Bothalia

A JOURNAL OF BOTANiCAL RESEARCH

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE PRETORIA

Obtainable from the South African National Biodiversity Institute, Private Bag X101, 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 South African National Biodiversity 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 sub-

jects 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 South African National Biodiverity 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.

From 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. Dicroidium, 1983, by J.M. & H.M.

Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium ), 1983, 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 36,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

national

biodiversity

institute

S A N B I

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

Oct. 2006

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Archer, Mrs C. South African National Biodiversity Institute, Pretoria, RSA.

Bredenkamp, Dr C.L. South African National Biodiversity Institute, Pretoria, RSA.

Burrows, J. P.O. Box 710, Lydenburg 1120, RSA.

Cowling, Prof. R.M. University of Port Elizabeth, RSA.

Dold, A. Selmar Schonland Herbarium, Albany Museum, Grahamstown, RSA.

Fish, Mrs L. South African National Biodiversity Institute, Pretoria, RSA.

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

Ihlenfeldt, Prof. H-D. Ostseestr. 40, D-24369 Waabs/Langholz, Germany.

Judd, Dr W.S. University of Florida, Gainesville, USA.

Klak, Ms C. University of Cape Town, RSA.

Leistner, Dr O.A., South African National Biodiversity Institute, Pretoria, RSA.

Linder, Prof. H.P. University of Zurich, Switzerland.

Manning, Dr J.C. South African National Biodiversity Institute, Cape Town, RSA.

Phillips, Dr S.M. The Herbarium, Royal Botanic Gardens, Kew, England, UK.

Pysek, Dr P. Academy of Sciences, Pruhonice, Czech Republic.

Rebelo, Dr A.G. South African National Biodiversity Institute, Cape Town, RSA.

Roux, Dr J.P., South African National Biodiversity Institute, Cape Town, RSA.

Singh, Ms Y. South African National Biodiversity Institute, Durban, RSA.

Snijman, Dr D.A. South African National Biodiversity Institute, Cape Town, RSA.

Stevens, Dr P.F. Missouri Botanical Garden, St Foius, USA.

Van Wyk, Prof. A.E. University of Pretoria, RSA.

Wiland-Syzmanska, Dr J. Department of Geobotany, Adam Mickiewicz University, Pozanan, Poland.

Williams, Dr P. Landcare Research, Nelson, New Zealand.

CONTENTS

Bothalia 36,2

1. Two new species of Spiloxene (Hypoxidaceae) from the northwestern Cape, South Africa. D.A.

SNIJMAN 133

2. New species of Iridaceae from the Hantam-Roggeveld Centre of Endemism, and the Bokkeveld, Northern

Cape, South Africa. J.C. MANNING and P. GOLDBLATT 139

3. Three new species of Lachencilia J.Jacq. ex Murray (Hyacinthaceae: Massonieae) from Western and

Northern Cape, South Africa. G.D. DUNCAN and T.J. EDWARDS 147

4. New species and taxonomic changes within Pentaschistis (Danthonioideae, Poaceae) from Western Cape,

South Africa. C.A. GALLEY and H.P. LINDER 157

5. Notes on African plants:

‘Forsyth ifl. 1835)’: a phantom botanical collector at the Cape of Good Hope explained. E.C.

NELSON 167

Asphodelaceae. Corrections to the eponymy and geographical distribution of Aloe vanrooyenii. G.F.

SMITH and N.R. CROUCH 174

Asteraceae. Arctotis decurrens (Arctotideae), the correct name for A. merxmuelleri and A. scullyi. R.J.

McKenzie, p.p.j. Herman and n.p. barker i 7 i

Malvaceae. A new species of Corchorus in southern Africa. M.J. MOEAHA 168

Mesembryanthemaceae. The identity of Ruschia promontorii. P.M. BURGOYNE 163

Pteridophyta: Pteridaceae. Cheilanthes de/toidea, a new locality in Gauteng, South Africa. R.R.

KLOPPER, P. LEMMER and J. NEL 173

6. Patterns of plant diversity and endemism in Namibia. P. CRAVEN and P. VORSTER 175

7. Phytogeography of Passerina (Thymelaeaceae). C.L. BREDENKAMP and A.E. VAN WYK 191

8. Comparisons of invasive plants in southern Africa originating from southern temperate, northern temperate

and tropical regions. L. HENDERSON 201

9. South African National Biodiversity Institute: administration and research staff 31 March 2006, publica-

tions 1 April 2005-31 March 2006. Compiler: B.A. MOMBERG 223

10. Guide for authors to Bothalia 243

1 1 . Book reviews 253

New species in Bothalia 36,2 (2006)

Corchorus argillicola M.J.Moeaha & P.J.D. Winter, sp. nov., 169

Ixia amethystina J.C. Manning & Goldblatt, sp. nov., 139

Lachenalia cernua G.D. Duncan, sp. nov., 150

Lachenalia lutea G.D. Duncan, sp. nov., 147

Lachenalia nardousbergensis G.D. Duncan, sp. nov., 152

Moraea marginata J.C. Manning & Goldblatt, sp. nov., 141

Pentaschistis clavata C. A. Galley, sp. nov., 159

Pentaschistis horrida C. A. Galley, sp. nov., 160

Pentaschistis trifida C.A. Galley, sp. nov., 157

Romulea singularis J.C. Manning & Goldblatt, sp. nov., 142

Spiloxene nana Snijman, sp. nov., 133

Spiloxene pusilla Snijman, sp. nov., 135

IV

Bothalia 36,2: 133-138 (2006)

Two new species of Spiloxene (Hypoxidaceae) from the northwestern

Cape, South Africa

D.A. SNIJMAN*

Keywords: Hypoxidaceae, new species, South Africa, Spiloxene Salisb., taxonomy, winter rainfall region

ABSTRACT

Newly described are two new species of Spiloxene Salisb.: S. nana Snijman from the Bokkeveld Escarpment, Northern

Cape Province, is a shade-loving plant with narrow, pale green leaves and small, white or rarely cream-coloured flowers; S.

pusilla Snijman from the Matsikamma, Gifberg and Pakhuis Mountains, Western Cape Province, resembles S. nana in habit

but the yellow- or white-tepalled flowers which are tetramerous or hexamerous have darkly coloured stamens and style and an

ovary with a short, solid, narrow prolongation at the apex. Inhabiting rock overhangs formed by quartzitic sandstone sheets,

both species are close allies of 5. scullyi (Baker) Garside from Namaqualand.

With about 30 species, Spiloxene Salisb. is the largest

genus of the family Hypoxidaceae in the Greater Cape

Floristic Region, which corresponds geographically with

the winter and all-year rainfall region of southern Africa.

The genus, as presently circumscribed, was first sepa-

rated from Hypoxis L. by Williams ( 1901 ) who reinstated

the name Ianthe Salisb. that was originally applied to just

three species (Salisbury 1866). Despite its early history,

the genus has nevertheless remained one of the most

poorly studied groups of Cape geophytes. The last com-

prehensive revision of the genus was that of Nel (1914)

who, having taken into account that the letters I and J are

interchangeable in the transliteration of the Greek letter

Tota’, re-introduced the spelling Jan the, which was first

used by Pax (1887) for a section of Hypoxis. Fourcade

(1932) then changed the name to Spiloxene following

the discovery that Ianthe Salisb. (Salisbury 1866) is

a later homonym of Janthe Griseb. (Grisebach 1844),

which has become a synonym of Celsia L. in the family

Scrophulariaceae.

In two recent regional treatments, the accounts of

Spiloxene included an undescribed species, which was

referred to as ‘sp.l’ (Snijman 2000a; Manning et al.

2002). In the short period following the preparation of

these publications, however, new material has been col-

lected that clarifies what was proposed as the new spe-

cies. It has become apparent that what was previously

regarded as one species actually comprises two species

of shade-loving plants that are separated from one anoth-

er geographically. The new species are described and

illustrated here as Spiloxene nana from the Bokkeveld

Escarpment near Nieuwoudtville, Northern Cape, and

S. pusilla from the Matsikamma-Gifberg massif near

Vanrhynsdorp and the Pakhuis Mountains in the northern

Cederberg, Western Cape.

Spiloxene nana Snijman , sp. nov., quoad habitum

et floribus parvis ad S. pusillae Snijman, sed ab ea differt

filamentis e basi libris, antheras et stigmatibus luteolis

(haud rubiginosis). Ab S. scullyi (Baker) Garside foliis

1. 5-5.0 mm (nec 3-14) mm latis, floribus 2 (nec 2-5),

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

MS. received: 2006-04-18.

albis raro cremeis (non luteis), ovario triloculare (non

uniloculare) et placentatio axilis (non parietali) facile

distinguitur. Figura 1.

TYPE. — Northern Cape, 3119 (Calvinia): Oorlogs-

kloof Nature Reserve, 773 m, (-AC), 11-10-200 W.A.J.

Pretorius 589 (NBG, holo.; K, MO, PRE).

Deciduous, entirely glabrous, softly herbaceous, cor-

mous plant, 95-350 mm high. Conns growing horizontal-

ly or geotropically; new corms somewhat globose, 4-10

mm diam., loosely covered by brown tunics; old conns

persisting for a few seasons as withered, flattened discs;

innermost tunics papery, outennost softly fibrous; fibres

fine, reticulate and free from conn base; roots dimorphic,

arising from near base of new conn, slender and spread-

ing, contractile and geotropic. Shoot arising laterally, sur-

rounded at base by a membranous cataphyll up to 45 mm

long. Foliage leaves synanthous, 2-6, 3-ranked in a basal

tuft, sheathing from ± 10-30 mm at base; blades suberect

to slightly arched, nanowly lanceolate, V-shaped in t/s,

35-350 x 1. 5-5.0 mm, the innermost narrowest, pale

green, soft and thin-textured, keeled abaxially almost to

apex; margins entire. Inflorescences 2 or more, erect, a

2-flowered umbel-like raceme; scape 30-150 x 0. 5-2.0

mm, laterally compressed, pale green with membranous

lateral edges; bracts 2, opposite, partially sheathing

pedicels proximally, lanceolate, 10-40 x 3-4 mm, folia-

ceous, shallowly keeled, inconspicuously nerved, pale

green, with translucent margins. Flowers stellate, usu-

ally opening sequentially, unscented; pedicels suberect

at anthesis thereafter spreading horizontally, triangular in

t/s, 35-80 x 1 mm, slender, pale green; tepals 6, reflexed

when fully open, narrowly lanceolate, 2.5— 7.0(— 1 2.0)

mm long, white or rarely cream-coloured, occasionally

flushed pink, backed with pale green in outer whorl,

outer tepals 1.5-2. 5 mm wide and mucronate, inner

tepals 1-2 mm wide and minutely mucronate. Stamens 6,

slightly spreading, unequal, outer whorl slightly shorter

than inner; filaments inserted at base of tepals and joined

to sinus between anther lobes, filiform, 0.5-1 .5 mm long,

yellow; anthers latrorse, oblong, 1. 5-2.0 mm long before

opening, yellow, apical and basal lobes slightly spread-

ing and ± 0.05 mm long; pollen yellow. Ovary inferior,

narrowly tubular to ellipsoidal, 2.5— 8.0(— 1 1 .0) x 0. 7-2.0

mm, 3-locular with axile placentation, narrowed distally

134

Bothalia 36,2 (2006)

FIGURE 1. — Spiloxene nanci: A, B, Pretorius 589; C-F, Rourke 2218. A, whole plant, including t/s leaf; B, corm; C, flower; D, stamens, style

column and stigmatic branches; E, capsule; F, seed. Scale bars; A, 5 mm; B, 2 mm; C, 2.5 mm; D, 1 mm; E, 3 mm; F, 0.2 mm. Artist: John

Manning.

Bothalia 36,2 (2006)

135

into an inconspicuous, solid, ± 0.5 mm prolongation that

sometimes extends up to 1 mm after anthesis; ovules

15-20 per locule; style column cylindrical, 0.3-0. 5

mm long, white; stigmatic branches 3, erect but even-

tually spreading, 1. 2-3.0 mm long, tapering upwards

from broad base, with edges of abaxial surface slightly

folded together, densely papillate on margins and edges

of adaxial surface, white. Capsule narrowly ellipsoidal,

2.5-11.0 x 1. 0-2.0 mm, thin-walled, shedding withered

ovary beak and perigone; dehiscence loculicidal, irregu-

larly longitudinal; placental ridges remaining contiguous

axially; septa disintegrating. Seeds depressed-ellipsoi-

dal, 0.53 x 0.43 mm; funicle stout, attached in chalazal

hemisphere; micropyle prominent; testa brittle, lustrous

black with wicker basket-like ornamentation, consisting

of ± 15 longitudinal rows of closely set, transversally

widened cells; outer periclinal walls each with a central,

conical, raised papilla. Figure 1.

Phenology. Spiloxene nana is spring flowering, mostly

from the middle of September to late October. The leaves

are green during the winter and spring but die back dur-

ing the onset of the dry conditions which extend through

summer and autumn.

Distribution and habitat : the species is known only

from the Bokkeveld Escarpment near Nieuwoudtville,

Northern Cape (Figure 2), where it was first collected

by Mrs L. Bolus in 1930. Surrounded by sheer quartzitic

sandstone cliffs belonging to the Nardouw Formation,

the Escarpment is deeply dissected by the Oorlogskloof

River which flows through a 1 00 m deep gorge. Spiloxene

nana is found on the Escarpment’s southwest-facing

slopes which are relatively moist and cool, unlike the

hotter and drier slopes that face northeast. Several popu-

lations, sometimes comprising as many as 500 plants in

each, have been recorded in and around the Oorlogskloof

Nature Reserve. Individuals are usually massed together

in shallow soil under damp, shaded rock ledges at

730-770 m. Although locally dominant in small patches,

other tiny, shade-loving plants may co-occur, especially

the softly herbaceous, annual species, Crassula strigosa.

FIGURE 2. — Known distribution of Spiloxene nana, •, in the

Northern Cape Province; and S. pusilla, ▲, in the Western Cape

Province.

Diagnostic features : as the epithet suggests, Spiloxene

nana is characterized by slender leaves, 1. 5-5.0 mm

wide, and relatively small flowers ranging from 5— 1 2(—

20) mm across. In addition, the flowers are inconspicu-

ously coloured — white or rarely cream-coloured with a

pink flush, but always with pale green on the backs of the

outer tepals — and the tepals usually reflex downwards

when fully mature. Although the leaves are fairly long

(up to ± 350 mm) they are characteristically soft and pale

green with a delicate appearance.

Its affinities seem to lie primarily with Spiloxene

pusilla, which closely resembles S. nana in habit.

Furthermore, the somewhat membranous leaves and

the thin-textured, pale green foliaceous bracts sug-

gest that S. nana and S. pusilla are allied to S. scullyi

(Baker) Garside, a species found in the granite hills of

Namaqualand and which belongs to Nel’s Aquaticae

group (Nel 1914). Although Nel described the corms of

the five species in this group as lacking persistent fibres

and the disc-like remnants of old corms, those of S. scul-

lyi may sometimes be softly fibrous towards the apex and

may retain a few old discoid corms laterally. Both these

features are shown, for example, in Pearson 6585 (BOL)

and Scully s.n. (Herbarium Normale Austro-Africanum

1381 in SAM). S. scullyi differs from S. nana and S.

pusilla, however, in generally being more robust. It has

broader leaves, 3— 9(— 14) mm wide, and considerably

larger flowers, 1 4— 28(— 38) mm across, which are plain

yellow above. Most importantly, the ovary in S. scullyi

is unilocular with parietal placentation. Like most other

Spiloxene species, the ovary in S. nana or S. pusilla is

trilocular with axile placentation.

The similarities and differences between Spiloxene

nana and S. pusilla are fully discussed below under S.

pusilla.

Other specimens examined

NORTHERN CAPE. — 3119 (Calvinia): stream feeding waterfall

± 10 miles out of Nieuwoudtville, (-AC), L. Bolus ( BOL19597 );

Oorlogskloof trail near Nieuwoudtville, (-AC), Esterhuysen s.n.

(NBG); 9 km SW of Nieuwoudtville on Groot Tuin 653, near hik-

ing trail, (-AC), Helme 3075 (NBG); Oorlogskloof Nature Reserve,

(-AC), W.A.J. Pretorius 279 (NBG), near Eland se Kliphuis, Rourke

2218 (NBG, PRE), Snijman 1865a (NBG, PRE), Annex Uitkomst 797,

± 425 m NE of Dolfontein camp, on a steep SSW-facing slope under a

damp, shaded rock ledge, R.C. Turner 1374 (NBG, PRE); Lokenburg,

under overhanging rocks, (-CA), Acocks 19725 (NBG, PRE); Uitkomst

Farm, SW of Nieuwoudtville, (-CA), W.F. Barker 10730 (NBG); Farm

Driefontein, 18 km S of Nieuwoudtville, (-CA), Goldblatt & Manning

19725 (NBG, PRE).

Spiloxene pusilla Snijman, sp. nov., habitu et tex-

tura S. nanae Snijman similis, praecipue differt filamen-

tis ad stylum connatis, ovario breviter rostrato, antherae

connective) et stylo rubiginoso. Figura 3.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): Drie-

koppen Peak, Gifberg Mtns, on floor of low overhangs,

(-DB), 6-9-1964, Esterhuysen 30747 (NBG, holo.; B,

BOL, K, MO, PRE, Z).

Delicate, deciduous, entirely glabrous, cormous herb,

70-190(-300) mm high. Corms growing horizontally or

geotropically, new corms somewhat globose, 4-7 mm

diam., loosely covered by dark brown tunics, withering

after fruiting and persisting for a few seasons as flat-

136

Bothalia 36,2 (2006)

FIGURE 3, — Spiloxene pusilla, Snijman 1623. A, whole plant; B, conn; C, t/s leaf; D, flower; E, t/s ovary; F, stamens, style column and stigmatic

branches; G, capsule; H, seed. Scale bars: A, 5 mm; B, 2 mm; C, D. G, 1 mm; E, F, 0.5 mm; H, 0.2 mm. Artist: Claire Linder Smith.

Bothalia 36,2 (2006)

137

tened discs; innermost tunics papery, outermost softly

fibrous; fibres reticulate and free from corm base; roots

dimorphic, arising from base of new corm, slender and

spreading, contractile and geotropic. Shoot arising lat-

erally, surrounded at base by a membranous cataphyll

up to 10 mm long. Foliage leaves synanthous, 2-6,

3-ranked in a basal tuft, sheathing for up to 10 mm at

base; blades suberect to slightly arched, narrowly lanceo-

late, V-shaped in t/s, 45—1 60(— 300) x 1.0-2. 5 mm, pale

green, soft and thin-textured, keeled abaxially almost to

apex; margins entire. Inflorescences 2 or more, erect, a

2(rarely 3)-flowered umbel-like raceme; scape 20-100

x 0.5-1. 5 mm, laterally compressed, sharp-edged, pale

green; bracts 2(3), opposite, partially sheathing pedicels

proximally, lanceolate, 15-25 x 1-3 mm, foliaceous,

shallowly keeled, inconspicuously nerved, pale green,

with translucent margins. Flowers stellate, usually open-

ing sequentially; pedicels suberect at anthesis and there-

after, terete, 20— 50(— 80) mm long, slender, pale green;

tepals 4 or 6, rarely 5, reflexed when fully open, narrowly

lanceolate, 3^1 mm long, yellow or white, backed with

pale green mostly in outer whorl, outer tepals 1 .5 mm

wide and mucronate, inner tepals 1.0-1. 2 mm wide and

minutely mucronate. Stamens as many as tepals, suberect

proximally to slightly spreading distally, unequal, outer

whorl slightly shorter than inner; filaments joined to

sinus between anther lobes, filiform, 1.4-2. 2 mm long,

yellow, maroon-tipped, outer whorl adnate to style base

for up to ± 0.5 mm. inner whorl adnate to style for up to ±

1 mm; anthers latrorse, oblong, 1.0-1. 4 mm long before

opening, apical and basal lobes ±0.17 mm long and

slightly spreading; thecae and pollen yellow; connective

dark red. Ch’ary > inferior, narrowly ellipsoidal, 2. 3-3.0

x 0.8-1. 2 mm, 2- or 3-locular with axile placentation,

narrowed distally into a solid, 0. 5-1.0 mm prolongation

that lengthens up to 1.5 mm after anthesis; ovules 15-20

per locule; style column cylindrical, 1.5 mm long, yel-

low; stigmatic branches 2 or 3, erect, 0.9-1. 5 mm long,

slightly broader than style column, with edges of abaxial

surface slightly folded together, densely papillate on

margins and edges of adaxial surface, dark red. Capsule

narrowly ellipsoidal, 2. 5-5.0 x 1.0-1. 5 mm, thin-walled,

partially topped with withered, persistent ovary beak and

perigone; dehiscence irregularly apical and longitudinal;

placental ridges remaining contiguous axially; septa

disintegrating. Seeds ovate, ± 0.5 x 0.4 mm; raphe con-

tinuous with a short, straight, persistent funicle attached

in chalazal hemisphere; micropyle slightly raised; testa

brittle, shiny black, of transversally widened cells, orna-

mented with ± 12 widely spaced, longitudinal ribs; outer

periclinal walls each with a central, blunt, raised papilla;

papillae closely aligned giving ribbed appearance. Figure 3.

Phenology r. flowering extends throughout spring from

early September to mid-October. The leaves remain

green for slightly longer until the onset of the summer

dry period, when they are shed. New leaves emerge each

year with the onset of winter rain which often starts to

fall in autumn. Most species of Spiloxene depend on

sunshine to open their flowers (lasting from ± 11:00 to

± 16:00), but those of S. pusilla are able to open in the

shade and to remain open throughout the day.

Distribution and habitat : Spiloxene pusilla is restrict-

ed to the northwestern Cape Fold Mountains (Figure

2), where populations are found on the relatively moist,

cool summit of the Gifberg and Matzikamma massif,

near Vanrhynsdorp, and on the Pakhuis Mountains east

of Clanwilliam, at 460-760 m. These mountains have

large areas of exposed, flat, quartzitic sandstone sheets

that are often waterlogged in winter and are extremely

dry in summer. The plants shelter under south-facing,

overhanging rocks in shallow, sandy soil, where they

escape the severe conditions of the exposed rock sheets.

5. pusilla is often the sole inhabitant of these cool, shady,

shelf-like habitats, where it forms dense mats having the

appearance of soft, pale green turf.

Diagnostic features : one of the most unusual features

of this species, and perhaps the most important reason

why it has remained unnamed for so long, despite sev-

eral collections having been made since its discovery by

E.P. Phillips in 1911, is the fusion of the inner filaments

to the style. Handwritten notes by the late Ms M.F.

Thompson (Mrs Rand), author of several papers on Cape

Hypoxidaceae (Thompson 1969, 1976a, b, 1978, 1979),

suggest that she regarded the fusion of the filaments to

the style to be sufficiently significant to delay formally

describing the species until its relationships were more

clearly understood. Fortunately, the availability of many

more collections of the newly described and apparently

closely related S. nana has made this description pos-

sible.

Morphologically Spiloxene pusilla bears a striking

resemblance to S. nana. Both are shade-loving plants

with delicate, narrow, pale green, extremely soft-textured

leaves and small flowers, often only 1 0 mm in diameter.

In addition, both species inhabit rock overhangs where

the plants form dense, turf-like patches.

The particular features that separate Spiloxene pusilla

from S. nana are those of the flower. The inner filaments

are shortly fused to the style (versus free from the style);

the anther connectives and stigmatic branches are dark

red (as opposed to yellow); and the ovary is constricted

into a short (0. 5-1.0 mm) but distinct solid, neck-like

prolongation at anthesis (compared with an indistinct

prolongation of up to ± 0.5 mm that sometimes lengthens

slightly after anthesis).

Elsewhere in the family a solid prolongation of the

ovary, most often referred to as the ovary beak, is found

in Empodium Salisb., Curculigo Gaertn., Molineria Colla,

Rhodohypoxis Nel and Saniella Hilliard & B.L.Burtt

(Hilliard & Burtt 1978), and although infrequent in

Spiloxene , it is well developed in S. alba (Garside 1950).

Moreover, the darkly coloured stamens and style, in

otherwise yellow or white flowers, are only known in

Spiloxene , particularly in the southwestern Cape species,

S. canaliculata Garside, S. capensis (L.) Garside and S.

serrata (Thunb.) Garside, which belong to a guild of

beetle pollinated plants (Steiner 1998). Undoubtedly the

flowers of S. pusilla are too small to support monkey

beetles, but their dark filaments and stigmas may attract

specific, albeit tiny, pollinators — an aspect which is yet

to be studied. Records show that this dark coloration is

constant throughout the distribution range but that popu-

lations on the Matsikamma Mountain and Gifberg con-

sistently have yellow tepals, whereas those in the north-

ern Cederberg have white or cream-coloured tepals.

138

Bothalia 36,2 (2006)

Although Figure 3 shows a plant with tetramerous

flowers, this is not constant in the populations, and

hexamerous flowers also frequently occur, occasionally

even on the same plant. Only very rarely has a pentamer-

ous flower been noted. This variation in the number of

floral parts is not unique and is also known in species of

Hypoxis and Empodium.

Lastly, the only other member of Hypoxidaceae that

has parts of the androecium fused to the style is Pauridia

Harv., a small Western Cape genus of two species.

Pauridia , however, is characterized by tepals that are

joined below to form a cup-shaped or tubular perigone;

an inner whorl of fertile stamens which is inserted in the

perigone throat; and an outer whorl of staminodes that

is fused to the style. Moreover, the stigmatic branches

are slender and distinctly shorter than the style column.

This is in contrast to Spiloxene pusilla, in which all six

stamens are fertile, the flowers are stellate, and the stig-

matic branches are distinctly broader and longer than the

style column. These features, together with the cormous

habit, are common to all the currently known species of

Spiloxene, which suggests that this northwestern Cape

species is best placed within Spiloxene.

Despite clear evidence from leaf anatomy and rbcL

sequence data that Spiloxene and Pauridia are closely

related (Thompson 1976a; Hilliard & Burtt 1978; Nordal

1998; Rudall et al. 1998; Burtt 2000; Judd 2000), both

genera have been consistently retained in local floras

(Garside 1950; Snijman 2000a, b). Should future studies

reveal, however, that Spiloxene and Pauridia comprise

one monophyletic genus, it nevertheless seems likely,

given the different relative positions of the structures

involved, that the fusion of filaments to style in S. pusilla

is not homologous with the fusion of staminodes to style

in P. minuta (L.f.) Durand & Schinz and P longituba

M.F.Thomps.

Other specimens examined

WESTERN CAPE. — 3118 (Vanrhynsdorp): Matsikamma Mtn, Op

de berg 314, Dreyers Kloof, (-DB), Helme 1358 (NBG); Matsikamma,

Farm Sewefontein, (-DB), Snijman 1623 (NBG, PRE), Snijman 1860

(NBG); Gifberg, (-DD), Compton 20847 (NBG), Esterhuysen 22062

(BOL, NBG, PRE), E.P Phillips 7569 (SAM); summit of Gifberg

Pass, near De Kom, (-DD), Snijman 1864 (NBG). 3219 (Wuppertal):

Pakhuis, N Cederberg, (-AA), Esterhuysen 21722 (BOL, NBG),

Esterhuysen 21910 (BOL, NBG, PRE); N Cederberg, Diagonal Kloof

N of Ribbokberg, (-AA), H.C. Taylor 11869 (NBG).

ACKNOWLEDGEMENTS

1 am most grateful to Claire Linder Smith and John

Manning for the accompanying illustrations and to Nick

Helme, Colin Paterson-Jones, Wessel Pretorius, Dave

and Betty Schlebusch, and Ross Turner for their kind

assistance in the field. The Northern Cape Department of

Nature and Environmental Conservation and the Western

Cape Nature Conservation Board are thanked for grant-

ing permits to collect specimens of the above species.

REFERENCES

BURTT, B.L. 2000. Saniella and its relation to other South African gen-

era of Hypoxidaceae. Edinburgh Journal of Botany 57: 63-70.

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

neighbouring divisions. Transactions of the Royal Society of

South Africa 21 : 75, 76.

GARSIDE, S. 1950. Spiloxene. In R.S. Adamson & T.M. Salter, Flora

of the Cape Peninsula : 214-217. Juta, Cape Town.

GRISEBACH, A.H.R. 1844. Spicilegium florae rumelicae et bithym-

icae 1 . Vieweg, Braunschweig.

HILLIARD, O.M. & BURTT, B.L. 1978. Notes on some plants from

southern Africa chiefly from Natal: VII. Notes from the Royal

Botanic Garden Edinburgh 36: 43-76.

JUDD, W.S. 2000. The Hypoxidaceae in the southeastern United States.

Harvard Papers in Botany 5: 79-98.

MANNING, J„ GOLDBLATT, P. & SNIJMAN, D. 2002. The color

encyclopedia of Cape bulbs. Timber Press, Oregon and Cam-

bridge.

NEL, G. 1914. Die afrikanischen Arten der Amaryllidaceae-Hypoxi-

deae. Botanische Jahrbiicher 5 1 : 287-340.

NORDAL, I. 1998. Hypoxidaceae. In K. Kubitzki, The families and

genera of vascular plants 111: flowering plants — Monocotyle-

dons Lilianae (except Orchidaceae): 286-294. Springer, Berlin.

PAX, F. 1887. Amaryllidaceae (Hypoxidoideae-Hypoxideae). In A.

Engler & K. Prantl, Die natiirlichen Pftanzenfamilien, Teil 2,

Abteiling 5: 121, 122. Engelmann, Leipzig.

RUDALL, P.J., CHASE, M.W., CUTLER, D.F., RUSBY, J. & DE

BRUIJN, A.Y. 1998. Anatomical and molecular systematics of

Asteliaceae and Hypoxidaceae. Botanical Journal of the Lin-

nean Society 127: 1 — 42.

SALISBURY, R.A. 1 866. The genera of plants: a fragment containing

part of Liriogamae. Van Voorst, London.

SNIJMAN, D.A. 2000a. Hypoxidaceae. In P. Goldblatt & J.C. Man-

ning, Cape Plants. A conspectus of the Cape flora of South Af-

rica. Strelitzia 9: 108-110. National Botanical Institute, Pretoria

& Missouri Botanical Garden, St. Louis.

SNIJMAN, D.A. 2000b. Hypoxidaceae. In O.A. Leistner, Seed plants

of southern Africa: families and genera. Strelitzia 10: 621-623.

National Botanical Institute, Pretoria.

STEINER, K.E. 1998. Beetle pollination of peacock moraeas (Iridaceae)

in South Africa. Plant Systematics and Evolution 209: 47-65.

THOMPSON, M.F. 1969. Spiloxene capensis and S. canaliculata. The

Flowering Plants of Africa 39: t. 1557A & B.

THOMPSON, M.F. 1976a. Studies in the Hypoxidaceae. I. Vegetative

morphology and anatomy. Bothalia 12: 111-117.

THOMPSON, M.F. 1976b. Empodium namaquensis. The Flowering

Plants of Africa 44: t. 1727.

THOMPSON, M.F. 1978. Studies in the Hypoxidaceae. II. Floral mor-

phology and anatomy. Bothalia 12: 429—435.

THOMPSON, M.F. 1979. Studies in the Hypoxidaceae. III. The genus

Pauridia. Bothalia 12: 621-625.

WILLIAMS, F.N. 1901. On Ianthe, a genus of Hypoxidaceae. Journal

of Botany 39: 289-294.

Bothalia 36,2: 139-145 (2006)

New species of Iridaceae from the Hantam-Roggeveld Centre of

Endemism, and the Bokkeveld, Northern Cape, South Africa

J.C. MANNING* and P. GOLDBLATT**

Keywords: Iridaceae, Ixia amethystina J.C. Manning & Goldblatt, Moraea marginata J.C. Manning & Goldblatt, new species, Romulea singularis

J.C. Manning & Goldblatt, South Africa, taxonomy

ABSTRACT

Three new species of Iridaceae are described from the Bokkeveld and Roggeveld Escarpments. Ixia amethystina, a

member of section Dichone, is endemic to the edge of the Roggeveld Escarpment. It shares an unusual, inclined spike that is

nodding in bud with I. trifolia but is distinguished by its blackish purple (not yellow) anthers, narrower leaves 1. 5-2.0 mm

wide, medium-textured corm tunics that form a distinct neck at the base of the stem, and short style branches 2. 0-2. 5 mm long.

Moraea marginata, another Roggeveld endemic, is a member of section Polyanthes and florally similar to M. fistulosa and

M. monticola but differs in its linear, channelled leaves 5-7 mm wide, with unusual, thickened margins. Romulea singularis,

from the edge of the Kobee River Valley in the Bokkeveld Mountains, is a member of section Ciliatae. It is unique in the

genus in its narrowly funnel-shaped, mauve to purple flowers with slender perianth tube 10-11 mm long, and unusually long

filaments, 8-9 mm long, inserted in the lower half of the tube.

INTRODUCTION

Plant geographers have been aware of the high levels

of endemism in the flora of the winter rainfall region

along the western margin of the Upper Karoo for almost

a century, culminating in its recent recognition as the

Hantam-Roggeveld Centre of Endemism (Van Wyk

& Smith 2001). The Hantam-Roggeveld occupies the

high-lying southwestern portion of the South African

interior plateau and includes the Hantams, Roggeveld

and Nuweveld Mountains. The vegetation is primarily

succulent karroid shrubland, with renosterveld at the

higher elevations and in moister areas. The region is

exceptionally rich in geophytes, which may account for

nearly 40% of the flora in some places (Snijman & Perry

1987). The richest and most interesting geophyte flora

is found along its western rim, on the main Roggeveld

Escarpment and on the Hantamsberg, which receives

more rain than the surrounding country. Among the

geophyte flora are some 90 species of Iridaceae, many

endemic to the region (Manning et al. 2002).

The Bokkeveld Mountains, which border the Hantam-

Roggeveld in the northwest, constitute the northern limit

of the Cape Floral Region (Goldblatt & Manning 2000).

This sandstone escarpment supports dry fynbos vegeta-

tion, with a narrow belt of renosterveld along its interior

margin. Although not recognized as a separate centre of

endemism by Van Wyk & Smith (2001), the region is a

centre of diversity for several geophyte genera (Manning

et al. 2002), and its flora includes numerous endemic

Iridaceae (Manning & Goldblatt 1 997).

Here we describe two new species from the Roggeveld-

Hantam plateau, Ixia amethystina and Moraea margina-

ta, and a third, Romulea singularis, from the Bokkeveld

Mountains. All three species appear to be most closely

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

** B.A. Krukoff Curator of African Botany, Missouri Botanical

Garden, P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2006-04-11.

allied to other endemics from their respective regions,

emphasizing the importance of local radiation in the

floras of these two regions. Two of these species, M.

marginata and R. singularis, were brought to our atten-

tion by botanist and ardent plant enthusiast, Nick Helme,

who has been responsible for the discovery of several

new species in the Cape region in the past few years

(Manning & Goldblatt 2001a; Van Jaarsveld & Thomas

2003; Goldblatt 2004).

Ixia amethystina J.C. Manning & Goldblatt, sp.

nov.

Species habitu floribusque Ixia trifolia G.J. Lewis

similis sed floribus pallide purpureis centro tuboque

atropurpureis, antheris atropurpureis, ramis stylorum

inconspicuis filamentis brevioribus 2. 0-2. 5 mm longis,

et foliis angustioribus 1. 5-2.0 mm latis differt.

TYPE. — Northern Cape, 3220 (Sutherland): west of

Farm Agterkop, near the top of Gannaga Pass, (-AA), 16

September 1997, P Goldblatt & J.C. Manning 10745 A

(NBG, holo.; MO, iso.).

Plants 150-300 mm high. Corms globose, 12-20

mm diam.; tunics of medium-textured, wiry, reticulate

fibres, extending up in papery neck 30-70 mm long.

Stem erect, unbranched or with up to two branches that

are erect below and then inclined, 1.0-1. 5 mm diam.

below base of main spike. Cataphylls submembranous,

flushed reddish brown, upper one reaching above ground

level and then papery and dark reddish brown. Leaves

3 or 4, all basal, uppermost completely sheathing lower

two thirds of stem, remainder suberect or lowermost

slightly falcate, 1 .5-2.0 mm wide, reaching to near top of

stem, firm-textured, margins and midrib hyaline, slightly

thickened, plane or slightly twisted in upper half, with

an additional one or two membranous, scale-like leaves

in upper third of stem, in axils of which lateral branches

may develop. Spike inclined, crowded, 5-7-flowered,

branches 1^1-flowered, inflorescences nodding in bud;

bracts scarious, translucent or flushed purple above.

140

Bothalia 36,2 (2006)

outer 5-7 mm long, acute or obscurely three-dentate,

inner bracts about as long as outer or slightly shorter,

bicuspidate, margins connate in lower 1.5 mm around

ovary. Flowers rotate, pale purple with small, dark purple

eye, faintly scented; perianth tube filiform and clasping

style for entire length, 2. 0-2. 5 mm long, widened only in

upper 1 mm; tepals obovate, somewhat narrowed below

into short claw, spreading and slightly cucullate distally,

12-13 x 7-8 mm. Filaments inserted at apex of tube

and occluding throat, blackish purple, free, diverging

above, 2.5 mm long; anthers oblong-sagittate, 4. 5-5.0 x

1.5 mm, erect, thecae narrowly separated by connective

and dehiscing laterally by narrow slits extending length

of thecae, blackish purple; pollen creamy yellow. Ovary

ovoid, 2. 5-3.0 mm long; style straight and erect, ± 2 mm

long, dividing at or just below mouth of tube, branches

involute-filiform and stigmatic at apex only, purple,

arching outward, 2. 0-2. 5 mm long. Flowering time : late

September to early October. Figure 1A-F.

Distribution and ecology: Ixia amethystina is known

from a small area southwest of Middelpos on the edge of

the Roggeveld Escarpment (Figure 2). The three known

collections were made within a few kilometres of each

other on the Farms Zoekop and Agterkop. Plants grow

in stony clay in renosterveld ( Elytropappus rhinocerotis)

FIGURE 1 . — Ixia amethystina,

NBG 192794 : A, conn and flo-

wering stem; B, flower, three-

quarter view; C, flower, side

view; D, floral bracts, outer

(left) and inner (right); E, sta-

mens and style branches; F,

stamen, abaxial view. Ixia tri-

folia, Helme 1662 (NBG): G,

flower, three-quarter view; H,

stamens and style branches.

Scale bars: A-C, G, 10 mm;

D, 5 mm; E, F, H, 2 mm.

Artist: John Manning.

Bothalia 36,2 (2006)

141

FIGURE 2. — Distribution of Ixia amethystina, •; Moraea margina/a,

O; and Romulea singiilaris , V

shrubland. I. amethystina appears to be less sensitive to

light and temperature than other species in the subgenus.

The flowers open widely around 08:00, even in cool,

overcast conditions and remain fully open until late

afternoon, whereas those of I. trifolia , like many other

species of section Dichone, will not open fully under

cooler conditions.

Diagnosis and relationships : Ixia amethystina is

an attractive species distinguished by its pale purple

flowers with small, dark eye, relatively broad, blackish

anthers that dehisce laterally so that the pollen forms a

contrasting pale margin to each anther, and short style

branches that are subequal to the filaments in length,

thus scarcely projecting from between the stamens. The

lovely amethyst-coloured flowers are borne on inclined

spikes so that they face directly upward in an elegant,

arching spray.

The filiform perianth tube clasping the style for its

whole length and the involute-filiform style branches

stigmatic only at the extreme apex, place Ixia amethys-

tina in section Dichone of subgenus Ixia (Goldblatt &

Manning 1999). Here it falls among a small group of spe-

cies that are endemic or near-endemic to the Roggeveld

Escarpment, including I. brevituba G.J. Lewis, I. trifolia

G.J.Lewis and 7. curvata G.J. Lewis. These species

share relatively unspecialized, longitudinally dehiscent

anthers, 4-5 mm long. The species of section Dichone

from the southwestern Cape below the Escarpment, in

contrast, fall into two groups defined by their derived

stamens, the one characterized by its very short anthers,

2-4 mm long and the other by their curious attachment,

resulting in their reclinate orientation.

Among the Roggeveld species of section Dichone,

Ixia amethystina appears to be most closely allied to

I. trifolia (Figure 1G, H) on the basis of their unusual,

inclined spikes, and lateral branches that are decurved

and nodding when young. All other species have spikes

that are erect from bud to fruit. Although not explicitly

mentioned in the original description of I. trifolia (Lewis

1962), this feature was later noticed and illustrated by

De Vos (1999). The two species both have a perianth

tube 2-3 mm long, which is longer than in I. brevituba

( 1 .0-1.5 mm) but shorter than in I. curvata (3-5 mm). I.

amethystina differs from I. trifolia in its blackish purple

anthers, purple perianth tube, which gives the flowers

a small, dark, central eye, consistently narrower leaves

1. 5-2.0 mm wide, and medium-textured corm tunics

drawn into a well-developed neck. 7. trifolia, like all

other species in section Dichone, has yellow anthers,

although the filaments may be pale mauve, and a pale

perianth tube, giving the flowers a well-defined, whit-

ish eye. The observation by Lewis (1962) and later De

Vos (1999) that 7 trifolia may occasionally have a dark

eye is not corroborated by examination of living plants.

The pale eye in this species is usually surrounded by

darker shading, which in pressed specimens may give

the eye a dark appearance. The leaves of 7 trifolia are

broader than in 7 amethystina, (2.5-)5.0-12.0 mm wide,

and the tunics are more coarsely fibrous, with the lower

fibres developed into woody claws but not drawn into a

neck above. In addition, the style branches in 7 trifolia

are longer than the filaments, 4-5 mm long, and project

conspicuously between them.

The unusual pale purple of the flowers of Ixia

amethystina occurs occasionally in both 7 trifolia and

7 brevituba (Lewis 1962) although the flowers in these

two species are more usually bright pink. Other species

of section Dichone invariably have pale to deep pink

flowers. 7 trifolia is more widely distributed along the

Roggeveld Escarpment than 7 amethystina and has been

collected from several places along the escarpment from

Uitkyk in the north to Komsberg in the south and thence

to Laingsburg and Tweedside below the escarpment. The

two species are parapatric, with 7 amethystina occur-

ring immediately to the north of the range of 7 trifolia.

The differences in their flowers appear to be related to

their pollination biology. 7 amethystina has flowers that

conform to the hopbine beetle pollination syndrome

(Goldblatt et al. 1998) and the beetle Kubousia axillaris

Burmeister has been collected on the flowers. All indi-

viduals of this insect carried pure loads of Ixia-type pol-

len on their dorsal thorax and frons (unpublished data).

In contrast, 7 trifolia is pollinated by solitary female

anthophorine bees (unpublished observations).

Etymology, named for the distinctive colour of the

flowers.

Other material examined

NORTHERN CAPE. — 3220 (Sutherland): Zoekop Farm, 3 km S

of entrance along road to Gannaga Pass, (-AA), 26 September 2002,

NBG192794 (NBG); Farm Zoekop, past ruins near edge of escarpment,

(-AA), 24 September 2002, Rosch 154 (NBG).

Moraea marginata J.C. Manning & Goldblatt, sp.

nov.

Species habitu floribusque Moraea fistulosa Goldblatt

affinis sed breviora 100-150 mm alta, foliis canaliculatis

et marginibus incrassatis hyalinis 20-25 x 5-7 mm, et

floribus parvioribus tepalis 9-13 mm longis differt.

TYPE. — Northern Cape, 3220 (Sutherland): Suther-

land, 200 m along Bo-Visrivier road south of town. Farm

142

Bothalia 36,2 (2006)

Tweefontein, stony flats in open karroid scrub, (-BC),

15 November 2005, J. Manning 2997 (NBG, holo.; K,

MO, iso.).

Plants 100-150 mm high. Conn globose, 15-20 mm

diam.; tunics of coarse black fibres. Stem erect, usually

with 1-4 branches at upper nodes, dull purplish where

exposed; spathes dry and papery at flowering, attenuate,

inner 25-30 mm long, outer about half as long. Foliage

leaf solitary, basal, longer than stem, trailing and slightly

twisted and coiled, firm-textured, greyish green with

maroon margins but dry at flowering, linear and chan-

nelled, 20-25 x 5-7 mm, margins conspicuously thick-

ened and cartilaginous, especially evident when dry;

cauline leaves 3 or 4, bract-like and entirely sheathing,

imbricate, dry and papery, attenuate. Flowers blue-violet;

tepals free but contiguous at base, oblanceolate, outer

larger, 10-13 x 3. 5-4.0 mm, inner 9-12 x 2. 5-3.0 mm,

shortly unguiculate, claws 1.0-1. 5 mm long, erect and

held against base of filaments, limbs spreading or slight-

ly reflexed, inner twisted slightly counter-clockwise

distally, propeller-like, with small, oblong, yellow nectar

guides 1. 5-2.0 mm long at base, unscented. Filaments

free but contiguous at base, 4-5 mm long, suberect,

mauve; anthers erect, 4. 5-5.0 mm long, yellow, curving

inwards above at anthesis. Ovary narrowly ellipsoid, 4

mm long; style erect, filiform, mauve, 3.5— 4.0 mm long,

style branches spreading to ascending between anthers,

filiform, 4 mm long. Capsules barrel-shaped, 6-8 x 5

mm. Seeds subglobose or angled by pressure, ± 2 mm

diam., reddish brown, testa surface rugulose. Flowering

time : November; flowers opening at ± 16:00 and wilting

at ± 20:00. Figure 3.

Ecology, known from a single small population on the

southern outskirts of the town of Sutherland at an eleva-

tion of around 1 550 m (Figure 2). Plants occur locally

in fine alluvium over shale at the foot of outcrops of

sandstone in open, succulent karroid scrub. The leaves

are quite dry and withered at flowering, and the attrac-

tive, blue-violet flowers, 20-25 mm in diameter, open in

early summer in the late afternoon for just a few hours,

withering around sunset.

Diagnosis and relationships'. Moraea marginata is

recognized by the combination of stellate, purplish

flowers with free stamens and filiform style branches,

and a solitary, linear, channelled leaf with conspicuous,

maroon margins that are conspicuously thickened when

dry. The distinctive flower form places M. marginata

with the two species previously segregated as the genus

Roggeveldia. This genus was established by Goldblatt

(1979) for R.fistulosa Goldblatt, a Moraea- like plant that

was anomalous in Moraea as then circumscribed in hav-

ing free stamens and filiform style branches that spread

between the stamens, rather than being opposite to and

± appressed to them as in typical species of Moraea. A

second species of Roggeveldia , R. montana Goldblatt,

very similar to R. fistulosa in morphology, was described

by Goldblatt ( 1 992) more than a decade later. At this time

he suggested that the relationships of the genus lay with

Moraea section Polyanthes, most particularly with M.

crispa Thunb., with which it shared a similar vegetative

and floral morphology, including a rotate, blue-violet

perianth, free stamens and narrow style branches without

style crests. The genus Roggeveldia was subsequently

included in Moraea sect. Polyanthes (Goldblatt 1998),

following a revision of the circumscription of Moraea.

Initially based on evidence from morphology, anatomy

and chromosome cytology, this decision received sub-

sequent support from molecular study (Goldblatt et at.

2002b).

Moraea marginata is indistinguishable from M. fistu-

losa (Goldblatt) Goldblatt (= Roggeveldia fistulosa) and

M. monticola Goldblatt (= R. montana) in its flowers but

differs sharply from them in its vegetative morphology.

M. fistulosa is characterized by an erect, fistulose leaf

about as tall as the stem and up to 3 mm in diameter,

whereas the smaller M. monticola is distinguished by its

trailing, filiform leaf, longer than the stem and twisted

and slightly coiled, and ± 1 mm in diameter. In both these

species, therefore, the leaf is terete/filiform. M. margi-

nata is thus unique in the group in its linear, channelled

leaf, 5-7 mm wide, with conspicuously thickened and

cartilaginous margin. The leaf of M. marginata , which

is longer than the stem, trailing, and slightly twisted and

coiled, represents a remarkable specialization in the M.

crispa alliance. Similar leaves with thickened margins

are also encountered in some plants of M. crispa from the

Roggeveld Plateau, including Bond 145 (NBG, SAM)

from the Farm Gunstfontein on the Klein Roggeveld,

and Snijman 774 (NBG) from near Williston. Although

these collections are vegetatively indistinguishable from

M. marginata, the partially fused filaments and flattened

style branches, bifid at the tips and with small style

crests, are entirely consistent with M. crispa and there is

no doubt that they represent that species.

All the species of the Roggeveldia group have a simi-

lar phenology, flowering in early summer, in October or

November, with individual flowers opening in the late

afternoon around 16:00 and withering in the evening

between 19:00 and 20:00. The species are concentrated

in the western Karoo, along the edge of the central

escarpment, and are generally poorly known and col-

lected. Moraea marginata and M. fistulosa are known

only from the type collections, both from the edge of the

Roggeveld Escarpment near Sutherland, and M. mon-

ticola from three collections, one in Namaqualand and

two from the southern margin of the Great Karoo. M.

crispa, in contrast, is widespread through the drier inte-

rior mountains of the Cape region and along the western

and southern edge of the interior escarpment (Goldblatt

1 986). M. crispa is probably plesiomorphic in its some-

what flattened and bifid style branches and thus most

likely sister to the Roggeveldia-groap.

Etymology, alluding to the unusual, thickened leaf

margin.

Romulea singularis J.C. Manning & Goldblatt,

sp. nov.

Inter species sectionis Romulea floribus malvinis ad

lilacinis fauce albo purpureo striato, tubo rubro-ochraceo

externe, tubo perianthi anguste infundibuliforme 10-11

mm longo, tepalis lanceolatis ±13x3 mm, filamentis

8-9 mm longis antheris longioribus ad basem puberulis

recedens.

Bothalia 36,2 (2006)

G

FIGURE 3. — Moraea marginata,

Manning 2997 (NBG): A,

whole plant; B, outer tepal; C,

inner tepal; D, floral details,

perianth removed; E, apex of

style branch; F, capsule; G,

seed. Scale bars: A-C, F, 10

mm; D, 2 mm; E, 0.2 mm; G,

1 mm. Artist: John Manning.

TYPE. — Northern Cape, 3119 (Calvinia): Oorlogskloof

Nature Reserve, Farm Uitkomst, 4 km east of Arrie se

Punt, overlooking Saaikloof, seasonally moist sandstone

pavement in Bokkeveld Sandstone fynbos, 830 m, (-CA),

18 September 2005, N.A. Helme 3564 (NBG, holo.).

Plants 200-400 mm high; stem subterranean. Corm

asymmetric with broad, crescent-shaped basal ridge,

10-12 mm diam.; tunics brown, woody, lower margins

splitting into fine, parallel fibrils 1. 5-2.0 mm long in

irregular clusters, drawn into coarse fibres 5-6 mm long

above. Leaves up to 6, lower 2 basal and longest, remain-

der cauline but appearing basal through contraction of

stem, blades spreading, narrowly 4-grooved, 15-35 x

0.75-1.25 mm. Inflorescence of up to 4 single-flowered

units; outer bracts ovate, subobtuse, green, with slender,

well-spaced veins and narrow, hardly visible translucent

margins, 7-10 mm long, inner bracts notched apically,

green in centre with broad translucent margins flecked or

flushed pale brown, slightly shorter than outer. Flowers

narrowly funnel-shaped, mauve to lilac with white throat

streaked with purple, tube reddish ochre on outside,

probably unscented, 12-15 mm diam.; perianth tube

narrowly funnel-shaped, 10-11 mm long, with lower,

narrow portion 3^4 mm long; tepals lanceolate, ±13x3

mm. Filaments inserted at top of lower, narrow portion of

tube, sparsely puberulous at base, 8-9 mm long, exserted

± 2 mm; anthers pale yellow, ± 3 mm long. Ovary ellip-

soid, 2 mm long; style dividing between base of anthers

and middle of anthers, branches ± 2 mm long, divided

for ± half of their length. Capsule and seeds unknown.

Flowering time : September. Figure 4.

Ecology, known from a single population on the

Bokkeveld Escarpment near Nieuwoudtville, on the east-

ern edge of the Kobee River Gorge in the Oorlogskloof

144

Bothalia 36,2 (2006)

Nature Reserve. The plants are rare and localized in

seasonally wet sand and moss on sandstone pavement

in arid fynbos vegetation. At the time that the species

was collected in mid-September, most of the plants had

finished flowering, suggesting that peak flowering takes

place in early September.

Diagnosis and relationships', although known from

just two plants, the flowers of Romulea singularis are so

unusual that they leave no doubt that the species is dis-

tinct. R. singularis is readily distinguished from all other

species by its narrowly funnel-shaped, mauve to purple

flowers with slender perianth tube, 10-11 mm long. The

unusually long filaments, 8-9 mm long and puberulous

at the base, are inserted in the lower part of the tube and

are shortly exserted, and the floral bracts are short, 8-10

mm long. The unusual length of the filaments and their

insertion in the lower part of the tube, at the top of the

basal, narrow portion, are unique among the handful of

long-tubed species of Romulea that are known (Manning

& Goldblatt 2001b). Those with a truly salver-shaped

flower [R. alba J.C. Manning & Goldblatt, R. hantamen-

sis (Diels) Goldblatt, R. stellata M.P.de Vos and R. syrin-

godeoflora M.P.de Vos] have the filaments inserted near

the mouth of the tube, but even R. kamisensis M.P.de

Vos, with a narrowly funnel-shaped flower, has the fila-

ments inserted in the upper part of the tube. In addition,

the filaments in all of these species are glabrous through-

out and 3-5 mm in length, thus not unusually long.

The oblique conn with crescent-shaped basal ridge

splitting into parallel fibrils, places Romulea singularis

in section Ciliatae of subgenus Romulea (Manning &

Goldblatt 2001b). Within the section, however, its rela-

tionships are more difficult to determine. The narrowly

funnel-shaped, mauve to purple flower at first appearance

suggests an atypically short-tubed form of R. kamisensis

M.P.de Vos, a species that is thus far known from central

Namaqualand and the northern rim of the Knersvlakte.

Closer examination, however, reveals that the resem-

blance is superficial. The bracts of R. kamisensis are

much longer, 13-22 mm long, and the filaments, which

are inserted in the upper part of the tube, are entirely

glabrous, just 4-5 mm long, and are included within the

tube. In all these respects R. kamisensis is quite unlike R.

singularis and it is therefore probable that the similarity

in flower form in the two species is convergent.

The unusually well-developed, basal ridge on the

corm, with the parallel fibrils grouped into small clus-

ters, may suggest a possible relationship with Romulea

toximontana M.P.de Vos. This species, which is also

endemic to seasonally wet sandstone pavement on the

Bokkeveld-Matsikamma-Gifberg Mountain complex,

is distinctive in section Ciliatae in the wide, fan-shaped

basal ridge on the corm. Its white, cup-shaped flowers,

with short perianth tube, although quite unlike those of

R. singularis in shape, share with it the darker streaks

in the throat and the pale yellow anthers, and the bracts

are typically short, 10-20 mm long, with similar slender

veins. Unfortunately these features are probably all ple-

siomorphic and it is therefore only the well-developed

rim on the corm that might actually signal a relationship

between the two species. Cytology may prove more

useful since the chromosome number, In = 28, in R.

FIGURE 4. -Romulea singula-

ris. Helme 3564 (NBG): A,

whole plant; B, tepal; C, outer

bract; D, inner bract; E, floral

details. Scale bars: A-D, 10

mm; E, 5 mm. Artist: John

Manning.

Bothalia 36,2 (2006)

145

toximontana is unique in section Ciliate, where 2 n = 24

is usual (De Vos 1972).

On the basis of the presence of secondary vascular bun-

dles in the leaves, and possibly also their shared chromo-

some number, 2 n = 26, which is rare in section Ciliatae,

it is likely that the long-tubed Romulea kamisensis shares

a short-tubed ancestor with R. namaquensis M.P.de Vos

(Manning & Goldblatt 2001b). R. singularis may be

similarly but independently derived from some other

short-tubed species and it does appear as if most of the

long-tubed species of Romulea are independently derived

within the genus in response to pollination by long-

tongued insects. The pollination biology of the genus

has been fairly well studied (Goldblatt et al. 2002a), and

most of the other long-tubed species have been shown

to be adapted to pollination by long-proboscid flies in

the family Nemestrinidae. A similar pollination system

is thus likely for R. singularis, and the species will most

probably prove to be another member of the Prosoeca

peringueyi pollination guild, which is well-developed in

the Bokkeveld Mountains (Manning & Goldblatt 1996).

Etymology, named for its unusual flowers.

ACKNOWLEDGEMENTS

We are most grateful to Nick Helme for bringing

Moraea marginata and Romulea singularis to our atten-

tion. Thanks are due also to Elizabeth Parker for facili-

tating the trip to collect the type material of M. margi-

nata. Material was collected under permits issued by the

Departments of Nature Conservation of the Northern and

Western Cape.

REFERENCES

DE VOS, M.R 1972. The genus Romulea in South Africa. Journal of

South African Botany, Suppl. vol. 9.

DE VOS. M.R 1999. Ixia. Flora of southern Africa 7,2,1: 1-87. Na-

tional Botanical Institute, Pretoria.

GOLDBLATT, P. 1979. Roggeveldia, a new genus of southern African

Iridaceae-Irideae. Annals of the Missouri Botanical Garden 66:

839-844.

GOLDBLATT, P. 1986. The moraeas of southern Africa. Annals of

Kirstenbosch Botanic Gardens 14. National Botanic Gardens,

Cape Town and Missouri Botanical Garden, St Louis.

GOLDBLATT, P. 1992. New species, chromosome cytology and notes

on the southern African Iridaceae-Irideae: Moraea. Roggeveldia

and Homeria. South African Journal of Botany 58: 209-214.

GOLDBLATT, P. 1998. Reduction of Barnardiella, Galaxia, Gynan-

driris, Hexaglottis, Homeria and Roggeveldia in Moraea (Irida-

ceae: Irideae). Novon 8: 371-377.

GOLDBLATT, P. 2004. A synoptic review of the African genus Hes-

perantha (Iridaceae: Crocoideae). Annals of the Missouri Bo-

tanical Garden 90: 390 — 443 .

GOLDBLATT, P„ BERNHARDT. P. & MANNING, J.C. 1998. Pol-

lination of petaloid geophytes by monkey beetles (Scarabaeidae:

Rutelinae: Hopliini) in southern Africa. Annals of the Missouri

Botanical Garden 85: 215-230.

GOLDBLATT, R, BERNHARDT, P. & MANNING, J.C. 2002a. Flo-

ral biology of Romulea (Iridaceae: Crocoideae): a progression

from a generalist to a specialist pollination system. Adansonia

24: 243-262.

GOLDBLATT, P. & MANNING, J.C. 1999. New species of Sparaxis

and Ixia (Iridaceae: Ixioideae) from Western Cape, South Af-

rica, and taxonomic notes on Ixia and Gladiolus. Bothalia 29:

59-63.

GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanical

Institute, Cape Town & Missouri Botanical Garden, St. Louis.

GOLDBLATT, P, SAVOLAINEN, V„ PORTEOUS, 0., SOSTARIC,

I., POWELL, M„ REEVES, G„ MANNING, J.C., BARRA-

CLOUGH. T.G. & CHASE, M.W. 2002b. Radiation in the Cape

flora and the phylogeny of peacock irises Moraea (Iridaceae)

based on four plastid DNA regions. Molecular Phylogeny and

Evolution 25: 341-360.

LEWIS, G.J. 1962. South African Iridaceae: the genus Ixia. Journal of

South African Botany 27: 45-195.

MANNING, J.C. & GOLDBLATT, P. 1996. The Prosoeca peringueyi

(Diptera: Nemestrinidae) pollination guild in southern Africa:

long-tongued flies and their tubular flowers. Annals of the Mis-

souri Botanical Garden 83: 67-86.

MANNING, J.C. & GOLDBLATT, P. 1997. Nieuwoudtville. South Af-

rican Wild Flower Guide 9. Botanical Society of South Africa,

Cape Town.

MANNING, J.C. & GOLDBLATT, P. 2001a. Three new species of

Tritoniopsis (Iridaceae: Crocoideae) from the Cape Region of

South Africa. Bothalia 31: 175-181.

MANNING, J.C. & GOLDBLATT, P. 2001b. A synoptic review of Rom-

ulea (Iridaceae: Crocoideae) in sub-Saharan Africa, the Arabian

Peninsula and Socotra, including new species, biological notes,

and a new infrageneric classification. Adansonia 23: 59-108.

MANNING, J.C., GOLDBLATT, P. & SNIJMAN, D A. 2002. The

color encyclopedia of Cape bulbs. Timber Press, Portland.

SNIJMAN, D.A. & PERRY, P. 1987. A floristic analysis of the Nieu-

woudtville Wild Flower Reserve, north-western Cape. South Af-

rican Journal of Botany 53: 445^154.

VAN JAARSVELD. E.J. & THOMAS, V. 2003. Freylinia helmei.

Flowering Plants of Africa 58: 112-116, t. 2197.

VAN WYK. A.E. & SMITH, G.F. 2001. Regions of floristic endemism

in southern Africa: a review with emphasis on succulents. Um-

daus Press, Hatfield, Pretoria.

■

Bothalia 36,2: 147-155 (2006)

Three new species of Lachenalia (Hyacinthaceae: Massonieae) from

Western and Northern Cape, South Africa

G.D. DUNCAN* and T.J. EDWARDS**

Keywords: Hyacinthaceae, Lachenalia J.Jacq. ex Murray, new species. South Africa

ABSTRACT

This is the sixth in a series of papers on Lachenalia, towards a revision of the genus. Three new species are described, L.

lutea from the southwestern part of the Western Cape, L. cernua from the southern Cape Peninsula and the Worcester Valley

of the Western Cape, andfr. nardousbergensis from the Bokkeveld Plateau of the Northern Cape, and the Nardousberge and

Middelburg Plateaus of the Western Cape.

INTRODUCTION

The horticulturally important and botanically diverse

genus Lachenalia J.Jacq. ex Murray is endemic to south-

ern Africa and comprises 120 species of deciduous geo-

phytes, almost all of which are winter growing (Duncan

et al. 2005). The distribution of Lachenalia extends

from southwestern Namibia into the western, southern,

eastern and central parts of South Africa, and the cen-

tre of diversity is in the Worcester grid (3319), divided

between the Succulent Karoo and Fynbos Biomes, in the

mountains and valleys of the winter rainfall region of the

Western Cape (Duncan 2005).

Species delimitation in Lachenalia is usually unam-

biguous, but in some instances there is gradation between

species. Other species display extensive variation which

has led to considerable taxonomic confusion due to over-

emphasis of minor morphological differences. Variation

within a species occurs in several macro-morphologi-

cal characters such as overall plant size, leaf number,

pedicel length, degree of stamen exsertion, flower size,

flower colour and orientation, and flowering period.

Variable species often display population stability in

features such as bulb and flower shape, and seed mor-

phology; however, a number of species are exceedingly

variable (Duncan et al. 2005).

In a cladistic analysis of morphological data, it was

concluded that a number of evolutionary pressures have

driven divergence of vegetative and floral characters in

Lachenalia. The convergent adaptation to conditions of

aridity appears to be the main reason for homoplasy in

whole sets of vegetative characters, and similarly, the

convergent modification of flowers to similar pollina-

tors is probably the main reason for homoplasy in whole

sets of reproductive characters (Duncan et al. 2005).

The genus was last revised by Baker (1897) in which

42 species were recognized. The new species described

here form part of a series of papers towards a revision

* South African National Biodiversity Institute, Kirstenbosch, Private

Bag X7, 7735 Claremont, Cape Town.

** Botany Department, University of KwaZulu-Natal, Private Bag

X01, 3209 Scottsville, Pietermaritzburg.

MS. received: 2006-01-13.

of the genus (Duncan 1993, 1996, 1997, 1998, Duncan

& Edwards 2002).

Lachenalia lutea G.D. Duncan, sp. nov.

Plantae 160-240 mm alta; bulbus globosus, 15-20

mm in diametro, folia 2, lanceolata, coriacea, patentia ad

suberecta, claro viridia, pagina superior immaculata vel

maculis atroviridibus, 90-140 x 12-30 mm, marginibus

coriaceis, inflorescentia spicata, erecta, densa, 70-110

mm longa, pedunculus vivido viridis vel immaculatus

maculis brunneo-purpureis, flores oblongo-campanu-

lati, suberecti, pallide ad claro viridi-flavi, demum

obscure sanguinei, peraromatici, perianthii tubum cyat-

hiforme, 3 mm longa, tepala exteriora ovata, 6-7 x 4

mm, gibbis claro viridibus vel atroflavis, tepala inte-

riora obovata, trans tepala exteriora bene exsertis, 8-9 x

4-5 mm, carinis claro viridibus vel atroflavis, stamina

inclusa vel trans perianthium parum exserta, filamenta

recta albida, 5-6 mm longa, ovarium ellipsoideum, 3 x

2.5 mm, stylum rectum, albidum, 5-6 mm longa, cap-

sula ellipsoidea, 8-9 x 4-5 mm, semina globosa, 0.9 x

0.8 mm, strophiolo inflato, 0. 5-0.6 mm longa.

TYPE. — Western Cape, 3418 (Simonstown):

Vergelegen Farm, Somerset West, low hillside near res-

ervoir, in heavy clay soil, (-BB), 7-10-1959, W.F. Barker

9088 (NBG, holo.!).

Deciduous, winter-growing geophyte 160-240 mm

high. Bulb globose, 15-20 mm diam., offset-forming,

white, surrounded by dark brown, spongy outer tunics;

cataphyll subterranean, translucent white, tightly sur-

rounding clasping leaf base, apex obtuse. Leaves 2,

lanceolate, 90-140 x 12-30 mm, spreading to suberect,

flat or weakly canaliculate, leathery, bright green or

greenish magenta, plain or marked with dark green spots

on upper surface, lower surface sometimes marked with

brownish purple spots; margins thickened; clasping base

10-30 mm long, greenish white to deep magenta, plain or

occasionally barred with brownish purple in upper part.

Inflorescence an erect, dense, few- to many-flowered

spike 70-110 mm long, with sterile tip 15-20 mm long;

peduncle erect, sturdy, 70-150 mm long, pale green or

brownish green, plain or marked with large, irregularly

scattered brownish purple blotches; rachis pale green,

shading to greenish yellow in upper third; bracts ovate

at base of inflorescence, becoming lanceolate above, 2-7

148

Bothalia 36,2 (2006)

x 1-5 mm, translucent white. Flowers sessile, suberect,

oblong-campanulate, sweetly scented; perianth tube cup-

shaped, 3 mm long, pale greenish yellow; outer tepals

ovate, 6-7 x 4 mm, pale to bright greenish yellow with

bright green gibbosities; inner tepals obovate, 8-9 * 4-5

mm, protruding well beyond outer tepals, apices suba-

cute, upper inner tepals overlapping, lower inner tepal

slightly recurved, pale greenish yellow with bright green

keels in upper half. Stamens usually well included within

perianth, rarely just exserted; filaments straight, usually

5-6 mm long, white, rarely up to 8 mm long; anthers

dull brownish maroon prior to anthesis, bright yellow at

anthesis, maturing to black after anthesis. Ovary ellip-

soid, 3 x 2.5 mm, pale yellowish green; style 5-6 mm

long, straight. Capsule ellipsoid, 8-9 x 4-5 mm. Seed

globose, 0.9 x 0.8 mm, shiny black, with inflated, smooth

strophiole 0.5-0. 6 mm long. Flowering time: late July to

early October. Figures 1A; 2.

Etymology, the specific epithet lute a is named for the

pale to bright yellow flowers.

History : the earliest known record of Lachenalia

lutea is that of J.F. Solly (Solly sub PRE11352) who col-

lected it in August 1915 at the foot of Sir Lowry’s Pass

east of Cape Town, where this species is common, and

has been collected on numerous occasions. It has since

been widely collected in the southwestern and southern

parts of the Western Cape.

Diagnostic features and affinities'. Lachenalia lutea

has for many years been overlooked as a distinct taxon.

It belongs to the group of species having sessile, oblong-

campanulate flowers with straight stamens, and most

closely resembles L. arbuthnotiae. L. lutea is recognized

by its dense inflorescence of suberect, heavily sweet-

scented, pale to bright yellow, oblong-campanulate

flowers with spreading inner tepals that protrude well

beyond the outer tepals (Figures 1A; 2). The outer tepals

have bright green gibbosities, the inner tepals have suba-

cute apices and the stamens are usually included within

the perianth or rarely slightly exserted. The sturdy

peduncle is pale green or brownish green and may be

plain or heavily marked with brownish purple blotches.

The two leathery, lanceolate leaves are spreading to

suberect and bright green or greenish magenta, with or

without darker green or greenish magenta blotches on

the upper surface. The globose seeds have a shiny black

testa and a smooth, inflated strophiole.

The similar oblong-campanulate flowers of

Lachenalia arbuthnotiae are also heavily sweet-scented

but are slightly curved and distinctly longer, with the

inner tepals only slightly longer than the outer ones,

and scarcely spreading. The inner tepal apices in this

species are obtuse and undulate, and flower orientation

is spreading to weakly suberect. It has much longer fila-

ments, 8-9 mm long, a much longer style, 9 mm long

and its anthers emerge at the mouth of the perianth or

are shortly exserted. The style protrudes conspicuously

beyond the perianth as the ovary enlarges. Like

those of L. lutea , the seeds of L. arbuthnotiae are

globose with a shiny testa, but its similar inflated.

FIGURE I. —Flowering specimens of three new species of Lachenalia. A, L. lutea, Duncan 430, in habitat, Elgin; B, L. cernua, Duncan 470. in

habitat, Wolseley; C, L. nardousbergensiS, Duncan 198, in habitat, Nieuwoudtville. Scale bars: 10 mm.

Bothalia 36,2 (2006)

149

FIGURE 2. — Lachenalia lutea, Duncan

430 (NBG). A, inflorescence and

leaves; B, bulb; C, single flower;

D, 1/s flower. Scale bars: A-D, 10

mm. Artist: Vicki Thomas.

smooth strophiole is slightly longer. As in L. lutea ,

L. arbuthnotiae grows in colonies but is an altogeth-

er larger plant occurring in a completely different habitat,

confined to the Cape Flats Lowlands from Wetton to

Faure, in isolated coastal fynbos remnants in seasonally

inundated, deep sandy soil (Duncan 1988a). The peak

flowering period for L. arbuthnotiae is mid-September,

whereas L. lutea generally flowers earlier, with a peak

period from mid-August to early September.

Distribution and habitat : Lachenalia lutea occurs

in the Fynbos Biome in the southwestern Cape, its dis-

tribution extending between Strand and Bot River, and

Tulbagh to Villiersdorp (Figure 3). It usually occurs

in stony, heavy clay soil in renosterveld, or rarely in

sandy soil, and is found in a variety of habitats includ-

ing seasonally moist, low-lying flats and hills, and on

shale bands of higher mountain slopes, growing as

scattered individuals or in small groups of two to three

plants within large colonies. It flowers particularly well

150

Bothalia 36,2 (2006)

following summer bush fires and is frequently seen

growing in association with L. orchioides (L.) Aiton

var. orchioides that flowers later in the season. L. Intea

is still a common species across most of its range but

its numbers have been much reduced by the establish-

ment of deciduous fruit orchards, and in recent years it

has been greatly reduced on the clay flats near Somerset

West, east of Cape Town, due mainly to industrial and

housing development.

Additional specimens examined

WESTERN CAPE.— 3318 (Cape Town): Klipheuwel, (-DA), 14-

8-1962, Lewis 6134 (NBG); Zuider Paarl, (-DB), 9-1917, Adendorff

17637 (PRE); Langverwacht Farm above Kuilsrivier, (-DC), 25-8-

1973, Oliver 4363 (NBG); Assegaaibosch, (-DD), 9-1969, Van der

Merwe 994 (PRE); Stellenbosch, (-DD), 27-8-1963, Taylor 5032

(PRE). 3319 (Worcester): Tulbagh Kloof, (-AC), 10-1920, Andreae

616 (PRE); 2 miles [3.2 km] west of Ceres, (-AC), 11-9-1968, Marsh

767 (PRE); 1 mile [1.6 km] south of Tulbagh Road Station, (-AC),

2-8-1967, Rourke 789 (NBG); Palmiet Valley, Wolseley, (-AC), Aug.

2003, Van Warmelo s.n. (NBG); Botha’s Halt near Worcester, (-CB),

30-8-1946, Compton 18283 (NBG); Wangenheim Farm, Rawsonville,

(-CB), 30-9-1963, Walters 952 (NBG). 3418 (Simonstown): road

from Strand to Gordon’s Bay, (-BB), 12-9-1947, Parker 4240 (NBG);

Gordon’s Bay, (-BB), 23-9-1958, Werdermann 137 (PRE); Sir Lowry’s

Pass, (-BB), 27-9-1958, Werdermann 210 (PRE); road to Sir Lowry’s

Pass (-BB), 12-10-1955, De Wet 930 (PRE); Sir Lowry’s Pass (-BB),

19-8-1915, Solly sub PRE1I352 (PRE); Steenbras River mouth, (-BB),

16-9-1951, Esterhuyser 18843 (PRE). 3419 (Caledon): Freshwoods,

Elgin, (-AA), 25-11-1998, Duncan 419 (NBG); Aprilskraal, Elgin,

(-AA), 7-10-2000, Duncan 430 (NBG); Theewaterskloof, Villiersdorp,

(-AA), 3-9-1975, Thomas s.n. (NBG); Happy Valley Farm, southeast

of Villiersdorp, (-AB), 30-9-1971, Barker 10840 (NBG); between

Hawston and Karwyderskraal, (-AC), 29-9-1967, Barker 10513

(NBG); Ysterklip Farm, Kleinmond, (-AC), 8-11-1967, Barker 10514

(NBG); Bot River, (-AC), 16-8-1982. Bot River-Hermanus Road,

(-AC), 30-8-1973, Barker 10884 (NBG); 2.5 km from turnoff at bridge

on Hermanus Road, (-AC), 30-8-1973, Barker 10886 (NBG); Bot

River Vlei, (-AC), 16-8-1982, O’Callaghan 249 (NBG); Kleinmond,

(-AC), 8-1963, Topper 149 (NBG); Greyton, (-BA), 15-8-1969,

Barker 10641 (NBG); Heuningkloof Farm beyond Caledon, (-BA),

7-9-1974, Barker 10917 (NBG); Middelplaas turnoff near Genadendal,

(-BA), 7-9-1974, Barker 10918 (NBG), Greyton Commonage, (-BA),

Hofmeyer s.n. (NBG); 46 km E of Caledon, (-BA), 3-9-1975, Thomas

s.n. (NBG).

Lachenalia cernua G.D. Duncan, sp. nov.

Plantae 150-270 mm altae, bulbus globosus, 15-25

mm in diametro, folium solitarium, interdum 2, lanceola-

tum, patens, viride que immaculatum, vel in pagina supe-

riore marroninum maculis plus atromarroninis, 110-260

x 1 0-28 mm, inflorescentia racemosa erecta vel suberec-

ta, 30-100 mm longa, pedunculus pallide viridis macu-

lis marronino-purpureis, 60-130 mm longa, pedicelli

suberecti, 2 mm longi, flores urceolati, tempore primo

ad medio florendi deinde cemui, in fructo patentes, pal-

lide cremeo-flavi, perianthii tuburn cyathiforme, 3 mm

longum, tepali exteriora ovata, base obscure caerulei

suffusi, gibbis flavo-viridibus, 6-8 x 5 mm, tepala inte-

riora obovata, trans tepalis exteriora bene exserta, carinis

flavis, marginibus recurvatis, 11-12 x 5 mm, stamina

trans perianthium breviter exserta, filamenta recta, alba,

11-13 mm longa, ovarium ellipsoideum, 4x3 mm, sty-

lum rectum, album, 12 mm longum, capsula ellipsoidea,

9-10 x 6 mm, semina globosa, 1.2 x 1.1 mm, strophiolo

inflato, 1 mm longo.

TYPE. — Western Cape, 3319 (Worcester): Palmiet

Valley Farm, hillside behind homestead, in open aspects

and semi-shade of sandstone boulders, (-AC), 21-9-

2002, Duncan 470 (NBG, holo., PRE, iso.).

Deciduous, winter-growing geophyte 150-270 mm

high. Bulb globose, shallow or deep-seated, 15-25 mm

diam., white with several layers of membranous, dark

brown outer tunics; cataphyll subterranean, translucent

white, loosely surrounding lowermost portion of clasping

leaf base, apex obtuse; clasping leaf base relatively long

depending on depth of bulb, usually completely subter-

ranean, occasionally emerging slightly above ground

level, 15-85 mm long, white, sometimes forming bulbils

along subterranean margins. Leaves usually solitary,

occasionally 2, spreading, narrowly lanceolate, weakly

canaliculate, 110-260 x 10-28 mm, uniformly pale to

dark green, or pale to dark maroon and sporadically or

heavily marked with darker maroon blotches on upper

surface. Inflorescence an erect or suberect, few- to many-

flowered raceme 30-100 mm long, with a short sterile

tip; peduncle erect or suberect, 60-130 mm long, pale

green, slightly to heavily marked with dark maroon or

maroonish purple blotches; rachis, 50-175 mm long, pale

green, immaculate or heavily blotched with maroon or

maroonish purple; bracts ovate in lower half of inflores-

cence, becoming lanceolate in upper half, 1-5 x 1-7 mm,

translucent white; pedicels suberect, 2 mm long, white.

Flowers urceolate, suberect in bud, cemuous from early

to mid-flowering, becoming spreading at late flowering

and fruiting stage, creamy white and pale yellow with

green markings; perianth tube cup-shaped, 3 mm long,

creamy white, occasionally tinged with dull blue in upper

half; outer tepals ovate, 6-8 x 5 mm, creamy white or

greenish yellow, occasionally tinged with dull blue at

base, with a yellowish green gibbosity near apex; inner

tepals obovate, protruding well beyond outer tepals, 11—

12x5 mm, creamy white or greenish white with pale to

dark yellow keels in upper half, apices slightly recurved.

Stamens ± straight, subequal, exserted 1-2 mm beyond

tip of perianth; filaments 11-13 mm long, white. Ovary

ellipsoid, 4x3 mm, bright green; style straight, white, 12

mm long. Capsule ellipsoid, 9-10 x 6 mm, olive green.

Seed globose, 1.2 x 1.1 mm, shiny black, with smooth,

inflated strophiole 1 mm long. Flowering time : late

September to mid-October. Figures IB; 4.

Etymology, the specific epithet cernua is named for

the slightly drooping orientation of its flowers during the

early and mid-flowering stage.

Bothalia 36,2 (2006)

151

FIGURE 4. — Lachenalia cernua ,

Duncan 470 (NBG). A. inflo-

rescence, leaf and bulb; B,

single flower; C, 1/s flower.

Scale bars: A, 10 mm; B, C, 5

mm. Artist: John Manning.

History : the earliest known record of Lachenalia

cernua is a sheet housed in the National Herbarium,

Pretoria, collected by the amateur botanist Dr F.Z.

van der Merwe in October 1937 ( Van der Merwe sub

PRE35699) on a hillside above the spa baths at Goudini

in the Worcester Valley of the southwestern Cape. Two

further collections in NBG were made at this locality

by J.W. Loubser in October 1971 and September 1972

respectively (Loubser s.n. 2181). In early October 2000,

flowering specimens were collected by the first author

some distance to the northwest of Goudini, on a hillside

just south of Wolseley (Duncan 428). In July 2001, leaf-

ing specimens of an unidentified Lachenalia were col-

lected by Adam Harrower of Kirstenbosch at the naval

base at Klavervlei near Simonstown in the southern Cape

Peninsula (Harrower 104). When one of these flowered

152

Bothalia 36,2 (2006)

in the Kirstenbosch nursery in October the same year,

it matched those of the Goudini and Wolseley collec-

tions. The discovery of the Simonstown population is

remarkable for a genus previously thought to have been

extensively documented in the southern Cape Peninsula,

and is the first record since 1949 of a new species in

this genus from the Cape Peninsula since the publica-

tion of the endemic Cape Peninsula species L. capensis

W.F.Barker, and L. variegata W.F.Barker, that occurs

from the Cape Peninsula to Clanwilliam (Barker 1949).

The restricted environment of the naval base no doubt

accounts for it having remained undetected there for so

long, and no other populations are known to occur on the

Cape Peninsula. In late September 2002, the Wolseley

locality was visited again, and the type collection was

made (Duncan 470).

Diagnostic features and affinities'. Lachenalia cer-

nua is a member of the group of species having small

pedicellate, urceolate flowers with straight stamens,

and includes L. peersii Marloth ex W.F.Barker, which it

most closely resembles. L. cernua is recognized by its

moderately dense inflorescence of urceolate, pale creamy

yellow flowers that are cemuous during early and mid-

flowering, becoming spreading during late flowering

and fruiting stage (Figures IB; 4). The inner tepals are

creamy white or greenish white with pale to dark yellow

keels in the upper half, and are well exserted beyond

the outer tepals, with slightly recurved apices. The outer

tepals are creamy white or greenish yellow, and have yel-

lowish green gibbosities. The stamens are exserted 1-2

mm beyond the tip of the perianth. The usually solitary

leaf is narrowly lanceolate and spreading, and varies in

colour from uniformly pale to dark green, to pale to dark

maroon with scattered darker maroon blotches on the

upper surface. The clasping leaf base is entirely subter-

ranean or occasionally slightly emerging above ground

level, and the globose seeds have a shiny black testa and

a long inflated strophiole, 1 mm long.

Lachenalia cernua resembles L. peersii in the shape

of its urceolate flowers with the inner tepals protrud-

ing well beyond the outer tepals, but the latter has pure

white, spreading flowers with included stamens, and

inner tepals that are distinctly recurved at their tips. L.

cernua lias longer inner tepals and its flowers emit a

weak, spicy scent, whereas those of L. peersii are strong-

ly carnation-scented. The peduncle of L. cernua is pale

green and slightly to heavily marked with dark maroon

or maroonish purple blotches, whereas the peduncle of

L. peersii is always immaculate. L. cernua usually has

a solitary, weakly canaliculate spreading leaf, often with

dark maroon blotches on the upper surface, whereas L.

peersii almost always has two, ± flat, lanceolate leaves

that are always unmarked. The flowering period of the

two species does not overlap as L. cernua starts flower-

ing in late September and ends in mid-October, whereas

L. peersii starts flowering in late October. The flowers of

L. cernua fade to dull red and become spreading during

the fruiting stage, whereas those of L. peersii fade to dull

pink and become suberect to erect at the fruiting stage.

Both species have globose seeds with inflated strophioles

1 mm long. The two species are geographically well

separated: L. cernua occurs on hillsides in the southern

Cape Peninsula, the Worcester Valley and near Wolseley,

whereas L. peersii is confined to flats and lower moun-

tain slopes along the southern Cape Atlantic coastline

stretching from Betty’s Bay to Caledon (Duncan 2003).

Distribution and habitat : Lachenalia cernua has a

restricted distribution in the southwestern Cape where

it is currently known from just three populations, one

from the western end of the Worcester Valley at Goudini,

another to the northwest of Goudini just south of

Wolseley, and the third near Simonstown in the south-

ern Cape Peninsula. The population at Goudini is the

closest spatially to the Simonstown population, a dis-

junction of more than 100 km (Figure 5). The Wolseley

population occurs on an east-facing hill slope in the

semi-shade of large sandstone boulders, as well as at a

slightly lower altitude in full sun. The plants growing

in semi-shade tend to occur in small groups and flower

erratically, whereas those in full sun usually occur sin-

gly and flower reliably every year. At the single known

population on the southern Cape Peninsula at Klavervlei

near Simonstown, plants grow under similar conditions,

mainly between large sandstone boulders on shaded,

east- and southeast-facing ridges at 363 m, and also in

open aspects at a slightly lower altitude, just above a

seasonal stream in which Moraea ramosissima occurs,

flowering at the same time of year. Other notable com-

panion species at this locality include Protea cynaroides

and Watsonia tabularis.

Additional specimens examined

WESTERN CAPE. — 3319 (Worcester): Palmiet Valley, (-AC), 7-

10-2000. Duncan 428 (NBG); Goudini, (-CB), 11-10-1971, Loubser

2181 (NBG); 27-9-1972, Loubser s.n. (NBG); Oct. 1937, Van der

Merwe sub PRE35699 (PRE). 3418 (Simonstown): Klavervlei,

Simonstown, (-AB), 19-10-2001, Duncan 463 (NBG); 17-7-2001,

Harrower 104 (NBG).

Lachenalia nardousbergensis G.D. Duncan, sp. nov.

Plantae 150-310 altae; bulbum subglobosum, 15-20

mm in diametro; folia 2, late lanceolata, prostrata, oli-

vacea, 100-180 x 25-55 mm, pagina superior venis

depressis longitudinalibus et pustulis magnis atroviridi-

bus, marginibus coriaceis albis; inflorescentia racemosa

erecta, 80-180 mm longa, floribus multis; pedunculus

robustus, in parte superiore inflatus, pallide viridis,

maculis brunneo-purpureis, 80-120 mm longus; pedicelli

suberecti, 2-6 mm longi; flores oblongo-campanulati,

cemui ad patentes, pallide ad atromagentei; perianthii

tubum cyathiforme, 1-2 mm longum; tepala exteriora

ovata, 7-8 x 4-5 mm, gibbis carinisque atromagenteis

vel viridibus; tepala interiora obovata, 8-9 x 4-5 mm,

trans tepala exteriora exserta, carinis atromagentis; stam-

ina trans perianthium bene exserta, filamenta declinata,

in parte 2/3 inferiore alba, in parte '/3 superiore magentea,

14-16 mm longum; ovarium obovoideum, 3x2 mm;

stylum declinatum, in parte 2/3 inferiore album, in parte

’/3 superiore magenteum, 13-14 mm longum; capsula

obovoidea, 6-8 x 5-7 mm; semen globosa, 1.2 x 1.3 mm;

strophiolo rudimentario, 0.6-0. 7 mm longo.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): road

to Nardousberge Plateau southeast of Klawer, in deep

Bothalia 36,2 (2006)

153

FIGURE 5. — Distribution of Lachenalia cemua,%\ L. nardousbergen-

sis. ■.

red sand, (-DD), 2-9-1945, W.F. Barker 3630 (NBG,

holo.!).

Deciduous, winter-growing geophyte, 150-310 mm

high. Bulb subglobose, 15-20 mm diam., clump-form-

ing or sometimes solitary, dark yellow, surrounded by

several layers of dark brown, spongy outer tunics; cata-

phyll subterranean, translucent white, tightly adhering

to clasping leaf base, apex obtuse. Leaves 2, broadly

lanceolate, 100-180 x 25-55 mm, prostrate, upper sur-

face olive-green with distinct depressed longitudinal

veins and large, irregularly scattered, dark green, flat-

tened pustules; margins white, coriaceous; clasping base

subterranean, 10-20 mm long, white. Inflorescence an

erect, many-flowered raceme 80-180 mm long, with a

short sterile tip; peduncle erect, sturdy, 80-120 mm long,

usually distinctly inflated in upper portion, pale green,

heavily marked with small to large, pale to dark brownish

purple blotches; rachis pale green, inflated in lower por-

tion, heavily marked with pale to dark brownish purple

blotches; pedicels suberect, 2-6 mm long, brownish green

to brownish magenta, shortest at base of inflorescence;

bracts ovate at base of inflorescence, becoming lanceo-

late above, 1-5 x 3-5 mm. Flowers oblong-campanulate,

nodding or spreading, pale to deep magenta; perianth tube

cup-shaped, 1-2 mm long, white throughout or dull blue

at base shading to white above; outer tepals ovate, 7-8

x 4-5 mm, pale magenta with darker magenta, brownish

magenta or green keels and gibbosities; inner tepals obo-

vate, 8-9 x 4-5 mm, apices subacute, slightly spreading,

protruding well beyond outer tepals, pale magenta with

prominent, dark magenta keels. Stamens well exserted

beyond perianth, declinate; filaments 14—16 mm long,

white in lower two thirds, shading to deep magenta in

upper third; anthers dull magenta prior to anthesis, yel-

low at anthesis. Ovary obovoid, 3x2 mm, pale green;

style declinate, well exserted beyond perianth, 13-14

mm long, white in lower one third, shading to pale to

deep magenta in upper two thirds. Capsule obovoid, 6-8

x 5-7 mm, brownish green. Seed globose, 1.2 x 1.3 mm,

shiny black with smooth, rudimentary strophiole, 0.6-0. 7

mm long. Flowering time: late August to early October.

Figures 1C; 6.

Etymology’: Lachenalia nardousbergensis is named

after the Nardousberge southeast of Klawer in the

Western Cape, where the type collection and several

other collections of this species have been made.

History: the earliest known collection of Lachenalia

nardousbergensis was made by W.F. Barker on 2

September 1945 along a road leading off the N7 to

the Nardousberge Plateau southeast of Klawer (Barker

3630). No further collections appear to have been made

until September 1968, when W. Chater collected it in the

same area ( Chater s.n ). It has since been recorded from

the Bokkeveld Plateau near Nieuwoudtville (Duncan

198 ) and the northern Cederberg (Nicklin 179 ; G.

Summerfield pers. obs.).

Diagnostic features and affinities: Lachenalia nar-

dousbergensis falls into the group of species having pedi-

cellate, oblong-campanulate flowers with well-exserted,

declinate stamens, and most closely resembles L. pur-

pureo-caerulea Jacq. L. nardousbergensis is recognized

by its usually distinctly inflated peduncle that is marked

with large brownish purple blotches, and its many-flow-

ered, moderately dense raceme of nodding or spreading,

pale to deep magenta, oblong-campanulate flowers with

well-exserted, declinate stamens, with the filaments

deep magenta in the upper third (Figures 1C; 6). Its two

prostrate, broadly lanceolate, olive-green leaves have

conspicuous longitudinal grooves along the upper sur-

face, and are covered with large, dark green, flattened,

oval pustules. The leaves are partially or completely

withered at flowering but remain green under cultivation

if plants are kept well watered in late winter and spring.

The flesh of the subglobose bulb is dark yellow and the

bulb is surrounded by several layers of dark brown, outer

tunics. Its globose, shiny black seeds have a rudimentary

strophiole, 0.6-0. 7 mm long.

The oblong-campanulate flowers of Lachenalia pur-

pureo-caerulea differ from those of L. nardousbergensis

in their deep purplish blue colour and in being larger

and more widely flared, with rounded, recurved, deep

purple inner tepal apices and a much shorter style 8 mm

long and stamens 9-10 mm long. The peduncle of L.

purpureo-caerulea is unmarked and non-inflated, and it

has two bright green, prostrate lanceolate leaves densely

covered with small green, dome-shaped pustules. Like

those of L. nardousbergensis, the leaves of L. purpureo-

caerulea are partially or completely withered at flower-

ing in the wild and the two species are geographically

widely separated, L. purpureo-caerulea having a highly

restricted distribution in the Darling/Mamre District of

the southwestern Cape, occurring on sandy gravel flats

in renosterveld, and flowering later in the season, from

mid-October to mid-November (Duncan 1988b). Like

those of L. nardousbergensis , the shiny black, globose

seeds of L. purpureo-caerulea fall into the group having

smooth, rudimentary strophioles.

Distribution and habitat: Lachenalia nardousber-

gensis has a limited distribution in the Fynbos Biome

in the northwestern part of the Northern Cape and the

northwestern part of the Western Cape, extending from

the Bokkeveld Plateau at Nieuwoudtville, southwest

to the Nardousberge Plateau southeast of Klawer, and

southeast to the Middelburg Plateau at the northern end

of the Cederberg (Figure 5). The plants occur in areas of

154

Bothalia 36,2 (2006)

FIGURE 6. — Lachenalia nardous-

bergensis, Duncan 198

(NBG). A, inflorescence and

leaves; B, bulb; C, single flo-

wer; D, 1/s flower. Scale bars:

A-D, 10 mm. Artist: Vicki

Thomas.

fairly level, high-lying ground, in deep red or yellowish

brown sand, growing as scattered individuals or in small

colonies in fynbos vegetation, among low succulent

undergrowth or restios.

Additional specimens examined

NORTHERN CAPE. — 3119 (Calvinia); Farm Oorlogskloof,

Nieuwoudtville, (-AC), 1-10-1973, Barker 10890 (NBG); Farm Glen

Ridge, Nieuwoudtville, (-AC), 12-9-1985, Duncan 198 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp); Nardousberge

Escarpment, (-DD), 17-9-1968, Chater s.n. (NBG). 3219 (Wuppertal):

2 miles [3.2 km] on road Brandewynrivier to Calvinia, (-AA), 27-9-

1970, Barker 10724 (NBG); Middelberg Plateau, (-AC), 4-8-1985,

Nicklin 179 (NBG).

ACKNOWLEDGEMENTS

We extend our thanks to Dr Ted Oliver for translating

the diagnoses into Latin, Ms Vicki Thomas and Dr John

Manning for preparing the line drawings, and Adam

Harrower and Gordon Summerfield for their assistance

in the field.

REFERENCES

BAKER, J.G. 1897. Lachenalia Jacq. In W.T. Thistleton-Dyer, Flora

capensis 6: 421-436. Reeve, London.

BARKER, W.F. 1 949. Planlae Novae Africanae. Journal of South Afri-

can Botany 15: 37-39.

DUNCAN, G.D. 1988a. Lachenalia arbuthnotiae. The Flowering

Plants of Africa 50: t. 1961.

DUNCAN, G.D. 1988b. Lachenalia purpureo-caerulea. The Lachena-

Bothalia 36,2 (2006)

155

lia handbook. Annals of Kirstenbosch Botanical Gardens 17: 56,

57. National Botanical Gardens, Cape Town.

DUNCAN, G.D. 1993. Lachenalia thomasiae. The Flowering Plants of

Africa 52: t. 2061.

DUNCAN, G.D. 1996. Four new species and one new subspecies of

Lachenalia (Hyacinthaceae) from arid areas of South Africa.

Bothalia 26:1-9.

DUNCAN. G.D. 1997. Five new species of Lachenalia (Hyacinthace-

ae) from arid areas of South Africa. Bothalia 21: 7-15.

DUNCAN, G.D. 1998. Five new species of Lachenalia (Hyacinthace-

ae) from arid areas of Namibia and South Africa. Bothalia 28:

131-139.

DUNCAN, G.D. 2003. Lachenalia peersii. Curtis’s Botanical Maga-

zine 20: 202-207.

DUNCAN, G.D. 2005. Character variation and a cladistic analysis of

the genus Lachenalia Jacqf. ex Murray (Hyacinthaceae: Mas-

sonieae). M.Sc. thesis, University of KwaZulu-Natal, Pieterma-

ritzburg.

DUNCAN, G.D. & EDWARDS, T.J. 2002. Anew species of Lachenalia

from Namaqualand, South Africa (Hyacinthaceae: Massonieae).

Bothalia 32: 190-192.

DUNCAN, G.D., EDWARDS, T.J. & MITCHELL, A. 2005. Character

variation and a cladistic analysis of the genus Lachenalia Jacq.f.

ex Murray (Hyacinthaceae). Acta Horticulturae 673: 113-120.

.

I .

Bothalia 36,2: 157-162 (2006)

New species and taxonomic changes within Pentaschistis (Danthonioi-

deae, Poaceae) from Western Cape, South Africa

C.A. GALLEY* and H.P. LINDER*

Keywords: Cape Floristic Region. Danthonioideae, fynbos, new species, Pentaschistis (Nees) Stapf, Poaceae, South Africa

ABSTRACT

Three new species of Pentaschistis (Nees) Stapf are described from the Cape Floristic Region, P trifida, P. clavata and P.

horrida. The former has been collected from inland ranges of the Cape Fold Belt, from the Cederberg to the Groot Swartberg,

the last two each from single sites in the Koue Bokkeveld: P. clavata on the wetter western border, and P. horrida on the

Baviaansberg. Pentaschistis juncifolia Stapf is re-instated, a species from the coastal plains (Hardeveld) between Bredasdorp

and Riversdale, which had been included in P. eriostoma (Nees) Stapf.

INTRODUCTION

The remarkable field work of Ms Esterhuysen resulted

in the description of many new species of Pentaschistis

(Nees) Stapf (Linder & Ellis 1990), but ongoing field-

work and taxonomic research on the grasses of the Cape

Floristic Region (CFR) (Goldblatt 1978) is resulting in

the occasional discovery of new grass species.

Pentaschistis comprises 66 recognized species and

is the most species-rich grass genus in the CFR (Linder

1989; Goldblatt & Manning 2000). Most species are

endemic to or centred in this region. Additionally there

are eleven species in the Drakensberg region, seven spe-

cies in the tropical east African mountains from Malawi

to Ethiopia, one species on Mt Cameroon, three species

in Madagascar, one endemic species in the Imatong

Mountains, Sudan, and one endemic species on St Paul

and Amsterdam Islands in the South Indian Ocean. A few

of the more drought-tolerant species occur in the drier

northwest of South Africa in the Greater Cape Floristic

Region (Jurgens 1991).

Despite the recent revision of the genus (Linder & Ellis

1990), there are still a number of taxonomic problems

remaining. Some species show enormous variation over

habitat and geographical range, such as the Pentaschistis

pictigluma complex in eastern Africa (Phillips 1994),

and the P pallida complex in the CFR. There have also

been a steady rate of discovery and descriptions of new

species in the genus over the past 1 5 years (Phillips 1986;

Linder & Ellis 1990; Phillips 1995). Here we describe a

further three new species. Pentaschistis trifida was found

by the first author on a recent collecting trip; P. clavata

was found by the late Mr Hugh Taylor and recognized

as new by Mrs Lyn Fish of PRE. P. horrida had already

been recognized as distinctive by Dr Roger Ellis and the

second author (Linder & Ellis 1990), but they included

it under P. rigidissima. After extensive field work we

were convinced that it is indeed a distinct species, for

the reasons given below. We also resurrect Pentaschistis

juncifolia Stapf to species level following observation of

differences in ecology and habit between this taxon and

P. eriostoma (Nees) Stapf.

* Institute of Systematic Botany, University of Zurich, Zollikerstrasse

107, CH-8008 Zurich, Switzerland.

MS. received: 2005-07-20.

Pentaschistis trifida C.A. Galley, sp. nov.,

P. trisetae (Thunb.) Stapf similis sed differt spiculis

unifloris, glandibus ellipticis, dimensionibus spiculorum

multo parvioribus: gluma inferiore 4. 5-5. 5 mm (non

15-18 mm) longa, lemma 2.4^4. 8 mm (non 6-8 mm)

longa, arista lemmatis 18-24 mm (non 25-35 mm)

longa, glumis omnino parce puberulis, lemmatibus inter

venas dense villosis, paleis glabris, et ramis floriferis

trifurcatis.

TYPE. — Western Cape, 3319 (Ceres): Baviaansberg,

north of the Hex River Mountains, 1 050 m, 33°12'

14.6"S, 19°37'04.5" E, (-BA), 11 Nov. 2004, C.A. Galley

577 (ZH, holo.; BOL, K, NBG, PRE).

Perennial; single or few stems. Culms 80-200 mm

tall; nodes glabrous; basal sheaths white shiny, persist-

ent; prophylls truncate, often bilobed, keels remaining

parallel to apex; innovation buds intravaginal. Glands

multicellular, elliptical, linear, common on inflorescence

branches. Leaves basal; sheaths red-purple, sparsely

puberulous; sheath mouth glabrous; blades puberulous;

ligules 0.25 mm long fringe of hairs; blades 20-50 x

0.5-1 mm, rolled; apex acute; margins scaberulous; old

blades persisting, entire. Inflorescence widely paniculate,

50-75 x 40-60 mm, open, trichotomously branched,

with 15-35 spikelets; pedicels mostly erect, longer or

shorter than spikelets; inflorescence branches longer than

spikelets, glabrous; nodes glabrous. Spikelets 1 -flowered.

Lower glume 4. 5-5. 5 mm long, longer than floret, acute,

1 -veined, puberulous, pale yellow with purplish base

and green tip; margins same texture as body of glume.

Upper glume similar to lower glume but slightly shorter.

Lemma 2.4-2. 8 mm long, hairs villous, scattered on

back of lemma between veins, veins 5; apex lobed, lobes

acute, 0.5 mm long, lobe setae 6-8 mm long; lemma awn

geniculate, 18-24 mm long, column twisted, 5.5 mm

long; veins 5. Palea linear to lorate, 3 x 0.4 mm, acute, as

long as lemma, keels parallel, glabrous. Callus up to 0.8

mm long, densely hairy with short hairs at base and long

hairs at top. Anthers 0.9-1. 2 mm long. Ovary stalked,

turbinate; styles two. Flowering time : late October to

early November. Figure 1A-H.

Etymology r. the specific epithet trifida is named after

the unique trichotomous branching pattern in parts of the

inflorescence; in the rest of the genus the branches are

paired.

158

Bothalia 36,2 (2006)

FIGURE I -Pentaschistis trifida , Galley 577: A, whole plant; B, glume pair; C, floret showing long callus; D, inflorescence stem showing glands;

E, lemma; F, palea with callus; G, anther; H, caryopsis. Pentaschistis clavata. Galley 567: I, whole plant; J, spikelet showing clavate hairs;

K, leaf showing tubercle-based hairs; L, lemma; M, palea; N, anther; O, ovary and styles. Scale bars: A, I, 10 mm; B, F-H, J-O, 1 mm; C,

E, 0.7 mm; D, 0.5 mm. Artist: Claire Linder-Smith.

Bothalia 36.2 (2006)

159

TABLE 1. — Comparison between Pentaschistis trifida and other species

Diagnostic characteristics : Pentaschistis trifida

resembles P. triset a (Thunb.) Stapf by having reduced

leaves and a long lemma awn in relation to the spikelet.

It resembles P. pusilla (Nees) H.P.Linder and P. clavata

(described here) by the single floret per spikelet, and P.

caulescens H.P.Linder, as both species occupy disturbed

habitats and have reduced leaves with red/purple leaf

sheaths. However, it can be easily distinguished from

these four species by the characteristics shown in Table

1. Unique to this new species is the elongated callus, the

trifurcating inflorescence branches and the elliptically

shaped linear-type glands on the inflorescence branches

(Figure ID), which contrast clearly with the purple

branches.

Distribution and habitat, this species was collected

from a shaded disturbed (pathside) habitat of deep, sandy

soil derived from Table Mountain sandstone in a fairly

arid area on the eastern side of the Baviaansberg (Figure

2). Although this is perhaps a more opportunistic habitat

typical of an annual species [e.g. P. airoides (Nees) Stapf

subsp. airoides habitat] this plant is a perennial; the

highly reduced leaves suggest that the water requirement

of this species is, however, probably low. Although

locally common, this was the only population found in

the vicinity.

Pentaschistis clavata C. A. Galley, sp. nov., ab P.

pusilla (Nees) H.P.Linder differt pilis clavatis lemmatis,

lamina foliorum setosa, involuta.

TYPE. — Western Cape, 3219 (Wuppertal): Koue

Bokkeveld Mountains south of Hexberg, on the Farm

De Boom, 1 212 m, 32°44'32.1"S, 19°1T35.2"E, (-BA),

FIGURE 2. — Distribution of Pentaschistis clavata , •; P. horrida, * : P.

trifida , ■; and P. juncifolia, A.

7 Nov. 2004, C.A. Galley 567 (ZH, holo.; BOL, E, G, K,

MO, NBG, NSW, NY, PRE, S, UPS, W).

Plants perennial, forming neat, rounded cushions, up

to 100 mm in diam. Multicellular glands absent. Culms

80-200 mm tall; nodes puberulous; basal sheaths white

shiny, persistent; prophylls truncate, often bilobed, keels

remaining parallel to apex; innovation buds intravaginal.

Leaves cauline; sheaths glabrous or with sparse, 2 mm

long tubercle-based hairs; sheath mouth with a ring of

2 mm long, stiff bristles; ligules 0.2 mm long fringes

of hair; blades 40-50 x 1 mm, expanded at base, with

sparse, scattered 2 mm long tubercle-based bristles; apex

keeled, acute; margins smooth; old blades persisting,

entire. Inflorescence widely paniculate, 20-35 x 15-35

mm, open, with 10-20 spikelets; pedicels mostly erect,

longer than spikelets; inflorescence branches longer

than spikelets, puberulous; nodes villous with long erect

hairs. Spikelets 1 -flowered, 2.5 mm long. Lower glume

2. 0-2. 5 mm long, as long as or shorter than floret, acute,

1 -veined, straw-coloured with purplish base; margins

same texture as body of glumes, glabrous. Upper glume

similar to lower glume, but somewhat shorter and nar-

rower. Lemma 2.2-2. 5 mm long, clavate hairs scattered

between veins on back, veins 7, not anastomosing, apex

finely tridentate. Pa/ea linear to lorate, truncate, 2.2 x 0.5

mm, glabrous or with few clavate hairs found between

the keels, as long as lemma; keels parallel, glabrous.

Lodicules without microhairs or bristles; obtriangular,

3-veined. Anthers 1.6-1. 9 mm long. Ovaiy stalked,

turbinate; styles two. Flowering time : November and

December. Figure 1 1— O.

Etymology, the species epithet clavata is named after

the clavate lemma hairs.

Diagnostic characteristics : Pentaschistis clavata

resembles P. pusilla in the single-flowered spikelet.

This is generally a rare feature in the Danthonioideae,

and within Pentaschistis is only found in two species, P.

pusilla (Linder & Ellis 1990) and P. trifida described in

this paper. P. clavata has a finely tridentate lemma apex,

similar to that of P. pusilla , and in addition the generally

soft, orthophyllous leaves and weakly perennial habit

are reminiscent of that found in P. pusilla. However,

there are several convincing differences (Table 2). The

new species has scattered, stout, clavate hairs on the

lemma back, the only known case of clavate hairs in

Pentaschistis. Clavate hairs occur in several other genera

of the Danthonioideae, such as Karroochloa (Conert

160

Bothalia 36,2 (2006)

TABLE 2. — Comparison between Pentaschistis clavata and P. pusilla

& Tiirpe 1969), Schismus (Conert & Ttirpe 1974) and

Tribolium (Linder & Davidse 1997), but these genera are

all rather distantly related to Pentaschistis.

Distribution and habitat : this new species is known

from a single locality above De Boom in the Koue

Bokkeveld, at an altitude of 1 270 m (Figure 2). The

species was first collected by Mr FI ugh Taylor in 1989,

describing the habitat as ‘streambank in moist soil with

moss’. Further investigations in 1998 and 2005 showed

that the species is quite common in the area, occurring

on damp sand derived from Table Mountain sandstone as

well as streambanks in wet moss. The habitat therefore

differs slightly from that of P. pusilla. It is possible that

the species is much more widespread — these mountains

are still poorly explored, and more populations may be

found in similar habitats.

Other specimen examined

WESTERN CAPE. — 3219 (Wuppertal): south of Hexberg, Koue

Bokkeveld Mountains, 1 270 m, (-CC), H.C. Taylor 12095 (PRE).

Pentaschistis horrida C.A. Galley, sp. nov., P.

rigidissimae Pilger ex H.P. Linder similis sed differt

longitudine majore culmorum et foliorum, ramificatione

caulium, indumento superficiei foliorum et dimensione

majore antherarum, 2. 1-2.8 mm (non 1.4-1. 8 mm).

TYPE. — Western Cape, 3319 (Ceres): Baviaansberg,

north of the Hex River Mountains, 1 900 m, (-BA), 26

Oct. 1997, H.P. Linder 6799 (ZH, holo.; BOL, E, G, K,

MO, NBG, NSW, PRE).

Perennial, caespitose or mat-forming; older plants

forming ‘fairy rings’ (with the centre of the ring dying),

up to several metres diam. Multicellular glands absent.

Culms 15CM100 mm tall; nodes glabrous; basal sheaths

white shiny, persistent; prophylls truncate, often bilobed,

upper margin ciliate or bristly, keels remaining parallel

to apex, scaberulous or dentate, extended into 20 mm

long awns; innovation buds intravaginal. Leaves cauline;

sheaths glabrous; sheath mouth glabrous; ligules 0. 5-1.0

mm long fringe of hairs; blades 150-200 x 1 mm,

rolled, rigid, with sparse, villous hairs at base of blade;

apex pungent; margins smooth; old blades persisting,

entire. Inflorescence widely paniculate, 70-90 x 25-50

mm, open at anthesis (soon closing again), with 30-60

spikelets; pedicels mostly patent, shorter than spikelets;

inflorescence branches as long as or longer than spike-

lets, scaberulous; nodes sparsely hairy, puberulous to

villous. Spikelets 2-flowered, 7.5 mm long. Lower glume

6. 5-7. 5 mm long, longer than florets, acute to acuminate,

1 -veined, pale green; margins scaberulous, same texture

as body of glumes; upper glume similar to lower glume.

Lemma 2.25-3.0 mm long, with scattered hairs on the

back; veins 7, anastomosing near apex; lemma lobes

acute, 1 . 1 mm long, shorter than the lemma body; lemma

awn geniculate, 8 mm long, column twisted, 3 mm long

thus as long as lemma lobe setae. Palea linear to lorate,

3.5 x 0.5 mm, apex rounded to acute or bi-lobed, longer

than lemma, glabrous; keels parallel, glabrous. Lodicules

without microhairs or bristles, obtriangular, 3-veined.

Anthers 2. 1-2.8 mm long. Ovaiy stalked, turbinate;

styles two. Flowering time : October to November. Figure

3.

Etymology’’, the species epithet horrida is named after

the prickly, pungent leaves.

Diagnostic characteristics’, the linear inflorescence,

pungent leaves and cushion habit associate this spe-

cies with Pentaschistis rigidissima Pilg. ex H.P.Linder.

Specimens of P. horrida were previously included within

this species and were considered as one extreme of a con-

tinuous gradient of size and spinescent variation (Linder

& Ellis 1990). There are several differences that separate

these species, including their growth form (Table 3).

Although both species may form ‘cushions’ the plant

base differs. The new species forms cushions that expand

in size over time by means of a branching culm system;

this effectively moves the living part of the plant out-

wards from the cushion centre, which eventually dies, so

that a whole plant forms a ‘fairy ring’. P. rigidissima, by

contrast, always forms small tufts, rather like a shaving

brush, and never forms rings. They have been found in

sympatry at several localities, and the morphological and

habit differences were consistently maintained. Lastly,

phylogenetic analysis of chloroplast data places these

two species in different clades (unpubl. data). P. horrida

is sister to P. rosea.

Distribution and habitat’. Pentaschistis horrida has

been collected from the inland ranges of the Cape Fold

Belt, from the Cederberg to the Great Swartberg (Figure

2). All populations occur in soils derived from sand-

stone. These areas are not only dry, but have a more

continental climate with severe frost in winter and fierce

heat and a long dry period in summer. The plants grow

in soil in open vegetation, sometimes on open plains

(such as on the summit of the Rooihoogte Pass over the

Waboomsberg, and on the summit plateau of Wolfberg in

the Cederberg), and sometimes on steep slopes (such as

on the Baviaansberg in the Koue Bokkeveld). P. rigidis-

sima by contrast is found more towards the coastal areas

on the coastal ranges of the Cape Fold Belt, where the

climate is more mesic. There are also microhabitat dif-

ferences. P. rigidissima is restricted to crevices in the

sandstone rocks and boulders, often in shady, cool and

protected places along the sides of boulders or even

underneath them. In contrast, P. horrida is found in

open areas, or at most between boulders, but never in

crevices.

Other specimens examined

WESTERN CAPE.— 3219 (Wuppertal): central Cederberg,

Sleeppad Hut to Sneeukop, 1 500-1 900 m (-AC), H P. Linder 4464

(BOL); Bokkeveld, Tafelberg 5500' (1 676m), (-CD), Esterhuysen

Bothalia 36,2 (2006)

161

FIGURE 3. — Pentaschistis horrida, Linder 6799: A, plant base and lower plant; B, upper plant and inflorescence; C, glume pair; D, florets; E,

lemma showing setae and awn; F, palea with callus; G, ovary and styles. Scale bars: A, B, 10 mm; C-G, 1 mm. Artist: Claire Linder-

Smith.

162

Bothalia 36,2 (2006)

TABLE 3. — Comparison between Pentaschistis horrida and P. rigidis-

sima

3931 (BOL). 3319 (Worcester): top of Rooihoogte Pass, 1 234 m,

33°36'12"S, 19°51'02" E, (-BD), C.A. Galley 374 (BOL, ZH). 3320

(Montagu): south of Matroosberg station, 3800' (1 158m), (-BD),

Acocks 19088 (BOL). 3321 (Ladismith): Towerkop, 1 750 m 33°27'

42"S, 21°13'01" E, (-AC), Linder, Hardy & Moline L7410 (BOL);

Towerkop, Swartberg, (-AC), Esterhuysen 26744 (BOL); Sewe Weeks

Poort, 2 000 m, (-AD), Linder 5486 (BOL); Montagu to Matroosberg,

3700' (1 128m), MRL 370 (BOL); Swartberg Pass, (-BD), Barker 999

(BOL): Great Swartberg, 1 800 m (-BD), Linder 4571 (BOL). 3322

(Prince Albert): Zwartbergen, 4000' (1 219m), (-AC), Bolus 11673

(BOL). 3323 (Uniondale): Kouga Mountains, 5500' (1 676m), (-DA),

Esterhuysen 27974 (BOL).

Pentaschistis juncifolia

Pentaschistis juncifolia was originally described by

Stapf in Flora capensis (1899), and is distinct from P.

eriostoma (Nees) Stapf based on the absence of densely

woolly or villous leaf sheaths. Linder & Ellis (1990)

sank this species into P. eriostoma on the basis of similar

spikelets and inflorescences, regarding it as a local fonn

of the very variable P. eriostoma. After recent fieldwork,

however, we are now convinced that these taxa are indeed

distinct, and that P. juncifolia should be recognized as a

separate species. The most striking difference between

the two taxa is the absence of a woolly indumentum at

the leaf sheath mouth in P. juncifolia. Aside from this,

P. juncifolia can be recognized as distinct from P. erios-

toma in the field as it forms much more neatly defined

tussocks and has a much paler inflorescence. The habitats

of the two taxa differ, with P. eriostoma occurring on

shales and sandstones and P. juncifolia associated with

eroded silcrete surfaces on the coastal plains between

Bredasdorp and Riversdale. They were observed in sym-

patry near a silcrete outcrop (Verkykerskop) along the

road from Malgas to Heidelberg (34°13'25" S, 20°43T4"

E) and were both abundant, maintaining morphological

and habitat distinctions. This morphological and eco-

logical evidence is backed up by phylogenetic analysis of

chloroplast data where P. eriostoma and P. juncifolia are

separate on the cladogram and P. juncifolia is sister to a

large clade that includes P. eriostoma (Galley & Linder

in review). P. juncifolia has been collected in flower in

early October.

Specimens examined

WESTERN CAPE. — 3420 (Bredasdorp): Swellendam Division.

Buffelsjachtsrivier, 1000-2000 ft (305-610 m), Zeyher 4545 (K!); road

from De Hoop to Malahas, 141 m, 34”21'44"S, 20°28'45" E, (-AD),

7 Oct. 2003, C.A. Galley 341 (BOL, ZH). 3421 (Riversdale): hills

near Zoetmelksrivier, Burchell 6761 (K! ); hills near Zoetmelksrivier,

BurcheU 6750 (K, lectotype!).

ACKNOWLEDGEMENTS

This research was funded by the University of Zurich, the

Swiss Science Foundation, the Swiss Academy of Sciences

and the ‘Georges und Antoine Claraz-Schenkungk Thanks

to L. Fish, and N. Bergh, A. Verboom and the Bolus her-

barium for assistance with field work. Also many thanks to

Claire Linder-Smith for the elegant illustrations.

REFERENCES

CONERT, H.J. & TURPE, A.M. 1969. Karroochloa , eine neue

Gattung der Gramineen (Poaceae, Arundinoideae: Danthonieae).

Senckenbergiana Biol ogle a 50: 289-318.

CONERT, H.J. & TURPE, A.M. 1974. Revision der Gattung Schismus

(Poaceae: Arundiniodeae: Danthonieae). Abhandlungen der

Senckenbergischen Naturforschenden Gesellschaft 532: 1-81.

GALLEY, C. & LINDER, H.P. (in review). The phylogeny of the

Pentaschistis clade (Danthonioideae, Poaceae) and the evolution

and loss of complex characters: multicellular glands, evolution.

GOLDBLATT, P. 1978. An analysis of the flora of southern Africa: its

characteristics, relationships, and origins. Annals of the Missouri

Botanical Garden 65: 369 — 436.

GOLDBLATT, P. & MANNING, J.C. 2000. Cape plants. A conspectus

of the Cape flora of South Africa. Strelitzia 9. National Botanical

Institute, Cape Town & Missouri Botanical Garden, St. Louis.

JURGENS, N. 1991. A new approach to the Namib Region. Vegetatio

97: 21-38.

LINDER, H.P. 1989. Grasses in the Cape Floristic Region: phyto-

geographical implications. South African Journal of Science 85:

502-505.

LINDER, H.P. & DAVIDSE, G. 1997. The systematics of Tribolium

Desv. (Danthonieae: Poaceae). Botanische Jahrbiiche 119:

445-507.

LINDER. H.P. & ELLIS, R.P. 1990. A revision of Pentaschistis

(Arundineae: Poaceae). Contributions from the Bolus Herbarium

12: 1-124.

PHILLIPS, S.M. 1986. Four new grasses from northeast tropical

Africa. Kew Bulletin 41: 1027-1030.

PHILLIPS, S.M. 1994. Variation in the Pentaschistis pictigluma com-

plex (Gramineae). In J.H. Seyani & A.C. Chikuni, Proceedings

of the XIII Plenary Meeting of AETFAT. Zomba. Malawi, 2-11

April 1991: 359-372. National Herbarium and Botanic Gardens

of Malawi, Zomba.

PHILLIPS, S.M. 1995. A new species of Pentaschistis (Gramineae)

from Ethiopia. Kew Bulletin 50: 615-617.

STAPF, O. 1899. Graminae. In W.T. Thiselton-Dyer, Flora capensis 1:

310-750. Reeve, London.

Bothalia 36,2: 163-174(2006)

Notes on African plants

VARIOUS AUTHORS

MESEMBRYANTHEMACEAE

THE IDENTITY OF RUSCHIA PROMONTORJI

INTRODUCTION

RELATIONSHIPS

The entity Ruschia promontorii, a Red Listed plant en-

demic to the Cape Peninsula, was first described by H.M.L.

Bolus (1929). Type material (Rohland NBG1 543/20) was

illustrated by Mary Page (unpublished plate, BOL). Much

later, Hartmann (1998b) placed this taxon in synonomy

under Amphibolia hutchinsonii , and upon discovering an

earlier homonym, Hartmann (2001) placed A. hutchinso-

nii as a synonym of Amphibolia laevis.

Desmet (2000) mentions this taxon as belonging to

Antimima , namely A. promontorii {'promontorii' and

‘ promontori' ) noting differences in distribution ranges

of Amphibolia laevis and this taxon. However, no formal

transfer was undertaken.

Initially I believed this taxon to be an Acrodon, but

upon closer examination concluded that certain features

mentioned below exclude it from that genus.

DISTRIBUTION AND ECOLOGY

This decumbent, mat-forming mesemb is found only

on the Cape Peninsula where it grows on the slopes of

Chapman’s Peak, Cape Point and a few other localities

(Figure 1). The plant colonizes steep, rocky scree slopes

often devoid of plant cover, with little competition from

other plants, but well above the salt spray of the ocean.

The lithology comprises sediments of the Cape

Supergroup, with sandstones of the Peninsula Formation

(up to 1 550 m thick) underlain by characteristic red-

dish sandstone and shale of the Graafwater Formation.

Granites of the Cape Peninsula Pluton intrude the pre-

Cape rocks in places (SACS 1980).

FIGURE 1. — Distribution of Ruschia promontorii.

The creeping, mat-forming habit (Figure 2 A) and

triquetrous leaves of Ruschia promontorii (Figure

2B) exclude it from Amphibolia, where it was placed

by Hartmann (1996, 2001) under the synonomy of

Amphibolia laevis. The latter is shrubby with decum-

bent branches and club-shaped leaves with rounded tips

(Hartmann 1996). Rectangular valve wings, a character

always present in capsules of Amphibolia , are absent in

R. promontorii. The type of R. promontorii, however,

lacks fruit, which has possibly led to the erroneous place-

ment of R. promontorii in the genus Amphibolia.

Ruschia promontorii was first thought to belong to

the genus Acrodon because of its mat-forming habit,

FIGURE 2. — Ruschia promontorii. A, creeping, mat-forming habit,

Burgoyne 6703 ; B, leaves with a smooth epidermis and epider-

mal thickening at margin. Scale bar: B, 5 mm.

164

Bothalia 36,2 (2006)

FIGURE 3. — Flowers of Ruschia promontorii, Burgoyne 6703, show-

ing filamentous staminodes gathered into a cone and petals with

a darker central stripe.

triquetrous leaves, bract and flower characters (Figure 3)

and white pollen (Table 1). However, its seeds lack bacu-

lae (Figure 4A) and microbaculae (Figure 4B) preventing

its transfer to Acrodon in which baculae and microbacu-

lae are always present (Glen 1986). Although seed char-

acters are of limited taxonomic importance, it has been

found that the presence of baculae and microbaculae is

constant within the genus Acrodon (Glen 1986).

Comparisons between the capsule morphology (Figure

5A-D) of Ruschia sarmentosa the type of the subgenus

Sarmentosae, R. promontorii, Acrodon and Amphibolia

are given in Table 1 .

Ruschia promontorii shares more characters with R.

sarmentosa than with any other taxon. The creeping,

sannentose habit and presence of similar closing bodies

and absence of valve wings in the capsule are charac-

ters shared with the subgenus Sarmentosae as a whole.

Closing devices present on the distal ends of the cover-

ing membranes are present in both R. promontorii and R.

sarmentosa.

Differences between these two species (Table 1)

include habitat preference, with Ruschia sarmentosa

growing in flat sandy areas close to the sea. The flowers

of R. sarmentosa are borne in cymes of three, whereas

those of R. promontorii are mainly single or in cymes

of three, the central one maturing while the other two

mostly abort. Leaves of R. sarmentosa are longer (30-45

mm long) and are clustered at the nodes, and the side

branches stand upright at the long intemodes, whereas

TABLE 1. — Comparison between Ruschia sarmentosa (subgenus Sarmentosae), R. promontorii , Acrodon and Amphibolia based on selected

characters

Bothalia 36,2 (2006)

165

FIGURE 4. — Seeds of Ruschia promontorii. A, no baculae, Bargoyne

6703; B. wax platelets on surface. Scale bars: A, 100 pm; B,

10 pm.

in R. promontorii the shorter leaves (18-35 mm long)

are usually borne one pair per node. Intemodes are much

shorter and side branches are mostly decumbent.

Herbarium specimens of the two species are easy

to distinguish, as once pressed and dried, the leaves

of Ruschia promontorii are pale grey resulting from a

prominent wax covering, whereas those of R. sarmentosa

remain dark green.

As previously mentioned, Desmet (2000) noted that

this taxon was better placed in Antimima. However, this

species does not belong to the genus Antimima , as the

criteria for inclusion to that genus, such as capsules open-

ing fully, and large closing bodies completely blocking

the distal ends of locules (Hartmann 1998a), are not met.

Capsules of Ruschia promontorii never open fully (Figure

5A), the closing bodies being hook-shaped and not large,

therefore excluding it from the genus Antimima.

It is interesting to note that the origin of the type

material of Ruschia sarmentosa described by Haworth

(1812) is from ‘Nova Hollandia’, a term used to describe

the Australian continent at that time. Salm Dyck (1840)

states: ‘M sarmentosum was sent to the good Haworth in

the year 1 806 by Mr Donn and it was said to have arisen

from Australian seeds. M. simile arose at Kew in 1819,

most certainly from Cape seed; from which [fact] doubt

remains in my mind as to which homeland M. sarmen-

tosum is to be attributed to. It is not possible to separate

these two plants, as is clear from the description' (trans-

lated from Latin by H.F. Glen).

The description by Haworth and an unpublished plate

done by J. Duncanson at Kew (numbered 127), dated

April 1826, predates the publication of Salm-Dyck

(1840) by 14 years and therefore would have been mate-

rial seen by Haworth, thus constituting the earliest type.

The holotype of Ruschia sarmentosa. therefore, is the

unpublished plate by J. Duncanson at Kew (numbered

127, dated April 1826) and not that of Salm-Dyck 1840:

§17 f 3.

CONSERVATION STATUS

This Red List species is restricted to the Cape Penin-

sula. It was first known only from the Cape of Good Hope

Nature Reserve (Smuts 1994) growing on cliffs below

the Cape Point lighthouse. Later its distribution was

extended from the lighthouse to further along the False

Bay coast also growing in rocky cliff situations. Listed

as endangered, using the old IUCN categories (Hilton

Taylor 1996), it has now been found growing on the

slopes of Chapmans Peak. This species has recently been

placed under further threat as the slopes of Chapman’s

Peak Drive have been cleared of loose material to make

it safer for motorists. It is not known how many plants

of Ruschia promontorii have been affected by this activ-

ity, but as R. promontorii tends to grow on scree slopes

which are naturally disturbed due to rock turnover, more

niches may have been created for these plants resulting

from the clearing of loose rock. The situation should be

monitored to observe the effect this activity has had on

the plant population.

The current Red List status of this species using the

latest criteria (IUCN 2001) is Endangered, B1 ab (I, II,

III, IV & V) +2ab (I, II, III, IV) (Klak & Victor 2001) as

it has an area of occupancy of less than 1 km2, while the

extent of occurrence is less than 100 km2.

Pressure from developers to build more homes along

the coast where this species grows is a constant threat to

the habitat. Only a few highly fragmented populations

exist, but plants are healthy and produce numerous flow-

ers that are pollinated, often producing an abundance of

seed. Consecutive years of low rainfall may affect the

number of flowers and seed produced. Fires have rav-

aged the area over the last few years but it is not known

whether this has influenced the population in any way.

All populations visited showed a healthy demography

with juvenile, mature and older plants present.

166

Bothalia 36,2 (2006)

FIGURE 5. — Fruit. A, Ruschia promontorii, Burgoyne 6703\ B, R. sarmentosa, Burgoyne 6702 ; C, Amphibolic hutchinsonii, Burgoyne, Smith &

Van Wyk 8160’, D, Acrodon purpureostylus, Burgoyne 6730. Scale bars: 10 mm. CB, closing bodies; CM. covering membranes; EK, expand-

ing keels; VC, valve covers; VW, valve wings.

CONCLUSION

Ruschia promontorii is better placed in the genus

Ruschia on the basis of the characters shared with R.

sarmentosa (subgenus Sarmentosae). However, it shares

a number of characters with Acrodon.

TAXONOMY

Ruschia promontorii L. Bolus, Notes on Mesembrian-

themum and allied genera 2: 121 (1929).

Mesembryanthemum pansifolium N.E.Br. (manuscript name)

Antimima promontorii Desmet 64 (2000), nom. illeg. as ‘promon-

toni’ and ‘promontori’.

Specimens examined

Ruschia promontorii

WESTERN CAPE. — 3418 (Simon’s Town): Simon’s Town, (-AB),

Burgoyne 6703, 7475 (PRE), Van Jaarsveld 3 1 63 , 3172 (PRE); Simon’s

Town, (-AD), Van Jaarsveld 3954, 12905 (PRE).

Ruschia sarmentosa

WESTERN CAPE. — 3318 (Cape Town): Bellville, (-DC), Hilton-

Taylor 4989 (PRE). 3418 (Simon’s Town): Simon’s Town, (-AB),

Burgoyne 6702 (PRE); Simon’s Town, (-AD), Van Jaarsveld 3633,

12921 (PRE).

Amphibolia laevis

WESTERN CAPE.— 3017 (Hondeklipbaai): Hondeklipbaai, (-

AD), Burgoyne 7542 (PRE), Burgoyne, Smith & Van Wyk 8160, (PRE).

3117 (Lepelfontein): Baievlei, (-BD), Van Rooyen 2231, (PRE).

3118 (Vanrhynsdorp): Doringbaai, (-CC), Boucher, 4049 (PRE).

3218 (Clanwilliam): Lambert’s Bay, (-AB), O’Callaghan, Van Wyk

& Morley 140 (PRE); Velddrif, (-CC), Marloth 7961 (PRE), Van

Jaarsveld 5685 (PRE). 3318 (Cape Town): Cape Town, (-CD), Van

Jaarsveld 3942 (PRE).

Acrodon purpureostylus

WESTERN CAPE.— 3320 (Montagu): Montagu, (-CC), Burgoyne

6730 (PRE).

ACKNOWLEDGEMENTS

The curator of Bolus Herbarium is thanked for

the opportunity to study specimens at the Herbarium

and for the use of colour plates. Western Cape Nature

Conservation and Annelise le Roux in particular are

thanked for issuing permits to collect Ruschia promon-

torii at short notice. Hugh Glen is thanked for translation

Bothalia 36,2 (2006)

167

of the Latin. Dick Brummit, Kew, is thanked for tracing

the N.E. Brown manuscript names.

REFERENCES

BOLUS, H.M.L. 1929. Notes on Mesembiyanthemum and allied gen-

era, part 2: 121

DESMET, P. 2000. Captions to slides for MSG. Mesemb Study Group

Bulletin 4: 64.

GLEN, H.F. 1986. Numerical taxonomic studies in the subtribe

Ruschiinae (Mesembryanthemaceae) — Astridia , Acrodon and

Ebracteola. Bothalia 16: 203-226.

HARTMANN, H.E.K. 1996. Miscellaneous taxonomic notes on

Aizoaceae. Bradleya 14: 20-56.

HARTMANN, H.E.K. 1998a. New combinations in Antimima

(Ruschioideae, Aizoaceae) from southern Africa. Bothalia 28:

67-82.

HARTMANN, H.E.K. 1998b. New combinations in Ruschioideae,

based on studies in Ruschia (Aizoaceae). Bradleya 16: 44-91.

HARTMANN, H.E.K. 2001. Illustrated handbook of succulent plants:

Aizoaceae A-E. Springer- Verlag, Berlin.

HAWORTH, A.H. 1812. Synopsis plantarum succulentarum: 283.

Taylor, London.

HILTON TAYLOR. C. 1996. Red Data List of southern African plants.

Strelitzia 4. National Botanical Institute, Pretoria.

IUCN 2001. IUCN Red List Categories and Criteria', version 3.1.

IUCN Species Survival Commission. IUCN, Gland, Switzerland

and Cambridge, UK.

KLAK, C. & VICTOR, J.E. 2001. Red List data. South African

National Biodiversity Institute, Pretoria. Unpublished.

S ALM-REIFFERSCHEIDT-D YCK, J. Duke of. 1836-1863. Monogra-

phia generum Aloe et Mesembryanthemi. I-V: Amz, Dusseldorf;

V, VI: Henry & Cohen, Bonn; VII: M. Cohen & fil., Bonn.

SMUTS, L.M. 1994. ISEP (Information System for Endangered Plants)

file. Cape Nature Conservation. Unpublished.

SOUTH AFRICAN COMMITTEE FOR STRATIGRAPHY (SACS)

1980. Stratigraphy of South Africa. Part 1 (compiled by L.E.

Kent). Lithostratigraphy of the Republic of South Africa, South

West Africa/Namibia, and the Republics of Bophuthatswana,

Transkei and Venda. Handbook of the Geological Survey of

South Africa No. 8: 201-235. Government Printer, Pretoria.

P.M. BURGOYNE*

* National Herbarium, South African National Biodiversity Institute,

Private Bag XI 01, 0001 Pretoria.

MS. received: 2000-10-27.

‘FORSYTH ( FL . 1835)’: A PHANTOM BOTANICAL COLLECTOR AT THE CAPE OF GOOD HOPE

EXPLAINED

At the instigation of Dr Peter Goldblatt, I have

attempted to establish the veracity of the inclusion of a

collector named Forsyth by Gunn & Codd (1981 : 157) in

their roster of botanical exploration of southern Africa.

This individual was credited as follows: ‘Collected at the

Cape 1835; also in Indonesia and Mascarenes Is.’. Those

authors cited Hedge & Lamond (1970) as their source.

In the index to collectors in the herbarium of the Royal

Botanic Garden, Edinburgh, Hedge & Lamond (1970)

noted that the Forsyth specimens were not dated and

that they were included in the herbarium of Archibald

Menzies.

Among African specimens associated with this

Forsyth are the type specimens of Fen-aria macro-

chlamys (Baker) Goldblatt & J.C. Manning (Goldblatt &

Manning 2004) and Babiana spiralis Baker (Goldblatt

& Manning 2005: 72). Both species are endemic to

Namaqualand, but no collector named Forsyth is known

to have visited Namaqualand before 1835.

Concerning the first taxon, Baker (1876: 338) noted

that he had seen only one specimen ‘without a note of

its special locality’, and gave the collection data in the

following form: ‘C. B. Spei. Forsyth in Herb. Bentham!’.

Regarding the second taxon, Baker (1892: 183) reported

that he had described it ‘from specimens in the herbar-

ium of Forsyth’, adding later (Baker 1896) that the two

specimens ‘without locality ... [were] in the Forsyth’s

herbarium, purchased by Mr. Bentham in 1835.’

Forsyth is not an unfamiliar name in British horti-

culture— Forsythia (Oleaceae) was named by Vahl to

honour William Forsyth (1737-1804), a distinguished

Scottish horticulturist and one of the founders of the

(Royal) Horticultural Society of London. His son, also

named William (1772-1835), was a nurseryman in

London throughout his life. Desmond & Ellwood (1994)

list several others with this surname, but none of these

individuals is recorded as having been a plant collector

at the Cape of Good Hope. There is no entry for Forsyth

in Lanjouw & Stafleu (1957).

The date 1835 and Baker’s allusion to specimens from

Forsyth’s herbarium purchased by Bentham in 1835 pro-

vided the clues to unravelling this enigma.

The younger William Forsyth, who died on 28 July

1835, ‘had an excellent horticultural library’ (Desmond

& Ellwood 1994: 257) which was sold by auction by

Sotheby in London during November 1835 (Sotheby

1835; Chalmers-Hunt 1976). An annotated copy of the

catalogue of the sale (Sotheby 1835) survives in the

Botany Library, The Natural History Musuem, London,

and indicates that 2 597 lots were auctioned includ-

ing this, the penultimate, one: ‘2596. A very extensive

and well-arranged hortus siccus contained in ten cases,

forming a range, with five mahogany doors’. William

Pamplin, publisher, bookseller and dealer in botanical

specimens, who annotated the catalogue, noted that it

was sold for £17 6s. 6d.

George Bentham’s manuscript diary, which survives

in the Archives of the Royal Botanic Gardens, Kew, con-

firms that Bentham attended this auction, buying a few

books as well as Forsyth’s entire herbarium:

18 November 1835: ‘... to Sotheby’s sale rooms to the auction of

Forsyth’s books, but got there too late for today’s sale ...’.

20 November 1835: ‘... to Forsyth’s sale where bid for several

books but they all went too dear ...’.

21 November 1835: ‘... then for the whole afternoon to Forsyth’s

sale where bought Vahl’s works & two or three others also, [and] the

herbarium which appears to contain a large quantity of W. Indian plants

for which I gave 16'A guineas’.

168

Bothalia 36,2 (2006)

Sixteen and a half guineas equal £17 6s. 6d. A copy of

Vahl's Symbolae botanicae (1790-1794), inscribed ‘Will

F. 1800’ on the verso of the title page, and with a book

plate ‘Presented by George Bentham Esq 1854’, is in the

Linnean Collection in the library at the Royal Botanic

Gardens, Kew (J. Flanagan pers. comm., 19 September

2005). Pamplin noted that it was sold for £1 17s. Od.

On 23 November Bentham went to Sotheby’s premis-

es ‘... to settle for my Saturday’s purchase which had

taken to the Hort Soc in order there to get rid of super-

fluous paper — then at the Flort Soc looking over sev-

eral parcels which appear very satisfactory’. Bentham

spent the following two weeks at the premises of the

Elorticultural Society, of which he was the Honorary

Secretary (Stevens 2003), sorting Forsyth’s herbarium;

by 28 November he had packed ‘25 large bundles’ which

he arranged to ‘bring home’. Bentham must subsequently

have shared duplicates with other botanists including

Archibald Menzies — that is the most likely explanation

for presence of ‘Forsyth 1835’ specimens in Menzies’

herbarium in the Royal Botanic Garden, Edinburgh.

It may safely be concluded, therefore, that annota-

tions on southern African specimens indicating Forsyth

and 1835 came from this miscellaneous collection which

Bentham purchased on 21 November 1835, and that they

should not be read as indicating a collector and collec-

tion date.

Who actually collected these specimens must remain

a matter of conjecture, unless other evidence, in the form

of contemporary annotations, occurs on the herbarium

sheets.

REFERENCES

BAKER. J.G. 1876. New Gladioleae. Journal of Botany, British and

foreign , new ser. 5: 333-339.

BAKER, J.G. 1 892. Handbook of the Irideae. Bell, London.

BAKER, J.G. 1896. Babiana spiralis. Jn W.T. Thiselton-Dyer, Flora

capensis 6: 111. Reeve, London.

CHALMERS-HUNT, J.M. 1976. Natural history auctions 1700-1972.

A register of sales in the British Isles. London.

DESMOND, R.G.C. & ELLWOOD, C. 1994. Dictionary of British and

Irish botanists and horticulturists. London.

GOLDBLATT, P. & MANNING, J.C. 2004. New species of Ixia

(Crocoideae) and Moraea (Iridoideae), and taxonomic notes on

some other African Iridaceae. Novon 14: 288-298.

GOLDBLATT, P. & MANNING, J.C. 2005. Taxonomic notes on

Babiana and Ferraria in arid southern Africa (Iridaceae).

Bothalia 35: 71-74.

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

HEDGE, I.C. & LAMOND, J.M. 1970. Index of collectors in the

Edinburgh Herbarium. Edinburgh.

LANJOUW, J. & STAFLEU, F.A. 1957. Index herbariorum, part 2

Collectors E-M. Regnum vegetabile 9.

SOTHEBY. 1835. Catalogue of the united libraries of William Forsyth

... and of the late William Forsyth ... together with his ... collec-

tion of unpublished manuscripts ... which will be sold by auction

by Mr. Sotheby and Son ... Strand on ... November 11th 1835

and nine following days. London.

STEVENS, P.F. 2003. George Bentham ( 1 800-1 884): the life of a bota-

nist’s botanist. Archives of natural history 30,2: 1 89-202.

E.C. NELSON*

* Tippitiwitchet Cottage, Hall Road, Outwell, Wisbech,

Cambridgeshire, PE 14 8PE, UK.

MS. received: 2005-10-12.

MALVACEAE

A NEW SPECIES OF CORCHORUS IN SOUTHERN AFRICA

INTRODUCTION

Corchorus L., a genus of Malvaceae s.l. (previously

in Tiliaceae) comprises ± 100 species widely distributed

throughout the tropics, subtropics and warm temper-

ate regions of the world (Wild 1984; Edmonds 1990;

Heywood 1993; Bayer & Kubitzki 2003). Species of

Corchorus are either annual or perennial herbs or small

shrubs, sometimes arising from a woody rootstock, with

simple to stellate hairs (Wild 1984; Edmonds 1990).

Corchorus species occupy a wide range of habitats

ranging from swamps and riverine conditions, to forest,

bushland, woodland and open savanna, and cultivated

fields. They also colonize diverse soil types ranging from

sand, clay, black turf and loam, on quartz and granite to

limestone (Edmonds 1990; Lebrun & Stork 2003).

Recent molecular phylogenetic analyses of the chloro-

plast genes atpB, rbcL and ndhF have placed the Tiliaceae

among the core Mai vales sensu lato (Angiosperm

Phylogeny Group 1998). Within this group, analyses

show Tiliaceae to be paraphyletic, with Corchorus

excluded from Tiliaceae sensu stricto (Alverson el al.

1998, 1999; Bayer et al. 1 999; Whitlock et al. 2001).

Since Linnaeus established the first four Corchorus

species, C. capsularis L., C. hirsutus L., C. olitorius

L., and C. siliquosus L. in Species pi ant arum (Linnaeus

1753) and Genera plantarum (Linnaeus 1754), (C. olitor-

ius being the type species of the genus), many additional

species have been described. The last floristic account

of Corchorus in southern Africa was published by Wild

in 1984 in which 15 species were recognized. Despite

this publication, confusion regarding species limits and

relationships within the genus is still not resolved or fully

understood. This is evident from the herbarium studies

which have shown that many specimens have been mis-

identified and that there are different opinions as to what

constitutes real species.

Current taxonomic treatments do not allow for effec-

tive identification of certain species, resulting in curator-

ial problems in all southern African herbaria. The genus

is currently under revision. A new species of Corchorus

was discovered during field excursions and the study of

morphological variation within the genus. The new spe-

cies together with C. trilocularis L. and C. confusus Wild

are very similar in floral and vegetative morphology and

Bothalia 36,2 (2006)

169

it is difficult to identify them from herbarium specimens

when capsules are not available. Corchorus trilocularis

L. and the related species form a distinct group distin-

guished by the ± straight capsules and fruiting pedicels.

Corchorus trilocularis is an annual, whereas C. confusus

and the new species are perennials. These three spe-

cies form a group that is confined to the eastern parts

of Africa extending up to Tanzania, with C. trilocularis

occurring in other tropical and subtropical regions of

tropical Africa, Madagascar, Asia, Australia and the

Middle East. DNA studies also support this grouping

(Moeaha et al. in press).

Corchorus argillicola M.J. Moeaha & PJ.D. Winter,

sp. nov.

TYPE.— Limpopo, 2328 (Mokopane): ± 2 km N of

Mapela, ± 30 km N of Mokopane (Potgietersrus), (-DD),

4 June 2002, P.J.D. Winter 5764 (PRE).

C. confuso Wild et C. triloculari L. capsuli pedicel-

lisque fructificantibus plusminusve rectis affinis sed cap-

suli omnino hirsutis non porcatis ab ambabus speciebus

differt.

Perennial herb with prostrate or spreading stems from

a woody rootstock; branches with pubescence all around

stem, or sometimes with line of short hairs on one side

of stem only. Leaf blade narrowly ovate, 30-50(-80)

x 1 0—1 8(— 25) mm, obtuse in basal leaves, crenate or

serrate-crenate, sometimes with pair of basal setae,

pubescent on both surfaces, especially on nerves, hairs

not tubercle-based; petiole up to 15 mm long, pubes-

cent; stipules up to 7 mm long, setaceous, pubescent.

Inflorescence of ( 1 )2^1-flowered cymes opposite upper

leaves; peduncles 3-5(-20) mm long, pubescent, pedi-

cels up to 3— 5(— 7) mm long, pubescent. Sepals narrowly

lanceolate, up to 8 mm long, pubescent abaxially. Petals

yellow, narrowly obovate, up to 8 x 5 mm, with short

ciliate claw, androgynophore ± 0.5 mm long, with an

annulus. Stamens numerous. Ovary cylindrical, densely

pubescent; style ± 2.5 mm long, slender, glabrous, stigma

cuspidate. Capsules up to 90 mm long; fruiting pedicel

straight, densely or sparsely pubescent, without longi-

tudinal ridges. Seeds numerous, black, ± 1.5 x 1 mm.

Figure 6.

Diagnostic characters and relationships', this species

was included in Corchorus confusus by Wild (1958), but

FIGURE 6. — Corchorus argillicola. A, flowering branch, x 1;B, base of plant showing smaller basal leaves with rounder apices, x 1; C, flower, x

2; D. base of leaf showing seta, x 4; E, capsules, x 1; F, apically oriented palmate hairs found on capsules, x 50. Artist: G. Condy.

170

Bothalia 36,2 (2006)

the comparison of live and herbarium material indicates

that two distinct taxa are involved and that the populations

from deep black turf or clay soils represent a distinct spe-

cies, ecologically separated from C. confusus. Corchorus

argillicola is related to C. confusus and C. trilocularis by

the more or less straight capsules and fruiting pedicels.

Herbarium specimens of these three species may easily

be confused. Wild (1958, 1984) hypothesized the con-

fusion of C. confusus and C. trilocularis in herbarium

material to the possibility that C. confusus is of hybrid

origin with C. trilocularis and C. asplenifolius as puta-

tive parents. However, field visits have indicated that

these species are quite different. The new species differs

mainly in the distribution of trichomes, which occur all

over the surface on the capsules, and in the lack of lon-

gitudinal ridges on the capsules at all stages of develop-

ment. In contrast, the capsules of C. confusus are angular,

and sparsely scabrous on the angles or glabrous, whereas

in C. trilocularis they have trichomes confined to the

ridges and on the angles, and are otherwise glabrous.

Furthermore, this new taxon is a perennial, whereas C.

trilocularis is an annual weed of cultivation.

Distribution and habitat'. Corchorus argillicola is

restricted to deep black turf and clayey soils below

1 300 m. It occurs mainly in the Limpopo basin, but

extends to the Chipinge District in SE Zimbabwe and

to Maputaland in northern KwaZulu-Natal (Figure 7).

It seems to be an ecologically specialized relative to C.

confusus , which occurs in a much wider range of condi-

tions including rocky and sandy habitats in savanna and

Grassland Biomes at altitudes up to 1 600 m.

Etymology, the specific epithet argillicola refers to the

clay soils to which this species is restricted.

Specimens examined

ZIMBABWE. — 1929 (Gweru): Shangani Dist., (-CD), Feiertag

s.n. (SRGH). 2032 (Chipinge): 7 km S of Chisumbanja, (-CC), Pope

& Miiller 1532 (SRGH); Chipinge Dist., Phipps 99 (SRGH); 6 km

S of Rusongo Hill, (-CD), Biegel, Pope & Russell 4903 (SRGH).

BOTSWANA. — 2226 (Serowe): 60 km NW of Serowe, (-AB). Wild &

Drummond 7286 (PRE, SRGH).

LIMPOPO. — 2328 (Baltimore): on Farm Wellust 426 LR. (-CA),

Schmidt 174 (PRU); ± 2 km N of Mapela, ± 30 km N of Mokopane

[Potgietersrus], (-DD), P.J.D. Winter 5764 (PRE). 2329 Polokwane

[Petersburg]: Ben-Lavin Nature Reserve, (-BB), James 262 (PRU);

near turn-off to Chuenespoort from Polokwane, (-CD), M.J. Moeaha

47 (UNIN); Turfloop Water Course, N of Suiferkuil Experimental

Farm, (-DC), M.J. Moeaha 13 (UNIN). 2428 (Nylstroom): S of

Rooisloot, (-BB), P.J.D. Winter s.n. (UNIN). 2429 (Zebediela): 8 km

W of Moletlane, (-AD), M.J. Moeaha 49 (UNIN). 2430 (Pilgrim’s

Rest): 1 mile N of Mokopane (-AA), Gillett 2807 (PRE); on road

between Tzaneen and Hoedspruit, above Selati River, (-BA), M.J.

Moeaha 27 (UNIN); Steelpoort, on Farms Driekop and Winterveld,

(-CA), Siebert & Du Plessis 1550 (PRU).

NORTH-WEST. — 2527 (Rustenburg): Brits, as exiting town area

to the east on R566, just north of Brits Abbatoir and south of railway

shunter, (-DB), S.P. Bester 6056 (PRE).

MPUMALANGA. — 2531 (Komatipoort): Nelspruit, on road be-

tween Malelane and Komatipoort, 5 km E of turn-off to Coopersdal,

(-BC), M.J. Moeaha 36 (UNIN); Nelspruit, in sugarcane fields ± 5 km

S of Komatipoort, (-BD), M.J. Moeaha 34 (UNIN).

SWAZILAND. — 2631 (Mbabane): Gollel, Hlatikulu, (-AD).

Compton 29425 (PRE).

KWAZULU-NATAL. — 2832 (Mtubatuba): False Bay Park, S sec-

tor, (-AB), Ward 7723 (PRE).

ACKNOWLEDGEMENTS

I would like to thank my supervisor Pieter Winter

for drawing my attention to the taxonomy of the genus

Corchorus and for his continuous encouragement and

support during the early stages of my research, and also

for his inspiration and constructive criticism, guidance,

enthusiasm, genuine interest in my work and for guid-

ance and participation with field work. I am very grateful

to the staff and curators of the herbaria mentioned above

for making their material available for this study. Special

thanks are due to Dr O.A. Leistner for the Latin transla-

tion of the diagnosis, Ms Gill Condy for line drawings

and Ms Hester Steyn for the distribution map. I am also

grateful to Dr J.C. Manning for his valuable comments

on the manuscript. A taxonomic analysis of Corchorus is

a registered M.Sc project at the University of Limpopo.

REFERENCES

ALVERSON, W.S., KAROL, K.G., BAUM, D.A., CHASE, M.W..

SWENSEN, S.M., McCOURT, K.J. & SYTSMA, K.J. 1998.

Circumscription of the core Malvales and relationships to other

Rosidae: evidence from rbcL sequence data. American Journal

of Botany 85: 857-887.

ALVERSON, W.S., WHITLOCK, B.A., NYFFELER, R„ BAYER, C.

& BAUM, D.A. 1999. Phylogeny of the core Malvales: evi-

dence from ndh F sequence data. American Journal of Botany

86: 1474-1486.

ANGIOSPERM PHYLOGENY GROUP (APG). 1998. An ordinal

classification for the families of flowering plants. Annals of the

Missouri Botanical Garden 82: 247-277.

BAYER, C„ FAY, M.F., DE BRUIJN, A.Y., SAVOLAINEN, V.,

MORTON, C.M., KUBITZKI, K„ ALVERSON, W.S. &

CHASE, M.W. 1999. Support for the expanded family con-

cept of Malvaceae within a recircumscribed order Malvales: a

combined analysis of plastid atpB and rbcL DNA sequences.

Botanical Journal of the Linnean Society 129: 267-303.

BAYER, C. & KUBITZKI, K. 2003. Malvaceae. In K. Kubitzki &

C. Bayer. The families and genera of vascular plants, vol. 5.

Springer, Berlin.

Bothalia 36,2 (2006)

171

EDMONDS, J.M. 1990. Herbarium survey of African Corchorus

L. species. Systematic and ecogeographic studies on crop

genepools 4. International Board for Plant Genetic Resources,

Rome.

HEYWOOD, V.H. 1993. Flowering plants of the world. Mayflower,

New York.

LEBRUN, J.P. & STORK, A.L. 2003. Tropical African flowering

plants. Ecology and distribution 1. Conservatoire et Jardin

Botaniques, Geneve.

LINNAEUS, C. 1753. Species plantarum, vol. 1. Holmiae,

Stockholm.

LINNAEUS, C. 1754. Genera plantarum, edn 5. Lugdoni Batavorum,

Stockholm.

MOEAHA. M.J, WINTER, P.J.D., VAN DER BANK, M. &

GROBLER, P. in press. Molecular phylogenetic analysis of

selected Corchorus species (Malvaceae s.l.) in southern Africa.

International Journal of Plant Sciences.

WHITLOCK, B.A., BAYER. C. & BAUM, D.A. 2001. Phylogenetic

relationships and floral evolution of the Byttnerioideae

(‘Sterculiaceae’ or Malvaceae s.l.) based on sequences of the

chloroplast gene, ndhF. Systematic Botany 26: 420-437.

WILD, H. 1958. Notes and new records of African flowering plants

(Tiliaceae). Bothalia 7: 422-424.

WILD, H. 1984. Tiliaceae. In O.A. Leistner, Flora of southern Africa

21: 32-42.

M.J. MOEAHA*

*National Herbarium, South African National Biodiversity Institute,

Private Bag X 101. Pretoria, 0001.

MS. received: 2005-08-11.

ASTERACEAE

ARCTOTIS DECURRENS (ARCTOTIDEAE), THE CORRECT NAME FOR A. MERXMUELLERI AND A. SCULLYI

The genus Arctotis L. comprises an estimated 50-60

species and belongs to the tribe Arctotideae. The repre-

sentatives of this genus are indigenous to the Flora of

southern Africa ( FSA ) region, with the highest concen-

tration of taxa in Western Cape. One species, A. venusta

Norl., has the widest distribution, extending to Angola

and southern Zimbabwe. Some species are in cultivation

and have been introduced into other countries, e.g. A.

stoechadifolia P.J.Bergius (Pope 1992).

The taxonomy of Arctotis is extremely chaotic, owing

largely to a great duplicity of published names and confu-

sion over the correct application of many names. The last

revision of the genus was undertaken by Lewin (1922),

who used mainly fruit, involucre and pappus characters to

distinguish between species. Unfortunately, he failed to

resolve many of the nomenclatural problems and added

to the confusion by misinterpreting some species and

publishing additional superfluous names. Furthermore,

recent morphological and molecular studies (McKenzie

et a!. 2005, 2006) indicate that the circumscription of the

genus also requires reappraisal.

A monographic revision of Arctotis is being under-

taken by the first author at Rhodes University, to resolve

the taxonomic disarray within the genus. In preparation

for the Conspectus of the Trans-Cape Succulent Karoo

Flora of South Africa (Snijman in prep.), two species,

A. merxmuelleri Friedrich and A. scullyi Diimmer, from

Northern Cape, and a third species, A. decurrens Jacq., of

uncertain provenance, were studied.

In a series of lavishly illustrated works published

around the turn of the nineteenth century, N.J. von

Jacquin and his son J.F. von Jacquin described 29 Arctotis

species (excluding those names now known to apply to

species in other genera). Of relevance to the present

investigation, Jacquin (1797) described A. decurrens

from cultivated material 'Ex Promontorio bonae Spei'

grown in the Royal Schonbrunn garden, Vienna. The

source, provenance and date of the original collection in

South Africa are unknown. The name A. decurrens Jacq.

has not been typified and application of the name by

previous workers has varied, resulting in uncertainty as

to what species the name applies to and. indeed, whether

it is a valid species. A search of authentic Jacquin

material in the Naturhistorisches Museum Herbarium.

Vienna (W) revealed two specimens (W0006629 and

0006630 ) determined as A. decurrens with labels bear-

ing 'Hort. Schonbr.’ (i.e. Hortus Schonbrunnensis) in

the lower left comer. A third specimen ( W0006628) bore

a label with 'Hort. Bot. Vindob.’ (i.e. Hortus Botanicus

Vindobonensis) in the lower left comer. The Hortus

Schonbrunnensis specimens are likely to have been pre-

pared prior to publication of the name A. decurrens by

Jacquin (1797) and can be considered as possible type

material. They are consistent in all important details

(particularly involucral bract and leaf morphology) with

the illustration and description of A. decurrens in Jacquin

( 1 797). Therefore in all likelihood Jacquin based his con-

cept of A. decurrens on material represented by the sheets

W0006629 and 0006630. Herein the sheet W0006630 is

designated as the lectotype, as it contains the most com-

plete material (three capitula, two also with upper stem

leaves, and six individual lower stem leaves). The sheet

W0006629, comprising a single flowering shoot without

lower stem leaves, is designated an isotype.

The Hortus Botanicus Vindobonensis specimen differs

from the Hortus Schonbrunnensis specimens in possess-

ing leaves with a dense lanate tomentum on the abaxial

surface, and the outer involucral bracts have shorter api-

cal appendages with lanate trichomes on the abaxial sur-

face. Given its origin from different cultivated material

from the Hortus Schonbrunnensis specimens, it cannot

be considered as type material.

Arctotis scullyi was described by Diimmer (1914) prior

to publication of Lewin's (1922) revision of Arctotis, but

Lewin stated that he did not see the publication until

after the end of the First World War. He therefore did not

include A. scullyi within his revision, but referred to the

172

Bothalia 36,2 (2006)

species in a postscript and commented that the informa-

tion on the involucre and cypselae was incomplete. The

precise locality of the holotype of A. scullyi ( W.C . Scully

221) is unknown, as the label only states ‘NamaTand

Minor’ (i.e. Little Namaqualand). For a long time, no

specimens of this species were lodged in the National

Herbarium, Pretoria (PRE). Only recently, a few speci-

mens collected between 1983 and 1998 were submitted,

identified and lodged as A. scullyi in PRE.

Friedrich (1980) described A. merxmuelleri from

material ( H.-C . Friedrich 498 ) collected from ‘sukkulen-

tenreiches Strandveld ostlich Port Nolloth bei 5 Miles’

in Little Namaqualand and cultivated in the Botanischer

Garten Mtinchen. In his description, he added beautiful

illustrations of inter alia the involucral bracts and cypse-

lae. The holotype of A. merxmuelleri is lodged in M, and

isotypes are lodged in BOL, K and PRE. No subsequent

collections have been lodged in PRE.

After close examination of the Jacquin specimens in

W, the illustrations by Friedrich (1980), concentrating

especially on the involucral bracts, and isotypes of A.

merxmuelleri and A. scullyi lodged in BOL and PRE, we

concluded that the names A. decurrens, A. merxmuelleri

and A. scullyi apply to the same species. The earliest

legitimate name is A. decurrens , to which A. merxmuel-

leri and A. scullyi are placed in synonymy herein.

Arctotis decurrens Jacq., Plantarum rariorum

horti caesarei Schoenbrunnensis descriptiones et icons

II. 20. t. 165 (1797). Type: ‘Hort. Schonbr.’ [Hortus

Schonbrunnensis], ex Hb. Jacq. (W, lecto & iso!), here

designated by R.J. McKenzie.

A. scullyi Diimmer: 152 (1914), syn. nov. Type: Northern Cape,

‘Nama’land Minor’ [Little Namaqualand], W.C. Scully 221 (BOL!,

holo; K (scanned image: http://www.kew.org/herbcat/!), PRE!).

A. merxmuelleri Friedrich: 13 (1980), syn. nov. Type: Northern

Cape, 2916BD Little Namaqualand, Port Nolloth, 8-12-1974, H.-C.

Friedrich 498 (M, holo; BOL!, K (scanned image: http://www.kew.

org/herbcat/!), PRE!).

The leaf tomentum appears to vary significantly. They

are most commonly sparsely to densely brownish or

FIGURE 8. — Known distribution of Arctotis decurrens in southern

Africa.

whitish scabrous on both sides and/or glandular-pilose,

but sometimes densely tomentose on both surfaces with

no glands.

This species is seemingly endemic to the Strandveld

of the West Coast (Namaqualand Sandveld) (Figure 8)

where it grows near the coast and further inland on sand

dunes in well-drained, sandy soil in full sun. It seems to

be quite frequent to abundant in this area but is not often

collected. At present it does not seem to be threatened but

its habitat is becoming increasingly under threat of open

mining activities, whereas overgrazing, land use changes,

informal housing and urban development appear to be of

a lesser threat.

Additional specimens examined

NORTHERN CAPE. — 2916 (Port Nolloth): about 10 km east from

Port Nolloth on road to Steinkopf, (-BD), 23-08-1998, Koekemoer

1175 (PRE); 16 km east of Port Nolloth, (-BD), 21-08-1980, A. le

Roux 2703 (BOL); 11 km east of Port Nolloth, (-BD), 14-09-1976, E.

J. van Jaarsveld 1423 (NBG). 2917 (Springbok): Oograbies, (-AC),

27-07-1993, L. van der Walt 274 (PRE); Kleinzee, Schulpfonteinpunt,

(-CA), 30-11-1983, G.K. Theron 3871 (PRE, PRU); Brazil, southeast

of Kleinzee, beside road to Swartlintjiesrivier, (-CC), 5-09-2005, R. J.

McKenzie 1302 (GRA). 3017 (Hondeklipbaai): Swartlintjies River

mouth, (-AD), 16-10-1980, A. le Roux & Parsons 34 (PRE); Hondeklip

Bay, at Aristea shipwreck, (-AD), 17-09-2003, L. Mucina 170903/20

(GRA); coast 4 miles south of Hondeklip Bay, (-AD), 10-1924, A IS.

Pillans 18133 (BOL); sandy hills round Hondeklip Bay, (-AD), 10-

1924, N.S. Pillans 18136 (BOL); ± 103 km from Springbok on sand

road to Hondeklipbaai (near coast), (-AD), 28-09-1986, D. Strydom 10

(PRE). 3117 (Lepelfontein): Brandsebaai, (-BD), 29-09-1992, M. W. van

Rooyen 2140 (PRE, PRU), 1-10-1992, M.W. van Rooyen 2248 (PRE,

PRU). Without precise locality: ‘Hort. Bot. Vindob.' [Hortus Botanicus

Vindobonensis], ex Hb. Jacq., Jacquin s.n. (W); cultivated in National

Botanic Gardens, 09-1920, Anon. (BOL); cultivated at Kirstenbosch

National Botanic Garden, no. 407/18, 04-1922, Phillimore s.n. (BOL);

cultivated at Kirstenbosch National Botanic Garden, no. 407/18, 07-

1922, Phillimore s.n. (BOL).

ACKNOWLEDGEMENTS

We thank the W herbarium (especially Ernst Vitek

and Anton Igersheim) for access to authentic Jacquin

specimens; curators of the BOL, NBG and PRU her-

baria for the loan of specimens; the Rhodes University

Joint Research Committee and the National Research

Foundation of South Africa (grant number 2069059

to N.P. Barker and a Postdoctoral Fellowship to R.J.

McKenzie) for financial support.

REFERENCES

DUMMER, R.A. 1914. A new Arctotis. Journal of Botany 52: 152,

153.

FRIEDRICH, H.-C. 1980. Arctotis merxmuelleri , eine neue Sippe

aus dem kleinen Namaqualand. Mitteilungen der Botanischen

Staatssammlung Miinchen 16, Beiheft: 11-17.

JACQUIN, N.J. VON 1797. Plantarum rariorum horti caesarei

Schoenbrunnensis descriptiones et icons, vol. 2. Wappler,

Viennae; White, Londini; & Luchtmans, Lugduni Batavorum.

LEWIN, K. 1922. Systematische Gliederung und geographische

Verbreitung der Arctotideae-Arctotidinae. Repertorium spe-

cierum novarum regni vegetabilis, Beihefte 11: 1-75.

McKENZIE, R.J., MULLER, E.M., SKINNER, A.K.W., KARIS, P.O.

& BARKER, N.P. 2006. Phylogenetic relationships and generic

Bothalia 36,2 (2006)

173

delimitation in subtribe Arctotidinae (Asteraceae: Arctotideae)

inferred by DNA sequence data from ITS and five chloroplast

regions. American Journal of Botany 93. In press.

mckenzie. r.j., samuel, j„ muller, e.m., skinner, a.k.w.

& BARKER. N.R 2005. Morphology of cypselae in sub-

tribe Arctotidinae (Compositae-Arctotideae) and its taxonomic

implications. Annals of the Missouri Botanical Garden 92:

569-594.

POPE, G.V. 1992. Compositae. In G.V. Pope, Flora zambesiaca 6,1:

1-264. Royal Botanic Gardens, Kew.

SNIJMAN, D. in prep. Conspectus of the Trans-Cape Succulent Karoo

Flora of South Africa.

R.J. McKENZIE*t, P.P.J. HERMAN** and N.P. BARKER*

* Molecular Ecology and Systematics Group, Department of Botany,

Rhodes University, P.O. Box 94, 6140 Grahamstown, South Africa.

** South African National Biodiversity Institute, Private Bag X101,

0001 Pretoria,

f Corresponding author.

MS. received: 2006-04-28.

PTERIDOPHYTA: PTERIDACEAE

CHE1LANTHES DELTOIDEA, A NEW LOCALITY IN GAUTENG, SOUTH AFRICA

Cheilanthes deltoidea Kunze is a small fern with a

short, creeping rhizome covered in narrowly lanceolate,

pale brown scales, thick black roots and old stipe bases.

Fronds are closely spaced and erect. The dark brown,

glabrous stipe is up to 75 mm long. The lamina, 18-100

x 16-110 mm, is triangular in outline, 2-pinnatifid to

deeply 3-pinnatifid and entirely glabrous with obscure

venation. Lamina margins are irregular and entire. Sori

form a marginal line around the lobe apices, with the

indusium continuous (Jacobsen 1983; Burrows 1990).

The species is endemic to southern Africa and occurs

in the western and northwestern parts of Northern Cape,

the Cederberg area in Western Cape and in southern

Namibia. An isolated record from the Waterberg in the

Limpopo Province is also known (Figure 9). It grows

only in sheltered rock crevices on southern aspects,

where it is completely sheltered from sunlight. Here it

forms ribbon-like clusters of densely crowded fronds.

It is found mostly on granite or gneiss rock formations.

During the dry season it shrivels up completely and

revives when sufficient moisture is available (Jacobsen

1983; Burrows 1990).

FIGURE 9. — Distribution of Cheilanthes deltoidea, adapted from the

map in Burrows (1990), with his kind permission: localities

based on specimens kept at PRE and NBG, •; new localities

in Gauteng, ©.

After above-average rainfall in January and early

February 2006, a population of C. deltoidea was discov-

ered in Centurion, Gauteng, on a property that is ear-

marked for development. The population consists of ± 13

plants in a relatively small area in the transition between

Carletonville Dolomite Grassland and Rocky Highveld

Grassland. Plants are very small, no more than 50 mm

tall and grow in mainly southwest-facing soil pockets

and rock crevices in chert rock. The chert outcrops are

associated with dolomite and form part of the Dolomite

Series of the Transvaal System. The fern has only been

found on the chert and not the associated dolomite.

Upon further investigation of a similar site adjacent

to the Rietvlei Nature Reserve, another population of

several plants was found. Unfortunately this site is also

earmarked for development. On this site, plants grow

in northwest-facing rock crevices on a single isolated

chert boulder. These plants seem to be somewhat smaller

and more stunted than those on the other site, perhaps

because they are to some extent more exposed. More

populations of this fern were subsequently found on two

other sites nearby that are to be developed.

Since the above-mentioned populations are threatened

by development, rocky outcrops in the Rietvlei Nature

Reserve were searched, with the hope of finding this spe-

cies inside the reserve. However, since the geology of the

reserve falls within the Pretoria Series of the Transvaal

System, the specific chert rock on which the fern grows

at the other sites was not found and no population of this

fem could be located in the reserve. Follow-up visits

will be carried out and one of the rangers has agreed to

keep searching for the fem in the reserve. Fortunately a

healthy population was later found on the grounds of the

Smuts House Museum.

Owing to its small size and the fact that it shrivels

up completely during dry times, it is possible that many

other populations of this fem have been overlooked in

the past. It is only now, after exceptionally good rains,

and while the plants are still green, that they are easily

spotted. It is unfortunate that much of the suitable habi-

tat, where populations could be found, has already been

destroyed through massive urban expansion.

The specimens from these new localities will be

examined further and compared to material from the

174

Bothalia 36,2 (2006)

distribution range of the species in Namaqualand, as well

as with the available specimens from the Waterberg. The

preliminary view is that it warrants description as a new

infraspecific taxon.

GAUTENG. — 2528 (Pretoria): Centurion, Portion 107 on Farm

Doomkloof 391-JR, (-CC), P. Lemmer 623 (PRE), Portion 198 (part of

remainder of Portion 335), R.R. Klopper, J. Nel & A. Nel 216 (PRE);

Centurion, Irene, Farm Doomkloof 391-JR, rocky ridges between

Smutskoppie and M57, (-CC), A.E. van Wyk 13630, 13653 (PRU);

Centurion, Irene, Smuts House Museum, on koppie close to monument,

(-CC), R.R. Klopper & A. IV. Klopper 217 (PRE); Centurion, Farm

Rietvallei 377-JR, adjacent to Rietvlei Nature Reserve, close to where

Olifantsfontein road crosses Sesmylspruit, (-CD), R.R. Klopper &A.W.

Klopper 215 (PRE).

ACKNOWLEDGEMENTS

The authors would like to thank the following people:

Reinhard Lemmer for finding the initial population at

Doomkloof; Koos Roux and John Burrows for con-

firming the identification of the Doomkloof specimen;

Matthys Dippenaar for confirming the rock type on

which the fern grows; John Burrows for providing per-

mission to use the distribution map of this species from

his book Southern African ferns and fern allies (1990);

Riaan Marais for giving permission to search for the fern

in the Rietvlei Nature Reserve; Cecil Labuschagne for

his time and for taking us to the rocky outcrops in the

Rietvlei Nature Reserve; Ate Berga for taking the time to

look for and find the fern at the Smuts House Museum;

Emelia Baumgartner for giving permission to collect a

specimen on the property of the Smuts House Museum.

We are also grateful to two anonymous referees for help-

ful comments on improving the manuscript.

REFERENCES

BURROWS, J.E. 1990. Southern African ferns and fern allies.

Frandsen Publishers, Sandton.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Afi-ica.

Butterworths, Durban.

R.R. KLOPPER*, P. LEMMER** and J. NEL***

* National Herbarium, South African National Biodiversity Institute,

Private Bag X101, 0001 Pretoria.

** Galago Ventures, P.O. Box 912878, 0127 Silverton, South Africa.

*** Fern Society of Southern Africa, P.O. Box 73125, 0040 Lynn-

wood Ridge, Pretoria.

MS. received: 2006-04-04.

ASPHODELACEAE

CORRECTIONS TO THE EPONYMY AND GEOGRAPHICAL DISTRIBUTION OF ALOE VANROOYENII

For the sake of providing appropriate recognition to

the discoverer of a plant, or those who draw the atten-

tion of systematists to the existence of new taxonomic

entities, taxa are often named for such key individuals.

However, not all relevant biographical information is

always documented by the nomenclaturist, much often

being presumed without regard for possible misinter-

pretation by subsequent botanists. Aloe greenii Baker

represents an example of a name about which uncertainty

lingers regarding the identity of the person commemo-

rated (Reynolds 1950; Van Wyk & Smith 2004), some

130 years following its circumscription (Baker 1880:

165). Accordingly, and as a necessary courtesy, we cor-

FIGURE 10. — Known distribution range of Aloe vanrooyenii.

rect the recent etymological explanation of the specific

epithet chosen for Aloe vanrooyenii Gideon F.Sm. &

N.R.Crouch (Smith & Crouch 2006). These authors

inadvertently gave a Mr Gert van Rooyen as the person

honoured; the plant actually commemorates Mr Pieter

van Rooyen of Greytown who stimulated and facilitated

further taxonomic studies of this unique maculate aloe.

Additionally, we provide a more complete distribution

map for this KwaZulu-Natal endemic (Figure 10), based

on specimens housed at NU and NH.

REFERENCES

BAKER. J.G. 1 880. A synopsis of Aloineae and Yuccoideae. Journal of

the Linnaean Society 18: 148-241.

REYNOLDS, G.W. 1950. The aloes of South Africa. The Trustees of

the Aloes of South Africa Book Fund, Johannesburg.

SMITH, G.F. & CROUCH, N.R. 2006. Asphodelaceae. Aloe vanrooy-

enii: a distinctive new maculate aloe from KwaZulu-Natal,

South Africa. Bothalia 36: 73-75.

VAN WYK, B-E. & SMITH, G. 2004. Guide to the aloes of South

Africa, edn 2. Briza Publications, Pretoria.

G.F. SMITH* & N.R. CROUCH**

* South African National Biodiversity Institute, Private Bag X101,

0001 Pretoria/ Schweickerdt Herbarium, Department of Botany,

University of Pretoria, 0002 Pretoria.

** Ethnobotany Unit, South African National Biodiversity Institute,

P.O. Box 52099, 4007 Berea Road / School of Chemistry, University

of KwaZulu-Natal, 4041 Durban, South Africa.

MS. received: 2006-08-11.

Bothalia 36,2: 175-189 (2006)

Patterns of plant diversity and endemism in Namibia

P. CRAVEN* and P. VORSTER**

Keywords: Namibia, phytogeography, plant endemism

ABSTRACT

Species richness, endemism and areas that are rich in both species and endemic species were assessed and mapped for

Namibia. High species diversity corresponds with zones where species overlap. These are particularly obvious where there

are altitudinal variations and in high-lying areas. The endemic flora of Namibia is rich and diverse. An estimated 16%

of the total plant species in Namibia are endemic to the country. Endemics are in a wide variety of families and sixteen

genera are endemic. Factors that increase the likelihood of endemism are mountains, hot deserts, diversity of substrates

and microclimates. The distribution of plants endemic to Namibia was arranged in three different ways. Firstly, based on

a grid count with the phytogeographic value of the species being equal, overall endemism was mapped. Secondly, range

restricted plant species were mapped individually and those with congruent distribution patterns were combined. Thirdly,

localities that are important for very range-restricted species were identified. The resulting maps of endemism and diversity

were compared and found to correspond in many localities. When overall endemism is compared with overall diversity,

rich localities may consist of endemic species with wide ranges. The other methods identify important localities with their

own distinctive complement of species.

INTRODUCTION

Species diversity was traditionally measured by count-

ing the number of different species recorded in a specific

area or grid (Linder 2001). The ‘weight' of the species

was not taken into consideration. Today various measures

of diversity have been proposed that give greater value to

species that are taxonomically, geographically, ecologi-

cally or economically distinct, but no ‘best’ method for

mapping species diversity has yet been found (Craven

2002b). The development of floristic databanks allows

quick and efficient retrieval of phytogeographic data

that can produce computerized distribution maps. This

study used the computerized data of specimens housed

in the National Herbarium of Namibia (WIND) and the

National Herbarium in Pretoria (PRE) to survey distribu-

tion patterns of plant diversity, overall endemism and

centres of endemism and diversity on a quarter-degree

scale in Namibia. The endemic flora were also analysed

and factors that may have contributed to the resulting

distribution patterns were discussed briefly when evi-

dent from the use of map overlays. Known centres of

endemism were not redefined, only species with similar

patterns of distribution were recorded.

Caldecott et al. (1996) separated the current knowl-

edge of biodiversity into global, regional, national,

ecoregional and site information. Patterns of diversity in

Namibia have been shown on continental-scale diversity

maps (Mutke et al. 2001) and included in region-based

studies such as those of Goldblatt (1978), Cowling et al.

(1989) and Gibbs Russell (1985, 1987). The account of

Linder (2001) on patterns of plant species endemism and

richness for the African flora does not include arid areas

such as most of Namibia.

The first national assessment and map of relative

species richness in Namibia was that of Maggs et al.

* To whom correspondence should be addressed: RO. Box 399,

Omaruru, Namibia.

** Botany Department, University of Stellenbosch, Private Bag XI,

7602 Matieland, Stellenbosch.

MS. received: 2004-07-29.

(1994). It was based on distributional data per magiste-

rial district following Merxmuller (1966-1972), as well

as other literature. Species diversity was re-assessed for

the Biodiversity Country and mapped on half-degree grid

squares (Maggs 1998; Maggs et al. 1998). This study

updates the underlying data used in Maggs (1998) on a

finer scale.

A taxon is endemic if confined to a particular area

(Major 1988) which may be large or small. Clearly, data

on endemism would be more useful if given by floristic

province rather than political divisions (Major 1988;

Van Wyk & Smith 200 1 ), but datasets between different

countries are seldom compatible in quality or quantity.

Due to the fact that Namibia is home to a considerable

number of endemics with adequate data, this assessment

only reviews species limited to Namibia (Figure 1).

An endemic is therefore defined here as a taxon that is

restricted to within the political borders of Namibia. Taxa

that extend marginally into another region, i.e. beyond

the political borders of Namibia are referred to as near-

endemic.

Centres of outstanding species diversity and endemism

such as the Kaokoveld and Gariep have long attracted

attention, but their boundaries, floristic elements and

origins remain fairly sketchy. Different approaches and

methodologies have also contributed to the centres

(sometimes with the same name) not being compatible

or comparable (Van Wyk & Smith 2001; Craven 2002b).

Stott (1981) suggested that the process consists of stages

i.e. after taxonomic study (essential for elucidating

closely related taxonomic units) using specimens, spe-

cies distributions are plotted and areas of congruence

identified. The plants are then arranged into recognizable

groups, which on further analysis would identify phyto-

geographic centres determined by a high concentration

of taxa with restricted distributions. This approach was

pioneered in southern Africa by Weimarck (1941) and is

found in taxonomic literature, for example in Nordenstam

(1969) and Hilliard (1994). The approach looks at the

geographical ranges of species regardless of their growth

form or other factors such as topography and present

176

Bothalia 36,2 (2006)

FIGURE 1. — Namibia, its neigh-

bours, capital city and some

important mountains and

towns.

climate, and identifies particular geographical areas

inhabited by species that are restricted to these ranges.

Once such a centre is recognized, explanations are sought

on how they may have arisen (e.g. past climate) and how

they are being maintained. This knowledge is fundamen-

tal to understanding the origin, migration, and speciation

of plants and is essential for developing strategies for

biological conservation. This study did not attempt to

redefine the centres presently known for Namibia, but

attributes a number of new species to the centres.

Maps presenting overall patterns of diversity and

endemism in Namibia have been used to identify regions

of importance for conservation (Simmons et al. 1998;

Mendelsohn et al. 2002). This paper shows two other

methods of mapping important areas for endemics, which

should also be taken into consideration.

MATERIALS AND METHODS

Database

Georeferenced specimens on the specimen database

(SPMNDB) in WIND provided the grid-diversity count.

This database has ± 120 000 specimens and includes

specimens housed in PRE that were collected in Namibia.

All specimens of higher plants were used in the evalua-

tion. A sizable proportion was georeferenced following

the quarter-degree square system of Edwards & Leistner

(1971). Gaps in coverage due to collecting biases and

database input errors were corrected where possible

(Craven 2002b). The number of species present in each

quarter-degree square was calculated and mapped.

Distribution data for endemic species were obtained

from the specimen dataset as well as literature sources.

Records for endemic species were found in 722 out of

over I 200 quarter-degree squares in Namibia. These

were variously arranged and mapped: 1 , endemic fami-

lies, genera and species individually; 2, the number of

endemic species per quarter-degree square; 3, the number

of quarter-degree squares in which species occurred; and

4, areas where species restricted to only one or two quar-

ter-degree squares were found.

Degree scale

Quarter-degree squares were chosen for the grid

scale in order to show patterns of distribution on as fine

a resolution as possible and allowing small or more

localized centres of diversity to be apparent. Sufficient

information at that scale is available for Namibia and

the total area or number of quarter-degree square grids

is manageable. Where no or few records were recorded

in a quarter-degree square, the grids were ‘revisited’

and any such ‘empty’ square individually rechecked and

improved. Records from keyword searches and literature

sources, i.e. checklists for specific areas such as those of

Rodin (1985), Giess & Snyman (1986), Hines (1992) and

Clarke (1999) as well as collections of the first author,

were incorporated.

GIS data

Shapefiles produced in ArcView [Environmental Sys-

tems Research Institute (ESRI) 2000] of Namibian fea-

tures such as soils, topography and rainfall by the Agro-

Ecological Zoning Programme (1996-ongoing) and

Atlas of Namibia Project (2002), as well as the positions

of important mountains (adapted from Irish 2002) were

superimposed onto the grid-based plant data. They were

used to draw accurate borders, define localities and help

pinpoint possible reasons for the variations in diversity.

Profiles showing altitude change across the country that

were taken at selected latitudes to cut through various

notable topographic features (Atlas of Namibia Project

Bothalia 36,2 (2006)

177

2002) were also overlaid. The vertical scales have been

exaggerated to highlight the changes in altitude.

The methodology and tables used, and examples of

all stages of the process, is described in more detail in

Craven (2002b).

Inventory

The inventory of species endemic to Namibia, taxo-

nomic limits and nomenclature of species follows Craven

(1999). The numerous sources for endemic status and

updates are listed in Craven (2002b). Genera listed

as endemic follow Leistner (2000). Over 600 species

(Appendix 1) were investigated and eventually, records

for ± 540 endemic spermatophyte species were avail-

able for the analysis and maps. Not all endemics were

included in the analysis because endemism in some

genera appears inflated due to numerous infraspecific

taxa (Maggs et al. 1 998), whereas other genera e.g. Cras-

sula (Crassulaceae), Euphorbia (Euphorbiaceae), Salsola

(Chenopodiaceae), Tetragonia (Aizoaceae) and some in

the family Mesembryanthemaceae need to be revised.

Species that are known to occur just over the border of

the country were not included, e.g. a number of species

in the genus Commiphora (Burseraceae) and family

Acanthaceae, particularly the genus Petalidium.

Maps

Arcview (ESRI 2000) was used to produce the maps.

Overall species richness and overall endemism could

therefore be superimposed to find possible geographical

correlation. Similarly, the shapefiles of the ± 540 species

endemic to Namibia for Craven (2002b) were overlaid

and the resulting maps scrutinized for congruent patterns.

Areas were identified and their species listed, because as

Van Wyk & Smith (2001) point out, such areas will have

their own distinctive complement of species.

Endemic species not included by previous authors

were assigned to known centres, i.e. Kaokoveld, Gariep,

Waterberg-Otavi and Namibia Central and Southern

Highland Centres, where possible. This required looking

at locality data of collected specimens for the last two

centres, as elements occur at higher elevations or on a

specific substrate. The large data set of georeferenced

specimens was also used to find subcentres or more

localized areas of significance.

RESULTS

Overall patterns of species richness and endemism

Species diversity is higher in localities where one veg-

etation type shifts to another. Variations in altitude and

the maximum altitude in any grid are also significantly

related to grid diversity (Figure 2). Only two localities in

the northeast region can attribute high species richness

to high rainfall. Where summer rainfall species overlap

with winter rainfall species, however, there is an increase

in diversity, e.g. the Rosh Pinah area. Another influenc-

ing factor is mist that occurs along the Namibian coast,

which may be responsible for more favourable micro-

climates and increased species richness inland, e.g. the

Namukluft near Rosh Pinah and Aus areas.

Namibian endemics were not found in the northeast,

which forms part of the Zambezian Domain of White

(1983) as this domain continues into countries further

north. The southeast is part of the Kalahari Desert, which

extends into Botswana, and no endemics were found

there because of the general paucity of species and the

12° 14

16° 18° 20° 22°

FIGURE 2. — Grid-diversity for Na-

mibia depicted in five classes

with altitude profiles showing

correlation between changes

in altitude and diversity.

178

Bothalia 36,2 (2006)

FIGURE 3. — Distribution and num-

ber of Namibian endemic gen-

era per quarter-degree grid.

artificial definition of endemic used here. In the rest

of Namibia, the map of overall distribution of endemic

species does not show any particular spatial pattern.

Localities where the quarter-degree squares with the most

endemics occur, are often associated with mountains in

Namibia. The best example is the Brandberg, which also

confirms that endemism increases when mountains are

located in deserts. Substrate-specific endemic plants are

well known and in Namibia, four Jamesbrittenia species

occur only in the limestone of the Waterberg-Otavi area

(Hilliard 1994). Hot deserts have very high endemism

in spite of their limited flora and vegetation and this has

been shown in Namibia by the number of endemic spe-

cies confined to the Namib (Craven 2002b). ArcView

shapefiles for physical features of Namibia, e.g. soils,

aspects of climate, did not show marked patterns of simi-

larity at this level of resolution.

Geographical comparison of the overall pattern of

diversity and that of endemism, as well as areas of

importance for localized endemics and that of overall

endemics, show a degree of congruence. Important local-

ized areas however, do not always coincide. Because

the count is based purely on the number of species or

endemics within that square, areas of richness cannot be

distinguished by a particular combination of plant spe-

cies or endemics. In addition, squares with associated

floral elements cannot be identified. Thus a particularly

rich area may consist of very widespread species.

Families endemic to Namibia

There are no families of higher plants restricted to the

political borders of Namibia. The most well-known fam-

ily that occurs only in southwest Angola and Namibia is

Welwitschiaceae.

Genera endemic to Namibia

The natural ranges of sixteen genera fall with-

in Namibia (Figure 3). They are listed in Table 1

with an indication of the number of quarter-degree

square grids in which they have been found. With the

exception of the genus Ondetia (Asteraceae), most

of the endemic genera occur in the central and west-

ern parts of Namibia. Arthraerua (Amaranthaceae),

Marlothiella (Apiaceae), Eremothamnus (Asteraceae),

Namibia, Synaptophyllum (Mesembryanthemaceae)

and Neoluederitzia (Zygophyllaceae) occur along the

coast, whereas Bavnesia (Apocynaceae) and Namacodon

(Campanulaceae) grow at higher altitudes. Manuleopsis

TABLE 1. — Genera endemic to Namibia and the number of quarter-

degree squares in which they occur

QDS, quarter-degree square.

Bothalia 36,2 (2006)

179

FIGURE 4. — Distribution and num-

ber ofNamibian endemic spe-

cies per quarter-degree grid.

(Scrophulariaceae) is fairly widespread, but generally

found on higher ground. Chamaegigas (Scrophulariaceae)

is a hydrophyte that inhabits pools in granite outcrops in

central Namibia.

All four genera in family Mesembryanthemaceae

occur in the winter rainfall region of southwest Namibia.

Family Apiaceae, with two endemic genera, is of par-

ticular interest as it has very few representatives in spe-

cies and number of individuals in Namibia. A recently

described genus, Baymesia (Bruyns 2000), attests to

the fact that new genera may still be described in

Namibia, especially in certain families that require revi-

sion, e.g. Mesembryanthemaceae. Ondetia occurs close

to Botswana and may eventually be found there (Craven

& Klaassen 1998). It is very closely related to Geigeria

(which is presently under revision) and often mistaken

for a Geigeria species in the field.

There are also four near-endemic genera, i.e. they occur

in Namibia and southwestern Angola, namely Antiphiona

(Asteraceae), Marcelliopsis (Amaranthaceae), and

Welwitschia (Welwitschiaceae). Volkiella (Cyperaceae)

has been recorded once in Zambia. Ruschianthemum

(Mesembryanthemaceae), which occurs just over the

border in the northern Cape, has been included in

Stoeberia (Chesselet & Van Wyk 2002).

Species endemic to Namibia

Approximately 600 of the nearly 4 000 indigenous

species recorded for Namibia are considered endemic

to within the borders of the country (Figure 4) and are

found in many different families and genera; 62 of the

157 families in Namibia have endemic species, whereas

231 genera out of 958 genera have endemic species.

The most important families are the Mesembryan-

themaceae, Asteraceae and Acanthaceae. The genera Aloe

(Asphodelaceae), Euphorbia (Euphorbiaceae), Herman-

nia (Sterculiaceae), Jamesbrittenia (Scrophulariaceae),

Petalidium (Acanthaceae), Salsola (Crassulaceae),

Stipagrostis, Eragrostis (Poaceae) and Zygophyllum

(Zygophyllaceae) have the most endemic species besides

Conophytum and Lithops (Mesembryanthemaceae),

which have numerous infraspecific taxa. Distributions

of about 600 endemic species and certain families and

genera are mapped in Craven (2002b). Two examples

showing distinctive patterns are illustrated here. Family

Mesembryanthemaceae (Figure 5) is mainly restricted

to the southwestern comer of Namibia which is also the

only area with winter rainfall. Figure 6 shows endemism

in Commiphora which occurs more commonly in the

north with few plants in the winter rainfall zone. Some

species have very widespread distributions and are well

represented in the collection, whereas others are limited

to one locality and one collection. One endemic was

recorded in 195 different quarter-degree squares, and

nine quarter-degree squares had more than 40 endemics.

The quarter-degree square with the highest number of

species were tabulated (Table 2) and the grid in which

the Brandberg occurs is shown to have the most endemic

species, followed by the Windhoek area.

Species endemic to localized areas are found mainly

in western Namibia, but also in the central regions

associated with high elevations. A number of localities

were found to house four or more very restricted-range

species.

The dominant life form of the endemics of the south-

west winter rainfall region is succulents, whereas further

inland, i.e. east of the Hunsberg, dwarf shrubs are more

180

Bothalia 36,2 (2006)

common. The only region with endemic trees is the

Kaokoveld. The distribution of endemic grasses shows

them to be widespread.

Namibian near-endemic species

Near-endemics are defined here as species that extend

marginally into another region, i.e. beyond the political

borders of Namibia. Two noteworthy areas for endem-

ics and near-endemics in Namibia have been identified

under the auspices of the IUCN Plant Conservation

Programme. These centres of exceptional species rich-

ness and endemism (Davis & Heywood 1994) are

the Kaokoveld in the northwest and the Gariep in the

southwest. Further analysis of near-endemics is needed,

including the species in this study. The number of species

per quarter-degree square in the northwest and south will

be higher.

Taxon phytogeographic centres

Kaokoveld Centre

Considering the topography and climate, it is no

wonder that mapped plant distributions show two main

subregions besides the Brandberg, namely the coastal

strip, which is affected by fog and cooler temperatures,

and the inland highlands. Endemics of the coast in-

clude Ectadium rotundifolium, Merremia multisecta,

Hermannia gariepina and grasses such as Chloris fla-

bellata and Sporobolus virginicus. Most of the endemic

taxa occurring on the highlands will not be found on the

coastal plains. They are more numerous and often con-

fined to mountainous areas.

Subcentres of importance are the Baynes Mountains

and the area around Sanitatis and Orupembe. Both areas

have six localized endemics each. Four endemics are

found in the Sesfontein area. The family with the most

endemics is the Acanthaceae with five representatives

and the other endemics are from a wide variety of fami-

lies.

Gariep Centre

Relatively numerous local endemics were found

around Aus, the Huib Hoch Plateau to Namus Mountains,

the Hunsberg and a section of the Warmbad District.

Species limited to these subcentres come from a variety

of families and vary from grasses and geophytes to suc-

culent Euphorbiaceae and Mesembryanthemaceae. Life

form type shows a certain degree of uniformity in some

of the subcentres. Dwarf shrubs such as Caesalpinia

merxmuellerana and Petalidium cymbiforme are charac-

teristic of the endemics of the Hunsberg, whereas more

succulent species, including three in Euphorbia , are

found on the western side of the Hunsberg in the Numas

Mountains. Three succulent mesembs are also endemic

to the Wannbad District, namely Antimima eendornensis ,

Schwantesia constanceae and S. succumbens. No spe-

cific life forms are characteristic of the Aus area, which

is characterized by varying substrates, or the Huib Hoch

Plateau areas. Although field work will probably result

in many of these species being recorded further afield,

some conspicuous plants such as Caesalpinia merxmuel-

lerana and Zygophyllum giessii have not been found to

be widespread despite intense searching.

Waterberg-Otavi Centre

Additional species restricted to this centre identified

here, but not necessarily only occurring on limestone, are

Heteromorpha papillosa (Apiaceae), Pentatrichia avas-

montana (Asteraceae), Plectranthus dinteri (Lamiaceae)

FIGURE 5. — Distribution of endem-

ic plants in Mesembryanthe-

maceae and diversity per quar-

ter-degree grid.

Bothalia 36,2 (2006)

181

which is also in the Windhoek region, and Thesium

xerophyticum (Santalaceae) also on the Gamsberg. In

addition, Thesium is in need of revision and although

both the genera Plectranthus and Heteromorpha have

been revised, little or no field work was carried out

in Namibia. Pentatrichia is presently under revision.

Species reported to date only from the Waterberg are

Dintera pterocaulis (Scrophulariaceae), Eriospermum

citrinum and E. lavranosii (Eriospermaceae).

Central Namibia and Southern Highland Centre

Fifteen Manuleae (Scrophulariaceae) were found to

occur in a highland centre (Hilliard 1994) that includ-

ed the Brandberg, Erongo, Khomashochland, Auas,

Gamsberg, Naukluft. Tiras, Karasberg and surround-

ing high ground, usually above ± 900 m. It excludes

the limestone area of the Waterberg, but may stretch

into Botswana. An analysis of mapped endemics and

habitat data show that numerous range-restricted spe-

cies are confined to higher elevations, as prescribed by

the definition of this centre by Hilliard (1994). The flo-

ristic elements range from shrubs ( Nicotiana africana)

and dwarf shrubs (Core horns merxmuelleri, Hermannia

merxmuelleri) to geophytes ( Haemanthus avasmontanus,

Lapeirousia avamontana) with a few very localized suc-

culents ( Ebracteola montis-moltkei , Euphorbia montei-

roi subsp. brandbergensis, Aloe viridiflora).

DISCUSSION

Numerous hypotheses have been proposed to explain

patterns of species diversity (Schmida & Wilson 1985),

but none have been found to apply well to all bodies of

data. Underlying datasets and resolution can also result

in different patterns of diversity. This is seen when

FIGURE 6. — Distribution of endem-

ic plants in Commiphora and

diversity per quarter-degree

grid.

Namibia is mapped continentally (Mutke et al. 2001),

regionally (Rebelo 1994) or nationally (Maggs et al.

1994). In spite of the finer resolution used here, the most

species-rich areas in Namibia do not differ much from

those first indicated (Maggs et al. 1994). The differ-

ences lie in better definition of the boundaries, additional

localities and important smaller localities. Examples are

the Naukluft. Windhoek and Aus regions. The inclusion

of the Naukluft as an important area by Maggs et al.

(1994) was suspected to be an artifact of high collecting

intensity and this was proved here to be the case. On

the other hand, species richness in the Windhoek area

is not necessarily an artifact of good collecting, despite

being close to a high population of potential collectors.

It includes the second highest mountain in Namibia (Irish

2002) , and a number of grasses (Klaassen & Craven

2003) , and other species that are not known from else-

where in Namibia are found here. It indicates that this

flora includes outliers from more distant areas and is in

agreement with Major (1988), who considers mountains

to be mesic refugial islands, which form ideal refuges in

times of climate change. Another area of high diversity,

near Aus, housed a concentration camp during World

War II where the interned German citizens botanized to

TABLE 2. — Quarter-degree squares (QDS) with the most endemic

species recorded

182

Bothalia 36,2 (2006)

pass the time. Factors that contribute to the richness of

this area include the diverse nature of the topography (the

start of the escarpment) and varying substrate (scattered

granite outcrops in the area) as well as the fog coming

inland from the coast.

Although the plant species diversity map is the most

detailed map of its kind for Namibia, care should be

taken before using it for management purposes or predic-

tions. Both the delimited areas and the numbers of spe-

cies predicted for the regions need further refinements.

Two aspects of conservation value that it does indicate

are: 1, that many areas are more diverse than may appear

during periods of harsh conditions, which may last for

years or decades; and 2, the relationship between areas

set aside as formal conservation areas and diversity. It is

apparent that certain areas of high species richness are

not afforded any formal protection.

The first map to show the overall distribution of

endemic species in Namibia (Maggs et al. 1994) was

based on 145 species. Endemics of southern Africa,

including Namibia, were mapped by Rebelo (1994).

Differences between the latter map (regional) and the

present one for Namibia (national) are noticeable because

of the scale, definition of the word endemic, and datasets

used. Maps published in Simmons (1998) and Simmons

et al. (1998) were based on updated data in Maggs et

al. (1997) using half-degree squares, because accord-

ing to Simmons et al. (1998), bird data show that this

scale reduces collecting bias. The results presented here

justify the use of quarter-degree, because working on a

national level, more quality control of the data is pos-

sible and other sources such as literature, field work and

shapefiles of physical features can be used. An undefined

‘escarpment’ area was said to be the main centre for

endemic plants by Simmons et al. (1998). Overlaying a

defined escarpment developed by the Atlas of Namibia

Project (2002) onto the endemic data indicates that this

is only partially correct. Although there is an association

between increased numbers of endemic species and the

escarpment, the area between the northern and southern

escarpments also show localities of importance, not only

in general, but also for those with limited ranges.

The maps resulting from this study are published by

the Atlas of Namibia Project (2002), where they are also

combined with maps of Namibian fauna. Mendelsohn

et al. (2002) conclude that the most notable zones of

high diversity for fauna and flora occur in the northeast,

in the Karstveld around Tsumeb, in highland areas in

the centre of Namibia, and in various scattered areas of

higher ground further west. Plant endemism was also

combined with that of animals, and Mendelsohn et at.

(2002) conclude that the overall patterns of endemism in

Namibia are quite different from those of overall diver-

sity. The greatest majority of endemics are found in the

dry, western and northwestern regions of Namibia. On

a regional scale of plants only (and lower resolution),

Rebelo (1994) reports a stronger correlation between

species richness and endemicity.

Namibian endemic plants, as in other parts of the

world, are usually associated with altitude, substrate,

or variations in geography, which provide numerous

microhabitats. The most important areas for species

richness in Namibia, however, are ‘transitional’ areas,

which Shmida & Wilson (1985) define as areas between

different ecological regions, i.e. zones in which species

overlap. Variations in altitude and the maximum altitude

in any grid are significant as reported for Africa as a

whole (Linder 1999). Superimposing altitudinal profiles

onto the shapefile of species richness shows this clearly

(Figure 2). It is also substantiated by studies which show

variations in species diversity with altitude for specific

sites or localities (Moisei 1982; Rutherford 1992).

Centres of diversity and endemism in Namibia, such

as the Kaokoveld and the Gariep Centre, although lacking

consistency in definition, have been discussed by numer-

ous authors. Volk (1964) proposed a Kaoko Element,

which was elaborated on by Nordenstam (1974). Hilliard

(1994) included two taxa, Jamesbrittenia canescens var.

laevior and J. heucherifolia (Scrophulariaceae), which

are confined to southern Angola and northern Namibia.

Other authors that recognized the Kaokoveld Centre are

Hilton-Taylor ( 1 994a), Maggs et al. ( 1 994), Maggs et al.

(1998), Van Wyk & Smith (2001) and Craven (2002a).

Hilton Taylor (1994b) considers the Gariep to be essen-

tially a geographic rather than a phytogeographic centre,

but both Nordenstam (1969) and Hilliard (1994) recog-

nize it as a taxon phytogeographic centre of importance

for numerous species.

Mapping all Namibian endemics has shown that

the distributions of many elements need to be reas-

sessed. Species presently regarded as e.g. Kaokoveld

elements, ( Welwitschia mirabilis , Acanthosicyos hor-

ridus, Cyphostemma currorii, Acacia robynsiana and

Moringa ovalifolia) may stretch far beyond what is

generally regarded as the centre. Another example is

the Brandberg, which is considered an outlier of the

Kaokoveld Centre by Nordenstam (1974) and Hilton

Taylor (1994a). Provisional results of the floristic ele-

ments of the Brandberg (Craven & Craven 2000) show

that numerous range-restricted species also occur on

other highlands further south and it is rather a part of the

Highland Centre as suggested by Hilliard (1994).

The Kaokoveld and the Gariep Centres are basically

geographic regions, so inclusion of species within the

centres was based on presence or absence in the area. This

is not possible with the Namibia Central and Southern

Highland Centre and the Waterberg-Otavi Centre as they

are identified by determinants such as habitat or substrate

specificity. These centres do not show a clear geographic

pattern of distribution on a map until altitude contours or

outlines of mountains are included. More endemic spe-

cies will probably be included in these centres once more

is known about such habitat requirements.

The name, Gariep, has also been used in large-scale

mapping by Jurgens ( 1991 ) despite a different approach,

i.e. including life form and climate in the analyses. Such

approaches must not be confused with that of taxon phy-

togeography as discussed here.

In general, the variable nature of most aspects of

Namibia’s rainfall, as well as the paucity of overall cli-

matic data, precludes using climate at this level of resolu-

tion. It is, however, true to say that Namibia differs from

Africa as a whole, because the most species-rich areas

Bothalia 36,2 (2006)

183

are in the wetter parts of the continent (Linder 1999).

This study highlighted the need for the microclimatic

conditions in which so many endemics thrive, to be stud-

ied and documented in a systematic way. Defining these

habitats is essential to understanding the distributions of

endemic plants. The same can be said for peculiar or iso-

lated substrata (serpentine, limestone, quartzite, calcare-

ous sands) which is a widespread phenomenon in some

areas (Major 1988; Cowling et al. 1992), and Namibia is

no exception.

Although distribution of endemics among life form

classes was not studied here in detail, it is evident that

life forms change with locality as recorded for endemic

species in general (Major 1988). This is ascribed to

climate, history of the flora and competition with the

associated flora (Major 1988).

Certain taxa require floristic study and field work, but

a provisional assessment suggests that a locality may

be home to a variety of species from various taxonomic

groups as suggested by Cowling et al. (1992).

Knowledge of local endemism will help create a bet-

ter basis for future policy (Brenan 1978). This study

does provide sufficient information on certain aspects of

Namibia’s endemic plants to start formulating conserva-

tion strategies, but there is still a need for satisfactory

taxonomic knowledge as well as more distribution data.

It is also imperative that the information is used correctly.

Simmons et al. (1998) concluded that another 11% of

the land area would be required to protect Namibia’s

endemic plants. If the endemic plants already found

within protected areas had been removed from the data-

set prior to the analysis, the resulting value would be

much lower. No matter how highly a species is regarded,

as soon as it is adequately conserved, it is no longer used

as an argument to conserve another area (Kirkpatrick

1983). Because species are not spread evenly around the

world and unique concentrations may occur in relatively

small areas, i.e. within the political borders of a country

like Namibia, the onus is on Namibia to protect these

species.

ACKNOWLEDGEMENTS

The National Herbarium of Namibia, especially the

database manager, Esmerialda Klaassen are thanked for

the use of SPMNDB. John Mendelsohn and the Atlas

team are also thanked for the final maps on diversity and

endemism produced for the Atlas of Namibia. The Agro-

Ecological Zoning Programme is thanked for the use of

shapefiles. The unpublished plant lists of Craig Hilton-

Taylor for the Kaokoveld and Gariep Centres is greatly

appreciated. The contribution made by SABONET, i.e.

the database and funding for further study, is greatly

appreciated.

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APPENDIX 1. — List of Namibian endemic plants used in the evaluation

= Species name in italics indicates a synonym used in the original evaluation.

* Endemic species not used in the evaluation due to lack of data or species described since the evaluation.

• Species that have been collected outside Namibia since the evaluation. Unless otherwise stated, species indicated

by • are near-endemic and mainly based on collections of Craven or Bruyns, particularly from Angola.

Acanthaceae

Barleria

•damarensis T. Anderson

dinteri Oberm.

jubata S. Moore

kaloxytona Lindau

lanceolata (Schinz) Oberm.

meeuseana P. G.Mey.

merxmuelleri P. G.Mey.

solitaria P. G.Mey.

Blepharis

ferox P. G.Mey.

fleckii P. G.Mey.

gigantea Oberm.

meyeri Vollesen

pruinosa Engl.

spinifex Menem.

Hygrophila gracillima (Schinz) Burkill

Justicia

cuneata Vahl subsp. hoerleiniana (P. G.Mey.) Immelman

guerkeana Schinz

•platysepala (S. Moore) P.G.Mey.

Monechma

calcaratum Schinz

callothamnum Munday

crassiusculum P.G.Mey.

desertorum (Engl.) C.B. Clarke

grandiflorum Schinz

leucoderme (Schinz) C.B. Clarke

serotinum P.G.Mey.

tonsum P.G.Mey.

Peristrophe

grandibracteata Lindau

hereroensis (Schinz) K.Balkwill

namibiensis K.BalkM’ill

subsp. brandbergensis K.Balkwill

subsp. namibiensis

Petalidium

canescens (Engl) C.B. Clarke

cymbiforme Schinz

giessii P.G.Mey.

lanatum (Engl.) C.B. Clarke

linifolium T.Anderson

luteo-album A.Meeuse

*ohopohense P.G.Mey.

pilosi-bracteolatum Merxm. & Hainz

ramulosum Schinz

rautanenii Schinz

suberispum P.G.Mey.

Rhinacanthus kaokoensis K.Balkwill & S. Williamson

Ruellia

aspera (Schinz) E. Phillips

brandbergensis Kers

Bothalia 36,2 (2006)

185

Aizoaceae

Aizoanthemum

dinteri (Schinz) Friedrich

galenioides (Fenzl ex Sond.) Friedrich

rehmannii (Schinz) H.E.K.Hartmann = Aizoanthemum membrum-

connectens Dinter ex Friedrich

Aizoon giessii Friedrich

Tetragonia

rangeana Engl.

•schenckii (Schinz) Engl.

Trianthema hereroensis Schinz

Alliaceae

Tulbaghia calcarea Engl. & K.Krause , insuff. known

Amaranthaceae

Arthraerua leubnitziae (Kuntze) Schinz

Calicorema squarrosa (Schinz) Schinz

Hermbstaedtia spathulifolia (Engl.) Baker

Marcelliopsis splendens (Schinz) Schinz

Amarvllidaceae

Ammocharis nerinoides (Baker) Lehmiller

Crinum

paludosum I. Verd.

rautanenianum Schinz

Haemanthus avasmontanus Dinter

*Namaquanula bruynsii Snijman

*Nerine pusilla Dinter

Strumaria

hardyana D.Miill.-Doblies & U.Miill.-Doblies

phonolithica Dinter

Anacardiaceae

Rhus

problematodes Merxm. & Roessler

volkii Suess.

Apiaceae

Anginon streyi (Merxm.) I. Allison & B.-E.van Wyk

Heteromorpha papillosa C.C. Towns.

Marlothiella gummifera H. Wolff

Phlyctidocarpa flava Cannon & W.L. Theob.

Polemanniopsis sp. = Merxmuller & Giess 32010

Apocynaceae

•Australluma peschii = Caralluma peschii Nel

Baynesia lophophora Bruyns

Brachystelma

blepharanthera H.Huber

codonanthum Bruyns

recurvatum Bruyns

schinzii (K.Schum.) N.E.Br.

schultzei (Schltr.) Bruyns

•Ceropegia dinteri Schltr.

Cynanchum meyeri (Decne.) Schltr.

Ectadium

latifolium (Schinz) N.E.Br.

rotundifolium (H.Huber) Venter & Kotze

•Gomphocarpus semiplectens K.Schum.

Hoodia

juttae Dinter

officinalis (N.E.Br.) Plowes subsp. delaetiana (Dinter) Bruyns

ruschii Dinter

triebneri (Nel) Bruyns

Huemia

hallii E.Lamb & B.M.Lamb

plowesii L. C. Leach

Larryleachia tirasmontana Plowes = Lavrania picta (N.E.Br.) Plowes

subsp. parvipunctata Bruyns

Lavrania haagnerae Plowes

Microloma

hereroense Wanntorp

penicillatum Schltr.

Orbea

albocastanea (Marloth) Bruyns

maculata (N.E.Br.) L.C.Leach

•subsp. kaokoensis Bruyns

subsp. rangeana (Dinter & A. Berger) Bruyns

Raphionacme

haeneliae Venter & R.L. Verh.

namibiana Venter & R.L. Verh.

Stapelia

kwebensis N.E.Br. = Stapelia longipedicellata (A. Berger) N.E.Br.,

not endemic

pearsonii N.E.Br

*remota R. A. Dyer

schinzii A. Berger & Schltr.

*var. bergeriana (Dinter) L.C.Leach

var. schinzii

•Stapeliopsis umiflora Lavranos

Stigmatorhynchus hereroensis Schltr.

Tridentea

marientalensis (Nel) L.C.Leach subsp. albipilosa (Giess) L.C.Leach

•pachyrrhiza (Dinter) L.C.Leach

Tromotriche ruschiana (Dinter) Bruyns

•Tylophora fleckii (Schltr.) N.E.Br.

Aponogetonaceae

Aponogeton azureus H.Bntggen

Asphodelaceae

Aloe

argenticauda Metxm. & Giess

asperifolia A. Berger

corallina I. Verd.

dewinteri Giess

•dinteri A. Berger

erinacea D.S. Hardy

*hereroensis Engl. var. lutea A. Berger

namibensis Giess

omavandae Van Jaarsv

pachygaster Dinter

sladeniana Pole Evans

viridiflora Reynolds

Bulbine

caput-medusae G. Will.

francescae G. Will. & Baijnath

namaensis Schinz

praemorsa = Bulbine tetraphylla Dinter, not endemic

*rhopalophylla Dinter

Trachyandra

ensifolia (Solch) Roessler

glandulosa (Dinter) Oberm.

lanata (Dinter) Oberm.

peculiaris (Dinter) Oberm.

Asteraceae

Amphiglossa thuja (Merxm.) Koekemoer

Anisopappus

pinnatifidus (Klatt) O.Hoffm. ex Hutch.

pseudopinnatifidus S. Ortiz & Paiva

Antiphiona

fragrans (Metxm.) Merxm.

pinnatisecta (S. Moore) Merxm.

Arctotis frutescens T.Norl.

Aspilia eenii S. Moore

Berkheya schinzii O.Hoffm.

Calostephane marlothiana O.Hoffm.

*Chrysocoma puberula Merxm.

Crassocephalum coeruleum (O.Hoffm.) R.E.Fr.

Dauresia alliariifolia (O.Hoffm.) B.Nord. & Pelser = Senecio alliari-

ifolius O.Hoffm.

Dicoma

cuneneensis Wild

dinteri S. Moore

*obconica S. Ortiz & Pulgar

Eremothamnus marlothianus O.Hoffm.

Eriocephalus

dinteri S. Moore

giessii M.A.N. Muller

kingesii Merxm. & Eberle

klinghardtensis M.A.N. Muller

pinnatus O.Hoffm.

Euryops

mucosus B.Nord.

walterorum Merxm.

Felicia

alba Grau

gunillae B.Nord.

smaragdina (S. Moore) Merxm.

*Garuleum schinzii O.Hoffm. subsp. crinitum (Dinter) Merxm.

Gazania thermalis Dinter

186

Bothalia 36,2 (2006)

Geigeria

odontoptera O.Hoffm.

omativa O.Hoffm. subsp. omativa var. filifolia (Mattf.) S. Ortiz &

Rodr.Oubiha = G. englerana Muschl. & Geigeria otaviensis

(Merxm.) Merxm.

pilifera Hutch.

plumosa Muschl.

*subsp. angustifolia S. Ortiz & Rodr.Oubiha

subsp. brachycephala S. Ortiz & Rodr.Oubiha = G. brachycephala

Muschl.

rigida O.Hoffm.

Gorteria diffusa Thunb. subsp. parviligulata Roessler

Helichrysum

amboense Schinz

deserticola Hilliard

erubescens Hilliard

marlothianum O.Hoffm.

Lasiopogon

ponticulus Hilliard

volkii (B.Nord.) Hilliard

Myxopappus hereroensis (O.Hoffm.) Kdllersjo

Nicolasia

heterophylla S. Moore

subsp. affinis (S. Moore) Merxm.

subsp. heterophylla

Nidorella nordenstamii Wild

Nolletia tenuifolia Mattf.

*Norlindhia aptera B.Nord.

Ondetia linearis Benth.

Osteospermum

montanum Klatt

muncatum E. Mey. ex DC. subsp. longiradiatum T.Norl.

Othonna

brandbergensis B.Nord.

clavifolia Marloth

•graveolens O.Hoffm.

sparsiflora (S. Moore) B.Nord.

Pegolettia

pinnatilobata (Klatt) O.Hoffm. ex Dinter

plumosa M.D.Hend.

Pentatrichia

avasmontana Merxm.

rehmii (Merxm.) Merxm.

*Philyrophyllum brandbergense P.P.J. Herman

Pentzia tomentosa B.Nord.

Pteronia

eenii S. Moore

polygalifolia O.Hoffm.

pomonae Merxm.

rangei Muschl.

spinulosa E. Phillips

Rennera eenii (S. Moore) Kdllersjo

Senecio

engleranus O.Hoffm.

giessii Merxm.

hermannii B.Nord.

windhoekensis Merxm.

Sphaeranthus wattii Giess ex Merxm.

Tripteris nervosa Hutch.

Ursinia frutescens Dinter

Vemonia

obionifolia O.Hoffm

subsp. dentata Merxm.

subsp. obionifolia

Boraginaceae

*Ehretia namibiensis Retief & A.E.van Wyk subsp. kaokoensis Retief &

A.E.van Wyk

Heliotropium albiflorum Engl.

*Trichodesma angustifolium Harv. subsp. argenteum Retief & A.E.van

Wyk

Brassicaceae

Heliophila

*deserticola Schltr. var. micrantha A.Schreib.

obibensis Marais

Burseraceae

Commiphora

*dinteri Engl.

giessii J.J.A.van der Walt

*kaokoensis W.Swanepoel

kraeuseliana Heine

•saxicola Engl.

•virgata Engl.

Campanulaceae

Namacodon schinzianum (Markgr.) Thulin

Wahlenbergia

densicaulis Brehmer

erophiloides Markgr.

*intricata (Dinter & Markgraf) P. Craven, ined. = Lightfootia dinteri

Engl, ex Dinter

subumbellata Markgr.

Capparaceae

Cleome

camosa (Pax) Gilg & Gilg-Ben.

foliosa Hookf. var. namibensis (Kers) Codd

labumifolia Roessler

suffruticosa Schinz

Chenopodiaceae

*Chenopodium amboanum (Murr) Aellen

Salsola

albisepala Aellen

arborea C.A.Sm. ex Aellen

*aroabica Botsch.

campyloptera Botsch.

cauliflora Botsch.

columnaris Botsch.

cryptoptera Aellen

denudata Botsch.

dinteri Botsch.

dolichostigma Botsch.

etoshensis Botsch.

*garubica Botsch.

gemmata Botsch.

giessii Botsch.

hoanibica Botsch.

hottentottica Botsch.

huabica Botsch.

kleinfonteini Botsch.

koichabica Botsch.

marginata Botsch.

mirabilis Botsch.

namibica Botsch.

okaukuejensis Botsch.

*omaruruensis Botsch.

*parviflora Botsch.

procera Botsch.

ptiloptera Botsch.

robinsonii Botsch.

*schreiberae Botsch.

scopiformis Botsch.

seminuda Botsch.

*seydelii Botsch.

spenceri Botsch.

swakopmundi Botsch.

ugabica Botsch.

unjabica Botsch.

Suaeda

articulata Aellen

salina B.Nord.

Colchicaceae

Androcymbium exiguum Roessler subsp. exiguum

Omithoglossum calcicola K.Krause & Dinter

Convolvulaceae

Convolvulus argillicola Pilg.

Merremia

bipinnatipartita (Engl.) Hallier f.

guerichii A.Meeuse

Crassulaceae

Adromischus

schuldtianus (Poelln.) Poelln.

*subsp. brandbergensis B.Nord. & Van Jaarsv.

subsp. juttae (Poelln.) Toelken

subsp. schuldtianus

Bothalia 36,2 (2006)

187

Crassula

aurusbergensis G. Will.

ausensis Hutchison

subsp. ausensis

subsp. giessii (Friedrich) Toelken

*subsp. titanopsis Pavelka

elegans Schonland & Baker f. subsp. namibensis (Friedrich)

Toelken

luederitzii Schonland

numaisensis Friedrich

Tylecodon

aridimontanus G. Will.

*aurusbergensis G. Will. & Van Jaarsv.

Cucurbitaceae

Citrullus rehmii De Winter

Cucumella clavipetiolata J.H.Kirkbr.

Cyperaceae

•Bulbostylis mucronata C.B. Clarke

Cyperus rehmii Merxm.

Ebenaceae

Euclea asperrima Friedr.-Holzh.

Eriospermaceae

Eriospermum

buchubergense Dinter , insuff. known

citrinum PL. Perry

flexum P.L. Perry

*graniticolum Dinter ex Poelln., insuff. known

halenbergense Dinter

lavranosii P.L. Perry

volkmanniae Dinter

Euphorbiaceae

Euphorbia

angrae N.E.Br.

baliola N.E.Br.

caperonioides R.A.Dyer & P.G.Mey.

chamaesycoides B.Nord.

•cibdela N.E.Br.

damarana L.C. Leach

friedrichiae Dinter

giessii L.C. Leach

insarmentosa P. G.Mey.

juttae Dinter

kaokoensis (A.C. White, R.A.Dyer & B.Sloane) L.C. Leach

lavrani L.C. Leach

leistneri R.H.Archer

mauritanica L. var. foetens Dinter ex A.C. White, R.A.Dyer &

B.Sloane

monteiroi Hook.f. subsp. brandbergensis B.Nord.

namibensis Marloth

namuskluftensis L.C. Leach

otj ipembana L. C. Leach

pergracilis P.G.Mey.

*pseudoduseimata A. C. White, R.A.Dyer & B.Sloane

rudis N.E.Br.

*spartaria N.E.Br.

*spinea N.E.Br

venenata Marloth

verruculosa N.E.Br.

volkmanniae Dinter

Phyllanthus dinteri Pax

Tragia

dinteri Pax

lancifolia Dinter ex Pax & K.Hoffm.

Fabaceae

•Acacia montis-usti Merxm. & A.Schreib.

Bolusia amboensis (Schinz) Harms

Caesalpinia

merxmuellerana A.Schreib.

pearsonii L. Bolus

Crotalaria

aurea Dinter ex Baker f.

colorata Schinz subsp. colorata

kurtii Schinz

Decorsea dinteri (Harms) Verde.

Elephantorrhiza

rangei Harms

schinziana Dinter

Eriosema harmsiana Dinter

Erythrina decora Harms

Haematoxylum dinteri (Harms) Harms

Indigofera

acanthoclada Dinter

anabibensis A.Schreib.

giessii A.Schreib.

hochstetteri Baker subsp. streyana (Merxm.) A.Schreib.

merxmuelleri A.Schreib.

pechuelii Kuntze

rautanenii Baker f.

Lebeckia

dinteri Harms

obovata Schinz

Lessertia

acanthorhachis (Dinter) Dinter

cryptantha Dinter

eremicola Dinter

Lotononis

bracteosa B.-E.van Wyk

mirabilis Dinter

pachycarpa Dinter ex B.-E.van Wyk

pallidirosea Dinter & Harms

schreiberi B.-E.van Wyk

Sesbania pachycarpa DC. subsp. dinterana J.B.Gillett

Tephrosia

griseola H.M.L. Forbes

monophylla Schinz

pallida H.M.L. Forbes

Frankeniaceae

Frankenia pomonensis Pohnert

Geraniaceae

Monsonia

deserticola Dinter ex R.Knuth

drudeana Schinz

ignorata Merxm. & A.Schreib.

trilobata Kers

Pelargonium

cortusifolium L 'Her.

mirabile Dinter

otaviense R.Knuth

paniculatum Jacq.

Sarcocaulon

inerme Rehm

marlothii Engl.

peniculinum Moffett

Hyacinthaceae

Albuca

amboensis (Schinz) Oberm.

*englerana K. Krause & Dinter

hereroensis Schinz

*karasbergensis PE. Glover

*reflexa Dinter & K. Krause

Drimia

namibensis (Oberm.) J.C. Manning & Goldblatt = Rhadamanthus

namibensis Oberm.

secunda (B.Nord.) J.C. Manning & Goldblatt = Rhadamanthus

secundus B.Nord.

Lachenalia

giessii W.F.Barker

klinghardtiana Dinter

namibiensis W.F.Barker

nutans G.D. Duncan

pearsonii ( P.E. Glover) W.F.Barker

Ledebouria scabrida Jessop

Omithogalum

candidum Oberm.

rautanenii Schinz

stapffii Schinz

tubiforme (Oberm.) Oberm.

Hypoxidaceae

Hypoxis dinteri Nel

Iridaceae

Babiana longicollis Dinter

•Ferraria schaeferi Dinter

Lapeirousia

avasmontana Dinter

gracilis Vaupel

188

Bothalia 36,2 (2006)

Moraea

garipensis Goldblatt

*graniticola Goldblatt

hexaglottis Goldblatt

namibensis Goldblatt

rigidifolia Goldblatt

Kirkin ceae

Kirkia dewinteri Merxm. & Heine

Lamiaceae

Acrotome fleckii (Giirke) Laimert

Aeollanthus namibiensis Ryding

Hemizygia floccosa Laimert

Plectranthus

dinteri Briq.

unguentarius Codd

Stachys dinteri Laimert

•Tetradenia kaokoensis Van Jaarsv. &A.E.van Wyk

Lobeliaceae

Lobelia hereroensis Schinz

Loranthaceae

Agelanthus discolor (Schinz) Balle

Lythraceae

Nesaea luederitzii Koelme var. hereroensis Koehne

Malvaceae

Hibiscus

dinteri Hochr.

discophorus Hochr.

fleckii Giirke

merxmuelleri Roessler

sulfuranthus Ulbr.

Pavonia rehmannii Szyszyl.

Mesembryanthemaceae

Amphibolia saginata (L. Bolus) H EX. Hartmann

Antimima

argentea (L. Bolus) H EX. Hartmann

aurasensis H. EX. Hartmann

buchubergensis (Dinter) H. EX. Hartmann

dolomitica (Dinter) H. EX. Hartmann

eendomensis (Dinter) H.E.XHartmann

modesta (L. Bolus) H.E.XHartmann

quarzitica (Dinter) H. EX. Hartmann

Astridia hallii L. Bolus

Brownanthus

•arenosus (Schinz) Ihlenf. & Bittrich

namibensis (Marloth) Bullock

pubescens (N.E.Br. ex Maasss) Bullock

Cephalophyllum

*compressum L. Bolus

confusum (Dinter) Dinter & Schwantes

Cheiridopsis caroli-schmidtii (Dinter & A. Berger) N.E.Br

Conophytum

halenbergense (Dinter & Schwantes) N.E.Br

klinghardtense Rawe

subsp. baradii (Rawe) S. A. Hammer

subsp. klinghardtense

quaesitum (N.E.Br.) N.E.Br. subsp. densipunctum (L. Bolus)

S. A. Hammer

ricardianum Loesch & Tischer

subsp. ricardianum

subsp. rubifiorum Tischer

taylorianum (Dinter & Schwantes) N.E.Br.

subsp. emianum (Loesch & Tischer) de Boer ex S. A. Hammer

subsp. taylorianum

Delosperma klinghardtianum Schwantes

Dinteranthus microspermus (Dinter & Derenb.) Schwantes subsp.

impunctatus N. Sauer

*Dracophilus delaetinus (Dinter) Dinter & Schwantes

Drosanthemum

nordenstamii L. Bolus

pauper (Dinter) Dinter & Schwantes

Eberlanzia clausa (Dinter) Schwantes

Ebracteola

derenbergiana (Dinter) Dinter & Schwantes

montis-moltkei (Dinter) Dinter & Schwantes

Fenestraria rhopalophylla (Schltr. & Diels) N.E.Br subsp. rhopalo-

phylla

Jensenobotrya lossowiana A.G.J.Herre

Juttadinteria

attenuata Walgate

*ausensis (L. Bolus) Schwantes

deserticola (Marloth) Schwantes

simpsonii (Dinter) Schwantes

Lithops

dinteri Schwantes

subsp. dinteri

* subsp. multipunctata (de Boer) D.T.Cole

*francisci (Dinter & Schwantes) N.E.Br

*gracilidelineata Dinter subsp. brandbergensis (de Boer) D.T.Cole

*hermetica D. T. Cole

julii (Dinter & Schwantes) N.E.Br subsp. julii

karasmontana (Dinter & Schwantes) N.E.Br

*subsp. bella ( N.E.Br. ) D.T.Cole

*subsp. eberlanzii (Dinter & Schwantes) D.T.Cole

*optica (Marloth) N.E.Br

pseudotruncatella (A. Berger) N.E.Br

*subsp. archerae (de Boer) D.T.Cole

*subsp. dendritica (Nel) D.T.Cole

*subsp. volkii (Schwantes ex de Boer & Boom) D.T.Cole

*ruschiorum (Dinter & Schwantes) N.E.Br

schwantesii Dinter

*subsp. gebseri (de Boer) D.T.Cole

subsp. schwantesii

*vallis-mariae (Dinter & Schwantes) N.E.Br

wemeri Schwantes ex H. Jacobsen

*Malephora engleriana (Dinter & A. Berger) Schwantes

Mesembryanthemum pellitum Friedrich

Namibia

cinerea (Marloth) Dinter & Schwantes = Namibia ponderosa

(Dinter & Schwantes) Dinter & Schwantes

pomonae (Dinter) Dinter & Schwantes ex Walgate

Psammophora

*nissenii (Dinter) Dinter & Schwantes

*saxicola H. EX. Hartmann

Psilocaulon

gessertianum (Dinter <& A. Berger) Dinter & Schwantes

•salicomioides (Pax) Schwantes

Ruschia

deminuta L. Bolus

*namusmontana Friedrich

*odontocalyx (Schltr & Diels) Schwantes

*pollardii Friedrich

*ruschiana (Dinter) Dinter & Schwantes

vulvaria (Dinter) Schwantes

Ruschianthus falcatus L. Bolus

Schwantesia

constanceae N.Zimm.

succumbens (Dinter) Dinter

Synaptophyllum juttae (Dinter & A. Berger) N.E.Br

*Titanopsis schwantesii (Schwantes) Schwantes

*Trichodiadema littlewoodii L. Bolus

Molluginaceae

*Corbichonia rubriviolacea (Friedrich) Jeffrey

Hypertelis caespitosa Friedrich

Mollugo walteri Friedrich

Suessenguthiella caespitosa Friedrich

Nyctaginaceae

Boerhavia deserticola Codd

Commicarpus

*decipiens Meikle

fruticosus Pohnert

Orobanchaceae

Alectra

pseudobarleriae (Dinter) Dinter

schoenfelderi Dinter & Melch.

Oxalidaceae

Oxalis

ausensis R.Knuth

hunsbergensis ined.

Bothalia 36,2 (2006)

189

Oxalis (cont.)

luederitzii Schinz

pseudo-cemua R.Ktmth

schaeferi R.Knuth

Passifloraceae

•Adenia pechuelii (Engl.) Harms

Pedaliaceae

Rogeria bigibbosa Engl.

Sesamothamnus leistneranus, ined. = De Winter & Leistner 5504

Sesamum

abbreviatum Merxm.

marlothii Engl.

Plumbaginaceae

Limonium dyeri Lincz.

Plumbago

pearsonii L. Bolus

wissii Friedrich

Poaceae

*Brachiaria schoenfelderi C.E.Hubb. & Schweick.

Eragrostis

aristata De Winter

kingesii De Winter

omahekensis De Winter

pygmaea De Winter

sabinae Launert

scopelophila Pilg.

stenothyrsa Pilg.

waited Pilg.

•Kaokochloa nigrirostris De Winter

Merxmuellera rangei (Pilg.) Conert

*Panicum pearsonii F.Bolus

Pennisetum foermeranum Leeke

Pogonarthria leiarthra Hack.

Setaria finita Launert

Sporobolus nebulosus Hack, (endemic to southern Africa)

Stipagrostis

•damarensis (Mez) De Winter

garubensis (Pilg.) De Winter

gonatostachys (Pilg.) De Winter

•hermannii (Mez) De Winter

lanipes (Mez) De Winter

namibensis De Winter

pellytronis De Winter

•ramulosa De Winter

sabulicola (Pilg.) De Winter

seelyae De Winter

Polygalaceae

•Polygala guerichiana Engl.

Portulacaceae

Anacampseros filamentosa (Haw.) Sims subsp. tomentosa (A. Berger)

Gerbaulet

Avonia dinted (Schinz) G. D. Rowley = Anacampseros dinteri Schinz

•Ceraria longipedunculata Merxm. & Podlech

Rubiaceae

Amphiasma

divaricatum (Engl.) Bremek.

merenskyanum Bremek.

Kohautia

amboensis (Schinz) Bremek.

azurea (Dinter & K. Krause) Bremek.

Santalaceae

Thesium xerophyticum A W.Hill

Scrophulariaceae

Anticharis

ebracteata Schinz

imbricata Schinz

inflata Marloth & Engl.

Aptosimum

arenarium Engl.

suberosum F.E. Weber

Chamaegigas intrepidus Dinter ex Heil

Cromidon pusillum (Roessler) Hilliard

Diclis tenuissima Pilg.

Dintera pterocaulis Stapf

Jamesbrittenia

acutiloba (Pilg.) Hilliard

barbata Hilliard

bicolor (Dinter) Hilliard

chenopodioides Hilliard

dolomitica Hilliard

fimbriata Hilliard

fleckii (Thell.) Hilliard

fragilis (Pilg) Hilliard

giessii Hilliard

hereroensis (Engl) Hilliard

lyperioides (Engl.) Hilliard

pallida (Pilg.) Hilliard

pilgeriana (Dinter) Hilliard

primuliflora (Thell.) Hilliard

sessilifolia (Diels) Hilliard

Manulea

dubia (Skan) Overkott ex Roessler

namibensis (Roessler) Hilliard

tenella Hilliard

Manuleopsis dinteri Thell.

Nemesia

karasbergensis L. Bolus

violiflora Roessler

•Phyllopodium hispidulum (Thell.) Hilliard

Selago

amboensis Rolfe

lepida Hilliard

nachtigalii Rolfe

Solanaceae

Lycium grandicalyx Joubert & A.M. Venter

Nicotiana africana Merxm.

Solanum

damarense Bitter

dinteri Bitter

rigescentoides Hutch.

Sterculiaceae

Dombeya rotundifolia (Hochst.) Planch, var. velutina 1. Verd.

Hermannia

amabilis Marloth ex K.Schum.

*complicata Engl.

elliottiana (Harv.) K.Schum.

engleri Schinz

glandulosissima Engl.

juttae Dinter & Engl.

merxmuelleri Friedr.-Holzh.

minimifolia Friedr.-Holzh.

solaniflora K.Schum.

Tecophilaeaceae

•Eremiolirion amboensis = Cyanella amboensis Schinz

Tiliaceae

Corchorus merxmuelleri Wild

Turneraceae

Tumera oculata Story var. paucipilosa Oberm.

Verbenaceae

Priva auricoccea A.Meeuse

Vitaceae

Cyphostemma

bainesii (Hook.f) Desc.

•juttae (Dinter & Gilg) Desc.

omburense (Gilg & M. Brandt) Desc.

Zygophyllaceae

Neoluederitzia sericeocarpa Schinz

Zygophyllum

applanatum Van Zyl

cylindrifolium Schinz

giessii Merxm. & A.Schreib.

hirticaule Van Zyl

longistipulatum Schinz

•stapffii Schinz , previously known as Z. orbiculatum in Angola

Bothalia 36,2: 191-199 (2006)

Phytogeography of Passerina (Thymelaeaceae)

C.L. BREDENKAMP*+ and A.E. VAN WYK**

Keywords: Afromontane, Cape Floristic Region, distribution, endemism, fynbos, grassland. Passerina L., phytogeography, Red Data List, relict,

Thymelaeaceae

ABSTRACT

Passerina L. comprises 20 species and four subspecies of microphyllous, wind-pollinated shrubs. Once considered cos-

mopolitan. the genus as currently defined, is endemic to southern Africa. Endemism within the genus is highest in the Cape

Floristic Region (CFR), where all members show morphological and anatomical adaptations to the winter rainfall and dry

warm summers of the Mediterranean or semi-Mediterranean climate of the region. The Western Cape is the centre of diversity

for Passerina, from where certain species extend to the west, north and east. The outlier populations of Passerina montana

Thoday on the interior plateau of South Africa and Zimbabwe, as well as the Auas Moutains in Namibia, most probably origi-

nated in the CFR and formed part of a previously wider northern temperate Afromontane grassland-dominated vegetation

during the Quartemary, of which relicts remained in the high mountain areas. P. burchellii Thoday and P. paludosa Thoday

have the most restricted distribution and are regarded as Vulnerable. All other species are either widespread or under no im-

mediate threat.

INTRODUCTION

Meisner (1840, 1857) redefined Passerina L. by clari-

fying the taxonomic position of 92 ‘species exclusae’,

thus changing the status of the genus from cosmopolitan

to endemic in southern Africa, an opinion also reflected

in the now outdated taxonomic revision of the group by

Thoday (1924). A recent monograph of Passerina recog-

nized 20 species and four subspecies (Bredenkamp 2002;

Bredenkamp & Van Wyk 2003); these are listed in Tables

1 and 2. All members of Passerina are woody, evergreen

shrubs or subshrubs with microphyllous leaves and small,

rather insignificant wind-pollinated flowers. It is the only

exclusively anemophilous genus of the Thymelaeaceae

and the plants usually grow gregariously.

Taking the most southerly distribution of Passerina

montivaga Bredenkamp & A.E. van Wyk into consid-

eration, no less than 18 species of Passerina occur

in the Cape Floristic Region (CFR). The CFR is also

acknowledged as the smallest of the world’s six floris-

tic kingdoms (Van Wyk & Smith 2001). Following a

taxonomic revision of Lachnaea L. (Thymelaeaceae),

Beyers (2001) reviewed the recognition of local centres

of endemism within the CFR, from the initial descrip-

tions by Weimarck (1941) up to those of Goldblatt &

Manning (2000). In this paper we follow the interpreta-

tion of Goldblatt & Manning (2000), which identifies

the following six principal local centres of endemism:

the Northwestern (NW), Southwestern (SW), Agulhas

Plain (AP), Karoo Mountain (KM), Langeberg (LB) and

Southeastern (SE) Centres.

Species of Passerina endemic to the CFR (Table 1) are

morphologically and anatomically adapted to the winter

rainfall and dry warm summers of the Mediterranean or

* South African National Biodiversity Institute, Private Bag X101, 0001

Pretoria.

^Student affiliation: Department of Botany, University of Pretoria, 0002

Pretoria.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany, Univer-

sity of Pretoria, 0002 Pretoria.

MS. received: 2004-11-19.

semi-Mediterranean climate in the region (Bredenkamp

& Van Wyk 1999, 2000, 2001). Most species in the CFR

are associated with fynbos, a sclerophyllous vegetation

type on oligotrophic soils derived mainly from quartzitic

Cape Supergroup rocks. These species are adapted to a

variety of habitats, e.g. high-mountain peaks above the

snowline, where plants are often surrounded by mist

(throughout the year) or covered by snow especially dur-

ing the winter months; forest and mountain fynbos; vleis

and marshes; coastal limestone deposits and limestone

hills; coastal fynbos, where the plants grow on sand

dunes and in sandy areas. Many species are pioneers

growing along roadsides and in disturbed places.

Species near-endemic to the CFR are more wide-

spread (Table 1 ). They have adapted to a wider amplitude

of environmental conditions: where ranges extend north

of the southern Cape mountain ranges, they are often

adapted to arid karroid vegetation and summer rainfall;

certain species are adapted to forest margins and others

tolerate periodic falls of snow at high altitudes.

The few species of Passerina endemic to the northern

Drakensberg or near-endemic to the Great Escarpment

of southern Africa (Table 2) are associated with the high

moisture levels prevalent on the eastern escarpment and

conditions of summer rainfall. These plants are often

found in the ecotonal belt between forest and grassland;

they also grow along streams and riverbanks and on

mountain slopes.

In this contribution we describe, for the first time

in one paper, the patterns of geographical distribution

shown by members of Passerina. Patterns are interpreted

in terms of geology, climate, vegetation type and histori-

cal change, and the conservation status of threatened taxa

is suggested.

MATERIAL AND METHODS

All infrageneric taxa of Passerina were studied during

extensive field work covering the complete geographi-

192

Bothalia 36,2 (2006)

10* 15* 20* 25" 30" 35" 40*

FIGURE 1 . — Known geographical distribution of the genus Passerina.

Distribution in Angola on Huila Plateau near Lubango and Chela

Mountains is shown in insert.

cal range of the genus. Live material was collected, as

far as possible, from at least five different localities

for every taxon. Habitat and distribution data from 22

national and international herbaria were compiled in

a Microsoft Access Database and integrated with the

Pretoria Computerised Information System (PRECIS),

from which distribution maps for all taxa were gener-

ated. For mapping purposes the degree square system

was used (Edwards & Leistner 1971). Categories used to

indicate Red List status of taxa are based on the criteria

of the IUCN Species Survival Commission (2000).

OBSERVATIONS

The combined distribution of all species of Passerina

is shown in Figure 1. The number of species per one-

degree square is indicated in Figure 2. In Passerina the

highest numbers of species per one-degree square are

concentrated in a belt including those grids between 33°

and 34°S and 18° to 27°E. The CFR (mainly Western

Cape) is clearly the centre of diversity for Passerina,

from where certain species extend to the west, north and

east (see also Bredenkamp & Van Wyk 200 1 ). The highest

numbers of species occur in the grids 3321 (Ladismith),

3322 (Oudtshoom) and 3419 (Caledon). The highest

FIGURE 2. Number of species of Passerina per one-degree grid square. Lines PQ and RS: boundaries between summer (A), intermediate (B)

and winter (C) rainfall areas. Line XY shows northern boundary of Cape Supergroup rock outcrops. Distribution on Huila Plateau and Chela

Mountains in Angola is shown in insert.

Bothalia 36,2 (2006)

193

diversity of species (six per half-degree square) occurs

in the False Bay area, from Seekoeivlei. including the

Cape Flats, to De Mond at the Palmiet River (3418B)

(Bredenkamp 2002). Levyns (1938) was the first to show

that the Caledon District is the centre of species richness

in the CFR with a reduction in numbers to the north

and east. Oliver et al. (1983) regard the quarter-degree

square 3418BB as the richest area in the CFR. Beyers

(2001), also working in the Thymelaeaceae, found that

the highest number of Lachnaea L. species occurred in

the quarter-degree square 3319AD (Worcester).

Species endemic or near-endemic to the CFR (Table 1)

Passerina shows a high percentage of endemism

within the CFR, as nine species out of 20 (45%), as well

as the three subspecies, are endemic to this region. High

percentages of species, 25%-65%, are also demonstrated

in each of the local centres of endemism. P. esterhuyse-

niae (Northwestern Centre), P. paludosa (Southwestern

Centre),/3, galpinii (Agulhas Plain Centre) and/3, pendula

(Southeastern Centre) are all endemic to one local centre

of endemism only.

The Northwestern Centre has a relatively high con-

centration of Passerina species, as 40% of the species

occur there. The occurrence of 65% of Passerina spe-

cies in the Southwestern Centre, confirms that the grids

3419 (Caledon) and 3418 (Simonstown) can be regarded

as centres of total species richness [Levyns (1938) and

Oliver el al. (1983)]. Geology and soils play an impor-

tant role in the species composition of the Agulhas Plain

Centre, where limestones extensively outcrop along the

southern coast from the Agulhas Peninsula to Mossel

Bay (Goldblatt & Manning 2000). Thirty percent of

Passerina species occur in this centre. The percentages

of Passerina species represented in the Karoo Mountain

Centre (25%) and the Langeberg Centre (35%) are

relatively low, and no Passerina species are endemic to

either of these Centres. The Southeastern Centre has a

relatively high concentration of Passerina species, with

40% of the species occurring there.

Species considered near-endemic to the CFR are

Passerina comosa and P. nivicola, distributed from the

CFR to the Northern Cape and P. falcifolia, P. rubra and

P. truncata subsp. truncata distributed from the CFR to

the Eastern Cape. Passerina nivicola, restricted mostly to

mountainous areas, is possibly still under-collected.

Species endemic to the Northern, Western and Eastern

Cape and KwaZulu-Natal (Table 1)

P. obtusifolia and P. corymbosa (= P. vulgaris) are so-

called Cape ubiquists (Weimark 1941) as they are very

common and adapted to a wide range of Cape habitats.

Their distributions currently include all the Centres with-

in the CFR and both occur in three other South African

provinces. Passerina rigida is confined to coastal areas

from South Africa’s western coast to the northeastern

coast of KwaZulu-Natal.

Species endemic or near-endemic to the Great Escarp-

ment of southern Africa

Endemism of Passerina drakensbergensis, P. mon-

tana and P. montivaga is indicated in Table 2.

Based on fossil pollen evidence, Scott et al. (1997)

regard the dryer forest types of East Africa and Australia

as the best apparent analogies for the palaeovegetation

of southern Africa during the terminal Cretaceous to the

early Tertiary. During the Neogene, plant communities

in southern Africa evolved into equivalents of modem

biomes of the subcontinent. Currently, temperate grass-

land is widespread on the interior plateau and includes

fynbos-like vegetation in moist higher-altitude areas

(O’Connor & Bredenkamp 1997). During the Quaternary,

highveld grassland expanded at the expense of woody

vegetation, coupled by a southward spread of relatively

dry mountain fynbos elements. Evidence for the presence

of such fynbos vegetation during the Holocene in the

contemporary Grassland Biome has been found as far

north as the Nyanga Mountains of Zimbabwe (Scott et

al. 1997). A phylogenetic study by Bredenkamp (2002)

indicates that Passerina ftliformis and P. paludosa (both

common in the Cape Peninsula) are probably the most

primitive extant members of the genus, and P. truncata,

P. montana and P. paleacea as the most advanced. We

hypothesize that P. montana probably originated from

an ancestor in the CFR and adapted to the environmental

conditions of the high-mountain Afromontane grassland,

which had a wider northerly distribution during the

Quaternary. Because of environmental changes since

the beginning of the Quaternary, the boundaries of the

Grassland and Savanna Biomes changed, resulting in

relicts of Afromontane grassland and fynbos elements in

high-altitude areas such as Nyanga, the Huila Plateau and

the Auas Mountains. In descriptions of the Afromontane

Region, White (1981, 1983) and Cowling & Hilton-

Taylor (1997) mention the significant outliers of this

phytochorion occurring on the high mountains of West

Africa, the Eastern Zimbabwe Highlands and Angola

(Huila Plateau). Although this broad pattern of distribu-

tion (Cape centre, eastern African extension with reduced

species) is common to other genera as well, Passerina

is unusual because it is represented in all the more

pronounced Afromontane refuges in southern Africa,

especially those isolated western outliers in Namibia and

Angola.

Rennie (1936) argued that the occurrence of certain

species, including species of Passerina, on the Auas

Mountains in Namibia could be interpreted as relicts

of the CFR. suggesting that northward expansion of at

least certain elements of that flora took place along the

west coast into present-day Namibia. Unfortunately the

Passerina specimens from the Auas Mountains avail-

able to him were sterile, resulting in their incorrect

identification as P. truncata (= P. glomerata). As the

most northerly known distribution of P. truncata at the

time was Steinkopf in Namaqualand, he concluded that

P. truncata once ranged further north through Namibia to

the Auas Mountains. However, the specimens from both

Auas and Huila are unmistakeably P. montana, a species

TABLE 1. — Endemism of Passerina species in Cape Floristic Region (CFR), as well as in various provinces of South Africa

194

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TABLE 1 . — Endemism of Passerina species in Cape Floristic Region (CFR), as well as in various provinces of South Africa (cont.)

Bothalia 36,2 (2006)

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Bothalia 36,2 (2006)

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Bothalia 36,2 (2006)

199

distributed mainly along the eastern Great Escarpment.

The present distribution of P. montana renders Rennie’s

interpretation rather improbable.

CONCLUSIONS

In Passerina the highest numbers of species per one-

degree square are concentrated in a belt between the 33°

and 34°S and 18° to 27°E, occurring in the grids 3321

(Ladismith), 3322 (Oudtshoom) and 3419 (Caledon).

The highest diversity of species occurs in the False Bay

area, from Seekoeivlei to the Palmiet River (3418B).

Hence the CFR (Western Cape) is the centre of diversity

for Passerina.

Passerina demonstrates a high degree of regional

endemism, with 45% of the species endemic to the

CFR. Of these endemics, 20% are endemic to one of

four centres of the CFR; 1 0% ( P . montana and P. monti-

vaga) are near-endemic to the Great Escarpment and 5%

(P. drakensbergensis ) is endemic to the Bergville District

in the northern Drakensberg.

Passerina species that are near-endemic or endemic

to the Great Escarpment probably originated in the CFR

and adapted to conditions associated with the high-

mountain Afromontane grassland, a vegetation type

which is hypothesized to have had a much wider dis-

tribution during the Quaternary. The widely disjunct

distribution of P. montana is probably due to subsequent

environmental changes. The boundaries of the Grassland

and Savanna Biomes shifted, resulting in relicts of

temperate grassland and fynbos elements (such as P.

montana) in isolated high-altitude refuges such as the

Waterberg Plateau in Fimpopo (South Africa), Nyanga-

Chimanimani Highlands in eastern Zimbabwe, Huila

Plateau in Angola and the Auas Moutains in Namibia.

ACKNOWLEDGEMENTS

The authors would like to thank Emsie du Plessis for

the technical editing of the manuscript and Sandra Turck

for the figures.

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I ' ' . ..

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Bothalia 36.2: 201-222 (2006)

Comparisons of invasive plants in southern Africa originating from

southern temperate, northern temperate and tropical regions

L. HENDERSON*

Keywords: alien invasive plants, comparisons, southern Africa, temperate and tropical origins

ABSTRACT

A subset of invasive alien plant species in southern Africa was analysed in terms of their history of introduction, rate of

spread, countries/region of origin, taxonomy, growth forms, cultivated uses, weed status and current distribution in southern

Africa, and comparisons made of those originating from south of the tropic of Capricorn, north of the tropic of Cancer and

from the tropics. The subset of 233 species, belonging to 58 families, includes all important declared species and some po-

tentially important species. Almost as many species originate from temperate regions (112) as from the tropics (121). Most

southern temperate species came from Australia (28/36), most tropical species from tropical America (92/121) and most

northern temperate species from Europe (including the Mediterranean) and Asia (58/76). Transformers account for 33% of

all species. More transformers are of tropical origin (36) than of northern temperate (24) and southern temperate origin ( 1 8).

However, 50% of southern temperate species are transformers, compared to 32% of northern temperate and 29% of tropical

species. Southern temperate transformer species are mainly woody trees and shrubs that were established on a grand scale as

silvicultural crops, barriers (hedges, windbreaks and screens) and cover/binders. Most aquatics, herbs, climbers and succulent

shrubs are from the tropics. Ornamentals are the single largest category of plants from all three regions, the tropics having

contributed twice as many species as temperate regions.

INTRODUCTION

All terminology relating to invasive plants such as

‘alien’, ‘invasive’, ‘naturalized’, ‘weed’, ‘environmental

weed’ and ‘transformer’ are according to Richardson et

al. (2000) unless stated otherwise in the text. More than

1 000 alien plant species are known to be naturalized in

southern Africa (Wells et al. 1986). A high proportion

of these species are herbaceous, ruderal and agrestal

weeds. This paper concentrates on a subset of 233 spe-

cies extracted from the book ‘Alien weeds and invasive

plants' by Henderson (2001) and which contains all the

major and some of the emerging environmental weeds.

Major invaders are those invasive alien species that are

well established, and which already have a substantial

impact on natural and semi-natural ecosystems; emerg-

ing invaders currently have less influence but have

attributes and potentially suitable habitat that could result

in increased range and consequences in the next few

decades (Nel et al. 2004). The list of species selected

for this study includes virtually all the declared plants

whose control, propagation and trade are subject to the

Conservation of Agricultural Resources Act, Act 43 of

1983 (CARA), as amended in 2001.

Southern Africa has had a long history of plant intro-

ductions from various parts of the world (Wells et al.

1986). This paper aims to compare the plants that have

originated from northern temperate, southern temperate

and tropical regions in terms of their history of introduc-

tion, rate of spread, countries/region of origin, taxonomy,

growth forms, cultivated uses, weed status and current

distribution in southern Africa.

* Agricultural Research Council: Plant Protection Research Institute,

c/o SANBI, Private Bag XI 01, 0001 Pretoria,

e-mail: henderson@sanbi.org

MS. received: 2005-09-08.

METHODS

The subset of 233 alien plant species selected for

this study includes all declared species under CARA,

excluding two hybrids that originated in South Africa

( Rubus xproteus and Psidium xdurbanensis), and two

eucalypts for which there is little evidence of their inva-

siveness, Eucalyptus paniculata and E. sideroxylon. The

regions of origin were checked against the United States

Department of Agriculture’s ARS Germplasm Resources

Information Network (GRIN) database, the Missouri

Botanical Garden’s MBG: W3TROPICOS database,

Mabberley (1997), and other literature sources.

Southern temperate species are defined as those

species whose region of origin is entirely south of, or

straddles, the tropic of Capricorn. This region includes

the South American countries of Uruguay, Argentina,

Chile and southern Brazil. It also includes New Zealand,

Tasmania and Australia (Australian Central Territory,

New South Wales, Victoria, South Australia, Western

Australia and southern Queensland).

Northern temperate species are defined as those spe-

cies whose region of origin is entirely north of, or strad-

dles, the tropic of Cancer. This region includes Europe,

North Africa, much of Asia, and North America.

Tropical species are defined as those species whose

region of origin occurs entirely within the tropics or

straddles either the tropics of Cancer or Capricorn. This

region includes tropical America (the northern half of

South America, Central America, Mexico and the West

Indies), tropical Africa and Asia (much of India, Thailand

and Malaysia), Indonesia, and tropical Australia.

The earliest dates of occurrence in southern Africa

were obtained from specimen records in the Pretoria

National Herbarium (PRE) and various literature sources.

The quarter-degree squares occupied and current natural-

ized distributions of the species were obtained from the

Southern African Plant Invaders Atlas (SAPIA) data-

202

Bothalia 36,2 (2006)

base and the Pretoria National Herbarium. The SAPIA

database which is housed at the Agricultural Research

Council (ARC) — Plant Protection Research Institute in

Pretoria, currently contains 50 000 locality records of

more than 500 alien plant species. The database includes

records from roadside surveys undertaken from 1 979—

1993 and from the SAPIA mapping project conducted

from January 1994 until December 1998 (Henderson

1998), with further ad hoc records added to the present.

Weed status was extracted from Henderson (2001).

The definitions of the various categories of environmen-

tal weeds are after Swarbrick (1991 ).

Environmental weeds

Transformers — plants which can dominate or replace

any canopy or subcanopy layer of a natural or seminatu-

ral ecosystem, thereby altering its structure, integrity and

functioning.

Potential transformers — plants that are already invad-

ing natural or seminatural habitats, and have the potential

to dominate any canopy or subcanopy layer but not yet

having a marked effect. They are either transformers

elsewhere in the world or showing signs of this ability in

southern Africa.

Special effect weeds — plants which can significantly

degrade the value or purpose for which a natural or

seminatural ecosystem is valued without necessarily

dominating it or greatly altering its vegetational structure

or functioning. Examples include weeds which compete

with and replace similar native plants, are of high visual

impact, poisonous, or have chemical irritants.

Minor weeds — plants that invade and persist in any

canopy or subcanopy layer of a natural or seminatural

ecosystem but cannot or do not dominate that layer or

seriously alter the vegetation structure or its functioning,

although the accumulation of several to many species

may do so.

Ruderal and agrestal weeds

Mostly annual or biennial plants which are primarily

weeds of waste places (ruderals) and cultivated lands

(agrestals).

The lists of species originating from southern temper-

ate, northern temperate and tropical regions are given in

Appendices 1, 2 & 3.

RESULTS AND DISCUSSION

History of introduction of invasive species

Only 15 species were introduced before 1800 and

all had their origins in northern temperate and tropical

regions (Figure 1). The earliest species to arrive before

the colonization of the Cape by the Dutch in 1652, were

Ricinus communis (castor-oil plant) and Achyranthes

aspera (burweed), believed to be of tropical African ori-

gin, and Catharanthus roseus (Madagascar periwinkle).

All three species are likely to have had a long association

with humans in Africa. Ricinus communis and C. roseus

would have been used for their medicinal value, whereas

A. aspera would have been dispersed by domestic live-

stock. There is evidence that R. communis was in the

Eastern Cape more than 1 200 years ago (Brink 1988)

and this begs the question whether it should be regarded

as indigenous and not alien. All three of the aforemen-

tioned species are widespread in southern Africa but have

not become major invaders.

The arrival of the Dutch at Cape Town in 1 652 marks

the start of the introduction of plant species from other

continents that would eventually become major invaders.

Seven species arrived between 1652 and 1700. Species

of northern temperate origin were: Nasturtium offici-

nale (watercress), Quercus robur (English oak), Salix

babylonica (weeping willow), Pinus pinaster (cluster

pine) and P. pinea (stone pine). Species of tropical ori-

gin were: Opuntia ficus-indica (sweet prickly pear) and

Datura stramonium (common thorn apple). A further five

species arrived before 1800. Arundo donax (giant reed)

was the only northern temperate species, whereas spe-

cies from the tropics were Canna indica (Indian shot),

Xanthium spinosum (spiny cocklebur), presumably an

accidental introduction, Opuntia monacantha (cochineal

prickly pear) and Psidium guajava (guava). Six of the

species introduced before 1800 ( Arundo donax, Opuntia

ficus-indica, O. monacantha, Pinus pinaster, Psidium

guajava and Salix babylonica) are, or were previously.

-♦-Southern temperate -"-Northern temperate Tropical

FIGURE 1 . — History of introduc-

tion of species from tropical,

northern temperate and south-

ern temperate regions.

Bothalia 36,2 (2006)

203

major invaders in southern Africa and have transformed

landscapes. They have either reached or are close to the

limits of their distribution in southern Africa. Opuntia

ficus-indica and O. monacantha had reached pest status

in South Africa by the early 1900s but following success-

ful biological control are no longer regarded as problems

in most parts of the country.

The greatest rate of arrival of species (1.45 species/

year) occurred from 1820-1899. It was during this period

that the first southern temperate species, Acacia longi-

folia (long-leaved wattle), was introduced in 1827 from

Australia (Stirton 1978), 170 years after the introduction

of the first northern temperate species. From the 1830s to

the 1880s many more Australian woody species, belong-

ing to the genera Acacia, Atriplex, Eucalyptus, Grevillea,

Hakea, Leptospennurn, Paraserianthes, Pittosporum and

Syzygium were introduced as sand-binders, hedges, fod-

der plants and for timber. The first southern temperate

species of South American origin to be introduced was

Opuntia aurantiaca (jointed cactus) as an ornamental

rockery plant in 1843.

Almost as many species arrived during the 1900s

(103 species) as in the 1800s (115 species). Plants intro-

duced prior to 1850 were largely utility plants, whereas

after 1850 a greater proportion of the species were of

ornamental value. This trend becomes more obvious in

the 1900s. Up to the 1840s there is not a vast differ-

ence between the numbers of species from each of the

three regions (southern temperate — 10, northern temper-

ate— 19, tropical — 15). After 1850 many more species of

tropical origin were introduced than of northern temper-

ate and southern temperate origin. The cumulative spe-

cies curves in Figure 1 show a surge in species arrival

during the 1850s. This may partly be an artefact of the

very detailed records of plants in the Cape Town Botanic

Gardens provided by McGibbon in 1858. Seventy one

species in cultivation in the Cape Town Botanic Garden

at this time are now on the list of 233 major and emerging

invaders. This includes some of the worst environmen-

tal weeds such as Chromolaena odorata (triffid weed),

Lantana camara (lantana), Opuntia aurantiaca (jointed

cactus) and Pereskia aculeata (pereskia).

Rate of spread

Only a very rough estimate of rate of spread (total

quarter-degree squares (QDS) divided by years since

arrival) can be determined with the available data

(Appendices 1, 2 & 3). This estimate is the average

rate of spread of the entire known history of a species

in southern Africa. One would not expect the rate to

be constant over this time period. Historical data from

the SAPIA database provides evidence that some spe-

cies have had a slow rate of spread over much of their

time period followed by exponential growth e.g. Azolla

filiculoides (red water fern) and Campuloclinium macro-

cephalum (pompom weed).

The species that have shown the fastest average rate

of spread in southern Africa are Azolla filiculoides (4.4 1 8

QDS/year, ornamental, tropical), Prosopis glandulosa

var. torreyana and hybrids (mesquite trees: 4.097 QDS/

year, agricultural crops, northern temperate), Populus

xcanescens (grey poplar: 3.945 QDS/year, cover/binder,

northern temperate), Acacia mearnsii (black wattle:

3.007 QDS/year, silvicultural crop, southern temperate),

Agave americana (American agave: 2.986 QDS/year,

barrier, tropical), Melia azedarach (seringa: 2.764 QDS/

year, ornamental, tropical) and Opuntia ficus-indica

(2.501 QDS/year, agricultural crop, tropical). While

the dispersal of all these species has been assisted by

humans, the current distributions of Agave americana

and Populus xcanescens are almost entirely attributed to

human-assisted dispersal. Populus xcanescens spreads

only vegetatively by suckering, whereas Agave ameri-

cana spreads mainly by suckering but also to a limited

extent by seed.

The earliest introductions from all regions of origin

have, on average, spread the furthest. This is shown in

Figure 2 which plots the mean QDS occupied in 2003

against mean residence time for each of the regions of

origin. The conclusion that can be drawn from this graph

is that most species still have a long way to go before

reaching their potential spread.

Countries/regions of origin

Almost as many species originated from temperate

regions (112) as from the tropics (121) (Table 1). Most

southern temperate species came from Australia (28/36),

most tropical species from tropical America (Central

and northern South America, Mexico and West Indies)

(93/121) and most northern temperate species from

Europe, the Mediterranean coastline of southern Europe

and North Africa, and Asia (57/76). Only nine species are

entirely of African and Madagascan origin.

Taxonomy

The subset of 233 species belongs to 58 families

(Table 1). Most families (41) are of tropical origin; 28

families are of northern temperate origin and 1 1 families

of southern temperate origin. The Fabaceae is by far

the largest family with 41 species. Only the Fabaceae,

Cactaceae and Poaceae have species from all three

regions of origin.

The top families, with the most number of species

in each of the regions are: Fabaceae, Myrtaceae and

Proteaceae from southern temperate regions; Rosaceae,

Fabaceae, Pinaceae, Oleaceae and Salicaceae from

northern temperate regions and Fabaceae, Asteraceae,

Solanaceae, Cactaceae and Myrtaceae from tropical

regions.

Most genera are of tropical origin (Table 1 ). However,

the largest genus, Acacia, with 13 species is from the

southern temperate region (Australia). Opuntia is the

only genus with species from all three regions (one

from southern temperate, three from northern temperate,

six from tropical regions). Few other genera are repre-

sented in more than one region ( Cortaderia , Cuscuta,

Eucalyptus, Myriophyllum, Oenothera, Pinus, Solanum,

Syzygium).

Ten of the 58 families ( 1 7%) are alien to southern Afri-

ca (Appendices 1, 2 & 3). One family (Myoporaceae)

is from southern temperate regions, three families

(Fagaceae, Liliaceae sensu stricto, and Pinaceae) from

northern temperate regions and six families (Agavaceae,

Aristolochiaceae, Cannaceae, Casuarinaceae, Papavera-

ceae, Pinaceae and Salviniaceae) from the tropics. The

204

Bothalia 36,2 (2006)

TABLE 1 . — Summary of taxonomy, growth forms, weed status, region of origin and cultivated uses of species originating from southern temperate,

northern temperate and tropical regions. Transformer species are given in bold

Characteristics Southern Northern Tropical Total

temperate temperate

Pinaceae with nine species is the largest alien family. If

it were not for Rhipsalis baccifera, the sole indigenous

cactus in southern Africa, the Cactaceae would also be

an alien family.

Ninety of 128 genera (70%) are alien to southern

Africa (Appendices 1, 2 & 3). Eleven alien genera are

of southern temperate origin, 33 alien genera are of

northern temperate origin and 53 alien genera are of

tropical origin. The remaining genera, with both alien

and indigenous species, have some of the major invaders

Bothalia 36,2 (2006)

205

FIGURE 2. — Mean quarter-degree

squares (QDS) occupied in

2003 against Mean residence

time (years).

e.g. Acacia, Azolla, Caesalpinia, Eichhomia, Lantana,

Rubus, Salix, Solatium, Sesbania.

Growth forms

Most aquatics, herbs, climbers and succulent shrubs

are from the tropics (61 species) compared to only 27

species from temperate regions. Climbers are almost

exclusively of tropical origin (22/25 species). Most

woody trees and shrubs are from temperate regions (80

species) compared to 53 from tropical regions. Equal

numbers of grasses (five species) originate from temper-

ate and tropical regions (Table 1).

Cultivated uses

Ornamentals are the single largest category of plants

from all three regions with 196/233 species having been

used for ornamentation. Of these species, 129 have been

used primarily (i.e. as a major use) as ornamentals, with

twice as many species from tropical regions (86) than

northern temperate (31) and southern temperate regions

(12).

Barrier plants (hedges, windbreaks and screens) are

the next largest category of cultivated plants. Thirty-one

species have been used primarily as barriers, with more

species from temperate regions (24 species) than the

tropics (7 species). Almost equal numbers of agricultural

crop species originated from temperate (14 species) and

tropical regions (11 species). Most silvicultural crops are

of temperate origin (12/15 species).

Weed status

Transformers account for 33% of all species. Thirty-

five transformers are of tropical origin compared to 24 of

northern temperate and 18 of southern temperate origin.

However, 50% of southern temperate species are trans-

formers, compared to 32% of northern temperate and

29% of tropical species. Southern temperate transformer

species are mainly woody trees and shrubs that were

established on a grand scale as silvicultural crops, barri-

ers and cover/binders.

Although ornamentals constitute the largest category

of cultivated plants, all the other categories (barriers,

crops, cover/binders) have a much higher percentage of

transfomer species. Sixty-six percent (18/27 species) of

silvicultural crops and cover/binders are transformers,

with seven species from southern temperate regions,

eight species from northern temperate regions and three

species from the tropics.

Thirty-one alien genera have transformer species; six

are from southern temperate regions; 10 from northern

temperate regions and 18 from tropical regions. Sixteen

genera that have both alien and indigenous species have

transformer species.

Current naturalized distributions

A visual examination of the current distributions of

all species showed that there are about eight major dis-

tribution patterns or zones. These zones are illustrated

in Figures 3 & 4. Further analysis of the species within

each of the three major regions of origin showed that

there was a concentration of species within certain zones

which correlate with the biomes of southern Africa as

defined by Rutherford (1997). The highest percentage

(36%) of northern temperate species occur in the central

high interior or Grassland Biome (Figure 5A which uses

Pyracantha angustifolia, yellow firethom, as an exam-

ple). Forty-four percent of southern temperate species

occur along the southern and southwestern seaboard,

which includes the whole of the Fynbos and Forest

Biomes (Figure 5B which uses Acacia saligna. Port

Jackson, as an example). Fifty-three percent of tropical

species are distributed along the eastern seaboard and

northeastern interior, which coincides with the Savanna

Biome (Figure 5C which uses Jacaranda mimosifolia,

jacaranda, as an example).

CONCLUSIONS

All three regions of origin have made large contribu-

tions to alien plant invasion in southern Africa. Almost

equal numbers of species, genera and families came from

temperate and tropical regions, with the least from the

southern temperate region and most from the tropics. The

earliest introductions from all three regions have spread

the furthest and most species still have a long way to go

before reaching their potential spread.

206

Bothalia 36,2 (2006)

FIGURE 3. — A, Zone 1 : western and central arid zone e.g. Prosopis species, mesquite trees; B, Zone 2A: southern ‘Mediterranean’ zone e.g. Pinus

pinaster, cluster pine; C, Zone 2B: southern and southwestern ‘Mediterranean’ zone e.g. Acacia cyclops, red eye/rooikrans; D, Zone 3:

southern and eastern cool, moist zone e.g. Acacia melanoxylon, Australian blackwood; E, Zone 4A: highveld zone e.g. Pyracantha angus-

tifolia , yellow firethom; F, Zone 4B: highveld zone with extension to seaboard e.g. Populus xcanescens, grey poplar.

Ornamentals are the single largest category of plants

from all three regions but the tropics has contributed

twice as many species as temperate regions. Temperate

regions have provided slightly more transformers than

the tropics and these are mainly plants that have been

cultivated for non-ornamental purposes. The southern

temperate region, with species mainly from Australia,

has provided a disproportionate number of transformers

(18/36 species or 50%), compared with 32% from north-

ern temperate and 29% from tropical regions.

The current distributions of invasive plants in south-

ern Africa are a reflection of the climatic zones of their

origin. Northern temperate species are concentrated in

the cold, high interior or Grassland Biome. Southern

temperate species are concentrated along the southern

and southwestern seaboard which includes the whole

Bothalia 36,2 (2006)

207

FIGURE 4. — A, Zone 5: eastern seaboard and escarpment e.g. Chromolaena odorata , triffid weed; B, Zone 6A: eastern seaboard, escarpment and

middleveld e.g. Jacaranda mimosifolia, jacaranda; C, Zone 6B: eastern seaboard and escarpment e.g. Lantana camara , lantana; D, Zone

7A: dry interior e.g. Opuntia imbricata, imbricate cactus; E, Zone 7B: dry interior and extension to moister areas e.g. Datura stramonium ,

common thorn apple; F, Zone 8: widespread e.g. Arundo donax, giant reed.

of the Fynbos and Forest Biomes. Tropical species are

concentrated along the eastern seaboard and northeast-

ern interior which coincides with the greater part of the

Savanna Biome.

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]□□□□□□ •□□□□£

:□□□□□□[

•□□□[

II u )l if

laaooQaDDaaaagonnDBDODDDDnnDDaDDnc

Forest

Grassland

Nama Karoo

Savanna

Succulent Karoo

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i). :i i in □ i jit 'i ii i r :u:;i i i

ci i i hi n i ij i ii

)[ : □□□□□□( 10 l ir □ JCX. I I

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APPENDIX 1.- — Southern temperate species: summary of information. Quarter-degree squares (QDS) were obtained from SAPIA database

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APPENDIX 2. — Northern temperate spceies: summary of information. Quarter-degree squares (QDS) were obtained from SAPIA database

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APPENDIX 3. — Tropical species: summary of information. Quarter-degree squares (QDS) were obtained from SAPIA database (cont.)

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‘Alien genera and families not indigenous in southern Africa; PRE, Pretoria National Herbarium; u, underestimated; #, primary use.

APPENDIX 3. — Tropical species: summary of information. Quarter-degree squares (QDS) were obtained from SAPIA database (cont.)

222

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Bothalia 36,2: 223-242 (2006)

South African National Biodiversity Institute: administration and

research staff 31 March 2006, publications 1 April 2005-31 March

2006

Compiler: B.A. Momberg

CHIEF DIRECTORATE (ADIR)

CAPE TOWN— PEARSON HOUSE

Huntley, Prof. B.J. M.Sc. Chief Executive Officer

Laidler, Mrs S.A. B.Sc.(Agric.)(Hons). Senior Provisioning Admin. Officer. Personal Assistant

Finca, Ms N.F. Specialist Cleaner

PRETORIA

Mabeba, Ms K.F. B.A. (Hons). Secretary to SANBI Board

WORKING FOR WETFANDS PROJECT (EDIR/WF)

Dini, J.A. B. Sc. (Hons). Programme Manager. Pretoria (contract worker)

Beetge, A. N. Dip. (Forestry). Regional Co-ordinator.

Mpumalanga (contract worker)

Buckle, J.D. B. Sc. (Hons). Technical Advisor, Southern

region. Port Elizabeth (contract worker)

Goge, M.C. M.Sc. (Environmental Science). Provincial

Coordinator, KwaZulu Natal (contract worker)

Mangqalaza, Ms S.M. M.Ed. Social Development &

Training Co-ordinator. Pretoria (contract worker)

Manyeza, Ms I.M. Provisioning Admin. Officer (con

tract worker)

Mokhutsane, T.J. Regional Co-ordinator, Limpopo.

Pretoria (contract worker)

Mukhoro, M. B.Sc.(Hons)(Environm.Managem.).

Project Manager, National Wetland Inventory

(contract worker)

Munzhedzi, T.E. B.A. (Ed.). Regional Co-ordinator,

North-West and Free State. Rustenburg (contract

worker)

GLOBAL INVASIVE SPECIES PROGRAMME (GISP)

CAPE TOWN

Jackson, Ms L.F. Ph.D. Director (contract worker)

Barnard, Ms P.E. Ph.D. Senior Specialist Scientist. Working Group Co-ordinator/Scientific and Technical Co-ordi-

nator (contract worker)

Matthews, Ms S.G. Communications Co-ordinator (contract worker)

Whiting, Ms D.M. Senior Administrator (contract worker)

MARKETING AND COMMUNICATION DIRECTORATE (AMAR)

PRETORIA

Director: vacant

Liebenberg, Mrs E.J.L. M.Sc. Control Agricultural Technician. Acting Manager (North)

CAPE TOWN

Van Aswegen, Ms C.E. HED. B.A. (History & Psych.). Principal Communications Officer. Admin, support

HUMAN RESOURCES DIRECTORATE (APER)

PRETORIA

Netshiombo. M.J. B. Admin. (Hons), DPLR(UNISA), AHR(UNISA). Chief Director

Baloyi, Ms I. N.Dip.(Manag. Assist.). Senior Secretary IV

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224

Bothalia 36,2 (2006)

CAPE TOWN

Kriel, Mrs G.A. Dip. (Sec.). Senior Secretary IV

Haupt, Mrs C.S. Specialist Cleaner. Guest house

Karlie, Ms F. Cleaner I

PERSONNEL— CAPE TOWN

Engelbrecht, B. N.Dip.(Hort.), N.Dip.(PRM), Dip. (Forestry). Deputy Director: Human resources management and

health & safety co-ordination

Staal, P.B. Dip.(Soc. Sci.), Cert. Indust. Relations. Assistant Director. Labour & staff relations

Claassen, Ms G.E. Senior Telkom Operator III. Admin.

Support, front line duties

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Dollie, Mrs N.J. Specialist Cleaner

Du Toit, Ms R. HED. Assistant Director. Training and

development

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Operator

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III. Recruitment Clerk

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ment equity

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& IOD admin.

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Singh, S.. B. Com. (Hons). Chief Director. Finance

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Retirement Fund

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tract worker)

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tor. Financial management: general ledger

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III. Creditors

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ENVIRONMENTAL EDUCATION DIRECTORATE (EDIR, EENT/GP)

PRETORIA

Qwathekana, Ms N.M. B. A. (Hons), B.A.(Eng., Geog. & Film Studies III), Dip. (Ed.), M.Phil.(Geog. & Envir. Sci.)

Director

Mpungose, J.E. B.Paed., B. A. (Hons) (Geog.), Advanced Postgrad.Cert.(Ed. Mngt.), M.Phil. (Geog.). Assistant

Director: Environm. Ed. Co-ordinator

Canham, B.J. B.A.(Sociol. & Eng.). Programme

Manager. Greening of the Nation (contract work-

er)

Eyssell, Ms A. B.Sc.(Hons)(Hort.). Senior Environm.

Ed. Officer. Outreach Horticulturist

Kutumela, M.S. (student)

Mahasha, Ms P.M. N.T.C.III(Hort.). (student)

Maphuta, Mrs M.S. Specialist Cleaner, Assistant to cen-

tre manager

Maseola, C.G. N. Dip. (Nat. Cons.) (contract worker)

Mashiyi, G. B.Com. Admin./Finance Officer. Greening

the Nation (contract worker)

Mathaba, T.C. Environm. Ed. Officer

Novellie, Mrs E. HED, B.Sc.(Hons)(Zoo. & Mammolo-

gy). Principal Environm. Ed. Officer

Pillay, Ms R. Senior Provisioning Admin. Officer

Ramabulane, S.A. N.T.C.III(Hort). (student)

Sikhauli, Ms N. B.Sc. (student)

CAPETOWN

Belle, E. (student)

Dlamini, Ms N. (student)

Motsoko, Ms N. (student)

Nketu, M.J. (student)

Bothalia 36.2 (2006)

225

EASTERN CAPE

Mchunu, Ms E.N. B.A. Project Officer. Butterworth (contract worker)

Mpongwana, Ms S.N. B.Bib. Project officer. Cradock (contract worker)

Zondani, V. B.A. (Hons). Regional Co-ordinator: Greening of the Nation (contract worker)

WESTERN CAPE (EECT/C)

Coe, W.F. Regional Co-ordinator (contract worker)

Katise, Ms T.C. (student)

Van Dayar, Ms M.M. HED IV (contract worker)

GOLDFIELDS CENTRE— CAPE TOWN (EECT)

Fullard, D. B.Sc.Ed., B. Ed. (Hons). Deputy Director. Environm. Ed. Co-ordinator

Boyana (Magija), Ms N.F. N.Dip.(Hort.), B. Tech. (Envi-

ronm. Managem.). Senior Environm. Ed. Officer.

Outreach greening

Ellman, Ms R.S. HED, B.Sc. Senior Environm. Ed.

Officer. Resource development

Hey, Ms S.J. HED, B.A.(Geog.). Senior Environm. Ed.

Officer. Garden-based programme

Matthews, M.Z. Specialist groundsman. Outreach

greening

Mgodeli, W.M. Driver II

Mjuleni, Ms L.M. N.Dip.(Hort.). Outreach greening

(student)

Mswazi, M.Q. B.Soc. (student)

September, Ms M. Senior Provisioning Admin. Clerk II.

Admin, support

HAROLD PORTER NBG— BETTY’S BAY

Xaba, P.A. N.Dip.(Hort.). Senior Environmental Education Officer. Overberg Useful Plants Project (YARP/CF)

(contract worker)

Williams, Ms L. N. Dip. (Nature Cons.) (student)

LOWVELD NBG— NELSPRUIT (EENT/MP)

Mamatsharaga, L.A. M.Sc.(Ed.). Assistant Director: Environm. Ed.

Hlalu, Ms X. N.Dip.(Hort). Outreach Horticulturist Mavimbela, Ms S.W. (student)

(contract worker) Nyathi, Ms G.S. N. Dip. (Pers. Assist.). Senior Provisio-

Matshaya, Ms N.N. N.Dip.(Hort.) (student) ning Admin. Clerk II

Randima, Ms G.D. Specialist Cleaner

FREE STATE NBG— BLOEMFONTEIN (EENT/FS)

Moletsane, M.E. B. Ed. (Hons). Principal Environm. Ed. Phangoa, M.P. (student)

Officer. Admin, support Tshabalala, Ms B.N.W. (student)

Ngena, Ms K.M.G. Provisioning Admin. Clerk I

WALTER SISULU NBG— ROODEPOORT (EENT/GW)

Konanani, L.N. (student) Moore, Mrs J.M. N.H.Dip.(Sec.). Senior Provisioning

Kondlo, Ms M. N.Dip.(Hort.), Advanced Cert.Envi- Admin. Clerk II. Admin, support

ronm.Ed. Outreach Horticulturist Vatsha, M.L. B.A., HED. Principal Environm. Ed.

Molefe, Ms K.E. Dip. (Nature Cons.). Senior Environm. Officer

Ed. Officer. Outreach education

BIODIVERSITY PROGRAMMES, POLICY AND PLANNING DIRECTORATE (DBIO)

PRETORIA

Maze, Ms K.E. M.Sc. Director. Biodiversity policy and planning

Matlala, Ms J. Senior Administrative Officer

BIOREGIONAL POLICY AND MONITORING UNIT— PRETORIA (DBIO/DD)

Driver, Ms A.L. M.A., M.B.A. Deputy Director. Bioregional policy and monitoring

Smith, Ms T.J. Ph.D. Bioregional Policy and Products Officer

226

Bothalia 36,2 (2006)

BIODIVERSITY, KRC— CAPE TOWN (DBIO/C)

Roberts, R. B. Sc. (Hons). Chief Information Technology Advisor (contract worker)

Rogers, Ms I.M. Supervisor: data ecoding and georeferencing (contract worker)

BIODIVERSITY PLANNING— PRETORIA (DBIO/PL)

Rouget, M.J.F. Ph.D. Biodiversity Planning Manager. Alien plant invasions, conservation planningJonas, Ms Z.R.

M.Sc.(Conserv. & GIS). Conservation Planner (Cape Town)

Mohamed, Ms B.M. B. Sc. (Hons). TSP GIS Specialist

Wistebaar, Ms N.P. B. Sc. (Hons), (student)

Tshitangano, N.F. B.Sc. (student)

EASTERN CAPE CO-ORDINATION UNIT— PORT ELIZABETH (DBIO/EC)

Cadman, M.J. Ph.D. Bioregional Programmes Co-ordinator: Eastern Cape (contract worker)

Cumming, T.L.C. B.Sc. (Hons). Project Developer (contract worker)

Hartmann, N.R. B.Sc.(Hons)(GIS). Intern Project Officer (contract worker)

Myles, Ms M.L. B.A. Senior Administrative Officer (contract worker)

BIODIVERSITY GIS (BGIS)— CAPE TOWN

Willoughby, S.W. M.A.(Geogr. Sci. ). Biodiversity GIS Project Manager (contract worker)

Cocks, M. M.Sc.(Bot.).Web Developer (contract worker)

Khatieb, Ms S. B.Sc. (Hons). GIS Technician (contract worker)

Potgieter-Haung, Ms W. Financial Admin.

THREATENED SPECIES PROGRAMME— PRETORIA (YDBR/TS)

Foden, Ms W.B. M. Sc. (Cons. Biol.). Programme Manager (contract worker)

Naidoo, Ms K. B.Sc. (Hons). Red List Officer (contract Rigala, Ms Z.C. N.Dip.(Hort.). Data Encoder (contract

worker) worker)

Potter, Ms L. M. Sc. (Cons. Biol.). Red List Officer (con- Victor, Ms J.E. M. Sc. (Plant Syst.), H.Dip.(Joum.). Con-

tract worker) trol Agricultural Scientist. Red List Scientist.

Taxonomy of Rutaceae, Asclepiadaceae

CUSTODIANS OF RARE AND ENDANGERED WILD FLOWERS (CREW) THREATENED PLANT PROJECT

Raimondo, Ms D.C. M.Sc. (Cons. Biol.). National Programme Manager (Pretoria)

Ebrahim, I. N.Dip.(Hort.). Fynbos Programme Manager (Cape Town)

Jacobs, L.E.O. Data Encoder. Cape Town (contract worker)

Von Witt, Ms C.G. Project Co-ordinator: Cape Floristic Region (contract worker)

SUCCULENT KAROO ECOSYSTEM PROGRAMME (SKEP)— CAPE TOWN (YDBR/SK)

Henderson, O.C. Programme Manager

Hartney, Ms D.J. B.A.(Hons)(Envir. & Geogr. Sci.) Programme Developer (contract worker)

Mathys, Ms C.N. Dip. (Journ.). Communications Intern (contract worker)

Williams, Mrs B. Dip. (Admin.). Programme Administrator (contract worker)

NATIONAL GRASSLANDS BIODIVERSITY PROGRAMME— PRETORIA (YDBR/GG)

Nazare, Ms F.C. M.Sc. Programme Co-ordinator (contract worker)

CAPE ACTION PLAN FOR PEOPLE (CAPE) PROJECT

CAPE TOWN

Sandwith, T. Programme Co-ordinator (contract worker)

Madaka, R. eForum Library Intern (contract worker

based at UCT)

Barnett, M. Ph.D. Programme Developer (contract

worker)

Bothalia 36,2 (2006)

227

Court, Ms S.J. N. Dip. (Computer Admin.). Finance and ment Intern (contract worker)

Procurement Manager (contract worker) Parker, Mrs A. B. A. (Hons). Project Developer (contract

Damons, Ms M.H. B.A.(Dev. & Env.). Project Develop- worker)

GARDENS DIRECTORATE— ADMIN STAFF (GDIR)

PRETORIA

Willis, C.K. M. Sc. (Cons. Biol.). Chief Director: Gardens and Horticultural Services

Bagus, Mrs J. N. Dip. (Account.). Senior State Accountant. Poverty Relief Projects

Heilgendorff, J.P. N.H.Dip.(Hort.). Gardens IT Manager

Els, Ms L. N. Dip. (Sec.). Senior Secretary IV

CAPE FLATS NATURE PROJECT— CAPE TOWN (YAPR/CF)

Goldman, Ms T. B.Soc.Sci.(Hons). Project Manager (contract worker)

Hathom, Ms P.M. B. A. (Hons), N.Dip.(Hort.). Capacity building Manager (contract worker)

Martin, Ms M. Cert. Masters Business Serv.(CMBS). Admin. Co-ordinator (contract worker)

URBAN CONSERVATION— CAPE TOWN (GDIR/UC)

Davis, G.W. Ph.D. Deputy Director: Communication. Project management and fund raising

Peter, L.M. Dip.(Hort.). Principal Communications Officer. Communications Manager. Edith Stevens Reserve

Phoswayo, Ms V. Senior Provisioning Admin. Clerk II. Admin, support

INTERPRETATION (GINN)

Roff, J. Cert.Envir.Interpr. & Ed. Communication Officer. Interpretation (Pietermaritzburg)

BUILDING PLANNING, MAINTENANCE & DEVELOPMENT— CAPE TOWN (BPMD)

Linde, D.C. N.T.C.III(Civil & Structural: Building), N.T.C.III (Inspector of Works: Building), M.S.A.I.D, Cert.

Estate Agency. Control Works Inspector

Abrahams. P. Handyman. Building maintenance Peck, W.I. Senior Handyman. Building maintenance

Manasse, S.P. Dip. (Masonry). Artisan Foreman. Buil- Tomlinson E.C. Handyman. Building maintenance

ding maintenance

CURATORS

Behr, Ms C.M. Curator: Pretoria NBGBritz, R.M. Curator: Lowveld NBG (Nelspruit)

Gavhi, M.P. Curator: Free State NBG (Bloemfontein)

Le Roux, PH. Deputy Director. Curator: Kirstenbosch NBG (Cape Town)

Oliver, I.B. Curator: Karoo Desert NBG (Worcester)

Tarr, B.B. Curator: Natal NBG (Pietermaritzburg)

Willcock (nee Turner), Mrs S.L. Curator: Walter Sisulu NBG (Roodepoort/Mogale City)

Xaba, Ms A.C. Curator: Harold Porter NBG (Betty’s Bay)

HAROLD PORTER NBG— BETTY’S BAY (GHPG)

Xaba, Ms A.C. N.Dip.(Hort.). Control Agricultural Technician. Curator

Abrahamse, F. Senior Foreman. Estate maintenance and

development

Arendse, L.P. Auxiliary Services Officer II. Access con-

trol

Arendse, Ms M. Auxiliary Services Officer II. Access

control

Bebe, Ms N. Cleaner I

Bezuidenhout, Mrs H.M. Chief Provisioning Admin.

Officer

Carolus, Ms B.J. B.Tech.(Hort.). Chief Agricultural

Development Technician. Horticulture

Forrester, Ms J.A. N.T.C.IIKHort.). Chief Agricultural

Development Technician. Horticulture

Julies, C.A. Provisioning Admin. Clerk

October, Ms R.P. Dip. (Ed.). Senior Auxiliary Services

Officer. Plant records and admin, support

Smith, E.J. Foreman. General Garden Maintenance

Van Wyk, Ms I. (student)

Van Wyk, A.B. Artisan. General maintenance

228

Bothalia 36,2 (2006)

KAROO DESERT NBG— WORCESTER (GKAR)

Oliver, I.B. N.Dip.(Hort.), N. Dip. (Parks Recrea. & Admin), N. Parks Dip. (Parks Recrea. & Management). Control

Agricultural Technician. Curator

Harris, Ms S. N.Dip.(Hort). Senior Agricultural Devel-

opment Technician. Scientific collections

Kwayimani, P. N.Dip.(Hort). Senior Agricultural

Development Technician. Garden management

Makubalo, F.N. Principal Foreman. Nursery

Mpeke, Ms E.N. Specialist Cleaner

Salonika, Ms A.S.D. Senior Provisioning Admin. Clerk

III. Admin, support

Sibozo, N.E. Driver II. Plant sales

Simani, D.K. Principal Foreman. Plant collections

Viljoen, D.M. N.Dip.(Hort.). Chief Agricultural Devel-

opment Technician. Records Officer

KIRSTENBOSCH NBG— CAPE TOWN (GKBC)

Le Roux, P.H. Dip. (Forestry), N.Dip.(Hort.), N. Dip. (Parks & Recr.), Cert.Turf Management. Deputy Director:

Garden Management. Curator

Adams, T.D. B.Tech.(Hort.). Senior Agricultural Development Technician. Supervisor: Greenhouse

Hitchcock, A.N. N.H.Dip.(Hort.). Control Agricultural Development Technician. Nursery Manager

Morkel, A.T. N. Dip. (Nature Cons.). Control Agricultural Development Technician. Estate Manager

Notten, Ms A.L. B.Sc., N.Dip.(Hort.). Chief Agricultural Development Technician. Interpretive Officer

Trautman, C.E. Artisan. Supervisor: Workshop

Adonis, A. Principal Foreman. Dell & ericas

Adonis, S.J. Senior Foreman. Alien vegetation control

Arends, Ms S.J. Principal Auxiliary Services Officer.

Plant records

Barnes, M. (student)

Bowler, M. Principal Foreman. Annuals

Brown, B.M. N.Dip.(Hort.). Agricultural Development

Technician. Seed room

Crowie, R.W. Principal Foreman. General garden

De Abreu, Ms P. (student)

Emms, P. Kirstenbosch Scholar 2006 (contract worker)

Duncan, G.D. M.Sc., N.Dip.(Hort.). Control Agricul-

tural Development Technician. Bulbs, systematics

of Lachenalia

Engelbrecht, F. Senior Provisioning Admin. Clerk II.

Stores

Engelbrecht, Mrs F.D. Control Auxiliary Services

Officer. Plant records

Fani, F.B. (student)

Grace, T. Senior Provisioning Admin. Clerk III. Stores

& admin, support

Harrower, A.D. B.Sc.(Bot. & Zoo.) Ball Agreement.

Project Manager

Hope, C.F. Senior Handyman. Construction

Jacobs, H.C. Principal Foreman. Plant production

Jansen, K. Principal Foreman. Drivers

Jodamus, Ms N.E. N.Dip.(Hort.). Chief Agricultural

Development Technician. Annuals, Rutaceae,

alpines and Cape endemics

Kamalie, Ms S. Senior Typist. Receptionist

Kayster, G.J. Principal Foreman. Construction

Kuscus, G.W. Principal Foreman. General maintenance

Lusithi, Ms X. (student)

Mathys, Mrs S.S.B. Senior Accounting Clerk III. Reve-

nue and garden statistics

Matthews, I.N. Principal Foreman. Estate & trails

Mbambezeli, N.G. N.Dip.(Hort). Agricultural Devel-

opment Technician. Trees & shrubs

Mitchells, G. Control Specialist Groundsman. Senior

Foreman. Casual staff projects

Morris, J.N.M. Senior Foreman. Proteas

Newman, W. Artisan. Mechanical workshop

Oliver, R.C. N.Dip.(Hort). Senior Specialist Grounds-

man. Nursery (student)

Picane, Ms S. Auxiliary Services Officer II. Tissue cul-

ture

Prins, F.B. Security Officer III

Rudolph, A. Security Officer 111

Shanks, G.R. Ball Agreement. Glass House Assistant

(contract worker)

Smith, Mrs A. Senior Provisioning Admin. Clerk II.

Admin, support

Solomons, T.C. Senior Security Officer II

Tamboer, J.S. Principal Foreman. Nursery services

Twine, Ms M. Chief Agricultural Development Techni-

cian. Proteas & restios

Van Gusling, E.J. Principal Foreman. Mowers

Van der Walt, Mrs L.E. N.Dip.(Hort.). Chief

Agricultural Development Technician. Herbaceous

collections

Van Jaarsveld, E.J. M.Sc., N.Dip.(Hort.). Control Agri-

cultural Technician. Succulents

Van Wyk, F. Principal Auxiliary Services Officer II.

Lable Maker

Viljoen, Ms C.C. N.Dip.(Hort). Chief Agricultural De-

velopment Technician. Plant production

Voigt, W.E. N.Dip.(Hort.). Chief Agricultural Develop-

ment Technician. Dell

Wall, Ms K.E. (student)

VISITORS CENTRE— CAPE TOWN (GKBC/VC)

Struys, Ms S. B.A.(Hons)(Directing), Postgrad. Dip. (Market. Manag.). Assistant Director: Communication. Events

& Centre Manager

Fredericks, Ms N.C.E. Senior Auxiliary Services

Officer. Visitors’ Centre. Information services

Jacobs, A.P. Chief Auxiliary Services Officer. Visitors’

Centre. Information services

Bothalia 36,2 (2006)

229

Malan, Ms C.E. B. Sc. (Hons). Principal Communication

Officer: Tour co-ordinator

Pekeur, Ms A. Senior Provisioning Administration Clerk

II: Events Co-ordinator

Phillips, R. Senior Provisioning Admin. Clerk. Facilities

Officer

Williams, G.C. Senior Auxiliary Services Officer.

Information

LOWVELD NBG — NELSPRUIT (GLOW)

Britz, R.M. N. Dip. (Forestry). Control Agricultural Technican. Curator

Froneman, W.C.F. N.T.C.III(Hort.), N. Dip. (Nature

Cons. & Man.), N. Dip. (Parks & Rec. Admin.),

N.T.C.III(Hort.). Control Agricultural Technician.

Nursery management & garden development

Hurter, P.J.H. B. Sc. (Hons). Control Agricultural Techni-

cian. Garden Manager. Cycad conservation

Le Roux, Ms L. N.H. Dip. (Nature Cons.). Chief Auxil-

iary Services Officer II. Interpretation

Maqungo, Ms V.L.B. Auxiliary Services Officer. Front

line Officer

Mathebula, Ms I.N. Senior Auxiliary Services Officer.

Front line Officer

Mathebula, Ms N.R. Senior Accounting Clerk I. Admin.

Support

Mlombo, Ms T.C. Foreman. Garden

Mukoma, T. Dip.(Hort.), B.Tech.(Agric. Managem.),

B.Tech.(Hort). Agricultural Development

Technician. Horticulturist

Ndlovu, L.D. Senior Foreman. Handyman

Ngwenya, PS. Senior Auxiliary Services Officer II.

Kiosk

Shongwe, V.P. Foreman. Garden

Sibanyoni, Ms S.M. Cleaner II

Van der Walt, Mrs G.A.M. Chief Provisioning Admin.

Clerk

Xozumti, M.M. Principal Foreman. Supervisor. Garden

KWAZULU-NATAL NBG— PIETERMARITZBURG (GKZN)

Tarr, B.B. N. Dip. (Parks & Rec. Admin.), Advanced Dip. (Adult Educ.). Control Agricultural Technician. Curator

Dlungwane, T.R. Principal Foreman. Garden mainte-

nance

Johnson, Ms I. HED, M.Sc. Control Agricultural

Development Technician

Ngiba, S.E. (student)

Nonjinge, S.H.B. N.T.C.III(Hort.). Chief Agricultural

Development Technician

Sibiya, Ms C.P.T. Cleaner II

Van der Merwe, Mrs M.E.H. Senior Provisioning

Admin. Clerk III

Zimu, M.J. Principal Foreman. Garden

FREE STATE NBG— BLOEMFONTEIN (GFSG)

Gavhi, M.P. N.Dip.(Hort.). Control Agricultural Technician. Curator

Barnard, Ms A.D. Senior Provisioning Admin. Clerk III

(part time)

Katise, Ms T.C. (student)

Lepitla, M.H. Senior Foreman. Garden

Mankazana, Ms N. (student)

May, T.S. Foreman. Garden

Ngalo, M.S. Senior Auxiliary Services Officer.

Interpretation

Nyuleka, Ms N.A. Senior Accounting Clerk I

Raditlhare, Mrs E.M. Cleaner II

Rambuwani, L.D. N.Dip.(Hort.). Senior Agricultural

Development Technician. Nursery

Sebolai, R.P.A.N. Senior Handyman. General mainte-

nance

PRETORIA NBG (GPTA)

Behr, Ms C.M. B. Sc. (Hons). Control Agricultural Development Technician. Curator

Baloyi, K.J. Senior Auxiliary Services Officer II.

Information Officer. Garden records

Baloyi, M.S. Dip. (IBM), Dip.(PTM), Dip. (Payroll

Admin.). Senior Provisioning Admin. Clerk I.

Leave records and H.R. support

Bell, Ms F.C. HED, N.Dip.(Hort.). Chief Agricultural

Development Technician.

Creighton, Ms D.D. Senior Provisioning Admin. Clerk

III. Admin, support

Difoloko, J.A. Dip. (Ed.), N.Dip.(Hort). Senior Agricul-

tural Development Technician.

Ferreira, Ms L. B. A. (Fine Art), N. Dip. (Nature Cons.).

Chief Auxiliary Services Officer. Information

Keyter, B.A. Senior Security Officer II

Kutama, B.T. Principal Foreman. Garden: hard land-

scape development and maintenance

Lithudza, E.F. Dip.(Hort). Chief Agricultural

Development Technician

Mabapa, K.I. Cleaner II

Mahange, M.J. B. Tech. (Public Managem. & Admin.).

Senior Provisioning Admin. Officer

Makgobola, Ms M.R. Auxiliary Services Officer II.

Reception & admin. Support

Mahlangu, J.F. Senior Foreman. Garden: machine oper-

ators and irrigation

Mahlangu, R.E. Cert. (Office Admin.), Cert.(Plater.).

Senior Artisan. Workshop and general mainte-

nance

Mangoale, F.L. Artisan. Building construction develop-

ment and maintenance

Masimula, Ms B.M. Specialist Groundsman.

Mkhasibe, Mrs N.S. Dip. (Office Admin.). Senior Pro-

visioning Admin. Clerk I. Leave records and H.R.

support

230

Bothalia 36,2 (2006)

Modisha, M.D. Cleaner II

Naidoo, D.A. N.Dip.(Hort.), Dip. (Fund & Managem.).

Control Agricultural Development Technician.

Ngcobo, Ms B.P (student)

Schiel, A. Cert. (Plater). Artisan. Building construction

development and maintenance

Sibiya, Ms T.R. Cleaner II

Singh, Mrs R. Senior Provisioning Admin. Clerk III.

Admin, support

Solomons, Ms C.V. Principal Auxiliary Services Officer.

Plant records clerk

WALTER SISULU NBG— ROODEPOORT (GSIS)

Willcock (nee Turner), Mrs S.L. B. Sc. (Hons), N.Dip.(Hort). Control Agricultural Technician. Curator

Aubrey, Mrs A.E. B.Tech.(Hort). Chief Agricultural

Development Technician. Plant records, interpreta-

tion, information (part time)

Baloyi, S.J. Handyman. Stores

Dlamini, M.D. N.Dip.(Hort.). Senior Agricultural De-

velopment Technician. Garden, nursery

Hankey, A.J. N.Dip.(Hort), B.Tech.(Hort). Control

Agricultural Development Technician. Garden,

estate, collections, nursery

Head, Mrs S.E. Dip. (Shorthand & Typing). Provisioning

Admin. Officer

Mabela, H.L. (horticultural student)

Mamosebo, M.A. Factotum

Manyikana, T.M. Factotum

Mmola, Mrs B.E. Cleaner II

Mtsweni, P. N.Dip.(Hort.). Senior Agricultural Devel-

opment Technician. Support services, estate

Ndou, A.P. Senior Auxiliary Services Officer II. Infor-

mation services

Ndzondo, Ms N.L. Senior Provisioning Admin. Clerk I

Ndzondo, Mrs P.G. Cleaner II

Nedambale, M.P. Senior Foreman. Garden

Nemalili, M.E. Senior Foreman. Machines and vehicles

Nenungwi, M.S. Senior Foreman. Nursery

Tiro, D.W. Senior Accounting Clerk II

RESEARCH DIRECTORATE (RDIR)

PRETORIA

Smith, Prof. G.F. Ph.D., F.L.S. Chief Director: Research & Scientific Services

Marais, Mrs A.C. Senior Provisioning Admin. Officer. Personal Assistant

Arnold, T.H. Head: Data Management (Pretoria)

Crouch, Prof. N.R. Head: Ethnobotany Unit (Durban)

Donaldson, Dr J.S. Director: Kirstenbosch Research Centre (Cape Town)

Koekemoer, Dr M. Curator: National Herbarium (Pretoria)

Leistner, O.A. D.Sc. F.L.S. Agricultural Scientist (contract worker)

Meyer, Mrs N.L. B. Sc. (Hons). Agricultural Development Technician (contract worker)

Roux, Dr J.P. Curator: Compton Herbarium (Cape Town)

Singh, Ms Y. Curator: Natal Herbarium (Durban)

Steenkamp, Ms Y. Assistant Director: SABONET Regional Project Co-ordinator (Pretoria)

Wolfson, Dr M.M. Director: Research Support Services

KWAZULU-NATAL HERBARIUM— DURBAN (RHED)

Singh, Ms Y. HED, M.Sc. Control Agricultural Development Technician. Taxonomy of Araceae, Hypoxidaceae.

Curator

Zulu Botanical Knowledge Project (contract

worker)

Ngwenya, M.A. Senior Agricultural Development

Technician. Herbarium Officer. Plant identifica-

tion and information, Zulu Botanical Knowledge

Project

Mazibuko, J.V.G. Senior Auxiliary Services Officer.

Herbarium Assistant

Noble, Mrs H-E. Chief Provisioning Admin. Clerk III

Parbhoo, Ms S. B. Sc. (Microbiol.). Data capturer (con-

tract worker)

ETHNOBOTANY UNIT— DURBAN (RETH)

Crouch, Prof. N.R. Ph.D. Deputy Director. Ethnobotany of southern African flora, bioprospecting

Douwes, E. B. Sc. (Hons), (student)

Apollos, Mrs C.E. Senior Provisioning Admin. Clerk II.

Marketing

Glen, H.F. Ph.D. Specialist Scientist. Taxonomy of

trees, cultivated plants; botanical history

Glen, Mrs R.P. M.Sc. Control Agricultural Technician.

Wetland plants of southern Africa

Hlongwane, Mrs N.C. Cleaner II & messenger

Keswa, V. B.Sc. Field worker. Zulu Botanical Know-

ledge Project (contract worker)

Magubane, M.M. Dip.(Agric.). Field work Supervisor.

Bothalia 36,2 (2006)

NATIONAL HERBARIUM— PRETORIA (RHEN)

Koekemoer, Ms M. Ph.D. Deputy Director. Herbarium management. Taxonomy of Asteraceae: Gnaphalieae

231

Bredenkamp, Mrs C.L. Ph.D. Control Agricultural Scientist. Assistant Curator: Public relations. Taxonomy

of Vitex, Passerina, Malvaceae, Sterculiaceae, and other related families

Fish. Mrs L. B.Sc. Principal Agricultural Scientist. Assistant Curator: Collections Manager. Taxonomy of

Poaceae

Herman, P.P.J. M.Sc. Principal Agricultural Scientist. Assistant Curator: Personnel. Taxonomy of Asteraceae

Mothogoane, M.S. Chief Auxiliary Services Officer. Assistant Curator: Herbarium assistants. Wing C

Sebothoma, P.N. Cert. Sec. Principal Auxiliary Services Officer. Assistant Curator: Service room. Plant iden-

tifications co-ordinator

Van Rooy, J. Ph.D. Control Agricultural Scientist. Assistant Curator: Technical staff. Taxonomy and bioge-

ography of mosses

Anderson, J.M. Ph.D. Specialist Scientist. Molteno

Palaeoflora, Gondwana Alive

Archer Mrs C. M.Sc. Principal Agricultural Scientist.

Taxonomy of Cyperaceae, monocotyledons (gen-

eral)

Archer, R.H. Ph.D. Principal Agricultural Scientist.

Taxonomy of mainly Celastraceae, Euphorbiaceae

Bester, S.P. M.Sc. Senior Agricultural Scientist. Taxon-

omy of Apocynaceae, Ericaceae, Rutaceae

Burgoyne, Ms P.M. M.Sc. Control Agricultural Scien-

tist. Mesembryanthemaceae and Crassulaceae

Gotzel, Ms A. Senior Provisioning Admin. Clerk III

Govender, Ms M. B.Sc. Senior Agricultural Develop-

ment Technician. Curation and plant ID in Wing C

Jordaan, Mrs M. M.Sc. Principal Agricultural Scientist.

Taxonomy of Celastraceae: Celastroideae, interac-

tive key to the trees of southern Africa

Kgaditsi, T.W. Senior Auxiliary Services Officer. Speci-

men mounter, general assistant

Klopper, Ms R.R. M.Sc. Senior Agricultural Scientist.

Pteridophyta and selected monocotyledonous

families

Makgakga, M.C. B.Sc. Agricultural Development

Technician. Curation and plant ID in Wing B

Makgakga, K.S. Principal Auxiliary Services Officer.

Herbarium Assistant. Encoding plant specimens,

data capturing, labels typist, curation of Wing D

Makholela, Ms T.M. Ph.D. Principal Agricultural Scien-

tist. Taxonomy of Acanthaceae and Rubiaceae

Maserumule, M.K. Principal Auxiliary Services Officer.

Curation of Wing B

Masombuka, Ms A.S. N. Dip. (Nature Cons.). Principal

Auxiliary Services Officer. Herbarium Assistant.

Curation of Wing A

Meyer, J.J. HED. Chief Agricultural Development Tech-

nician. Bioprospecting Project

Mothapo, M.A. H.Cert.Off.Admin.(DMS). Principal

Auxiliary Services Officer. Label typist

Mpongo, T. B.Sc. Senior Agricultural Development

Technician. Curation and plant ID in Wing A

Nkoane, Ms G.K. Principal Auxiliary Services Officer.

Loans, exchanges, gifts, parcelling, stores

Phahla, T.J. Senior Auxiliary Services Officer. Specimen

mounter of cryptogams, packer, general assistance

Phephu, Ms N. B.Sc. Agricultural Development Tech-

nician. Mosses (contract worker)

Ready, Mrs J.A. N.Dip.(HorL). Principal Auxiliary Ser-

vices Officer. Plant identifications, Helichrysum.

Curation of Wing D

Retief, Ms E. Ph.D. Principal Agricultural Scientist.

Taxonomy of Boraginaceae, Verbenaceae, Lamia-

ceae, Asteraceae, Rubiaceae, Geraniaceae,

Oxalidaceae, Vitaceae

Smithies, Mrs S.J. M.Sc., Dip. Ed. (Moray House). Chief

Agricultural Development Technician. Taxonomy

of Scrophulariaceae sens, lcit., Pedaliaceae, Big-

noniaceae, Lentibulariaceae, Gesneriaceae, Mar-

tyniaceae, Orobanchaceae

Steyn, Ms C.C. Principal Auxiliary Services Officer.

Scientific support

Swelankomo, Ms N. B.Sc. (Hons). Senior Agricultural

Development Technician. Curation and plant ID in

Wing D

Welman, Ms W.G. M.Sc. Principal Agricultural Scien-

tist. Taxonomy of Convolvulaceae, Solanaceae,

Cucurbitaceae, Asteraceae: Senecioneae, Acan-

thaceae

Winter, P.J.D. M.Sc. Principal Agricultural Scientist.

Taxonomy of mainly Apiaceae

AFRICAN PLANTS INITIATIVE [API] (CEPF)

PRETORIA

Rampho, Ms E.T. B.Sc. Chief Agricultural Development Technician. Project Co-ordinator (RHEN)

Chiliza, S.B. Senior Herbarium Assistant (contract

worker)

Grunyuza, Ms T. N. Dip. (Fine Art). Senior Herbarium

Assistant (contract worker)

Khumalo, Ms A.N. Senior Herbarium Assistant (con-

tract worker)

Madlala, E.N. Senior Herbarium Assistant (contract

worker)

Mashua, Ms T.J. (student)

Mnengwane, Ms J. J.J. Senior Herbarium Technician

(contract worker)

Moeaha, Ms M.J. Senior Herbarium Technician.

Poaceae Project (contract worker)

232

Bothalia 36,2 (2006)

Mudau, Ms A.C. Senior Herbarium Technician. Poaceae Nthungeni, N. Senior Herbarium Assistant (contract

Project (contract worker) worker)

Nembudani, M.T. B.Sc. Senior Herbarium Technician. Tshidada, Ms N.J. B.Sc. Senior Herbarium Assistant

Poaceae Project (contract worker) (contract worker)

COMPTON HERBARIUM— CAPE TOWN

Arendse, S. M.Sc. Project Co-ordinator (contract wor- Mannie, Ms L. M.Sc. (contract worker)

ker) Vlotman, Ms L.R. Data Capturer (contract worker)

Davids, Ms N. Data Capturer (contract worker) Williams, Mrs V.J. Dip. (Ed.). Data Capturer (contract

Smith, Ms M. Data Capturer (contract worker) worker)

DURBAN

Majola, S.B. Data Capturer (contract worker) Marimuthoo, Ms D. Data Capturer (contract worker)

DATA MANAGEMENT— PRETORIA (RPDC)

Arnold, T.H. M.Sc. Principal Data Technologist. Assistant Director. Computer database application especially in

taxonomy

Boman, Ms M.J. PRECIS data typist

Botha, Mrs A.G. Chief Auxiliary Services Officer.

Administrative Assistant

De Wet, Mrs B.C. B.Sc.(Comp. Sci.), B.A., H.D.L.S.

Principal Agricultural Datametrician. Chief

PRECIS programmer (contract worker)

Mashilo, M.B. B.Sc. (Info. Technol.), IT Support Officer:

API medicinal plants (contract worker)

Montshonyane, Ms E.M. Senior Herbarium Assistant:

API medicinal plants (contract worker)

Mostert (nee Joubert), Mrs R.E. B.Sc. (Hons). Agricul-

tural Scientist. PRECIS Information Officer

Mphephu, T.A. Scientific Officer (contract worker)