ISSN 0006 8241 = Bothalia

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Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 31,1

May 2001

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

Obtainable from the National Botanical Institute, Private Bag X101, Pretoria 0001, Republic of

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

BOTHALIA

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

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

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

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

published annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

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

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

or of economic value for illustration.

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 ), 1989, by J.M.

& H.M. Anderson.

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

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

RSA.

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

by J.M. Anderson (ed.)

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 31,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NA Si

NATIONAL

Botanical

INSTITUTE

2 Cussonia Avenue, Brummeria, Pretoria

Private Bag X101, Pretoria 0001

ISSN 0006 8241

May 2001

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J. Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, Netherlands

Acknowledgements to referees

Anderberg, Dr A.A. University of Stockholm, Sweden

Archer, Mrs C. National Botanical Institute, Pretoria, RSA

Archer, Dr R. National Botanical Institute, Pretoria, RSA

Baijnath, Prof. H. University of Durban-Westville, RSA

Beaumont, Ms A.J. University of Natal, Pietermaritzburg, RSA

Beyers, Dr J.B.P. National Botanical Institute, Cape Town, RSA

Bremer, Prof. K. University of Uppsala, Sweden

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

Campbell, Prof. B.M. Jalan CIFOR, Bogor Barat, Indonesia

Carter, Ms S. Royal Botanic Gardens, Kew, UK

Crouch, Dr N.R. National Botanical Institute, Durban, RSA

Ellis, Dr R.P. Grassland Research Centre, Agricultural Council, Pretoria, RSA

Faden, Dr R.B. Smithsonian Institution, Washington DC, USA

Fish, Ms L. National Botanical Institute, Pretoria, RSA

Gibson, Prof. A. M.E. Mathias Botanical Garden, Los Angeles, USA

Glen, Dr H.F. National Botanical Institute, Pretoria, RSA

Hanna, Dr W. Coastal Plain Experimental Station, USDA, Tifton, USA

Herman, P.P.J. National Botanical Institute, Pretoria, RSA

Hoffman, Dr M.T. National Botanical Institute, Cape Town, RSA

Hutchings, Ms A. University of Zululand, RSA

Kativu, S. National Herbarium & Botanic Garden, Harare, Zimbabwe

Koopman, Prof. A. University of Natal, Pietermaritzburg, RSA

Lubke, Prof. R.A. Rhodes University, Grahamstown, RSA

Martins, Dr E. Instituto de Investigacao Cientifica Tropical, Lisbon, Portugal

McDonald, Dr D.J. National Botanical Institute, Cape Town, RSA

Miller, Dr J.S. Missouri Botanical Garden, St Louis, USA

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

Nordal, Prof. I. University of Oslo, Norway

Nordenstam, Prof. R.B. Naturhistoriska Riksmuseet, Stockholm, Sweden

Paton, Dr A. Royal Botanic Gardens, Kew, UK

Retief, Ms E. National Botanical Institute, Pretoria, RSA

Steyn, Dr E.M.A. National Botanical Institute, Pretoria, RSA

Stieperaere, Dr H. National Botanical Garden, Meise, Belgium

Smith, Prof. G.F. National Botanical Institute, Pretoria, RSA

Timberlake, Dr J.R. Biodiversity Foundation for Africa, Bulawayo, Zimbabwe

Vorster, Dr P. University of Stellenbosch, RSA

Williamson, Dr G. Cape Town, RSA

CONTENTS

Volume 31,1

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

I.M. OLIVER 1

2. The genus Ehretia (Boraginaceae: Ehretioideae) in southern Africa. E. RETIEF and A.E. VAN WYK 9

3. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 11. F. zuurberg-

ensis, a new species from Eastern Cape and new records for the area. S.M. PEROLD 25

4. Notes on African plants:

Amaryllidaceae. A new species of Cyrtanthus (Cyrtantheae) from the southern Cape, South

Africa. D. A. SNIJMAN 31

Amaryllidaceae. A new species of Brunsvigia (Amaryllideae) from Western Cape, South

Africa. D.A. SNIJMAN 34

Anthericaceae. The correct name in Chlorophytum for Anthericum longistylum. C. ARCHER

and S. KATIVU 40

Asphodelaceae. Aloe greatheadii var. davyana (Alooideae), a new locality in KwaZulu-Natal,

South Africa. G.F. SMITH and N.R. CROUCH 31

Asphodelaceae. Jodrellia fistulosa, a new generic record for the Flora of southern Africa region.

C. ARCHER 39

Asteraceae. A new species of Oedera from Western Cape, South Africa. J.B.P. BEYERS 41

Asteraceae. A new species of Emilia (Senecioneae) from South Africa. J.C. MANNING and

P. GOLDBLATT 46

Commelinaceae. Rediscovery of Triceratella drummondii, and comments on its relationships

and position within the family. N.P. BARKER, R.B. FADEN, E. BRINK and A.P. DOLD . 37

Fabaceae. First distribution record for Brachystegia spiciformis (Caesalpinioideae) in South

Africa. P.J.H. HURTER and E. VAN WYK 43

Fossombroniaceae. Fossombronia nyikaensis, a new species from Malawi. S.M. PEROLD ... 48

Lamiaceae. Plectranthus hilliardiae subsp. australis, a new taxon from Eastern Cape, South

Africa. E.J. VAN JAARSVELD and A.E. VAN WYK 44

5. Leaf anatomy of the genus Passerina (Thymelaeaceae): taxonomic and ecological significance.

C.L. BREDENKAMP and A.E. VAN WYK 53

6. A lexicon of plants traded in the Witwatersrand umuthi shops. South Africa. V.L. WILLIAMS,

K. BALKWILL and E.T.F. WITKOWSKI 71

7. Thirty years of change in the fynbos vegetation of the Cape of Good Hope Nature Reserve, South

Africa. S.D.J. PRIVETT, R.M. COWLING and H.C. TAYLOR 99

8. Patterns of endemism within the Karoo National Park, South Africa. F. RUBIN, A.R. PALMER and

C. TYSON 117

9. Vegetation-environment relationships in a catchment containing a dambo in central Zimbabwe.

I. MAPAURE and M.P. McCARTNEY 135

10. Miscellaneous notes:

Poaceae. The basic chromosome number of the genus Pentameris (Arundinoideae). J.J. SPIES

and R. ROODT 145

11. Book reviews 147

New species, subspecies and combinations in Bothalia 31,1 (2001)

Brunsvigia elandsmontana Snijman, sp. nov., 34

Chlorophytum recurvifolium (Baker) C. Archer & Kativu , comb, nov., 40

Cyrtanthus debilis Snijman , sp. nov., 32

Ehretia alba Retief & A.E.van Wyk , sp. nov., 18

Ehretia rigida (Thunb.) Druce subsp. nervifolia Retief & A.E.van Wyk, subsp. nov., 20

Ehretia rigida (Thunb.) Druce subsp. silvatica Retief & A.E.van Wyk, subsp. nov., 20

Ehretia namibiensis Retief & A.E.van Wyk, sp. nov., 17

Ehretia namibiensis Retief & A.E.van Wyk subsp. kaokoensis Retief & A.E.van Wyk, subsp. nov., 18

Emilia hantamensis J.C. Manning & Goldblatt, sp. nov., 46

Erica anemodes E.G.H.Oliv., sp. nov., 3

Erica limnophila E.G.H.Oliv., sp. nov., 6

Erica richardii E.G.H.Oliv., sp. nov., 1

Erica viminalis E.G.H.Oliv., sp. nov., 5

Fossombronia nyikaensis Perold, sp. nov., 48

Fossombronia zuurbergensis Perold , sp. nov., 25

Oedera epaleacea Beyers, sp. nov., 41

Plectranthus hilliardiae subsp. australis Van Jaarsv. & A.E.van Wyk, subsp. nov., 44

iv

Bothalia 31,1: 1-8 (2001)

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

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

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

ABSTRACT

Four new species of Erica L. from the mountains of Western Cape are described: E. richardii E.G.H.Oliv., rare and

localized on quartzite cliffs in the foothills of the Great Swartberg north of Klaarstroom, E. anemodes E.G.H.Oliv., and E.

viminalis E.G.H.Oliv., both restricted to the Hex River and adjacent mountains, and E. limnophila E.G.H.Oliv., rare around

high altitude marshes in the Wemmershoek and Dutoitskloof Mountains.

1. Erica richardii E.G.H.Oliv., sp. nov., E.flocci-

florae Benth. et E. saxigenae Dulfer affinis propter for-

mam seminum sed a prima floribus albis corolla puberu-

la, habitu tenello non crasso lignoso et a secunda floribus

albis corolla minore et habitu tenello non crasso lignoso

differt. Figura 1 .

TYPE. — Western Cape, 3322 (Oudtshoom): Klaar-

stroom area, Witberg southeast of Farm Droekloof, north-

ern slopes near summit, 1 140 m, (-BC), 12 July 2000,

Oliver 11529 (NBG, holo.; BM, K, MO, NY, PRE, S).

Shrubs varying from very small, ± 10 x 50 mm and

prostrate, to erect and divaricate, up to 250 x 300 mm,

woody, single-stemmed reseeder. Branches : few to many

main branches ( 10— )20— 30(— 60) mm long terminating in a

florescence, no secondary branches, with ± 3 new branch-

es formed each year just below each florescence, all cov-

ered with dense short retrorse white hairs. Leaves 3-nate,

erect, imbricate, linear, 3.5(— 4.8) x 0.8 mm, adaxially

slightly rounded, abaxially rounded, both sides with very

short scattered hairs becoming glabrous, margins subacute

with non-sticky glands in young stages, apex acute some-

times reddish, sulcus narrow open at base; petiole ± 0.6

mm long, glabrous, ciliate. Inflorescence : flowers 3-nate

in 1 whorl at apex of main branches; pedicel ± 7 mm long,

red, covered with dense short white hairs, these somewhat

clustered together in stellate-like groups with few, non-

sticky, very short-stalked glands; bract partially recaules-

cent V3-V2 way up pedicel, oblong to oblanceolate, ± 2.4 x

0.8 mm, covered with short scattered hairs and few, short,

non-sticky glands on margins, creamish white with red-

dish apex, sulcus narrow ± V2 as long as bract; bracteoles

2, placed a short distance above bract, otherwise like bract.

Calyx 4-partite; lobes not laterally imbricate, elliptic to

obovate, ± 2.5 x 1.5 mm long, flattened base, white to

creamish, with semitransparent margins, with stouter

cream-coloured, sometimes reddish, sulcate upper portion,

sparsely and shortly hairy, margins minutely serrated with

occasional short, non-sticky, gland-tipped hairs, sulcus

narrow ± V3 length of sepal. Corolla 4-lobed, globose

urceolate, ± 4.5 x 3.8 mm, covered with very short hairs,

white; lobes ±1x1 mm, rounded entire. Stamens 8, free.

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

7735 Claremont, Cape Town.

MS. received: 2000-09-12.

included; filaments oblong broad, ± 2.0 x 0.3 mm, with

distinct apical S-bend, glabrous, white; anthers bipartite,

basally attached, lanceolate in adaxial view, appendicu-

late; thecae erect, ± 1.1 x 0.6 mm and ovate in side view,

with long pale red hairs between thecae otherwise

glabrous, red-brown with darker apex, appendages obo-

vate, ± 0.4 x 0.2 mm, pink-white, edged with long white

hairs; pollen as tetrads. Ovary 4-locular, globose, ± 1.8 x

2.0 mm, slightly tapered at apex, glabrous, green, soon

turning purplish with well-developed dark red nectaries

around base; ovules ± 20 per locule, spreading laterally

from centrally placed placenta; style ± 2 mm long, mani-

fest, becoming exserted in fruiting stage, green with pur-

ple apex, glabrous; stigma truncate. Fruit a dehiscent

loculicidal capsule, ± 3.5 x 3.5 mm, very hard and woody;

valves splitting halfway open but suberect and spreading

very little, septa mainly on the valves very thin and papery

portion on columella. Seeds asymmetrically angular-

obovoid with folds and ridges, ± 0.9 x 0.6 mm, yellow

occasionally tinged red, testa thickened, deeply reticulate,

cells 50-75 x 40-60 pm, anticlinal walls straight abaxial-

ly, undulate adaxially, inner periclinal walls with nume-

rous small pits. Figure 1.

This new species is related to E. flocciflora and E. sax-

igena , and to a lesser extent E. affinis Benth., mainly

because of the very similar seeds, which are unusual in

the genus with their asymmetrical shape covered with

irregular ridges and folds. The capsules are also similar in

being very woody and not opening very much. Other

shared characters are the long hairy pedicels with

coloured bract and bracteoles, the glabrous, globose

ovary with a short style and truncate stigma, and the

anthers with broad appendages and filaments. It seems to

be closest to E. flocciflora and E. saxigena because all

three possess hairy corollas and sepals, and have similar

leaves. E. flocciflora differs in having the hairs noticeably

long and woolly on the pedicel, bract, bracteoles and

sepals, the flower colour being greenish yellow, the

appendages broader and serrate (not ciliate) and in the

shrubs being extremely woody and erect up to 1 m tall. E.

saxigena differs in having no hairs on the anthers, erose

appendages, a velvety, hairy, much larger crimson corol-

la and a stout woody growth.

Erica affinis can be distinguished by the glabrous

bright pink corolla and sepals, the latter being broad and

2

Bothalia 31.1 (2001)

FIGURE 1. — Erica richardii. A, flowering branch, x 1; B, flowering branchlet; C, stem with leaves removed; D, leaf; E, flower; F, flower, cut

open laterally to show position of internal organs; G, bract; H, bracteole; I, sepal; J, stamen, front, side and back views; K, gynoecium, cut

longitudinally; L, capsule, with one valve removed; M, seed; N, testa cells (outer portion of anticlinal walls ± straight, inner portion jig-

sawed). All drawn from type, Oliver 11529. Scale bars: B, 4 mm; C-G, K, L, 2 mm; M, 1 mm; N, 50 pm.

petaloid and as long as the corolla, and by the broader

bract and bracteoles which are adpressed to the calyx. It

also produces much larger shrubs up to 1.5, rarely 2 m tall.

Superficially the new species looks like the small-

flowered forms of E. spectabilis Klotzsch ex Benth., E.

eustacei L. Bolus, E. tragulifera Salisb. and E. formosci

Thunb. in the shape and colour of the flower, the broad

filaments, hairs on the anthers, and the long hairy pedi-

cel with similar bract and bracteoles. The seeds of these

species, however, are quite different from those of E.

richardii, being rounded and symmetrically shaped and

simply reticulate. All four of these species occur in the

mountains in and around the Little Karoo to the south

and west of Klaarstroom.

Erica richardii has only recently been discovered in a

habitat that is most unlikely for Ericaceae. It grows only

on the north side of cliffs, boulders and rocky outcrops of

quartzites of the Witteberg Series on the boundary of the

dry arid Great Karoo (Figure 2). This habitat occurs in

the low mountain ridges north of the Great Swartberg

FIGURE 2. — Known distribution of

Erica richardii, ■; and E. ane-

modes, •.

Bothalia 31.1 (2001)

3

Range at Klaarstroom where only a few remnants of fyn-

bos vegetation manage to survive, namely one small

patch of a species of Restionaceae and also some plants

of Erica discolor , Phylica axillaris var. microphyllum

(Rhamnaceae) and Agathosma ovalifolia (Rutaceae) on

the moister, cooler, south-facing side of the summit

ridge, but with no species of Proteaceae present. E. floc-

ciflora also grows in arid habitats, but in the dry south-

ern foothills of the Kouga Mountains and E. saxigena in

dry rocky habitats on Anthoniesberg and the far eastern

Swartberg, whilst E. affinis grows at high altitudes in

moister mountain fynbos along the Kouga and

Baviaanskloof Mountains to the Groot Winterhoek area.

This species was sent to us in May 2000 by Richard

Taylor of the Technicon at Saasveld, George, a keen

climber and supporter of the Protea Atlas Project, who

was trying to locate the most northern outliers of

Proteaceae in the low foothills north of the Great

Swartberg Range. He led us to the locality where we

managed to locate 25 plants spread for some 4 km along

the north-facing cliffs and upper part of the summit

ridge. The plants were still in flower in early July with

older fruiting flowers and some buds also present on

some plants.

The plants of E. richardii grow only in crevices in the

rocks and vary considerably in size. The younger plants

and those in very small rock crevices on vertical faces are

small and prostrate, whereas the older ones that have

managed to grow in larger rock crevices or on more hor-

izontal rock faces, are divaricate — the largest one 300

mm tall. Most seem to have a very woody basal stem,

which would indicate that they are all of considerable age.

However, the branches are surprisingly rather thin and

delicate with leaves confined to the upper parts, which is

in stark contrast to the extremely woody, stoutly branched

and leafy shrubs of E. flocciflora and E. saxigena.

Paratype material

WESTERN CAPE. — 3322 (Oudtshoom): Klaarstroom area. Witberg

southeast of Farm Droekloof, summit ridge, (-BC), 1 150 m, 12-07-

2000, frt, Oliver 11535 (NBG. PRE); ibid., 1 040 m, 30-04-2000,

Taylor in NBG 169521 (NBG).

2. Erica anemodes E.G.H.Oliv ., sp. nov., E. caly-

cinae L. (formae Cedromontanae) similissima sed habitu

parvo compacto, calcaribus antherarum angustioribus,

pilis plumosis paucioribus in ramulis pedicellisque, stig-

mate capitato-peltato differt. Figura 3.

TYPE. — Western Cape, 3319 (Worcester): Ceres Dist.,

Waaihoek Peak, west slopes at summit, 1 920 m, (-AD),

12 October 1985, Oliver 8793 (NBG, holo.; K, NY, PRE).

Compact erect shrublets, 200-500 mm tall, single-

stemmed reseeder. Branches : densely and frequently

branching with many main branches ± 20-30 mm long

ending in an inflorescence, occasional secondary branch-

FIGURE 3. — Erica anemodes. A, flowering branch, x 1; B, stem with leaves removed; C, leaf; D, flower; E, bract; F, bracteole; G, sepal, abaxial

view (left) and adaxial view (right); H. corolla; I, stamen, back, side and front views; J, gynoecium; K, ovary, with one side removed; L,

fruit; M, capsule, with one valve removed; N, seed; O, testa cells. All drawn from type, Oliver 8793. Scale bars: B-N, 1 mm; O, 100 pm.

FIGURE 4. — Erica viminalis. A, flowering branch, x 1: B, stem with leaves removed; C, leaf; D, flower; E, bract; F, bracteole; G, sepal; H, sta-

men, back, front and side views; I. gynoecium; J, ovary, with one side removed; K, capsule; L, seed; M, testa cells. All drawn from type,

Esterhuysen 28100. Scale bars: B-G, 2 mm; FI-L, 1 mm; M, 100 pm.

es 2-10 mm long ending in an inflorescence; stems cov-

ered with dense short retrorse simple hairs, no infrafoliar

ridges. Leaves 3-nate, slightly imbricate, erect to sub-

spreading, elliptic-oblong, ± 4.2 x 1.0 mm, adaxially

flattened, abaxially rounded margins subacute, glabrous,

minutely ciliate, when young with few, small, non-sticky

glands on margins, narrowly sulcate, sulcus open at base;

petiole ± 0.7 mm long, adpressed, adaxially puberulous,

minutely ciliate. Inflorescence : 3 flowers in 1(2) whorls

at ends of main and secondary branches; pedicel 3 mm

long, curved, with dense, short, retrorse, simple hairs,

occasionally a few plumose hairs intermixed; bract par-

tially recaulescent in middle of pedicel, ovate to elliptic,

±2x1 mm, acute, glabrous, margins with small non-

sticky glands and basally short hairs, white, sulcus nar-

row in upper '/4; bracteoles 2, just above bract, elliptic to

obovate, ± 1.5 x 0.9 mm, obtuse to subacute, otherwise

like bract. Calyx 4-partite; lobes ovate to elliptic, ± 2.0 x

1 .3 mm, laterally imbricate, adpressed to corolla, apex

subacute and cucullate, white, glabrous, margins with

small, sessile, non-sticky glands and basally few short

hairs, sulcus V4-V3 length of sepal. Corolla 4-lobed,

broadly campanulate, 2.5 x 2.5 mm, glabrous, white;

lobes erect to spreading, ± '/2 length of corolla, ± 1.3 x

1.3 mm, triangular subacute, entire. Stamens 8, free,

included to manifest; filaments ± 2 mm long, linear-

oblong, with strong apical S-bend, glabrous, white;

anthers dorsifixed near base, bilobed, quadrate in adaxi-

al view, appendiculate; thecae erect, adpressed, ± 1.1 x

0.6 mm, ellipsoid in lateral view, dark brown, aculeate on

inner and adaxial margins, appendages pendulous, lance-

olate, ± 0.5 x 0.2 mm, irregularly serrate and with few

hairs, brown; pore ± V2 length of theca; pollen in tetrads.

Ovary 4-locular, broadly ovoid to ellipsoid, 1.0 x 1.5

mm, obtuse, glabrous, with reduced nectaries around

base; ovules ± 12 per locule, subpendulous from large

placenta in upper half; style exserted, ± 2.3 mm long;

stigma capitate-peltate, ± 0.7 mm across. Fruit a dehis-

cent capsule, broadly ovoid, ± 1.1 x 1.8 mm, valves split-

ting almost to the base but only slightly spreading, sep-

tum ± 90% on the woody columella and 10% on valve.

Seeds ovoid-ellipsoid, ± 0.7 x 0.5 mm, occasionally

semi-angular, very shallowly reticulate, shiny, brown;

testa not very hard, 70-100 x 50-70 pm, anticlinal walls

straight to mostly slightly undulate, inner periclinal wall

with numerous small pits. Figure 3.

This new species is characterized by its compact,

dense habit, glabrous leaves, white flowers with erect to

spreading lobes, pedicel mainly with simple hairs and

very few plumose hairs admixed and a capitate-peltate

stigma. It is a high altitude species. It is most similar to

the widespread and common E. calycina , especially the

Cederberg form, which differs in having larger

appendages on the anthers (about twice the size), many

more plumose hairs on the stem and pedicel, a hairy

ovary with truncate stigma, finely hairy leaves, and in

growth is woody and erect up to 1 m tall. Several of the

collections of the new species were originally identified

as being E. calycina.

Bothalia 31,1 (2001)

5

It is also similar to E. laxa [= E. lucida var. laxa\

which differs in having dense plumose hairs on the stem,

leaves and pedicel and pink flowers. It is a coastal endem-

ic on recent sands of the west coast, north of Cape Town.

The typical E. lucida is widespread along the mountains

of the western region from Cape Town to the Kamiesberg

in Namaqualand, and forms sparsely branched, fastigiate

shrubs, up to 1.5 m tall.

Erica anemodes is confined to slopes near the sum-

mits of the highest peaks in the Hex River Mountain

complex and adjoining Keeromsberg between Ceres and

Worcester, mostly in the range of 1 500-1 900 m in alti-

tude (Figure 2). Stokoe’s collection from 1 200 m is the

lowest recorded. The type collection was made near the

summit of Waaihoek (= windy comer), hence the epithet

(Greek, anemodes = windy). This latter locality was on a

rather dry, stony, west-facing slope as opposed to the

records of Esterhuysen and McDonald from southern

slopes, either rocky and/or peaty or cliffs. Only three of

the collections recorded the habit of the species — low

compact shrublets with McDonald noting that the

species was locally common. The type collection came

from a population which contained numerous small

shrublets just in that area, and was made on an excursion

with Ms Esterhuysen.

A comment given by Esterhuysen on her collection

from the Keeromsberg is significant — ‘growing with E.

calycina ’ which would indicate that she noted the dis-

tinctness of this new species in the wild.

It would appear that the species is tending towards

wind pollination since the nectaries are very reduced and

almost non-existent and the stigma is expanded.

Unfortunately the recording of pollen discharge was not

noted when Oliver 8793 was collected.

Paratype material

WESTERN CAPE. — 3319 (Worcester): Milner Ridge Peak, cliffs S

side, 5000-6000 ft [1 520-1 830 m], (-AD), 11-11-1943, Esterhuysen

9383 (BOL); ibid.. S slopes, 5000 ft [1 520 m], 11-11-1943, Ester-

huysen 9381 (BOL); Michell’s Peak, 4000 ft [1 220 m], (-AD), 12-

1920, Stokoe 65 (BOL): Brandwacht Peak, 6000 ft [1 830 m], (-CB),

26-11-1944, Esterhuysen 11024 (BOL); Horseshoe Ridge Peak, rocky

S slopes, 5500 ft [1 680 m], (-CB), 1-11-1953, Esterhuysen 22214

(BOL); Fonteintjiesberg, steep SE slopes, 5000-6000 ft [1 520-1 830

m], (-CB). 20-10-1963, Esterhuysen 30427 (BOL); Keeromsberg,

cliffs, S side, 5000-6000 ft [1 520-1 830 m], (-DA), 8-11-1943,

Esterhuysen 9291 (BOL); ibid., SE slopes overlooking Patryskloof,

1 400 m, 30-10-1988. McDonald 1713 (NBG).

3. Erica viminalis E.G.H.Oliv., sp. nov., bene dis-

tincta propter habitum elatum 3 ad 5 pedum viminalem,

folia quadrinata. flores glabros albos ad roseos calcaribus

antherarum longis tenuibus, et ovulis ± 30 in quoque

loculo. Figura 4.

TYPE — Western Cape, 3319 (Worcester): Worcester

Dist., Hex River Mountains, Milner Peak, on wet shaded

cliffs at base of amphitheatre, E side, 5000 ft [1 520 m],

(-AD), 4 January 1959, Esterhuysen 28100 (BOL, holo.;

BM, E, G, K, L, MO, NBG, NY, P, PRE, S, W).

Erect, robust, willowy shrub, up to 1.5 m tall, single-

stemmed reseeder. Branches', main branches fast-grow-

FIGURE 5. — Known distribution of Erica viminalis, ■: and E.

limnophila. •.

ing with continuous apical growth, secondary branches

occasional to numerous but not at every node, 10-50 mm

long, mostly terminating in an inflorescence, occasional-

ly with continuous apical growth, tertiary branchlets

occasional, intemodes short, 1-2 mm long, sparsely

puberulous with simple spreading hairs. Leaves 4-nate,

subspreading, linear-oblong, 3. 0-5.0 x 0.4 mm, rounded

in t/s, subobtuse, a few short hairs adaxially, sparsely and

shortly ciliate, narrowly sulcate, open at base; petiole ±

0.8 mm long, glabrous, adpressed. Inflorescence : 4 flow-

ers in 1 whorl terminal on secondary, rarely tertiary

branchlets, erect; pedicel ± 3.5 mm long, upper half

glabrous, lower half with few, short, spreading hairs;

bract partially recaulescent ± ]/4 way up pedicel, lanceo-

late, ± 1.3 x 0.3 mm, glabrous, ciliate, sulcus small, nar-

row at apex; bracteoles 2, ± half way up pedicel, nar-

rowly lanceolate, ± 0.8 x 0.2 mm, esulcate, glabrous, cil-

iate. Calyx 4-partite; lobes lanceolate-ovate, ± 1.5 x 0.8

mm, adpressed to corolla, not laterally imbricate,

glabrous, ciliate, sulcus in upper '/3 narrow, pinkish

white. Corolla 4-lobed, ovoid, ± 3.5 x 2.0 mm, white to

pinkish, glabrous; lobes rounded, ± 0.7 x 0.7 mm, entire,

erect to subspreading. Stamens 8, included, free; fila-

ments linear, ± 1 .6 mm long, with apical S-bend, thin and

delicate, glabrous, white; anthers bilobed, lanceolate in

adaxial view, dorsifixed near base, appendiculate; thecae

erect, adpressed, ± 0.5 x 0.3 mm, broadly ovate in lateral,

view, ± smooth, light brown; appendages pendulous, nar-

rowly oblong, ± 0.4 mm long, pilose with spreading

hairs; pore ± 'A of theca; pollen in tetrads. Ovary 4-loc-

ular, broadly ellipsoid to obovoid, ± 1.2 x 1.0 mm, emar-

ginate, glabrous, nectaries around base; ovules ± 30 per

locule, spreading from centrally placed placenta; style ±

1.5 mm long, included; stigma capitate. Eruit a thin-

walled dehiscent capsule, broadly obovoid, ± 1.4 x 1.6

mm, valves splitting ± 2A their length, and to ± 30°, septa

on valves only. Seeds ellipsoid, ± 0.6 x 0.4 mm, slightly

reticulate, pale yellow-brown, testa thin with elongate

6

Bothalia 31,1 (2001)

cells ± 100 x 50 pm, anticlinal walls finely undulate,

inner periclinal walls densely and finely pitted. Figure 4.

Erica viminalis is characterized by its tall growth (3 to

5 ft according to Esterhuysen) with fast-growing wil-

lowy branches, (hence the epithet, viminalis = bearing

long flexible shoots/branches or willowy), 4-nate leaves

with 0-2 sclereids around the midvein, the numerous (±

30) ovules per locule (a large number for a small-flow-

ered species) and the long thin anther appendages.

It is most similar to several species in the section

Orophanes. It shares with E. gracilis the 4-nate leaves,

the similar corolla and sepal shape and size, and the

glabrous ovary and leaves with few sclereids, but that

species has much smaller leaves and darker pink flowers

and forms a dense, small, delicately branched shrub.

With E. leucantha it shares the white flowers, similar

anther appendages, glabrous ovary, similar stigma type

and 4-nate leaves having few sclereids, but it does not

have the glabrous leaves and stems, and the narrow

cyathiform corolla. It is also similar to E. sitiens in many

of the above characters but that species has much larger

flowers with a short pedicel and only 6 ovules per locule

and forms smaller fastigiate shrublets up to 1 m tall.

The species is known only from the type collection

made by Elsie Esterhuysen in 1959. Her locality is at the

base of the ‘amphitheatre’ on the east side of Milner Peak

in the Hex River Mountains (Figure 5). This is the side

of the range that faces down into the Hex River Valley

and is very dissected with a veritable amphitheatre of

very large rocky cliffs and deep gullies. She notes that

the plants grew on wet shaded cliffs and that they were

conspicuous due to their size. Fortunately, some older

flowers with sufficiently matured fruits were also present

on the very floriferous branches, which she collected.

Although we have not seen the species in the wild, the

material reminds us of the rapid growth pattern and habit

of two other species we have recently described, namely

E. oakesiorum from the Riviersonderend Mountains and

E. magnisylvae from near Gansbaai.

4. Erica limnophila E.G.H.Oliv., sp. nov., bene

distincta, habitu semiprostrato, foliis tematis, corolla

pilosa rosea, calcaribus antherarum longis tenuibus, sed

praesertim inflorescentis crescentia apicali extensis, vena

media foliorum, bractearum, bracteolarum, sepalo-

rumque crassa sine callo apicali dignoscenda. Figura 6.

TYPE. — Western Cape, 3319 (Worcester): Paarl Div.,

Wemmershoek Peak, edge of swamp below summit on

east side, 5800 ft [1 770 m], (-CC), 31 December 1944,

Esterhuysen 11239 (BOL, holo.; K, NBG, NY, PRE).

Sprawling, tangled shrublets, ± 150 mm tall, 200-300

mm across, single-stemmed reseeder. Branches: thin del-

icate main branches (30— )50(— 1 00) mm long with con-

tinuous apical vegetative growth; few secondary branch-

es ( 1 0— )20(— 30) mm long with continuous apical growth

or subterminal inflorescence; with simple, short, spread-

ing hairs and with or without short-stalked glands inter-

mixed. Leaves 3-nate, often imbricate, subspreading to

spreading, slightly recurved, elliptic-oblong, 2. 5-3.4 x

0.8- 1.0 mm, adaxially flattened, abaxially rounded, sub-

acute, margin subacute, pilose with few, longer, stouter,

gland-tipped hairs on abaxial surface and margins, sulcus

slightly open revealing thick prominent main vein, and

open basally; petiole 1.0-1. 5 mm long, adpressed,

glabrous, ciliate with short simple hairs and few, longer,

gland-tipped hairs. Inflorescence : 3 flowers in single

whorl terminal or subterminal below vegetative growth

on main and secondary branches, central axis continuing

growth; pedicel 1. 5-2.0 mm long, pilose; bract partially

recaulescent in middle of pedicel, lanceolate, ± 2.8 x 0.8

mm, with foliaceous upper half, coloured pinkish below,

sulcate ± '/3 of its length with prominent midvein, indu-

mentum as in leaf; bracteoles 2, approximate to calyx,

lanceolate, ± 2.8 x 0.6 mm, otherwise like bract. Calyx 4-

partite; lobes not laterally imbricate, ovate-lanceolate, ±

3x1 mm, apex recurved, with green upper 2/3, coloured

below, sulcus ± V3 length of sepal with prominent vein

thickened in upper V3 adaxially, pilose, ciliate with long,

stouter, gland-tipped hairs intermixed. Corolla 4-lobed,

urceolate, 5. 0-5. 5 x 3. 5-5.0 mm, pilose, pale pink; lobes

erect, rounded to subacute, ± 1.5 x 1.2 mm, entire to

suberose. Stamens 8, free, included; filaments linear-

oblong, tapering towards apex, ± 3.0 x 0.2 mm, with api-

cal S-bend, glabrous; anthers bilobed, subquadrate in

adaxial view, dorsally attached near base, appendiculate;

thecae erect, adpressed with diverging apices, ± 0.8 x 0.3

mm, lanceolate in lateral view, orange-brown, strigulose,

appendages pendulous subspreading laterally ± 0.7 x 0. 1

mm, minutely spiculate or with very short teeth; pore ±

'/4 length of theca; pollen in tetrads. Ovary 4-locular, glo-

bose, ±1x1 mm, emarginate, pilose, with large nec-

taries around base; ovules ± 15 per locule, spreading

from placenta in upper half of locule; style ± 2.5 mm

long, straight, glabrous; stigma included, capitellate.

Fruit a dehiscent capsule, septa only on valve. Seeds

immature but testa cells ± 30 x 50 pm, anticlinal walls

undulate and periclinal walls pitted. Figure 6.

The relationships of this new species are rather vague

with characters shared with several different groups of

species. The nearest allies seem to be the E. leptocla-

da-E. flacca complex because of the similar prominent

thickened midrib especially on the sepals — this vein

enlarges considerably in the complex and forms a dis-

tinctive apical callosity which is missing in E. limnophi-

la. The hairiness with glands intermixed, the slightly

widened filament and the low spreading habit, all agree

with this complex, but the differences are the short pedi-

cel, recaulescent bract and bracteoles approximate to the

calyx. In the complex, the bract is characteristically non-

recaulescent (i.e. placed on the main branch) and leaf-

like and the two bracteoles are placed right at the base of

a pedicel which is long, relative to the flower. They also

have pollen in monads which is not shared by E.

limnophila.

Another group to which the new species is similar is

the E. trichoclada-E. cederbergensis group with their

often sprawling habit, stalked glands on the leaves and

stems, hairy corolla, ovary and appendages and the cap-

itate to truncate stigma; even the transverse section of the

leaves is similar with the very narrow epidermis and

numerous sclereids surrounding the midvein — not a very

common feature in the genus. The main difference of the

Bothalia 31,1 (2001)

7

FIGURE 6. — Erica limnophila. A, flowering branch, x 1; B, flowering branchlet; C, stem with leaves removed; D, leaf; E, flower; F, bract; G,

bracteole; H, sepal, side, abaxial and adaxial views; I, stamen, back, side and front views; J, gynoecium; K, ovary, with one side removed;

L, capsule, with one valve removed; M, testa cells. All drawn from type, Esterhuysen 11239. Scale bars: B, 4 mm; C-H, J-L, 2 mm; I, 1

mm; M, 50 pm.

new species lies in the fine very short pubescence as

opposed to the long villous pubescence in the group.

The glands and hairs on the leaves, corolla and ovary,

similar stigma and anther appendages and the similar

transverse section of the leaf is shared with the E. pubes-

cens-E. sphaeroidea group but the species in this group

form erect stout plants and grow in dry habitats and have

multi-whorled inflorescences.

The features of E. limnophila are prostrate, spreading

plants, 3-nate leaves, pilose corolla, long, thin anther

appendages, short pubescence with long, stout, gland-

tipped hairs intermixed, inflorescence continuing with

vegetative growth often before the flowers have opened

and the thickening of the midvein in the leaves, bracts,

bracteoles and sepals, but with no apical callosity. The

thickening of the vein in the sepals and the single-

whorled inflorescence which continues vegetative

growth, are features not seen in any other species of

Erica.

Erica limnophila is another new high altitude species,

but in this case, is confined to the Dutoitskloof/Wem-

mershoek Mountains (Figure 5). It has been recorded

from only two localities about 8 km apart. In both local-

ities its habitat has been recorded as being around the

edge of a marsh/swamp/pan, hence the choice of epithet,

limn - = pertaining to standing water, pools, lakes, -philus

= loving. In this habitat, the species would undoubtedly

be a single-stemmed reseeder. Due to the extensive fires

in this mountainous region during February /March 1999,

fresh material and mature fruits could not be collected.

This will have to wait for at least another five years.

Bothalia 31,1 (2001)

8

Paratype material

WESTERN CAPE. — 3319 (Worcester): Paarl Dist., Wemmershoek

Peak, 5200-5500 ft [1 580-1 680 m], (-CC), 15-12-1968, Esterhuysen

32095 (BOL, NBG); ibid., 5750 ft [1 750 m], 12-1944, Lewis & Stokoe

in SAM 577 85 (NBG, SAM); Worcester Dist., between Goudini

Sneeukop and Deception Peak, 5000 ft [1 520 m], (-CC), 16-12-1973,

Esterhuysen 33446 (BOL, NBG).

Bothalia 31.1: 9-23 (2001)

The genus Ehretia (Boraginaceae: Ehretioideae) in southern Africa

E. RETIEF* and A.E. VAN WYK**

Keywords: Boraginaceae, Ehretia. Ehretiaceae, key, morphology, palynology, pollen, southern Africa, taxonomy

ABSTRACT

The genus Ehretia P.Browne in southern Africa is revised. Six species and five subspecies of Ehretia are currently rec-

ognized in southern Africa, of which E. alba Retief & A.E. van Wyk, E. namibiensis Retief & A.E. van Wyk subsp. namibi-

ensis. E. namibiensis subsp. kaokoensis Retief & A.E. van Wyk, E. rigida subsp. silvatica Retief & A.E. van Wyk and E.

rigida subsp. nervifolia Retief & A.E.van Wyk are newly described. The genus is widely distributed in the region and occurs

in a variety of habitats, ranging from the forests of the Eastern Cape to the hot, arid, semidesert parts of Namibia. Members

of Ehretia in southern Africa are predominantly multistemmed shrubs or small trees. Characters of the leaf, the trichome com-

plement, inflorescence and corolla are used to distinguish between the different species. A key to the species, distribution

maps and illustrations of various micro- and macromorphological as well as palynological features are provided.

CONTENTS

Abstract 9

Introduction 9

Materials and methods 10

Historical outline 10

Morphological characters of taxonomic significance 10

Habit 10

Leaf 10

Inflorescence 11

Pollen 12

Fruit 14

Taxonomic treatment 15

Ehretia P.Browne 15

Key to species 16

1. E. amoena Klotzsch 16

2. E. coerulea Giirke 17

3. E. obtusifolia Hochst. ex DC 17

4. E. namibiensis Retief & A.E.van Wyk 17

5. E. alba Retief & A.E.van Wyk 18

6. E. rigida (Thunb.) Druce 19

Specimens examined 21

Acknowledgements 22

References 22

INTRODUCTION

The genus Ehretia was described by Browne (1756).

In 1759 Linnaeus validated the generic name with the

binominal E. tinifolia. Ehretia, with about 33 species in

the tropics of both the Old and New World, is found par-

ticularly in Africa and Asia, with a few species in tropi-

cal America and the West Indies. Lebrun & Stork (1997)

recognized 12 species in tropical Africa, three of which

extend to southern Africa (Herman 1993). The genus is

widely distributed in southern Africa, occurring in a vari-

ety of habitats ranging from lush forests of the Eastern

Cape to hot, arid parts of Namibia. Members of Ehretia

* National Herbarium, National Botanical Institute, Private Bag X101,

0001 Pretoria.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany, Univer-

sity of Pretoria, 0002 Pretoria.

MS received: 2000-06-12.

in southern Africa are predominantly multistemmed

shrubs or small trees, with rigid, arching or drooping,

often entangled branches. The inflorescences are

cymose, the individual cymules usually scorpioid, termi-

nal on young shoots and/or lateral at apices of abbreviat-

ed branchlets. The style is terminal and bifid, whereas the

fleshy fruit is subglobose with four 1 -locular pyrenes.

The genus belongs to the family Boraginaceae s.l. of

mainly herbs, characterized by inflorescences of scorpi-

oid or helicoid cymes, terminal or gynobasic styles and

fruit usually consisting of four nutlets.

Boraginaceae is sometimes treated as two separate

families, Boraginaceae s. str. and Ehretiaceae (Hutchin-

son 1969). Ehretiaceae is taken to include predominantly

woody taxa such as Cordia and Ehretia but also includes

the herbaceous, monotypic genus Coldenia L. (C.

procumbens L.). However, various characters, including

woodiness, pollen morphology and also the structure of

the inflorescence, show that the two families are closely

related and that the recognition of two separate families

is not justified (Retief & Van Wyk 1999). Since Martius

(1827) proposed the family Ehretiaceae, a view followed

by Lindley (1830), the circumscription of the Boragi-

naceae has been a matter of controversy. Many authors

regarded the two families as a natural entity, and treated

Boraginaceae in a wide sense, e.g. Giirke (1897), Chanda

& Mukherjee (1978), Sahay (1978), Cronquist (1981),

Verdcourt (1991), Brummitt (1992), Thome (1992),

Takhtajan (1997) and the Angiosperm Phylogeny Group

(1998). On the other hand, fewer workers favoured two

separate families, e.g. Hutchinson (1969), Dahlgren

(1989) and Chant (1993). In the present paper we recog-

nise Boraginaceae s.l. with five subfamilies (following

Riedl 1997): Wellstedioideae, Ehretioideae, Cordioideae,

Heliotropioideae and Boraginoideae.

The purpose of this paper is to present a taxonomic

revision of the genus Ehretia in southern Africa.

Diagnostic characters, an identification key, full descrip-

tions of existing and new taxa, illustrations and distribu-

tion maps are provided. The generic description of

Ehretia and also species descriptions are based on occur-

rence in southern Africa and adjacent Flora zambesiaca

10

Bothalia 31,1 (2001)

regions. This paper forms a part of a revision of the

Boraginaceae for the Flora of southern Africa which is

currently in progress.

MATERIALS AND METHODS

Herbarium specimens of Ehretia in BLFU, BM, BOL,

COI, E, GRA, J, K, KNP, NBG, NH, NMB, NU, PRE,

PRU, SAM, UNIN and WIND were used to gather data on

morphological characters, flowering time and distribution.

Extensive fieldwork was done to study live plants in their

natural habitat. Both untreated and acetolysed pollen

grains were studied. Pollen was acetolysed according to

the standard method of Erdtman (1960). For scanning

electron microscopy studies samples were coated with

gold and studied with an ISI-SX-25 SEM. Although ace-

tolysis is the basic technique used by virtually all palynol-

ogists, untreated pollen grains were also studied because

certain characters are destroyed by this treatment.

HISTORICAL OUTLINE

After Browne established the genus Ehretia in 1756,

De Candolle (1845) enlarged the concept of the genus

significantly by publishing about 58 species within the

genus. Many of these species are now treated as segre-

gate genera, for example Bourreria, Carmona , Roche-

fortia and Rotula (Thulin 1987; Miller 1989). In De

Candolle’s revision only Ehretia hottentotica , a species

described by Burchell (1824), is mentioned for South

Africa. De Candolle recognized four sections within

Ehretia and placed E. hottentotica in the section

Bourreria, a taxon characterized by four pyrenes.

Thunberg, the ‘father of Cape botany' (Gunn & Codd

1981), collected a specimen in the Cape which he named

Capraria rigida in his Prodromus plantae capensis of

1800. To him this specimen showed similarity to two

species of Capraria, a genus now placed under Freylinia

(Scrophulariaceae). Certain specimens collected by

Drege (1843) were regarded as belonging to Grumilea , a

genus subsequently sunk under Psychotria (Rubiaceae).

Drege (1847) applied the name Ehretia zeyheriana, a

nomen nudum of Buek (1796-1878), to specimens col-

lected by Ecklon, Zeyher and himself. In 1859 Harvey

also used this name when describing a new species dif-

fering from ‘£. hottentotica' in having larger, less obo-

vate and thinner leaves. He stated that differences ‘may

be owing to a freer growth in better soil, and eventually

the two varieties will perhaps be united’. Harvey men-

tioned that the taxon E. eckloniana (another manuscript

name of Buek) is also closely related to E. hottentotica.

Wright (1904) published a revision of Ehretia in

South Africa in which he used the name E. hottentotica

of Burchell, although he gave Thunberg’s Capraria rigi-

da as a synonym. Druce (1917) had to arrange and exam-

ine a large number of African and Australian plants and

consulted rather critically the Floras of these regions. He

published the name E. rigida (Thunb.) Druce for this

taxon. The method of nomenclature of the Hookerian

school differed from the continental plan in not insisting

upon the permanence of the specific epithet (trivial

name) when transferred to a different genus. Wright

(1904) thus did not use Thunberg’s epithet, but followed

the so-called Kew Rule. He also published another

species for the region, E. amoena Klotzsch, with Galpin

1242 as the only cited specimen.

In a revision of the Boraginaceae of South West

Africa (the present Namibia), Friedrich-Holzhammer

(1967) recognized three microfamilies, namely Helio-

tropiaceae, Boraginaceae and Wellstediaceae. Ehretia,

represented by E. amoena and E. rigida, was placed in

Heliotropiaceae together with Heliotropium and Cordia.

Martins (1990, 1993) revised Boraginaceae for Flora

zambesiaca ( FZ ) and Flora de Mozambique. He record-

ed E. amoena, E. obtusifolia Hochst. ex DC. and E. rigi-

da for the FZ region. These three species names were

also listed by Herman (1993). The present revision, the

first since Wright (1904) published his account of

Ehretia for the Flora of southern Africa ( ESA ) region,

provides descriptions of two new species and the rein-

statement of E. coerulea Giirke. Six species and five sub-

species are accepted, namely E. alba Retief & A.E.van

Wyk, E. amoena Klotzsch, E. coerulea Giirke, E. nami-

biensis Retief & A.E.van Wyk subsp. namibiensis, E.

namibiensis subsp. kaokoensis Retief & A.E.van Wyk, E.

obtusifolia Hochst. ex DC., E. rigida (Thunb.) Druce

subsp. rigida, E. rigida subsp. silvatica Retief & A.E.van

Wyk and E. rigida subsp. nerx’ifolia Retief & A.E.van

Wyk.

MORPHOLOGICAL CHARACTERS OP TAXONOMIC

SIGNIPICANCE

Habit

Species of Ehretia are predominantly shrubs, occa-

sionally small or straggling trees. However, suffrutices

no taller than up to 0.4 m are known to occur in E.

obtusifolia and E. rigida. We support the observation by

Burtt Davy (1922) that the gap between trees and herbs

is bridged by forms of every gradation in size, from those

that may be classed as either trees or shrubs to those

sometimes defined as woody herbs, suffrutices or

‘obscurely shrubby’ plants. The ‘suffrutex type’ of habit,

occurring in the species mentioned, can be regarded as

transitional to the many herbaceous growth forms in the

family. However, shrubs occur in Lobostemon, a genus

more or less endemic to the fynbos of South Africa and

also a member of the mainly herbaceous subfamily

Boraginoideae. The occurrence of suffrutices supports

the view that there is no clear division of Boraginaceae

into two families, Ehretiaceae and Boraginaceae, on the

basis of habit. Various other growth forms (ecotypes) are

found in E. rigida. Plants of subsp. rigida are usually

stunted in habit, whereas those of subsp. silvatica and

subsp. nervifolia are more lush, with branches often

straggling, entangled and arching at the top. E. rigida

subsp. nervifolia, a large, deciduous or evergreen, multi-

stemmed shrub, is sometimes characterized by long,

drooping, precocious branches with pale mauve flowers

appearing early in spring together with the new leaves.

Leaf

Various leaf characters are important in distinguishing

between taxa. Leaves of southern African species of

Bothalia 31,1 (2001)

FIGURE 1. — Outline of leaf blade of Ehretia spp.: A, E. rigida subsp. rigida , Olivier 1208; B, E. rigida subsp. silvatica, Flanagan 605; C, E.

rigida subsp. nervifolia, Jacobsen 4862; D, E. rigida subsp. nervifolia , Compton 29914; E, E. amoena, Codd & Dyer 4497; F, E. amoena ,

Ward 2980; G, E. coendea. Miller B1136; H & I, E. obtusifolia. Barnard 200; J, E. rigida subsp. nervifolia , Bcdsinhas 3595; K. E. alba ,

Smith 2140; L, Seydel 1233; M, E. namibiensis subsp. namibiensis, Midler 997; N, E. obtusifolia, Lang TRV32211. Scale bar: 20 mm.

Illustrations by Gillian Condy.

Ehretia are usually obovate to broadly obovate or ellip-

tic, displaying quite some variety in size and venation

(Figure 1 ). The size of the blade varies from 6-95 x 3-56

mm. the largest ones found in E. amoena (Figure IE, F)

in the relatively moist eastern regions of the Northern

Province, Mpumalanga and Swaziland, to the smallest in

E. alba (Figure IK), a species of the drier central regions

in southern Africa. Reduction in leaf size is regarded as

an adaptation for reducing transpiration under stress con-

ditions. The venation in all species is brochidodromus

and is usually prominent on the lower surface (especial-

ly on dried specimens) in most species, but in E. alba and

E. rigida subsp. ner\>ifolia, occurring under more arid

climatic conditions, the tertiary veins are obscured on the

lower surface.

Three trichome types are found on leaves and other

parts of Ehretia: (a) 2- or 3-celled short hairs, (b) setae

with a 1 -layered, multicellular base, and (c) simple, mul-

ticellular, capitate glandular trichomes (Figure 2A-C).

The indumentum varies from sparse to dense (Figure

2D-F). The blade is sometimes glabrous except for setae

occurring along the margin, a condition characteristic of

various boraginaceous species (Figure 2D). These setae

are often bent and usually appressed. Leaves of E. amoe-

na, for example, are often described as rough (like sand

paper) owing to short, stiff setae orientated at an angle to

the upper surface. E. alba which occurs under relatively

arid climatic conditions, has numerous stomata sunken in

a prominently ribbed epidermal surface (Figure 2G), in

contrast to the smoother surface of E. rigida from a moist

habitat (Figure 2H). Domatia are sometimes present in

the axils of principal lateral veins on the lower surface

(Figure 21), but may be present or absent on leaves of the

same plant. Domatia are not specific for Ehretia in

Boraginaceae; they also occur in Cordia , for example.

The pit domatia (which are usually hairy) are believed to

provide shelter to beneficial mites.

Inflorescence

Inflorescences of the Boraginaceae are characteristical-

ly scorpioid or helicoid cymes, often coiled at the apex,

uncoiling and elongating at maturity. Southern African

species of Ehretia have inflorescences with scorpioid

cymules, borne apically on young shoots and/or at the

apex of abbreviated branchlets (brachyblasts) (Figure 3A,

D). On the basis of the composition of the inflorescence,

local members of Ehretia can be divided into different

12

Bothalia 31,1 (2001)

FIGURE 2. — Ehretia leaf surface. A, E. rigida subsp. rigida, Geldenhuys 1226, multicellular hair; B, E. namibiensis subsp. kaokoensis, De Winter &

Leistner 5435, seta; C, E. rigida subsp. nen’ifolia, Compton 26649, glandular trichome; D, E. alba. Venter, Hahn & Archer 1108, blade

glabrous, setae along margin; E, E. rigida subsp. silvatica. Cooper 77, setae spaced; F, E. amoena. Chase 4160, blade densely hairy; G, E.

alba. Venter, Hahn & Archer 1108, epidermis wrinkled; H, E. rigida subsp. rigida, Geldenhuys 1226, epidermis not strongly wrinkled; I, E.

alba. Venter, Hahn & Archer 1108, domatia. Scale bars: A, 20 pm; B, 48 pm; C, 259 pm; D, 141 pm; E, F, 300 pm; G, H, 9.2 pm; I, 233 pm.

groups. E. amoena and E. coerulea have much-branched

cymose panicles, whereas the other species are characterized

by cymose inflorescences with less branching (Figures 3A,

B; 4 A, B). Inflorescences of Ehretia rigida subsp. rigida ,

however, are borne mainly apically on young shoots where

several occur together in a cluster (or if only one inflores-

cence is present, the peduncle is rather thick, up to 1 .5 mm

diam.), forming a corymb-like structure (Figures 3C, D; 4C,

D). The peduncle of the inflorescence is sometimes very

short, with flowers clustered at the end of an abbreviated

branchlet, a condition found in E. ohtusifolia. An inflores-

cence often develops at the end of a new branch, whereupon

new growth of the branch continues sympodially.

Trichome complements of the inflorescences (and

calyces) differ and can be used to distinguish between

species. E. amoena , E. rigida and E. alba usually do not

have simple, multicellular, capitate glandular trichomes,

whereas E. coerulea, E. ohtusifolia and E. namibiensis are

characterized by this type of trichome, especially on the

inflorescence, calyx and midrib of the leaf. Hybridization

between species is suspected; the presence of unexpected

glandular trichomes may signify this phenomenon.

Pollen

Pollen grains of southern African Ehretia species

(Figure 5A-F), are tricolporate, isopolar, oblate-spher-

oidal with P = 20-31 pm, E = 17-29 pm (immature

pollen grains are prolate, P = 13-22 pm, E = 17-29 pm).

The grains are ± circular in polar view and hexagonal.

Bothalia 31.1 (2001)

13

FIGURE 3. — Ehretia inflorescences and abbreviated branchlets. A, E. rigida subsp. nervifolia, Verdoom 603, terminal and lateral; B, E. rigida

subsp silvatica, Thode STEU6495, inflorescence less compact; C, E. rigida subsp. rigida, Germishuizen 1490, inflorescence terminal,

corymb-like; D, E. rigida subsp. rigida, Germishuizen 1490, abbreviated branchlets. Scale bars: A, 18 mm; B, 10 mm; C, 4 mm; D, 9 mm.

elliptic or rectangular in equatorial view. Apertures are

long, broad or rather narrow in diameter and mesocolpi-

um centres (‘pseudocolpi’) are present. The tectum is

reticulate to rugulose. Orbicules are present, covering the

surface of the thecae. A grain of E. obtusifolia was found

to have four apertures instead of the normal three (Figure

5C). Two pollen types are often recognized in Ehretia.

Pollen of Ehretia is usually hexagonal with relatively

broad apertures and mesocolpium centres (Figure 5A).

Similar pollen grains are known in Wellstedia (Wellste-

dioideae) which is sometimes placed in its own family,

Wellstediaceae and in members of the closely related

family Hydrophyllaceae. Palynological and also morpho-

logical similarities between members of Ehretia , Wellste-

dia and Hydrophyllaceae may point to a common ances-

tor or even to their placement in an entity of their own.

However, elliptic or rectangular pollen grains with nar-

row apertures and ‘furrow-like’mesocolpium centres also

occur (Figure 5D). This ‘pollen type’ shows some simi-

larity to that of members of Heliotropium (Helio-

tropioideae) regarded as part of Boraginaceae s. str. if two

families are recognized, but it is actually only a stage in

the development of the pollen grain to maturity. The retic-

ulate or rugose tectum ornamentation of Ehretia (Ehre-

tioideae) which bears resemblance to some pollen grains

of Cordia (Cordioideae) may be cited in support of the

14

Bothalia 31,1 (2001)

FIGURE 4. — Ehretia inflorescences.

A, E. amoena , Chase 4160',

B, E. rigida subsp. nen’ifolia ,

Pole Evans 1480', C, E. rigida

subsp. nervifolia, Wearne 50;

D, E. rigida subsp. rigida ,

Germishuizen 1522. Scale

bar: 20 mm. Illustrations by

Gillian Condy.

recognition of the separate family Ehretiaceae with the

subfamilies Ehretioideae and Cordioideae.

Fruit

The family Boraginaceae is characterized not only by

scorpioid or helicoid cymes, but its fruit also displays

characters of importance for identification. Most mem-

bers of the Boraginaceae in southern Africa have a fruit

of four nutlets. Outer nutlet surfaces vary in texture and

sculpturing, a useful feature distinguishing between taxa.

The fruit of Ehretia , however, is drupaceous with a thin

to leathery exocarp, fleshy to mucilaginous mesocarp

and bony endocarp. The colourful, bright red or orange

fruits attract birds, which are responsible for seed disper-

sal. The genus can be divided into two groups on the

FIGURE 5.— Ehretia pollen. A-D, pollen grain: A, E. rigida subsp. rigida, Germishuizen 3366 , equatorial view; B, E. rigida subsp. nervifolia,

Ward 7223, polar view; C. E. ohtusif'olia, Verdoorn 2355, with four apertures; D, E. rigida subsp. nervifolia, Hanekom 1859, immature. E,

E. rigida subsp. rigida, Germishuizen 3366, tectum; F, E. rigida subsp. rigida, Germishuizen 3366, granular ecto-aperture with lolongate

endo-apcrture area. Scale bars: A, 4.2 pm; B, 3.4 pm; C, D, 5 pm; E, 1.6 pm; F, 1.8 pm.

Bothalia 31,1 (2001)

15

FIGURE 6. — Ehretia fruit, E. amoena, Brenan 14180. A, pyrene; B, C, surface. Scale bar; A, 459 pm; B, 125 pm; C, 99 pm.

basis of fruit structure; species of the Old World includ-

ing the southern African taxa have a drupe of four

pyrenes with areolate surfaces and with some depressed

areas (Figure 6), whereas species of the New World and

Asia have one or two pyrenes per drupe with a verrucose

surface. The local species show no significant differ-

ences in outer surface pattern of the pyrenes. The differ-

ence in fruit structure between members of Ehretioideae

and Boraginoideae, i.e. drupe versus nutlet, is often used

to support the recognition of a separate family, Ehreti-

aceae. However, this is not entirely true, as Coldenia and

Tiquilia Pers. (Ehretioideae) have nutlets and Tourne-

fortia L. (Heliotropioideae) has drupaceous fruits.

TAXONOMIC TREATMENT

Ehretia P.Browne , The civil and natural history of

Jamaica: 168 (1756); L.: 936 (1759); A.DC.: 502 (1845);

Benth.: 840 (1876); Boiss.: 125 (1879); Giirke: 87 (1897);

C.H. Wright; 4 (1904); Sim; 279 (1907); E.Phillips: 627

(1951); Riedl; 7 (1967); Kazmi: 145 (1970); Taton: 19

(1971); R. A. Dyer: 509 (1975); E.S.Martins: 75 (1990);

J.S.Mill.: 20 (1991); Verde.: 33 (1991); E.S.Martins: 34

(1993); Riedl: 91 (1997); Relief: 181 (2000). Type: E.

tinifolia L.

Suffrutices, shrubs or small trees, usually multi-

stemmed and deciduous. Trie home complement consis-

ting of simple, 2- or 3-celled, short hairs, setae with 1-

layered multicellular bases and simple, multicellular,

capitate, glandular trichomes; young stems, inflores-

cence axes and calyx with the same trichome comple-

ment. Branches rigid, arching or drooping, often entan-

gled. Bark greyish white. Leaves simple, alternate on

young shoots or young branches or clustered at apex of

an abbreviated branchlet, petiolate; leaf blade somewhat

discolorous, obovate to elliptic, apex retuse, truncate,

obtuse, cuspidate or acute, base asymmetric or cuneate,

margin usually entire, ciliate, upper and lower surface

variously hairy or, if glabrous, then still with setae usu-

ally densely arranged along margin; petiole short in pro-

portion to length of midrib. Inflorescence a cymose pan-

icle or a dichotomously branched cyme with cymules

mostly scorpioid, usually ebracteate. Flowers bisexual,

regular, sessile or pedicellate, usually precocious, short-

lived, sometimes fragrant. Calyx deeply divided, usual-

ly hairy outside but glabrous inside; tube shallow; lobes

5, ovate or triangular, apex obtuse to acute, ciliate.

Corolla white, cream-coloured, pale mauve, blue, lilac

or purple, usually 5-lobed, imbricate; tube funnel-

shaped or cylindric; lobes oblong, reflexed, shorter than,

as long as or longer than tube. Stamens as many as

corolla lobes; filaments linear, arising from throat of

corolla tube, exserted; anthers oblong, lobed at base.

Ovary 4-locular, with a single ovule at or above the mid-

dle of each locule; style terminal, flattened, bifid, stig-

mas capitate. Fruit subglobose, drupaceous, breaking up

at maturity into four 1 -locular pyrenes, usually glabrous.

Seed elliptic, convex on back, with a deep cavity on

face.

Species ± 33, in the tropics of both Old and New

World, particularly in Africa and Asia, a few in tropical

America and West Indies; six in southern Africa, wide-

spread (Figure 7).

FIGURE 7. — Distribution of Ehretia in southern Africa.

16

Bothalia 31,1 (2001)

Key to species

la Corolla tube widely funnel-shaped, ± as long as calyx; corolla lobes and tube white; lower surface of leaf blade

prominently veined, densely to sparsely hairy, setae appressed, setae on veins conspicuously orientated at

right angles to veins 1 . E. amoena

lb Corolla tube ± cylindric, gradually widening into a throat, longer than calyx; lobes white, mauve, blue, lilac or

purple, tube white or coloured; leaf blade with venation prominent or obscure on lower surface, glabrous,

except for setae along margin or differently hairy:

2a Inflorescence axes and calyces densely covered with glandular trichomes, often intermixed with slender setae:

3a Inflorescence a much-branched cymose panicle; corolla blue or purplish blue, occasionally white; lower sur-

face of leaf blade prominently veined, densely to sparsely hairy, setae appressed, setae on veins con-

spicuously orientated at right angles to veins 2. E. coerulea

3b Inflorescence a simple cymose panicle; corolla purple to mauve, tube can be white; lower leaf surface not

with setae appressed and conspicuously orientated at right angles to veins, densely glandular pubescent

or pilose:

4a Leaf blade densely hairy throughout with glandular trichomes and long, slender setae or only densely setu-

lose, with glandular trichomes only present on midrib and veins; suffrutex, shrub or small tree of shade

or full sun, rocky ridges, thomveld, woodland or open flats 3 . E. obtusifolia

4b Leaf blade pilose, only midrib with scattered glandular trichomes; shrub of dry river beds, riverine wood-

land, calcrete and rocky outcrops 4. E. namibiensis

2b Inflorescence axes with setae and/or multicellular hairs, axes can appear to be glabrous because multicellular

hairs are short and appressed:

5a Corolla tube and lobes white, young buds mauve; leaf blade glabrous except for setae along margin or scat-

tered multicellular hairs, venation of lower surface obscure or only midrib and lateral veins prominent

when leaves are young; leaf surface wrinkled with numerous sunken stomata when blade thickens under

relatively arid climatic conditions; shrub of open and riverine woodland, dry river beds, rocky hillsides,

red sand 5. E. alba

5b Corolla tube usually white, lobes pale mauve, lilac or purple; venation of lower surface prominent, under

more arid conditions tertiary venation sometimes obscured; leaf surface not wrinkled to somewhat so;

shrub or shrubby tree of valley bushveld, forest, riverine vegetation, kloofs, hillsides, grassland, often

part of a bush clump 6. E. rigida

1. Ehretia amoena Klotzsch in Peters, Naturwis-

senschaftliche Reise nach Mossambique: 248 (1861);

Giirke: 335 (1895); C.H. Wright: 5 (1904); Baker &

C.H.Wright: 24 (1905); Codd: 161 ( 1951); J.H.Ross: 297

(1972); Coates Palgrave: 802 (1977); P.van Wyk: 239

(1984); E.S. Martins: 76 (1990); Verde.: 34 (1991);

E.S. Martins: 36 (1993); Beentje: 576 (1994); Retief &

Herman: 351 (1997); Van Wyk & Van Wyk: 162 (1997).

Type: Mozambique, surroundings of Sena, Peters (B,

holo.; K!).

E. mossambicensis Klotzsch: 249 (1861). Type: Mozambique, Tete,

Rios de Sena, Peters (B, holo.)).

E. stuhlmannii Giirke: 336 (1895). Types: Tanzania, Uzaramo Dist.,

Dar es Salaam, Stuhlmann 57 (B, syn. t); Nyika, Mtindi Market to Kwa

Mkembe, Volkens 35 (B, syn.f).

E. goetzei Giirke: 311 (1900), Type: Tanzania, Morogoro Dist.,

Ukutu, Megeta River, near Kisaki, Goetze 128 (B, holo.f; K!).

E. corymbosa sensu Fosberg: 260 (1974).

E. obtusifolia sensu Pooley: 438 (1993); sensu P.van Wyk: 239

(1984).

Shrub or small tree, single or multistemmed, up to 5

m high, branches arching at top. Leaf blade occasional-

ly elliptic, 45— 95(— 1 14) x 23— 56(— 75 ) mm, apex obtuse,

cuspidate, margin entire, sometimes with one or more

teeth, widely spaced, on either side of apex, veins promi-

nently raised on lower surface, lower surface of leaf

blade prominently veined, densely to sparsely hairy,

setae appressed, setae on veins conspicuously orientated

at right angles to veins, upper surface less hairy, setae

often short and stiff at an angle to surface of leaf blade,

giving a rough texture to blade; petiole 4-10 mm long,

length of petiole : length of midrib = ±1:14. Inflores-

cence a branched cymose panicle, terminal on a young

shoot or terminal on a short young shoot at apex of an

abbreviated branchlet; axes and calyx clothed with setae

and multicellular hairs. Calyx lobes narrowly triangular,

2-3 x 1 mm, apex acute. Corolla white; tube widely fun-

nel-shaped, as long as calyx; lobes reflexed, usually as

long as calyx or longer, apex obtuse, margin not ciliate.

Fruit glabrous; pyrenes 3-4 mm long. Flowering time :

October to December.

Distinguishing characters : corolla white, tube funnel-

shaped; inflorescence a corymbose panicle, setae and

multicellular hairs on axes. Distribution-, eastern part of

southern Africa, extending northwards into Zimbabwe

and further eastwards into Mozambique (Figure 8); also

in Malawi, Zambia, Kenya and Tanzania (Verdcourt

1991 ). Habitat-, margin of coastal forest, bushveld, wood-

land, mopane veld, along watercourses, sandy soil. Com-

mon names: sandpaper bush; skurweblaarbos (Afrikaans);

FIGURE 8. — Distribution of Ehretia amoena in southern Africa.

Bothalia 31,1 (2001)

17

FIGURE 9. — Distribution of E. coerulea, A; and E. obtusifolia , ■. in

southern Africa.

mutupe (Sotho); umklele-omkhulu (Zulu). Utilization:

fruit edible, not very tasty; browsed by game; wood

strong, used for fence posts and stampers; grows quickly

from seed (Pooley 1993). Illustrations : Codd: 161

(1951); Palmer & Pitman: 1946 (1972); Pooley: 438

(1993); Van Wyk & Van Wyk: 163 (1997).

2. Ehretia coerulea Giirke in Botanische Jahr-

biicher Heft 3: 312 (1900); Baker & C.H.Wright: 24

(1905) pro parte; White: 364 (1962). Type: Tanzania,

Iringa Dist., Lukose River, Goetze 484 (B, holo.; K!).

E. coerulea Giirke var. glandulosa Suess. in Suess. & Merxm.: 116

(1951). Type: Marandellas, bush by the river through compound, Dehn

697 (M, holo.!).

Tall shrub or small tree, up to 4(-8) m high. Leaf

blade occasionally asymmetric, 40-80 x 20-60 mm,

apex obtuse to truncate, cuspidate, margin entire or den-

tate in upper half of blade, veins raised on lower surface,

setae perpendicular to veins, appressed, glandular tri-

chomes present upper surface with setae only or glandu-

lar pubescent; petiole 4-10 mm, length of petiole : length

of midrib = 1 : 5-6. Inflorescence a cymose panicle, axes

and calyx with setae and glandular trichomes. Calyx

lobes narrowly triangular, ± 3-4 mm long, apex acute.

Corolla blue to purplish blue, occasionally white; tube

cylindric, occasionally funnel-shaped, ± 4-5 mm long;

lobes shorter than or as long as tube, 3^1 mm long. Fruit

glandular hairy; pyrenes ± 4 mm long. Flowering time :

November to December.

Distinguishing characters : corolla blue to purplish

blue; tube ± cylindric, occasionally funnel-shaped; inflo-

rescence a cymose panicle with setae and glandular tri-

chomes. Distribution: Namibia and Botswana (Figure 9);

also in Zambia, Zimbabwe, Malawi and Tanzania.

Habitat: savanna, woodland, mopane veld, veld adjacent

to riverine vegetation, termitaria, sandstone outcrops.

3. Ehretia obtusifolia Hochst. ex DC., Prodromus

9: 507 (1845); A.Rich.: 83 (1850); C.B.Clarke: 142

(1883); Hiem: 716 (1898); Riedl: 48 (1967); Kazmi: 146

(1970); E.S. Martins; 78 (1990); Verde.: 35 (1991). Type:

Ethiopia, Tigre, Medschara, near Gapdia, Schimper 652

(G, holo.; K!).

E. fischeri Giirke: 336 (1895). Types: Tanzania, Mwanza Dist.,

Kagehi, Fischer 323 (B, syn. t) & Biharamulo/Mwanza Dist., Usinga

near French Mission at Usambiro, Stuhlmann 850 (B, syn. f).

Suffrutex, shrub or small tree, up to 3 m high. Leaf

blade 17-25 x 10-12 mm, apex usually obtuse, margin

entire, venation prominent on lower surface, upper sur-

face with setae and scattered glandular trichomes, lower

surface densely covered with setae and glandular tri-

chomes; petiole up to 2-4 mm long, length of petiole :

length of midrib =1:6. Inflorescence a branched cyme,

one or two at apex of a short abbreviated branchlet,

densely covered with setae and glandular trichomes.

Calyx lobes narrowly ovate to triangular, 2.5-3. 0 mm

long, apex obtuse to acute. Corolla tube cylindric, white,

5 mm long; lobes mauve or blue, ± 4 mm long; margin at

apex ciliate. Fruit glabrous, pyrenes 3. 0-3. 5 mm long.

Flowering time: September to December.

Ehretia obtusifolia is closely related to E. amoena and

E. rigida subsp. nervifolia. Usually they are readily distin-

guished, but in areas where the species are sympatric, spec-

imens with intermediate features occur (Martins 1990).

Distinguishing characters: corolla cylindric, tube

white, lobes mauve or blue; inflorescence axes and calyx

glandular hairy. Distribution: Northern Province, Mpuma-

langa, Gauteng, Swaziland and KwaZulu-Natal (Figure

9); also northwards into Africa, up to Ethiopia; also

found in Iran, Afganistan, Pakistan and India (Riedl

1967). Habitat: bushveld, woodland, grassveld, hill

slopes, bushclumps, stony soil, sandy loam.

4. Ehretia namibiensis Retief & A.E.van Wyk, sp.

nov., E. obtusifoliae affinis sed lamina folii pilosa, sparse

pubescenti cum trichomatibus glandulosis sparsis solum

in costa, setis paginae superioris brevioribus quam illis in

pagina inferiori differt (costa laminae folii non dense

glanduloso-pubescenti); distributione geographica etiam

differt.

TYPE. — Namibia, 2416 (Maltahohe): Farm Bullsport,

(-AB), Liebenberg 5087 (WIND, holo.; K, PRE, iso.).

E. amoena sensu Friedr.-Holzh.: 3 (1967) pro parte.

Shrub, up to 3.5 m high. Stems with scattered setae

and glandular trichomes. Bark light grey. Leaves alter-

nate or clustered on older branches, pilose, petiolate; leaf

blade broadly obovate or broadly elliptic, 20-50 x 10-35

mm, apex mostly retuse, occasionally acute, obtuse or

emarginate, margin entire or shallowly toothed, sparsely

hairy with setae on upper surface shorter than on lower

surface, scattered glandular trichomes present on midrib

of lower surface, dull yellowish green; petiole 3-5 mm

long, length of petiole : length of midrib = 1:5-10.

Inflorescence lateral, single at apex of very abbreviated

branchlet or terminal, one or two together at apex of

branchlet; axes densely hairy with scattered setae and

glandular trichomes. Calyx lobes narrowly triangular, 3

mm long, apex obtuse to acute, margin ciliate. Corolla

lavender-blue or pale mauve, 8-9 mm long; lobes slight-

18

Bothalia 31,1 (2001)

sis, ■; and E. namibiensis subsp. kaokoensis. A, in southern

Africa.

ly shorter than tube. Fruit subglobose, 5-6 x 4—5 mm

long, glabrous or with glandular trichomes.

Distinguishing characters', corolla lavender-blue or

pale mauve, tube cylindric; leaf blade sparsely hairy with

setae and scattered glandular trichomes on midrib only.

Distribution : occurs over a large area of Namibia and

Botswana (Figure 10); according to Giess (1971) it is

known from mopane, thombush and highland savanna in

Namibia.

Key to subspecies

Apex of leaf blade mainly retuse; leaf blade 20-25 x 1 0-20

mm; fruit glabrous 4a. subsp. namibiensis

Apex of leaf blade mainly cuspidate; leaf blade 40-45 x 28-35

mm; fruit glandular hairy 4b. subsp. kaokoensis

4a. subsp. namibiensis

Shrub, 2-3 m high. Leaf blade 20-25 x 10-20 mm,

apex mainly retuse, margin entire, indumentum pilose

with setae and multicellular hairs present; petiole up to 4

mm long, length of petiole : length of midrib = 1 : 5 or 6.

Corolla lavender-blue to mauve or purple; tube ± 5 mm

long; lobes ± 4 mm long. Fruit glabrous. Flowering time :

October to May.

Distribution: Namibia and Botswana (Figure 10).

Habitat: dry river beds, riverine woodland on shallow

soil, calcrete and rocky outcrops. Utilization: extract of

roots drunk by man in hope of enhancing conception by

wife ( Woollard & Bethea 2168). Common name:

' mogorometsa' (Woollard & Bethea 2168).

4b. subsp. kaokoensis Retief & A.E.van Wyk, subsp.

nov., subsp. namibiensi affinis sed magnitudine laminae

folii (40-45 x 28-25 mm non 20-25 x 10-20 mm), apice

plerumque cuspidato (non plerumque retuso), margine

interdum dentibus vadosis, fructu cum trichomatibus

glandulosis sparsis, non glabro, differt.

TYPE. — Namibia, 1813 (Sesfontein); 67.2 km S of

Ombombo on road to Sesfontein, (-DD), De Winter &

Leistner 5823 (PRE, holo.; WIND, iso.).

Shrub 3^4 m high. Leaf blade 40^15 x 28-35 mm,

apex usually cuspidate, margin shallowly toothed; peti-

ole up to 5 mm long, length of petiole : length of midrib

= 1 : 8-10. Corolla pale mauve; tube ± 5 mm long; lobes

± 3 mm long. Fruit with scattered glandular trichomes.

Flowering time: April to May.

Distribution: endemic to Namibia in the Kaokoland

Centre of Plant Endemism (Van Wyk & Van Wyk 1997,

Anderson & Van Wyk 1999), characterized by desert

grassland and shrubland, arid savanna and desert annuals;

probably also present in adjacent southwestern Angola

(Figure 10). Habitat: sandy banks of watercourses.

5. Ehretia alba Retief & A.E.van Wyk, sp. nov., E.

rigidae (Thunb.) Druce affinis sed sic differt: lobi corol-

lae albi (non malvini indici vel purpurei); carina et venae

laterales in pagina inferiora foliorum juvenicumpromi-

nentes (venae tertiariea praeterea prominentes); lobi

calycis anguste triangulares (non triangulares et ovati).

TYPE. — Namibia, 2218 (Gobabis): Breitenberg,

Gobabis Dist., (-DC), Merxmiiller 1071 (WIND, holo.;

PRE, iso.).

Shrub up to 4 m high. Branches rigid. Leaf blade 6-25

x 3-13 mm, apex acute or obtuse; glabrous except for

appressed setae along margin, thin or thick in texture,

drying reddish brown when pressed or blade with lateral

veins obscure, drying the same colour as midrib; midrib

and secondary veins on lower surface prominent; petiole

up to 3 mm long, length of petiole : length of midrib =

1 : 10-15. Inflorescence usually single at apex of abbre-

viated branchlet; axes and calyces with multicellular hairs

and scattered setae; peduncle short. Flowers sometimes

fragrant. Calyx lobes narrowly triangular, up to 3 mm

long; apex acute to acuminate. Corolla white, fading to

cream-coloured; tube 3 mm long; lobes ± 3 mm long, as

long as tube or slightly shorter. Style white, sometimes

tinged purple or lilac. Flowering time: August to October.

Distinguishing characters: corolla tube and lobes

white, young buds pale mauve; leaves thickening when

older and also under unfavourable climatic conditions,

glabrous except for setae along margin and scattered mul-

ticellular hairs, leaf surface wrinkled, with numerous

sunken stomata. Distribution: Namibia, Botswana, Free

State, North-West, Northern Cape (Figure 11); it is found

in different biomes (Rutherford & Westfall 1994), namely

Grassland, Nama-Karoo, Succulent Karoo and Savanna;

according to Acocks (1988), E. alba occurs in Kalahari

Thomveld, Vryburg Shrub Bushveld and Typical Orange

River Broken Veld in South Africa. Habitat: riverine

woodland, open woodland, thomveld and dry river beds,

stony places and red sand. Utilization: grazed. Common

name: bergwitboom(bos) (Afrikaans) — according to Smith

(1966) the flowers produced in great abundance before the

leaves unfold, give the plants a whitish (Afrikaans: wit)

appearance when seen from a distance; stems and branch-

es also of a pallid greyish colour.

Bothalia 31,1 (2001)

19

FIGURE 1 1 . — Distribution of Ehretia alba in southern Africa.

6. Ehretia rigida ( Thunb .) Druce in Report of the

Botanical Society and Exchange Club, British Isles 1916:

620 (1917); Palmer & Pitman: 1943 (1972); Coates

Palgrave: 803 (1977); Venter & Venter: 172 (1996); Van

Wyk & Van Wyk: 163 (1997). Type: South Africa, Cape

of Good Hope, Thunberg UPS14330 (UPS, holo.; PRE,

microfiche!).

Capraria rigida Thunb.: 103 (1800). Type: as above.

Much-branched shrub or multistemmed tree, up to 12

m high. Leaf blade 20-60 x 9-45 mm, apex slightly

retuse, obtuse to acute, occasionally cuspidate, venation

on lower surface distinct, lower surface glabrous or hairy

with setae and multicellular hairs along veins, upper sur-

face glabrous or hairy with setae or multicellular hairs

scattered along veins, coriaceous or thin; petiole 2-10

mm long, length of petiole : length of midrib = 1 : 3-11.

Inflorescence single or more than one terminal on a

prominent new shoot and/or terminal on an abbreviated

branchlet. densely clothed with multicellular hairs and

scattered setae; peduncle varies in thickness. Calyx lobes

ovate to triangular, 2. 0-2. 5 mm long, apex obtuse to

acute. Corolla tube cylindric, 7-10 mm long, usually

white; lobes pale mauve, lilac or purple, shorter or as

long as tube. Fruit glabrous; pyrenes ± 3 mm long.

Distinguishing characters : venation of lower surface

prominent; inflorescence and calyx clothed with setae

and/or multicellular hairs; calyx ovate; corolla tube

cylindric, white; lobes pale mauve, lilac or purple.

Distribution : eastern parts of southern Africa; north-

wards into Zimbabwe and eastwards into Mozambique

(Figure 12). Habitat, valley bush, hillsides, forest, bush

clumps, thomveld, woodland or open flats.

Key to subspecies

la Inflorescences terminal, one or several in a ‘head’ on young

shoots, much-branched; peduncles and pedicels usually

thick, up to 1.5 mm diam.; leaf blade usually somewhat

coriaceous; usually a stunted shrub with branches rigid

and entangled; more or less confined to Albany Centre

of Endemism 6a. E. rigida subsp. rigida

lb Inflorescences terminal on very short young shoots or occa-

sionally terminal on short young shoots at apices of

FIGURE 12. — Distribution of Ehretia rigida subsp. rigida. ■; E. rigi-

da subsp. silvatica. A and E. rigida subsp. nervifolia. ▲, in

southern Africa.

abbreviated branchlets, once- or twice-branched; pedun-

cles and pedicels usually thinner, up to 1.0 mm diam.;

leaf blade thin in texture; known from Bathurst east-

wards to KwaZulu-Natal, further north into Free State,

North-West, Gauteng, Mpumalanga and Northern

Province:

2a Tree or scandent shrub of forest in Eastern Cape and south-

ern KwaZulu-Natal 6b. E. rigida subsp. silvatica

2b Shrub of savanna in KwaZulu-Natal, Free State, North-

West, Gauteng, Mpumalanga and Northern Province

6c. E. rigida subsp. nervifolia

6a. subsp. rigida

Ehretia hottentotica Burch.: 147 (1824); C.H. Wright: 5 (1904) pro

parte. Type: Graaff-Reinet, Burchell 2117.

E. zeyheriana H.Buek ex Harv.: 5 (1859). Syntypes: Eastern Cape,

near Uitenhage, Zeyher 161 (PRE, iso.). Prior PRE40572 (PRE, iso.);

Albany, T. Williamson s.n. (TCD).

Shrub or tree, often stunted, up to 4 m high. Leaf

blade 20-35 x 12-23 mm, apex acute or rounded,

glabrous or main veins with setae and others with scat-

tered setae, coriaceous or thin; petioles 3-8 mm, length

of petiole : length of midrib = 1 : 4—7. Inflorescence ter-

minal, one or several in a ’head’ on young shoots, many-

flowered, peduncles and pedicels thick, up to 1.5 mm

diam., multicellular hairs or long, thin setae present.

Calyx lobes triangular, ± 2. 0-2. 5 mm long, apex obtuse

to acute. Corolla cylindric, 8-10 mm long; tube usually

white; lobes lilac, blue, bluish purple or purple, shorter

than tube. Fruit ± 3 mm long. Flowering time : October

to March.

Distribution: Eastern and Western Cape, more or less

confined to the Albany Centre of Plant Endemism (Van

Wyk & Van Wyk 1997; Anderson & Van Wyk 1999)

(Figure 12). Habitat: scrub, bushclumps. Euphorbia thick-

et, valley bushveld, dune bush. Common names: Cape

lilac — plants bear masses of lilac to lilac-blue flowers and

in general appearance are suggestive of the European lilac

(Smith 1966). Illustrations: Batten & Bokelmann: t. 99, 3

(1966); Palmer & Pitman: 1944 (1972); Shearing & Van

Heerden: 123 (1994).

20

Bothalia 31,1 (2001)

6b. subsp. silvatica Retief & A.E.van Wyk, subsp.

nov., subsp. rigidae affinis sed habitu effuso non pumilo,

lamina folii maiori tenuiori, inflorescentia non corym-

bosa, differt. Incolit sylvas, margines sylvae, non

dumeta.

TYPE. — South Africa, 3327 (Peddie): East London,

(-BB), Thode STEU6495 (PRE, holo.).

Scandent shrub or multi-stemmed tree up to 12 m

high. Leaf blade : broadly obovate or elliptic, (35-)

40 — 60(— 110) x (22-)30-45(-54) mm, apex obtuse or

acute; lower surface glabrous or with setae along veins,

upper surface glabrous or with setae on veins only, thin

in texture; petioles glabrous, with multicellular hairs or

setae, 5-20 mm long, length of petiole : length of midrib

= 1 : 6-11. Inflorescence terminal on a young shoot or

occasionally terminal on short young shoot at apex of

abbreviated branchlet, few-flowered, peduncles and

pedicels thin, up to 1.0 mm diam. Calyx ± 2.5 mm long.

Corolla lobes pale mauve or pale lilac, 4—6 mm long, as

long as tube. Fruit 4 mm long. Flowering time: July to

October.

Distribution: confined to Eastern Cape and southern

KwaZulu-Natal (Figure 12). Habitat: forest, forest scrub,

forest margin.

6c. subsp. nervifolia Retief & A.E.van Wyk, subsp.

nov., subsp. rigidae affinis sed inflorescentiis singulis

vel binis lateralibus in ramulis brevibus secus ramos vet-

eriores, vel terminalibus in surculis brevibus in apice

ramuli brevis, floribus paucis, pedunculis gracilibus

(inflorescentiis non in surculis juvenibus terminalibus,

singulis vel pluribus, corymbosis, pedunculis crassis), et

distributione differt. Incolit graminosos, nemores vel

savannam maritimam, non dumeta.

FIGURE 13. — Ehretia rigida (Thunb.)

Druce subsp. nennfolia, pub-

lished as E. hottentotica,

Pole Evans 1929. A, flower-

ing branch, x 0.8; B, single

flower, x 3.2; C, corolla from

6-merous (unusual) flower

laid open, x 2.5; D, pistil, x

4.9. Illustrations by Cythna

Letty.

Bothalia 31,1 (2001)

21

TYPE. — Mpumalanga, 2431 (Acomhoek), 8.8 km

from Skukuza on Lower Sabie road, (-DC), Codd & De

Winter 5031 (PRE, holo.; K, iso.).

Ehretia hottentotica sensu Pole Evans: t. 353 (1929).

E. rigida sensu J.H.Ross: 297 (1972); Venter & Venter: 172 (1996)

pro parte; sensu Fabian & Germish.: 338 (1997) pro parte; sensu Retief

& Herman: 352 (1997) pro parte; sensu Van Wyk & Van Wyk: 163

( 1 997 ) pro parte.

Multistemmed shrub, up to 5 m high. Branches entan-

gled, arching or drooping. Leaf blade obovate to broadly

elliptic, ( 1 1—) 1 6 — 35(— 50) x (4— )7-15(-25) mm, apex

acute, obtuse or emarginate, margin entire, usually with

setae along margin, upper surface glabrous, lower sur-

face with scattered setae along midrib and veins, with

coriaceous or thin texture; yellowish green, greyish

green to dark green; petiole 1. 5-5.0 mm long, length of

petiole : length of midrib = 1 : 3-7. Inflorescences one or

two lateral on apices of abbreviated branchlets along

older branches or terminal on very short young shoot or

terminal on a short young shoot at apex of abbreviated

branchlet, few-flowered; axes and calyx with multicellu-

lar hairs and/or setae; peduncles thin, only slightly elon-

gating in fruit. Calyx 5-merous, sometimes 6-merous;

lobes 1. 5-2.0 mm long. Corolla 5-merous, sometimes 6-

merous; tube white, 5-8 mm long; lobes mauve, 2-6 mm

long. Fruit ± 4 mm long. Flowering time : September to

December. Figure 13.

Distinguishing characters ; corolla lobes pale mauve;

branches arching prominently or conspicuously drooping

when plants are growing in a more arid environment; leaf

blade usually glabrous except for appressed setae along

margin or pubescent with setae along midrib and veins;

multicellular hairs present on inflorescence and calyx;

tertiary veining not prominent on lower surface but sec-

ondary veins so, usually three on either side of midrib,

raised. Distribution-. North-West, Northern Province,

Mpumalanga, Swaziland and KwaZulu-Natal (Figure

12); extends into Mozambique and Zimbabwe; sym-

patric with E. amoena and E. obtusifolia. Habitat : shrub

of savanna, dry thombush, coastal scrub. Common

names: puzzle bush; morobe (Tswana); deurmekaarbos

(Afrikaans) — according to Smith (1966) this name was

given to Ehretia known from Griqualand West to the

Waterberg. Utilization: tea made from dried leaves,

stalks and berries ( Sutton 785); fruit edible. Illustrations:

Pole Evans: t. 353 (192 9); Van Wyk & Malan: 253

(1997); Fabian & Germishuizen: 339 (1997).

SPECIMENS EXAMINED

(southern Africa only)

Specimens held at PRE, unless otherwise indicated.

The numbers in brackets signify the identity of the spec-

imens; (1) E. amoena ; (2) E. coerulea\ (3) E. obtusifolia ;

(4a) E. namibiensis subsp. namibiensis ; (4b) E. namibi-

ensis subsp. kaokoensis\ (5) E. alba-, (6a) E. rigida subsp.

rigida\ (6b) E. rigida subsp. silvatica\ (6c) E. rigida

subsp. nervifolia.

Abner 5 (2). Acocks 93 (5) K. PRE: 724 (5) BOL. PRE; 2211 (5) BOL.

K. PRE; 2471 (5) BOL, PRE; 11856 (6a). Adams 163 (5) NBG. An-

derson L2. L4, L7 (5); L14 (6c); 227 (5). Apuuli 27 (6a) GRA.

Archibald 3717 (6b) GRA, PRE; 5868 (6b).

Badenhorst 79 (5). Balarin 29 (6a) GRA. Balsinhas 3456 (6c) K, PRE;

3595 (6c). Barker 103, 108 (5); 741 (6c). Basson 127 (5). Barnard 145

(4a); 200 (3). Bayliss 482 (6b); 1326 (4b), Boshoff 1646 (5). Bouwer

2188 (5). Bradfield 103 (5). Braun 197 (6c). Bredenkamp & Van

Vuuren 183 (6c) PRE, UNIN; 345 (3) PRE, UNIN. Bredenkamp 126

(6c) PRE, PRU; 381 (3) PRU; 647, 833, 1121 (6c) PRE, PRU; 2015 (3)

PRE, PRU; 2073 (3) PRU. Brenan 14180 (1). Brink 374 (6a) GRA,

PRE. Britten 439 (6a); 1852 (6a) GRA; 1926 (6a) GRA, PRE; 2269

(6a). Brown PRE40569 (5). Brueckner 936 (5). Buerger 1058, 1142

(4a); 1313 (6c). Buitendag 187 ( 3) PRE, NBG; 799 (1) K, NBG, PRE;

863 (6c) NBG, PRE. Burgoyne 3332, 3483 (5). Burrows 2119 (6a);

2771. 3698, 4609 (6a) GRA.

Carr 4 (5). Catto 44, 147 (6c) PRU. Codd 661, 810, 2240 (6c); 2741

(3); 4976 ( 1); 5677(3) BM, PRE; 5659 (1) K, PRE. Codd & De Winter

3126 (1); 4976 (1) K. PRE; 5001, 5031 (6c) K, PRE. Codd & Dyer

4497 ay, Coetzee 1097 (6c); 256 (6c) PRE, PRU. Cole 339 (5); 990

(6c); 1075 (4a). Comins 1261, 1303 (6a) GRA, PRE. Compton 26649

(3) NBG, PRE; 28019 (6c) K, NBG, NH, PRE; 28208 (1) NBG, PRE;

28422 (6c) NH, NBG, PRE; 29354 (1) K, NH, PRE; 29914 (6c); 30316

(6c) NBG, NH, PRE; 30398 ( 6c) K, NBG, NH, PRE; 31022 (6c);

31136 (1) K, NBG, NH, PRE. Cooper 77, 84 (6b) NH. PRE. Crawley

PRE40559 (3). Crous 36 (6c) GRA. Culverwell 279 (3); 498 (6c); 1089

(6c); 1355 (1). Curson 265, 381 (6c). Curtis 203 (6a) GRA.

Dahlstrand 811 (6a) GRA. Dale 2143 (1). Daly 790 (6a) GRA. De

Winter 369, 404, 5918A, 5966 (6c). De Winter & Leistner 5435, 5823

(4b). De Winter & Wiss 4291 (2) K, PRE. WIND. Devenish 1465 (6c)

K, PRE. Dieterlen 867 (6c). Dinter 297 (4a) SAM. Dlamini PRE40557

(6c); PRE40583 (1). Dold 2394 (6a) GRA. Dold & Johnson 2394 (6a)

GRA. Downing 483 (6c) PRE, NU. Du Preez 191 (6c) BLFU. Du Toit

166 (6c); 826 (6c) K, PRE; 2800 (6c) NH.

Ecklon & Zeyher PRE12110, PRE21252 (6a). Edwards 910, 1267 (6c)

NU, PRE. Ellery 314 (6c). Engelbrecht 534, 579 (6c) NMB.

Fairall 2 (6c) NBG. Fanshaw & Bradley 109 (6a) GRA. Flanagan 605

(6b) BOL, GRA, NH; 778 (6a) PRE, SAM. Forest Dept. PE 135 (6a)

GRA. Fouche PRE40548 (6c). Fourie 127, 2251 (6c); 2848 (3).

Fourcade 3065, 5413 (6a) NBG. Francis 13 (6a) GRA. Fries &

Norlindh 629, 836 (6a).

Galala 119 (3). Galpin 7 (6a); 454, 499, 500 (3); PRE10690 (6a); 1242

( 1 ) K, PRE; 14783, 14788 (4a); 3322 (6a) GRA. Geldenhuys 887, 1226

(6a). Gemmell BLFU9030 (6b). Genis 95 (6c). Germishuizen 1490 (6a)

PRE, K; 1503, 1522 (6a); 2219 (3); 3366, 5093, 7750A (6c). Gerstner

736, 6085 (6c). Gibbs Russell 3343 (6a); 3656, 3726 (6a) PRE, GRA;

3841. 3902 (6a) BOL, GRA, PRE; 3974 (6a). Giess 9536 (5); 9581 (2).

Giess & Wiss 3317 (5). Giffen 1493 ( 6a). Gillett 504 (6c) K; 1068,

1068A (6c); 2452 (6a) NBG; 2600 (6c) NBG; 4593 (6a) GRA, PRE.

Gilliland 789 (1); 800 (3). Goldblatt 4940 (6a). Goossens 97 (6c); 667,

991, 1296 (5). Green 43, G51 (6c). Grobler 16 (5) PRE, WIND.

Hafstrom H^7 1 (5). Hall 45 (6a) GRA. Hall-Martin 1244 (1) PRU.

Haller 17 (6a) GRA. Hancock 49 (6c) NH, PRE. Hanekom 971 (5) K,

PRE; 175 A (4a) PRE, WIND; 176a (5) PRU; 1637, 1859 (6c) K, PRE.

Hahn 243(1) PRU. Hansen 3177, 3243 (5); 3476 (3) K, PRE. Hardy &

Scott-Smith 1526 (6c) K, PRE. Hemm 240 (6c). Henderson 104 (6c).

Henrici 1834, 2555 (5). Herman 759 (1); 1101 (5). Herman, Welman,

Pienaar & Crosby 683, 694 (6c). Heydom 31 (6c). Hildyard 192 (6c).

Hillary & Robertson 525 (5). Hilner 327 (6a) GRA. Hines 114, 133 (5)

PRE, WIND. Hitchins 61 (6c) NH, NU. PRE. Hobson 968, 1145 (6a)

GRA. Hoffman 40 (6c); 1019 (6a) GRA. Hofmeyr 50, 62 (6c). Holland

270 (6a) GRA. Holt 146 (3). Holzhammer 715 (5). Hoole L18 (4b)

GRA. Huntley 1208 (3); 1390, 1909 (6c). Hutchinson 1534 (6a) NBG;

7939 (6c) NBG.

Immelman 188 (3). Irvine 1.19 (6c). Ivy 2 (3).

TV. Jacobsen 102, 1539 (6c), 1934, 2623 (3). W.J. Jacobsen 1874, 2560,

4862, 4871 (6c), 4999 (3). Jacot Guillarmod 9657 (6a) GRA. James

256 ( 1) PRU. Joffe 316 (6c). Johnson 1057 (6b) GRA, PRE. Jooste 185

(5). Junod TRV24405.

Keet 1385 (6c); 1653 (5) PRE, WIND; 5988 (5). Kerfoot 7967 (4a) J,

PRE. Kerfoot & Falconer 32 (4a) J, PRE. Kinges 1357A (6c). Flap-

per 61 A, 124/AJ12 (6c). Kok & Pienaar 1303 (1) PRE, PRU. Kreulen

537 ( 5).

22

Bothalia 31,1 (2001)

Lang TRV30172 (1); 32107 (3); TRV32211 (3). Lawn 334 (3) NH; 600

(6c) NH. Le Roux 64 (6c) NMB; 556 (6c); 1030 (2) PRE, WIND, 1173

(4b) WIND. Leach & Bayliss 11528 (3). Leendertz 284 (6c) GRA; 506

(6c) PRU; 769, 801 (3); 905 (6c); PRE59261 (3); TRV5314 (6c);

TRV5550 (3); TRV9747 (6c). Leistner 894, 2061 (5) PRE. K. Letty 22

(6c); PRE8257 (6a). Liebenberg 849 (6a); 5087 (6a) K, PRE; 8634 (6)

K, PRE; 8740 (6c). Liengme 348 ( 1); 440 (1) K, PRE. Louw 387, 1023

(6c). Lubbe 1774 (6a) GRA. Lubbe, Everard & Avis 2735 (6b) GRA.

MacClean 974 (6c) NH, PRE. MacDonald 141 (6c) PRU; 455 (6c) E,

K, NU, PRE. MacOwan 93 (6a) GRA, K, PRE; 353 (6a) BM. SAM.

Maguire 575, 630 (6a) NBG; 1333 (6c) NBG; Makgakga 58 (3).

Mansvelt Marlothl3487 (6c). Markotter NHU 17842 (6c). Marloth 815

(5); 2074 (6c); 4274 (6a). Marsh 937 (6a). Mason 2606 (5). Matthee

1092 (6c) PRU. McFerren 129 (2). McMurtry 2018 (1); 2715 (6c); 2879

(1). Merxmuller 1071 (5) PRE, K. Miller B1136 (2). Mogg 15195A,

15195B, 15864, 16593, 19843, 20072 (6c); 21143 (6c) GRA; 23571,

31006, 37156 , 37230 (6c). Moll 753 (6c); 1079 (6c) NU. PRE; 1128 (6c)

K, NU, PRE; 1805 (6c): 2169 (6c) NU, PRE; 2204 (3) NU, PRE; 4123

(6c); 4143 (6c) K, NH. PRE; 4244 (6c) NH, PRE; 4258 (6c) E, K, NH,

PRE; 4390 (1). Moll & Morris 625 (6b) NU, PRE. Moran PRE40563

(5). Morris 800 (6b); 596 (6c) NH, PRE; 1025 (6c) K, NU, PRE; 1197

(5a). Morris & Engelbrecht 30, 1118 (5). Moss & Rogers 273 (1) NBG.

Mott 356, 769 (6c). Muller 897, 1073 (6c) NMB; 1637 (5) NMB;

PRE40558 (5). Muller 997 (4a). Murdoch 53 (1).

Nel 69 ( 1 ) K, NBG, PRE; 119 (6c); 238 ( 1 ) NBG. PRE; 264 ( 1 ) K, PRE.

Netshiungani 1624 (1). Ngwenya 36 (3) NH; 379 (6c). Nicholson 705,

724 (6b); 901, 1330 (6c); 1459, 1552 (6b); 1566 (3); 1617 (6b). Nicol

60 (6a) NBG. Noel 629 (6a) GRA. Norrgrann 241 (3).

Oates 61, 65 (3); 95 (1); 105 (3); 308 (5); 357 (1). Obermever 750 (6c);

828 (1) K, PRE; 1246 (1); TRV29257 (1); TRV33565 (6c). Obermever,

Schweickerdt & Verdoorn 44, 45 (6c). O'Callaghan 1012 (6b) GRA.

Olivier 1208 (6a); 2336 (6a) PRU; 2821 (6a). Onderstall 1077 (6c).

Osborne 151 (6a) GRA.

Palmer 1167 (6a). Paterson 2026 (6a). Pauw 127 (6c) PRU. Pearson

4399 (5); 8260 (5) BM, PRE. Peeters, Gericke & Burelli 11 (5). Pegler

110 (6b). Pentz 324 (6c). Penzhorn 6442 (6a) GRA. Peyper 1287 (5)

NMB; 2043 (6c) NMB. Pahl 3.40 (6a) GRA. Phelan 703 (6c). Phillips

42 (5). Phillipson 365 (6a) K, PRE; 3193 (6a) GRA, PRE. Pienaar 643,

673 (6c). Pole Evans 1480, PRE40545 (6c); PRE40592 (1). Pott 171

(6a) GRA. Potts 3216 (5). Prinsloo 45 (3). A. Prior PRE40566 (6a). J.

Prior 397 (1).

Raal 1383 (3). Ramovha 23 (6c) PRU. Reed 8 (6a) GRA. Renny 97 (1).

Repton 4 (6c). E.Retief 619 (6c). I. M. Retie f 278, 547 (1). Retief &

Strauss 2167 (3). Reyneke 410 (5). Roberts 1843A, 5365 (6c). Rodin

4203 (6c). Rogers TRV2603 (1) BOL, PRE; 12891 (1) K, PRE; 17134

(6a) K; 18220 (1) BM, PRE; 21832 (6c). Rose lnnes 9, 189 (6c). Ross

2009, 2014 (6c) NH, PRE. Roux 94 (5). Rowland 169 (3).

SAGP/SAAB 1/ 34 (6c) PRE, PRU; 2/44 (6c); 2/49 (6c) PRU; 3/93,

4/75 (6c) PRE, PRU. Salisbury 61 (6a); 138 (6a) GRA. Scharf 1481

(6a); 1681 (6a) NBG, PRE; 1159 (6a). Scheepers 1735 (5). Schlechter

3294 (6c) BM, BOL, GRA, NH, PRE. Schrire 1169, 1504 (3) NH.

Schmidt 84 (6b) GRA. Schweickerdt 1014 (6c). Seagrief 149.6 (6a)

GRA. Seaman 67, 97 (6c) NMB. Seydel 239, 1233 (5); 1239 (5) COI,

PRE. Shackleton 566 ( 1) PRU. Sidey 2393 (6c); 3134 (6a); 3723 (6c).

Smit 219, 1256 (1 ) PRU. Smith 254, 398 (1); 693 (6c); 739, 1090, 2140

(5); 2149 (1) K, PRE; 2592 (5) 5195, 3245, 7017 (6c). Smuts & Gillen

2104, 2126, 2136, 3447 (3). Snyman & Noailles 233 (6c). Spies 11(1).

Spreeth 6 (6c) PRU. Stayner 64 (6a) GRA, PRE. Stephan 823 (6c).

Stewart 88 (6c) NH, PRE. Steynberg 986 (3). Stirton 8854 (6c) K, PRE.

Story 4592 (5); 4942 (5) PRE, K; 5504 (4a). Straub 122 (3). Strey 1136

(6c); 2505 (6a); 4853 (3) NH, PRE; 5874 (6c); 9021 (6c) NH, NU,

PRE; 9969 (6c) NH, PRE. Strey & Moll 3734 ( 1) NH, PRE. Sutton 785

(6c). Swartz 33 (6a). Swierstra TRV4383 (3).

Tainton 2 (6c). Tanaka 21 (5). Taylor 946, 1286A (6a). Taylor &

Edwards 8861 (6b) GRA. G.C. Theron 236, 447 (6a); 1455 (6c). J.J.

Theron 2048 (3) J, PRE. Thode A598 (6a); 2713A (6b) NH, PRE; 6495

(4b). Thompson 918, 1884 (6a) NBG, PRE. Thorncroft 3018 (1).

Thome SAM35767 (4a) SAM. Tinley 332 (1) K, NU, PRE; 494 (6c)

NH, NU, PRE. Tinley & Ward 30 (6c). Tolken & Hardy 716 (5).

Troughton 108 (6a) GRA. PRE. Turner 20 (6c). Tyson 2201 (6a) GRA,

SAM; 2662 (6a) SAM; Marloth Herb 8502 (6a).

Vahrmeijer 376A (6c); 1500 ( 1 ) K. PRE. Van Dam TRV18826 (6a). Van

den Berg 3 (1). Van der Schijff 618 (3); 782 (1) KNP, PRE; 904 (6c) K,

KNP, PRE; 915 (3); 1213 ( 1 ) KNP, PRE; 2273 (6c) K, PRE; 3306 (6c);

3406, 3917, 4075 (6c) K, PRE. Van der Schijff & Marais 3639 (1) KNP,

PRE. Van der Spuy 36 (5). Van Graan & Hardy 494 (6c). Van Greuning

525 (1) PRE, PRU; 500, 580 (1) PRU. Van Heerden 743 (1). Van

Rooyen & Bredenkamp 4, 99 (5) PRU. Van Rooven & Schultz 4318 (6c)

PRU. Van Rooyen 1012 (1) PRU; 1053, 1144A, 1266 (1) PRU; 1948,

1989 (6c) PRE, PRU; 4421( 6c) PRU. Van Vuuren 549, 1583 (6c). A.E.

van Wyk 392 (6a) NBG, PRE; 460 (6c); 2615 (1) PRU; 10462 (1) PRU;

3029 (6c) PRE, PRU; 3547 (6c) PRU; 4019, 5156, 5918, 5954 (1) PRE,

PRU. P. van Wyk BSA352 (1 ) PRE, PRU; BSA1552, BSA3110 ( 1 ) PRU.

Van Wyk & Theron 4719 (3) PRE, PRU. Venter 1945 (3) BLFU, PRE;

1108, 2813 (6c); 5148, 7216,8583 (1); 12173, 13052 (1) UNIN; 13056

(6c) NU; 13598 (6c) UNIN. Venter, Hahn & Archer 41 (5); 93 (2).

Verdoorn 50, 603 (6c); 1121 (5) BOL, NMB, PRE; 2110. 2353 (3);

2354 (6c) K, PRE; 2355, 2357 (3); 2358 (6c). Viljoen 88 (6c). Von

Gadow 451 (6a) GRA, PRE.

Wager PRE59522 (3); 77? V 22463 (1); TRV236 72 (3). Walter 29 (6a)

GRA. Wanntorp 93 (5). Ward 3 (2); 1206 (1) NH, PRE; 2980 (1); 3751

(6b) NU, PRE; 7223 (6c) NH, PRE. Wearne 50 (6c) NU, PRE. Webber

7742A (6a) GRA. Wentzel N15 (1). Werdermann & Oberdieck 1057

(6a); 1948 (6c) K, PRE; 2286 (5). Werger 1465 (6c). West 399, 489

(6c); 1549 (6c) K, NH, PRE. Westphal TRV29118 (1). White 2 (6a)

GRA. Wild 5082 (5). Williamson 81 (6c). Wilman 14 (5). Winter 2417

(3) UNIN; 2695 (6c) UNIN; 2892 (3) UNIN; 2966 (6c) UNIN; 3082 (3)

UNIN. Wood 10919, 11500 (3) NH. Woollard & Bethea 2168 (4a).

Young 504A (6c); 584A (3); 2516 (6c).

Zambatis 264, 713, 722 (3); 748 (1); 1204 (3), 1224, 1233 (3). Zeyher

161 (6a) BOL, GRA, K, NBG, PRE. Zietsman 290, 981 (6c) NMB;

1341 (6c) NMB, PRE; 1373 (5) PRE, NMB; 3445 (5) NMB. Zwanziger

164 (6c).

ACKNOWLEDGEMENTS

The assistance of Drs H. Glen, O.A. Leistner, S. Perold

and J. van Rooy, Mmes G. Condy, N. Netnou, E. du

Plessis, A. Romanowski, J. Ready and H. Snyman of the

National Botanical Institute in preparing the manuscript,

is gratefully acknowledged. A word of special apprecia-

tion to Patricia Craven, Norbert Hahn, Mervyn Lotter,

Coleen Mannheimer and Pieter Winter for their friendly

assistance and expert advice. The directors, curators and

staff of the cited herbaria are thanked for allowing the

authors to study their material and for sending specimens

on loan.

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Bothalia 31,1: 25-29 (2001)

Studies in the liverwort genus Fossombronia (Metzgeriales) from

southern Africa. 11. F. zuurbergensis , a new species from Eastern

Cape and new records for the area

S.M. PEROLD*

Keywords: Eastern Cape. Fossombronia Raddi, F. zuurbergensis, Hepaticae, Metzgeriales, new records, new species, southern Africa

ABSTRACT

Fossombronia zuurbergensis Perold, sp. nov. from Eastern Cape is described. It is distinguished by having leaves and

pseudoperianths with markedly dentate margins; the spore ornamentation on the distal face consists of ridges which can be

quite variable, although, at least some of them run parallel to each other in straight or curved, short or long ridges, others

occurring at right angles to the former. New records for F. capensis var. capensis and F. crispa are reported for Eastern Cape.

Fossombronia zuurbergensis Perold , sp. nov.

Plantae aggregatae, virides, dioicae; masculae parvae

vel mediocres, feminae maiores. Folia patentia, margin-

ibus valde dentatis, saepe in plantis masculis minoribus

supra subbiloba. Rhizoidea purpurea. Antheridia serie

uno dorsaliter secus caulem disposita, a bracteis perigo-

nialibus protecta, plerumque lateraliter juncta et cum 3-8

processis spinosis. Pseudoperianthium campanulatum,

folio proxime juxta idem paulo brevius, breviter stipita-

tum, supra late expansum, margine in ore valde convolu-

to dentatoque, cum usque ad 33 processibus spinosis,

basin versus cum appendicibus lateralibus versiformibus.

Sporae 37.5^15.0 pm diametro, superficie distali cum

7-9 cristis transversis brevibus vel longis, ± 5 pm dis-

tantibus, 2.5 pm altis, paucis rectis, aliis curvatis, inter se

parallelis, gregibus nonnullis ad illas perpendicularibus;

superficie proximali cum processibus irregularibus coni-

cis vel spinosis, papillisque minoribus tecta. Elateres

75-125 pm longi, 5.0-12.5 pm lati, tenue papillosi, alii

2- vel 3-spirales, alii medio 3-spirales, apicibus bispi-

rales.

TYPE. — Eastern Cape, 3325 (Port Elizabeth): along

Zuurberg Pass, (-BC), at altitude 670 m, John J. Perold

19 (PRE. holo.).

Plants in crowded stands, often growing together with

Riccardia species, in persistently damp areas, green;

shoots prostrate, smallish to medium-sized in male

plants, up to 9 mm long, ± 1.5 mm high, ± 3 mm wide;

in female plants larger, 10-12 mm long, ± 2 mm high, 3

mm wide, at pseudoperianth up to 5 mm wide, simple or

once apically furcate, segments moderately divergent, ±

2 mm long. Stems tapering distally, chlorophyllose,

plano-convex in cross section, in male plants apically

(Figure IN) up to 170 pm (9 or 10 cell rows) high. 220

pm wide, basally (Figure 10) ± 180 pm high, 260 pm

wide, in female plants apically (Figure IP) 200-210 pm

(9 cell rows) high, 380 pm wide, basally (Figure IQ) 270

pm high, 350 pm wide. Rhizoids purple, 12.5-22.5 pm

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

MS. received: 2000-10-02.

wide. Leaves spreading, not ruched above, margins

markedly dentate, overlapping basally by ± V3 their

width, sometimes more, obliquely and succubously

inserted, often somewhat bilobed above in smaller male

plants (Figure 1A-E), 1250-1625 x 1200-1575 pm, in

female plants (Figure 1F-J), margins with 10-15 or 16

spinous processes, 2 cells long and 1 cell wide, topped by

apical papilla (Figure IF), sometimes with basally wider,

triangular processes, up to 5 cells and a papilla long,

leaves irregular in shape, often as in a partly opened fan,

1425-2000 pm long, 1000-2500 pm wide above,

1000-1325 pm wide below, a larger leaf lateral to

pseudoperianth (Figure IK), up to 2375 x 2875 pm wide

above, ± 1400 pm wide below, marginally with as many

as 24 processes. Leaf cells mostly thin-walled, no appre-

ciable differences between those of male and female

plants, at upper margin rectangular across or 4- or 5-

sided, 25.0-37.5 x 50.0-57.5 pm; upper laminal cells 5-

or 6-sided, 57.5-85.0 x 45-55 pm; middle laminal cells

75.0-127.5 x 50.0-62.5 pm; basal cells 90-125 x 60-70

pm. Oil bodies (Figure 1M) 9-24 per cell, round or oval,

shiny, smooth, 2. 5-5.0 pm diam.; chloroplasts numer-

ous, ± 5 pm diam.

Dioicous. Antheridia in a row dorsally along stem

(Figure 2A), short-stalked, globose, 220-250 pm diam.,

shielded by perigonial bracts (Figure 1R-V), mostly lat-

erally joined, 400-590 x 320-440 pm, with 3-8 spinous

processes, each with an apical papilla, marginal cells 4-

sided, 50-70 x 25.0-37.5 pm, internal cells long-rectan-

gular, 50-85 x 30.0-37.5 pm. Archegonia in a single row

or in pairs, opposite or subopposite, dorsally along stem,

140-200 pm long, one recently fertilized, (Figure 2E).

Pseudoperianth shortly proximal to apex of stem (Figure

2B) or nearer to base (Figure 2C), rarely 2 per shoot

(Figure 2D), campanulate (Figure IX), slightly shorter

than large leaf immediately lateral to it, raised on a short

stalk, 700-875 pm wide at base, then widely flaring

above (Figure 2F), 2000-2250 pm long, up to 2750 pm

wide across mouth, margin highly convolute and dentate,

with up to 33 spinous processes (Figure 1W), 87.5-500.0

pm or 3-5 cells and an apical papilla long, generally

raised on a gradually widening base, cells comparable in

shape and size to those of leaves; toward base with sev-

26

Bothalia 31,1 (2001)

FIGURE 1. — Fossombronia zuurbergertsis. A-E, male leaves; F-J, female leaves; K, large leaf next to pseudoperianth; L, detail of female leaf

margin; M, median leaf cells with oil bodies (solid lines) and chloroplasts (dotted lines); N, c/s male stem apex; O, c/s male stem base; P,

c/s female stem apex; Q, c/s female stem base; R-V, perigonial bracts; W, opened pseudoperianth; X, pseudoperianth from side; Y, c/s seta;

Z, cells in inner capsule wall. A-E, H, J-M, P, Q, W-Z, John PerolcI 19\ F, G, I, N, O, R-V, John Perold 24. Scale bars: A-K, W, X, 500

pm; L, Y, 100 pm; M, Z, 50 pm; N-V, 250 pm.

Bothalia 31,1 (2001)

27

FIGURE 2. — Fossombronia zuurbergensis. A, simple male plant with perigonial bracts along stem; B, simple female plant with pseudoperianth

near apex of shoot; C, apically furcate female plant with pseudoperianth and capsule more proximally; D, simple female plant with 2

pseudoperianths in acropetal sequence; E, female plant with fertilized archegonium and vestigial pseudoperianth (indicated by arrow); F,

pseudoperianth partly from above, appendages near its base. A, John Perold 24: B-F, John Perold 19. A, x 24.3; B, X 13.3; C, x 9; D, x

8.7; E, x 20; F, x 16.5.

eral, variously shaped, narrow or wider, appendages,

1000-1200 pm long (Figure 1WU W2). Capsules glo-

bose, 260-300 pm diam., wall bistratose, cells of inner

layer triangular, quadrate or rectangular, 37.5-55.0 x

27.5-50.0 pm, each cell wall with 1-3 dark brown,

nodular thickenings, rarely with semi-annular thicken-

ings (Figure 1Z). Seta up to 3.25 mm long, ± 250 pm

diam., 9 cells across (Figure 1Y). Spores brown,

37.5-45.0 pm diam., including ridges projecting at mar-

gin, hemispherical; distal face (Figure 3A-D) convex,

with 7-9 short or long ridges across, ± 5 pm apart and

2.5 pm high, a few straight, others curved, parallel to

FIGURE 3. — Fossombronia zuurbergensis. A-E, spores; F, elater. A-C, distal face; D, side view of distal face; E, proximal face. A, D-F, John

Perold 19: B, John Perold 5; C, John Perold 113 (Baviaanskloof). A, x 990; B, x 1 138; C, x 1064; D, x 1097; E, x 967; F, x 2707.

28

Bothalia 31,1 (2001)

FIGURE 4. — Distribution of Fossombronia zuurbergensis, •. New localities of F. capensis var. capensis, CD; and F. crispa. A, in Eastern Cape.

each other, but some groups running at right angles to the

former, occasionally interlinked to form complete or

incomplete alveoli, truncate or spinous processes some-

times interspersed in between; proximal face (Figure 3E)

lacking triradiate mark, flat to slightly concave, covered

with irregular conical or spinous processes and smaller

papillae, around spore periphery 21-25 projecting

‘spines’, i.e. the ends of ridges which extend over the

sides of the distal face, some joined by an incomplete

membranous wing. Elaters yellow, 75-125 x 5.0-12.5

pm, tapering to tips and ending in loop, or not, finely

papillose (Figure 3F), bispiral or trispiral, others trispiral

in centre, tips bispiral, very rarely once furcate.

Fossombronia zuurbergensis has been named for the

Zuurberg in the Eastern Cape (Figure 4), where it was

collected at several localities along the Pass, some only in

small stands, others in larger clumps. A few collections

had to be kept in cultivation to allow the spores to ripen.

The vegetation type of vascular plants in this area is

Mountain Fynbos (Low & Rebelo 1996). The soil is

derived from quartzitic sandstone shale of the Witpoort/

Weltevrede formation of the Witteberg group of the Cape

Supergroup (Visser 1984). It was recently also collected

further to the west at Baviaanskloof (3324DA) along the

banks of the Wit River, at altitude ± 900 m. These areas

have fairly sparse rain, both in winter and in summer.

On account of the markedly dentate margins of the

leaves and pseudoperianths, it was originally thought

that these plants, although noticeably larger, belonged to

F. spinifolia Steph., which had only been collected twice

by Breutel in 1853 at Genadendal in the Western Cape

(Perold 1997a). However, on comparing the ornamenta-

tion on the distal face of F. zuurbergensis spores with

those of F. spinifolia, the former has 7-9, longer and

sometimes parallel, as well as smoother ridges running

across it. In F. spinifolia spores the 6 or 7 ridges on the

distal face, besides being shorter, are also much less reg-

ular and not smooth; occasionally with a few large papil-

lae interspersed between them. F. densilamellata

S.W.Amell (1952) also has spores with parallel ridges on

the distal spore face, but they are closer together and

more numerous, i.e. 12-16 and are sometimes connected

with slender threads between them (Perold 1997b).

F. zuurbergensis can be distinguished by the many

spinose processes at the margins of the leaves and

pseudoperianths, by being dioicous with smaller male

plants, by mostly composite perigonial bracts and by the

spore ornamentation with some smooth, parallel ridges

on the distal face.

NEW RECORDS OF F. CAPENSIS VAR. CAPENSIS

FROM EASTERN CAPE

F. capensis var. capensis (Perold 1 999) was collected

for the first time in Eastern Cape at several localities

along the Zuurberg Pass and at Baviaanskloof. Sporulat-

ing specimens are easy to recognize because of the poor-

ly formed elaters.

NEW RECORDS OF F. CRISPA FROM EASTERN CAPE

F. crispa is widespread in southern Africa (Perold

1997c), but has not previously been reported from

Eastern Cape. I collected the specimens in October 1999,

at Nerina Bush Camp along the stream bank, on soil in

the Zuurberg National Park. The spores have the usual

incompletely reticulate ornamentation on the distal face.

The plants, however, are clearly dioicous. In Perold

( 1997c), I, stated that F. crispa is ‘monoicous, sometimes

seemingly dioicous’.

SPECIMENS EXAMINED

Held at PRE. Bracketed numbers after collectors' name

and number refer to the species in the text, namely: F.

zuurbergensis (1), F. capensis var. capensis (2) and F.

crispa (3).

Bothalia 31,1 (2001)

29

John Perold 1, 2, 6, 8-10 (2); 11 (1); 13, 16 (2); 19 (1) (holotype); 28,

35, 37 ( 2); 38 (1); JSM7 (2): 42 (1); 46, 75 (2); 79 (1); 80 ( 2); 102, 113.

145 (1) (the last two collections are from Baviaanskloof (3324DA), the

others from the Zuurberg).

S.M. Perold 4147, 4148 (3).

ACKNOWLEDGEMENTS

I sincerely thank my nephew, John Perold, and his

wife, Audrey, for their hospitality during my visit to

them at Addo, in October 1999 and for John’s help,

interest and enthusiasm in collecting liverworts in

Eastern Cape and for delivering them to me. For his

helpful suggestions Dr H. Stieperaere is thanked and

also Dr H.F. Glen for translating the diagnosis into

Latin. Furthermore, I extend my gratitude to Ms G.

Condy for the drawings, to Mrs A. Romanowski for

developing and printing the photographs and to Ms D.

Maree for typing the manuscript.

REFERENCES

ARNELL, S.W. 1952. Hepaticae collected in South and West Africa

(1951). New and little known species. Botaniska Notiser 1952:

307-329.

LOW, A.B. & REBELO, A.G. (eds). 1996. Vegetation of South Africa,

Lesotho and Swaziland. Department of Environmental Affairs

& Tourism, Pretoria.

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 3. An amendment to F.

spinifolia. Bothalia 27: 39—42.

PEROLD. S.M. 1997b. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 2. An amendment to three

species from Western Cape, described by S.W. Amell. Bothalia

27: 29-38.

PEROLD, S.M. 1997c. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 4. A re-examination of F.

crispa, F. leucoxantha and F. tumida. Bothalia 27: 105-115.

PEROLD, S.M. 1999. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 7. F. capensis var. spiralis,

a new variety from Western Cape. Bothalia 29: 1—4.

V1SSER, D.J.L. Compiler. 1984. Geological map of the Republics of

South Africa, Transkei, Bophuthatswana, Venda and Ciskei and

the Kingdoms of Lesotho and Swaziland. Government Printer.

.

Bothalia 31,1: 31-52 (2001)

Notes on African plants

VARIOUS AUTHORS

ASPHODELACEAE

ALOE GREATHEADII VAR. DAVYANA (ALOOIDEAE), A NEW LOCALITY IN KWAZULU-NATAL, SOUTH ALRICA

The grass aloes (A. sect. Leptoaloe A. Berger) and

maculate aloes (A. sect. Pictae Salm-Dyck) are notorious

for the intergradation of their constituent species, com-

plicating accurate field and herbarium identification. In

southern Africa, Glen & Hardy (1987, 2000) have sub-

stantially reduced the number of species recognized,

while Carter (1994) sank the eastern African A. gramini-

cola Reynolds into A. lateritia Engl, as var. graminicola

(Reynolds) S. Carter.

The second author recently came across a small pop-

ulation of a maculate aloe in the Pietermaritzburg

District in south-central KwaZulu-Natal which, at first

sight, did not fit in comfortably with the description of

any of the known species in Aloe sect. Pictae. Specimens

from this population were characterized by the distinctly

conical shape of the young and mature inflorescences,

the prominence of the floral bracts and the comparative

robustness of the plants. However, following careful

consideration of the combination of these and other char-

acters, we regard the specimens as belonging to the

northern, predominantly grassland and savanna taxon, A.

greatheadii Schonland var. davvana (Schonland) Glen &

D.S. Hardy, and therefore, this extends its distribution

range in a southeasterly direction by 120 km (Figure 1).

Aloe greatheadii var. davy’ana is exceptionally com-

mon throughout its distribution range and is not threat-

ened in any way. However, some populations of this vari-

ety, such as the one south of Pietermaritzburg reported

here, may become locally extinct as a result of urban

expansion.

For a description of the taxon see Glen & D.S. Hardy

(1987). For a complete synonymy see Glen & Hardy

(2000).

KWAZULU-NATAL. — 2930 (Pietermaritzburg): valley bushveld

adjacent to Ashburton Township, (-CB), N.R. Crouch 821 (NH).

LIGURE 1. — Distribution map of Aloe greatheadii var. davvana, based

on Van Wyk & Smith (1996) and specimens kept in PRE and

NH. New locality in KwaZulu-Natal, O.

RELERENCES

CARTER, S. 1994. Aloaceae. In R.M. Polhill, Flora of tropical East

Africa'. 1-60.

GLEN, H.F. & HARDY, D.S. 1987. Nomenclatural notes on three

southern African representatives of the genus Aloe. South

African Journal of Botany 53: 489-492.

GLEN, H.F. & HARDY, D.S. 2000. Aloaceae (First part): Aloe. Flora

of southern Africa 5, 1,1: 1-167.

VAN WYK. B-E. & SMITH, G.F. 1996. Guide to the aloes of South

Africa. Briza Publications, Pretoria.

G.F. SMITH* and N.R. CROUCH**

* National Botanical Institute. Private Bag X 1 0 1 . 0001 Pretoria.

** Ethnobotany Unit, National Botanical Institute, P.O. Box 52099,

4007 Berea Road, Durban.

MS. received: 2000-06-30.

AMARYLLIDACEAE

A NEW SPECIES OF CYRTANTHUS (CYRTANTHEAE) FROM THE SOUTHERN CAPE. SOUTH AFRICA

INTRODUCTION

Cyrtanthus Aiton, a genus of about 55 species, is

found from East Africa to Western Cape, South Africa.

Currently 21 Cyrtanthus species are known from the

Cape Floristic Region, five of which have been described

in the last 25 years (Dyer 1977, 1980; Snijman & Van

Jaarsveld 1995; Snijman 1999). The vegetation of the

Cape Region, like all the other mediterranean-type

shrublands of the world, is fire prone, and fire-stimulated

flowering is common to many of its plants. This trait is

particularly well developed in Cyrtanthus, not only in

species from the Cape, but also in several from the grass-

lands to the north.

32

Bothalia 31,1 (2001)

Extensive fires swept across the western and southern

Cape mountains at the end of 1999 and the effects engen-

dered considerable debate on the role of fire in maintain-

ing fynbos biodiversity (Saunders & Saunders 2000).

The fires, however, provided a rare opportunity to study

populations of a pink-flowered Cyrtanthus, which Mr

J.H. Vlok had reported on the Outeniqua Mountains in

the southern Cape. Field studies of these delicate plants,

first collected by Miss D.M. Gemmell in 1936, con-

firmed that they belong to yet another new species of

Cyrtanthus , formally described here as C. debilis.

Cyrtanthus debilis Snijman , sp. nov., foliis hyster-

anthis. floribus roseis tubaeformibus, staminibus fascicu-

latis declinatis, filamentis 15-30 mm longis, et stylo tri-

furcato, a speciebus nobis notis bene distincta. Figura 2.

FIGURE 2. — Cyrtanthus debilis : A, bulb with young leaf and inflores-

cence; B, bulb and lateral view of inflorescence; C, flower with

two lateral tepals and stamens removed; D, apex of style and

stigma; E, infructescence; F, seed; G, mature leaves; H, trans-

verse section through leaf blade. Drawn from Snijman & Vlok

1717 by Claire Linder Smith. Scale bars: A, B. E, G, 10 mm; C,

F, 5 mm; D, 1 mm; H, 0.5 mm.

TYPE. — Western Cape, 3322 (Oudtshoom): Oute-

niqua Mountains, near summit of Robinson Pass, (-CC),

21-11-1999, Snijman & Vlok 1717 (NBG, holo.; K, PRE).

Deciduous bulbous herb, 8-22 mm tall when flower-

ing. Bulbs solitary, hypogeal, ovate to ellipsoidal, up to

20^30 mm long, 10-25 mm diam., with a slender neck up

to 20 mm long; outer tunics brown and papery; inner

tunics cream-coloured and fleshy. Leaves (2)3-5, linear,

hysteranthous, sometimes 1 emerging at flowering,

7-180 x 1-2 mm, suberect to spreading, glabrous, three-

angled in cross section with 1 median vein, dark green but

reddish proximally. Inflorescence l-2(-4)-flowered;

scape erect, 35-140 x 2-4 mm, tapering distally, brown-

ish green without a grey bloom, hollow; spathe valves 2,

equitant, narrowly lanceolate, erect, 25-35 x 3-5 mm,

maroon-pink, margins inrolled; bracteoles up to 4, fili-

form, up to 23 mm long; pedicels erect, 7-20 x 1 mm,

brownish green. Flowers slightly spreading, 35-70 mm

long, trumpet-shaped, rose-pink with darker pink on

perigone tube, tepal sinuses in throat and median bands

on tepals, unscented; tube slightly curved in proximal

half, 1 8 — 48 mm long, 2 mm diam. at base, widening grad-

ually and evenly to 8-15 mm at throat; tepals oblong-

lanceolate, 18-30 x 6-10 mm, spreading regularly, usual-

ly 5-veined; outer whorl prominently mucronate, as wide

or slightly wider than the inner; margins sometimes

slightly undulate. Stamens biseriate, tightly clustered

together against lower tepals; filaments 15-30 mm long,

inserted ± midway up tube, upturned apically, white,

outer whorl inserted 2-3 mm below the inner, inner whorl

extended beyond the outer by up to 10 mm; anthers dor-

sifixed, ± 2 mm long, yellow. Ovary ellipsoidal, 5-8 x

2-4 mm, greenish; ovules axile, ± 16 per locule. Style

clustered with stamens, extended beyond inner stamens

by ± 5 mm; upturned apically, white; stigma 3-branched,

up to 1.5 mm long, penicillate apically. Capsule narrow-

ly ellipsoidal, 10-20 x 5-9 mm, 3-valved. Seeds black,

flattened, irregularly angled, 5-6 x 3^1 mm.

Phenology

The flowering time of the particular species of

Cyrtanthus , referred to as fire lilies, depends on the advent

of fire, then rain. In the case of C. debilis the bulbs may

flower any time between November and April. Flowering

in a population is brief and the entire period lasts ± 10

days. The leaves of C. debilis emerge in May and begin to

die back as the dry season approaches in summer.

Thereafter the bulbs enter a prolonged vegetative phase

until fire sweeps through the area again. In the Outeniqua

Mountains this happens once every 12 to 18 years (J.H.

Vlok pers. comm.). A photograph by Mr W. Werner of

flowering pot plants of C. debilis in Reid & Dyer (1984:

fig. 9, named C. clavatus) suggests that the bulbs respond

to cultivation. However, the conditions under which these

plants flowered in containers is not known.

Diagnostic features

Cyrtanthus debilis is easily distinguished from other

Cyrtanthus species by its pink, trumpet-shaped flowers

and the characteristic length and position of the stamens.

In particular, the filaments are 15-30 mm long, well

Bothalia 31,1 (2001)

33

FIGURE 3. — Known distribution of Cyrtanthus debilis, ©; C. ventricosus , •; and C. clavatus , ▲.

exserted from the throat and tightly clustered together

against the lower tepals. Although the species is distinc-

tive, its affinities with other members of the genus are

not so readily apparent.

Reid & Dyer (1984) regarded the specimens of C.

debilis , known to them from the National Herbarium at

Pretoria, as a darker salmon-pink-flowered form of C.

clavatus , a species of open grassland found east of Port

Elizabeth in the Eastern Cape (Figure 3). They, never-

theless, recommended further field studies. Herbarium

specimens show the flared perigone, the stamens and

style that arch downwards to the lower tepals, and the

three-branched stigma that are common to C. clavatus

and the four other species recognized by Reid & Dyer

(1984) in this group, namely C. loddigesianus (Herb.)

R. A. Dyer, C. speciosus R. A. Dyer, C. helictus Lehm. and

C. smithiae Watt, ex Harv. The pressings, however,

obscure several features evident in the living plants, par-

ticularly flower colour and stamen position. The flowers

of C. debilis are pale pink throughout with a darker red-

dish pink tube and median bands on the tepals, whereas

those of C. clavatus are white to cream, usually with pink

median bands on the tepals. Furthermore, the stamens of

C. clavatus are short (± 7 mm), more or less included in

the perigone tube and, although somewhat declinate, are

set apart from each other. The two species also differ

with respect to vegetative phenology. Cyrtanthus debilis

has hysteranthous leaves and the bulbs flower only after

fire, whereas in C. clavatus the bulbs flower freely after

good summer rains, usually with their leaves present.

In Reid & Dyer’s (1984) key to the southern African

species of Cyrtanthus, C. debilis — given as C. clavatus

(colour form) — was grouped with species in which the

perigone tube is narrowest at the base, but flares widely

to the throat, so that the upper portion is campanulate.

Species with this floral form and characterized by linear

leaves and pink flowers are C. galpinii Baker and C.

thomcroftii C.H. Wright from the northern provinces of

South Africa, and C. sanguineus (Lindl.) Walp. subsp.

salmonoides (RR.O. Bally & Carter) Nordal and C. san-

guineus subsp. minor Nordal from East Africa (Nordal

1979). Unlike these northern species, the perigone tube

in C. debilis flares open smoothly from base to throat,

without an abrupt change in shape between the lower and

upper halves. Moreover, these northern species display

subtle differences in floral markings. The tepals are uni-

formly coloured, the perigone tube is usually green in the

lower part and darker stripes, if present, lead down into

the throat from between the tepals, rather than from the

backs of the tepals as in C. debilis.

Cyrtanthus debilis also shows a likeness to C. ventri-

cosus (Jacq.) Willd., a hysteranthous-leaved species known

from the western Cape and the Baviaanskloof Mountains

in the southeastern Cape, but not yet recorded on the

intermediate mountain ranges of the southern Cape

(Figure 3). The flowers in both species are fire dependent

and they have well-exserted, unusually long stamens

(25^-5 mm in C. ventricosus ) that arise in the lower half

of the tube. Cyrtanthus ventricosus is distinguished from

C. debilis , however, by its stamens which arch against

the upper tepals and by an undivided stigma. Typically

the somewhat flared flowers of C. ventricosus are red,

but populations on the Cape Peninsula have pale salmon

flowers with a darker tube and median bands on the

tepals, very like the markings of C. debilis.

As yet there is no phylogeny for the genus, so the sig-

nificance of floral features for grouping C. debilis , C.

ventricosus and C. clavatus is not clear. Studies in

Cyrtanthus still need to draw on additional data to indi-

cate which of the floral features reflect natural affinities

and which, if any, are the result of convergent evolution.

Distribution and habitat

Cyrtanthus debilis is endemic to the Outeniqua Moun-

tains of the southern Cape, between George and Oudts-

hoom (Figure 3). The populations occur on seasonally

moist southern and northern slopes in stony, sandy soil

amongst mountain fynbos. Since flowering is restricted to

the first summer after fire the plants are rarely seen.

Nonetheless, the species is known from several popula-

tions within the Outeniqua Nature Reserve where there

are no apparent threats to their future survival.

Etymology

The species epithet, debilis , reflects the apparent

frailty of the flowers when they appear from the black-

ened, newly burnt veld.

34

Bothalia 31,1 (2001)

WESTERN CAPE. — 3321 (Ladismith): Attakwas-

kloof, near summit of old Voortrekker Pass, (-DD),

Oliver 4134 (NBG, PRE). 3322 (Oudtshoorn): northern

slopes of Outeniqua Mountains, along Groot Doomrivier,

(-CC), Viviers & Vlok 370 (NBG); Outeniqua Mountains,

near summit of Robinson Pass, Snijman & Vlok 1717 (K,

NBG, PRE); Ruytersbosch, Gemmell sub BLFU5032

(BLFU, PRE); Jonkersberg on south-facing slopes of

Bolleberg, Vlok 814 (PRE).

ACKNOWLEDGEMENTS

I am grateful to the Western Cape Nature Conservation

Board for granting me permission to collect plants in the

wild; Jan and Anne Lise Vlok for their generous help and

hospitality; and Colin Paterson-Jones for invaluable

assistance in the field. Claire Linder Smith is sincerely

thanked for'the line drawing.

REFERENCES

DYER. R.A. 1977. Cyrtanthus montanus. The Flowering Plants of

Africa 44: t. 1756.

DYER, R.A. 1980. A new species of Cyrtanthus from the Baviaans-

kloof, southeastern Cape. Bothalia 13: 135.

NORDAL, I. 1979. Revision of the genus Cyrtanthus (Amaryllidaceae)

in East Africa. Norwegian Journal of Botany 26: 183-192.

REID, C. & DYER, R.A. 1984. A review of the southern African species

of Cyrtanthus. American Plant Life Society, La Jolla.

SAUNDERS, R. & SALINDERS, R. 2000. Does fynbos need to bum?

Veld & Flora 86: 76-78.

SNIJMAN, D.A. 1999. New species and notes on Cyrtanthus in the

southern Cape, South Africa. Bothalia 29: 258-263.

SNIJMAN, D.A. & VAN JAARSVELD, E.J. 1995. Cyrtanthus flam-

mosus. Flowering Plants of Africa 54: 100-103.

D.A. SNIJMAN*

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

7735 Claremont, Cape Town.

MS. received: 2000-08-10.

A NEW SPECIES OF BRUNSVIGIA (AMARYLLIDEAE) FROM WESTERN CAPE, SOUTH AFRICA

INTRODUCTION

Brunsvigia Heist., a genus of about 23 species, is

endemic to southern Africa. Although the genus is wide-

spread, the highest concentration of species is in the

northwest region of Western Cape and the midlands of

KwaZulu-Natal (Vorster 1999). When last reviewed.

Dyer (1950, 1951) recognized 17 species, one of which

was later formally described by Barker (1963). Since

then studies in the genus have been limited to the winter

rainfall region. Goldblatt (1972) and D. & U. Miiller-

Doblies (1994) transferred two further species to Bruns-

vigia after their respective re-assessments of Nerine

Herb, and Boophone Herb. In addition, D. & U. Muller-

Doblies (1994) described two new Brunsvigia species

from Namaqualand and the western Cape.

Independent cladistic analyses of morphological and

molecular data based on nrDNA ITS sequences have

indicated that Brunsvigia is closely allied to Nerine

(Snijman & Linder 1996; Meerow & Snijman 2000).

Both genera are recognized by their pink, rarely red or

white, more or less zygomorphic flowers, in which the

filaments are fused into a short basal tube. Brunsvigia is

distinguished from Nerine by its specialized fruiting

heads. These have long, stiff pedicels that radiate out-

ward in all directions and carry large, transversally

veined capsules which are tardily dehiscent. When dry,

the infructescence detaches from the bulb at ground level

and seed is dispersed while the entire head tumbles in the

wind. Other specialized features, shared by many, but not

all Brunsvigia species, are the brittle, yellowish brown

bulb tunics and the flattened leaves that are pressed to the

soil surface. The possible adaptive significance of the

prostrate-leaved habit, a particularly characteristic

growth form of many geophytes in southern Africa’s

winter rainfall region, has been fully discussed by Esler

et al. (1999).

The current re-assessment of species for the IUCN

Red List of threatened plants has stimulated renewed

interest in the critically transformed Coastal Renoster-

veld, which is now largely confined to the Elandsberg

Private Nature Reserve between Wellington and Tulbagh

(Rebelo 1996). Recent collections from this botanically

rich site have generated many new and unexpected

records. Most significant for the family Amaryllidaceae

has been the discovery of an unknown species of Bruns-

vigia, which is formally described here as B. elands-

montana.

Brunsvigia elandsmontana Snijman , sp. nov.,

species insignis perigonio actinomorpho, tubo perigonii

(2-5 mm longo), staminibus centralis, a speciebus nobis

notis bene distincta; differt a B. marginato (Jacq.) Aiton

floribus roseis vividis et staminibus brevibus (10-16

mm). Figura 4.

TYPE. — Western Cape, 3319 (Worcester); Wellington,

Elandsberg Private Nature Reserve, on well-drained peb-

bly flats, (-AC), 14-3-2000, Snijman 1731 (NBG, holo.;

E, K, MO, PRE).

Deciduous bulbous herb, 110-200 mm tall when flow-

ering. Bulb solitary, hypogeal, ± ovate, 30-45 mm diam.,

extended into a stout neck 30-50 mm long; outer tunics

tan-coloured and brittle; inner tunics cream-coloured and

fleshy. Foliage leaves 4—6, usually absent at flowering,

distichous, prostrate, overlapping at first becoming fal-

cate and outspread when mature, elliptical, up to 1250 x

25-70 mm, adaxial surface dark green, somewhat rough,

abaxial surface pale green with darker green veins,

smooth, margin cartilaginous, pink, crisped, apex obtuse

or somewhat acute. Inflorescence 6-18-flowered, initial-

ly compact, ultimately hemispherical, 80-130 mm diam.,

enlarging to 200 mm diam. in fruit; scape erect, elliptical

in cross section, 50-140 x 5-10 mm, dull rose pink,

breaking at ground level when dry; spathe valves 2,

oblong-lanceolate, 20-35 x 8-20 mm, leathery, spread-

ing, greenish pink; bracteoles at base of flowers linear,

20-35 mm long; pedicels straight, ± radiating, 20-45 mm

Bothalia 31,1 (2001)

35

FIGURE 4. — Brunsvigia elcmds-

montana : A, bulb and mature

leaves; B, inflorescence; C,

flower with two lateral tepals

and stamens removed; D, in-

fructescence. Drawn from

Snijman 1731 by John Man-

ning. Scale bar: A, B, D, 10

mm; C, 0.5 mm.

long, elongating to 40-75 mm in fruit, greenish pink,

turning pinkish when dry. Perigone actinomorphic,

17-23 x 20-30 mm. vivid rose-pink, throat usually con-

colorous with tepals, abaxial surface with paler midrib,

unscented; tube 2-5 mm long, expanding slightly to 2-3

mm diam. at throat; tepals oblong-lanceolate, 15-20 x

5-7 mm, separate or slightly overlapping at base, out-

spread, sometimes somewhat recurved towards apex,

margin plain or slightly undulate; stamens 6, centrally

arranged, the inner almost as long as tepals, the outer

somewhat shorter; filaments 10-16 mm long, pale and

swollen at base, otherwise slender and pink, tightly clus-

tered in lower third then slightly spreading and upturned

towards apex; anthers dorsifixed, 4 mm long, dusky pink

before opening; pollen cream-coloured. Ovary turbinate

and 3-angled, 4—5 mm across, greenish pink; ovules axile,

5 or 6 per locule; style 20-25 mm long, slender, central,

slightly upturned towards apex when mature; stigma tri-

fid, papillate. Capsule inflated, turbinate, 10-25 x 10-15

mm. 3-angled with papery, transversally veined walls, the

angles thinly ribbed, rounded near apex, loculicidally

dehiscent in upper two thirds, pinkish to straw-coloured.

Seeds non-dormant, greenish. ± 5 mm diam.

Diagnostic features

Brunsvigia elandsmontana is distinguished from

other species in the genus by the combination of its

bright pink flowers, an actinomorphic perigone with an

elongated tube, and centrally arranged filaments which

are upturned towards the apex.

Only two other species of Brunsvigia have an actin-

omorphic perigone: B. pulchra (W.F. Barker) D. &. U.

Miill.-Doblies from northern Namaqualand and B. mar-

ginata (Jacq.) Aiton from Western Cape, between the

Kouebokkeveldberge and Franschhoekberge (Figure 5).

With respect to the widely flared tepals, B. elandsmon-

tana is most like the highly ornamental B. marginata.

This scarlet-flowered species, however, is characterized

by a long perigone tube (5-10 mm) and well-exserted

stamens (30^15 mm long), whereas in B. elandsmon-

tana,, the perigone tube reaches only 2-5 mm long and

the 10-16 mm long stamens never exceed the tepals.

Both species occupy fire-prone fynbos habitats but at dif-

ferent elevations. Brunsvigia marginata , a montane

species, is found at 450-1 200 m above sea level, where-

as B. elandsmontana is restricted to the lowlands at alti-

tudes of ± 100 m.

Most species in the genus have pink flowers. Field

studies in the Western Cape suggest that colour intensity

is often variable within a species while the floral mark-

ings are usually constant. Thus in B. bosmaniae

Leighton, B. minor Lind, and B. striata (Jacq.) Aiton, the

flowers vary from pale to dark pink but the tepal colour

36

Bothalia 31,1 (2001)

on the adaxial surface is always broken by darker veins

or borders. In contrast, the flower colour of B.

elandsmontana is pure, and the adaxial tepal surface is

unbroken by darker streaking.

It is well established that the major pollinator of the

red-flowered B. marginata is the big brown butterfly,

Aeropetes tulbaghia (L.) (Johnson & Bond 1994). Less is

known about the pollination ecology of the pink-flow-

ered species of Brunsvigia. Preliminary work has shown

that the scented flowers of B. bosmaniae are regularly

visited at dusk by noctuid moths and two species of hawk

moth: Agnus convolvuli (L.) and Hippotion celerio (L.)

(Raimondo 1998). Occasionally noctuid moths visit the

unscented flowers of B. striata but they mainly attract

butterflies and the anthophorid bee, Amegilla niveata

(Friese) (J.C. Manning & J.C. Paterson-Jones pers.

comm.). As yet nothing is known about the pollination

biology of B. elandsmontana. However, the lack (to

humans) of any scent and the almost regular floral form

suggest that the flowers are likely to attract generalist,

diurnal, nectar-feeding insects.

Phenology

The population flowers between March and May, for

± two weeks. Within a month after flowering the

infructescence dries out, breaks loose and scatters seed

as it tumbles across the ground in the wind. The leaves

usually emerge in May after flowering, but they are shed

as soon as the dry season approaches, usually towards

the end of September. Flowering appears to be most pro-

lific after fire, not only in the first season, but for as long

as the habitat remains open.

Distribution and habitat

About 700 mature plants of B. elandsmontana are

known from a single population in the Elandsberg

Private Nature Reserve, northeast of Wellington (Figure

5). The Reserve, which extends from the western slopes

of the Elandsberg to the adjacent flats, covers a variety of

soil types and moisture regimes. These provide diverse

habitats within a mosaic of vegetation types, mainly fyn-

FIGURE 5. — Distribution of Brunsvigia elandsmontana , ©; and B.

marginata, •

bos, renosterveld and sandveld. Brunsvigia elandsmon-

tana is typically confined to marginal fynbos on well-

drained, pebbly flats. The habitat is remarkably rich in

geophytes, among which is a pink-flowered form of

Haemanthus sanguineus that extends eastwards into the

Breede River Valley. When the vegetation is mature the

overstorey is dominated by Clijfortia ruscifolia,

Leucadendron corymbosum, L. lanigerum, and Serruria

acrocarpa.

The population’s area of occupancy is extremely

small (less than 1 km2). Consequently, the IUCN Red

Data status of B. elandsmontana has been assessed as

Vulnerable D2.

Etymology

Brunsvigia elandsmontana is named to honour the

Elandsberg Private Nature Reserve, which apart from pro-

tecting the endangered geometric tortoise, Psammobates

geometricus (L.), is home to many rare plant species. Some

of the vulnerable or endangered plants protected here are

Aristea lugens, Haemanthus pumilio, Lachenalia polyphyl-

la, Lampranthus scaber , Moraea villosa subsp. elandsmon-

tana, Oxalis natans. Protea mucronifolia, Serruria candi-

cans, and Watsonia dubia.

Specimens examined

WESTERN CAPE— 3319 (Worcester): Wellington, Elandsberg

Private Nature Reserve, (-AC), Parker 141 (NBG); Parker sub

NBG167399 (NBG); Snijman 1678 (NBG); Snijman 1731 (NBG, E, K,

MO, PRE); Steyn A54 (NBG).

ACKNOWLEDGEMENTS

I am grateful to Dale and Elizabeth Parker for their

generous help and hospitality, and to John Manning and

Colin Paterson-Jones for their kind assistance with illus-

trations and photographs. Western Cape Nature

Conservation Board is thanked for granting me permis-

sion to collect wild plants in Western Cape.

REFERENCES

BARKER, W.F. 1963. Two new species of Amaryllidaceae. Journal of

South African Botany 29: 163-166.

DYER, R.A. 1950. A review of the genus Brunsvigia. Plant Life 6:

63-83.

DYER, R.A. 1951. A review of the genus Brunsvigia Heist. Plant Life

7: 45-64.

ESLER, K.J., RUNDELL, P.W. & VORSTER, P. 1999. Biogeography

of prostrate-leaved geophytes in semi-arid South Africa:

hypotheses on functionality. Plant Ecology 142: 105-120.

GOLDBLATT, P. 1972. Chromosome cytology in relation to classifi-

cation in Nerine and Brunsvigia (Amaryllidaceae). Journal of

South African Botany 38: 261-275.

JOHNSON, S.D. & BOND, W.J. 1994. Red flowers and butterfly pol-

lination in the fynbos of South Africa. In M. Arianoutsou & R.

Groves, Plant-animal interactions in Mediterranean-type

ecosystems: 137-148. Kluwer Academic Publishers, Dordrecht.

MEEROW, A.W. & SNIJMAN, D.A. 2000. Abstract: Phylogeny of Ama-

ryllidaceae tribe Amaryllideae based on nrDNA ITS sequences.

Botany 2000: new frontiers in botany. American Society of

Plant Taxonomists Conference, Portland, Oregon.

MULLER-DOBLIES, D. & MULLER-DOBLIES, U. 1994. De Lilii-

floris notulae 5. Some new taxa and combinations in the

Amaryllidaceae tribe Amaryllideae from arid southern Africa.

Feddes Repertorium 105, 5, 6: 331-363.

Bothalia 31,1 (2001)

37

RA1MONDO. D.C. 1998. Evolution of floral morphology in Bruns-

vigia and Crossyne (Amaryllidaceae). B.Sc. Hons project, Botany

Department, University of Cape Town. Unpublished.

REBELO. A.G. 1996. West Coast Renosterveld. In A.B. Low & A.G.

Rebelo, Vegetation of South Africa, Lesotho and Swaziland : 66.

Department of Environmental Affairs & Tourism. Pretoria.

SNIJMAN, D.A. & LINDER, H P. 1996. Phylogenetic relationships,

seed characters, and dispersal system evolution in Amaryllideae

(Amaryllidaceae). Annals of the Missouri Botanical Garden 83:

362-386.

VORSTER, P. 1999. The geographic distribution of Amaryllidaceae in

the Flora of southern Africa area. In J. Timberlake & S. Kativu,

African plants: biodiversity, taxonomy and uses: 171-177.

Royal Botanic Gardens, Kew.

D.A. SNIJMAN*

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

7735 Claremont, Cape Town.

MS. received: 2000-08-28.

COMMELINACEAE

REDISCOVERY OF TRICERATELLA DRUMMONDll, AND COMMENTS ON ITS RELATIONSHIPS

AND POSITION WITHIN THE FAMILY

The monotypic genus Triceratella Brenan was placed

in a separate tribe Triceratelleae in recent classifications

of the Commelinaceae (Faden & Hunt 1991; Faden

1998). However, Triceratella is still poorly understood

because it is known only from the type collection. The

genus and sole species, T. drummondii Brenan — named

in honour of the collector, R.B. Drummond — were

described by Brenan (1961a). based on a 1958 gathering

from southern Zimbabwe. The holotype of T. drum-

mondii is housed at Kew (K) with isotypes at PRE and

SRGH (Brenan 1961a), and the New York Botanical

Garden (NY, Faden pers. obs).

The new collection

In order to learn more about Triceratella , three inten-

sive searches of the type locality of T. drummondii have

been made since 1996, all without success (Faden pers.

obs.). Drummond reported to the second author that he

re-collected the species at the same locality ± 10 years

after his initial collection. The specimens, however, can-

not be located.

On 5 July 1997, a specimen of Triceratella was col-

lected in Mozambique during an environmental impact

assessment (EIA) survey in Moebase, Quelimane District,

Zambezia Province, 220 km northeast of Quelimane (38°

45 1 00" E 17° 5' 00" S). Initially, the specimen was difficult

to identify to genus, but it was keyed out by the third

author using Dyer (1976). The specimen, Dold 3227 (col-

lected with T. Avis and R. Lubke) is housed at GRA.

The new collection of Triceratella came from the sea-

ward margin of seasonal vlei in the dune slack, ± 800 m

west of the shoreline, and 100 m east of a forest of Icuria

dunensis Wieringa. The latter is a new genus and species

of Fabaceae: Caesalpiniodeae (Wieringa 1999) that was

also collected during the same trip. The habitat in which

these species were found may be described as coastal

dune scrub/woodland to forest that occurs along a narrow

coastal strip. Triceratella was growing in open wet sand

immediately behind the coastal dunes, with Digitaria

eriantha and Fimbristylis hispida in association with the

woody Garcinia livingstonei and Strychnos spinosa.

Depressions in the sand held freshwater wetlands domi-

nated by Eragrostis ciliaris, Xyris anceps and Utri-

cularia (sp. indet.). Only one population of about 20

plants of Triceratella was found during this survey. A

single plant was collected.

This new locality for Triceratella is over 1 000 km from

the type locality. This great distance, the differences in

ecology — at the type locality T. drummondii was recorded

as growing in moist sand on Forest Sandstone in associa-

tion with Bacopa floribunda , Fuirena leptostachya , Xyris

rubella, Torenia spicata and other species — and apparent

differences in the seeds of the new collection (when com-

pared to those of the original description and illustration)

led us to speculate that the new specimen might be a sec-

ond species of the genus. The isotype of T. drummondii

housed at PRE was borrowed so that the identity and status

of the new specimen could be determined. It was conclud-

ed that the new collection was conspecific with the type of

T. drummondii.

The recent collection provided the opportunity to

study and describe some features of this species in

greater detail than had been done previously. Particular

attention was given to the seeds and to the structure of

the hairs on the leaves and sepals. These examinations

were carried out by means of a JEOL JSM 840 Scanning

Electron Microscope (SEM) at the Rhodes University

Electron Microscope Unit. All samples were already dry,

and thus were not further treated before being sputter-

coated with gold-palladium.

Seed morphology

Seed morphology is important in the classification of

Commelinaceae (Bruckner 1926, 1930; Brenan 1961b;

Faden 1998). Such features as the position of the embry-

otega (or embryostega or operculum) and shape of the

hilum have been used to characterize the genera and

some subtribes (Faden 1998). Because the morphology

of the seeds in the new collection and in the type of T.

drummondii initially seemed different, seeds of both

specimens were examined by means of SEM. The seeds

were found to be identical in shape and surface morphol-

ogy. Figure 6A, B shows a seed from the specimen from

Mozambique. The seed shape and ribbed nature agree

with the illustration by Brenan (1961a). The details of the

colliculate surface ornamentation are clarified by our

SEM micrographs.

One point that confused us initially was the presence

or absence of a distinct embryotega on the seed. The pho-

tomicrographs do not show the clear circular or elliptic

outline that normally demarcates the embryotega on the

testa of Commelinaceae seeds. Yet according to Brenan’s

38

Bothalia 31,1 (2001)

FIGURE 6. — Scanning electron micrographs of seed, leaf and sepal surfaces of Triceratella drummondii ( Dold 3227). A, surface of seed, show-

ing colliculate and ribbed nature in greater detail; B, whole seed, placed face down on hilum with embryotega uppermost, note absence of

micropylar collar; C, four-celled clavate hairs on adaxial leaf surface; D, four-celled clavate hairs on abaxial sepal surface; E, close-up

view of apical cells of clavate hairs found on sepal. Scale bars; 100 mm.

type description, the embryotega is dorsal and prominent

in Triceratella. The explanation of this apparent discrep-

ancy is that the seeds lack a micropylar collar. Thus the

prominent point of the seed is the embryotega, but with-

out a micropylar collar as there is no circular or elliptic

ring surrounding it on the surface. This morphology is

also found in Cartonema R.Br. (Grootjen 1983), and sup-

ports the relationship of Triceratella with Cartonema

and their segregation as subfamily Cartonematoideae.

Leaf hairs

The abundant hairs on the leaves of Triceratella drum-

mondii were described as ‘very uniform, (3)4(5)-celled,

uniseriate, unbranched, usually with a 3-celled terminal

stalk; terminal cell clavate, very delicate walled’ by

Tomlinson (1964: 208). These hairs are similar to glandu-

lar microhairs, which are present in nearly all Com-

melinaceae, terminating in a clavate cell. Moreover, the

terminal cell of occasional hairs in Triceratella are greatly

enlarged, thereby resembling glandular microhairs more

closely (Tomlinson 1964). The hairs in Triceratella differ

from glandular hairs in being mainly 4-celled, instead of

regularly 3-celled, and in the terminal cell usually being

less differentiated from the cells basal to it. Although

Tomlinson avoids describing these hairs as glandular, he

notes that the terminal cell in each hair is thin-walled and

has denser contents than the cells more basal to it. This

suggests that the terminal cell may be secretory and the

hairs thereby glandular, which Faden (1998) considers

them to be. Our photomicrograph (Figure 6C) shows foliar

hairs identical to those illustrated by Tomlinson, except

that the cells have collapsed. We believe that this is a result

of drying in the preparation of the herbarium specimen,

and that the hairs shown by Tomlinson were either ide-

alised or rehydrated. The terminal cell does not appear dif-

ferentiated from the other cells of the hair, except in shape.

We were unable to perceive the density of the cellular con-

tents as the SEM micrographs do not show cellular con-

tents through the cell wall.

In discussing the possible relationships of Tricera-

tella, Tomlinson ( 1964) mentions the resemblance of its

Bothalia 31,1 (2001)

39

hairs to the distinctly glandular hairs in Cartonema. The

hairs in Cartonema philydroides F.Muell. are similar to

those of Triceratella but differ in being longer, thicker,

more tapered from the base, with a smaller, but distinct-

ly enlarged terminal cell (Faden unpublished). When pre-

sent in other genera, e.g. Tinantia , such hairs are normal-

ly confined to the reproductive parts, such as the sepals,

pedicels and inflorescence axes. The presence of such

glandular hairs on the vegetative parts of Cartonema and

Triceratella is exceptional in Commelinaceae, and is a

defining character of subfamily Cartonematoideae, as is

the absence of true glandular microhairs (Faden

1998).The resemblance of some hairs in Triceratella to

glandular microhairs, supports the conclusion of

Tomlinson (1964) that Triceratella forms a link between

Cartonema and the rest of the Commelinaceae.

Sepal hairs

The hairs on the sepals (Figure 6D, E) closely resem-

ble those on the vegetative parts. While they do not

exactly match the distinctly glandular hairs on some

Cartonema species, e.g. C. philydroides , they closely

resemble hairs on the vegetative parts of some undeter-

mined Cartonema collections in the US National

Herbarium (US).

In conclusion, the similarity of unusual seed and tri-

chome morphologies in Triceratella and Cartonema

strongly supports their relationship and their treatment as

the separate subfamily Cartonematoideae. This study does

not, however, provide the final verdict as to whether the

hairs observed in Triceratella are glandular in nature or not.

Although they come from different habitats, the new

specimen and the type collection both were growing in

wet sand. This information might prove valuable in help-

ing to identify other areas where this species might be

found. Until, and unless additional populations of this

species are found, it must be considered to be Critically

Endangered according to the new IUCN Red Data List

categories (IUCN 1994).

ACKNOWLEDGEMENTS

Mr R. Cross and Ms S. Pinchuck of the Rhodes

University Electron Microscopy Unit are thanked for

their assistance in use of the scanning electron micro-

scope. The curator of PRE is thanked for the loan of the

isotype, and for permission to examine a seed from this

specimen by means of SEM.

REFERENCES

BRENAN. J.P.M. 1961a. Triceratella, a new genus of Commelinaceae

from southern Rhodesia. Kirkia 1: 14—19.

BRENAN, J.RM. 1961b. The classification of Commelinaceae.

Journal of the Linnean Society, Botany 59: 349-370.

BRUCKNER. G. 1926. Beitrage zur Anatomie. Morphologie und

Systematik der Commelinaceae. Botanische Jahrbiicher 61,

Beiblatt 137: 1-70.

BRUCKNER, G. 1930. Commelinaceae. In A. Engler, Die natiirlichen

Pflanzenfamilien, edn 2, 15a: 159-181. Wilhelm Engelmann,

Leipzig.

DYER. R.A. 1976. The genera of southern African flowering plants 2:

910. Department of Agricultural Technical Services, Pretoria.

FADEN, R.B. 1998. Commelinaceae. In K. Kubitzki, The families and

genera of vascular plants 4: 109-128. Springer- Verlag, Berlin.

FADEN, R.B. & HUNT. D.R. 1991. The classification of the Com-

melinaceae. Taxon 40: 19-31.

GROOTJEN, C.J. 1983. Development of ovule and seed in Cartonema

spicatum R.Br. (Cartonemataceae). Australian Journal of

Botany 31: 297-305.

IUCN 1994. IUCN Red List categories. IUCN, Gland, Switzerland.

TOMLINSON, P.B. 1964. Notes on the anatomy of Triceratella

(Commelinaceae). Kirkia 4: 207-212.

WIERINGA, J.J. 1999. Monopetalanthus exit. A systematic study of

Aphanocalyx, Bikinia, Icuria, Michelsonia and Tetraberlinia

(Leguminosae, Caesalpinioideae). Wageningen Agricultural

University Papers 99: 1-320.

N.P. BARKER*, R.B. FADEN**, E. BRINK*** and A.P. DOLD*

’ Department of Botany, Rhodes University, 6140 Grahamstown, South

Africa.

Department of Botany, National Museum of Natural History,

Smithsonian Institution, Washington DC 20560-0166, USA.

**' Selmar Schonland Herbarium, P.O. Box 101, 6140 Grahamstown.

South Africa.

MS. received: 2000-05-16.

ASPHODELACEAE

JODRELLIA FISTULOSA. A NEW GENERIC RECORD FOR THE FLORA OF SOUTHERN AFRICA REGION

Jodrellia Baijnath, described in 1978 (Baijnath

1978), is a small genus of about three species. Currently

it is regarded as having an uncertain generic circum-

scription (Kativu 1996). It was initially segregated from

Bulbine Wolf on the basis of tepal nervation and rela-

tive width: in Bulbine they are 1 -nerved, whereas in

Jodrellia the inner ones are 1 -nerved and the outer ones

3-5-nerved; in Bulbine the inner ones are wider than

the outer, whereas in Jodrellia the reverse occurs

(Baijnath 1978). Recent DNA studies have shown that

Jodrellia may in fact be related to Trachyandra Kunth

rather than Bulbine (Baijath pers. comm., cited by

Kativu 1996).

The new record is from the extreme eastern part of the

Caprivi Strip, Namibia, botanically a poorly known area

of the Flora of southern Africa region. It matches well the

PRE specimens of Jodrellia fistulosa (Chiov.) Baijnath

from Tanzania, Zimbabwe and Zambia, which have non-

inflated fruits. The type, Chiovenda 557, is from the

apparently disjunct northern Ethiopian/Eritrean popula-

tions. This specimen is in poor condition and it is impos-

sible to decide whether the fruits are mature, and if not,

whether they would be inflated at maturity. There is thus

a possibility that the southern populations may represent

a distinct entity. Kativu (1996) provides a distribution

map of Jodrellia fistulosa for the Flora zambesiaca area.

40

Bothalia 31,1 (2001)

The new locality (Figure 7) is not very far from the west-

ernmost record cited by Kativu (1996), i.e. Machili,

Zambia (16°5f S; 25°07' E). The habitat is fairly typical

for the species, i.e. Colophospermum mopane (Benth.)

J. Leonard woodland, in sandy soil.

Specimen examined

NAMIBIA. — 1724 (Katima Mulilo): 29 km SE of Katima Mulilo

on Ngome road, (-DA), Venter, Hahn & Archer 128 (PRE, WIND).

REFERENCES

BAIJNATH, H. 1978. Jodrellia, a new genus of Liliaceae from tropi-

cal Africa. Kew Bulletin 32: 571-578.

KATIVU, S. 1996. Asphodelaceae of the Flora zambesiaca area.

Kirkia 16: 27-53.

C. ARCHER*

FIGURE 7. — Distribution of Jodrellia fistulosa (Chiov.) Baijnath in * National Botanical Institute, Private Bag X101, 0001 Pretoria,

southern Africa. MS. received: 2000-08-08.

ANTHERICACEAE

THE CORRECT NAME IN CHLOROPHYTUM FOR ANTHER1C UM LONGISTYLUM

Until recently, the generic limits defined by Ober-

meyer (1962) for the southern African species of

Anthericum L. and Chlorophytum Ker Gawl. have re-

mained unchallenged. A study of these genera by Nordal

et al. (1990) showed that the genus Anthericum is mainly

European, and in Africa should be applied in a restricted

sense only, with about three species south of the Sahara

(Nordal & Thulin 1993), necessitating the transfer of the

remaining African species south of the Sahara to

Chlorophytum. Kativu (1993) thus made 23 new combi-

nations for taxa in the Flora zambesiaca area, including

those with ranges extending into southern Africa.

One of the species recognized by Obermeyer (1962)

is Anthericum longistylum Baker (1876), a distinctive

taxon occurring in the northern and eastern parts of

the Flora of southern Africa region (Figure 8) plus

Zimbabwe. Obermeyer lists as a synonym Anthericum

FIGURE 8. — Distribution of Chloropytum recurvifolium (Baker) C. Archer

& Kativu in southern Africa.

polyphyllum Baker (1896). Due to the pre-existing

Chlorophytum longistylum Poelln. (1946), Kativu could

not transfer Anthericum longistylum to Chlorophytum ,

but he was apparently unaware of the pre-existing C.

polyphyllum (Baker) Poelln. (1946), based on Dasy-

stachys polyphylla Baker (1898), now considered by

Nordal et al. ( 1997) to be a synonym of Chlorophytum

suffruticosum Baker, an East African species. Kativu

illegitimately transferred Anthericum polyphyllum

Baker (1896) as Chlorophytum polyphyllum (Baker)

Kativu. A new combination in Chlorophytum is there-

fore required for our species. Fortunately there is anoth-

er epithet available for this taxon: Obermeyer (1962)

places Anthericum recurvifolium Baker as a synonym of

Anthericum longistylum. The new combination is affect-

ed here.

Chlorophytum recurvifolium (Baker) C. Archer

& Kativu , comb. nov. Type: Zimbabwe, Mashonaland,

6-Mile Spruit near Salisbury [now Harare], Cecil 143

(K, holo.).

Anthericum recurvifolium Baker in Kew Bulletin 1906: 28 (1906).

Anthericum longistylum Baker: 305 (1876); Baker: 381 (1896);

Obermeyer: 686 (1962). Type: South Africa, ‘Transvaal’, without pre-

cise locality, Baines s.n. (K, holo.).

Anthericum polyphyllum Baker: 383 (1896). Chlorophytum poly-

phyllum (Baker) Kativu: 63 (1993), non C. polyphyllum (Baker)

Poelln. (1946). Type: South Africa, Mpumalanga, near Barberton,

Gal pin 1149 (K, holo.; GRA, PRE).

REFERENCES

BAKER, J.G. 1876. Revision of the genera and species of Anthericeae

and Eriospermae. Journal of the Linnean Society 15: 253-363.

BAKER, J.G. 1896. Liliaceae. In W.T. Thiselton-Dyer, Flora capensis

6: 253-384. Reeve. London.

BAKER. J.G. 1898. Liliaceae. In W.T. Thiselton-Dyer, Flora of trop-

ical Africa 7: 421-568. Reeve, Kent.

Bothalia 31,1 (2001)

41

BAKER, J.G. 1906. Anthericum (Phlangium) recurvifolium Baker

(Liliaceae-Asphodeleae). In Diagnoses Africanae XIV. Kew

Bulletin 1906: 15-30.

KATIVU, S. 1993. In S. Kativu & I. Nordal, New combinations of

African species in the genus Chlorophytum (Anthericaceae).

Nordic Journal of Botany 13: 59-65.

NORDAL, I.. ERIKSEN, T.E. & FOSBY, M. 1990. Studies in the

generic delimitations in Anthericaceae. Mitteilungen aus dem

Institut fur Allgemeine Botanik Hamburg 23(b): 535-559.

NORDAL, I.. KATIVU. S. & POULSEN, A.D. 1997. Anthericaceae. In

R.M. Polhill, Flora of tropical East Africa. Balkema, Rotterdam.

NORDAL, I. & THULIN, M. 1993. Synopsis of Anthericum and

Chlorophytum (Anthericaceae) in the Horn of Africa, including

the description of three new species. Nordic Journal of Botany

13: 257-280.

OBERMEYER. A. A. 1962. A revision of the South African species of An-

thericum, Chlorophytum and Trachyandra. Botlmlia 7: 669-767 .

VON POELLNITZ, K. 1946. Die Chlorophytum- arten Tanganyikas.

Portugaliae acta biologica, ser. B, 1: 255-383.

C. ARCHER* and S. KATIVU**

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

** Department of Biological Sciences, University of Zimbabwe, PO.

Box MP 167, Mount Pleasant, Harare, Zimbabwe.

MS. received: 2000-08-08.

ASTERACEAE

A NEW SPECIES OF OEDERA FROM WESTERN CAPE, SOUTH AFRICA

Oedera epaleacea Beyers , sp. nov. Oederae fove-

olatae affinis sed receptaculo epaleaceo floribus disci

femineo-sterilibus stylo modo breviter fisso differt.

TYPE. — Western Cape, 3219 (Wuppertal): Swartrug-

gens, on Farm Zeekoe-gat, southwest of the farm road

about 500 m from the turnoff on the road to Katbakkies

Pass, 1 030 m, (-DC), 5-09-2000. Beyers 272 (NBG,

holo.; K, PRE, UPS, iso.).

Erect, moderately to densely branched, rounded

shrub, up to 0.6 m tall, with a stout trunk up to 50 mm

diam. Branches ascending-erect, glabrous, leafy, becom-

ing nude and marked with leaf scars. Leaves alternate,

spreading, sessile, entire, midribbed, narrowly oblong,

4—12 x 0.8-1. 5 mm, glabrous, densely and distinctly

glandular-punctate and foveolate, mucronulate; semi-

terete in cross section. Capitula solitary, sessile, termi-

nal, heterogamous, radiate. Involucre campanulate, 4—5

mm wide. Involucral bracts 5- or 6-seriate, 36^47,

imbricate, firm and hard basally, scarious apically,

midribbed, glabrous; outer ovate, inner gradually longer,

narrowly elliptic and apically brown, innermost narrow-

ly obovate or narrowly oblong with a spreading, brown

apical limb, up to 10.3 x 1.7 mm, acute. Receptacle

slightly convex, shortly squamose, epaleate. Ray florets

female, fertile, 13-18; tube cylindrical, 2-3 mm long,

with ± stalked glands; lamina spreading, narrowly obo-

vate, 6-10 x 1.3-2. 2 mm, yellow with a dorsal, brown

stripe, 4-veined, apically minutely 3-lobed; style terete,

bifid, 2.5— 4.0 mm long; style branches erect-spreading,

linear, 0.7-1. 5 mm long, glabrous, stylopodium not dis-

tinct; cypselas almost terete or angular, narrowly elliptic,

1.8-2. 6 x 0.5-0. 7 mm, densely pilose; pappus crown-

like, of ± connate scales, up to 0.6 mm long. Disc florets

38 — 42, female sterile; corolla 3. 7-5. 5 x 0.6-0. 9 mm,

divided into a lower cylindrical tube with scattered, ±

stalked glands on the outside and upper narrowly cyathi-

form, glabrous limb; corolla lobes 5, spreading, ovate-

triangular. 0. 3-0.7 mm long, dorsally gland-dotted; style

terete, 2. 5-5. 3 mm long, shallowly cleft, style branches

up to 0.3 mm long, apically rounded and papillate; sty-

lopodium distinct; anthers linear, 1.4— 2.2 mm long, with

sterile, flat, ovate apical appendage and sterile, flat, sub-

ulate, entire tails, 0.2-0.4 mm long; cypselas almost

terete, narrowly elliptic to narrowly oblong, 1.5-2. 7 x

0.3-0. 6 mm, glabrous or sparsely pilose, may be scat-

tered with ± stalked glands; pappus crown-like, of ± con-

nate scales, up to 0.6 mm long. Figure 9.

Diagnostic characters and relationships

Oedera foveolata (Bremer) Anderb. & Bremer and O.

sedifolia (DC.) Anderb. & Bremer are among those

species, treated in the taxonomic revision of Relhania

L’Her. by Bremer (1976), which were transferred to

Oedera by Anderberg & Bremer (1991). O. epaleacea is

related to the above two species, which are recognized

by their glandular-punctate leaves. As in O. epaleacea,

O. foveolata, a species only known from Karoo Poort,

the leaves are distinctly glandular-punctate and foveo-

late, and the cypselas of the ray florets are densely pilose.

However, in O. epaleacea the receptacle is epaleate and

the disc florets are female sterile. O. epaleacea grows

intermingled with O. sedifolia, one of the more common

species in the genus, but can easily be distinguished from

it by the glabrous branches and leaves and the female

sterile disc florets. The leaves of O. epaleacea, when

crushed have a sulphury odour. In O. sedifolia the

branches and leaves are tomentose becoming glabrous

later, the disc florets are perfect and leaves when crushed

give off a resinous odour.

Distribution and ecology

Oedera epaleacea is known only from the Farm

Zeekoe-gat in the Swartruggens area (Figure 10). It

occurs in arid fynbos where it grows in crevices between

rock sheets, in sandstone-derived sandy soils, in full or

partial sun. Flowering specimens seen from June to

September.

Etymology

The specific epithet refers to the epaleate receptacle.

Other specimens examined

WESTERN CAPE. — 3219 (Wuppertal): Zeekoe-gat 137, west of

Riet River, in ravine. 16-06-2000, Stobie 2 (NBG); ibid., on hill slope,

22-07-2000, Stobie sub NBG1 70457 (NBG).

FIGURE 9. — Oedera epaleacea , Beyers 272 (NBG). A, portion of plant; B, leaf (abaxial view and cross section). C-E, involucral bracts; C, outer;

D. middle; E, innermost. F, disc floret; G, stamen of disc floret; H, style of disc floret; I. ray floret; J, cypsela of ray floret; K, style of ray

floret.

FIGURE 10. — Geographical distribution of Oedera epaleacea.

ACKNOWLEDGEMENTS

I would like to thank Mrs Mary Stobie, a keen ama-

teur botanist, for bringing the material in for naming and

also for accompanying me to their farm to collect the

type material, Prof. Kare Bremer for his expert advice.

Dr Ted Oliver for the Latin diagnosis, and Mrs Inge

Oliver for the line drawings.

REFERENCES

ANDERBERG, A. A. & BREMER, K. 1991. Parsimony analysis and

cladistic reclassification of the Relhania generic group (Aster-

aceae-Gnaphalieae). Annals of the Missouri Botanical Garden

78: 1061-1072.

BREMER, K. 1976. The genus Relhania (Compositae). Opera Botan-

ica 40: 1-85.

J.B.P. BEYERS*

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

7735 Claremont, Cape Town.

MS. received: 2000-10-04.

Bothalia 31,1 (2001)

43

FABACEAE

FIRST DISTRIBUTION RECORD FOR BRA CHYSTEGIA SPICIFORMIS (CAESALPINIOIDEAE) IN SOUTH AFRICA

Brachystegia spiciformis Harms is a characteristic

tree constituent of the so-called miombo woodlands, a

colloquial name for a broad vegetation type covering

more than 2.7 million km2 in southern, central and east-

ern Africa (Millington et al. 1994; Campbell 1996).

These woodlands form the dominant element of the

Zambezian Region, a phytogeographical unit that covers

an area of about 3.8 million km2 in south-central Africa

(White 1983). Although the Zambezian Region extends

southwards across the Limpopo River Valley to include

most of the Northern Province of South Africa, B. spici-

formis has hitherto been conspicuous by its absence from

South Africa. Previously known only from pollen

records dating to 19 000 years BP from near Naboom-

spruit, some 350 km south of its current main distribu-

tion area, the absence of B. spicifonnis from South Africa

has been the subject of much speculation (Scott 1982;

Frost 1996).

Recent botanical explorations in the remote northeast-

ern Soutpansberg of the Northern Province, a region

known as Venda, have now brought to light a relict

miombo-like plant community with B. spiciformis as the

dominant canopy tree. Although the presence of B. spici-

formis in Venda has hitherto been unknown to botanists,

it is well known to the local Vhavenda; its Tshivenda

name is Mutsiwa. The locality of the newly discovered

populations of B. spiciformis , the extant distribution of

this species elsewhere in southern Africa and the nearest

known site of fossil Brachystegia pollen, are shown in

Figure 11.

This surprising discovery, which extends the distribu-

tion range of B. spiciformis southwards across the arid

Limpopo Valley, is of considerable biogeographical sig-

nificance. It is believed that during geological times,

miombo woodlands have expanded and contracted in

response to climatic change (Campbell 1996). The popu-

FIGURE 11. — Brachystegia spiciformis. New record, ▲; previously

known distribution in southern Africa, based on collections at

PRE, •. Site from which fossil pollen attributed to Brachy-

stegia has been recorded, ■.

lation of B. spiciformis in Venda may well be a relict,

dating from a once much further southward extension of

miombo woodland, thus supporting palaeopalynological

evidence suggesting the former presence of this vegeta-

tion type as far south as Naboomspruit. In this respect, B.

spiciformis recalls the occurrence of relict and widely

disjunct populations of the tropical African Bracken-

ridgea zanguebarica Oliv., Millettia stuhlmannii Taub.

and Oxytenanthera abyssinca (A. Rich.) Munro in Venda.

Because of their considerable cultural significance to the

local Venda people, it was speculated in the past that the

presence of the latter three species in Venda might have

been due to human introduction (Netshiungani & Van

Wyk 1980). However, with the discovery of B. spici-

formis in the same general area, it is tempting to implicate

one or more natural vicariant events in the establishment

of these isolated outlier populations at the southern end

of their ranges in Africa.

The newly discovered B. spiciformis community com-

prises several thousand trees occurring as fragmented

archipelago-like islands within the prevailing Soutpans-

berg Arid Mountain Bushveld (Van Rooyen & Breden-

kamp 1998). Mature trees are 15-20 m tall. The slender,

unbranched trunks indicate that they have not been sub-

jected to damage by humans or larger animals in the

recent past. Seed dispersal in B. spiciformis is by explo-

sive dehiscence of the pod and the seed is seldom dis-

persed far from the mother tree (Chidumayo & Frost

1996). This is reflected by the age structure of the new B.

spiciformis community; more mature (tallest) trees are

concentrated towards the centre of the vegetation islands.

The understorey is conspicuously species-poor and

include such taxa as Lagynias dryadum and Leptactina

delagoensis, with suppressed saplings (or suckers) of B.

spiciformis. Surrounding vegetation is dominated by

Combretum collinum subsp. gazense , C. vendae, Albizia

adianthifolia, Afzelia quanzensis and Burkea africana.

In view of the localized occurrence of B. spiciformis

in Venda, the uniqueness of its habitat and the scientific

importance of this outlier population, it is hoped that the

conservation authorities concerned will afford official

protection to the site.

Voucher specimens

NORTHERN PROVINCE.— 2230 (Messina): Thengwe, (-DA),

PJ.H. Hurter 124 (PRE, PRU), E. van Wyk & PJ.H. Hurter 5 (PRE),

E. van Wyk 65 (PRE, K).

ACKNOWLEDGEMENTS

We are indebted to Prof. A.E. van Wyk for confirming

the identity of herbarium material and for constructive

comments on the manuscript. We also thank Mrs Emsie

du Plessis for improvements to the text, Dr Hugh Glen

for assistance with the distribution map and Mr Mervyn

Lotter for access to literature on miombo woodlands.

44

Bothalia 31,1 (2001)

REFERENCES

CAMPBELL. B. (ed.) 1996. The miombo in transition: woodlands and

welfare in Africa. Centre for International Forestry Research,

Bogor.

CHIDUMAYO, E. & FROST, P. 1996. Population biology of miombo

trees. In B. Campbell, The miombo in transition: woodlands

and welfare in Africa : 59-71. Centre for International Forestry

Research, Bogor.

FROST. P. 1996. Palaeohistory of miombo woodlands. In B. Campbell,

The miombo in transition: woodlands and welfare in Africa: 6.

Centre for International Forestry Research, Bogor.

MILLINGTON, A.C.. CR1TCHLEY, R.W., DOUGLAS, T.D. & RYAN,

P. 1994. Estimating woody biomass in sub-Saharan Africa. The

World Bank. Washington DC.

NETSHIUNGANI, E.N. & VAN WYK, A.E. 1980. Mutavhatsindi:

mysterious plant from Venda. Veld & Flora 66: 87-90.

SCOTT, L. 1982. A late Quaternary pollen record from the Transvaal

bushveld. South Africa. Quaternary Research 17: 339-370.

VAN ROOYEN, N. & BREDENKAMP, G. 1998. Soutpansberg Arid

Mountain Bushveld. In A.B. Low & A.G. Rebelo, Vegetation of

South Africa. Lesotho and Swaziland: 21. Department of En-

vironmental Affairs and Tourism, Pretoria.

WHITE, F. 1983. The vegetation of Africa. Natural Resources Research

No. 20. A descriptive memoir to accompany the UNESCO/

AETFAT/UNSO vegetation map of Africa. UNESCO. Paris.

P.J.H. HURTER* and E. VAN WYK **

* Lowveld National Botanical Garden, PO. Box 1024, 1200 Nelspruit.

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

MS. received: 2000-10-04.

LAMIACEAE

PLECTRANTHUS HILLIARDIAE SUBSP. AUSTRALIS. A NEW TAXON FROM EASTERN CAPE, SOUTH AFRICA

Plectranthus hilliardiae subsp. australis Van

Jaarsv. & A.E. van Wyk , subsp. nov., a subspecie typica

statura majora, foliis ellipticis, 85-160 x 30-50 mm,

nitentibus, paene glabris, pagina adaxali purpurea, mar-

ginibus serrato-dentatis differt.

TYPE. — Eastern Cape, 3129 (Lusikisiki): northern

Transkei, Mbotyi Forest, forest floor, (-BC), Van

Jaarsveld, Bellstedt, Dekker, F. van Wyk & Adams 16345

(NBG, holo.). Figure 12.

Erect to decumbent, soft, semi-succulent, herbaceous,

aromatic perennial up to 650 mm tall; stems 4-angled, up

to 4 mm diam, moderately branched, antrorse pilose with

tufts of long, multicellular, purple-tinged hairs at nodes.

Leaves elliptical, 85-160 x 30-50 mm, dark green, shiny,

subglabrous to sparsely strigose on upper surface, dark

purple to mottled purplish, reticulate-veined on lower

surface, the veins sparsely strigose and dotted with small,

transparent, sessile glands, apex acute, base cuneate,

margin serrate-dentate, ciliate; petiole 15-35 mm long,

appressed glandular-pilose. Inflorescence a simple or

loose-branched panicle, up to 250 mm tall; rachis glan-

dular-tomentose; bracts linear-lanceolate, up to 2 mm

long, persistent. Flowers in sessile 3-flowered cymes

forming 6-flowered verticillasters, 10-30 mm apart;

pedicel 4-5 mm long. Calyx 3-4 mm long at flowering

stage, enlarging to 10 mm in fruiting stage; upper lip

ovate, erect, lower lip 4-toothed, deltoid-subulate.

Corolla pale bluish, rarely white; lobes purple-flecked;

tube 23-27 x 4 mm, parallel-sided and narrowing up to 3

mm at throat; upper lip erect, up to 6 mm long, bilobed

at apex with lateral ear-like lobes; lower lip shallow

cymbiform, up to 4 mm long, horizontal or deflexed.

Stamens 4, free, lower pair longer, up to 8 mm long. Style

horizontally exserted by 8-10 mm. Nutlets dark brown,

1.8 mm diam. Figure 13.

The new taxon was first noticed and recorded from

Magwa Falls in 1992 (Van Jaarsveld & Powrie 12226).

An evaluation of the southern forms of P. hilliardiae

from between Port St Johns and Lupatane for their horti-

cultural potential, revealed a consistent variation in leaf

shape, size, dentation and colour of the undersurface of

the leaves. During a recent collecting trip (April 2000)

the northern limits of the range of the new subspecies

were traced to the Noyokaan Gorge, just south of

Lupatane Gorge. Plants from the more open Lupatane

Gorge represented the typical subspecies hilliardiae.

Both subspecies of P. hilliardiae are confined to the

Pondoland Centre of Endemism, a region congruent with

surface outcrops of quartzitic sandstone belonging to the

Msikaba Formation.

Subsp. australis is characterized by somewhat larger,

elliptic, shiny leaves, 85-160 x 30-50 mm, with an

almost glabrous, purple adaxial surface and serrate-den-

tate margin. It occurs in deep shade in well- watered

gorges, but also in tree-fall openings of climax coastal

escarpment forests (Mbotyi Forest). The leaves of subsp.

hilliardiae are smaller, broadly elliptical to elliptical-

obovate, 55-90 x 40-60 mm, sparsely strigose above,

the surface not shiny and the margins crenate-dentate.

The typical subspecies tends to occur in more open and

exposed terrain and slightly drier situations, such as

FIGURE 12. — Known distribution of Plectranthus hilliardiae subsp.

australis in Eastern Cape, •; typical subspecies, O.

Bothalia 31,1 (2001)

45

FIGURE 13. — Plectranthus hilliar-

diae subsp. australis, x 0.75.

Artist: Vicki Thomas.

along forest margins and the fringes of stream banks

(Lupatane, Mkambati and Umtamvuna River Gorges).

Subsp. australis exhibits a more vigorous growth in cul-

tivation, retaining its ornamental purple leaf colouring

and floriferous nature. In 1994. a specimen of subsp.

australis from Magwa Falls was hybridized with P. sac-

catus var. longitubus (from Umtamvuna River Gorge)

producing a very attractive cultivar.

Other specimens examined

EASTERN CAPE. — 3129 (Lusikisiki): Fraser Falls, (-BC), Van

Jaarsveld, Bellstedt, Dekker, F. van Wyk & Adams 16381 (NBG);

Magwa Falls, (-BC), Van Jaarsveld & Powrie 12226 (NBG); Myokane

Gorge, (-BD), Van Jaarsveld 16339 (NBG).

ACKNOWLEDGEMENTS

Our thanks go to Mr G. Germishuizen and Mrs E. du

Plessis for editing the text and Dr O.A. Leistner for trans-

lating the diagnosis into Latin.

E.J. VAN JAARSVELD* and A.E. VAN WYK**

* National Botanical Institute, Kirstenbosch, Private Bag X7, 7735

Claremont.

** H.G.W.J. Schweickerdt Herbarium, Department of Botany,

University of Pretoria, 0002 Pretoria.

MS. received: 2000-10-31.

46

Bothalia 31,1 (2001)

ASTERACEAE

ANEW SPECIES OF EMILIA (SENECIONEAE) FROM SOUTH AFRICA

INTRODUCTION

Emilia Cass, is a genus of tribe Senecioneae compris-

ing almost 100 species distributed through tropical

Africa and Asia (Jeffrey 1986; Bremer 1994). It is dis-

tinguished by its annual or perennial herbaceous habit,

scanty indumentum, ecalyculate involucre and a basic

chromosome number n = 5 (Nordenstam 1978; Jeffrey

1986). Although about 60 species are currently recog-

nized from mainland Africa, especially Tanzania, only

six are known from southern Africa. The species de-

scribed here was collected as part of a botanical survey

of the flora of the Nieuwoudtville area and was identified

as an unnamed species of Emilia by Prof. B. Nordenstam

of the Swedish Museum of Natural History, an expert in

the genera of the tribe. It represents the first record of the

genus from the winter rainfall region of South Africa and

an unusual disjunction in the distribution of this essen-

tially tropical genus.

Emilia hantamensis J.C. Manning & Goldblatt ,

sp. nov., a speciebus alteribus Emiliae differt capitulis

floribus radiatis, foliis dentatis vel pinnatisectis et capit-

ulis magnis solitariisque, involucris 6-10 mm diam.

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

ville Dist., Farm Glenlyon, along trekpath, on doleritic

clay, (-AC), 5-10-2000, Goldblatt 11619 (NBG, holo.;

K, MO, PRE, S, iso.). Figure 14.

Annual herb 150-300 mm high, branching mainly

near base; stems sparsely leafy, very thinly cobwebbed at

first, later glabrescent. Leaves mainly basal but lower

soon withering, alternate, oblanceolate, 30-70 x 6-30

mm, narrowed to petiole-like base, leathery, thinly cob-

webbed at first, especially in axils, later glabrous or near-

ly so, margins and adaxial surface minutely papillate -

scabridulous, lower leaves especially with midrib more

conspicuously papillate-scabridulous abaxially and

decurrent on stems as softly scabridulous ridges; lower

leaves with expanded part oblanceolate to elliptic,

coarsely and sharply dentate, becoming more deeply

divided to pinnatifid upwards; upper leaves pinnatisect

with lobes oblong and coarsely serrate. Capitula het-

erogamous, radiate, solitary on long peduncles with few,

scattered, subulate bracts. Involucre broadly cup-shaped,

ecalyculate, 6-10 mm diam.; involucral bracts uniseriate,

10-13, narrowly oblong-ovate, acute, leathery and most-

ly with one or both margins distinctly subscarious, cili-

ate-puberulous at tips, 3-5-veined, 7-8 x 2-3 mm.

Receptacle Hat, punctate, glabrous. Ray florets female,

5-8; tube cylindrical, abaxial (outer) side densely ad-

pressed-hairy, 2. 5-3.0 mm long; lamina deflexed-spread-

ing, elliptic-oblong, ± 5-veined, 10-12 x 2. 5-3.0 mm,

yellow. Ovary narrowly obovoid, triangular in transverse

section, abaxial (outer) face largest, convex, densely

adpressed-hairy with apical hairs longest, inner faces

narrower, glabrous but separated by an adpressed-hairy

ridge; style terete with slightly swollen base on a distinct

stylopodium, branches narrowly oblong, 0.8 mm long,

lateral margins stigmatic, apices truncate and shortly

papillate. Cypselas heteromorphic, those of ray florets

narrowly ellipsoid, reddish brown, arcuate, triangular in

transverse section with outer face largest, convex and

densely covered with adpressed, acute, white, myxo-

genic hairs, inner faces narrower, flat or concave,

glabrous but intervening ridge also densely hairy, ± 5

mm long. Pappus bristles pluriseriate, numerous, white,

barbellate, connate at base into short collar, persistent,

4-5 mm long. Disc florets bisexual, numerous, ± 6 mm

long, glabrous, yellow; tube cylindrical, 3. 5^1.0 mm

long, limb obconical, 2.5 mm long, 5-lobed; lobes nar-

rowly triangular, 2.0 x 0.8 mm, with submarginal veins

and median resin duct. Anthers 2.5 mm long including

ovate, somewhat keeled apical appendage; anther base

obtuse, ecaudate; filament collar somewhat swollen

towards base. Ovary narrowly obovoid, densely

adpressed-hairy; ovary wall crystals rhomboidal; style

terete with swollen base on distinct stylopodium, style

branches with margins stigmatic, apices shortly conical

with central tuft of longer papillae, outer surface with

fringe of papillae at apex. Cypselas narrowly oblong, 4-

or 5-angled, black, densely covered with adpressed,

acute, white, myxogenic hairs, ± 5 mm long. Pappus

bristles pluriseriate, numerous, white, barbellate, con-

nate at base into short collar, persistent, 4-5 mm long.

Flowering time : September to November or early

December. Figure 15.

FIGURE 14. — Distribution of Emilia hantamensis in Northern Cape.

FIGURE 15. — Emilia hantamensis. A, whole plant. B E, ray floret: B, floret; C, style; D, stigmatic branches; E, cypsela (inner face). F-J, disc

floret: F, floret; G, two stamens; H, style; I, stigmatic branches; J, cypsela. Scale bars: A, 10 mm; B, C, E-G, I, J, 2 mm; D, H, 1 mm.

Artist: John Manning.

Distribution and biology, the species is restricted to

clay soils derived from dolerite and is at present known

only from the dolerite outcrops just east ofNieuwoudt-

ville in the Northern Cape (Figure 14). This region,

which marks the westernmost limit of the dolerite silk is

known locally as the Flantam (Manning & Goldblatt

1997) and its doleritic soils become very glutinous and

spongy in the wet season, drying out in the summer.

This habitat supports several other local edaphic

endemics, especially geophytes and annuals. Among the

annuals which are restricted to doleritic clay soils along

the Bokkeveld and Roggeveld escarpment are

Heliophila collina (Brassicaceae), Diascia cardiose-

pala, Alonsoa unilabiata and Cromodon varicalyx

(Scrophulariaceae).

History, first collected by botanists Dee Snijman and

Pauline Perry as part of their botanical survey of the

Nieuwoudtville Wildflower Reserve, E. hantamensis was

initially thought to be an undescribed species of Othonna

(Snijman & Perry 1987). This opinion was revised when

material that was sent to Bertil Nordenstam of the

Swedish Museum of Natural History, an expert in the

genera of the tribe Senecioneae, was returned as a new

species of Emilia. The species was again collected in

1996, when it was photographed for inclusion in a

48

Bothalia 31,1 (5001)

regional wildflower guide (Manning & Goldblatt 1997),

at which time its status was once again confirmed by

Nordenstam.

Diagnosis and relationships', this distinctive species is

the first record of Emilia in the winter rainfall region of

South Africa, but it has all the essential morphological

features of the genus (Bremer 1994) and there seems to

be little doubt that it is correctly placed in that genus.

Emilia hantamensis is distinguished from all other south-

ern African species by its large, radiate capitula with the

involucre 6-10 mm diam. The southern African species,

which are distributed across the continent from Namibia,

through Botswana and into KwaZulu-Natal, have discoid

capitula. The other African species with radiate capitula

typically have the involucre no more than 5 mm diam. E.

hantamensis appears to belong in section Spathulatae on

account of its radiate, yellow-flowered capitula and

exappendiculate style branches, although the rather nar-

row corolla lobes of the disc florets are not typical of the

section. No other species of sect. Spathulatae are known

further south than Zimbabwe and the location of E. han-

tamensis in the Northern Cape represents a most unusu-

al disjunction.

Other material examined

NORTHERN CAPE. — 3119 (Calvinia): Nieuwoudtville Dist., Nie-

woudtville Reserve, flats in spongy red clay soil, 8-09-1983, (-AC),

Perry & Snijman 2354 (NBG); Farm Charlies Hoek, abundant in red,

spongy doleritic soil, (-AC), 13-10-1996, Snijman 1564 (NBG); dole-

rite outcrops just east of town, (-AC), 2-12-1996, Manning s.n. (NBG).

REFERENCES

BREMER, K. 1 994. Asteraceae. Cladistics and classification. Timber

Press.

JEFFREY, C. 1986. The Senecioneae in East tropical Africa. Kew Bul-

letin 41: 873-943.

MANNING, J. & GOLDBLATT, P. 1997. Nieuwoudtville. South Afri-

can Wildflower Guide 9. Botanical Society of South Africa.

NORDENSTAM, B. 1978. Taxonomic studies in the tribe Senecioneae

(Compositae). Opera Botanica 44: 1-84.

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

ville Wild Flower Reserve, northwestern Cape. South African

Journal of Botany 53: 445^154.

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

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

7735 Claremont, Cape Town.

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

den, PO Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2000-10-23.

FOSSOMBRONIACEAE

FOSSOMBRON1A NYIKAENS1S, A NEW SPECIES FROM MALAWI

Fossombronia nyikaensis Perold , sp. nov.

Plantae aggregatae, plerumque smaragdinae, medioc-

res vel robustulae, dioicae. Folia supra subrotundata,

saepe plicata, plerumque brevioria quam lata. Plantae

masculae rarissimae, plerumque quam feminae minores.

Antheridia a bracteis perigonialibus forma irregulari pro-

tecta. Archegonia raro etiam bractea versiformi protecta.

Pseudoperianthium irregulariter 4-lobatum, substipita-

tum, folios proximos fere aequans. Sporae 27.5-37.5 pm

diametro, superficie distali cum 16 vel 17 papillis vel

cristis papillosis transversis; superficie proximali cum

papillis multis grossis irregularibus, inter eas tenuiter

granulata, ± 60 papillis brevibus e superficie distali cir-

cum margines procurrentibus. Elateres 107.5-150.0 pm

longi, 7.5-10.0 pm lati medio, apices versus angustati,

omnino bispirales vel partim bispirales, partimque trispi-

rales.

TYPE. — Malawi, 1033: Nyika National Park, south-

ern circular route at Chelinda Bridge, (-DB), on soil

under rock overhang, at altitude 2 227 m, Koekemoer

1792 (PRE, holo.), with Calycutaria crispula Mitt, and

Fissidens sp.

Plants in close stands, clear green, some leaves with

yellowed margins, others tinged with red, medium-sized

to fairly robust; male plants very rare; shoots smaller, up

to 5 mm long, ± 1 .5 mm high, 1 .8-2.0 mm wide; female

plants larger, 9.5-14.0 mm long, 1.6 mm high, 2. 8-3.0

mm wide; mostly simple, occasionally apically furcate.

Stems prostrate, old stems giving rise to new growth

from their apices, chlorophyllose, with globose nodules

near base, or with short, + 1 mm long, fleshy branches

with young leaves arising laterally near apices or bases;

in cross section plano-convex, in male plants apically

(Figure 16M) 260 pm (8 cell rows) high, 400 pm wide,

basally 300 pm high, 300 pm wide; in female plants api-

cally (Figure 16N) 240 pm (9 cell rows) high, 350 pm

wide, at base (Figure 160) 270 pm high, 240 pm wide.

Rhizoids purple, 12.5-17.5 pm wide. Leaves suberect to

spreading, overlapping, undulating, frequently plicate,

somewhat rounded above, succubously inserted on stem,

mostly shorter than wide, young apical leaves smaller,

margins with 3-8 sessile papillae, ± 15 x 22.5 pm, some-

times broken, basally brownish, wedged between 2 mar-

ginal cells (Figure 16L); in male plants (Figure 16A-D)

leaves smaller, 875-1250 x 1075-1325 pm; in female

plants (Figure 16E-K) leaves larger, 1175-1750 pm

long, 1150-2250 pm wide above, nanowing below to

1150-1500 pm. Leaf cells thin-walled, in male plants

rather smaller than in female plants, where at upper mar-

gins rectangular across or 5- or 6-sided, rarely isodia-

metric, 20-50 x 35.0-57.5 pm, at lower lateral margins

long-rectangular, 105-150 x 20-25 pm, upper laminal

cells 5- or 6-sided, 50.0-67.5 x 35.0-42.5 pm, middle

laminal cells 4— 7-sided, 62.5-100.0 x 37.5-50.0 pm,

basal cells 67.5-125.0 x 30.0^12.5 pm. Oil bodies no

longer present; chloroplasts numerous, round or oval, ±

3 pm diam.

Dioicous. Antheridia dorsal on stem, in I or 2 row(s),

short-stalked, globose, !200-210 pm diam., shielded by

irregularly shaped perigonial bracts (Figure 17A, B),

Bothalia 31,1 (2001)

49

FIGURE 16. — Fossombronia nyikaensis. A-D, male leaves; E-K, female leaves; L, detail of leaf margin with papilla; M, c/s male stem apex; N,

c/s female stem apex; O, c/s female stem base; P-S, perigonial bracts; T, pseudoperianth from side, showing outline of capsule within; U,

opened pseudoperianth; V, cells in inner capsule wall. A-V, Koekemoer 1792. Scale bars: A-K, T, U, 500 pm; L, 100 pm; M-O, P-S, 250

pm; V, 50 pm.

50

Bothalia 31,1 (2001)

FIGURE 17. — Fossombronia nyikaensis. A, simple male shoot with perigonial bracts; B, close-up of bracts; C, simple female shoot with archego-

nia and young pseudoperianth at ± midlength; D, close-up of same; E, mature pseudoperianth with lateral outgrowths, from side; F,

pseudoperianth and capsule from above. A-F, Koekemoer 1792. A, x 11.2; B, x 19; C, x 7.5; D, x 12.5; E, x 15.8; F, x 22.6.

either single, 25CM100 x 120-220 (jm, or, generally with

fused lobes (Figure 16P-S), 490-520 x 270^450 pm,

apical projections each topped by a papilla, marginal

cells 30.0-52.5 x 25.0-37.5 pm, cells in interior

52.5-67.5 x 30.0-32.5 pm. Archegonia dorsally along

stem (Figure 17C, D), in a row, spaced, or sometimes

crowded together, ± 250 pm long, rarely shielded by var-

iously shaped bract. Pseudoperianth (Figures 16T, U;

17E, F) usually less than 1 mm proximal to apex of

shoot, but sometimes near to midlength, irregularly

lobed, slightly raised on stalk, ± 375 pm long, almost

same length as adjoining leaves and 1250-1600 pm long,

± 1750 pm wide across mouth, lobes up to 675 pm wide,

with upper 500 pm or more free, marginally with 3 or 4

triangular processes, up to 130 pm long, some acutely

pointed, others short and basally wider, slit toward base

FIGURE 18. — Fossombronia nyikaensis. A-E, spores; F, elater. A, distal face; B, distal face partly from side; C, detail of part of distal face; D,

side view of distal face; E, proximal face. A-F, Koekemoer 1792. A, x 1200; B, x 1 105; C, x 2018; D, x 1265; E, x 1234; F, x 917.6.

Bothalia 31,1 (2001)

51

along side, with several lateral outgrowths, up to 600 pm

long, ceils not appreciably different in shape and size

from those in leaves. Capsules almost sessile, globose, ±

675 pm diam., wall bistratose, cells of inner layer

(Figure 16V) rectangular to polygonal, 35-40 x 25.0-

42.5 pm, each cell wall with 1 or 2 dark brown nodular

and occasional semi-annular thickenings. Spores light

brown, hemispherical, 27.5-37.5 pm diam.; distal face

(Figure 18A-D) convex, ornamentation papillose, with

16 or 17 small papillae and ridges across, papillae ± 2.5

pm long and +1.3 pm wide, some of them remaining

discrete, others linked together to form short or some-

times longer, straight or curved, rather uneven, low

ridges, less than 1.5 pm apart; proximal face (Figure

18E) slightly concave, lacking triradiate mark, with

numerous, coarse, irregular papillae and in between dust-

ed with fine granules, ± 60 low papillae from distal face

projecting around periphery. Elaters (Figure 18F) yel-

low, 107.5-150.0 pm long, 7.5-10.0 pm wide in centre,

tapering to tips and ending in a loop, bispiral throughout

or partly bispiral and partly trispiral.

DISCUSSION

Fossombronia nyikaensis has been named for the

locality where it was collected (Figure 19), and is known

only from the type specimen and another one without

spores, but still easily recognizable by the similarity

between the thalli of the two samples. These collections

were recently made, growing close together on the Nyika

Plateau, Malawi, by Ms Marinda Koekemoer, Curator of

the National Herbarium (PRE), on a Southern African

Botanical Diversity Network (SABONET) Expedition to

the region. Specimen Koekemoer 1792 was divided into

two samples: 1, most of the collection was placed in a

small paper bag and allowed to air dry; 2, a small living

sample with five or six almost ripe sporangia and adher-

ent soil was kept damp in a small plastic dish lined with

wet filter paper and covered with a lid. During trans-

portation over many kilometers of extremely rough dirt

roads, the soil and plants were rolled into small balls

inside the dishes. The capsules of this new Fossombronia

species had ripened by the time that the expedition had

returned to South Africa, and the spores could be studied.

Although female plants were fairly numerous in the

specimens, those with ripe capsules were few. The cap-

sules appeared to be almost sessile. Male plants were

very rare.

The unique ornamentation on the distal face of F.

nyikaensis spores, immediately distinguished it from

other African species with which it had been compared,

particularly those that had been collected by members of

the British Bryological Society (BBS) on a visit to

Mount Mulanje and elsewhere in Malawi, in 1991. The

spores from the latter specimens are reticulately orna-

mented or have lamellae; none are papillate. The orna-

mentation of F. nyikaensis spores is a little like that of F.

cerebriformis (Scott & Pike 1984) from Victoria,

Australia, but in the latter the lamellae are less densely

crowded and there are no discrete papillae. The spores of

F. nyikaensis are also much smaller than those of F. cere-

briformis, which are 51-90 pm diam. and dark brown,

with 30-50 peg-like spines around the margin. In F.

FIGURE 19. — Locality of F. nyikaensis in Malawi.

cerebriformis the leaves are 2^1-lobed and strongly

ruched, while the upper margin is crenulate with project-

ing cells and occasional papillae. The pseudoperianth has

a lightly lobed upper margin.

The Nyika Plateau is situated in northern Malawi,

southwest of the northernmost tip of Lake Malawi. It is a

massive granite and granitic gneiss upfaulted block. The

vegetation consists of Afromontane forest patches, large-

ly confined to valley head sites, that lie in a matrix of

grassland dominated by Themeda triandra (Meadows &

Linder 1993). The climate is influenced by the Tropical

Easterly Wind Belt. Rainfall comes with the monsoon,

wet conditions occurring from March to May and from

mid-October to December; dry conditions prevail from

December to March.

Specimens examined

Held at PRE.

Koekemoer 1786 (a), 1792 (holotype).

ACKNOWLEDGEMENTS

My sincere thanks to Ms M. Koekemoer for kindly col-

lecting bryophytes in Malawi, among which was this

new Fossombronia species. Dr H. Stieperaere is thanked

for his helpful suggestions and I am grateful to Dr H.F.

Glen for translating the diagnosis into Latin. I also

extend my gratitude to Ms G. Condy for the drawings, to

Mrs A. Romanowski for developing and printing the

photographs and to Ms D. Maree for typing the manu-

script. The BBS is thanked for the loan of Fossombronia

52

Bothalia 31,1 (2001)

species collected in Malawi, which had to be returned for

a monographic study of Fossombroniaceae by Stotler,

Crandall-Stotler and their students.

REFERENCES

MEADOWS, M.E. & LINDER, H.R 1993. A palaeoecological per-

spective on the origin of Afromontane grasslands. Journal of

Biogeograpliy 20: 345-355.

SCOTT, G.A.M. & PIKE, D.C. 1984. New species of Fossombronia

from Australia. Journal of the Hattori Botanical Society 56:

339-349.

S.M. PEROLD*

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

MS. received: 2000-10-02.

Bothalia 31.1: 53-70(2001)

Leaf anatomy of the genus Passerina (Thymelaeaceae): taxonomic and

ecological significance

C.L. BREDENKAMP* and A.E. VAN WYK**

Keywords: anatomy, epidermis, mesophyll, Passerina , sclerenchymatous sheath, southern Africa, Thymelaeaceae, vascular bundles

ABSTRACT

A comparative anatomical study was made of the genus Passerina comprising 20 species and four subspecies, most of

which are endemic to southern Africa. It showed that anatomical variation is useful in species recognition and classification.

Anatomical characters typical of Thymelaeaceae and displayed in Passerina include isobilateral leaves, a papillate cuticular

membrane, mucilaginous epidermal cell walls, a parenchymatous bundle sheath and extraxylary sclerenchyma fibres.

Vascular bundles of the leaf lack intraxylary phloem. Characters common to Passerina are inverse-dorsiventral and epis-

tomatic leaves, inverted palisade parenchyma and an abaxial hypodermal sclerenchymatous sheath. Orientation of the main

vascular bundle in relation to the epidermis and mesophyll allows the recognition of four leaf structural types and ten states,

according to which all species can be characterized and grouped. Functionally many anatomical features of the leaf in

Passerina are interpreted as adaptations to the Mediterranean climate of the Cape Floristic Region, where most species occur.

CONTENTS

Introduction 53

Material and methods

Light microscope (LM) studies 55

Scanning electron microscope (SEM) studies .... 55

Transmission electron microscope (TEM) studies . 55

Measurements of leaf in transverse section (t/s) . . 55

Terminology 56

Phylogeny 57

Results

Macromorphology of leaf 57

Leaf anatomy 57

Leaf structural types 63

Key to leaf structural types 63

Leaf structural type A 63

Leaf structural type B 63

Key to subtypes of leaf type B and relevant taxa 63

Leaf structural type C 64

Key to species 64

Leaf structural type D 64

Key to species 64

Comparative leaf anatomy at infrageneric level . . 64

Discussion and additional observations

Leaf structure 65

Epidermal tissue 65

Mesophyll 65

Leaf structural types: orientation and structure of

main vascular bundle in relation to epidermis

and mesophyll 67

Sclerenchyma 68

Less important taxonomic characters 68

Phylogenetic considerations 69

Taxonomic significance 69

Conclusions 70

Acknowledgements 70

References 70

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

** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University

of Pretoria, 0002 Pretoria.

MS. received: 2000-05-09.

INTRODUCTION

This paper emanates from a comparative leaf anatomi-

cal survey of the genus Passerina L., undertaken as part

of a monographic study of the group. Work undertaken

thus far has revealed at least four new species and four

new subspecies, to be added to the 16 existing species, of

which most species are endemic to southern Africa

(Thoday 1924; Bond & Goldblatt 1984). Despite the now

outdated revision by Thoday (1924), boundaries of infra-

generic taxa in Passerina remain a problem, mainly

owing to the apparent lack of marked morphological dif-

ferences between the species. This paper explores the

importance of leaf anatomy in Passerina as a source of

potential taxonomic evidence.

Physiography and climate are important in the distri-

bution of Passerina (Table 1). Most species of Passerina

are endemic to the Cape Lloristic Region. The climate of

this region is mostly Mediterranean or semi-Mediterra-

nean. Winter rainfall occurs in the west and along the

south coast, complemented by some summer rain, which

increases eastwards. The western Karoo and Nama-

qualand (Succulent Karoo Biome) are characterized by

winter precipitation and summer drought. Only two

species, P. sp. nov. 4 and P. montana are distributed to

the east and north along the eastern mountains and east-

ern escarpment of southern Africa, areas that receive pre-

dominantly summer rainfall.

Anatomical research in Passerina was initiated by

Pick ( 1 882a, b), who studied the effect of light on the ori-

entation of assimilation tissue and discussed the inverse-

dorsiventral leaf. Gilg (1891) published a taxonomic

account on the Thymelaeaceae, indicating that the

anatomical structure of stems was similar throughout the

family and subsequently of no taxonomic value; he fol-

lowed Endlicher (1847) in distinguishing the subtribe

Passerininae on the basis of floral morphology. Van

Tieghem (1893) described the anatomy of the root, stem

and leaf for all the genera of the Thymelaeaceae known

54

Bothalia 31,1 (2001)

TABLE 1. — Distribution and habitat of Passerina species (order of taxa according to leaf structural type referred to throughout paper)

at the time and classified the family into three groups,

namely Drapetees, Thymelees and Aquilariees, with

Passerina in the Thymelees. In his key, the genera

Lachnaea L., Cryptadenia Meisn., Gnidia L. and

Chymococca Meisn. are grouped on the basis of gela-

tinized epidermal cells, whereas Passerina is distin-

guished by the absence of these cells. Subsequent work

by Bredenkamp & Van Wyk (1999, 2000) has shown the

presence of these cells in Passerina. Supprian (1894)

studied the stem and leaf anatomy of the Thymelaeaceae,

describing the epidermis (‘Hautsystem’), the mechanical

system (‘Mechanisches system’), the assimilation system

(‘Assimilationsystem’), the vascular structure (‘Leitungs-

system’), the aeration system (‘Durchliifdtungssystem’)

and excretion management (‘Excretbehalter’) for all the

known genera. He classified the family into the subfam-

ilies Aquilarioideae and Daphnoideae and placed

Passerina into the latter, under the tribe Euthymeleae.

Gilg (1894) studied the relationships in the Thyme-

laeaceae, using mainly floral anatomy, and classified the

family into the subfamilies Aquilarioideae, Phaleri-

oideae, Thymelaeoideae and Drapetoideae. Passerina is

classified in the subfamily Thymelaeoideae, tribe Daph-

neae, subtribe Passerininae. In the same article, he gave

a full account of the anatomical method applied by Van

Tieghem (1893) and Supprian (1894), concluding that

many of the characters used by them were not constant,

that there would always be criticism against the anatomi-

cal method and that floral characters were more reliable

in the delineation of the Thymelaeaceae.

Bothalia 31,1 (2001)

55

Solereder (1899, 1908) summarized the anatomical

work on the Thymelaeaceae up to the beginning of the

twentieth century. The most prominent subsequent

anatomical study in the Thymelaeaceae was done by

Leandri (1930), who delimited the Thymelaeoideae on

the basis of intraxylary phloem in the stem. He classified

Passerina in the tribe Daphneae, and because of the

absence of floral nectaries, into the subtribe ‘Passe-

riniinae’, thus combining anatomical and floral charac-

ters. Hereafter the most significant works on the

Thymelaeaceae comprised the compilation of all the

available knowledge. Domke (1934) proposed a widely

adopted classification system for the family based on

previous anatomical and floral morphological evidence,

classifying Passerina under the subfamily Thyme-

laeoideae, tribe Gnidieae and subtribe Passerininae.

Finally Metcalfe & Chalk (1950) and Metcalfe (1979,

1983) published accounts of the Thymelaeaceae in their

standard works on the anatomy of the dicotyledons.

With the genera in Thymelaeaceae delineated on the

basis of anatomy and floral morphology, the focus

changed to generic revisions. Anatomical work on

Passerina was done by Thoday (1921), who described

the structure and behaviour of the ericoid leaves of P. fil-

iformis L. and P. cf.falcifolia C.H. Wright under drought

conditions and supplied some notes on their anatomy.

Kugler (1928) described the inverse-dorsiventral leaves

of P. filiformis (= P. pectinata Hort.). Recent comparative

studies on Thymelaeaceae in southern Africa include

those on leaf anatomy of the genera Lachnaea and

Cryptadenia (Beyers 1992; Beyers & Van der Walt 1995)

and on leaf and involucral bract morphology of system-

atic use in Gnidia (Beaumont et al. 1994). Previous stud-

ies of leaf anatomy identified mucilagination of the epi-

dermal cells as being of possible taxonomic importance.

Recently Bredenkamp & Van Wyk (1999) clarified the

structure of these epidermal cells and the origin of the

mucilage, indicating that this character is of taxonomic

importance mainly at the family level. The present com-

parative anatomical study of the leaves in Passerina is

the most comprehensive to date. Our primary objective

has been to assess the infrageneric taxonomic signifi-

cance of leaf anatomy in the genus.

The most outstanding anatomical feature of the

Thymelaeaceae, namely the presence of sclerenchyma

fibres, may well be an adaptation of members of the fam-

ily to their environments. However, sclerenchyma fibres

in the leaves of Passerina have never been studied at infra-

generic level before, and this study meets that need. The

presence of tough sclerenchyma fibres in the stems of

Passerina , is well known among indigenous people, who

use especially the bark for making ropes, straps and whips.

The wide distribution of Passerina in the Cape

Floristic Region, the southern and eastern coastline of

South Africa and along the eastern escarpment of south-

ern Africa to Zimbabwe, and with outliers as far north as

Tanzania, provides an opportunity to study the possible

leaf anatomical adaptations of these plants to a wide

range of habitats, experiencing both Mediterranean and

summer rainfall conditions. For example, we suspect the

decreasing rainfall from the eastern escarpment to the

northwestern parts of the Northern Cape to be reflected

by adaptations in the leaf structure of the group. The pre-

sent paper provides a description of anatomical charac-

ters in Passerina as well as an assessment of their taxo-

nomic and ecological significance.

MATERIAL AND METHODS

Leaves, both fresh and from herbarium material, of all

infrageneric taxa in Passerina were studied. Additional

Passerina specimens examined since Bredenkamp &

Van Wyk (2000: 70) are listed in Table 2. Leaves from

herbarium material were rehydrated in water for 5 min-

utes at boiling point. All leaf material was fixed and

stored in a 0. 1 M phosphate-buffered solution at pH 7.4,

containing 2.5% formaldehyde, 0.1% glutaraldehyde and

0.5% caffeine [modified Kamovsky fixative; Kamovsky

(1965)]. Whenever possible, material from at least five

different localities was included.

Light microscope (LM) studies

The light microscope was used for general leaf anato-

my and epidermal studies. Unless otherwise stated, the

tenth leaf from the growing point of a twig was used in

all comparative studies. To prepare semi-thin transverse

sections, a 1 mm wide segment of leaf material was cut

from the centre of each leaf, thus including the main vein

as well as both leaf margins. Samples were dehydrated,

embedded in glycol methacrylate (GMA) and sectioned

according to the methods of Feder & O’Brien (1968).

Sections were stained in toluidine blue ‘O’, subjected to

the periodic acid-Schiff’s (PAS) reaction and mounted in

Entellan (Art. 7961, E. Merck, Darmstadt).

The following three methods were followed in the

study of the cuticles (Bredenkamp & Van Wyk 2000):

1 . GMA sections of leaves in transverse section were

stained with 1% Sudan Black B dissolved in 70% ethanol.

2. Macerated cuticular mounts were stained with a 1%

aqueous safranin solution.

3. Epidermal mounts, obtained by removing small

pieces of ad- and abaxial epidermis manually and by

making paradermal hand sections, were stained with 1%

safranin dissolved in 50% ethanol.

Scanning electron microscope (SEM) studies

The scanning electron microscope was used to study

the epidermal surface features (including epicuticular

waxes) and to verify the structure of the cuticle (Bre-

denkamp & Van Wyk 2000).

Transmission eclectron microscope (TEM) studies

The transmission electron microscope was used for

the study of the structure of mucilaginous epidermal cell

walls in Passerina (Bredenkamp & Van Wyk 1999).

Measurements of leaf in transverse section (t/s)

Using LM, all measurements were done by using a

calibrated eyepiece.

56

Bothalia 31,1 (2001)

TABLE 2. — Additional Passerina specimens examined and taxonomical changes made since Bredenkamp & Van Wyk (2000: 70)

Species

Voucher specimen Locality

comosa

filiformis subsp. filiformis

filiformis subsp. nov.

glomerata subsp. glomerata

glome rata subsp. nov.

sp. nov. 1

sp. nov. 3

sp. nov. 4

Thoday 212 *

Boucher 2833'

Bredenkamp 1039

Schlechter 5125'

Taylor 1542*

Bredenkamp 988*

Bredenkamp 994

Bredenkamp 1002

Bredenkamp 984, 985

Bredenkamp 97T

Bredenkamp 973

Stokoe 8040*

Esterhuysen 2858T

Goldblatt & Manning 8627

Stokoe 9302'

Schlechter 5846’

Esterhuysen 10734’

Esterhuysen 28006 *

Killick 238

Bredenkamp 1016 \ 1017*

Van Wyk & Bredenkamp 7,

1012 '

WESTERN CAPE. — 3320 (Montagu): Montagu Dist.. near Concordia, (-CD).

WESTERN CAPE. — 3118 (Van Rhynsdorp): Clanwilliam Dist., Diepkloof S of

Verlorevlei, (-AD).

WESTERN CAPE. — 3318 (Cape Town): Signal Hill, (-CD).

WESTERN CAPE. — 3218 (Clanwilliam): Alexander's Hoek, (-BC).

WESTERN CAPE. — 3218 (Clanwilliam): Malmesbury Div., Vredenburg,

Steenberg’s Cove, (-CC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains, Eikeboom, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains, Kromrivier, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains, Algeria, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Citrusdal, Piekenierskloof Pass, (-CD).

WESTERN CAPE. — 3219 (Wuppertal): Groenfontein, (-DC).

WESTERN CAPE.— 3319 (Worcester): Tulbagh, (-AC).

WESTERN CAPE. — 3219 (Wuppertal): Cederberg Mountains.

WESTERN CAPE. — 3319 (Worcester): Hex River Mountains.

WESTERN CAPE. — 3220 (Sutherland): Roggeveld Escarpment. (-AB).

WESTERN CAPE. — 3322 (Oudtshoorn): Swartberg Pass, Prince Albert area, (-AC).

WESTERN CAPE. — 3322 (Oudtshoorn): Montagu Pass, (-CD).

EASTERN CAPE. — 3323 (Willowmore): Kouga Mountains, (-DA).

EASTERN CAPE. — 3324 (Steytlerville): Cockscomb, Uitenhage area, (-BD).

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Table Mountain, (-CB).

KWAZULU-NATAL.— 3030 (Port Shepstone): Oribi Gorge. (-CB).

KWAZULU-NATAL. — 3130 (Port Edward): Umtamvuna River Bridge, (-AA).

Bredenkamp 1327

Gillett 4537

Keet s.n.

KWAZULU-NATAL. — 3130 (Port Edward): Mkambati Nature Reserve, (-AC).

WESTERN CAPE. — 3422 (Mossel Bay): en route to Knysna from George, (-BB).

WESTERN CAPE.— 3423 (Knysna): Knysna, Redlands, (-AA).

* Material used for the SEM study of the ad- and abaxial epidermis.

All specimens are housed at PRE.

Thickness of the main vascular bundle

The shortest distance between the outermost points of

the ad- and abaxial epidermis through the main vascular

bundle, was measured.

Leaf width

Measurements from leaf margin to leaf margin would

be unreliable considering the cymbiform shape of the

leaf, the involute nature of the leaf margins, and the fact

that leaf margins correspond to changes in turgor pres-

sure. Leaf width was therefore measured as the shortest

distance between the outer epidermal walls of the abaxi-

al epidermis, parallel to the adaxial epidermis, but per-

pendicular to the axis of the main vascular bundle.

Terminology

Epidermal structure

Epidermal structure was described by Bredenkamp &

Van Wyk (1999, 2000).

Cuticle

Following Jeffree (1986), we distinguish the cuticle

proper, the cuticular layer and the cell wall.

Cuticular ornamentation ( LM and SEM)

We follow Wilkinson (1979) in our choice of termi-

nology.

Epicuticular wax

The recognition of soft waxes is based on the criteria

proposed by Amelunxen et al. (1967). Further interpreta-

tion of epicuticular waxes and crystals was done accord-

ing to Wilkinson (1979) and Barthlott et al. (1998).

Leaf anatomy

The definitions of dorsiventral and isobilateral leaves

(Eseau 1965; Metcalfe 1979; Fahn 1982; Mauseth 1988)

are accepted in the present study. However, we follow

Kugler (1928) who coined the term ‘inverse-dorsiven-

tral’ for leaves where the palisade parenchyma develops

abaxially, because the abaxial epidermis is exposed to

the environment. In addition to leaf symmetry, the works

of the above-mentioned authors as well as those of Sole-

reder (1908) and Metcalfe & Chalk (1950) have been

used for the interpretation of leaf anatomy.

Crystals

Crystals were interpreted according to Metcalfe

(1983). Unstained GMA sections of leaves were used to

Bothalia 31,1 (2001)

57

identify the presence and position of crystals by means of

polarized light, after which they were tested for the pres-

ence of flavonoid glycosides, such as diosmin, according

to the method of Jackson & Snowdon (1990). The iden-

tity of the crystals was finally confirmed using energy

dispersive spectrometry (EDS).

Phylogeny

Speculations on phylogeny are based on the prevail-

ing family characters representing the ancestral state and

derived characters at genus and species levels considered

as possibly advanced. A cladistic analysis based on

anatomical and morphological characters in Passerina is

scheduled for the final stages of the study.

RESULTS

Macromorphology of leaf

Leaf arrangement decussate, sometimes imbricate,

closely adpressed to stem or spreading at an angle of

5°-20°(-60°); spreading of leaves often prominent in

juvenile plants. Lamina inversely ericoid; adaxial surface

concave, villous, often forming a groove facing the stem;

abaxial surface convex, orientated more or less acroscop-

ically, thus exposing a large surface area to the environ-

ment; cuticle often amber-coloured (in herbarium materi-

al) and outline of epidermal cells often macroscopically

visible. Leaf shape cymbiform, falcate or cigar-shaped;

plane shape linear, oblong, lanceolate, or narrowly trul-

late. Leaf base sessile or cuneate. Leaf apex truncate and

hump-backed, obtuse, rounded, acuminate or acute to al-

most spine-tipped. Margins sometimes ciliate. Size (1.5-)

2.5-4.0(-8) x (0.8— )1 .2— 2.0(— 3.0) mm. Figure 1.

Leaf anatomy

Trichomes

Adaxial surface of leaf villous, with uniseriate tri-

chomes forming a felty layer over adaxial epidermis; tri-

chomes bordering leaf margin often conspicuous.

Abaxial surface of leaves mostly glabrous, young leaves

tomentose to sparsely hairy in a few species (Table 3;

Figure 2A).

Epidermis

Lamina epistomatic. Adaxial epidermis uniseriate,

following concave leaf surface; stomata anomocytic;

cuticular membrane 2-5 pm thick; epidermal cells irreg-

ularly shaped, outer periclinal wall convex in t/s, polyg-

onal in surface view. Abaxial epidermis uniseriate, con-

vex (following leaf surface curvature), glabrous or

sparsely hairy; cuticular membrane well developed,

( 10-)20-30(-70) pm thick in t/s; epicuticular waxes

coating entire surface, crystalloids, wax platelets and

plates present or absent (Figure 2B-D). Anatomical

structure and taxonomic value of abaxial epidermis are

discussed by Bredenkamp & Van Wyk (2000), and cor-

related with leaf structural types in Table 3.

TABLE 3. — Correlation between leaf structural type and epidermal characters (Bredenkamp & Van Wyk 2000) in Passerina

58

Bothalia 31,1 (2001)

FIGURE 1. — Camera lucida drawings of leaves of Passerina species arranged according to leaf structural type (Table 3), correlating leaf shape to

outline of leaves in t/s: A, P. sp. nov. 1, Bredenkamp 1044: B, P. burchellii , Bolus 684: C, P. pendula, Bredenkamp 908 ; D, P. comosa,

Andreae 1288: E, P. paludosa , Bredenkamp 1035 ; F, P. galpinii, Bredenkamp 946: G, P. drakensbergensis, Bredenkamp 1019: H. P. eri-

coides , Bredenkamp 962 ; 1, P. sp. nov. 2, Esterhuysen 26859: J, P. sp. nov 3, Esterhuysen 28006: K, P. rubra, Bredenkamp 914: L, P.fdi-

formis subsp .fdiformis, Bredenkamp 896: M, P. obtusifolia, Bredenkamp 919: N, P. glomerata subsp. glomerata, Bredenkamp 984: O. P.

falcifolia, Bredenkamp 917: P, P sp. nov. 4, Bredenkamp 1016: Q, P. montana, Bredenkamp 889: R, P. paleacea, Bredenkamp 960: S, P.

rigida, Bredenkamp 911: T, P. vulgaris, Bredenkamp 901 . Leaf size A-T x 20. Scale bar: 10 mm; leaves in t/s not to scale.

Tanniniferous substances

Tanniniferous substances (Figure 2C, D) present in

ad- and abaxial epidermis, mesophyll, bundle sheaths,

parenchyma abaxial of vascular tissue in vascular bun-

dle, staining homogenously or with vesicular appear-

ance.

Crystals

Calcium oxalate crystals (Figure 2E, F) present in mes-

ophyll, clustered crystals (druses) in parenchyma, frag-

mented calcium oxalate crystals, resembling crystal sand,

present in intercellular spaces; flavonoid glycosides (dios-

min) absent.

Bothalia 31.1 (2001)

59

FIGURE 2. — A, B, SEM micrographs of trichomes and epicuticular waxes: A, P. falcifolia, Bredenkamp 915; B, P. rigida, Bredenkamp 1013. C-H,

LM photographs of leaf: C, P. galpinii, Bredenkamp 946; D, P. pendula. Bredenkamp 909, stomatal apparatus in adaxial epidermis stained

with Sudan Black B; E, P. paleacea, Pillans 3783, calcium oxalate crystals and druses in polarized light; F, P. falcifolia, Tyson 1449, cluster

crystal from ruptured palisade parenchyma in polarized light; G, P. drakensbergensis, Bredenkamp 1019, mesophyll in leaf margin; H, P.

glomerata subsp. glomerata, Bredenkamp 977, mesophyll and tracheid in leaf margin, ab, abaxial epidermis; ad, adaxial epidermis; bs, bun-

dle sheath; c, cambial cells; CM, cuticular membrane; co, calcium oxalate crystals; cy, coOenchyma; d, druse; e, epidermal cell; ef, extraxy-

lary sclerenchyma fibres; gc, guard cell; il, inner ledge; m, mucilage; mb, median vascular bundle; ol, outer ledge; ph, phloem; pi, plates; pp,

palisade parenchyma; pr, peristomatal rim; ps, platelets; s, sclerenchymatous hypodermal sheath; sb, secondary vascular bundle; sp, spongy

parenchyma; st, stomata; t, trichome; ta, tanniniferous compounds; tr, tracheid; X, xylem. Scale bars: A, B, D, 10 pm; C, E-H, 100 pm.

60

Bothalia 31,1 (2001)

FIGURE 3. — LM photographs of leaf structural types A and B. Type A: A, P. sp. nov. 1 , Bredenkamp 1044. Type B 1 : B, P. pendula, Bredenkamp

90S. Type B2: C, P. paludosa, Bredenkamp 1035. Type B3: D, P. ericoides , Bredenkamp 962. Type B4: E, P.fdifprmis subsp .filiformis,

Bredenkamp 1039. Type B5: F, P. glomerata subsp. glome rata, Bredenkamp 984 , with secondary tissue in vascular bundle; G, P. obtusi-

folia, Bredenkamp 919. Type B6: H, P. rigida, Bredenkamp 1013. Abbreviations as for Figure 2. Scale bars: A-H, 100 pm.

Bothalia 31,1 (2001)

61

FIGURE 4. — LM photographs showing leaf structural types C and D. Type C: A, P . falcifolia, Bredenkamp 917\ B, P. sp. nov. 4, Bredenkamp

1016. Type Dl: C, P. montana. Bredenkamp 889. Type D2: D. P. paleacea, Bredenkamp 960\ E, P. rigida, Bredenkamp 962\ F, P. vul-

garis, Bredenkamp 901. Abbreviations as for Figure 2. Scale bars: A-F, 100 pm.

Leaf structure, mesophyll, vascular and sclerenchvma

tissue in t/s

Leaf isobilateral or inverse-dorsiventral. Outline

variable, narrowly transversely elliptic in flatter leaves,

transversely elliptic (Figures IF; 3A; 4C, E) or trans-

versely oblong (Figure 1R) in cymbiform leaves, also

canaliculate (Figures 1G; 3H) or carinate (Figures

lO-P; 4A, B, F); adaxial epidermis slightly concave in

flatter leaves or leaf lamina and margins strongly

upturned to involute forming a central groove or fur-

row. Margins filled with palisade parenchyma extend-

ing to adaxial epidermis, abruptly becoming irregular

adaxially, conforming to shape and size of spongy

parenchyma (Figure 2G); terminal vein endings often

present, ultimately consisting of a single tracheid

(Figure 2H). Width (570-)880(-1480) pm. Midrib often

raised below owing to supporting sclerenchyma fibres,

cymbiform leaf folding along reinforced midrib; thick-

ness of main vein (260-)440(-560) pm. Mesophyll pal-

isade-like and homogeneous or inverted — spongy

parenchyma situated adaxially and palisade parenchyma

abaxially. Palisade parenchyma horseshoe-, U- or V-

shaped, 1- or 2-layered, or 2- or 3-layered, (3)4 or 5(6)

cells per 50 pm; cells narrowly elliptic to elliptic in iso-

bilateral leaves or elongated in inverse-dorsiventral

leaves, containing chloroplasts, tanniniferous deposits

and druse crystals. Spongy parenchyma in isobilateral

leaves ± uniform in shape, cells narrowly elliptic to

elliptic, densely arranged with larger intercellular

spaces in centre of leaf, aerenchymatic, meso- or xero-

morphic in inverse-dorsiventral leaves, often resem-

bling palisade parenchyma adaxial to veins, rounded,

pentagonal or heptagonal with lobes connecting neigh-

bouring cells; cells either loosely arranged with large

intercellular spaces or densely arranged with small.

62

Bothalia 31,1 (2001)

Key to tissues

Chlorenchyma

Spongy parenchyma

Palisade parenchyma

Parenchyma

Xylem

Phloem

Sclerenchyma

Epidermis

Cuticular membrane

Collenchyma

FIGURE 5. — Diagrammatic representation of leaf structural types A, B, C and D. also indicating increasing xeromorphic gradient within types B

and D, as well as for type B to D.

centrally orientated intercellular spaces in xeromorphic

leaves; elongated cells connecting bundles laterally,

constituting a paraveinal mesophyll; chloroplasts and

tanniniferous deposits present; calcium oxalate crystals

present in crystalliferous cells (presence in intercellular

spaces possibly due to processing) (Figure 2E, F). Main

vascular bundle centrally arranged in isobilateral

leaves (Figure 3A), variously orientated in relation to

mesophyll of inverse-dorsiventral leaves (Figures 3-5).

Shape ovate, widely ovate, very widely ovate, oblate or

Bothalia 31,1 (2001)

63

obovate. Bundle sheath completely enveloping vascular

bundle (Figure 3A-H) or present adaxially only (Figure

4A-F); number of cells ( 1 2—) 14— 28(— 32), mostly one

layer of parenchymatous cells or irregularly two-lay-

ered; cells larger than other tissues in vascular bundle,

roundish or often longitudinally or transversely elon-

gated; tanniniferous deposits ample. Vascular tissue

collateral with adaxial xylem and abaxial phloem, sur-

rounded by a layer of colourless parenchymatous cells,

becoming tanniniferous or sclerenchymatous abaxially,

separating vascular tissue from abaxial sclerenchyma

fibres. Xylem arranged in a ± semilunar band or in ± 7

radial tiers, alternating with xylem parenchyma

(Figures 2C; 3F-G; 4A, F), often separated from

phloem by thin-walled parenchyma cells. Phloem

arranged in shallow band, sieve tubes and companion

cells interspersed with phloem parenchyma. Secondary

growth indicated by cambial cells and thin-walled

derivatives arranged in rows between secondary xylem

and secondary phloem; primary xylem situated in most

adaxial position between large parenchymatous cells

and primary phloem bordering on sclerenchyma fibres,

conspicuous in P. glomerata subsp. glomerata (Figure

3F). Extraxylary fibres bordering phloem tissue abaxi-

ally, ± separated by irregular layer of parenchyma often

containing tanniniferous substances and becoming scle-

renchymatous (Figures 2C; 3F, H); enclosed in bundle

sheath or extending beyond bundle sheath and palisade

parenchyma up to abaxial epidermis. Secondary vascu-

lar bundles (5)6-1 3(-l 9) corresponding to main vascu-

lar bundles in orientation, shape, bundle sheath and

arrangement of vascular tissue. Sclerenchymatous

hypodermal sheath formed by sclerenchyma fibres

extending paradermally, often up to leaf margins (‘wan-

dering’ fibres sensu Thoday 1921), connecting with

fibres from main and secondary vascular bundles in

P. paleacea , P. rigida , and P. vulgaris (Figures 4D-F;

5); sclerenchyma fibres in mesophyll (‘Spicularzellen’

sensu Supprian 1894) absent.

Leaf structural types

The isobilateral and inverse-dorsiventral states of

the leaf dictate the orientation of the main vascular

bundle. The main vascular bundle is central in isobilat-

eral leaves (Figure 3A), and either close to the adaxial

epidermis, situated centrally, or variously arranged in

relation to the abaxial epidermis in inverse-dorsiven-

tral leaves (Figure 5). In this transformation series,

four leaf structural types and ten states are identified

(Tables 3; 4).

Key to leaf structural types

la Leaf isobilateral type A (Figures 3 A; 5 A)

lb Leaf inverse-dorsiventral:

2a Bundle sheath completely enveloping main vascular bun-

dle, extraxylary sclerenchyma fibres enclosed in bun-

dle sheath type B (Figures 3B-H; 5B)

2b Bundle sheath capping main vascular bundle adaxially, ±

absent abaxially, extraxylary sclerenchyma fibres not

enclosed in bundle sheath:

3a Main vascular bundle bordering on palisade parenchy-

ma, extraxylary sclerenchyma fibres fitting into V-

shaped palisade parenchyma . . . type C (Figures 4 A, B; 5C)

3b Main vascular bundle extending beyond palisade paren-

chyma, extraxylary sclerenchyma fibres in contact

with abaxial epidermis:

4a Sclerenchymatous hypodermal sheath absent

type D1 (Figures 4C; 5D)

4b Sclerenchymatous hypodermal sheath present

type D2 (Figures 4D-F; 5D)

Leaf structural type A

Leaf isobilateral. Main vascular bundle central; bun-

dle sheath completely enclosing vascular tissue. Second-

ary vascular bundles close to one another, with bundle

sheaths adhering, forming a central plate of veins.

Sclerenchymatous hypodermal sheath absent (Tables 3;

4; Figures 3A; 5A). Species represented; P. sp. nov. 1

(Figure 3A).

Leaf structural type B

Leaf inverse-dorsiventral. Palisade parenchyma

horseshoe-shaped. Main vascular bundle variously orien-

tated in relation to mesophyll: adhering to adaxial epider-

mis (type Bl), centrally arranged with strands of spongy

parenchyma (type B2), touching palisade parenchyma

abaxially (type B3), sunken into palisade parenchyma

(type B4), extending beyond palisade parenchyma with

collenchyma wedged between main vascular bundle and

abaxial epidermis (type B5) or adhering to abaxial epi-

dermis (type B6). Shape ovate, widely ovate, very wide-

ly ovate, oblate or obovate. Bundle sheath completely en-

veloping main and secondary vascular bundles. Extra-

xylary sclerenchyma fibres enclosed in bundle sheath.

Sclerenchymatous hypodermal sheath absent (Tables 3; 4;

Figures 3B-H; 5B).

Key to subtypes of leaf type B and relevant taxa

la Main vascular bundle close to or adhering to adaxial epidermis type Bl : P. burchellii, P. pendula (Figure 3B)

lb Main vascular bundle central or abaxial:

2a Main vascular bundle central . . . type B2:

3a Mesophyll inverse-dorsiventral, spongy parenchyma more homogenous and palisade-like:

4a Main vascular bundle surrounded by aerenchymatic spongy parenchyma, strands of spongy parenchyma connect-

ing main vascular bundle to adaxial epidermis, palisade parenchyma and secondary vascular bundles

P. paludosa (Figure 3C)

4b Main vascular bundle surrounded by mesomorphic spongy parenchyma, secondary vascular bundles closely arranged

P. comosa

3b Mesophyll inverse-dorsiventral, palisade and spongy parenchyma clearly distinguished:

5a Mesophyll xeromorphic P. obtusifolia , P. glomerata subsp. glomerata

5b Mesophyll aerenchymatic or mesomorphic:

6a Outline of leaf in t/s transversely elliptic P. galpinii (Figure 2C), P. montana (Figure IQ)

6b Outline of leaf in t/s transversely oblong P. paleacea (Figure 1R)

2b Main vascular bundle abaxially arranged or interfering with palisade parenchyma:

64

Bothalia 31,1 (2001)

7a Main vascular bundle touching palisade parenchyma abaxially . . . type B3:

8a Abaxial epidermal cells large, strongly mucilaginous, periclinal x anticlinal dimensions in t/s 30-65 x (-35)55-60

(-70) pm:

9a Palisade parenchyma U-shaped P. paleacea

9b Palisade parenchyma horseshoe-shaped . . . P. galpinii , P. drakensbergensis , P. ericoides (Figure 3D), P. rigida, P. montana

8b Abaxial epidermal cells exceptionally large, abundantly tanniniferous, mucilagination minimal, periclinal x anti-

clinal dimensions in t/s (20— )30— 45(— 50) x (25-)30-75(-105) pm:

10a Outline of leaf in t/s transversely elliptic to cordiform

. . P. obtusifolia (Figure 1M), P. glomerata subsp. glomerata (Figure IN), P.filiformis subsp .filiformis (Figures 1L)

10b Outline of leaf in t/s not transversely elliptic, depressed obovate or canaliculate respectively

P. sp. nov. 2 (Figure II), P. sp. nov. 3 (Figure 1J)

7b Main vascular bundle sunken into palisade parenchyma, causing specialization of, or extending beyond palisade

parenchyma:

11a Palisade parenchyma indented because of sunken main vascular bundle . . . type B4:

12a Abaxial epidermal cells strongly mucilaginous, periclinal x anticlinal diam. in t/s 30-65 x 45-60 pm:

13a Leaf outline in t/s canaliculate; secondary vascular bundles ± 6 on each side of main bundle . . . P. rubra (Figure IK)

13b Leaf outline in t/s transversely oblong, margins erect; secondary vascular bundles 3 or 4 on each side of main

bundle P. paleacea (Figure 1R)

12b Abaxial epidermal cells strongly tanniniferous, periclinal x anticlinal cell diam. in t/s (35-)40-45 x 45-75

(-105) pm P. filiformis subsp. filiformis (Figure 3E), P.filiformis subsp. nov., P. obtusifolia

lib Palisade parenchyma abaxial of main vascular bundle specialized, or main vascular bundle extending beyond pal-

isade parenchyma:

14a Main vascular bundle separated from abaxial epidermis by collenchyma containing ample amounts of tanninife-

rous substances type B5: P. glomerata subsp. glomerata (Figure 3F), P. obtusifolia (Figure 3G)

14b Main vascular bundle extending beyond palisade parenchyma, bordering on abaxial epidermis, abaxial cells of

vascular bundle sheath often collenchymatous . . . type B6:

15a Main vascular bundle abaxially orientated, shape ovate to very widely ovate

P. filiformis subsp. nov., P. glomerata subsp. nov.

15b Main vascular bundle exceptionally large, situated close to adaxial epidermis, reaching and touching abaxial

epidermis, shape elliptic P. rigida (Figure 3H)

Leaf structural type C

Leaf inverse-dorsiventral. Palisade parenchyma V-

shaped. Main vascular bundle bordering on palisade

parenchyma abaxially; shape obovate; bundle sheath 1 or

2 layers of parenchymatous cells capping main vascular

bundle adaxially, ± absent abaxially; cells rounded or

longitudinally lobed, containing ample amounts of tan-

niniferous substances. Extraxylary sclerenchyma fibres

not enclosed in bundle sheath, bordering on and fitting

into the V-shaped palisade parenchyma. Sclerenchyma-

tous hypodermal sheath absent (Figures 4A, B; 5C).

Key to species

la Bundle sheath cells longitudinally lobed, radiating outwards

adaxially, containing ample amounts of tanniniferous

substances Pfalcifolia (Figure 4A)

lb Bundle sheath cells rounded P. sp. nov. 4 (Figure 4B)

Leaf structural type D

Leaf inverse-dorsiventral. Palisade parenchyma

horseshoe-, U- or V-shaped. Main vascular bundle

extending beyond palisade parenchyma, ultimately in

contact with abaxial epidermis. Shape obovate. Bundle

sheath 1 or 2 layers of parenchymatous cells capping

main vascular bundle adaxially, ± absent abaxially; cells

rounded or lobed. Extraxylary sclerenchyma fibres

extending beyond bundle sheath, running through pal-

isade parenchyma up to abaxial epidermis; not enclosed

in bundle sheath (ultimate stage in P. montana, type Dl).

Sclerenchymatous hypodermal sheath formed by hypo-

dermal fibres extending paradermally in direction of leaf

margins, often connecting with fibres from main and sec-

ondary vascular bundles. Present in all species of leaf

structural type D2, absent in P. montana (type Dl)

(Figures 4C-F; 5D).

Key to species

la Sclerenchymatous hypodermal fibres absent

type Dl: P. montana (Figure 4C)

lb Sclerenchymatous hypodermal fibres present . . . type D2:

2a Outline of leaf in t/s transversely oblong; parts of lamina

extending beyond adaxial epidermis narrow, contain-

ing terminal vascular tissue only; margins upturned;

central furrow wide and shallow; palisade parenchy-

ma U-shaped, density 5 cells per 50 pm

P. paleacea (Figures 1R; 4D)

2b Outline of leaf in t/s transversely elliptic or carinate; parts

of lamina extending beyond adaxial epidermis varying

in width, each containing (2)3(— 6) secondary vascular

bundles; margins diverging outward or involute; cen-

tral groove deep; palisade parenchyma horseshoe- or

V-shaped, density (3)4(5) cells per 50 pm:

3a Outline of leaf in t/s transversely elliptic; margins invo-

lute; palisade parenchyma horseshoe-shaped

P. rigida (Figure 4E), less often P. vulgaris

3b Outline of leaf in t/s carinate; margins diverging outward

or involute; palisade parenchyma V-shaped

20. P. vulgaris (Figure 4F)

Intermediate states, with main vascular bundle orien-

tated in more than one position in relation to the ad- and

abaxial epidermis and mesophyll, were recorded in

P. glomerata subsp. glomerata, P. obtusifolia, P. montana,

P. paleacea and P rigida. All states were accounted for in

Tables 3 and 4, as well as in the construction of keys.

Comparative leaf anatomy at infrageneric level

In order to facilitate the interpretation of existing data

for all infrageneric taxa, data are summarized in Tables 3

and 4.

Bothalia 31,1 (2001)

65

DISCUSSION AND ADDITIONAL OBSERVATIONS

Leaf structure

Prevailing characters in Thymelaeaceae

Leaf structure in Thymelaeaceae exhibits a transfor-

mation series from mainly dorsiventral, the prevailing

state in the family, to isobilateral or centric in Diarthron

Turcz., Pimelea Banks & Sol. and Thymelaea Juss.

(Leandri 1930; Metcalfe & Chalk 1950), all these states

being present in Lachnaea and Cryptadenia (Beyers

1992; Beyers & Van der Walt 1995) and the inverse-dor-

siventral state prominent in Passerina. Thoday (1921)

recorded the isobilateral state in juvenile leaves in select-

ed species of Passerina , possibly reflecting its primitive

status in the genus. Leaves of P. sp. nov. 1 (Figures 3A;

5A) are isobilateral, whereas the leaves of all the other

species are inverse-dorsiventral, possibly representing

the more advanced state (Figures 3B-H; 4A-F; 5B-D).

Most species of Passerina are adapted to the dry,

warm summers and humid winters of the winter rainfall

area of the Cape Floristic Region, others survive in the

arid conditions of the Karoo, some grow in a range of

habitats along the eastern escarpment and some are

adapted to maritime conditions along the southern

African coast (Table 1). What appears to be adaptations

to these varying environments are reflected in the leaf

structure of the various species. Weiglin & Winter

(1991), studying the morphological-anatomical features

of perennial halophytes, pointed out the importance of

curvature of outer epidermal cell walls, epicuticular

waxes, mesophyll orientation, enrolled leaves and forti-

fication of tissue — characters present in most Passerina

species. Other important leaf adaptations in Passerina

are the decussate and appressed arrangement, the cymb-

iform shape and the inverted palisade parenchyma,

developing on that side of the leaf which is exposed to

the highest light intensity. Structural support is ren-

dered by the presence of sclerenchyma in the main and

secondary vascular bundles and in some Western Cape

species these fibres proliferate beyond the vascular

bundle sheath to join with hypodermal fibres, forming

a sclerenchymatous hypodermal sheath. The impor-

tance of many of these characters is further discussed

below.

Epidermal tissue

Prevailing characters in Thymelaeaceae

According to Metcalfe & Chalk (1950) the cuticular

membrane (CM) of the leaf is usually smooth. Epi-

dermal cells are arched outwards in Linostoma Wall, ex

Endl. and papillose on the lower surface in species of

Daphne L.

Speculations on functions and ecological aspects of

the leaf epidermis in Passerina have been dealt with by

Bredenkamp & Van Wyk (1999, 2000). Leaf arrange-

ment in Passerina causes the abaxial epidermis to be

largely exposed to the atmosphere, resulting in epider-

mal characters being more affected by environmental

change. However, the arrangement of the epidermal

cells and the ornamentation of the cuticular membrane

(CM) correlate well with leaf structural type (Table 3).

Species with epidermal cells arranged randomly and

with smooth or papillate cuticular ornamentation, all

have leaf structural type B. With the exception of P.

montana (type Dl) with epidermal cells arranged in

rows and smooth or papillate cuticular ornamentation,

structural types C and D correlate with epidermal cells

arranged in rows with striate cuticular ornamentation.

Finally, structural type D is also characterized by a

well-developed sclerenchymatous hypodermal sheath

(Table 4), possibly representing the more advanced

state.

Mesophyll

Prevailing characters in Thymelaeaceae

Palisade cells are generally short. Mesophyll, includ-

ing irregular sclerenchymatous fibres, is found in species

of Daphne L., Daphnopsis C.Mart., Enkleia Griff.,

Gyrinops Gaertn., Peddiea Harv. and Stephanodaphne

Baill. (Metcalfe & Chalk 1950).

In the isobilateral leaf of P. sp. nov. 1, mesophyll is

palisade-like and homogeneous. All other species of

Passerina display the possibly more advanced state in

which the mesophyll is inverted, with elongated, abaxial

palisade parenchyma and adaxial spongy parenchyma.

The orientation of the mesophyll in relation to leaf shape,

sclerenchyma and vascular tissue (Table 4), forms the

basis of various leaf structural types distinguished in this

study. The palisade parenchyma is horseshoe-shaped in

all species with narrowly transversely elliptic, cordiform

or canaliculate leaves in t/s and with leaf structural type

B (Figure 3B-H). In P. paleacea, the leaf is transversely

oblong in t/s and the palisade parenchyma is U-shaped

(Figures 1R; 4D). Palisade parenchyma is V-shaped

(Figure 4A, B, F) in species with structural types C and

D, in which the leaves are typically carinate in t/s. In the

most xeromorpic state, sclerenchyma extends through

the V-shaped palisade parenchyma, joining other hypo-

dermal fibres to form a sclerenchymatous hypodermal

sheath (Figure 4F). Irregular sclerenchymatous fibres are

absent in the mesophyll of leaves in Passerina.

The mesophyll of the leaf is seemingly adapted to sur-

vive arid conditions and high light intensity, thus becom-

ing xeromorphic in most species. This is reflected by the

palisade parenchyma which usually occurs in 1-3 layers,

quite densely arranged with 3-6 cells per 50 pm.

Palisade cells contain large numbers of chloroplasts,

ample amounts of tanniniferous substances and crystals

of calcium oxalate. In contrast, the spongy parenchyma

is usually adaxially arranged and aerenchymatous in

most species, corresponding to the epistomatic state of

the leaves. With spongy parenchyma around the stomata,

molecules of carbon dioxide would penetrate deep into

the leaf and the large intercellular spaces of aerenchyma

surrounding the vascular bundles would possibly have a

moist atmosphere critical to physiological processes

such as photosynthesis, respiration and transpiration.

TABLE 4. — Selected morphological and anatomical characters of all leaf structural types in Pcisserina

66

Bothalia 31,1 (2001)

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Bothalia 31,1 (2001)

67

Ecological significance

The xeromorphic character of the mesophyll can be

expressed in terms of the number of cell layers and the

density (reflected by the number of cells per 50 pm) of

the palisade parenchyma as well as the appearance of the

spongy parenchyma (Table 4). Possible adaptation to

physiological drought is shown by the homogeneous and

palisade-like mesophyll of P. sp. nov. 1, which grows on

the high mountains of the Karoo, where it is often cov-

ered by snow in winter. Among the remainder of the

species, the most xeromorphic state is found in P. glom-

erata subsp. glomerata (Figure 3F), P. obtusifolia

(Figure 3G) and P. comosa , all growing in the NW parts

of Western Cape; their leaves have (1)2 or 3 layers of

cells in the palisade parenchyma, a density of 4 or 5 cells

per 50 pm and the spongy parenchyma is densely

arranged. The mesophyll of P. ericoides (Figure 3D),

P. paleacea (Figure 4D) and P. rigida (Figures 3H; 4E)

can also be considered xeromorphic, possibly in

response to the effect of salt spray, as these species grow

on the sea shore. A xeromorphic mesophyll is also indi-

cated for P. montana (Figure 4C), which occupies a wide

range of habitats along the eastern escarpment. On the

other hand, P. paludosa (Figure 3C), a rare species from

marshy areas in the Cape Peninsula, has an inverse dor-

siventral leaf with one layer of palisade parenchyma, a

density of 3 or 4 cells per 50 pm and a large aerenchy-

matic spongy parenchyma, corresponding to the general

trend towards aerenchymatic tissue in marsh plants.

Leaf structural types: orientation and structure of

main vascular bundle in relation to epidermis and

mesophyll

Prevailing characters of vascular bundles in

Thymelaeaceae

hi Thymelaeaceae, large portions of vascular bundles are

often occupied by sclerenchyma and surrounded by a sheath

of parenchymatous cells containing tannin (Van Tieghem

1893; Gilg 1894; Supprian 1894; Metcalfe & Chalk 1950).

Both these characters are present in Passerina. Intraxylary

phloem in the petiole and midrib was recorded by Leandri

(1930) and cited by Domke (1934) for many genera,

excluding Passerina (= Chymococca). The lack of intraxy-

lary phloem in leaves of Passerina was confirmed by the

present study (Tables 3; 4; Figure 5).

Xeromorphic gradient

The progressive change in orientation of the main

vascular bundle in relation to the mesophyll shows a

xeromorphic gradient (Figure 5). Leaf structural type A

was defined on the basis of the isobilateral leaf, which is

probably an ancestral state. The xeromorphic character

of the leaf is strongly supported by the associated

homogenous mesophyll and central vascular system. The

leaves are inverse-dorsiventral in all other structural

types. A xeromorphic gradient is clearly expressed in leaf

structural type B. In types B1 and B2 the mesophyll is

aerenchymatic and the main vasular bundle adaxially or

centrally arranged. An increase in the density, the num-

ber of cells and layers and specialization of tissues takes

place in types B3 and B4, with B5 and B6 representing

the most xeromorphic forms. A similar increase in xero-

morphism can be shown in structural types C and D 1 and

D2, with the abaxial arrangement of the main vascular

bundle, the increase in sclerenchyma tissue and the ulti-

mate formation of the sclerenchymatous hypodermal

sheath, in type D2, as the most xeromorphic state.

Ecological significance

Replacement of the main vascular bundle from the ad-

to the abaxial position with the sequential increase in

xeromorphism suggests an adaptive strategy (Figure 5).

In type B the main vascular bundle is close to the adaxial

epidermis and stomata, possibly enhancing transpiration.

It is furthermore completely surrounded by aerenchyma.

Vascular tissue in close contact with aerenchyma provides

water, causing a moist atmosphere and a high water

potential in the large intercellular spaces. Inorganic and

organic substances are transported by the xylem and

phloem for various physiological processes taking place

in the mesophyll of the leaf. The more xeromorphic

arrangement mechanically strengthens the leaf, allows

enough moisture for photosynthesis and respiration, but

possibly retards loss of water through transpiration.

Increasing xeromorphism is illustrated by the vascular

bundle becoming abaxially orientated and finally by scle-

renchyma tissue of the vascular bundle abaxially project-

ing beyond the vascular bundle sheath and reaching up to

the abaxial epidermis, leaving only the adaxial part of the

vascular bundle in contact with aerenchyma. Vascular tis-

sue, providing moisture for the critical physiological

processes, is thus separated from aerenchyma by one or

two layers of the parenchymatous bundle sheath cap.

These adaptations, associated with geographical distribu-

tion (Table 1), can clearly be illustrated at species level

and for all the leaf structural types.

Type A: the main and secondary vascular bundles are

closely arranged with bundle sheaths adhering, forming

a central plate of veins surrounded by two or three layers

of palisade-like mesophyll, possibly acting as a protec-

tive sheath against the minimum temperatures, which are

often below freezing point in the habitat of P. sp. nov. 1

(Table 1; Figure 3A).

Type Bl; P. burchellii and P. pendula (Figure 3B) are

both mountainous species often surrounded by mist. The

main vascular bundle is more or less against the adaxial

epidermis close to the stomata and is completely sur-

rounded by aerenchyma, possibly enhancing transpira-

tion and aeration of the leaf.

Types B2. B3 and B4: these are the most common leaf

types, found in ± two-thirds of Passerina species (Table

4; Figure 3C-E). The possible adaptive significance of

these structural types remains much the same as in Bl,

except that the leaf becomes sequentially more xeromor-

phic as the main vascular bundle borders on or sinks into

the palisade parenchyma, possibly curtailing water loss.

Type B5; xeromorphism is enhanced in P. glomerata

subsp. glomerata and P. obtusifolia (Figure 3F, G), both

growing in the warm, arid Karoo. The main vascular

68

Bothalia 31,1 (2001)

bundle is abaxially embedded in palisade parenchyma

which differentiates into collenchyma and the spongy

parenchyma surrounding the main vascular bundle

which is more densely arranged with smaller intercellu-

lar spaces. This more xeromorphic arrangement mechan-

ically strengthens the leaf, allows moisture for physio-

logical processes but possibly retards loss of water.

Type B6: found in P. glomerata subsp. nov. occurring

on mountain tops from the Cederberg to the Cape

Peninsula and P. filiformis subsp. nov. which grows

between Malmesbury and Vredendal. In this xeromor-

phic leaf the main vascular bundle is adaxially surround-

ed by spongy parenchyma, enhancing aeration, and is

abaxially strengthened by palisade parenchyma and the

vascular bundle sheath which differentiates into col-

lenchyma. In P. rigida (Figure 3H), which grows in salt

spray along the coast, the exceptionally large main vas-

cular bundle is close to the adaxial epidermis and borders

on the tanniniferous abaxial epidermis, with the possible

adaptive advantage of strengthening the leaf and allow-

ing transpiration and associated physiological processes

at the same time.

Type C: found in P.falcifolia (Figure 4A) and P. sp.

nov. 4 (Figure 4B), growing in relatively moist environ-

ments (Table 1). The abaxial surface of the carinate leaf

is in contact with the atmosphere, but the obovate main

vascular bundle, situated abaxially, is well protected in

the V-shaped palisade parenchyma. The adaxial

parenchymatous bundle sheath cap and the vascular tis-

sue are in close contact with the aerenchyma, providing

moisture for the various physiological processes.

Type D 1 : in P. montana (Figure 4C) the extraxylary

sclerenchyma fibres touch the abaxial epidermis, but

hypodermal fibres are absent. This arrangement indicates

a high degree of mechanical strengthening and xeromor-

phism, possible adaptations to the wide range of habitats

along the eastern escarpment where these plants grow

(Table 1).

Type D2: present in P. paleacea (Figure 4D), P. rigida

(Figure 4E) and P. vulgaris (Figure 4F), all growing in

Western Cape, the centre of diversity for Passerina and

from where certain species extend west-, north- and east-

wards. Orientation and structure of the main vascular bun-

dle are the same as for type C, except that the sclerenchyma

tissue of the vascular bundle projects beyond the vascular

bundle sheath and reaches up to the abaxial epidermis form-

ing a sclerenchymatous hypodermal sheath, thus strength-

ening the leaf and making it more xeromorphic. Vascular

tissue remains in close contact with the aerenchyma, pro-

viding moisture for the various physiological processes.

Sclerenchyma

Prevailing characters in Thymelaeaceae

Van Tieghem (1893) described extraxylary fibres,

specially mentioning those without lignification in

Daphne mezereum L. and with lignification in D. cneo-

rum L. Supprian (1894), mentioned the presence of

fibres in the mesophyll of the leaves, which he called

‘Spicularzellen’ and regarded as a constant taxonomic

character. In a subsequent paper, Gilg (1894), critically

discussed the anatomical method applied by the two pre-

vious workers, doubting the constant taxonomic value of

‘Spicularzellen’. Thoday (1921) described a sclerenchy-

matous hypodermal sheath extending to the margins of

the leaves in P . filiformis and P. cf. falcifolia , introduc-

ing the term ‘wandering fibres’. Metcalfe & Chalk

(1950) acknowledged the previous works, also mention-

ing the presence of bundles of sclerenchymatous ele-

ments supporting leaf margins in species of Daphnopsis

C.Mart., Dicranolepis Planch, and Passerina.

During this study variation concerning leaf structural

types was taken into consideration and amply document-

ed. Considering the wide distribution of especially P.

montana and P. rigida , variation in leaf structural type

could be expected. In P. montana (Figure 4C), hypoder-

mal fibres are absent although lignified fibres project

beyond the vascular bundle sheath and reach the abaxial

epidermis (type Dl). Hypodermal sclerenchyma fibres

have been recorded in P. paleacea and P. rigida with leaf

structural types B2, B3 and B4 (Tables 3; 4), but without

the development of a hypodermal sclerenchymatous

sheath. The hypodermal sclerenchymatous sheath is usu-

ally associated with leaf structural type D2 as in the fol-

lowing description. The state in which the main vascular

bundle is abaxially orientated, the sclerenchyma extending

beyond the vascular bundle sheath, through the inverted

palisade parenchyma, reaching the abaxial epidermis (type

D) and connecting with the hypodermal sclerenchymatous

fibres to form a hypodermal sclerenchymatous sheath,

often reaching up to the leaf margins. Leaf structural type

D2 is regarded by the present authors as the ‘ultimate’

adaptation, in Passerina , to the Mediterranean climate of

the Cape Floristic Region. Figures 4D-F; 5.

Less important taxonomic characters

Leaf width

Considering the cymbiform, canaliculate or carinate

shape of leaves as well as the movement of the lamina

due to turgor pressure in the leaf, leaf width can at most

be used to interpret leaf shape, but is not regarded as taxo-

nomically significant.

Crystals

Calcium oxalate crystals or lime crystals were consid-

ered as taxonomically valuable in the Thymelaeaceae

and certain species of Passerina by Supprian (1894).

Solereder (1908), Metcalfe & Chalk (1950) and Metcalfe

(1983) report the presence of both druses and crystal

sand in the Thymelaeaceae, but do not consider these

crystals of much taxonomic value. In the present study,

druses were recorded in the parenchyma cells of the mes-

ophyll in all taxa of Passerina. Calcium oxalate crystals

in the intercellular spaces are regarded as fragments of

druses resulting from processing.

Tanniniferous substances

The substances are abundantly present in the epider-

mis, mesophyll as well as vascular bundle sheath and

parenchyma in all taxa of Passerina. No significant

interspecific variation was recorded.

Bothalia 31,1 (2001)

69

Phylogenetic considerations

Leaf structural type correlated with epidermal structure

A phylogenetic gradient for the leaf structural types

cannot be shown, as various characters probably evolved

separately. Thoday (1921) reported that juvenile leaves

in Passerina were isobilateral in transverse section, indi-

cating the possible ancestral state of leaf structural type

A. The central arrangement of the vascular bundle in leaf

structural type B is probably a primitive state, as re-

arrangement of the vascular bundle and consequent dif-

ferentiation of tissues probably represent derived states.

Species with epidermal cells arranged randomly and

smooth or papillate cuticular ornamentation (Group A in

Table 3) which correlate with leaf structural type B,

probably represent a primitive state. The hypodermal

sclerenchymatous sheath could have evolved as an adap-

tation to the Mediterranean climate of the Cape Floristic

Region and is therefore considered to be a derived char-

acter. In conjunction with leaf structural type D, this state

is possibly advanced. Epidermal cells arranged in rows

with striate cuticular ornamentation (Group B) and asso-

ciated with leaf structural type D possibly represent the

most derived state.

Mesophyll

Parenchyma cells are palisade-like and homogeneous

in the isobilateral leaf. All other species possibly repre-

sent the derived state in which the mesophyll is inverted,

with elongated, abaxial palisade parenchyma, horseshoe,

U- or V-shaped and with adaxial spongy parenchyma.

Sclerenchyma

Sclerenchyma (notably extraxylary fibres) in the

leaves of Passerina could possibly be regarded as prim-

itive as it is a prevailing state in the Thymelaeaceae.

Within Passerina the hypodermal sclerenchymatous

sheath in certain species could have evolved as an adap-

tation to the Mediterranean climate of the Cape Floristic

Region and in conjunction with leaf structural type D.

regarded as a possible advanced state.

Taxonomic significance

The present study has clarified the taxonomic signifi-

cance of various anatomical leaf characters at the fol-

lowing levels:

Family level

Most family characters prevail in Passerina in their

unmodified forms, e.g. the epidermis with a papillate

cuticular membrane (CM) and mucilaginous epidermal

cells, as well as the presence of extraxylary sclerenchy-

ma in the vascular bundle. Other characters have been

modified, for example the parenchymatous bundle

sheath cap in leaf structural types C and D (Tables 3, 4)

and the absence of intraxylary phloem in the leaves.

Irregular sclerenchymatous fibres present in the meso-

phyll of leaves in many species, have been modified into

a sclerenchymatous hypodermal sheath in Passerina.

Genus level

In most genera of Thymelaeaceae, leaves are isobilat-

eral, dorsiventral or inverse-dorsiventral (Kugler 1928).

P. sp. nov. 1 (leaf structural type A) is characterized by

isobilateral leaves. In all other species, the inversely eri-

coid leaves are arranged close to the stem, with the abax-

ial epidermis exposed to the environment. The palisade

parenchyma develops abaxially, the mesophyll is invert-

ed and the leaf is regarded as inverse-dorsiventral

(Kugler 1928).

Species level

Based on leaf epidermal characters, two groups (A and

B) of species are recognized, in which many species could

be classified to species level (Bredenkamp & Van Wyk

2000). The correlation of epidermal characters with the

leaf structural types has confirmed the authenticity of both

these character sets (Table 3). On the basis of anatomical

evidence the delimitation of the various species and infra-

specific taxa was confirmed (Tables 3, 4). P. sp. nov. 1,

growing on high mountains in the Ceres Karoo, is charac-

terized by isobilateral leaves and leaf structural type A.

P. glomerata subsp. glomerata (types B2, B3, B5), com-

mon to a large range of Karoo habitats, could be distin-

guished from P. glomerata subsp. nov. (type B6), growing

on mountain tops from the Cederberg range to the Cape

Peninsula. P. filifonnis L. was previously considered a

taxon with a wide distribution from Western and Eastern

Cape, along the eastern escarpment of southern Africa,

possibly up to Tanzania. The present study and the mor-

phological study indicate that plants growing in southern

parts of Western Cape, Eastern Cape and the escarpment

can clearly be distinguished from those of Western Cape.

Consequently P. filifonnis was split into P. filifonnis

subsp. filifonnis (types B3 and B4), P. filifonnis subsp.

nov. (types B4, B6) and P. sp. nov. 4 (type C). P. filifonnis

subsp. filifonnis is common in the Cape Peninsula, and

distributed from Piquetberg, across the Hex River Moun-

tains, to Attaquaskloof in the southern parts of Western

Cape. P. filifonnis subsp. glutinosa occurs in the area

between Malmesbury and Vredendal. P. sp. nov. 4 has a

wide distribution from Mossel Bay and Oudtshoom to

Eastern Cape and along the escarpment northwards to

Zimbabwe, with outliers in Tanzania. Furthermore, the

study of the leaf structural types revealed four leaf struc-

tural types and ten states, according to which all species

can be characterized and grouped. Leaf type A occurs in

P sp. nov. 1 ; type B in P. burchellii Thoday, P. pendula

Eckl. & Zeyh., P. comosa C.H.Wright, P. paludosa

Thoday, P galpinii C.H.Wright, P drakensbergensis

Hilliard & B.L.Burtt, P. ericoides L., P. sp. nov. 2, P. sp.

nov. 3, P. rubra C.H.Wright, P. filifonnis L. subsp. fili-

formis, P. filifonnis L. subsp. nov., P. obtusifolia Thoday,

P. glomerata Thunb. subsp. glomerata , and P. glomerata

Thunb. subsp. nov.; type C in Pfalcifolia C.H.Wright, P. sp.

nov. 4; type D in P. montana Thoday, P. paleacea Wikstr.,

P. rigida Wikstr. and P. vulgaris Thoday.

70

Bothalia 31,1 (2001)

CONCLUSIONS

Leaf anatomical evidence proved extremely useful in

the classification of Passerina. Four leaf structural types

and ten states are associated with specific habitats and

geographical distribution, illustrating a xeromorphic gra-

dient. On the basis of leaf structural types, four new

species and four subspecies were identified. Certain phy-

logenetic tendencies were proposed and the systematic

value of the various characters at family, genus and

species levels were assessed, thus enabling the anatomi-

cal characterization of all infrageneric taxa in Passerina.

This study illustrates prevailing Thymelaeaceae char-

acters as well as their modification and newly evolved

tendencies in Passerina. Observations correlate with

those obtained from other studies in the genus. Based on

the secondary reticulum of the polyforate pollen grain of

Passerina , the subtribe Passerininae Endl. was raised to

the tribe Passerineae (Endl.) Bredenkamp & Van Wyk

(1996). The unique leaf structural types and states iden-

tified by the present leaf anatomical study, provide more

evidence in support of the tribe Passerineae.

ACKNOWLEDGEMENTS

We are indebted to Mmes H. du Plessis and C. Steyn

and Dr E. Steyn for assistance with the LM, as well as Mrs

A. Romanowski for developing and printing many excel-

lent photographs. We are grateful to Prof. J. Coetzee, Mr

C.F. van der Merwe and Mr A.J. Botha, of the University

of Pretoria, for assistance with the SEM, TEM and EDS.

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Bothalia 31,1: 71-98 (2001)

A lexicon of plants traded in the Witwatersrand umuthi shops, South

Africa

V.L. WILLIAMS*, K. BALKWILL* and E.T.F. WITKOWSKI*

Keywords: families, medicinal plants, plant parts, survey, trade, umuthi , vernacular names

ABSTRACT

At least 5 1 1 medicinal plant species are traded commercially in 50 Witwatersrand umuthi shops. The plants are listed

alphabetically by genus and common (vernacular) name. The orthographic vernacular names, as well as the orthographic

variations in these names, are incorporated into the list. Annotations include the plant family, the number of umuthi shops

stocking the species, the language of the common name, and the plant part traded. The plant family in the region which has

the highest number of species and infraspecific taxa in trade is Liliaceae sensu lato., followed in descending order by

Fabaceae, Asteraceae, Euphorbiaceae and Amaryllidaceae. Approximately 88.6% of the vernacular names are in Zulu. The

mean number of umuthi shops per species is 12.3, ranging from 1 to 41 . Three hundred and fifty three species (69.2%) occur

in the four northern provinces, and 23 species are listed as threatened on the Red Data List.

INTRODUCTION

In 1994, a semiquantitative survey of 50 Witwaters-

rand umuthi shops was undertaken. Prior to the survey,

no attempt had been made to describe the commercial

market for medicinal plants and parts in the region. The

dynamics of the medicinal plant trade in Witwatersrand

umuthi shops have now been quantitatively described in

terms of the species used, sources and suppliers of plants,

the adequacy of the sample size, the diversity and equi-

tability of species being traded and the probability of cer-

tain taxa being utilized based on their availability in the

southern African flora and biomes (Williams et al. 2000).

In addition, that paper lists the species most commonly

available in at least two-thirds of the umuthi shops on the

Witwatersrand. The purpose of this paper is to present a

checklist of species in trade on the Witwatersrand. In

addition, a list of all orthographic vernacular names and

their variants referred to in this study is provided, with

cross references to the appropriate botanical names. The

list will be useful to researchers wanting to identify

species in trade.

METHODS

A stratified random sample of 50 umuthi shops was

surveyed on the Witwatersrand in 1994. The research

participants selected (including traditional healers who

owned shops) were proportionately representative of the

geographical distribution, ethnicity and gender of the

herb traders on the Witwatersrand at the time of the sur-

vey (Williams et al 1997). The vernacular names for the

plants sold in each umuthi shop were recorded — either

from a label on the shelf, or from the name cited by the

research participant. The language of the plant names

was also recorded to aid identification. It was not deter-

mined whether the plants sold were used as medicine or

for charm purposes.

* Department of Animal, Plant and Environmental Sciences, Uni-

versity of the Witwatersrand, Private Bag 3, WITS 2050, South Africa.

MS. received: 1999-02-19.

Forty-three references were used to identify most of

the plants (Watt & Breyer-Brandwijk 1932, 1962;

Gerstner 1938a, b, 1939a, b, c, 1941a, b; Miller 1941;

Ferreira 1949; Watt 1956; Bryant 1966; Hanekom 1967;

Stayt 1968; Jacot Guillarmod 1971; Immelman et al.

1973; Netshiungani & Van Wyk 1980; Liengme 1981;

Netshiungani 1981; Von Breitenbach 1981a, b, 1984,

1986, 1989, 1991; Arnold & Gulumian 1984; Cun-

ningham 1988; Scott-Shaw 1990; Hutchings 1992, 1996;

Moll 1992; Veale et al. 1992; Pooley 1993; Loxton et al.

1994; Hahn 1994; Brandt et al. 1995; Estes 1995;

Nichols 1995; Van Wyk et al. 1997; Kroon 1999). In

addition, species were identified from descriptions of

aerial plant parts given by the herb traders, or from spec-

imens purchased at the market and identified at the C.E.

Moss Herbarium (J), University of the Witwatersrand.

An incomplete set of voucher specimens was purchased,

and is presently being incorporated into the traditional

medicine collection at the Adler Museum of the History

of Medicine, University of the Witwatersrand.

Species identification from plant parts recorded in a

trade survey is a problem, especially when tracing the

plants through their vernacular names in the literature.

Errors in identification are sometimes made by the re-

corders of this information, and there are likely to be

citation errors of the vernacular names given by the

traders. However, the scientific names allocated to these

plants are, for the most part reliable, especially for the

names recorded in the Zulu vernacular for which an

extensive body of literature exists.

Following species identification, the data were

entered into a relational database designed in the pro-

gram DATAEASE by modifying the herbarium manage-

ment system of the C.E. Moss Herbarium. This obviated

entry of every scientific name encountered. The data cap-

ture format was designed specifically for the entry of the

survey records, and the data were entered under some of

the following fields for each species recorded in an

umuthi shop, including: 1, genus and species number

(Genspec No., following Arnold & De Wet 1993); 2,

genus and species; 3, common names; 4, language of the

72

Bothalia 31,1 (2001)

common name, and 5, trader information and code. Herb

traders were given codes denoting the region and order in

which they were surveyed, for instance ‘JB30’ for

‘Johannesburg Shop 30'. The number of records entered

for the fifty herb traders totalled 6 285.

In the commercial medicinal plant market, the names

cited for plants are generally not orthographically cor-

rect. Quite often in Zulu, for example, the prefix of the

stem word is omitted or shortened. Therefore, to identify

plants from Zulu names from some species lists requires

searching for stem words that are orthographically or

phonetically similar. In order to produce a ‘user friendly’

list, both the orthographic names and their variants were

captured on this database. The correct, or orthographic,

form of the vernacular names in this paper was checked

by A. Ngwenya from the Natal Herbarium (NH) in Durban.

Following data capture, a complete checklist of all

families, genera, species, plant parts, orthographic vernac-

ular names and orthographic variants, was extracted from

the database, as well as the number of citation records per

species; authors of the botanical names are provided in

this list (Appendix 1). Citation records were extracted

according to the vernacular names, thus it was not possi-

ble to discriminate between the species. In Appendix 1, the

number of shops cited is for the applicable vernacular

name, not necessarily the precise species (e.g. seven

species of Helichrysum called imphepho are cited as being

recorded in 35 shops, as imphepho was recorded from 35

shops). The checklist was compared with the Red Data

List (Hilton-Taylor 1996a, b, 1997) to compile a list of red

data taxa traded medicinally on the Witwatersrand. The

checklist was also compared with the flora found in the

four northern provinces (Retief & Herman 1997) to exam-

ine the extent to which species may be harvested in areas

other than KwaZulu-Natal, where most of the harvesting

occurs (Williams et al. 2000).

RESULTS AND DISCUSSION

Five hundred and eleven species from 328 genera and

119 families were identified as being traded on the

Witwatersrand (Appendix 1). The most commonly traded

families in terms of the number of genera (in parenthe-

TABLE 1. — Broad floristic analysis of taxa traded on the Witwaters-

rand

* only families with > 6 genera are included in this listing.

* including species, subspecies and varieties.

ses), are: Fabaceae (Leguminosae) (28), Liliaceae sensu

lato (23), Asteraceae (Compositae) (20), Euphorbiaceae

(14) and Rubiaceae (13) (Table 1). The largest families

traded, in terms of the number of species and infraspecif-

ic taxa, are: Liliaceae s.l. (57), Fabaceae (Leguminosae)

(38), Asteraceae (Compositae) (34), Euphorbiaceae (28)

and Amaryllidaceae (15) (Table 1; Figure 1 ).

A mean of 126 ± 65.9 (SD) species was recorded per

umuthi shop, ranging from a minimum of 10 species to a

maximum of 294 species in a shop (Williams et al. 2000).

On average, a species was recorded in 12.3 ± 11.3 shops.

Seventy eight species were recorded in only one shop, and

no species was recorded in more than 42 out of a maxi-

mum 50 umuthi shops (Williams et al. 2000). Drimia spp.

had the highest recorded frequency of occurrence (82%),

followed by Eucomis autumnalis and Scilla natalensis

(78% each) (Williams et al 2000). Thirty six species were

found in more than 33 of the shops surveyed.

A comparison with the flora of the four northern

provinces (Retief & Herman 1997) shows that 353 of the

510 species (69.2%) are found in the Northern Province,

North-West, Gauteng and/or Mpumalanga. A mean of

Bothalia 31,1 (2001)

73

27.6% of the species are known to be harvested from

this region, compared to 42.1% for KwaZulu-Natal

(Williams el al. 2000). Therefore, the pressure on plant

resources in KwaZulu-Natal could be relieved if com-

mercial gatherers harvested plants in one of the four

northern provinces. However, the predominance in uti-

lization, demand and trade of and for Zulu medicinal

plants in the region, and the familiarity (or lack thereof)

of other ethnic groups with the Zulu plant pharma-

copoeia are constraints to accessing and harvesting these

resources from elsewhere.

Twenty one species (4.1%) from 17 genera and 16

families represent alien plant species. These species are

naturalized in the regions harvested by commercial gath-

erers. In addition to the exotics harvested in southern

Africa, there are at least five plants imported from India

by Asian traders.

A total of 23 species (4.7%) traded on the Wit-

watersrand are threatened according to the Red Data List

(Hilton-Taylor 1996a, b; 1997) (Table 2). A further 28

species are on the Red Data List, but are characterized as

‘not threatened'. The global conservation status of

Siphonochilus aethiopicus is listed in Table 2 as ‘Not

Threatened', even though subsequent corrections to the

Red Data List (Hilton-Taylor 1997) have listed the

species as extinct in the wild in KwaZulu-Natal. The

plant is only obtainable from farmers in KwaZulu-Natal,

or collected in the veld in Mpumalanga and Swaziland.

Schlechterina mitostemmatoides is also reported to be

threatened (A. Hutchings pers. comm.).

The common names of medicinal plants in trade are

predominantly in the Zulu vernacular. Eighty-nine per-

cent of the plant names recorded were Zulu (Table 3)

(Appendix 2). Despite the inclusion of Sotho, Venda,

Shangaan and Swazi traditional healers (who were shop

owners) in the survey, the proportion of plant names trad-

ed in these languages was low. This shows how tradi-

tional healers of other ethnic groups have adapted to the

trade and become familiar with the names of plants in the

language that dominates the trade, i.e. Zulu.

There is practical value in listing both the orthograph-

ic plant names and the orthographic variants in the plant

checklist. Firstly, species identification is made easier,

and secondly, the checklist provides a guide by suggest-

ing the correct form of vernacular names to be used in

the future. The justification for using capitals to separate

the prefix from the stem of the noun, while not strictly

correct in Zulu, is incorporated in the paper for the ben-

efit of those using this list who are familiar with the

structure of Zulu nouns. Most Zulu dictionaries enter

nouns under the stem, but for readers not being able to

identify the stem it is useful to capitalize it (A. Koopman

pers. comm.). It is the authors’ experience that this sepa-

ration makes species identification from vernacular

TABLE 2. — Global conservation status of taxa traded on the Witwatersrand, according to the Red Data List for southern Africa

(Hilton-Taylor 1996a, b; 1997)

* The old IUCN categories (Davis et al. 1986) rather than the new categories (IUCN 1994) were used by Hilton-Taylor because many of the southern

African data are still too sparse to apply the new criteria and categories (Hilton-Taylor 1996a).

± Hilton-Taylor (1996a) listed Siphonochilus as Endangered (E) in KwaZulu-Natal and the former Transvaal and Indeterminate (I) in Swaziland.

The overall global conservation status of the species was listed as Not Threatened (nt). A subsequent correction to the conservation status of the

species in KwaZulu-Natal listed Siphonochilus as Extinct (Ex) in the province (Hilton-Taylor 1997). However, no changes to the global con-

servation status of the taxon was recommended, hence the species is cited as Not Threatened in the above table.

74

Bothalia 31,1 (2001)

TABLE 3. — Language of common names

names in the literature easier. Entering the Zulu name

alphabetically under both the initial vowel of the prefix

and the stem of the noun is also an approach to accom-

modate a variety of users of the list.

CONCLUSION

The aim of this paper is to provide a checklist of the

flora traded in Witwatersrand umuthi shops. The value of

this list is that it can also be used by researchers in other

regions to identify plants used medicinally and traded

commercially. Knowing what species are traded com-

mercially is the foundation for identifying taxa threat-

ened by the trade, and comparisons with other regional

and national medicinal plant markets.

ACKNOWLEDGEMENTS

We thank: Fanie Mokgaba from the Johannesburg

Zoological Gardens for his assistance with the survey;

the NRF, Endangered Wildlife Trust and TRAFFIC

southern Africa for financial support; C. Prentice and R.

Archer from NBI Pretoria for help with updating the

nomenclature; A.E. van Wyk , N. Hahn and M. Mander

for help with plant identification; A. Hutchings, N.

Crouch and A. Koopman for valuable comments on the

manuscript; Alfred Ngwenya from the Natal Herbarium

(NH) in Durban for providing the correct Zulu format

and checking all the vernacular names; and finally the

research participants who took part in the survey.

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APPENDIX 1.— CHECKLIST OF TAXA TRADED MEDICINALLY ON THE WITWATERSRAND

Taxa are arranged alphabetically according to genus and species. After the author(s) of species name, the sequence

of annotation is as follows:

plant family;

number of herb traders stocking the species, maximum of 50 herb traders;

vernacular name(s): orthographic/correct Zulu names are in italics ;

language of vernacular name: Z = Zulu, S = Sotho, V = Venda, Sh = Shangaan, T = Tswana, Ts = Tsonga, X =

Xhosa, Sw = Swazi, E = English, A = Afrikaans or combination/variation thereof;

plant part traded: bk = bark, bb = bulb/tuber/corm, fl = flower, fr = fruit, 1 = leaf, rt = root, sd = seed, st = stem,

wpl = whole plant.

*exotic taxa, including naturalized taxa.

+Apiaceae = Umbelliferae; Asteraceae = Compositae; Clusiaceae = Guttiferae; Fab. = Fabaceae = Leguminosae;

Lamiaceae = Labiatae; Poaceae = Graminae.

Abrus precatorius L. subsp. africanus, Verde., Fab.VPapilionoideae, 2

imkoka (Z) sd; umKhokha (Z) sd

Acacia

caffra (Thunb.) Willd., Fab.*/Mimosoideae, 6

mthole (Z) bk; umTholo (Z) bk

karroo, Hayne, Fab.VMimosoideae, 2

umuNga (Z) bk

robusta Burch.

subsp. clavigera (E.Mey.) Brenan, Fab.VMimosoideae, 1

umgamazi (Z) bk; umkomasi (Z) bk; umNgamanzi (Z) bk

subsp. robusta, Fab.VMimosoideae, 1

umgamazi (Z) bk; umkomasi (Z) bk; umNgamanzi (Z) bk

sieberiana DC. var. woodii (Burtt Davy) Keay & Brenan, Fab.+/

Mimosoideae, 2

mkhamba (Z) bk; umKhamba (Z) bk

xanthophloea Benth., Fab.VMimosoideae, 35

kanyakude (Z) bk; mkanyakude (Z) bk; umKhanyakude (Z) bk

Acalypha

glandulifolia Buchinger ex Meisn., Euphorbiaceae, 8

bonisele (Z) wpl; gibonisele (Z) wpl; ibonisele (Z) wpl; unGibonisele

(Z) wpl

peduncularis E.Mey. ex Meisn., Euphorbiaceae, 4

sununundu (Z) rt; uSununundu (Z) rt

punctata Meisn., Euphorbiaceae, 4

sununundu (Z) rt; uSununundu (Z) rt

schinzii Pax, Euphorbiaceae, 4

sununundu (Z) wpl; uSununundu (Z) wpl

villicaulis Hochst., Euphorbiaceae, 22

byere (Sh) rt; mpendulo (Z) rt; umPendulo (Z) rt

Achyrocline stenoptera (DC.) Hilliard & B.L.Burtt, Asteraceae*, 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) 1/st

Achyropsis avicularis (E.Mey. ex Moq.) Cooke & Wright, Amaranth-

aceae, 11

bohlololo (Z) wpl; iBohlololo (Z) wpl; isigiba-nyongo (Z) wpl;

isiNamasenyongo (Z) wpl; sinama- senyonga (Z) wpl; sipha-

senyuko (Z) wpl; umSiphawenyoka (Z) wpl

Acokanthera

oblongifolia (Hochst.) Codd, Apocynaceae, 6

hlungunyembe (Z) 1/rt; iHlungunyem.be (Z) 1/rt; nhlungunyembe (Z) 1/rt;

oppositifolia (Lam.) Codd, Apocynaceae, 9

hlungunyembe (Z) 1/rt; iHlungunyembe (Z) 1/rt; khwangu (Z) 1/rt;

mgwangu (Z) 1/rt; mhlungunyembe (Z)l/rt; mkwangu (Z) 1/rt;

nhlungunyembe (Z) 1/rt; umKhwangu (Z) 1/rt

Acorus calamus L.*, Araceae, 20

iKalamuzi (Z) rt; kalmoes (Z) rt

Acridocarpus

natalitius Juss.

var. linearifolius Launert, Malpighiaceae, 28

mabophe (Z) rt; mahbhope (Z) rt; uMabopha (Z) rt

var. natalitius, Malpighiaceae, 28

mabophe (Z) rt; mahbhope (Z) rt; uMabopha (Z) rt

Adenia gummifera (Harv.) Harms var. gummifera, Passifloraceae, 34

impendewa shaye (Z) st; imPindamshaye (Z) st; mpindamshaye (Z) st;

phindamshye (Z) st; pindamshaye(Z) st

Adenostemma

caffra DC., Asteraceae*, 15

mahogwe (Z) 1/st; uMahogwe (Z) 1/st

viscosum J.R. & G.Forst., Asteraceae*, 15

mahogwe (Z) 1/st; uMahogwe (Z) 1/st

Afzelia quanzensis Welw., Fab.VCaesalpinioideae, 3

dlavusa (Z) bk; inkehli (Z) bk; mdlavusa (Z) bk; nkehle (Z) bk; um-

Dlavuza (Z) bk

Agapanthus

africanus (L.) Hoffm., Liliaceae s.L, 20

ubane (Z) rt; uBani (Z) rt; uhlakahla (Sw) rt

campanulatus Leighton, Liliaceae s.L, 10

ubane (Z) rt; uBani (Z) rt

praecox Willd. subsp. orientalis (Leighton) Leighton, Liliaceae s.l.. 10

ubane (Z) rt; uBani (Z) rt

Agathosma ovata (Thunb.) Pillans, Rutaceae, 17

mahirisaka, white (Z/E) rt; maisaka, white (Z/E) rt; mayisake, white

(Z/E) rt; mysaka, white (Z/E) rt; uMahesakomhlope (Z) rt

Alberta magna E.Mey., Rubiaceae, 3

mchumane (Z) bk; umCumane (Z) bk

Albizia adianthifolia (Schumach.) W. Wight, Fab.VMimosoideae, 33

katankawu (Z) bk; mbhando (V) bk; mdlanlothi (Z) bk; mgadankawu

(Z) bk; mhlandloti (X) bk; mkadankawu (Z) bk; mslangoti (Z)

bk; solo (Z) bk; umGadankawu (Z) bk; umHIadlothi (X) bk;

umKadankawu (Z) bk; uSolo (Z) bk; zinhla (Z) bk

Albuca

fastigiata ( L.f. ) Dryand, Liliaceae s.l.. 9

maphipha ntelezi (Z) bb; uMaphiphintelezi (Z) bb

nelsonii N.E.Br., Liliaceae s.l., 9

maphipha ntelezi (Z) bb; uMaphiphintelezi (Z) bb

76

Bothalia 31,1 (2001)

Albuca (cont.)

setosa Jacq., Liliaceae s.l., 20

inGcino (Z) bb; ncino (Z) bb; ngcina (Z) bb; ngcino (Z) bb

Alepidea

amatymbica Eckl. & Zeyh. var. amatymbica. Apiaceae+, 34

iKhathazo (Z) rt; kataza (Z) rt; kathazo (Z) rt

longifolia E.Mey., Apiaceae+, 34

iKhathazo (Z) rt; kataza (Z) rt; kathazo (Z) rt

pilifera WWm., Apiaceae*, 1

lesokwana (S) rt

setifera N.E.Br., Apiaceae*, 1

lesokwana (S) rt

Aloe

aristata Haw., Liliaceae s.l., 16

mathithibala (Z) wpl; uMathithibala (Z) wpl

maculata All., Liliaceae s.l., 5

cena (Z) 1; iCena (Z) 1

micrantha Haw., Liliaceae s.l., 1

sekgopha (S) 1

sp., Liliaceae s.l., 5

cena (Z) 1; iCena (Z) 1; inqgene (Z) 1; marobadibogale (S) 1

Andrachne ovalis (Sond.) Miill.Arg., Euphorbiaceae, 19

mbeza (Z) rt; uMbesa (Z) rt; uMembeza (Z) rt

Anemone

caffra Eckl. & Zeyh., Ranunculaceae, 14

manzemnyama (Z) rt; uManzamnyama (Z) rt

fanninii Harv. ex Mast., Ranunculaceae, 14

manzemnyama (Z) rt; uManzamnyama (Z) rt

Ansellia africana Lindl., Orchidaceae, 11

imFeyenkawu (Z) rt

Anthospermum rigid um Eckl. & Zeyh. subsp. pumilum (Sond.) Puff,

Rubiaceae, 19

mlomo mnandi (Z) rt; umLomomnandomncane (Z) rt

Apodytes dimidiata E.Mey. ex Arn. subsp. dimidiata, Icacinaceae, 4

dakane (Z) rt; indakane (Z) rt; umDakane (Z) rt

Aptenia cordifolia (L.f.) Schwant. var. cordifolia, Mesembryanthe-

maceae, 4

bohlololo (Z) wpl; iBohlololo (Z) wpl

Argyrolobium tuberosum Eckl. & Zeyh., Fab.VPapilionoideae, 2

tsoetla-e-nyenyane (S) rt; tswetlane (S) rt

Artemisia afra Jacq. ex Willd., Asteraceae*, 19

lengana (S) 1/st; hlonyane (Z) 1/st; mhlonyane (Z) l/st; umHlonyane (Z)

1/st

Asclepias

cucullata (Schltr.) Schltr., Asclepiadaceae, 9

delawena (Z) l/rt; delunina (Z) 1/rt; uDelunina (Z) l/rt

spp., Asclepiadaceae, 23

iShongwe (Z) rt; shongwe (Z) rt

Asparagus

africanus Lam., Liliaceae s.l., 19

isiGoba (Z) rt

asparagoides (L.) W.Wight, Liliaceae s.l., 20

buthe (Z) rt; iButha (Z) rt

falcatus L., Liliaceae.?.!., 19

isiGoba (Z) rt

laricinus Burch., Liliaceae s.l., 19

buthe (Z) rt; iButha (Z) rt

ramosissimum Baker, Liliaceae s.l., 20

buthe (Z) rt; iButha (Z) rt

setaceus (Kunth) Jessop, Liliaceae s.l., 19

buthe (Z) rt; iButha (Z) rt

sp., Liliaceae s.l., 19

isiGoba (Z) rt

virgatus (Baker) Oberm., Liliaceae s.l., 19

buthe (Z) rt; iButha (Z) rt

Aspilia natalensis (Sond.) Wild , Asteraceae*, 4

pamapuce (Z) 1/st; phamaphuce (Z) 1/st; ubuHlungwana (Z) 1/st;

uPhamephuce (Z) 1/st

Aster bakerianus Bur/t Davy ex C.A.Sm., Asteraceae+, 30

dlutjane (Z) rt; dlutshani (Z) rt; idlutshane (Z) rt; lutjane (Z) rt;

uDlutshana (Z) rt

Balanites maughamii Sprague, Balanitaceae, 28

gobandlovu (Z) bk; ipamba (Z) bk; iPhamba (Z) bk; uGobandlovu (Z)

bk

Baphia racemosa (Hochst.) Baker, Fab.VPapilionoideae, 1

isiFithi (Z) bk

Barringtonia racemosa (L.) Roxb., Lecythidaceae, 6

mhuluka (Z) rt; mululuka (Z) rt; uMuluIuka (Z) rt

Basella paniculata Volkens, Basellaceae, 1

ndaba-ugehlele (Z) 1/st; unDabingehlele (Z) 1/st

Bauhinia bowkeri Harv., Fab.VCaesalpinioideae, 17

dlandlovu (X) rt; mdlandlovu (X) rt; umDIandlovu (X) rt

Begonia

dregei Otto & Dietr., Begoniaceae, 3

iDlula (Z) rt

homonyma Steud., Begoniaceae, 3

iDlula (Z) rt

Behnia reticulata (Thunb.) Didr., Liliaceae s.l., 18

isiGoba (Z) rt; iZaza (Z) rt

Belamcanda spp., Iridaceae, 1

inDawoluthi (Z) rt; ndauluti (Z) rt

Berchemia discolor (Klotzsch) Hemsl., Rhamnaceae, 7

madlozana (Z) rt; mahlozane (Z) rt; uMadlozana (Z) rt

Berkheya

multijuga (DC.) Roessl., Asteraceae*, 5

iKhakhasi (Z) rt; ugashi (Z) rt; ukashi (Z) rt

onopordifolia (DC.) O.Hojfm. ex Burtt Davy var. onopordifolia.

Asteraceae*, 1

mohato (S) rt

Bersama

lucens (Hochst.) Szyszyl., Melianthaceae, 32

isiNdiyandiya (Z) bk; sindiyandiya (Z) bk

stayneri E. Phillips, Melianthaceae, 32

indiyaza (Z) bk; isiNdiyandiya (Z) bk; ndiyaza (Z) bk; sindiyandiya (Z)

bk; unDiyaza (Z) bk

swinnyi E. Phillips, Melianthaceae, 32

indiyaza (Z) bk; isiNdiyandiya (Z) bk; ndiyaza (Z) bk; sindiyandiya (Z)

bk; unDiyaza (Z) bk

tysoniana Oliv., Melianthaceae, 32

indiyaza (Z) bk; isiNdiyandiya (Z) bk; ndiyaza (Z) bk; sindiyandiya (Z)

bk; unDiyaza (Z) bk

Blighia uni jugata Baker, Sapindaceae, 2

iDlebendlovu (Z) bk; mhlabelo (Z) bk; ndlebendtovu (Z) bk; umHIa-

belo (Z) bk; umHIabeloweqolo (Z) bk

Boophane disticha (L.f.) Herb., Amaryllidaceae, 27

iNcotho (Z) bb; incoto (Z) bb; ngoto (Z) bb; ngubozohlonya (Z) bb

Boscia

albitrunca (Burch.) Gilg & Gilg-Ben., Capparaceae, 14

inYokiziphinda (Z) rt; mvithi (Z) rt; nogimpinda (Z) rt; nyokaspinda (Z)

rt; umnyokiziphinda (Z) rt; umvete (Z) rt; umVithi (Z) rt

foetida Schinz subsp. rehmanniana (Pest.) Toelken, Capparaceae, 10

mvithi (Z) rt; umvete (Z) rt; umVithi (Z) rt

Bowiea volubilis Harv. ex Hook.f. Liliaceae s.l., 32

gibisile (Z) bb; uGibisila (Z) bb

Brachylaena discolor DC. subsp. discolor var. discolor. Asteraceae*,

23

isiPhahluka (Z) l/rt; spahluka (Z) l/rt; spatluga (Z) l/rt; umPhahla (Z) l/rt

Brackenridgea zanguebarica Oliv., Ochnaceae, 5

mutavhatsindi (V) rt; ntabazimbi (V) rt; tabazimbi (V) rt; thabatsindi (V) rt

Bridelia

cathartica Bertol.f, Euphorbiaceae, 3

gazine (Z) bk; ngazine (Z) bk; unGazine (Z) bk

micrantha (Hochst.) Bail/., Euphorbiaceae, 20

hlalamakwababa (Z) bk; mutshutshunga (V) bk; umHIalamagwababa

(Z) bk; umhlalimakwaba (Z) bk

Brunsvigia sp., Amaryllidaceae, 35

maime (Z) bb; mayime (Z) bb; uMayime (Z) bb

Buddie. ja salviifolia (L.) Lam., Loganiaceae, 2

iLoshana (Z) rt; loshana (Z) rt

Bothalia 31,1 (2001)

77

Bulbine

alooides (E.) Willd., Liliaceae s.l., 2

iBhucu (Z) wpl; ibuco (Z) wpl; ibuqu (Z) wpl

asphodeloides (L.) Willd., Liliaceae s.l., 2

iBhucu (Z) wpl; ibuco (Z) wpl; ibuqu (Z) wpl

frutescens (L.) Willd., Liliaceae s.l., 2

iBhucu (Z) wpl; ibuco (Z) wpl; ibuqu (Z) wpl

latifolia (L.f.) Roem. & Schult., Liliaceae s.l., 2

iBhucu (Z) wpl; ibuco (Z) wpl; ibuqu (Z) wpl

Burchellia bubalina [L.f. ) Sims, Rubiaceae, 2

intshwalabenyoni (Z) bk; iqongqo (Z) bk; tshwalabenyoni (Z) bk;

umqonqo (Z) bk; uQongqo (Z) bk; uTshwalabenyoni (Z) bk

Buxus

macowanii Oliv., Buxaceae, 1

galagala (X) I/fl; umGalagala (X) 1/fl; umPhicamaguma (Z) 1/fl

natalensis [Oliv.) Hutch., Buxaceae, 1

galagala (X) 1/fl; umGalagala (X) 1/fl

Cadaba natalensis Sond., Capparaceae, 4

amangwe, black (Z/E) bk; amaNgwamnyama (Z) bk

Callilepis laureola DC., Asteraceae+, 27

hlamvu (Z) rt; iHlamvu (Z) rt; imPila (Z) rt; mpila (Z) rt

Calodendrum capense [L.f.) Thunb., Rutaceae, 29

memezi, white (Z/E) bk; uMemezomhlope (Z) bk

Canthium inerme [L.f. ) Kuntze, Rubiaceae, 2

umVuthwemini (Z) rt; vutwamini (Z) rt

Capparis

brassii DC., Capparaceae, 15

cwaningi (Z) rt; iQwaningi (Z) rt; mabusane (Z) rt; moapatladi (S) rt;

moopatladi (S) rt : uMabusane (Z) rt

tomentosa Lam., Capparaceae, 15

cwaningi (Z) rt; iQwaningi (Z) rt; mabusane (Z) rt; moapatladi (S) rt;

moopatladi (S) rt; uMabusane (Z) rt

Carpobrotus edulis [L.) L. Bolus, Mesembryanthemaceae, 8

iKhambilamabulawo (Z) 1/st

Casearia gladiiformis Mast., Flacourtiaceae, 1

juluka (Z) rt; mjuluka (Z) rt; umJuluka (Z) rt

Cassia abbreviata Oliv. subsp. beareana (Holmes) Brenan, Fab.*/

Caesalpinioideae, 1

nomanyama (Sh) bk

Cassine peragua L. subsp. peragua, Celastraceae, 4

maqadini (X) bk

Cassipourea

flanaganii (Schinz) Alston, Rhizophoraceae, 29

memezi, red (Z/E) bk; uMemeziobomvu (Z) bk

gerrardii ( Schinz ) Alston, Rhizophoraceae, 29

memezi, red (Z/E) bk; mqonga (X) bk; uMemeziobomvu (Z) bk; um-

Qonga (X) bk

Catha edulis (Vahl) Forssk. ex Endl. , Celastraceae, 2

mhlwazi (Z) bk/1; shikwane (S) bk/1; umHIwazi (Z) bk/i

Catunaregam spinosa (Thunb.) Tirveng. subsp. spinosa, Rubiaceae, 5

maqadani (Z) fr; uMaqadini (Z) fr

Celosia trigyna L., Amaranthaceae, 8

uVelabahleke (Z) wpl; velabahleka (Z) wpl

Celtis mildbraedii Engl., Ulmaceae, 3

uZinhla (Z) bk; zinhla (Z) bk

Cephalaria humilis (Thunb.) Roem. & Schult., Dipsacaceae, 22

dira-ga-dibone (S) rt; mpheabonwe (S) rt; mpikayiboni (Z) rt; pigabone

(Z) rt; pigayibone (Z) rt; pikayiboni (Z) rt; raadeboni (S) rt;

uMpikayiboni (Z) rt

Cheilanthes hirta Swartz, Adiantaceae, 1 1

iNkomankoma (Z) rt/1; khomankhoma (Z) rt/1; nkomankoma (Z) rt/1

Chenopodium

ambrosioides L. *, Chenopodiaceae, 5

hlahlabadimo (S) 1/st; insukumbili (Z) 1/st; ntsukumbili (Z) 1/st;

slaslabadiem (S) 1/st; sukumbili (Z) 1/st; tlhatlhabadimo (T)

wpl; uN sukumbili (Z) 1/st

sp., Chenopodiaceae, 1

phate-ea-ngaka (S) rt; phateyangaka (S) rt

Chlorophytum comosum (Thunb.) Jacq., Liliaceae s.l, 5

iPhamba (Z) wpl; pamba (Z) wpl; phamba (Z) wpl

Chrysanthemoides monilifera (L.) Norl. subsp. monilifera. Astera-

ceae+, 1

inKupheyana (Z) wpl; nqupulwane (Z) wpl

Cinnamomum

camphora (L.) J.Presl*, Lauraceae, 31

loselina (Z) bk; roselina (Z) bk; uLoselina (Z) bk; uRoselina (Z) bk

zeylancium Nees.*, Lauraceae, 4

mondi (Z) bk; uMondi (Z) bk

Cissampelos torulosa E.Mey. ex Harv., Menispermaceae, 2

kalimele (Z) 1/st; uKhalimele (Z) 1/st

Clausena anisata (Willd.) Hook.f. ex Benth., Rutaceae, 3

nukelamabiba (Z); umnukela mbiba (Z); umNukelambiba (Z); unukam-

biba (Z)

Clivia

miniata (Lindl.) Regel var. miniata. Amaryllidaceae, 35

maime (Z) wpl; mayime (Z) wpl; uMayime (Z) wpl

nobilis Lindl., Amaryllidaceae, 35

maime (Z) wpl; mayime (Z) wpl; uMayime (Z) wpl

spp., Amaryllidaceae, 35

maime (Z) wpl; mayime (Z) wpl; uMayime (Z) wpl

Clutia

hirsuta E.Mey. ex Sond., Euphorbiaceae, 2

ngwedlane (Z) 1/rt; uNgwaleni (Z) 1/rt

platyphylla Pax & K.Hoffm., Euphorbiaceae, 2

ngwedlane (Z) 1/rt; uNgwaleni (Z) 1/rt

pulchella L., Euphorbiaceae, 2

ngwedlane (Z) 1/rt; uNgwaleni (Z) 1/rt

spp., Euphorbiaceae, 1

umBethe (Z) rt

Coddia rudis (E.Mey. ex Harv.) Verde., Rubiaceae, 1

dondweni (Z) rt; umDondwani (Z) rt

Combretum

caffrum (Eckl. & Zeyh. ) Kuntze, Combretaceae, 6

mdubu (Z) rt; umDubu (Z) rt

erythrophyllum (Burch.) Sond., Combretaceae, 6

mdubu (Z) rt; umDubu (Z) rt

hereroense Schinz, Combretaceae, 6

mdubu (Z) rt; umDubu (Z) rt

kraussii Hochst., Combretaceae, 6

mdubu (Z) rt; umDubu (Z) rt

Commiphora

africana (A. Rich.) Engl, Burseraceae, 1

uMinyela (Z) bk

harveyi (Engl.) Engl., Burseraceae, 1

uMinyela (Z) bk

woodii Engl, Burseraceae, 1

umDewehlati (Z) bk

Convolvulus sagittatus Thunb., Convolvulaceae, 15

uVimbukhalo (Z) rt; vimbokhalo (Z) rt

Corbichonia decumbens (Forssk.) Exell, Aizoaceae, 4

iNyongwane (Z) rt; nyongwane (Z) rt

Corchorus

asplenifolius Burch., Tiliaceae, 34

bangalala (Z) rt; uBangalala (Z) rt

confusus Wild, Tiliaceae, 10

bonisele (Z) rt; gibonisele (Z) rt; ibonisele (Z) rt; unGibonisele (Z) rt

Cotula anthemoides L., Asteraceae+, 1

hlonyane (Z) 1/st; mhlonyane (Z) 1/st; umHlonyane (Z) 1/st

Crabbea hirsuta Harv., Acanthaceae, 23

musa (Z) rt; ntantanyana (Z) rt; uMusa (Z) rt

Crinum

bulbispermum (Burm.f.) Milne-Redh. & Schweick., Amaryllidaceae,

12

mduze (Z) bb; umduze (Z) bb; umNduze (Z) bb

delagoense I.Verd., Amaryllidaceae, 12

mduze (Z) bb; umduze (Z) bb; umNduze (Z) bb

macowanii Baker, Amaryllidaceae, 12

mduze (Z) bb; umduze (Z) bb; umNduze (Z) bb

78

Bothalia 31,1 (2001)

Crinum (cont.)

moorei Hook.f , Amaryllidaceae, 12

mduze (Z) bb; umduze (Z) bb; umNduze (Z) bb

sp., Amaryllidaceae, 12

mduze (Z) bb; umduze (Z) bb; umNduze (Z) bb

Crocosmia

aurea (Pappe ex Hook.) Planch., Iridaceae, 12

dwendweni (Z) bb; mlunge (Z) bb; udwendweni (Z) bb; umLunge (Z)

bb; unDwendweni (Z) bb

panicuiata (Klatt) Goldblatt. Iridaceae, 12

dwendweni (Z) bb; mlunge (Z) bb; udwendweni (Z) bb; umLunge (Z)

bb; unDwendweni (Z) bb

Crotalaria

globifera E.Mey., Fab.VPapilionoideae, 12

bhubhubhu (Z) rt; bububu (Z) rt; uBhubhubhu (Z) rt

spp.. Fab.VPapilionoideae, 30

uVelabahleke (Z) rt; velabahleka (Z) rt

Croton

gratissinius Burch, var. subgratissimus (Prain) Burtt Davy, Euphorbi-

aceae, 31

iLethi (Z) bk; mahlabekufeni (Z) bk; mahlabufeni (Z) 1/st; uMahla-

bekufeni (Z) bk

sylvaticus Hochst, Euphorbiaceae, 30

umaHlabekufeni (Z) bk; dubanhlosi (Z) bk; indumbadlozi (Z) bk;

indumbahlozi (Z) bk; mahlabekufeni (Z) bk; mahlabufeni (Z)

bk; minya (Z) bk; moolologa (S) bk; mzilanyoni (Z) bk; ndum-

badlozi (Z) bk; uMinya(Z) bk; umZilanyoni (Z) bk

Cryptocarya

latifolia Sond., Lauraceae, 25

mdlangwenya (Z) bk; mkhondweni (Z) bk; ndlangwenya (Z) bk; uMdlang-

wenya (Z) bk; umKhondweni (Z) bk

myrtifolia Stapf, Lauraceae, 25

mkondweni (Z) bk; umKhondweni (Z) bk

Cunonia capensis L., Cunoniaceae, 5

lulama (Z) bk; malulek (Z) bk; mlulama (Z) bk; umLulamomkhulu (Z)

bk

Cupressus sp.*, Cupressaceae, 19

abaNqonqosi (Z) bk; abaqonqosi (Z) rt

Curtisia dentata (Burm.f.) C.A.Sm., Cornaceae, 19

inJundumlahleni (Z) bk; jundu (Z) bk; mlahleni (Z) bk; tnlahlenisefile

(Z) bk; pepelangeni (Z) bk; umLahleni (Z) bk; umLahlenisefile

(Z) bk; umlatleni (Z) bk; uPhephelelangeni (Z) bk;

Cussonia spicata Thunb., Araliaceae, 4

msenge (Z) bk/st; musenzhe (V) bk/st; umSenge (Z) bk/st; usenge (Z)

bk/st

Cyanotis speciosa (L.f.) Hassk., Commelinaceae, 1

makhotigobile (Z) rt; uMakhothigobile (Z) rt

Cyathea dregei Kunze, Cyathaceae, 1 1

iNkomankoma (Z) rt; khomankhoma (Z) rt; nkomankoma (Z) rt

Cyathula sp., Amaranthaceae, 2

uBulawu (Z) bk

Cycnium racemosum Benth., Scrophulariaceae, 6

hlabalangane (Z) rt; shlabahlangane (Z) rt; uHlabahlangane (Z) rt

Cymbopogon

excavatus (Hochst.) Stapf ex Burtt Davy, Poaceae*, 4

isiQunga (Z) 1/st/rt; siqunga (Z) 1/st/rt

marginatus (Steud.) Stapf ex Burtt Davy, Poaceae+, 4

isiQunga (Z) 1/st/rt; siqunga (Z) 1/st/rt

plurinodis (Stapf) Stapf ex Burtt Davy, Poaceae+, 4

isiQunga (Z) 1/st/rt; siqunga (Z) 1/st/rt

spp., Poaceae+, 19

ihozawoza (Z) 1/st; iWozawoza (Z) 1/st; sukasamba (Z) 1/st; sukasiham-

ba (Z) 1/st; uSukasihambe (Z) 1/st; wozawoza (Z) 1/st

validus (Stapf) Stapf ex Burtt Davy, Poaceae+, 4

isiQunga (Z) 1/st/rt; siqunga (Z) 1/st/rt

Cyrtorchis arcuata (Lindt.) Schltr., Orchidaceae, 2

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt; phamba (Z) rt; phamba (Z)

Dalbcrgia obovata E.Mey., Fab.VPapilionoideae, 1

umZungulu (Z) st

Datura

metel L.*, Solanaceae, 1

iLoyi (Z) 1/st

stramonium L.*, Solanaceae, 1

iLoyi (Z) 1/st

Deinbollia oblongifolia (E.Mey. ex Am.) Radik., Sapindaceae, 31

iPhengulula (Z) rt; lulama (Z) bk; mlulama (Z) bk; pengulula (Z) rt;

phengulula (Z) rt; umLulamaomkhulu (Z) bk

Dianthus

basuticus Burtt Davy, Caryophyllaceae, 4

hlokwaletsela (S) wpl; hlokwana-latesela (S) wpl; ilokwalatsela (S)

wpl; tlokwana la tsela (S) wpl

crenatus Thunb., Caryophyllaceae, 2

iNingizimu (Z) wpl; mningizinru (Z) wpl

mooiensis F.N. Williams, Caryophyllaceae, 32

chambeswe (Z) wpl; chanibeswe (Z) wpl; hlokwaletsela (S) wpl; hlok-

wana-latesela (S) wpl; ilokwalatsela (S) wpl; tjanibeswe (Z)

wpl; tlokwana la tsela (S) wpl; tshanibezwe (Z) wpl; tshokha-

lasile (S) wpl; uTshanibezwe (Z) wpl; utyanibswe (Z) wpl

zeyheri Sond.

subsp. natalensis Hooper, Caryophyllaceae, 2

iNingizimu (Z) wpl; mningizimu (Z) wpl

subsp. zeyheri, Caryophyllaceae, 2

iNingizimu (S) wpl; mningizimu (S) wpl

Diaphananthe millarii (Bolus) H.P.Linder, Orchidaceae, 4

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt; phamba (Z) rt

Dicoma anomala Sond., AsteraceaeV 7

hloenya (S) rt; hlwenya (Z) rt; iNyongwane (Z) rt; nyongwane (Z) rt

Dierama pendulum (L.f.) Baker, Iridaceae, 11

dwendweni (Z) bb; isiDwa (Z) bb; udwendweni (Z) bb; unDwendweni

(Z) bb

Dietes iridioides (L.) Sweet ex Klatt, Iridaceae, 27

isiGqiki-sikatokoloshe (Z) 1/st; isiqunga sikatokoloshe (Z) 1/st; siqunga

si tokoloshe (Z) 1/st

Dioscorea

dregeana (Kunth) T.Durand & Scltinz var. hutchinsonii Burkill ined.,

Dioscoreaceae, 21

dakwa (Z) rt; mokamba (V) rt; sitshetsane (V) rt; ttDakwa (Z) rt

rupicola Kunth, Dioscoreaceae, 1

iMpiyampinya (Z) rt; mpiyampinya (Z) rt

sylvatiea (Kunth) Eckt. , Dioscoreaceae, 28

iNgwevu (Z) rt; sekolopatas (S) rt; skilpad (A) rt; skulpati (A) rt; uFudu

(Z) rt

Diospyros

galpinii (Hiern) De Winter, Ebenaceae, 26

dodemnyama (Z) rt; iMpishimpishi (Z) rt; inDodemnyama (Z) rt; ndo-

demnyama (Z) rt

villosa (L.) De Winter var. villosa, Ebenaceae, 26

dodemnyama (Z) rt; inDodemnyama (Z) rt; ndodemnyama (Z) rt

Disa spp., Orchidaceae, 3

hlamvu (Z) rt; iHlamvu (Z) rt

Dombeya rotundifolia (Hochst.) Planch, var. rotundifolia, Sterculi-

aceae, 20

ilukuluku (Z) bk; inHliziyonkulu (Z) bk; nhliziyonkulu (Z) bk

Drimia

elata Jacq., Liliaceae s.L, 41

isiKlenama (Z) bb; ndongwana zimbomvana (Z) bb; sekanama (S) bb;

skanama (S) bb; unDonganazibomvana (Z) bb

robusta Baker, Liliaceae s.l., 41

isiKlenama (Z) bb; ndongwana zimbomvana (Z) bb; sekanama (S) bb;

skanama (S) bb; unDonganazibomvana (Z) bb

Dryopteris

athamantica (Kunze) Kuntze, Aspidiaceae, 1 1

iNkomankoma (Z) rt; khomankhoma (Z) rt; nkomankoma (Z) rt

inaequalis (Schltdl.) Kuntze, Aspidiaceae, 11

iNkomankoma (Z) rt; khomankhoma (Z) rt; nkomankoma (Z) rt

Dumasia villosa DC. var. villosa, Fab.+/Papilionoideae, 15

kalimela (Z) 1/st; uKhalimele (Z) 1/st

Eclipta prostrata (L.) L.*, AsteraceaeV 10

iPhamephuce (Z) wpl; phamaphuce (Z) wpl; pamapuce (Z) wpl

Bothalia 31,1 (2001)

79

Ehretia rigida (Thunb.) Druce, Boraginaceae, 2

umHlele (Z) rt; umKele (Z) rt

Ekebergia capensis Sparrm., Meliaceae, 26

imanaye (Z) bk; isiManaye (Z) bk: mahlunzidintaba (Z) bk; ronyamati

(Z) bk: simanaye (Z) bk; uMathunzini{ Z) bk; uMathunzini-

wezintaba (Z) bk: umNyamathi (Z) bk

Elaeodendron

croceum (Thunb.) DC., Celastraceae. 15

isiNama (Z) bk; isiThundu (Z) bk; sahlula-amanye (Z) bk; sahlula-

manye (Z) bk; sehlulamanye (Z) bk; sithundu (Z) bk; snama (Z)

bk; umBovana (Z) bk; uSahlulamanye (Z) bk; uSehlulamanye

(Z) bk

transvaalense (Burtt Davy) R.H. Archer , Celastraceae, 37

gwavuma (Z) bk: iNgwavwna (Z) bk; mulumananma (V) bk; ngwavu-

ma (Z) bk

Elephantorrhiza elephantina (Burch.) Skeels, Fab.VMimosoideae, 28

dabu (Z) rt (It); inTolwane (Z) rt (It); mdabu (Z) rt (It); modabo (S) rt

(It); mosehlana (S) rt (It); ntolwane(Z) rt (It); umDabu (Z) rt (It)

Embelia ruminata (E.Mey. ex A. DC.) Mez , Myrsinaceae. 1

binini (Z) rt; isiBhinini (Z) rt

Encephalartos sp.. Zamiaceae, 2

isiGqikisomkhovu (Z) fL/st; isiKotshonkovu (Z) fl/st: kotsonkovu (Z)

fl/st

Entada rheedii Spreng., FabV/Mimosoideae, 12

inTindili (Z) sd; ntindili (Z) sd

Equisetum ramosissimum Desf. , Equisetaceae, 2

iShobalehashi (Z) rt; iShobalenyathi (Z) rt; shoba lenyathi (Z) rt

Eragrostis plana Nees , PoaceaeV 10

mvithi (Z) rt; umvete (Z) rt; umVithi (Z) rt

Eriosema

cordatum E.Mey.. Fab.VPapilionoideae, 13

qontsi (Z) rt; uQonsi (Z) rt

salignum E.Mey., Fab.VPapilionoideae. 7

bangalala (Z) rt: iQonsi (Z) rt; qontsi (Z) rt; uBangalala (Z) rt; uQonsi

(Z) rt

Eriospermum

cooperi Baker, Liliaceae 5./., 5

iNcamashela (Z) rt; ncamashela (Z) rt

luteo-rubrum Baker, Liliaceae s.l., 9

mathinta (Z) rt; matinta (Z) rt; uMathinta (Z) rt

mackenii (Hook.f.) Baker. Liliaceae s.l., 21

iNshlansula (Z) rt; ntsulansula (Z) rt

ornithogaloides Baker, Liliaceae s.l., 5

iNcamashela (Z) rt; ncamashela (Z) rt

spp.. Liliaceae s.l., 21

iNsulansula (Z) rt; ntsulansula (Z) rt

Erythrophleum lasianthum Corbishley, Lab.VCaesalpinioideae, 27

khwangu (Z) bk; mgwangu (Z) bk; mkhwangu (Z) bk; umKhwangu (Z)

bk

Eucalyptus sp. *, Myrtaceae, 16

dlavusa (Z) bk; dlebe (Z) bk; imPisikayihlangulwa (Z) bk; mdelebe (Z)

bk; mhlavusa (Z) bk; mpiskayi hlangulwa (Z) bk; mslavusa (Z)

bk; umDIavusa (Z) bk; umDlebe (Z) bk

Euclea

crispa (Thunb.) Guerke subsp. crispa, Ebenaceae, 1

iDungamuzi (Z) rt

divinorum Hiem, Ebenaceae, 9

mhlangula (Z) rt; mhlangulwa (Z) rt; mshekisani (Z) rt; sakchaan (Z)

rt; shekisane (Z) rt; umHlangula (Z) rt; umShekisane (Z) rt

natalensis A. DC., Ebenaceae, 2

iNkunzemnyama (Z) rt; nkunzi, black (Z/E) rt; nkunzi-emnyama (Z) rt;

nzimane (Z) rt

spp., Ebenaceae, 2

dungamuzi (Z) rt; iDungamuzi (Z) rt

undulata Thunb., Ebenaceae, 7

mshekisani (Z) rt; sakchaan (Z) rt; shekisane (Z) rt; umShekisane (Z) rt

Eucomis

autumnalis (Mill.) Chitt., Liliaceae s.l., 39

mathunga (Z) bb; uMathunga (Z) bb

bicolor Baker, Liliaceae s.l., 10

imBola (Z) bb; mbola (Z) bb; imbola (Z) bb

Eulophia

cucullata (Afzel. ex Sw.) Steud., Orchidaceae, 6

amabelejongosi (Z) rt; dwendweni (Z) rt; udwendweni (Z) rt; mabele-

jongosi (Z) rt; uMabelejongosi (Z) rt; unDwendweni (Z) rt

spp., Orchidaceae, 6

amabelejongosi (Z) rt; dwendweni (Z) rt; udwendweni (Z) rt; mabele-

jongosi (Z) rt: uMabelejongosi (Z) rt; unDwendweni (Z) rt

Euphorbia

bupleurifolia Jacq., Euphorbiaceae, 2

kalmasan (Z) wpl; inKamamasane (Z) wpl

natalensis Bernh., Euphorbiaceae, 2

kalmasan (Z) wpl; inKalamasane (Z) wpl

pulvinata Marl., Euphorbiaceae, 2

kalmasan (Z) wpl; inKamamasane (Z) wpl

woodii N.E.Br., Euphorbiaceae, 20

iNIdehle (Z) wpl; sihlenhle (Z) wpl

Faurea

macnaughtonii E. Phillips, Proteaceae, 1 1

isiSefu (Z) bk; sefo (Z) bk; sisefo (Z) bk

saligna Harv., Proteaceae, 11

isiSefu (Z) bk; sefo (Z) bk; sisefo (Z) bk

Ficus sur Forssk., Moraceae, 4

iNtombi kayibhinci (Z) bk; mkiwane (Z) bk; ntombi-kayibhinci (Z) bk;

umKhiwane (Z) bk

Foeniculum vulgare Mill.*, Apiaceae+, 13

iMbozisa (Z) 1/st; iMboziso (Z) 1/st; mbozisa (Z) 1/st; mboziso (Z) 1/st

Garcinia

gerrardii //an', ex Sim, Clusiaceae+, 8

rnbinda (Z) bk; wnBinda (Z) bk

livingstonei T.Anderson, Clusiaceaep 13

isaphulo (Z) bk; mphimbi (Z) bk: mupimbi (V) bk; saphulo (Z) bk;

sepula (Z) bk; umPhimbi (Z) bk

Gardenia

thunbergia Thunb., Rubiaceae, 1

umValasangweni (Z) rt; velasangweni (Z) rt

volkensii K.Schum. subsp. spatulifolia (Stapf & Hutch.) Verde., Rubi-

aceae, 1

umValasangweni (Z) rt; velasangweni (Z) rt

Gasteria croucheri (Hook.f.) Bake, r, Liliaceae s.l., 1

imPundu (Z) 1; pundu (Z) 1

Gerbera piloselloides (L.) Cass., AsteraceaeV 8

moya-wezwe (Z) st; uMoyawezwe (Z) st

Gerrardina foliosa Oliv., Flacourtiaceae, 31

iLethi (Z) bk; lulama (Z) bk; maleleka (X) bk; mlulama bk; umaLuleka

(Z) bk; umLulama (Z) bk

Gladiolus

dalenii Van Gee!, Iridaceae, 12

dwendweni (Z) bb; udwendweni (Z) bb; unDwendweni (Z) bb

sericeo-villosus Hook.f. subsp. sericeo-villosus, Iridaceae, 12

dwendweni (Z) bb; isiDwa (Z) bb; mlunge (Z) bb; udwendweni (Z) bb;

umLunge (Z) bb; unDwendweni (Z) bb

Gloriosa superba L., Liliaceae s.l., 3

hlamvu (Z) rt; iHlamvu (Z) rt

Glycyrrhiza glabra L. *, Fab. VPapilionoideae, 19

mlomo mnandi (Z) rt; umLomomnandomncane (Z) rt

Gnidia

burchellii ( Meisn .) Gilg, Thymelaeaceae, 13

dikili (Z) rt; isiDikili (Z) rt; mongadi (S) rt; sidikili (Z) rt

cuneata Meisn., Thymelaeaceae, 13

dikili (Z) rt; isiDikili (Z) rt; mongadi (S) rt; sidikili (Z) rt

kraussiana Meisn. var. kraussiana, Thymelaeaceae, 24

dikili (Z) rt; isiDikili (Z) rt; selengwe (Z) rt; sidikili (Z) rt; silawengwe

(Z) rt; umSilawengwe (Z) rt

spp., Thymelaeaceae, 14

dikili (Z) rt; isiDikili (Z) rt; sidikili (Z) rt

Gomphocarpus

fruticosus (L.) W.T.Aiton, Asclepiadaceae, 2

msingala salugazi (Z) rt; singala salugazi (Z) rt; uSingalwesalukazi (Z) rt

physocarpus E.Mey., Asclepiadaceae, 2

msingala salugazi (Z) rt; singala salugazi (Z) rt; uSingalwesalukazi (Z) rt

80

Bothalia 31,1 (2001)

Greyia sutherlandii Hook. & Har\>., Greyiaceae, 1

ubande (Z) bk; umBande (Z) bk

Gunnera perpensa L. , Haloragaceae, 3 1

gobo (Z) rt; gopho (Z) rt; tangazane (S) rt; uGobho (Z) rt; ugopo (Z) rt

Gymnosporia buxifolia (L.) Szyszyl ., Celastraceae, 34

bangalala (Z) rt; uBangalala (Z) rt

Haemanthus

albiflos Jacq., Amaryllidaceae, 2

uZeneke (Z) bb; zaneke (Z) bb

coccineus L., Amaryllidaceae, 2

uZeneke (Z) bb; zaneke (Z) bb

deformis Hook.f. , Amaryllidaceae, 2

uZeneke (Z) bb; zaneke (Z) bb

spp.. Amaryllidaceae, 2

uZeneke (Z) bb; zaneke (Z) bb

Halleria lucida L., Scrophulariaceae, 1

tshwalabenyoni (S) 1/st

Harpagophytum procumbens (Burch.) DC. ex Meisn., Pedaliaceae, 3

mziwalapho ungaphi (Z) sd; sumuso owalapha wayongaphi (Z) fr/rt;

umZiwalaphungaphi (Z) fr/rt

Harpephyllum caffrum Bernh. ex Krauss , Anacardiaceae, 1 1

gwenya (Z) bk; ngwenya (Z) bk; umGwenya (Z) bk

Haworthia

fasciata (Willd.) Haw.. Liliaceae s.l., 16

mathithibala (Z) wpl; uMathitliibala (Z) wpl

limifolia Marloth , Liliaceae s./., 16

mathithibala (Z) wpl; uMathithibala (Z) wpl

Helichrysum

acutatum DC., Asteraceae*, 9

uZangume (Z) rt; zangume (Z) rt

cymosum (L.) D.Don subsp. calvum Hilliard, Asteraceae+, 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) L/st

decorum DC., Asteraceae*, 35

iMphepho (Z) L/st; mpepo (Z) 1/st; mphepho (Z) 1/st

epapposum Bolus, Asteraceae+, 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) 1/st

gymnocomum DC., Asteraceae+, 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) 1/st

natalitium DC., Asteraceae+, 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) 1/st

nudifolium (L.) Less., Asteraceae+, 35

iMphepho (Z) 1/st; mpepo (Z) l/st; mphepho (Z) 1/st

odoratissimum (L.) Sweet, AsteraceaeL 35

iMphepho (Z) 1/st; mpepo (Z) 1/st; mphepho (Z) 1/st

Helinus integrifolius (Lam.) Kuntze, Rhamnaceae, 12

bhubhubhu (Z) rt; bububu (Z) rt; uBhubhubhu (Z) rt

Heliophila subulata Burch, ex DC., Brassicaceae, 8

moya wezwe (Z) rt; uMoyawezwe (Z) rt

Hermannia depressa N.E.Be, Sterculiaceae, 7

phate-ea-ngaka (S) rt; phateyangaka (S) rt

Heteropyxis natalensis Harv., Myrtaceae, I

thathayane (T) rt; umKhuze (Z) rt

Hippobromus pauciflorus (L.f) Radik., Sapindaceae, 25

fazi-tetayo (Z) rt; iqhume (Z) rt; isiPhahluka (Z) rt; lilathile (Z) rt;

mfazi-othethayo (X) rt; mfazitetio (Z) rt; qhume (Z) rt; spahlu-

ka (Z) rt; spatluga (Z) rt; ucume (Z) rt; umFazothethayo (Z) rt;

uQhume (Z) rt; uTiye (Z) rt

Hippocratea longipetiolata Oliv., Celastraceae, 34

bangalala (Z) rt; uBangalala (Z) rt

Homalium dentatum (Harv.) Warb., Flacourtiaceae, 1

iDlebeyendlovu (Z) bk; ndlebendlovu (Z) bk

Hydnora africana Thunb., Hydnoraceae, 21

mafumbuka (Z) rt; uMavumbuka (Z) rt

Hypericum aethiopicum Thunb. subsp. aethiopicum, Clusiaceae+, 7

isiVumelwane (Z) wpl; ntsukumbili (Z) wpl; sukumbili (Z) wpl;

unSukumbili (Z) wpl; uNsukumbili (Z) wpl; vumelewana (Z)

wpl

Hypoxis

colchicifolia Baker, Hypoxidaceae, 33

iLabatheka (Z) bb; iLabathekelimnyama (Z) bb; labateka (Z) bb

gerrardii Baker, Hypoxidaceae, 4

ikomfe (Z) bb; iNkomfe (Z) bb

hemerocallidea Fisch. & C.A.Mey., Hypoxidaceae, 4

ikomfe (Z) bb; iNkomfe (Z) bb

sp., Hypoxidaceae, 33

iLabatheka (Z) bb; labatheka (Z) bb

Ilex mitis (L.) Radik, var. mitis, Apocynaceae, 21

iPhuphuma (Z) st; isiDumo (Z) bk; phuphuma (Z) st; sdumo (Z) bk;

sedumo (T) bk; sidumo (Z) bk; tshidzungu (Sh) bk

Ipomoea

crassipes Hook., Convolvulaceae, 15

uVimbukhalo (Z) rt; vimbokhalo (Z) rt

spp., Convolvulaceae, 22

bhoqo (Z) rt; boqo (Z) rt; uBoqo (Z) rt; uBhoqo (Z) rt

Jatropha

hirsuta Hochst., Euphorbiaceae, 6

godide (Z) rt; uGodide (Z) rt

zeyheri Sond., Euphorbiaceae, 6

godide (Z) rt; uGodide (Z) rt

Justicia capensis Thunb., Acanthaceae, 11

iKhokhela (Z) rt; khokhela (Z) rt; kokela (Z) rt

Kalanchoe

crenata (Andr.) Haw., Crassulaceae, 15

mahogwe (Z) 1/st; uMahogwe (Z) 1/st

sp., Crassulaceae, 35

mvuthuza (Z) 1/st; umVuthuza (Z) 1/st; vuthuza (Z) 1/st

Kigelia africana (Lam.) Benth., Bignoniaceae, 13

mvongoti (Z) fr; umVongothi (Z) fr

Kniphofia uvaria (L.) Oken, Liliaceae s.l., A

iCacane (Z) rt

Knowltonia bracteata Harv. ex Zahlbr., Ranunculaceae, 35

mvuthuza (Z) rt; umVuthuza (Z) rt; vuthuza (Z) rt

Kohautia amatymbica Eckl. & Zeyh. , Rubiaceae, 3

umFanozacile (Z) rt

Ledebouria

cooperi (Hook.f.) Jessop, Liliaceae s.l., 1

cubudwana (Z) bb; iCubudwana (Z) bb

ovatifolia (Baker) Jessop, Liliaceae s.l., 1

cubudwana (Z) bb; iCubudwana (Z) bb

revoluta (L.f.) Jessop, Liliaceae s.l., 1

cubudwana (Z) bb; iCubudwana (Z) bb

Lichtensteinia interrupta (Thunb.) Sond., Apiaceae*, 4

iMbozisa (Z) rt; kapalanga (Z) rt; mbozisa (Z) rt; mkapalanga (Z) rt;

umKhalaphanga (Z) rt

Lippia javanica (Burm.f.) Spreng., Verbenaceae, 6

msuzwane (Z) 1/st; suzwane (Z) 1/st; umSuzwane (Z) 1/st; usuzwane (Z)

1/st

Lithospermum cinereum DC., Boraginaceae, 1

moshala-shuping (S) bb

Littonia modesta Hook., Liliaceae s.l., 2

hlamvu (Z) rt; iHlamvu (Z) rt

Lonchocarpus capassa Rolfe, Fab.VPapilionoideae, 1

impandezoviyo (Z) bk; wnBhandu (Z) bk; umPhanda (Z) bk

Lycopodium clavatum L., Lycopodiaceae, 1

nwelele (Z) wpl; unWele (Z) wpl

Macaranga capensis (Bail/.) Benth. ex Sim, Euphorbiaceae, 10

mphumalelo (Z) bk; umphumeleli (Z) bk; unOmpumelelo (Z) bk

Maesa lanceolata Forssk., Myrsinaceae, 17

denda (Z) rt; isiDenda (Z) rt; isiThende (Z) rt; maguqu (Z) rt; maququ

(Z) rt; sdenda (Z) rt; uMaguqu (Z) rt

Malva

parviflora L.*, Malvaceae, 2

phate-ea-ngaka (S) rt; phateyungaka (S) rt

verticcllata L. var. verticellata*, Malvaceae, 2

moshala-shuping (S) rt

Bothalia 31,1 (2001)

81

Margaritaria discoidea (Baill.) Webster, Euphorbiaceae, 2

khwangu (Z) rt; mgwangu (Z) rt; mkwangu (Z) rt; umKhwangu (Z) rt

Maytenus

acuminata (L.f.) Loes., Celastraceae, 31

isiNama (Z) bk; lulama (Z) bk; mlulama (Z) bk; snama (Z) bk; um-

Lulama-omkhulu (Z) bk

undata (Thunb.) Blakelock, Celastraceae, 30

dabulovalo (Z) bk; inDabulaluvalo (Z) bk; ndabula lovalo (Z) bk; nda-

bulovalo (Z) bk

Mentha aquatica L., Lamiaceae+, 1

iMbozisa (Z) 1/st; mbozisa (Z) 1/st

Mesembryanthemum sp.. Mesembryanthemaceae, 8

gibleweni (Z) st; ngebeleweni (Z) st; uGebeleweni (Z) st; iKhambi-

lamabulawo (Z) 1/st

Microgramma lycopodioides (L.) Copel., Polypodiaceae, 13

iKhambilezintwala (Z) wpl; ikhumbile-twala (Z) wpl; kambi le ntwala

(Z) wpl; khambi-lentwala (Z) wpl

Millettia sutherlandii Harw, Fab.VPapilionoideae, 1

umKhunye (X) rt

Mimosa

pigra L., Fab.VMimosoideae, 11

iMbuna (Z) 1/st: mbuna (Z) 1/st

pudica L. var. hispida Brenan, Fab.VMimosoideae, 1 1

iMbuna (Z) 1/st; mbuna (Z) 1/st

Mimusops

caffra E.Me y. ex DC.. Sapotaceae, 3

amaSethole (Z); masethole (Z) bk

obovata Sond., Sapotaceae, 3

amaSethole (Z); masethole (Z) bk

sp., Sapotaceae, 1

thunzikulu (Z) bk

Momordica foetida Schumach., Cucurbitaceae, 2

inTshungu (Z) 1/st

Monanthotaxis caffra (Sond.) Verde., Annonaceae, 6

mazwende (Z) rt; mgogentilanya (S) rt; uMazwende (Z) rt; umGogawe-

zinhlanva (Z) rt

Mondia whitei (Hook.f.) Skeels. Periplocaceae, 2

mondi (Z) rt; uMondi (Z) rt

Monsonia natalensis Knuth, Geraniaceae, 1

inagile (Z) rt; uNakile (Z) rt

Moraea spathulata (L.f.) Klatt, Iridaceae, 3

hlamvu (z) rt; iHlamvu (Z) rt

Mucuna coriacea Baker subsp. irritans (Burn Davy) Verde., Fab.V

Papilionoideae. 1

shiritsi (V) rt

Mundulea sericea (Willd.) A.Chev., Fab. '/Papilionoideae, 2

mukundandou (V) rt: masimba ndlovu (Z) rt; umSindandlovu (Z) rt

Myrica serrata Lam., Myricaceae, 30

iFethi (Z) bk; lulama (Z) bk; mlulama bk; umLulama (Z) bk

Myrothamnus flabellifolius (Sond.) Welw., Myrothamnaceae, 26

uVukwababa (Z) 1/st; uVukwabafile (Z) 1/st; vugamafde (Z) 1/st; vuka-

kwa-bafile (Z) 1/st; vukakwabafileyo (Z) 1/st

Mystacidium

capense (L.f.) Schltr., Orchidaceae, 3

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt; phamba (Z) rt

spp., Orchidaceae, 3

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt; phamba (Z) rt

Noltea africana (L.) Rchb.f, Rhamnaceae, 1

maleleka (X) bk; umaLuleka (Z) bk

Nuxia floribunda Benth., Loganiaceae, 30

lulama (Z) bk; mlulama (Z) bk; umLulama (Z) bk

Nymphaea

nouchali Burm.f, Nymphaeaceae, 6

iZibu (Z) wpl

spp.. Nymphaeaceae, 6

iZibu (Z) wpl

Ochna

holstii Engl., Ochnaceae, 11

espangu (Z) bk; isiBhanku (Z) bk; sibanku (Z) bk; spanku (Z) bk

natalitia (Meisn.) Walp., Ochnaceae, 9

isiThundu (Z) rt; madlozana (Z) rt; mahlozana (Z) rt; sithundu (Z) rt;

uMadlozana (Z) rt

Ocotea hullata (Burch.) E.Mey., Lauraceae, 33

nukani (Z) bk; uNukane (Z) bk

Olea woodiana Knobl., Oleaceae, 14

mhlwazimamba (Z) rt; mlwazi mamba (Z) rt; umHlwazimamba (Z) rt

Olinia

radiata J.Hofmeyr & E. Phillips, Oliniaceae, 12

mzaneno (Z) bk; umZaneno (Z) bk; zaneno (Z) bk

ventosa (L.) Cufod., Oliniaceae, 3

iN gobamakhosi (Z) rt

Ornithogalum longibracteatum Jacq., Filiaceae s.l., 18

mababaza (Z) bb; uMababaza (Z) bb

Osteospermum imbricatum L. subsp. nervatum (DC.) Norl. var.

nervatum, Asteraceae+, 8

inKuphuIana (Z) 1/st; mokuphule (S) 1/st; nkuphulana (Z) 1/st; nku-

phulwane (Z) 1/st; sebelele (S) 1/st

Osyridocarpus schimperianus (Hochst. ex A. Rich.) A. DC., Santala-

ceae, 8

malala (Z) 1/st; marobalo (S) 1/st; uMalala (Z) 1/st

Othonna natalensis Sch.Bip., Asteraceae\ 2

iNcama (Z) rt

Ozoroa spp., Anacardiaceae, 1

isiFice (Z) bk/ 1/ rt

Pachycarpus

rigidus E.Mey., Asclepiadaceae, 23

iShongwe (Z) rt; poho-tsehla (S) rt; pootshetla (S) rt; shongwe (Z) rt

spp., Asclepiadaceae, 23

iShongwe (Z) rt; shongwe (Z) rt

Pappea capensis Eckl. & Zeyh., Sapindaceae, 31

uVumebomvu (Z) rt; vuma, red (Z/E) rt

Parinari curatellifolia Planch, ex Benth., Chrysobalanaceae, 1

uBulawu (Z) bk

Pelargonium luridum (Andr.) Sweet, Geraniaceae, 7

eshaka (Z) rt: iShaqa (Z) rt; isihaga (Z) rt; shaqa (Z) rt; unYawol-

wenkukhu (Z) rt; uVendle(Z) rt; vendhle (Z) rt

Pellaea

calomelanos (Sw.) Link, Adiantaceae, 1 1

iNkomankoma (Z) rt/1; khomankhoma (Z) rt/1; nkomankoma (Z) rt/1

rufa A.F.Tryon, Adiantaceae, 4

iKhambi (Z) rt

sp., Adiantaceae, 1 1

iNkomankoma (Z) rt; khomankhoma (Z) rt; nkomankoma (Z) rt

Peltophorum africanum Sond., Fab.VCaesalpinioideae, 12

mosehla (T) bk; mosetla (T) bk; msehla (Z) bk; umSehle (Z) bk

Pentanisia prunelloides (Eckl. & Zeyh.) Walp. subsp. prunelloides,

Rubiaceae, 30

cimamlilo (Z) rt; geimamlilo (Z) rt; iCimamlilo (Z) rt; iCishamlilo (Z)

rt; setima mollo (S) rt

Peucedanum

magalismontanum Sond., ApiaceaeV 28

bheka (Z) wpl; bheka-mina (Z) wpl; bhekaminagedwa (S) wpl; iBheka

(Z) wpl

thodei Arnold, Apiaceae+, 4

iMpondovu (Z) 1/st; mpondofu (Z) 1/st

Phyllanthus meyerianus Miill.Arg., Euphorbiaceae, 31

iLethi (Z) bk

Phytolacca

dioica L., Phytolaccaceae, 5

mzimuka (Z) rt; umZimuka (Z) rt; zimuka (Z) rt

heptandra Retz., Phytolaccaceae, 1

poho-tsehla (S) rt; pootshetla (S) rt

Piliostigma thonningii (Schum.) Milne -Redh., Fab.VCaesalpinioideae, 1

nkolokotso (Sh)

82

Bothalia 31,1 (2001)

Pimpinella caffra Harv., Apiaceae+, 28

bheka (Z) wpl; bheka-mina (Z) wpl; iBheka (Z) wpl

Pinus sp.*, Pinaceae, 15

abaPhaphe (Z) bk

Pittosporum viridiflorum Sims, Pittosporaceae, 30

fusamfu (Z) bk; mfusamvu (Z) bk; umFusamvu (Z) bk

Plectranthus

grallatus Briq., Lamiaceae\ 10

mnyama we mpunzi (Z) rt; umNyamawempunzi (Z) rt

sp., Lamiaceae+, 14

kungwini (Z) rt; nkungwini (Z) rt; uNkungwini (Z) rt

Pleurostylia capensis (Turcz.) Oliv., Celastraceae, 21

morulela (S) bk; mthunyelelwa (Z) bk; mtunyelwa (Z) bk; murulela (V)

bk; rnurumelelwa (V) bk; umThumelela (Z) bk; umThunyelelwa

(Z) bk

Plumbago auriculata Lam., Plumbaginaceae, 5

mashinishini (Z) rt; mashwileshwile (Z) rt; shwilieshwili (Z) rt;

uMaswelisweli (Z) rt; uMatshintshine (Z) rt

Podocarpus henkelii Stapf ex Dallim. & Jacks.. Podocarpaceae, 19

abaNqonqosi (Z) bk; abaqonqosi (Z) bk; vulakuvaliwe (Z) bk

Polygala

fruticosa P.J.Bergius, Polygalaceae, 2

iThethe (Z) rt

gerrardii Chod., Polygalaceae, 12

ndegandhlela (Z) wpl; nqangandlela (Z) wpl; uNgqengendlela (Z) wpl

hottentotta Presl. Polygalaceae, 10

uZekane (Z) rt; zagane (Z) rt; zekane (Z) rt

macowanii Paiva. Polygalaceae, 11

ndegandhlela (Z) wpl; nqangandlela (Z) wpl; uNgqengendlela (Z) wpl

marensis Burn Davy , Polygalaceae, 12

ndegandhlela (Z) wpl; nqangandlela (Z) wpl; uNgqengendlela (Z) wpl

ohlendorfiana Eckl. cfi Zeyh., Polygalaceae, 12

ndegandhlela (Z) wpl; nqangandlela (Z) wpl; uNgqengendlela (Z) wpl

serpentaria Eckl. <£ Zeyh.. Polygalaceae, 12

ndegandhlela (Z) wpl; nqangandlela (Z) wpl; uNgqengendlela (Z) wpl

sp., Polygalaceae, 2

iThethe (Z) rt

Portulaca sp., Portulacaceae, 4

chisizwe (Z) st; shisizwe (Z) st; uShisizwe (Z) st

Priva cordifolia (L.f ) Druce var. abyssinica (Jaub. & Spach) Moldenke,

Verbenaceae, 31

isiNama (Z) rt; snama (Z) rt

Proboscidea fragrans (Lindl.) Decne., Martyniaceae, 1

umZungulwane (Z) fr

Protea gaguedi J.F.Gmel., Proteaceae, 1

Tshidzungu (V) fl/rt

Protorhus longifolia (Bernh.) Engl.. Anacardiaceae, 3

isiFico (Z) bk; isiFico-sehlathi (Z) bk; nhlangothi (Z) bk; umHIuthi (Z)

bk; umKhumizo (X) bk; umuThibomvu (Z) bk; unHkmgothi (Z)

bk

Prunus africana (Hook.f. ) Kalkm. , Rosaceae, 1 1

inYazangomelinmyama (Z) bk; nyazangoma, black (Z/E) bk; yazango-

ma, black (Z/E) bk

Psammotropha myriantha Sond., Aizoaceae, 4

iMphephotshani (Z) rt; mpepeshwane (Z) rt

Psoralea pinnata L., Fab.VPapilionoideae, 1

umHlonishwa (Z) it

Ptaeroxylon obliquum (Thunb.) Radik., Ptaeroxylaceae, 5

mthathi (Z) bk; umThathe (Z) bk

Pterocarpus angolensis DC., Fab.+/Papilionoideae, 2

umuvungazi (Z) bk; umVangazi (Z) bk; uvangasi (Z) bk

Pterocelastrus

echinatus N.E.Br., Celastraceae, 28

sahlula-amanye (Z) bk; sahlulamanye (Z) bk; sehlulamanye (Z) bk;

uSahlulamanye (Z) bk; uSehlulamanye (Z) bk

rostratus (Tliunb.) Walp., Celastraceae, 28

sahlula-amanye (Z) bk; sahlulamanye (Z) bk; sehlulamanye (Z) bk;

uSahlulamanye (Z) bk; uSehlulamanye (Z) bk

tricuspidatus (Lam.) Sond., Celastraceae, 28

sahlula-amanye (Z) bk; sahlulamanye (Z) bk; sehlulamanye (Z) bk;

uSahlulamanye (Z) bk; uSehlulamanye (Z) bk

Puliearia scabra (Thunb.) Druce, Asteraceae’, 2

isiThaphuka (Z) rt; stapuga (Z) rt

Ranunculus multifidus Forssk., Ranunculaceae, 1

ixapuzi (Sw) 1/st; izapuzi (Z) 1/st; uXhaphozi (Z) L/st

Rapanea melanophloeos (L.) Mez, Myrsinaceae, 37

inHluthe (Z) bk; mahpipa (Z) bk; maphipha (Z) bk; uMaphipha (Z) bk;

uMaphiphakluibalo (Z) bk; uVukakwabafilikhubalo (Z) bk

Raphionacme sp., Periplocaceae, 25

matanjana (Z) rt; uMathanjana (Z) rt

Rauvolfia caffra Sond., Apocynaceae, 5

hlabamanzi (Z) bk; khaba (Z) bk; khabamanzi (Z) bk; umHlambamanzi

(Z) bk; umkhabamasi (Z) bk

Rhamnus prinoides L’He'r., Rhantnaceae, 8

umYenye (Z) rt; uNyenya (Z) rt

Rhipsalis baccifera (J.S.Mill.) Steam, Cactaceae, 8

gibleweni (Z) st; ngebeleweni (Z) st; uGebeleweni (Z) st

Rhoicissus

rhomboidea (E.Mey. ex Harv.) Planch., Vitaceae, 1

umThwazi (X) rt

tridentata (L.f.) Wild & Drum, subsp. tridentata. Vitaceae, 26

isiNwazi (Z) rt; sinwazi (Z) rt; snwazi (Z) rt; umThwazi (Z) rt

Rhus chirindensis Baker f., Anacardiaceae, 24

dwendwenlencuba (Z) bk; hlokoshiyane (Z) bk; inHlokoshiyane (Z)

bk; nyazangoma, red (Z/E) bk; uDwendwelencuba (Z) bk;

umYazangomobomvu (Z) bk; yazangoma, red (Z/E) bk

Rhynchosia

nervosa Benth. & Harv. var. nervosa, Fab.VPapilionoideae, 1

madilika (Z) bk; uMadilika (Z) bk

spp., Fab.VPapilionoideae, 34

bangalala (Z) rt; kalimela (Z) 1/st; uBangalala (Z) rt; uKhalimele (Z)

1/st

Ricinus communis L.*, Euphorbiaceae, 1

mokhura (S) 1/st

Rubia cordifolia L. subsp. conotricha (Gaud.) Verde., Rubiaceae, 24

inTwalalubombo (Z) rt; mabone (Z) rt; ntalibombo (Z) rt; ntwalubom-

bo (Z) rt; rooi-storm (Afrikaans) rt

Rubus

pinnatus Willd., Rosaceae, 2

iJingijolo (Z) rt; jingijolo (Z) rt

rigidus J.E.Sm., Rosaceae, 2

iJingijolo (Z) rt; jingijolo (Z) rt

Rumex

crispus L.*, Polygonaceae, 4

dololenkonyan (Z) rt; iDololenkonyane (Z) rt

lanceolatus Thunb., Polygonaceae, 2

dololenkonyane (Z) rt; iDolo lenkonyane (Z) rt; ndolenkonyane (Z)

Salacia kraussii (Harv.) Harv., Celastraceae, 34

bangalala (Z) rt; uBangalala (Z) rt

Sandersonia aurantiaca Hook., Liliaceae s.i, 2

hlamvu (Z) rt; iHlamvu (Z) rt

Sarcophvte sanguinea Sparrm. subsp. sanguinea. Balanophoraceae,

20

mafumbuka (Z) rt; mavumbuka (Z) rt; uMavumbuka (Z) rt

Scabiosa columbaria L., Dipsacaceae, 28

bheka (Z) wpl; bheka-mina (Z) wpl; bhekaminagedwa (S) wpl; iBheka

(Z) wpl

Scadoxus puniceus (L.) Friis & Nordal, Amaryllidaceae, 5

mphompho (Z) bb; umPhompho (Z) bb

Schlechterina mitostemmatoides Harms, Passifloraceae, 7

hlahlanyosi, white (Z/E) wpl; hlalanyosi (Z) wpl; iHlalanyosi (Z) wpl

Scholia hrachypetala Sond., Fab.VCaesalpinioideae, 22

iHluze (Z) bk; umGxamu (Z) bk; uVovo (Z) bk

Bothalia 31,1 (2001)

83

Scilla

natalensis Planch., Liliaceae s.L, 39

guduza (Z) bb; inGuduza (Z) bb; nguduza (Z) bb

nervosa (Burch.) Jessop. Liliaceae s.l., 21

inGcino (Z) bb; ncino (Z) bb; ngcina (Z) bb; ngcino (Z) bb

Sclerocarya birrea (A. Rich.) Hochst. subsp. caffra (Sond.) Kokwaro,

Anacardiaceae, 28

mganu (Z) bk; umGanu (Z) bk

Scolopia mundii (Eckl. & Zeyh.) Warb., Flacourtiaceae. 10

mdwendwelencuba (Z) bk; uDwendwelwegcuba (Z) bk

Secamone gerrardii Harv. ex Benth., Asclepiadaceae, 22

iPhophoma (Z) rt; phophoma (Z) rt; popoma (Z) rt

Securidaca longepedunculata F resen., Polygalaceae, 2

iPhuphuma (Z) st; phuphuma (Z) st

Senecio

coronatus (Thunb.) Harv.. Asteraceae+, 24

balebatsa (S) rt; bulibazi (Z) rt; lepatsi (S) rt; libazi (Z) rt; molebatsa

(S) rt; ubuLibazi (Z) rt

gregatus Hilliard. Asteraceae*, 19

ntsukumbili (Z) 1/st; sukumbili (Z) 1/st; unSukumbili (Z) 1/st;

uNsukumbili (Z) 1/st

speciosus Willd.. AsteraceaeL 4

bohlololo (Z) wpl; iBohlololo (Z) wp

Sideroxylon inerme L.. Sapotaceae, 2

amaSethole (Z) bk; masethole (Z) bk; uMaphipha (Z) bk

Silene bellidioides Sond.. Caryophyllaceae. 13

gwayana (Z) wpl; isigwayana (Z) wpl; uGwayana (Z) wpl

Siphonochilus aethiopicus (Schweinf) B.L.Burtt. Zingiberaceae, 12

isiPhephetho (Z) rt; siphephetho (Z) rt; sphephetho (Z) rt

Sisyranthus sp., Asclepiadaceae, 3

wnFanozacile (Z) rt

Smilax anceps Willd.. Liliaceae s.l.. 5

inGqaqabulani (Z) rt; xaxabulane (Z) rt

Solanum

aculeastrum Dun., Solanaceae, 19

inTuma (Z) fr; ntuma (Z) fr; umThuma (Z) fr

hermannii Dun., Solanaceae, 19

inTuma (Z) fr; ntuma (Z) fr; umThuma (Z) fr

mauritianum Scop.*, Solanaceae, 13

bhoqo (Z) rt; boqo (Z) rt; uBoqo (Z) rt; uBhoqo (Z) rt; uGwayana (Z)

bk/st

panduriforme E.Mey., Solanaceae, 19

inTumemncane (Z) fr; ntuma (Z) fr

Spermacoce natalensis Hochst.. Rubiaceae, 27

inSulansula (Z) rt; isiNdiyandiya (Z) rt; mabophe (Z) rt; mahbhope (Z)

rt; ndyandiya (Z) rt; ntsulansula (Z) rt; sindiyandiya (Z) rt;

sulansula (Z) rt; uMabophe (Z) rt

Spilanthes mauritiana (Pers.) DC., Asteraceae+, 1

isiSinini (Z) 1/st; sinini (Z) 1/st

Spirostachys africana Sond., Euphorbiaceae, 15

mthombothi (Z) bk; tomboti (Z) bk; umThombothi (Z) bk

Stachys nigricans Benth., Lamiaceae+, 5

musa (Z) rt; uMusa (Z) rt

Stangeria eriopus (Kuntze) Baill., Stangeriaceae, 27

fingo (Z) rt (It); imFingo (Z) rt (It); mfingo (Z) rt (It)

Stapelia gigantea N.E.Br., Asclepiadaceae, 14

isililo (Z) wpl; maliliza (Z) wpl; uZililo (Z) wpl

Strophanthus

gerrardii Stapf, Apocynaceae, 1

hlungubendlovu (Z) fr; ubuHlungubendlovu (Z) fr

luteolus Codd, Apocynaceae, 1

hlungubendlovu (Z) fr; ubuHlungubendlovu (Z) fr

petersianus Klotzsch, Apocynaceae, 1

hlungubendlovu (Z) fr; ubuHlungubendlovu (Z) fr

speciosus ( Ward & Harv. ) Reber, Apocynaceae, 1

hlungu (Z) fr; isiHlungu (Z) fr; uhlungu (Z) fr

Strychnos

decussata (Pappe) Gilg, Loganiaceae, 1

lahlankosi (Z) bk; mlahlankosi (Z) bk; mpafa (Z) bk; umLahlankosi (Z)

bk

henningsii Gilg, Loganiaceae, 8

mkwalothi (Z) bk; nkawulothi (Z) bk; nkwalothi (Z) bk; umNono (Z)

bk; uManono (X) bk; umQalothi (Z) bk

mitis S. Moo re, Loganiaceae, 7

mkwalothi (Z) bk; nkawulothi (Z) bk; nkwalothi (Z) bk; umNono (Z)

bk; uManono (X) bk; umQalothi (Z) bk

spinosa Lam., Loganiaceae, 1

mhlahla (Z); mslasa (Z); umHIala (Z)

Synadenium cupulare (Boiss.) L.C.Wheeler, Euphorbiaceae, 3

mdlebe (Z) bk; umDlebe (Z) bk

Synaptolepis kirkii Oliv., Thymelaeaceae, 31

uVumomhlope (Z) rt; vuma, white (Z/E) rt

Tabernaemontana elegans Stapf. Apocynaceae, 2

Muhatu (V) rt/fr

Talinum caffrum (Thunb.) Eckl. & Zeyh., Portulacaceae, 33

punyu (Z) rt; punyuka (Z) rt; uMpunyu (Z) rt

Teedia lucida Rudolphi, Scrophulariaceae, 1

hloenya (S) rt; hlwenya (Z) rt

Tephrosia

marginella H.M. Forbes, Fab.'/Papilionoideae, 1

isiDala (Z) 1/st; sidala (Z) 1/st

spp., Fab.+/Papilionoideae, 1

iLozana (Z) rt; lozlana (Z) rt

Terminalia

phanerophlebia Engl. & Diels, Combretaceae, 9

amaNgwansundu (Z) rt; amaNgwamhlophe (Z) rt

sericea Burch, ex DC., Combretaceae, 9

amaNgwamhlophe (Z) rt

spp., Combretaceae, 9

amaNgwe (Z) rt

Tetradenia riparia (Hochst.) Codd, LamiaceaeL 1

iBoza (Z) 1/st

Thesium pallidum A. DC., Santalaceae, 35

mahesaka, red (Z/E) rt; maisaka, red (Z/E) rt; maysaka, red (Z/E) rt;

mysaka, red (Z/E) rt; uMahesakobomvu (Z) rt

Thespesia acutiloba (Baker f.) Exell & Mendonca, Malvaceae, 1

iPhuphuma (Z) st; phuphuma (Z) st

Tieghemia quinquenervia (Hochst.) Balle, Loranthaceae, 8

iPhakama (Z) wpl; phakama (Z) wpl

Toddaliopsis bremekampii I.Verd., Rutaceae, 1

iNtana (Z) bk

Tragia

meyeriana Miill.Arg., Euphorbiaceae, 34

bangalala (Z) rt; imBabazane (Z) rt; mbabazane (Z) rt; uBangalala (Z) rt

rupestris Sond., Euphorbiaceae, 2

imBabazane (Z) rt; mbabazane (Z) rt

Trema orientalis (L.) Blume, Ulmaceae, 2

muvungazi (Z) bk; umuvungazi (Z) bk; umVangazi (Z) bk; uvangasi

(Z) bk

Tribulus terrestris L, Zygophyllaceae, 1

sekanama (T) bb

Tricalysia lanceolata (Sond.) Burn Davy, Rubiaceae, 6

msuzwane (Z) 1/st; suzwane (Z) 1/st; umSuzwane (Z) Pst; usuzwane (Z) Pst

Trichilia

dregeana Sond., Meliaceae, 29

mkhuhlu (Z) bk; mutuhu (V) bk; uMathunzi (Z) bk; umKhuhlu (Z) bk;

vungulamoya (Z) bk; vungulomo (Z) bk

emetica Vahl, Meliaceae, 29

mkhuhlu (Z) bk; mutuhu (V) bk; uMathunzi (Z) bk; umKhuhlu (Z) bk;

vungulamoya (Z) bk; vungulomo (Z) bk

Tridactyle

bicaudata (Lindl.) Schltr., Orchidaceae. 4

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt; phamba (Z) rt

84

Bothalia 31,1 (2001)

Tridactyle (cont.)

tridentata (Harv.) Schltr., Orchidaceae, 4

ipamba (Z) rt; iPhamba (Z) rt; pamba (Z) rt: phamba (Z) rt

Trimeria

grandifolia ( Hochst .) Gilg, Flacourtiaceae, 5

iDlebelendlovu (Z) bk; ndlebelendlovu (Z) bk

trinervis Harv., Flacourtiaceae, 3

iDlebelendlovu (Z) bk; ndlebelendlovu (Z) bk

Triumfetta rhomboidea Jacq., Tiliaceae, 1

ndolenkonyane (Z) rt; inDolencane (Z) rt

l\ilbaghia

alliacea L.f., Liliaceae s.l., 25

iShaladilezinvoka (Z) bb; moilela (Z) bb; mwelela (Z) bb; umWelela

(Z) bb;

ludwigiana Harv., Liliaceae s.l., 22

moilela (Z) bb; mwelela (Z) bb; umWelela (Z) bb

simmleri P.Beauv., Liliaceae s.l., 22

moilela (Z) bb; mwelela (Z) bb; umWelela (Z) bb

spp., Liliaceae s.l., 3

iShaladilezinyoka (Z) bb

Turbina oblongata (E.Mey. ex Choisy) A. Meeuse, Convolvulaceae, 22

bhoqo (Z) rt; boqo (Z) rt; uBoqo (Z) rt; uBhoqo (Z) rt

Turraea

florihunda Hochst., Meliaceae, 25

lulama (Z) bk; madlozana (Z) rt; mahlozana (Z) rt; mlulama (Z) bk;

uMadlozana (Z) rt; umLulama (Z) bk; umVuma (Z) rt

obtusifolia Hochst., Meliaceae, 11

swazi (Z) 1/st; uMadlozanomncane (Z) 1/st; uSwazi (Z) 1/st; iKhambi

(Z) rt

Tyiophora flanaganii Schltr., Asclepiadaceae, 5

iNhlanhla (Z) rt; nhlanhla (Z) rt

Typha capensis (Rohrb.) N.E.Br., Typhaceae, 2

IBhuma (Z) rt

Urginea

altissima (L.f.) Baker, Liliaceae s.l., 27

isiKlenama (Z) bb; mahlokolosa (Z) bb; sekanama (S) bb; skanama (S)

bb; uMahlogolosi (Z) bb

delagoensis Baker, Liliaceae s.l., 28

isiKlenama (Z) bb; mahlanganisa (Z) bb; sekanama (S) bb; skanama

(S) bb; uMahlanganisa (Z) bb

macrocentra Baker, Liliaceae s.l., 27

isiKlenama (Z) bb; sekanama (S) bb; skanama (S) bb

sanguinea Schinz, Liliaceae s.l., 27

isiKlenama (Z) bb; sekanama (S) bb; skanama (S) bb

spp., Liliaceae s.l., 31

magoga (T) bb; mahlokolosa (Z) bb; skanama, white (S/E) bb; umahl-

ogolosi (Z) bb; uMahlongolozi (Z) bb; umHlabelo (Z) bb

Uvaria

caffra E.Mey. ex Sond., Annonaceae, 3

mazwende (Z) rt; uMazwendomnyama (Z) rt

lucida Benth. subsp. virens (N.E.Br.) Verde., Annonaceae, 3

mazwende (Z) rt; uMazwendomnyama (Z) rt

Vangueria infausta Burch, subsp. infausta. Rubiaceae, 1

ntswila (Ts) st

Vernonia

adoensis Sch.Bip. ex Walp., Asteraceae+, 29

inNyathelo (Z) rt; inYathelo (Z) rt; nyathelo (Z) rt; slonyane (Z) rt;

uHlonyane (Z) rt

natalensis Sch.Bip. ex Walp., AsteraceaeL 1

lahlankosi (Z) 1/rt; umLahlankosi (Z) 1/rt

tigna Klatt, Asteraceae+, 22

hlunguhlungu (Z) rt; uHlungu-lungu (Z) rt; umHlunguhlungu (Z) rt

Viscum verrucosum Harv., Viscaceae, 8

iPhakama (Z) wpl; phakama (Z) wpl; uphagama (Z) wpl

Vitellariopsis

dispar (N.E.Br.) Aubrev., Sapotaceae, 2

pumbulu (Z) bk; umPhumbulu (Z) bk

marginata (N.E.Br.) Aubrev., Sapotaceae, 2

pumbulu (Z) bk; umPhumbulu (Z) bk

Vitex

rehmannii Guerke, Verbenaceae, 13

mluthu (Z) 1; umLuthu (Z) 1

wilmsii Guerke var. reflexa (H. Pearson) Pieper, Verbenaceae, 13

mluthu (Z) 1; umLuthu (Z) 1

Wahlenbergia sp., Campanulaceae, 1

manolo (S) rt

Warburgia salutaris (Bertol.f.) Choiv., Canellaceae, 32

isiBhaha (Z) bk; sbaga (Z) bk; sbhaha (Z) bk

Widdringtonia nodiflora (L.) Powrie, Cupressaceae, 2

thaululo (V) rt

Withania somnifera (L.) Dun., Solanaceae, 5

bovimba (Z) rt; ubovimba (Z) rt; ubuVimbo (Z) rt

Wrightia natalensis Stapf, Apocynaceae, 2

Musunzi (V) rt

Ximenia

americana L., Olacaceae, 1

ntsekele (Sh) rt

caffra Sond. var. natalensis Sond., Olacaceae, 5

izimpande zomthunduluka (Z) rt; mthunduluka (Z) rt; umThunduluka

(Z) rt

Xysmalobium undulatum (L.) W.T. Alton, Asclepiadaceae, 23

isinga (Z) bk; iShinga (Z) rt; iShongwe (Z) rt; poho-tsehla (S) rt; poot-

shetla (S) rt; shongwe (Z) rt

Zantedeschia albomaculata (Hook. ) Bail!, subsp. albomacuiata. Ara-

ceae, 3

iLabathekelimhlope (Z) bb; labatheka, white (Z/E) bb

Zanthoxylum

capense (Thunb.) Harv., Rutaceae, 26

lunguntabele (Z) rt; mlungumabele (Z) rt; mnokwana (S) rt; mnung-

wane (Z) rt; nungwane (Z) rt; senokontaropa (S) rt; umNungu-

mabele (Z) rt; umNungwane (Z) rt

davyi (l.Verd.) Waterm., Rutaceae, 26

lunguntabele (Z) rt; mlungumabele (Z) rt; mnokwana (S) rt; mnung-

wane (Z) rt; nungwane (Z) rt; umNungumabele (Z) rt; umNung-

wane (Z) rt

Ziziphus mucronata Willd. subsp. mucronata, Rhamnaceae, 17

lahlankosi (Z) bk; mlahlankosi (Z) bk; mpafa (Z) bk; umHlahlankosi

(Z) bk; umPhafa (Z) bk

APPENDIX 2.— LEXICON OF VERNACULAR NAMES USED FOR PLANT SPECIES TRADED

MEDICINALLY ON THE WITWATERSRAND

After the vernacular name, the sequence of annotations is as follows:

abbreviation for language of vernacular name in brackets;

genus and species for vernacular name;

plant part used: bk = bark, bb = bulb, tuber, corm, 11 = flower, fr = fruit, 1 = leaf, rt = root, sd = seed, st = stem, wpl

= whole plant;

unknown 1-90 = vernacular names not identified.

*exotic taxa, including naturalized taxa.

Bothalia 31,1 (2001)

85

abaNqonqosi (Z)

see Nqonqosi, aba

abaPhaphe (Z)

see Phaphe, aba

abaqonqosi (Z)

Cupressus sp.* bk

Podocarpus henkelii bk

amabelejongosi (Z)

Eulophia cucullata rt

E. spp. rt

amaNgwamhlope (Z)

see Ngwamhlope, ama

amaNgwamnyama (Z)

see Ngwamnyama, ama

amaNgwansundu (Z)

see Ngwansundu, ama

amangwe, black (Z)

Cadaba natalensis bk

amaNgwe (Z)

see Ngwe, ama

amaSethole (Z)

see Sethole, ama

Babazane, im (Z)

Tragia meyeriana rt

T. rupestris rt

balebatsa (S)

Senecio coronatus rt

Bande, um (Z)

Greyia sutherlandii bk

bangalala omhlope (Z)

unknown 1 rt

Bangalala, u (Z) (also bangalala)

Corchorus asplenifolius rt

Eriosema saligna rt

Gymnosporia buxifolia rt

Hippocratea longipetiolata rt

Rhynchosia spp. rt

Salacia kraussii rt

Tragia meyeriana rt

Bani, u (Z)

Agapanthus africanus rt

A. campanulatus rt

A. praecox subsp. orientalis rt

Belejongosi, uma (Z)

Eulophia cucullata rt

E. spp. rt

Bethe, um (Z)

Clutia spp. rt

Bhaha, isi (Z)

Warburgia salutaris bk

Bhandu, um (Z)

Lonchocarpus capassa bk

Bhanku, isi (Z)

Ochna holstii bk

bheka-mina (Z)

Peucedanum magalismontanum wpl

Pimpinella caffra wpl

Scabiosa columbaria wpl

Bheka, i (Z) (also bheka)

Peucedanum magalismontanum wpl

Pimpinella caffra wpl

Scabiosa columbaria wpl

bhekaminagedwa (S)

Peucedanum magalismontanum wpl

Scabiosa columbaria wpl

Bhinini, isi (Z)

Embelia ruminata rt

Bhoqo, u (Z) (also bhoqo)

Ipomoea spp. rt

Solanum mauritianum rt

Turbina oblongata rt

Bhubhubhu, u (Z) (also bhubhubhu)

Crotalaria globifera rt

Helinus integrifolius rt

Bhucu, i (Z)

Bulbine alooides wpl

B. asphodeloides wpl

B. frutescens wpl

B. latifolia wpl

Bhuma, i (Z)

Typha capensis rt

Binda, um (Z)

Garcinia gerrardii bk

binini (Z)

Embelia ruminata rt

Bohlololo, i (Z) (also bohlololo)

Achyropsis avicularis wpl

Aptenia cordifolia var. cordifolia wpl

Senecio speciosus wpl

Bola, um (Z)

Eucomis bicolor bb

bonisele (Z)

Acalypha glandulifolia wpl

Corchorus confusus rt

Boqo, u (Z) (also boqo)

Ipomoea spp. rt

Solanum mauritianum rt

Turbina oblongata rt

Bovana, um (Z)

Elaeodendron croceum bk

bovimba (Z)

Withania somnifera rt

Boza, i (Z)

Tetradenia riparia 1, st

bububu (Z)

Crotalaria globifera rt

Helinus integrifolius rt

Bulawu, u (Z)

Parinari curatellifolia bk

Cyathula sp. bk

bulibazi (Z)

Senecio coronatus rt

Butha, i (Z) (also buthe)

Asparagus laricinus rt

A. setae eus rt

A. virgatus rt

Myrsiphyllum asparagoides rt

M. ramosissimum rt

byere (Sh)

Acalypha villicaulis rt

Cacane, i (Z)

Kniphofia uvaria rt

Cena, i (Z) (also cena)

Aloe maculata I

A. sp. 1

chambeswe (S)

Dianthus mooiensis subspp. wpl

chanibeswe (S)

Dianthus mooiensis subspp. wpl

chisizwe (Z)

Portulaca sp. rt

Cimamlili, i (Z) (also cimamlilo)

Pentanisia prunelloides subsp. prunel-

loides rt

Cishamlilo, i (Z)

Pentanisia prunelloides subsp. prunel-

loides rt

Cubudwana, i (Z) (also cubudwana)

Ledebouria cooperi bb

L. ovatifolia bb

L. revoluta bb

Cumane, um (Z)

Alberta magna bk

ewaningi (Z)

Capparis brassii rt

C. tomentosa rt

Dabingehlele, un (Z)

Basella paniculata

Dabu, um (Z) (also dabu)

Elephantorrhiza elephantina rt

Dabulaluvalo, in (Z)

Maytenus undata bk

dabulamava (Z)

unknown 2 wpl

dabulovalo (Z)

Maytenus undata bk

Dakane, um (Z) (also dakane)

Apodytes dimidiata subsp. dimidata rt

Dakwa, u (Z)

Dioscorea dregeana rt

Dala, isi (Z)

Tephrosia marginella 1, st

Dawoluthi, in (Z)

Belamcanda spp. rt

delawena (Z)

Asclepias cucullata 1, rt

Delunina, u (Z) (also delunina)

Asclepias cucullata rt

Denda, isi (Z) (also denda)

Maesa lanceolata rt

Dewehlati, um (Z)

Commiphora woodii bk

Dikili, isi (Z) (also dikili)

Gnidia burchellii rt

G. cuneata rt

G. kraussiana var. kraussiana rt

G. spp. rt

dira-ga-dibone (S)

Cephalaria humilis rt

Diyaza, un (Z)

Bersama stayneri bk

B. swinnyi bk

B. tysoniana bk

Dlandlovu, um (X) (also dlandlovu)

Bauhinia bowkeri rt

Dlavusa, um (Z) (also dlavusa)

Afzelia quanzensis bk

Eucalyptus sp.* bk

Dlebe, um (Z) (also dlebe)

Eucalyptus sp.* bk

Synadenium cupulare bk

Dlebelendlovu, i (Z)

Trimeria grandifolia bk

T. trinervis bk

Dlebeyendlovu, i (Z)

Blighia unijugata bk

Homalium dentatum bk

Dlololenkonyane, i (Z)

Rumex crispus rt

R. lanceolatus rt

Dlula, i (Z)

Begonia dregei rt

B. homonyma rt

dlutjane (Z)

Aster bakeranus rt

Dlutshana, u (Z)

Aster bakeranus rt

dlutshani (Z)

Aster bakeranus rt

Dodemnyama, in (Z) (also dodemnyama)

Diospyros galpinii rt

Diospyros villosa var. villosa rt

Dolencane, in (Z)

Triumfetta rhomboidea rt

dolenkonyane (Z)

Triumfetta rhomboidea rt

dololenkonyan (Z)

Rumex crispus rt

R. lanceolatus rt

Dondwani, um (Z) (also dondweni)

Coddia rudis rt

Dongana-zibomvana, un (Z)

Drimia elata bb

D. robusta bb

dubanhlosi (Z)

Croton sylvaticus bk

Dubu, um (Z)

Combretum caffrum rt

C. erythrophyllum rt

C. hereroense rt

C. kraussii rt

duganeswe (Z)

unknown 3 bk

dumaphantsi (Z)

unknown 4 bk

Dumbahlozi, in (Z)

Croton sylvaticus bk

86

Bothalia 31,1 (2001)

Dumo, isi (Z)

Ilex mitis var. mins bk

dungamuzi (Z)

Euclea spp. rt

Dungamuzi, i (Z)

Euclea crispa subsp. crispa rt

Euclea spp. rt

Dwa, isi (Z)

Gladiolus sericeo-villosus forma sericeo-

villosus bb

Dwendwelencuba, u (Z)

Rhus chirindensis bk

Dwendwelwegcuba, u (Z)

Scolopia mundii bk

Dwendweni, un (Z) (also dwendweni)

Crocosmia aurea bb

C. paniculata bb

Dierama pendulum bb

Eulophia cucullata rt

E. spp. rt

Gladiolus dalenii bb

G. sericeo-villosus forma sericeo-villosus

bb

dwendwenlencuba (Z)

Rhus chirindensis bk

eshaka (Z)

Pelargonium luridum rt

espangu (Z)

Ochna holstii bk

Fanozacile, urn (Z)

Kohautia amatymbica rt

Sisyranthus sp. rt

fazi-tetayo (Z)

Hippobromus pauciflorus rt

Fazothethayo, urn (Z)

Hippobromus pauciflorus rt

Feyenkawu, im (Z)

Ansellia africana rt

Fice, isi (Z)

Ozoroa spp. bk, 1, st

Fico, isi (Z)

Protorhus longifolia bk

Ficosehlati, isi (Z)

Protorhus longifolia bk

Fingo, im (Z) (also fingo)

Stangeria eriopus rt

Fithi, isi (Z)

Baphia racemosa bk

Fudu, u (Z)

Dioscorea sylvatica rt

Fusamvu, um (Z) (also fusamfu)

Pittosporum viridiflorum bk

Gadankawu, um (Z)

Albizia adianthifolia bk

Galagala, um (X) (also galagala)

Buxus macowanii 1, fl

B. natalensis 1, fl

Ganu, um (Z)

Sclerocarya birrea subsp. caffra bk

garpe (S)

unknown 5 bb

gashi (Z)

unknown 6 bb

Gazine, un (Z) (also gazine)

Bridelia cathartica bk

gcimamlilo (Z)

Pentanisia prunelloides subsp. prunel-

loides rt

Gcino, in (Z)

Albuca pachychlamys bb

Scilla nervosa bb

Gebeleweni, u (Z)

Mesembryanthemum sp. 1, st

Rhipsalis baccifera st

Gibisila, u (Z) (also gibisile)

Bowiea volubilis bb

gibleweni (Z)

Mesembryanthemum sp. 1, st

Rhipsalis baccifera st

gibonisele (Z)

Acalypha glandulifolia wpl

Corchorus confitsus rt

Gibonisele, u (Z)

Acalypha glandulifolia wpl

Gibonisele, un (Z)

Corchorus confusus rt

Goba, isi (Z)

Asparagus africanus rt

A. falcatus rt

A. sp. rt

Belmia reticulata rt

Gobandlovu, u (Z) (gobandlovu)

Balanites maughamii bk

Gobho, u (Z)

Gunnera perpensa rt

gobo (Z)

Gunnera perpensa rt

Godide, u (Z) (also godide)

Jatropha hirsuta rt

J. zeyheri rt

Gogawezinhlanya, um (Z)

Monanthotaxis caffra rt

gopho (Z)

Gunnera perpensa rt

Gqaqabulani, in (Z)

Smilax anceps rt

Gqikisikatokoloshe, isi (Z)

Dietes iridioides 1, st

Gqikisomkhovu, isi (Z)

Encephalartos sp. fl, st

Guduza, in (Z) (also guduza)

Scilla natalensis bb

gwavuma (Z)

Elaeodendron transvaalense bk

gwayana (Z)

Silene bellidioides wpl

Gwayana, u (Z)

Silene bellidioides wpl

Solanum mauritianum bk, st

Gwayana, i (Z)

Silene bellidioides wpl

Gwenya, um (Z) (also gwenya)

Harpephyllum caffrum bk

Gxamu, um (Z)

Scholia brachypetala bk

Hlabahlangane, u (Z)

Cycnium racemosum rt

hlabalangane (Z)

Cycnium racemosum rt

hlabamanzi (Z)

Rauvolfia caffra bk

hlabehlangan (Z)

Cycnium racemosum rt

Hlabekufeni, uma (Z)

Croton sylvaticus bk

Hlabelo, um (Z)

Urginea spp. bb

Hladlothi, um (X)

Albizia adianthifolia bk

hlahlabadimo (S)

Chenopodium ambrosioides * 1, st

Hlahlankosi, um (Z)

Ziziphus mucronata subsp. mucronata bk

hlahlanyosi, white (Z)

Schlechterina mitostemmatoides wpl

hlala kwabaFileyo (Z)

unknown 7 wpl

Hlala, um (Z)

Strychnos spinosa bk

Hlalamagwababa, um (Z)

Bridelia micrantha bk

hlalamakwahaba (Z)

Bridelia micrantha bk

hlalanyosi (Z)

Schlechterina mitostemmatoides wpl

hlalanyosi, red (Z)

unknown 8 rt

Hlalanyosi, i (Z)

Schlechterina mitostematoides wpl

Hlambamanzi, um (Z)

Rauvolfia caffra bk

Hlamvu, i (Z) (also hlamvu)

Callilepis laureola rt

Disa spp. rt

Gloriosa superba rt

Littonia modesta rt

Moraea spathulata rt

Sandersonia aurantiaca rt

Hlangothi, un (Z)

Protorhus longifolia bk

Hlangula, um (Z)

Euclea divinorum rt

Hlele, um (Z)

Ehretia rigida rt

Hliziyonkulu, in (Z)

Dombeya rotundifolia var. rotundifolia bk

hloenya (S)

Dicoma anomala rt

Teedia lucida rt

Hlokoshiyane, in (Z) (also hlokoshiyane)

Rhus chirindensis bk

hlokwaletsela (S)

Dianthus basuticus subspp. wpl

D. mooiensis subspp. wpl

hlokwana-Iatesela (S)

Dianthus basuticus subspp. wpl

D. mooiensis subspp. wpl

Hlonishwa, um (Z)

Psoralea pinnata rt

Hlonyane, u (Z)

Vernonia adoensis rt

Hlonyane, um (Z) (also hlonyane)

Artemisia afra 1, st

Cotula anthemoides 1, st

Hluguwendlovu, u (Z)

Strophanthus luteolus fr

Hlungu, isi (Z) (also hlungu)

Strophanthus speciosus fr

Hlungu-lungu, u (Z)

Vernonia tigna rt

hlungubendlovu (Z)

Strophanthus gerrardii fr

S. luteolus fr

S. petersianus fr

Hlungubendlovu, u (Z)

Strophanthus sp. fr

Hlungubendlovu, ubu (Z)

Strophanthus gerrardii fr

S. petersianus fr

hlunguhlungu (Z)

Vernonia tigna rt

Hlunguhlungu, um (Z)

Vernonia tigna rt

Hlungunyembe, i (Z) (also hlungunyembe)

Acokantliera oblongifolia rt, 1

A. oppositifolia rt, 1

Hlungwana, ubu (Z)

Aspilia natalensis l/st

Hluthi, um (Z)

Protorhus longifolia bk

Rapanea melanophloeos bk

Hluze, i (Z)

Schotia brachypetala bk

Hlwazi, um (Z)

Catha edulis bk, 1

Hlwazimamba, um (Z)

Olea woodiana rt

hlwenya (Z)

Dicoma anomala rt

Teedia lucida rt

iltheka (Z)

see Bheka, i

iBIiucu (Z)

see Bhucu, i

ilthuma (Z)

see Bhuma, i

Bothalia 31,1 (2001)

87

iBohlololo (Z)

see Bohlololo, i

ibonisele (Z)

Acalypha glandulifolia wpl

Corchorus confusus rt

iBoza (Z)

see Boza, i

ibuco (Z)

Bulbine alooides wpl

B. asphodeloides wpl

B. frutescens wpl

B. latifolia wpl

ibuqu (Z)

Bulbine alooides wpl

B. asphodeloides wpl

B. frutescens wpl

B. latifolia wpl

iButha (Z)

see Butha, i

iCacane (Z)

see Cacane, i

iCena (Z)

see Cena. i

iCimamlili (Z)

see Cimamlilo, i

iCishamlilo (Z)

see Cishamlilo, i

iCubudwana (Z)

see Cubudwana, i

iDlebelendlovu (Z)

see Dlebelendlovu, i

iDIebeyendiovu (Z)

see Dlebeyendlovu. i

iDlula (Z)

see Dlula, i

idlutshane (Z)

Aster bakeranus

iDololenkonyane (Z)

see Dlololenkonyane, i

iDungamuzi (Z)

see Dungamuzi, i

iGwayana (Z)

see Gwayana, i

iHlalanyosi (Z)

see Hlalanyosi, i

iHlamvu (Z)

see Hlamvu, i

iHlungunyembe (Z)

see Hlungunyembe, i

iHluze (Z)

see Hluze, i

ihozawoza (Z)

Cymbopogon spp. rt, st

ijikantambo (Z)

unknown 9 sd

Uingijolo (Z)

see Jingijolo, i

ikalamuzi (Z)

see Kalamuzi, i

iKhakhasi (Z)

see Khakhasi, i

iKhambi (Z)

see Khambi, i

iKhambilamabulavvo (Z)

see Khambilamabulawo, i

iKhambilezintwala (Z)

see Khambilezintwala, i

iKhathazo (Z)

see Khathazo, i

iKhokhela (Z)

see Kkokhela, i

ikhumbile-twala (Z)

Microgramma lycopodioides wpl

ikomfe (Z)

Hypoxis gerrardii bb

H. hemerocallidea bb

iLabatheka (Z)

see Labatheka, i

iLabathekelimhlophe (Z)

see Labathekelimhlophe, i

iLabathekelimnyama (Z)

see Labathekelimnyama, i

iletha (Z)

unknown 10

iLethi (Z)

see Lethi, i

ilokwalatsela (S)

Dianthus basuticus subspp. wpl

D. mooiensis subspp. wpl

iLoshana (Z)

see Loshana, i

iLoyi (Z)

see Loyi, i

iLozana (Z)

see Lozana, i

ilugulugu (Z)

unknown 1 1 bk

ilukuluku (Z)

Dombeya rotundifolia var. rotundifolia bk

imanaye (Z)

Ekebergia capensis bk

imBabazane (Z)

see Babazane, im

imbola (Z)

Eucomis bicolor bb

iMbozisa (Z)

see Mbozisa, i

iMboziso (Z)

see Mboziso, i

iMbuna (Z)

see Mbuna, i

imFeyenkawu (Z)

see Feyenkawu, im

iMfingo (Z)

see Fingo, im

imkoka (Z)

Abrus precatorius subsp. africanus sd

impandezoviyo (Z)

Lonchocarpus capassa bk

impendewa shaye (Z)

Adenia gummifera var. gummifera st

iMphepho (Z)

see Mphepho, i

iMphephotshani (Z)

see Mphephotshani, i

imPila (Z)

see Pila, im

imPindamshaye (Z)

see Pindamshaye, im

iMpishimpishi (Z)

see Mpishimpishi, i

imPiskayihlangulwa (Z)

see Piskayihlangulwa, im

iMpiyampinya (Z)

see Mpinyampinya, i

iMpondovu (Z)

see Mpondovu, i

imPundu (Z)

see Pundu, im

inagile (Z)

Monsonia natalensis rt

iNcama (Z)

see Ncama, i

iNcamashela (Z)

see Ncamashela, i

iNcotho (Z)

see Ncotho, i

incoto (Z)

Boophane disticha bb

inDabulaluvalo (Z)

see Dabulaluvalo, in

indakane (Z)

Apodvtes dimidiata subsp. dimidiata rt

inDawoluthi (Z)

see Dawoluthi, in

Indian kalmoes (E)

unknown 12 rt

Indian pills (E)

unknown 13 sd

indiyaza (Z)

Bersama stayneri bk

B. swinnyi bk

B. tysoniana bk

indlaba baloyi (Z)

unknown 14 bk

indlebendlovu (Z)

unknown 15 bb

inDodemnyama (Z)

see Dodemnyama, in

inDolencane (Z)

see Dolencane, in

indumbadlozi (Z)

Croton sylvaticus bk

inDumbahlozi (Z)

see Dumbahlozi, in

inGcino (Z)

see Gcino, in

iNgobamakhosi (Z)

see Ngobamakhosi, i

inGqaqabulani (Z)

see Gqaqabulani, in

inGuduza (Z)

see Guduza, in

iNgvvavuma (Z)

see Ngwavuma, in

iNgwevu (Z)

see Ngwevu, i

inhlanhla mpedi (Z)

unknown 16 rt

iNhlanhla (Z)

see Nhlanhla, i

iNhlehle (Z)

see Nhlehle, i

inHIiziyonkulu (Z)

see Hliziyonkulu, in

inHlokoshiyane (Z)

see Hlokoshiyane, in

iNingizimu (Z)

see Ningizimu, i

injundumlahleni (Z)

see Jundumlahleni, in

inKalamasane (Z)

see Kalamasane, in

inKamamasane (Z)

see Kamamasane, in

inkehli (Z)

Afzelia quanzensis bk

iNkomankoma (Z)

see Nkomankoma, i

iNkomfe (Z)

see Nkomfe, i

iNkunzemnyama (Z)

see Nkunzemnyama, i

inKupheyana (Z)

see Kupheyana, in

inKuphuIana (Z)

see Kuphulana, in

inNyathelo (Z)

see Nyathelo, in

inqgene (Z)

Aloe sp. 1

iNsulansula (Z)

see Nsulansula, i

iNtana (Z)

see Ntana, in

inTindili (Z)

see Tindili, in

inTolwane (Z)

see Tolwane, in

iNtombikayibhinci (Z)

see Ntombikayibhinci, i

inTshungu (Z)

see Tshungu, in

intshwalabenyoni (Z)

Burchellia bubalina bk

Bothalia 31,1 (2001)

inTuma (Z)

see Tuma, in

inTumemncane (Z)

see Tumerrmcane, in

inTwalalubombo (Z)

see Twalalubombo, in

inYathelo (Z)

see Yathelo, in

inYazangomelimnyama (Z)

see Yazangomelimnyama, in

inYokiziphinda (Z)

see Yokiziphinda, in

iNyongwane (Z)

see Nyongwane, i

ipamba (Z)

Balanites maughamii bk

Chlorophytum comosum wpl

Cyrtorchis arcuata rt

Diaphananthe millarii rt

Mystacidium capense rt

M. spp. rt

Tridactyle bicaudata rt

T. tridentata rt

iPhakama (Z)

see Phakama, i

iphamba lephanzi (Z)

unknown 17 bb

iPhamba (Z)

see Phamba, i

iphambala (Z)

unknown 18 bk

iPhengulula (Z)

see Phengulula, i

iPhophoma (Z)

see Phophoma, i

iPhuphuma (Z)

see Phuphuma, i

iqhume (Z)

Hippobromus pauciflorus rt

iQongqo (Z)

see Qongqo, i

iQonsi (Z)

see Qonsi, i

iQwaningi (Z)

see Qwaningi, i

isaphulo (Z)

Garcinia livingstonii bk

iShaladilezinyoka (Z)

see Shaladilezinyoka, i

iShaqa (Z)

see Shaqa, i

iShinga (Z)

see Shinga, i

iShobalehashi (Z)

see Shobalehashi, i

iShongwe (Z)

see Shongwe, i

isiBhaha (Z)

see Bhaha, isi

isiBhanku (Z)

see Bhanku, isi

isiBhinini (Z)

see Bhinini, isi

isiDala (Z)

see Dala, isi

isiDenda (Z)

see Denda, isi

isiDikili (Z)

see Dikili, isi

isiDumo (Z)

see Dumo, isi

isiDwa (Z)

see Dwa, isi

isiFice (Z)

see Fice, isi

isiFico (Z)

see Fico, isi

isiFicosehlathi (Z)

see Ficosehlati, isi

isiFithi (Z)

see Fithi, isi

isigiba-nyongo (Z)

Achyropsis avicularis wpl

isiGoba (Z)

see Goba, isi

isiGqikisikatokoloshe (Z)

see Gqikisikatokoloshe, isi

isiGqikisomkhovu (Z)

see Gqikisomkhovu, isi

isihaga (Z)

Pelargonium luridum rt

isiHlungu (Z)

see Hlungu, isi

isiKIenama (Z)

see Klenama, isi

isiKotshonkovu (Z)

see Kotshonkovu, isi

isililo (Z)

Stapelia gigantea wpl

isilumi (Z)

unknown 19 rt

isiManaye (Z)

see Manaye, isi

isiNama (Z)

see Nama, isi

isiNamasenyongo (Z)

see Namasenyongo, isi

isiNdiyandiya (Z)

see Ndiyandiya, isi

isinga (Z)

Xysmalobium undulation bk

isiNwazi (Z)

see Nwazi, isi

isiPhahluka (Z)

see Phahluka, isi

isiPhephetho (Z)

see Phephetho, isi

isiqunga sikatokoloshe (Z)

Dietes iridioides 1, st

isiQunga (Z)

see Qunga, isi

isiSefu (Z)

see Sefu, isi

isiSinini (Z)

see Sinini, isi

isiThaphuka (Z)

see Thaphuka, isi

isiThende (Z)

see Thende, isi

isiThundu (Z)

see Thundu, isi

isiVumelwane (Z)

see Vumelwane, isi

iThethe (Z)

see Thethe, i

iWozawoza (Z)

see Wozawoza, i

ixapuzi (Sw)

Ranunculus multifidus 1, st

izapuzi (Z)

Ranunculus multifidus 1, st

iZaza (Z)

see Zaza, i

iZibu (Z)

see Zibu, i

izimpande zomthunduluka (Z)

Ximenia caffra var. natalensis rt

Jingijolo, i (Z) (also jingijolo)

Rubus pinnatus rt

R. rigidus rt

Juluka, um (Z) (also juluka)

Casearia gladiiformis rt

jundu (Z)

Curtisia dentata bk

Jundumlahleni, in (Z)

Curtisia dentata bk

junundu (Z)

unknown 20 bk

Kalamasane, in (Z)

Euphorbia natalensis wpl

Kalamuzi, i (Z)

Acorus calamus rt

kalimele (Z)

Cissampelos torulosa 1, st

Dumasia villosa var. villosa 1, st

Rhynchosia spp. I, st

kalmasan (Z)

Euphorbia bupleurifolia wpl

E. natalensis wpl

E. pulvinata wpl

kalmoes (Z)

Acorus calamus rt

Kamamasane, in (Z)

Euphorbia bupleurifolia wpl

E. natalensis wpl

E. pulvinata wpl

kambi le ntwala (Z)

Microgramma lycopodioides wpl

kanyakude (Z)

Acacia xanthophloea bk

kapalanga (Z)

Lichtensteinia interrupta rt

katankawu (Z)

Albizia adianthifolia bk

kataza (Z)

Alepidea amatymbica var. amatymbica rt

A. longifolia rt

kathazo (Z)

Alepidea amatymbica var. amatymbica rt

A. longifolia rt

Kele, um (Z)

Ehretia rigida rt

khaba (Z)

Rauvolfia caffra bk

khabamanzi (Z)

Rauvolfia caffra bk

Khakhasi, i (Z)

Berkheya multijuga rt

Khalimeie, u (Z)

Cissampelos torulosa 1, st

Dumasia villosa var. villosa 1, st

Rhynchosia spp. 1, st

Khamba, um (Z)

Acacia sieberiana var. woodii bk

khambi-lentwala (Z)

Microgramma lycopodioides wpl

Khambi, i (Z)

Pellaea rufa rt

Turraea obtusifolia rt

Khambilamabulawo, i (Z)

Carpobrotus edulis 1, st

Mesembryanthemum sp. 1, st

Khambilezintwala, i (Z)

Microgramma lycopodioides wpl

Khanyakude, um (Z)

Acacia xanthophloea bk

Khaphalapanga, um (Z)

Lichtensteinia interrupta rt

Khathazo, i (Z) (also khatazo)

Alepidea amatymbica var. amatymbica rt

A. longifolia rt

khawulani (Z)

unknown 21 1, st

Khiwane, um (Z)

Ficus sur bk

Khokha, um (Z)

Abrus precatorius subsp. africanus sd

Khokhela, i (Z) (also khokhela)

Justicia capensis rt

khomankhoma (Z)

Clieilanthes hirta rt, 1

Cyathea dregei rt

Dryopteris athamantica rt

D.jnaequalis rt

Pellaea calomelanos rt, 1

P. sp. rt

Bothalia 31,1 (2001)

89

Khondweni, um (Z)

Cryptocarya latifolia bk

C. myrtifolia bk

Khuhlu, um (Z)

Trichilia dregeana bk

T. emetica bk

Khumizo, um (X)

Protorhus longifolia bk

Khunye, um (X)

Millettia sutherlandii rt

Khuze, um (Z)

Heteropyxis natalensis rt

Khwangu, um (Z) (also khwangu)

Acokanthera oppositifola 1, rt

Erythrophleum lasianthum bk

Margaritaria discoidea rt, bk

Klenama, isi (Z)

Drimia elata bb

D. robusta bb

Urginea altissima bb

U. delagoensis bb

U. macrocentra bb

U. sanguinea bb

kokela (Z)

Justicia capensis rt

komba (Z)

unknown 22 bb

Kotshonkovu, isi (Z)

Encephalartos sp. fl, st

kotsonkovu (Z)

Encephalartos sp. fl, st

kungwini (Z)

Plectranthus sp. rt

Kupheyana, in (Z)

Chrysanthemoides monilifera subsp.

monilifera wpl

Kuphulana, in (Z)

Osteospermum imbrication subsp. netva-

tum 1, st

labateka (Z)

Hypoxis colchicifolia bb

H. sp. bb

labatheka, white (Z)

Zantedeschia albomaculata subsp. albo-

maculata bb

Labatheka, i (Z)

Hypoxis colchicifolia bb

H. sp. bb

Labathekelimhlophe, i (Z)

Zantedeschia albomaculata subsp. albo-

maculata bb

Labathekelimnyama, i (Z)

Hypoxis colchicifolia bb

iahlankosi (Z)

Gerrardina foliosa bk

Vemonia natalensis 1, rt

Ziziphus mucronata subsp. mucronata bk

Lahlankosi, um (Z)

Gerrardina foliosa bk

Vemonia natalensis 1, rt

Lahleni, um (Z)

Curtisia dentata bk

Lahlenisefile, um (Z)

Curtisia dentata bk

lebone (Z)

unknown 23 rt

lehwama (S)

unknown 24 bb

lengana (S)

Artemisia afra 1, st

lengwana (S)

unknown 25 1, st

lepatsi (S)

Senecio coronatus rt

lesokwana (S)

Alepidea pilifera rt

A. setifera rt

Lethi, i (Z)

Croton gratissimus var. subgratissimus bk

Gerrardina foliosa bk

Myrica serrata bk

Phyllanthus meyerianus bk

letswelane (S)

unknown 26 rt

Libazi, ubu (Z) (also libazi)

Senecio coronatus rt

lilathile (Z)

Hippobromus pauciflorus rt

Lomomnandomncane, um (Z)

Anthospemium rigidum subsp. pumilum rt

Glycyrrhiza glabra rt

Loselina, u (Z) (also loselina)

Cinnamomum camphora bk

Loshana, i (Z) (also loshana)

Buddleja salviifolia rt

Loyi, i (Z)

Datura metel 1, st

D. stramonium * 1, st

Lozana, i (Z)

Tephrosia spp. rt

lozlana (Z)

Tephrosia spp. rt

lucky beans (E)

unknown 27 sd

lulama (Z)

Cunonia capensis bk

Deinbollia oblongifolia bk

Gerrardina foliosa bk

Maytenus acuminata var. acuminata bk

Myrica serrata bk

Nuxia floribunda bk

Turraea floribunda bk

Lulama, um (Z)

Deinbollia oblongifolia bk

Maytenus acuminata var. acuminata bk

Myrica serrata bk

Nuxia floribunda bk

Lulama-omncane, um (Z)

Turraea floribunda bk

Lulama-womfula, um (Z)

Gerrardina foliosa bk

Lulamomkhulu, um (Z)

Cunonia capensis bk

Luleka, uma (Z)

Gerrardina foliosa bk

Noltea africana bk

Lunge, um (Z)

Crocosmia aurea bb

C. paniculata bb

Gladiolus sericeo-villosus forma sericeo-

villosus bb

lungumabele (Z)

Zanthoxylum capense rt

Z. davyi rt

Luthu, um (Z)

Vitex rehmannii 1

V. wilmsii var. reflexa 1

lutjane (Z)

Aster bakeranus rt

Mababaza, u (Z) (also mababaza)

Omithogalum longibracteatum bb

Mabelejongosi, u (Z) (also mabelejongosi)

Eulophia cucullata rt

E. spp. rt

mabone (Z)

Rubia cordifolia subsp. conotricha rt

Mabopha, u (Z) (also mabophe)

Acridocarpus natalitius var. natalitius rt

A. natalitius var. linearifolius rt

Spermacoce natalensis rt

Mabusane, u (Z) (also mabusane)

Capparis brassii rt

C. tomentosa rt

Madilika, u (Z) (also madilika)

Rhynchosia nervosa var. nervosa bk

madiophalana (S)

unknown 28 rt

Madlozana, u (Z) (also madlozana)

Berchemia discolor rt

Ochna natalitia rt

Turraea floribunda rt

Madlozanomncane, u (Z)

Turraea obtusifolia rt

mafumbuka (Z)

Hydnora africana rt

Sarcophyte sanguinea subsp. sanguinea rt

magoga (T)

Urginea spp. bb

Maguqu, u (Z) (also maguqu)

Maesa lanceolata rt

mahbhope (Z)

Acridocarpus natalitius var. natalitius rt

A. natalitius var. linearifolius rt

Spermacoce natalensis rt

mahesaka, red (Z)

Thesium pallidum rt

Mahesakobomvu, u (Z)

Thesium pallidum rt

Mahesakomhlope, u (Z)

Agathosma ovata rt

mahirisaka, white (Z)

Agathosma ovata rt

Mahlabekufeni, u (Z) (also mahlabekufeni)

Croton gratissimus var. subgratissimus bk

C. sylvaticus bk

mahlabufeni (Z)

Croton gratissimus var. subgratissimus bk

mahlabufeni (Z) (cont.)

C. sylvaticus bk

Mahlanganisa, u (Z) (also mahlanganisa)

Urginea delagoensis bb

mahlokolosa (Z)

Urginea altissima bb

U. spp. bb

Mahlongolozi, u (Z)

Urginea altissima bb

U. spp. bb

mahlozana (Z)

Berchemia discolor rt

Ochna natalitia rt

Turraea floribunda rt

mahlunzidintaba (Z)

Ekebergia capensis bk

Mahogwe, u (Z) (also mahogwe)

Adenostemma caffra 1, st

A. viscosum 1, st

Kalanchoe crenata 1, st

mahpipa (Z)

Rapanea melanophloeos bk

maime (Z)

Brunsvigia sp. wpl

Clivia miniata var. miniata wpl

C. nobilis wpl

C. spp. wpl

maisaka, red (Z)

Thesium pallidum rt

maisaka, white (Z)

Agathosma ovata rt

maisaka, black (Z)

unknown 29 rt

Makhothigobile, u (Z) (also makhotigobile)

Cyanotis speciosa rt

Malala, u (Z) (also malala)

Jasminum angulare 1, st

Osyridocarpus schimperianus 1, st

maleleka (X)

Gerrardina foliosa bk

Noltea africana bk

malidalanga (Z)

unknown 30 bk

maliliza (Z)

Stapelia gigantea wpl

malubakupekwa (Sh)

unknown 3 1 rt

malulek (Z)

Cunonia capensis bk

90

Bothalia 31,1 (2001)

Manaye, isi (Z)

Ekebergia capensis bk

mankunkuniku (Z)

unknown 32 bk

manolo (S)

Wahlenbergia sp. rt

Manono, u (X) (also manono)

Strychnos henningsii bk

S. mitis bk

Manzamnyama, u (Z) (also

manzemnyama)

Anemone caffra rt

A. fanninii rt

maphipha ntelezi (Z)

Albuca fastigiata rt

A. nelsonii rt

Maphipha, u (Z) (also maphipha)

Rapanea melanophloeos bk

Maphiphakhubalo, u (Z)

Rapanea melanophloeos bk

Maphiphintelezi, u (Z)

Albuca fastigiata rt

A. nelsonii rt

maqadini (X)

Cassine peragua subsp. peragua bk

Maqadini, u (Z) (also maqadani)

Catunaregam spinosa subsp. spinosa fr

maququ (Z)

Maesa lanceolata rt

marobadibogale (S)

Aloe sp. 1

marohalo (S)

Osyridocarpus schimperianus 1, st

masethole (Z)

Mimusops caffra bk

M. obovata bk

Sideroxylon inerme bk

mashinishini (Z)

Plumbago auriculata rt

mashwileshwile (Z)

Plumbago auriculata rt

masimba ndlovu (Z)

Mundulea sericea rt

masunungule (V)

unknown 33 bk

Maswelisweli, u (Z)

Plumbago auriculata rt

matanjana (Z)

Raphionacme sp. rt

Mathanjana, u (Z)

Raphionacme sp. rt

Mathinta, u (Z) (also mathinta)

Eriospermum luteo-rubrum rt

Mathithibala, u (Z) (also mathithibala)

Aloe aristata wpl

Haworthia fasciata wpl

PI. limifolia wpl

Mathunga, u (Z) (also mathunga)

Eucomis autumnalis bb

Mathunzini, u (Z)

Ekebergia capensis bk

Mathunziwezintaba, u (Z)

Ekebergia capensis bk

matinta (Z)

Eriospermum luteo-rubrum rt

Matshintshine, u (Z)

Plumbago auriculata rt

Mavumbuka, u (Z)

Hydnora africana rt

Sarcophyte sanguinea subsp. sanguinea rt

Mayime, u (Z) (also mayime)

Brunsvigia sp. wpl

Clivia miniata var. miniata wpl

C. nobilis wpl

C. spp. wpl

mayisake, white (Z)

Agathosma ovata rt

maysaka, red (Z)

Thesium pallidum rt

mazwende (Z)

Monanthotaxis caffra rt

Uvaria caffra rt

U. lucida subsp. virens rt

Mazwende, u (Z)

Monanthotaxis caffra rt

Mazwendomnyama, u (Z)

Uvaria caffra rt

Uvaria lucida subsp. virens rt

mbabazane (Z)

Tragia meyeriana rt

T. rupestris rt

Mbesa, u (Z) (also mbeza)

Andrachne ovalis rt

mbhando (V)

Albizia adianthifolia bk

mbinda (Z)

Garcinia gerrardii bk

mbola (Z)

Eucomis bicolor bb

Mbozisa, i (Z) (also mhozisa)

Foeniculum vulgare 1, st

Lichtensteinia interrupta rt

Mentha aquatica 1, st

Mboziso, i (Z) (also mboziso)

Foeniculum vulgare 1, st

Mbuna, i (Z) (also mbuna)

Mimosa pigra 1, st

M. pudica var. hispida 1, st

mcaka (Z)

unknown 34 1, st

mchumane (Z)

Alberta magna bk

mdabu (Z)

Elephantorrhiza elephantina rt

mdlandlovu (X)

Bauhinia bowkeri rt

mdlangwenya (Z)

Cryptocarya latifolia bk

Mdlangwenya, u (Z)

Cryptocarya latifolia bk

mdlanlothi (Z)

Albizia adianthifolia bk

mdlavusa (Z)

Afzelia quanzensis bk

Eucalyptus sp.* bk

mdlebe (Z)

Eucalyptus sp. * bk

Synadeniwn cupulare bk

mdubu (Z)

Combretum cajfrum rt

C. erythrophyllum rt

C. hereroense rt

C. kraussii rt

mduza (Z) (also mduze)

Crinum delagoense bb

C. macowanii bb

C. moorei bb

mdwendwelencuba (Z)

Scolopia mundii bk

Membeza, u (Z)

Andrachne ovalis rt

memeza, white (Z)

Calodendrum capense bk

memezi, red (Z)

Cassipourea flanaganii bk

C. gerrardii bk

Memeziobomvu, u (Z)

Cassipourea flanaganii bk

C. gerrardii bk

Memezomhlope, u (Z)

Calodendrum capense bk

mf'azi-othethayo (X)

Plippobromus pauciflorus rt

mfazitctio (Z)

Plippobromus pauciflorus rt

mfingo (Z)

Stangeria eriopus rt

mfusamvu (Z)

Pittosporum viridiflorum bk

mgadankawu (Z)

Albizia adianthifolia bk

mganu (Z)

Sclerocarya birrea subsp. caffra bk

mgogentilanya (S)

Monanthotaxis caffra rt

mgwangu (Z)

Acokanthera oppositifola 1, rt

Erythropldeum lasianthum bk

Margaritaria discoidea rt, bk

mhlabelo (Z)

Blighia unijugata bk

mhlahla (Z)

Strychnos spinosa bk

mhlandloti (X)

Albizia adianthifolia bk

mhlangula (Z)

Euclea divinorum rt

mhlangulwa (Z)

Euclea divinorum rt

mhlavusa (Z)

Eucalyptus sp. * bk

mhlonyane (Z)

Artemisia afra 1, st

Cotula anthemoides 1, st

mhlwazi (Z)

Catha edulis bk, 1

mhlwazimamba (Z)

Olea woodiana rt

mhuluka (Z)

Barringtonia racemosa rt

minya (Z)

Croton sylvaticus bk

Minya, u (Z)

Croton sylvaticus bk

Minyela, u (Z)

Commiphora africana bk

C. harveyi bk

mjuluka (Z)

Casearia gladiiformis rt

mkadankawu (Z)

Albizia adianthifolia bk

mkanyakude (Z)

Acacia xanthophloea bk

mkapalanga (Z)

Lichtensteinia interrupta rt

mkhamba (Z)

Acacia sieberiana var. woodii bk

mkhapalanga (Z)

Lichtensteinia interrupta rt

mkhondweni (Z)

Cryptocarya latifolia bk

C. myrtifolia bk

mkhuhlu (Z)

Trichilia dregeana bk

T. emetic a bk

mkhwangu (Z)

Acokanthera oppositifola 1, it

Erythrophleum lasianthum bk

Margaritaria discoidea rt, bk

mkiwane (Z)

Ficus sur bk

mkwalothi (Z)

Strychnos henningsii bk

S. mitis bk

mkwangu (Z)

Acokanthera oppositifola 1, rt

Erythrophleum lasianthum bk

Margaritaria discoidea rt, bk

mlahleni (Z)

Curtisia dentata bk

mlahlenisefile (Z)

Curtisia dentata bk

mlomo ninandi (Z)

Anthospermum rigidum subsp. pumilum rt

Glycyrrhiza glabra rt

Bothalia 31,1 (2001)

91

mlulama (Z)

Cunonia capensis bk

Deinbollia oblongifolia bk

Gerrardina foliosa bk

Maytenus acuminata var. acuminata bk

Myrica serrata bk

Nuxia floribunda bk

Turraea floribunda bk

mlunge (Z)

Crocosmia aurea bb

C. paniculata bb

Gladiolus sericeo-villosus forma sericeo-

villosus bb

mlungendodeni (Z)

unknown 35 bb

mlungumabele (Z)

Zanthoxylum capense rt

Z. davyi rt

mluthu (Z)

Vite.x rehmannii 1

V. wilmsii var. reflexa 1

mlwazi mamba (Z)

Ole a woodiana rt

mmolate (S)

unknown 36 bk

mningizimu (Z)

Dianthus crenatus wpl

D. zeyheri subspp. wpl

mnokwana (S)

Zanthoxylum capense rt

Z. davyi rt

mnungwane (Z)

Zanthoxylum capense rt

Z. davyi rt

mnyama we mpunzi (Z)

Plectranthus grallatus rt

mnyamati (Z)

Ekebergia capensis bk

moakaila ( V )

unknown 37 st

moapatladi (S)

Capparis brassii rt

C. tomentosa rt

mobade (V)

unknown 38 bk

modabo (S)

Elephantorrhiza elephantina rt

mohato ( S )

Berkheya onopordifolia var. onopordifolia

rt

moilela (Z)

Tulbaghia alliacea bb

T. ludwigiana bb

T. simmleri bb

mokamba (V)

Dioscorea dregeana rt

mokhura (S)

Ricinus sp. 1, st

mokuphule (S)

Osteospermum imbrication subsp. nervatum

1, st

molebatsa (S)

Senecio coronatus rt

momotheka (S)

unknown 39 rt

Mondi, u (Z) (also mondi)

Cinnamomum zeylancium bk

Mondia whitei bk

mongadi (S)

Gnidia burchellii rt

moolologa (S)

Croton sylvaticus bk

moopatladi (S)

Capparis brassii rt

C. tomentosa rt

morensoeng (S)

unknown 40 rt

morulela (S)

Pleurostylia capensis bk

mosaye (S)

unknown 41 bk

mosehla (T)

Peltophorum africanum bk

mosehlana (S)

Elephantorrhiza elephantina rt

mosetla (T)

Peltophorum africanum bk

moshala-shuping (S)

Lithospermum cinereum rt

Malva verticellata var. verticellata rt

Moyawezwe, u (Z) (also moya-wezwe)

Gerbera piloselloides st

Heliophila subulata rt

mpafa (Z)

Ziziphus mucronata subsp. mucronata bk

mpendulo (Z)

Acalypha villicaulis rt

mpepeshwane (Z)

Psammotropha myriantha rt

mpepo (Z)

Achyrocline stenoptera 1, st

Helichrysum cymosum subsp. calvum 1, st

H. decorum 1, st

H. epapposum 1, st

H. gymnocomum 1, st

H. natalitium 1. st

H. nudifolium 1, st

H. odaratissimum 1, st

mpheabonwe (S)

Cephalaria humilis rt

Mphepho, i (Z) (also mphepho)

Achyrocline stenoptera 1, st

Helichrysum cymosum subsp. calvum 1, st

H. decorum 1, st

H. epapposum 1, st

H. gymnocomum 1, st

H. natalitium 1, st

H. nudifolium 1, st

H. odaratissimum 1, st

Mphephotshani, i (Z)

Psammotropha myriantha rt

mphimbi (Z)

Garcinia livingstonei bk

mphompho (Z)

Scadoxus puniceus bb

mphumalelo (Z)

Macaranga capensis bk

Mpikayiboni, u (Z) (also mpikayiboni)

Cephalaria humilis rt

mpila (Z)

Callilepis laureola rt

mpindamshaye (Z)

Adenia gummifera var. gummifera st

Mpinyampinya, i (Z)

Dioscorea rupicola rt

Mpishimpishi, i (Z)

Diospyros galpinii rt

mpiskayi hlangulwa (Z)

Eucalyptus sp. * bk

mpiyampinya (Z)

Dioscorea rupicola rt

Mpondovu, i (Z) (also mpondofu)

Peucedanum thodei 1, st

Mpunyu, u (Z)

Talinum caffrum rt

mqonga (X)

Cassipourea gerrardii bk

msehla (Z)

Peltophorum africanum bk

msenge (Z)

Cussonia spicata bk, st

mshekisani (Z)

Euclea divinorum rt

E. undulata rt

msingala salugazi (Z)

Asclepias fruticosa rt

A. physocarpa rt

mslangoti (Z)

Albizia adianthifolia bk

mslasa (Z)

Strychnos spinosa bk

mslavusa (Z)

Eucalyptus sp. * bk

msuzwane (Z)

Lippia javanica 1, st

Tricalvsia lanceolata 1, st

mthathi (Z)

Ptaeroxylon obliquum bk

mthole (Z)

Acacia caffra bk

mthombothi (Z)

Spirostachys africana bk

mthunduluka (Z)

Ximenia caffra var. natalensis rt

mthunyelelwa (Z)

Pleurostylia capensis bk

mtoto (Z)

unknown 42 bk

mtunyelwa (Z)

Pleurostylia capensis bk

muangata (V)

unknown 43 rt

muhatu (V)

Tabemaemontana elegans rt, fr

mukundandou (V)

Mundulea sericea rt

Mululuka, u (Z) (also mululuka)

Barringtonia racemosa rt

mulumananma (V)

Elaeodendron transvaalense bk

munyu (Z)

unknown 44 rt

mupimbi (V)

Garcinia livingstonei bk

murulela (V)

Pleurostylia capensis bk

murumelelwa (V)

Pleurostylia capensis bk

Musa, u (Z) (also musa)

Crabbea hirsuta rt

musenzhe (V)

Cussonia spicata bk, st

musinde (V)

unknown 45 bk

musunzi (V)

Wrightia natalensis rt

mutavhatsindi (V)

Brackenridgea zanguebarica rt

mutshutshunga (V)

Bridelia micrantha bk

mutuhu (V)

Trichilia dregeana bk

mutzene (V)

unknown 46 bk

muvhungo (V)

unknown 47 rt

muvungazi (Z)

Trema orientalis bk

mvhakata (Sh)

unknown 48 rt

mvithi (Z)

Boscia albitrunca rt

B. foetida subsp. rehmanniana rt

Eragrostis plana rt

mvongoti (Z)

Kigelia africana fr

mvuthuza (Z)

Kalanchoe sp. 1, st

Knowltonia bracteata 1, st

mwelela (Z)

Tulbaghia alliacea bb

T. ludwigiana bb

T. simmleri bb

mysaka, red (Z)

Thesium pallidum rt

92

Bothalia 31,1 (2001)

mysaka, white (Z)

Agathosma ovata rt

mzaneno (Z)

Olinia radiata bk

mzilanyoni (Z)

Croton sylvaticus bk

mzimuka (Z)

Phytolacca dioica rt

mziwalapha ungaphi (Z)

Harpagophytum procumbens rt, sd

mzunya (Z)

unknown 49 rt

Nakile, u (Z)

Monsonia natalensis rt

Nama, isi (Z)

Elaeodendron croceum bk

Maytenus acuminata bk

Priva cordifolia var. abyssinica rt

Namasenyongo, isi (Z)

Achyropsis avicularis wpl

Ncama, i (Z)

Othonna natalensis rt

Ncamashela, i (Z) (also ncamashela)

Eriospermum cooperi rt

E. omithogaloides rt

ncino (Z)

Albuca pachychlamys bb

Scilla nervosa bb

Ncotho, i (Z)

Boophane disticha bb

ndaba-ugehlele (Z)

Basella paniculata 1, st

ndabula lovalo (Z)

Maytenus undata bk

ndabulovalo (Z)

Maytenus undata bk

ndauiuti (Z)

Belamcanda spp. rt

ndengandhlela (Z)

Polygala confusa rt

P. gerrardii rt

P. marensis rt

P. ohlendorfiana rt

P. serpentaria rt

Ndiyandiya, isi (Z)

Bersama lucens bk

B. stayneri bk

B. swinnyi bk

B. tysoniana bk

Spermacoce natalensis bk

ndiyaza (Z)

Bersama stayneri bk

B. swinnyi bk

B. tysoniana bk

ndlangwenya (Z)

Cryptocarya latifolia bk

ndlebelendlovu (Z)

Trimeria grandifolia bk

T. trinervis bk

ndlebendlovu (Z)

Homalium dentatum bk

ndodemnyama (Z)

Diospyros galpinii rt

D. villosa var. villosa rt

ndolenkonyane (Z)

Triumfetta rhomboidea rt

ndongwana zimbomvana (Z)

Drimia elata bb

D. robusta bb

ndumbadlozi (Z)

Croton sylvaticus bk

Nduze, um (Z)

Crinum bulbispermum bb

C. delagoense bb

C. macowanii bb

C. moorei bb

C. sp. bb

Nga, uiTiu (Z)

Acacia karroo bk

Ngamanzi, um (Z)

Acacia robusta subsp. clavigera bk

A. robusta subsp. robusta bk

ngazine (Z)

Bridelia cathartica bk

ngcina (Z)

Albuca pachychlamys bb

Scilla nervosa bb

ngcino (Z)

Albuca pachychlamys bb

Scilla nen’osa bb

ngebeleweni (Z)

Mesembryanthemum sp. 1, st

Rhipsalis baccifera st

Ngobamakhosi, i (Z)

Olinia ventosa rt

ngoto (Z)

Boophane disticha bb

Ngqengendlela, u (Z)

Polygala confusa rt

P. gerrardii rt

P. marensis rt

P. ohlendorfiana rt

P. serpentaria rt

ngubozohlonya (Z)

Boophane disticha bb

nguduza (Z)

Scilla natalensis bb

ngwahma (X)

unknown 50 bk

Ngwaleni, u (Z)

Clutia hirsuta 1, rt

C. platyphylla 1, rt

C. pulchella 1, rt

Ngwamhlope, ama (Z)

Terminalia phanerophlebia rt

T. sericea rt

Ngwamnyama, ama (Z)

Cadaba natalensis bk

Ngwansundu, ama (Z)

Terminalia phanerophlebia rt

Ngwavuma, in (Z) (also ngwavuma)

Elaeodendron transvaalense bk

Ngwe, ama (Z)

Terminalia spp. rt

ngwedlane (Z)

Clutia hirsuta 1, rt

C. platyphylla 1, rt

C. pulchella 1, rt

ngwenya (Z)

Harpephyllum caff rum bk

Ngwevu, i (Z)

Dioscorea sylvatica rt

nhlangothi (Z)

Protorhus longifolia bk

nhlanhla omhlope (Z)

unknown 51 rt, st

Nhlanhla, i (Z) (also nhlanhla)

Tylophora flanaganii rt

Nhlehle, i (Z)

Euphorbia woodii wpl

nhliziyonkulu (Z)

Dombeya rotundifolia var. rotundifolia bk

nhlungunyembe (Z)

Acokanthera oblongifolia rt, 1

A. oppositifolia rt, 1

Ningizimu, i (Z)

Dianthus crenatus wpl

D. zeyheri subspp. wpl

nkawulothi (Z)

Strychnos henningsii bk

S. mitis bk

nkehle (Z)

Afzelia quanzensis bk

nkolokotso (Sh)

Piliostigma tlionningii bk

Nkomankoma, i (Z) (also nkomankoma)

Cheilanthes hirta rt. 1

Cyathea dregei rt

Dryopteris athamantica rt

D. inaequalis rt

Pellaea calomelanos rt, 1

P. sp. rt

Nkomfe, i (Z)

Hypoxis gerrardii bb

H. hemerocallidea bb

Nkungwini, u (Z) (also nkungwini)

Plectranthus sp. rt

Nkunzemnyama, i (Z)

Euclea natalensis

nkunzi, black (Z)

Euclea natalensis rt

nkunzi-emnyama (Z)

Euclea natalensis rt

nkuphulana (Z)

Osteospermum imbrication subsp. nen’atum

1, st

nkuphulwane (Z)

Osteospermum imbricatum subsp. neivatum

1, st

nkwalothi (Z)

Strychnos henningsii bk

S. mitis bk

nogimpinda (Z)

Boscia albitrunca rt

nokoloka (Sh)

unknown 52 rt

nomanyama (Sh)

Cassia abbreviata subsp. beareana bk

nomuhlahli (Z)

unknown 53 rt

Nono, um (Z)

Strychnos henningsii bk

S. mitis bk

nqangandlela (Z)

Polygala confusa rt

P. gerrardii rt

P. marensis rt

P. ohlendorfiana rt

P. serpentaria rt

Nqonqosi, aba (Z)

Cupressus sp.* bk

Podocarpus henkelii bk

nqupulwane (Z)

Chrysanthemoides monilifera subsp. monili-

fera wpl

Nsukumbili, u (Z)

Chenopodium ambrosioides 1, st

Hypericum aethiopicum subsp. aethiopicum

1, st

Senecio gregatus 1, st

S. serratuloides var. gracilis 1, st

Nsulansula, i (Z)

Eriospermum mackenii rt

E. sp. rt

Spermacoce natalensis rt

nswila (Sh)

unknown 54 st

ntabazimbi (V)

Brackenridgea zanguebarica rt

ntalibombo (Z)

Rubia cordifolia subsp. conotricha rt

Ntana, in (Z)

Toddaliopsis bremekampii bk

ntantanyana (Z)

Crabbea hirsuta rt

ntemulula (Z)

unknown 55 bk

ntindili (Z)

Entada rlieedii sd

ntolwane (Z)

Elephantorrhiza elephantina rt

Ntombikayibhinci, i (Z) (also ntombi-kay-

ibhinci)

Ficus sur bk

ntsekele (Sh)

Ximenia americana rt

Bothalia 31,1 (2001)

93

ntsontane (Z)

unknown 56 rt

ntsukumhili (Z)

Chenopodium ambrosioides* 1, st

Hypericum aethiopicum subsp. aethiopicum

1, st

Senecio gregatus 1, st

S. serratuloides var. gracilis 1, st

ntsulansula (Z)

Eriospennum mackenii rt

E. sp. rt

Spermacoce natalensis rt

ntswila (Ts)

Vangueria infausta subsp. infausta st

ntuma (Z)

Solarium aculeastrum fr

S. hermannii fr

5. panduriforme fr

ntwalubombo (Z)

Rubia cordifolia subsp. conotricha rt

Nukane, u (Z) (also nukani)

Ocotea bullata bk

nukelamabiba (Z)

Clausena anisata bk

Nukelambiba, um (Z)

Clausena anisata rt

Nungumabele, um (Z)

Zanthoxylum capense rt

Z. davyi rt

Nungwane, um (Z) (also nungwane)

Zanthoxylum capense rt

Z. davyi rt

Nwazi, isi (Z)

Rhoicissus tridentata subsp. tridentata rt

nwelele (Z)

Lycopodium clavatum wpl

Nyamathi, um (Z)

Ekebergia capensis bk

Nyamawempunzi, um (Z)

Plectranthus grallatus rt

Nyathelo, in (Z) (also nyathelo)

Vemonia adoensis rt

nyazangoma, black (Z)

Primus africana bk

nyazangoma, white (Z)

unknown 57 bk

nyazangoma, red (Z)

Rhus chirindensis bk

Nyenya, u (Z)

Rhamnus prinoides rt

nyokaspinda (Z)

Boscia albitrunca rt

Nyongwane, i (Z) (also nyongwane)

Corbichonia decumbens rt

Dicoma anomala rt

nzimane (Z)

Euclea natalensis rt

Ompumelelo, un (Z)

Macaranga capensis bk

pahte-ea-ngaka (S)

Hermannia depressa rt

pamapuce (Z)

Aspilia natalensis 1, st

Eclipta prostrata wpl

pamba (Z)

Balanites maughamii bk

Chlorophytum comosum wpl

Cyrtorchis arcuata rt

Diaphananthe millarii rt

Mystacidium capense rt

M. spp. rt

Tridactyle bicaudata rt

T. tridentata rt

payabashemane (S)

unknown 58 rt

Pendulo, um (Z)

Acalypha villicaulis rt

pengulula (Z)

Deinbollia oblongifolia rt

pepelangeni (Z)

Curtisia dentata bk

Phafq, um (Z)

Ziziphus mucronata subsp. mucronata bk

Phahla, um (Z)

Brachylaena discolor subsp. discolor 1, rt

Hippobromus pauciflorus rt

Phahluka, isi (Z)

Brachylaena discolor subsp. discolor 1, rt

Hippobromus pauciflorus rt

Phakama, i (Z) (also phakama)

Tieghemia quinquenervia wpl

Viscum verrucosum wpl

phamaphuce (Z)

Aspilia natalensis 1, st

Eclipta prostrata wpl

Phamba, i (Z) (also phamba)

Balanites maughamii bk

Chlorophytum comosum wpl

Cyrtorchis arcuata rt

Diaphananthe millarii rt

Mystacidium capense rt

M. spp. rt

Tridactyle bicaudata rt

T. tridentata rt

Phamepuce, u (Z)

Aspilia natalensis 1, st

Eclipta prostrata wpl

Phanda, um (Z)

Lonchocarpus capassa bk

Phaphe, aba (Z)

Pinus sp*. bk

phate-ea-ngaka (S)

Chenopodium sp. rt

Hermannia depressa rt

Malva parviflora * rt

phateyangaka (S)

Chenopodium sp. rt

Hermannia depressa rt

Malva pan’iflora * rt

Phengulula, i (Z) (also phenguluia)

Deinbollia oblongifolia rt

Phephelelangeni, u (Z)

Curtisia dentata bk

Phephetho, isi (Z)

Siphonochilus aethiopicus rt

Phicamaguma, um (X)

Buxus macowanii 1, fl

B. natalensis 1, fl

Phimbi, um (Z)

Garcinia livingstonei bk

phindamshye (Z)

Adenia gummifera var. gummifera st

Phompho, um (Z)

Scadoxus puniceus bb

Phophoma, i (Z) (also phophoma)

Secamone gerrardii rt

Phumbulu, um (Z)

Vitellariopsis dispar bk

V. marginata bk

Phuphuma, i (Z) (also phuphuma)

Ilex mitis st

Securidaca longipedunculata st

Thespesia acutiloba st

pigabone (Z)

Cephalaria humilis rt

pigayibone (Z)

Cephalaria humilis rt

pikayiboni (Z)

Cephalaria humilis rt

Pila, im (Z)

Callilepis laureola rt

Pindamshaye, im (Z) (also pindamshaye)

Adenia gummifera var. gummifera st

pindamuva (Z)

unknown 59 rt

Piskayihlangulwa, im (Z)

Eucalyptus sp. * bk

poho-tsehla (S)

Pachycarpus rigidus rt

Phytolacca heptandra rt

Xysmalobium undulatum rt

pootshetla (S)

Pachycarpus rigidus rt

Phytolacca heptandra rt

Xysmalobium undulatum rt

popoma (Z)

Secamone gerrardii rt

pumbulu (Z)

Vitellariopsis dispar bk

V. marginata bk

Pundu, im (Z) (also pundu)

Gasteria croucheri 1

punyu (Z)

Talinum caff rum rt

punyuka (Z)

Talinum caffrum rt

Qalothi, um (Z)

Strychnos henningsii bk

S. mitis bk

Qhume, u (Z) (also qhume)

Hippobromus pauciflorus rt

Qonga, um (X)

Cassipourea gerrardii bk

Qongqo, u (Z)

Burchellia bubalina bk

Qongqo, i (Z)

Burchellia bubalina bk

Qonsi, i (Z)

Eriosema salignum rt

Qontsi, u (Z) (also qontsi)

Eriosema cordatum rt

E. salignum rt

Qunga, isi (Z)

Cymbopogon excavatus rt, 1, st

C. marginatus rt, 1, st

C. plurinodis rt, 1, st

C. validus rt, 1, st

Qwaningi, i (Z)

Capparis brassii rt

C. tomentosa rt

raadeboni (S)

Cephalaria humilis rt

rooi-storm (A)

Rubia cordifolia subsp. conotricha rt

Roselina, u (Z) (also roselina)

Cinnamomum camphora bk

Sahlulamanye, u (Z) (also sahlulamanye)

Elaeodendron croceum bk

Pterocelastrus echinatus bk

P. rostratus bk

P. tricuspidatus bk

sakchaan (Z)

Euclea divinorum rt

E. undulata rt

sakhe yedwa (S)

unknown 60 bk

saphulo (Z)

Garcinia livingstonii bk

sbaga (Z)

Warburgia salutaris bk

sbhaha (Z)

Warburgia salutaris bk

sdenda (Z)

Maesa lanceolata rt

sdumo (Z)

Ilex mitis var. mitis bk

sebelele (S)

Osteospermum imbricatum subsp. nervatum

1, st

sebelete (S)

unknown 61 rt

sedumo (T)

Hex mitis var. mitis bk

sefo (Z)

Faurea mcnaughtonii bk

F. saligna bk

94

Bothalia 31.1 (2001)

Sefu, isi (Z)

Faurea mcnaughtonii bk

F. saligna bk

Sehle, um (Z)

Peltophorum africanum bk

Sehlulamanye, u (Z) (also sehlulamanye)

Elaeodendron croceum bk

Pterocelastrus echinatus bk

P. rostratus bk

P. tricuspidatus bk

sekanama (S)

Drimia elata bb

D. robusta bb

Tribulus terrestris bb

Urginea altissima bb

U. delagoensis bb

U. macrocentra bb

U. sanguinea bb

sekgopha (S)

Aloe microcantha 1

sekolopatas (S)

Dioscorea sylvatica rt

sekunda mahose (V)

unknown 62 rt

selengwe (Z)

Gnidia kraussiana var. kraussiana rt

selepane (S)

unknown 63 rt

Senge, um (Z)

Cussonia spicata bk, st

senokomaropa (S)

Zanthoxylum capense rt

sepula (Z)

Garcinia livingstonei bk

sesembane (Sh)

unknown 64 rt

Sethole, ama (Z)

Mimusops caffra bk

M. obovata bk

Sideroxylon inerme bk

setima mollo (S)

Pentanisia prunelloides subsp. prunelloides

rt

Shaladilezinyoka, i (Z)

Tulbaghia alliacea bb

T. spp. bb

Shaqa, i (Z) (also shaqa)

Pelargonium luridum rt

Shekisane, um (Z) (also shekisane)

Euclea divinorum rt

E. undulata rt

shikwane (S)

Catha edulis bk, 1

shilwane (S)

unknown 65 st

Shinga, i (Z)

Xysmalobium undulatum rt

shiritsi (V)

Mucuna coriacea subpp. irritans rt

Shisizwe, u (Z) (also shisizwe)

Portulaca sp. rt

shlabahlangane (Z)

Cycnium racemosum rt

shoba lenyathi (Z)

Equisetum ramosissimum rt

Shobalehashi, i (Z)

Equisetum ramosissimum rt

Shongwe, i (Z) (also shongwe)

Asclepias spp. rt

Pachycarpus rigidus rt

P. spp. rt

Xysmalobium undulatum rt

shwilieshwili (Z)

Plumbago auriculata rt

sibanku (Z)

Ochna holstii bk

sibindi sengwenya (Z)

unknown 66 bk

sidala (Z)

Tephrosia marginella 1, st

sidikili (Z)

Gnidia burchellii rt

G. cuneata rt

G. kraussiana var. kraussiana it

G. spp. rt

sidumo (Z)

Ilex mitis var. mitis bk

sihlenhle (Z)

Euphorbia woodii wpl

Silawengwe, um (Z) (also silawengwe)

Gnidia kraussiana var. kraussiana rt

silepe (Z)

unknown 67 rt

simanaye (Z)

Ekebergia capensis bk

sinama-senyonga (Z)

Achyropsis avicularis wpl

Sinandlovu, um (Z)

Mundulea sericea it

sindiyandiya (Z)

Bersama lucens bk

B. stayneri bk

B. swinnyi bk

B. tysoniana bk

Spermacoce natalensis bk

singala salugazi (Z)

Asclepias fruticosa rt

A. physocarpa rt

Singalwesalukazi, u (Z)

Asclepias fruticosa rt

A. physocarpa rt

Sinini, isi (Z) (also sinini)

Spilanthes mauritiana 1, st

sinwazi (Z)

Rhoicissus tridentata subsp. tridentata rt

sipha senyuko (Z)

Achyropsis aviculris wpl

Siphawenyoka, um (Z)

Achyropsis avicularis wpl

siphephetho (Z)

Siphonochilus aethiopicus rt

siqunga (Z)

Cymbopogon excavatus rt, 1, st

C. marginatus rt, 1, st

C. plurinodis rt, 1, st

C. validus rt, 1, st

siqunga si tokoloshe (Z)

Dietes iridioides 1, st

sisefo (Z)

Faurea mcnaughtonii bk

F. saligna bk

sithundu (Z)

Elaeodendron croceum bk

Ochna natalitia rt

sitshctsane (V)

Dioscorea dregeana it

sizo (Z)

unknown 68 bk

skanama (Z)

Drimia elata bb

D. robusta bb

Tribulus terrestris bb

Urginea altissima bb

U. delagoensis bb

U. macrocentra bb

U. sanguinea bb

skanama, white (S)

Urginea sp. bb

skilpad (Z)

Dioscorea sylvatica rt

skulpati (Z)

Dioscorea sylvatica rt

slaslahadiem (S)

Chenopodium ambrosioides * 1, si

slonyane (Z)

Vernonia adoensis rt

snama (Z)

Elaeodendron croceum bk

Maytenus acuminata bk

Priva cordifolia var. abyssinica rt

snwazi (Z)

Rhoicissus tridentata subsp. tridentata rt

Solo, u (Z) (also solo)

Albizia adianthifolia bk

spahluka (Z)

Brachylaena discolor subsp. discolor rt. 1

Hippobromus pauciflorus rt

spanku (Z)

Ochna liolstii bk

spatluga (Z)

Brachylaena discolor subsp. discolor 1, rt

Hippobromus pauciflorus rt

spele (T)

unknown 69 bb

sphephetho (Z)

Siphonochilus aethiopicus rt

stapuga (Z)

Pulicaria scabra rt

sukasamba (Z)

Cymbopogon spp. rt, st

Sukasihambe, u (Z) (also sukasihamba)

Cymbopogon spp. rt, st

Sukumbili, un (Z) (also sukumbili)

Chenopodium ambrosioides* 1, st

Hypericum aethiopicum subsp. aethiopicum

1, st

Senecio gregatus 1, st

S. serratuloides var. gracilis 1, st

sulansuia (Z)

Eriospermum mackenii rt

E. sp. rt

Spermacoce natalensis rt

sumuso owalapha wayongaphi (Z)

Harpagophytum procumbens fr, rt

Sununundu, u (Z) (also sununundu)

Acalypha peduncularis rt

A. punctata rt

A. schinzii wpl

Suzwane, um (Z) (also suzwane)

Lippia javanica 1, st

Tricalysia lanceolata 1, st

swazi (Z)

Turraea obtusifolia 1, st

Swazi, u (Z)

Turraea obtusifolia 1, st

T. emetica bk

tabazimbi (V)

Brackenridgea zanguebarica rt

tangazane (S)

Gunnera perpensa rt

thabatsindi (V)

Brackenridgea zanguebarica rt

Thaphuka, isi (Z)

Pulicaria scabra rt

thathayane (T)

Heteropyxis natalensis rt

Thathe, um (Z)

Ptaeroxylon obliquum bk

thaululo (V)

Widdringtonia nodiflora rt

Thende, isi (Z)

Maesa lanceolata rt

Thethe, i (Z)

Polygala fruticosa rt

Thibomvu, umu (Z)

Protorhus longifolia bk

Tholo, um (Z)

Acacia caffra bk

Thombothi, um (Z)

Spirostachys africana bk

Thuma, um (Z)

Solatium aculeastrum fr

S. hermannii fr

Bothalia 31,1 (2001)

95

Thumelela, um (Z)

Pleurostylia capensis bk

Thundu, isi (Z)

Elaeodendron croceum bk

Ochna natalitia rt

Thunduluka, um (Z)

Ximenia caffra var. natalensis rt

Thunyelelwa, um (Z)

Pleurostylia capensis bk

thunzikulu (Z)

Mimusops sp. bk

Thwazi, um (Z)

Rhoicissus rhomboidea rt

R. tridentata subsp. tridentata rt

Tindili, in (Z)

Entada rheedii sd

titikwane (V)

unknown 70 bb

Tiye, u (Z)

Hippobromus pauciflorus rt

tjanibeswe (S)

Dianthus mooiensis subspp. wpl

tlhatlhabadimo (T)

Chenopodium ambrosioides * 1, st

Tlokwana la tsela (S)

Dianthus basuticus subspp. wpl

D. mooiensis subspp. wpl

Tolwane, in (Z)

Elephantorrhiza elephantina rt

tomboti (Z)

Spirostachys africana bk

tshanibezwe (Z)

Dianthus mooiensis subspp. wpl

tshidzungu (Sh)

Ilex mitis var. mitis bk

Protea gaguedi fl, rt

tshokhalasile (S)

Dianthus mooiensis subspp. wpl

tshukwane (S)

unknown 7 1 rt

Tshungu, in (Z)

Momordica foetida 1, st

Tshwalabenyoni, u (Z) (also tshwalabenyoni )

Burchellia bubalina bk

tshwetshwe (Z)

unknown 72 rt

tsoetla-e-nyenyane (S)

Argyrolobium tuberosum rt

tswetlane (S)

Argyrolobium tuberosum rt

Tuma, in (Z)

Solanum aculeastrum fr

S. hermannii fr

Tumemncane, in (Z)

Solanum panduriforme fr

Twalalubombo, in (Z)

Rubia cordifolia subsp. conotricha rt

ubande (Z)

Greyia sutherlandii bk

ubane (Z)

Agapanthus africanus rt

A. campanulatus rt

A. praecox subsp. orientalis rt

uBangaiala (Z)

see Bangalala, u

uBani (Z)

see Bani, u

uBhoqo (Z)

see Bhoqo, u

uBhubhubhu (Z)

see Bhubhubhu, u

uBoqo (Z)

see Boqo, u

ubovimba (Z)

Withania somnifera rt

ubuHlungubendlovu (Z)

see Hlungubendlovu, ubu

ubuHlungwana (Z)

see Hlungwana, ubu

uBulawu (Z)

see Bulawu, u

ubuLibazi (Z)

see Libazi, ubu

ubuVimbo (Z)

see Vimbo, ubu

ucume (Z)

Hippobromus pauciflorus rt

uDakwa (Z)

see Dakwa, u

uDelunina (Z)

see Delunina, u

uDlutshana (Z)

see Dlutshana, u

uDwendwelencuba (Z)

see Dwendwelencuba, u

uDwendwelwegcuba (Z)

see Dwendwelwegcuba, u

udwendweni (Z)

Crocosmia aurea bb

C. paniculata bb

Dierama pendulum bb

Eulophia cucullata rt

E. spp. rt

Gladiolus dalenii bb

G. sericeo-villosus forma sericeo-villosus bb

uFudu (Z)

see Fudu, u

ugashi (Z)

Berkheya multijuga rt

uGebeleweni (Z)

see Gebeleweni, u

uGibisila (Z)

see Gibisila, u

uGibonisele (Z)

see Gibonisele, u

ugini (Z)

unknown 73 st

uGobandlovu (Z)

see Gobandlovu, u

uGobho (Z)

see Gobho, u

uGodide (Z)

see Godide, u

ugopo (Z)

Gunnera perpensa rt

uGwayana (Z)

see Gwayana, u

uHlabahlangane (Z)

see Hlabahlangane, u

uhlakahla (Sw)

Agapanthus africanus rt

uHlonyane (Z)

see Hlonyane, u

uHluguwendlovu (Z)

see Hluguwendlovu, u

uhlungu (Z)

Strophanthus speciosus fr

uHlungu-lungu (Z)

see Hlungu-lungu, u

uHlungubendlovu (Z)

see Hlungubendlovu, u

ukashi (Z)

Berkheya multijuga rt

uKhalimele (Z)

see Khalimele, u

ukhathwa (Z)

unknown 74 rt

uLoselina (Z)

see Loselina, u

uMababaza (Z)

see Mababaza. u

umaBelejongosi (Z)

see Belejongosi, umu

uMabelejongosi (Z)

see Mabelejongosi, u

uMabopha (Z)

see Mabopha, u

uMabusane (Z)

see Mabusane, u

uMadilika (Z)

see Madilika, u

uMadlozana (Z)

see Madlozana, u

uMadlozanomncane (Z)

see Madlozanomncane, u

uMaguqu (Z)

see Maguqu, u

uMahesakobomvu (Z)

see Mahesakobomvu, u

uMahesakomhiope (Z)

see Mahesakomhlope, u

umaHlabekufeni (Z)

see Hlabekufeni. uma

uMahlabekufeni (Z)

see Mahlabekufeni, u

uMahlanganisa (Z)

see Mahlangqnisa, u

umahlogolosi (Z)

Urginea altissima bb

U. spp. bb

uMahlongolozi (Z)

see Mahlongolozi, u

uMahogwe (Z)

see Mahogwe, u

uMakhothigobile (Z)

see Makhothigobile, u

uMalala (Z)

see Malala. u

umaLuleka (Z)

see Luleka, uma

uManono (X)

see Manono, u

uManzamnyama (Z)

see Manzamnyama, u

uMaphipha (Z)

see Maphipha, u

uMaphiphakhubalo (Z)

see Maphiphakhubalo, u

uMaphiphintelezi (Z)

see Maphiphintelezi, u

uMaqadini (Z)

see Maqadini, u

umasangana (Z)

unknown 75 st

uMaswelisweli (Z)

see Maswelisweli, u

uMathanjana (Z)

see Mathanjana, u

uMathinta (Z)

see Mathinta, u

uMathithibala (Z)

see Mathithibala, u

uMathunga (Z)

see Mathunga, u

uMathunzini (Z)

see Mathunzini, u

uMathunziniwezintaba (Z)

see Mathunziwezintaba, u

uMatshintshine (Z)

see Matshintshine, u

uMavumbuka (Z)

see Mavumbuka, u

uMayime (Z)

see Mayime, u

uMazwende (Z)

see Mazwende, u

uMazwendomnyama (Z)

see Mazwendomnyama, u

umBande (Z)

see Bande, um

uMbesa (Z)

see Mbesa, u

umBethe (Z)

see Bethe, um

umBhandu (Z)

see Bhandu, um

96

Bothalia 31,1 (2001)

umBinda (Z)

see Binda, um

umbitchi khubalo (Z)

unknown 76 bk

umBola (Z)

see Bola, um

umBovana (Z)

see Bovana, um

umchambalala (Z)

unknown 77 rt

umCuniane (Z)

see Cumane, um

umDabu (Z)

see Dabu, um

umDakane (Z)

see Dakane, um

umDewehlati (Z)

see Dewehlati, um

umDIandlovu (X)

see Dlandlovu, um

uMdlangwenya (Z)

see Mdlangwenya, u

umDlavusa (Z)

see Dlavusa, um

umDIebe (Z)

see Dlebe, um

umDondwani (Z)

see Dondwani, um

umDubu (Z)

see Dubu, um

umduze (Z)

Crinum bulbispermum bb

C. delagoense bb

C. macowanii bb

C. moorei bb

C. sp. bb

uMembeza (Z)

see Membeza, u

uMemeziobomvu (Z)

see Memeziobomvu, u

uMemezomhlope (Z)

see Memezomhlope, u

umFanozacile (Z)

see Fanozacile, um

umFazothethayo (Z)

see Fazothethayo, um

umFusamvu (Z)

see Fusamvu, um

umGadankawu (Z)

see Gadankawu, um

umGalagala (X)

see Galagala, um

umgamazi (Z)

Acacia robusta subsp. clavigera bk

A. robusta subsp. robusta bk

umGanu (Z)

see Ganu, um

umGogawezinhlanya (Z)

see Gogawezinhlanya, um

umGwenya (Z)

see Gwenya, um

umGxamu (Z)

see Gxamu, um

umhlabelo-we-qolo (Z)

Blighia unijugata bk

umHIabclo (Z)

see Hlabelo, um

umHIadlothi (X)

see Hladlothi, um

umHIahlankosi (Z)

see Hlahlankosi, um

umHIala (Z)

see Hlala, um

umHIalamagwababa (Z)

see Hlalamagwababa, um

umhlalimakwaba (Z)

Bridelia micrantha bk

umHIambamanzi (Z)

see Hlambamanzi, um

umHlangula (Z)

see Hlangula, um

umHlele (Z)

see Hlele, um

umHlonishwa (Z)

see Hlonishwa, um

umHlonyane (Z)

see Hlonyane, um

umHlunguhlungu (Z)

see Hlunguhlungu, um

umHluthi (Z)

see Hluthi, um

umHlwazi (Z)

see Hlwazi, um

umHIwazimamba (Z)

see Hlwazimamba, um

uMinya (Z)

see Minya, u

uMinyela (Z)

see Minyela, u

umjuluka (Z)

see Juluka, um

unikadankawu (Z)

Albizia adianthifolia bk

umKele (Z)

see Kele, um

umkhabamasi (Z)

Rauvolfia caffra bk

umKhalapanga (Z)

see Khaphalapanga, um

umKhamba (Z)

see Khamba, um

umKhanyakude (Z)

see Khanyakude, um

umKhiwane (Z)

see Khiwane, um

umKhokha (Z)

see Khokha, um

umKhondweni (Z)

see Khondweni, um

umKhuhlu (Z)

see Khuhlu, um

umKhumizo (Z)

see Khumizo, um

umKhunye (X)

see Khunye, um

umKhuze (Z)

see Khuze, um

umKhwangu (Z)

see Khwangu, um

unikomasi (Z)

Acacia robusta subsp. clavigera bk

A. robusta subsp. robusta bk

umLahlankosi (Z)

see Lahlankosi, um

umLahleni (Z)

see Lahleni, um

umLahlenisefile (Z)

see Lahlenisefile, um

umlatleni (Z)

Curtisia dentata bk

umLomomnandomncane (Z)

see Lomomnandomncane, um

umLulama (Z)

see Lulama, um

umLulama-onincane (Z)

see Lulama-omncane, um

umLulama-womfula (Z)

see Lulama-womfula, um

umLulaniomkhulu (Z)

see Lulamomkhulu, um

umLunge (Z)

see Lunge, um

umLuthu (Z)

see Luthu, um

umNduze (Z)

see Nduze, um

umNgamanzi (Z)

see Ngamanzi, um

umNono (Z)

see Nono, um

umnukela mbiba (Z)

Clausena anisata rt

umNukelambiba (Z)

see Nukelambiba, um

umNungumabele (Z)

see Nungumabele, um

umNungwane (Z)

see Nungwane, um

umNyamathi (Z)

see Nyamathi, um

umNyamawempunzi (Z)

see Nyamawempunzi, um

umnyiso (Z)

unknown 78 rt

umnyokiziphinda (Z)

Boscia albitrunca rt

uMondi (Z)

see Mondi, u

unioya wovungu (Z)

unknown 79 bk

uMoyawezwe (Z)

see Moyawezwe, u

umPendulo (Z)

see Pendulo, um

umPhafa (Z)

see Phafa, um

umPhahla (Z)

see Phahla, um

umPhanda (Z)

see Phanda, um

umPhicamaguma (Z)

see Phicamaguma. um

umPhimbi (Z)

see Phimbi, um

umPhompho (Z)

see Phompho, um

umPhumbulu (Z)

see Phumbulu, um

umphumeleli (Z)

Macaranga capensis bk

uMpikayiboni (Z)

see Mpikayiboni, u

uMpunyu (Z)

see Mpunyu, u

umQalothi (Z)

see Qalothi, um

umQonga (X)

see Qonga, um

umqonqo (Z)

Burchellia bubalina bk

umSehle (Z)

see Sehle, um

umSenge (Z)

see Senge. um

umShekisane (Z)

see Shekisane, um

umSilawengwe (Z)

see Silawengwe, um

umSinandlovu (Z)

see Sinandlovu, um

umSiphawenyoka (Z)

see Siphawenyoka, um

umSuzwane (Z)

see Suzwane, um

umThathe (Z)

see Thathe, um

umTholo (Z)

see Tholo, um

umThombothi (Z)

see Thombothi, um

umThuma (Z)

see Thuma, um

uniThumelela (Z)

see Thumelela, um

umThunduluka (Z)

see Thunduluka, um

Bothalia 31,1 (2001)

97

umThunyelelwa (Z)

see Thunyelelwa, um

umThwazi (X)

see Thwazi, um

uMululuka (Z)

see Mululuka, u

umuNga (Z)

see Nga, umu

uMusa (Z)

see Musa, u

umuThibomvu (Z)

see Thibomvu, umu

umuvungazi (Z)

Pterocarpus angolensis bk

Trema orientalis bk

umValasangweni (Z)

see Valasangweni, um

umVangazi (Z)

see Vangazi. um

umvete (Z)

Boscia albitrunca rt

B. foetida subsp. rehmanniana rt

Eragrostis plana rt

umVithi (Z)

see Vithi, um

umVongothi (Z)

see Vongothi. um

umVuma (Z)

see Vuma, um

umVuthuza (Z)

see Vuthuza, um

umVuthwemini (Z)

see Vuthwemini, um

umWelela (Z)

see Welela, um

umYazangomobomvu (Z)

see Yazangomobomvu, um

umYenye (Z)

see Yenye, um

umZaneno (Z)

see Zaneno, um

umZilanyoni (Z)

see Zilanyoni, um

umZimuka (Z)

see Zimuka, um

umZiwalapha-ungaphi (Z)

see Ziwalapha-ungaphi, um

umZungulu (Z)

see Zungulu, um

umZungulwane (Z)

see Zungulwane, um

uNakile (Z)

see Nakile, u

unDabingehlele (Z)

see Dabingehlele, un

unDiyaza (Z)

see Diyaza, un

unDonganazibomvana (Z)

see Donganazibomvana, un

unDwendweni (Z)

see Dwendweni, un

unGazine (Z)

see Gazine. un

unGibonisele (Z)

see Gibonisele, un

uNgqengendlela (Z)

see Ngqengendlela, u

uNgwaleni (Z)

see Ngwaleni, u

unHlangothi (Z)

see Hlangothi, iu

uNkungwini (Z)

see Nkungwini, u

unOmpumelelo (Z)

see Ompumelelo, un

unSukumbili (Z)

see Sukumbili, un

uNsukumbili (Z)

see Nsukumbili, u

unukambiba (Z)

Clausena anisata rt

uNukane (Z)

see Nukane, u

unWele (Z)

see Wele, un

unYawolwenkukhu (Z)

see Yawolwenkukhu, un

uNyenya (Z)

see Nyenya, u

uphagama (Z)

Tieghemia quinquenervia wpl

Viscum verrucosum wpl

uPhamepuce (Z)

see Phamepuce, u

uPhephelelangeni (Z)

see Phepheielangeni, u

uQhume (Z)

see Qhume, u

uQongqo (Z)

see Qongqo. u

uQontsi (Z)

see Qontsi, u

uquama mahlombe (Z)

unknown 80 bk

uRoselina (Z)

see Roselina, u

uSahlulamanye (Z)

see Sahlulamanye, u

uSehlulamanye (Z)

see Sehlulamanye, u

usenge (Z)

Cussonia spicata bk, st

uShisizwe (Z)

see Shisizwe, u

uSingalwesalukazi (Z)

see Singalwesalukazi, u

uSolo (Z)

see Solo, u

uSukasihambe (Z)

see Sukasihambe, u

uSununundu (Z)

see Sununundu, u

usuzwane (Z)

Lippia javanica 1, st

Tricalysia lanceolata 1, st

uSwazi (Z)

see Swazi, u

uTiye (Z)

see Tiye, u

uTshwalabenyoni (Z)

see Tshwalabenyoni, u

utyanibswe (Z)

Dianthus mooiensis subspp. wpl

uvalo (Z)

unknown 81 bb

uvangasi (Z)

Pterocarpus angolensis bk

Trema orientalis bk

uVelabahleke (Z)

see Velabahleke, u

uVendle (Z)

see Vendle, u

uVimbukhalo (Z)

see Vimbukhalo, u

uVovo (Z)

see Vovo, u

uVukakwabafdikhubalo (Z)

see Vukakwabafrlikhubalo, u

uVukwababa (Z)

see Vukwababa, u

uVukwabafile (Z)

see Vukwabafile, u

uVumebomvu (Z)

see Vumebomvu, u

uVumomhlope (Z)

see Vumomhlope, u

uvungalumoya (Z)

unknown 82 bk

uXhapozi tZ)

see Xhapozi, u

uZangume (Z)

see Zangume, u

uZekane (Z)

see Zekane, u

uZeneke (Z)

see Zeneke, u

uZililo (Z)

see Zililo, u

uZinhla (Z)

see Zinhla. u

Valasangweni, um (Z)

Gardenia thunbergia rt

G. volkensii subsp. spatulifolia rt

Vangazi, um (Z)

Pterocarpus angolensis bk

Trema orientalis bk

Velabahleke, u (Z) (also velabahleka)

Celosia trigyna wpl

Crotalaria spp. rt

velasangweni (Z)

Gardenia thunbergia rt

G. volkensii subsp. spatulifolia rt

velasmoyeni (Z)

unknown 83 rt, st

Vendle, u (Z) (also vendhle)

Pelargonium luridum rt

Vimbo, ubu (Z)

Withania somnifera rt

Vimbukhalo, u (Z) (also vimbokhalo)

Convolvulus sagittatus subspp. rt

Ipomoea crassipes rt

Vithi, um (Z)

Boscia albitrunca rt

B. foetida subsp. rehmanniana rt

Eragrostis plana rt

Vongothi, um (Z)

Kigelia africana fr

Vovo, u (Z)

Scholia brachypetala bk

vugamafile (Z)

Myrothamnus flabellifolia 1, st

vuka kubafde (Z)

unknown 85 bk

vuka khubalo (Z)

unknown 84 bk

vuka-kwa-bafde (Z)

Myrothamnus flabellifolia 1, st

vukakwabaFdeyo (Z)

Myrothamnus flabellifolia 1, st

Vukakwabafdikhubalo, u (Z)

Rapanea melanophloeos bk

Vukwababa, u (Z)

Myrothamnus flabellifolia 1, st

Vukwabafile, u (Z)

Myrothamnus flabellifolia 1, st

vulakuvaliwe (Z)

Podocarpus henkelii bk

vuma, white (Z)

Synaptolepis kirkii rt

vuma, red (Z)

Pappea capensis rt

Vuma, um (Z)

Turraea floribunda rt

Vumebomvu, u (Z)

Pappea capensis rt

Vumelwane, isi (Z) (also vumelewana)

Hypericum aethiopicum subsp. aethiopicum

wpl

Vumomhlope, u (Z)

Synaptolepis kirkii rt

vungulamoya (Z)

Trichilia dregeana bk

T. emetica bk

vungulomo (Z)

Trichilia dregeana bk

T. emetica

98

Bothalia 31,1 (2001)

Vuthuza, um (Z) (also vuthuza)

Kalanchoe sp. 1, st

Knowltonia bracteata 1, st

Vuthwemini. um (Z) (also vutwamini)

Canthium inerme rt

Wele, un (Z)

Lycopodium clavatum wpl

Welela, um (Z)

Tulbaghia alliacea bb

T. ludwigiana bb

T. simmleri bb

wezifo zonge (Z)

unknown 86 rt

Wozawoza, i (Z) (also wozawoza)

Cymbopogon spp. rt, st

xaxabulane (Z)

Smilax anceps rt

Xhaphozi, u (Z)

Ranunculus multifidus 1, st

xidomeja (Sh)

unknown 87 rt

Yathelo, in (Z)

Vemonia adoensis rt

Yawolvvenkukhu, un (Z)

Pelargonium luridum rt

yazangoma, red (Z)

Rhus chirindensis bk

yazangoma, black (Z)

Primus africana bk

Yazangomelimnyama, in (Z)

Primus africana bk

Yazangomobomvu, um (Z)

Rhus chirindensis bk

Yenye, um (Z)

Rhamnus prinoides rt

Yokiziphinda, in (Z)

Boscia albitrunca rt

zagane (Z)

Polygala hottentotta rt

zaneke (Z)

Haemanthus albiflos bb

II. coccineus bb

H. defonnis bb

H. spp. bb

Zaneno, um (Z) (also zaneno)

Olinia radiata bk

Zangume, u (Z) (also zangume)

Helichryswn acutatum rt

Zaza, i (Z)

Behnia reticulata rt

Zekane, u (Z) (also zekane)

Polygala hottentotta rt

Zeneke, u (Z)

Haemanthus albiflos bb

H. coccineus bb

H. defonnis bb

H. spp. bb

Zibu, i (Z)

Nymphaea noucliali wpl

N. spp. wpl

Zilanyoni, um (Z)

Croton sylvaticus bk

Zililo, u (Z)

Stapelia gigantea wpl

Zimuka, um (Z) (also zimuka)

Phytolacca dioica rt

Zinhla, u (Z) (also zinhla)

Celtis mildbraedii bk

Ziwalaphungaphi, um (Z)

Harpagophytum procumbens fr, rt

zomaga (Z)

unknown 88 bk

zukela (Z)

unknown 89 1, st

Zungulu, um (Z)

Dalbergia obovata st

Zungulwane, um (Z)

Proboscidea fragrans fr

zwane (Z)

unknown 90 rt

Bothalia 31,1: 99-115 (2001)

Thirty years of change in the fynbos vegetation of the Cape of Good

Hope Nature Reserve, South Africa

S.D.J. PRIVETT *t, R.M. COWLING* and H.C. TAYLOR**

Keywords: fire effects, fynbos, succession, temporal change, vegetation monitoring

ABSTRACT

This study used permanently marked 50 m2 sites, surveyed at a 30 year interval, to provide a descriptive account of the

temporal change in the fynbos vegetation of the Cape of Good Hope Nature Reserve, South Africa. Management records

were used to examine the role of post-fire age, fire frequency and intensity, as well as biotic interactions (competition from

overstorey proteoids and alien plants) in influencing vegetation composition over this time period. The mean similarity in

species composition of sites between surveys was 62%, indicating an average of nearly 40% turnover in species over the 30

year period. The main causes of this change included differences resulting from different stages in the post-fire succession

as well as the impact of differential fire regimes (especially frequency effects). Competition from serotinous Proteaceae,

which proved highly mobile after fire, as well as invasive Australian acacias also impacted on the composition of the veg-

etation over time. The study demonstrated that fynbos communities are temporally dynamic and that the changes over time

in species composition are caused by a variety of processes. The study also provided evidence for the role of temporal diver-

sity in contributing to the high species diversity in fynbos systems.

CONTENTS

Introduction 99

Study area 101

Methods 101

Results and discussion 102

1 . Fire effects 102

1.1. Post-fire succession 102

1.1.1. Early successional changes 103

1.1.2. Successional changes after long fire-free

intervals in upland fynbos 103

1.1.3. Successional changes after long fire-free

intervals in coastal fynbos 105

1.2. Fire frequency 105

1.2.1. Short intervals between fires 106

1.2.2. Long intervals between fires 106

2. Dynamics of serotinous Proteaceae 107

2. 1 . Causes of decline of serotinous Proteaceae . . 108

2.2. Causes of increase of serotinous Proteaceae . 108

2.3. The impact of serotinous Proteaceae on under-

storey species diversity 109

3. Alien plants 110

3.1. A comparison of the alien threat in 1966 and

1996 110

3.2. Impact of aliens ’on indigenous vegetation . . . 110

3.2.1. Sites invaded between surveys and sub-

sequently cleared 110

3.2.2. The impact of existing stands of alien plants 1 1 1

Conclusions 112

Acknowledgements 112

References 112

Appendix 1 114

Appendix 2 115

* Institute for Plant Conservation, Botany Department, University of Cape

Town, Private Bag, 7700 Rondebosch, Cape Town.

** Late, of the National Botanical Institute, Private Bag X7, 7735 Clare-

mont, Cape Town.

t Present address: Grootbos Nature Reserve, PO. Box 148, 7220 Gans-

baai, South Africa.

MS. received: 2000-02-15.

INTRODUCTION

'The ecological phenomena that we study often oper-

ate on temporal scales longer than our own existence

and certainly longer than the time span of a research

grant’

Wiens et al. (1986)

A major challenge facing vegetation scientists is to

determine the resilience of communities, as well as indi-

vidual species, to the varying components of the distur-

bance regime (Cowling 1987). Fire and invasion by alien

plants are the major disturbance factors acting on the

fynbos vegetation of the southern Cape Peninsula, South

Africa. For practical reasons, studies on the effects of

these disturbances on fynbos dynamics have always been

temporally restricted. Studies on succession following

fire have monitored recovery after a single fire event.

Similarly, the impacts of the various components of the

fire regime on recruitment have looked at parent to

seedling ratios at the same site following a single fire

event (Bond 1980; Van Wilgen 1981; Hoffman et al.

1987; Cowling & Gxaba 1990). These studies are limit-

ed as they base their results on single fire events. Most

studies on the impact of alien plants on indigenous veg-

etation have compared adjacent invaded versus non-

invaded sites (Richardson et al. 1989; Holmes & Cow-

ling 1997), but see Richardson & Van Wilgen (1985) for

an exception. There is always the problem with this

approach that differences in stands may reflect pre-exist-

ing differences in the physical environment and not the

effects of fire or alien plants.

While many studies on decade-scale vegetation

change have been undertaken internationally (e.g. Fojt &

Harding 1995; Rose et al. 1995; Minnich et al. 1995;

Dodd et al. 1995) and some local research has explored

long-term changes in the semi-arid Karoo (Hoffman &

Cowling 1990; O'Connor & Roux 1995), no work on

medium-term temporal dynamics has been carried out in

fynbos. Thus, while spatial patterns in the complex and

diverse fynbos vegetation have been extensively re-

100

Bothalia 31,1 (2001)

FIGURE 1.— The Cape of Good

Hope Nature Reserve

study area showing loca-

tion of 100 original vege-

tation sites surveyed by

Taylor (1969). Sites 5, 6,

7, 11, 20, 26, 32, 33, 35,

41, 43, 51-53, 69, 77, 85,

93 and 96 were not relo-

cated and therefore were

excluded from this study.

Source: 1:10 000 ortho-

photo, Chief Directorate,

surveys.

Bothalia 31.1 (2001)

101

searched over the last few decades (Taylor 1978; Kruger

1984; Campbell 1985; Cowling & Holmes 1992), no

studies have explored the contribution of the temporal

component to this complexity. Consequently little is

known about the resilience of fynbos to change over time

or how repeated disturbances influence patterns in vege-

tation distribution at the community and landscape scale.

This study explored the resilience of fynbos vegetation in

the Cape of Good Hope Nature Reserve (CGHNR) to 30

years of variable disturbance, both natural and managed.

The study used mostly a descriptive approach to explore

changes in the composition of permanent vegetation sites

established in 1966. It attempted to invoke explanations

for the observed change from historical fire and alien

plant records by asking the following questions:

1. How much change has occurred in the vegetation of

the CGHNR between 1966 and 1996?

2. How important was post-fire age as a determinant of

species composition and diversity?

3. What effects have long fire-free intervals had on veg-

etation dynamics in upland and coastal fynbos?

4. What impact have unusually short and long fire fre-

quencies had on vegetation composition?

5. What impact has high intensity fire following long

fire-free intervals had on vegetation composition

and structure?

6. What has been the dynamics of the common seroti-

nous overstorey species Leucadendron laureolum,

and what impact has it had on understorey species

diversity?

7. How has the abundance and range of invasive alien

plants changed over the last 30 years, and what

effect have these changes had on the indigenous

flora of the Reserve?

STUDY AREA

The area encompassed by the Cape of Good Hope

Nature Reserve (CGHNR) was used for this study

(Figure 1). A detailed description of the physical setting

and historical background of the Reserve is document-

ed in Taylor (1984a). The Reserve had certain attributes

that made it suitable for this study: high species diver-

sity and complexity; the existence of permanent moni-

toring sites; and detailed management records including

fire histories (Appendix 1) for the period since the es-

tablishment of the sites. The vegetation of the CGHNR

has been well documented and a comprehensive list of

1 073 species compiled (Taylor 1969, 1983, 1984a, b;

Fraser & McMahon 1995; Fraser in press.). The vege-

tation communities were described by Taylor (1984b)

who emphasized the variability within communities and

suggested that historical effects relating to disturbance

had influenced within-community species heteroge-

neity.

METHODS

Taylor (1969) established 100 permanently marked

vegetation quadrats of 50 m2 (5 x 10 m) during 1965/66

(Figure 1). These provided the baseline species dataset

against which a contemporary species list could be com-

pared. The management records of the Reserve provided

detailed information on both fire history and alien vege-

tation from 1960 to the present. Eighty-one of the 100

quadrats were relocated using Taylor’s original site

markers that demarcated the southwestern comer of each

quadrat (Figure 1). All species present in each quadrat

were noted with the exception of annuals and seasonally

apparent geophytes, both of which were not perennially

identifiable. Much effort went into ensuring that sites

were precisely relocated and that species were accurate-

ly identified. This was crucial for achieving the objec-

tives of the study. The nomenclature of all species was

corrected according to Arnold & De Wet (1993). The

original species lists of Taylor (1969) were used to assist

with identification.

The number of individuals of each species in a site

was counted (estimates were made when distinction

between individuals was difficult, e.g. resprouting Res-

tionaceae) and later converted to a five-category system

that corresponds to a simplified form of Acocks’s (1975)

system of abundance symbols (Table 1). This was done

in order to be compatible with Taylor’s (1969) study that

used Acocks’s (1975) symbols for assessing abundance.

Hugh Taylor assisted with initial field work and was

satisfied that the methodology corresponded with that

used in his initial survey.

The changes in floristic composition within sites over

the thirty year interval between surveys was explored

using the Sorenson’s coefficient (Kent & Coker 1992) in

the form of the following equation:

(a + b)

where Ss is the Sorenson coefficient as a percentage; a is

the number of species present at a site in 1966; b is the

number of species present at the same site in 1996; and c

is the number of species present at both sampling times.

This coefficient was used throughout the study to

quantify temporal changes in species composition. These

results, together with the two temporally separated

species data sets and matched photographs were used to

explore established hypotheses relating to determinants

of vegetation dynamics and temporal change in fynbos.

We do not attempt to provide a detailed analysis of any

aspect of fynbos dynamics in this study. We use specific

examples of sites which have undergone changes relating

to recorded fire or alien plant impacts to emphasize the

role of these factors in structuring contemporary vegeta-

tion patterns.

TABLE 1. — Abundance categories used in this study

102

Bothalia 31,1 (2001)

FIGURE 2. — Site 43 on western seaboard near Olifantsbos Point, was

covered by asteraceous fynbos in 1966. Over the last 30 years

the entire area has been inundated by shifting dune sands,

resulting in the local extinction of all species with the exception

of the pioneer grass Ehrharta villosa. Wreck of Thomas T.

Tucker is visible in background of both photographs.

RESULTS AND DISCUSSION

Nineteen of the original 100 sites were not included in

this study due to problems with accurate re-location or

through loss to infrastructure development. The 81 sites

used in this study showed considerable variability in

their floristic similarity between the two surveys

(Appendix 1). The mean similarity for all sites was 61.8

± 8.9%, while the lowest value was 43.8% and the high-

est 84.3%. The potential for extreme environmental

change to alter the composition of a site was graphically

illustrated by site 43 occurring near Olifants Point on the

western seaboard of the Reserve. This site had to be

excluded from the analyses as mobile dune sands have

inundated it during the last 30 years (Figure 2).

1 . Fire effects

1.1. Post-fire succession

Post-fire succession in fynbos is noted for a rapid ini-

tial establishment of all components of the community

including those (fire ephemerals) that only appear in the

first years after fire. Therefore, species diversity is great-

est in the immediate post-fire environment and there is a

marked decrease in species diversity as the cover of

overstorey shrubs increases (Kruger 1987; Cowling &

Pierce 1988; Cowling & Gxaba 1990). The rate of de-

cline in the successional sequence differs between areas

with different environmental conditions. Growth in dry

lowland fynbos sites is generally slower than in moister

mountain fynbos and suppression of understorey species

is predicted to start later, probably only after 30 years

(Hoffman et al. 1987).

We explored the rate of post-fire succession for the

Reserve as a whole as well as the role of post-fire age in

influencing species diversity and composition. By

including the data sets for both surveys it was possible to

examine the effect of the full spectrum of vegetation ages

on species diversity. Differences in the successional

sequence between communities complicates the trend

and therefore only the fire-prone mesic oligotrophic pro-

teoid community was included. This community occurs

on the well-drained rocky hills and is the dominant veg-

etation type of the Reserve (Privett 1998).

There was a significant negative relationship between

vegetation age and species diversity within this commu-

nity (Figure 3). The initial peak in species diversity was

followed by a gradual decline over time. The significant-

ly steeper negative slope of the 1966 data set (-1.19 ±

0.24 as against -0.54 ± 0.1; F = 5.97, df = 1, P < 0.02)

resulted from the inclusion of a number of young sites

(< 4 years post-fire) which had high species diversity.

The mean number of perennial species recorded in 1966

was 38.2 ± 17.4 per site, the maximum was 79 and the

minimum was six. In 1996 there were on average fewer

species per site (35.6 ± 15.5). This suggests that there is

a rapid decline from an initial post-fire peak and that the

majority of post-fire ephemerals have seeded and died

within four years after fire. Thereafter species are lost

from the community at a slower rate.

FIGURE 3. — Relationship between vegetation age and species diver-

sity in 50 mesic oligotrophic proteoid sites at the two sampling

times. Fitted curves are negative exponential, circles represent

1966 sites and triangles 1996 sites.

Bothalia 31,1 (2001)

103

FIGURE 4. — Site 15 near Groot Rondevlei, has shown a marked

decline in species diversity between surveys as a result of suc-

cessional effects. The low, white flowering shrub in the 1966

photograph (three years post-fire) is the post-fire ephemeral

Edmondia sesamoides which was not present in the older veg-

etation in 1996 (nine years post-fire). The 1966 vegetation is

characterized by the visible dominance of the resprouting

graminoid guild, whereas in the older 1996 vegetation, the non-

sprouting shrubs. Leucadendron laureolum and Serruria villosa

are dominant. Note the invasion between surveys of the

Witsands dune field (headland in middle, background of photo-

graph) by the invasive alien. Acacia cyclops.

1.1.1. Early successional changes

Figure 3 illustrates the initial decline in species diver-

sity during the first four years following fire. One of the

relocated sites (15) provided an opportunity for explor-

ing the differences in vegetation composition between

very young and slightly older vegetation (Figure 4). The

impact of fire season and frequency on the site’s vegeta-

tion between surveys should be negligible as the site was

burnt twice in summer wildfires at intervals of twelve

and then eleven years. In 1966 this site was only three

years old and included 66 species, whereas in the nine-

year-old vegetation in 1996 only 27 species were record-

ed. Despite only six years difference in the post-fire age

at the two sampling times, successional effects have

resulted in major compositional changes in this site

between surveys (Ss = 49.46%). Only four of the 27

species recorded in 1996 were new to this site suggesting

that the compositional change which has occurred was as

a direct result of losses due to successional effects. The

43 species lost include a number of fire ephemerals such

as Aspalathus abietina, A. linguiloba, A. retroflexa,

Roella ciliata, R. triflora , Selago spuria, Thesium cari-

natum and T. acuminatum as well as the predominantly

early successional species such as Edmondia sesamoides

and Ursinia crithmifolia (Asteraceae).

The stage in the post-fire succession of fynbos vege-

tation influences the composition and diversity of com-

munities. This has important implications for ecologists

developing community-environment models and wher-

ever possible successional effects should be controlled

by only sampling in similar-aged stands.

1.1.2. Successional changes after long fire-free intervals

in upland fynbos

Fynbos is a fire-prone vegetation whose component

species have a variety of fire-regeneration traits. Fire is

an important process for the regeneration of many

species and in its absence these species senesce and

eventually die (Bond 1980; Van Wilgen 1981). With long

fire-free intervals species diversity declines but at the

same time bird-dispersed species are introduced as perch

sites develop (Manders & Richardson 1992; Cowling et

al. 1997). The late stages of fynbos succession are char-

acterized by autogenic changes and follow the facilita-

tion model of Connell & Slayter (1977). Seeds of forest

and thicket species are dispersed into shrublands, and

seedlings sometimes establish in mature fynbos commu-

nities, especially in sheltered sites (Cowling et al 1997).

The slow-growing seedlings of thicket species are usual-

ly killed by fire before thicket patches have developed

sufficiently to exclude fires. However, where fire-free

intervals exceed approximately fifty years, dense thick-

ets form which eventually coalesce into closed-canopy

vegetation that excludes fire (Manders & Richardson

1992; Richardson & Van Wilgen 1985, Cowling et al.

1997). In drier lowland fynbos the rate at which species

senesce and thicket develops presumably takes longer

than for wet mountain fynbos (Hoffman et al. 1987;

Privett 1998). Five of the sites enumerated in 1966 had

not burnt once at the time of the second survey (74, 81,

95, 99 and 100). These provided an opportunity to deter-

mine the rate of senescence and thicket development in

the relatively dry fynbos of the southern Cape Peninsula.

The mean of the Sorenson coefficient for these upland

fynbos sites was 58.5 ± 7%; the lowest value was for site

95 (46%) and the highest was for site 81 (66.7%). The

diversity of all sites, with the exception of site 99, which

remained constant, declined between surveys. Two of

these sites have not burnt for more than fifty years and

have similar habitat characteristics (74, Figure 5 and 81,

Figure 6). They are located on relatively steep slopes at

approximately 90 m above sea level and adjacent to the

coast. The one is situated on an east-facing slope above

False Bay (74), while site 81 is located on a west-facing

slope above the Atlantic Ocean. Site 74 is therefore

exposed to the strong salt-laden southeasterly winds dur-

ing the summer months, whereas site 8 1 is more protect-

ed from wind but still exposed to strong northwesterlies

in winter.

Predictably, both sites were relatively species poor.

However, neither had experienced significant thicket

development (Figures 5, 6). Both sites had lost several

104

Bothalia 31,1 (2001)

FIGURE 5. — Site 74, oligotrophic proteoid fynbos on the southern

slopes of Paulsberg overlooking False Bay, is exposed to strong

southeast winds during the dry summer months. This vegeta-

tion was at least 20 years old in 1966 and has not been burnt

between surveys. There has been very little change in species

composition or structure over the last 30 years. While there has

been some senescence among both the sprouting and non-

sprouting species, there has been no colonization by thicket

species despite fifty years without fire.

species, most of which were shorter-lived non-sprouters

such as Anthospermum aethiopicum, Chironia baccifera.

Erica mammosa, Leucospermum conocarpodendron (a

weak sprouter), Metalasia muricata, Struthiola ciliata

and Serruria villosa. However, the species that had been

lost from these sites also included some sprouting spe-

cies such as Adenandra villosa, Corymbium africanum.

Erica cerinthoicles and Liparia parva. Therefore, a vari-

ety of species with different regeneration modes had a

lifespan long enough to survive the initial twenty years

without fire, but were unable to survive for fifty years.

Even within the Reserve, differences exist in the

potential for, and rate of, thicket colonization. The habi-

tat of site 81 appears to be more suitable for the devel-

opment of thicket than that of site 74. The thicket

species, Euclea racemosa, Myrsine africana, Olea

capensis, Rhus lucida and R. laevigata were already pre-

sent at this site in 1966, whereas site 74 supported no

thicket species. The better early colonization of site 81 is

probably a result of its taller structure, thereby providing

more perch sites, and its peripheral location to a nearby

thicket. Since the initial survey, neither site has demon-

strated a notable increase in thicket species. The bird-dis-

persed Rhus lucida and Pterocelastrus tricuspidatus col-

onized sites 74 and 81 respectively. Erica tristis, a tree

FIGURE 6. — Site 81, oligotrophic proteoid fynbos above large coastal

thicket at Gifkommetjie, is situated on the upper slopes of a fire-

protected amphitheatre above the Atlantic coast. The vegetation

was at least 20 years old in 1966 and has not burnt between

surveys. In 1966 the site contained some seedlings of thicket

species. These have not developed into adult trees over the last

30 years. The vegetation has remained very stable with only the

loss of a few non-sprouting species and the addition of some

short-lived colonizers between surveys. Despite a fire-free peri-

od of ± 50 years, and its close proximity to a well-established

thicket, this site has not developed into a thicket. Some indi-

viduals of Leucospermum conocarpodendron are visible in

both photographs suggesting that this species has a lifespan in

excess of 50 years.

erica associated with fire-free sites, colonized site 81.

The few other species, which have colonized the sites,

are mostly small wind-dispersed shrubs and graminoids.

There has been a small increase in the number of indi-

viduals of thicket species present at site 81, but they have

shown virtually no increase in relative dominance over

the last 30 years. After 30 years Olea capensis is still

only present as suppressed seedlings, whereas seven of

the nine Euclea racemosa individuals are also seedlings

or suppressed saplings.

The physical environment of the southern Cape

Peninsula appears unsuitable for the rapid development

of thicket or forest. Thicket colonization in this area is an

extremely slow process, which, under favourable condi-

tions appears to require fire-free intervals in the order of

centuries rather than decades. Even under the long natur-

al fire regimes of the past, the vegetation of the moun-

tainous southern Reserve probably comprised an assem-

blage of longer-lived fynbos plants, rather than a thicket

community.

Bothalia 31,1 (2001)

105

FIGURE 7. — Site 80, coastal scree asteraceous fynbos near Black Rocks

on the False Bay coast, was already very old in 1966 and has not

been burnt since. The species composition has remained stable

between surveys. There has been an increase in the cover and

biomass of Scirpus nudosus (graminoid in the middle of the site).

This is most probably the result of an improvement in soil mois-

ture since stands of Acacia cyclops were removed above the site

between surveys. Evidence for the longevity of some shrubs is

provided by the persistence of a single individual of the resprout-

ing Asparagus capensis in the bottom left hand comer of both

photographs. This individual is well over 30 years old.

1.1.3. Successional changes after long fire-free intervals

in coastal fynbos

Unlike the upland fynbos, coastal asteraceous and

coastal thicket communities are less frequently burnt

owing to their location adjacent to the sea. It can there-

fore be hypothesized that species occurring in these

communities are not fire adapted, and that they are more

stable in the long-term absence of fire.

Taylor estimated the ages of sites 80 (Figure 7 ) and 90

(Figure 8) as very old in 1966 and neither have experi-

enced fire between the two surveys. These two sites are

quite different in their floristic composition. Both are sit-

uated adjacent to the False Bay coastline. However, site

80 is dwarf scrub on shallow, rocky and damp soils,

while site 90 is a typical coastal dune thicket on deep

well-drained calcareous sands. The Sorenson coeffi-

cients for these sites were 71.1% and 79.0% respective-

ly. They were therefore amongst the most stable in this

study. They showed no change in the relative proportion

of growth form or regeneration mode between surveys.

Numbers of species only decreased slightly between sur-

veys. The most obvious change in site 80 was the large

FIGURE 8. — Site 90, coastal dune thicket on the False Bay coast at

Buffelsbaai. This vegetation type has remained very stable in

composition over the last 30 years. It is dominated by the thick-

et species Sideroxylon inenne, Cassine peragua and Rhus

species. All of these species are long-lived and, as a result, the

community is stable in the absence of fire. The mountain peaks

in the background are from left to right: Paulsberg, Die Boer

and Judas Peak. The person in both photographs is Hugh

Taylor, who initially established the permanent sites in 1966.

increase in the sedge Scirpus nudosus (Figure 7). This

could be attributed to the eradication of stands of Acacia

cyclops above this site, and the subsequent improvement

in the hydrological regime. At site 90 there has been vir-

tually no change in species composition and dominance.

The coastal thicket species Sideroxylon inerme, Cassine

peragua and Rhus species have remained dominant with

approximately the same relative cover.

It can be concluded that the vegetation of the coastal

communities, especially the dune thicket, is very stable

in the absence of fire. These communities tend to be

dominated by long-lived, slow-growing species that are

able to persist for many decades (or centuries) in the

absence of fire.

1.2. Fire frequency

Fires in fynbos occur with intervals of between four

and 40 years (Le Maitre & Midgley 1992). The natural

fire frequency for fynbos in the Cederberg Mountain

Range, some 200 km to the north, was estimated at

between 11 and 15 years (Brown et al. 1991). Owing to

its peninsular geography, the vegetation of the southern

Cape Peninsula would probably have burnt less fre-

106

Bothalia 31,1 (2001 )

latitude

FIGURE 9. — The relationship between mean fire interval and latitude

in the Cape of Good Hope Nature Reserve.

quently than landlocked areas further inland. An analysis

of the last fifty years’ fire records for the Reserve revealed

an average fire interval of 23.2 years. This was calculat-

ed as the average of the frequencies for all sites from the

fire prior to Taylor’s survey and included prescribed

burns. The pre-colonial fire interval was probably even

longer. The area was inhabited by the Peninsular Khoi-

khoi who had little incentive to start fires as they sub-

sisted on marine resources rather than livestock products

(Elphick 1977).

There was a significant positive relationship between

latitude in the Reserve and the mean fire interval (Figure

9). Sites in the southern areas of the reserve, where the

Cape Peninsula narrows to less than a few kilometres,

have had a longer average fire frequency than those in

the northern areas. Fires at unusually short, or long inter-

vals, can alter the species composition of fynbos (Van

Wilgen 1981; Van Wilgen & Forsyth 1992; Vlok 1996;

Schwilk et al. 1997). Frequent burning can alter species

composition by eliminating the large non-sprouting

shrubs, especially Proteaceae, as they do not have suffi-

cient time to reach maturity between fires (Van Wilgen &

Kruger 1981; Van Wilgen 1981). Fast maturing and

sprouting species should be favoured by a short fire fre-

quency (Vlok 1996). Increased competition from the

sprouters also results in a decline in non-sprouters, and a

decline in overall diversity of a site (Vlok 1996).

Where fire is excluded for more than ± 30 years,

senescence occurs in some species and seed stores

become depleted. Poor regeneration and even local

extinction may follow such long intervals between fires

(Bond 1980). A further consequence of long intervals

between fires is that fuel loads increase, resulting in

higher fire intensities. These fires have been shown to

increase seedling recruitment of myrmecochorous

Proteaceae (Bond et al. 1990), as well as to cause higher

mortality among certain resprouting species (Le Maitre

1986).

1.2.1. Short intervals between fires

Only one site in this study was burnt at a very short

frequency (site 14) and could be used to infer the impacts

of short fire intervals on the vegetation (no photograph

available for 1966). Between 1970 and 1988 site 14 on

the eastern boundary of the Reserve was burnt on a three-

year rotation as one of three tracer belts making up a

boundary fire break. These fires were carried out under

cool conditions during late autumn or winter. At the time

of Taylor’s original survey the vegetation on site 14 was

only three years post-fire, whereas in 1996 it was eight

years old.

This site has undergone considerable compositional

change over the last 30 years. The Sorenson coefficient

of only 55% was below average for the data set and only

48% of the original species were still present on the site.

The changes in species composition were generally as

predicted for short fire intervals. Species diversity

decreased: there were 12 fewer species in the second sur-

vey. This decline was as a result of a combination of the

impact of repeated short fire intervals and the later stage

in the post-fire succession in the 1996 survey. Some of

the locally extinct species are relatively slow-maturing

non-sprouters and were probably lost as a direct conse-

quence of the short fire intervals. These include Erica

plukenetii , Psoralea imbricata and Leucadendron laure-

olum; the last-mentioned being common in the vegeta-

tion adjacent to the old fire break. Other species lost from

this site such as Roella ciliatum , R. prostratum and

Thesium spicatum , are early successional species, which

under a normal fire regime are unlikely to be present in

vegetation of eight years post-fire age. The changes were

not restricted to the non-sprouters as only 54% of the

original sprouting species were present in the second sur-

vey. It would appear that the short fire frequencies might

have also been detrimental to some sprouting species.

However, the relative dominance of sprouting species

has increased between the surveys with species such as

Hypodiscus aristatus and Chondropetalum nudum show-

ing considerable increases in cover.

Evidence from this single site supports the findings of

Vlok (1996) that short fire frequencies result in an

increase in the coverage of sprouting species, and as a

result, a decline in non-sprouters and in overall diversity.

The most plausible explanation for this change is that

short fire frequencies eliminate non-sprouting species (as

was the case at this site) thereby reducing competition

and favouring sprouters.

1.2.2. Long intervals between fires

Eighteen of the sites have experienced at least one

long (> 30 year) fire interval since the fire prior to

Taylor’s survey and have not been invaded by alien veg-

etation. On average these sites showed a higher similari-

ty in species composition between surveys (65.2%) than

the average for the complete data set. These sites were

used to examine the effects of long intervals between

fires on the numbers of individuals of serotinous and

myrmecochorous Proteaceae.

Bothalia 31,1 (2001)

107

TABLE 2. — Comparison of response of serotinous and myrmeco-

chorous Proteaceae from 18 (5 x 10 m2) sites sampled in 1966

and 1996, which experienced a long interval (> 30 years)

between fires in Cape of Good Hope Nature Reserve

The response of serotinous and myrmecochorous

species to long fire intervals differed in the Reserve

(Table 2). Serotinous non-sprouting Proteaceae such as

Leucadendron coniferum, L. xanthoconus and the com-

mon L. laureolum showed a tendency to decline in sites

that have experienced long fire-free intervals (> 30

years). This supports the findings of Bond (1980) that fire

in senescent fynbos results in smaller daughter popula-

tions or even local extinction of serotinous species. As

predicted, myrmecochorous species showed an increase

in abundance and even colonized some sites after high

intensity fires. This supports the hypothesis that seedling

recruitment in these species is strongly and positively

related to local fire intensity (Bond et al. 1990). Adult

mortality is, however, often high after these intense fires.

A random survey of individuals of the myrmecochorous

species Leucospermum conocarpodendron and Mimetes

fimbrifolius in the vicinity of site 92 (Figure 10) showed

that 97% (n = 66) of the adult trees were killed in forty-

five year old vegetation during the 1991 summer wildfire.

Because of its sea-bound geography, the southern

Peninsula experiences unusually long fire intervals. This

has favoured species with unusual life histories such as

the myrmecochorous tree proteoids, Leucospermum

conocarpodendron and Mimetes fimbrifolius and the tree

erica, Erica tristis. It would appear that the myrmeco-

chorous tree Proteaceae are adapted to long fire-free

intervals which facilitate high seedling regeneration and

subsequent maturation after long-interval, intense fires.

Although adult survival is much higher in low intensity

fires, seedling recruitment is low and the species can be

expected to decline following repeated short intervals

between fires (Midgley et al. 1998).

There was no significant change in the proportion of

sprouting versus non-sprouting species recorded at the

two sample times (%2 = 2.8 x 10'6, P = 0.99 NS), nor the

number of occurrences of sprouting versus non-sprout-

ing species {yf- 0-37, 1 d.f., P - 0.53 NS). This suggests

that long fire-free intervals have not differentially

favoured either regeneration strategies in the Reserve.

It can be concluded that high intensity fires after long

(> 30 year) intervals have only minor effects on species

composition but may result in a decline of serotinous,

and an increase in myrmecochorous Proteaceae. The fre-

quency of occurrence and average abundance of L. cono-

carpodendron and M. fimbrifolius for all sites in the

Reserve has remained constant between surveys (fre-

quency; 1966, 11 sites, 1996, 10 sites, mean abundance;

1966, 1.8, 1996, 1.9).

2. Dynamics of serotinous Proteaceae

The species data sets collected for this study provide

an opportunity to explore the long-term (30 year) spatial

and temporal dynamics of Leucadendron laureolum , a

common serotinous species in the Reserve. We attempt

to invoke exogenous explanations for its population fluc-

tuations in terms of fire regime over the past 30 years.

Serotinous Proteaceae are well known for their tendency

for density fluctuations after fire. A variety of hypotheses

have been proposed to explain these fluctuations (see

Midgley 1989 for a review).

Leucadendron laureolum is very widely distributed in

the Reserve and its densities are known to fluctuate over

time (Le Maitre 1988; Cowling & Gxaba 1990). This

species is a habitat generalist (Williams 1972) which

Taylor (1969) described as a ‘weedy’ species capable of

forming dense stands in most of the fynbos communities

FIGURE 10. — Site 92, oligotrophic proteoid fynbos near Platboom:

vegetation was 45 years old when it burnt in an intense summer

fire in 1991. The species turnover between sampling times was

50%. A large proportion of this change was as a result of the

colonization of post-fire (asteraceous) species in 1996. The

intense fire resulted in high mortality of Leucospermum cono-

c carpodendron (tall proteoid bush in the middle of the 1966 pho-

tograph), but seedling recruitment of this species was high in

the area. The thicket of Acacia cyclops in the background in

1966 was destroyed by the 1991 fire and the post-fire seedling

cohort removed by clearing teams.

108

Bothalia 31,1 (2001)

TABLE 3. — Comparison of range and abundance of serotinous pro-

teoid species, Leucadendmn laureolum for 81 (5 x 10 m) sites

surveyed in 1966 and 1996 in Cape of Good Hope Nature

Reserve. Sites colonized: species absent in 1966 and present in

1996. Sites extinct: species present in 1966 and absent in 1996

on sandstone-derived soils of the southern Cape

Peninsula. A comparison of the distribution and density

of L. laureolum in 1966 and 1996 showed that at the

landscape scale this species has remained remarkably

constant over the last 30 years (Table 3).

However, at the scale of the 50 m2 site most popula-

tions have fluctuated considerably over this time. While

the number of sites occupied and overall abundance has

remained constant, the species has undergone coloniza-

tion and extinction at many sites (Table 3). L. laureolum

has become extinct on nearly 30% of the sites occupied

in 1966, whereas it has colonized a similar number of

new sites in the same period. Furthermore, its abundance

has remained constant at only nine out of the 26 sites in

which it was recorded in both surveys.

2. 1 . Causes of decline of serotinous Proteaceae (Figure 1 1)

Fire season influences the regeneration of serotinous

Proteaceae. Autumn fires favour regeneration of seroti-

nous species, while winter and spring fires result in poor

regeneration (Le Maitre 1988). Of the 15 sites at which

L. laureolum either declined or went extinct, 40% were

burnt at least once in winter. The winter fire was the last

in the fire sequence at all except one of these sites. Short

fire intervals (< 4 years) were probably responsible for

local extinction at a further two sites (14 and 48).

Senescence after long fire-free intervals may also have

contributed to the decline of this species at some sites.

However, seven sites only experienced summer and/or

autumn fires. Therefore nearly 50% of the sites at which

L. laureolum declined were exposed to fires during theo-

retically favourable seasons. The decline in species at

these sites could not be explained by fire regime and was

more likely related to dispersal and/or competitive

effects. Three of these sites had moderately dense stands

(2 000 to 10 000 stems per hectare) of L. laureolum pre-

sent on them in 1966. Population declines at these densi-

ties could therefore have resulted from high levels of

intraspecific competition. Cowling & Gxaba (1990) sug-

gested that the most plausible explanation for the shifting

abundance of L. laureolum is the dispersal of its small,

winged seeds by persistently strong southeasterly winds

in the summer months after fire. Recent research into the

desiccation tolerance of newly emerged Proteaceae

seedlings demonstrated that L. laureolum had only mod-

erate resistance to post-emergence desiccation (Mustart

& Cowling unpublished data). It would appear that for at

FIGURE 11. — Site 62, oligotrophic proteoid fynbos on the sandstone

hill above Brightwater. The 1966 photograph shows Leuca-

dendron laureolum as a common overstorey species at this site.

It was burnt in a natural autumn fire in 1979 and a prescribed

winter burn in 1988. The combined effects of these two fires

resulted in the local extinction of this species from site 62. It

would appear from the photographs that the autumn fires have

favoured the survival of the fire-tolerant adult Leucospermum

conocarpodendron’s (large proteoid bushes in 1996 photo-

graph) at this site.

least four sites in this study, climatic events were the

most likely explanation for population declines.

2.2. Causes of increase of serotinous Proteaceae (Figures

12, 13)

All sites, which have been colonized or shown an

increase in cover, have experienced at least one autumn or

summer fire (24 sites). Sixty seven percent of these were

burnt exclusively in summer fires and only one had a win-

ter fire as its last burn. Season of burn, therefore, appears

to be an important determinant of post-fire population

increases in L. laureolum: both summer and autumn fires

favour increases in density. Nine of the eleven sites colo-

nized were burnt exclusively in summer, while the

remaining two were last burnt in an autumn fire.

Colonization relies on effective dispersal of seeds into

uninhabited sites. The high proportion of sites colonized

following summer fires is most likely the result of disper-

sal of seeds into new sites by the strong SE winds during

the summer months. Prescribed burns tend to be under-

taken during the relatively windless autumn months

resulting in less wind dispersal of seeds. Rapid germina-

tion following winter rains also results in the lower

mobility of the species following autumn or winter burns.

Bothalia 31,1 (2001)

109

FIGURE 12. — Site 29, an unusual form of wet restioid fynbos on the

Smitswinkel Flats which Taylor (1984b) described as ‘a dis-

tinctive woodland-like variety of seepage scrub with scattered

Psoralea pinnata above a dense layer of grasses and Restion-

aceae’. This site has undergone considerable changes over the

last 30 years (only 56% similarity). While the composition of

the graminoid field layer has remained similar, a dense cover of

Leucadendron laureolum has replaced the scattered overstorey

of P. pinnata.

FIGURE 13. — Site 73, oligotrophic proteoid fynbos near Kanonkop.

These matched photographs demonstrate the highly mobile

nature of the non-sprouting serotinous species, Leucadendron

laureolum. This species only recruits after fire. In the 1966 sur-

vey this species was not recorded in the vicinity of site 73.

Between surveys the site burnt twice, the second time in 1983

was during the favourable autumn months. As a result in 1996,

the species was common throughout the area. Changes such as

this, which result in compositional shifts in understorey

species, were common at many sites between surveys.

2.3. The impact of L. laureolum overstorey on under

storey species diversity

The highly mobile nature of this species poses ques-

tions as to what effect dense stands have on understorey

species composition. The majority of work on the impact

of proteoid overstories has shown reductions in local

richness of understorey species under dense stands

(Campbell & Van der Meulen 1980; Esler & Cowling

1990; Cowling & Gxaba 1990). Cowling & Gxaba

(1990) found that stands with sparse overstories of L.

laureolum in the reserve had double the number of

understorey species per square metre than dense stands.

However, Vlok (1996) showed that high densities

(> 3000 plants per hectare) of overstorey proteoids

before a fire, enriched the local diversity levels of under-

storey species after a fire. Vlok (1996) noted that the spa-

tial scale at which local diversity was measured affected

the results of these studies. We used sites, which have

undergone changes in the density of their proteoid over-

storey to explore the consequence of these changes on

understorey community composition.

A linear regression model was used to explore the rela-

tionship between changes in the abundance of L. laure-

olum between surveys and the corresponding changes in

species diversity. Both changes in abundance and in diver-

sity were calculated by subtracting the values for 1996

from those of 1966. As succession can influence diversity,

only sites with vegetation age > 4 and < 20 years at both

sampling times were included in the analysis.

There was no significant relationship between changes

in overstorey cover and changes in species diversity in

this study (Figure 14). The 50% reduction in diversity

recorded by Cowling & Gxaba (1990) was in small

quadrats of 1 mr. This study utilized much larger qua-

drats (50 m2 ) thus allowing for greater heterogeneity and

patchiness in the overstorey and consequently better sur-

vival opportunities for a range of understorey shrubs.

The lack of pattern in this study could be the result of

there being insufficient sites with high densities of L.

laureolum. The increase in understorey diversity record-

ed by Vlok (1996) was under stands with densities of

greater than 3000 stems. ha1. It would appear that com-

petition from overstorey proteoids only becomes impor-

tant at very high densities (>3000 stems. ha1)- Light to

moderate densities of proteoid overstorey do not signifi-

cantly impact species diversity at the community scale.

The timespan of this study provides an indication that

over the medium term, low to medium densities of over-

110

Bothalia 31,1 (2001)

Change in abundance of overstorey shrub,

Leucadendron laureolum

FIGURE 14. — The relationship between fluctuations in the density of

L. laureolum over the last 30 years, and changes in species

diversity. The abundance values can be converted to approxi-

mate densities per hectare as follows: 1 = 1-1 000 stems. ha1;

2 = 1 000-2 000 stems. ha1; 3 = 2 000-10 000 stems.ha1.

storey proteoid shrubs have little effect on the diversity

of understorey species. Populations of overstorey pro-

teoids demonstrated high mobility at the site level, yet

remained remarkably constant at the landscape scale

over time and appear to only influence understorey

diversity at high densities.

taken in the Reserve since 1981. These measures have

probably been the most intensive continual clearing

operation undertaken anywhere in the world. As the den-

sity of alien plants on each site was recorded at both sam-

pling times, we were able to determine changes in densi-

ty of alien plants across the Reserve as well as to explore

the impacts of invasion on the indigenous vegetation.

3.1. A comparison of the alien threat in 1966 and 1996

The effectiveness of the Reserve’s alien control pro-

gramme was evident from the low occurrence and densi-

ties of alien plants in all except one site in the second sur-

vey (Table 4). In 1966 eight of the 81 sites contained

alien plants, whereas in 1996 only five sites were infest-

ed. In addition there were a further four sites (4, 25, 90

and 92) which showed evidence of having been invaded

by alien trees after the 1966 survey and that have subse-

quently been cleared prior to the second survey in 1996.

Acacia cyclops is the only species still posing a major

threat to the indigenous flora of the Reserve and is now

concentrated on the steep slopes in the extreme south-

west. This area has been included in future alien eradi-

cation planning. The only site that included a dense cover

of alien plants was site 97, which is located in this area

of the Reserve. Outside of this area, alien plants were

only encountered as scattered juveniles. The CGHNR

eradication programme uses a system of covering the

entire Reserve every two years to ensure that these

seedlings will not mature. Pinus pinaster and Acacia

saligna, which were both abundant at the time of the first

survey, have been brought almost completely under con-

trol, with only occasional juveniles recorded in 1996.

3. Alien plants

Much of the fynbos biome has been impacted by

dense stands of alien vegetation (Macdonald &

Richardson 1986; Richardson et al. 1992). The estab-

lishment of an overstorey of alien trees and shrubs

markedly reduces the richness of fynbos communities

(Richardson & Van Wilgen 1985; Richardson etal. 1989;

Holmes & Cowling 1997). The rate of fynbos recovery

following the removal of alien thickets declines with

increasing period of invasion (Holmes & Cowling 1997).

The CGHNR has a long history of invasion by alien trees

(Taylor & Macdonald 1985; Taylor et al. 1985;

Macdonald et al. 1989). Intensive mechanical control

measures, involving initial felling of stands and subse-

quent follow-up on a 2-year rotation have been under-

The alien eradication programme in the CGHNR has

been effective at considerably reducing the threat of alien

plants on the indigenous flora of the Reserve.

3.2. Impact of aliens on indigenous vegetation

3.2.1. Sites invaded between surveys and subsequently

cleared

The two sites (25 and 95) which were densely invad-

ed by alien trees in the period between surveys and sub-

sequently cleared have regained less than 50% of their

original species since clearing (mean Sorenson’s coeffi-

cient of 45.28 ±2.1). Sites 4, 18, 90 and 92 were only

moderately infested between surveys and have shown

TABLE 4. — Record of alien species in the 50 m2 sites in 1966 and 1996

Bothalia 31.1 (2001)

FIGURE 15. — Site 25, coastal dune thicket within 20 m of the Atlantic

coastline at Olifantsbos Bay. In 1966 the site was characterized

by a well-developed thicket. A. cyclops subsequently invaded

the site. These trees were removed in 1986 and the natural veg-

etation has since started to recover. The sprouting guild is

recovering well, while most of the non-sprouting species pre-

sent in 1966 have been lost and have been replaced by short-

lived post-disturbance species.

better recovery since clearing (mean Sorenson’s coeffi-

cient of 58.66 ± 13.6). Site 25 provided an opportunity to

explore the impacts of dense alien invasion on coastal

dune vegetation (Figure 15).

This site is situated on coastal dunes within 20 m of

the Atlantic coastline. In 1966 the vegetation was a dense

coastal dune thicket of at least 30 years post-fire age. It

was dominated by the thicket species Euclea racemosa,

Rhus laevigata and R. glauca with a lower abundance of

Pterocelastrus tricuspidatus and Cassine peragua. The

graminoids Ficinia ramosissima and Ischyrolepis

eleocharis dominated the ground layer. At that time there

were no alien plants on the site but moderate densities of

Acacia cyclops within a 200 m radius (Taylor 1969). The

site was invaded by this species during the following

twenty years, becoming densely infested before clearing

in 1986 (CGHNR, unpublished records).

Seven of the 14 species present in 1966 still occupied

the site in 1996. Of these, only Helichrysum dasyanthum

was a non-sprouter. The typical coastal non-sprouters

present in the initial survey such as Passerina paleacea,

Coleonema album and Metalasia muricata had become

locally extinct. Clearing of the alien trees has created

gaps which have permitted colonization by a group of

weedy, post-disturbance non-sprouting species including

FIGURE 16. — Site 97, coastal scree asteraceous fynbos at Platbank on

the False Bay coast near Cape Point. In 1966 the alien thicket

was outside of the site (dark trees in the top of the photograph).

Note the indigenous thicket of Tarchonanthus camphoratus in

the bottom right hand comer (lighter trees). The site was invad-

ed by 1973, and by 1996 Acacia cyclops covered the entire

area. The result is that the cover of indigenous species has

declined dramatically and most of the species have been elimi-

nated (e.g. the skeletons of T. camphoratus are evident in the

foreground of the 1996 photograph).

Cineraria geifolia, Senecio purpureus and Cullumia

squarrosa as well as the sprouting pioneer graminoids

Ehrharta villosa , Stenotaphrum secundatum and Penta-

schistis pallida. None of these species were present in

the 1966 thicket and, consequently, diversity on this site

has temporarily increased as a result of clearing tree

invaders. The six sprouting species that have persisted on

the site included E. racemosa, R. laevigata and R. glau-

ca. These species were again dominating the site in 1996.

The less abundant thicket species have all been lost as a

consequence of invasion.

The invasion of this site by dense stands of A. cyclops

had a major impact on the natural vegetation, especially

non-sprouting species. The indigenous species are, how-

ever, recovering well since the aliens were eradicated.

3.2.2. The impact of existing stands of alien plants

Only one of the sites in this study has received no

attention from clearing teams since the 1966 survey. Site

97 is situated below steep cliffs on the False Bay coast,

close to Cape Point (Figure 16). In 1966 it contained

only seven species, being dominated by a few low shrubs

and succulent species of the family Aizoaceae. There

112

Bothalia 31,1 (2001)

were no aliens on site, but by 1973, H. Taylor reported

that the site had become densely invaded. In 1996 A.

cyclops formed an impenetrable thicket over the entire 50

nr quadrat.

While the diversity of the site has remained constant

(seven species), the six indigenous taxa now cover less

than 5% of its area. The only species to persist between

surveys were Felicia fruticosa and Exornis microphylla,

which grow on the coastal edge of the site and the tree

species Tarchonanthus camphoratus. However, most of

the original T. camphoratus thicket which grew just out-

side of the site has been eliminated as a direct result of

invasion by A. cyclops (Figure 16).

Within less than 30 years A. cyclops has virtually

eliminated the vegetation of this site and much of this

geographically restricted community in the south of the

Reserve. If it were not for the intensive and highly effec-

tive alien management programme administered by

Reserve stalf over the last 1 5 years, a far greater propor-

tion of the Reserve’s vegetation would have been simi-

larly impacted by alien plants.

CONCLUSIONS

The vegetation of the CGHNR has undergone consid-

erable change over the last 30 years. On average there

has been a temporal turnover in species of 40% across

the entire Reserve. There was a negative relationship

between post-fire vegetation age and species diversity in

both the 1966 and 1996 data sets. In proteoid fynbos the

early post-fire succession was characterized by a pre-

dominance of resprouting graminoids, and ephemeral

species which live for less than five years. After five

years the vegetation composition stabilizes and is gener-

ally dominated by an overstorey proteoid layer. While

predictions from moister inland localities suggest that

fire-free intervals in excess of 30 years should result in

the establishment of thicket patches, fynbos communities

persisted even after 50 years in the Reserve. Thicket and

forest development is much slower in the dry and windy

southern Cape Peninsula than in the wetter mountain

ranges. Coastal thicket communities maintain a stable

composition and relative cover in the absence of fire.

Short fire frequencies (< 5 years) favoured fast-maturing

and sprouting species, while long fire-free intervals (> 30

years) resulted in high intensity fires which favoured

myrmecochorous Proteaceae and negatively impacted

serotinous Proteaceae. The common non-sprouting

serotinous proteoid shrub, Leucadendron laureolum

showed a high degree of spatial mobility over the sam-

pling period. However, despite its considerable changes

in distribution, the abundance and number of sites occu-

pied by this species remained constant at the landscape

scale over the last 30 years. Stands of alien plants have

impacted large areas of natural vegetation in the Reserve.

However, evidence from this study suggests that the nat-

ural flora is recovering well after clearing of alien stands.

The threat posed by alien plants on the natural vegetation

has been considerably reduced over the last 30 years.

Thus, while some of this change can be traced to

human interference (prescribed burning and alien

plants), most is a result of natural colonization and

extinction. Temporal diversity resulting from high levels

of colonization and extinction after patchy fires con-

tributes to the high species diversity in fynbos systems.

ACKNOWLEDGEMENTS

Our grateful thanks are due to the Western Cape

Regional Services Council for permission to undertake

research in the Cape of Good Hope Nature Reserve.

Thanks also to the management staff of the CGHNR,

especially Mr R. Emtzen who provided past manage-

ment records and useful interpretative discussions. Many

thanks to W. Bond, B. Lamont and R Zedler for com-

ments on an earlier version of this manuscript. This

research was financially supported by the Foundation for

Research and Development and Pew Charitable Trusts.

Footnote

Since the submission of this manuscript, Hugh Taylor

has passed away. His co-authors would like to acknowl-

edge Hugh’s pioneering research in fynbos, particularly

his meticulous research in the Cape of Good Hope

Nature Reserve, and his foresight in establishing the per-

manent plots used in this study.

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114

Bothalia 31,1 (2001)

APPENDIX 1. — Comparison of species composition in 81 sites in 1966 and 1996. Vegetation types abbreviated as follows: WET, wetlands; WRP,

wet restioid fynbos; MOP, rnesic oligotrophic proteoid fynbos; SED, sedgeland; SPF, sandplain proteoid fynbos; DAF, dune asteraceous

fynbos: CSA, coastal scree asteraceous fynbos

Bothalia 31,1 (2001)

115

APPENDIX 2. — The fire histories of all sites used in this study

Bothalia 31,1: 117-133 (2001)

Patterns of endemism within the Karoo National Park, South Africa

F. RUBIN*. A.R. PALMER**, and C. TYSON***

Keywords: checklist, endemic patterns. Karoo National Park, South Africa

ABSTRACT

A checklist of 864 plant taxa representing 355 genera and 93 families is given for the Karoo National Park. A total of

121 species are endemic to the Nama-Karoo, with Asteraceae the most common with 33 taxa, followed by

Mesembryanthemaceae with 26 taxa. Phytochorological affinities indicate that 19.8% of the species are from the Nama-

Karoo Biome only, another 19.8% are distributed over two biomes, and 41% of the species have a widespread distribution.

The Karoo National Park conserves 30% of the recognized endemics of the Nama-Karoo Biome.

INTRODUCTION

The Karoo National Park, ± 75 000 ha and established

in 1979, is situated in the Nama-Karoo (Rutherford &

Westfall 1986), ± five km WNW of Beaufort West

(Figure 1), 32°11'S-32°23'S and 22°15'E-22035'E. The

general landscape in which the Park occurs, extends from

the Great Karoo, across the Nuweveld Mountain Range,

which is part of the Great Escarpment, to the Upper

Karoo (King 1942), or Southern Interior Basin (Kruger

1983). Elevation in the Park ranges from 820 to 1 620 m

asl and mean annual rainfall varies from 175 mm in the

west to 254 mm in the central area, and 406 mm on top of

the Nuweveld Mountain Range (Dent et al. 1987).

Elements of four phytochoria are represented in the

Park. These are the Tongaland-Pondoland succulent thick-

ets (Moll & White 1978), Karoo-Namib elements (Werger

1978), Afromontane remnants (White 1983), and Sudano-

Zambesian grasslands (Werger & Coetzee 1978). A phy-

tosociological study described fifteen dominant plant

communities in the original 33 000 ha of the Karoo

National Park (Rubin & Palmer 1996). The two Veld Types

represented in the original section of the Park are the

Karroid Broken Veld of the Great Karoo (Acocks 1988:

Veld Type 26), and the Karroid Merxrnuellera Mountain

Veld replaced by Karoo vegetation (Veld Type 42).

The vegetation of the upper plateau, on top of the

Nuweveld Mountain Range, consists of montane grass-

land on undulating rocky dolerite. These higher areas

contain many taxa known to occur in the winter rainfall

area of the southwestern Cape, whereas areas with soils

derived from doleritic origin on the middle plateau sup-

port numerous summer rainfall species (Figure 2). This

combination has allowed for succulents of considerable

interest (P. Bruyns pers. comm.). The upper plateau of

the Nuweveld Mountain Range falls away in a northerly

* South African National Parks, Scientific Services, P.O. Box 176,

6573 Sedgefield, South Africa.

** ARC Range and Forest Institute, P.O. Box 101, 6140 Grahamstown,

South Africa.

*** Geodatec, Bartholomew Street, 6140 Grahamstown, South Africa.

MS. received: 1999-10-01.

direction (Figure 2), with a rapid change in vegetation

from the mesic vegetation to a xeric dwarf shrub land-

scape, typical of the lower elevations south of Beaufort

West. In these areas, dolerite-derived substrate and deep

soils are replaced by more fertile sandstone, siltstone and

mudstone-derived soils.

On the dolerite of a slightly lower middle plateau

(Figure 2), montane grassland is replaced by a grassy

shrubland. On the low-lying plains a strong aridity gra-

dient can also be detected from east to west, with spars-

er, grassy, succulent taxa in the west and dwarf shrubs

dominating to the east.

Rubin & Palmer (1996) created a baseline inventory

of the homogeneous plant communities of the Park. This

study explores the floristic composition of the Karoo

National Park, relative to the floristic richness of the rest

of the Karoo Biome, and the floristic relationship with

other biomes.

METHODS

Plant species data of the Karoo National Park have

been collected by many people since the proclamation of

the Park in 1979. Valuable contributions were made by

B. Randall, a botanist who was resident in the Park from

1987 to 1988, D. Shearing, who worked on a field guide

for Karoo plants, P. Bruyns, who concentrated on succu-

lents and geophytes, and C. van Ginkel who started the

phytosociological survey. A checklist is provided with

information included after each taxon, i.e. : author name,

collector’s name and number, specimen location, biome

distribution, and growth form.

Phytochorological affinities and patterns of

endemism of the taxa in this list were determined from

literature held at the Selmar Schonland Herbarium,

Grahamstown, namely Hutchinson & Phillips (1917),

Von Poellnitz (1933), Phillips & Dyer (1934), Jacobsen

(1960), Obermeyer (1961a-c), Nordenstam (1968),

Tolken (1969), Dyer (1975), Hilliard (1977), Ober-

meyer (1978), Court (1981), Hilliard (1981a, b), Bond

& Goldblatt (1984), Kiillersjo (1985), Kallersjo (1988),

118

Bothalia 31,1 (2001)

FIGURE 2. — The topography of the Karoo National Park against the Nuweveld Mountain Range,

Bothalia 31.1 (2001)

119

TABLE 1. — Number of species and their distribution in one. two or

three biomes in the Karoo National Park

BK. Bushmanland Karoo; F, Fynbos; G. Grassland; GK, Great Karoo;

KNP, Karoo National Park; NK, Nama-Karoo: S. Savanna; SK.

Succulent Karoo; ST. Subtropical Thicket; UK, Upper Karoo.

Kallersjo (1991), Van Wyk (1991), and Gerbaulet (1991a,

b). The species were assigned to various categories

based on the biome distinctions as originally mapped by

Rutherford & Westfall (1986), namely G. Grassland. S,

Savanna, ST, Subtropical Thicket, F, Fynbos, SK,

Succulent Karoo (Table 1). The Nama-Karoo Biome

(NK) was subdivided, according to rainfall and eleva-

tion, into three: the Bushmanland Karoo (BK), adjacent

to the Succulent Karoo, the more mesic Upper Karoo

(UK), north of Beaufort West, and the more xeric Great

Karoo (GK), south of Beaufort West (Figure 1; Table 1)

(Palmer & Hoffman 1997). The categories used in clas-

sifying the species' phytochorological affinities are: 1,

widespread (W), including distribution ranges extending

outside South Africa and exotics; 2, transgressor species

with ranges extending across three biomes, the most

common category being NK/SK/F; 3, transgressor

species with ranges extending across two biomes, the

most common category being NK/SK; 4, species whose

range is restricted to one biome only, the most common

category being NK; 5, endemics belonging to the Nama-

Karoo subdivisions: Bushmanland Karoo, Upper Karoo,

Greater Karoo and Karoo National Park Endemics

(Table 1).

Grass species were divided into C3 and C4 categories

with the help of Watson & Dallwitz (1992), and for the

rest of the plant species list, the growth form and possi-

ble synonyms for each species were obtained from the

PRECIS database. National Botanical Institute.

RESULTS

A total of 864 species and infraspecific taxa (see

checklist), representing 355 genera and 93 families (Table

2) was recorded in the study area. Two veld types are rep-

resented by the original section of the Karoo National

Park (Figure 3), namely the Karroid Broken Veld (VT 26)

which covers approximately 66% and Karroid Merx-

muellera Mountain Veld replaced by Karoo (VT 42)

which covers approximately 33% of the original 33 000

ha of the park (Acocks 1988). A total of 121 species

(14%) are endemic to the Nama-Karoo (Table 1). Species

of Asteraceae are the most common with 33 taxa, fol-

Loxton Road

Acocks

ZZm CENTRAL UPPER KAROO

| | MERXMUELLERA MOUNTAIN VELD REPLACED BY KAROO

flgl CENTRAL LOWER KAROO

f~H KARROID BROKEN VELD

| | MOUNTAIN RENOSTERBOSVELD

5 0 5 10 15 20 25 Kilometers

FIGURE 3. — The expansion of the

Karoo National Park and the

location of the Acocks (1988)

Veld Types in and surround-

ing the park.

120

Bothalia 31,1 (2001)

TABLE 2. — Number of genera and species in the plant families collected in the Karoo National Park. Families in bold type encompass >50%

of spp.

Montiniaceae

Rosaceae

Fabaceae

Geraniaceae

Oxalidaceae

Linaceae

Zygophyllaceae

Polygalaceae

Euphorbiaceae

Anacardiaceae

Celastraceae

Sapindaceae

Melianthaceae

Rhamnaceae

Tiliaceae

Malvaceae

Sterculiaceae

Tamaricaceae

Flacourtiaceae

Cactaceae

Thymelaeaceae

Myrtaceae

Onagraceae

Araliaceae

Apiaceae

Primulaceae

Ebenaceae

Oleaceae

Loganiaceae

Gentiaceae

Apocynaceae

Asclepiadaceae

Convolvulaceae

Boraginaceae

Verbenaceae

Lamiaceae

Solanaceae

Scrophulariaceae

Selaginaceae

Bigoniaceae

Pedaliaceae

Lentibulariaceae

Acanthaceae

Plantaginaceae

Rubiaceae

Dipsacaceae

Cucurbitaceae

Campanulaceae

Lobeliaceae

Asteraceae

TOTAL

1

2

14

4

1

3

2

3

1

5

1

2

1

2

1

6

1

2

1

2

15

2

9

2

7

6

11

4

1

I

5

1

3

1

3

1

54

1

2

34

28

5

2

9

15

9

2

1

7

20

1

7

1

2

1

7

1

5

2

3

1

2

24

5

10

5

13

18

31

12

2

1

11

1

7

1

6

2

I

170

93 355 864 26

lowed by Mesembryanthemaceae with 26 taxa. The only

species with Red Data status (Hilton-Taylor 1996) are

Cliffortia arborescens which is considered to be rare, and

Anisodontea malvastroides which is threatened.

Of the 864 species listed in the Karoo National Park

(Table 2), 356 species (41.1%) have a widespread distri-

bution, i.e. over more than three biomes (Table 1). Of the

remaining species, 170 species (19.8%) were from the

Nama-Karoo Biome only, another 171 species were dis-

tributed over two biomes, of which the Nama-Karoo-

Succulent Karoo (NK/SK) mix was the highest with 56

species, followed by Nama-Karoo-Fynbos (NK/F) and

Nama-Karoo-Savanna (NK/S) both with 25 species

(Table 1). Of the species distributed over three biomes,

the Nama-Karoo-Succulent Karoo-Fynbos (NK/SK/F)

combination is the highest, with 37 species, followed by

Nama-Karoo-Fynbos-Savanna (NK/F/S) (23 species)

(Table 1). Twenty-three of the species listed are natural-

ized exotics (Table 2).

Bothalia 31,1 (2001)

121

DISCUSSION

The Nama-Karoo Biome includes 21 Veld Types

(Acocks 1988), and ±21 47 taxa, 18% of which are

endemic (Gibbs Russell 1987). Although the Karoo

National Park only effectively conserves representative

samples of one of these 21 Veld Types, namely the

Karroid Broken Veld (VT 26), it nevertheless protects

30% of the recognized flora of the Nama-Karoo Biome.

The Park is therefore an important conservation area.

Nine families (in bold. Table 2) encompass over 50% of

the species in the Park. A similar pattern was noted in the

Karoo Nature Reserve near Graaff-Reinet (Palmer

1989), and as with Gibbs Russell (1987) no differential

occurrence could be found for important families of the

Nama-Karoo Biome. Therefore, unlike the other five

major biomes, the Nama-Karoo Biome does not appear

to be floristically distinct (Gibbs Russell 1987).

The major biomes fall into two groups, namely those

with winter rainfall and those with summer rainfall

(Gibbs Russell 1987). The Nama-Karoo Biome, which is

distinguished by extreme temperature, (i.e. very hot sum-

mers and very cold winters,) and autumn and spring rain-

fall peaks (Rutherford & Westfall 1994), shares half of

its species with that of the Savanna Biome. A further

30% of the Karoo Biome species are shared with the

Grassland Biome and a third with the Fynbos Biome

(Gibbs Russell 1987).

The Nama-Karoo Biome remains one of the least stud-

ied of southern Africa’s biomes (Palmer & Hoffman

1996). The recent (1997-2000) extensions of the Park

incorporate further sections of the Karroid Broken Veld

(Veld Type 26), and to a lesser extent, Merxmuellera

Mountain Veld replaced by Karoo vegetation (Veld Type

42), as well as small areas of previously excluded Veld

Types, namely Central Upper Karoo (Acocks 1988: Veld

Type 27) and Mountain Renosterbosveld (Veld Type 43)

(Figure 3). Central Upper Karoo or Upper Nama-Karoo

(50) in Low & Rebelo (1996), occupies the central part of

the upper plateau of the Upper Karoo, while the Mountain

Renosterbosveld or Escarpment Mountain Renosterveld

(60) in Low & Rebelo (1996), is located on the slopes and

foothills of the Nuweveld Mountains. Very little is known

about this form of Renosterveld and the conservation sta-

tus is very poor (Low & Rebelo 1996).

The only other Veld Type adjacent to the Park is the

Central Lower Karoo (Acocks 1988: Veld Type 30), or

Central Lower Nama-Karoo (54) in Low & Rebelo

(1996). The area supporting this Veld Type is separated

from the Park by the national road (Nl), is flat and stony

and plant species given in Acocks (1988) appear to be

similar to those in the Park. Nevertheless, this Veld Type

was given a high conservation priority rating by Hilton-

Taylor & Le Roux (1989), presumably because it was not

included in a conservation area, and should be consid-

ered for inclusion in the Park.

ACKNOWLEDGEMENTS

The National Botanical Institute is thanked for the use

of data from the National Herbarium, Pretoria (PRE)

Computerized Information System (PRECIS), as well as

the staff of the National Herbarium for the many speci-

mens that were identified. Many other specialists have

contributed to identification and collection: P. Bruyns

(Bolus Herbarium, UCT), E. Brink (Selmar Schonland

Herbarium, Grahamstown), D. Shearing (Layton Her-

barium, now Southern Cape Herbarium), and previous

personnel, students or inhabitants of the Karoo National

Park, i.e. B. Randall, B.K. Bengis, T. Braak and C. van

Ginkel, and visitors such as B.A. Dyer, G.E. Gibbs-

Russell, M.B. Bayer, and R. Rishworth.

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CHECKLIST OF THE VASCULAR PLANTS OF THE KAROO NATIONAL PARK

The nomenclature and arrangement of the checklist follow Arnold & De Wet ( 1993), with synonyms/name changes

provided by the National Botanical Institute. The species name (in roman type) is followed by the author name (in

italics) according to Brummitt & Powell (1992), collector name and number(s) (in italics), specimen location, biome

distribution and growth form. The categories are separated by commas. Lichens and fungi do not have biome distri-

bution and growth form details. In grasses, C3 and C4 depict the photosynthetic pathways.

Abbreviations used:

BOL, Bolus Herbarium; BK, Bushmanland Karoo; cl, climber; Cp, Cape; dw, dwarf shrub; E, Endangered; F,

Fynbos; FS, Free State; g, geophyte; G, Grassland; GK, Great Karoo; gr, graminoid; GRA, Grahamstown Herbarium;

h, herb; hy, hydrophyte; K, Insufficiently known; KRNP, Karoo National Park; N, KwaZulu-Natal; NK, Nama-

Karoo; p, parasite; R, Rare; S, Savanna; sc, scrambler; SCH, Layton Herbarium; sh, shrub; SK, Succulent Karoo; ST,

Subtropical thicket; su, succulent; t, tree; UK, Upper Karoo; W, widespread; ?, unknown distribution; *, exotic; +,

endemic; [ J, occasional growth form.

LICHENS

PARMELIACEAE

Xanthoparmelia sp. ( Parmelia hottentotta ), Shearing s.n., SCH, NK/F/S

USNEACEAE

Usnea undulata. Shearing s.n., SCH, W

FUNGI

TULOSTOMATACEAE

Podaxis pistillaris, Shearing s.n., SCH, W

PTERIDOPHYTA

OPHIOGLOSSACEAE

Ophioglossum polyphyllum A. Braun, Brctack 483, SCH, ?, h, g

MARSILEACEAE

Marsilea sp., Bruyns s.n., BOL, W

ADIANTACEAE

Adiantum capillus-veneris L., Shearing s.n., SCH, W, h, [cl]

Cheilanthes

capensis (Thunb.) Sw., Palmer & Van Ginkel 2908, GRA, W, h

eckloniana ( Kunz.e ) Mett., Braack 348, SCH, W, h

hirta Sw. var. hirta, Shearing s.n., Bruyns 3323, 3400, Rubin 40, BOL,

KRNP, W, h

induta Kunz.e, Braack 348, SCH, KRNP, Cp, h

quadripinnata ( Forssk .) Kuhn, Rubin s.n., KRNP, W, h

Pellaea

calomelanos (Sw.) Link var. calomelanos, Braack 491, SCH, W, h

rufa A.F.Tryon, Braack 49, SCH, Cp, h

ASPLENIACEAE

Asplenium

trichomanes L., Bruyns 3345 , BOL. SCH, W, h

ceterach cordalum (Thunb.) Desv., Palmer & Van Ginkel 1844, 2805,

GRA, W, h

ANGIOSPERMAE

MONOCOTYLEDONAE

TYPHACEAE

Typha capensis (Rohrb.) N.E.Br, Randall 27414, SCH, W, h, hy

POACEAE

Andropogon

chinensis (Nees) Merr., NK/G/S, C4

schirensis A. Rich., S/G, C4

Cymbopogon

dieterlenii Stapfex E. Phillips, Rubin 26766, KRNP, W, C4

excavatus (Hochst.) Stapf ex Burtt Davy, Palmer & Van Ginkel 2713,

2755, GRA, W, C4

plurinodis ( Stapf) Stapf ex Burtt Davy, W, C4

prolixus (Stapf) E. Phillips. Rubin 65, 3205, SCH, KRNP, W, C4

validus (Stapf) Stapf ex Burtt Davy, Shearing 856, SCH, W, C4

Bothalia 31,1 (2001)

123

Hyparrhenia hirta ( L .) Slap/, Braack 422, Rubin 3172, 36914, KRNP,

W, C4

Heteropogon contortus ( L .) Roem. & Schult., Palmer & Van Ginkel

1846, GRA, W. C4

Themeda triandra Forssk.. Palmer & Van Ginkel 2719, Rubin 26773,

GRA. KRNR W, C4

Digitaria

argyrograpta ( Nees ) Stapf Palmer & Van Ginkel 2682, 2842, Rubin

26908, GRA. KRNR NK/G/S, C4

eriantha Steud., Palmer & Van Ginkel 1845, 2726, 2735, GRA. W, C4

Panicum

deustum Thunb., Shearing 27405, SCH. NK/S/STh, Cj/C3

maximum Jacq ., Shearing s.n., SCH. W, C4/C3

Setaria verticillata ( L .) P.Beauv., Shearing s.n., SCH, W. C4

Pennisetum setaceum (Forssk.) Chiov.. Rubin s.n., KRNP. W, *

Cenchrus ciliaris L.. Palmer & Van Ginkel 2717, 2794, 2839, Rubin

26720, GRA. KRNP, W. C4

Ehrharta calycina Sm. var. calycina. Palmer & Van Ginkel 2752, Rubin

3209. GRA. KRNP. W, C3

Helictotrichon sp.. Shearing 855, SCH, ?, C3

Merxmuellera

disticha (Nees) Conert, Palmer & Van Ginkel 2752, Rubin 26910,

GRA. KRNP. NK/F/G, C3, [hy]

stricta (Schrad.) Conert, Rubin 26777, KRNP. NK/F/G. C3

Karroochloa purpurea (L.f.) Conert & Tiirpe, Rubin 2657, KRNP. W. C3

Pentaschistis

airoides (Nees) Stapf

subsp. airoides. Palmer & Van Ginkel 2893, 2916, GRA, NK/SK/F. C3

subsp. jugorum (Stapf) H.P.Linder, Rubin 29, G, C3

ecklonii (Nees) McCIean, Rubin 3208, KRNP. F, +, C3

patula (Nees) Stapf, Palmer & Van Ginkel 2919. GRA, SK/F, C3

setifolia (Thunb.) McCIean, Rubin 3211, 26768, KRNP, G, C3

Chaetobromus dregeanus Nees, Rubin 3213, KRNP. NK/SK/F. C3

Arundo donax L., pers obs.. W, C3

Phragmites australis (Cav.) Steud., Shearing s.n., SCH, W, C3

Agrostis lachnantha Nees var. lachnantha. Shearing 911, SCH, W, C3

Polypogon

monspeliensis (L.) Desf. Bengis 450, SCH. W, C3, *

viridis (Gouan) Breistr., Palmer & Van Ginkel 2469, GRA. W. C3, *

Stipagrostis

ciliata (Desf.) De Winter var. capensis (Trin. & Rupr.) De Winter,

Palmer & Van Ginkel 2769, Rubin 26915, GRA. KRNP,

NK/SK/S. C4

namaquensis (Nees) De Winter, Shearing s.n., SCH. NK/SK/S. C4, sh

obtusa (Delile) Nees, Braack 65, Rubin 26904, KRNP. NK/SK/S, C4

Aristida

adscensionis L., Shearing s.n., SCH, W, C4

congesta Roem. & Schult. subsp. congesta, Palmer & Van Ginkel

1868, 2733, GRA. W. C4

diffusa Trin. subsp. diffusa. Palmer & Van Ginkel 1830, Rubin 26909,

GRA. KRNP. NK/G/S, C4

Tragus

berteronianus Schult., Palmer & Van Ginkel 1831, GRA, W, C4

koelerioides Asch., Palmer & Van Ginkel 2672, GRA. NK/G/S, C4

racemosus (L.) All., Shearing s.n., SCH, W, C4

Sporobolus fimbriatus (Trin.) Nees, Palmer & Van Ginkel 2809, GRA.

W, C4

Eragrostis

bergiana (Kunth) Trin., Rubin 53, NK (GK/UK), +, C4

chloromelas Steud., Palmer & Van Ginkel 2748, GRA, NK/G/S, C4

curvula (Schrad.) Nees, Palmer & Van Ginkel 2090, 2723, Rubin

3174, 3206, GRA. KRNP, W, C4

lehmanniana Nees var. lehmanniana. Palmer & Van Ginkel 1812,

GRA, W. C4

obtusa Munro ex Ficalho & Hiem, Palmer & Van Ginkel 1810, Rubin

26996, GRA, KRNP. W, C4

porosa Nees, Rubin 89, KRNP. NK/S (GK/UK), C4

x pseud-obtusa De Winter, NK/S (UK), C4

Cynodon

dactylon (L.) Pers., Shearing s.n., SCH. W, C4

incompletus Nees, Randall 27343, KRNP, W, C4

Oropetium capense Stapf, NK/G/S, C4

Diplachne fusca (L.) P.Beauv. ex Roem. & Schult., Bengis 462, SCH,

W, C4

Enneapogon

cenchroides (Roem. & Schult.) C.E.Hubb., Palmer & Van Ginkel 2873 ,

GRA. W, C4

desvauxii P.Beauv., Palmer & Van Ginkel 2487, GRA, W, C4

scaber Lehm., Palmer & Van Ginkel 1809, 2676, Rubin 26907, 26916.

GRA. KRNP, NK (BK), +, C4

scoparius Stapf, Palmer & Van Ginkel 2718, 2892, GRA, W. C4

Fingerhuthia

africana Lehm., Palmer & Van Ginkel 1811, 2814, Rubin 26929,

GRA, KRNP, W, C4

sesleriiformis Nees, Palmer & Van Ginkel 2906, GRA, NK/G/S, C4

Koeleria capensis (Steud.) Nees, Rubin 3207 , KRNP. W, C3

Melica

decumbens Thunb., Bengis 367, Rubin 3210, KRNP. NK/G, C3

racemosa Thunb., Shearing 853, Rubin 3244, KRNP, W, C3

Festuca scabra Vahl, Bengis 383, KRNP, W, C3

Bromus

leptoclados Nees. Bengis 396, KRNP. W, C3

pectinatus Thunb., W, C3

sp., Rubin 3213, KRNP, NK/G/S. C3

Brachypodium bolusii Stapf, NK/G, C3

Hordeum marinum Huds. subsp. gussoneanum (Pari.) Theli, Rubin

3243, KRNP. F/SK/G, C3

Schismus inermis (Stapf) C.E.Hubb., Sugden s.n., Cp, C3

CYPERACEAE

Carex zuluensis C.B. Clarke, Sugden s.n., W, h, gr

Cyperus

bellus Kunth, Shearing s.n., SCH, W, h, gr

laevigatus L., Shearing s.n., SCH, NK/F/S, h, gr

longus L. var. longus. Palmer & Van Ginkel 2091, GRA, W. h, gr

marginatus Thunb., Shearing s.n., SCH, W, h, gr

rupestris Kunth, Sugden s.n., W, h, gr

tenellus L.f. var. tenellus. Palmer & Van Ginkel 2102, GRA, NK/F/S,

h, gr

usitatus Burch, var. usitatus, Rubin 28, SCH, W, h, gr, g

Mariscus capensis (Steud.) Schrad., Rubin 106, SCH, W, h, gr

Ficinia

lateralis (Vahl) Kunth, Palmer & Van Ginkel 2096, GRA, F/STh, h, gr

ramosissima Kunth, Rubin 3257, KRNP, NK/F/S (GK), h, gr

Scirpoides dioecus (Kunth) Browning, Palmer & Van Ginkel 2093,

Rubin 3199, GRA. KRNP. W, gr, [h]

Pseudoschoenus inanis (Thunb.) Oteng-Yeb., Bengis 465, Palmer &

Van Ginkel 2092, GRA, NK/SK/STh, h, gr

Isolepis prolifer (Rottb.) R.Br., Bengis 464, KRNP. W, h, gr

RESTIONACEAE

Restio bifidus Thunb., Sugden s.n., Cp, h, gr

Rhodocoma fruticosus Thunb., Sugden s.n., Cp/N. h, gr

COMMELLNACEAE

Commelina sp., Bruyns 3393, BOL, W, h

JUNCACEAE

Juncus

acutus L., Sugden s.n., Cp, h

dregeanus Kunth, Shearing s.n., SCH, F/S/G, h

exsertus Buchenau subsp. exsertus, Randall 27356, SCH, W, h, hy

rigidus Desf, Bengis 461, SCH, W. h, hy

rupestris Kunth., Sugden s.n., Cp, h, hy

punctorius L.f, Shearing s.n., SCH, NK/SK/F, h, hy

COLCHICACEAE

Androcymbium melanthioides Willd. var. melanthioides. Shearing s.n.,

SCH, W, g

Omithoglossum undulatum Sweet, Shearing s.n., SCH, F, g

ASPHODELACEAE (Part A)

Bulbine

abyssinica A. Rich., Bengis 430, Rubin 26776, KRNP, W, h, su

filifolia Baker, Bruyns 3413. BOL. W, g, su

frutescens (L.) Willd., Bengis 335, SCH, W, h, su

narcissifolia Salm-Dyck, Bengis 361, SCH, W, h, su

Trachyandra

acocksii Oberm., Shearing s.n., SCH, NK, +, dw

saltii (Baker) Oberm. var. saltii, Bruyns 3343, BOL, W, h

thyrsoidea (Baker) Oberm., Shearing s.n., SCH, F, h

HYACINTHACEAE (Part A)

Schizobasis intricata (Baker) Baker, Palmer & Van Ginkel 2702, GRA,

W, g, [cl]

ERIOSPERMACEAE

Eriospermum

capense (L.) Thunb. subsp. capense, Bruyns 3983, BOL, NK/SK, g

paradoxum (Jacq.) Ker Gawl.. Bruyns 3394. BOL, NK/SK/F, g

124

Bothalia 31,1 (2001)

ASPHODELACEAE (Part B)

Kniphofia

caulescens Baker , Bengis 381 , SCH, NK/G (GK), h

uvaria (L.) Oken, Palmer & Van Ginkel 2913, SCH, GRA, W, h

Aloe

aristata Haw., Bruyns s.n. , BOL, W, h, su

broomii Schonland var. broomii, Bengis 489. SCH, NK/G (GK/UK),

dw, h, su

claviflora Burch., Shearing s.n., SCH, NK, +, h, su

longistyla Baker, Bruyns 3424, BOL, GK, +, h, su

microstigma Salm-Dyck, Shearing s.n., SCH, NK/STh (GK), sh, su

microstigma x claviflora, BOL, NK/STh (UK), h, su

striata Haw. subsp. striata. Shearing s.n., SCH, NK/F, sh, dw, su

striatula Haw. var. striatula. Palmer & Van Ginkel 2495, GRA, NK/G,

sh, su

variegata L., Shearing s.n., BOL, W, h, su

Gasteria disticha (L.) Haw., Bruyns 3974, BOL, NK/F, h, su

Astroloba herrei Uitewaal, Bruyns 3419, BOL, Cp, su

Haworthia

marumiana Uitewaal, Cp, su

nigra (Haw.) Baker, Bruyns 3381, BOL, GK, +, su

semiviva ( Poelln .) M.B.Bayer, Bruyns 3993, BOL, NK/STh, su

venosa (Lam.) Haw. subsp. tessellata (Haw.) M.B.Bayer s.n., Bruyns

3379, BOL, Cp, su

sp„ Bruyns 3383, BOL, NK/SK, su

ALLIACEAE

Tulbaghia

capensis L., Bruyns 4028, BOL, F, h

leucantha Baker, Bruyns 3350, SCH, BOL, NK/SK/F/G, h

HYACINTH ACEAE (Part B)

Albuca

juncifolia Baker, Braack 81 , KRNP, NK/F (GK), g

setosa Jacq., Shearing s.n., SCH, NK (GK/UK), +, g

Urginea sp.. Bruyns 3977, BOL. ?, g

Drimia anomala (Baker) Benth., Bruyns 3423, BOL, Cp/N , g

Rhadamanthus piatyphyllus B.Nord., BOL, NK/S, g

Dipcadi viride (L.) Moench, Palmer & Van Ginkel 2608, GRA, W, g

Scilla sp., ?, g

Omithogalum

hispidum Homem. subsp. hispidum, Bruyns 3987, BOL, NK/SK/F

(GK), g

juncifolium Jacq., Shearing s.n., SCH. W, g

tenuifolium F.Delaroche subsp. aridum Oberm., Shearing s.n., SCH,

NK/S, g

zebrinellum U.& D.MiiU.-Doblies, Bruyns 3985, BOL, NK/SK (GK), g

sp., Bruyns 3372, BOL, ?, g

Ledebouria sp., Rishworth 53, KRNP27653, ?, g

Veltheimia capensis (L.) DC., Shearing s.n., SCH, NK/SK (GK), g

Lachenalia sp., Shearing s.n., SCH, ?, g

Massonia

echinata L.f., Palmer & Van Ginkel 2915, GRA. NK/F, g. [su]

pustulata Jacq., Bruyns 3358, BOL, NK/SK/F, g, [su]

ASPARAGACEAE

Asparagus

acocksii Jessop, Shearing s.n., SCH, W, sh (cl)

aethiopicus L., Palmer & Van Ginkel 1861, Rubin 26743, GRA,

KRNP, F/STh, cl

africanus Lam., Shearing s.n., SCH, W, cl

burchellii Baker, Shearing s.n., SCH. NK/SK/F, sh

capensis L. var. capensis, Palmer & Van Ginkel 1850, 2519, SCH,

GRA, NK/F/S, sh

cooperi Baker, Rubin 88, KRNP. W, sh, h, cl

crassicladus Jessop, Palmer & Van Ginkel 2122, 2690, GRA, Cp, su

exuvialis Burch, forma exuvialis, Bayer 145, Palmer & Van Ginkel

1836, GRA, NK/S/STh, sh

glaucus Kies, Rubin 26739, KRNP, NK, +, sh

kraussianus ( Kunth ) J.F.Macbr., Bruyns 3357, BOL, F, cl

laricinus Burch., W, sh

microraphis (Kunth) Baker, Rubin 26790, KRNP, W, sh

mucronatus Jessop, Shearing s.n., SCH, NK/SK/F/G south of 28, sh

retrofractus L., Shearing s.n., SCH, NK/SK, cl

striatus (L.f) Thunb., Shearing s.n.. Rubin s.n., KRNP, SCH, NK/STh, sh

suaveolens Burch., Rubin 97, KRNP, W, sh

AMARYLLIDACEAE

Haemanthus

humilis Jacq. subsp. humilis, Shearing s.n,, SCH, W, g

sp.. KRNP27402, 7, g

Boophane disticha (L.f.) Herb.. Shearing s.n., SCH, W, g

Strumaria gemmata Ker Gawk, Shearing s.n., SCH, NK/SK/F/S, g

Gethyllis sp., Rubin 41, KRNP, NK/SK, g

Ammocharis coranica (Ker Gawl.) Herb., Shearing s.n., SCH, W, g

Cyrtanthus smithiae Watt ex Harv., Bruyns 3989, BOL, ST, h, g

HYPOXIDACEAE

Empodium sp., Braack 134, ?, g

IRIDACEAE

Romulea macowanii Baker var. macowanii, Braack 79, Palmer & Van

Ginkel 2164, GRA, NK/G (UK), h, g

Moraea

ciliata (L.f.) Ker Gawl., Braack 80, KRNP, W, h, g

crispa Thunb., Bruyns 3356, BOL, NK/F, h, g

polystachya (Thunb.) Ker Gawl., Shearing s.n., SCH, W, h, g

Homeria

cookii L. Bolus, Bruyns 3361 , BOL, W, h, g

miniata (Andrews) Sweet. Shearing s.n., SCH, NK/SK/F (GK), h, g

Hesperantha radiata (Jacq.) Ker Gawl., Bruyns 3346, BOL, W, h, g

Ixia marginifolia (Salisb.) G.J. Lewis, Bengis 375, KRNP, Cp, h, g

Tritonia karooica M.P.de Vos, Shearing s.n., SCH, NK/SK/F, h, g

Babiana hypogea Burch, var. ensifolia G.J. Lewis, Shearing s.n., SCH,

NK, +, h, g

ORCH1DACEAE

Holothrix

grandiflora (Solid.) Rchb.f, Bruyns 3983, BOL. SK, h. g

villosa Lindl. var. villosa, Bengis 444, KRNP, NK/F, h, g

Anochilus flanaganii (Bolus) Rolfe, Bengis 404, KRNP, G, g

DICOTYLEDONAE

SALICACEAE

Salix mucronata Thunb. subsp. capensis (Thunb.) Immelman, Shearing

s.n., SCH, W, t

Populus alba L., Sugden s.n., W, t, *

URTICACEAE

Urtica urens L., Shearing s.n., SCH, W, h

Forsskaolea Candida L.f, Rubin 26798, 26923, KRNP, NK/SK, h

LORANTHACEAE

Moquiniella rubra (A.Spreng.) Bade. Rubin 26997, KRNP, W, sh, p

Septulina glauca (Tlninb.) Tiegh.. Rubin 26935, KRNP, NK/SK/F, sh, p

VISCACEAE

Viscum

capense L.f. subsp. capense, Randall 27336, W, sh, p

continuum E.Mey. ex Sprague, Palmer & Van Ginkel 2476, GRA,

NK/F/S (GK), sh, p

rotundifolium L.f, Palmer & Van Ginkel 1815, GRA, W, sh, p

SANTALACEAE

Thesium

acutissimum A. DC., Rubin 26770, KRNP. W, h, p, [sh]

gnidiaceum A. DC. var. gnidiaceum, Bengis 343, KRNP. G/STh, h, p,

[dw]

hystricoides A. W. Hill, Bengis 459, KRNP. SK, dw, p, [sh]

imbricatum Thunb., Rubin 3225, KRNP, W. dw, p, [sh]

lineatum L.f., Palmer & Van Ginkel 1816, 2801, Rubin 26732, GRA,

KRNP, NK/SK, sh, p, [dw]

oresigenunt Compton, KRNP. NK/F/G (GK), sh, p, [h]

triflorum Thunb. ex L.f, Shearing s.n., SCH, NK/F/S, h, p, [cl]

POLYGONACEAE

Rumex crispus L., Rubin 24, KRNP, W, sc, *

CHENOPODI ACEAE

Chenopodium

album L., Palmer & Van Ginkel 2492, 2689, 2727, 2785, GRA,

KRNP. W. h

ambrosioides L., Palmer & Van Ginkel 2798, GRA, W, h

foliosum Asch., Rubin 3245, KRNP, GK, +, h

glaucum L, Rubin 3255, KRNP, W, h

Atriplex

lindleyi Moq.

subsp. inflata (F.Muell.) Paul G. Wilson, Shearing s.n., SCH. W, h

subsp. quadripartita Paul G. Wilson, STh, h

nummularia Lindl. subsp. nummularia, Shearing s.n., SCH, NK/SK/G,

sh, *

semibaccata R.Br. var. semibaccata, Shearing s.n., SCH. W, h, sh

Bothalia 31.1 (2001)

125

Atriplex (cont.)

vestita ( Thunb .) Aellen

var. appendiculata Aellen, Rubin 26748 , KRNP, W, sh

var. inappendiculata Aellen, Palmer & Van Ginkel 2483, GRA, W, sh

Bassia

diffusa (Thunb.) Kuntze, Shearing s.n., GK, +, h

salsoloides (Fenzl) A. J. Scott, Palmer & Van Ginkel 2788, GRA, NK/

SK/STh, dw

Suaeda fruticosa ( L .) Forssk., Palmer & Van Ginkel 2474, GRA. KRNR

NK/S. dw

Salsola

aphylla L.f, Shearing s.n., SCH, NK/SK/G, sh

calluna Fenzl ex C.H.Wright, Shearing s.n., SCH, NK, +, dw

dealata Botsch., Shearing s.n., SCH, NK, +, dw

kali L„ Bengis 514, Randall 27415, KRNP. W, h

minutifolia Botsch., Bengis 331, KRNP, NK, +, dw

smithii Botsch., Shearing s.n., SCH, NK, +, dw

tuberculata ( Moq .) Fenzl, Rubin s.n., KRNP, NK, +, dw

AMARANTHACEAE

Amaranthus praetermissus Brenan, Palmer & Van Ginkel 2100, Rish-

worth 40, Rubin 26730, GRA, KRNP. NK/SK/G, h

Sericocoma avolans Fenzl. Palmer & Van Ginkel 2838, Rishworth 4.

Rubin 26710, GRA, KRNP. NK/SK (GK). dw

NYCTAGINACEAE

Boerhavia cordobensis Kuntze, Palmer & Van Ginkel 2769, GRA, W, h

AIZOACEAE

Limeum

aethiopicum Burnt, subsp. aethiopicum var. aethiopicum, Rubin s.n.,

KRNP, NK/SK/STh (GK). h, [dw]

humifusum Friedrich, GRA, NK. +, dw

Gisekia

phamacioides L. var. phamacioides. Palmer & Van Ginkel 2787,

Rubin 26928, GRA. KRNP, W, h

sp„ KRNP, ?, h

Phamaceum

albens L.f, Rubin 3227, NK/SK, dw

dichotomum L.f, Rubin s.n., KRNP, W, h, [dw]

lineare L.f, Palmer & Van Ginkel 2855, GRA, SK/F. h

Hypertelis

bowkeriana Sond., Palmer & Van Ginkel 2767, 2877, GRA, W, h

salsoloides (Burch.) Adamson var. salsoloides, Rishworth 60. KRNP27660.

KRNP. W, dw

Trianthema triquetra Rottler ex Willd. subsp. triquetra var. triquetra. Shear-

ing s.n., SCH. W. h

Plinthus karooicus I.Verd., Rubin s.n., KRNP, NK/S, dw

Galenia

africana L. var. africana, Randall 27397, KRNP, W, dw

fruticosa (L.f.) Sond.

var. fruticosa. Shearing s.n., SCH. NK/SK. dw

var. prostrata Adamson, Rubin s.n., KRNP, NK/SK, dw

papulosa ( Eckl . & Zeyh.) Sond. var. papulosa, Bengis 456, KRNP,

NK/SK, h

procumbens L.f, Palmer & Van Ginkel 2105, 2694, GRA, NK/

SK/STh, dw

pubescens (Eckl. & Zeyh.) Druce var. pubescens. Palmer & Van

Ginkel 2040, 2447, GRA. W, dw

sarcophylla Fenzl. Palmer & Van Ginkel 2841, Rubin 26729, GRA,

KRNP, NK/SK/G, dw

Aizoon canariense L., Rubin 26922, KRNP. W, h

Tetragonia

echinata Aiton, Palmer & Van Ginkel 2454, Rubin 26716, GRA,

NK/STh, h

fruticosa L., Braack s.n., KRNP200, NK/SK/STh. dw

sarcophylla Fenzl var. sarcophylla. Shearing s.n., SCH, NK/SK/S, dw

MESEMBRYANTHEMACEAE

Arenifera spinescens (L.Bolus) H.E.K. Hartmann, GK, +, su

Aridaria sp.. Shearing s.n., SCH, ?, su, h

Chasmatophyllum musculinum (Haw.) Dinter & Schwantes, Shearing

s.n., SCH, W, su

Delosperma

floribundum L.Bolus, Rubin 38, KRNP. G, su, dw

pruinosum (Thunb.) J.W.Ingram, Cp, su, dw

pubipetalum L.Bolus, Rubin 3194, KRNP, GK, +, su, dw

subincanum (Haw.) Schwantes, Rubin s.n., KRNP, GK, +, su, dw

versicolor L.Bolus, Shearing s.n., SCH, KRNP, Cp, su, dw

Drosanthemum

archeri L.Bolus, Shearing s.n., SCH, SK/F, su

dejagerae L.Bolus, Bengis 397. KRNP, +, su

hispidum (L.) Schwantes var. platypetalum (Haw.) Schwantes, Bengis

332, Palmer & Van Ginkel 2449, GRA, NK/SK (GK), su

lique (N.E.Br.) Schwantes, Shearing s.n., SCH, GK, +, su

vespertinum L.Bolus var. vespertinum, Braack 57, KRNP, GK, +, su

Faucaria sp., Rubin 30, KRNP. NK/STh, su

Hereroa

concava L.Bolus, Bruyns 3375, BOL, KRNP, +, su

stanleyi (L.Bolus) L.Bolus, Shearing s.n., SCH, NK, +, su

Malephora

crocea (Jacq.) Schwantes var. crocea, Rubin s.n., KRNP, NK

(UK/BK), +, su

lutea (Haw.) Schwantes, Palmer & Van Ginkel 2482, GRA, GK, +, su

Mesembryanthemum

guerichianum Pax, Bengis 507, Shearing s.n., SCH, F/SK/STh, su

sp.. Shearing s.n., SCH, ?, su

Nananthus sp., Rubin 26943, KRNP, ?, su

Pleiospilos

compactus (Aiton) Schwantes

subsp. canus (Haw.) H.E.K. Hartmann & Liede, Shearing s.n., SCH,

GK, +, su

subsp. compactus, Bruyns 3428, BOL, GK, +, su

Prenia tetragona (Thunb.) Gerbaulet. Bengis 508, KRNP. BK, +, su

Psilocaulon

absimile N.E.Br., Palmer & Van Ginkel 2481, 2791, GRA, SK, su

articulatum (Thunb.) Schwantes, Bengis 429, Randall 27400, KRNP,

NK/F (UK/GK), su

utile L.Bolus, Rubin s.n., KRNP, SK/S, su

Rhinephyllum obliquum L.Bolus, Bruyns 3370, BOL, BK, +, su

Ruschia

dejagerae L.Bolus, Bruyns 3348, BOL, KRNP, +, su

hamata (L.Bolus) Schwantes, Bengis 362, Rubin 26771, KRNP,

NK/G/S (BK), su

intricata H.E.K. Hartmann & Stober, Braack s.n., KRNP14, GK, +, su

karrooica (L.Bolus) L.Bolus, Bruyns 3403, BOL, UK, +, su

parvifolia L. Bob, Shearing s.n., SCH, SK/F, su

pauciflora L.Bolus, Shearing s.n., SCH, KRNP. +, su

spinosa (L.) Dehn, Bengis 452, Palmer & Van Ginkel 2683, GRA, NK,

+, su

Sceletium

anatomicum (Haw.) L.Bolus, Bengis 493, KRNP, UK, +, su

dejagerae L.Bolus, Shearing s.n., SCH, KRNP, +, su

emarcidum (Thunb.) L.Bolus ex H. Jacobsen. Shearing s.n., SCH, SK,

su

expansum (L.) L.Bolus, Palmer & Van Ginkel 2504, GRA, GK, +, su

Stomatium sp.. Bruyns 3349. BOL, NK, +, su

Trichodiadema

attonsum (L.Bolus) Schwantes, Bengis 352, SCH, NK/G, su

barbatum (L.) Schwantes, Rubin s.n., KRNP, NK, +, su

densum (Haw.) Schwantes, Bengis 473, KRNP, GK, +, su

setuliferum (N.E.Br.) Schwantes var. setuliferum, Rishworth 12,

KRNP27638, SCH, GK, +, su

PORTULACACEAE

Talinum

caffrum (Thunb.) Eckl. & Zeyh., Shearing s.n., SCH, NK/G/S, h, su

sp., Bruyns 3415, BOL, ?, h, su

Anacampseros

albidiflora (Haw.) Sims, Rishworth 38, KRNP27638, Rubin 84,

NK/SK. h, su

lanceolata (Haw.) Sweet subsp. lanceolata, Randall 27408, KRNP, W.

h, su

marlothii Poelln., Bruyns 3432, BOL, Cp, h, su

subnuda Poelln. subsp. subnuda. W. h, su

telephiastrum DC., Bengis 453, Palmer & Van Ginkel 2836, GRA, W,

h, su

ustulata E. Meyer. Bruyns 3380, BOL, SCH, NK/S, h, su

Portulaca oleracea L., Palmer & Van Ginkel 2793, GRA, W, h

CARYOPHYLLACEAE (Part A)

Cerastium capense Sond., Bengis 385, KRNP, W, h

ILLECEBRACEAE

Pollichia campestris Aiton, Shearing 866, SCH. W, h

CARYOPHYLLACEAE (Part B)

Silene

burchellii Otth var. burchellii. Shearing s.n., SCH, W, h

undulata Aiton, Rubin 3235, KRNP, W, h

126

Bothalia 31,1 (2001 )

Dianthus

micropetalus Ser., Rubin 26774 , KRNP, W, h

namaensis Schinz var. namaensis, Rubin 14, KRNP, Cp, h

thunbergii Hooper forma thunbergii. Palmer & Van Ginkel 1847, GRA,

NK/STh (GK), h

RANUNCULACEAE

Clematis brachiata Thunb., Braack 97, KRNP, W, cl

Ranunculus multifidus Forssk., Bengis 448, SCH, W, h

MENISPERMACEAE

Cissampelos capensis L.f., Palmer & Van Ginkel 2499, 2730, GRA,

NK/F, h, cl

PAPAVERACEAE

Argemone ochroleuca Sweet subsp. ochroleuca. Shearing s.n., SCH,

NK. +, h

Papaver aculeatum Thunb., Braack 848, SCH, W, h

FUMARIACEAE

Cysticapnos pruinosa ( Bemh .) Liden ( Phacocapnos pruinosus ), Rubin

3250, KRNP, NK/S, h

BRASSICACEAE

Heliophila

camosa (Thunb.) Steud., Bengis 351, Rubin 26778, SCH, KRNP, W, dw

latisiliqua E.Mey. ex Sond. var. latisiliqua. Palmer & Van Ginkel

2471, GRA, NK/F, h

suavissima Burch, ex DC., Palmer & Van Ginkel 2885, SCH, GRA,

W, dw

Lepidium africanum (Bunn.f.) DC. subsp. divaricatum ( Aiton ) Jonsell, W, h

Sisymbrium

burchellii DC. var. burchellii. Shearing s.n., SCH. NK/F, h

capense Thunb. x S. turczaninowii Sond., Shearing s.n., SCH, KRNP, G, h

orientale L., Bengis 426, KRNP, W, h

CAPPARACEAE

Cadaba aphylla (Thunb.) Wild, Braack 163, Rubin 26912, KRNP, W,

sh. [t]

CRASSULACEAE

Cotyledon

??cuneata Thunb., Bruyns photo. BOL, NK/SK/F. dw, su

orbiculata L. var. orbiculata, Bengis 503, SCH. W, sh, su

papillaris L.f., Bruyns 3412, BOL, W, dw, su

Tylecodon

reticulatus (L.f.) Toelken subsp. reticulatus, Bruyns 3426, BOL, NK/

SK (GK), dw, su

wallichii (Harv.) Toelken

subsp. ecklonianus (Harw) Toelken, Shearing s.n., SCH, NK/SK

(GK), dw, su

subsp. wallichii, Bruyns 3430, BOL, NK/ SK (GK), dw, su

Crassula

atropurpurea (Haw.) D.Dietr.

var. anomala (Schonland & Baker /.) Toelken, Bruyns 3409, BOL,

NK/SK/F, dw, su

var. watermeyeri ( Compton ) Toelken, Bruyns 3392, BOL, SK, dw, su

barbata Thunb. subsp. barbata, Bengis 394, KRNP, NK/F (GK), h, su

campestris (Eckl. & Zeyh.) Endl. ex Walp., Rubin 33, KRNP, W, h, su

capitella Thunb.

subsp. capitella, Rubin 80, KRNP, NK/G/STh, h, su

subsp. thyrsiflora (Thunb.) Toelken, Palmer & Van Ginkel 2123,

2700, BOL, GRA, W, h, su

corallina Tluinb. subsp. corallina, Bruyns 3338, BOL, NK/G/S, h, su

deltoidea Thunb., Bruyns 3389, BOL, UK/BK, +, h, su

dependens Bolus, Palmer & Van Ginkel 22896, 2918, BOL, GRA,

NK/G/S, dw, su

exilis Harv. subsp. cooperi (Regel) Toelken, Bruyns 3374, BOL,

NK/G (GK), su, h

expansa Dryancl. subsp. expansa var. expansa. Palmer & Van Ginkel

2498, 2588, SCH, BOL, GRA, W, su, h

hemisphaerica Thunb., Bruyns 3384, BOL, NK (GK/BK), +, h, su

montana Thunb. subsp. quadrangularis (Schonland) Toelken, Bruyns

3386, BOL, GK, +, h, su

muricata Thunb., Rubin 48, KRNP, NK/F (GK), dw, su

muscosa L.

var. muscosa, Bruyns 3378, BOL, W, h, su

var. parvula (Eckl & Zeyh.) Toelken, Rubin 3223, BOL, F/STh/G,

h, su

nemorosa (Eckl. & Zeyh.) Endl. ex Walp., Bruyns s.n., BOL, W, h, su

nudicaulis L. var. platyphylla (Harv.) Toelken, Bruyns 3387, BOL,

GK, +, dw, su

papillosa Schonland & Baker f, Bruyns 3388, BOL, NK/STh (GK/

UK), h, su

pyramidalis Thunb., Shearing s.n., NK (GK/UK), +, h, su

rupestris Thunb. subsp. rupestris. Shearing s.n., Bruyns s.n., BOL,

NK/F/S, dw, su

subaphylla (Eckl. & Zeyh.) Harv. var. subaphylla, Bruyns 3382, BOL.

NK/SK/STh, dw, su

tetragona L.

subsp. acutifolia (Lam.) Toelken, Shearing 873, NK/SK/STh, dw, su

subsp. connivens ( Schonland ) Toelken, Bruyns 3418, BOL, NK/

STh, dw, su

subsp. rudis ( Schonland & Baker /.) Toelken, Bruyns 3367, BOL,

NK/G, dw, su

subsp. tetragona, Bruyns 3391, BOL, NK/SK, dw, su

Adromischus

filicaulis (Eckl. & Zeyh.) C.A.Sm. subsp. ftlicaulis, Bruyns 3431,

BOL, SK, dw, su

humilis (Marloth) Poelln., Bruyns 4029, BOL, NK, +, dw, su

sp„ Rubin 82, KRNP, ?

sphenophyllus C.A.Sm., Bruyns 3429, BOL, NK/F/S, dw, su

triflorus (L.f.) A. Berger, Shearing s.n., BOL, GK, +, dw, su

MONTINIACEAE

Montinia caryophyllacea Thunb., Randall 27349, NK/SK/F, sh, [t]

ROSACEAE

Rubus ludwigii Eckl. & Zeyh. subsp. ludwigii, Randall 27313, Rubin

3353, SCH. KRNP. W. sh

Cliffortia arborea Marloth, Palmer & Van Ginkel 2740, GRA. BK, +, R, t

FABACEAE

Acacia karroo Hayne, Randall 27383, SCH, W, t, sh

Prosopis glandulosa Torr. var. glandulosa, Randal! 27387, SCH, W, t, sh

Lotononis

azureoides B.-E.van Wvk, Shearing s.n., SCH, NK. +, dw

caerulescens (E.Mey.) B.-E.van Wyk, Shearing s.n., SCH, G/STh, sh, [dw]

cf. carnosa (Eckl. & Zeyh.) Benth. subsp. camosa, Bengis 442, SCH,

G/STh, sh, dw '

decumbens (Thunb.) B-E. van Wyk subsp. decumbens, Sugden s.n., Cp

laxa Eckl. & Zeyh., Shearing s.n., SCH, S/G, h

parviflora (P.J.Bergius) D.Dietr., Bengis 474, KRNP, SK/F, h

pungens Eckl & Zeyh., Palmer <£ Van Ginkel 2699, 2881, GRA, W, h

sp„ Rubin 26721, KRNP, ?,

tenella (E.Mey.) Eckl. & Zeyh., Shearing s.n., SCH, NK/SK/F/S, h

Aspalathus acicularis E.Mey. subsp. acicularis, Rubin 26764, KRNP,

W, sh, dw

Dichilus gracilis Eckl & Zeyh., Shearing s.n.. SCH, NK/S. h. dw

Melolobium

candicans (E.Mey.) Eckl. & Zeyh., Palmer & Van Ginkel 1855, GRA,

W, sh, dw, h

microphyllum (L.f.) Eckl. & Zeyh., Palmer & Van Ginkel 2157, Rubin

26759, GRA, KRNP, W, sh, dw

Argyrolobium

collinum Eckl. & Zeyh., Shearing s.n., SCH, F/SK/S, dw

ftliforme Eckl. & Zeyh., KRNP, F, h

pauciflorum Eckl. & Zeyh. var. pauciflorum, Rubin 26754, SCH.

KRNP, W, h

pumilum Eckl. & Zeyh., Shearing s.n., SCH, NK/F (GK), dw

Medicago laciniata (L.) Mill, Shearing s.n., SCH, W, h, *

Melilotus indica (L.) All., Shearing s.n., SCH, W, h, *

Trifolium

africanum Ser. var. africanum. Shearing s.n., SCH, W, h

burchellianum Ser., Sugden s.n.. W, h

Indigastrum argyraeum (Eckl. & Zeyh.) Schrire, Shearing s.n., SCH, W, h

Indigofera

alternans DC. var. alternans, Rubin 26740, KRNP, W, h

denudata L.f, Sugden s.n., Cp

heterophylla Thunb., Palmer & Van Ginkel 1856, GRA, NK/F (GK),

dw

meyeriana Eckl, & Zeyh., Rubin 3214, KRNP, W, sh, dw

nigromontana Eckl. & Zeyh., Rishworth s.n., KRNP27666, SCH,

NK, +

pungens E.Mey., KRNP, W, dw, sh

sessiliflora DC., Palmer & Van Ginkel 2737, Rubin 3191, GRA,

KRNP, NK/STh (GK), h

Sutherlandia

frutescens (L.) R.Br, Palmer & Van Ginkel 2516, 2914, SCH, GRA,

NK/SK/F, sh, dw

humilis E. Phillips & R.A.Dyer, NK/S, dw, sh

Bothalia 31,1 (2001)

127

Sutherlandia (cont.)

microphylla Burch, ex DC., Shearing s.n., SCH, NK/SK/G, sh

Lessertia

cf. annularis Burch., Shearing s.n., SCH, NK/SK/F. h

inflata Harv., Bengis 364, SCH, UK, +, h

GERANIACEAE

Geranium

harveyi Briq., Palmer & Van Ginkel 2144, 3261, GRA, KRNP, G, h

incanum Burm.f var. incanum. Shearing s.n., SCH, W. h

Sarcocaulon

camdeboense Moffett, Rishworth s.n., KRNP27663, Rubin 26931,

NK. +, dw, su

crassicaule Rehm, Shearing s.n., SCH. NK/SK. dw, su

patersonii (DC.) G.Don, NK/STh, dw, su

salmoniflorum Moffett, Braack 40, SCH, NK/S, dw, su

vanderietiae L.Bolus, KRNR NK/STh (GK), dw. su

Erodium cicutarium (L.) L'He'r., Bengis 77, 371, SCH, W, h

Pelargonium

abrotanifolium (L.f.) Jacq., Braack 154, Rubin 26909, KRNP, W, sh, dw

altemans J.C.Wendl., Shearing s.n., SCH. NK/SK/F. dw, su

althaeoides ( L .) L'He'r., Rubin 3249, KRNP, W, dw

aridum R. A. Dyer, Bruyns 3347, BOL, S/G/STh, dw

articulatum ( Cav .) Willd., Bruyns 3331, BOL, NK/SK/F, g

denticulatum Jacq., Bengis 435, F, +, sh

exhibens Vorster, Shearing s.n., SCH. NK/S. R , dw

glutinosum (Jacq.) L'He'r., Shearing s.n., SCH, NK/F/S, sh

griseum R.Knuth, Bengis 355, Braack 191, Rubin 26761, KRNP, NK

(GK/UK), +, dw

grossularioides (L.) LHer., Bengis 372, Braack 186, SCH, W, h

hypoleucum Turcz., Bruyns 3324, BOL, F, h

karooicum Compton & PE. Barnes, Bruyns 3751, BOL, SCH,

NK/SK, dw, su

cf. leucophyllum Turcz., NK/S, h

minimum (Cav.) Willd., Bengis 478, Rubin 27, KRNP, W, h

multicaule Jacq. subsp. multicaule, Bruyns 3327, BOL, W, dw

nanum L Her., Bruyns 3325, BOL, W, h

odoratissimum (L.) L Her., Braack 152, Shearing 846, SCH, W, dw

ramosissimum (Cav.) Willd., Shearing s.n., KRNP, NK/SK (UK), dw

senecioides L’Her., Shearing s.n., SCH, F/SK/S, h

tetragonum (L.f.) LHer., Shearing s.n., SCH, NK/SK/F/S, su, sc

OXALIDACEAE

Oxalis

lanata L.f. var. lanata, Bengis 400, F, g

commutata Sond. var. commutata. Shearing s.n., SCH. SK/F. g

melanosticta Sond. var. melanosticta. Palmer & Van Ginkel 2704,

GRA, NK/SK/F, g

cf. obtusa Jacq., Braack 171. KRNP. NK/SK/F. g

pes-caprae L. var. pes-caprae. Shearing s.n., SCH, NK/SK/F, g

LINACEAE

Linum

aethiopicum Thunb., Palmer & Van Ginkel 2674, 2813, GRA,

NK/F/S (GK), h, [dw]

thunbergii Eckl. & Zeyh., Shearing 841, W, h

ZYGOPHYLLACEAE

Zygophyllum

chrysopteron Retief Rubin 103, NK, +, sh, dw

gilfillanii N.E.Br.. Palmer & Van Ginkel 2104, 2902, Rubin 26727,

GRA, KRNP. Cp/FS, sh, dw, [su]

incrustatum E.Mey. ex Sond., Palmer & Van Ginkel 2779, Rubin

26706, GRA. KRNP, NK, +, sh, [dw]

lichtensteinianum Cham. & Schltdl., Palmer & Van Ginkel 2490,

Rubin 26797, GRA, KRNP, NK/F, dw

microcarpum Licht. ex Cham. & Schltdl., Shearing s.n., SCH,

NK/SK/F, sh. dw

microphyllum L.f, Palmer & Van Ginkel 2861, GRA, SK, dw

retrofractum Thunb., Palmer & Van Ginkel 2087, 2784, Rubin 26785,

GRA, KRNP. NK/SK/F, dw

Augea capensis Thunb., Shearing s.n., SCH, NK/SK, dw, su

Tribulus terrestris L., Randall 27389, SCH, W. h

POLYGALACEAE

Polygala

asbestina Burch., KRNP27657, Rishworth 57, Rubin 26746, GK, +,

dw, [h]

bowkerae Harv., Shearing s.n., SCH, NK, +, K, dw

ephedroides Burch., Rubin 26738, 26756, SCH, KRNP, W, sh, [dw]

hottentotta C.Presl, W, h, [dw]

leptophylla Burch., Palmer & Van Ginkel 2688, 2848, GRA, W, dw, [h]

serpentaria Eckl. & Zeyh., Rubin 77, KRNP, G, h

virgata Thunb. var. virgata. Shearing s.n., SCH, W, sh, [dw]

sp„ Rubin 267017, KRNP, ?

Muraltia macrocarpa Eckl. & Zeyh., Palmer & Van Ginkel 3260, SCH,

GRA, GK, +, sh, [dw]

EUPHORBIACEAE

Clutia

marginata E.Mey. ex Sond., Rubin 3224, KRNP, NK/F, sh, dw

thunbergii Sond., Palmer & Van Ginkel 2888, GRA, NK/SK (GK), dw

Euphorbia

braunsii N.E.Br., Shearing s.n., SCH, NK (GK/UK), +, dw, su

burmannii E.Mey. ex Boiss., Shearing s.n., SCH, NK/SK/F/S, dw, su

clavarioides Boiss. var. clavarioides, Bruyns 3366, BOL, W, dw, su

decepta N.E.Br., Shearing s.n., SCH, NK/S, dw, su

decussata E.Mey. ex Boiss., Bruyns s.n., Braack 72, SK, dw, su

hypogaea Marloth, Dyer 4109. GRA, GK, +, dw, su

mauritanica L. var. mauritanica, Rubin s.n., Bruyns 3245, BOL, W,

sh, dw, su

rectirama N.E.Br., Braack s.n., KRNP131, BOL, NK/S, dw, su

stellispina Haw., Shearing s.n., SCH. BK, +, sh, dw, su

stolonifera Marloth, Bruyns 3355, BOL, GK, +, dw, su

wilmaniae Marloth, Bruyns 3976, BOL, NK/S (UK), dw, su

sp., Bengis 490, ?,

Chamaesyce inaequilatera (Sond.) Sojak, Palmer & Van Ginkel 2812,

GRA, W, h

ANACARDIACEAE

Rhus

burchellii Sond. ex Engl., Palmer & Van Ginkel 1853, 2725, GRA, W,

sh, [t]

dregeana Sond., Randall 27362, SCH, W, sh, [t]

erosa Thunb., W, sh, [t]

lancea L.f, Bengis 420, SCH, W, t, [sh]

longispina Eckl. & Zeyh., Shearing s.n., SCH, NK/F/S (GK), sh, [t]

lucida L. forma lucida. Palmer & Van Ginkel 2760, GRA, F/STh/G,

sh, [t]

pyroides Burch, var. pyroides, Shearing s.n., SCH, W, t, sh

refracta Eckl. & Zeyh., Shearing s.n., SCH, NK/STh (GK), sh, [t]

undulata Jacq., Bengis 327, KRNP, W, sh, [t]

CELASTRACEAE

Maytenus

heterophylla (Sond.) Marais, Palmer & Van Ginkel 1828, 2082,

GRA, KRNP, W, t, sh

undata (Thunb.) Blakelock, Palmer & Van Ginkel 2518, GRA, W, t, sh

SAPINDACEAE

Cardiospermum sp., Rubin 3178, KRNP, ?, cl

MELIANTHACEAE

Melianthus comosus Vahl, Palmer & Van Ginkel 2098, 2140, GRA, W,

sh, [t]

RHAMNACEAE

Rhamnus prinoides L’Her., Rubin 3259, KRNP, W, t, sh

TIL1ACEAE

Grewia robusta Burch.. Palmer & Van Ginkel 1841, 2759, GRA, W, sh

MALVACEAE

Abutilon sonneratianum (Cav.) Sweet, Palmer & Van Ginkel 2894,

SCH, GRA, W, sh, [h]

Anisodontea

capensis (L.) Bates, Rubin 19, SCH, NK (GK/UK), +, sh

malvastroides (Baker f ) Bates, Palmer & Van Ginkel 2134, SCH,

GRA. Cp. sh

triloba (Thunb.) Bates, Bengis 440, Rubin 26796, KRNP, NK/SK, sh

Malva parviflora L. var. parviflora. Shearing s.n., SCH, W, h, *

Hibiscus

aridus R. A. Dyer, Sugden s.n., Cp,

pusillus Thunb., Palmer & Van Ginkel 2721, GRA, W, h

Radyera urens (L.f.) Bullock, Bengis 498, SCH, W, sh, [h]

STERCULIACEAE

Hermannia

abrotanoides Schrad, Sugden s.n., Cp,

alnifolia L„ Braack 33, KRNP, NK/SK/F, dw

althaeifolia L., Braack 156, SCH, NK/SK/F, h

angularis Jacq., Palmer & Van Ginkel 2807, GRA, NK/F (GK), dw

burkei Burtt Davy, S, h, [cl]

cemua Thunb. subsp. jacobeifolia (Turcz.) De Winter, Shearing s.n.,

SCH, NK/S, h

128

Bothalia 31,1 (2001 )

Hermannia (cont.)

coccocarpa ( Eckl . & Zeyh.) Kuntze, Braack s.n., KRNP91, Rubin

3198, SCH, W, h"

comosa Burch, ex DC., Bengis 415 , W, h

cuneifolia Jacq. var. cuneifolia, Braack 23, Rubin 26711, KRNP,

SCH, W, dw

depressa N.E.Br., Palmer & Van Ginkel 2811. GRA, W, h

desertorum Eckl. & Zeyh., Palmer & Van Ginkel 1822, 2865, Rubin

26741, GRA, KRNP, NK/SK/F, dw

erodioides (Burch, ex DC.) Kuntze, Palmer & Van Ginkel 1843,

Rubin 267191, 26924, KRNP, NK/S. h

filifolia L.f. var. filifolia, Palmer & Van Ginkel 2708, 2709, Rubin

26702, 26745, SCH, GRA, KRNP, W, dw

filifolia L.f. var. grandicalyx I.Verd., Shearing, SCH, NK/F/S, dw

grandiflora Aiton, Shearing s.n., GRA, KRNP, S, sh, [dw]

holosericea Jacq., Shearing s.n., SCH, NK/F/S (GK), dw, [sh]

johanssenii N.E.Br., Shearing s.n., SCH, SK, dw

linearifolia Harv., Shearing s.n., SCH, W, dw, [sh]

pulchella j L.f, Braack 12, NK/STh, dw

spinosa E.Mey. ex Harv., Palmer & Van Ginkel 2460, Rubin 26733,

GRA. KRNP. NK, +, dw

vestita Thunb., Palmer & Van Ginkel 1866, 2820, SCH, GRA, NK

(GK/UK), +, dw

TAMARICACEAE

Tamarix usneoides E.Mey. ex Bunge, Shearing s.n., SCH, SK, sh, [t]

FLACOURTIACEAE

Kiggelaria africana L., Bengis 309, SCH, W, t, [sh]

CACTACEAE

Opuntia ficus-indica (L.) Mill., Shearing s.n., SCH, W, t, sh, su, *

THYMELAEACEAE

Gnidia

capitata L.f, Rubin 26925, KRNP. W, sh, [dw]

cuneata Meisn., Palmer & Van Ginkel 2862, 2837, GRA, W, sh, [dw]

deserticola Gilg. Braack 38, SCH, NK/F/S, sh, [dw]

meyeri Meisn., Bengis 457, SCH, SK/F , sh, dw

polycephala (C.A.Mey.) Gilg, Shearing s.n., SCH, NK/SK/F/G/S, dw, h

Passerina

montana Thoday, Rubin 27034, KRNP, W, sh, [dw]

vulgaris Thoday, Rubin s.n.. Shearing s.n., KRNP, W, sh, [dw]

MYRTACEAE

Myrtus communis L., Braack 164, W, sh, *

ONAGRACEAE

Epilobium capense Buchinger ex Hochst., Shearing s.n., SCH, W, h

Oenothera rosea L'Her. ex Aiton, Shearing s.n., SCH, W, h, *

ARALIACEAE

Cussonia panicuiata Eckl. & Zeyh. subsp. paniculata, Bengis 495, SCH, W, t

APIACEAE

Conium chaerophylloides (Thunb.) Sond., Bengis 401, W, h

Heteromorpha

arborescens (Spreng.) Cham. & Schltdl.

var. arborescens. Shearing s.n., SCH, F/SK/S, W, sh

var. abyssinica (A. Rich.) H.Wolff, Rubin 2693, 26933, KRNP, W, sh

Deverra denudata ( Viv .) Pfisterer & Podlech subsp. aphylla (Cham. &

Schltdl.) Pfisterer & Podlech, Braack s.n., KRNP128, W, dw,

[h]

Berula erecta (Huds.) Coville subsp. thunbergii (DC.) B.L.Burtt, Randall

27320, Shearing 910, SCH, Cp, h, hy

Annesorhiza filicaulis Eckl. & Zeyh., Shearing 857, SCH, Cp, h

Peucedanum sp., Shearing s.n., SCH, ?, h

PR1MULACEAE

Samolus sp.. Shearing s.n., Hobson 27385, W, h

EBENACEAE

Euclea

coriacea A. DC., Palmer & Van Ginkel 2872, GRA, NK/G (GK/UK),

sh, [t]

crispa (Thunb.) Giirke subsp. crispa, Shearing s.n., SCH, W, t, sh

undulata Thunb. var. undulata, Randall 27393, SCH, W, t, sh

Diospyros

austro-africana De Winter var. austro-africana, Palmer & Van Ginkel

2148, 2743, Rubin 26749, GRA, SCH, W, sh

lycioides Desf. subsp. lycioides, Bengis 315, SCH, W, sh

OLEACEAE

Olea europaea L. subsp. africana (Mill.) PS. Green, Shearing s.n., SCH,

W, t, sh

Menodora juncea Harv., Rubin 3195, 26901, KRNP, NK/SK (GK), sh, dw

LOGANIACEAE

Gomphostigma incomptum (L.f.) N.E.Br.. Palmer & Van Ginkel, GRA,

NK, +, sh

Buddleja

glomerata H.L.Wendi, Palmer & Van Ginkel 2891, Rubin 26913,

SCH, GRA, KRNP, NK (GK/UK), +, sh

salviifolia (L.) Lam., Palmer & Van Ginkel 2143, GRA, KRNP, W, t, sh

GENTIANACEAE

Sebaea pentandra E.Mey. var. pentandra, Braack s.n., KRNP90, SCH,

NK/SK/S/G, h

APOCYNACEAE

Carissa haematocarpa (Eckl.) A. DC., Bengis 337, SCH, KRNP,

NK/F/S, sh

Pachypodium succulentum (L.f.) Sweet, Bengis 308, SCH, NK/STh, sh, su

ASCLEPIADACEAE

Microloma armatum (Thunb.) Schltr. var. armatum, Palmer & Van

Ginkel 2757, Rubin s.n., GRA. KRNP, NK/SK, sh, dw

Xysmalobium gomphocarpoides (E.Mey.) D.Dietr. var. parvilobum

Bruyns, Bruyns 3364, BOL, NK (GK/UK), +, h

Schizoglossum

aschersonianum Schltr. var. aschersonianum, Bruyns 3362, BOL, Cp, h

bidens E.Mey. subsp. bidens, Bruyns 3329. BOL. W, h

Aspidoglossum sp., BOL, NK, +,

Gomphocarpus

filifonnis (E.Mey.) Dietr., Randall 27339, SCH, Cp, h

fruticosus (L.) W.T.Aiton, Randall 27321, SCH, W, h

Sarcostemma viminale (L. ) R.Br. Palmer & Van Ginkel 2747, BOL,

GRA, W, cl

Brachystelma circinatum E.Mey., Bruyns 3706, BOL, W, g

Hoodia

gordonii (Masson) Sweet ex Decne., Shearing s.n., SCH, SK, su

flava (N.E.Br) Plowes, Shearing s.n., SCH, NK/SK, su

Piaranthus comptus N.E.Br., Bruyns 3665, BOL, NK (GK/UK), +, su

Duvalia

angustiloba N.E.Br., Bruyns 3417, BOL, KRNP, +, su

caespitosa (Masson) Haw. var. caespitosa, Shearing s.n., SCH, NK/

SK, su

corderoyi (Hook.f.) N.E.Br., Bruyns 3376, BOL, NK/S (GK/UK), su

Stapelia

engleriana Schltr., Bruyns 3411, BOL, GK, +, su

grandiflora Masson, Shearing s.n., SCH, NK/S, su

olivacea N.E.Br., Bruyns 3652, BOL, NK (GK/UK), +, su

sp., Rishwortli 62, ?, su

Tridentea

gemmiflora (Mass.) Haw., Bruyns 4030, BOL, NK (GK/UK), +, su

jucunda (N.E.Br.) L.C. Leach, Palmer & Van Ginkel s.n., GRA, GK,

+, su

parvipuncta (N.E.Br) L.C. Leach subsp. parvipuncta, Bruyns 3990,

BOL, NK/SK, su

Huemia barbata (Masson) Haw., Bruyns 4221, BOL, NK/S, su

Fockea comaru (E.Mey.) N.E.Br., Bruyns s.n., BOL, NK/SK/F, g

CONVOLVULACEAE

Cuscuta campestris Yunck., Shearing s.n., SCH, S/G, h. p. *

Convolvulus

arvensis L., Shearing s.n., SCH, W, h, cl, *

dregeanus Choisy, Shearing 904, SCH, W, h

sagittatus Thunb., Shearing s.n., SCH, W, h

thunbergii Roem. & Schult., Palmer & Van Ginkel s.n., W, h

BORAGINACEAE

Ehretia rigida (Thunb.) Druce, Bengis 329, SCH, W, sh, [t]

Heliotropium ciliatum Kaplan, Shearing s.n., SCH, W, h

Trichodesma africanum (L.) Lehm., Bengis 463, NK/SK, h

Cynoglossum lanceolatum Forssk.. Bengis 387, W, h

Lappula capensis (DC.) Gorke, Shearing s.n., SCH, NK/SK/STh, h

Anchusa capensis Thunb., Bengis 365, SCH, NK/SK/F. h

Myosotis

arvensis (L.) Hill, Bengis 398, NK/F, h

graminifolia DC., Braack 187, UK, +, h

Lithospermum scabrum Thunb., Bengis 382, NK/SK/STh, h

Lobostemon argenteus (P.J.Bergius) H.Buek, Bengis 419, SCH,

NK/SK, sh

Bothalia 31,1 (2001)

129

VERBENACEAE

Lantana rugosa Thunb., Shearing s.n., SCH, W, sh

Chascanum

cuneifolium (L.f. ) E.Mey., Rubin 92, KRNR W, dw

garipense E.Mey., Palmer & Van Ginkel 2703, 2817, GRA, W, h

pinnatifidum (L.f.) E.Mey. var. pinnatifidum. Shearing 1378, SCH, ?, h

pumilum E.Mey., Shearing s.n., SCH, NK/SK/S. h

LAMLACEAE

Teucrium trifidum Retz., Shearing s.n., SCH. W. h

Leonotis

leonurus (L.) R.Br., Palmer & Van Ginkel 2770, GRA, W, sh

ocymifolia (Bunn.f.) Iwarsson var. ocymifolia. Shearing s.n., SCH,

W, sh

Lamium amplexicaule L., Shearing s.n., S, h, *

Ballota africana ( L .) Bentli., Bengis 469, SCH, W, dw

Stachys

cuneata Banks ex Benth., Shearing s.n., SCH, NK/SK. sh

grandifolia E.Mey. ex Benth., Palmer & Van Ginkel 2099, GRA,

NK/G. h ’

rugosa Alton, Palmer & Van Ginkel 2133, 2825, 2741, GRA. KRNP,

NK/SK/G, sh

Salvia

albicaulis Benth.. Rubin 26713. KRNR W, h

disermas L., Shearing 865, SCH. W, sh, h

stenophylla Burch, ex Benth., Braack 223, Palmer & Van Ginkel

2477, GRA, W, h

verbenaca L., Randall 27322, SCH. W. h

Becium burchellianum (Benth.) N.E.Br.. Rubin s.n.. KRNP. NK/STh

(UK), sh

Mentha aquatica L., Sugden s.n., W, h

SOLANACEAE

Nicandra physalodes (L.) Gaertn., Palmer & Van Ginkel 2167, GRA,

W. sh, dw, h, *

Lycium

cinereum Thunb. sensu lato, Palmer & Van Ginkel 1818, 1820, 2441,

GRA, W, sh, dw

hirsutum Dunal, Rubin 26938. KRNP. W, sh, dw

oxycarpum Dunal, Palmer & Van Ginkel 2799, 2845, Rubin 26291.

26937, GRA. KRNP, GK. +, t, sh, dw

prunus-spinosa Dunal, Palmer & Van Ginkel 2696, GRA, W, sh, dw

schizocalyx C.H. Wright, Bengis 334, SCH, W, sh, dw

of. villosum Schinz, Braack 27, NK, +, sh, dw

Withania somnifera ( L .) Dun., Shearing 851, SCH, W, sh, dw, h

Solanum

capense L., Palmer & Van Ginkel 1851. 2768, Randall 27370, SCH,

W, sh. dw

coccineum Jacq., Rubin s.n., KRNP, W, sh, dw

elaeagnifolium Cav., Shearing s.n., SCH, W, sh, dw, *

nigrum L., Braack 183, SCH, W, h, *

retroflexum Dunal, Shearing s.n., SCH, W. h

rigescens Jacq., Rubin 26715, KRNP, W, sh

supinum Dunal var. supinum. Shearing s.n., SCH, NK/SK/F, sh, dw, h

tomentosum L., Palmer & Van Ginkel 1867, 2728, 2101, GRA. W, sh,

[dw, h]

Datura stramonium L., Shearing s.n., SCH, W, h, [sh], *

Nicotiana glauca Graham, Randall 27304, SCH, W, t, sh, *

SCROPHULARIACEAE (Part A)

Aptosimum

indivisum Burch, ex Benth., Bengis 458, Palmer & Van Ginkel 2116,

GRA. W, dw

procumbens (Lehm.) Steud., Rubin 61, Palmer & Van Ginkel 2154,

SCH. GRA. W, h

spinescens (Thunb.) Weber, Palmer & Van Ginkel 2110, Rubin 26704,

GRA. KRNP. NK/SK. dw

sp.. Shearing s.n., SCH. ?, h

Peliostomum leucorrhizum E.Mey. ex Benth., Shearing s.n., SCH, NK/

SK, dw

Diascia

alonsooides Benth., Bengis 370, SCH. NK, +, h

capsularis Benth., Shearing s.n., SCH, W, h, [dw]

Nemesia

floribunda Lehm., Bengis 391, KRNP, W, h

fruticans (Thunb.) Benth., Rubin 3240, 3251, 3252, KRNP. W, h

versicolor E.Mey. ex Benth., Braack 30, SCH. SK/F, h

Manulea fragrans Schltr.. Braack 36, SCH, NK, +, h

Sutera

affinis (Bernh.) Kuntze, Rubin s.n., KRNP, F, h

argentea (L.f.) Hiem, Rubin 79, W, dw

campanulata (Benth.) Kuntze, Palmer & Van Ginkel 2697, GRA, W, sh

cephalotes Kuntze, Palmer & Van Ginkel 2852, GRA, F, h, sh

halimifolia (Benth.) Kuntze, Palmer & Van Ginkel 2161, 2693, Rubin

26942, GRA, W, h

macrosiphon (Schltr.) Hiern, Bengis 342, NK/G (UK), dw

marifolia (Benth.) Kuntze, Rubin s.n., KRNP, F, h

pinnatifida (Benth.) Kuntze, Palmer & Van Ginkel 1840, 2731, 2901,

GRA, W. h

rotundifolia (Benth.) Kuntze, Gibbs-Russell 4276, GRA, NK, +, dw

uncinata (Desr.) Hilliard, Shearing s.n., SCH, NK/SK/F, sh

Jamesbrittenia

albiflora (I.Verd.) Hilliard, Shearing s.n., SCH, W, h

atropurpurea (Benth.) Hilliard subsp. atropurpurea. Palmer & Van

Ginkel 2815. GRA, W. sh, dw

aurantiaca (Burch.) Hilliard, Shearing s.n., SCH, W, h

tysonii (Hiern) Hilliard, Shearing s.n., SCH, NK/S, h, dw

Zaluzianskya

crocea Schltr., Bengis 406, NK/G, h

ovata (Benth.) Walp., Bengis 405, W, h

violacea Schltr., Braack 31, NK/F , h

Limosella grandiflora Benth., Shearing s.n., SCH, NK/G/S, hy

SELAGINACEAE

Hebenstretia

parviflora E.Mey., Bengis 374, SCH. NK/SK/F, h

ramosissima Jaroscz, Braack 20, SCH, F, h

robusta E.Mey., Shearing s.n., SCH, NK/SK, h, sh

Selago

albida Choisy, Palmer & Van Ginkel 2886, Rubin 3221, 26911, SCH,

GRA. KRNP, NK/SK/STh, sh

corymbosa, Sugden s.n., Cp,

forbesii Rolfe, Sugden s.n., Cp,

punctata Rolfe, Palmer & Van Ginkel 2153, 2168, GRA, G, sh

speciosa Rolfe, Shearing s.n., SCH, G, sh

Walafrida

geniculata (L.f. ) Rolfe, Palmer & Van Ginkel 2114, 2684, 2829, GRA,

W, dw

gracilis Rolfe, Shearing s.n., SCH, F, sh, dw

micrantha ( Choisy ) Rolfe, Rubin 3236, KRNP, W, sh

nachtigalii (Rolfe) Rolfe, Braack 68, W, h

saxatilis (E.Mey.) Rolfe, Palmer & Van Ginkel 2130, 2503, 2742,

Rubin 3201, SCH, GRA, KRNP. W, h

Cromidon sp. nov. Compton, Bengis 341, ?,

SCROPHULARIACEAE (Part B)

Veronica anagallis-aquatica L.. Shearing s.n., SCH, W, h

Harveya

purpurea (L.f. ) Harv., Shearing s.n., SCH, F/S/G, p

bolusii Kuntze, Shearing s.n., SCH, F, p

randii Hiern, Rubin 26767, KRNP, W. p

BIGNONIACEAE

Rhigozum

obovatum Burch., Palmer & Van Ginkel 1837, GRA, W, sh

trichotomum Burch., Braack s.n., KRNP10, W, sh

PEDALIACEAE

Sesamum

capense Bunn.f. Shearing s.n., SCH, W, h

triphyllum Welw. exAsch. var. triphyllum, Rubin 26737, KRNP, W, h

LENTIBULAR1ACEAE

Utricularia bisquamata Schrank, Shearing 107, SCH, W, hy, h

ACANTHACEAE

Barleria

pungens L.f, Shearing s.n., SCH, F/STh, dw, h

stimulans E.Mey. ex Nees, Palmer & Van Ginkel 1835, GRA, NK/S,

dw, h

Blepharis

capensis ( L.f. ) Pers.

var. capensis, Rubin s.n., SCHv, NK/SK/F, sh, dw

var. latibracteata Obertn., Rubin 94, Cp, sh, dw

mitrata C.B. Clarke, Rubin 104. SCH. NK, +, sh, dw. h

villosa (Nees) C.B. Clarke, Bengis 509, NK, +, dw, h

Acanthopsis disperma Nees, Shearing s.n., SCH, NK/SK, h

Justicia orchioides L.f. subsp. orchioides, Shearing s.n., SCH, NK/F/S/G,

sh, dw

Monechma

divaricatum (Nees) C.B. Clarke, Palmer & Van Ginkel 1821, GRA,

W, sh, dw, h

130

Bothalia 31,1 (2001)

Monechma (cont.)

incanum (Nees) C.B. Clarke, Palmer & Van Ginkel 2103, 2783, Rubin

26919, GRA, KRNP, NK/S, sh, dw

spartioides (T.Anderson) C.B. Clarke, Palmer & Van Ginkel 1834,

2488, 2738, SCH, GRA, Cp, sh, dw

PLANTAGINACEAE

Plantago lanceolata L., Shearing s.n., SCH, W, h, *

RUBIACEAE

Anthospermum

aethiopicum L., Palmer & Van Ginkel 2912, GRA, KRNP, NK/F/S, sh

dregei Sond. subsp. dregei, Rubin s.n., SCH, KRNP, SK/F, dw

rigidum Eckl. & Zeyh. subsp. pumilum (Sond.) Puff, Bengis 475, W, dw

spathulatum Spreng. subsp. spathulatum, Rubin 26751, KRNP, W, sh, dw

Nenax microphylla (Sond.) Salter, Rubin 26726, KRNP, NK/G/STh, dw

Galium

capense Thunb. subsp. capense, Rubin 8237, 26762, SCH, KRNP,

NK/F/S (GK), h

tomentosum Tlumb., Rubin 26940, KRNP, W, h

DIPSACACEAE

Scabiosa columbaria L., Rubin 3258, 26729, 27008, KRNP, W, h

CUCURBITACEAE

Kedrostis

africana (L.) Cogn., Bruyns 3427, BOL, W, cl

crassirostrata Bremek., NK/S, h, cl

Citrullus lanatus (Thunb.) Matsum. & Nakai, Shearing s.n., SCH, W, h, cl

Cucumis

africanus L.f, Rubin 26735, KRNP, W, h, [cl]

myriocarpus Naudin subsp. myriocarpus. Shearing s.n., SCH, W, h, [cl]

zeyheri Sond., Shearing s.n., SCH, W, h, [cl]

CAMPANULACEAE

Wahlenbergia

cemua (Thunb.) A. DC., Bengis 421, F/STh/G, h

nodosa (H.Buek) hammers. Palmer & Van Ginkel 2821, 2900, Rubin

26753, 26944, GRA, KRNP, W, sh

LOBELIACEAE

Lobelia

erinus L., Sugden s.n., W, h

thermalis Thunb., Shearing s.n., SCH, W, h

ASTERACEAE

Pteronia

adenocarpa Harv., Palmer & Van Ginkel 1863, GRA, NK, +, sh

cinerea L.f, NK/F, sh, dw

glauca Thunb., Palmer & Van Ginkel 2514, 2849, Rubin 26787,

GRA, KRNP, NK/SK, sh, dw

glomerata L.f, Braack 221, Rubin 3185, SCH, KRNP, NK/F, sh, [dw]

inflexa Thunb. ex L.f, Rubin s.n., SK, sh

membranacea L.f, Bengis 433, Rubin 3184, SCH, KRNP. NK/F/S, sh

mucronata DC., Rubin s.n.. KRNP, NK, +, sh, dw

pallens L.f., Shearing s.n., SCH, NK/F, sh, dw

paniculata Thunb., Shearing s.n., SCH, NK/SK/F/S, sh, dw

punctata E. Phillips, Braack s.n., KRNP96, KRNP, NK. +, sh

scariosa L.f, Rubin 102, GK, +, sh

sordida N.E.Br., Palmer & Van Ginkel 2151, 2868, GRA, KRNP,

NK/SK, dw, [sh]

spinulosa E. Phillips, BK, +, dw, [sh]

staehelinoides DC.. Palmer & Van Ginkel 2515, 2806, GRA, NK/F/S,

sh, dw

tricephala DC., Rubin 51, GK, +, sh

viscosa Thunb., Bengis 423, Rubin 3187, 26718, SCH, NK, +, sh, dw

Ameilus tridactylus DC. subsp. tridactylus. Palmer & Van Ginkel s.n.,

GRA, NK/SK/F, h

Aster squamatus (Spreng.) Hieron., Palmer & Van Ginkel 2095, SCH,

W, h, *

Erigeron sp„ Randall 27412, SCH, KRNP, ?, h

Felicia

fascicularis DC.. Braack 24, Rubin 26783, GRA, KRNP, W, sh

filifolia (Vent.) Burn Davy subsp. filifolia. Palmer & Van Ginkel

1858, 2111, Rubin 26715, 26934, SCH, GRA, KRNP, W, sh

hirsuta DC., Palmer & Van Ginkel 1854, 2850, Rubin 26717, KRNP,

W, sh

lasiocarpa DC., Shearing s.n., GRA, NK, +, sh

muricata (Thunb.) Nees subsp. muricata, Palmer & Van Ginkel 2129,

KRNP, W, sh

namaquana (Harv.) Merxm., Braack 5, KRNP, NK/SK, h

ovata (Thunb.) Compton, Braack s.n., KRNP 122, Rubin 26782, GRA,

W, sh

zeyheri ( Less. ) Nees

subsp. linifolia (Harv.) Grau, Palmer & Van Ginkel 1850, 2673,

2789, GRA. W. sh

subsp. zeyheri. Palmer <£ Van Ginkel 2714, 2843, 2851, NK, +, sh

Conyza

scabrida DC., Shearing s.n., SCH, W, sh

sp„ Braack 185, GRA, KRNP, W, h

Chrysocoma oblongifolia DC, Bengis 345, Palmer & Van Ginkel 2112,

Rubin 26758, SCH, KRNP. W, sh, dw

Tarchonanthus camphoratus L, Bengis 447, Rubin 3176, KRNP, W, sh, [dw]

Trichogyne decumbens (Thunb.) Less., Shearing s.n., SCH, NK/SK/G, h

Ifloga glomerata (Harv. ) Schltr., Shearing s.n., SCH, NK/SK/F/G/S, sh, [h]

Vellereophyton niveum Hilliard, Shearing 909, SCH, NK/F (GK), sh, dw

Pseudognaphalium undulatum (L.) Hilliard & B.L.Burtt, Shearing s.n.,

SCH, KRNP, W, sh, dw

Helichrysum

dregeanum Sond. & Harv., Rubin 3182, 3183, GRA, W, sh, dw

hamulosum E.Mey. ex DC., Palmer & Van Ginkel 2754, SCH, W, sh,

[dw]

lineare DC., Rubin s.n., SCH, GRA, KRNP, W, h

lucilioides Less., Palmer & Van Ginkel 2818, SCH, NK/SK (GK/

UK), sh, [h]

pumilio (O.Hoffm.) Hilliard & B.L.Burtt

subsp. fleckii (S. Moore) Hilliard. Bengis 346, SCH, NK/SK/G, sh, dw

subsp. pumilio. Shearing s.n., SCH, NK/SK, sh, dw

revolutum (Thunb.) Less., Rubin s.n., SCH, KRNP. SK/F, sh, dw

rosum (P.J.Bergius) Less. var. rosum. Shearing s.n., SCH, NK/F/S,

sh, [dw]

rugulosum Less., Shearing 843, SCH, W, h

scitulum Hilliard & B.L.Burtt, Shearing s.n.. SCH, NK, +, sh, [h]

trilineatum DC., Bengis 377, GRA, G, sh, dw

zeyheri Less., Palmer & Van Ginkel 2156, 2711, 2887, SCH, W, sh, dw

Elytropappus rhinocerotis (L.f.) Less., Palmer & Van Ginkel 2137,

2745, Rubin 26936, KRNP, W, sh, dw

Pterothrix spinescens DC., Randall 27395, W, sh

Relhania sp., Bengis 412, GRA, KRNP, ?, h

Rosenia

humilis (Less.) K. Bremer, Palmer & Van Ginkel 1857, 2508, Rubin

3197, 26742, GRA, W, sh

oppositifolia (DC.) K. Bremer, Palmer & Van Ginkel 2486, SCH, UK,

+, sh

spinescens DC., GRA, GK. +, sh

Ley sera

gnaphalodes (L.) L., Rubin 49, GRA, NK/SK/F, dw

tenella DC., Palmer & Van Ginkel 1832, SCH, GRA, KRNP, NK/

SK/F, h

Pegolettia retrofracta (Thunb.) Kies, Palmer & Van Ginkel 2489, 2512,

Rubin 26903, SCH, W, dw

Geigeria filifolia Mattf., Bengis 480, SCH, W, h

Xanthium spinosum L., Shearing s.n., GRA, W, h, *

Schkuhria pinnata (Lam.) Cabrera, Palmer & Van Ginkel 1731, W. h. *

Tagetes minuta L., Shearing s.n., KRNP, W, h, *

Eriocephalus

cf. aspalathoides i, GRA, KRNP, W. sh

decussatus Burch, (ex Shearing), Shearing s.n., SCH, NK/SK, sh

ericoides (L.f.) Druce, Palmer & Van Ginkel 1824, 1825, 1848, SCH,

GRA, KRNP, W, sh

pubescens DC., Shearing s.n., SCH, W, sh

punctulatus DC., Palmer & Van Ginkel 2106, Rubin 3241, 26784,

KRNP, F, sh

spinescens Burch., Rubin 26712, KRNP, NK/SK/S, sh, [dw]

sp.. Shearing s.n., SCH, ?, sh

tenuifolius DC., Rubin s.n., GK, +, sh

Athanasia

cf. linifolia Burm., GRA, Cp, sh

microcephala (DC.) D.Dietr., Shearing s.n., SCH, NK/SK/F, sh

minuta (L.f.) Kallersjo subsp. minuta. Shearing s.n., SCH. NK/SK,

dw, [sh]

Phymaspermum

aciculare (E.Mey. ex Harv.) Benth. & Hook, ex Jacks., Shearing s.n.,

SCH, NK, +, sh

cf. leptophyllum (DC.) Benth. & Hook, ex Jacks., Palmer & Van

Ginkel 2729, GK, +, sh

parvifolium (DC.) Benth. & Hook, ex Jacks., Shearing s.n., SCH,

NK/STh, dw, [sh]

schroteri Compton, Shearing s.n., SCH, SK, +, sh

sp., Randall 27331, SCH. ?, sh

Bothalia 31.1 (2001)

131

Cotula

coronopifolia L., Shearing s.n., SCH, W, h. [hy]

heterocarpa DC., Sugden s.n., Cp, h

sororia DC.. NK/STh (UK), h

microglossa {DC.) O.Hoffm. & Kuntze ex Kuntze, Bengis 413, W, h

Artemisia afra Jacq. ex Willd., Randall 27324, SCH, GRA, KRNR W,

h, [sh]

Pentzia

globosa Less., Pabner & Van Ginkel 2822, 2823, 2858, GRA. KRNP,

W, sh

incana (Thunb.) Kuntze. Palmer & Van Ginkel 1838, 2826. Rubin

26708, SCH, GRA. W, sh

punctata Harv.. Palmer & Van Ginkel 1842, 2150, 2883, SCH. NK/F. sh

quinquefida {Thunb.) Less., Bengis 511. Rubin 26998, KRNP. NK/G

(UK/BK), sh

sphaerocephala.DC., Braack s.n., KRNP 129, Rubin 27031, SCH. W, sh

spinescens Less.. Bengis 512, SCH, UK. +, sh

Oncosiphon piluliferum (L.f.) Kallersjd, Shearing s.n., SCH, NK/SK/

F/S, h

Cineraria

aspera Thunb.. Bengis 399. KRNP. W, h

lobata L'He'r., Shearing s.n., SCH. S/F, h, [dw]

mollis E.Mey. ex DC., Rubin 3230, 3247, KRNP. W, h

Senecio

abbreviates S. Moore, BOL, NK, +, cl

achilleifolius DC., Shearing s.n., SCH, G, h, [sh]

acutifolius DC., Rubin 26918, KRNP. NK/G (GK/UK), sh, dw

articulates (L.) Sch.Bip., BOL, NK/STh, h, su

asperates DC., Palmer & Van Ginkel 2909, SCH. GRA, W. h

burchellii DC.. Bengis 437. W. sh, dw, h

carroensis DC., Rubin s.n.. GK, +, h

cotyledonis DC., Palmer & Van Ginkel 2511, GRA, NK/SK. sh, dw, su

erysimoides DC., Shearing s.n., SCH. SK/G. h

hallianus G.D. Rowley, BOL. Cp, h, su

harveianus MacOwan, Shearing s.n., SCH. G, h, [dw]

inaequidens DC.. Palmer & Van Ginkel 2128, 2753, GRA. W. h. [sh]

leptophyllus DC., Palmer & Van Ginkel 2472, GRA. NK/F/S, sh, dw, h

linifolius L., Bruyns 3975, BOL, F/STh. sh

madagascariensis Poir., Shearing s.n., SCH. W, h

pearsonii Hutch., BOL, Cp, h

radicans {L.f.) Sch.Bip., Palmer & Van Ginkel 2112, GRA, W, h, su

striatifolius DC., Braack s.n.. KRNP136, W. h

vimineus DC., Shearing s.n., SCH. G/S. h

Kleinia longiflora DC., Palmer & Van Ginkel 1833, GRA. STh, sh, su.

Euryops

annae E. Phillips, Palmer & Van Ginkel 2741, 2897. GRA. KRNP, W. sh

anthemoides B.Nord. subsp. anthemoides, Pabner & Van Ginkel

2844, GRA. NK/STh (UK), dw

cuneatus B.Nord., Braack 19, GK, +, sh

empetrifolius DC., Braack 18, W, sh

imbricates {Thunb.) DC., Bengis 470. Rubin 26781, 26939, SCH.

KRNP. GK. +, sh

laterifloras {L.f.) DC., Pabner & Van Ginkel 2904, GRA. W, sh

nodosus B.Nord., Rubin 50. KRNP. UK/G, sh, dw

oligoglossus DC.

subsp. oligoglossus, Pabner & Van Ginkel 2136, 2749, SCH. GRA.

KRNP. NK. +, sh

subsp. racemosus (DC.) B.Nord., Rubin 70, NK, +, sh

subcamosus DC.

subsp. subcamosus DC., Braack 35, SCH, NK/SK, sh

subsp. vulgaris B.Nord., Shearing s.n., SCH. NK/ SK/G. sh

sulcatus (Thunb.) Harv., Rubin 69. KRNP. NK, +, sh

trifidus {L.f.) DC., Bengis 389. NK, +, sh

Othonna

auriculifolia Licht. ex Less., Pabner & Van Ginkel 2162, 2910. GRA.

NK, +, h

camosa Less. var. camosa, Bengis 505, W, sh, su.

furcata ( Lindl .) Druce, Gibbs-Russell 335, GRA, NK, +, sh, su.

pavonia E.Mey., Shearing s.n., SCH. NK (GK/UK), +, sh

protecta Dinter, Bruvns 3958, BOL, NK/SK (GK), dw, h, su

sedifolia DC., Palmer & Van Ginkel 2500. BOL, GRA. NK/SK (BK),

dw, su

Oligocarpus calendulaceus (L.f.) Less.. Braack 3, SCH, KRNP, W, h

Dimorphotheca

cuneata (Thunb.) Less., Bengis 358, SCH, NK/SK/STh, sh

sp., Braack 172, ?,

Garaleum bipinnatum (Thunb.) Less., Palmer & Van Ginkel 1826, 2800,

GRA, NK/SK, sh, [h]

Osteospermum

leptolobum (Harv) Norl., Rubin 26714 , KRNP. NK (GK/UK), +, sh

microphyllum DC., Pabner & Van Ginkel 2450, 3168, GRA, NK, +, sh

rigidum Aiton var. rigidum, Rubin s.n., KRNP, NK/SK, sh, [dw]

spinescens Thunb., Shearing s.n., SCH. NK/G/S, sh

Tripteris

aghillana DC. var. aghillana, Pabner & Van Ginkel 2517, Rubin 26747,

GRA. KRNP. NK/SK/F/G, h

sinuata DC. var. sinuata, Pabner & Van Ginkel 2827, 2863, Rubin

26703. GRA. KRNP, NK/SK, sh

Ursinia nana DC. subsp. nana, Bengis 476, SCH, W, h

Arctotis

adpressa DC., Shearing s.n., SCH, NK/SK, h

arctotoides (L.f.) O.Hoffm., Pabner & Van Ginkel 2874, GRA, W, h

erosa (Harv.) PBeauv., Bengis 380, SCH, SK , h

leiocarpa Harv., Braack 4, SCH, NK (GK/UK), +, h

microcephala (DC.) PBeauv., Shearing s.n., SCH, W, h

sulcocarpa Lewin, Shearing s.n., SCH, NK/SK/F, h

Arctotheca calendula (L.) Levyns, Shearing s.n., SCH. SK/F/S/G, h

Haplocarpha sp.. Braack s.n., KRNP201, ?, h

Gazania

heterochaeta DC., Rubin s.n., NK/SK, h

krebsiana Less.

subsp. arctotoides (Less.) Roessler, Palmer & Van Ginkel 2685,

GRA, W, h

subsp. krebsiana, Rubin 42, KRNP, W. h

lichtensteinii Less., Randall 27388, SCH. NK/SK/F. h

linearis (Thunb.) Druce var. linearis, Palmer & Van Ginkel 2473, GRA,

KRNP. w. h

Hirpicium alienatum (Thunb.) Druce. Rubin 26905, SCH. KRNP. NK/

SK, dw

Berkheya

carlinifolia (DC.) Roessler subsp. carlinifolia, Rubin 3246, KRNP,

NK/SK, sh

decurrens (Thunb.) Willd., Palmer & Van Ginkel 2094, GRA, F/STh. h

glabrata (Thunb.) Fourc.. Bengis 449, KRNP, W, h

heterophylla (Thunb.) O.Hoffm.

var. heterophylla. K/STh, h

var. radiata (DC.) Roessler, Shearing s.n., SCH, F/S, h

pinnatifida (Thunb.) Thell. subsp. pinnatifida, Rubin 6, SCH. G/STh, h

spinosa (L.f.) Druce, Bengis 403, GK. +, sh

spinosissima (Thunb.) Willd. subsp. spinosissima, SCH, NK. +, sh

Cuspidia cemua (L.f.) B.L.Burtt subsp. cemua, Bengis 481, Pabner &

Van Ginkel 2828. Rubin 26728. GRA, KRNP, NK/STh (GK), h

Cirsium vulgare (Savi) Ten., Randall 27319, W, h, *

Dicoma

capensis Less., Randall 27410. KRNP, SCH, NK/SK, h

spinosa (L.) Druce, Rubin 26709, 26792. KRNP. NK, +, dw

Cichorium intybus L., Shearing s.n., SCH, NK/G , h, *

Tragopogon porrifolius L., SCH, W, h, *

Sonchus wilmsii R.E.Fr., Rubin s.n., G/S, h

Lactuca capensis Thunb., Bengis 379, W, h

Abutilon, 127

Acacia, 126

Acanthaceae, 129

Acanthopsis, 129

Adiantaceae, 122

Adiantum. 122

Adromischus, 126

INDEX OF FAMILIES AND GENERA

Agrostis, 123

Aizoaceae, 125

Aizoon, 125

Albuca, 124

Alliaceae, 124

Aloe, 124

Amaranthaceae, 125

Amaranthus, 125

Amaryllidaceae, 124

Amellus, 130

Ammocharis, 124

Anacampseros, 125

Anacardiaceae, 127

Anchusa, 128

Androcymbium, 123

Andropogon, 122

Angiospermae, 122

Anisodontea, 127

Annesorhiza, 128

Anochilus, 124

Anthospermum, 130

132

Bothalia 31,1 (2001)

Bothalia 31,1: 135-143 (2001)

Vegetation-environment relationships in a catchment containing a

dambo in central Zimbabwe

I. MAPAURE* and M.P. MCCARTNEY**

Keywords: Canonical Correspondence Analysis, dambo, plant-water relations, vegetation classification, Zimbabwe

ABSTRACT

Seasonally saturated wetlands, known as dambos, are a common landscape element throughout much of southern

Africa. The diversity of species composition within catchments containing dambos is widely attributed to hydrological con-

ditions, but plant-water relationships are poorly established. In this paper a detailed classification and a vegetation map are

presented for a small catchment in central Zimbabwe containing a dambo. Canonical Correspondence Analysis has been

applied to explore the link between vegetation composition and environmental variables. This confirms that water is a key

influence in species distribution and small-scale patterning of vegetation within the catchment.

INTRODUCTION

Many factors influence the composition of vegetation,

including the supply of nutrients, micro-climate, soil

moisture, intraspecific competition, grazing and manage-

ment practices (Gimingham 1972). In some environ-

ments, the relationship between water and vegetation has

been shown to be particularly important. For example,

Gumell (1981) and Gumell & Gregory (1987) illustrated

that the vegetation composition of heathland areas in tem-

perate climates is particularly ‘fine-tuned’ to the underly-

ing soil moisture regime. In a vegetation survey of Zim-

babwe, Rattray (1957) concluded that soil moisture

‘appeared to be the most important single factor affecting

the distribution of grasses.’ Dye & Walker (1980) illus-

trated how the influence of soil properties on the infiltra-

tion of water had a marked effect on floristic distribution

in areas of Zimbabwe where sodic soils are prevalent.

Dambos (sometimes termed vleis) are seasonally water-

logged, predominantly grass covered, shallow de-

pressions bordering headwater drainage lines. They are a

widespread land feature in southern Africa. In Zimbabwe

they occupy about 4% of the land surface (Whitlow

1984). Field observations indicate that soil moisture is

one of the major variables influencing floristic variation

within catchments containing dambos. Perera (1982) sur-

mised that the most important controls on the distribution

of plant communities within dambos are micro-relief, the

water table and the physical and chemical properties of

the soil. Whitlow (1985) states that within dambos ‘veg-

etation generally comprises a mosaic of plant communi-

ties which changes in character from the margins to the

central portions of the dambo dependent upon the degree

and duration of waterlogging.’

The current study is believed to be one of the first to

statistically validate the relationship between vegetation

* Tropical Resource Ecology Programme, Department of Biological

Sciences, University of Zimbabwe, P.O. Box MP 167, Mount Pleasant,

Harare, Zimbabwe.

** Institute of Hydrology, Crowmarsh Gifford, Wallingford,

Oxfordshire 0X10 8BB, UK.

MS. received: 1999-05-31.

distribution and environmental factors for a dambo.

There were two elements to the study. Firstly, a detailed

vegetation classification and a vegetation map, based on

floristic dominance and physiognomy, was derived for a

small catchment in the highveld of central Zimbabwe.

Secondly, the link between species composition and

environmental variables was investigated independently

of the mapping exercise using Canonical Correspond-

ence Analysis (Ter Braak 1988a). The results indicate

that vegetation associations may provide a basis for iden-

tifying characteristic soil moisture regimes in highveld

catchments containing dambos.

STUDY AREA

The study area (Figure 1), is located in the headwaters

of the Manyame River at the Grasslands Research

Station, near Marondera, 70 km southeast of Harare.

Relief over the catchment is low (gradients less than 4%)

and the elevation varies from 1 611 to 1 654 m above sea

level. In Zimbabwe, land at elevations greater than 1 200

m is known as highveld. The catchment area of 3.33 km2

comprises two parts: the upland region, known as the

interfluve (2.12 km2) and a single dambo (1.21 km2).

The dambo extends the whole length of the catchment

and incorporates a narrow spur on the northern slope

(Figure 1). This spur crosses the catchment boundary and

joins another dambo in a neighbouring catchment. The

main road and railway line from Harare to Mutare pass

through the eastern end.

Rainfall is seasonal and largely dependent on the

movement of the Intertropical Convergence Zone over

southern Africa. Average annual rainfall is 859 mm, but

varies considerably from year to year. Rainfall occurs pre-

dominantly during the wet summer (October to April),

and the winter months (May to September) are usually

dry. During the winter, the mean temperature is about

12.5°C, although it is not uncommon for night tempera-

tures to drop below freezing. During the summer, the

mean temperature is 19.5°C. Average potential evapora-

tion is about 1 700 mm.

136

Bothalia 31,1 (2001)

FIGURE 1. — Grasslands research catchment area showing the dambo, which is at the head waters of Manyame River, near Marondera, 70 km southeast of Harare.

Bothalia 31,1 (2001)

137

25

20

_E 15-

c

o

>

Q

W 10-

5

1 I I I I I I ! I I I I I I | I

0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500

r Relative to top of the weir at the catchment outlet Distance from Plot 1 (m)

FIGURE 2. — Cross section showing soil profile, location of vegetation sampling plots and dipwells along transect across the grasslands research

catchment, near Marondera.

Soils within the catchment have been derived pre-

dominantly from the underlying granite. However, in

places, soil mineralogy and texture have been influenced

by the presence of dolerite dykes. A distinct catena exists

on the slopes of the catchment (Figure 2). The interfluve

soils are acidic, strongly leached and have reached an

advanced stage of weathering. They are moderately

deep, coarse to medium-grained loamy sands overlying

sandy clay loams. The dambo soils are hydromorphic

with high organic matter content in the topsoils and gley-

ing of the subsoils (Whitlow 1985). At shallow depth

within the dambo, there is a well-defined clay wedge of

irregular shape that is embedded within a lens of sandy

clay. It is the presence of these heavy textured subsoils

that impedes vertical drainage and results in the wet sea-

son saturation of the dambo.

METHODS

Two approaches were employed, one for vegetation

mapping and the other for detailed environmental corre-

lation. For vegetation mapping, inventory and classifica-

tion, the method employed by Timberlake et al. (1993)

was followed. Black-and-white aerial photographs (scale

1 : 25 000. 1983) were interpreted using a mirror stereo-

scope, and the vegetation was stratified on the basis of the

textural differences on the photographs. To see if there

were any significant changes since 1983, aerial colour

photographs were taken in the wet season of 1995.

Sampling was done in both the dry and wet seasons. A

total of 22 samples were randomly located in the stratified

areas (stratified random sampling). At each sampling

point all species were identified and their cover-abun-

dance values assessed according to the Braun-Blanquet

scale (Mueller-Dombois & Ellenberg 1974). Since the

number of samples was few and the size of the catchment

area is small, it was possible to produce a floristic classi-

fication directly from the field sheets, in conjunction with

both the black-and-white and the colour aerial pho-

tographs, without computer-aided classification.

For environmental correlations, detailed inventories

were made of species within a radius of 5 m around each

of 16 markers laid out along a transect across the middle

of the catchment. The markers ran from the interfluve on

the southern slope to the interfluve on the northern slope

of the catchment crossing the valley bottom immediately

to the east of the stream channel (Figure 1). Species pres-

ence/absence data were obtained, and later used to relate

species composition directly to environmental variables

measured within each plot.

Environmental factors were determined at each of the

16 plots. Soil characteristics determined for the surface

soil horizon were: bulk density (BDEN), pH (H20), elec-

trical conductivity (EC) and percentage carbon (%C).

Dry bulk density was determined from the weight of

undisturbed core samples oven dried at 105°C for 24

hours. Soil pH and EC were measured electrometrically

in a 1:5 suspension in distilled water. The percentage of

carbon in the soil was determined by the colorimetric

method using potassium dichromate (Chapman 1987).

Within each plot, depth to the water table was mea-

sured daily from November 1995 to October 1996 in nar-

row diameter (32 mm) dipwells and the mean depth was

calculated for each month. At each location, the mean

value for February 1996 (February water table depth —

FWTD) was taken as an index of the wet season water

level, and the mean level for September 1996 (Sep-

tember water table depth — SWTD) was taken as an

index of the end of dry season water level. These were

used as surrogate indicators of the soil moisture regime

within each plot.

Species and environmental data from the 16 plots were

subjected to Canonical Correspondence Analysis (CCA)

(Ter Braak 1988b) to elucidate the relationships between

vegetation composition and the six measured environmen-

tal variables. CCA is a direct ordination technique com-

monly used in community ecology studies. It integrates

regression and ordination techniques into a method of mul-

tivariate direct gradient analysis that is used to detect uni-

modal relationships between species and environmental

variables (Ter Braak 1986, 1987, 1988a). The relationships

can be shown in an ordination diagram by vectors for the

environmental variables with lengths proportional to their

138

Bothalia 31,1 (2001)

TABLE 1. — Summary of land cover in the Grasslands Research Catchment

Bothalia 31,1 (2001)

139

importance and directions showing their correlation with

each axis. The statistical validity of the ordination was test-

ed using an unrestricted Monte-Carlo permutation test

available in CANOCO (Ter Braak 1988b).

RESULTS AND INTERPRETATION

Vegetation classification

The results of the vegetation classification are shown in

Table 1 and mapped in Figure 3. The vegetation was classi-

fied into 10 types based on floristic dominance and phys-

iognomy. Physiognomically, the types range from wood-

land through wooded grassland to grassland and plantations

of exotic timber species. Physiognomic classes follow Pratt

et al. (1966). A woodland is defined as a stand of trees with

canopy cover of 20-80%, a grassland is land dominated by

grasses, sometimes with widely scattered herbs with canopy

cover not exceeding 2%, and a wooded grassland as grass-

land with scattered or grouped trees with canopy cover less

than 20% (Pratt et al. 1966). Woody species and grass

nomenclature follow Drummond (1975) and Bennett

(1980), respectively, with a few species name updates in

both cases. Herb nomenclature follows various Flora

zambesiaca volumes dealing with specific plant families.

Grassland is the most extensive of all the vegetation

types in the catchment, covering 48% of the area. Six dis-

tinct types can be recognized, based on differing floristic

dominance (Table 1). The most extensive is Hyparrhenia

filipendula-dommdXed grassland. This is a medium-height

grassland of up to 1.2 m high which extends from the

channel grassland to the woodland and wooded grassland

on the upper slopes. Very localized dominance of other

grass species (i.e. Setaria incrassata , Sporobolus pyrami-

dalis and Loudetia simplex) is also evident.

In the dambo itself, along the stream channel, there is

a narrow belt, up to 10 m wide, characterized by grass

and sedge species that can tolerate periodic waterlog-

ging. Species dominance is not uniform, occurring in

mosaics of local dominance of different species. The

most common aquatic macrophyte is Potamogeton thun-

bergii which is quite abundant in pools in the channel.

Two patches of short grassland up to 1 m tall, domi-

nated by Sporobolus subtilis , occur in the midslopes of

the dambo area. Grass cover is generally high, with over

80% ground cover. The areas where this grassland is

found become waterlogged during the rainy season and

may remain so for long periods.

A patch of grassland dominated by Aristida junci-

formis occurs in the western part of the catchment area.

Hyparrhenia fdipendula is an important component of

this grassland, becoming more or less co-dominant in

some places. The grasses are generally short, most being

less than 1 m, and the cover is less than 70%.

Sporobolus pyramidalis grassland is found on both

well-drained soils on the upper slopes and on dark,

heavy, periodically waterlogged soils closer to the valley

bottom. It is composed of medium-height grasses of up

to 1 m with more than 90% ground cover. In the wetter

areas, Arundinella nepalensis becomes co-dominant.

Two patches of wooded grassland occur in the west of

the catchment area. Larger trees, Brachystegia spiciformis,

Julbernardia globiflora, Syzygium guineense subsp.

guineense and Acacia sieberiana , are mainly confined to

termitaria. The most common grass on termitaria is

Cynodon dactylon.

Natural woodland occurs on the interfluve and covers

about 12% of the catchment. Much of the miombo wood-

land is dominated by Brachystegia spiciformis which

forms almost pure stands in places. The canopy cover is

about 60%. The canopy layer is up to 10 m high and con-

sists of more or less even-aged trees. Scattered tall emer-

gent trees are found in places. The shrub layer is sparse

with occasional bare patches of ground.

To Marondera

To Harare

Key

□

Hyparrhenia ftlipendula grassland

Mixed dambo channel grassland

Sporobolus subtilis grassland

Aristida junciformis grassland

Sporobolus pyramidalis grassland

Mixed wooded grassland

Miombo woodland

Mixed woodland

Eucalyptus plantations

Cultivation and /or settlements

■ Railway

— Road

T rack

S^"**** Dambo margin

....... Catchment boundary

FIGURE 3. — Map showing vegetation types in grasslands catchment area near Marondera.

140

Bothalia 31,1 (2001)

One small patch of miombo woodland occurring in the

northwest of the catchment area is somewhat different in

character. Julbernardia globiflora is co-dominant with

Brachystegia spiciformis and there are a significant num-

ber of large Parinari curatellifolia trees on the edges. The

area is heavily disturbed by livestock and the shrub layer

is very poor. The herb layer is characterized by high den-

sities of Achyranthes aspera and Bidens pilosa.

A mixed woodland is found on the sandy soils in the

northwestern part of the catchment area. It is a sparse

woodland of up to 8 m with a canopy cover of 30-40%.

In some places it assumes the structure of a wooded

grassland and comprises a mixture of species whose

dominance varies from place to place. The most common

trees in this woodland are Combretum molle, Strychnos

spinosa, Burkea africana, Albizia antunesiana and

Vangueria infausta.

Cultivated land and settlements occupy just over 16%

of the catchment area. Much of the cultivation takes

place either side of the railway line embankment, with

other small areas close to settlements between the road

and the railway (Figure 3). There is also some cultivation

on the interfluve in the southwestern part of the catch-

ment area. These fields are used by the Grasslands

Research Station for experimental trials. In all the culti-

vated'areas, the main crop is maize.

About 24% of the catchment, primarily either side of

the dambo at the eastern end of the catchment, compris-

es plantations of exotic trees, particularly Eucalyptus

camaldulensis and other Eucalyptus species. These are

old trees (estimated to be at least 20 years) which have

grown to heights of up to 30 m.

Vegetation-environment relationships

A total of 63 species were encountered in the 16

detailed inventory plots. Each of the 16 plots was

assigned to one of the 10 vegetation types identified in

Table 1 . Several plots (i.e. 2, 4 and 9 to 12) were located

on or near the mapped lineation between vegetation

types (i.e. in ecozone regions), but in all cases it was pos-

sible, through observation, to place each of them in one

of the types. Of the natural vegetation types, only

TABLE 2. — Plot numbers of the vegetation types

Aristida junciformis and mixed wooded grassland were

not represented by any plots (Table 2).

The results of the CCA technique applied to the six

environmental variables and 63 species, indicates that

Axis 1 accounts for 84.6% of the observed variation in

vegetation, whereas Axes 2, 3 and 4 account for 54.3%,

48.6% and 38.7% respectively. Relationships between

plots and Axes 1 and 2 are shown in Figure 4. The 1 6

plots were separated, by eye, into seven groups, two con-

sisting of one plot each, four consisting of two or three

plots each, and one with four plots.

There is greater separation of the groups along Axis 1

than along Axis 2. Group I are the plots in the dambo

centre. Group II are the plots situated away from the

dambo centre, but otherwise in wetter parts of the

dambo. Group III contains the plots in drier parts of the

dambo generally closest to the dambo-interfluve margin,

and includes plot 14, which is located in the dambo spur.

Group IV and V are the plots found in a more or less tran-

sitional zone between the woodland on the interfluve

crest and the dambo grassland on the northern and south-

ern slopes respectively. Groups VI and VII are both

found in woodland on the interfluve crests, but differ in

species composition. These groups are defined by char-

acteristic indicator species (Table 3), generally conform-

ing to those described by Whitlow (1985).

The analysis shows strong positive correlation

between bulk density (BDEN) and September water

table depth (SWTD) (r = 0.71), electrical conductivity

(EC) and pH (r = 0.70), and percentage carbon (%C) and

EC (r = 0.79). There is strong negative correlation

C\J

■1.0 -0.5 0.0 0.5 1.0 1.5 2.0 2.5 3.0

FIGURE 4. — Canonical Correspond-

ence Analysis ordination

showing the separation of the

16 plots.

CCA Axis 1

Bothalia 31,1 (2001)

141

TABLE 3. — Indicator species in identified groups

Group Plots Location Indicator species

Arundinella nepalensis, Aristida junciformis, Sporobolus subtilis

Sporobolus pyramidalis, Hyparrhenia filipendula

Sporobolus pyramidalis, Monocymbium ceresiiforme

Sporobolus subtilis, Hyparrhenia filipendula

Hyparrhenia filipendula, Melinis repens, Cynodon dactylon

Brachystegia spiciformis, Dicerocaryum senecioides

Julbernardia globiflora, Parinari curatellifolia

between BDEN and EC (r = -0.76), and BDEN and %C

(r = -0.94). SWTD and February water table depth

(FWTD) both show strong positive correlation with Axis

1 (r = 0.87 and r = 0.96, respectively). %C and EC show

positive correlations with Axis 2 (r = 0.88 and r = 0.73,

respectively). BDEN shows a strong negative correlation

with Axis 2 (r = -0.83). The Monte-Carlo permutation

test shows significant differences in floristic composition

in relation to environmental variables ( p < 0.01).

However, the test is not significant for the first canonical

axis alone. Plots 1, 2, 3 and 16 are more influenced by

levels of the water table and the bulk density of the soil,

whereas %C, pH and EC are more important determi-

nants for the remaining plots, particularly those close to

the dambo centre. The depth of the water table is much

larger on the interfluve, where woodland predominates,

than in the dambo. where grassland predominates

(Figure 5).

DISCUSSION

Vegetation classification

The vegetation associations described in this paper

represent two major structural types, woodland and

grassland. The crests of the interfluves comprise wood-

land with a transition to grassland downslope. The

dambo is entirely grassland. Ten vegetation types were

defined and mapped on the basis of species dominance

and composition.

FIGURE 5. — Comparison of mean February water table depth

(FWTD) and September water table depth (SWTD) for the

seven vegetation groups shown in Figure 4.

A spatial successional sequence was observed along

the catena from the woodland areas on the upper slopes

down to the dambo channel. Definitive indicator species

include Monocymbium ceressiiforme marking the dambo

margin with woodland areas, while Sporobolus subtilis,

Andropogon eucomus and Aristida junciformis dominate

the wetter areas. The dambo centre vegetation is, howev-

er, different from that described by Whitlow (1985) and

in the wetter regions, particularly towards the catchment

outlet, it is characterized by Arundinella nepalensis.

Verbena bonariensis and Euphorbia cyparissoides.

It is evident that the vegetation structure and compo-

sition have been influenced by past and present manage-

ment practices in the catchment, particularly livestock

grazing and land clearance for crop farming. Intensive

grazing leads to changes in species composition with a

tendency towards increasing dominance of unpalatable

tuff grasses such as Sporobolus pyramidalis (Noy-Meir

1981; O'Connor 1995; Kennan 1969). It is, therefore,

hypothesized that the current composition of the Spo-

robolus pyramidalis grassland is related to past manage-

ment practices, and that it could have resulted from a

gradual conversion of Hyparrhenia filipendula grassland

owing to the impacts of grazing. Presently, cattle are

stocked at about 3-4 hectares per livestock unit on a rota-

tional basis (L. Lungu, late researcher at the Grasslands

Research Station, pers. comm.). However, there is a ten-

dency to concentrate animals in the paddocks close to the

channel. Localized degradation effects by cattle on veg-

etation is further seen in the prevalence of weeds such as

Bidens bitemata, Tagetes minuta, Achyranthes aspera

and Conyza albida in large sections of the grassland and

under the trees in the miombo woodland.

The miombo woodland shows evidence of manage-

ment. Campbell et al. (1995) described a miombo site in

the area and noted that much of the woodland cover was

removed after the establishment of the Grasslands

Research Station in 1929. Subsequently, the area has been

protected for the last 30 years which could explain the

even-aged structure of trees noted in the present study.

Vegetation-environment relationships

There is a general trend of increasing soil depth to the

water table from left to right along CCA Axis 1, which is

also associated with increasing species woodiness. This

is the key determinant separating dambo and interfluve

vegetation; the former lying at the negative end and the

latter lying at the positive end of Axis 1 . The only excep-

tion is Group IV which contains plots lying immediately

142

Bothalia 31,1 (2001)

up and down slope of the dambo spur on the northern

side of the catchment. Although located on the interfluve

this group falls on the negative end of Axis 1. However,

in terms of water table regime this group is intermediate

between the dambo and interfluve proper (Figure 5).

Within the dambo it is not possible to directly link

floristic composition to February water table depth

(FWTD) and September water table depth (SWTD). This

may be because FWTD and SWTD have been derived

from a single year’s data which are atypical and do not

reflect subtle long-term differences in the water table

regime across the dambo. Interpretation is further com-

plicated by management practices, which, as discussed

above, have altered vegetation composition within the

dambo. Nevertheless, a link between vegetation compo-

sition and soil moisture regime is revealed through the

interrelationship between the identified groups and the

other environmental variables used in the analysis.

Figure 4 shows that the groups of dambo vegetation are

spread along CCA Axis 2. Since there is a trend of

increasing soil pH, EC and %C from the negative to the

positive end of Axis 2, these are the key environmental

variables associated with differences in species composi-

tion within the dambo. However, pH and EC are highly

correlated with organic matter content (expressed as %C)

which is largely governed by the extent and duration of

waterlogging (Savory 1965). It is therefore probable that,

over long periods of time, the variation in floristic com-

position would correlate with differences in water table

regime. Thus, while not directly identified by FWTD and

SWTD, it is very likely that differences in vegetation

composition within the dambo are influenced by subtle

variation in the water table regime over time.

Considering the whole catchment, it is evident that the

amount of soil moisture and extent of waterlogging are

major factors influencing vegetation composition.

Comparison of Tables 2 and 3 reveals that in several

of the groups identified in the CCA, plots located in

more than one of the natural vegetation classes identified

independently in the classification and mapping exercise.

For instance. Group 1 comprises Plots 9 and 10 which

occur in the region classified as mixed dambo channel

grassland, as well as Plot 1 1 which is located in region of

Hyparrhenia filipendula grassland. Similarly, in Group

II, Plots 6 and 8 lie in a region classified as Sporobolus

pyramiclalis grassland, but Plot 12 occurs in a region

classified as Sporobolus subtilis grassland, while in Group

III, plots 4, 5 and 7 are in a region classified as Spo-

robolus pyramidatis grassland, but Plot 14 is located in a

region of Sporobolus subtilis grassland. Nevertheless,

within each group, the majority of the plots lie within

one type, as shown in Table 4.

TABLE 4. — Links between vegetation type and group

These results indicate that there is a strong, but not

unambiguous, correspondence between the mapped veg-

etation classes and the environmental variables identified

by the CCA which are associated with floristic composi-

tion. The lack of a clearer relationship can be attributed

to two factors. Firstly, the common link between plots

within a group are the indicator species rather than the

dominant species which identify the class. Secondly,

plots are effectively spot samples and may not be truly

representative of the vegetation of the area within which

they are situated. This second factor in particular ex-

plains the lack of correspondence for those plots that lie

in ecozones, where the vegetation is transient between

vegetation types.

CONCLUSION

Although land management policies have influenced

vegetation composition to some extent, there is never-

theless a clear relationship between vegetation composi-

tion and environmental variables within the catchment.

The amount of soil moisture and extent of waterlogging

(as indicated by the levels of the water table in the wet

and dry seasons) is the overriding factor distinguishing

the floristic composition of the interfluve and the dambo.

Furthermore, within the dambo, the study findings strong-

ly suggest that the extent and duration of waterlogging is

a key influence in determining the presence/absence of

indicator species.

The results strongly support qualitative observations

made elsewhere in Zimbabwe. Subtle differences in soil

moisture resulting from slight variation in water table

regime can result in noticeable differences in vegetation

composition. The relationship between the identified

vegetation classes and the environmental variables used

in the CCA suggests that with additional data collection

and analysis, vegetation associations may provide a basis

for identifying characteristic soil moisture regimes in

similar highveld catchments containing dambos.

ACKNOWLEDGEMENTS

This research was partly funded by the UK Natural

Environment Research Council through the Dambo

Processes Integration Experiment. Alfred Maroyi of the

National Herbarium, Harare, helped with botanical field-

work. Luckmore Chigwaze of the Horticultural Research

Centre, Marondera assisted with field measurement of

the environmental variables.

REFERENCES

BENNETT, K.E. 1980. Key to Zimbabwean grass species. Kirkia 11:

169-286.

CAMPBELL, B.M., CUNLIFFE, R.N. & GAMBIZA, J. 1995. Vege-

tation structure and small-scale pattern in rniombo woodland,

Marondera, Zimbabwe. Bothalia 25: 121-126.

CHAPMAN, D.J. 1987. Soil survey: methods of analyses for soils.

Unpublished report to Department of Research and Specialist

Services, Soils & Chemistry Research Institute, Harare, Zimbabwe.

DRUMMOND. R.B. 1975. A list of trees, shrubs and woody climbers

indigenous or naturalised in Rhodesia. Kirkia 10: 229-285.

DYE, PJ. & WALKER, B.H. 1980. Vegetation-environment relations

on sodic soils of Zimbabwe Rhodesia. Journal of Ecology 68:

589-606.

Bothalia 31,1 (2001)

143

GIMINGHAM, C.H. 1972. Ecology of heathlands. Chapman & Hall,

London.

GURNELL, A.M. 1981. Heathland vegetation, soil moisture and dyna-

mic contributing area. Earth Surface Processes and Landfonns

6: 553-570.

GURNELL, A.M. & GREGORY, K.J. 1987. Vegetation characteris-

tics and the prediction of runoff: analysis of an experiment in

the New Forest. Hampshire. Hydrological Processes 1: 125 —

142.

KENNAN, T.C.D. 1969. A review of research into cattle-grass rela-

tionship in Rhodesia. Proceedings of the veld management con-

ference'. 5-28. Government Printers, Salisbury.

MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and me-

thods of vegetation ecology. John Wiley, New York.

NOY-MEIR, I. 1981. Responses of vegetation to the abundance of

mammalian herbivores. In P.A. Jewel & S. Holt, Problems in

management of locally abundant wild animals: 233-246.

Academic Press, New York.

O’CONNOR. T.G. 1995. Transformation of a savanna grassland by

drought and grazing. Journal of Range and Forage Science 1 2:

53-60.

PERERA, N.P. 1982. The ecology of wetlands (dambos) of Zambia and

their evolution for agriculture — a model for the management of

wetlands in subhumid eastern and southern Africa. International

Journal of Ecology and Environmental Science 8: 27-38.

PRATT. D.J., GREENWAY. PJ. & GWYNNE, M.D. 1966. A classifi-

cation of East African rangeland, with an appendix on termi-

nology. Journal of Applied Ecology 3: 369-382.

RATTRAY, J.M. 1957. The grasses and grass associations of Southern

Rhodesia. Rhodesia Agriculture Journal 54: 197-234.

SAVORY, R.M. 1965. The relationship of soil development to water

table levels in a granitevlei at Marandellas, Southern Rhodesia.

Unpublished M.Sc. (Agriculture) thesis, University of London.

TER BRAAK, C.J.F. 1986. Canonical Correspondence Analysis: a new

eigenvector technique for multivariate direct gradient analysis.

Ecology 67 : 1167-1179.

TER BRAAK, C.J.F. 1987. The analysis of vegetation-environment

relationships by Canonical Correspondence Analysis. Vegetatio

69: 69-77.

TER BRAAK. C.J.F. 1988a. CANOCO— an extension of DECORANA

to analyse species environment relationships. VegetatiolS: 159,

160.

TER BRAAK, C.J.F. 1988b. CANOCO — a Fortran program for cano-

nical community ordination by [partial] [detrended] [canoni-

cal] correspondence analysis, principal components analysis

and redundancy analysis (Version 3.1). Agricultural Mathe-

matics Group. Technical report, Wageningen.

TIMBERLAKE, J„ NOBANDA, N. & MAPAURE, I. 1993. Vegeta-

tion survey of the communal lands — north and west Zimbabwe.

Kirkia 14: 171-270.

WHITLOW. J.R. 1984. A survey of dambos in Zimbabwe. Zimbabwe

Agricultural Journal 81: 129-138.

WHITLOW, J.R. 1985. Dambos in Zimbabwe: a review. Zeitschrift fur

Geomorphologie Supplementband 52: 1 15-146.

Bothalia 31.1: 145-146 (2001)

Miscellaneous notes

POACEAE

THE BASIC CHROMOSOME NUMBER OF THE GENUS PENTAMERIS ( ARUNDINOIDEAE)

The genus Pentameris Beauv. contains nine species and

is endemic to Western Cape, South Africa (Barker 1993).

In his revision of Pentameris , Barker (1993) described

chromosome numbers for P distichophylla (Lehm.) Nees

(2n = 6x = 36) and P. thuarii Beauv. (2n = 2x = 12). Both

numbers suggest a basic chromosome number of 6 for this

genus. Unfortunately his report did not contain any photo-

graphic evidence of these chromosome numbers.

During this study, meiotic chromosomes of the fol-

lowing species were studied according to the method

described by Spies & Du Plessis (1985):

P. thuarii , Spies 6160: Western Cape, 3322 (Oudtshoom): Montagu

Pass, (-CD).

P oreophila N.P.Barker, Spies 6166: Western Cape, 3322 (Oudts-

hoom): Swartberg Pass, (-AC).

P. macrocalycina (Steud.) Schweick., Spies 3644: Western Cape.

3319 (Worcester): Franschhoek Pass, (-CC).

At least 20 well-spread meiotic cells, where chromo-

some configuration could be recognized, were studied

per specimen.

These results clearly indicate that P. thuarii (Figure

1 A) and P. oreophila (Figure IB) have gametic chromo-

some numbers of n = x = 7, and P. macrocalycina (Figure

1C, D) has n = 3x = 21. The most interesting finding was

the high number of multivalents observed in the P.

macrocalycina specimen. The lowest number observed

in a cell was five and most cells contained seven quadri-

valents. This possibly indicates that the particular speci-

men is an auto-alloploid with a possible AAAABB

genomic constitution.

The results of this study unequivocally indicate that

the basic chromosome number of Pentameris should be

x = 7. This supports the phylogenetic analysis, based on

ITS sequences of the Arundinoideae, of Hsiao et al.

(1998), where Pentameris forms a clade with Penta-

schistis and Prionanthium , both having a basic chromo-

some number of seven (De Wet 1954, 1960; Tateoka

1965; Davidse et al. 1986; Davidse 1988; Du Plessis &

Spies 1988; Spies & Du Plessis 1988; Du Plessis & Spies

1992; Morton 1993; Spies et al. 1994; Visser & Spies

1994; Klopper et al. 1998).

ACKNOWLEDGEMENTS

The University of the Orange Free State and the

National Research Foundation are thanked for financial

assistance during this study.

FIGURE 1. — Meiotic chromosomes in

the genus Pentameris. A, P. thua-

rii, Spies 6160, late anaphase I

with 7 chromosomes in each

pole; B, P. oreophila , Spies

6166, early anaphase I. C, D, P.

macrocalycina, Spies 3644: C,

metaphase I with 6iv9n: D,

diakinesis with 7iV7n. Scale bar:

6.4 pm.

146

Bothalia 31,1 (2001)

REFERENCES

BARKER, N.R 1993. A biosystematic study of Pentameris (Arundi-

neae, Poaceae). Bothalia 23: 25-47.

DAVIDSE, G. 1988. A revision of the genus Prionanthium (Poaceae:

Arundineae). Bothalia 18: 143-153.

DAVIDSE, G„ HOSHINO, T. & SIMON. B.K. 1986. Chromosome

counts of Zimbabwean grasses and an analysis of polyploidy in

the grass flora of Zimbabwe. South African Journal of Botany

52: 521-528.

DE WET, J.M.J. 1954. The genus Danthonia in grass phylogeny.

American Journal of Botany 41: 204—211.

DE WET, J.M.J. 1960. Chromosome numbers and some morphological

attributes of some South African grasses. American Journal of

Botany 47: 44—49.

DU PLESSIS, H. & SPIES, J.J. 1988. Chromosome studies on African

plants. 8. Bothalia 18: 119-122.

DU PLESSIS. H. & SPIES, J.J. 1992. Chromosome numbers in the ge-

nus Pentaschistis (Poaceae, Danthonieae). Taxon 41: 709-720.

HSIAO, C„ JACOBS. S.W.L., BARKER, N.P. & CHATTERTON, N.J.

1998. A molecular phylogeny of the grass family (Poaceae) based

on sequences of rDNA. Australian Systematic Botany 11: 41-52.

KLOPPER, K.C., SPIES, J.J. & VISSER, B. 1998. Cytogenetic studies

in the genus Pentaschistis (Poaceae: Arundinoideae). Bothalia

28: 231-238.

MORTON, J.K. 1993. Chromosome numbers and polyploidy in the

flora of Cameroon Mountain. Opera Botanica 121: 159-172.

SPIES, J.J. & DU PLESSIS, H. 1985. The genus Rubus in South

Africa. I. Chromosome numbers and geographical distribution

of species. Bothalia 15: 591-596.

SPIES. J.J. & DU PLESSIS. H. 1988. Chromosome studies on African

plants. 6. Bothalia 18: 1 11-114.

SPIES. J.J., LINDER, H.P., LABUSCHAGNE, I.F. & DU PLESSIS, H.

1994. Cytogenetic evidence for the species delimitation of

Pentaschistis airoides and P patula (Poaceae: Arundineae).

Proceedings of the XIII'1' Plenary Meeting of AETFAT, Zomba,

Malawi 1: 373-383.

TATEOKA, T. 1965. Chromosome numbers of some East African

grasses. American Journal of Botany 52: 864—869.

VISSER, N.C. & SPIES, J.J. 1994. Cytogenetic studies in the genus

Tribolium (Poaceae, Danthonieae). V. Section Acutiflora. South

African Journal of Botany 60: 285-292.

J.J. SPIES* and R. ROODT*

* Department of Botany and Genetics (106), University of the Orange

Free State, P.O. Box 339, 9300 Bloemfontein.

MS. received: 2000-02-17.

Bothalia 31,1: 147-150 (2001)

Book Reviews

FLORA OF AUSTRALIA VOL. 17A. PROTEACEAE 2: GREVIL-

LEA, by R.O. MAKINSON. 2000. CSIRO Publishing , P. O. Box 1 139,

Collingwood, 3066 Victoria, Australia. Hard cover ISBN 0 643 05969

5. price US$ 89.95; paperback ISBN 0 643 05970 9, price US$ 69.95.

Pp xx + 524, frontispiece + 64 colour figures, 40 b/w figures, 452 dis-

tribution maps.

By common consent (see, for example, Schmid 2000) Flora of

Australia is among the best examples of this genre in the world; with-

out hesitation, this reviewer joins the chorus of praise for the latest

instalment. This volume treats only one genus, namely Grevillea.

However, that genus encompasses no less than 357 species and 95

infraspecific taxa, of which 43 taxa are newly described here, making

this the third-largest genus of Australian flowering plants. Two other

volumes of similar size are needed to complete the account of

Australian Proteaceae. That these already exist is a tribute to the orga-

nization and productivity of Australian taxonomists.

There is the common misconception in South Africa that the fami-

ly Proteaceae is resident only in the winter rainfall area of the Western

Cape. Meyer et al. (1997) indicate that in the southern African flora we

have 360 species and 32 infraspecific taxa of Proteaceae, in other

words, about the same diversity as in a single Australian genus of the

family. Admittedly, Grevillea is their largest genus by quite a wide mar-

gin. A strictly family to family comparison would reveal that Australia

has 46 genera and 1118 species to our 16 genera and 360 species (fyn-

bos trumpeters please take note).

The present volume reveals some features which one would be

delighted to see more of in other floras. For example, a dedication to

previous workers in the group is not unheard-of. but it is a respectful

gesture to one's predecessors, and, a frontispiece reproducing a water-

colour painting of a suitably decorative species appearing in the vol-

ume is also not unknown (as, for example, in earlier volumes of Flora

zambesiaca), yet is very attractive. In the case of the latter item, the

choice must have been difficult indeed, in view of the number of spec-

tacular species in the genus treated. Many more (but still barely one in

five) species are illustrated in a section of colour photographs, an idea

one would like to see copied in other floras. A relatively short intro-

ductory essay gives details of the history and current state of the study

of this genus, possible phylogeny, biogeography and ecology, mor-

phology, and poisonous and irritant properties. The species are keyed

out not only in taxonomic groups, but with keys to species occurring in

Northern Territory, South Australia, Queensland, New South Wales and

Victoria (these placed between the main taxonomic text and the maps).

There are no keys for Western Australia or Tasmania, as the former

state is home to the overwhelming majority of Grevillea species, so a

state key would almost echo the main one, and the latter state boasts

only one species, making a key superfluous. The text for each species

is in every way up to the high standard one has come to expect of this

Flora.

Why should a South African welcome this work so enthusiasti-

cally? Obviously, it is a model of what a 21st century Flora ought to

be, and so an inspiration to all taxonomists. Closer to home, the

species treated have considerable horticultural potential in a climate

like ours, and sooner or later one is going to be called on to identify

material of this genus. Already, 15 species, one hybrid and several

cultivars of Grevillea are known to be in cultivation in southern

Africa. More worryingly, one of these is reported to be starting to

invade Mpumalanga; in this case a clue to possible control measures

is given by the presence, no more than three metres from the work-

station at which this review is being written, of a cabinet made of the

wood of G. robusta. The account reviewed here informs me (with a

reference to literature) that this species is recorded as containing an

irritant in the wood.

In summary, this volume can only enhance the already high regard

in which the Flora of Australia is held, and one feels that the author and

each member of the production team have earned the great Strine com-

pliment: 'Good on yer, mate!’.

REFERENCES

MEYER, N.L., MOSSMER. M. & SMITH, G.F. (eds). 1997. Taxon-

omic literature of southern African plants. Strelitzia 5. National

Botanical Institute, Pretoria.

SCHMID. R. 2000. Review: Flora of Australia. Taxon 49: 612.

H.F. GLEN*

* National Herbarium, National Botanical Institute, Private Bag X101,

0001 Pretoria.

PHOTOGRAPHIC GUIDE TO TREES OF SOUTHERN AFRICA, by

BRAAM VAN WYK, PIET VAN WYK and BEN-ERIK VAN WYK.

2000. Briza Publications , P.O. Box 56569, Arcadia, 0007 Pretoria. Pp.

356. Soft cover: ISBN 1-875093-24-9, price R169.95.

This is another book on southern African plants published by Briza

in their now well-established series of illustrated books to promote

knowledge of indigenous plants among the general public (Van

Oudtshoom 1992; Van Wyk & Smith 1996; Venter & Venter 1996; Van

Wyk et al. 1997; Smith et al. 1998; Smith 1999; Van Wyk & Gericke

2000).

The authors have used a different approach to that of Van Wyk &

Van Wyk (1997) in that it is aimed mainly at the interested layman. It

is less scientific with numerous visual aids for easy identification. The

authors stated: ‘Our main aim was to show the interesting and charac-

teristic crown shapes and branching patterns of trees, with less focus on

the detailed features of the leaves and flowers by which we would usu-

ally identify trees. The book is intended as a visual guide to the identi-

fication of trees within the characteristic southern African landscape, of

which they are an integral part.’ Southern Africa here (as in Van Wyk

& Van Wyk 1997) comprises Namibia, Botswana, Zimbabwe,

Mozambique south of the Zambezi River, South Africa, Swaziland and

Lesotho. The colour coding on the distribution maps is also similar to

that in Van Wyk & Van Wyk (1997).

There are a number of brief (one page long) introductory chapters

on Biomes and the habitat of trees. Fruit trees (referring specifically to

indigenous trees). Furniture and craftwork from trees. Firewood trees.

Medicinal and poisonous trees. Trees and animals. Each of these chap-

ters is richly illustrated with a full page of beautiful colour pho-

tographs. The next chapter describes the Architecture of trees accord-

ing to a number of models, and a few examples of southern African

trees are mentioned under each. The chapter on Identifying trees pre-

sents one with guidelines for characters to use in the identification of

trees. This is followed by a page providing an Explanation of icons and

codes used in the descriptions of the trees.

Three hundred species of mostly indigenous trees are described in

alphabetical order according to species, with a few exotics also includ-

ed. Each page deals with one species. The description consists of the

scientific name, English and Afrikaans common names, main features

such as size of the tree, shape of the crown, and the bark, leaf, flower

and fruit characters, reference to similar species, habitat description

and uses. Each species is depicted by excellent, clear, colour pho-

tographs: a large one of the habit (growth form) and smaller ones of the

flowers, fruits and/or bark, a colour-coded distribution map illustrating

the range of distribution in southern Africa as well as coloured icons,

the meaning of each explained in the beginning of the book, all of

which are clear tools to assist in the identification of a specific tree. As

the purpose of the book is identification of a tree by means of its more

obvious features, the photographs and icons occupy about half of each

page and the description the other half. I was about halfway through the

book when I realized that the family name to which the tree belongs, as

well as some synonyms and the RSA and Zimbawean tree numbers are

also supplied in small print at the bottom of each page for those inter-

ested, and for the real botanists! Many of the photographs are repeti-

tions from Van Wyk (1993, 1996) and Van Wyk & Van Wyk (1997).

148

Bothalia 31,1 (2001)

A relatively simple key is provided in the Guide to the species. This

key is based on the icons provided in each tree description. It is

designed to help one in identifying an unknown tree. There is also an

illustrated glossary in which some terms are explained, followed by a

page of references for further reading, and an index to scientific, fami-

ly and common names. The final page contains a summary of the icons

used and a ruler in millimetres, should one need to take measurements

in the field.

So many books on south(-ern) African trees have appeared on the

shelf in the past few years, that one wonders why another book on

southern African trees should be considered as really necessary (Van

Wyk 1993; Moll & Moll 1994; Van Wyk & Van Wyk 1997; Pooley

1998, 1999). The same basic information is repeated with little addi-

tional information added. This book could have had a new approach if

an attempt was made to classify the trees according to the different

architectural models described by the authors in one of the introducto-

ry chapters. However, it possibly fills a niche in the market for the lay-

man who wants to know and identify southern African trees.

REFERENCES

MOLL, E. & MOLL, G. 1994. Common trees of southern Africa, edn

2. Struik Publishers, Cape Town.

POOLEY, E. 1998. Trees: southern African green guide. Southern Book

Publishers, Halfway House.

POOLEY. E. 1999. Saso! first field guide to trees of southern Africa.

Struik Publishers, Cape Town.

SMITH, G.F., CHESSELET, P„ VAN JAARSVELD, E.J., HARTMANN,

H„ HAMMER, S„ VAN WYK, B-E., BURGOYNE, P, KJLAK,

C. & KURZWEIL, H. 1998. Mesembs of the world. Illustrated

guide to a remarkable succulent group. Briza Publications,

Pretoria.

SMITH. N. 1999. Guide to the acacias of South Africa. Briza Publi-

cations, Pretoria.

VAN OUDTSHOORN, F.P. 1992. Guide to grasses of South Africa.

Briza Publications, Pretoria.

VAN WYK, B. & VAN WYK, P. 1997. Field guide to trees of southern

Africa. Struik Publishers, Cape Town.

VAN WYK, B-E. & GERICKE, N. 2000. People's plants. A guide to

useful plants of southern Africa. Briza Publications, Pretoria.

VAN WYK, B-E. & SMITH, G. 1996. Guide to the aloes of South

Africa. Briza Publications, Pretoria.

VAN WYK. B-E., VAN OUDTSHOORN, B. & GERICKE, N. 1997.

Medicinal plants of South Africa. Briza Publications, Pretoria.

VAN WYK, P. 1993. Southern African trees: a photographic guide.

Struik Publishers, Cape Town.

VAN WYK, P. 1996. Field guide to the trees of the Kruger National

Park , edn 3. Struik Publishers, Cape Town.

VENTER, F. & VENTER, J-A. 1996. Making the most of indigenous

trees. Briza Publications, Pretoria

P.P.J. HERMAN*

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

THE WORLD OF HAWORTHIAS Vol. 2. Descriptions, by INGO

BREUER. 2000. Ingo Breuer and Arbeitskreis fiir Mammillarien-

freunde e. V. (AfM), Niederzier and Homburg/Saar, Kirchstr. 36, 52382

Niederzier, Germany. Pp. 518. Soft cover: ISBN 3-926573-13-9. Price:

DEM 84 (± R300).

A copy of this book was donated to the Mary Gunn Library of the

National Botanical Institute in Pretoria by the author, Ingo Breuer, and

as editor of this journal and in appreciation of the gesture, I thought it

only fitting to tell the reader something about this publication.

Volume 1, presenting all the relevant data necessaiy for original

research and the status of the names, appeared in 1998 (Breuer 1998)

and consists of five chapters. Volume 2 consists of chapter 6, the pagi-

nation continuing from Vol. 1, and deals with descriptions and the

nomenclature of close to 500 names used for taxa in Haworthia to date.

In the Foreword, Prof. Gideon Smith, Director of Research and

Scientific Services, National Botanical Institute of South Africa, states:

‘This treatment of Ingo Breuer comes as a refreshing change. It allows

the reader to make his or her own decisions on the application of some

of the controversial names, and therefore assists in maintaining an

acceptable level of stability in the over 400 names that have been vari-

ously upheld for the more or less 80 species of Haworthia.'

Descriptions of taxa are arranged in alphabetical order under the

following headings: Reference, Illustrations, Latin diagnosis. Descrip-

tion. Further names included. Origin. Type. Distribution, Additional

notes and Remarks. The reference includes the literature reference

number from Volume 1, Chapter 1 and the complete literature quotation

with the page number of the beginning of the description. New litera-

ture since Volume 1 is listed alphabetically according to author and

appears towards the end of this volume. The Latin diagnosis is given

only if it was included in the original publication. The description of

each taxon contains the original description in its original language.

English translations are provided for descriptions in German, French,

Portuguese, Dutch, Norwegian or Latin.

The author endeavoured to obtain illustrations of most of the type

material to accompany each description so that a better image of the

taxon could be presented. Furthermore, where possible, black-and-

white photographs of specimens from type locations or locations with-

in the known distribution areas, are provided. A 24-page section of

colour photographs depicting various Haworthia taxa is found near the

middle of the volume.

For many of the Haworthia names, a type was not available at the

time of description and had to be designated afterwards. In this volume,

there are a further 28 lectotypes and 16 neotypes designated and listed.

Since the publication of the first volume in 1998, several new taxa have

been described, three new species and 31 new varieties, mostly by

Bayer (1999). A holotype is cited for all taxa and the new names are

listed in alphabetical order. The place of discovery, the collector of the

original material and collecting number of the herbarium material and,

if available, the isotype and its depository are also included. In the case

of types of living plants, the botanical garden where the plant is cur-

rently cultivated is indicated.

As Prof. Smith says ‘Ingo Breuer has established himself as a

leader in the field of accessing information on taxon definition in the

popular succulent plant genus Haworthia. The success of what he has

begun as a scholarly and meticulous librarian and researcher will no

doubt continue and in years to come serve as a most worthwhile basis

for further refinement of our understanding of the classification of and

relationships in this taxonomically and morphologically important

group.’

The book and its companion volume provide valuable tools for

understanding the world of haworthias.

REFERENCES

BAYER, M.B. 1999. Haworthia revisited. A revision of the genus.

Umdaus Press, Hatfield, South Africa.

BREUER, I. 1998. The world of haworthias, Vol. 1. Bibliography and

annotated index. Breuer and Arbeitskreis fiir Mammillarien-

freunde e. V. (AfM). Niederzier, Germany.

G. GERM1SHUIZEN*

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

THE CAMBRIDGE ILLUSTRATED GLOSSARY OF BOTANICAL

TERMS, by MICHAEL HICKEY and CLIVE KING. 2000. Cambridge

University Press, The Edinburgh Building, Cambridge, CB2 2RU, UK.

Pp. xii + 208, 297 x 210 mm. Soft cover: ISBN 0 521 79401 3, price

£18.95; Hard cover: ISBN 0 521 79080 8, price (not known).

B.D. Jackson, whose monumental A glossary of botanic terms com-

prises some 25 000 terms, quotes Dr Samuel Johnson: ‘Every other

author may aspire to praise, the lexicographer can only hope to escape

reproach.’ So how do you do justice to the poor compilers of glossaries,

how do you assess their work fairly? Do you count the number of

entries, do you compare il to other works in the field, or do you just live

with it for a while and see what it does for you? I did a bit of all that.

Bothalia 31,1 (2001)

149

To begin with, I was surprised at the format of the book (by the

way, I have only seen a soft cover copy). I always thought that profes-

sional publishers disliked the A4 format, presumably for aesthetic rea-

sons, and that this size was favoured rather for practical reasons by peo-

ple not professionally in the book business. As will appear later on,

there are other elements about this book that make you think it was pro-

duced rather for the love of the subject than for the financial gains that

might be derived from it. That is not to say that I think it will not sell

well. On the contrary, it could well be a bestseller.

It has a bright, cheerful cover: the front, in the palest avocado

green, shows a horse-chestnut leaf, complete with conkers, ready for

action, and the bottle-green back has ferny fronds and blurb. In the

foreword, S.M. Walters points out that this entirely new book is the

product of the collaboration of many years between Michael Hickey

and Clive King in the Cambridge University Botanic Garden. The

authors state that they included all the terms most commonly used in

describing vascular plants, as well as some elementary histological,

cytological and genetical terms. Chemical terms have been largely

excluded. In the notes to readers, which include hints on the use of hand-

lens and microscope, it is pointed out that the number in square brack-

ets after the definition of a term refers to the page on which the term is

illustrated. A list of commonly used symbols includes ‘±’ in the sense

of ‘more or less’, which the mathematicians get all worked up about.

This is followed by lists of Latin and Greek prefixes (general and

numerical) as well as suffixes, measurements and taxonomic ranks.

Lists of ranks may be particularly useful, now that ranks above order

tend to be relegated to the realm of fantasy. Perhaps the suffixes of

ranks could have been added directly and not listed separately. Pages 1

to 46 present definitions of some 2 400 terms. Pages 47 to 208 are

devoted to illustrations. I found the definitions generally concise and

clear, and having illustrations to back up many of them is a major

advantage. Latin plurals are dealt with globally on p.l. This saves

space, something that is not particularly evident in the rest of the work,

but is considered less user-friendly. Contrasting terms are not given. I

was somewhat disappointed when I looked in vain for a number of

terms I had come across recently, admittedly mainly in the older litera-

ture. Some examples, all of which I found in both Jackson and my

Millennium Collins, are: alveolate, anticous (posticous is given, but as

‘postichous’), arillode, condyle, dextrorse and sinistrorse, dissepiment,

idioblast, incrassate, mamillate, mono- and polyphyletic, muticous,

putamen, and tristichous. I hasten to add that I did not find them either

in the glossary of the latest volume of the European garden flora. One

may well argue that the best way of getting rid of outdated or superflu-

ous terms is not to use them and to refer the information seeker to

Jackson. And seek we must, because the old literature, often containing

uncommon terminology, is still almost our daily bread. The word

‘cladistics’ is explained but cladistic jargon is sold short.

For me the heart of the matter are the illustrations which are spread

out in a rather lavish manner over roughly three-quarters of the book.

The blurb states that the glossary is comprehensive and beautifully

illustrated. Concerning the illustrations, I would certainly regard them

as comprehensive. I can not think of a glossary more extensively illus-

trated. But I can certainly think of glossaries with more information per

square decimeter of paper — this old-fashioned unit of measurement is

not in the glossary but ‘mm Millemetre(s)’ is. Looking at the three

pages of inflorescence illustrations, I find them no more informative

than the single page on the subject in Heywood’s Flowering plants of

the world. There is much to be said for large clear illustrations but

using 28 pages for 14 headings and 14 drawings with a purely decora-

tive function, which are repeated later, seems unnecessary luxury for a

book that appears to have been designed primarily for the use of stu-

dents and others in need of an affordable practical handbook. As the

blurb informs us ‘the illustrations are grouped in sections according to

specific features such as leaf shape and flower structure, so allowing

comparison of different forms at a glance. In addition to supporting the

definitions, the illustrations therefore also provide a unique compila-

tion of information that can be referred to independently of the defini-

tions’. I would not describe the illustrations as beautiful but rather as

informative or effective, and this is obviously what their main attributes

should be in a work of this nature. Numerous subjects and themes that

are rarely, if ever, depicted in such glossaries, are shown: annulate,

photosynthetic and contractile roots, turions, droppers or sinkers;

stages in germination of Rhizophora; jaculators in Ruellia', Raunkiaer’s

life forms; leaf fall in a woody dicotyledon; thermonasty, nyctinasty,

geocarpy, etiolation and hydrotropism; Fibonacci numbers in phyl-

lotaxy; Beltian bodies, myrmecophilous plants; anthocarp, carpophore;

limen in Passiflora\ flower structure of Canna and Strelitzia. the haft in

Iris', hinge cells in the leaf of Ammophila, bursicles in Dactylorhizw, the

etaerio of Rubus. the balausta of Punica, the hesperidium of Citrus, the

coenocarpium of Ananas, the galbuli of Juniperus, the velum of

Isoetes, and massulae of Azolla. As you page through the illustrations

you notice, al least in my copy, that on p.105, and less so on p.106

behind it, there is very little bottom margin.

Comparing the work to Jackson one comes to the obvious conclu-

sion that for general every day use it is much more user-friendly but

that the old classic is by no means ready for the recycler. Comparing it

to a work like Vascular plant systematics by Radford et al. it scores

heavily in the ease of access: if you are looking for the definition of a

given term you have only one alphabet to run through, or if you are

looking for an illustration of a specific feature you have a choice of

only 14 well-defined sections.

I foresee that this book will prove to be very popular, especially

among students, and that it will greatly facilitate their venture into the

jungle of botanical terminology.

O.A. LEISTNER*

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

CONTEMPORARY PLANT SYSTEMATICS. edn 3, by DENNIS W.

WOODLAND. 2000. Andrews University Press, 213 Information

Services Building. Berrien Springs. Michigan 4 9104-1700, USA. Hard

cover: ISBN 1-883925-25-8, 275 x 180 mm, price: $64.99.

I approached this book with high expectations: it has a serviceable,

plastic-covered case (my younger daughter lovingly rubbed it against

her cheek: ‘nice and smooth"), a visually striking, coloured cover

design with flowers, fern fronds and a CD against a pinaceous back-

ground; it is the third edition of a richly illustrated, and presumably

popular handbook called ‘a winner’ by Taxon, and recommended by

Systematic Botany for introductory plant systematic courses (according

to the blurb on the back cover), and it comes complete with a free CD

showing 4 700 plant images.

Let me get my reasons for a certain disappointment off my chest

right at the start. One tends to think that book reviewers who dwell on

printing errors, slips of the pen, misguided forefingers — call them what

you like — have little else to say. But in this case I find them too fre-

quent and too blatant not to dwell on them. To give but a few of the

most glaring examples: on p. 34, comprising less than 200 words, there

are three spelling errors. And if ‘pendent’ was not correctly spelt on the

facing page, on which there are two spelling errors, you might think it

was not merely a spelling error. In the glossary, on p. 538, four terms

are spelt incorrectly, and in such a way that this may not be obvious to

the newcomer. Further on in the glossary ‘repand’ refers you to

‘unulate’, where again the d is missing, and under ‘upland theory’ the

word ‘banitats’ presumably stands for ‘habitats’. The term ‘synconi-

um’, which I have found in no other reference work, is used through-

out the book for ‘syconium’. Edward Lee Greene, correctly spelt on p.

465 is given twice as ‘Green’ on p. 7. Talking about author names,

Authors of plant names by Brummitt & Powell (1992), given in one of

several most useful literature lists, is not followed, at least not consis-

tently: John Torrey is abbreviated to ‘Torrey’ and J.D. Hooker to

‘Hook.’. Brummitt (1997) is cited on p. 386 but is not included in the

references for this chapter on pp. 485 to 487. (It should be Taxon 46:

723-734). The less said about the spelling of foreign words the better:

in the single word frangaise in the title of Lamarck’s famous flora men-

tioned on p. 13, there are three spelling mistakes. I should perhaps men-

tion that in Stafleu & Cowan (1979) there is also one mistake in that

word, whereas in Stafleu (1967) it is correct. I noticed only a few

spelling mistakes in scientific names, such as Nicotiana tubacum (also

in edn 2), Tropaeolum majas on the CD and Rhynchacalycaceae on pp.

206 and 566.

In the glossary, the author often defines adjectives as though they

were nouns: acaulescent: a naked stem; awl-shaped: a small... ever-

green leaf; coalescent: sepals united; or the other way round: natural-

ization: established completely after introduction from another region.

Some definitions I would not go along with, or would try to improve,

e.g. genus: the first word of the species name of a plant or animal; used

150

Bothalia 31,1 (2001)

as a noun and capitalized; hot spot: a rising plume of magma that flows

out through an opening in the earth’s crust; radical: a root or arising

from that root; type: a specimen designated by the author of a species

as the type or morphological example for that taxon (syn. holotype).

Enough of the gripes. Chapter 1, Significance of systematics, deals

with taxonomy, biosystematics, nomenclature and the 'green revolu-

tion’. 2, How plants get their names, introduces the reader to common

or vernacular names, Latin names and the Principles of the Code of

Botanical Nomenclature. In the discussion of the Code, verbatim

quotes would sometimes have been preferable to the wording given.

Oversimplifications such as ‘trees are given as feminine’ occur, and

misleading phrases such as ‘type specimens of similar taxa'. The con-

struction of dichotomous keys is thoroughly explained in 3, How plants

are identified, but multi-entry keys are merely mentioned. 4,

Collecting, handling and preserving specimens, is described in detail. I

wonder how widely the ‘flexostat’ is used in this process in other insti-

tutions. Chapter 5, Families of ferns and their associated plants

(Pteridophytes), deals with 24 of the 37 families listed in the classifi-

cation of pteridophytes on pp. 30 and 31. Every family is dealt with on

a separate page with several line drawings and text divided into para-

graphs with headings such as General description. Stems, Leaves,

Sporangia, Class, Gametophytes, Distribution, Economic value.

Chromosomes/cytology, Classification and Fossil record. The basic

structure of ferns and their typical reproductive cycle are described and

illustrated at the beginning of the chapter. This chapter, much expand-

ed since the previous edition, is to be welcomed because so many basic

systematic texts, as listed on p. 396, deal only with flowering plants,

sometimes even when it says ‘vascular plants' in the title. Chapter 6,

Families of Pinophyta, treats gymnosperms in a similar way.

Terminology of flowering plants (Magnoliophyta) in chapter 7 includes

illustrated glossaries of stem structures, leaf morphology, surface fea-

tures, including pubescence, inflorescences, flower structures and

fruits. In chapter 8, Families of flowering plants I. Magnoliopsida

(Dicots), 193 families are dealt with, grouped into six subclasses

according to the Cronquist classification. This system was chosen

‘because students have detailed references for further information [and

because] it also provides an organized sequence to follow, whether or

not one agrees with the classification'. Again, most families are treated

on one page, about half of which is devoted to line drawings. The text

is also divided into paragraphs with bold headings from General

description to Fossil record. The descriptions are brief and, generally,

somewhat simplified. Chapter 9 deals similarly with 42 families of the

Liliopsida (Monocots). 10, History and development of classification,

discusses form systems of plants from the times of Theophrastus to

Camerarius, the sexual systems, with an outline of the 24 classes used

by Linnaeus, the early ‘natural’ systems from Adanson to Bentham and

Hooker, including an outline of de Jussieu’s classification, the post-

Darwinian ‘natural’ classification systems by Eichler, Engler,

Wettstein, Bessey (with some of his ‘dicta’) and Hutchinson. Under

contemporary classification systems (1950 to present) mainly the fol-

lowing are discussed: Takhtajan, Cronquist, Thome and Dahlgren. The

author concludes that their systems have more similarities than differ-

ences. He also states that ‘most younger botanists prefer to address less

speculative issues [than] developing classification schemes’. Under

Contemporary phenetic methods, he deals at some length with numer-

ical taxonomy, and four pages on contemporary phylogenetic methods

deal with cladistics and the Angiosperm Phylogeny Group. The chap-

ter ends with the ordinal cladogram of APG and the statement ‘How

well the new ideas will be accepted remains to be seen... yet, one sys-

tematic text has embraced the new ideas completely and applied them

to an instructional level (Judd et al. 1999: Plant systematics. A phylo-

genetic approach. Sinauer Assoc., Inc., Sunderland, MA). Seventeen of

the great classifiers are depicted, mostly by courtesy of the Royal

Botanic Gardens, Kew. Chapter 11, Literature of systematics, consists

largely of separate lists of literature on plant systematics, terminology

and dictionaries, specific and comprehensive guide books, botanical

journals (including South African Journal of Botany) and basic sys-

tematic texts. Chapter 1 2 deals with Contemporary views of the origin

of vascular plants. The next chapter, 13, Contemporary methods of

studying plants, is one of the highlights of the book. It consists mainly

of contributions by a number of experts on subjects such as plant anato-

my, morphology, palynology, systematics, molecular systematics,

cytology and genetics, biogeography and ecology, and the way they are

applied to plant systematics. Chapter 14, Preserving plant biodiversity,

discusses the value of plant species to humans, how species become

endangered, and our current knowledge about endangered plants. In

this chapter Small & Warwick, of Agriculture and Agri-Food, Canada,

discuss genetic engineering and biodiversity. Chapter 15 deals with The

role of botanical gardens in society and presents a well-illustrated

overview of the world’s leading botanical gardens, including Kirsten-

bosch, with a photo by Fiona Getliffe Norris. Appendix I lists selected

bibliographic references cited in the book, chapter by chapter.

Appendix II Floras of the world lists references by region. Southern

Africa is represented by Adamson & Salter’s Flora of the Cape Penin-

sula, Flora of Swaziland by Compton, Dyer’s Genera, the FSA, Merx-

miiller’s Prodromus, Trees of southern Africa by Palgrave, Flora of

Natal by Ross and two works by the arboriphilistic Van Wyks: Trees of

the Kruger National Park and Field guide to the wild flowers of the

Witwatersrand, both attributed to B. van Wyk. Who can blame a for-

eigner for getting confused, especially with the convertible initials?

Appendix III gives the classification by Cronquist. A glossary of some

1 5 pages, and the index complete the book, apart from a page about the

CD, and the CD itself in a plastic pocket, which contains over 4 700

images representing more than 270 families and 1 150 genera of vas-

cular plants. On average, I found the images informative, but not of a

particularly high standard. Most of the many line drawings in the book

I would assess similarly. They appear to be drawn with too thick a line

for their size, with the result that lines often flow into each other creat-

ing solid black areas. This is much less worrying in edition 2, where the

same drawings are printed the same size, but far neater.

I can imagine that this work, written with enthusiasm and a love of

nature, served and still serves as a source of inspiration for many stu-

dents brought up on the past editions and studying with the help of the

present one, and others to come (see p. 398). Unfortunately the many

blemishes prevent me from recommending the work unreservedly to

the uncritical reader.

REFERENCES

BRUMMITT, R.K. & POWELL, C.E. (eds). 1992. Authors of plant

names. Royal Botanic Gardens, Kew.

STAFLEU, F A. 1967. Taxonomic literature. International Bureau for

Plant Taxonomy and Nomenclature, Utrecht, The Netherlands.

STAFLEU, F.A. & COWAN, R.S. 1979. Taxonomic literature, edn 2,

vol. 2. Bohn, Scheltema & Holkema, Utrecht.

O.A. LEISTNER*

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

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BOTHALIA

Volume 31,1 May 200 1

CONTENTS

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

I.M. OLIVER 1

2. The genus Ehretia (Boraginaceae: Ehretioideae) in southern Africa. E. RETIEF and A.E. VAN WYK 9

3. Studies in the liverwort genus F ossombronia (Metzgeriales) from southern Africa. ll.L zuurberg-

ensis, a new species from Eastern Cape and new records for the area. S.M. PEROLD 25

4. Notes on African plants:

Amaryllidaceae. A new species of Cyrtanthus (Cyrtantheae) from the southern Cape, South

Africa. D.A. SNIJMAN .’ , ... . . 31

Amaryllidaceae. A new species of Brunsvigia (Amaryllideae) from Western Cape, South

Africa. D.A. SNIJMAN 34

Anthericaceae. The correct name in Chlorophytum for Anthericum longistylum. C. ARCHER

andS. KATIVU : 40

Asphodelaceae. Aloe greatheadii var. davyana (Alooideae), a new locality in KwaZulu-Natal,

South Africa. G.F. SMITH and N.R. CROUCH 31

Asphodelaceae. Jodrelliafistulosa, a new generic record for the Flora of southern Africa region.

C. ARCHER 39

Asteraceae. A new species of Oedera from Western Cape, South Africa. J.B.P. BEYERS 41

Asteraceae. A new species of Emilia (Senecioneae) from South Africa. J.C. MANNING and

P. GOLDBLATT 46

Commelinaceae. Rediscovery of Triceratella drummondii, and comments on its relationships

and position within the family. N.P. BARKER, R.B. FADEN, E. BRINK and A.P. DOLD . 37

Fabaceae. First distribution record for Brachystegia spiciformis (Caesalpinioideae) in South

Africa. P.J.H. HURTER and E. VAN WYK 43

Fossombroniaceae. F ossombronia nyikaensis, a new species from Malawi. S.M. PEROLD ... 48

Lamiaceae. Plectranthus hilliardiae subsp. australis, a new taxon from Eastern Cape, South

Africa. E.J. VAN JAARSVELD and A.E. VAN WYK 44

5. Leaf anatomy of the genus Passerina (Thymelaeaceae): taxonomic and ecological significance.

C.L. BREDENKAMP and A.E. VAN WYK 53

6. A lexicon of plants traded in the Witwatersrand umuthi shops, South Africa. V.L. WILLIAMS,

K. BALKWILL and E.T.F. WITKOWSKI 71

7. Thirty years of change in the fynbos vegetation of the Cape of Good Hope Nature Reserve, South

Africa. S.D.J. PRIVETT, R.M. COWLING and H.C. TAYLOR 99

8. Patterns of endemism within the Karoo National Park, South Africa. F. RUBIN, A.R. PALMER and

C. TYSON 117

9. Vegetation-environment relationships in a catchment containing a dambo in central Zimbabwe.

I. MAPAURE and M.P. McCARTNEY 135

10. Miscellaneous notes:

Poaceae. The basic chromosome number of the genus Pentameris (Arundinoideae). J.J. SPIES

and R. ROODT 145

11. Book reviews 147

Abstracted, indexed or listed in • AETFAT Index • AGRICOLA • AGRIS • BIOSIS: Biological Abstracts/RRM • CABS • CABACCESS • CAB

ABSTRACTS • ISI: Current Contents, Scisearch, Research Alert • Kew Record of Taxonomic Literature • Taxon: reviews and notices.

ISSN 006 8241

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