderals Verbena bonariensis and

Polygonum hystriculum, the grass Miscanthus jimceus,

the sedge Mariscus congestus and the woody exotic tree

Acacia mearnsii (species group S).

This subcommunity is subjected to a degree of distur-

bance but this is moderate as the grass species are more

dominant than the ruderals.

3.2. Conyza albida-Hyparrhenia taniba Severely Dis-

turbed Floodplain Subcommunity

Diagnostic species: the weeds Tagetes miniita, Bidens

pilosa and Gymnopentzia bifurcata (species group J).

Dominant species: the woody exotic tree Acacia mearn-

sii (species group L) and the grass Eragrostis plana

(species group S).

Common species: The grass Paspaliim urx’illei, the

sedge Mariscus congestus, and the forbs Verbena bonar-

iensis and Polygonum hystriculum (species group S).

The presence of ruderals is an indication of the

severely disturbed nature (in terms of grazing and tram-

pling) of this floodplain subcommunity. Standing water

is common on these floodplains during the summer

months, but they dry up during winter.

4. Elionurus muticus-Heliclirysum argyrophyllum Over-

grazed Floodplain Community

Diagnostic species: the grass Elionurus muticus and the

forbs Heliclirysum argyrophyllum and Senecio retrorsus

(species group M).

Dominant species: the grasses Elionurus muticus

(species group M) and Eragrostis plana (species group S).

Common species: the forbs Verbena bonariensis and

Polygonum hystriculum (species group S).

This community, is located on gentle footslopes and

floodplains and is subjected to a high degree of grazing

pressure.

4. 1 . Elionurus muticus-Senecio retrorsus Wet Over-

grazed Floodplain Subcommunity

Diagnostic species: the grass Elionurus muticus and the

forbs Heliclirysum argyrophyllum and Senecio retrorsus

(species group M), with the absence of species group N.

Dominant species: the grasses Elionurus muticus

(species group M) and Eragrostis plana (species group S).

Common species: the forbs Verbena bonariensis and

Polygonum hystriculum (species group S).

This subcommunity is found on floodplains with gen-

tle gradients of 0-2°. Water collects on these floodplains

and attracts livestock which causes grazing pressure

(Low & Rebelo 1996). Where standing water occurs, the

hydrophilic Polygonum hystriculum (species group S) is

very prominent.

4.2. Elionurus muticus-Themeda triandra Drier Over-

grazed Floodplain Subcommunity

Diagnostic species: the grass Themeda triandra (spe-

cies group N), with the absence of species group R

Dominant species: the grasses Themeda triandra (spe-

cies group N) and Elionurus muticus (species group M).

Common species: the grass Eragrostis curx’ula (species

group S) and the forb Heliclirysum argyrophyllum (spe-

cies group M).

This subcommunity shows less evidence of moisture

than the Elionurus muticus-Senecio retrorsus Subcom-

munity, and these floodplains are consequently under

less severe grazing pressure. The forbs Senecio retrorsus

and Heliclirysum argyrophyllum and the grass Elionurus

muticus (species group M) are less common and less

dominant, and there is a greater presence of other grass-

es such as Themeda triandra and Tristachya leucothrix

(species group N). Few woody species were recorded in

this subcommunity. The soils are loamy sand, with mod-

erate to rapid drainage. The gradient is from 3°-5° and

this explains the greater runoff of water compared to the

Elionurus muticus-Senecio retrorsus Subcommunity.

5. Themeda triandra-Helichrysum aureonitens Grassy

Floodplain Community

Diagnostic species: the grasses Themeda triandra,

Tristachya leucothrix, Harpochloa falx, Monocymbiurn

cereciiforme, Eragrostis racemosa and the forbs Heli-

chrysum pilosellum, Oxalis depressa and Vernonia natal-

ensis (species group N).

Dominant species: Themeda triandra (species group N).

Common species: the grasses Tristachya leucothrix,

Eragrostis racemosa, (species group N) and E. curvula

(species group S), and the forb Heliclirysum aureonitens

(species group S).

This community contains many more grasses than the

other wetlands. The position of these wetlands is usually

between a gentle midslope and a valley bottom or river.

Grazing is apparent in much of the community but is

usually not extensive, either because the land is well

managed or the site falls within a nature reserve. How-

ever, a small number of the grassy wetlands sample plots

showed minimal disturbance. Two subcommunities are

classified on the basis of moisture content.

5.1. Themeda triandra-Monopsis decipiens Wet Grassy

Floodplain Subcommunity

Diagnostic species: the grass Eragrostis capensis, the

forbs Senecio venosus, Oxalis obliquifolia, Monopsis deci-

piens, Commelina africana and Hypoxis rigidula (spe-

cies group P).

Dominant species: the grass Themeda triandra (species

group N).

Common species: the grasses Tristachya leucothrix,

Eragrostis racemosa (species group N) and E. curvula

(species group S), and the forb Heliclirysum aureonitens

(species group S).

Footslopes with a high moisture content are the site of

this subcommunity, during the rainy season shallow pans

may be formed here. Herbaceous plants are associated

Bothalia 30.2 (2000)

183

with this marshy ground and no woody species were

recorded. Disturbance is low on these footslopes, either

because of good management or (more frequently) the

occurrence of the sample site in a nature reserve. The

principal nature reserves for the conservation of these

wet footslopes are Craigie Bum, Coleford and Mt Currie.

5.2. Tliemeda triandra-Tristachya leucothrix Drier

Grassy Floodplain Subcommunity

Diagnostic species: Themeda triandra (species group

N) and the absence of species group O and P.

Dominant species: the grasses Themeda triandra

(species group N) and the forb Helichryswn aureonitens

(species group S).

Common species: the grasses Eragrostis cunnda (spe-

cies group S) and Setaria pallide-fusca (species group

R).

No woody species were recorded in this subcommuni-

ty. Signs of disturbance are minimal, this is indicated by

the relative absence of weedy species. The soils of these

drier footslopes are sandier than those of the Themeda

triandra-Monopsis decipiens Subcommunity, and drain-

age is more rapid. This accounts for the absence of such

moisture-loving forbs such as Monopsis decipiens.

6. Hyparrhenia hirta-Eragrostis plana Grassy Wetland

Community

Diagnostic species: the tall grass Hyparrhenia hirta

(species group Q).

Dominant species: Hyparrhenia hirta (species group Q).

Common species: the grasses Eragrostis plana and

Paspaliim dilatation (species group S).

The extent of standing water in these wetlands is min-

imal, due to the relatively rapid drainage through the

stony soils of this community. H. hirta is a grass com-

monly associated with sandy, stony soils (Gibbs Russell

et al. 1991) and this community is relatively species-

poor. There are signs of grazing in this community but no

signs of overgrazing.

7. Rubiis ciineifolius— Verbena bonariensis Disturbed Wet-

land Community

Diagnostic species: the exotic bramble Rubiis cunei-

folius (species group R) with the absence of species

group Q.

Dominant species: the grasses Paspalum dilatatiim and

Eragrostis cun’ula and the sedge Mariscus congestiis

(species group S).

Common species: the exotic weeds Verbena bonarien-

sis and Cirsiiim vidgare (species group S).

This community is found in a variety of habitats, with

a gradient of moisture presence, but all showing signs of

alien encroachment and inadequate management of this

encroachment.

8. Juncus ejfusus-Polygonum hystricuhim Wetland Com-

munity

Diagnostic species: the forb Polygonum hystriculum

and the sedge Juncus ejfusus (species group S).

Dominant species: the sedge Juncus ejfusus and the

grasses Paspalum unillei and Eragrostis curvula (spe-

cies group S).

Common species: the weed Verbena bonariensis and

the grass Anmdinella nepalensis (species group S).

This vegetation is restricted to the habitat typically

found in shallow water in the littoral zone of marshes or

pans where there is extensive standing water. There is lit-

tle evidence of overgrazing in these wetlands.

DISCUSSION

Many of the vegetation communities described in the

present study can be related to communities previously

described by various authors. The Rhus dentata-Leu-

cosidea sericea thickets described by Eckhardt (1993)

and Fuls et al. ( 1993) in the southeastern Free State were

also found in the present study area in similar conditions:

moist, steep, south-facing slopes. These thickets represent

one of the vegetation types (Type 10) described by

O'Connor & Bredenkamp (1997).

The Passerina montana Afromontane Fynbos class

(O’Connor & Bredenkamp 1997) is not well represented

in the present survey, as the fynbos vegetation grows at

high altitude (1 800-2 800 m) (Killick 1963; Edwards

1967; Kruger 1979). However, the present survey includ-

ed releves at sufficiently high altitude (1 900 m) to

include some of the lower altitudinal limit of the heath-

land. These releves were sufficient to create a separate

subcommunity (Table 1). Du Preez & Bredenkamp

( 1991 ) also refer to the Passerina montana Afromontane

Fynbos, while a comparable fynbos community was

described from the eastern Cape mountains (Hoare 1997).

Riparian vegetation at lower altitudes (below ± 1 700

m) in the study area tends to be more disturbed as it falls

outside the conserved area of the Drakensberg Parks, and

is often trampled by livestock approaching drinking

points. Regular flooding also contributes to disturbance

of the riparian vegetation, and pioneer species and

exotics such as Verbena bonariensis, Tagetes minuta.

Datura fero.x, Oenothera rosea and Melinis repens are

distinctive of these areas. Grasses lining river banks

include Leersia hexandra, Anmdinella nepalensis,

Paspalum dilatatum, P. urx’illei and Miscanthus capen-

sis. Woody species commonly include the exotic wattle

Acacia mearnsii, the exotic willow Salix babylonica, the

indigenous willow S. mucronata, Rhus species, Rubus

ludwigii, Leucosidea sericea and Euclea species. Similar

riparian vegetation in the Free State grasslands was

described by Eckhardt (1993) and Smit (1992).

In open, still water Aponogeton and Polygonum species

frequently occur, with Polygonum spp. often growing in

the littoral zone.

The vegetation of marshes and bottomlands is often

dominated by one or a few species (Bloem 1988; Fuls et al.

1993). In the present study these species were usually

Phragmites australis, Typha capensis, Mariscus congestus.

184

Bothalia 30,2 (2000)

Conyza sp., or Cyperus spp. In still, shallow water or

marshy land, forbs such as Monopsis decipiens, Lobelia

flaccida and Pseiidognaphalium luteo-album are common.

Although the species diversity of a particular marsh might

be low, nearby marshes often support a very different plant

species composition. This phenomenon is also mentioned

by Smit (1992) and Fuls et al. (1993). The importance of

wetland conservation is underlined by this phenomenon as

species diversity might be easily overlooked if one marsh

is taken to be representative of wetland species diversity.

Other reasons for wetland conservation, besides preserva-

tion of biodiversity, include their functions as water storage

systems, stream flow regulators, flood attenuators, water

purifiers, erosion control agents, and specialized habitats

for various animals, as well as the the exploitation of wet-

land plants for ethnobotanical uses, especially as a source

of fibre for mat-making, baskets and thatching. Wetlands

also contain medicinal plants that are selectively removed

(Begg 1986; Walmsley 1988).

While the water catchment areas of the high

Drakensberg are adequately conserved by the KwaZulu-

Natal Parks Board (this being one of the prime reasons for

Drakensberg conservation), the wetlands downstream and

outside the park are poorly conserved and often over-

grazed and trampled. Burning of wetlands is also a com-

mon agricultural practice, decreasing the moisture and

organic material in the wetland (Begg 1986). Their func-

tions as outlined above are thus compromised. Affore-

station of wetland habitat has continued, despite numer-

ous commissions of enquiry and academic research

reports, most of them advising against wetland afforesta-

tion (Begg 1986). Other wetlands are drained for a vari-

ety of purposes, consistent with the widespread attitude

that wetlands are ‘wastelands’ (Begg 1986). Franklin Vlei

itself is suiTOunded by a town and railway yard, both of

which cause pollution, and is subject to drainage by vari-

ous farmers. Enforcement of the various regulations con-

cerning wetlands is difficult because of fragmented con-

trol, legal loopholes and inadequate definitions of words

such as ‘marsh’, ‘swamp’, ‘vlei’, ‘wetland’ and ‘drainage’

(Begg 1986). The downstream consequences of wetland

loss, in terms of flood attenuation, water storage and pol-

lution control are impossible to quantify. The vegetation

described in the present study can only be said to be ten-

tatively representative of these disturbed wetlands, as a

great deal of floristic change is expected to have occurred

due to the various disturbances.

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Bothalia 30,2: 187-200(2000)

Wetland vegetation in the North-eastern Sandy Highveld, Mpumalanga,

South Africa

P.M. BURGOYNE* , G.J. BREDENKAMP** and N. VAN ROOYEN**

Keywords: conservation, endangered habitats, Mpumalanga, phytosociology, plant communities. South Africa

ABSTRACT

The wetland vegetation of the high mountain grasslands of Mpumalanga was sampled by using stratification based on

geology and land types. Floristic data were classified by TWfNSPAN procedures and refined by using the Braun-Blanquet

method. This resulted in the recognition of four major wetland plant communities which are subdivided into eleven minor

plant communities. The major communities include the Phragmites australis Wetland occurring in relatively deep water,

the Miscanthus jimceus Wetland from moist river banks and wet drainage lines, the Eragrostis biflora-Stiburus

allopeciiroides Moist Grassland restricted to moist, poorly drained soils with a high water table, and Amndinella nepalensis

Moist Grasslands on black vertic soils.

INTRODUCTION

South Africa is rapidly approaching the position of

maximum exploitation of it’s natural water resources

(Walmsley 1988). Due to the increased demands that

people place on the natural habitat for the basic require-

ments of life, the natural resources are slowly but surely

being depleted and will not be able to provide for the

needs of the people (Bayer 1970; Allen 1972; Scheepers

1975; Asibey 1977; Mentis & Huntley 1982). Many wet-

lands have been destroyed beyond rehabilitation (Cowan

1991 ), making them one of the most endangered ecosys-

tem types in South Africa (Walmsley 1988). The Sabie,

Elands and Crocodile Rivers are fed by smaller rivers

originating from wetlands in and around the study area.

These wetlands are therefore important for maintaining

the flow of the rivers. Wetlands are therefore considered

as a scarce resource which should enjoy high conserva-

tion priority (Eckhardt et al. 1993; Smit et al. 1995). It is

vital that these natural wetland resources be maintained,

if not improved, to be able to cope with future pressures

being placed upon them.

The degradation of wetlands in this area affects the

continued health of these river systems (Walmsley

1988). This degradation is often caused by dessication of

the soil by pine forests in the catchment areas. An esti-

mated 30-35% of this study area is currently under cul-

tivation, with forestry being the most abundant practice

(Matthews et al. 1993). Apart from the exotic plantations

there is also a considerable problem with black wattle

(Acacia mearnsii) and silver wattle (Acacia dealbata) in

these drainage systems. The removal of these trees will

enable the natural vegetation to re-establish itself

(Mueller-Dombois & Ellenberg 1974).

An important conclusion of Myburgh et a/. ( 1 995 ) is that

individual wetlands in the Grootvlei area, Mpumalanga, are

poor in plant species, but that the different wetlands are

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

** Department of Botany, University of Pretoria, 0002 Pretoria.

MS. received: 1996-02-15.

floristically and ecologically quite distinct. Bloem et al.

(1993) similarly recognized a large variety of wetland com-

munities, each with unique species composition but with

low plant species richness, Ifom the Verlorenvallei Nature

Reserve in the North-eastern Sandy Highveld (Acocks

1988). This implies that a variety of wetland community

types will have to be included in a conservation programme

in order to preserve wetland biodiversity.

The phytosociological study of the wetlands of this

study area will not only contribute to the knowledge of

South African wetland vegetation and diversity, but can

be incorporated into the Grassland Biome Classification

Project (Du Preez & Bredenkamp 1991). Other compa-

rable vegetation surveys of the area include those of

Deall (1985), Deall et al. (1989), Bloem (1988), Turner

(1989) and Matthews (1991).

An important site rich in wetland communities is the

Lakenvalei, north of Belfast. A recommendation to pre-

serve this site is made, thus preserving many of the wet-

land communities of this study.

STUDY AREA

Physiography

The study area comprises two parts (Figure 1), both

situated at high altitudes in Mpumalanga, the average

being above 1 800 m. The area is mountainous and

rugged with deep ravines and steep cliffs in places and

gently sloping plains in others. This area has the highest

rainfall in the northern provinces, resulting in an unique

flora (Stevens 1989). The altitude ranges from 900-

2 331 m (Die Berg), which is the highest mountain in

Mpumalanga. Mount Anderson (2 284 m) lies just out-

side the bounds, to the north of the smaller study area.

Gradients of up to 57° are not uncommon, and many ver-

tical cliffs are to be found.

The geology of the study area was described in detail

(Burgoyne 1995) and only a brief summary is given here.

188

Bothalia 30,2 (2000)

12 14 16 18 20 22 24 26 28 30 32 34 36

FIGURE 1. — Locality, rainfall in mm and topography of the study area.

Nomenclature follows the South African Committee for

Stratigraphy (SACS 1980).

The Pretoria Group is dominated by quartzite and

shale, combined with some conglomerate and also some

chemical members. Volcanic eruptions occurred inter-

mittently and were normally localized. The soils derived

from sedimentary rocks are generally very shallow and

poor in nutrients, whereas the soils derived from the

lavas are richer in nutrients and are generally deeper

(SACS 1980). Fine-grained homfels together with silt

and sandstone, with minor layers of carbonaceous

(dolomite) and siliceous (chert) rocks, are to be found

throughout the study area in a broken, broad band from

the north to the south. These rock types give rise to soils

that are rich in minerals but the soils themselves are not

deep. Diabase sills and dykes can be found at intervals

all over the study area. These sills and dykes can some-

times be noted from aerial photographs by the difference

in the vegetation growing on them.

Bothalia 30,2 (2000)

189

The Ecca Series of the Karoo Supergroup consists of

sediments that were laid down in a freshwater basin

(Truswell 1970). Vegetation growing in the swamps gave

rise to the formation of coal which is extracted by means

of open cast mining. This is detrimental to the ecology of

the area because much of the topsoil is lost or ruined,

while seepage from open casts and waste dumps threat-

ens wetland systems.

Recent Quartemary Deposits (Pleistocene) occur in

the study area in most of the riverbeds where the prod-

ucts of many years of erosion have been deposited

(SACS 1980). Some wetland communities occur specif-

ically on these deposits along the rivers.

The Fa and Ac land types were studied in this area

(Land Type Survey Staff 1984). These two land types

constitute the land covered by grassland in the escarp-

ment area. Other land types were not studied for the rea-

son that they are not grassland, but forest or savanna, as

well as the fact that they are considered as lowlands. The

Fa and Ac land types differ from each other in terms of

microclimate, terrain form and geology (Land Type

Survey Staff 1979). The Ac land type represents the high

altitude plateau of the escarpment and occurs on sedi-

ments of various geological groups. Slopes are generally

shallow and may reach considerable lengths, forming

systems of catenas, creating a gently undulating plateau

landscape. The soils are defined as red-yellow and

apedal, with free drainage, and are dystrophic or meso-

trophic. Soil forms Hutton, Griffin, Clovely and Inanda

are frequent, although others may be present to a lesser

extent (Land Type Survey Staff 1979). The Fa land type

represents the rugged slopes with deep gorges and val-

leys of the escarpment, linking the grassland plateau with

the low-lying bushveld vegetation (Figure 1). It is prin-

cipally found on Vermont sandstone, Magaliesberg shale

and Steenkampsberg quartzite where the main soil-form-

ing process is weathering of the parent material, resulting

in the formation of relatively young soils with orthic top-

soils. The B horizons are formed by clay illuviation.

Although Glenrosa and Mispah Forms are the most com-

mon soil forms, other forms make up 7% of this land

type.

Climate

The climate of this region, according to the classifica-

tion of Koppen (Schulze 1947), is a temperate, rainy cli-

mate with a dry winter season (summer rainfall). There

is a C-type humidity province (subhumid), with grass-

land as the characteristic vegetation, together with a pre-

cipitation (P)/evaporation (E) index of 32 to 63 (Schultze

1947; Schulze & McGee 1978). The area is also classi-

fied as a w-type, which indicates a moisture deficiency in

the winter.

Lydenburg (1 439 m) and Waterval Boven (1 430 m)

have similar temperature ranges. Lydenburg records an

extreme minimum of-7.8°C and Waterval Boven records

the highest maximum temperature of 38.8°C (Weather

Bureau 1968). Sabie, however, has a maximum average

of 30.2°C, which is higher than the other towns in the

area. The temperatures recorded for Belfast are lower due

to its situation on an open, exposed, high-altitude plain

where free airflow occurs. Dullstroom (2 100 m) is

expected to be colder than the other towns in the area.

Frost occurs generally in the winter months in this

area and is common on all slopes, crests, as well as in the

valleys. It has been known to persist into the months of

September and October in the southern parts of the study

area. Frost causes teracettes on slopes in this region (Van

Zinderen Bakker & Werger 1974) by the daily frost-thaw

process. These terracettes are often colonized by pio-

neers because they are basically disturbed areas. During

rainy seasons the teracettes can be eroded, thus losing

valuable topsoil. The formation of frost-heaved tussocks

is also an important process in this region (Sigafoos &

Hopkins 1951; Hopkins & Sigafoos 1954).

The higher parts of the Steenkampsberg receive more

rainfall than surrounding areas (Figure 1). This is partly

due to the higher elevation coupled to the fact that these

mountains act as a barrier against which the rain falls.

The same can be observed in the northeastern part of the

Escarpment which receives a higher rainfall than the

areas surrounding it. The high rainfall in the area might

have contributed to the higher species diversity than in

most other parts of the country (MacArthur 1972; Huston

1979; Stevens 1989; Matthews et al. 1993).

From the Walter Climate Diagrams (Figure 2) it can

be seen that Sabie has a considerably higher rainfall

than the other towns in the area. Precipitation from mist

and fog supplements the rainfall rather significantly.

Mist is a common occurrence especially during the

months of October to February. From the data (Figure

3) it can be seen that the most mist occurs in Belfast,

but other areas that do not have ample weather stations

for recording this phenomenon, may have much higher

occurrences of mist. Areas above 2 000 m in altitude

are shrouded in mist for most of the summer months,

and mist can cover the entire escarpment area for three

weeks at a time.

Snowfalls are uncommon in this area, the last falls at

Dullstroom were recorded in 1974. The Steenkampsberg

Range is well above 2 000 m and it forms the tail-end of

the Drakensberg Range, thus any extreme weather con-

ditions caused by cold fronts moving northwards during

winter months would cause snow on these mountains

(Weather Bureau 1986).

METHODS

The geological and land type maps, 2530 Barberton

(scale 1:250 000) were used as units of statification.

Within each stratification unit, wetland sites were

chosen subjectively, as originally postulated in the

Braun-Blanquet approach (Braun-Blanquet 1932). This

was done to enable a survey of the limited and scattered

wetlands throughout the area. Thus a thorough recon-

naissance of the study area had to be done to choose the

wetland sites so that all wetland communities represent-

ed in the area were sampled. Within a particular wetland,

samples were placed randomly. Releves were compiled

in 39 sample plots.

190

Bothalia 30,2 (2000)

SABIE dlOBm) 17,3 1133.7 mm

WATERVAL BOVEN (1A30m) 16.7 8<,9mm

Based on a species-area curve of the vegetation of the

Verlorenvalei Nature Reserve (Bloem 1988), a sample

site of 200 was used, due to the low species diversity

of wetlands in high mountain grassland. This is also the

same size quadrat as recommended by Deall (1985).

For each quadrat the following data were recorded:

• location by farm name or nearest beacons and stand

co-ordinates (Land type series, 2530 Barberton 1979,

e.g. 2530 AA)

• altitude in metres was noted from the nearest contour

on the topocadastral map (Land type series, 2530

Barberton 1979)

• geology

• land type

• geomorphology position (Munnik et fl/. 1984)

• degree of disturbance, including disturbance due to

road construction, exotic invaders, soil erosion, over-

burning, trampling and power line construction, using

the following scale: 0, no visible erosion; 1, sheet ero-

sion; 2, donga erosion; 3, sheet and donga erosion

• degree of grazing, using the following scale: 0, no

recent grazing; 1, selectively grazed; 2, evenly

grazed; 3, heavily grazed

• soil depth (cm)

• soil texture was noted using the method as prescribed

by the FSSA(1980)

• soil form and soil series were noted following

Mac Vicar et al. ( 1 977)

• water depth and permanency were noted

FIGURE 2.— Walter Climate Dia-

grams for four towns in the

study area (Walter 1991).

• Braun-Blanquet cover-abundance values for every

species present (Westhoff & Van der Maarel 1978);

taxon names in accordance with Arnold & De Wet

(1993)

• height of vegetation

Classification of the floristic data was accomplished

by using a default TWfNSPAN (Two-way INdicator

SPecies ANalysis, Hill 1979). Both the releves and the

species are grouped into a specific order (Noy-Meir

1973; Gauch & Whittaker 1981). TWINSPAN produced

Months of the year

FIGURE 3. — The occurrence of mist at three stations in the study area.

Bothalia 30.2 (2000)

191

a table that was then refined by Braun-Blanquet proce-

dures (Werger 1974) as was proposed by Bredenkamp &

Bezuidenhout (1995). Type releves were chosen as the

most representative releve for a community by compar-

ing releves from the Braun-Blanquet tables (Table 1 ).

RESULTS

The floristic composition of the wetland plant com-

munities is given in Table 1. General characteristic

species for the wetlands (species group T) were identi-

fied by comparing this table with phytosociological

tables from other plant communities of the area

(Burgoyne 1995). Species which occur throughout all

wetland communities include the grasses Anmdinella

nepalensis, Alloteropsis sp., Aristida aquiglumis, Pemii-

setum thunbergii, Agwstis lachnantha, Hemarthria

altissima and Setaria pallide-fusca, the sedges Fuirena

piibescens, Mariscus congestus, M. sumatrensis, Eleo-

charis palustris, Pycreiis nilidus. and Kyllinga erecta,

the forbs Pycnostachys reticulata, Sebaea sedoides,

Meritha aquatica. Lobelia flaccida, Helichrysum dijficile

and H. mimdtii, and the monocot Eriocaidon dregei.

The species with the highest constancy values are

Fuirena pubescens (66%) and Anmdinella nepalensis

(66%), followed by Mariscus congestus (39%) and

Schoenoplectus corymbosus (32%).

The analysis of the wetland vegetation resulted in the

recognition of four communities and 1 1 subcommuni-

ties, which are classified as follows:

1. Phragmites australis Deep Wetland Community

1.1. Phragmites australis-Ficinia acuminata Deep

Wetland Subcommunity

1.2. Phragmites australis-Senecio microglossus Deep

Wetland Subcommunity

2. Miscanthus junceus Wetland Community

2.1. Alepidea amatymbica-Miscanthus junceus Moist

River Bank Subcommunity

2.2. Agrostis gigantea-Miscanthus junceus Moist

Grassland Subcommunity

2.3. Panicum schinzii-Miscanthus junceus Shallow

Wetland Subcommunity

2.4. Carex cognata-Miscanthus junceus Wetland Sub-

community

2.5. Ischaemum fasciculatum— Miscanthus junceus

Wetland Subcommunity

3. Eragrostis biflora-Stiburus allopecuroides Moist

Grassland Community

3.1. Helichysum aureonitens-Eragrostis biflora-Sti-

burus allopecuroides Moist Grassland Subcom-

munity

3.2. Disa patula-Eragrostis biflora-Stiburus allope-

curoides Moist Grassland Subcommunity

4. Anmdinella nepalensis Moist Turf Grassland Com-

munity

4.1. Hypericum aethiopicum-Arundinella nepalensis

Moist Turf Grassland Subcommunity

4.2. Imperata cylindrica-Arundinella nepalensis

Moist Turf Grassland Subcommunity

1 . The Phragmites australis Deep Wetland Community

Species group: C (Table 1 ).

Diagnostic species: the tall-growing reed, Phragmites

australis and bulrush Typha capensis.

Dominant species: hygrophylic forbs such as

Zantedeschia albomaculata, Berkheya speciosa and

the fern Thelypteris confluens which can grow in

almost pure stands.

This community occurs predominantly in deep water,

mostly deeper than 0.2 m and often deeper than 1.1m.

The soils are high in organic matter, representing the

Champagne Form.

This community is divided into two subcommunities

namely the Phragmites australis-Ficinia acuminata

Deep Wetland Subcommunity in water 0.2-0. 7 m deep,

mostly at altitudes of 2 000 m and lower, and the

Phragmites australis-Senecio microglossus Deep

Wetland Subcommunity in water deeper than 1.1 m,

mostly at altitudes higher than 2 000 m above sea level.

1.1. Phragmites australis-Ficinia acuminata Deep

Wetland Subcommunity

Type releve: 307.

Av. no. spp. per releve: 32.

Max. no. spp. per releve: 47.

Min. no. spp. per releve: 10.

Altitude: 980-2 000 m.

Geology: predominantly from the Magaliesberg and the

Steenkampsberg Formations although some shale from

the Strubenkop and Lakenvalei Formations is also pre-

sent.

Soil form: Champagne and deep alluvial soils high in

organic matter.

Height of vegetation: 2.6 m.

Species group: A (Table 1 ).

Diagnostic species: the sedge Ficinia acuminata, the geo-

phytes Dierama pendulum, Ornithogalum monophyl-

lum and Bnmsvigia radulosa, the forbs Wahlenbergia

sp., Cycnium racernosum, Cyphia stenopetala, Ascle-

pias dissona, Vigna vexillata, Vernonia hirsuta, V.

sutherlandii and Helichrysum pilosellum.

Dominant species: the reed Phragmites australis', the

reedbeds form important bird breeding sites (Figure 4).

Stands of these deep wetlands may cover a surface

area of up to 5 ha. This community is found where sea-

sonal fluctuation of the water level occurs. Only in the

most dry of seasons (as in the spring of 1992) is relative-

ly deep surface water not present.

192

Bothalia 30,2 (2000)

TABLE 1 . — Phytosociological table showing wetland communities

1 2 3 4

1.1 1.2 2.1 2.2 2.3 2.4 2.5 3.1 3.2 4.14.2

33111 2321222 130 03 32 130 3113 01321 23221 30 2

00453 4052480 725 41 00 716 1311 59139 41454 11 1

75651 3941740 206 19 87 301 3792 24591 48950 19 5

Species Group A

Ficinia acuminata

Dierama pendulum

Hah! enbergi a sp.

Cycnium racemosum

Cyphia stenopeta / a

Brunsi^igi a radu/osa

Asc I epi as dissona

V i gna vexi 1 1 ata

Orni thoga ! urn monophy H urn

Vernonia hirsuta

Vernoni a suther ! and i i

He I i chrysum pi lose I turn

Species Group B

Senecio microglossus

Polygonum mei sneri anum

Kniphofia multi flora

Polygonum sp.

Eragrostis cy I i ndri f I ora

Species Group C

++ + I

+++ I

++ + 1

++ I

+ + j

+ + I

+ + 1

+ I

+ + I

+ + 1

+ +

R

+

R

1+1+1 I

I++ II

I + + i I

I A| I

1+ I +1

Phragmi tes australis

Zantedeschi a a! bomacu! ata

Typha c a pens is

T helypteri s conf I uens

Berkheya sped os a

Species Group 0

Geranium mu 1 1 i sectum

Alep idea amatymbica

Pelargonium alchemi I loides

Did is rotund i f o! i a

Kyllinga paucif lora

Adiantum capi 1 1 i s-vener i s

Di heteropogon amp! ectens

Aeschynomene rehmanni i

Asc I epi as cultriformis

Cepha I ar i a attenuata

Species Group E

Sder/'a d i eter leni i

Agrostis gigantea

Sphagnum truncatum

Senecio stri at i fol ius

Rorippa nasturt i um-aquat i cum

Aristea sp.

Disperis cooper i

Koeleria capensis

Comme / / na afri cana

Hypoxis rigidula

Alepidea setifera

Aloe ecklonis

Tu I bachi a nutans

He ! i chrysum subg ! omeratum

Peucedanum sp.

MB 1B| 3A1 + 1I I I

I++ + I ++ R I I++I

I R R| + 1R |R I I

1+ + I ++ I II

I

1

I

+ I I +

I 1 +

I

1 I

I

Species Group F

Panicum schinzii

Aponogeton junceus

Crassula pel lucida

Lolium mu it if lorum

Habenari a sp.

Species Group G

Epilobium salignum

Si urn repandum

kni phof i a I i near i f o! i a

Hdcus lanatus

Eucomus comosa

1+ I I nil I

I +1 +I++I++I I

I I +1 IR+I I

+ I I I I++I I

I I I I++I I

+ -r I+ + + 1 |+ + +j++|++|

+ +1 + + I +++ I ++ I ++ I

R I++ I++ I IRRI+

1 II 1 I II Mil

+ I + 1 I I IRRI

+

1

Bothalia 30,2 (2000)

193

TABLE 1. — Phytosociological table showing wetland communities (cont.)

1 2 3 4

1.1 1.2 2.1 2.2 2.3 2.4 2.5 3.1 3.2 4.14.2

33111 2321222 130 03 32 130 3113 01321 23221 30 2

00453 4052480 725 41 00 716 1311 59139 41454 11 1

75651 3941740 206 19 87 301 3792 24591 48950 19 5

Species Group H

Nerine angust i f o I i a

Carex cognata

Disa cooper i

Eu tophi a ova! is

Butbine abyss i ni ca

Eutophia t eontog i ossa

Species Group I

Carex austro-af ricana

Rumex I anceo I atus

Gunner a per pens a

Festuca caprina

Satyr i urn haitackii

Senecio serratu ! i odes

kniphofia fiuviatiiis

Leers i a hexandra

Species Group J

i schaemum fascicuiatum

Doiichos faiciformis

Conyza pinnata

Digitaria eyiesii

Penni setum macrourum

Digitaria ftaccida

Pycreus sp.

Atys i carpus rugosus

Satyrium parvifiorum

He! i chrysum opacum

Habenari a I i thophi I a

Eragrost i s curvu t a

Berkheya echinacea

Species Group K

M iscanthus junceus

Ranunculus meyeri

Ranunculus multi fidus

Juncus exsertus

Species Group L

He ! i chrysum aureoni tens

Hypoxis filiform is

Agrostis eri antha

Asclepias multicaul is

Sebaea leiostyl a

Juncus dregeanus

Oxalis obliqui folia

Anther i cum cooper i

Ficinia sp.

Species Group M

Buchner a glabrata

Habenar i a davata

Disa aconi toides

Andropogon eucomi s

Chironia purpurescens

Species Group N

Disa patuta

He I i ctotr i chon turgidulum

Orni thogalum tenui f o! ium

Eragrost i s c a pens is

Bulbostylis sp.

Plectranthus sp.

Alep idea gracilis

+ I I I +R|

I I I A|

I I I +1

1 I I +1

I I I +1

IB+BB+I 4

1+ + 1+ +

I 4 B I I AA I 44 1 111 +

I 1I++I++I +111 I

I++ I 1 1 +1 ++I+

I +I++I++I I ++I

I 1

I I

I 1 +

I I

I

1

I

I I I I 111 l+AA A|

I I I I I I+++++I

1+11 I++I + I I 1 A A|

I I I I +1 IR + +1

I III I I ++ +1

I I++I I I I++ + I

I I I I I I + +1

I I I I I I R R|

I I I I I I 1 I

+ I I

++ M I

B I I 1

194

Bothalia 30,2 (2000)

TABLE 1. — Phytosociological table showing wetland communities (cont.)

Species Group 0

Drosera madagascar i ens i s

Eragrostis bi flora

Asco ! epi s capons i s

Diorama sp.

Polygala uncinata

Utricular! a prohonsilis

Just id a pot i o! ari s

Species Group P

SchoonopI octus corymbosus

Mariscus konionsis

Juncus oxycarpus

Bulbostyl i s burcho! I i i

Lodobouri a coopori i

Gladiolus longicollis

Denokia capons is

Satyrium ! ongi cauda

OldonI andi a horbacea

Species Group Q

Hypor i cum aothi opi cum

Crassula sarcocau I i s

Glad id is ocklonis

Lodobouri a sp.

Species Group R

Pycrous macranthus

Imporata cy! i ndrica

Senocio I at i fo! i us

1 +

++I

+ I

R R I

I++

I++1

I++I

I

1

I

1

I

1 +

Species Group s

Bothalia 30,2 (2000)

195

1.2. Phragmites australis-Seuecio microglossiis Deep

Wetland Subcommunity

Type releve: 243.

Av. no. spp. per releve: 36.

Max. no. spp. per releve: 36.

Min. no. spp. per releve: 32.

Altitude: above 2 000 m.

Geology: quartzite of Steenkampsberg and Lakenvalei

Formations.

Soil form: Champagne.

Height of vegetation: 2.6 m.

Species group: B (Table 1).

Diagnostic species: the forbs Senecio microglossiis.

Polygonum meisnerianiim, Polygonum sp. and the

grass Eragrostis cylindri flora, together with the char-

acteristic and conspicuous monocot Kniphofia multi-

flora, which can be seen from afar when flowering.

Dominant species: the tall-growing reed Phragmites aus-

tralis.

This high altitude community occurs in the central

parts of the deep wetlands and the surface area covered

may be up to 15 ha. Lakenvalei between Belfast and

Dullstroom is a good example of this community. These

wetlands are periodically burned and according to some

of the farmers in the region this is beneficial to the breed-

ing of birds like the wattled crane, who prefer open areas

where they can clearly see any foes approaching

(Tarboton 1981).

2. Miscanthus junceus Wetland Community

Species group: K (Table 1 ).

Diagnostic species: the robust grass Miscanthus junceus

dominant, and the forbs Ranunculus meyeri and

Ranunculus multifidus, together with the sedge

Juncus exertus. A shrinking of the Miscanthus

junceus wetlands was observed over a period of two

years, possibly aggravated by low rainfall.

These wetlands are represented by moist river bank

communities to wet drainage lines. No trees occur and

FIGURE 4. — The Phragmites autra-

lis deep wetlands, showing

the dominance of the reed

Phragmites australis.

rockiness is confined to the presence of small pebbles in

the sediments. The moisture regime varies from moist soil

which releases free water when trodden on, to surface

water of a depth of 0.7 m. The area covered by these wet-

lands is generally shrinking, due to management practices.

2.1. Alepidea amatymbica-Miscanthus junceus Moist

River Bank Subcommunity

Type releve: 172.

Av. no. spp. per releve: 18.

Max. no. spp. per releve: 30.

Min. no. spp. per releve: 9.

Altitude: 1 200-2 000 m.

Geology: mainly from quartzites and shales of the

Pretoria group.

Soil form: Champagne, deep, dark and rich in organic

content, Arcadia may also be present.

Height of vegetation: 0.15-0.65 m.

Species group: D (Table 1).

Diagnostic species: the forbs Geranium multisectum,

Alepidea amatymibica, Pelargonium alchemilloides,

Diclis rotundifolium, Aeschynomene rehmannii, Ascle-

pias dissona and Cephalaria attenuata, together with

the sedge Kyllinga paucifolia, the fern Adiantum capil-

lis-veneris and the grass Diheteropogon amplectens.

Dominant species: the tall grass Miscanthus junceus.

This subcommunity is found on wet soils. The mois-

ture status is such that when trodden on, water is released

to the surface.

2.2 Agrostis gigantea-Miscanthus junceus Moist Grass-

land Subcommunity

Type releve: 41 .

Av. no. spp. per releve: 45.

Max. no. spp. per releve: 53.

Min. no. spp. per releve: 29.

Altitude: 1 400-2 200 m.

Geology: shale and diabase from various geological for-

mations.

196

Bothalia 30,2 (2000)

Soil form; Champagne, deep, rich dark loam, high in

organic matter.

Height of vegetation: 0.4 m.

Species group; E (Table 1).

Diagnostic species: the moss Sphagnum africanum,

which grows in mats, the remains of which may form

a peat bog, and the forbs Senecio striatifolius, Ro-

rippa nasturtium-aqiiaticuni (much eaten by cattle),

Commelina africana, Alepidea setifera and Peuce-

danum sp. Also the geophytes Aristea sp. Hypoxis

rigidula. Aloe ecklonis, and Tulbaghia nutans togeth-

er with the orchid Disperis cooperii, the grass Koe-

leria capensis and the asteraceous forb Helichrysum

subglomeratum.

Dominant species; the grasses Scleria dieterlenii and

Agrostis gigantea.

This community is represented by numerous small

patches of shallow wetlands that are a result of a raised

water table caused by the lithological formations in the

area. This wetland type may also be formed by the dry-

ing up of larger wetlands through destructive practises.

The moisture status is such that visible water is pre-

sent on the soil surface but never deeper than 0.05 m.

The vegetation is extensively grazed especially in the dry

season when it is still green, as compared to the adjacent

drier grassland vegetation.

The species richness (53) in this community is higher

than in any other wetland community and should there-

fore be considered important to conserve.

2.3 Panicum schinzii-Miscanihus junceus Shallow

Wetland Subcommunity

Type releve: 207.

Av. no. spp. per releve: 22.

Max. no. spp. per releve; 34.

Min. no. spp. per releve: 9.

Altitude: 1 200-2 000 m.

Geology: varies greatly but quartemary deposits are domi-

nant.

Soil form; Champagne, deep, rich dark loam, high in or-

ganic matter.

Height of vegetation: 0.7 m.

Species group: F (Table 1).

Diagnostic species; the grasses Panicum schinzii and

Loliurn multiflonun, the semisucculent forb Crassula

pellucida, which may form mats of pure stands in

patches, possibly due to it’s seed dispersal mecha-

nism, the orchid Habenaria sp. and the water plant

Aponogeton junceus.

Dominant species: Miscanthus junceus.

This subcommunity covers a larger area than the

Agrostis gigantea-Miscanthus junceus Moist Grasslands

Subcommunity (2.2) and is also found along the periph-

ery of the larger wetlands, where the water depth is up to

0.8 m. Moisture levels in these wetlands vary greatly and

can become quite dry, especially in the winter when rain-

fall is low.

General species occurring in all the above-mentioned

Miscanthus communities (1.1. to 2.3.) are: the forbs

Epilobium salignum, Sium repandum, the geophytes

Eucomis comosa, Kniphofia linearifolia and the grass

Holcus lanatus.

2.4 Carex cognata-Miscanthus junceus Wetland Com-

munity

Type releve: 61.

Av. no. spp. per releve: 26.

Max. no. spp. per releve: 34.

Min. no. spp. per releve; 17.

Altitude: 1 600-1 900 m.

Geology: loose Quartemary deposits.

Soil form: Champagne, deep, rich dark loam, high in or-

ganic matter.

Height of vegetation: 0.6 m.

Species group; H (Table 1).

Diagnostic species: the sedge Carex cognata, the geo-

phytic orchids Disa cooperii, Eulophia leontoglossa,

E. ovalis and the geophytes Nerine angustifolia and

Bulbine abyssinica.

Dominant species: Miscanthus junceus.

Large and perennial, these wetlands contain ± 0.7 m

water all year and are fed by fountains forming the

beginnings of the river systems of Mpumalanga. The

only time when they may have little or no water in them

is during severe drought.

Species group I represents general species occurring

in communities 1.1 to 2.4 and include the sedge Carex

austro-qfricana, the forbs Riimex lanceolatus, the con-

spicuous Gunnera perpensa with its large round leaves,

Senecio serrulatuloides, the orchid Satyrium hallackii,

the monocot Kniphofia fluviatilis and the grasses

Festuca caprina and Leersia hexandra.

2.5 Ischaemum fasciculatum-Miscanthus junceus Wet-

land Community

Type releve: 137.

Av. no. spp. per releve: 19.

Max. no. spp. per releve: 36.

Min. no. spp. per releve: 9.

Altitude: 1 200-2 000 m.

Geology: Quartzites of the Pretoria Group.

Soil forni: vertic soils of the Arcadia soil form.

Height of vegetation: 0.7 m.

Species group: J (Table 1 ).

Diagnostic species: the grasses Ischaemum fasciculatum,

Digitaria flacida, D. eyelsii, Pennisetum macrourum

and Eragrostis cunnila, the orchid Satyrium pan’iflo-

rum, and the sedge Pycreus sp.

Dominant species: Stiburiis alopecuroides and Ischae-

mum fasciculatum.

These communities cover an area of less than three

hectares and are subjected to periods of drying out. The

water is also never as deep as the wetlands of the previ-

ous community (2.4) and is only ± 0.4 m deep. The

slopes occupied by these wetlands are also steep (± 12°)

as opposed to the wetlands in the previous community

where a gradient of 3° is the steepest noted. Running

water is therefore a feature of this community.

Bothalia 30,2 (2000)

197

Forbs in this community include Dolichos falciformis,

Conyza pinnata, Berkheya echinacea, Alysicarpus nigo-

sus and Helichrysum opacum. Species group K (Table 1 )

are found throughout the previously mentioned commu-

nities (1.1 to 2.5) and include the sedges Miscanthus

jiinceiis and Jiinciis exsertus, together with the forbs

Ranunculus meyeri and R. midtifidus.

It is interesting to note that the robust grass Mis-

canthus junceus is relatively absent in the Senecio

microglossus-Phragniites australis Community, where-

as it is most dominant in all the other communities. The

explanation for this may be that Phragmites australis and

Carex austro-africana out-compete this species to such

an extent that it cannot survive.

3 . Eragrostis biflora-Stiburus alopecuroides Moist Grass-

land Community

Species group; O (Table 1).

Diagnostic species: the insect trapping plants Drosera

madagascariensis and Utricularia prehensilis, the

forbs Polygala uncinata and Justicia petiolaris, the

sedge Ascolepis capensis, the geophyte Dierama sp.

and the grass Eragrostis biflora. This grass is also vis-

ibly prominent due to its fine leaves and light pink,

fluffy inflorescence.

These wetlands are found in poorly drained soils. In

most cases the lithology is impenetrable, resulting in a

raised water table. Palatable grasses are predominant,

affording much grazing.

This community is divided into two subcommunities:

Helichysum aureonitens— Eragrostis biflora-Stiburus

alopecuroides Moist Grassland Subcommunity which is

found on soils of diabase origin, and Disa patula-

Eragrostis alopecuroides Moist Grassland Subcom-

munity, found on soils of sedimentary origin.

3.1. Helichysum aureonitens-Eragrostis biflora— Stibunis

alopecuroides Moist Grassland Subcommunity

Type releve: 194.

Av. no. spp. per releve; 31.

Max. no. spp. per releve: 37.

Min. no. spp. per releve: 23.

Altitude; 950-2 200 m.

Geology; igneous rock, particularly Transvaal diabase.

Soil form: Champagne, 0.3 m deep, relatively rich in

organic matter, stones may occur in this community

but their size never exceeds 0.05 m diam.

Height of vegetation; 0.4 m.

Species group; L (Table 1 ).

Diagnostic species; the geophytes Hypoxis fdiformis and

Oxalis obliquifolia, the forbs Helichrysum aureo-

nitens, Sebaea leiostyla, Asclepias multicaulis and

Anthericum cooperii and the sedges Juncus dregea-

nus and Ficinia sp. together with the grass Agrostis

eriantha.

Dominant species: Stibunis alopecuroides, Fuirena pu-

bescens and Eragrostis biflora.

This community is found on all aspects and slopes,

and the area covered by stands of this community never

exceeds 1.5 ha. There are thus small areas where the

water is trapped by the geological strata. No visible water

is present but, during the rainy season, water may seep

out when trodden on. Grazing is moderate to heavy due to

the presence of palatable grasses in this community.

Species group M represents a group of species com-

mon to the Ischaemum fasciculatum-Miscanthus junceus

Wetland Subcommunity (2.5) and the Helichrysum aureo-

nitens-Eragrostis biflora-Stiburus alopecuroides (3.1)

Wetland Subcommunity. These include the forbs

Buchnera glabrata and Chironia purpurescens, the

orchids Habenaria clavata and Disa aconitodes together

with the grass Andropogon eucomis.

3.2. Disa patula-Eragrostis biflora-Stiburus alopecu-

roides Moist Grassland Subcommunity

Type releve: 244.

Av. no. spp. per releve: 19.

Max. no. spp. per releve: 36.

Min. no. spp. per releve: 9.

Altitude: 1 200-2 000 m.

Geology: quartzites of the Pretoria Group.

Soil form; Champagne or Mispah.

Height of vegetation: 0.4 m.

Species group: N (Table 1 ).

Diagnostic species: the orchid Disa patula, the forbs

Plectranthus sp. and Alepidea gracilis, the geophyte

Ornithogalum tenuifolium, the sedge Bulbostylis sp.

and the grasses Helictotrichon turgidulurn and Era-

grostis capensis.

Dominant species: the grass Stibunis alopecuroides domi-

nates the vegetation and especially in the months when

it flowers, the light purple heads are prominently visi-

ble. Also when mist is present this community stands

out visibly from those next to it due to the dew drops

that are caught in the hairs of the leaves of this plant.

In this community depressions with poor drainage are

periodically flooded in the wet season, thus forming wet-

lands, which dry out if further rainfall ceases or is insuffi-

cient. The sandy loam soil is dark, rich in organic materi-

al and is not deeper than 0.4 m, having a rock base which

is normally not penetrable to water, thus the water table is

raised, resulting in moist conditions. Rocks may be found

in this community but are not larger than 0.05 m diam.

An affinity exists between community number 3.1

and 3.2 through the common species shared in species

group M. Species group P, present in communities

1.2-3. 2 represents species common to these communi-

ties and include the following: the sedges Schoeno-

plectiis corymbosus, Mariscus keniensis, Juncus oxycar-

piis and Bulbostylis burchellii, the forbs Denekia capen-

sis and Oldenlandia herbacea, the geophytes Ledebouria

cooperi and Gladiolus longicollis and the orchid Saty-

rium longicauda. The presence of species in more than

one community shows environmental affinities which

exist between the communities. These environmental

factors have yet to be ascertained and a combination of

factors may be responsible for the distribution of species.

198

Bothalia 30,2 (2000)

4. Arundinella nepalensis Moist Turf Grassland Com-

munity

This major community is fairly poor in species com-

position, is often dominated by the widespread Anin-

dinella nepalensis (species group T, Table 1). No diag-

nostic species group could be recognized.

It is divided into two subcommunities namely the

Hypericum aethiopicum-Arundinella nepalensis Moist

Turf Grassland Subcommunity and the Imperata cylin-

drica-Arimdinella nepalensis Moist Turf Grassland Sub-

community. Both these subcommunities have the vertic

Arcadia soil form in common, though the origins of the

soil differ.

4. 1 . Hypericum aethiopicum-Arundinella nepalensis Moist

Turf Grassland

Type releve: 311.

Av. no. spp. per releve: 20.

Max. no. spp. per releve: 30.

Min. no. spp. per releve: 10.

Altitude: 900-2 100 m.

Geology: mainly Transvaal diabase.

Soil form: vertic, clay rich loams of the Arcadia soil

form.

Height of vegetation: 0.8 m or higher due mainly to the

presence of the robust grass Arundinella nepalensis.

Species group: Q (Table 1 ).

Diagnostic species: the succulent Crassula sarcocaulis,

the forbs Hypericum aethiopicum and Senecio lati-

folius and the geophytes Ledebouria sp. and Gladio-

lus ecklonis.

Dominant species: Hypericum aethiopicum and Arundi-

nella nepalensis.

This community is found on all aspects and on slopes

of moderate inclination ( 15°-30°). There are rock sheets

present which may lie exposed at the surface or are cov-

ered by a thin (± 0.30 m) layer of soil. Because the soil

is not deep, the roots of the plants grow so closely togeth-

er that a mat is formed. The moisture status gives rise to

a semipermanent soggy layer of soil underlain by an

impenetrable layer of solid or weathered rock. During

the dry season the soil may dry out completely.

4.2. Imperata cylindrica-Arundinella nepalensis Moist

Turf Grassland Subcommunity

Type releve: 215.

Av. no. spp. per releve: 30.

Max. no. spp. per releve: 30.

Min. no. spp. per releve: 30.

Altitude: 900-1 900 m.

Geology: alluvial deposits of Quarternary origin.

Soil form: vertic soils of the Arcadia soil form.

Height of vegetation: 0.6 m.

Species group: R (Table 1 ).

Diagnostic species: the grass Imperata cylindrica which

may grow in nearly pure stands, the sedge Pycreus

macranthus and the forb Senecio latifolius.

Dominant species: the grass Imperata cylindrica.

This community is found on very gentle slopes, nor-

mally in shallow valleys. During wet periods much water

is held in the soil and during dry seasons the soil may be

cracked and dry.

Species Group S present in communities 2. 1-4.2 but

absent entirely in community 2.2, represents species that

are flexible in their habitat requirements, thus inhabiting

a wide variety of environmental conditions within the

parameters of the wetland. These include the grasses

Stiburus alopecuroides, Helictotrichon hirtellum and

Paspalum urvillei, the forbs Monopsis decipiens,

Wahlenbergia virgata, Hypericum lalandii and Cepha-

laria zeylieriana and the sedge Xyris capensis.

DISCUSSION

Wetlands are fragile ecosystems and mismanaging

them can result in a shrinkage of the area covered by the

wetland or a total disappearance of some species in the

wetland (Walmsley 1988; Eckhardt et al. 1993), for

example, the Miscanthus junceus wetlands (2.1 to 2.5)

are intensively grazed by cattle and sheep, as they con-

tain plant species that are highly desireable in the winter

months when greens are scarce for grazing animals.

Burning takes place at intervals of between one and five

years and this, accompanied by grazing of the green

shoots and trampling by livestock, may cause a serious

depletion of the water. With continued grazing and tram-

pling and increased evaporation, the surface area of the

wetland shrinks until only moist grassland is left.

The wattled crane {Grus carunculata), which has its

breeding grounds in this area, is an endangered bird

species (Batchelor et al. 1982). The Verlorenvalei Nature

Reserve has been set aside specifically as a breeding

ground for wattled cranes. According to some local farm-

ers, they breed only in wetlands that have been burned

the previous season. Under-burning can have as serious

an effect on the condition of the wetland and grassland as

can over-burning (Trollope 1989). Thus if wetlands are

left unburned, the cranes will either move to a more suit-

able site or would simply not breed that year due to

unfavourable conditions. Most cranes, however, return to

their territories each year to breed. Not burning the wet-

lands may be a possible explanation for their low num-

bers. According to Heyns (1985), wetlands of the

Verlorenvalei Nature Reserve should be burned every

third year to remove dead plant material.

Grasslands found in the Belfast, Lydenburg and Dull-

stroom Districts are of fire climax vegetation as opposed

to climatic climax vegetation (Tainton 1981). Here it can

be seen that fire, together with grazing, maintains the

grass cover (Burkhart 1975), and prevents the establish-

ment of shrubs and trees. Thus fire started naturally

(lightning) or by man, together with grazing, have creat-

ed pressures with which the grasslands of Africa have co-

evolved (Daubenmire 1974; Grubb 1977; Owen &

Wiegert 1981; Tainton 1981; Mentis & Huntley 1982).

Fire has always been used by pasturalists for the manage-

ment of the vegetation, but abuse of this tool can result in

deterioration in the vegetation (Trollope 1989). Such mis-

use of fire is to be found in the study area where sheep

Bothalia 30,2 (2000)

199

farmers bum the veld out of season to obtain green

growth for grazing. This causes a reduction of canopy

cover, basal cover and a reduction of vigour (Tainton

1981; Trollope 1989) as well as an increased mn-off area

for rain which in turn causes soil erosion (Trollope 1989).

The great variety of wetland plant communities iden-

tified in this study, most of them being relatively poor in

species richness, emphasizes the results of Myburgh et

al. (1995) and Bloem (1986). Therefore a variety of wet-

land communities should be conserved in order to pre-

serve the wetland biodiversity in Mpumalanga.

ACKNOWLEDGEMENTS

We wish to thank the Foundation for Research De-

velopment (now NRF) for funding.

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Miscellaneous notes

POACEAE

CHROMOSOME STUDIES ON AFRICAN PLANTS. 14. PANICOIDEAE

Various papers from our laboratories presented cytoge-

netic data on chromosome numbers of South African rep-

resentatives of the grass subfamily, Panicoideae (Spies

1982; Spies & Du Plessis 1986a. b; Spies & Du Plessis

1987a. b; Spies et al. 1991, 1994). In this paper miscella-

neous chromosome numbers of the subfamily Panicoideae

are presented. The numbers reported present mainly new

localities for the species studied. In this way it will con-

tribute to our understanding of the geographical distribu-

tion of polyploidy in the species studied.

MATERIALS AND METHODS

Eor this study cytogenetic material was collected and

fixed in the field. The material used and their localities are

listed in Table 1 . Voucher specimens are housed in the Geo

Potts Herbarium. Department of Botany and Genetics,

University of the Orange Free State. Bloemfontein

(BLEU), or in the National Herbarium, Pretoria (PRE).

Anthers were squashed in aceto-carmine and meioti-

cally analysed (Spies et al. 1996). Chromosome numbers

are presented as gametic chromosome numbers to con-

form with previous papers on chromosome numbers in

this journal (Spies & Du Plessis 1986b).

RESULTS AND DISCUSSION

Fifty-seven specimens, representing 25 species (as

well as four infraspecific taxa) and 15 genera, were stud-

ied (Table 1). The species will be discussed alphabetical-

ly under the appropriate tribe.

Tribe Andropogoneae

The genus Bothriochloa consists of 35 species

(Clayton & Renvoize 1986) worldwide, with three in-

digenous species in southern Africa (Gibbs Russell et al.

1990). The somatic chromosome number of 2n = 20

(Table 1 ) is the lowest ploidy level yet observed for this

species in South Africa. The formation of bivalents only

in this specimen with 2n = 4x = 20 indicate that this

species is either an alloploid, or it has a secondary basic

chromosome number of ten, as is observed in most rep-

resentatives of the tribe Andropogoneae (De Wet 1954,

1958; De Wet & Anderson 1956; Celarier & Harlan

1957; Olorode 1975; Dujardin 1978; Spies & Du Plessis

1986a, b, 1987a. b; Sinha et al. 1990; Spies et al. 1994).

The genus Cymbopogon consists of 40 species

(Clayton & Renvoize 1986) worldwide, with six indige-

nous species in southern Africa (Gibbs Russell et al.

1990). The C. plurinodis specimen is diploid (2n = 2x =

20) with normal meiotic chromosome behaviour.

Diploid, tetraploid and octoploid levels in this species

have been reported (De Wet 1954; Spies & Du Plessis

1987a; Du Plessis & Spies 1988; Spies et al. 1994).

The genus Diclianthium consists of 20 species

(Clayton & Renvoize 1986) worldwide, with one indige-

nous and one naturalized species in southern Africa

(Gibbs Russell et al. 1990). The specimen of D. arista-

turn is tetraploid with n = 2x = 20 (Figure lA, B). The

formation of bivalents only indicates that this species is

an alloploid. Previous reports indicated diploid,

tetraploid and hexaploid levels (De Wet 1954; Spies &

Du Plessis 1986b; Sinha et al. 1990).

The genus Elionurus comprises 15 species (Clayton

& Renvoize 1986) worldwide. Two of these species are

indigenous to southern Africa (Gibbs Russell et al.

1990), with only E. muticus representing the genus in

this study. Both specimens studied have a gametic chro-

mosome number of n = 3x = 15 (Figure IC-H). Up to

two putative B-chromosomes are present, resulting in

laggards or late segregating chromatids during anaphase

(Figure IF, G). Although several laggards during ana-

phase I were observed, no micronuclei during telophase

I (Figure 2H) were observed. This is the highest ploidy

level yet described for this species in South Africa. The

formation of bivalents in these specimens with 30 somat-

ic chromosomes, clearly indicate that five is the basic

chromosome number for this species.

The genus Eulalia consists of 30 species (Clayton &

Renvoize 1986) worldwide. Two of these species are

indigenous to southern Africa (Gibbs Russell et al.

1990). The E. villosa specimen has a gametic chromo-

some number of n = x = 5. This is, to the best of our

knowledge, the first chromosome number report on this

species. This chromosome number confirms the basic

number of five for the tribe Andropogoneae.

The genus Miscanthus consists of 20 species (Clayton

& Renvoize 1986) worldwide, with two indigenous

species in southern Africa (Gibbs Russell et al. 1990).

The specimen of M. capensis has a chromosome number

of n = 15 (Figure II). There were no meiotic abnormali-

ties observed in this specimen. This appears to be the

first chromosome number report for this species.

The genus Trachypogon consists of three species

(Clayton & Renvoize 1986) worldwide, with one indige-

nous species in southern Africa (Gibbs Russell et al.

1990). The specimen of T. spicatus has a chromosome

number of n = 10 (Figure IJ, K) and exhibited bivalent

formation during meiosis, indicating that the species is

either an alloploid or has a secondary basic chromosome

number of ten.

202

Bothalia 30,2 (2000)

TABLE 1 . — Gametic chromosome numbers (n) of representatives of subfamilies Panicoideae in southern Africa with their voucher specimen numbers and

specific localities. Species are listed alphabetically and the localities are presented according to the system described by Edwards & Leistner (1971)

Bothalia 30,2 (2000)

203

FIGURE 1. — Meiotic chromosomes in some specimens of the genera Dicanthium, Eliomirus, Miscanthus and Trachypogon. A, B, Dicanthiiim

aristalwn, Spies 2007, n = 20. anaphase I with a 20-20 chromosome segregation. C-H, Eliomirus muticus, both specimens with n = 3x =

15. C-E, G, H, Spies 2117a-. C. D. metaphase I with 15n; E, anaphase I with a 15-15 chromosome segregation; G, anaphase I with one chro-

mosome laggard; H. telophase I. E, Spies 2115a. anaphase I with laggards. I, Miscanthus capensis. Spies 5242, n = 15, anaphase I with a

15-15 chromosome segregation. J. K. Trachypogon spicatus, Spies 2107, n = 10, metaphase I with lOn. Scale bar: 40 pm.

All the specimens studied have either a basic chromo-

some number of five or a secondary basic number of ten,

conforming with previous reports on these genera (De

Wet 1954; De Wet & Anderson 1956; Celarier & Harlan

1957; De Wet 1958; Olorode 1975; Dujardin 1978; Spies

& Du Plessis 1986a, b, 1987a, b; Sinha et al. 1990; Spies

et al. 1994). The fact that the chromosome numbers of

two of the species studied (28.6%) are published for the

first time and that two new ploidy levels (28.6%) are

described for the seven species studied, indicate that

more cytogenetic studies of the Andropogoneae are

needed (57.1% of the studied specimens presented new

data). More specimens should be studied to determine

the ploidy levels present and the geographical distribu-

tion of these ploidy levels.

Tribe Arundinelleae

The genus Loudetia consists of 26 species (Clayton &

Renvoize 1986) worldwide, six of which are indigenous to

southern Africa (Gibbs Russell et al. 1990). L. simplex is

tetraploid with n = 2x = 18 (Figure 2A-C). Laggards and

micronuclei were observed (Figure 2B, C). Chromosome

numbers based on 6 (De Wet 1958) and 10 were reported

previously (Moffett & Hurcombe 1949; Dujardin &

Beyne 1975; Dujardin 1978). The basic chromosome

numbers observed in this study (x = 9) differ from those

previously reported. However, the results of this study can

also be interpreted as x = 6 with a high ploidy level (hexa-

ploid). More studies are needed to determine the real basic

chromosome number of this species and whether those

studied included an aneuploid specimen.

The genus Tristachya consists of 22 species (Clayton

& Renvoize 1986) worldwide. Six of these species are

indigenous to southern Africa (Gibbs Russell et al.

1990). The specimens of T. leucothrix (Figure 2D-I) are

tetraploid with n = 2x = 12. Meiosis was normal and only

bivalents were observed, indicating that the species is an

allotetraploid. This is a new ploidy level for this species.

Previously an octoploid specimen was observed (Spies

& Du Plessis 1987a).

The overwhelming presence of x = 6 observed in

Tristachya suggests that the basic chromosome number

204

Bothalia 30,2 (2000)

FIGURE 2. — Meiotic chromosomes in one specimen each of the genera Loudetia and Tristachya. A-C, L. simplex. Spies 1981h, n = 18: A, diaki-

nesis with 18n; B, anaphase I with two laggards; C, telophase 1 with micronuclei. D, E, T. leucothrix, H. dii Plessis 152, n = 12; D, diakine-

sis with 12ii; E, anaphase I with a 12-12 chromosome segregation. F, G, T. leucothrix. Spies 5148, n = 12: F, diakinesis; G, metaphase I with

12n. H, T. leucothrix, Spies 5061, n = 12, metaphase I with 12n. I, T. leucothrix. Spies 4711, n = 12, metaphase I with 12n. Scale bar: 40 pm.

of the Arundinelleae should most probably be six, which

conforms to some previous reports on these genera

(Moffett & Hurcombe 1949; De Wet 1958; Spies & Du

Plessis 1987a). In this tribe 100% of the species studied

proved to be either a new chromosome number or a new

ploidy level for the species.

Tribe Paniceae

The genus Alloteropsis consists of five to eight species

worldwide and three species are indigenous to southern

Africa (Clayton & Renvoize 1986; Gibbs Russell et al.

1990). Alloteropsis semialata is diploid (n = x = 9) with

normal meiosis (Figure 3 A, B) and this confirms previous

reports (Moffett & Hurcombe 1949; De Wet & Anderson

1956; De Wet 1958). Ahexaploid level has also been report-

ed (Moffett & Hurcombe 1949; De Wet & Anderson 1956).

The genus Anthephora consists of 12 species (Clayton

& Renvoize 1986) worldwide, with four indigenous to

southern Africa (Gibbs Russell et al. 1990). The A.

pubescens specimens are n = 2x = 18 (Figure 3C) which

conforms to previous reports on this genus and confirms

the tetraploid level in the specimens studied (Spies 1982;

Spies & du Plessis 1986a).

The genus Axonopus has 110 species (Clayton &

Renvoize 1986) worldwide, with one naturalized species

in southern Africa (Gibbs Russell et al. 1990). The

Axonopus ajfinis specimens are n = 2x = 20 (Figure 3D)

which conforms to previous reports (Burton 1942;

FIGURE 3. — Meiotic chromosomes

in some specimens of Allote-

ropsis. Anthephora, Axonopus

and Brachiaria. A, Alloterop-

sis semialata. Spies 4715, n =

9, metaphase I with 9n. B, A.

semialata. Spies 5068, n = 9,

diakinesis with 9n. C. Anthe-

phora pubescens. Spies 5500,

n = 18, diakinesis with 18n.

D, Axonopus affinis. Spies

2561, n = 20, metaphase with

20||. E, Brachiaria serrata.

Spies 5075, n = 9, metaphase

I with 9ii. F, B. xantholeuca.

Spies 4812, n = 9, diakinesis

with 9ii. Scale bar: A, 26.6

pm; B, D-F, 40pm; C, 32

pm.

Bothalia 30,2 (2000)

205

FIGURE 4. — Meiotic chro-

mosomes. A, Paspa-

lum un’illei. Spies

1980. n = 20, me-

taphase I with 20ii. B,

P. un’illei, Saayman

67, n = 20, metaphase

I with 20ii. C, Setaria

pallide-fusca. Spies

2772, n = 18, me-

taphase I with 18ii.

Scale bar: 40 pm.

Brown 1948; Hickenbick 1975; Mehra 1982). However,

chromosome numbers of 2n = 54 and n = 25 have been

previously reported (Brown 1946; Mehra 1982). Diploid

levels for this species have also been observed (Burton

1942; Brown 1948; Hickenbick 1975; Mehra 1982).

The genus Brachiaria consists of 100 species world-

wide (Clayton & Renvoize 1986), with 19 indigenous

and one naturalised species in southern Africa (Gibbs

Russell et al. 1990). Five of these species are included in

this study (Table 1 ). All the specimens of B. bovonei, B.

brizantha, B. nigropedata, B. serrata (Figure 3E) and B.

xantholeuca (Figure 3F) are n = x = 9, which conforms

with previous results (Moffet & Hurcombe 1949; De Wet

& Anderson 1956; Nath & Swaminathan 1957; De Wet

1958, 1960; Nassar 1977; Dujardin 1978, 1979; Sharma

& Kaur 1980; Basappa & Muniyamma 1981; Mehra

1982; Spies & du Plessis 1986a, b, 1987b; Sinha et al.

1990; Spies et al. 1991), the exception being B. .xant-

holeuca, for which this is the first chromosome number

report. To the best of our knowledge, this is the first

report on the presence of B-chromosomes in Brachiaria.

Chromosome numbers of n = 18, 27 or 45 have also been

previously observed in B. nigropedata and B. serrata

(Dujardin 1978, 1979; Sharma & Kaur 1980; Mehra

1982; Spies & du Plessis 1986b, 1987b; Sinha et al.

1990; Spies et al. 1991).

The genus Paspalum consists of 330 species (Clayton

& Renvoize 1986) worldwide, with three indigenous

species and three naturalised species in southern Africa

(Gibbs Russell etal. 1990). All the specimens of P. dilata-

tum, P. scrobiculatiim and P. urxhllei, except Spies 2362,

are tetraploids, with n = 2x = 20 (Figure 4A, B). Only

bivalents were observed, indicating that these species are

alloploids. Meiotic abnormalities such as laggards and

micronuclei were observed in some specimens of P.

dilatatiim. Laggards were also observed in a few cells of

one specimen of P. itn'illei. Chromosome numbers of n =

21, 25 or 27 and 2n = 50 or 63 have been previously

reported in P. dilatatum and P. scrobiculatiim (Tateoka

1954; Bashaw & Forbes 1958; Singh & Godward 1960;

Mehra & Sharma 1975; Mehra & Chaudhary 1976, 1981;

Dandin & Chennaveeraiah 1977, 1983; Rao Sindhe 1977;

Sharma a/. 1978; Mehra 1982; Hickenbick ct a/. 1987;

Spies & Du Plessis 1987a).

The last genus included in this study, Setaria, consists

of 110 species (Clayton & Renvoize 1986), with 19

indigenous and two naturalised species in southern

Africa (Gibbs Russell et al. 1990). Five of these species

are included in this study (Table 1 ). All the studied spec-

imens of S. incrassata, S. nigrirostris, S. pallide-fusca

(Figure 4C) and S. plicatilis are diploid with n = x = 9

which conforms with previous reports (Krishnaswamy &

Ayyangar 1935; De Litardiere

1948; Moffett & Hurcombe 1949; Krishnaswamy 1951;

De Wet 1954; Singh & Godward 1960; Olorode 1975;

Gupta & Singh 1977; Sharma a/. 1978; Dujardin 1979;

Mehra 1982; Bir & Sahni 1984; Spies & du Plessis

1986a, b). The S. sphacelata specimens studied have

diploid, tetraploid and hexaploid levels, which conforms

to previous reports (Spies & du Plessis 1986b, 1987a).

Meiotic abnormalities, particularly laggards during

anaphase I, were observed in one S. sphacelata var.

splendida specimen. All the other specimens of S.

sphacelata var. sericea, var. sphacelata, var. splendida

and var. torta exhibited normal bivalent formation during

meiosis I. No chromosome numbers have been previous-

ly reported for S. sphacelata var. sphacelata and var.

splendida.

This study provided new reports for three species and

two interspecific taxa (10%) of the tribe Paniceae and

helped to confirm the basic chromosome number of

Axonopis as 10. It is also the first report on the presence

of B-chromosomes in Brachiaria. This indicates that

chromosome number reports on grasses should be done

more frequently and such studies will contribute to the

knowledge of these grasses. At this stage the knowledge

is insufficient to determine the correlation between

ploidy level and geographical distribution.

This study confirms a basic chromosome number of 5

and a secondary basic chromosome number of 10 for the

tribe Andropogoneae, a basic chromosome number of 6

for the tribe Arundinelleae and 10 for the genera

Axonopus and Paspalum of the tribe Paniceae.

ACKNOWLEDGEMENTS

The University of the Orange Free State and the

Foundation for Research and Development are thanked

for financial assistance during this study.

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A. STRYDOM*, J.J. SPIES* and S.M.C. VAN WYK*

* Department of Botany and Genetics (106), University of the Orange

Free State, P.O. Box 339, 9300 Bloemfontein.

MS. received: 1999-06-30.

ALOACEAE

THE CONSERVATION STATUS OF ALOE IN SOUTH AFRICA: AN UPDATED SYNOPSIS

INTRODUCTION

The Aloaceae is a medium-sized family of about 510

species of succulent-leaved, petaloid monocots, geo-

graphically restricted to the Old World (Smith & Van

Wyk 1998a). The area of highest species diversity is

southern Africa, particularly South Africa (Van Wyk &

Smith 1996a), with other centres of diversity occurring

in the East- Afro Arc of tropical southern Africa (West

1974), eastern Africa (Carter 1994; Lavranos 1995),

Saudi Arabia (Collenette 1985), Yemen (Wood 1982),

and Madagascar (Reynolds 1966).

The genus Aloe L., which is by far the largest genus

in the Aloaceae, is an important component of the South

African flora from taxonomic (Reynolds 1950), phyloge-

netic (Smith & Van Wyk 1991), ethnomedicinal (Van

Wyk & Smith 1996b), chemical/chemotaxonomic

Bothalia 30,2 (2000)

207

(Viljoen & Van Wyk 1996), ecotourism (Smith & Van

Wyk 1998b) and horticultural (Smith & Van Wyk 1996,

1998c) perspectives. The wide-ranging interest in the

genus amongst many scientists and collectors has affect-

ed the survival of wild populations.

Until recently, contributions on the threatened species

of South African Aloe were restricted to single-species

narratives, such as A. bowiea Schult. & Schult.f. (Smith

1989, 1991; Smith & Van Wyk 1990), A. micracantha

Haw. (Smith 1993a) and A. vossii Reynolds (Willis &

Willis 1995). Hilton-Taylor & Smith (1994) were the first

to provide a comprehensive synopsis of the conservation

status of southern African Aloe and its generic relatives.

For example, they cite that, apart from illegal collecting

activities, other factors that lead to the decline of some

populations of Aloe species include overgrazing, mining

and industrial activities, commercial forestry, insect pre-

dation, urbanization, agricultural development and frag-

mentation effects on narrow distribution ranges. Forty-

nine Aloe species occurring in the Flora of southern Africa

(FSA) region (Botswana, Lesotho, Namibia, South Africa

and Swaziland) were assigned Red List status (Hilton-

Taylor & Smith 1994). Of these species, 44 occur in South

Africa. All species of Aloe appear on CITES (Convention

on International Trade in Endangered Species of Wild

Fauna and Flora) Appendices. However, of the southern

African species only A. albida (Stapf) Reynolds, A. pil-

lansii L. Guthrie, A. polyphylla Schonland ex Pillans, A.

thomcroftii Pole Evans and A. vossii Reynolds are includ-

ed in Appendix I (CITES 1998).

Plant Red Data Lists document extinct, threatened

and potentially threatened plant species (including sub-

species, varieties, and taxonomically undescribed groups

recognized as being distinct) that have been assigned cat-

egories of urgency for conservation according to the

World Conservation Union’s Species Survival Commis-

sion (lUCN-SSC) (lUCN 1994). Red Data Lists enable

decision-makers, research specialists and field managers

to set more clearly defined goals and priorities. Aloe, a

prominent and conspicuous component of the southern

African flora, is a flagship genus for conservation efforts

and its protection will therefore set a standard for the

protection of lesser known plant species.

Although species of Aloe occur throughout southern

Africa, the subtropical eastern seaboard of South Africa

contains a significant number of taxa, many of which are

threatened for a variety of reasons. The conservation status

of most of the species of Aloe occurring in KwaZulu-Natal

were recently assessed by Scott-Shaw (1999). Those

species that are under some sort of threat in KwaZulu-

Natal, but which are plentiful in other provinces are not

accorded a national conservation status here. These include

the traditionally utilized A. aristata Haw., which also

occurs in the Tree State, and Eastern and Western Cape.

Hilton-Taylor & Smith (1994) applied the ‘old’ cate-

gories in their account of the conservation status of the

Aloe species of the FSA region. Shortly thereafter, the

lUCN-SSC adopted a new set of Red List guidelines

which are intended to be more objective, widely applic-

able to terrestrial and aquatic biota, and useful to Red

List compilers and field managers alike (lUCN 1994).

Milestones in the improvement of these guidelines

include their applicability at different geographical

scales — national, regional and subregional (Gardenfors

et al. 1999), and a review of the lUCN (1994) Red List

categories and their defining criteria (lUCN-SSC

Criteria Review Working Group 1999). The review arose

out of a need to assess harvested species, long-lived ani-

mal species (such as elephants and marine turtles) and

the status of some small and narrowly distributed endem-

ic plants and vertebrates. Revised guidelines may be offi-

cially adopted by the lUCN-SSC in October 2000

(Gardenfors a/. 1999).

This paper updates and summarizes the current conser-

vation status of species of Aloe in South Africa. It provides

conservation perspectives on the genus over the past five

years, with emphasis on Red Data Lists (Hilton-Taylor &

Smith 1994; Hilton-Taylor 1996), using the ‘new’ lUCN

Red List categories (lUCN 1994). The legal, protection

and conservation data used by Hilton-Taylor & Smith

( 1994) have remained more or less unchanged and are still

applicable. This information is therefore not repeated here.

The main objective of this paper is to contribute towards

determining which Aloe species are in need of conserva-

tion, and to make broad recommendations on future stud-

ies and in situ monitoring.

MATERIAL AND METHODS

The taxonomic treatment of Aloe follows Van Wyk &

Smith (1996a). Red List categories are assigned in accor-

dance with guidelines and concepts set by the lUCN-

SSC (lUCN 1994), and are based on herbarium speci-

mens housed at PRE, published information, field obser-

vations and known distribution ranges of species of Aloe.

In a few instances the category assigned represents a best

estimate based on available information.

Quantitative criteria are used to place a species in a par-

ticular Red List category (for details see lUCN 1994). The

term ‘threatened’ is used to describe species which are

Critically Endangered, Endangered or Vulnerable. The cri-

teria rely on data derived from estimates, projections, infer-

ences or quantitative analyses. These data include percent-

age of decline in population or distribution, the number of

mature individuals and suspected/anticipated future popu-

lation declines.

Criterion A was used where we could estimate per-

centage decline over the past ten years. Ten years was

preferred rather than three generations, because it is diffi-

cult to estimate generation time reliably without compre-

hensive field work or autecological studies. At least a

20% decline had occurred in this period of time (placing

the taxon in the VU category). In some cases it may well

be higher, but more reliable data would be necessary to

estimate a 50% or higher decline. Criterion B was used in

cases where the taxon had a continuing decline along with

a restricted distribution range and either a small number

of locations or severe fragmentation. Criterion C could

not be used, as it was too difficult to estimate population

sizes. All taxa which were not experiencing a decline, but

had a restricted distribution range (<100 km^) were clas-

sified as Vulnerable according to Criterion D2.

208

Bothalia 30,2 (2000)

The Lower Risk categories are assigned to taxa that

do not satisfy the criteria for being threatened, and such

taxa are not listed in Table 2, but reported separately. The

Data Deficient category is assigned to taxa for which

there is not enough information to do an assessment.

RESULTS AND DISCUSSION

During the present study the global conservation sta-

tus of 68 Aloe taxa occurring in South Africa was

assessed. Our results showed that the data for 38 of these

Aloe species meet the criteria for one of the categories of

threat, namely Critically Endangered, Endangered or

Vulnerable (Figure 5). In order to draw attention to

species in the Data Deficient category, they are listed

with the threatened species in Table 2. Although the

number of species for which no assessment could be

made are proportionately low (Figure 5), they do occur

in four of the eight provinces of South Africa (Figure 6)

and their unknown status should concern conservation

agencies, field botanists and researchers. At present, no

threatened Aloe species are known from the Free State.

A. peglerae, a striking species, occurs in the North-West

and in Gauteng. Continual census work and monitoring

of this species need to be implemented. KwaZulu-Natal

and Northern Cape each have 1 1 species of threatened

aloes (Figure 6). Further updates of, and refinements to

TABLE 2. — lUCN Red List status of Aloe species in South Africa. Author citations are contained in Smith et al. (1997)

* EC, Eastern Cape; G. Gauteng; KN, KwaZulu-Natal; M, Mpumalanga; NC, Northern Cape; NP, Northern Province; NW. North-West; WC.

Western Cape,

Bothalia 30,2 (2000)

209

B Critically Endangered (CR)

B Endangered (E)

^ Vulnerable (VU)

I I Data Deficient (DD)

26 (60%)

FIGURE 5. — lUCN status of Aloe species listed in Table 2. Numbers

and percentages of species placed in the relevant lUCN cate-

gories are in relation to the total number of 43 listed.

the data presented here, will be recorded in a database

held at the National Herbarium (PRE).

The following 19 taxa were listed by Hilton-Taylor &

Smith { 1994) as being not threatened, and here similarly,

they are not assigned a threatened status, but are catego-

rized as Lower Risk (least concern) i.e. LR(lc). These

are: Aloe ajfinis, A. angelica, A. chortolirioides var.

chortolirioides, A. chortolirioides var. woolliana, A.

dewetii, A. dominella, A. dyeri, A. greatheadii var.

davyana, A. greenii, A. hlangapies, A. integra, A. kraus-

sii, A. linearifolia, A. minima, A. riipestris, A. striata

subsp. karasbergensis, A. sujfidta, A. thorncroftii and A.

vanbalenii.

The status of three species previously listed as threat-

ened by Hilton-Taylor ( 1996) has been changed. A. vry-

heidensis which was listed as Rare, is now assigned a

status of Lower Risk (near threatened) i.e. LR(nt)

because of its extremely wide distribution range (see Van

Wyk & Smith 1996a: 71) and lack of known threats at

present. A. arenicola was previously listed as Vulnerable

and is now listed as LR(lc) because of its wide distribu-

tion range along the west coast of the Western Cape and

Northern Cape. The third is A.falcata which is now clas-

sified as LR(nt). Although previously classified as

Vulnerable due to illegal collecting and agricultural

activities, this widespread species is probably not under

threat at present, as the Aloe craze of the 70’s has lost

momentum in recent years.

Aloe kniphofioides, A. mudenensis and A. thraski

were assessed by Scott-Show (1999) but not by Hilton-

Taylor (1996). These species are categorized here as

LR(lc). While the status of the former did not change, the

latter two were categorized by Scott-Shaw as LR(nt), but

our data do not support this interpretation.

Two species of Aloe, A. polyphylla from Lesotho and

A. bowiea, are the most threatened in southern Africa.

Of these two, A. bowiea with its narrower geographical

distribution range is more threatened. With less than

2 000 individuals thought to be left in nature, one of

three known populations extinct and another threatened

by urban and industrial expansion, this species may

soon join the ranks of those taxa that are extinct in the

wild. Although Scott-Shaw (1999) refers to the uncon-

firmed occurrence of A. polyphylla in the KwaZulu-

Natal Drakensberg, this has yet to be substantiated by a

herbarium or other record. The species is therefore still

only known from and endemic to Lesotho. Prom an eth-

nomedical perspective, Marshall (1998) rates A. poly-

phylla as one of the three most threatened taxa out of

102 species shortlisted as being of conservation con-

cern in the 17 East and southern African countries sur-

veyed.

Through the efforts of Reynolds (1950, 1966) and

Jeppe (1969), Aloe species became popular in domestic

and amenity horticulture in South Africa, contributing to

the ‘aloe craze’ of the 1970s. In his benchmark publica-

tion on South African aloes, Reynolds (1950) gave

detailed locality information for the known species,

thereby facilitating the often illegal collecting of speci-

mens of a number of species and the wanton destruction

of some populations. An international and often illegal

trade in South African aloes is ongoing, and has recently

been documented by Newton & Chan (1998). Some Aloe

species are difficult to cultivate away from their natural

habitats and large numbers of individuals undoubtedly

ended up on compost heaps. Along with the introduction

of species from wild populations, a number of natural

enemies of Aloe, especially snout beetles, white and

brown scale insects and spider mites, entered domestic

gardens (De Villiers & Schoeman 1988; Myburgh 1990)

and even some previously uninfected natural areas

(Williamson 1998). These pests eventually led to the

Data Deficient

^ Vulnerable

I I Endangered

I Critically Endangered

FIGURE 6. — Distribution of 43 Aloe

species, listed in Table 2, in

the nine provinces of South

Africa.

210

Bothalia 30,2 (2000)

demise of many cultivated plants and, concurrently, a

decline in the popularity of aloes as garden plants.

However, with the increasing realization that water-

wise gardening techniques should become part of southern

African gardening practices (Fourie 1984; Honig et al.

1998, 1999), species of Aloe are again becoming popular

in cultivation. In the past 30 years a number of species and

strains of some species have fortunately proven them-

selves to be resilient against insect and mite infestations

and superior strains are commercially available. In partic-

ular, the non-threatened A. arborescens Mill, has shown

itself to be exceptionally suitable as a striking landscape

plant. Many gardeners currently realize that it is better to

have a smaller selection of well-grown subjects than a

comprehensive collection of sickly plants.

Plant use is playing an increasingly important role in

determining the conservation status of Aloe species. In

some instances, such as A. pratensis Baker and A. aris-

tata, specimens are being transported over long distances

to established ethnomedicinal marketing sites where

whole plants are being sold for as little as US$0.50. The

reasons for this trend may be various; it may indicate a

specific preference for the species, or it could mean that

the plants are becoming rarer closer to the market, or

even that gardeners purchasing through the muthi mar-

kets are creating a demand for taxa that are otherwise dif-

ficult or relatively costly to obtain from nurseries and

gardening shops. However, not all Aloe species traded in

the markets are threatened. Numerous non-threatened

species such as the widely grown barrier plants A. striat-

ula Haw. and A. arborescens are also available in the

market, as are the dry leaves of A./croxMill. and A. mar-

lothii Berger, which are sold as snuff components.

Ironically, the traditional healer community may, whilst

destructively harvesting some Aloe taxa, be simultane-

ously promoting the ex situ conservation of other select-

ed A/o^ species. Crouch & Hutchings (1999) inventoried

the plants grown in five healer gardens in KwaZulu-

Natal and found (of 198 taxa catalogued) the Aloaceae to

be the most cultivated family, with 17 taxa represented,

including 1 1 from the genus Aloe. Both the extra-provin-

cial A. brevifolia Mill, and A. striatula Haw. var. caesia

Reynolds were noted in cultivation.

The grass-leaved aloes of South Africa present an

interesting challenge for conservationists in general

(Craib 1996). These species have an effective defence

mechanism (camouflage). In their natural grassland habi-

tats, casual succulent plant collectors are likely to mistake

their leaves for those of pooid species. However, as a

result of difficulties experienced in locating grass-leaved

aloes in the wild, especially when they are not in flower,

entire populations could easily be exterminated through

sheer ignorance of their presence. This is particularly

applicable in the Grassland Biome and mist belt of the

eastern Drakensberg Escarpment of South Africa, which

are subject to extensive agricultural activity and commer-

cial forestry. As is the case with species of the related

alooid genera, Haworthia (Craib 1990) and Chortolirion

(Hargreaves 1989; Smith 1993b), it is unlikely that grass-

leaved aloes will recolonize a habitat once it has been dis-

turbed. Where possible, localities of graminoid Aloe

.species should be monitored for decline or expansion.

Very few species of Aloe take kindly to invasive alien

plant infestations, A. greatheadii Schonland var. davyana

(Schonland) Glen & D.S. Hardy being a notable excep-

tion. This species seems to thrive in the shade and profuse

leaf litter of Australian Eucalyptus trees. However, in

Eastern Cape the habitats of A. africana Mill., A. bowiea,

A. ferox, A. pluridens Haw. and A. speciosa Baker are

increasingly threatened by jointed cactus, Opuntia auran-

tiaca Lindl.; prickly pear, O. ficus-indica (L.) Mill.;

‘rooikrans,’ Acacia cyclops A.Cunn. ex G.Don; and Port

Jackson willow. Acacia saligna (Labill.) H.L.Wendl.

With the exception of A. bowiea, these species are fortu-

nately exceedingly common and not threatened by the

alien invaders.

CONCLUSION

Now, more than ever, it is important to inform con-

servation authorities of those taxa that are threatened and

in dire need of protection. A commitment to Red Data

Lists will be necessary to achieve a solution for the con-

servation of Aloe species. Conservation authorities

should be at the forefront in this endeavour. Indeed, with-

out appropriate conservation measures in place to ensure

the long-term viability of natural populations (Hilton-

Taylor 1997), future survival prospects look bleak for a

number of Aloe species. It is hoped that the listing of the

conservation status of Aloe species provided here will

stimulate and support in situ conservation efforts and

species recovery programmes.

ACKNOWLEDGEMENTS

We thank Messrs. C.K. Willis and E.J. van Jaarsveld

for useful discussions on the conservation status of Aloe

species in South Africa.

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23-26.

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LOP. C. MACE, G., MOLLIR, S. & POSS, S. 1999. Draft

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HARGREAVES, B.J. 1989. Succulents in the snow. Succulents of

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22-25; 81-86.

HILTON-TAYLOR, C. 1996. Red Data List of southern African plants.

Strelitzia 4. National Botanical Institute, Pretoria.

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G.F. SMITH*. E.M.A. STEYN*, J.E. VICTOR*.

N.R. CROUCH**, J. GOLDING*t and C. HILTON-TAYLOR^

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

** Ethnobotany Unit, National Botanical Institute, P.O. Box 52099,

4007 Berea Road, Durban.

*t SABONET, National Botanical Institute, Private Bag XlOl, 0001

Pretoria.

t lUCN/SSC UK Office, 219c Huntingdon Road, Cambridge CB3

ODL, United Kingdom.

MS. received: 2000-06-20.

APOCYNACEAE

CHROMOSOME STUDIES ON AFRICAN PLANTS. 15. PERIPLOCOIDEAE

The subfamily Periplocoideae currently forms part of

the family Apocynaceae (Venter & Verhoeven 1997).

This subfamily was previously classified as a section

(Brown 1810) or subfamily (Schumann 1895) of the As-

clepiadaceae, or as a separate family, the Periplocaceae

(Schlechter 1914). The Asclepiadaceae has been studied

extensively by especially Albers (Albers 1979, 1983;

Albers & Delfs 1983) but as far as can be ascertained,

almost no cytogenetic data have been published on the

Periplocoideae. The aim of this paper is to contribute to

the cytogenetic knowledge of the Apocynaceae in gener-

al, and the Periplocoideae in particular.

MATERIALS AND METHODS

Cytogenetic material was collected and fixed in the

field (Spies & Du Plessis 1986). The material used and

their localities are listed in Table 3. Voucher specimens

are housed in the Geo Potts Herbarium, Department of

Botany and Genetics, University of the Orange Free

State, Bloemfontein (BLEU).

Anthers were squashed in aceto-carmine and meioti-

cally analysed (Spies etal. 1996). Chromosome numbers

are presented as gametic chromosome numbers to con-

form to previous papers on chromosome numbers in this

journal (Spies & Du Plessis 1986).

RESULTS AND DISCUSSION

Twenty specimens, representing 10 species and four

genera, were studied (Table 3). All specimens proved to

be diploid (2n = 2x = 22) with a basic chromosome num-

ber X = 1 1 .

212

Bothalia 30,2 (2000)

TABLE 3.-Gametic chromosome numbers of specimens of Periplocoideae, with voucher specimen numbers and specific localities

Taxon Voucher n Collecting locality

Tribe Periploceae Bartl.

Tacazzea apiculata Oliv. Venter 9248 1 1

Venter 9250 1 1

Venter 9251, 9252, 9253, 1 1

9322

KWAZULU-NATAL.— 2732 (Ubombo): Makane’s Drift, (-AB).

KWAZULU-NATAL.— 2732 (Ubombo): Sordwana, (-DA).

KWAZULU-NATAL. — 2832 (Mtubatuba): Richards Bay next to

Mzingazi Lake, (-CC).

FREE STATE. — 2826 (Brandfort): Soetdoring Nature Reserve, (-CC).

FREE STATE. — 2926 (Bloemfontein): Rustfontein Dam, (-BC).

KWAZULU-NATAL. — 2732 (Ubombo): Lebombo Mountains near

Josini, (-AA).

FREE STATE. — 2827 (Senekal): Evening Star, Clocolan, (-CD).

FREE STATE. — 2926 (Bloemfontein): Brandkop Racing Track, (-AA).

MPUMALANGA. — 2430 (Pilgrim’s Rest): Blyderivier Canyon, (-DB).

GAUTENG. — 2527 (Rustenburg): Rustenburgkloof, (-CA).

GAUTENG. — 2528 (Pretoria): Wonderboom Reserve, (-CA).

KWAZULU-NATAL.— 2732 (Ubombo): Makane’s Drift, (-AB).

NAMIBIA. — 2615 (Luderitz): Luderitz, (-CA).

NAMIBIA. — 2013 (Unjab Mouth): 22 km south of Torra Bay, (-AA).

NAMIBIA. — 2817 (Viool.sdrif): 133 km west of Noordoewer village

along the Jan Haak road, (-AA).

NORTHERN CAPE.— 2816 (Vioolsdrif): Richtersveld, (-BD).

Tribe Periploceae

Tacazzea apiculata is the only species being studied

cytogenetically (Table 3).

Tribe Gymnanthereae

Chromosome numbers are reported for the first time

in Raphionacme dyeri, R. galpinii, R. Inrsuta, R. procum-

hens and R. zeyheri. This number of 2n = 2x = 22 is con-

firmed for R. flanaganii Schltr., which was previously

studied by F. Albers (pers. comm.).

Tribe Cryptolepideae

This is the first report for any African species in this

tribe. The chromosome number of 2n = 2x = 22 found in

Cryptolepis cryptolepioides, C. ohtiisa, Ectadium lati-

folium, E. rotundifolium and E. virgatiim (Figure 7), cor-

responds with those chromosome numbers previously

noted for certain Asian species of this tribe, i.e. Cryp-

tolepis buchananii Roem. & Schult. (Sharma 1970;

Navaneetham 1982), C. grandiflora Wight (Navanee-

tham 1982; Navaneetham & Sampathkumar 1984) and

C. sinensis (Lour.) Merr. [= C. elegans Wall.] (Navanee-

tham 1981).

ACKNOWLEDGEMENTS

The University of the Orange Free State and the

National Research Foundation are thanked for financial

assistance during this study.

FIGURE 7. — Meiotic chromosomes

(2n = 2x = 22) in representa-

tives of the Apocynaceae. A,

Tacazzea apiculata, Venter

9252, anaphase I. B, C, T.

apiculata. Venter 9322: B,

anaphase I; C, anaphase II. D,

Raphionacme Inrsuta, Venter

9309. early anaphase I; E,

C ryptolepis cryptolepiodes.

Venter 9181, metaphase 1: F,

Ectadium rotundifolium. Ven-

ter 9313, metaphase I; G, E.

virgatum. Venter 9237, diaki-

nesis. Scale bar: A-F, 6.4 pm;

G, 8.3 pm.

Bothalia 30,2 (2000)

213

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J.J. SPIES*. H.J.T. VENTER* and S.M.C. VAN WYK*

* Department of Botany and Genetics, University of the Orange Free

State, P.O. Box 339, 9300 Bloemfontein,

MS. received: 2000-06-30.

PICKING UP THE PIECES: RED DATA LISTS IN SOUTHERN AFRICA

The Southern African Botanical Diversity Network

(SABONET) is essentially a regional network of botani-

cal institutions (Huntley et al. 1998). One of it’s key

activities is the Southern African Plant Red Data List

Project which started in May 1999. The Project aims to

produce a comprehensive account of plant species threat-

ened or potentially threatened with extinction in

SABONET’s ten member countries. South Africa’s

National Botanical Institute is responsible for the overall

management and administration of SABONET. The

Southern African Plant Red Data List Project is funded

by the NETCAB Programme (Regional Networking and

Capacity Building Programme) of the World Conserv-

ation Union’s Regional Office of Southern Africa

(lUCN-ROSA). It is co-funded by the Global Environ-

ment Facility (GEE) which is implemented by the United

Nations Development Programme (UNDP).

The countries participating in the Southern African

Plant Red Data List Project are Angola, Botswana,

Lesotho, Malawi, Mozambique, Namibia, South Africa,

Swaziland, Zambia and Zimbabwe. The Project relies

on information from people working for State, paras-

tatal and non-governmental institutions, from plant scien-

tists (ecologists and taxonomists) within and outside

the region, as well as from amateur botanists, many of

whom have extensive knowledge of southern Africa’s

threatened flora. This collaborative approach is impor-

tant as there is frequently a fragmented understanding

of the region’s plant species that are threatened or

potentially threatened with extinction, the key threats

involved, and the environmental, social and economic

consequences thereof

In addition to the publication of Red Lists for the ten

countries, a secondary objective is to establish a network

of southern African threatened plant professionals, com-

petent to undertake Red Data List assessments, which

will lead to more informed conservation and research

decisions, and increased knowledge and awareness.

A task faced by botanists in many countries and

regions throughout the world is to assess plant species

according to the lUCN Red List Categories and Criteria

which were developed by the lUCN’s Species Survival

Commission (SSC) and adopted by lUCN Council in

1994 (lUCN 1994). The 1994 Red List Categories are

standardized categories with quantitative criteria that

are used to determine the threatened status (Red Data

List status) of species. The criteria help to provide

some justification and transparency to the assignment

of a particular threatened status to a species. See the

lUCN’s web site for the lUCN Red List Categories at

http://www.iucn.org/themes/ssc/redlists. At the first

World Conservation Congress held in Montreal in 1996 a

resolution was adopted by the lUCN membership

requesting the SSC to review the Red List Categories and

Criteria, and in particular, to see if they were applicable to

all organisms, especially marine species and those which

were targets of management programmes. Since 1997, the

SSC has conducted an intensive review of the Red List

Categories and Criteria, culminating in a revised system,

which was formally adopted by the lUCN Council in

February 2000. This revised system, which will be made

available later in 2000, arose out of a wide consultation

process conducted under the auspices of the lUCN/SSC

Red List Programme. The review involved the 7000

members of the SSC network and the direct participation

of more than 70 scientists (including the 22 members of

the Criteria Review Working Group) in seven workshops.

The revised system includes changes to the Red List

Categories and their definitions; revised definitions of

certain terms used to improve clarity; substantial changes

to the criteria including the thresholds used; clarification

of conceptual issues, with particular emphasis on the use

of uncertainty in assessments for poorly known or under-

stood species; and guidance on the use of the system at

national or regional levels.

The activities of the Southern African Plant Red Data

List Project include identifying country endemics, con-

solidating existing information through collaboration

and training (regional and national) as a basis for under-

taking Red List assessments. The anticipated project out-

puts include: 1, published Red Data Lists for the ten

SABONET-member countries; 2, a trained cohort of

214

Bothalia 30,2 (2000)

southern Africans knowledgeable about threatened

plant issues; 3, electronic databases of threatened plants

and associated regional human resources; 4, regular

articles on issues concerning threatened plants in

SABONET’s newsletter, SABONET News, published in

April, August and December; and 5, a web site dedicat-

ed to southern African threatened plant issues at

http://www. sabonet. org.

Global plant Red Data Lists have been compiled by

the lUCN and the World Conservation Monitoring

Centre — the 1997 lUCN World List of Threatened Plants

(Walter & Gillett 1998) and The World List of Threat-

ened Trees (Oldfield et al. 1998). Future listings need to

consider over-utilized species, country endemics, species

known from only the type locality and those known from

only single herbarium collections. In particular, Angola

and the Flora zamhesiaca countries require urgent atten-

tion. The Flora of southern Africa countries have been

afforded more comprehensive accounts owing to the rel-

atively long history vested in Red Data Lists (Hall et al.

1980; Hall & Veldhuis 1985; Hilton-Taylor 1996a, b,

1997; Scott-Shaw 1999).

Red Data Lists reflect to some extent, changes in plant

population dynamics, and therefore, attempts to monitor

and document these changes need to form part of an ongo-

ing process. If Red Data Lists are not compiled, then many

extinctions, including those of potential economically and

socially important plants, may occur before effective con-

servation action can be taken. lUCN Red Lists are widely

known for their objective and rigorous scientific

approaches for evaluating the extinction risk faced by

plants and animals. It is in this spirit that the Southern

African Plant Red Data List Project aims to lay the foun-

dations for showing the reality of plant biodiversity degen-

eration and the potential extinction crisis in the region.

REFERENCES

HALL, A.V., DE WINTER, M., DE WINTER, B., & VAN OOSTER-

HOUT, S.A.M. 1980. Threatened plants of southern Africa.

South African National Scientific Programmes Report No. 45.

CSIR, Pretoria.

HALL, A.V. & VELDHUIS, H.A. 1985. South African Red Data Book

plants: Fynbos and Karoo Bioines. South African National

Scientific Programmes Report No. 117. CSIR, Pretoria.

HILTON-TAYLOR, C. 1996a. Red Data List of southern African

plants. Strelitzia 4. National Botanical Institute, Pretoria.

HILTON-TAYLOR, C. 1996b. Red Data List of southern African

plants. 1. Corrections and additions. Bothalia 26: 177-182.

HILTON-TAYLOR, C. 1997. Red Data List of southern African plants.

2. Corrections and additions. Bothalia 27: 195-209.

HUNTLEY, B.J., MATOS, E.M., AYE, T.T., NERMARK, U., NAGEN-

DRAN, C.R., SEYANl, J.H., DA SILVA, M.A.C., IZIDINE, S.,

MAGGS, G.L., MANNHEIMER, C., KUBIRSKE, R., SMITH,

G.F., KOEKEMOER, M., DLAMINI, G.M., PHIRl, P.S.M.,

NOBANDA, N. & WILLIS, C.K. 1998. Inventory, evaluation

and monitoring of botanical diversity in southern Africa: a re-

gional capacity and institution building network (SABONET).

Southern African Botanical Diversity Network Report No. 4.

lUCN. 1994. lUCN Red List categories. Prepared by the Species

Survival Commission. lUCN, Gland, Switzerland.

OLDFIELD, S., LUSTY, C. & MACKINVEN, A. 1998. The World list of

threatened trees. World Conservation Press, Cambridge, United

Kingdom.

SCOTT-SHAW, C.R. 1999. Rare and threatened plants of KwaZulu-

Natal and neighbouring regions. KwaZulu-Natal Nature

Conservation Service, Pietermaritzburg, South Africa.

WALTER, S.K. & GILLETT, H.J. (eds). 1998. 1997 lUCN Red List of

Threatened Plants. Compiled by the World Conservation

Monitoring Centre. lUCN — The World Conservation Union,

Gland, Switzerland and Cambridge, United Kingdom.

J.S. GOLDING*

* SABONET, National Botanical Institute, Private Bag XlOl, 0001

Pretoria.

MS. received: 2000-06-01.

Bothalia 30,2: 215-224 (2000)

OBITUARIES

OTTO HEINRICH VOLK ( 1903-2000)

Otto Heinrich Volk (Figure 1) was bom in Richen,

near Heidelberg, Germany, on 6th December 1903. He

was the son of Pastor Volk, who died in 1913 (Herre

1971; Wiss 1983). He went to school in Heilbronn,

where he matriculated in 1923. After military service, he

studied natural science at the Universities of Munich,

Vienna and Heidelberg, graduating Dr.Phil.Nat. His

examiners were Prof. L. Joost and Prof. H. Walter, whose

names will be familiar to all students of ecology and veg-

etation science. In 1930 he was appointed assistant to

Prof. H. Burgeff at the University of Wurzburg, and in

1936 he became a lecturer at the same university. After

the Second World War, he returned to Wurzburg, where

he became Professor of Botany in 1950. Soon after this

he set out for Afghanistan, where he lectured in Botany

for two years at the University of Kabul. He returned to

Wurzburg to head the newly-founded Institute for

Pharmacognosy, and from 1957 to 1963 was the head of

the Academic Foreign Office, appointed through the

Senate of the University of WUrzburg. He retired in

1972. Retirement did not, however, diminish his botani-

cal interests and activities for many years. Eventually he

moved to an old-age home near Tubingen where he died

on 28th January 2000.

Not long after his return to Germany after the Second

World War, he developed an interest in medicinal plants

and pharmacognosy. Apart from leading him to high

office in his university, this interest allowed him to

FIGURE 1. — Otto Heinrich Volk (1903-2000), photographed in about

1966. Photo from the National Botanical Institute archives.

travel to many interesting and obscure places around the

world in search of medicinal plants. These expeditions

included one to Central and South America in the early

1970’s, and one of several months duration in Australia

in 1979. In 1958 he visited Peru for a congress of South

American botanists. He visited Peshawar in Afghanistan

in 1960, in connection with a UNESCO conference on

medicinal plants.

He first visited South West Africa [Namibia] in 1937,

and was interned at Windhoek and later at Andalusia (now

Jan Kempdorp in North-West, South Africa) from 1940

until 1944, during the Second World War. Here he taught

botany to a remarkable group of his fellow internees,

including Messrs Willy Giess and Rudolf Strey. After the

war, these two gentlemen became the driving forces of

Windhoek Herbarium (now the National Botanical

Research Institute of Namibia) and Natal Herbarium,

respectively. Other well-known names who studied with

Dr Volk at this stage were H. Kinges (later Professor of

Biology at Worms), and H.-J. Wiss (who returned to

Namibia to farm, became President of the SWA

Wissenschaftliche Gesellschaft 1960-61, and contributed

numerous botanical and botanic-historical items to that

Society’s newsletter). In addition to botany, various lan-

guages, agriculture, zoology, chemistry, physics and geol-

ogy were taught. Examination certificates from Andalusia

were recognised as being of university standard in

Germany after the War (Herre 1975). At Andalusia, Vofk

( 1944) was the prime mover in the production of a curious

and now very rare book entitled Bestimmiingsschliissel fur

QBEiHmmunqsfdifiiffEt

flic [liaiDEitQ'ftitam'fciie tficasgaitunriEn

oon Pr. O. DoIE

Do3«nt )flc Botonlt on 6(i Unlottlltot tDGtjburfl

^aljldinltte Don CD. ®it8

1 9 k k

FIGURE 2. — Title page of the book Volk produced while interned at

Andalusia. Digital reproduction of a photocopy in the posses-

sion of the senior author.

216

Bothalia 30,2 (2000)

siidwestafrikanische Grasgathmgen [Identification key

for South West African grass genera] (Figure 2). The type

for this book was made suneptitiously out of used tooth-

paste tubes (which at that time were made of lead), and the

illustrations were engraved by Mr Giess on pieces of

tomato-box wood. Some of the type and engravings are

presently on display in the Swakopmund Museum.

Notwithstanding his experiences in the internment

camp, Volk returned to Namibia in 1956 to continue his

researches there. The liverworts he collected on this

expedition are the subject of a paper by Amell ( 1957). In

1963 he and his wife again visited the territory, where he

carried out ecological studies, returning to East and

South Africa and Namibia for six months in 1972. Other

visits to our area were in 1968, 1981, 1984 and 1985. On

one of the earlier expeditions to Namibia, Volk attempt-

ed to collect a specimen of one or another species of suc-

culent Euphorbia, always a dangerous operation. On this

occasion he received a drop of latex in his eye. Quick

thinking and fast action by his companions saw his eye

bathed in large quantities of milk while he was taken to

the nearest hospital, some distance away. This action

probably saved the sight in that eye.

^6^ ^^4- ^ c,, p rX^ — ^

xa SrG-t 60/< -C-j ^

y^-tusw-) Ow I 'ckk

— I-

I ^ J? ^ C-^ Ct

^ <9(5 yyu ^ 'hn'i Ck>-i) <JV —

^ o>c. i) , ^

! ,

/A 0.0^ -

9^ be, l-x. jAo X. ^A>-c /

c>6 -A oA Axr-

FIGURE 3. — Part of a letter written by Dr Volk to the second author.

Black ink and pencil on blue paper, reproduced digitally.

conjunction with Volk, and several articles by Prof.

Wolfram Hartung (1987, 1992, 1994), attest to this.

His first wife. Rose, a botanical artist, contributed

about 70 excellent line drawings of Namibian plants,

especially grasses, to the classic work by Walter & Volk

( 1954) on pasture management in that country. She bore

him two daughters. In 1950 he married Miss Irene

Rubrow. There were no children from this marriage.

Volk collected over 6 000 specimens in southern

Africa and ± 1 600 in Afghanistan; these are housed in M,

PRE, NH, WIND and WBM (abbreviations according to

Holmgren et al. 1990). It is reported that on each expe-

dition to southern Africa at least, he started a new regis-

ter at number 1; therefore, it is essential to cite a date

when referring to any Volk specimen. He is commemo-

rated in the genus name Volkiella Merxm. <fe Czech (Cyper-

aceae) and the specific epithets Acacia volkii Suesseng.,

Rhus volkii Suesseng., Riccia volkii S.W.Amell, and others.

He was for many years a corresponding member of

the SWA Wissenschaftliche Gesellschaft.

His main studies were in plant geography and materia

medica, and these are the theme of most of his publica-

tions. An early fascination with succulents matured into

an interest in plants of arid areas in general, as can be seen

from the publications list below. Later, he started working

on liverworts, especially Riccia, and wrote (Figure 3) sev-

eral papers on this genus. He also possessed the ability to

arouse enthusiasm for his particular plants in others. The

publications on chromosomes in Riccia and other liver-

worts by Bomefeld (1984, 1987, 1989), as well as the

series of papers on Riccia by the second author, some in

ACKNOWLEDGEMENTS

We are grateful to Dr O.A. Leistner for reading drafts

of this obituary, and offering helpful comment and infor-

mation. We also wish to thank Dr Bomefeld for sending

us a list of Prof Volk’s publications.

REFERENCES

ARNELL. S. 1957. Hepaticae collected in South West Africa by Prof.

Dr O.H. Volk. Mitteihmgen der botanischen Staatssammlung,

MUnchen 2: 262-272.

BORNEFELD, T. 1984. Chromosomanalyse der Gattung Riccia von

Slid- und SW-Afrika und allgemeine Bemerkungen zur Zyto-

genetik der Lebeimoose. Nova Hedwigia 40: 313-328.

BORNEFELD, T. 1987. The natural system of Marchantiales based

upon cytogenetic and morphological evidence. Nova Hedwigia

45: 41-52.

BORNEFELD, T. 1989. The Riccia species of S- and SW-Africa.

Chromosome numbers and composition of the chromosome

sets. Nova Hedwigia 48: 371-382.

HERRE, A.G.J.H. 1971. The genera of the Mesembryanthemaceae.

Tafelberg, Cape Town.

HERRE, A.G.J.H. 1975. Erinnerungen an die Lagerzeit in Andalusia. In

R. Kock. Erinnenmg an die Internienmgszeit (1939-1946) und

zeitgeschichtliche Ergdnzungen: 51-107. Andalusia, Windhoek.

HOLMGREN, P„ HOLMGREN, N. & BARNETT, L.C. 1990. Index

Herbarionim, edn 8. New York Botanical Garden, New York.

VOLK, O.H. 1944. Bestimmimgsschliissel fur siidwestafrikanische

Grasgattungen. SK-Druck, Andalusia.

WALTER, H. <fe VOLK, O.H. 1954. Grundlagen der Weidewirtschaft

in Siidwestafrika und Bestimmungsschlussel fiir die siidwest-

afrikanischen Grasgattungen. Ulmer, Stuttgart.

WISS, H.-J. 1983. Prof. Dr. Emeritus Otto Heinrich Volk zum 80.

Geburtstag 1983. Mitteihmgen der SWA Wissenschaftliche

Gesellschaft 24,8, 9: 7-10.

PUBLICATIONS BY O.H. VOLK

ABOU-MANDOUR, A. A. & VOLK. O.H. 1970a. Untersuchungen

iibcr Cytokinine parasitischer Herkunft: 1. Cytokinin-Aktivitat

in Blattem von Betula pendula Roth als Folge eines Befalles

mit Stigmella argentipedella Zeller (Lep.). Zentralhlatt fiir

Bakteriologie, Parasitenkunde, Infektionskrankheiten und

Hygiene 125: 77-84.

ABOU-MANDOUR, A. A. <& VOLK, O.H. 1970b. Untersuchungen

iiber Cytokinine parasitischer Herkunft: II. Versuche iiber die

Bothalia 30.2 (2000)

217

Auswirkungen eines cytokininartigen Faktors aus mit Stigmella

argentipedella Zeller (Lep.) befallenen Birkenblattern auf das

photosynthetische '■'C-Markierungsmuster bei Ankistrodesmus

braunii. Zentralblatt fiir Bakteriologie, Pamsitenkunde, Infek-

tionskrankheiten iind Hygiene 125: 295-302.

ABOU-MANDOUR. A. A. & VOLK. O.H. 1971. Nachweis von

Cytokinin-Aktivitat in rost-infizierten Pe/argonlnm-Blattern,

Zeitschrift fiir Pflanzenphysiologie 65: 240-247.

ABOU-MANDOUR. A. A., VOLK, O.H. & REINHARD, E. 1968.

Uber das Vorkommen eines cytokininartigen Faktors in

Cusciita reflexa. Planta {Berlin) 82: 153-163.

BORNEFELD, T. & VOLK. O.H. In press. Bemerkungen zu einer

Aufsmmlung von Lebermoosen (Hepaticae, Marchantiales) bei

Omaruru im Friihjahr 1995. Dinteria.

BORNEFELD, T. VOLK, O.H. & WOLE. R. 1996. Exormotheca bul-

bigena sp. nov. (Hepaticae. Marchantiales) and its relation to E.

hohiii in southern Africa. Bothalia 26: 159-165.

CZYGAN. F.-C., KRUGER, A., SCHIER. W. & VOLK, O.H. 1977.

Pharmazeutische-biologische Untersuchung der Gattung Har-

pagophytum (Bruch.) DC. ex Meissn. 1. Mitteilung: Phytochem-

ische Standardisierung von Tubera Harpagophyti. Deutsche

Apotheker Zeitung 117: 1431-1434.

HARTUNG, W.. HELLWEGE, E.M. & VOLK. O.H. 1994. The func-

tion of abscisic acid in bryophytes. Journal of the Hattori

Botanical Laboratory 76: 59-65.

HARTUNG, W„ WEILER. E.W. & VOLK. O.H. 1987. Immunochemical

evidence that abscisic acid is produced by several species of

Anthocerotae and Marchantiales. The Bryologist 90: 393^00.

HAUTUM, MULLER & VOLK. O.H. 194L Eine Flupregulierung.

Deutsche Wasserwirtschaft 36.

HAUTUM. MULLER. VOLK, O.H. & K(DHLER. 1939. Die Staustufe

Schlopfurt. Deutsche Wasserwirtschaft 34: 1-8.

HELLWEGE. E.M., DIETZ. K.-J.. VOLK, O.H, & HARTUNG, W.

1994. Abscisic acid and the induction of desiccation tolerance

in the extremely xerophilic liverwort Exormotheca holstii.

Planta 194: 525-531.

HELLWEGE, E.M.. VOLK. O.H. & HARTUNG, W. 1992. A physio-

logical role of abscisic acid in the liverwort Riccia fluitans L.

Journal of Plant Physiology 140: 553-556.

HERRE. A.G.J. & VOLK. O.H. 1948. Mesembryanthemaceae Herre &

Volk, familia nova. Siikkulentenkunde (Zurich) 2: 38.

HERRE, A.G.J. & VOLK, O.H. 1961. Key to the genera of mesem-

bryanthemums. In H. Jacobsen. Handbook of succulent plants

3: 965-976. Blandford, London.

HERRE. A.G.J. & VOLK. O.H. 1971. Key to the genera of the

Mesembryanthemaceae. In A.G.J. Herre. The genera of the

Mesembryanthemaceae: 15-24. Tafelberg. Cape Town.

JACOBSEN. H. & VOLK, O.H. 1961. The form and mode of life of

the mesembryanthemums. In H. Jacobsen, A handbook of suc-

culent plants 3: 923-927.

JACOBSEN. H.. VOLK, O.H. & HERRE. A.G.J. 1950. Mesembryan-

themaceae. Ulmer, Ludwigsburg.

PEROLD. S.M. & VOLK. O.H. 1988a. Studies in the genus Riccia

(Marchantiales) from southern Africa. 8. R. campbelliana (sub-

genus Rica'a), newly recorded for the region. Bothalia 18: 37^2.

PEROLD. S.M. & VOLK. O.H. 1988b. Studies in the genus Riccia

(Marchantiales) from southern Africa. 9. R. nigrella and the sta-

tus of R. capensis. Bothalia 18: 43-49.

POFILMANN. G & VOLK. O.H. 1959. Tylose, ein wasserldsliches

Abdruckmterial fiir Oberflachenuntersuchungen. Zeitschrift fiir

wissenschaftliche Mikroskopie und mikroskopische Technik 64:

252-256.

REINHARDT, E.. CORDUAN, G. & VOLK. O.H. 1967. Die Scopo-

letinabgabe von Nicotiana ta/jaciun-Gewebekulturen an das

Medium in kontinuierlichen Suspensionskulturen. Planta Medica

15: 357-360.

REINHARDT, E., CORDUAN, G. & VOLK. O.H. 1968a. Uber

Gewebekulturen von Ruta graveolens. Planta Medica 16: 8-16.

REINHARDT, E.. CORDUAN, G. & VOLK. O.H. 1968b. Nachweis

von Harmin in Gewebekulturen von Peganum harmala.

Phytochemistry 7: 503. 504.

ROCKENSTEIN, E. & VOLK. O.H. 1971. Beitrage zur Kenntnis der

Digitalis purpurea L. sardinischer Herkunft. Planta Medica 19:

270-278.

SCHAPKA. U. & VOLK. O.H. 1979. Ein Verzeichnis von in Afghani-

stan gebrauchlichen Pflanzennamen. Afghanistan Journal 6: 3-14.

SCHIPPER. A. & VOLK. O.H. 1960. Beitrage zur Kenntnis der

Alkaloide von Peganum harmala. Deutsche Apotheker Zeitung

100: 255-258.

VOLK. O.H. 1931. Beitrage zur Okologie der Sandvegetation in der

oberrheinischen Tiefebene. Zeitschrift fiir Botanik 24: 81-185.

-1934. Ein neuer fUr botanische Zwecke geeigneter Lichtmesser.

Berichte der Deut.schen Botanischen Gesellschaft 52: 195-202.

-1935. Kalk- und Gipspflanzen, ein Beitrag zu dem Kapitel Boden und

Pflanzen. Berichte der Deutschen Botanischen Gesellschaft 53:

796-806,

-1937a. liber einige Trockenrasengesellschaften des WUrzburger

Wellenkalkgebietes. Beiheft zum Botanischen Centralblatt 57:

577-598.

-1937b. liber das Verhalten der Pflanzen bei Trockenheit. Berichte der

Physikalisch-medizinischen Gesellschaft zu Wurzburg Neue Folge

60: 16-23.

-1937c. Untersuchungen Uber das Verhalten der osmotischen Werte

von Pflanzen aus Steppenartigen Gesellschaften und lichten

Waldem des mainfrankischen Trockengebietes. Zeitschrift fiir

Botanik 32: 65-149.

-1939. Rooi Slangkop — Der Schlagkopf (Urginea sanguinea). Siid West

Persianer-Markt. Kalkfeld SWA.

-1940. Soziologische und dkologische Untersuchungen an der

Auenvegetation im Churer Rheintal und Domleschg. Jahres-

bericht der Naturforschenden Gesellschaft Graubiindens,

1938/3916: 1-51.

-1944. Bestimmungsschliissel fiir siidwestafrikanische Grasgattungen.

SK-Druck, Andalusia.

-1949a. Zur Kenntnis der Pollinien der Asclepiadaceen. Berichte der

deutschen botanischen Gesellschaft 62: 68-72.

-1949b. Review: Obemdorfer, Pflanzensoziogische Exkursionsflora

fur Sudwestdeutschland und die angenzenden Gebiete.

Vegetatio 1: 385.

-1950. Vegetationseindrucke in Afghanistan. Vegetatio, Acta Geo-

botanica 3: 210-212.

-195 1 . Zur Kenntnis der Stapelienbluten. Sukkulentenkunde (Zurich) 4:

46-59.

-1952. Pilzgarten und Klima-Anlagen im Termitenbau. Orion, naturwis-

senschaftliche-technische Zeitschrift fiir Jedennann: 41-^3.

-1954a. Landwirtschaftliche Probleme Afghanistans. Mitteilungen des

Instituts fiir Auslandsbeziehungen 4: 233-236.

-1954b. Klima- und Pflanzenverbreitung in Afghanistan. Vegetatio,

Acta Geobotanica 5-6: 422-433.

-1955a. Sukkulentenformen. In H. Jacobsen. Handbuch der sukkulen-

ten Pflanzen 3: 9-13. Fischer, Jena.

-1955b. Geographische Verbreitung. In H. Jacobsen. Handbuch der

sukkulenten Pflanzen 3: 20-24.

-1955c. Bodenverhaltnisse. In H. Jacobsen, Handbuch der sukkulenten

Pflanzen 3: 24-27.

-1955d. Bluten und Frucht der Mesembryanthemaceae. In H. Jacobsen.

Handbuch der sukkulenten Pflanzen 3: 46-51.

-1955e. Afghanische Drogen. Planta Medica 3: 129-146.

-1955f. Materia medica Afghanistans. Phannazeutische Zentralhalle 94.

-I955g. Verschiedene Beitrage zur Okologie und Verbreitung der

Mesembryanthemaceae. In H. Jacobsen, Handbuch der sukku-

lenten Pflanzen 3. Fischer. Jena.

-1956a. Die Weideverhaltnisse in Sudwest. Allgemeine Zeitung,

Windhoek nr. 113.

-1957. Beobachtungen in neuerer Zeit uber Verfalschungen und Un-

terschiebungen bei pflanzlichen Drogen. Deutsche Apotheker

Zeitung 97: 421^23,

-1958. Trockenrasen aus der Umgebung von Pavia. Archivio Botanico

e Biogeografico Italiano 34: 31-49.

-1960a. Geographical distribution. In H. Jacobsen, Handbook of suc-

culent plants 3: 928-930. Blandford, London.

-1960b. Soil conditions. In H. Jacobsen, Handbook of succulent plants

3: 931, 932.

-1960c. Flowers and fruits of the mesembryanthemums. In H.

Jacobsen. Handbook of succulent plants 3: 945-950.

-1961a. A survey of Afghan medicinal plants. Pakistan Journal of

Scientific and Industrial Research 4: 232-238.

-1961b. Aus der Okologie des Lebensraumes im Mittleren Osten. In R.

Paret. Die Welt des Islam und die Gegenwart: 171-178. Kohl-

hammer Verlag, Stuttgart.

-1963. Eine Geographie von Afghanistan. Die Erde, Zeitschrift der

Gesellschaft fiir Erdkunde 94: 73-76.

-1964a. Zur Kenntnis von Harpagophytum procumbens DC. Deutsche

Apotheker-Zeitung 104: 573-576.

-1964b. Die afro-meridional-occidentale Floren-Region in SWA.

Beitrage zur Phytologie zu Ehren Prof. H. Walters 65 Geburt-

stages. Ulmer, Stuttgart.

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tropischen SUdwestafrika. In K. Buchwald, W. Lendholt & K.

Meyer, Beitrage zur Landespflege 2: 108-131.

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Siidwestafrika (Namibia). I. Mitteilimgen der botanischen

Staatssanunlung, Miinchen 15: 223-242.

-1981. Beitrage zur Kenntnis der Lebermoose (Hepaticae) aus

Siidwestafrika (Namibia) II. Mitteilimgen der botanischen

Staatssammhmg, Miinchen 17: 245-252.

-1983. Vorschlag fiir eine Neugliederung der Gattung Riccia L.

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453^65.

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afrika/Namibia IV. Zur Biologie. Nova Hedwigia 39: 117-143.

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2771-2775.

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hoeker Bergland, Siidwestafrika. Journal der SWA Wissen-

schaftUche Gesellschafl 25: 5^4.

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Riccia (Marchantiales) from the south-west Cape. Bothalia 15:

117-124.

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Africa. 1. Two new species of the section Pilifer: R. duthieae

and R. alatospora. Bothalia 15: 531-539.

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Africa. 2. A new species of the section Pilifer: R. sarcosa.

Bothalia 16: 25-21 .

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Africa. 3. R. schelpei, a new species, in the new subgenus

Chartacea. Bothalia 16: 29-33.

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Africa. 4. Three endemic species, R. natalensis, R. microcilia-

ta sp. nov. and R. mammifera sp. nov. Bothalia 16: 169-180.

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Africa. 5. R. rosea, a new species. Bothalia 16: 181-185.

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Africa. 6. R. hirsuta, a new species, in a new section. Bothalia

16: 187-191.

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Africa. 15. R. hirsuta and R. tomentosa, sp. nov., two distinct

species previously treated as one. Bothalia 20: 23-30.

VOLK, O.H., PEROLD, S.M. & BORNEFELD, T. 1988. Studies in the

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new white-scaled species of the group ‘Squamatae’: R. argen-

teolimbata and R. albornata. Bothalia 18: 155-163.

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M. 1972. M. Linosid A und B, zwei neue Flavon-C-glykoside aus

Linum maritimum L. Zeitschrifl fiir Naturforschung 27b:

808-812.

WALTER, H. & VOLK, O.H. 1954. Grundlagen der Weidewirtschaft

in Siidwestafrika und Bestimmungsschlussel fiir die siidwest-

afrikanischen Grasgattungen. Ulmer, Stuttgart.

H.F. GLEN* and S.M. PEROLD*

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

IN MEMORY OF S.W. ARNELL, HEPATICOLOGIST (1895-1970)

Sigfrid Wilhelm Arnell (Figure 4) was born in Gavle,

Sweden on the 5th March 1895. He studied medicine,

first in Uppsala and later in Stockholm, where in 1922,

he obtained the ‘medicine licentiat’, a Swedish degree

which permitted him to practice medicine. His main

interest in medicine was in the rapidly developing field

of X-ray diagnostics and most of his medical career was

devoted to heading the X-ray department of the hospital

in his native city. In 1959 he publicly defended his thesis

for the degree of doctor of medicine and received this

degree in the same year in Stockholm.

It was only at the relatively late age of 45, that Arnell

really became interested in liverworts and began to

work with this group of plants consistently, inasmuch

as his free time permitted. Originally, he studied

Fennoscandian hepatics (Arnell 1956), but later he

became very interested in the hepatics of the southern

hemisphere, and in particular, those of South Africa.

For his contributions to hepaticology, the degree 'doc-

tor honoris causa' was conferred on him in Uppsala in

1958. Hassel de Menendez (1976), the eminent

Argentinian hepaticologist, remarked that Arnell’s

(1963a) Hepaticae of South Africa was a useful start to

the study of the liverworts of that part of the world. She

conceded, however, that very much more work re-

mained to be done there.

Because Amell’s name and particulars were unfortu-

nately left out in Gunn & Codd’s ( 1981 ) Botanical explo-

ration of southern Africa, the second author had tried over

the years to procure some information about him. and was

later infomied that an obituary (freely refeiTed to in the

above paragraphs), as well as a list of his hepaticological

publications, were published by Olle Martensson in 1972.

As it will be the 30th anniversary of Arnell’s death on

26 October 2000, it seems fitting to pay tribute to his

botanical efforts in southern Africa, which he visited in

1951 and which culminated in many papers and the

above-mentioned book. Without Arnell’s input, which

was done in his spare time and at his own expense, there

would have been little to build on for any South African

hepaticologist who followed him. His work has been

criticized, sometimes quite severely, but one should also

bear in mind that, as a medical doctor, he well knew what

Bothalia 30.2 (2000)

219

FIGURE 4. — Sigfrid Amell beside Lake Mavasjaure. northern Sweden

in the summer of 1960.

the prognosis for his Parkinson’s disease would be, and

how limited his time was. Furthermore, his eyesight was

also failing. Apparently, after 1965 he became ‘so dis-

abled that he was no longer able to deal with scientific

work’ (Martensson 1972).

A friend of Amell’s, Mr Gillis Een, writes the follow-

ing personal recollections about him, in dedication to his

memory:

Sigfrid Amell was a second generation bryologist. His

father, Hampus Wilhelm Amell 1848-1932, was a pro-

fessional botanist and teacher, whereas Sigfrid was a

medical doctor and an amateur botanist. He had inherit-

ed from his father not only his interest in hepaticology,

but also an old microscope that he had used all his life. I

remember seeing it in his study — an old-fashioned mo-

nocular brass model.

The first time I met Sigfrid Amell was very special. It

was in the summer of 1954 and I was botanising in

Abisko with Hemian Persson and William Steere. Early

one fine morning, Sigfrid and Olle Martensson turned up

outside our hotel. They had been hiking all night in order

to climb Mount Nissontjarro during the cool hours,

which is a very sensible thing to do so far north of the

Arctic Circle, where the sun never sets in the summer.

They had been looking for Scapania kaurinii Ryan (S:

B25018), an arctic-alpine species, and they had found

plenty of it, as well as Scapania spitsbergensis (Lindb.)

Miill.Erib. (S: B25019). Sigfrid gave me samples of both,

which for many years were very cherished items in my

private collection of bryophytes. The day before, I had

coWected Arnellia fennica (Gottsche) Lindb. (S; B25020)

below the hotel, belonging to a genus that was named

after Sigfrid ’s father.

When Sigfrid retired from the medical profession in

1960 at the age of 65, he moved from Gavle to Bromma,

a suburb of Stockholm, and we became neighbours with-

in a few blocks of each other. His retirement gave Sigfrid

an opportunity to spend more time on bryology. We met

rather often to discuss Scandinavian hepatics, and gradu-

ally we began to make plans for two collecting journeys,

one towards the Arctic Circle and one towards the Tropic

of Cancer.

He was still very fit and in the summer of 1960 we

organised a collecting trip to northern Sweden, more pre-

cisely, to Lake Mavasjaure in Arjepluog parish in Pite

Lappmark. In preparation for the journey, we pitched

Sigfrid’s old tent on the lawn of the garden of the Arnell

house, which attracted some attention, but we found that

it could still be used. Sigfrid’s wife called us boy scouts

who had never grown up!

Our luggage was on the heavy side and we were not

interested in long hikes. For those reasons we hired a taxi

aeroplane, which landed on the lake itself, where we

hired a small rowing boat in the summer camp of the

reindeer herdsmen. Air taxi-ing was rather unusual in

Sweden at that time, but brought us into an area where

few botanists had been before.

Sigfrid was extremely active in the field and was

always clean-shaven, even in the rather primitive camp-

ing conditions. He insisted on doing the rowing while I

did the bailing. He waded through the bogs and climbed

the mountains. He wore glasses with two pairs of lens-

es— one pair could be flipped up or down in order to

change the focal distance. His collecting was rather dis-

criminating and he carefully avoided all the common

species.

It so happened that 1960 was a lemming year in the

Scandinavian mountains. These small rodents were

everywhere. When we met them on a reindeer path, com-

ing in the opposite direction, they sat down on their

behinds and barked at us, obviously expecting us to leave

the path free for them. Sigfrid, who had a kind heart, had

problems every time we pitched our tents, because he did

not want to harm them or block their way. The lemmings

began to migrate and in so doing, they attempted to swim

across the lake. Thousands of them died in the cold water

and were washed ashore, where their small bodies

formed long ridges.

Sigfrid was incredibly even-tempered. I remember a

rainy day in Pite Lappmark. Sigfrid’s old-fashioned, tall

tent was used as a kitchen in bad weather. That particu-

lar day, I managed to turn over both the little spirit stove

and the kettle with boiling tea-water. The tent became

very damp and unpleasant. He did not say one harsh

word to me, neither then, nor the next morning, when we

had sunshine and could dry out his tent.

220

Bothalia 30,2 (2000)

Sigfrid never boasted, but he was certainly very proud

of his achievements. I remember when we wrote our

names in the guest book of the same summer camp at

Mavasjaure. He wrote after his name ‘Dr. med. and Dr.

phil. hon. c.’ I do not know whether the reindeer herds-

men, who managed the camp, were impressed or not, but

other visiting Swedes probably were.

In December 1961 we took a chartered flight to

Egypt. For the first two weeks we were sightseeing,

accompanied by very competent guides, one of whom

was our botanical host in Cairo, Prof. Vivi Tackholm.

She led us around the historical museum, where she told

us about all the plants that were put in the graves with the

Pharaohs’ embalmed bodies, their scientific names and

what they signified. She also took us to the herb market,

where she told us what kind of food the spices were used

in, and what herb to buy if one wanted to poison one’s

neighbours!

Sigfrid had a very broad education and many inter-

ests. He enjoyed every part of the sightseeing tour and

was good company. He was particularly interested in tex-

tiles and one of his hobbies was that of knotting mats

which are known as rya rugs.

Vivi organised all our field trips during the weeks that

followed. She was a remarkable woman in many ways —

she came to Cairo as the wife of a Swedish professor of

botany. When he died rather young, she simply took over

the job and made it an incredible success. She published

a Flora of Egypt, and also made friends everywhere.

Sometimes she joined us, together with some of her stu-

dents, many of whom were women, unusual for that

time.

Our visit to Egypt took place only a few years before

the completion of the big dam in Aswan. The Nile still

had its yearly pulse and in December the water level

was retreating. The mud-banks were drying up and

cracking. I have vivid memories of Sigfrid sliding in

mud, looking for Riccias. He found many species of

that particular genus, such as Riccia frostii Austin (S:

B24340). 1 took photographs of the beautiful patterns

they sometimes formed on the mud, slides which are

now deposited in NRM/KBO {K00547, K00548,

K00549 & K00550). This is just another bryophyte

habitat that has been completely destroyed in our life-

time. Sigfrid published some of his findings a few years

later (Arnell 1963b).

In 1962 1 was professionally engaged in the technical

development of certain processes in the cane sugar

industry. 1 made a long journey that brought me to Natal

in South Africa, to Mauritius and Reunion in the Mas-

carenes and to Queensland and NSW in Australia. 1 col-

lected as many bryophytes as I could on weekends and

in my spare time. At that time Sigfrid had almost fin-

ished his Hepaticae of South Africa, which was pub-

lished the next year. Before 1 left for South Africa, he

and Herman Persson impressed upon me, that for a very

long period of time, no bryologist had visited the

Mascarenes, so 1 decided to take a week off from my

work, in order to do some more serious collecting on the

two main islands.

I always sent my extra Scandinavian collections to

Herman Persson at NRM in Stockholm as it saved me a

lot of trouble with customs. This time, Sigfrid took pos-

session of the collection as soon as it arrived, extracted

all the hepatics and started to identify them. The result

was a publication (Arnell 1965) which was to be his

last.

What I did not know then, but learned much later, was

that Sigfrid’s eyesight was failing him very rapidly. That

may explain why he was so eager to finish the work

while he still could, and also why many hepaticologists,

who at later dates have studied my samples, have dis-

agreed with some of his determinations.

Sigfrid also studied my collections from Queens-

land, resulting in the description of two new species

(Arnell 1963c). The type specimens were deposited in

the herbarium of Uppsala University. For many years

they could not be found there. The explanation is prob-

ably that Sigfrid moved to Uppsala, where he died in

1970, and his private herbarium was donated to the

University, where it was kept apart. It was only recent-

ly that Lars Soderstrom was able to locate the two type

specimens.

This is how I remember Sigfrid Arnell. Dates and

other details may be slightly inaccurate, but I have tried

to paint a true picture of a very dear friend.

REFERENCES

For a complete list of hepaticological works by S.W.

Arnell, see Martensson: 240-242 (1972).

ARNELL. S.W. 1956. Illustrated moss Flora of Fennoscandia.

Gleemp. Lund. Sweden.

ARNELL. S.W. 1963a. Hepaticae of South Africa. Swedish Natural

Science Council. Stockholm.

ARNELL. S.W. 1963b. Some hepatics new to Egypt. Botaniska Notiser

116: 7-10.

ARNELL. S.W. 1963c. Two new hepatics from Queensland. Svensk

Botanisk Tidskrift 57: 190-192.

ARNELL, S.W. 1965. Hepaticae collected by Mr. Gillis Een in

Mauritius and Reunion in 1962. Svensk Botanisk Tidskrift 59:

65-84.

GUNN. M. & CODD. L.E. 1981. Botanical exploration of southern

Africa. Balkema. Cape Town.

HASSEL DE MENENDEZ. G.G. 1976. Taxonom.ic problems and

progress in the study of the Hepaticae. Journal of the Hattori

Botanical Laboratory 41: 19-36.

MARTENSSON. O. 1972. Obituary: Sigfrid W. Arnell 1895-1970.

Journal of Bryology 7: 239-242.

G. EEN* and S.M. PEROLD**

* The Swedish Museum of Natural History. Box 50007. SE-104 05

Stockholm. Sweden, email: gillis. een(®nrm.se

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

email: smp@nbipre.nbi.ac.za

Bothalia 30.2 (2000)

221

ROSEMARY CHARLOTTE HOLCROFT (1942-2000)

Rosemary Holcroft (nee Temperley) (Figure 5) was

bom in Nairobi Kenya, on 12 December 1942. Her

father. Dr B.N. Temperley, was a geologist working in

Tanzania. After two years the family returned to

England, but being fed up with the vagaries of the

English weather and the post war situation in the coun-

try, the family returned to East Africa, this time to Kenya.

Rosemary completed her schooling at Msongari Loreto

Convent in Nairobi. From an early age her parents

instilled in her a love of nature and taught her to draw.

In 1960 Rosemary left Kenya and enrolled at the

University of Cape Town Ballet School under the direc-

tion of Dulcie Howes, where she obtained a Diploma in

Dance Teaching in 1963. She taught ballet for seven

years having her own studios, first in Pretoria and then in

Cape Town. In 1964 she married Michael Roos, a well-

known artist. Her son, Marc, was born in 1972. While in

Cape Town her interest in botanical illustration first mani-

fested itself: in 1965 she prepared a set of pen and ink

illustrations of ferns for Professor E.A.C.L.E. Schelpe of

the University of Cape Town.

Rosemary moved to Pretoria in 1972 and lived with

her parents who had settled there after her father’s retire-

ment. After seeing the Botanical Research Institute’s

exhibition of botanical art at the Pretoria Art Museum, in

August 1973, Rosemary became inspired to enroll for a

B.A. (Fine Arts) course at the University of South Africa,

which unfortunately she did not complete. In 1974 she

participated in a group exhibition of botanical illustra-

tions held at the Association of South African Arts

gallery in Pretoria on the occasion of the eightieth birth-

day of Cythna Letty, doyenne of South African botanical

artists. In the same year she exhibited at the Creative Arts

exhibition and the annual central show of the South

African Aloe and Succulent Society.

In 1975 Rosemary joined the Botanical Research

Institute as a temporary half-day Senior Technician and

this was the beginning of a 10-year stint as a highly com-

petent and productive botanical artist. She contributed to

most of the Institute’s publications including 100 plates

for The Flowering Plants of Africa, 67 of which have

been published, plus 150 plates for the Flora of Pretoria,

which has yet to be published. She also contributed to

several external publications such as the Cactus and

Succulent Journal (of the USA). In 1976 she married for

a second time, this time to Leslie Holcroft, a computer

consultant.

In 1980 I decided to write a Field guide to the flora

of the Natal Drakensberg (published in 1990) and for

this I needed 48 plates illustrating over 200 species

(Figure 6). I enlisted the help of Rosemary who did the

drawings whenever she had time to spare. To obtain

material for illustrations we visited the Berg on two

occasions. The second trip was written up by Rosemary

in Veld & Flora (December 1985) in an article entitled

‘A birthday to remember’. The first two paragraphs

illustrate vividly Rosemary’s intense love of nature

which so markedly influenced her work. The paragraphs

read as follows:

Tt was December 12, 1984 and I thought how lucky I

was to be listening to such a scintillating symphony on

my birthday. I was sitting in the veld beside a tributary

that fed into the main river of the Ndedema Gorge, about

1 800 m above sea level in the Cathedral Peak area of the

Natal Drakensberg. The gurgle of water running over and

between smooth pebbles, the rush of water over slabs of

rocks and the splash of water over the edge and into rock

pools, made a harmony of sound punctuated by the

melodies of birds singing in the bushes and trees above

the mountain stream.

The sky was clear and the air was cool, with a gentle

breeze occasionally blowing the grass and veld flowers

surrounding me in the foreground. The steep slopes of

the gorge, the dark patches of forests, the rounded spurs

forming the Little Berg and the massive blocks of basalt

that formed the jagged skyline of the Dragon Mountains

were quiet and still. The perfect silhouette of Monk’s

Cowl, Cathkin Peak and Champagne Castle reminded

me of a previous visit to this land of ponderous awe and

wonder’.

Rosemary resigned from the Institute in 1985 largely

due to ill health — possibly the incipient stages of Crohn’s

Disease which afflicted her later. Shortly afterwards she

purchased a Toyota Hilux 4x4 vehicle in which she

travelled to Namibia, Botswana, Lesotho and other

places, taking photographs and writing articles for maga-

zines. She described this period of ten years as ‘the most

creative and fulfilling of her life’. Apart from travel.

FIGURE 5. — Rosemary Charlotte Holcroft (1942-2000).

222

Bothalia 30,2 (2000)

FIGURE 6.— One of the line drawings done by Rosemary on p. 97 of the book by D.J.B. Killick entitled, A //eW guide to the flora of the Natal

Drakensberg ( 1 990).

Bothalia 30.2 (2000)

223

Rosemary did a certain amount of free-lance art work.

She and Les were divorced in 1985.

In 1988 Rosemary worked at Optima, the South

African Council for the Blind in Pretoria as hostel super-

visor. In 1994 after five operations due to Crohn’s

Disease she was medically boarded from work and put

on a disability pension. She then bought a cottage at

Southport on the KwaZulu-Natal South Coast near Port

Shepstone and stayed there until she died on 4 February

2000. A memorial service for Rosemary was held in the

Botanical Garden of the National Botanical Institute,

Pretoria, on 7 March and was attended by relatives,

friends and former colleagues. And so we took leave of

an accomplished botanical artist and a good friend.

D.J.B. KILLICK

465 Sappers Contour. Lynnwood. 008 1 Pretoria.

WERNER RAUH (191 3-2000), ONE OF THE WORLD’S MOST PROLIFIC AUTHORS ON SUCCULENT PLANTS

Professor Dr Werner Rauh, (Eigure 7) one of the most

knowledgeable cactologists and succulent enthusiasts,

passed away peacefully in his sleep on 7 April 2000, a few

weeks before his 87th birthday. Wemer was bom on 16

May 1913 in Niemegk. near Bitterfeld in Saxony, one of

the 1 6 states of Germany. He passed his university entrance

examinations in 1932 at the local gymnasium. Even as a

youngster he displayed an interest in geography and biolo-

gy, which led him eventually to study botany, zoology,

chemistry and geology at the Universities of Leipzig,

Innsbmck and Halle. At the last-mentioned he studied

under Prof. Wilhelm Troll, one of the world’s greatest mor-

phologists. This exposure to an exceptional plant specialist

and keen observer no doubt further fired his interest in

plant form, and the wonderful shapes of cacti, succulents

and bromeliads became an almost full-time passion of his.

After receiving a D. Phil, in 1937 and an appointment to a

professorship in 1 939, he moved to Heidelberg as an assis-

tant to A. Seybold. For the following 61 years his name

became inseparably associated with the University of

Heidelberg. Tme to his classical botanical training, his aca-

demic activities were very much aligned towards taxono-

my, systematics, morphology and biogeography. In 1947

he was appointed as lecmrer at this University, in 1953 as

Associate Professor, in 1957 as Extraordinary Professor,

and then in 1 960 as Professor and Director of the Institute

for Systematic Botany and Plant Geography, and the asso-

ciated Botanical Garden. After his retirement at the age of

68, he remained at the Institute as Professor Emeritus (Dorr

1997; Mortimer 2000).

Werner's first intercontinental excursions were to the

Atlas Mountains in North Africa, but soon he was confi-

dently travelling to more remote areas such as relatively

unknown or unexplored parts of Peru and Ecuador (see

Schwartz 1987 for a list of Prof Rauh’s expeditions). His

appointment as an Extraordinary Professor at the

University of Heidelberg in 1957 coincided with his first

expedition to the island of Madagascar. At the same time he

was offered the directorship of the world-famous Botanical

Garden and Botanical Museum in Berlin-Dahlem. He,

however, declined the offer. The Garden was one of his pri-

maiy passions. From a small facility with three greenhous-

es at the time, it expanded to include no fewer than 15

greenhouses on his retirement. These were all filled with an

amazing array of botanical treasures, many of which he

collected on his countless expeditions to the world’s tropi-

cal and subtropical regions. Today the Botanical Garden of

the University of Heidelberg, under the directorship of

Werner’s successor. Prof Peter Leins, is a world-famous

research, education and training and display facility.

Indeed, Werner’s efforts to establish the Garden as a lead-

ing instimtion will enable it to rise to the challenges of the

new millennium.

Throughout his botanical career the cacti and other suc-

culent plants of the arid regions of southern and northern

America and southern Africa were his first loves as far as

research was concerned. The material that he brought back

from his numerous visits to remote and previously unex-

plored areas was put to good scientific use and extensively

researched. For example, he was the first to show that rep-

resentatives of the Didiereaceae and the Cactaceae could

be successfully intergrafted (Rauh & Dinklage 1972, 1978;

FIGURE 7. — Prof. Dr Wemer Rauh with a specimen of Tillandsia

fendleri Griseb. Photograph taken ± 1974 by W. Barthlott.

224

Bothalia 30,2 (2000)

Rowley 1992). He was also fascinated by the bromeliads of

the South American rain forests and the plants of the high

mountains of the tropics. These and other topics were

addressed in more than 300 scientific and semi-scientific

papers, as well as several lectures and some two dozen

popular-scientific books. From a biogeographical point of

view, Werner reserved a special place in his heart for the

botanical world of Peru and, of course, Madagascai'. On his

regular collecting trips and expeditions he discovered

numerous species new to science. Some of these were

named after him, such as the genera Rauhia Traub.

(Amaryllidaceae), Raiihocereus Backeb. (Cactaceae) from

Peru, Rauhiella Pabst & Braga (Orchidaceae) from Brazil,

and Werauhia J.R. Grant (Bromeliaceae). A number of

species were also named after him, including TiUandsia

raithii L.B.Sm. (Bromeliaceae), an enormous pineapple

plant from northern Peru, and the beautiful miniature Aloe

raiihii Reynolds (Aloaceae) from Madagascar. Especially

the last-named of these has become a popular species in

numerous collections of succulent plants. The original set

of herbarium specimens that he collected during his many

expeditions are kept in HEID. Duplicates have been

deposited in K. MO, MPU, MSUN, R PRE, and TAN

(Gunn & Codd 1981; Dorr 1997).

Eor generations to come, professional botanists and

interested amateurs alike will find the published legacy left

behind by Wemer, indispensable reference sources. His last

two books on the Red Island, Madagascar (Rauh 1995,

1998), were exceptionally well produced by Strawberry

Press, and although rather expensive, represent essential

reading material for anyone interested in the natural histo-

ry of this biological paradise. These volumes are excellent

snapshots in time of a magnificent natural flora and fauna

suffering from the mthless transformation caused by human

intervention. Wemer was also an accomplished photogra-

pher, his photographs adorning the pages of many of his

books. In some instances the picmres in his books are the

only published ones of the species, and are often so good

that they can be used to identify flowering or even non-

flowering specimens.

Professor Rauh justifiably received extensive recogni-

tion for his research accomplishments. He was an honorary

member of numerous societies as well as the Vice-

President of the International Organization for Succulent

Plant Study from 1976 to 1982 (Supthut 1999) and

President in 1983. The Republic of Pern and the Princi-

pality of Monaco awarded him medals — on the occasion of

his 65th birthday. Princess Grace of Monaco presented him

with the first ever Cactus d’Or, conferred by the lOS, in

Monte Carlo. This was the first and only Cactus d'Or made

of gold (Rowley 2000). During the course of his tenureship

as Director of the University of Heidelberg Botanical

Garden, he was awarded the golden Veitch Memorial

Medal by the Royal Horticultural Society of London and

the Willdenow Medal by Berlin. In 1999, a year before he

passed away, the Republic of Madagascar, through its

Ambassador in Germany, His Excellency Rabesa, made

him a Knight of the National Order, and in November 1999

he received the Eederal Distinguished Service Cross with

Ribbon from the mayor of Heidelberg, Beate Weber. Since

1968 Wemer was a regular member of the Academy of

Sciences and Literature in the city of Mainz and since 1980

he was a corresponding member of the Heidelberg

Academy of Sciences.

In 1 997 he suffered the loss of his wife, Hilde, while his

own health was also failing rapidly. However, he remained

mentally focused on the numerous manuscripts that he was

working on and still paid regular visits to the Botanical

Garden.

He was indeed a remarkable human being and scientist

and with his death, an era of almost charismatic botanical

exploration of tropical regions has come to an end. Interest

in succulents from the tropics were, for the past 50 years,

strongly influenced by Wemer Rauh’s numerous plant

introductions, published and illustrated papers and lectures

(Rowley 1997). Many scientists and hobbyists will remem-

ber him as an amazingly prolific author on these fascinat-

ing plants. Werner’s death came as a shock, but was not

unexpected, as he had been seriously ill with breathing

problems for some time. His funeral on 28 April in

Heidelberg was attended by family and close friends.

REFERENCES

DORR. L.J. 1997. Plant collectors in Madagascar and the Comoro

Islands. Royal Botanic Gardens. Kew.

GUNN. M. & CODD. L.E. 1981. Botanical exploration of southern

Africa. Balkema. Cape Town.

RAUH. W. 1995. Succulent and xerophytic plants of Madagascar. Vol.

1. Strawberry Press. Mill Valley.

RAUH. W. 1998. Succulent and xerophytic plants of Madagascar. Vol.

2. Strawberry Press. Mill Valley.

RAUH. W. & DINKLAGE. W, 1972. Grafting succulents. Cactus &

Succulent Journal (U.S.) 44: 139-154.

RAUH. W. & DINKLAGE. W. 1978. A new method of propagation of

Didierea madagascariensis H.Bail. Cactus & Succulent

Journal (U.S.) 50: 132. 133.

ROWLEY. G.D. 1992. Didiereaceae. ‘Cacti of the Old World’. British

Cactus & Succulent Society, Richmond,

ROWLEY, G.D. 1997. A historv of succulent plants. Strawberry Press,

Mill Valley.

ROWLEY, G.D, & MORTIMER. K. 2000. Wemer Rauh, 1913-2000.

British Cactus & Succulent Journal 18: 68, 69.

SCHWARTZ, H. 1987. Werner Rauh: a life in botany. The Euphorbia

Journal 4: 8-10.

SUPTHUT, D. 1999. A short chronicle of 50 years of lOS. lOS Bulletin

1: 6-17.

W. BARTHLOTT* and G.F. SMITH**

* Botanisches Institut der Universitat Bonn, Meckenheimer Alice 170,

D-53115 Bonn, Germany.

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

Bothalia 30,2: 225-227 (2000)

Book Reviews

RARE AND THREATENED PLANTS OE KWAZULU-NATAL AND

NEIGHBOURING REGIONS, by ROB SCOTT-SHAW. 1999.

KwaZulu-Natal Nature Conser\ aticm Serx’ice. P. O. Box 13053, Cascades,

3202 Pietermaritzburg, South Africa. Hard cover: ISBN 0-620-24688-X,

price ZARIOO.OO.

Red Data Lists seem to be the feature of the month at the time this

review is being written, with a project commencing to produce a book

of this format for all ten countries of SABONET (Southern African

Botanical Diversity Network), and involving almost all the scientists of

the National Herbarium, together with many others. This project cer-

tainly has an excellent example in Rob Scott-Shaw’s new book.

After minimal preliminary matter, the meat of the book com-

mences. This is a species-by-species account of the 682 rare and threat-

ened plants known in KwaZulu-Natal. Each report includes an assess-

ment according to the new lUCN guidelines. Endemic status is given,

as are a thumbnail description of the plant, distribution, habitat, popu-

lation biology (if known), rarity, legal status, economic potential,

threats, urgency for conservation, future needs and ’remarks' — in fact,

all that conservationists and decision-makers need to know. After the

tables explaining the lUCN categories of threat, ecosystems and plant

communities, the plants are listed by class of threat, quarter-degree grid

location and names of protected areas. Lists are also given of threat-

ened plant hotspots (centres of endemism), threatened plants in tradi-

tional use and species afforded legal protection. There is an analysis of

trends, a glossary, a bibliography and an index. One additional feature

lifts the present book from the ranks of the merely excellent to the sub-

lime. In the centre of the book are eight pages of colour photographs

(92 pictures) of some of the more attractive species under discussion.

A further 96 species are illustrated with line drawings by Heather

Borchers in the text. An embarrassingly large number of these pictures

illustrate plants never before figured in print.

It would be helpful if the meanings of the sun symbol in each

species text, and the flower symbol in some texts, were described more

explicitly than they are. It seems that the sun symbol precedes coded

notes of bioclimatic regions, biomes or veld types in which the plants

occur. The codes are explained in Table 3 on page 149. and we are

informed in the section on ‘Habitat and ecology', on page ii. of the

existence of such coded information, but nowhere could I find a note

stating that it is marked with a sun sign. Evidently the flower indicates

notes on populations protected in KwaZulu-Natal nature reserves.

Unfortunately this, too, is not made explicit.

There are two points which concern me about this book and the

SABONET Red Data Project. The first concerns the exposure of sensi-

tive data to unscrupulous users. One reads in the introduction to this

book that ’to avoid unscrupulous use of wild plants by illegal plant col-

lectors. precise localities are not generally given". This assurance

seems to be relatively well kept for some plants of high monetary value

(cycads). However, one reads in the discussion of one species of ‘a dis-

junct population in [a named, small] Nature Reserve'. The reserve in

question is traversed by a public road open 24 hours a day, and scan-

dals concerning illegal harvesting of cycads, though frequent, do not

represent more than a small fraction of this illicit trade. My disquiet

increases on seeing maps indicating which quarter-degree square/s the

various species inhabit. Match a good (1:50 000) map — cheaply

obtained from the Government Printer — with the notes in the book on

habitat, add the lack of moral .sense and scruples seen by collectors on

television and ... exit one species? I hope not. But the dilemma persists:

conservationists need this information, and in order to raise funds they

need to demonstrate activity by publishing; but how does one keep the

plants one is supposed to be protecting safe from those who. like a cer-

tain Bruchus. would ’gather golden fruit from the sale of fruitless flow-

ers' (Eerrari 1633; Glen 1991) unhindered by any regard for the law?

The second concerns the choice of taxa. The decision as to what to

include or leave out must necessarily be subjective; the southern

African flora is simply too diverse for any other course to be viable. A

subjective choice of candidates is not necessarily wrong; indeed my

limited knowledge suggests that in almost all cases the author of the

book reviewed here has made the right choice. But in one area I find

his choice inexplicable. The account of Rhizophoraceae is graced by

the inclusion of Bruguiera gymnorrluza and Rhizophora mucronata.

two of the commonest and most widespread trees on the shores of the

Indian and Pacific Oceans. Although both are over-exploited through-

out their range, they are still among the most widespread of mangroves

in KwaZulu-Natal and are given a rating of Lower Risk (conservation

dependent), which is arguably correct. Yet Ceriops tagal (Rhizo-

phoraceae) and Lumnitzera racemosa (Combretaceae), which are

almost as widespread globally as the first-named two species, but only

extend as far as Kosi Bay in KwaZulu-Natal, are not mentioned at all.

Red Data lists are necessarily in a permanent state of flux, as they

reflect ongoing changes in plant populations. Because Red Data List

assessments are based on the most recent available information, it is

important to incorporate new information in future updates. It is a point

of concern that nowhere does this book invite comment for changes

and amendments.

Normally one would end a book review by considering who should

acquire or read the book. In this case I would rather, in view of the

doubts expressed above, ask who should be granted the privilege of

being allowed access to this information. Con.servationists and those

charged with the making and enforcement of conservation laws, each

need at least one copy within easy reach of their workplace — on the

desk or in the police vehicle rather than on the bookshelf. On the other

hand, the information presented here is so good that it needs to be kept

locked, bolted and barred from ’enthusiast growers' like Bruchus in the

Eerrari story referred to above. Congratulations are due to both author

and publisher on an excellent first contribution to a comprehensive

understanding of KwaZulu-Natal’s threatened flora.

REEERENCES

EERRARI. G.B. 1633. De florum cultura libri IV. Stephanus Paulus.

Rome.

GLEN. H.E. 1991. The do-it-yourself mythology of Giovanni Battista

Ferrari, Trees in South Africa 43: 8-11.

H.E. GLEN*

* National Herbarium, National Botanical Institute, Private Bag XlOl,

0001 Pretoria.

TAXONOMY OF CULTIVATED PLANTS, edited by SUSYN

ANDREWS, ALAN LESLIE and CRINAN ALEXANDER. 1999.

Royal Botanic Gardens, Kew. Richmond. Surrey TW9 3AE. England.

Pp. xix + 553, 250 x 158 mm. Hard cover: ISBN 1 900347 89 X, price

GBP 27.00.

Sagan (1997) makes the point that 10 000 years ago, when there

were no cultivated plants and no man-induced weed problems, the

maximum carrying capacity of the world was about 10 million humans.

Now there are over 6000 million people, which means that over 99.9%

of us owe our lives directly or indirectly to cultivated plants. Yet when

I moved into the taxonomy of cultivated plants in 1992, it was consid-

ered something to be done in private between consenting adults, and

certainly not Respectable. The collection of cultivated plant specimens

was PRE's equivalent of a Siberian salt mine, and no decent person

went there voluntarily. Fortunately, times have changed.

Just how much they have changed is shown by tbe volume under

review. It is the proceedings of the third symposium on the taxonomy

of cultivated plants, held in 1998 in Edinburgh, and sponsored by the

Royal Botanic Gardens in Edinburgh and Kew. and the Royal

Horticultural Society. The symposium attracted about 130 delegates

from all around the world, including two South Africans, and was one

of the best-organised and most informative events this reviewer has

ever attended. This excellence is reflected in the proceedings.

The attention to detail and creative thoughtfulness that was such a

feature of the symposium is seen even in the dust-jacket of the pro-

ceedings volume. Dust-jackets are important sales features in modern

publishing, and this one is surely one of the best come-ons that has ever

226

Bothalia 30,2 (2000)

graced a botanical tome. Rows of garden pelargoniums stretch to a

curved horizon (the scope of the proceedings is world-wide), and in the

foreground, on a stake, is a sturdy garden label bearing the title of the

book.

Opening the book does not disappoint. The very first paper is by

H.A. McAllister reminding all and sundry of the importance of living

collections for taxonomy. He includes all plants in his comments, but

curiously omits the most three-dimensional and least amenable to

pressing of all, the mesembs, from his survey. Other authors in this sec-

tion of the proceedings bring to our attention the importance of ex situ

conservation of rai'e and endangered species, and of national collec-

tions of various plant groups. To take but one example, it would trans-

form the identification of waterlilies from a nightmare to a practical

possibility if we had access to a national collection like that described

by B.J. Davies. This section also contains methods which many taxon-

omists would do well to consider. J.D. Twibell describes the value of

vapour profiling in elucidating the taxonomy of Artemisia. But why

should this method be u.sed only by horticultural taxonomists? Ion

Williams (pers. comm.) told a group of which I was a member years

ago that he could identify species of Diosmeae (Rutaceae) in the field

by smell alone, and I am sure that the same is possible in Apiaceae and

Lamiaceae, to name but two obvious examples. Capturing this infor-

mation in a form intelligible to others is problematic, and it seems to

me that the choice lies between following TwibeU’s example or degen-

erating into a form of qualitative ‘winespeak’.

In the present climate where anything not immediately profitable is

likely to disappear forever, it may behove those who wish to preserve

our wild flora by breeding and selling selections, to read the sections on

nomenclature in ornamental plants, intellectual property rights and reg-

istration of plant names. Most of the lawyers contributing to these sec-

tions are American and Swiss, but the principles are governed by inter-

national conventions to which South Africa is a signatory. Mrs Sadie’s

paper on cultivar registration in South Africa shows that the system

operating in this country is a model that other, nominally more

advanced, countries may aspire to emulating.

Naming cultivated plants can be a difficult process. Most ornamen-

tals are surprisingly close to the wild forms of the species to which they

belong, and the problems these present are firstly to persuade the

library to buy the necessary set of world floras, revisions and mono-

graphs to use as tools, and secondly to know which one to use in each

case. But some plants, of which beets and brassicas are outstanding

examples, resemble no known wild plants. How does one attach names

to these? Cultivated-plant taxonomists have not one but two codes

under which to work (Greuter et al. 1994 and Trehane et al. 1995),

which raises the problem of which code applies when (addressed by

R.D. Spencer in this volume). Hetterscheid and his co-workers explore

in three papers the consequences of regarding cultivated plants not as

evolving biological entities but as industrial products to be named as

such. One can see their culton concept working in the numerous man-

made forms of beets, cabbages, kohl-rabi, pak-choy, cauliflower and

similar crops, but Stirton (also in this volume) shows that it does not

tell the whole story in the case of invasive garden escapes such as

Lantana caniara. and this reviewer’s impression is that while the cul-

ton concept has its place in understanding the origin, naming and

derivation of cultigens, it has little if anything to add to one’s under-

standing of the majority of taxa a horticultural taxonomist such as

myself actually sees.

Contributions not mentioned specifically in this review are numer-

ous, and include all the 53 posters, as well as discussions of techniques

in breeding and taxonomy, case studies and Prof. Steam’s invited paper

on early introductions from Japan into European gardens. I cannot,

however, resist the thought that if certain breeders in this country had

ever heard of the necessity of DUS (Distinctness, Uniformity, Stability)

trials before cultivars can be formally registered, the National

Herbarium’s collections would not be cluttered with specimens from

the only plant ever of hybrids named for no discernible purpose other

than vanity.

A large book (over 500 pages) packed with information useful to

many more workers than only horticultural taxonomists, deserves a

long review, and so I make no apology for the length of this one.

Indeed, it would take twice as many words to do justice to the good

things found between these covers. Buyers using southern African cur-

rencies may be horrified at the cover price of this volume, but calculat-

ed per page it is close to the cost of an illicit photocopy (about 50 South

African cents per page at the time of writing). For this, one gets a hefty,

well-printed and excellently bound volume.

The question any review should attempt to answer is, who needs

this book? Horticultural taxonomists, undoubtedly. As 1 have indicated

above, wild-plant taxonomists will also find useful information here on

methods that they would not readily discover elsewhere.

Conservationists need to examine the studies of national collections.

Taxonomists interested in the philosophy of their science will find

much of interest in the writings of the Hetterscheid school here.

Horticulturalists and garden administrators needing to manage a breed-

ing program should examine the property rights section. I am tempted

to say that there is something here for almost everybody who thinks

seriously, on entering a garden, a farm, or a plantation.

Here are the reviewer’s obligatory quibbles, neither of which

detract from the value of the work. First, one could wish for a longer

account of the International Association for Cultivated Plant Taxonomy

(lACPT) than the two lines it received in the introduction. However,

inside information offers the reviewer a suggestion as to why more

extensive coverage of lACPT was probably not possible. Second, is a

certain paper on Lantana in the book really the same as that presented

at the symposium? Both title and abstract differ from those in the sym-

posium handout. Again, I can think of a valid and honourable reason for

any divergence, but cannot help wondering what the limits of diver-

gence between verbal presentation and printed paper should be. In this

case, one is left looking forward to both the next symposium and the

next instalment of Lantana.

In conclusion, one can do no less than to praise Crinan Alexander,

Susyn Andrews and their team for a flawlessly organised symposium

and, to quote Sir Ghillean Prance in the preface: T commend the ... edi-

tors of this volume for producing a milestone publication which will be

of considerable use to all interested in the naming of cultivated plants’.

REFERENCES

GREUTER, 'W., BARRIE, F.R., BURDET, H.M., CHALONER, W.G.,

DEMOULIN, V„ HAWKSWORTH, D.L., J0RGENSEN, P.M.,

NICOLSON, D.H.. SILVA, PC., TREHANE, P. & MCNEILL,

J. 1994, International Code of Botanical Nomenclature {Tokyo

Code). Koeltz, Konigstein.

SAGAN, C. 1997. Billions and billions: thoughts on life and death at

the brink of the millennium. Headline, London.

TREHANE, P, BRICKELL, C.D., BAUM, B.R., HETTERSCHEID,

W.L.A., LESLIE, A.C., MCNEILL, J„ SPONGBERG, S.A. &

VRUGTMAN, F. 1995. International Code of Nomenclature

for Cultivated Plants — 1995. Quarterjack, Wimborne.

H.F. GLEN*

* National Herbarium, National Botanical Institute, Private Bag XlOl,

0001 Pretoria.

OXYANTHUS (RUBIACEAE-GARDENIEAE-GARDENIIAE) EN

AFRIQUE CENTRALE: ETUDE SYSTEM ATIQUE, by B. SONKE.

1999. Opera Botanica Belgica 8. National Botanic Garden of Belgium,

Meise. Pp. 106. Soft cover: ISBN 90-72619-39-0, ISSN 0775-9592,

price 900 Belgian francs.

The genus O.xyanthus, described by De Candolle in 1 807, is endem-

ic to Africa. According to Sonke, Oxyanthiis comprises 35 species, of

which 17 are described in the publication. One species, O. latifolius

Sond., and different infraspecific taxa of O. pyriformis (Hochst.) Skeels

and O. speciosiis DC. (type species of genus) occur in southern Africa

(Herman 1993). Only O. speciosus subsp. stenocarpus known from the

Northern Province and Mpumalanga in South Africa, is found in the

study area covered by the author. Sonke used the difference in infio-

rescence structure as the main character to divide the genus into two

groups. An outstanding feature of the actinomoiphic, pentamerous, her-

maphroditic fiowers is the corolla with tubes long and narrowly cylin-

dric and contorted lobes. There is practically no placenta hull in the

mature dry-walled fruits, a character which is aberrant in the

Gardenieae-Gardeniiae.

The publication presents a classical taxonomic revision, based on

field observations and a study of herbarium material of the genus in

central Africa. It is divided into two main sections. The first section

deals with the history, taxonomic position, anatomy, biology and

chorology of the genus. A formal taxonomic treatment of 17 recognized

central African species of O.xyanthus forms the second section. Two

Bothalia 30,2 (2000)

227

identification keys permitting determination of species by flowering or

fruiting material are given. A concise treatment of each species supple-

mented with information relating to important references, type collec-

tions, synonymy, morphological observations, notes on infraspecific

variability, ecology, distribution, citation and localization of specimens

examined, are given.

Black and white photographs, various line drawings, graphs and

tables are used to explain the different morphological features dis-

cussed in the first part. Three of the 17 species in the taxonomic treat-

ment, are accompanied by accurate line drawings, done by two artists.

Distribution maps are also provided for each taxon described and are

conveniently placed with its description.

A discussion of problematic material and imperfectly known

species, a list of excluded names and a list of the 35 species recognized

for the entire genus Oxyanthus are also provided at the end of the pub-

lication. This work adds to the list of several other publications on the

Rubiaceae that appeared in Opera Botanica Belgica (see Retief 1999),

and is a valuable contribution to our knowledge of Oxyanthus endem-

ic to the African mainland.

REFERENCES

CANDOLLE, A.P. DE. 1807. Oxyanthus. Annales du Museum

National d'Histoire Naturelle 9. Paris.

HERMAN, P.P.J. 1993. Rubiaceae. In T.H. Arnold & B.C. de Wet,

Plants of southern Africa: names and distribution. National

Botanical Institute, Pretoria.

RETIEE, E. 1999. Review; The African species of Ixora (Rubiaceae-

Pavetteae), by P. de Block, 1998. Bothalia 29; 216.

E. RETIEF*

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

Bothalia 30,2: 229-242 (2000)

National Botanical Institute South Africa:

administration and research staff 31 March 2000,

publications 1 April 1999-31 March 2000

Compiler: B.A. Momberg

CHIEF DIRECTORATE

CAPE TOWN

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Laidler, Mrs S.A. Secretary

MARKETING AND COMMUNICATIONS SUBDIRECTORATE

CAPE TOWN

Moeng, S. M.A. (Languages). Deputy Director

Davis, G.W. Ph.D. Assistant Director

Malan, Mrs C.E. B. Sc. (Hons). Principal Communications Officer

Malcolm, Mrs N.J. B.Journ.(Hons). Principal Communications Officer. Media

Mulder, Mrs G.P Secretary/Typist III

Mafu, Ms N.N. Senior Administration Clerk II

Glass, Ms B. IMM Dip. Media Officer (contract worker)

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Davidson, D.C. B.A. (Hons.), PRISA Final Dip. Principal Communications Officer

Loedolff, Mrs J. B.Sc.dnd.Tech.). Senior Industrial Technician. Photographer/Illustrator

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230

Bothalia 30,2 (2000)

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Madikane, M.S. Senior Accounts Clerk I

Mcontsi, Ms N. Accounts Clerk II. Finance

Mirkin, Ms Y.A. Senior Secretary II

Neuwirth, Ms E.V. Senior Accountant. Salaries

Paulse, Mrs D.W.S. Senior Accounts Clerk II

Rawoot, N.A. Internal Auditor

Yeomen, Mrs I.N. Senior Accounts Clerk III. Creditors

GARDENS DIRECTORATE— ADMIN STAFF

CAPE TOWN

Vacant — Director: Gardens

Winter, J.H.S. N.Dip.(Hort.). Deputy Director: Gardens & Horticultural Services

Woodward, Ms YJ. Senior Administration Clerk III. Secretary

Lewis, N.I. Engraver II

Behr, Ms C.M. Curator: Harold Porter NBG

Britz, R.M. Curator: Lowveld NBG

Chaplin, P.J. Curator: Witwatersrand NBG

Heilgendorff, J.P. Curator: Pretoria NBG

Le Roux, PH. Deputy Director. Curator: Kirstenbosch

NBG

Mogale, A.O. Curator: Free State NBG

Oliver, I.B. Curator: Karoo NBG

Tarr, B.B. Curator: Natal NBG

PLANNING, MAINTENANCE & DEVELOPMENT— CAPE TOWN

Linde, D.C. N.T.C.III(Technician and Inspector of Works). M.S.A.I.D. Cert. Estate Agency.

Control Works Inspector

Arendse, D.S. Artisan’s Assistant II. Building maintenance

Manasse, S.P. Dip. (Masonry). Foreman. Building maintenance

Peck, W.I. Artisan’s Assistant I. Building maintenance

HAROLD PORTER NBG— BETTY’S BAY

Behr, Ms C.M. B. Sc. (Hons). Control Technician

Bezuidenhout, Mrs H.M. Senior Administration Clerk III

Forrester, Ms J.A. N.T.C.III(Hort.). Chief Technician. Horticulture

Samuels, Ms D.C. Cleaner II

Bothalia 30,2 (2000)

KAROO NBG— WORCESTER

Oliver, I.B. N.Dip.(Hort.)(PRA). Control Technician

231

Ashworth, Mrs E.H. Senior Administration Clerk III

Mpeke, Ms E.N. Cleaner II

Viljoen, D.M. N.Dip.(Hort.). Chief Technician. Collections

Voigt, W.E. Technician

KIRSTENBOSCH NBG— CAPE TOWN

Le Roux, PH. N.Dip.(Hort.). Deputy Director: Garden. Head: Agricultural Support Services

Goldschmidt, S.M. Assistant Director. Personnel Practitioner. Management

Breedt, C. Control Technician. Estate Manager

Hitchcock. A.N. N.H.Dip.(Hort.). Chief Technician. Supervisor: Nursery

Jaques, R.F. Chief Technician. New plant introductions

Notten, Ms A.L. Chief Technician. Supervisor: Seed room

Trautman, C.E. Artisan. Supervisor: Workshop

Adams, T.D. Greenhouse Supervisor

Arends, Ms S.J. Administration Clerk. Plant records

Coerecius, Mrs R. Senior Administration Clerk III

Crane, Ms A.H. Data Capturer (contract worker)

Crous, H.T. Chief Technician. Tissue culture

Duncan, G.D. N.Dip.(Hort.). Chief Technician. Bulbs

Engelbrecht, Mrs L.D. Technician. Plant records

Geduldt, D.C. Accounts Clerk II. Plant records

Grace, T. Senior Storeman III

Jamieson, Mrs H.G. N. Dip. (Parks & Rec.). Chief Tech-

nician. Restio/Asparagus

Jodamus, Ms N.L. Senior Technician: Annuals

Lawrence, E. Senior Technician: Dell & Ericas

Manuel, I.P. Senior General Eoreman. Seed room

Mathys, Mrs S.S.B. Senior Accounts Clerk III. Gates

Picane, Ms S. Auxiliary Services Officer II. Tissue Culture

Prins, E.B. Security II

Rudolph, A. Security II

Shaide, Ms A.C. Senior Communications Officer. Out-

reach Programme

Smith, Mrs A. Typist II

Solomons, T.C. Security Officer I

Townsend, D.J. N.Dip.(Hort.). Chief Technician. Trees

& shrubs

Van der Walt, Mrs L.E. N.Dip.(Hort.). Chief Technician.

Herbaceous collections

Van Jaarsveld, E.J. M.Sc., N.Dip.(Hort.). Control Techni-

cian. Succulents

Williams, G.C. Gateman. Security II

LOWVELD NBG— NELSPRUIT

Britz, R.M. N. Dip. (Forestry). Control Technican

Froneman, W.C.F. N.Dip. (Nature Cons. & Man.), N.Dip.

(Parks & Rec. Admin.), N.T.C.IIKHort.). Chief Tech-

nician

Hurter, P.J.H. B. Sc. (Hons). Chief Scientific Officer. Re-

search

Maqungo, Ms V.L.B. Accounts Clerk II. Kiosk

Mathebula, Ms N.R. Accounts Clerk I. Kiosk

Van der Walt, Mrs G.A. Senior Administration Clerk III

NATAL NBG— PIETERMARITZBURG

Tarr, B.B. N.Dip. (Parks & Rec. Admin.), Advanced Dip. (Adult Education). Control Technician

Gates, Mrs J.E. N.Dip. (Parks & Rec. Admin.), N.Dip.

(Hort.) Intermed.Dip.(Marketing Man.). Chief Tech-

nician. Kniphofia, forest spp.

Nonjinge, S.H.B. Senior Scientific Officer

Sibiya, Ms C.P.T. Cleaner I

Van der Merwe, Mrs M.E.H. Senior Administration Clerk

III

Zuma, Mrs K.K. Administration Aid II

FREE STATE NBG— BLOEMFONTEIN

Mogale, A.O. Control Technician

Eysele, Mrs J.P Senior Administration Clerk III

Harris, Ms S. Technician

Lumley, M.J. Chief Scientific Officer. Nursery

Raditlhare, Mrs E.M. Administration Aid II

Thaele, Mrs M.E. Administration Aid II

232

Bothalia 30,2 (2000)

PRETORIA NBG

Heilgendorff, J.P. H.N.Dip.(Hort.). Control Technician

Baloyi, K.J. Senior Auxiliary Services Officer II. Garden

records

Baloyi, M.S. Dip. (IBM) Auxiliary Services Officer II.

Garden records

Creighton. Ms D.D. Administration Clerk II

Eyssell, Ms A. B.Sc.(Agric.). Technician. Production

and Sales nursery

Keyter, B.A. Senior Security Officer I

Klapwijk, N.A. N.Dip.(Hoit.), N.Dip.(Plant Prod.), N.Dip.

(Diesel Pitting). Chief Technician. Planning and

development. Index Nursery, New Plant Company

Law, C. Senior Technician

Mariri, Ms M.A. Cleaner I

Myburgh, P. Auxiliary Services Officer II

Ramatsetse, M.P. Security II

Swartz, Ms P.P. M.Sc. Chief Horticulturist. Scientific

and horticultural curation of living collections of

succulents, orchids, medicinal and rare and en-

dangered plants; garden planning and development;

Madagascan plants; tour groups and visitors

Venter, W.A. N.T.C.II. Senior General Poreman. Main-

tenance

WITWATERSRAND NBG— ROODEPOORT

Chaplin, P.J. N.T.C.III(Hort.). Control Technician

Aubrey, Mrs A.E. N.Dip.(Hort.). Senior Horticulturist.

Garden, plant records, seedstore

Hankey, A.J. N.Dip. (Hort.). Chief Horticulturist. Garden,

estate, collections

Head, Mrs S.E. Senior Administration Clerk III

Manjati, Mrs N.L. Accounts Clerk II. Shop Assistant

Mmola, Mrs B.E. Administration Aid II. Cleaner

Ndzondo, Mrs P.G. Administration Aid II. Cleaner

Tebeile, Ms Z.M. Accounts Clerk II. Receptionist

Turner, Ms S.L. B. Sc. (Hons), N.Dip. (Hort.) Chief Horti

culturist. Nursery, garden, information

RESEARCH DIRECTORATE

PRETORIA

Smith, Prof. G.P. Ph.D., F.L.S. Director: Research

Rutherford. M.C. Ph.D., Dip.(Datamet.). Deputy Director: Ecology and Conservation (Cape Town)

Wolfson, Mrs M.M. Ph.D. Deputy Director: Education and Research Support

Meyer, Mrs N.L. B.Sc.(Hons) Technician (contract worker) Steyn, Dr E.M.A. D.Sc. Principal Scientist. Embryology,

Marais, Mrs A.C. Senior Secretary III anatomy, taxonomy

PLANT SYSTEMATICS SUBDIRECTORATE

PRETORIA

Smith, Prof. G.F. Ph.D., F.L.S. Systematics of succulents and rosulate, petaloid monocots

Arnold, TH. Head: Data Management (Pretoria)

Crouch, N.R. Head: Ethnobotany Unit (Durban)

Koekemoer, Miss M. Curator: National Herbarium (Pretoria)

Leistner, O.A. D.Sc. F.L.S. Scientist (contract worker)

Rourke, Dr J.P. Curator: Compton Herbarium (Cape Town)

Williams, Ms R. Curator: Natal Herbarium (Durban)

Willis, C.K. Deputy Director. Regional Project Co-ordinator: SABONET

(Pretoria)

COMPTON HERBARIUM— CAPE TOWN

Rourke, J.P. Ph.D., F.M.L.S., F.R.S.S.Af. Assistant Director. Systematics of

southern African Proteaceae, Stilbaceae

Baatjes, Ms A. Data Capturer (SABONET contract

worker)

Beyers, Mrs J.B.P. M.Sc. Principal Scientist. Assistant

Curator: Collections. Taxonomy of the Gnidieae

(Thymelaeaceae)

Chesselet, Ms P.C.M. M.Sc. Chief Scientific Officer

Conrad, Ms C. (SABONET contract worker)

Cupido, C.N. B. Sc. (Hons). Scientific Officer

Cupido, Mrs C.S. Auxiliary Services Officer II. Techni-

cal Assistant

Bothalia 30.2(2000)

233

Davidse, Mrs E. Auxiliary Services Officer II. Herba-

rium Assistant

Engelbrecht. Ms M. (SABONET contract worker)

Eoster, Mrs S.E. Senior Secretary III

Kurzweil, H. Ph.D. Specialist Scientist. Systematics of

southern African terrestrial orchids

Leith, Mrs J. Senior Administration Clerk II

Manning, J.C. Ph.D. Specialist Scientist. Systematics of

Iridaceae and Orchidaceae; anatomy

Marinus, Ms E.D.A. Principal Auxiliary Services Officer.

Herbarium Assistant

Oliver, E.G.H. Ph.D. Principal Scientist. Taxonomy of

the Ericoideae (Ericaceae)

Oliver, Mrs I.M. (contract worker)

Patenson-Jones, Mrs D.A. (nee Snijman) Ph.D. Principal

Scientist. Systematics of Amaryllidaceae; cladis-

tics

Roux, J.P. N.T.C.III(Hort.), F.L.S., Ph.D. Principal Scien-

tist. Systematics of Pteridophyta

Steiner, K.E. Ph.D. Specialist Scientist. Systematics of

Scrophulariaceae and evolutionary interactions

between oil-secreting flowers and oil-collecting

bees (contract worker)

Williams, Mrs V.J. (SABONET contract worker)

NATAL HERBARIUM— DURBAN

Williams, Ms R. B. Sc. (Hons), H.D.E. Chief Scientific Officer

Arumugam, Ms N. Chironia systematics (SABONET

contract worker)

Crouch, N.R. Ph.D. Principal Scientist. Ethnobotanist

Hlongwane, Mrs C. Administration Aid II. Cleaner &

messenger

Ngwenya. A.M. Principal Auxiliary Services Officer.

Herbarium Assistant. Plant identification, plant

information

Ngcobo, PS. Auxiliary Services Officer II

Noble, Mrs H-E. Senior Administration Clerk III

Ntuli, Mrs N. Data Capturer (SABONET contract work-

er)

Singh, Ms Y. M.Sc., H.E.D. Chief Scientific Officer. Taxon-

omy of Araceae, Hypoxis, plant identifications

Tomalin, Ms M. B. Sc. (Hons), B.Iurius. Data Capturer

(SABONET contract worker)

NATIONAL HERBARIUM— PRETORIA

Koekemoer, Miss M. M.Sc. Assistant Director. Herbarium management.

Taxonomy of Poaceae, Asteraceae: Gnaphalieae

Bredenkamp, Mrs C.L. M.Sc. Principal Scientist. Assistant Curator: Public relations. Taxon-

omy of Vitex, Phylica, Rhamnaceae, Sterculiaceae and other related families

Herman, P.P.J. M.Sc. Principal Scientist. Assistant Curator: Personnel. Taxonomy of Aster-

aceae, Flora of Transvaal

Heymann, Mrs M.Z. T.E.Dip., B. A. (Education & History), B.Ed. Principal Auxiliary Ser-

vices Officer. Assistant Curator: Services, loans, gifts and exchanges

Anderson, H.M. Ph.D. Principal Scientist. Palaeobotany,

palaeogeography

Anderson, J.M. Ph.D. Specialist Scientist. Palaeobotany,

palaeogeography

Archer, R.H. Ph.D. Senior Scientist. Taxonomy of main-

ly Celastraceae, Euphorbiaceae

Archer Mrs C. M.Sc. Senior Scientist. Taxonomy of

Cyperaceae, Restionaceae, Orchidaceae

Burgoyne, Ms PM. M.Sc. Senior Scientist. Mesembry-

anthemaceae

Cloete, Mrs M. Dip. (Typing). Senior Provisioning Clerk

III. Specimen label typist (part time)

Fish, Mrs L. B.Sc. Chief Scientific Officer. Taxonomy

of Poaceae. Plant collecting programme; supervising

mounters

Glen, H.F. Ph.D. Principal Scientist. Taxonomy of trees,

herbarium for cultivated plants, and botanical col-

lectors

Glen, Mrs R.P M.Sc. Chief Scientific Officer. Taxonomy

of ferns, water plants

Jordaan, Mrs M. M.Sc. Chief Scientific Officer. Taxonomy

of Casuarinaceae-Connaraceae, Maytenus

Kgaditsi, W.T. Senior Auxiliary Services Officer. Mount-

er, general assistant in cultivated plants section

Makgakga, M.C. Senior Auxiliary Services Officer. Her-

barium assistant. Wing B

Makgakga, K.S. Senior Auxiliary Services Officer. Mount-

er of vascular plants

Makwarela, A.M. B.Sc. (Hons). Scientific Officer. Wing B

Masombuka, Ms A.S. Auxiliary Services Officer II. Her-

barium assistant

Meyer, J.J. N. Dip. (Teaching). Scientific Officer. Wing C

Mmakola, E.K. Data Capturer (SABONET contract work-

er)

Mothogoane, M.S. Auxiliary Services Officer II. Herba-

rium Assistant

Naicker, K. Sales & Marketing Management Certificate.

Senior Administration Clerk I

Netnou, Ms N.C. B.Sc. (Hons). Scientific Officer. Wing D

Nkoana, L.S. B.Sc. (Hons). Senior Scientific Officer.

SABONET project

Nkoane, Ms G.K. Auxiliary Services Officer II. Herbarium

Assistant. Parcelling, pressing, general assistance

Nkonki, Mrs T. Scientific Officer. Wing B

Perold, Mrs S.M. Ph.D. Taxonomy of Hepaticae (contract

worker)

Phahla, T.J. Senior Auxiliary Services Officer. Mounter

of bryophytes and vascular plants

234

Bothalia 30.2 (2000)

Ready, Mrs J.A. N.D.(Hort.). Senior Auxiliary Services

Officer. Herbarium assistant, Wing D

Retief, Miss E. M.Sc. Principal Scientist. Pollen studies

of Boraginaceae. Taxonomy of Boraginaceae, Ver-

benaceae, Lamiaceae, Asteraceae, Rubiaceae

Riddles, L.M.D. B.Sc. Scientific Officer. Wing A

Sebothoma, P.N. Auxiliary Services Officer II. Plant

identifications co-ordinator

Smithies, Mrs S.J. M.Sc., Dip. Ed. (Moray House). Chief

Scientific Officer. Taxonomy of Scrophulariaceae,

Selaginaceae, Lobeliaceae

Steyn, Ms C.C. Principal Auxiliary Services Officer.

Wing B

Van Rooy, J. M.Sc. Senior Scientist. Taxonomy and

biogeography of mosses; supervising bryophyte

mounter

Van Wyk, E. M.Sc. Scientific Officer. Seedbank mana-

ger, Kew Millenium Seedbank Project (contract

worker)

Victor, Ms J.E. M.Sc., H.Dip.Journ. Senior Scientific

Officer. Taxonomy of Rutaceae, Asclepiadaceae

Welman, Miss W.G. M.Sc. Principal Scientist. Taxonomy

of Convolvulaceae, Solanaceae, Cucurbitaceae,

Campanulaceae, Asteraceae, Acanthaceae

DATA MANAGEMENT— PRETORIA

Arnold, T.H. M.Sc. Assistant Director. Computer application especially in taxonomy

Botha, Mrs A.G. Principal Auxiliary Services Officer.

Secretary (part time)

De Wet, Mrs B.C. B.Sc. (Computer Science), B.A.,

H.D.L.S. Principal Datametrician

Harris, Mrs B.J. Principal Auxiliary Services Officer.

Encoding, quality control

Hawker, Mrs L.C. Scientist (contract worker)

Mbedzi, M.D. Auxiliary Services Officer II

Smit, G.C. Principal Network Controller

Snyman, Mrs E.E. N.Dip.(Comp. Data Proc.) Senior

Scientific Officer

Steyn, Ms H.M. Botanical Information Officer

(SABONET contract worker)

Tleane, Ms L.R. Data Support Officer (contract worker)

Van Rooyen, Mrs V.H. Senior typist

SABONET

PRETORIA

Willis, C.K. M.Sc. (Cons. Biol.). Deputy Director. Regional Project Co-ordinator

Golding, Ms J.S. B.Sc. (Hons). Southern African Red Mossmer, Ms M. B.Sc. (Hons). Editing of publications.

Data List Co-ordinator (contract worker) website management (contract worker)

Haasbroek, Ms C.M. Finances (contract worker) Noko, Ms N.R. Administrative Officer (contract worker)

EDUCATION AND RESEARCH SUPPORT— PRETORIA

Wolfson, Mrs M.M. Ph.D. Deputy Director. Physiology/Ecophysiology of Poaceae, carbon uptake

metabolism, allocation in response to environmental and management stress

Adams, Ms E.M. Adsministration Officer III

Liebenberg, Mrs E.J.L. Manager: Research Support Services and Publications

Potgieter, Mrs E. Principal Librarian

EDUCATION

GOLD FIELDS CENTRE— CAPE TOWN

Louw, Ms G.C. Assistant Director. Communication

Cupido, Ms M. Senior Administration Clerk. Centre co-ordinator

Hitchcock, Mrs W.A. Principal Communications Officer. Adult education

Huet, Mrs H. Senior Administration Officer II

Mgodeli, W.M. Bus driver

Mkefe, T.X. SPTD. Principal Communications Officer

Tyhokolo, Ms S.E. SPTD. Senior Communications Officer

BLOEMFONTEIN

Masilo,T. Education Officer (contract worker)

Bothalia 30,2 (2000)

235

PIETERMARITZBURG

Roff, J. Education Officer

PRETORIA

Symonds, Ms A.M. N. Dip. (Nature Cons.), H.D.E. Assistant Director. Communication

Novellie, Mrs E. Education Officer (contract worker)

Terblanche, Ms A.J. Principal Communications Officer

WITWATERSRAND

Moore, Mrs J.M. Senior Administration Clerk I (contract worker)

Van der Westhuizen, Mrs S. M.Sc.(Bot.). Principal Communications Officer

INTERPRETATION— PRETORIA

Joffe, Mrs H. B.Sc. Chief Garden Utilization Officer

RESEARCH SUPPORT SERVICES AND PUBLICATIONS— PRETORIA

Liebenberg, Mrs E.J.L. M.Sc. Chief Scientific Officer. Cytotaxonomy. Manager

Brink, Mrs S.S. Dip. (Typing). Chief Typesetter. Typeset-

ting, layout, word processing

Condy, Ms G.S. M.A. Chief Industrial Technician. Botan-

ical artist

Du Plessis, Mrs E. B.Sc. (Hons), S.E.D. Technical editor.

Editing, translating, layout

Germishuizen, G. M.Sc. Assistant Director. Editor

Ledwaba, Mrs D.M. Senior Registry Clerk I

Mapheza. T.P. Administration Clerk III. Bookshop

Momberg, Mrs B.A. B.Sc. (Entomology & Zoology).

Technical editor. Editing, layout, ISBN and ISSN

allocations (part time)

Maree, Ms D.J. H.O.D. Computer Operator

Nkosi, PB. Administration Clerk II. Bookstore

Pretorius, Ms M.A. Senior Administration Clerk II

Romanow.ski, Mrs A.J. Dip. (Photography). Chief Indus-

trial Technician (Photography). Scientific photo-

grapher

Tloubatla, J.M. Courier/Photocopy Machine Operator II

Turck, Mrs S. B.A. (Information Design). Senior Indus-

trial Technician. Graphic design

MARY GUNN LIBRARY— PRETORIA

Potgieter, Mrs E. B.Libr. Principal Librarian

Eourie, Mrs A. B.A., H.D.Libr.Sci. Senior Librarian (part time)

ECOLOGY AND CONSERVATION SUBDIRECTORATE

CAPE TOWN

Rutherford, M.C. Ph.D., Dip.(Datamet.). Deputy Director: Research

Hunter, D.A. Senior Administration Clerk III. Personal Assistant to Deputy Director: Research

Parenzee, Ms H.A. Senior Administration Clerk III

Powrie, L.W. M.Sc. Chief Scientific Officer. Spatial modelling, databases

CLIMATE CHANGE

Rutherford, M.C. Ph.D., Dip.(Datamet.). Specialist Scientist. Modelling, global change

Arnolds, J.L. Senior Auxiliary Services Officer. Laboratory

Midgley, G.E. Ph.D. Specialist Scientist. Ecophysiology, modelling

Motete, Ms N. Scientist

Musil, C.F. Ph.D. Specialist Scientist. Ecophysiology, modelling

Snyders, S.G. Auxiliary Services Officer II. Greenhouse, maintenance

36

Bothalia 30,2 (2000)

CONSERVATION BIOLOGY

Donaldson, J.S. Ph.D. (Zoology) Assistant Director. Supervisor: Conservation Farming Project. Cycad biology

Rosenberg, J.de Wet. B. Sc. (Hons) Chief Scientific Officer. Cycad biology, conservation farming

Cader, Ms L. Secretary. Protea Atlas Project (contract worker)

Charlton, Ms V.J. Secretary. Protea Atlas Project (contract worker)

Ebrahim, I. Assistant. Protea Atlas Project (contract worker)

McDonald, D.J. Ph.D. Principal Scientist. Vegetation science

Rebelo, AG. Ph.D. (Zoology) Principal Scientist. Protea Atlas Project

CONSERVATION FARMING PROJECT

Niinni, Ms I. B.Sc; H.D.E. Chief Scientific Officer. Project Co-ordinator

Allsopp, Ms J. (contract worker)

Millar, Ms D.L. (contract worker)

DESERTIFICATION

Hoffman, M.T. Ph.D. Senior Specialist Scientist. Disturbance and historical ecology

Cloete, M.J. Eieldwork (contract worker)

Petersen, Ms A. B.Sc. (Hons). Senior Scientific Officer. Land use and vegetation mapping

Roberts, R.D. (contract worker)

Solomon, Ms A.M. B.Sc. (Hons). M.Sc. student. Impact of fuel wood collection (contract worker)

HORTICULTURAL RESEARCH

Brown, N.A.C. Ph.D. Specialist Scientist. Seed research

Botha, PA. N.H.Dip.(Hort.). Chief Scientific Officer. Seed research

Higgins, S. Ph.D. Scientist. Molecular systematics

Jacobs, E.C. Auxilliary Services Officer

Prosch, D. M.Sc. Seed research (contract worker)

INFORMATION TECHNOLOGY

O’Callaghan, M.G. Ph.D. Information Technology Manager. Information management and development

Evans, N. IT Support Officer (contract worker)

Jamie, Ms N. IT Administration Officer (contract worker)

Pratt, Ms C. Network Controller (contract worker)

SUPPORT SERVICES

Bardien-Overmeyer, Ms S. Manager (contract worker)

Bowler, Mrs M. Administration Aid II. Assistant: teas and functions

De Witt, D.M. Tradesman (B-Group). Assistant: maintenance

HARRY MOLTENO LIBRARY

Reynolds, Ms PY. M.A.dnf.Sc.), B.Proc. Principal Librarian, NBI Web Site Manager

Jagger, B.W. B.A. Senior Library Assistant. Inter-library loans; circulation control

Ovens, Dr C.S.H. Ph.D.dnf.Sc.) Dip.Datametrics (contract librarian)

Bothalia 30.2 (2000)

111

PUBLICATIONS BY THE STAFF

1 April 1999-31 March 2000

ANDERSON, H.M. 1999. Looking for ‘ancient’ plants. PlaiilLife 21 : 40.

ANDERSON, J.M. 1999a. South Africa. In T.R Jones & N.R Rowe,

Fossil plants and spores: modern techniques, Rart 10, Sixty

international laws: collecting, transporting and ownership of

fossils: 328, 329. The Geological Society, London.

ANDERSON, J.M. 1999b. In C. Willis, Challenges facing southern

African botany in the new millenium. SABONET News 4: 185.

ANDERSON, J.M. (ed.). 1999c. Towards Gondwana alive. Promoting

biodiversity and stemming the Si.xth Extinction. National

Botanical Institute, Rretoria.

ANDERSON, J.M., ANDERSON. H.M. & MACRAE, C.S. 1999.

Freezing cold to searing heat. Riant and insect life of the Karoo

Basin. In C. Macrae, Life etched in stone: 1 40-1 66. The Geo-

logical Society of South Africa, Johannesburg.

ANDERSON, J.M., ANDERSON, H.M., ARCHANGELSKY, S.,

BAMFORD, M., CHANDRA, S., DETTMANN, M., HILL, R.,

MCLOUGHLIN, S. & R(DSLER, O. 1999. Rattems of Gon-

dwana plant colonisation and diversification. Journal of African

Earth Sciences 28: 145-167.

ARCHER. C. 1999. Review: Gladiolus in southern Africa, by Reter

Goldblatt & John Manning. 1998. Bothalia 29: 215, 216.

ARCHER, C. & ARCHER, R.H. 1999. A new species of Ornlthogalum

subgenus Urophvllon (Hyacinthaceae) from central South Africa

and southern Namibia. South African Journal of Botanv 65:

431-433.

ARCHER, R.H. 1999. Rare succulent species in the Eastern Cape and

Little Karoo: some preliminary observations. S.4BONET News

4: 35-37.

ARCHER. R.H. & CONDY, G. 1999a. Crinum macowanii. Flowering

Plants of Africa 56: 30-35.

ARCHER. R.H. & CONDY. G. 1999b. Crinum acaule. Flowering

Plants of Africa 56: 36^0.

ASHWELL, A. 1999a. Environmental education in botanic gardens: a

training eourse for botanic gardens educators in Africa. Environ-

mental Education Bulletin 17: 13-15.

ASHWELL, A. 1999b. Cape fox at Kirstenbosch. In N. Malcolm, NBl

news. Veld & Flora 85: 111.

ASHWELL, A. 1999c. Kirstenbosch reflections. In N. Malcolm, NBI

news. Veld & Flora ?>5: 156, 157,

BALARIN, M.G., BRINK, E. & GLEN, H.F. 1999, Itinerary and spec-

imen list of M.A. Rocock’s botanical collecting expedition in

Zambia and Angola in 1925. Bothalia 29: 169-201.

BANGANI, V, CROUCH, N.R. & MULHOLLAND, D A. 1999. Ho-

moisoflavanones and stilbenoids from Scilla nervosa. Phyto-

chemistry 5 1 : 947-95 1 .

BOSENBERG, J.D. 1999. Help save the Albany cycad. Encephalartos

60: 21,22.

BOTHA. D. 1999. In C. Willis, Challenges facing southern African botany

in the new millenium. SABONET News 4: 184.

BOTHA. C.J., SCHULTZ, R. A., VAN DER LUGT, J.J., RETIEF, E. &

LABUSCHAGNE, L. 1999. Neurotoxicity in calves induced by

the plant, Nierembergia hippomanica Miers var. violacea Millan

in South Africa. Onderstepoort Journal of Veterinary Research

66: 237-244.

BRITS, G.J., CALITZ, F.J. & BROWN, N.A.C. 1999. Heat desiccation

as a seed scarifying agent in Leucospenmim spp. (Rroteaceae) and

its effects on the testa, viability and germination. Seed Science &

Technology 21 : 163-176.

BROWN, N.A.C. 1999. The role of fire in enhancing regeneration: the

Cape Floral region. In B.G. Bowes. A colour atlas of plant pro-

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44. Prolea Atlas Project, Kirstenbosch, Cape Town.

REBELO, A.G. & REBELO, P. (eds). 1999c. Protea Atlas Newsletter

45. Protea Atlas Project, Kirstenbosch, Cape Town.

REBELO, A.G. & REBELO, P. (eds) 2000. Protea Atlas Newsletter 46.

Protea Atlas Project, Kirstenbosch, Cape Town.

RETIEF, E. 1999. Review: The African species of Ixora

(Rubiaceae-Pavetteae), by P. de Block, 1998, Bothalia 29: 216.

RETIEF, E. & VAN WYK, A.E. 1999. Taxonomic significance of

pollen apertures in some African Boraginaceae. In K. Heine,

Palaeo-ecology of Africa and the surrounding islands 26.

Proceedings of the Third Conference on African Palynology,

Johannesburg, 14—19 September 1997: 207-218. Balkema,

Rotterdam,

ROHDE, R., HOFFMAN, M.T. & COUSINS, B, 1999. Experimenting

with the commons. Land reform and agrarian change in south-

ern Africa. Occasional Paper Series 12: 1-32. Programme for

Land and Agrarian Studies, School of Government, University

of the Western Cape.

ROURKE, J.P. 1999a. The botanical art of Cynthia Tail. Discovering a

forgotten artist. Veld & Flora 85: 172-174.

ROURKE, J.P. 1999b. Obituary: Elise Bodley (1921-1997). Bothalia

29: 211-213.

ROURKE, J.P. 1999c. A new species of Serruria from the southern

Cape, South Africa (Proteaceae). Bothalia 29: 263-266.

ROUX, J.P. 1999a. Polypodium ensiforme, the correct name for

Microsorum ensiforme (Polypodiaceae: Polypodioideae).

Bothalia 29: 103-107.

ROUX, J.P. 1999b. Review: Flora Malesiana. Series II — ferns and fern

allies, Vol. 3, edited by C. Kalkman & H.P. Nooteboom, 1998.

South African Journal ofBotanv 65: 185.

RUTHERFORD, M.C. 1999. In C. Willis, Challenges facing southern

African botany in the new millenium. SABONET News 4: 181.

RUTHERFORD, M.C., POWRIE, L.W. & SCHULZE, R.E. 1999.

Climate change in conservation areas of South Africa and its

potential impact on iloristic composition: a first assessment.

Diversity and Distributions 5: 253-262.

SCHOLES, R.J., MIDGLEY, G.F, & WAND, S.J.E. 1999. South

African Climate Change Countty Studies: vulnerability and

adaptation of rangelands. National Research Foundation, Divi-

sion of Water, Environment and Forestry Technology, CSIR,

Pretoria. (Mar.).

SINGH, Y. 1999a. Hypoxis. Yellow stars of horticulure, folk remedies

and conventional medicine. Veld & Flora 85: 123-125.

SfNGH, Y. 1999b. In C. Willis, Challenges facing southern African

botany in the new millenium. SABONET News 4: 186.

SMITH, G.F. 1999a. From the President. lOS Bulletin 1: 1,2.

SMITH. G.F. 1999b. Know your Board members: Philip Downs to

head the lOS Conservation Section. lOS Bulletin 7: 3, 4.

SMITH, G.F. 1999c. Recent progress with the Species plantarum: Flora

of the World Project. SABONET News 4: 52-55.

SMITH, G.F. 1999d. The genus Peperomia in southern Africa: the final

words? SABONET News 4: 124.

SMITH, G.F. I999e. In C. Willis, Challenges facing southern African

botany in the new millenium. SABONET News 4: 181.

SMITH, G.F. I999L Documenting plant diversity on a global scale:

recent progress with the Species plantarum: Flora of the World

Project. South African Journal of Science 95: 55, 56.

SMITH, G.F. I999g. Monetary management for the new millenium.

How will biodiversity benefit? In J.M. Anderson, Towards

Gondwana alive: 134, 135. Gondwana Alive Society, Pretoria.

SMITH, G.F. 1999h. Conservatism (more of the same) versus

Liberalism (thinking big): aspects of botany in the new milleni-

um. SABONET News 4: 181-183.

SMITH, G.F. 2000. Plant nomenclature into the 21st centui7: what hap-

pened in St Louis at the Nomenclature Section of the XVIth

Intemational Botanical Congress? South African Journal of

Science 96: 5, 6.

SMITH, G.F., BROWN, N.A.C., BOTHA, D.J., RUTHERFORD,

M.C., DONALDSON, .I.S., DE LANGE, J.H. & DAVIS, G.W.

1999. Horticultural research in the National Botanical Institute

of South Africa: past achievements and future thrusts. South

African Journal of Science 95: 344—348.

SMITH, G.F. & BURGOYNE, P. 1999. Vygie splendour. South African

Gardening, September: 32, 33.

SMITH, G.F., BURGOYNE, P.M., CHESSELET, P. & CONDY, G.

1999. Delosperma cooperi fa. cooperi. Flowering Plants of

Africa 56: 46-52.

SMITH, G.F. & CROUCH, N.R. 1999. Mesembs in the muthi-market:

Lithops lesliei as an ethnomedicinal plant. British Cactus and

Succulent Journal 17: 133-137

SMITH, G.F., CROUCH, N.R. & CONDY, G. 1999a. Aloe pruinosa.

Flowering Plants of Africa 56: 2-6.

SMITH, G.F., CROUCH, N.R. & CONDY, G. 1999b. Rhipsalis bac-

cifera subsp. mauritiana. Flowering Plants of Africa 56: 94—98.

SMITH, G.F, & FORSTER, PI. 1999. Astroloba corrugata: a new

name for an ‘old’ species. Haworthiad 13(4): 130-132.

SMITH, G.F. & GERMISHUIZEN, G. 1999a. Progress with the trial

phase for registration of new plant names. Bothalia 29:

207-209.

SMITH, G.F. & GERMISHUIZEN, G, 1999b. Progress with the trial

phase for registration of new plant names. Forum Botanicum

36,1: 11, 12.

SMITH, G.F., IHLENFELDT, H-D., THIEDE, J., EGGLI, U. & MET-

ZING, D. 1999. The Intemational Organization for Succulent

Plant Study (lOS): its role and potential services to the interna-

tional scientific community. Taxon 48: 715-720.

SMITH, G.F. & RETIEF, E. 1999. Notes on the coastal distribution of

the genus Crassula (Crassulaceae) in southern Africa. Aloe 36:

60, 61.

SMITH, G.F. & STEYN, E.M.A. 1999a. A first record of Agave decip-

iens naturalised in southern Africa. South African Journal of

Botany 65:249-252.

SMITH, G.F. & STEYN, E.M.A. 1999b. Agave vivipara: the correct

name for Agave angustifolia (Agavaceae). Bothalia 29: 100.

SMITH, G.F., STEYN, E.M.a! & BOTHA, D.J, 1999. Gardens of the

North. Our inland national botanical gardens. Veld c& Flora 85:

158-162.

SMITH, G.F., STEYN, E.M.A. & COETZEE, J. 1999. Morpho-

anatomical leaf features of Aloe suzannae Decary

( Asphodelaceae). In .1. Timberlake & S. Kativu, Afi-ican plants:

biodiversitv, taxonomy and uses. Royal Botanic Gardens, Kew.

SMITH, G.F., STEYN, E.M.A. & CONDY, G. 1999. Senecio abbre-

viatus. Flowering Plants of Africa 56: 110-114.

SMITH, G.F. & SWARTZ, P, 1999. Re-establishing X/oe in

Madagascar. Part 3. The next chapter. British Cactus & Succu-

lent Journal 17: 45^9.

SMITH, G.F. & VAN WYK, A.E. 1999a. Endemic succulents compre-

hensively treated in a new book. lOS Bulletin 1: 35, 36.

SMITH, G.F. & VAN WYK, A.E. 1 999b. The Jardin Exotique de Monaco:

one of the great succulent plant gardens of the world. Aloe 36:

55-59.

SMITH, G.F., WALKER, C.C. & CONDY, G. 1999. Senecio row-

leyanus. Flowering Plants of Africa 56: 1 16-120.

SMITH, G.F. & WILLIS, C.K. 1999a. Herbaria are forever: notes on

the Sehlabathebe National Park Herbarium, Lesotho. Aloe 36:

49-51.

SMITH, G.F. & WILLIS, C.K. 1999b. Systematic biologists in South

Africa join forces. South African Journal of Science 95:

156-158.

SMITH, G.F. & WILLIS, C.K. 1999c. Systematic biologists in south-

ern Africa join forces. SABONET News 4: 27-30.

SMITH, G.F. & WILLIS, C.K. 1999d. The Sehlabathebe Herbarium,

survival of the fittest. Plant Talk 19: 38.

SMITH, G.F. & WILLIS, C.K. 1999e. Conquering Crassula in

Lesotho’s Sehlabathebe National Park. SABONET News 4: 1 57,

158.

SMITH, G.F., WILLIS, C.K. & MOSSMER, M. 1999a. SABONET

Report no. 6 published. SABONET News 4: 89.

SMITH, G.F., WILLIS, C.K. & MOSSMER, M. 1999b. Index herbar-

iortim: southern .African supplement, edn 2. Southern African

Botanical Diversity Network Report No. 8..

SMITH, G.F., WILLIS, C.K. & MOSSMER, M. 1999c. Southern

African herbarium needs assessment. Southern African

Botanical Diversity Network Report No. 6.

SNIJMAN, D.A. 1999a. New species and notes on Cyrtanthus in the

southern Cape, South Africa (Amaryllidaceae: Cyrtantheae).

Bothalia 29: 258-263.

SNIJMAN, D.A. 1999b. Growth periodicity, tlowering and phylogeny

of the African Amaryllideae (Amaryllidaceae). In J. Timberlake

& S. Kativu, African plants: biodiversity taxonomy and uses:

1-16, Royal Botanic Gardens, Kew.

SNIJMAN, D.A. 1999c. The discovery of a new species of Amatyllis

(Amaryllidaceae) in the Richtersveld. PlantLife 20: 17, 18.

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SNIJMAN, D.A. 1999d. A new species, notes on subgeneric taxa, and

new synonyms in Hessea ( Amaryllidaceae: Amaryllideae) from

South Africa. Novon 9: 107-110.

SNIJMAN, D.A., MANNING, J.C. & GOLDBLATT, P. 1999. Anew

Rhadamanthus species (Hyacinthaceae) from the northwestern

Cape, South Africa. Novon 9: 111-113.

STEINER, K.E. 1999. A new species of Diascia (Scrophulariaceae)

from the Eastern Cape (South Africa), with notes on other mem-

bers of the genus in that region. South African Journal of

Botany 65: 223-23 1 .

STEYN, E.M.A. 1999. lOPB chromosome data 15. lOPB Newsletter

31: 15.

STEYN, E.M.A. & SMITH, G.F. 1999. Species plantarum: Flora of

the World. Part J. Welwitschiaceae. Australian Biological

Resources Study, Canberra.

STEYN, E.M.A., SMITH, G.F. & HILL, K.D. 1999. Species plan-

tarum: Flora of the World. Part 2. Stangeriaceae: 1-9.

Australian Biological Resources Study, Canberra.

STEYN, E.M.A., SMITH, G.F. & CONDY, G. 1999. Greyia flana-

ganii. Flowering Plants of Africa 56: 86-92.

SWARTZ, P.P. 1999. Green gold of Africa. Envirotour 2, 4: 4-7.

SWARTZ, PP 2000. Woude en hulle omkringende grasvelde Deel 1:

die onbekende Pongola-woud. Veld & Flora 86: 30-32.

SYMMONDS, R. & CROUCH, N. 2000. Propogating the Pepper-bark

tree {Warburgia salutaris): the Silverglen experience. PlantLife

22: 24-26.

TODD, S.W. & HOFFMAN, M.T. 1999. A fence-line contrast reveals

effects of heavy grazing on plant diversity and community com-

position in Namaqualand, South Africa. Plant Ecology 142:

169-178.

VAN JAARSVELD, E.J. 1999a. Biological control of indoor pests.

Environmentally friendly pest control in the Botanical Society

Conservatory. Veld & Flora 85: 130, 131.

VAN JAARSVELD. E.J. 1999b. Indigenous house plants. Bringing

bulbs indoors. Veld & Flora 85: 132.

VAN JAARSVELD, E.J. 1999c. Geckos in the greenhouse. Using rep-

tiles to control garden pests. Veld & Flora 85: 175-177.

VAN JAARSVELD. E.J. 1999d. Indigenous climbers for the garden.

Fe/f/(£F'/ora85: 182, 183.

VAN JAARSVELD, E.J. 1999e. In: Questions and answers. Veld &

Flora 85: 187.

VAN JAARSVELD, E.J. 1999L Daisy chain. SA Gardening,

November: 29-31.

VAN JAARSVELD, E.J. 1999g. Salvia thermara, a new species from

the Western Cape, South Africa (Lamiaceae). Bothalia 29:

100-102.

VAN JAARSVELD, E.J. 1999h. The reintroduction of Gasteria baylis-

siana — a check up. British Cactus and Succulent Journal 17:

119-121.

VAN JAARSVELD, E.J. 1999i. In C. Willis, Challenges facing southern

African botany in the new millenium. SABONET News 4: 1 87.

VAN ROOY, J. 1999. Introduction to bryology in southern Africa. 6.

What to look for in mosses. PlantLife 2 1 : 28-30.

VAN ROOY, J. 2000. Introduction to bryology in southern Africa. 7.

Classification and diversity. PlantLife 22: 35, 36.

VICTOR, J. 1999. In C. Willis, Challenges facing southern African

botany in the new millenium. SABONET News 4: 187, 188.

VICTOR. J.E. & VAN WYK, A.E. 2000 (1999). Pollen morphology of

Adenandra (Rutaceae: Diosminae) and its taxonomic implica-

tions. Grana 38: 1-11.

VICTOR. J.E. & VAN WYK, A.E. 2000 (1999). Pollen morphology of

Diosma and Coleonema (Rutaceae: Diosminae) and its taxo-

nomic implications. Grana 38: 12-19.

VOIGT, W.E. 1999. What flower is that? A soprophytic fungus at

Kirstenbosch. Veld & Flora S5: 167.

WAND, S.J., ESLER, K.J., RUNDEL, PW. & SHERWIN, H.W. 1999.

A preliminary study of the responsiveness to sesonal atmos-

pheric and rainfall patterns of wash woodland species in the

arid Richtersveld. Plant Ecology 142: 149-160.

WAND. S.J.E., MIDGLEY, G.F. & STOCK, W.D. 1999. Predicted

responses of C4 grass-dominated southern African rangelands to

rising atmospheric CO2 concentrations. Vlth International

Rangeland Congress Proceedings 2: 922, 923.

WAND, S.J.E., MIDGLEY, G.F., JONES, M.H. & CURTIS, PS. 1999.

Responses of wild C4 and C3 grass (Poaceae) species to elevated

atmospheric CO2 concentration: a meta-analytic test of current

theories and perceptions. Global Change Biology 5: 723-741 .

WELMAN, W.G. 1999a. Aspects of Ipomoea. The moonflower and the

Commissioner General. Veld & Flora 85: 70, 71.

WELMAN, W.G. 1999b. Eendagskoonblomme. Vier eendagskoon-

blomme in die grasveld van die somerreenvalstreek. Veld &

Flora 85: 72, 73.

WELMAN, W.G. 1999c. Three new records of Solatium section

Oliganthes in southern Africa (Solanaceae). Bothalia 29: 98,

99.

WELMAN, W.G. 1 999d. Notes on Dichondra and Xenostegia in south-

ern Africa (Convolvulaceae). Bothalia 29: 253, 254,

WELMAN, W.G. 1999e. Colophospermum mopane is the correct name

for the ‘mopane’. SABONET News 4: 188-195.

WELMAN, W.G., CRAIB, C. & CONDY, G. 1999. Ipomoea adenioides

var. adenioides. Flowering Plants of Africa 56: 100-104.

WILLIAMS, R. 1999. News from South Africa. Natal Herbarium.

SABONET News 4: 76, 77.

WILLIAMSON, G. & VAN JAARSVELD, E. 1999. Tylecodon nigri-

caiilis (Crassulaceae), a new species from Namaqualand. Aloe

36: 43^5.

WILLIS, C.K, 1999a. Editorial. SABONET News 4: 2, 3.

WILLIS, C.K. 1999b. Editorial. SABONET News 4: 82, 83.

WILLIS, C.K. 1999c. SABONET courses: database and herbarium

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WILLIS, C.K. 1999d. Review: Lepidoptera of southern Africa. Host

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WILLIS, C.K. 1 999e. SABONET — A regional southern African capac-

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WILLIS, C.K. 1999g. SABONET courses: Database and Herbarium

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WILLIS, C.K. & ARNOLD, T.H. 1999. Computerisation of southern

African herbaria. SABONET News 4: 207-2 1 1 .

WILLIS, C.K., BOTHA, D. & WINTER, J. 1999. Southern African

Botanical Gardens needs assessment update. SABONET News

4: 9-22.

WILLIS, C.K., GLEN, H.F. & FISH, L. 1999. The paper chase.

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WILLIS, C.K. & HUNTLEY, B.J. 1999. SABONET— A regional

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WILLIS, C.K. & SMITH, G.F. 1999b. Further progress with updating

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WILLIS, C.K. & SMITH, G.F. 1999c. Threatened knowledge in south-

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WILLIS, C.K., SMITH, G.F. & KOSE, L. 1999. Sehlabathebe National

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2^

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■X ■ ' A/'

Bothalia 30,2: 243-252 (2000)

Guide for authors to Bothalia

This guide is updated when necessary and includes an

index. Important points and latest additions appear in

bold type.

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 Bota-

nical 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 pub-

lished annually.

1 Editorial policy

1.1 Bothalia welcomes original papers dealing with

flora and vegetation of southern Africa and related sub-

jects. Full-length papers and short notes, as well as book

reviews and obituaries of botanists, are accepted. The edi-

tor should be notified that an article is part of a series

of MSS; please submit a list of the parts of a series; all

parts should preferably be published in one journal.

1.2 Submission of a manuscript to Bothalia implies

that it has not been published previously and is not

being considered for publication elsewhere.

1.3 Authors whose first language is not English are

requested to have their MS edited by an English speak-

er before submission.

1.4 Articles are assessed by referees, both local and

overseas. Authors are welcome to suggest possible refer-

ees to judge their work. Authors are responsible for the

factual correctness of their contributions. Bothalia main-

tains an editorial board (see title page) to ensure that

international standards are upheld.

1 .5 Page charges: As stated in our notification includ-

ed in volume 23,1 (May 1993), MSS submitted for pub-

lication in Bothalia are subject to payment of page

charges of R 125,00 per printed page, VAT included. The

following are exempt from these charges; 1, NBl mem-

bers; 2, persons/institutions who have been granted

exemption by the Executive Committee of the NBI; 3,

authors of contributions requested by the Editor; 4, con-

tributors to the column "FSA contributions’. The Editor’s

decision on the number of pages is final. An invoice will

be sent to the author, who must arrange for payment as

soon as possible to NBI, Publications Section, Private

Bag XlOl, Pretoria 0001 .

2 Requirements for a manuscript

2. 1 The original manuscript should be typed on one side

of A4-size paper, double-spaced throughout (including

abstract, tables, captions to figures, literature references,

etc.) and have a margin of at least 30 mm all round. Three

photocopies (all pages photocopied on both sides of the

paper, including figures, to reduce weight for postage) of

all items, including text, line drawings, tables and lists

should be submitted, and the author should retain a com-

plete set of copies. Three photographs (or high quality

photocopies) of each photograph/photograph mosaic

should be submitted for review purposes. If the article

was generated on a computer, a copy of the diskette should

be submitted with the final (accepted) version (see 3).

2.2 Papers should conform to the general style and layout

of recent issues of Bothalia (from volume 26 onwards).

2.3 Material should be presented in the following

sequence: Title page with title, name(s) of author(s), key-

words, abstract (and information that should be placed in

a footnote on the title page, such as address(es) of

author(s) and mention of granting agencies.

2.4 The sequence continues with Introduction and aims.

Contents (see 8), Material and methods. Results, Inter-

pretation (Discussion), Specimens examined (in revi-

sions and monographs). Acknowledgements, References,

Index of names (recommended for revisions dealing with

more than about 15 species). Tables, Captions for figures

and figures. In the case of short notes, obituaries and book

reviews, keywords and abstract are superfluous.

2.5 All pages must be numbered consecutively begin-

ning with the title page to those with references, tables,

captions for figures and figures.

2.6 Eor notes on the use of hyphens and dashes see

3.10 to 3.12.

2.7 Special character; use your own word or code that

is unique and self-explanatory, enclosed between

ANGLE BRACKETS, e.g. <mu>m for pm. Please sup-

ply us with a list of the codes.

3 Requirements for diskettes/stiffies

(to be submitted only with final/accepted version)

3. 1 data must be IBM compatible and written in ASCII,

or in Word 97 for Windows 95. An rtf file is prefere-

able because it retains the formatting.

3.2 the original printout of the diskette should be sup-

plied in double line spacing.

3.3 tables need not be placed on the diskette — a clearly

laid out hard copy is adequate.

3.4 use a non-breaking space to keep two elements to-

gether on the same line, e.g. 3 500.

3.5 DO NOT JUSTIFY LINES.

3.6 do not break words, except hyphenated words.

3.7 all lines, headings, keys, etc., should start flush at

the margin, therefore NO INDENTATIONS, FOOT-

NOTES, TABS OR STYLES of any kind.

3.8 in Word, italics and bold should be used where

necessary.

3.9 paragraphs and headings are delineated by a car-

riage return (ENTER) but no indentation.

3.10 a hyphen is designated as one dash, with no space

between the letter and the dash, e.g. ovate-lanceolate.