Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 28,1

May 1998

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

Obtainable from the National Botanical Institute, Private Bag XI 01, Pretoria 0001, Republic of

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

BOTHALIA

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

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

ed to the furtherance of botanical science. The main fields covered are taxonomy, ecology, anato-

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

lished annually.

Two booklets of the contents (a) to Vols 1-20 and (b) to Vols 21-25, 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 well known botanical artists have

contributed to the series, such as Cythna Letty (over 700 plates), Kathleen Lansdell, Stella Gower,

Betty Connell, Peter Bally, Fay Anderson, Ellaphie Ward-Hilhorst and Gillian Condy. 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 infor-

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

available:

Molteno Formation (Triassic) Vol. I . Introduction. Dicroidium, by J.M. & H.M. Anderson.

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

Anderson.

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

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

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 28,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

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2 Cussonia Avenue, Brummeria, Pretoria

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ISSN 0006 8241

May 1998

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

CONTENTS

Volume 28,1

New species and combinations in Bothalia 28,1 (1998) iv

1. Studies in the liverwort genus Fossombronia (Metzgeriales) from southern Africa. 5. A new species

from Northern and Western Cape. S.M. PEROLD 1

/ 2. A taxonomic revision of Maurocenia (Celastraceae), a Western Cape monotypic endemic. R.H.

ARCHER & A.E.VAN WYK 7

3. FSA contributions 10: Trapaceae. B. VERDCOURT 11

/ 4. A revision of Ledebouria (Hyacinthaceae) in South Africa. 1. Two new species. S. VENTER &

T.J. EDWARDS 15

5. Revised key to Ipomoea (Convolvulaceae) in southern Africa. W.G. WELMAN & A.D.J. MEEUSE . . 19

/ 6. Taxonomic notes on the genus Kliadia (Mesembryanthemaceae/Aizoaceae). P. CHESSELET, H.E.K.

HARTMANN, N. HAHN, P. BURGOYNE & G.F. SMITH 25

7. FSA contributions 1 1 : Zingiberaceae. R.M. SMITH 35

8. Notes on African plants:

y/ Asphodelaceae: Alooideae. Astroloba corrugata : description of a long-known species in a

southern African endemic alooid genus. N.L. MEYER & G.F. SMITH 60

V Asteraceae. A note on the Brachylaena discolor complex. P.P.J. HERMAN 42

w Aytoniaceae (Hepatophyta). Asterella abyssinica newly reported from South Africa and Malawi.

S.M. PEROLD ’. 53

Cyperaceae. A new combination in Isolepis. C. ARCHER 41

^ Fabaceae. A new species of Pearsonia from Mpumalanga, South Africa. G. GERMISHUIZEN . . 57

Fabaceae. A new species of Rhynchosia from South Africa. G. GERMISHUIZEN 58

Hyacinthaceae. A new species of O rnithogalum from the Richtersveld, South Africa. G. WIL-

LIAMSON 62

Metzgeriales-Fossombroniaceae. Fossombronia rwandaensis, a new species from tropical Africa.

S.M. PEROLD 45

i/ Oxalidaceae. Taxonomic delimitation of Oxalis engleriana. L.L. DREYER & A.E. VAN WYK . . 65

,/ Thymelaeaceae. A new species of Lachnaea endemic to the southeastern mountains of the Western

Cape. J.B.P. BEYERS 49

9. New combinations in Antimima (Ruschioideae. Aizoaceae) from southern Africa. H.E.K. HARTMANN 67

10. Meiotic chromosome behaviour in Cenchrus ciliaris (Poaceae: Panicoideae). N.C. VISSER, J.J.

SPIES & H.J.T. VENTER 83

11. Invasive alien woody plants of the southern and southwestern Cape region. South Africa. L. HEN-

DERSON 91

12. Miscellaneous notes:

New editor for Bothalia. G.F. SMITH 113

13. Obituaries:" John Denzil Carr (1916-1997). H.F. GLEN 117

Hermanus Phillipus van der Schijff (1921-1997). G.K. THERON 118

^ Michiel (Mike) Adriaan Niklaas Muller (1948-1997). GILLIAN MAGGS & G. GER-

MISHUIZEN 123

14. Book reviews 125

15. Errata in Bothalia 27,2 (1997) 129

New species and combinations in Bothalia 28,1 (1998)

Antimima addita (L.Bolus) H.E.K. Hartmann, comb, nov., 70

Antimima amoena (Schwantes) H.E.K.Hartmann, comb, nov., 70

Antimima androsacea (Marloth & Schwantes) H.E.K.Hartmann , comb, nov., 70

Antimima argentea ( L.Bolus ) H.E.K.Hartmann, comb, nov., 70

Antimima biformis (N.E.Br.) H.E.K.Hartmann, comb, nov., 70

Antimima bina (L.Bolus) H.E.K.Hartmann, comb, nov., 70

Antimima bracteata (L.Bolus) H.E.K.Hartmann, comb, nov., 70

Antimima brevicarpa ( L.Bolus ) H.E.K.Hartmann, comb, nov., 70

Antimima brevicollis ( N.E.Br. ) H.E.K.Hartmann , comb, nov., 71

Antimima buchubergensis (Dinter) H.E.K.Hartmann , comb, nov., 71

Antimima compacta ( L.Bolus ) H.E.K.Hartmann, comb, nov., 71

Antimima compressa (L.Bolus) H.E.K.Hartmann, comb, nov., 71

Antimima concinna (L.Bolus) H.E.K.Hartmann, comb, nov., 71

Antimima condensa (N.E.Br.) H.E.K.Hartmann, comb, nov., 71

Antimima crassifolia (L.Bolus) H.E.K.Hartmann, comb, nov., 71

Antimima dasyphylla (Schltr.) H.E.K.Hartmann, comb, nov., 71

Antimima defecta (L.Bolus) H.E.K.Hartmann, comb, nov., 71

Antimima dekenahi ( N.E.Br. ) H.E.K.Hartmann, comb, nov., 72

Antimima distans ( L.Bolus ) H.E.K.Hartmann, comb, nov., 72

Antimima dolomitica (Dinter) H.E.K.Hartmann, comb, nov., 72

Antimima eendornensis (Dinter) H.E.K.Hartmann, comb, nov., 72

Antimima elevata ( L.Bolus ) H.E.K.Hartmann, comb, nov., 72

Antimima emarcescens ( L.Bolus ) H.E.K.Hartmann, comb, nov., 72

Antimima erosa (L.Bolus) H.E.K.Hartmann, comb, nov., 72

Antimima evoluta (N.E.Br) H.E.K.Hartmann , comb, nov., 72

Antimima exsurgens (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima fenestrata (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima fergusoniae (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima gracillima ( L.Bolus ) H.E.K.Hartmann, comb, nov., 73

Antimima granitica (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima hallii (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima hamatilis (L.Bolus) H.E.K.Hartmann, comb, nov., 73

Antimima herrei ( Schwantes ) H.E.K.Hartmann, comb, nov., 73

Antimima intervallaris (L.Bolus) H.E.K.Hartmann, comb, nov., 74

Antimima ivori (N.E.Br.) H.E.K.Hartmann , comb, nov., 74

Antimima karroidea (L.Bolus) LI. E.K. Hartmann, comb, nov., 74

Antimima klaverensis (L.Bolus) H.E.K.Hartmann, comb, nov., 74

Antimima koekenaapensis (L.Bolus) H.E.K.Hartmann, comb, nov., 74

Antimima komkansica ( L.Bolus ) H.E.K.Hartmann, comb, nov., 74

Antimima lawsonii (L.Bolus) H.E.K.Hartmann, comb, nov., 74

Antimima leipoldtii (L.Bolus) H.E.K.Hartmann, comb, nov., 74

Antimima leucanthera ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima limbata ( N.E.Br. ) H.E.K.Hartmann, comb, nov., 75

Antimima lodewykii ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima loganii ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima lokenbergensis ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima luckhoffii (L.Bolus) H.E.K.Hartmann, comb, nov., 75

Antimima maleolens ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima maxwellii ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima menniei ( L.Bolus ) H.E.K.Hartmann, comb, nov., 75

Antimima mesklipensis (L.Bolus) H.E.K.Hartmann, comb, nov., 76

Antimima meyerae (Schwantes) H.E.K.Hartmann, comb, nov., 76

Antimima minima (Tischer) H.E.K.Hartmann, comb, nov., 76

Antimima minutifolia (L.Bolus) H.E.K.Hartmann, comb, nov., 76

Antimima modesta (L.Bolus) H.E.K.Hartmann, comb, nov., 76

Antimima mucronata (Haw.) H.E.K.Hartmann, comb, nov., 76

Antimima mutica ( L.Bolus ) H.E.K.Hartmann, comb, nov., 76

Antimima nobilis (Schwantes) H.E.K.Hartmann, comb, nov., 77

Antimima nordenstamii (L.Bolus) H.E.K.Hartmann, comb, nov., 77

Antimima oviformis (L.Bolus) H.E.K.Hartmann, comb, nov., 77

Antimima papillata (L.Bolus) H.E.K.Hartmann, comb, nov., 77

iv

New species and combinations in Bothalia 28,1 (1998) (cont.)

Antimima paucifolia (L.Bolus) H.E.K. Hartmann, comb, nov., 77

Antimima pauper (L.Bolus) H.E.K. Hartmann, comb, nov., 77

Antimima peersii (L.Bolus) H.E.K. Hartmann, comb, nov., 77

Antimima perforata (L.Bolus) H.E.K. Hartmann, comb, nov., 77

Antimima persistens (L.Bolus) H.E.K. Hartmann, comb, nov., 78

Antimima pilosula (L.Bolus) H.E.K. Hartmann, comb, nov., 78

Antimima piscodora (L.Bolus) H.E.K. Hartmann, comb, nov., 78

Antimima prolongata (L.Bolus) H.E.K. Hartmann, comb, nov., 78

Antimima propinqua (N.E.Br.) H.E.K. Hartmann, comb, nov., 78

Antimima prostrata (L.Bolus) H.E.K. Hartmann, comb, nov., 78

Antimima pumila (Fedde & Schuster) H.E.K. Hartmann, comb, nov., 78

Antimima pusilla (Schwantes) H.E.K. Hartmann, comb, nov., 79

Antimima pygmaea (Haw.) H.E.K. Hartmann, comb, nov., 79

Antimima quarzitica ( Dinter) H.E.K. Hartmann, comb, nov., 79

Antimima roseola (N.E.Br.) H.E.K. Hartmann, comb, nov., 79

Antimima saturata (L.Bolus) H.E.K. Hartmann, comb, nov., 79

Antimima saxicola (L.Bolus) H.E.K. Hartmann, comb, nov., 79

Antimima schlechteri (Schwantes) H.E.K. Hartmann, comb, nov., 79

Antimima simulans (L.Bolus) H.E.K. Hartmann, comb, nov., 79

Antimima sobrina (N.E.Br.) H.E.K. Hartmann, comb, nov., 80

Antimima solida (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima stayneri (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima stokoei (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima subtruncata (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima triquetra (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima tuberculosa (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima turneriana (L.Bolus) H.E.K. Hartmann, comb, nov., 80

Antimima vanzylii (L.Bolus) H.E.K. Hartmann, comb, nov., 81

Antimima varians (L.Bolus) H.E.K. Hartmann, comb, nov., 81

Antimima ventricosa (L.Bolus) H.E.K. Hartmann, comb, nov., 81

Antimima verruculosa (L.Bolus) H.E.K. Hartmann, comb, nov., 81

Antimima watermeyeri ( L.Bolus ) H.E.K.Hartmann, comb, nov., 8 1

Antimima wittebergensis (L.Bolus) H.E.K.Hartmann, comb, nov., 81

Astroloba corrugata N.L.Mey. & G.F.Sm., sp. nov., 61

Fossombronia cederbergensis Perold, sp. nov., 1

Fossombronia rwandaensis Perold , sp. nov., 45

Isolepis leucoloma (Nees) C. Archer, comb, nov., 42

Lachnaea sociorum Beyers, sp. nov., 49

Ledebouria atrobrunnea S. Venter sp. nov., 15

Ledebouria dolomiticola S.Venter, sp. nov., 16

Ornithogalum decus-montium G.Will., sp. nov., 63

Pearsonia hirsuta Germish., sp. nov., 57

Rhynchosia atropurpurea Germish., sp. nov., 58

v

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Bothalia 28,1: 1-5 (1998)

Studies in the liverwort genus Fossombronia (Metzgeriales) from

southern Africa. 5. A new species from Northern and Western Cape

S.M. PEROLD*

Keywords: Fossombronia. F. cederbergensis. Hepaticae, Metzgeriales, new species, southern Africa

ABSTRACT

A new species of Fossombronia. F. cederbergensis, from the winter rainfall area of the Northern and Western Cape is

described It is charactenzed by ‘ruched’ leaves that are mostly wider than long, by tuberous stem apices that enable the

plants to survive the hot, dry summers and by completely or incompletely reticulate spores.

Fossombronia cederbergensis Perold, sp. nov.

Plantae prostratae, crebrae in coloniis. Folia imbrica-

ta. valde convoluta, forma irregulari, plerumque superne

lobata, valde latiora quant longiora. Rhizoidea ad apicem

ventralem caulis interdum hyalina, proximaliter pur-

purea. Caules apicem versus tuberosi. Dioicae. Plantae

masculae rarissimae, parvulae; antheridia bracteis irregu-

lariter formatis tecta. Plantae femineae maiores, pseudo-

perianthio campanulato, orificio undulato, subplicato vel

partim reflexo. Sporae 37.5-50.0 pm diametro, orna-

mentis variis, superficie distali cum reticulo 11 vel 12

areolulis trans diametrum, sed saepe imperfecte reticula-

ta; lamellis tenuibus, interdum subsinuatis, plerumque in

nodis prominentibus; superficie proximali cum nota tri-

radiata plerumque indistincta vel carenti, cum cristis bre-

vibus irregularibus tenuibus vel pergrossis ramosis.

Elateres 140-250 x 7.5-10.0 pm. extremitates versus

decrescentes, ±5 pm lati, bis vel ter spirales in centro,

interdum leviter papillosi.

TYPE. — Northern Cape, 3219 (Wuppertal): 2 km

south of Algeria Forest Station, at sandstone rock out-

crop, on sandy soil, (-AC), S.M. Perold 2359 (PRE,

holo.). Western Cape, 3218 (Clanwilliam): 21 km N of

Citrusdal, on sandstone rock outcrops above Olifants

River, Hex River Estates, (-BD), S.M. Perold 2380

(PRE. para.).

Plants in dense mats, fresh apical growth generally

entirely green, except for some smaller young leaflets

which are often stained throughout with various shades

of purple, otherwise only leaf margins and scattered inte-

rior cells purple, proximal leaves frequently dying,

becoming translucent and crumpled, sometimes only

their basal remnants remaining in 2 oblique lines forming

a row of disconnected ‘Vs’ dorsally along stem; shoots

simple (Figure 2A) or once/twice furcate, branches

sometimes developing from lateral buds; male plants

very rare, rather small, up to 3.5 mm long, 0.75 mm high,

± 1.3 mm wide; female and sterile plants 9-13 mm long,

1.5 mm high, 1.75 mm wide, successive years’ growth

from apex of old stem, if branched, terminal segments

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

MS. received: 1997-05-15.

4- 5 mm long, narrowly (Figure 2B) to moderately diver-

gent. Stems prostrate, green, but sometimes dorsally pink

or purple toward swollen apex, in cross section plano-

convex, in male plants (Figure 1M) 280 pm (11 cell

rows) high, 500 pm wide, becoming strongly arched and

tapering toward base, 250 pm high, 400 pm wide; in

female plants apically (Figure IN) 370 pm (13 cell rows)

high, 580 pm wide, basally (Figure 10) 250 pm high,

320 pm wide. Rhizoids at apex of stem sometimes all

hyaline, then becoming mixed with purple rhizoids,

proximally all rhizoids purple, until next apex, giving

rise to new growth, where some are again hyaline,

12.5-20.0 pm wide. Leaves in male plants overlapping,

spreading, succubously inserted on stem, irregularly

shaped, ‘ruched’ above (Figure 1A-E), 600-1000 x

650-850 pm, margins with ± 4 unicellular slime papillae,

20.0- 22.5 x 15.0-17.5 pm, mostly along sides; in female

plants leaves (Figure 1F-J) densely overlapping, very

frilly (Figure 2C), shape irregular, generally shorter than

wide, 575— 700(— 1 625) x 650-1 375(-l 875) pm, often

lobed above, lobes 250-320 x 280-450 pm, margins

with up to 7 well-spaced papillae. Leaf cells not appre-

ciably different in male and female plants, thin-walled, at

upper margins subquadrate to rectangular across (Figure

IK), 25-45 x 27.5-45.0 pm, at lateral margins long-rec-

tangular, 22.5-72.5 x 20.0-52.5 pnV, upper laminal cells

5- or 6-sided, 35-70 x 37.5-42.5 pm; middle laminal

cells 50-75 x 30^45 pm; basal cells 45.0-87.5 x 30-45

pm. Oil bodies in young leaves (Figure 1L) smooth,

round, 7-19 per cell, variable in size, from minute to 2.5

pm diam.; chloroplasts oval or round, 2. 5-5.0 pm diam.

Dioicous. Antheridia dorsal on stem, in a row, short-

stalked, globose, ± 230 pm diam., mostly soon shed;

bracts (Figure 1P-R) remaining, shape irregular,

575-700 x 650-1375 pm, with up to 3 finger-like

processes and 2 or 3 marginal papillae, cells in interior

30.0- 37.5 x 25.0-27.5 pm, basal cells 47.5-60.0 x

30.0- 32.5 pm. Archegonia well spaced, in 1 or 2 irregu-

lar rows dorsally along stem (Figure 2D), 250-370 pm

long, mostly obscured by strongly ‘ruched’ leaves.

Pseudoperianth (Figures IT, U; 2E, F) campanulate,

proximal to stem apices, 1375-1625 pm long, projecting

± 875 pm above leaves, from narrow base widely flaring

above, 1950-2575 pm wide across mouth, margin undu-

lating to somewhat plicate or partly reflexed, occasional-

ly with winged outgrowth at side; cells comparable in

2

Bothalia 28,1 (1998)

Bothalia 28,1 (1998)

3

FIGURE 2. — Fossumbrunia cederbergensis. A, arched terminal segment from side, fresh young growth at apex; B, shoot with two narrowly diver-

gent terminal segments; C, very frilly leaves in female plant; D, archegonia in 2 rows on stem, between leaves. E, F, pseudoperianth: E,

from side; F, from above. A, D, Koekemoer 1209: B, C, E, F, S.M Perold 2359. A, x 18; B, x 17; C, x 83; D, x 16; E, x 31; F, x 22.

shape and size to those of leaves, except for rather longer,

thin-walled basal cells, 90.0-137.5 x 37.5^47.5 pm.

Capsules globose, ± 700 pm diam., wall bistratose, cells

of inner layer polygonal, 25.0-42.5 x 25-35 pm, each

cell wall (Figure IV) with 1-3 brown nodular and occa-

sionally semi-annular thickenings. Seta 1.4-5. 5 mm

long, ± 170 pm or 10 cell rows diam. (Figure IS). Spores

light to dark brown, hemispherical, (37.5-)40.0-47.5

(-50.0) pm diam., including spines projecting at periph-

ery (Figure 3D); distal face convex, ornamentation with

a mesh of 1 1 or 12 small, up to 5 pm wide, complete are-

olae (Figure 3C) across diam., but often incompletely

reticulate (Figure 3A, B), lamellae thin, sometimes

slightly sinuous, mostly raised at the nodes; proximal

face with triradiate mark (Figure 3E) generally indistinct

or lacking, with short, irregular, fine to very coarse,

FIGURE 3. — Fossombronia cederbergensis. A-E, spores: A-C, distal face; D, detail of areolae on distal face; E, proximal face. F, elater. A, B, E,

S.M. Perold 2359\ C, D, F, S.M. Perold 2380. A, x 771 ; B, x 990; C, x 693; D, x 1560; E, x 918; F. x 994.

4

Bothalia 28,1 (1998)

FIGURE 4. — The distribution of F. cederbergensis in Northern and

Western Cape.

branched ridges, sometimes a few papillae present in

between, around spore periphery ± 25 low spines pro-

jecting ± 2.5 pm, occasionally more, (but difficult to

count their number as only some seen ‘end-on’), others

with sides of lamellae showing. Elaters (Figure 3F) yel-

low-brown, 140-250 x 7.5-10.0 pm, tapering to tips, ± 5

pm wide, bi- or sometimes trispiral in centre, occasion-

ally finely papillose.

F ossombronia cederbergensis has been named for the

Cederberg, where I first collected it. The species is

known from several localities in the winter rainfall areas

of Northern and Western Cape (Figure 4), with vegeta-

tion types. Northwestern Mountain Renosterveld and

Mountain Fynbos (Low & Rebelo 1996). The plants

often grow on slighly acid, rather sandy soil overlying

sandstone outcrops, but sometimes on fine-grained,

moisture retaining clay, at roadsides, dried seeps, stream

banks or pond margins. Of the 12 specimens referred

here, only three had spores, and they show rather wide

variation in the ornamentation, from completely reticu-

late in S.M. Perold 2380 to incompletely reticulate in

S.M. Perold 2359 and Powrie 182. The specimens that

lack spores could be identified by vegetative characters.

The swollen tuberous apices of the stems help the plants

to survive the hot, dry summers. In some plants up to

three seasons’ growth could be detected. A sample of the

specimen Koekemoer 1209, quickly resumed growth

from the stem apices, nine months after collection, when

kept wet in a closed plastic container for a few days.

The species can be distinguished by the ‘ruched’,

lobed leaves, which are mostly wider than long, the

tuberous stem apices, the completely or incompletely

reticulate spores and by the occasional to frequent pres-

ence of some hyaline rhizoids. The tuberous apices of the

stems bear some resemblance to those of F. spinifolia

(Perold 1997b), which has markedly dentate leaves in the

male plants, spores with 6 or 7 short irregular ridges, and

which is only known from Genadendal.

Scott & Pike (1987) described many new species from

Australia, one of which, F. tesselata, has spores similar-

ly reticulate to those of F. cederbergensis, but they are a

deep maroon, rather smaller at 30-41 pm diam. and the

elaters are mostly shorter than those of the southern

African species; its rhizoids are crimson throughout. F.

vermiculata with purple and hyaline rhizoids mixed on

the same stem was newly described by Scott & Pike

(1984) and in the same publication they mention that in

F. scrobiculatci the rhizoids are hyaline on most plants

but crimson on some, whereas in F. punctata they state

that the rhizoids are ‘(? always) hyaline, not crimson’. A

species recently described from southern Africa, F.

straussiana (Perold 1997a), always has hyaline or

brownish rhizoids and in the European species, F. hus-

notii, the rhizoids are characteristically hyaline, although

Scott & Pike (1988) claim that they are not all so.

Whether having purple and hyaline rhizoids mixed on the

same stem is really of taxonomic significance is not yet

known. However, failure to observe the presence of hya-

line or pale brown rhizoids ‘may cause confusion’ (Paton

1973). On some spore similarities, but rather poorly por-

trayed in their figures, Scott & Pike (1988: figs 13, 14)

have placed F. hamato-hirta Steph. which has ‘radicellis

pallidis’ (Stephani 1894) in synonymy under F. won-

draczekii which has purple rhizoids. Perhaps this should

be investigated further. The spore micrographs which I

took of the type and only known specimen of F. hamato-

hirta , H.I. Gordon 108 (G) from Ascension Island, bear

quite a close resemblance to those of F. straussiana and

as mentioned above, both have hyaline rhizoids. It is,

however, not possible to make a definite decision on their

synonymy with the available material of F. hamato-hirta.

SPECIMENS EXAMINED

Held at PRE, unless otherwise indicated.

Koekemoer 1209.

S.M. Perold 2093, 2197 (with Riccia villosa Steph ), 2323, 2359*

(holotype), 2364. 2380 (paratype)*, 3336 (with F. leucoxantha Lehm),

3339 (with F. leucoxantha Lehm.), 3355 (with F. leucoxantha Lehm.).

Perold, Koekemoer & Smook 3044. E. Powrie 182* (BOL).

* sporulating specimens.

ACKNOWLEDGEMENTS

I sincerely thank Dr H. Stieperaere (BR) for referee-

ing this article and his helpful suggestions; also my col-

leagues at NBI for their help with fieldwork and Dr H.F.

Glen for translating the diagnosis into Latin as well as

suggesting the specific epithet for the new species F.

cederbergensis. The curators of BOL and G are thanked

for the loan of specimens, and I extend my gratitude to

Ms G. Condy for the drawings, Mrs Romanowski for

developing and printing the photographs and to Ms D.

Maree for typing the manuscript.

REFERENCES

LOW, A B & REBELO, A. 1996. Vegetation of South A frica, Lesotho

and Swaziland. Dept. Environmental Affairs & Tourism, Pretoria.

PATON, J.A. 1973. Taxonomic studies in the genus Fossombronia

Raddi. Journal of Bryology 7: 243-252.

PEROLD, S.M. 1997a. Studies in the liverwort genus Fossombronia

( Metzgeriales) from southern Africa. I Three new species from

Northern Province, Gauteng and Mpumalanga. Bothalia 27:

17-27.

Bothalia 28,1 (1998)

5

PEROLD, S.M. 1997b. Studies in the liverwort genus Fossombronia

(Metzgeriales) from southern Africa. 3. An amendment to F.

spinifolia. Bothalia 27: 39-42.

SCOTT. G.A.M. & PIKE. D C. 1984. New species of Fossombronia

from Australia. Journal of the Hattori Botanical Laboratory 56:

339-349.

SCOTT, G.A.M. & PIKE, D.C. 1987. Studies on Fossombronia in

Australia II. Fourteen more new species. Journal of the Hattori

Botanical Laboratory 62: 367-386.

SCOTT, G A M. & PIKE, DC. 1988. Revisionary notes on

Fossombronia. The Biyologist 91: 193-201.

STEPHANI, F. 1894. Hepaticarum species novae. V. Hedwigia 33: 1-11.

Bothalia 28,1: 7-10 (1998)

A taxonomic revision of Maurocenia (Celastraceae), a Western Cape

monotypic endemic

R.H. ARCHER* and A.E. VAN WYK**

Keywords: Cassinoideae, Celastraceae, Maurocenia Mill., taxonomy. South Africa

ABSTRACT

A taxonomic account is given of the monotypic genus Maurocenia Mill. Maurocenia frangula Mill, has a restricted

range and is endemic to the Cape Peninsula and the West Coast National Park, Western Cape. Maurocenia frangularia (L.)

Mill., the species name and author citation widely used in the past, is incorrect. It is characterized by, among others, pen-

dulous ovules and gynodioecy, rare states in the Celastraceae. Maurocenia is apparently most closely related to the south-

ern African genus Lauridia Eckl. & Zeyh.

INTRODUCTION

The genus Maurocenia is fairly similar in many

anatomical and morphological features to Lauridia Eckl.

& Zeyh. (Archer & Van Wyk 1997). However, the pen-

dulous ovules in Maurocenia set it apart from the other

genera in the southern African Cassinoideae, which have

erect ovules (Archer & Van Wyk 1996). The occurrence

of pendulous ovules is relatively rare in the Celastraceae.

Perhaps the genus is, on account of its ovule orientation,

more closely related to the tropical American Tetrasiphon

Urb. and Gyminda Sarg. (Loesener 1942; Brizicky 1964)

than to the remaining southern African genera.

Maurocenia frangula was evidently not uncommon in

cultivation in Europe in the 18th century, in view of the

abundance of illustrations of the species. It was first

introduced in 1690 to the Royal Botanic Gardens in

London (Aiton 1811). In the pre-Linnaean period the

species was described and illustrated by Plukenet (1691),

Petiver (1702). Boerhaave (1710) and Dillenius (1732).

The genus and phrase name of Dillenius (1732),

Maurocenia frangula sempervirens, folio rigido subro-

tundo was taken up by Linnaeus (1737, 1743). However,

in 1753 Linnaeus reduced Maurocenia to synonymy

under Cassine L. The name Maurocenia was subse-

quently validly published when the genus was reinstated

by Miller (1754), in an attempt to rectify what he thought

was an error on Linnaeus’s part. The epithet frangula

was provided later (Miller 1768). being taken up from

the phrase name. Willdenow (1798) first introduced the

incorrect form of the epithet, frangularia, which is today

in general use in herbaria and in most recent literature.

The author citation in general use for the taxon, i.e. (L.)

Mill. (Arnold & De Wet 1993), is also incorrect.

Maurocenia Mill., The gardener’s dictionary

[abridg. edn 4, 1754]: 859, facs. edn, Cramer (1969);

Adans.: 303 (1763), as Maurocena ; Sond.: 465 (1860);

Kuntze; 147 & 149 (1891) p.p., excl. subg. Triceros Kuntze;

Thonner: 331 (1915); Davison; 343 (1927); Loes.: 179

(1942); R.A.Dyer: 335 (1975). Type: M. frangula Mill.

* National Botanical Institute, Private Bag X 101, 0001 Pretoria.

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

versity of Pretoria, 0002 Pretoria.

MS. received: 1997-04-23.

Maurocenia Mill. subg. Eumaurocenia Kuntze: 149

(1891).

Small evergreen, gynodioecious shrub to spreading

tree up to 8 m high, unarmed, glabrous, without elastic

threads in leaves and bark; bark greyish with layers of

powdery yellow pigment in exposed rhytidome, exfoliat-

ing in thin scales, surface cracked in grid-like pattern.

Branchlets terete or slightly flattened, greyish brown to

brown. Leaves opposite; lamina widely elliptic to circu-

lar, dark green above, grey-green below, (25— )35— 60

(-85) x (20-)30-55(-65) mm, base rounded to cuneate,

apex rounded to retuse, margin entire, revolute, coria-

ceous and rigid; venation slightly raised above and more

prominent below, reticulation inconspicuous, brochido-

dromous, midrib and petiole very conspicuous below

owing to whitish green colour contrasting with grey-

green of rest of lamina in dried leaves; petiole 2-3 mm

long; stipules minute, ± 1 mm long, ± triangular, brown-

ish black, marcescent. Inflorescences sessile, ± irregular-

ly dichasial, axillary towards apices of branchlets, 4-10-

flowered; bracts minute. Flowers often unisexual with

staminodes (female), or bisexual, pentamerous, ± 2 mm

diam.; pedicels 2-4 mm long. Sepals rudimentary, ± 0.3

mm long. Petals white, ovate, 1 .3 x 1.0 mm, sessile, apex

rounded, margin ± deflexed. Stamens ± erect; inserted

below margin of disc; filaments of perfect flowers 3-4

mm long, well exserted above flowers; filaments of

female flowers rudimentary (staminodes); anthers of per-

fect flowers 0.8 mm long, basifixed, introrse, dehiscing

by longitudinal slits. Disc fleshy, entire, undulate. Ovary

± V3 immersed in and adnate to disc, 2(3)-locular with 2

pendulous collateral ovules per locule, ovules epi-

tropous, dorsal; styles ± 0.25 mm long; stigma in female

flowers often conspicuously 2-lobed, papillate, stigma

inconspicuous in perfect flowers. Fruit baccate, pale red,

spheroid, 7-10 mm diam., mesocarp fleshy. Seed 1 or 2

per locule (up to 5 per fruit recorded), brown, ± spheroid,

post-chalazal vascular bundles not observed, fleshy

endosperm present; embryo erect, with cotyledons

fleshy, widely elliptic, base unequal (Figure 1).

Maurocenia frangula Mill., The gardener's dic-

tionary edn 8.: no page number (1768); Willd.: 1493

(1798), in syn. as frangularia', Pers.: 327 (1805);

Marloth: 153 (1925); Davison: 343 (1927); Loes.: 179

Bothalia 28,1 (1998)

AMSTfcusP-

FIGURE 1. — Maurocenia frangula.

A, branch with female flow-

ers, x 0.7; B, female flower, x

7; C, branch with perfect

flowers, x 0.7; D, perfect

flower, x 7. Drawn from

Marloth 5978 (PRE) by Anne

Stadler.

(1942); Adamson: 567 (1950); Levyns: 190 (1966); Von

Breitenbach: 640 (1965); Palmer & Pitman: 1324

(1973); Coates Palgrave: 515 (1977); Bond & Goldblatt:

224 (1984). Type: Dillenius, Hortus Eltham. t. 146, f.

147 (1732), lecto., here designated.

Cassine maurocenia L.: 269 (1753); 385 (1762); Willd. : 1493

(1798); Thunb.: 52 (1794); Thunb.: 225 (1818); Thunb.: 268 (1823);

W.T.Aiton: 170 (1811); Roem. & Schult.: 466 (1819); Spreng.: 939

(1824); Eckl & Zeyh.: 128 (1834/5); Hook.: t. 552 (1843); Sond.: 465

(1860); Hook.f.: 363 (1862); Szyszyl.: 33 (1888).

leones: Pluk : t 158, f. 2 (1691); Petiver: t. 57, f. 4 (1702), Boerh.:

244(1727); Dill.: t 146, f. 147 (1732); Hook.: t. 552 (1843); Marloth:

t 51 (1925); Davison: t. 19 (1927); Von Breitenbach: 641 (1965).

Maurocenia is a monotypic genus of mountain kloofs

and coastal bush on rocky sea shores in the Cape

Peninsula and near Stellenbosch (Figure 2). M. frangula

is a gynodioecious shrub or tree, frequently encountered

along the footpaths in the kloofs of Table Mountain. The

wood is hard and has a fine grain, and has formerly been

used in the manufacturing of musical instruments

(Pappe 1854). Vernacular names are ‘Hottentot’s cherry’

and ‘Hottentotskersie’, referring to the fruit which is edi-

ble and was eaten by Khoikhoin (Hottentot). The genus

name commemorates the Italian horticulturist, F.

Mauroceni.

Although the species is described here as gynodioe-

cious, it may well be dioecious. Further study is required

to confirm the functional sexuality of the flowers. The

long slender stamens would suggest an adaptation to

wind pollination which is supported by the almost com-

plete absence of sepals and deflexed petals. However,

Marloth (1925) described the flowers as sweetly scented,

visited by flies and bees.

Bothalia 28,1 (1998)

9

Specimens examined

WESTERN CAPE. — 3318 (Cape Town): West Coast National

Park, Postberg Peninsula, (-AA), Van Wyk BSA 142 (PRU); 'Montis

tabularis’ [Table Mountain], (-CD), Marloth 97 (PRE); Camps Bay,

(-CD), Marloth 5978 (PRE), Maude s.n. (BM), Prior PRE-47256

(PRE, Z); Table Mountain, Grotto Ravine, (-CD), Marloth 12044

(PRE); Stellenbosch, ridge behind Swartboskloof, adjoining Haelkop,

(-DD), Taylor 7301 (PRE). 3418 (Simonstown): Muizenberg, near

False Bay, (-AB), MacOwan 580 (BM, BOL, PRE, Z); slopes above

Bakoven Bay, (-AB), Marais 601 (BOL, PRE): Blackburn Kloof near

Hout Bay, (-AB), Marais 606 (PRE); Jagersfontein, (-AB), Pole Evans

4349 (PRE); Hout Bay, (-AB), Rehmann 1609 (Z), Smuts 1154 (PRE);

Boyes Drive, opp. St James, between Muizenberg and Kalkbaaiberg,

(-AB), Stauffer & Esterhuysen 5123 (PRE, Z); boulder near summit of

Steenberg, (-AB), Taylor 3296 (PRE); Cape of Good Hope Nature

Reserve, Booiseskerm, (-AB), Taylor 10531 (C. PRE); Cape

Peninsula, West Coast opposite Antipolis shipwreck, (-AB), Van

Jaarsveld 3154 (PRE); Chapmans Peak, (-AB), Van Jaarsveld 3171b

(PRE); Hout Bay, (-AB). Wolley Dod 1045 (BM); Simonstown,

Smitswinkelbaai, (-AD), Schlechter 700 (Z); eastern side of Hangklip

Peak, facing Blesberg, (-BD), Rourke 1855 (NBG, PRE).

Species excluded or insufficiently known

M. americana Mill. (1768)

M. arguta (Lindl.) Kuntze: 149 (1891) = Turpinia sp.

(Staphyleaceae)

M. cerasus Mill. (1768)

M. cochinchinensis (Lour.) Kuntze: 150 (1891) = Tur-

pinia sp.

M. heterophylla (Ruiz & Pav.) Kuntze: 150 (1891) = Tur-

pinia sp.

M. insignis (Kuntze) Kuntze: 149 (1891) = Turpinia sp.

M. occidentalis (Sw.) Kuntze: 150 (1891) = Turpinia sp.

M. phylliraea Mill. (1768)

M. pinnata (Schiede ex Schltdl.) Kuntze: 150 (1891) =

Turpinia sp.

M. pomifera (Roxb.) Kuntze: 147 & 149 (1891) =

Turpinia sp.

M. schinziana Loes.: 194 (1894) = Cassine parvifolia

Sond.

M. simplicifolia (Gardner & Champ.) Kuntze: 149 (1891) =

Turpinia sp.

M. sphaerocarpa (Hassk.) Kuntze: 147 & 150 (1891) =

Turpinia sp.

M. zollingeri Kuntze: 147 (1891) = Turpinia sp.

REFERENCES

ADAMSON, R.S. 1950. Celastraceae. In R.S. Adamson & T.M. Salter,

Flora of the Cape Peninsula : 564-568. Juta, Cape Town.

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

AITON, W.T. 1811. Hortus Kewensis, 2nd edn, Vol. 2. Longman,

London.

ARCHER, R.H & VAN WYK, A.E. 1996. Generic delimitation of sub-

family Cassinoideae (Celastraceae) in Africa. In L.J.G. van der

Maesen, X.M. van der Burgt & J.M. Medenbagh de Rooy, The

biodiversity of African plants: 459-463. Kluwer Academic

Publishers, The Netherlands.

ARCHER, R.H. & VAN WYK, A.E. 1997. A taxonomic revision of

Lauridia Eckl. & Zeyh. (Cassinoideae: Celastraceae). South

African Journal of Botany 63: 227-232.

ARNOLD, T.H. & DE WET, B.C (eds). 1993. Plants of southern

Africa: names and distribution. Memoirs of the Botanical

Survey of South Africa No. 62: 1-825.

BOERHAAVE, H. 1710. Index plantarum. Reprint 1728. Boutestein,

Leiden.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Flora. Journal

of South African Botany , Suppl. Vol. 13.

BRIZICKY, G.K. 1964. The genera of Celastrales in the southeastern

United States. Journal of the Arnold Arboretum 45: 206-234.

COATES PALGRAVE, K. 1977. Trees of southern Africa, 1995

impression. Struik, Cape Town.

DAVISON, J.D. 1927. Celastraceae. Bothalia 2: 289-346.

DILLENIUS, J.J 1732. Hortus elthamensis seu plantarum rariorum.

London.

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

Vol. 1 . Department of Agricultural Technical Services, Pretoria.

ECKLON, C.F. & ZEYHER, K.L.P. 1834/1835. Enumeratio plantarum

Africae australis extratropicae. Pars I. Perthes & Besser,

Hamburg.

HOOKER, J.D. 1862. Celastnneae. In G Bentham & J.D Hooker,

Genera plantarum 1: 357-371. Reeve, London.

HOOKER, W.J. 1843. leones plantarum 6. Bailliere, London.

KUNTZE, O. 1891 . Revisio generum plantarum. Pars 1. Felix, Leipzig.

LEVYNS, MR B 1966. A guide to the flora of the Cape peninsula,

2nd edn. Juta, Cape Town.

LINNAEUS, C. 1737. Hortus cliff ortianus. Amsterdam.

LINNAEUS, C. 1743. Genera plantarum, 3rd edn. David, Paris.

LINNAEUS, C. 1753. Species plantarum, 1st edn. Impensis Laurentii

Salvii, Holmiae.

LINNAEUS, C. 1762. Species plantarum, 2nd edn Impensis direct.

Laurentii Salvii, Holmiae.

LOESENER, L.E.T. 1942. Celastraceae. In A. Engler, H. Harms & J

Mattfeld, Die natiirlichen Pflanzenfamilien, 2nd edn, 20b:

87-197. Duncker & Humblot, Berlin.

MARLOTH, R. 1925. The Flora of South Africa, Vol. 2. Darter Bros.,

Cape Town.

MILLER, P. 1754. The gardener's dictionary [abndg. edn 4], facs. edn:

859 (1969). Cramer, London.

MILLER, P. 1768. The gardener’s dictionary, 8th edn. London.

PALMER, E. & PITMAN, N. 1973. Trees of South Africa, Vol. 2.

Balkema, Cape Town.

PAPPE, L. 1854. Silva capensis. Van De Sandt & De Villiers, Cape Town.

PERSOON, C.H. 1805. Synopsis plantarum Part 1 Cramerum, Paris

& Cottam, Tubingen.

PETIVER, J. 1702. Gazophylacii naturae & artis decas. London.

PLUKENET, L. 1691. Phytographia, Part 1. London.

ROEMER, J.J. & SCHULTES, J.A. 1819. Systema vegetabilium 5.

Cottae, Stuttgardtiae.

SONDER, O.W. 1860. Celastnnea;. In W.H. Harvey & O.W Sonder,

Flora capensis 1: 451-472. Hodges, Smith, Dublin.

SPRENGEL, K.P.J. 1824. Systema vegetabilium 1. Gottingae [Gottingen],

SZYSZYLOWITZ, I 1888. Polypetalae disciflorae rehmanianae.

Universatatis Jagellonicae.

THONNER, F. 1915. The flowering plants of Africa. Dulau, London.

10

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THUNBERG, C.R 1794. Prodromus plantarum capensium. Part 1. THUNBERG, C.P. 1823. Flora capensis, 2nd edn. Schultes, Stuttgard

t * i it VOM RRPITPNRAfH F 1Q6S Thp inrli ppumis trpp.x of xnutheri

Edman, Uppsala.

THUNBERG, C.P. 1818. Flora capensis , 1st edn. Gerhardum

Bonnierum, Hafniae.

VON BRE1TENBACH, F. 1965. The indigenous trees of southern

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W1LLDENOW, C.L. 1798. Species plantarum 1,2. Nauk, Berlin.

Bothalia 28,1 : 11-14 (1998)

FSA contributions 10: Trapaceae

B. VERDCOURT*

Aquatic floating herbs. Leaves dimorphic: floating

leaves alternate, in rosettes, only present at upper nodes,

petiole spongy and more or less inflated about the mid-

dle; lamina rhombic to deltoid, the upper margins

toothed, stipules small, scarious, divided to the base and

thus apparently more than two per leaf; submerged

leaves opposite, sessile, linear, entire, caducous. Ad-

ventitious roots developing from leaf scars of submerged

leaves, containing chlorophyll, pinnatisect into many fil-

iform segments; in those from upper nodes, segments

shorter or absent. Flowers hermaphrodite, regular, 4-

merous, solitary, from upper axils, pedunculate. Sepals 4.

Petals 4, white. Stamens 4, perigynous. Ovary half-infe-

rior. bilocular; ovules pendulous, 1 per locule; stigma

capitate. Fruit a 1 -seeded, top-shaped drupe; pericarp

soon disappearing; endocarp hard, variously 2-4-horned;

horns derived from persistent sepals; testa woody;

cotyledons very unequal, 1 almost vestigial and not de-

veloping.

A single recent genus often included in the

Onagraceae and seemingly closely related but with very

distinct morphology. Miki (1959) has suggested deriva-

tion from Lythraceae and Corner (1976: 274) states that

seed structure supports such an alliance and mentions

that Maheshwari claims that embryological evidence

denies close relationship with Onagraceae. Hemitrapa

Miki. first described from the Miocene of Japan, is clear-

ly distinct from Trapa and belongs with Trapella in

Trapellaceae (~ Pedaliaceae). The adventitious roots

have also been considered to be leaves but all the evi-

dence (Couillault 1973) supports the other view.

5829000 TRAPA

Trapa L., Species plantarum 1: 120 (1753); L.: 56

(1754); Nakano: 440^58 (1913); A.Arber: 245 (1920);

Gams: 884 (1925); Gams: 39-41, Karten 25-27 (1927);

Steenis: 43 (1949); V.N.Vassil.: 638 (1949); Brenan: 1

(1953); Gams: 108-115, figs 1-6 (1958); Miki: 289-294

(1959); Nakano: 159-167 (1964); Sculthorpe: 328

(1967); C.D.K.Cook et ah: 537 (1974); Cronquisl: 638

(1981); C.D.K.Cook: 217 (1990). Type species: Trapa

natans L.

Characters of the family.

About 70 species have been described, most of them

from Russia and adjacent areas. Some 25-30 species are

still maintained by Russian botanists but most others

recognise only a few or even only one polymorphic

species with numerous varieties. The latter appears to be

the best course and is followed here, although Szafer

* Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK.

This account is largely based on that prepared by J.P.M. Brenan for

Flora zximbesiaca.

MS. received: 1985-01-14.

(1954) argues the contrary. Daniel et al. (1983: 596)

would not even recognise varieties. There are many fos-

sil species, the genus having occurred throughout the

Tertiary and extending into the Upper Cretaceous. The

generic name is a contraction of calcitrappa (Latin), a

caltrop, a defensive weapon at one time used chiefly

against cavalry and somewhat like the fruits of the plant.

Trapa natans L., Species plantarum 1: 120

(1753); Gams: 884, fig. 2263-2269, t. 190/1 (1925);

Brenan: 1, fig. 1 (1953): Roessler: 1 (1966); Tutin: 303

(1968); D.F.Chamb.: 196 (1972); C.D.K.Cook et al.: fig.

262 (1974); Brenan: 346. fig. 85 (1978); Verde.: 448

(1986). Type: S Europe. Italy, Mantua, L. Superiore,

Fiori 471 (K, neo.l).

Annual with slender main axis 0. 5-4.0 m long accord-

ing to depth of water, unbranched or with 2-14 branches

each ending in a floating rosette 150-500 mm wide.

Leaves', blades often broader than long, 10-65 x 10-80

mm, glabrous above and beneath or hairy on nerves or

densely hairy all over; petiole up to 210 mm long. Sepals

lanceolate to narrowly triangular, 4-7 mm long. Petals

oblanceolate to obovate, 8-16 mm long. Fruit up to ± 25

mm wide, very variably 2-4-horned.

Two varieties are distinguished in southern Africa:

Fruit 2-horned ... var. bispinosa

Fruit 4-horned ... var. pumila

5829000-1 var. bispinosa (Roxb.) Makino in

Iinuma, Somoku-Dzusetsu ('/Iconography of plants in

Japan) edn 3, 1: 137 (1907); Dubois: 399 (1954); Brenan:

3 (1953); R.Fern. & A.Fern.: 203 (1970); Brenan: 348

(1978). Type: India. Roxburgh (BM, lecto.l).

T. bispinosa Roxb. 3: 29, t. 234 (1815); Rech.f. 127: 1 (1978). T.

bicornis Osbeck var. bispinosa (Roxb.) Nakano: 165 (1964).

T. cochinchinensis Lour., Flora cochinchinensis: 86 (1790). T. bicor-

nis Osbeck var. cochinchinensis (Lour.) C.M.H. Gluck ex Steenis: 43

(1949); Backer & Bakh.f.: 265 (1963). Type: N Vietnam, Hanoi,

Balansa 4526 (P. neo.).

T. chinensis Lour.: 86 (1790). Type: as for T. bicornis L.f., Plum.: t.

67 bottom (1693) (holo. !).

T. austroafricana V.N.Vassil.: 192, t. 12 (1965). Type: Zambia,

Chingola, Funshawe 2535 (K, holo.!; SRGH).

T. congolensis V.N.Vassil.. op. cit. 184. figs 8, 9 (1965). Type: Zaire,

Leopoldville, Couteaux 1014 (BR, holo.).

T. insperata V.N.Vassil., op. cit. 178, fig. 3 (1965). Type: Angola, R.

Cubango, Gossweiler 2087 (P, holo.).

Leaves sometimes purplish beneath. Fruit ± 30-50

mm wide overall, 2-horned, horns arising from upper

angles, erecto-patent to arcuate-ascending or almost hor-

izontal, straight or somewhat curved, conical, or attenu-

ate above, ± 10-18 mm long and ± 4-7 mm wide near

12

Bothalia 28,1 (1998)

FIGURE 1 — A-L, Trapa nutans var. bispinosa: A, habit, x 0.8; B, part of leaf margin, lower surface, x 3.8; C, nodes showing stipules, x 2.5;

D, flower, x 2.5; E, I s. flower, x 3.8; F, two sepals, x 3.8; G, petal, x 3.8; H, anther, two views, x 7.6; I, ovary and disc, x 7.6. J-L, fruit:

J, endocarp, x 1 2; K, side view, x 1 .2; L, apex of horn, x 2.5. M-O, fruit of T. nutans var. pumila: M, endocarp, x 1.2; N, side view, x

1.2; O, apex of horn, x 2.5. A-D, Smith 2987', E-I, Riegel & Gibbs Russell 3714', J, K, Gibbs Russell 2786', L, Ellis 3073', M-O, Wager

37369.

Bothalia 28,2 (1998)

13

FIGURE 2. — Distribution of Trapa natans var. bispinnsa in southern

Africa.

5829000-2 var. pumila Nakano ex Verde, in Kew

Bulletin 41: 448 (1986). Type: Vietnam, Hanoi, Balansa

4721 (P, holo. ! ; K, iso.!).

T. natans L. var. incisa Makino in Botanical Magazine, Tokyo 22:

172 (1908); Nakano: 454, t. 2/33-34 (1913) ‘grosse Formen’; T. natans

L. var. pumila Nakano: 166 (1964); Van Cuong & J.E. Vidal: 46, t.

7/5-6 (1973) based on var. incisa ‘grosse Formen’ but no type cited,

nom. invalid.

Fruit ± 40 mm wide overall, 4-horned, 2 fairly slen-

der horns slightly ascending from upper angles 10-15

mm long and 2 strongly deflexed curved inferior horns

mostly ± 10 mm long, all sharp and usually with reflexed

barbs. Sometimes 2 additional short projections above

inferior horns. Figure 1M-0.

In KwaZulu-Natal, Tongoland, Ingwavuma (Figure

3), also in Cameroon ( Saxer 275), Japan, Indochina,

Vietnam, India? Mainly in swamps and pans; 0-100 m.

Vouchers: Moll 1762 (PRE); Musil 46 (Univ. Nat ); Wager NH

37369 (NH, PRE).

base, sharp at point and usually reflexedly barbed for a

little way below it; an additional inferior tubercle or short

reflexed blunt horn may be present on each side and

occasionally 2 further small projections above it. Figure

1A-L.

In NE Namibia along the Okavango River, N

Botswana (Okavango Swamp) and Northern Province —

formerly northeastern Transvaal (Figure 2). Also in W

Africa, Zaire, Sudan, Tanzania, Zambia, Zimbabwe,

Malawi, Mozambique and Angola, throughout India,

Indochina, China and Japan, also in Java. Rooted in soft

mud in slowly flowing water, reedswamps, permanent

pans etc.; 940-1 030 m.

Vouchers: Biegel & Russell 3714 (K; SRGH); De Winter & Marais

4473 ; 4765 (K, PRE): Dinter 7198 (K, PRE); Ellis 3073 (PRE).

Very likely T. quadrispinosa Roxb. (Type locality:

Bangladesh, Sylhet) is this variant, but neither a speci-

men nor a drawing has been found; Vassiljev has chosen

Stewart 3351‘/2 from Kashmir as ‘lectotype’ although

presumably neotype was meant. It is also what Gluck

was going to call T. incisa var. quadricaudata forma lae-

vigata and forma tuberculosa in his proposed mono-

graph. T. natans var. africana has much longer more

slender ± equal spines.

The various forms of water chestnut are much culti-

vated, particularly in E Asia, and extensively used in

Chinese cuisine. Nakano’s careful work over half a cen-

tury has shown these are well-marked taxa but I have

preferred to treat them as varieties. Unfortunately he did

not follow normal taxonomic procedure of typification.

The form with additional blunt horns (Van Cuong &

Vidal loc. cit. 1973: t. 6/9) corresponds to var. jinumai

Nakano.

REFERENCES

ARBER, A. 1920. Water plants. Cambridge University Press, Cam-

bridge.

BACKER, C.A. & BAKHUIZEN VAN DEN BRINK, R.C. 1963.

Flora of Java 1: 264, 265.

BRENAN, J.P.M. 1953. Trapaceae. Flora of tropical East Africa: 1-3.

BRENAN, J.P.M. 1978. Trapaceae. Flora zambesiaca 4: 346-348.

CASPER, S.J. & KRAUSCH, H.-D. 1981. Ptendophyta und Antho-

phyta 2. Die Siisswasser- Flora Mitteleuropas 24: 659, fig. 177.

CHAMBERLAIN, D.F. 1972. Trapaceae. Flora of Turkey and the east

Aegean islands 4: 196.

COOK, C.D.K. 1990. Aquatic plant book. SPB Academic publishing,

The Hague.

COOK, C.D.K. et al. 1974. Water plants of the world. Junk, The

Hague.

CORNER, E.J.H. 1976. The seeds of dicotyledons 1: 274. Cambridge

University Press, Cambridge.

COUILLAULT, J. 1973. Organisation de l’appareil conducteur de

Trapa natans L. Bulletin de la Societe Botanique de France

119: 177-198.

CRONQUIST, A. 1981. An integrated system of classification of flow-

ering plants. Columbia, New York.

DANIEL, P, VAJRAVELU, E. & THIYAGARAJ, J.G. 1983. Consid-

erations on Trapa natans from peninsular India. Journal of

Economic and Taxonomic Botany 4: 593-601.

DUBOIS, L. 1954. Une plante curieuse. Zooleo 26: 399, 400.

FERNANDES, R. & FERNANDES, A. 1970. Trapaceae. Conspectus

florae angolensis 4: 203.

GAMS, H. 1925. Trapaceae. Hegi, lllustrierte Flora von Mitteleuropa

5, 2: 884.

FIGURE 3. — Distribution of Trapa natans var. pumila in South Africa.

14

Bothalia 28,1 (1998)

GAMS, H. 1927. Die Gattung Trapa L. Pflanzenareale, Reihe 1, Heft

3: 39-41, Karten 25-27.

GAMS, H 1958. Beitrag zur Variabililat derFriichte von Trapa natans.

Veroffentlichungen des Geobotanischen Institutes Riibel in

Zurich 33: 108-115, figs 1-6.

LINNAEUS, C. 1745. Flora suecica. Laurentius Salvius, Stockholm.

LINNAEUS, C. 1753. Species plantarum I: 120. Laurentius Salvius,

Stockholm.

LINNAEUS, C. 1754. Genera plantarum, edn 5. Laurentius Salvius,

Stockholm.

LOUREIRO, J. DE 1790. Flora cochinchinensis. Academy, Lisbon.

MAKINO, T 1907. In Iinuma, Somoku-Dzusetzu (Iconography ... plants

... Japan), edn 3,1: 137.

MAKINO, T. 1908. Observations on the flora of Japan (continued).

Botanical Magazine, Tokyo 22: 155-176.

MIKI, S. 1959. In Proceedings of the Academy of Japan 35: 289-294.

NAKANO, H. 1913. Beitrage zur Kentnis der Variationen von Trapa in

Japan. Botanische Jahrbiicher 50: 440^-58.

NAKANO, H. 1964. Further studies on Trapa from Japan and its adja-

cent countries. Botanical Magazine, Tokyo 77: 159-167.

PLUMIER, C. 1693. Description des plantes de TAmerique : t. 67 bottom.

RECHINGER, K.H. 1978. Trapaceae. Flora iranica 127: 1, 2.

ROESSLER, H. 1966. Trapaceae. Prodromus einer Flora von

Siidwestafrika 96: 1

ROXBURGH, W. 1815. Plants of the coast of Coromandel 3: 29, t.

234. East India Company, London.

SCULTHORPE, C D. 1967. The biology of aquatic vascular plants.

Arnold, London.

SZAFER, W. 1954. On some living and fossil forms of Trapa L. Acta

Societatis Botanicorum Poloniae 23: 117-142.

TUTIN, T.G. 1968. Trapaceae. Flora europaea 2: 303.

VAN CUONG, V. & VIDAL, J.E. 1973. Trapaceae. Flore Cambodge,

Laos, Viet-nam 14: 40^17.

VAN STEENIS, C.G.G.J 1949. Hydrocaryaceae. Flora malesiana 1,

4: 43, 44.

VASSILJEV, V.N. 1949. Hydrocaryaceae. Flora of the USSR 15: 638.

VASSILJEV, V.N. 1965. Novye Afrikanskie vidy roda Trapa L. Novosti

Sistematiki Vysshikli Rastenii 1965: 175-194.

VERDCOURT, B. 1986. Notes on Trapa for the ‘Flora of southern

Africa’. Kew Bulletin 41: 447-449.

Bothalia 28,1: 15-17 (1998)

A revision of Ledebouria (Hyacinthaceae) in South Africa. 1. Two

new species

S. VENTER* and T.J. EDWARDS**

Keywords: Hyacinthaceae, Ledebouria , new species. South Africa, taxonomy

ABSTRACT

A revision of Ledebouria in South Africa has revealed a number of undescribed species. This paper deals with two new

species, Ledebouria atrobrunnea S.Venter and L. dolomiticola S. Venter. Both species possess cylindrical bulbs, erect

leaves (rare in Ledebouria ) and prominent shoulders on the carpel apices. These synapomorphies are also seen in L. viscosa

Jessop, and its allies.

INTRODUCTION

Ledebouria Roth includes some highly variable spe-

cies and this has led to a proliferation of synonyms with-

in the genus. Species plasticity, however, has also result-

ed in distinct taxa being overlooked in previous revisions

(Jessop 1970). The species described in this account ( L .

atrobrunnea and L. dolomiticola) are based on characters

which do not vary significantly between populations or

in cultivated specimens. They form a natural group with

L. viscosa, sharing cylindrical bulbs, erect leaves (rare in

Ledebouria) and prominent shoulders on the carpel

apices. All three species are exceedingly rare with very

narrow distributions, features which make them particu-

larly vulnerable to extinction.

1. Ledebouria atrobrunnea S.Venter, sp. nov.

similis L. viscosae, sed foliis 4—6 (non solum 1-3), attenu-

atis (haud oblanceolatis usque spathulatis), non-viscosis

tortisque et squamarum apicibus duris distincte differt.

TYPE. — North-West, 2527 (Rustenburg): Kroondal,

Farm Zuurplaat 337, (-CD), Venter 13460 (PRE, holo.;

NU, UNIN).

Plants solitary. Bulb hypogeal, 30-60 x 20-30 mm,

cylindrical, dead bulb scales purplish brown, very hard,

attenuate, apices truncate, live bulb scales loose with

threads when torn, bulb white and often purple-spotted.

Leaves partly emerged at anthesis, 4-6, erect, spirally

twisted, linear-lanceolate, 60-80 x 3-10 mm, with

threads when torn, fleshy, dull glaucous, venation

obscure; margin undulate in lower half, smooth above;

base canaliculate, apex acute. Inflorescences 1-4, flac-

cid, lax, 30-60-flowered, longer than leaves; peduncle

compressed at base, purple with darker spots, 25-60 mm

long; rachis longitudinally ridged; raceme lax, oblong,

20-50 x 25-30 mm; bracts and bracteoles always pre-

sent, slightly fleshy, 0.8-1. 0 x 0.25-0.5 mm, linear to

bifurcate, grey-white. Pedicels spreading, 6-7 mm

long, speckled or pink. Tepals recurved, subequal, linear-

* Botany Department, University of the North, Private Bag 1106, 0727

Sovenga. Present address:

** Department of Botany, University of Natal, Private Bag X01,

Scottsville 3209.

MS. received: 1997-04-16.

oblong, 4. 0-5.0 x 1. 5-2.0 mm, pink to purple, keel

green; apex acute, slightly cucullate. Stamens erect, 3-4

mm long; filaments pink, base slightly flattened,

epitepalous; anthers 0.75-1.0 mm long, violet. Ovary

depressed, obtusely 6-lobed, 1.0-1. 5 x 2.0-2. 5 mm,

lobes obtusely debate, apical shoulders present, basal

lobes present; stipe 0.25 x 0.25 mm; style 2. 0-2. 5 mm

long, triangular in section, glabrous, purple. Capsule

FIGURE 1. — Ledebouria atrobrunnea. Venter 13460. A, habit, x 0.9;

B, section through lamina, x 3; C, tepal apex, x 12; D, bract, x

12; E, flower, x 7; F, tepal with stamen, x 7; G, ovary lateral

view, x 9. Drawing modified from Burrows (1993).

16

Bothalia 28,1 (1998)

FIGURE 2. — Geographical distribution of L. viscosa , •; L. atrobrun-

clavate, base truncate. Seed 4-5 mm long, drop-shaped,

strongly wrinkled, reddish brown. Figure 1.

This species is known only from Magaliesberg

Quartzites of the Pretoria Group (SACS 1980) in the

foothills of the Magaliesberg in the Kroondal area near

Rustenburg (Figure 2). It occurs in shallow red-brown

lithosols derived from coarse-grained recrystallised rock.

Vegetation of the area is Bankenveld and consists of low

closed deciduous Protea caffra-Lannea discolor Wood-

land (Acocks 1988).

The degree of leaf twisting and the prominence of

bulb scales vary within L. atrobrunnea. Plants from

exposed situations tend to have more prominent bulb

scales and leaves more twisted than shaded individuals.

The species flowers from September to late December.

L. atrobrunnea is related to L. viscosa Jessop which

also has cylindrical bulbs and erect leaves. It is distin-

guished by its linear-lanceolate, non-viscid, twisted

leaves and its hard purplish brown (hence the specific

epithet) bulb scales. In L. viscosa the leaves are viscid,

untwisted and oblanceolate to spathulate.

NORTH-WEST. — 2527 (Rustenburg): Kroondal, Farm Zuurplaat

337, (-CD), Venter 13460 (NU, PRE, UNIN); Venter 13483 (PRE,

UNIN).

2. Ledebouria dolomiticola S.Venter, sp. nov.

similis L. viscosae et L. atrobrunneae, sed ab ambobus

differ! bulbis epigaeis. Praeterea ab ilia foliis non-visco-

sis et ab hac bulbi squamis mortuis membranaceis (non

duris) recedit.

TYPE. — Northern Province, 2429 (Zebediela):

Strydpoort Mountain, Donkerkloof, Farm Rivierplaats

354, (-BA), Venter 13089a (PRE, holo.; NU, UNIN).

Bulbs epigeal, 40-100 x 15-30 mm, ellipsoid to

cylindrical, in dense groups; dead bulb scales thinly

membranous, brown, apices truncate, without threads

when torn, live bulb scales tightly appressed, white

inside, bulblets often present. Leaves fully developed at

anthesis, 3-5, erect, lanceolate, 80-100 x 8-20 mm,

without threads when torn, fleshy, glaucous, immaculate,

venation obscure; margin smooth; base canaliculate,

apex acute. Inflorescence solitary, flaccid, lax, 30-40-

flowered, longer than leaves; peduncle smooth, terete at

base, green, 60-90 mm long; rachis smooth; raceme lax,

cylindric, 40-60 x 20-30 mm; bracts with bracteoles,

membranous, 1.50 x 0.25 mm, linear-lanceolate, grey to

white. Pedicels spreading horizontally, 8-10 mm long,

white to purple. Tepals recurved, subequal, oblong, 5 x

1.0-1 .5 mm, pink to purple, keel green. Stamens erect, 3

mm long; filaments maroon, epitepalous; anthers 0.5 mm

long, yellow. Ovary spheroidal, 1.5 x 2.5 mm, lobes nar-

rowly transversely oblong; apex shoulders raised, basal

lobes present; stipe 0.25-0.50 mm long; style 3 mm long,

triangular in section, glabrous, purple above and white

below. Capsule subglobose, base truncate. Seed 4 mm

long, globose, strongly wrinkled, brown. Figure 3.

L. dolomiticola occurs in the Strydpoort Mountains

near Pietersburg, Northern Province (Figure 2). Plants

are limited to the Eccles Formation of the Chuniespoort

Group (SACS 1980). The steep dolomitic slopes and

cliffs on which L. dolomiticola grows experience. high

temperatures, especially during the summer months.

Plants occur commonly in rock fissures and lithosols.

Flowering occurs from January to April, as in L. viscosa.

FIGURE 3. — Ledebouria dolomiticola.. Venter 13089a. A, habit, x

0.3; B, section through lamina, x 1.3; C, bract with bracteole, x

4.4; D, tepal apex, x 6.3; E, tepal with stamen, x 3.8; F, flower,

x 3.8; G, ovary, lateral view, x 6.3. Drawing modified from

Burrows (1993).

Bothalia 28, 1 (1998)

17

L. dolomiticola is a close relative of L. atrobrunnea

and L. viscosa which also possess cylindrical bulbs, erect

leaves and prominent shoulders on the carpel apices. L.

dolomiticola is distinguished from both by its epigeal

bulbs and from L. viscosa by its non-viscous leaves and

from L. atrobrunnea by its membranous dead bulb scales.

Specimens of L. viscosa examined

NORTHERN PROVINCE. — 2427 (Thabazimbi): Kransberg, (-BC),

Meease 10493 (PRE); Farm Waterval, Dyer & Ehrens 4201 (PRE);

Farm Buffelshoek 446, (-DA), Venter 13455 (UNIN).

ACKNOWLEDGEMENTS

Financial assistance from the Natal University

Research Fund and the FRD is gratefully acknowledged.

The cited herbaria are thanked for the loan of material

examined during this study. Mr M. Lambert is thanked

for translating the Latin diagnoses. Lastly Mr I. Ter-

blanche and the Department of Nature Conservation,

Lebowa are thanked for granting permission to collect

material in their reserves. A major part of this research

was undertaken at the Department of Botany, University

of the North, Private Bag 1 106, Sovenga 0727.

REFERENCES

ACOCKS, J.P.H 1988. Veld Types of South Africa, 3rd edn. Memoirs

of the Botanical Survey of South Africa No. 57.

BURROWS, S. 1993. Illustrations in S. Venter, A revision of the genus

Ledebouria in South Africa. M.Sc. Natal University.

JESSOP. J.P. 1970. Studies in the bulbous Liliaceae: 1. Scilla,

Schizocarphus and Ledebouria. Journal of South African

Botany 36: 233-266.

SOUTH AFRICAN COMMITTEE FOR STRATIGRAPHY (SACS)

1980. Stratigraphy of South Africa, Part 1. Handbook of the

Geological Survey of South Africa.

Bothalia 28,1: 19-24 (1998)

Revised key to Ipomoea (Convolvulaceae) in southern Africa

W.G. WELMAN* and A.D.J. MEEUSE**

Keywords Convolvulaceae, Ipomoea, key, southern Africa

ABSTRACT

A revised key to the 57 indigenous and naturalised species of Ipomoea L. in southern Africa is presented.

INTRODUCTION

During the preparation of the Convolvulaceae for the

Flora of southern Africa (FSA), Vol. 28, part 1, it became

evident that the key to Ipomoea L. published by Meeuse

(1957) in his revision of the South African Convol-

vulaceae had become outdated. As publication of this

part of the FSA is not expected in the near future, it will

be useful to publish a revised key now.

Meeuse (1957) listed 47 species of Ipomoea occur-

ring naturally or naturalised in the southern African

region. He also included Stictocardia Hallier f. (one

species in southern Africa which can readily be confused

with Ipomoea) and Turbina sensu Meeuse (seven species

in southern Africa) in his key to Ipomoea. The list of

Ipomoea in the region compiled by Welman (1993) con-

tained 54 species, including I. shupangensis Baker and I.

urbaniana (Dammer) Hallier f., both from tropical

Africa; these are now excluded. The treatment of

Ipomoea for FSA Vol. 28,1 will contain 57 species. Four

are new records for the region namely I. fanshawei

Verde., I. fulvicaulis (Hochst. ex Choisy) Boiss. ex

Hallier f. vat. fulvicaulis, I. ochracea (Lindl.) G.Don, all

from northern Botswana and I. consimilis Schulze-Menz

from northern KwaZulu-Natal. The endemic I. bisavium

A. Meeuse was described after the publication of

Meeuse’s revision: t. 1360 (1961).

In Meeuse & Welman (1996) some name changes are

listed and explained. Six of the southern African species

of Turbina sensu Meeuse are returned to Ipomoea. name-

ly, I. holubii Baker [T. holubii (Baker) A. Meeuse], /.

oblongata E.Mey. ex Choisy [T. oblongata (E.Mey. ex

Choisy) A. Meeuse, I. atherstonei Baker], I. oeno-

theroides (L.f.) Raf. ex Hallier f. [T. oenotheroides (L.f.)

A.Meeuse], I. robertsiana Rendle [T. robertsiana

(Rendle) A.Meeuse], I. stenosiphon Hallier f. [T.

stenosiphon (Hallier f.) A.Meeuse], and I. suffruticosa

Burch. [T. suffruticosa (Burch.) A.Meeuse]. The seventh

species of Turbina sensu Meeuse in southern Africa is

now Paralepistemon shirensis (Oliv.) Lejoly & Lisowski

[71 shirensis (Oliv.) A.Meeuse] and is not included in this

key. Two other name changes have taken place, namely,

/. dichroa Choisy (/. arachnosperma Welw.), and I. indi-

ca (Burm.f.) Merr. (I. congesta R.Br.). Three varieties

and one subspecies have been sunk, namely, I. bathycol-

pos Hallier f. (/. bathycolpos var. sinuatodentata Hallier

f.), I. magnusiana Schinz [/. magnusiana var. eenii

(Rendle) A.Meeuse], I. obscura (L.) Ker Gawl. [/.

obscura var. fragilis (Choisy) A.Meeuse], and I. bolu-

siana Schinz (I. bolusiana subsp. pinnatipartita Verde.).

In Meeuse & Welman (1996) it is also mentioned that I.

carnea Jacq. subsp. fistulosa (Mart, ex Choisy) D. Austin

(/. fistulosa Mart, ex Choisy) should be excluded from

the list of Ipomoea species in the region. However, it has

recently become clear that this poisonous exotic plant

often occurs as an escape from cultivation and it is there-

fore included in the revised key; see Welman (1997).

Some more exotic species have escaped from cultiva-

tion and are naturalised; these are also included in the

key. Several other exotic species of Ipomoea are culti-

vated in especially the warmer parts of the region and

may rarely occur as culture relics or garden escapes and

are not included in the key, notably the following: I.

arborescens (Kunth) G.Don (small tree with up to 50

mm long white flowers), I. batatas (L.) Lam. (‘Sweet

Potato’), I. coccinea L. (annual twiner with scarlet flow-

ers), I. horsfalliae Hook, (perennial twiner with pink

flowers), I. intrapilosa Rose (shrub or tree with up to 95

mm long white flowers), I. quamoclit L. (annual twiner

with pinnate leaves and scarlet flowers), and I. tricolor

Cav. (perennial twiner with predominantly blue flowers).

KEY TO INDIGENOUS AND NATURALISED SPECIES

naturalised species indicated by *

la (lb: p. 23) Leaves simple, entire, shallowly lobed, emarginate or toothed, but not pinnately, palmately or

pedately lobed nearly to base nor compound:

2a Flowers in pedunculate heads, surrounded by single large bicuspidate bract:

3a Sepals lanceolate, acute, glabrous or sparsely hairy, mainly at tips and along margins; corolla distinctly

funnel-shaped, 30-50 mm long, limb 30-50 mm across; Northern Province .... 27. /. involucrata P.Beauv.

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

** Voorstraat 41, NL-1931 AH Egmond aan Zee, The Netherlands.

MS. received: 1997-09-25.

20

Bothalia 28,1 (1998)

3b Sepals oblong-spathulate to oblong, obtuse, glabrous below tips inside; corolla tube slender, subcylin-

dric, ± 20 mm long, limb spreading, 15-30 mm across; N Namibia, N Botswana, Mpumalanga ....

28. /. pileata Roxb.

2b Flowers not in heads, or, if so, heads not subtended by one large and bicuspidate, boat-shaped bract:

4a (4b: p. 23) Corolla narrowly to widely funnel-shaped:

5a Leaves with minute black dots beneath; sepals ± 8 mm long, much-imbricate, subequal, glabrous, obo-

vate, obtuse; KwaZulu-Natal Stictocardia laxiflora (Baker) Hallier f.

5b Leaves without black dots (sometimes minutely pitted or pellucidly glandular):

6a Corolla small, under 18 mm long and/or its limb less than ± 15 mm across; flowers in fascicles, clus-

ters or solitary, sometimes in pedunculate 1 to few-flowered cymes or heads:

7a Leaves when dry pellucidly glandular when seen in transmitted light, crisped along margin; sepals

very unequal, outer ones ovate from cordate base, obtuse, markedly accrescent, crisped along

margin; corolla 12-14 mm long, mauvish pink; capsule and seeds hairy; Namibia, Botswana,

Northern Province, Northern Cape 6.1. hackeliana (Schinz) Hallier f.

7b Leaves not pellucidly glandular:

8a Flowers in distinctly pedunculate, bracteate, dense heads; peduncle over 20 mm long; leaves entire,

densely white tomentose beneath; corolla up to 15 mm long, pale yellow; erect to trailing annual;

N Namibia, Botswana, Northern Province 22. I. chloroneura Hallier f.

8b Flowers not in pedunculate bracteate heads; if fascicled or clustered, peduncles never longer than

20 mm:

9a Leaves hastate at base with entire or toothed basal auricles, very rarely all lanceolate; flowers soli-

tary, rarely fasciculate; pedicels up to 12 mm; sepals linear, acuminate, hairy, 7-1 1 mm long;

corolla pinkish mauve with darker centre, occasionally white; capsule hirsute; Northern Pro-

vince, North-West, Gauteng, Mpumalanga 5.1. gracilisepala Rendle

9b Leaves cordate, cuneate or truncate at base, never hastate:

10a Leaves usually 3-lobed, albotomentose beneath; flowers mauve-purple, in pedunculate

dichasial 3-1 1-flowered cymes; N Namibia, N Botswana, Northern Province, Mpumalanga,

Swaziland 19. /. dichroa Choisy

10b Leaves entire, not albotomentose:

1 la Leaves cuneate to truncate-rounded, rarely subcordate at base; flowers very small, sessile;

corolla white, hardly longer than the usually bristly calyx; N Namibia, Botswana, Swaziland,

N South Africa 1. /. coscinosperma Hochst. ex Choisy

1 lb Leaves distinctly cordate at base:

1 2a Outer sepals cordate or auricled at base, attenuate into an acute apical portion; corolla about

twice as long as calyx; peduncles distinct, over 10 mm long; inflorescences cymosely 1-3-

flowered; Namibia, Botswana, Swaziland, N and E South Africa

. . . 8. /. sinensis (Desr.) Choisy subsp. blepharosepala (Hochst. ex A. Rich.) Verde, ex A.Meeuse

12b Outer sepals not as above; corolla slightly longer than calyx:

13a Leaf apex obtuse and minutely mucronate; petiole up to 20 mm long; infloresences axillary,

1-3-flowered, subsessile; Caprivi, N Botswana 2.1. leucanthemum (Klotzsch) Hallier f.

1 3b Leaf apex acute, subacute or acuminate; petioles up to 80 mm long:

14a Flowers usually in clusters; sepals very hairy, ovate-lanceolate with linear-acuminate,

spreading tips; capsule hairy; seeds glabrous; Botswana, Northern Province, Gauteng,

Mpumalanga 2.1. eriocarpa R.Br.

14b Flowers solitary or in 2-5-flowered cymes; sepals obtuse or subacute, ciliate or nearly

glabrous; capsule glabrous or with a few bristly hairs; seeds densely pubescent; N

Namibia, N Botswana, Northern Province, Mpumalanga, Swaziland, KwaZulu-Natal

4.1. plebeia R.Br. subsp. africana A.Meeuse

6b Corolla larger to very large, over 18 mm long; flowers usually pedunculate:

15a Leaves bilobed at apex, coriaceous or subcoriaceous, whole plant glabrous; stems stout, trailing;

growing only on sandy beaches; KwaZulu-Natal, Western and Eastern Cape

21.1. pes-caprae (L.) R.Br. subsp. brasiliensis (L.) Ooststr.

15b Leaves not bilobed (sometimes emarginate) at apex, or not found on sandy beaches:

16a Leaves cordate-ovate in outline, shallowly 3-lobed, or entire, cordate, cordate-ovate or cordate-

deltoid, covered with thin white cobwebby tomentum beneath (occasionally reduced to a few

scattered floccose patches); flowers mauve or magenta; bracteoles and sepals lanceolate or lin-

ear-lanceolate, acute to acuminate-aristate:

17a Flowers in dense globose pedunculate heads; corolla 20-40 mm long; Northern Province,

Mpumalanga, KwaZulu-Natal, Eastern Cape 20. I. wightii (Wall.) Choisy

17b Flowers in few- to many-flowered dichasia or cymes, sometimes solitary:

18a Corolla 18-25 mm long; N Namibia, N Botswana, Northern Province, Mpumalanga, Swazi-

land 19. /. dichroa Choisy

18b Corolla 40-60 mm long; Swaziland, KwaZulu-Natal, Eastern Cape 21. /. ficifolia Lindl.

16b Leaves either not with white cobwebby tomentum or bracts and sepals different (and if so, corol-

la white or pale mauve):

Bothalia 28, 1 (1998)

21

19a Leaves linear or lanceolate with narrow or sometimes somewhat rounded base, entire or some-

times toothed, repand, lobed or ± emarginate; plants erect to prostrate, often suffruticose:

20a Vegetative parts pubescent; calyx tomentose or pubescent:

21a Vegetative parts (at least on younger shoots) and calyx usually covered with an adpressed,

short and silvery pubescence:

22a Erect, much-branched shrub; leaves linear or oblanceolate, sessile or subsessile, narrowed

at base; flowers solitary on very short peduncles; Namibia, North-West, Free State, KwaZulu-

Natal, Northern and Eastern Cape; endemic 13. /. oenotheroid.es (L.f.) Raf. ex Hallier f.

22b Prostrate or suberect perennial; leaves usually distinctly petiolate, rounded or subcordate at

base; flowers solitary or 2 together; peduncles 10—50 mm long; Namibia, North-West,

Northern Cape; endemic 15. /. suffruticosa Burch.

21b Vegetative parts usually not with adpressed short and silvery tomentum:

23a Sepals unequal; plant hairy; Namibia, Botswana, N and E South Africa, Swaziland, Lesotho

9.1. crassipes Hook.

23b Sepals equal or subequal:

24a Sepals acuminate to aristate; leaves usually long-attenuate at base and well over 50 mm

long, usually also some toothed or pinnatisect, hairy or glabrous above; pubescence soft,

white; N Namibia, Botswana, Northern Province, North-West, Gauteng

7. 1. oenotherae (Vatke) Hallier f.

24b Sepals acute or acuminate; leaves not as above:

25a Leaves under 50 mm long, entire, on a short petiole or subsessile, glabrous above; pubes-

cence soft, white; Northern Province, Mpumalanga; endemic 14. /. robertsiana Rendle

25b Leaves usually over 50 mm long, entire, hairy above and below; pubescence usually stiff,

adpressed on leaves, yellowish or brownish; Namibia, Botswana, central and E south-

ern Africa • 24.1. oblongata E.Mey. ex Choisy

20b Vegetative parts (at least the leaves) glabrous; stems occasionally minutely scabrid-hirsutulous:

26a Corolla white, 20-35 mm long; plant stemless or with short stem; leaves entire or with few

teeth; mainly E southern Africa, not Swaziland; endemic 38. /. simplex Thunb.

26b Corolla magenta or pale mauve with darker centre, over 25 mm long:

27a Leaves usually over 5 mm wide; corolla pale mauve with darker centre; stems usually stout,

sometimes minutely scabrous or hirsutulous; Namibia, Botswana

39.1. welwitschii Vatke ex Hallier f.

27b Leaves usually less than 5 mm wide; corolla magenta; stems often slender, glabrous; main-

ly N southern Africa 40. /. bolusiana Schinz

19b Leaves broader, not linear or lanceolate or, if narrow, broadly truncate, hastate, sagittate or cor-

date at base:

28a (28b: p. 23) Annual or perennial plants, herbaceous, prostrate or climbing, sometimes suffru-

ticose and suberect, but if so, under 500 mm high, never tall and woody:

29a Stems thick, trailing, hollow or spongy, rooting at nodes; in moist, marshy, inundated locali-

ties or even completely aquatic, floating; N Namibia, N Botswana, N KwaZulu-Natal ....

36.1. aquatica Forssk.

29b Stems not thick, hollow or spongy or rooting at nodes and plant not usually found in very

moist localities:

30a (30b: p. 22) Leaves distinctly cordate or sagittate at base:

31a Corolla either yellow and concolorous, or white or yellow with a purple or brown centre;

sepals subequal, 4-8 mm long;

32a Corolla up to 25 mm long; N and E southern Africa

29.1. obscura (L.) Ker Gawl. var. obscura

32b Corolla 27 mm long or longer; Botswana 36.1. ochracea (Lindl.) G.Don var. ochracea

31b Corolla pink, mauve, magenta, purple to blue or pure white, if white or yellowish with

purple or magenta centre; sepals very unequal and over 8 mm long:

33a Sepals entirely glabrous:

34a Petiole short, under 15 mm long; leaves usually under 50 mm long; plant usually hairy

in some part other than calyx; corolla with small tufts of hairs protruding from mid-

petaline areas; Northern Province, North-West, Gauteng, Mpumalanga

31. 1. transvaalensis A.Meeuse

34b Petioles 15-200 mm long:

35a Leaves cordate-elliptic or rounded to subreniform, obtuse with minute cusps, basal

sinus narrow and deep; sepals 12-20 mm long; corolla 35-70 mm long; mainly N

South Africa; endemic 32.1. bathycolpos Hallier f.

35b Leaves cordate-deltoid or cordate-ovate, usually suddenly acuminate from broad,

usually irregularly and coarsely few-toothed base into an entire apex, basal sinus

broad; sepals 5-9 mm long; corolla 20-35 mm long; Botswana, Swaziland, N South

Africa 33.1. papilio Hallier f.

33b Sepals thinly to densely hairy or tomentose:

22

Bothalia 28,1 (1998)

36a Sepals long-attenuate or linear-acuminate at apex, 14-25 mm long; twining herbaceous

plants with retrorse hairs on stems and petioles; leaves often 3-lobed:

37a Outer sepals lanceolate at base with long and linear acumen, patently hirsute in basal

portion, 17-25 mm long; cultivated and naturalised; N Namibia 16.* I. nil (L.) Roth

37b Outer sepals lanceolate to ovate-lanceolate, gradually attenuate towards apex, with

adpressed pubescence, 14-22 mm long; cultivated and naturalised; Gauteng, Mpu-

malanga, Swaziland, KwaZulu-Natal, Eastern and Western Cape

18.* /. indica (Burm.f.) Merr.

36b Sepals acute, obtuse or mucronate but not with long and narrow acumen, if sepals long-

acuminate, plant not twining and without retrorse hairs on stems and petioles:

38a Calyx with bristly patent hairs in basal portion, glabrous towards apex; twining annu-

al with retrorse hairs on stems and petioles; cultivated and naturalised; N and E

South Africa 17.* /. purpurea (L.) Roth

38b Calyx not with bristly patent hairs in basal portion and glabrous towards apex:

39a Twining plant, growing in moist places (river banks, marshes, pools), often among

reeds, sedges etc.; N Namibia, N Botswana 48. /. rubens Choisy

39b Prostrate to suberect plants, or erect shrubs:

40a Plants prostrate to suffruticose, if suberect never more than 500 mm high; corolla

usually magenta:

41a Bracts usually close to calyx, never foliaceous; sepals subequal:

42a Bracts ovate, acuminate; Caprivi, N Botswana

12. /. fulvicaulis (Hochst. ex Choisy) Boiss. ex Hallier f. var .fulvicaulis

42b Bracts linear or lanceolate, rarely broader; Namibia, Botswana, central and E

southern Africa 24. /. oblongata E.Mey. ex Choisy

41b Bracts usually distant from calyx, often foliaceous; sepals very unequal, outer

ones ovate-acuminate, inner ones linear; Namibia, Botswana, N and E South

Africa, Swaziland, Lesotho 9.1. crassipes Hook.

40b Plant shrubby, erect; stems woody; sepals very broad, unequal; corolla pale mauve

with magenta centre; Namibia, Botswana, Northern Province, Mpumalanga ....

52.1. holubii Baker

30b (30a: p. 21 ) Leaves not distinctly cordate or sagittate at base:

43a Sepals entirely glabrous, sometimes rugose or muriculate:

44a Sepals 5-12 mm long:

45a Sepals 5-9 mm long; corolla magenta; leaves acuminate from broad base, coarsely

toothed in lower half; Botswana, Swaziland, N South Africa 33.1. papilio Hallier f.

45b Sepals 8-12 mm long; corolla pale mauve with darker centre; leaves linear-oblong to

elliptic, entire, emarginate or 3-Iobed, not broad or toothed at base; Namibia,

Botswana >. 39.1. welwitschii Vatke ex Hallier f.

44b Sepals 12-25 mm long:

46a Leaves rhomboid-lanceolate in outline, margin entire to lobed; sepals 15-18 mm long;

corolla pale purple or mauve; Botswana 46. 1. fanshawei Verde.

46b Leaves oblong, elliptic, ovate to linear-oblong, entire; sepals 12-25 mm long; corolla

magenta; Namibia, Botswana, central and E southern Africa

24. /. oblongata E.Mey. ex Choisy

43b Sepals more or less hairy to tomentose:

47a Plant shrubby, erect, over 500 mm high; sepals very hroad, suborbicular, rounded,

unequal, inner ones larger; Namibia, Botswana, Northern Province, Mpumalanga . .

52.1. holubii Baker

47b Plant prostrate; sometimes suffruticose, suberect, but if so, under 500 mm high and sepals

not very broad and rounded:

48a (48b: p. 23) Flowers in pedunculate, few to many-flowered heads (heads occasionally

binate):

49a Bracteolcs and sepals linear, narrow, bristly with long yellowish hairs; Mpumalanga,

Swaziland, E Free State, KwaZulu-Natal, Eastern Cape; endemic

10. /. pellita Hallier f.

49b Bracteoles and sepals lanceolate to ovate:

50a Leaves lanceolate or ovate-lanceolate from broad base to long-triangular, gradually

narrowed towards apex; corolla bright magenta, 30-50 mm long; Botswana, North-

ern Province, North-West, Gauteng, Mpumalanga, Free State 23.1. ommaneyi Rcndlc

50b Leaves usually oblong to ovale and not often gradually tapering to apex from broad

base:

51a Corolla magenta, 50 mm or longer; Namibia, Botswana, central and E southern

Africa 24. /. oblongata E.Mey. ex Choisy

51b Corolla mauve or purple, 25-50 mm long; Caprivi, N Botswana

12. 1. fulvicaulis (Hochst. ex Choisy) Boiss. ex Hallier f. var. fulvicaulis

Bothalia 28.1 (1998)

23

48b (48a: p. 22) Flowers solitary or 2 or 3 together, or not in heads but in cymes:

52a Sepals very unequal; outer ones ovate or lanceolate from broad base, inner ones lin-

ear; bracts usually distant from the calyx and often foliaceous; Namibia, Botswana,

N and E South Africa, Swaziland, Lesotho 9.1. crassipes Hook.

52b Sepals equal or not so distinctly unequal:

53a Leaves up to 45 mm long, broad at base, often subcordate; bracteoles very minute;

calyx 7-8 mm long; corolla bright magenta-rose, 20-40 mm long with small tufts

of hairs protruding from midpetaline areas (especially conspicuous in older buds);

Northern Province, North-West, Gauteng, Mpumalanga

31. /. trcinsvaalensis A.Meeuse

53b Leaves, bracts, sepals or corolla longer or corolla, if hairy on midpetaline areas,

without protruding hair tufts:

54a Bracteoles distant from, more rarely contiguous to calyx; leaves quite glabrous

above; corolla thinly hairy to glabrous on midpetaline areas; Northern Province,

Mpumalanga; endemic 14. I. robertsiana Rendle

54b Bracteoles contiguous to calyx; leaves usually hairy on both sides; corolla silky on

midpetaline areas; Eastern Cape; endemic 34. /. crispa (Thunb.) Hallier f.

28b (28a: p. 21) Tall perennial woody climbers or shrubs over 500 mm high:

55a Calyx glabrous:

56a Sepals orbicular to oblong, obtuse; leaves usually lobed, rarely entire; KwaZulu-Natal . . .

49. I. mauritiana Jacq.

56b Sepals ovate, cuspidate to acute; leaves entire, usually appearing after flowers; Caprivi, N

Botswana, Northern Province 50.1. shirambensis Baker

55b Calyx at least partly hairy, pubescent or tomentose:

57a Sepals broadly elliptic or obovate to orbicular, obtuse, 5-16 mm long, two inner ones larg-

er than outer ones; shrubs; leaves not longer than 250 mm:

58a Leaves cordate-ovate, usually under 50 mm long; erect shrub with ultimate twigs some-

times flexuous or twining; corolla 40-50 mm long; Namibia, Botswana, Northern

Province, Mpumalanga 52.1. holubii Baker

58b Leaves ovate to lanceolate, 100-250 mm long; erect to scrambling shrubs to 3 m high;

sepals suborbicular, 5-6 mm long; corolla 50-90 mm long; cultivated and seminatu-

ralised; Northern Province, Mpumalanga, Swaziland, KwaZulu-Natal

54. */. camea Jacq. subsp .fistulosa (Mart, ex Choisy) D.F.Austin

57b Sepals not as above; plants climbing; leaves mostly much larger:

59a Two outer sepals cordate-triangular, 14 mm long; young parts and peduncles not densely

hairy; Northern Province; endemic 11./. bisavium A.Meeuse

59b Two outer sepals not as above; young parts densely hairy, as are peduncles:

60a Young parts densely white-tomentose:

61a Young leaves with very conspicuous reticulate, somewhat floccose tomentum on main

veins below, older ones glabrescent; calyx at first tomentose, at least near base; corol-

la white, 60-90 mm long; Northern Province, Mpumalanga, Swaziland, KwaZulu-

Natal 55.1. albivenia (Lindl.) Sweet

61b Young leaves with complete tomentum below, tomentum usually persistent; calyx

tomentose; corolla white, rose-purple or mauve with darker throat, 60-100 mm long;

Namibia, N Botswana 56.1. verbascoidea Choisy

60b Young parts densely yellow-villous; sepals linear-lanceolate; N KwaZulu-Natal

57. /. consimilis Schulze-Menz

4b (4a: p. 20) Corolla salver-shaped, white or white with inside of tube purple or magenta:

62a Plant herbaceous and twining:

63a Corolla tube up to ± 40 mm long; leaves ovate to lanceolate, entire, cuneate to rounded at base; sepa-

ls not awned; N Namibia, Botswana, Northern Province, Mpumalanga, Swaziland

55.1. lapathifolia Hallier f.

63b Corolla tube at least 70 mm long; leaves cordate at base, entire or 3-lobed; sepals awned; cultivat-

ed and naturalised; Mpumalanga, KwaZulu-Natal, Eastern Cape 41. *1. alba L.

62b Plant not herbaceous and twining:

64a Plant shrubby, erect; calyx densely silky outside; Namibia, Botswana, Northern Province, Gauteng,

Mpumalanga 51./. adenioides Schinz

64b Plant woody and climbing; calyx glabrous; sepals obtuse; Northern Province

55.1. stenosiphon Hallier f.

lb (la: p. 19) Leaves deeply lobed (nearly to base), dissected or compound:

65a Leaves albotomentose below with prominent veins, palmately 3-5- or almost 7-lobed; Namibia,

Botswana, Swaziland, mainly central and N South Africa 25.1. magnusiana Schinz

65b Leaves glabrous or sometimes hairy but not albotomentose:

66a Flowers in dense pedunculate and bracteate heads; leaves hairy; N Namibia, Caprivi, N Botswana,

Mpumalanga 26.1. pes-tigridis L. var. pes-tigridis

24

Bothalia 28, 1 (1998)

66b Flowers solitary, fascicled or in cymes:

67a Corolla small, white tinged with mauve, 10-15 mm long; leaves palmately 5-7-fid with dentate or ser-

rate to pinnatisect segments; Namibia, N Botswana, mainly N South Africa

41. I. coptica (L.) Roth ex Roem. & Schult.

67b Corolla usually larger, rarely white; leaf segments not dentate or serrate, entire or basal ones with

lateral lobe:

68a Large glabrous perennial climber; leaves palmately 3-9-lobed, 60-150 x 60-200 mm, not pseudo-

stipulate; KwaZulu-Natal 49. /. mauritiana Jacq.

68b Annuals or prostrate perennnials, leaves much smaller, usually under 80 mm across and often pseu-

dostipulate by small leaves of developing or suppressed axillary shoots:

69a Leaves usually pseudostipulate, distinctly palmately or pedately 5-9-fid, quite glabrous; climbing

or occasionally prostrate annuals:

70a Calyx 8-10 mm long, outer sepals saccate at base; corolla yellow with mauve tube, 30-60 mm

long and across; leaves usually biternately pedate often with 9 segments; N Namibia, N

Botswana 42. /. tuberculata Ker Gawl.

70b Calyx up to ± 7 mm long, outer sepals not saccate; corolla mauve, purple or white; leaves

palmately partite with usually 5 segments:

7 1 a Corolla 30-60 x 40-60 mm, mauve, rarely white; NE southern Africa 43. /. cairica (L.) Sweet

71b Corolla up to ± 25 mm long:

72a Peduncles very slender, filiform; corolla often under 16 mm long, purplish mauve; N Namibia,

Northern Province, Mpumalanga 45. /. tenuipes Verde.

72b Peduncles rather stout, not filiform; corolla 15-25 mm long, purplish mauve or white;

Namibia, Botswana, Northern Province, Gauteng, Mpumalanga, Swaziland

44. 1. hochstetteri House

69b Leaves not pseudostipulate, pinnately dissected, tridentate or palmately compound; stems prostrate

or rarely suberect from perennial tuberous rootstock; corolla usually over 30 mm long:

73a Young parts and sepals with soft white hairs; sepals very acute; basal leaves, if present, linear,

cauline leaves tridentate to pinnatisect or all leaves pinnatisect; flowers solitary on short pedun-

cles, deep pink to magenta; N Namibia, Botswana, Northern Province, North-West, Gauteng . .

1.1. oenotherae (Vatke) Hallier f.

73b Sepals glabrous:

74a Leaves palmately 3-9-fid with linear segments under 5 mm wide (often very narrow); corolla

magenta; stems often rather slender, glabrous; mainly N southern Africa

40. /. bolusiana Schinz

74b Leaves trisect with linear, lanceolate or oblong segments usually over 5 mm wide; corolla pale

mauve with magenta centre 30-80 mm long; stems usually stout; Namibia, Botswana ....

39. /. welwitschii Vatke ex Hallier f.

REFERENCES

MEEUSE, A.D.J. 1957. The South African Convolvulaceae. Bothalia

6: 641-792.

MEEUSE, A.D.J 1961. lpomoea bisavium. The Flowering Plants of

Africa 34: t. 1360.

MEEUSE, A.D.J. & WELMAN, W.G. 1996. New records, name

changes and a new combination in southern Africa

(Convolvulaceae). Bothalia 26: 46-50.

WELMAN, W.G. 1993. Convolvulaceae. In T.H. Arnold & B.C.de Wet,

Plants of southern Africa: names and distribution. Memoirs of

the Botanical Survey of South A frica No. 62: 592-594.

WELMAN, W.G. 1997. Two exotic taxa of lpomoea and Merremia in

southern Africa (Convolvulaceae). Bothalia 27: 141.

Bothalia 28,1:25-33(1998)

Taxonomic notes on the genus Khadia (Mesembryanthemaceae/

Aizoaceae)

P. CHESSELET*, H.E.K. HARTMANN1, N. HAHN+t, P. BURGOYNE** and G.F. SMITH**

Keywords: Aizoaceae, Khadia , Mesembryanthemaceae, Ruschioideae, South Africa, succulents, taxonomy

ABSTRACT

The taxonomic status of the five species that comprise Khadia N.E.Br. is clarified on the basis of macro- and micro-

morphology. Distribution data, descriptions and a taxonomic key to the species are provided In Khadia we recognise the

following species: K. acutipetala (N.E.Br.) N.E.Br., K. alticola Chesselet & H.E.K.Hartmann, K. beswickii (L.Bolus)

N.E.Br., K. borealis L.Bolus and K. carolinensis (L.Bolus) N.E.Br.

CONTENTS

Introduction 25

Historical background 25

Taxonomy 27

Key to species 27

1. K. acutipetala 27

2. K. alticola 28

3. K. beswickii 29

4. K. borealis 31

5. K. carolinensis 31

Discussion 31

Uses 32

Specimens examined 33

Acknowledgements 33

References 33

INTRODUCTION

Khadia N.E.Br. is one of the few genera of the large,

primarily southern African succulent plant family

Mesembryanthemaceae Fenzl, which occurs in the north-

eastern summer rainfall area of South Africa. Other

mesem genera represented in this region include the

widespread Delosperma N.E.Br. (± 30 species), Lithops

N.E.Br. (3 species), and Frithia N.E.Br. (1 species).

Khadia occurs in three broad areas: 1, the Northern

Province; 2, Gauteng and the North-West; and 3,

Mpumalanga extending into northern KwaZulu-Natal

(Figure 1). All known species of Khadia occur at alti-

tudes above 1 400 m in quartzitic soils of the Sout-

pansberg Group, the Steenkampsberg Formation, the

Magaliesberg Formation and the Witwatersrand Super-

group.

A study of herbarium specimens of Khadia indicated

obvious problems in species identification. The major

reason for these problems is poor initial species diagno-

sis attributable to inadequate sampling of populations.

* Compton Herbarium, National Botanical Institute, Kirstenbosch,

Private Bag X7, 7735 Claremont, Cape Town.

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

1 Institut fur Allgemeine Botamk, Ohnhorststrasse 18, D-22609

Hamburg, Germany.

tf P.O.Box 1734, 0920 Louis Trichardt, South Africa.

MS. received: 1997-02-06.

The intrinsic complexity of some of the taxa as well as

the outdated revisionary status of the genus contributed

to the confusion. In this paper we have attempted to facil-

itate identification of species, improve species diagnosis,

bring systematics up to date and examine the character-

istics of the species accommodated in the genus and

hence its integrity. We have provided distribution data,

descriptions and a taxonomic key to the species. A recent

discovery by Pieter Winter (P. Winter in prep., University

of the North, Sovenga) suggests the existence of an unde-

scribed taxon, closely allied to K. borealis L.Bolus,

which is not included in this paper.

HISTORICAL BACKGROUND

Mesembryanthemum acutipetalum N.E.Br. was the

first species to be described in this group (Brown 1908).

A note on the label of the type specimen detailed its use

in making ‘Khadi’ (see below under USES), prompting

N.E. Brown, in his dismantling of Mesembryanthemum

L., to formulate the genus name Khadia in a taxonomic

key to genera of the ‘Mesembryanthema’ (Brown 1930).

By 1931, four species were assigned to the newly cir-

cumscribed Khadia: K. acutipetala (N.E.Br.) N.E.Br., K.

nationiae (N.E.Br.) N.E.Br., K. beswickii (L.Bolus)

N.E.Br., and K. nelsonii N.E.Br., all of which were dis-

covered in the surroundings of Johannesburg (Brown

12 14 16 18 20 22 24 26 28 30 32 34 36

FIGURE L— Known distribution of. Khadia acutipetala, A; K. alti-

cola, •; K. beswickii, O; K. borealis, ▼; K. carolinensis, O.

Bothalia 28,1 (1998)

1931). Bolus’ (1936) overview of the genus included six

species names, with K. carolinensis (L. Bolus) N.E.Br.

and the then newly described K. borealis completing the

picture. A short overview of Khadia, highlighting K.

beswickii and providing a key to species, appeared in The

Flowering Plants of Africa (Bolus 1958). Treatment of

the genus in Jacobsen (1960, 1981) was largely based on

published information. No further attempts were made to

resolve taxonomic problems in this genus until Chesselet

& Hartmann (1995) described K. alticola.

The identity of K. beswickii has been problematic

since its original description from a plant cultivated in

an Eastern Cape garden. The name has been misapplied

in herbarium identifications to the recently described K.

alticola (Chesselet & Hartmann 1995) and to K. acu-

FIGURE 2. — Growth forms of

Khadia spp.: A, K. borealis :

long internodes distinguish

this species from all other

Khadia species; B, K. alticola

with tightly packed leaf bases

and internodes not visible.

Scale bar: 10 mm. Drawn by

P. Chesselet.

tipetala specimens with six sepals. Bolus (1922) distin-

guished K. beswickii from K. acutipetala by its nearly

isophyllous habit, glaucous leaves, six-partite flowers

and pale rose petals. Brown (1931) was not able -to dis-

tinguish K. beswickii from K. acutipetala in dried spec-

imens in flower, and emphasized that distinction is pos-

sible only when fruit are available. He described the

capsule of K. beswickii as similar to that of K. acu-

tipetala but with only six locules. Brown (1926) placed

K. beswickii in synonymy under K. acutipetala but rein-

stated it five years later (Brown 1931). Bolus (1936)

considered K. beswickii and K. nationiae (N.E.Br.)

N.E.Br. to be close allies of K. acutipetala but empha-

sized that the number of sepals on all flowers of the

type of K. beswickii was six. Without live material at

his disposal, Brown used the more conspicuous ‘streaks

FIGURE 3. — Leaf anatomy of Kha-

dia: A. K. borealis, Chesselet

& Burgoyne 3, PRE: t/s leaf

epidermal layer with palisade

layer and parenchymatous

cortex, both containing en-

larged idioblast cells filled

with tannins, x 17. B, D, K.

carolinensis, Chesselet &

Burgoyne 5, PRE: B, leaf sur-

face scrape showing anomo-

cytic stomata, x 170; D, detail

of collateral vascular bundle,

x 170. C, K. alticola. Chesse-

let & Burgoyne 4, PRE: detail

of tannin-containing subepi-

dermal idioblast commonly

known as ‘pellucid dots’, x

170.

Bothalia 28,1 (1998)

27

and dots’, possibly phlobaphenes. on the dried petals of

K. nationiae to distinguish it from K. acutipetala , and

seems to have considered this characteristic sufficient

to keep them distinct. He later distinguished K. nationi-

ae from K. acutipetala and K. beswickii by its longer

leaves abruptly rounded to a point in side view, the

longer pedicel and the longer, leaf-like calyx lobes

(Brown 1931).

K. nelsonii is known from a single collection and

Brown (1931) in describing it, considered it to be very

similar to K. acutipetala. This specimen was collected

near Boksburg, on the outskirts of Johannesburg. Its eco-

logical requirements were thought to differ from those of

K. acutipetala as it had been found in a ‘strong, greasy,

loam country'. It displayed elongate growth with longer

leaves, two longer leaf-like sepals, and six stigmata.

K. borealis is the northernmost ranging species in the

genus. It was distinguished by its elongate branches with

long internodes, and the absence of a closing body in the

capsule (Bolus 1936). Until recently, this species had not

been collected since its discovery in the 1930s. The new

taxon of P. Winter mentioned in the introduction, closely

resembles K. borealis, but lacks the distinctive long

internodes on the branches.

TAXONOMY

Mesembrvanthemaceae Fenzl in Annalen des

Wiener Museums der Naturgeschichte 1: 349 (1836),

emend. Herre & Volk in Sukkulentenkunde 2: 38 (1948).

Ruschioideae Schwantes in Ihlenfeldt, Schwantes

& Straka in Taxon 11: 54 (1962), emend. H.E.K. Hart-

mann & Bittrich.

Khadia N.E.Br. in The Gardeners’ Chronicle 88:

279 (in clave) (1930). Type species: K. acutipetala

(N.E.Br.) N.E.Br. [= Mesembryanthemum acutipetalum

N.E.Br.: 407 (1908)].

Perennial dwarf succulent, compact to mat-forming,

more or less anisophyllous. Rootstock thickened, bearing

± horizontal underground ‘branches’, divided at apices

into branches with internodes mostly reduced and

obscure or elongated up to 40 mm with adventitious

roots at nodes (Figure 2A, B). Leaves opposite with 2 (3,

in cultivation up to 4) new leaf growths per branch per

season; connate at base, triquetrous distally with acumi-

nate tips, semiterete basally and broader than thick, dark

green or glaucous; subepidermal enlarged tanniniferous

cells (pellucid dots) concentrated along keel and margins

of leaves, embedded in subepidermal palisade layer and

in parenchymatous pith; stomata anomocytic; vascular

bundles collateral (Figure 3). Flowers solitary, pedicel-

late. Calyx 5-6-lobed, length of calyx lobes variable.

Petals white to deep magenta, with shades of pink.

Staminodes filamentous, white to purple, forming a dis-

tinct cone in newly opened flowers, spreading and bend-

ing backwards in time. Stamens', filaments variable in

colour and length; pollen grains tricolpate, exine struc-

ture semitectate; exine sculpture reticulate; muri thick,

with supratectal spinules (Figure 41). Nectary forming a

dark green ring (lophomorphic). Ovary (4)5-1 1-locular,

upper surface convex or centrally depressed (Figure 5);

placenta parietal; stigmata as many as locules, some-

times longer than filaments, adnate to upper surface of

ovary. Fruit a loculicidal capsule with as many locules as

ovary, reddish when young, dark brown later; pedicel

persistent or breaking off, resulting in a tumble fruit; top

of capsule raised in centre, or flat; base hemispherical to

funnel-shaped; valves without wings but with enlarged

rims, bases of expanding keels widely separated; cover-

ing membranes complete, rather firm, filled ± with a

whitish spongy tissue, with low rims at distal ends; clos-

ing device at distal rim of capsule reduced to a tiny bud

or knob formed by placenta or by an outgrowth of

expanding sheet over some protrusion from endocarp

(Figure 6). Seeds ± 1 mm long; arrangement and orienta-

tion of testa cells of ‘centrospermous’ type, with sinuous

or straight anticlinal walls, covered by granular or rod-

shaped epicuticular ornamentation, micropylar end

curved and elongated or short and straight (Figure

4A-H).

Key to species

la Plants with some trailing stems and intemodes up to 40 mm

long 4. K. borealis

lb Plants with intemodes always shorter than leaves, mostly

obscure:

2a Fruit of a population mostly (6)8-10(1 l)-locular (at least

50 % of all fruit) 1 . K. acutipetala

2b Fruit of a population mostly (5)6-locular (approximately

70 % of all fruit):

3a Rowers white, opening in the afternoon .... 5 . K. carolinensis

3b Flowers pink to magenta, petal bases often paler, open-

ing in sunlight:

4a Leaves 25-40 mm long, glaucous K. beswickii

4b Leaves 1 1-14 mm long, dark green 2. K. alticola

1. Khadia acutipetala ( N.E.Br. ) N.E.Br in The

Gardeners’ Chronicle 89: 279 (1931).

Mesembryanthemum acutipetalum N.E.Br.: 407 (1908). Type: Gau-

teng, near Johannesburg, Turner 4448 (K, holo.!).

Khadia nationiae (N.E.Br.) N.E.Br.: 279 (1931). Mesembry-

anthemum nationiae N.E.Br.: 46, 161, 162 (1926). Type: North-West,

near Rustenburg, Nation s.n. (K, holo.!).

Compact dwarf succulent, variable in degree of aniso-

phylly; rootstock a thick tap root with underground

branches, dense and obliquely to vertically orientated;

aerial branches up to 100 mm long in old specimens, with

persistent old leaf bases. Leaves triquetrous near acumi-

nate tips and semiterete towards base, variable in length

(15-35 mm) and breadth (4—6 mm), glaucous or dark

green. Flower up to 40 mm diam. when open; pedicel

15-50 mm long. Sepals 5 or 6: two longer foliose ones

with acutely pointed tips, one more rounded than the

other, 11-19 mm long and 3 or 4 membranous ones with

long pointed appendages at tips 5.5-10.5 mm long. Petals

in 3 or 4 whorls, deep magenta to light pink; petaloid sta-

minodes in a single whorl, pale pink, ± 8 mm long.

Androecium: staminodes in 2 or 3 whorls; filaments white

or pale pink, inner ones 4.0-7. 5 mm long, middle ones

5. 0-9. 5 mm long, outer ones 8. 0-9. 5 mm long (Figure

4A), all papillate but outer ones only slightly so; anthers

rich yellow; pollen faintly sweetly scented. Ovary with

(6)8-10(11) locules, roofs of locules rising sharply; stig-

mata as many as locules, subulate, 3-8 mm long, growing

with age, tips recurved. Fruit 8-12 mm diam. with flat-

28

Bothalia 28,1 (1998)

FIGURE 4. — Micromorphological characters of the seed (A-H) and pollen (I) of Khadia: A, K. acutipetala , Chesselet & Burgoyne I , PRE, x 46;

B, K. alticola, Chesselet & Burgoyne 4 , PRE, x 41; C, K. borealis, Chesselet & Burgoyne 3 , PRE, x 53; D, K. carolinensis, Chesselet &

Burgoyne 5, PRE, x 34; E, K. acutipetala, Chesselet & Burgoyne I, PRE, x 256; F, K. beswickii Ross Frames 17256, BOL, x 277; G, K

borealis Chesselet & Burgoyne 3 , PRE, x 2705; H, K. carolinensis, Chesselet & Burgoyne 5, PRE, x 2800; I, K. alticola, Chesselet &

Burgoyne 4, PRE, x 2050.

tish, hemispherical, smooth-walled, shallow undersurface

and a prominent raised lid; placenta parietal, extending to

base of capsule; covering membranes opaque with

recurved distal margins, raised centrally at columella or

flat and folding centrifugally; valves wedge-shaped, curl-

ing outwards with ridge along centre of valve on inner

surface; large phlobaphene-containing cells situated near

tips of valves on inner side; ‘closing bodies’ minute, pla-

cental, or callous-like, with numerous funicles arising

from them, or rod-like, and well defined; pedicel breaks

off when fruit are ripe, forming tumble fruit, Hat base

facilitates upright position of fruit when detached from

mother plant (Figure 6A-D). Seeds from capsules with

centrally raised covering membranes mostly red, from

capsules with flattish covering membranes mostly brown-

ish, often quite pale; with straight testa cell walls, rodlet-

shaped epicuticular sculpturing, micropylar end curved

and elongated (Figure 4A, E).

On the Magaliesberg K. acutipetala occurs in rocky

grassland near summits (1 600-1 700 m) in well-drained,

loamy or gritty sand derived from Magaliesberg

quartzites. Populations are found in full sun on gentle

north-facing slopes in sparse grassland vegetation domi-

nated by Tristachya biseriata. Urbanisation threatens this

species in a number of habitats — the Hospital Hill popu-

lation near the city centre of Johannesburg has long since

disappeared and the Northcliff Ridge populations in the

suburbs of Johannesburg, are presently threatened by

habitat destruction. K. acutipetala is, however, fairly

abundant in existing nature reserves and other fortuitous-

ly protected areas in Gauteng (Figure I ).

Note; the K. acutipetala species complex is charac-

terised by morphological variation between and within

populations, particularly of the capsule, but also of leaf

colour and shape, seed and flower colour, and length of

foliose sepals. Although differences may seem quite

striking, close examination of individuals reveals a con-

tinuous gradient of character combination, making it

impractical to define groups.

Vouchers; Behr 781 (PRE); Bredenkamp 875 (PRE); Chesselet &

Burgoyne 1, 2 (PRE).

2. Khadia alticola Chesselet & H.E.K. Hartmann

in Aloe 32: 48 (1995). Type: Mpumalanga, Steenkamps-

berg, Hartmann & Chesselet 32729 (PRE, holo. ! ).

Bothalia 28,1 (1998)

29

FIGURE 5. — Floral morphology of

Khadia'. A, K. acutipetala,

Chesselet & Burgoyne 1,

PRE; B, K alticola, Ches-

selet & Burgoyne 4, PRE; C,

K. borealis, Chesselet & Bur-

goyne 3, PRE; D, K. caroli-

nensis, Chesselet Bur-

goyne 5, PRE. Scale bar; 5

mm. No material of K. bes-

wickii was available. Drawn

by P. Chesselet.

Mat-forming, anisophyllous, succulent perennial with

main tap root and multiple branching occurring just

under soil surface, adventitious roots produced at nodes;

new growth develops at periphery of mats anisophyllous.

Leaves triquetrous, 8.8-16.7 x 5. 3-6. 8 mm, bright green

to dark green; old leaf bases persist longer than in other

Khadia species (Figure 2B); usually two leaf pairs per

seasonal growth and a maximum of 3 in habitat; up to 4

in cultivation. Flowers up to 40 mm diam. when open.

Petals pink fading towards base with age. Sepals 6.

Androeciunv. filamentous staminodes present; outer row

of filaments 4.5 mm long, middle row 2.3 mm long and

inner row 1.6 mm long (Figure 5B). Ovary with 6

locules; stigmata as many as locules, 4. 9-6.0 mm long;

placenta basal to parietal. Fruit opening incompletely

and not closing again completely; closing bodies very

small and brown, spongy tissue only in distal third of

covering membranes, ending in a rather sharp ledge

(Figure 6E-G). Seeds'. 15 or 16 seeds per locule; testa

cells isodiametric to slightly elongate with sinuous anti-

clinal walls covered with granular epicuticular ornamen-

tation; micropylar end straight, hilum aligned parallel

just below it (Figure 4B).

K. alticola is common at altitudes above 2 000 m in

Mpumalanga and KwaZulu-Natal in shallow, sandy,

humus-rich soil pockets and crevices between rock plates

(Figure 1 ). At the Steenkampsberg site it occurs among

Flelichrysum galpinii in bouldery patches together with

the geophyte Brachystelma stellatum, as well as grasses

and forbs. These eastern populations are exposed to a

high rainfall regime and mists.

Vouchers: Archer 1050 (PRE); N.J. Devenish 692 (PRE); Hartmann

& Chesselet 32729 (PRE); Hilliard & Burn 9173 (BOL).

3. Khadia beswickii (L.Bolus) N.E.Br. in The Gar-

deners’ Chronicle 89: 279 (1931).

Mesembryanthemum beswickii L.Bolus: 124 (1922). Type: Gauteng,

near Johannesburg, Beswick NBG 218/20 (BOL, holo.!).

Khadia nelsonii N.E.Br.: 279 (1931 ). Type: Gauteng, Witte Koppies

near Boksburg, Nelson s.n. (K, holo.!).

Mat-forming, anisophyllous, succulent perennial with

short root system. Leaves weakly, triquetrous, roundish

below, glaucous. Flowers with petals pink, white at base,

often quite pale. Fruit with 6 locules, base of capsule

bell-shaped and top low, rounded; rims of valves turned

backwards, covering membranes completely spongy,

distal closing ledge very inconspicuous to absent; clos-

ing body consisting only of placenta and a layer of

expanding sheet tissue. Seeds with anticlinal walls of

testa cells sinuous (Figure 4F); testa cells covered with

granular epicuticular ornamentation.

K. beswickii is known from Nigel on the East Rand in

Gauteng where it was collected in 1942 by A. A. Mauve

at two localities on outcrops of the Witwatersrand

Supergroup (Figure 1). The first, on the north side of the

Spaarwater Dam, where it occurred together with

Lithops lesliei , is now inaccessible due to the establish-

ment of an informal settlement at the site. Whether these

plants have survived there is uncertain. Specimens were

collected by H.E.K. Hartmann at this site during the

1980’s. At the second locality, outside Nigel, on the road

30

Bothalia 28,1 (1998)

FIGURE 6. — Capsule morphology of Khadia: A-D, K. acutipetalu, Chesselet & Burgoyne 1, PRE; E-G, K. alticola, Chesselet & Burguyne 4,

PRE; H-J, K borealis, Chesselet & Burgoyne 3, PRE; K-M, K. carolinensis, Chesselet & Burgoyne 5, PRE. Scale bar: 5 mm. No mater-

ial of K. beswickii was available. Drawn by P. Chesselet.

Bothalia 28.1 (1998)

31

to Balfour, we noted large holes in microhabitats where

plants may have grown, indicating possible harvesting.

The species may be under threat of extinction, or extinct.

Unlike the other Khadia species, which have been intro-

duced into cultivation, live plants of this species were not

available for study or propagation.

Vouchers: Mauve s.n. (PRE). Ross Frames 17256 (BOL, PRE).

4. Khadia borealis L. Bolus in Notes on Me-

sembryanthemum and allied genera 3: 6 (1936). Type:

Northern Province, Ventersdorp Farm. Crundall s.n. (BOL,

lecto!, here designated); Northern Province, Zout-

pansberg, Franzhoek Farm, Vogts s.n. BOL 21638 (BOL,

paralecto.!).

Perennial, slightly anisophyllous succulent with two

kinds of branches: 1. creeping growth with rosettes at

nodes on branches with distinct internodes, 1 5 — 40 mm

long; and 2, tufted growth in branches where internodes

are reduced; old leaf bases persisting; root system with

thick tap roots and adventitious roots arising at nodes

(Figure 2A). Leaves trigonous in cross section, with pel-

lucid dots concentrated along margins, glaucous, dark

green, leaf tips taper abruptly into hyaline point; leaf

pairs curve in same direction. Flowers small, ± 15 mm

diam. Petals white or pink. Calyx : with 5 sepals. 2 long

and 3 short with membranous margins. Androecium :

petaloid staminodes present; filaments in 3 rows, outer,

3. 5-5.0 mm long, papillate or epapillate, middle, ± 3.6

mm long, inner. 2. 1-2.7 mm long (Figure 5C); nectary

dark green, continuous and crenulate. Ovary with

(4)5— 7(8) locules, stigmata as many as locules, ± 3.5 mm

long. Fruit with lower part distinctly concave, top rather

low, rims recurved, placenta forms protrusions or pla-

cental ‘closing body’, closing ledges present (Figure

6H-J). Seeds orange; testa cells more or less isodiamet-

ric to elongate with smooth, convex periclinal walls;

anticlinal walls straight; epicuticular ornamentation con-

sisting of rodlets; micropylar end elongate with hilar

bulge small and indistinct.

In the Northern Province, the creeping K. borealis

grows in shallow soil pockets along flat lying strata with-

in the mist belt at approximately 1 400 m, near the sum-

mit of the Soutpansberg (Figure 1). It grows in full sun,

in grassland vegetation.

Vouchers: Chesselet & Burgoyne 3 (PRE). Hahn 430. 539, 604 (Herb.

Sout.).

5. Khadia carolinensis (L.Bolus) L.Bolus in Notes

on Mesembryanthemum and allied genera 3: 6 (1936).

Rabiea carolinensis (L.Bolus) N.E.Br.: 53 (1931). Mesembry-

anthemum carolinense L.Bolus: 128 (1922). Type: Mpumalanga,

Carolina. F.A. Rogers s.n. NBG 1833/17 (BOL, lecto.!, here designat-

ed); Mpumalanga, Carolina, C.L. Leipoldt s.n. NBG 2819/17 (BOL,

paralecto.!).

Plants very slightly anisophyllous, forming large mats

up to 1 m diam.; thick tap roots branching underground;

stems thickly covered with dead leaf bases. Leaves sub-

terete to outside, flattened on inner side, dark green.

Flowers ± 35 mm diam., opening in afternoon; pedicels

±10 mm long. Petals white (at times recorded as yellow

owing to slight yellow tinge on outer surface), in 3 rows,

± 10 mm long. Calyx with 6 sepals, foliose and membra-

nous, of similar length. Androecium: petaloid staminodes

± 6.5 mm long; stamens ± 7 mm long, in 3 tiers, outer

epapillate ± 8 mm, middle papillate at base ± 6 mm,

inner papillate 5 mm (Figure 5D). Ovary’ with 6 locules;

stigmata as many as locules, subulate, ± 7.5 mm long,

protruding beyond stamens, orange. Fruit with broad

closing ledges and tent-shaped covering membranes;

closing bodies large (Figure 6K-M). Seeds: testa cells

with sinuous anticlinal walls; epicuticular ornamentation

granular; micropylar end curved inwards towards

upward curving hilar bulge (Figure 4D, H).

K. carolinensis is plentiful in sandy loam among

quartzite rocks near Carolina in Mpumalanga, at alti-

tudes of ± 1 740 m (Figure 1).

Vouchers: Burgoyne 2388 (PRE); Du Toit 390 (PRE); Strey 3426

(PRE).

DISCUSSION

The growth form of Khadia, with a thick underground,

upward-branching root system and a mat of succulent

leaves above ground is not uncommon in other mesem

genera such as Hereroa (Schwantes) Dinter & Schwantes

and Ebracteola Dinter & Schwantes. Khadia is formally

classified in the now outdated subtribe Nananthinae

Schwantes, comprising plants forming ‘rosettes or

clumps', and including Nananthus N.E.Br., Aloinopsis

Schwantes, Titanopsis Schwantes, Khadia N.E.Br. and

Rabiea N.E.Br. (Herre 1971). Bolus (1958) considered

Khadia to be closely allied to typical Ruschia Schwantes,

differing in the variable number of stigmata and in the

absence of bracts, and in some species by the ‘placental

tubercle’. The character combination present in Khadia

makes its systematic placement difficult amongst the rus-

chioid genera (Hartmann 1991. 1993). It was tentatively

placed in the Ruschia- Group and thought to be closest to

Acrodon (Hartmann 1991), which is endemic or near

endemic to the Little Karoo Centre (Hartmann 1994).

Recent examinations of the ‘closing body', the structure

that blocks the opening of the locules in the capsules of

this genus, have shown that it is not homologous with

such structures recognised in other genera (Chesselet &

Hartmann 1995). In a detailed study of capsules of the

‘Mesembryanthema’, Hartmann (1988) distinguished

endocarpal closing bodies from similar structures derived

from the placenta. In the case of Khadia the so-called

‘closing bodies’ are of placental or more complex com-

position. Due to the unusual nature of this type of closing

device, further investigation is required before a decision

can be taken regarding the systematic affinities of

Khadia. Our present postulate, based on capsule charac-

teristics, is that its affinities may lie with Ebracteola.

Although the genus concept of Khadia has not been

critically evaluated, it is generally considered sufficient-

ly distinct from that of other mesem genera. Internal fea-

tures of the capsule have been highlighted as being typi-

cal for the genus (Chesselet & Hartmann 1995). The

composition of the ‘closing body’ is the single most

important of these features and has been used to distin-

guish species in the group. In her studies of leaf anatomy

in the mesems, Dupont (1968) suggests that in a well-

32

Bothalia 28, 1 (1998)

defined genus all species have the same type of stomata,

or proportions of the different types. In Khadia all

species have anomocytic stomata, which is the primitive

state in the mesems ( K . beswickii was not investigated

anatomically as we had no fresh material). Nevertheless,

there is no single apomorphy that can serve to delimit

Khadia. Hence the cohesion of Khadia is unashamedly

one of convenience, based on shared characteristics of

uncertain homology and favoured by the need for taxo-

nomic stability. Recently, Chesselet & Hartmann (1995)

separated Khadia into two groups: Group 1 which

included K. acutipetala and Group 2 which included K.

beswickii, K. borealis and K. carolinensis. Data present-

ed here suggest otherwise (Table 1). We propose two or

possibly three groups which may represent subgenera or

even separate genera: 1 , The K. acutipetala species com-

plex excluding or including 2, K. borealis, and 3, the dis-

tinct eastern group which includes K. beswickii, K. alti-

cola and K. carolinensis, and is united by features of the

flower, capsules and seed. K. borealis shares seed char-

acters, particularly the shape and ornamentation of testa

cells, with K. acutipetala but the capsule of K. borealis is

distinct (Figure 6).

Flower, fruit and seed morphology have been useful

sources of taxonomic information at the species level and

in establishing informal groups within Khadia. Further

investigations into capsule and floral anatomy, as well as

chemical characterisation may elucidate some of the

problems of delimitation outlined here but are unlikely to

make taxonomic decisions any easier.

USES

'Khadi' is a Tswana/South Sotho name for beer

brewed traditionally using the fleshy roots of a variety of

TABLE 1. — Character distribution in Khadia

Bothalia 28,1 (1998)

33

taxa. A number of mesem species, including Delosperma

mahonii (N.E.Br.) N.E.Br. and Trichodiadema stellatum

(Mill.) Schwantes are reported to have been used in

brewing alcoholic beverages (Juritz 1906; Brown 1926).

Researching the beer making process, Juritz (1906) pre-

dicted that any starchy root would suffice as long as

mouldy dough could be made of it. It is of interest to note

that brewing or possession of these beers was punishable

in terms of the Liquor Law amendment act, No. 28 of

1898 (Juritz 1906). Whether the roots of K. acutipetala

are still used today is uncertain. Khadia is reported to be

used medicinally by Manyika people of the Umtali

District of Zimbabwe (Watt & Breyer-Brandwijk 1962)

suggesting the distribution of this genus beyond South

Africa into Zimbabwe.

SPECIMENS EXAMINED

Archer 1050 (2) PRE; 2093 (4) PRE.

Behr 781 (1) PRE. Beswick s.n. NBG 218/20 (3) BOL. Bredenkamp

875 (1) PRE. Bryant D63, D79 (1) BOL. Burgoyne 2388 (5)

PRE.

Chesselet & Burgoyne 1 , 2 (1) PRE; 3 (4) PRE; 4 (2) PRE; 5 (5) PRE.

Codd 524 (5) PRE; s.n. PRE 55073 (2) PRE; s.n. (2) BOL.

Crandall s.n. (4) BOL.

Devenish 692 (2) PRE. Du Toit 390 (5) PRE.

Fuller 83 (1) BOL.

Gulp in 21659. s.n. (5) BOL; s.n. NBG 1278/31 (1) BOL. Gilfillan 90

(1) PRE. Gilmore 237 (1) BOL.

Hahn 430. 539, 604 (4) (Herb Sout). Hardy 441 (1 ) PRE. Hartmann &

Chesselet 32729 (2) PRE. Hilliard & Bunt 9173 (2) BOL.

Jacobsen 756 (1) PRE.

Leendertz. 2462 (1) BOL; 7714 (1) PRE; 9166 (5) PRE. Liebenberg

12332 S.U.G (5) BOL; 12334, s.n., s.n. (1) BOL.

Mauve s.n. (3) PRE. Mogg 14778 , 14902 (1) BOL. PRE.

Nation 372 (1) BOL; s.n. (1) K. Nelson s.n. (3) K.

Obermeyer s.n (1) PRE.

Repton 444 (1) PRE. Rogers s.n. NBG 1833/17 (5) BOL. Rose Innes

210 ( 1) PRE. Ross Frames 17256 (3) BOL. PRE.

Steyn 839 (5) BOL. Strey 3426 (5) PRE.

Thode A1317 (1) PRE. Turner 4448 (1) K, PRE.

Van Jaarsveld 14234 (4) PRE. Vogt s.n. (4) BOL. Vogts 21534. s.n. (1)

BOL; s.n. PRE 55056 (1) PRE.

Watson-Dobie 414, s.n. (1) BOL.

Young 3019, 3054 ( 1) PRE.

ACKNOWLEDGEMENTS

We thank the Curators of BOL and Kew for loan of

type material to PRE, Bruce Bayer and Dr Otto Leistner

for valuable comments on the manuscript, Dr Sarie

Perold for SEM micrographs, Antoinette Botha for

anatomical preparation of material, Adela Romanowski

for photographic work, Marinda Koekemoer for encour-

aging two of us (P.C & P.B.) in our fieldwork and Tony

Hitchcock for cultivating and propagating the live plant

material. The map was generated by Hannelie Snyman

using MAPPIT (available from NBI, Pretoria).

REFERENCES

BOLUS, H.M.L. 1922. Novitates africanae. Annals of the Bolus

Herbarium 3; 124—128.

BOLUS, H.M.L. 1936. Notes on Mesembryanthemum and allied gen-

era, part 3: 6-11. University of Cape Town, Cape Town.

BOLUS, H.M.L. 1958. Khadia beswickii. The Flowering Plants of

Africa 32: t. 1263.

BROWN, N.E. 1908. Diagnoses africanae: XXV. Kew Bulletin 1908 :

407.

BROWN, N.E. 1926. Ficoidaceae (Vyg-bosje Family). In J. Burtt

Davy, A manual of flowering plants and ferns of the Transvaal

with Swaziland, South Africa. 159-163. Longmans, Green,

London.

BROWN, N.E. 1930. Mesembryanthemum and some genera separated

from it. The Gardeners' Chronicle 88: 278, 279.

BROWN, N.E. 1931. Mesembryanthemum and some genera separated

from it. The Gardeners' Chronicle 89: 53, 279.

CHESSELET, P. & HARTMANN, H.E.K. 1995. Khadia alticola

Chess. & H.E.K. Hartm., spec. nov. (Mesembryanthema, Aizoa-

ceae). Aloe 32: 46-49.

DUPONT, 1968. Epidermes et plantules des Mesembryanthemacees.

Systematique. Evolution. Bulletin de la Societe d'Histoire

Naturelle de Toulouse 104: 7-64.

FENZL, E. 1 836. Monographic der Mollugineen und Steudelieen,

zweier Unterabtheilungen der Familie der Portulacaceen.

Annalen des Wiener Museums der Naturgeschichte 1: 349.

HARTMANN, H.E.K. 1988. Fruit types in Mesembryanthema.

Beitrdge zur Biulogie der Pflanzen 63: 313-349.

HARTMANN, H.E.K. 1991. Mesembryanthema. Contributions from

the Bolus Herbarium 13: 75-157.

HARTMANN, H.E.K. 1993. Aizoaceae. In K. Kubitzki, J.G. Rohwer

& V. Bittrich, The families and genera of vascular plants 2:

37-69. Springer Verlag, Berlin.

HARTMANN, H.E.K. 1994. On the phytogeography and evolution of

Mesembryanthema (Aizoaceae). In J.H. Seyani & A.C.

Chikuni, Proceedings of the XllI'1' Plenary Meeting AETFAT.

Malawi 2: 1165-1180.

HARTMANN, H.E.K. & BITTRICH, V. 1991. Typification of supra-

generic names — some nomenclatural changes in Aizoaceae.

South African Journal of Botany 57: 74.

HERRE, H. 1971. The genera of the Mesembryanthemaceae. Tafel-

berg. Cape Town.

HERRE, H. & VOLK, O H. 1948. Mesembryanthemaceae Herre et

Volk, familia nova Sukkulentenkunde 2: 38.

IHLENFELDT, H.-D., SCHWANTES, G. & STRAKA, H. 1962. Die

hoheren Taxa der Mesembryanthemaceae. Taxon 11: 52-56.

JACOBSEN, H. 1960. A handbook of succulent plants 3. London.

JACOBSEN, H. 1981. Das Sukkulentenlexikon. Gustav Fischer,

Stuttgart.

JURITZ, C.F. 1906. Kaffir beers, their nature and composition. The

Agricultural Journal of the Cape of Good Hope 48: 35-47.

WATT, J M. & BREYER-BRANDWIJK, M.G. 1962. Medicinal and

poisonous plants of southern and eastern Africa. Livingstone,

London.

.

Bothalia 28,1: 35-39 (1998)

FSA contributions 11: Zingiberaceae

R.M. SMITH*

Editor’s note: Hedychium coronarium and Siphonochilus kirkii have been added by Mrs C. Archer, with the author’s consent.

Rhizomatous herbs; rhizome usually fleshy, sympodi-

al, each element terminating in a leafy or flowering

shoot. Leaf shoots with few to many blades, either

arranged spirally and with tubular sheaths (Costoideae)

or distichously arranged, sheaths then usually open on

side opposite lamina (Zingiberoideae). Inflorescence

occasionally surrounded by sterile bracts, terminal on

leaf shoot or borne directly on rhizome at base of leaf

shoot or remote from it. Flowers zygomorphic, her-

maphrodite or (rarely) unisexual, solitary' in axils of

bracts or in cincinni, with or without bracteoles. Calyx

tubular, often unilaterally split. Corolla tube usually

slender; petals 3, often subequal. Labellum : (anterior sta-

minode) often large and showy; lateral staminodes usu-

ally present, either as conspicuous petaloid organs

(Zingiberoideae/Hedychieae) or as small subulate teeth

or swellings (Zingiberoideae/Alpineae). Fertile stamen

one only; anther more or less sessile or with a distinct fil-

ament; thecae parallel or slightly divergent, connective

sometimes developing into an appendage (anther crest).

Ovary’ inferior, unilocular with parietal placentation or

trilocular, or incompletely so with axile placentation;

style filiform, upper part usually held between thecae;

stigma usually expanded. Epigynous glands (stylodes)

forming erect outgrowths on top of ovary in Zingi-

beroideae; in Costoideae, three septal nectary glands

arise towards top of ovary. Fruit a dehiscent capsule or

fleshy berry. Seeds arillate.

Over 40 genera and about 1000 species occurring

mainly in the tropics of the Old World but with some rep-

resentatives in South and Central America. Four genera,

all belonging to the subfamily Zingiberoideae, are found

in the African continent but only one, Siphonochilus

(tribe Hedychieae), is native in the FSA area. Three

species of Hedychium (tribe Hedychieae) and Alpinia

zerumbet (tribe Alpineae) have become naturalised in cer-

tain areas.

The Zingiberaceae is notable for its spice plants and

includes ginger. Zingiber officinale', cardamom, Elettaria

cardamomum ; and turmeric, Curcuma longa. Cultivated

races of Curcuma are common in the Indian markets of

KwaZulu-Natal and are not infrequent in gardens; they

are used medicinally or powder from the tubers is used

as a cosmetic.

la Inflorescence borne separately from leaf shoot; flowers pur-

ple-pink, blotched with yellow in centre of labellum,

sometimes unisexual 1. Siphonochilus

lb Inflorescence terminal on leaf shoot; flowers yellow and

white or red and yellow, hermaphrodite:

* Royal Botanic Garden, Edinburgh, UK.

MS. received: 1985-03-14.

2a Inflorescence erect; corolla tube 50-90 mm long, slender;

lateral staminodes petaloid 2. Hedychium

2b Inflorescence pendulous; corolla tube under 20 mm long;

lateral staminodes reduced to small subulate teeth . . .

3. Alpinia

1. SIPHONOCHILUS

Siphonochilus J.M.Wood & Franks in Medley

Wood, Natal plants 6,3: t. 560, 561 (1911a); J.M.Wood &

Franks: 274 (1911b); B.L.Burtt: 369, t. 1 (1982). Type

species: S. natalensis J.M.Wood & Franks [= S. aethiopi-

cus (Schweinf.) B.L.Burtt].

Cienkowskia Schweinf.: t. 1 (1867), non Regel & Rach:

48 (1859).

Kaempferia subgen. Cienkowskia K.Schum.: 67 (1904).

Cienkowskiella Y.K.Kam: 8 (1980).

Inflorescence borne separately from leaves, some-

times precocious, long pedunculate or peduncle very

short. Flowers usually hermaphrodite, rarely unisexual,

each subtended by a bract; bracteoles absent. Calyx 3-

lobed. unilaterally split. Petals linear-lanceolate. Label-

lum large, showy, connate with petaloid lateral stamin-

odes for at least half its length. Stamen', anther crest

petaloid, often longer than parallel thecae. Stigma cup-

shaped or two-lipped. Epigynous glands short, stub-like.

Ovary trilocular with axile placentation.

Distributed from KwaZulu-Natal northwards to

Ethiopia and the Nile lands and across the continent to

Nigeria and the Gambia; also in equatorial West Africa.

About 20 species have been described but the genus is

badly in need of revision and the number is probably

rather less. Two species are recorded from southern Africa.

Siphonochilus was raised partly on account of the

unisexual flowers found in Kaempferia natalensis. This

in itself is insufficient to warrant generic distinction but

recent research has shown the separation of the African

species from Kaempferia to be entirely justifiable.

Spearing & Mahanty (1964) report that the African

plants have a basic chromosome number of 14, that of

Asiatic Kaempferia is 11. Morphologically, Siphono-

chilus differs in the absence of bracteoles, and in hav-

ing lateral staminodes that are connate to the labellum

for about half their length, non-ciliate rimmed stigma

and stub-like rather than needle-shaped epigynous

glands.

Key to species

Flowers on an elongated peduncle 2. S. kirkii

Flowers on very short peduncle concealed by basal bracts . . .

1.5. aethiopicus

36

Bothalia 28,1 (1998)

m s.

FIGURE 1 . — Siphonochilus aethiapicus : A, habit, x 0.8; B, root tuber, x 0.8; C, calyx, x 0.8; D, labellum and lateral staminodes spread out show-

ing position of stamen, x 0.6; E, stamen, x 1 .2; F, stigma, x 3.5; G, I s. of base of corolla tube and ovary, showing thickened walls of tube

and epigynous glands, x 1.2; H, ovary in t.s., x 2; I, female flower, x 0.8. A-H, from Hilliard & Burtt 6884A, Cult. R.B.G. Edinburgh; I,

redrawn from Wood & Franks, t. 560.

1. Siphonochilus aethiopicus (Schweinf) B.L.

Burtt in Notes from the Royal Botanic Garden Edinburgh

40; 372 (1982); Lock: 20 (1985). Syntypes: Ethiopia; am

Khor el Sherif und bci Famaka in Fesoghlu, 23-V-1848,

Cienkowski ; an der Gandua zwischen Wochni und

Metemme, 9-vi- 1 862, Steudner, bei Wochni im nordwest

Abyss., 3 1 -v- 1 862, 3-vi- 1 862, Steudner (K!).

Cienkowskia aethiopica Schweinf.; t. 1 (1867).

Kaempferia aethiopica (Schweinf.) Benth.: 642 (1883).

K. ethelae J.M.Wood: 94, t. 34 (1898); Wright: 314 (1913); Marloth:

167 (1915). Type: Cult. B.G. Durban, xi-1897, V1W7667(NH), orig-

inally from Mozambique, Vila de Manica (Massikessi), Benningfield.

K. natalensis Schlecht. & K.Schum. in K.Schum.: 72, fig. lOe, f

( 1 904); J.M.Wood & Franks: 1 1 2 ( 1 9 1 1 c); Wright: 315(1913); Marloth:

167, t. 53 (1915). Type: KwaZulu-Natal, Inanda, comm, viii-1879,

Wood 544 (K!).

Siphonochilus natalensis (Schlecht. & K.Schum.) J M.Wood &

Franks: t 560, 561 (1911a); J.M.Wood & Franks: 274 (1911b); B.L.

Burtt: 372, t. 1 (1982).

Cienkowskiella aethiopica (Schweinf.) Y.K.Kam: 10, t 3 (1980).

Bothalia 28,1 (1998)

37

Rhizome aromatic; fibrous roots bearing narrowly

elongate tubers. Leaf shoots 300-1000 mm. Leaves

4-12, sessile, 300-400 x 50-90 mm (those at base of leaf

shoot smaller), narrowly lanceolate, acuminate,

glabrous; ligule 3-10 mm, membranous, entire; sheaths

striate, glabrous. Inflorescence borne separately from

leaf shoot, often precocious, basal part embedded in soil.

Flowers 2-6, on a 10-20 mm bract-covered peduncle;

floral bracts 25-30 x ± 15 mm, obtuse, glabrous; pedi-

cels 10-20 mm long. Hermaphrodite flowers : calyx

20-30 mm. unilaterally split, shallowly 3-lobed, lobes

rounded, sometimes with aristate point. Corolla tube

white, 30-40 mm long, thick-walled. Petals white with

pink tips, 60-80 mm long, lanceolate-acuminate. La-

bellum: purple-pink, yellow at throat in centre, free part

± 60-80 x 60-70 mm, bifid for up to 2 3/4 of its length,

lower part connate with lateral staminodes for 50-60 mm

and encircling stamen (‘split-tube’ of Wood & Franks);

lateral staminodes (the free part) 60-80 x 40-50 mm.

erect. Stamen 50-80 mm; filament 10-15 mm; thecae

15-20 mm, parallel; connective prolonged into a 30-50

mm petaloid, irregularly dentate reflexed crest. Stigma

more or less cup-shaped. Epigynous glands 3-4 mm,

stub-shaped, often embedded in fleshy corolla wall.

Ovary 15x6 mm. glabrous, trilocular with axile placen-

tation. Female flowers: calyx, corolla tube and petals as

above. Lateral staminodes and labellum connate into an

80 mm closed tube; segments 4-6, 2 or 3, alternate ones

40 x 20 mm, oblong obtuse, the other 2 or 3, 40 x 5-7

mm, narrower. Stamen absent. Gynoecium as in her-

maphrodite flower. Fruit unknown. Figure 1.

The description of the female flower is taken from

Wood & Franks (1911a).

Occurs in Northern Province, Mpumalanga,

Swaziland and KwaZulu-Natal (Figure 2). Widespread

throughout tropical Africa. It seems likely that the

species never occurred naturally in the Flora area but

that it was introduced from tropical Africa and widely

cultivated (Williams et al. 1996). In 19th century Natal,

S. aethiopicus was often dug up and the tubers sold as

horse medicine; nowadays it is found only in the vicini-

ty of Zulu dwellings, where it was presumably original-

ly planted. In traditionally Zulu practice the plant is used

in the treatment of colds and chest complaints, to ward

off snakes and lightning, and as a remedy for malaria. It

is known as the Natal Ginger, Sherungulu , Indungulu.

Vouchers: Hilliard & Burn 68S4A (E); Holt, NH 28507 (NH); Medley

Wood , NH 11250 (NH).

S. aethiopicus is a very polymorphic species, size,

colour and depth of the lobing of the labellum may vary

within a single population and considerable variation

occurs in tuber length and in the size of the ligule.

The type plant of Kaempferia natalensis had only

female flowers; Wood & Franks (1911a), when estab-

lishing Siphonochilus , recorded hermaphrodite flowers

to be rare. In other respects the Natal plants cannot be

separated from their more northerly counterparts.

2. Siphonochilus kirkii (Hook.f) B.L.Burtt in

Notes from the Royal Botanic Garden Edinburgh 40: 372

(1982); Lock: 15, t. 4 (1985). Type: Tanzania, cult, at

Kew, Kirk s.n. (K, holo.).

Cienkowskia kirkii Hook.f.: t. 5994 (1872).

Kaempferia kirkii (Hook.f.) Wittm. & Perring: 57, t. 1364 (1892);

Baker: 294(1898); K.Schum.: 68 (1904).

Cienkowskiella kirkii (Hook.f.) Y.K.Kam: 11 (1980).

K. rosea Baker: 295 (1898); F.W.Andrews: 255 (1956). Syntypes:

Sudan, Schweinfurth 1946 (K, PRE!) and numerous other specimens

from eastern Africa.

K. montagui F.M.Leight.: 57 (1932). Syntypes: Zimbabwe, nr Mazoe

[= Mazowe], Montagu s.n. sub NBG888/21 (NBG); Wise s.n. sub

NBG33/26 (NBG).

Roots bearing small fusiform tubers ± halfway along

length. Leaf shoots 200-400 mm. Leaves 5-7, glabrous;

lamina undeveloped in basal 1-3 or ovate to elliptic,

acuminate, up to 170-310 x 55-100 mm, tapering at base

into a false petiole up to 250 mm long (including leaf

sheath); ligule obsolete; sheaths sulcate when dry.

Inflorescences 1^1, borne separately from leaf shoot;

7-15(-20)-flowered; peduncle 200-350 mm long, terete,

glabrous; bracts greenish, oblong to narrowly obovate,

obtuse, the lower up to 65 x 20 mm, the upper very much

smaller, up to 22 x 12 mm. Flowers hermaphrodite.

Calyx campanulate, 8-15 mm long, shallowly 3-lobed,

each lobe with a subterminal subulate projection ± 1 mm

long. Corolla tube ± 8 mm long; petals narrowly obovate

to narrowly oblong, acute, 22-26 mm long, whitish,

tinged with green or mauve. Labellum 3-lobed, lateral

lobes rhomboid, mauve, ± 30 x 20 mm, median lobe

broadly spathulate, emarginate, ± 45 x 45 mm, mauve

with a central yellow mark with or without a dark purple

mark on each side. Stamen ± 25 mm long; thecae ± 5 mm

long, curved; connective prolonged into oblong petaloid

crest, ±15x6 mm. Ovary ± 6 mm long, trigonous; stig-

ma peltate. Fruit obovoid, trigonous, winged at angles,

crowned with remains of calyx. Seed whitish, trigonous,

± 5.5 x 3.0 mm, with basal elaiosome.

Note: the above description is based on Lock (1985).

Occurs in the Caprivi Strip, Namibia (Figure 2).

Widespread throughout tropical Africa.

Vouchers: Hardy 7110 (PRE); Killick & Leistner 3016 (PRE).

FIGURE 2. — Distribution of Siphonochilus aethiopicus, O; S. kirkii,

•; and Hedychium coronarium, ■

38

Bothalia 28,1 (1998)

2. HEDYCHIUM J.Konig

Hedychium J.Konig in Retz., Observationes botani-

cae 3: 73 (1783); Benth. & Hook.f.: 642 (1883); K.Schum.:

40 (1904). Type species; H. coronarium J.Konig.

Plants up to 3 m tall. Inflorescence terminal on a frond-

like leaf shoot; bracts broad and imbricating and conceal-

ing main axis or narrow and enfolding flowers. Flowers

1-6 per bract, each subtended by a tubular bracteole.

Corolla tube long, slender, usually exceeding calyx. Petals

strap-shaped. Labellum : narrowed at base with a conspic-

uous, usually bilobed limb; lateral staminodes petaloid;

filament usually long and slender; anther connective

ecristate. Ovary trilocular with axile placentation. Fruit

globose or oblong, often bright red or orange within.

About 45 species, mainly eastern Himalayan, but a

few known from southern India, Thailand, Malaysia and

Indonesia. Several are widely cultivated throughout trop-

ical and subtropical zones and are valued for their per-

fume and prolific flowers.

Key to species

la Bracts broad and imbricating, concealing main axis; petaloid

parts white or yellow; filament white or yellow:

2a Flowers yellow; stamen longer than labellum

2. H. flavescens

2b Rowers pure white, base of labellum usually yellow-

green; stamen shorter than labellum I . H. coronarium

lb Bracts narrow, not concealing main axis; petaloid parts yel-

low; filament bright red 3 H. gardnerianum

1. Hedychium coronarium J.Konig in Retz., Ob-

servationes botanicae 3: 73 (1783); Sims: t. 708 (1804);

K.Schum.: 44 (1904). Type: Malaya, Konig s.n. (f).

Plant up to 3 m. Leaves sessile, up to 600 x 110 mm,

lanceolate-acuminate, glabrous with a few silky hairs on

and near midrib on abaxial surface; ligule papyraceous,

white or brown, up to 30 mm high. Inflorescence up to 200

x 1 10 mm; bracts 40-60 x 20-30 mm, ovate-obtuse, sub-

coriaceous, green, overlapping, each subtending 2-6 flow-

ers. Flowers white, intensely fragrant. Calyx tubular, uni-

laterally split, glabrous, hidden by bract. Corolla tube up

to 70 mm; petals 3, linear-lanceolate, equal, declined.

Staminodes: 30-50 mm, oblong-lanceolate; labellum ± 35

x 50 mm, usually yellow-green in centre, deeply 2-lobed.

Stamen white, shorter than labellum; anther 12 mm long.

Ovary glabrous, or slightly to densely hairy at flowering

stage, 5 mm long. Fruit a capsule; seeds rounded, greyish,

± 5 mm long.

Note: the above description is partly based on Smith

(1984), partly on Turrill (1914) and partly on new observa-

tions.

Vouchers: Rhincl s.n. (PRE); Schlieben & Mendelsohn 12618 (culti-

vated) (PRE).

II. coronarium is widely cultivated in the tropics and

subtropics. It has become naturalised in the Pietermaritz-

burg area (Figure 2), the seeds being dispersed by birds.

2. Hedychium flavescens Roscoe, Monandrian

plants of the order Scitamineae: t. 50 (1825); Lourteig:

123 (1972). Type; Roscoe, Monandr. t. 50.

Plants 2-3 m high. Leaves sessile, up to 600 x 80 mm,

lanceolate-acuminate, lightly pubescent below; ligule

10-20 mm, entire, pubescent. Inflorescence up to 200 x

80 mm; bracts 50 x 35 mm, broadly ovate, rather obtuse,

membranous at margins, unevenly pubescent. Flowers

yellow. Calyx up to 45 mm, pubescent, unilaterally split.

Corolla tube ± twice length of calyx. Petals linear, ± 40

x 20 mm. Labellum: broadly obcordate, narrowed at

base, bilobed above, 80 x 25 mm at broadest part; later-

al staminodes 25-30 mm, spathulate. Stamen up to 40

mm long. Ovary silky pubescent. Fruit ?

H. flavescens is native to the eastern Himalayas. As a

garden escape it has become naturalised in many coun-

tries and has been found in the Pietermaritzburg area.

3. Hedychium gardnerianum Ker Gawl. in

Botanical Register 9: t. 774 (1 Feb. 1824); Roscoe : t. 62

(before 21 April 1824). Type: Hort. Liverpool, Sheppard.

Plants 2-3 m high. Leaves shortly petiolate, 240-400 x

100-150 mm, lanceolate-acuminate, glabrous; ligule

20-40 mm entire. Inflorescence up to 350 mm long; bracts

30-50 mm, convolute, remote. Flowers yellow. Corolla

tube ± 50 mm long. Petals linear. Labellum: 25-30 x

10-20 mm, obovate, emarginate; lateral staminodes 30-35

mm long, narrowly oblanceolate; filament bright red.

Fruit red within.

This very handsome species is also a native of the

eastern Himalayas. The sweetly scented inflorescence is

the largest found in Hedychium. It is naturalised around

Kloof, inland from Durban.

3. ALPINIA Roxb.

Alpinia Roxb. in Asiatic Researches 11: 350(1810)

norm cons.; Benth. & Hook.f.: 648 (1883); K.Schum.:

308 (1904). Type species: A. galanga (L.) Willd.

Catimbium Lestib.: 346 (1841), non Juss. (1789).

Languas Small: 307 (1913).

Plants up to 120 mm tall, more usually 2 -4 m.

Inflorescence terminal on a frond-like leaf shoot. Flowers

borne singly or in cincinni; bracts and/or bracteoles pre-

sent or not. Labellum: often showy, lateral staminodes pre-

sent as small subulate teeth or reduced to small swellings

or absent; anther connective crested or not. Fruit usually

spherical.

A large genus of at least 200 species widely distributed

throughout SE Asia with representatives in Queensland

and Japan.

1. Alpinia zerumbet (Pers.) B.L.Burtt & R.M.Sm.

in Notes from the Royal Botanic Garden Edinburgh 31:

204, t. 10 (1972). Type: Wendland, t. 19 (1798).

Costus zerumbet Pers.: 3 (1805).

Zerumbet speciosum Wendf: 3, t. 19 (1798).

Alpinia speciosa (Wendl.) K.Schum.: 334 (1904), non (Bl.) D.Dietr :

13 (1839).

Catimbium speciosum (Wendl ) Holll : 152 (1950).

Bothalia 28,1 (1998)

39

Plants up to 3 m. Leaves sessile or shortly petiolate, up

to 600 x 100 mm, lanceolate acuminate, margins pubes-

cent, undersurface sometimes so; ligule pubescent.

Inflorescence pendulous, up to 300 mm long, axis pubes-

cent. Flowers white red and yellow, borne in 2-flowered

cincinni; bracts absent; bracteoles glistening white with

pink tips, open to base but encircling flower buds, quickly

deciduous. Corolla tube shorter than 20 mm calyx. Petals

white, dorsal much broader than laterals. Staminodes :

labellum up to 40 mm long, broadly ovate, bifid at apex,

yellow, mottled and striped with red; lateral staminodes

slender, subulate, up to 10 mm long; anther massive,

ecristate. Ovary densely pubescent. Fruit spherical, up to

20 mm diam., orange.

A. zerumbet, the Shell Ginger, is found in tropical gar-

dens all over the world. It is probably truly native to NE

India, Burma and Indo-China. It is reported to have

escaped from gardens in the Hluhluwe area of Zululand.

Excluded species

Kaempferia stenopetala K.Schum. in Das Pflan-

zenreich Heft 20 Zing. 69 (1904); Wright: 314 (1913).

This name was based on an unlocalised collection

from Natal ( Medley Wood 1942, K ! ). It is almost certain-

ly the Asiatic K. rotunda L. widely cultivated in the east

for its medicinal properties, and presumably brought to

Africa by the Indians.

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LOCK, J.M. 1985. Zingiberaceae. Flora of tropical east Africa: 1 — 40.

Balkema, Netherlands.

LOURTEIG, A. 1972. Le genre Hedychium a Madagascar (Zingi-

beraceae). Adansonia , ser. 2, 12,1:121-127.

MARLOTH, R. 1915. Zingiberaceae. The flora of South Africa 4:

165-167, fig. 53. Darter Bros, Cape Town.

PERSOON, C.H. 1805. Synopsis plantarum 1: 3.

REGEL, E A. VON & RACH, L.T. 1859. Index seminum hard botani-

ci Petropolitani.

ROSCOE, W. 1824-1828. Monandrian plants of the order Scitami-

neae, chiefly drawn from living specimens in the Botanic

Gardens at Liverpool. Liverpool.

ROXBURGH, W. 1810. Descriptions of several of the monandrous

plants of India. Asiatic Researches 1 1 : 318-362. Calcutta.

SCHUMANN, K. 1904. Zingiberaceae. In A. Engler, Das Pflanzen-

reich Heft 20: 40-67, 308. Leipzig.

SCHWEINFURTH, G.A. 1867. Cienkowskia. Beitrag zur flora aethi-

opiens: t.l. Georg Reimer, Berlin.

SIMS, J. 1804. Hedychium coronarium. Curtis’s Botanical Magazine

19, ser. 1: t. 708.

SMALL, J.K. 1913. Alpiniaceae. Flora of the southeastern United

States, edn. 2: 307. New York.

SMITH, R.M. 1984. Zingiberaceae. European Garden Flora 2: 120-129.

SPEARING, J.K. & MAHANTY, H.K. 1964. The relationship of the

African species of Kaempferia to those found in Asia. Abstract

of the 10th International Botanical Congress, Edinburgh: 478.

TURRILL, W.B. 1914. Hedychium coronarium and allied species.

Kew Bulletin 1914: 368-372.

WENDLAND, J.C. 1798. In H.A. Schrader & J.C. Wendland, Sertum

hannoveranum, fasc. 4: t. 3, 19. Goettingae.

WILLIAMS, R„ CROUCH, N. & PETTIT, G. 1996. Geophytes of

KwaZulu-Natal. 2. Underworld plants with a special appeal.

IBSA Bulletin 44: 26-31.

WITTMACK, M.C.L. & PERRING, W. 1892. Kaempferia ( Cien -

kowskya) kirkii Hook. Gartenflora 41: 57-59, t. 1364.

WOOD, J.M. 1898. Kaempferia ethelae. The Gardeners’ Chronicle, 3rd

ser., 23: 94. London.

WOOD, J.M. & FRANKS, M. 1911a. Siphonochilus. In J. Medley Wood,

Natal plants 6,3: 560, 561. Bennet & Davis, Smith Street, Durban.

WOOD, J.M. & FRANKS, M. 1911b. Plantarum novarum in herbario

horti regii conservatarum. Kew Bulletin 1911: 274.

WOOD, J.M. & FRANKS, M. 1911c. The Naturalist. Journal of the

Natal Scientific Society 1,3: 112-115.

WRIGHT, C.H 1913. Kaempferia L. In W.T. Thiselton-Dyer, Flora

capensis 5,3: 314-316.

.

Bothalia 28,1: 41-65 (1998)

Notes on African plants

VARIOUS AUTHORS

CYPERACEAE

A NEW COMBINATION IN ISOLEPIS

Since its description by Nees von Esenbeck in 1836,

the generic affinities of the small annual Cyperus leu-

coloma Nees from the Western Cape have remained

unquestioned. Nees and subsequent authors compared

and contrasted it with the superficially similar C. tenellus

L.f. on account of its ‘distichous’ glumes. The type was

not specified by Nees, but he stated that he had seen it ‘in

Herb. Reg. Ber.’. Boeckeler (1867/1868) cited a Bergius

specimen in B, and Clarke (1897) and Kiikenthal (1936)

both cited Bergius 174 in B. Fortunately it was possible

to borrow this sheet (Figure 1).

can be clearly seen that, while otherwise conforming

exactly to Nees’ description, the glumes in the holotype

of C. leucoloma Nees appear to be subdistichous due to

having been collected at a young stage and flattened by

the collector. Glume scars on spikelet axes of dissected

material ( Barker 243) show clearly that the glumes are

spirally arranged. C. leucoloma is undoubtedly a species

of Isolepis : it is probably related to /. incomptula Nees

which has a similar spikelet and glume morphology. The

genus is currently under revision. Meanwhile the new

combination is effected below.

Typification

The sheet consists of two specimens, one in the cap-

sule (Figure 1A) without locality or collector informa-

tion, and another (Figures IB; 2), Bergius 174, collected

at ‘Prom, bon sp.’. Prof. Hiepko, Director of the

Herbarium, B, is of the opinion that the two specimens

were remounted onto one sheet in the latter half of the

19th century, when a larger size herbarium sheet was

brought into use. It is highly likely that the specimen in

the capsule is the one seen by Nees, as the note (Figure

1C) which was folded inside the capsule is almost cer-

tainly in Nees’ handwriting (Burdet 1977: 189, 190).

This specimen is therefore taken to be the holotype. The

label (Figure ID) of the specimen (Figure IB) is written in

Schlechtendal’s hand, excepting the ‘N. ab Es.’ (P.

Hiepko pers. comm.). K.H. Bergius was an apothecary

and naturalist at the Cape from 1815 until his death in

1818, following which his specimens were sent to Berlin

by Von Chamisso (Gunn & Codd 1981: 93). Schlech-

tendal was at that time and up until 1833 ‘supervisor of

the public collection of herbs’ (curator) at B (Hiepko

1987: 221).

Further evidence that this is the original sheet are the

annotations thought to be by Boeckeler (Figure IE), and

those by Clarke (not shown) and Kiikenthal (Figure IF).

Additionally, according to Hiepko (1987: 251), the types

of Cyperus s.l. escaped the destruction during World War

II because they had been moved to the basement of the

Museum.

*

* i B

<U- n. ■ 7 |

r"--' '•

/ ^ 4-^ ^ ■'■j < . ■ i+4-

c

The correct generic placement of the species

There is currently much disagreement on the infrafa-

milial classification (at all levels) of Cyperaceae.

However, it is generally agreed that Cyperus s.str. has

distichous glumes and flattened spikelets, whereas Iso-

lepis R.Br. has spiralled glumes and rounded spikelets. It

F

FIGURE 1. — Isolepis leucoloma (Nees) C. Archer. A, ?holotype; B, spec-

imen; C, note in ?Nees’ handwriting; D, label in Schlechtendal’s

handwriting; E, sketch annotated by ?Boeckeler; F, annotation by

Kiikenthal. Lowermost portion of sheet not shown. Photograph:

R.H. Archer.

42

Bothalia 28,1 (1998)

FIGURE 2. — Enlargement of Figure IB, Bergius 174 to show mor-

phology of species, x 2. Photograph: R.H. Archer.

Isolepis leucoloma (Nees) C. Archer, comb. nov.

Cyperus leucoloma Nees in Linnaea 10: 133 (1836); Kunth: 50

(1837); Boeck.: 506 ( 1867/1868); C.B.Clarke: 566 (1894); C.B.Clarke:

165 (1897); Kiik.: 298 (1936); Levyns: 99 (1950); Bond & Goldblatt:

39 (1984). Type: not specified [most probably: Prom. bon. sp., Bergius

174 ] (B, specimen in capsule!).

Specimens examined

WESTERN CAPE. — 3219 (Wuppertal): on road to Heuningvlei,

(-AA), Erndon 129 p.p. (NBG); Porterville, on plateau of Voorberg

(above Dasklip Pass) on road to Forestry Centre, Zuurvlakte Farm,

(-CC), Barker 243 (PRE). 3318 (Cape Town): Malmesbury, (-BC),

Schlechter 16336 (K, P). 3319 (Worcester): Paarl Division, French

Hoek Peak, (-CC), Stokoe s.n. sub SAM 59230 (SAM). 3418

(Simonstown): mountain at Kalk Bay, (-AB), Levyns 831 (K).

ACKNOWLEDGEMENTS

Prof. P. Hiepko, Director of the Herbarium, Bota-

nischer Garten und Botanisches Museum Berlin-Dahlem,

is thanked for information and specimens borrowed. Dr

O.A. Leistner, National Botanical Institute, Pretoria, is

thanked for the translation, as far as was possible, of

Nees’ note.

REFERENCES

BOECKELER, J O. 1867/1868. Die Cyperaceen des Koniglichen

Herbariums zu Berlin Linnaea 35: 397-612.

BOND, P. & GOLDBLATT, P. 1984. Plants of the Cape Bora: A

descriptive catalogue. Journal of South African Botany Suppl.

Vol. 13.

BURDET, H.M. 1977. Autographes de botanistes sous forme de fiches.

X. Candollea 32: 165-206.

CLARKE, C.B. 1894. Cyperaceae. In T. Durand & H. Schinz,

Conspectus florae africae 5: 526-692.

CLARKE, C.B. 1897. Cyperaceae. In W.T. Thiselton-Dyer, Flora

capensis 7 : 1 49-3 1 0 .

GUNN, M & CODD, L.E. 1981. Botanical exploration of southern

Africa. Balkema, Cape Town.

HIEPKO, P. 1987. The collections of the Botanical Museum Berlin-

Dahlem (B) and their history. [See also: Appendix A, Appendix

B ] Englera 7: 219-252.

KUKENTHAL, G. 1936. Fam. IV, 20. Cyperaceae-Scirpoideae-

Cypereae. In A. Engler, Das Pflanzenreich 101.

KUNTH, C.S. 1837. Enumeratio plantation 2. Stuttgart

LEVYNS, M R. 1950. Cyperaceae. In R.S. Adamson & T.M. Salter,

Flora of the Cape Peninsula: 97-132.

NEES VON ESENBECK, C.G. 1836. Cyperaceae capenses secundum

novissimas Ecklonii collectiones. Linnaea 10: 127-207.

C. ARCHER*

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

MS. received: 1994-07-25.

ASTERACEAE

A NOTE ON THE BRACHYLAENA DISCOLOR COMPLEX

In their revision of the South African species of

Brachylaena, Phillips & Schweickerdt (1937) recognised

nine species, including B. uniflora Harv., B. transvaalen-

sis E. Phillips & Schweick. (a new species) and B. discol-

or DC. The key character by which they distinguished

the latter three species was the number of flowers in the

male capitula (1-3 in B. uniflora-, 7-50 in B. trans-

vaalensis and B. discolor). This is unfortunately not suf-

ficient for identification purposes because female plants

are not taken into account. B. transvaalensis was distin-

guished from B. discolor by the size of the capitula (less

than 10 mm long in B. transvaalensis and over 10 mm

long in B. discolor). Paiva (1972) reduced B. trans-

vaalensis to a subspecies of B. discolor [B. discolor DC.

subsp. transvaalensis (E. Phillips & Schweick.) Paiva]

and divided B. discolor DC. subsp. discolor into two

varieties (var. discolor and var. mossambicensis Paiva).

He did not include B. uniflora in his publication as this

species fell outside his study area, but a label on a speci-

men in PRE indicates that he recognised B. uniflora as a

separate entity. Hilliard & Burtt (1971) studied these

three species and came to the conclusion that they should

be regarded as members of a complex comprising either

a single species showing clinal variation in the number of

flowers per capitulum, loosely linked to an ecological

cline from coastal dunes to inland forest, or representing

two species which have met and are now hybridising.

They did find a general correlation between small, few-

flowered capitula with short involucres and much-

branched synflorescences, whereas larger capitula with

longer involucres tend to correlate with less branched

synflorescences. They did not find any conspicuous veg-

etative differences between these three species, a view

supported by my studies of herbarium material. Hilliard

(1977) did not agree with Paiva (1972) in the division of

B. discolor into subspecies and varieties, a view also sup-

ported by Pope (1992) and myself. Cilliers (1993) pub-

lished a synopsis of the genus Brachylaena in southern

Africa. He followed Paiva in upholding the subspecies of

B. discolor but sank B. uniflora Harv. under B. discolor

Bothalia 28, 1 (1998)

43

TABLE 1. — A summary of diagnostic features and distribution of three Brachylaena species

DC. subsp. transvaalensis (E. Phillips & Schweick.)

Paiva. The illustration of B. discolor suhsp. transvaalen-

sis in Cilliers (1993: fig. 15), appears to depict a combi-

nation of B. uniflora and B. transvaalensis: the branch

with leaves and female capitula (A) and the female capit-

ulum (C) look like those of B. uniflora ; the male capitu-

lum (B) like that of B. transvaalensis. Pope (1992) in his

treatment of the genus Brachylaena for the Flora zambe-

siaca area, recognised B. discolor without any subspecies

and regarded B. transvaalensis as a separate species, as

originally described by Phillips & Schweickerdt (1937).

He did not mention B. uniflora , as this species did not

fall within his study area. I agree with Phillips &

Schweickerdt (1937) who kept the three species distinct

and undivided, a view also subscribed to by Pope, as far

as he studied the complex.

Not any of the above-mentioned authors used the

shape of the involucre as a distinguishing character. The

involucres of the three species are either infundibuliform,

cyathiform or globose. It seems to be a constant charac-

ter which can be used easily. Pope (1992) used the char-

acter of the capitula being stalked or not. This feature is

not always easy to distinguish in South African material.

Table 1 gives a summary of the diagnostic features of

the three species B. discolor, B. transvaalensis and B.

uniflora. Using the shape of the involucre, they can gen-

erally be easily separated in the herbarium provided good

fertile material of either sex is available. Distribution of

the species (Figure 4) is listed in Table 1.

On the basis of the shape of the involucre, the three

species can be separated as follows (measurements of

involucre length include the stalk, where applicable, and

width is measured across the widest part):

la Involucre infundibuliform:

2a Male and female capitula small, synflorescences very

dense, involucre of both male and female capitula

narrowly infundibuliform, slender (Figure 3G-J),

2.5— 4.0(-5.5) x 1-3 mm in male capitula and (3-)

4 — 7(— 1 0) x 1-5 mm in female capitula; inner bracts

narrowly triangular to almost linear, acuminate, stiffly

erect; leaves oblanceolate to obovate, rounded or some-

times briefly acuminate at apex 3. B. uniflora

2b Female capitula large, mostly with distinct stalks with

only a few scattered bracts, involucres of female

capitula widely infundibuliform to cyathiform at

base (Figure 3B), (7—) 1 0— 1 8(— 23) x (3. 5-)5. 0-10.0

(-12.0) mm; bracts narrowly ovate with apex acute

and somewhat spreading, synflorescence less dense

than in B. transvaalensis: leaves similar to those of B.

uniflora but more constant in male than female plants

1 . B. discolor

lb Involucre globose to cyathiform:

3a Male capitula large, mostly with distinct stalks with only a

few scattered bracts, involucres of male capitula

(5.5— )6.0— 13.0(— 1 5.0) x (2.5-)4.0-6.0(-8.5) mm (Fig-

ure 3A); bracts narrowly ovate with apex acute, some-

what spreading, synflorescence less dense than in B.

transvaalensis: leaves oblanceolate to obovate, round-

ed or sometimes briefly acuminate at apex but more

constant in male than female plants 1 . B discolor

3b Male and female capitula small, mostly sessile or subses-

sile with involucral bracts extending to the base of the

stalk, involucres (3.5— )4.0— 6.5(— 7.0) x ( 1 .5— )2.5— 3.5

(-4.0) mm in male capitula and (4.0-)5.5-7.0(-8.5) x

(2.0— )3.0 — 4.5(— 8.5) mm in female capitula (Figure

3C-F), synflorescence very dense; leaves narrowly

elliptic, acute and ± acuminate at apex but more con-

stant in male than female plants 2. B. transvaalensis

Specimens examined

Abbott 3958 (3) NH. Abrahams s.n. (1) PRE. Archibald 3632/32 ,

3690 ( 1) PRE. Aubrey s.n. NU34158 , NU 34172 (1) NU.

Balkwill 372 (3) NU. Bayer 554 (1) NU. Bayliss BRI.B 1499 , 1516

(3); 1 540 ( 1 ) PRE. Biegel 2258 (2) NU. Boocock 26 (3) PRE. Borcherds

52 (2) PRE. D.J. Botha 654 (2) PRE. W.M. Botha 5286 (2) PRE.

Bourquin 46 (2), 150 (1) NU. Breyer s.n. Tvl. Mus. 24388 (2) PRE.

Britten 773 , 1837, 2300, 2412 (1) PRE. Bruce 7(1) PRE Brummitt

12457 (2) PRE. Bruton 72 (1) PRE. Burger 525 (2) PRE. Bunt Davy

2392 (1) PRE.

Coleman 335 (3) NH. Comins 1255A (1) PRE. Compton 26061,

26994, 29070, 29073 (2) PRE. Cooper 1240 (1) PRE. Culverwell 961,

975 (1) PRE. Cunningham 856 (3) NU.

Davis 82 (3) NH. De Winter 8359a (1 ) PRE Dlamini s.n. PRE31785 ,

PRE31787, PRE43310 (2) PRE. Du Toit 1265 (1) NH, PRE.

Edwards 1529 (3) NU, PRE. Evans 5398 (2) PRE.

44

Bothalia 28,1 (1998)

FIGURE 3. — Shape of involucres of d and 9 capitula of Brachylaena

spp. A, B, B. discolor A, d , x I , Von Wyk 6120 & Mogg 38407',

B, 9, x 1, Sirey 8789 & Schweickerdt 1384. C-F, II. trans-

vaalensis: C, d, x I ; D, d, x 3, Galpin 451 & Van Vuuren 1278',

E, 9, x 1 ; F, 9, x 3, O’Connor 1471 & Forester Evans 5398.

G-J, B. uniflora'. G, d, x 1; H, d, x 3, Kotze 432 & Medley

Wood 12670. I, 9, x 1 ; J, 9, x 3, Bayliss BRI BI499 & Boocock

26. Artist: Gillian Condy.

Fegen 2060 (1) PRE. Flanagan 860. 861 (1) PRE. Forbes 330 (1)

NH. Forbes & Obermeyer 28 (1) NH, PRE. Forestry Dept., Pta. s.n.

Tvl. Mus. 9703 (2) PRE.

Galpin 451, 12114, 13995 ( 2) PRE. Gerrard & McKen 1017 ( 1), 1866

(3) NH. Gerstner 3013 (3), 4892 (2), s.n. PRE43300 (3) PRE. Gibbs

Russell 3879 (1) PRE. Giffen 254 (1) PRE. Goodman 304 (1) NU.

Gordon 162 (3), 348 (1) NH. Gordon-Gray 596 (1) NU; 607 (3) NH;

1580 (3) NU. Grenfell 1105 (2) PRE. Guy & Jarman 2 (2) NU.

Hafstrbm & Acocks 1529 (2) PRE. Hall-Martin 1867 (1) PRE. Halse

s.n. (3) NU. Hanekom 2312 (2) PRE. Harrison 29 (1) PRE. Hemm 268

(male plant) (2) PRE. Hillary 392 (2) NU. Hilliard 5068, 5069 (3) NU.

Hubbard 6103 (1) PRE.

N. Jacobsen 1810, 2963 (2) PRE. W. Jacobsen 5329 (2), 4712 (1)

PRE. Jones 55 (2) PRE. Jordaan 87 (2) PRE.

Karsten s.n. PRE31786, PRE43311 (2) PRE. Keet 1130 (2) PRE.

Kluge 173, 1070 (2) PRE. Kotze 89, 450 (1), 432 (3), 2835 (2) PRE.

Krige 19 (2) PRE.

Lansdell 66 (1) NH, PRE; s.n. NH 34283 (1) NH. Law 71 (2) NU.

Lawn 643 (3) NH. Letty 222 (1 ) PRE.

Macleod 19 (2) NU. Marloth 4174 (1) PRE. Marriott s.n. (1) PRE.

McCollum 1532 (2) PRE. McMurtry 2190 ( 1 ) PRE. Meyer 6144(1) PRE.

Mills 342 (3) PRE; 469 ( 1) NH, PRE. Mogg 33761 (3), 38407, 38443 (1)

PRE. Moll 1816 ( 1) NU, PRE; 4501 (2) NH. Munro PS. 2(1), 35, 213 (2)

PRE.

Nel 258 (2) PRE. Nicholas 1637(1) NH, PRE. Nichols 424 (1) NH.

Nicholson 808 ( 3) PRE. Nixon 24 ( 1 ) NU.

Obermeyer s.n. Tvl. Mus. 35830 (2) PRE. Obermeyer, Schweickerdt &

Verdoorn 349 (2) PRE. O'Connor 1471 , 3559 (2) PRE Oranje 2 (2) PRE.

Pegel NDO 119 NH59807 (3) PRE. Pole Evans HI 57 19 (2) PRE. A.

Prior s.n. PRE30046 (1) PRE. J. Prior 103, 304 (2) PRE.

Rauh & Schlieben 9721 (2) PRE. Ross 2115, 2118 (1) NH, PRE;

2383, 2388 ( 1) PRE. Rudatis 653 (3), 1089 (1) PRE.

Scheepers 692 (2) PRE. Schlec liter 2886 (1) PRE. Schonland 3304 ( 1 )

PRE. Schrire, Van Wvk & Abbott 1782 (3) NH. Schweickerdt 1367 (1)

PRE; 1368 (2) NH, PRE; 1384 (1) NH, PRE. Sikhakhane 110(1) NH.

Sim 19260 (1) PRE. Smuts s.n. PRE43373 (1) PRE. Stephen 385 (1)

PRE. Stephen & Van Graan 1266 (1) PRE. Stewart s.n. PRE61399 (2)

PRE. Stielau 23 ( 1) PRE; 192 (1) NH, PRE. Story 1289 (1) PRE. Strey

6894, 7658, 8753, 8783, 8789, 8790, 8791, 8792, 8793, 8798, 9909(1)

PRE; 8822, 8858, 8859, 9907 ( 1 ) NH, PRE; 8779 ( 1 ) NH; 8795 ( 1 ) NU,

PRE; 8788 (1) NH, NU, PRE; 8754, 8797 (2) PRE; 8751, 8752, 8787,

8805, 8819, 8841 (3) PRE; 8784, 8804, 8820, 8824, 10973 (3) NH,

PRE; 8823 (3) NU, PRE. Strey & Huntley 4292 (1) NH, PRE. Stubbings

4B (3) PRE.

Taylor 206 ( 1) NU. Thode 2927, 2936 (3) PRE; AI518, A1532 (1)

PRE. Thorncroft s.n. Tvl. Mus. 2773, Tvl. Mus. 11174 (2) PRE. Thorns

FIGURE 4. — Distribution of B. discolor, O, B. transvaalensis, A; and

B. uni flora. •.

Bothalia 28,1 (1998)

45

sub Strey 5785 (3) NU, PRE. Tosh s.n. NU 30704 (1) NU. Tustin 3555 ,

3556 (2) PRE. Tyson 8575, 12571 (1) PRE.

Van Jaarsveld 748 (2) PRE. Van Vuuren 1278 (2) PRE. Van Wyk 6120

(1) PRE. Van Wyk & Theron 4600, 4741 (2) PRE. F. Venter 980 (3)

PRE. H.J.T. Venter 1927 (1) PRE.

C.J. Ward 5810, 6624, 7137 ( 1 ) PRE; 2687, 6623, 7138 ( 1 ) NH, PRE;

22 (1) NU; 2630 (3) NH, PRE. M.C. Ward 25 (1) NU; 801 (1) NH,

PRE; 504, 2090 ( 1) NH; 1542 (2) NH. Willox 1 (1) PRE. Wirminghaus

520 (1) NU. Wood 585 (3) NH; 4907(1). 12287, 12670 (3), 12670a (1),

12670b (3) PRE. Wright 114 (1) PRE.

Problem specimen

Hemm 268 (female plant, PRE), identified as B. dis-

color, is a very peculiar specimen totally out of the dis-

tribution range of the species and with smaller capitula

than the usual B. discolor, but with stalked capitula and

an involucre similar to that of B. discolor. Vegetatively it

looks very similar to Hemm 268 (male plant, PRE), iden-

tified as B. transvaalensis.

ACKNOWLEDGEMENTS

The Curators of the Natal Herbarium and the

Herbarium of the University of Natal are thanked for the

loan of specimens. Discussions with Prof. A.E. van Wyk

and Dr O.A. Leistner are gratefully acknowledged. Ms

Gill Condy is thanked for the illustrations.

REFERENCES

CILLIERS, S.S. 1993. Synopsis of the genus Brachylaena (Asteraceae)

in southern Africa. Bothalia 23: 175-184.

HILLIARD, O.M. 1977. Compositae in Natal. University of Natal

Press, Pietermaritzburg.

HILLIARD, O.M. & BURTT. B.L. 1971. Notes on some plants of

southern Africa chiefly from Natal; 2. Notes from the Royal

Botanic Garden Edinburgh 31,1: 3-6.

PAIVA, J A R. 1972. New and little known species from the Flora

zambesiaca area. 22. Notes on Inuleae. 1. Tarchonanthinae.

Boletim da Sociedade Broteriana 46: 355-384.

PHILLIPS, EP & SCHWEICKERDT, H.G. 1937. A revision of the

South African species of Brachylaena R. Brown. Bothalia 3:

205-221.

POPE, G.V. 1992. 97. Compositae. Flora zambesiaca 6,1: 5-8.

P.PJ. HERMAN*

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

MS. received: 1997-04-09.

METZGERIALES-FOSSOMBRONIACEAE

FOSSOMBRONIA RWANDAENS1S, A NEW SPECIES FROM TROPICAL AFRICA

INTRODUCTION

In a previous paper (Perold 1997), reference was made

to some specimens of Fossombronia, notably De Sloover

185.74, (BR, NAM) from Rwanda, which had been

referred to F. capensis S.W.Arnell. The two taxa, F.

capensis and the fertile Rwanda specimen placed here,

have several characters in common; both are dioicous and

have large perigonial bracts subtending the antheridia in

the male plants and their elaters are similar in that they are

poorly formed. The spore ornamentation in F. capensis

and the Rwanda specimen is, however, completely differ-

ent and in some capsules of the Rwanda species (particu-

larly those held at BR) it even exhibits a degree of dimor-

phism. Since spore ornamentation is regarded as the most

important character in distinguishing between species of

Fossombronia, it must be assumed that we are here deal-

ing with two different taxa. The records of F. capensis in

tropical Africa (Vana et al. 1979; Wigginton & Grolle

1996) should therefore be deleted, as its distribution is

seemingly restricted to a smallish area in the southern

Cape winter rainfall region (Perold 1997); F. rwandaensis

is described as a new species.

Fossombronia rwandaensis Perold, sp. nov.

Plantae prostratae, crebrae in coloniis. Folia imbrica-

ta, late patentia, irregulariter rectangularia. Rhizoidea

purpurea. Dioicae. Plantae masculae communes;

antheridiis bracteis magnis tectis, saepe binatim connatis.

Pseudoperianthium late campanulatum, supra basin cra-

teriforme plicatum vel fimbriatum. Sporae 42.5-52.5 pm

diametro, superficie distali plerumque cum cristis crassis,

rarissime reticulata. Elateres 62.5-77.5 pm longi,

debiliter formati. Gametophyta aliquantum similes sunt

illis F. capensis (ut hae plantae olim agnotae sunt), sed

sporarum ornamentis valde differunt.

TYPE. — Rwanda, prefecture de Gisenyi, sous le poste

minier de Gikungu, alt. env. 2 000 m. Sur talus tres

humide, J.L. De Sloover 18.574 (NAM, holo.!; BR, iso.!).

Plants predominantly male, in dense overlying mats,

green; shoots large, simple (Figure 6C) or once furcate

(Figure 6A), rarely twice, up to 20 mm long, 1. 8-2.0 mm

high, 3.2— 4.0 mm wide (up to 4.7 mm wide at pseudoperi-

anth in female plants), apical segments moderately diver-

gent, 2-8 mm long. Stems prostrate, sometimes tapering

proximally, chlorophyllose, occasionally with a lateral

bud, plano-convex in cross section, in male plants apical-

ly (Figure 5M) 200-250 pm (10 cell rows) high, 370^150

pm wide, basally (Figure 5N) 200-260 pm high, 280-350

pm wide, in female plants apically (Figure 50), 250-330

pm high, 350-540 pm wide, basally (Figure 5P) 200-300

pm high, 350-450 pm wide. Rhizoids purple, 17.5-25.0

pm wide. Leaves overlapping, widely spreading, suc-

cubously inserted, irregularly rectangular longitudinally or

transversely, upper margin and sometimes ‘leading’ edge ±

rounded, with low protrusions here and there, ‘trailing’

edge occasionally sloping obliquely, with up to 6 uneven-

ly spaced marginal slime papillae, ± 17.5 x 20.0 pm, ses-

sile or raised on a basal cell; in male (Figure 5A-E) and

46

Bothalia 28,1 (1998)

FIGURE 5. — Fossombronia rwandaensis, De Sloover 18.574. A-E, male leaves; F-J, female leaves; K, detail of trailing leaf margin; L, median

leaf cells with chloroplasts (dotted lines), oil bodies no longer present in material studied; M, cross section of apical part of male stem; N,

cross section of basal part of male stem; O, cross section of apical part of female stem; P, cross section of basal part of female stem; Q-U,

perigonial bracts; V, opened pseudoperianth; W, pseudoperianth from side; X, cells in capsule wall; Y, Z, elaters. Scale bars: A-J, V, W,

500 pm; K, 100 pm; L, X, 50 pm; M-U, 250 pm; Y, Z, 25 pm. Drawn by G. Condy.

Bothalia 28.1 (1998)

47

FIGURE 6. — Fossombronia rwandaensis, De Sloover 18.574. A, moderately divergent terminal segments with rows of perigonial bracts sub-

tending antheridia; B, detail of rows of perigonial bracts; C, apical part of female shoot; D, archegonium near apex (indicated by arrow);

E, pseudoperianth from side; F, pseudoperianth from above. A, x 14.4; B, x 24; C, x 12; D, x 33.6; E, x 22.8; F, x 23.4.

female plants (Figure 5F-J) not markedly different,

(107 5-) 1500-2 125 x 1850-2250(-2900) pm, sometimes

narrower above, up to 1625 pm. wide. Leaf cells thin-

walled, at upper margins subquadrate to rectangular

across, 17.5-32.5 x 30.0-45.0 pm, at lateral margins

(Figure 5K), long-rectangular, 62.5-87.5 x 32.5—42.5 pm,

upper laminal cells 5- or 6-sided, 45.0-57.5 x 35.0-47.5

pm, middle laminal cells 50.0-75.0 x 40.0-50.0 pm, basal

cells 55.0-95.0 x 50.0-67.5 pm. Oil bodies lost in the

material examined; chloroplasts clumped together at cell

margins (Figure 5L), ± 3 pm long.

Dioicous. Antheridia dorsal on stem, mostly in 2

crowded rows, short-stalked, ± ovoid, 340 x 300 pm,

shielded by perigonial bracts (Figure 6A, B), which are

often in fused pairs (Figure 5R-T), 600-800 x 550-600

pm, when single (Figure 5Q, U), ± 340 pm wide, mar-

gins with pointed or obtuse projections, up to 300 pm

long, marginal cells ± rectangular or subquadrate,

25.0-42.5 x 25.0-37.5 pm, cells in interior 4- or 5-sided,

37.5-67.5 x 30.0^10.0 pm. Archegonia in l(Figure 6D)

or 2 scattered rows along stem, naked; sometimes 2 in

close proximity becoming fertilised. Pseudoperianth

(Figures 5V, W; 6E, Fj ± 1.5 mm proximal to apex, wide-

ly flaring and pleated or frilled above bowl-shaped base,

about as tall as leaves, up to 2100 pm long and 5500 pm

wide across mouth, at margin numerous projections,

some rounded, others sharply pointed, with ± 28 slime

papillae, mostly sessile, one-celled, 20 x 20 pm, a few

raised on 3 tiers of basal cells, 42.5-50.0 x 37.5-50.0

pm, below with winged lateral outgrowths or append-

ages; cells not appreciably different in shape and size

from those in leaves. Capsules globose, ± 1000 pm diam.,

wall bistratose, inner layer (Figure 5X) rather pale, cells

irregularly shaped, polygonal or roughly triangular to

quadrangular, 37.5-62.5 x 30.0-42.5 pm, each cell wall

with 2 or 3 brown, nodular thickenings, entirely lacking

semi-annular bands. Seta almost absent, as all capsules

examined ± sessile. Spores golden brown, with ridges,

loops, blobs and very rarely reticulations deeper brown,

hemispherical to occasionally somewhat irregularly

shaped, a few still in tetrads (Figure 7H), 42.5-52.5 pm

diam., including spines projecting at periphery; distal

face (Figure 7A-D) convex, with thick ridges, irregular-

ly branching and curving or unbranched, short or long,

straight or bent or looped, sometimes nodular, very rarely

in some capsules only, reticulate (Figure 7G, I), with up

to 13 small areolae across face; proximal face (Figure 7E,

F) with triradiate mark rather indistinct to distinct, occa-

sionally with nodules aligned along its arms, otherwise

nodules scattered irregularly, size variable, up to 7.5 pm

wide, in between dusted with fine granules, around spore

periphery 25-29 conical or truncate spines, ± 3 pm long,

unconnected by a perispore. Elaters (Figure 5Y, Z) few,

rather delicate and poorly formed, often adhering to a

spore, 62.5-77.5 pm long, 10 pm wide throughout or 15

pm wide centrally and tapering to 10 pm wide tips, dif-

ficult to distinguish the spirals, but apparently 3-spiral.

Fossombronia rwandaensis has been named for the

country where the type and only specimen with spores, De

Sloover 18.574, was collected at high altitude in very

humid conditions. Some maps that were consulted, give

the spelling of the ‘prefecture’ as Kisenyi, not Gisenyi.

The other De Sloover collections that were determined as

F. capensis by Vana, De Sloover 13.345, 19.118, 19.169

(the latter from neighbouring Burundi) lack spores and

can, therefore, not be identified with certainty, even

though 13.345 has large perigonial bracts in the male

plants. F. rwandaensis can be distinguished by its dioicity,

by large bracts subtending the antheridia, usually in 2

crowded rows dorsally along the stems of the male plants,

by the cells in the inner wall of the capsules lacking semi-

annular bands, by the coarsely ridged spores and by poor-

48

Bothalia 28,1 (1998)

FIGURE 7. — Fossombronia rwandaensis, De Sloover IS. 574. Spores. A-C, distal face; D, detail of ridges near margin of spore; E, side view; F,

proximal face; G, several spores in one field, some ndged, others reticulate; H, reticulate spores in partially broken tetrad; I, reticulate

spore. A, x 1200; B, x 995; C, x 1212; D, x 2160; E, x 1260; F, x 1030; G, x 190; H, x 545; I, x 1055 (reticulate spores in G indicated by

arrows).

ly formed elaters. As mentioned above, some capsules also

contained a few spores that had reticulate ornamentation,

either still in tetrads or already separated. Out of a total of

758 spores examined (from various samplings), only 24 or

3% had reticulate ornamentation, whereas all the rest, i.e.

97%, were ridged. The reticulate spores may be aberrant or

still somewhat immature. As far as could be ascertained,

they appeared to be absent from capsules that had been

intact in the holotype specimen held at NAM. I think,

however, that contamination of the capsules with occa-

sionally reticulate spores, can be ruled out as a possible

explanation for their presence. Paton (1973) observes that,

‘Aberrant spores occur occasionally in this [i.e.

Fossombronia] and other genera. Sometimes it appears

that after the spore has attained full size, the development

of the sculpturing has been arrested prior to final cutinisa-

tion. In such cases lamellae may be poorly developed or

fragmentary’. Whether this would apply to the reticulate

spores occasionally encountered in F. rwandaensis

remains a matter of conjecture.

Fossombronia rwandaensis shares several characters

with F. capensis, namely dioicity, large perigonial bracts

and poorly formed elaters. The distribution of the latter is,

however, confined to a smallish area in the southern Cape

winter rainfall region (Perold 1997). The lamellate spore

ornamentation in F. capensis is also quite different. The

records of F. capensis in tropical Africa (Vana et al. 1979;

Wigginton & Grolle 1996) should, therefore, be deleted.

SPECIMENS EXAMINED

De Sloover 13.345 (BR, NAM), Rwanda, foret de Gishwati, le long

de la route Gisenyi-Kibuye. De Sloover 18.574 (NAM, holotype; BR,

isotype), Rwanda, prefecture de Gisenyi, sous le poste minier de

Gikungu, le long de la Bikoneko. De Sloover 19.118 (BR, NAM),

Rwanda, prefecture de Cyangugu, Foret de Rugege, entre Gisakura et

Karantba. De Sloover 19.169 (BR), Burundi, au km 30 de la route

Bujumbura-Ijenda, Matara.

ACKNOWLEDGEMENTS

I sincerely thank the Director of BR and Prof. J.L. De

Sloover for the loan of specimens. I also extend my grat-

itude to Dr H. Stieperaere for refereeing this article, Dr

H.F. Glen for the Latin translation, Ms G. Condy for the

drawings, Mrs A. Romanowski for developing and print-

ing the photographs and Mrs J. Vcldman for typing the

manuscript.

Bothalia 28,1 (1998)

49

REFERENCES

PATON, J.A. 1973. Taxonomic studies in the genus Fossombronia

Raddi. Journal of Bryology 7: 243-252.

PEROLD, S.M. 1997. 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.

VANA. J„ POCS, T. & DE SLOOVER, J.L. 1979. Hepatiques

d’Afrique Tropicale. Lejeunia 98: 1-23.

WIGGINTON, M.J. & GROLLE, R., supplemented by A. Gyarmati.

1996. Bryophytorum Bibliotheca 50: 1-267.

S.M. PEROLD*

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

MS. received: 1997-08-04.

THYMELAEACEAE

A NEW SPECIES OF LACHNAEA ENDEMIC TO THE SOUTHEASTERN MOUNTAINS OF THE WESTERN CAPE

Lachnaea sociorum Beyers, sp. nov., structura

inflorescentiae florisque L. ericoidis similis, sed sepalis

tomentosis, stigmate penicellato, foliis lanceolatis ad

anguste ellipticis, abaxiale leviter 3-costatis, apice acuto,

glabris.

TYPE. — Western Cape. 3321 (Ladismith): Langeberg,

Bergfontein, S-facing midslopes of Koksposberg, 379 m,

(-DC), 1 8-0 1-1991, McDonald 2059 (NBG. holo.; BOL,

K. NY, PRE, iso.).

Erect, compact, moderately branched shrub up to 0.6

m tall, multistemmed at base, resprouting after fire.

Branches erect, slender, ridged below leaf bases, covered

with long, straight, adpressed hairs admixed with

crooked ones, densely leafy, becoming naked with

prominent leaf scars. Leaves alternate, ascending or

occasionally inclined to patent, with apex curled inward,

imbricate, sessile, lanceolate to narrowly elliptic or occa-

sionally elliptic to obovate, 3. 6-6.0 x 0.9-2. 4 mm, cori-

aceous, green, those below inflorescence sometimes

tinged maroon, glabrous, adaxially concave, smooth,

dull, ± white-punctate, abaxially convex, faintly 3-ribbed

or subkeeled towards apex, glossy, subpapillate, apex

acute, base cuneate. Inflorescence a terminal, sessile,

ebracteate, 6— 14-flowered umbel with up to 9 open flow-

ers at a time, on main flowering branches. Flowers sub-

actinomorphic, white, skunk-scented. Pedicel 0.3-0. 7

mm long, sericeous. Hypanthium 2. 5-3. 7 mm long, cir-

cumscissile, three-fifths to three-quarters from base,

upper portion funnel-shaped, tomentose outside,

adpressed hirsute within, basal portion oblong, glabrous,

rarely tomentose outside, glabrous within. Sepals 4,

patent, subequal, widely ovate or subrotund, 1. 7-3.0 x

1.7-3. 2 mm, apex rounded or obtuse, adaxially and abax-

ially tomentose. Petals absent. Stamens 4 + 4, exserted,

outer, antisepalous whorl inserted at base of sepals,

0.7-1. 5 mm long, inner, antipetalous whorl inserted at

rim of hypanthium, 0.5-1. 2 mm long (inner stamens the

shortest or equal to the shortest of outer stamens);

anthers widely ellipsoid, 0.4— 0.6 mm long, abaxially

without broad connective tissue. Scales 8, exserted,

inserted at mouth of hypanthium immediately below

antipetalous stamens, obovoid or subglobose, 0.3-0. 5

mm long, glabrous, translucent-white when fresh. Ovary’

ellipsoid or obovoid, 0.9-1. 4 mm long, glabrous. Style

linear-obconical, 2. 0-2. 9 mm long, with straight,

adpressed or incurled hairs in upper half to two-thirds.

Stigma penicillate. Chromosome number. 2n = 18

(Beyers 176). Figure 8.

Leaf ana tomy

Blade dorsiventral, epistomatic, glabrous, in trans-

verse section slightly concave to more or less plane

adaxially, convex abaxially (Figure 9). Cuticle well

developed, thicker abaxially, surfaces papillate, with

flaky wax deposits, particularly adaxially (Figure 10).

Epidermis uniseriate, adaxial epidermal cells much

smaller than abaxial ones, usually periclinally elongat-

ed; abaxial epidermal cells square to radially elongated,

with outer tangential walls thicker than inner tangential

walls (before gelatinisation), inner tangential cell walls

of some cells gelatinised; adaxial epidermal cells

incompletely and weakly gelatinised; stomata sunk

below adjacent epidermal cells. Mesophyll differentiat-

ed into palisade and spongy parenchyma; palisade abax-

ial, 1- or 2-seriate; spongy parenchyma adaxial, cells

palisade-like, loosely arranged with fairly large intercel-

lular spaces. Vascular bundles comprising a large mid-

bundle, flanked by 2 smaller lateral bundles on either

side, with small intermediate bundles in between these

two types; each bundle capped abaxially by large

extraxylary fibres which are particularly well developed

in the mid- and lateral bundles; bundle sheath present,

uniseriate, parenchymatous, incomplete, interrupted by

extraxylary fibre cap. Crystals of diosmin (potassium

hydroxide test: Jackson & Snowdon 1990) present in

adaxial epidermis and mesophyll, occurring as shaero-

crystalline to somewhat dendritic masses, pale yellow;

crystals and tanniniferous deposits mutually exclusive.

Tanniniferous deposits present in all abaxial, and in

some adaxial epidermal cells.

Vouchers: Beyers 176; McDonald 2059.

Pollen morphology

Pollen grains shed as monads, spheroidal, pantopo-

rate, 39-41 pm diam. Tectum reticulate with triangular

supratectal subunits, (crotonoid pattern sensu Erdtman

1966); supratectal subunits triangular in surface view,

with either a single central spinule or with usually four

spinules, lateral sides of subunits straight or emarginate,

surface around spinules striate to granular-striate, with

muri (ridges) cross-linked in subunits with lateral sides

emarginate. Figure 1 1 .

Vouchers: Beyers 176; Oliver 10524.

50

Bothalia 28,1 (1998)

FIGURE 8. — Lachnaea sociorum. A, portion of plant; B, branchlet (enlarged); C, leaf (abaxial view and cross section); D, leaf (lateral view); E,

flower; F, flower split longitudinally with gynoecium removed; G, gynoecium; H, portion of plant; I, flower; atypical pubescent form. A-G,

McDonald 2059 ; H, I, Beyers 176.

Distribution, ecology and variation

Recorded from the eastern end of the Langeberg

Mountains, from Perdeberg in the west eastwards to

Gavelsbos and the southern slopes of the Attakwa

Mountains below Perdekop. Plants occur in stony soil on

the southern and northern slopes, at altitudes between

300 and 1 100 m (Figure 12). Flowering recorded from

August to January.

The population on the northern side of the Langeberg

at Tygerberg, west of Huisrivier, is very localised and rea-

sonably old. These plants are single-stemmed at the base,

branching close to the ground and are open at the crown

Bothalia 28,1 (1998)

51

B

FIGURE 9. — Lachnaea sociorum, McDonald 2059 , leaf anatomy, transverse sections. A, leaf lamina, note outline of shape, tanniniferous epider-

mal cells (particularly abaxially), sunken stomata (arrowed) and vascular bundles with prominent caps of extraxylary fibres; B, portion of

lamina showing tanniniferous abaxial epidermal cells, adjacent palisade layer and extraxylary fibres, note masses of diosmin crystals

(arrowed) in palisade cells. Scale bars: A, 100 pm; B, 50 pm .

with decumbent branches. Whether they would resprout

after a fire remains to be seen. Two collections were made

from this area, viz. Beyers 176 and McDonald 1780. In all

the material collected by McDonald, the basal portion of

the hypanthium is glabrous as in all collections from other

localities. However, in the material collected by Beyers

the basal portion of the hypanthium on the outside is

tomentose like the rest of the flower. This variation in

hairiness needs further investigation before it can perhaps

be formally expressed taxonomically.

Etymology

The specific epithet, sociorum , is derived from the

Latin socius, meaning associate or colleague. This

species is dedicated to colleagues Ted Oliver and Dave

McDonald, who on field trips always scouted for fresh

Lachnaea material for the first author.

Diagnostic characters and relationships

Lachnaea sociorum is closely related to L. ericoides

Meisn. Both have a similar inflorescence and floral struc-

ture. In both species the lower portion of the hypanthium

is glabrous (except for the one collection of L. sociorum ,

Beyers 176 , where it is tomentose). In L. sociorum the

leaves are usually lanceolate to narrowly elliptic, abaxi-

ally faintly 3-ribbed, with an acute, glabrous apex, the

sepals adaxially tomentose and the stigma penicillate. In

L. ericoides, on the other hand, the leaves are narrowly

52

Bothalia 28,1 (1998)

FIGURE 10. — Lachnaea sociorum, McDonald 2059. SEM micrographs of leaf surfaces illustrating papillate cuticular relief: A, adaxial surface

with dense flake-like wax deposit; B, abaxial surface with sparse flake-like wax deposit. Scale bar: 2 pm .

FIGURE 1 1 — Lachnaea sociorum. SEM micrographs of acetolized pollen grains illustrating crotonoid tectum. A, supratectal subunits with single

spinule, sides emarginate and with surface striate, note cross-links between muri (ridges), indicated by an arrow, McDonald 2059: B, supra-

tectal subunits, each with usually four spinules, sides straight and surface granular-striate, Oliver 10524 . Scale bar: 1 pm .

FIGURE 12. — Known distribution of L. sociorum.

elliptic to obovate, abaxially keeled, with a rounded, tuft-

ed apex, the sepals adaxially sericeous-tomentose and the

stigma capitate with elongated papillae. The leaf anato-

my of these two species is very similar except that in

transverse section the leaves of L. ericoides are deeply

concave adaxially and the extraxylary fibre strands cap-

ping the lateral vascular bundles are not as prominent as

in L. sociorum (Beyers & Van der Walt 1995: fig. 8a).

Specimens examined

WESTERN CAPE. — 3321 (Ladismith): Langeberg, Paardeberg,

1 100 m, (-CD), 1 1 -08-1977, Haynes 1401 (NBG); E of Garcia’s Pass

on northern side of Langeberg, Tygerberg, west of Huisrivier, 300 m,

(-DC), 15-12-1989, Beyers 176 (BOL, K, NBG, NY, PRE, Z), 17-12-

1988, McDonald 1780 (BOL, NBG, PRE); Langeberg, Bergfontein

area, lower slopes of Koksposberg, 425 m, (-DC), 30-10-1990,

Bothalia 28,1 (1998)

53

McDonald 1963 (PRE, NBG); Langeberg, Bergfontem, S-facing mid-

slopes of Koksposberg, 379 m. (-DC), 18-01-1991, McDonald 2059

(BOL, K, NBG, NY. PRE); Riversdale Dist., Langeberg, W of

Witelsberg, upper Witelsrivierkloof, 610 m, (-DC). 01-09-1994, Oliver

10524 (BOL, NBG, PRE); Langeberg, lower SW slopes of Witelsberg,

N of Koksposberg, 488 m, (-DC), 01-09-1994, Oliver 10536 (BOL, K,

NBG, PRE, Z); Mossel Bay Road to Perdekop before forest plantation,

600 m, (-DD), 17-10-1978, Bond 1578 (NBG). 3421 (Riversdale):

Riversdale, Farm Gavelsbos, mountain slope, SW aspect, stony soil, 307

m, (-AB), 08-09-1984, Bohnen 8479 (NBG, PRE). Grid ref. unknown:

Riversdale Flower Show, 01-09-1994, comm. Oliver (NBG); Riversdale

Div„ Langeberg, 01-10-1923, Muir 2824 (BOL), Muir 3053 (PRE).

ACKNOWLEDGEMENTS

The authors would like to thank Mrs Inge Oliver for

the line drawings, and Mr E.G.H. Oliver for translating

the diagnosis into Latin.

REFERENCES

BEYERS, J.B.P. & VAN DER WALT, J.J.A. 1995. The generic delimi-

tation of Lachnaea and Cryptadenia (Thymelaeaceae). Bothalia

5: 65-85.

ERDTMAN, G. 1966. Pollen morphology and plant taxonomy.

Angiosperms. Almqvist & Wiksell, Stockholm.

JACKSON, B.P. & SNOWDON, D.W. 1990. Atlas of microscopy of

medicinal plants, culinary herbs and spices. Belhaven Press,

London.

J.B.P. BEYERS* and A.E. VAN WYK**

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

7735 Claremont, Cape Town.

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

University of Pretoria, 0001 Pretoria.

MS. received: 1997-07-10.

AYTONIACEAE (HEPATOPHYTA)

ASTERELLA ABYSSINICA NEWLY REPORTED FROM SOUTH AFRICA AND MALAWI

A specimen of A. abyssinica, Duckett & Matcham

6049, from Monk’s Cowl, Drakensberg, Kwazulu-Natal.

collected in April 1997, has recently come to hand by the

generosity of Prof. J.G. Duckett, of the University of

London. Besides this, I also collected this species in

1991 in Nyika National Park, Malawi ( S.M . Perold 2664 ,

2672), from where it has not yet been recorded. The

species is described and illustrated here. It is referred to

the subgenus Brachyblepharis (Gottsche et al.) Grolle,

because the short tips of the segments of the pseudoperi-

anth separate at maturity. This is also the first report of

the subgenus Brachblepharis in both South Africa and

Malawi.

Asterella abyssinica ( Gottsche ) Grolle in Vanden

Berghen: 170 (1972).

Fimbriaria abyssinica Gottsche in Gottsche et al.: 569 (1846);

Hypenantron abyssinicum (Gottsche) Trevis.: 441 (1877); Steph.: 122

(1899). Type: Abyssinia cum Targionia elongata, in caespite Un. itin.

n. 500 a Kotschy lecto pauca specimina inventa (not seen).

Thalli smallish to medium-sized, rather delicate and

occasionally somewhat spongy, dorsally flat, green to

light olive-green, crystalline when fresh, margins hyaline

or tinged with mauve to deep purple, outlines of subdor-

sal air chambers faintly visible from above, medianly

elongate and apically directed, laterally in radiating, ±

parallel rows, air pores not visible, encircling cells very

slightly raised when wet; thallus margins flat or raised to

partly inflexed when dry; in crowded, overlying mats,

simple or once pseudodichotomously furcate (Figure

13A), with apical (often immediately distal to foot of

stalk) or latero-ventral innovations from a tapering stipi-

tate base. Branches ligulate to obcordate, widening grad-

ually from a narrow base, sometimes irregularly shaped

and abruptly constricted, when simple, up to 15 mm long,

when branched, total length ± 18 mm long, with terminal

branches ± 5 mm long and moderately divergent, 2. 3-5.0

mm wide, 400^150(-630) pm thick over midrib, laterally

thinning out into attenuate wings, apex slightly notched,

with appendages of few ventral scales recurved over

edge; margins acute, thin, weakly scalloped, slightly

undulate; ventrally the flanks of only the distal part pur-

ple, remainder green, but occasionally entirely purple,

medianly keeled, midrib often green throughout, with

row of purple scales on either side usually only distally

present (Figure 13B). but sometimes along whole length.

Dorsal epidermis unistratose, containing chloroplasts,

cells 5- or 6-sided when seen from above, thin-walled,

(40.0-)45.0-65.0(-72.5) x 22.5-37.5 pm, their orientation

changing from apically directed medianly to outwardly

sloping laterally, in transverse section 25.0-32.5 pm thick

(Figure 13E, F), toward margin occasionally containing an

oil body; marginal cells (Figure 13H) mostly in 1(2)

row(s), long- or short-rectangular, sometimes rather irreg-

ularly shaped, 22.5-30.0 x 12.5-22.5 pm; air pores hard-

ly raised, simple, small, ± 12.5 x 10.0 pm, 80.0-137.5 pm

distant from each other, bounded by innermost circle of

remains of collapsed cells (not shown), and outwardly sur-

rounded by 2 intact, partly overlapping concentric rings of

6 or 7 ± wedge-shaped cells in each (Figure 13G), inner

ones smaller, 10.0-15.0 x 15.0-22.5 pm, outer ones

15.0- 20.0 x 20.0-37.5 pm; assimilation tissue 215-260

pm thick, with small, empty air chambers (Figure 1 3E),

30.0- 100.0 pm wide, in 2 or 3 storeys, toward margins

elongating and sloping obliquely, chlorophyllose cells in

bounding walls rounded or elongated, 32.5-42.5 x

27.5- 30.0 pm; storage tissue confined to keel, ± 220 pm

thick, cells angular, isodiametric, 17.5-27.5 pm wide,

closely packed together, occasional cells with an oil body;

rhizoids arising from ventral epidermis of keel, smooth,

15.0- 20.0 pm wide, or pegged, 12.5-15.0 pm wide.

Scales in 2 forwardly directed ventral rows, one on either

side of midrib (Figure 13B), in different shades of mauve

to purple, obliquely triangular, with a single appendage

(Figure 131), body of scale up to 800 pm long, 650-750

pm wide at base, sometimes crescentic, cells 4—6 sided,

57.5- 65.0 x 27.5-40.0 pm, with up to 11 smaller, colour-

less cells containing remains of oil body; appendage

oblong to elliptical, 420-430 x 300-310 pm, sometimes

constricted at base, ± 200 pm wide, tapering above to a

54

Bothalia 28,1 (1998)

FIGURE 13. — Asterella abyssinica. A, dorsal view of branched thallus with gametoecia on separate branches of same plant; B, ventral view of

thallus; C, carpocephalum raised on stalk; D, ventral view of carpocephalum; E, t/s of thallus; F, t/s of air pore and air chamber; G, air pore

and surrounding cells from above; H, marginal cells of thallus; I, ventral scale; J, t/s of stalk; K, paleae. A-J, Duckett & Matcham 6049\

K, S.M. Perolcl 2672. Scale bars: A-E, 1 mm; F-H, 50 pm; I, J, 250 pm; K, 100 pm. Artist: G. Condy.

Bothalia 28,1 (1998)

55

FIGURE 14. — SEM micrographs of Asterella abyssinica, Duckett & Matcham 6049. A-E, spore: A, B, distal face; C, side view; D, proximal face;

E, much enlarged areolae on proximal face, basally highly porate. F, elater. A, x 543; B, x 586; C, x 590; D, x 594; E, x 4104; F, x 679.

conical apical cell, marginal cells subquadrate, ± 32.5 x

35.0 pm, or rectangular, ± 57.5 x 27.5 pm. inner cells

angular, 32.5-50.0 x 22.5^42.5 pm, only one cell contain-

ing an oil body, ± 27.5 x 25.0 pm,

Autoicous, but androecia and archegoniophores

sometimes on separate plants. Androecia extending back-

wards medianly, occasionally furcate, at apex of branch

or more proximally at constriction (Figure 13 A),

antheridia immersed in sessile, elongated or oval cush-

ions, ± 1375 x 750 pm, lacking scales and opening above

via stout, raised, conical papillae, ± 200 pm long.

Archegoniophores proximal to apical notch of main

branch (Figure 13C) or just proximal to stipitate innova-

tion of apical branch (Figure 13 A), single or occasional-

ly paired at apices of 2 forking branches, almost sessile

and hemispherical when young, paleae hidden (Figure

13 A). Carpocephala at maturity raised on stalk, arising ±

1.7 mm proximal to apex of branch in apical notch,

length 4-5 mm, whitish green or streaked with purple,

widening slightly toward base, in transverse section at

midlength (Figure 13J), ± 520 x 450 pm. weakly ribbed,

with a single rhizoid furrow containing pegged rhizoids,

cortical cells not well differentiated, except for their

outer walls being slightly thickened and rounded, vari-

able in size, 12.5-22.5 x 12.5-15.0 pm. medullary cells

angular, 15.0-35.0 x 15.0-27.5 pm; scattered along

length of stalk a few hyaline or purple paleae, 400-500 x

160-180 pm, at its summit numerous paleae (Figure

13K), hyaline or purple, up to 820 pm long, basally 2 or

3 cells wide, above 3-5 cells wide, 65.0-85.0 x

35.0-47.5 pm, apical cell 42.5-62.5 x 17.5-22.5 pm;

disc green, rather flat to weakly convex, suborbicular

(Figure 1 3C), 3. 5-4.0 mm diam., ±1.1 mm thick, margin

undulate, air chambers covered above by slightly bulging

walls and opening via small, compound air pores, below

(Figure 13D) with membranous, bistratose involucres

partly covering capsules like a flap, margin entire and

unistratose, capsules mostly 2, rarely 1 or 3(4), obovate,

1200 x 950 pm, wall green, unistratose, cells elongated,

4- or 5-sided, 45.0-75.0 x 25.0-37.5 pm, thin-walled,

toward apex of capsule rounded, 35.0-47.5 x 37.5-42.5

pm, walls with trigones at corners; pseudoperianths

exserting horizontally from beneath rim of car-

pocephalum (Figure 13C) for ± 1000 pm, colourless,

split into 8-10 segments, these up to 1375 pm long and

340 pm wide at base, tapering to slender tips, which are

initially attached, becoming free before capsule dehis-

cence. Spores 62.5-72.5 pm diam., triangular-globular,

pale yellow to yellow, translucent, wing undulate, up to

10 pm wide, margin crenulate, distal face (Figure 14A,

B) convex, reticulate, with network of ± 6 larger, prima-

ry areolae across, ± 10-15 pm wide and extending over

wing, their delimiting ridges only slightly raised and not

clearly defined among the numerous criss-crossing walls

of small, subsidiary areolae, areolar floor highly porate;

proximal face with prominent triradiate ridge (Figure

14C, D), its arms continuous from pole to wing, but not

clearly extending across, each of 3 facets with numerous

areolae, larger at pole and smaller toward wing, but not

clearly arranged in primary and subsidiary areolae, floor

also highly porate (Figure 14E). Elaters yellow, 135-150

pm long, 10 pm wide in centre, slightly tapering to

rounded ends, ± 7.5 pm wide, with coiled, bispiral thick-

enings throughout (Figure 14F).

DISCUSSION

Asterella abyssinica is widespread in tropical Africa

and has been reported by various authors from Burundi,

Cameroun, Ethiopia, Rwanda, Sierra Leone, Tanzania,

Zaire and Zimbabwe (Wigginton et al. 1996). These are

the first records, however, for South Africa (Figure 15)

and Malawi. The species is generally found along stream-

56

Bothalia 28,1 (1998)

Amended key to the southern African species of Asterella (see Perold 1994)

la Pseudoperianth with segments apically free at maturity subgenus Brachyblepharis 1. A. abyssinica

lb Pseudoperianth with segments remaining apically attached at maturity Subgenus Phragmoblepharis:

2a Thalli spongy, with tall air chambers mostly in one storey, and then not subdivided by supplementary partitions, each opening dor-

sally by a stellate pore; ventral scales occasionally fimbriate at single lanceolate appendage; carpocephalum round or umbonate

and lacking paleae at summit of stalk; pseudoperianth extending ± 300 pm beyond involucre and subdivided into 12 or 13 seg-

ments; spores 75-95 pm diam., dark brown, ornamentation with irregular zig-zagging ridges 2. A. muscicola

2b Thalli compact, firm, with small, low air chambers in several storeys, only some top ones opening above by a dorsal, non-stellate

pore; ventral scales with 1 (or 2) lanceolate or ovate appendages, margin ± entire; carpocephalum round or umbonate, papillose

or ± smooth, with paleae at summit of stalk; pseudoperianth extending more than 1000 pm beyond involucre and subdivided into

14—16 segments; spores more than 100 pm diam., yellow or brown, ornamentation with larger areolae generally containing sub-

sidiary areolae:

3a Thalli smallish to large; carpocephalum covered with distinct papillae, projecting ± 200 pm; paleae at summit of stalk shaggy, dense,

pale mauve or colourless, up to 3000 pm long and 4 or 5 cells wide at base; ventral scales with lanceolate appendage; spores

elaborately ornamented with 6-8 areolae across distal face, 20-30 pm wide and ridges extending across wing, containing

numerous subsidiary areolae (common, mostly summer rainfall species) 3. A. bachmannii

3b Thalli medium-sized to very large; carpocephalum ± smooth or with low papillae; paleae at summit of stalk colourless or purple,

length variable; ventral scales with lanceolate or ovate appendages, spores less elaborately ornamented:

4a Thalli medium-sized; carpocephalum with umbonate head; paleae at summit of stalk mostly colourless, some very long, more than

8000 pm in length, ± 4 cells wide at base; ventral scales with 1 or 2 lanceolate appendages; spores on distal face with (4 — )6— 9

areolae across, ± 32 pm wide, very high ridges seldom extending across wing, usually containing small subsidiary areolae

(winter rainfall species) 4. A. marginata

4b Thalli large to very large; carpocephalum with rounded head, distinctly lobed below; paleae at summit of stalk almost colourless

to purple, 2000-3000 pm long, some up to 7 cells wide at base; ventral scales with single, large-celled, ovate appendage, con-

stricted at base; spores on distal face with 5-8 areolae across, 25-30 pm wide and extending to wing margin, almost empty

of subsidiary areolae and hollow (summer rainfall, mostly Afromontane species) 5. A. wilmsii

banks or on damp rocks. The thalli are rather thin and

delicate and seemingly not adapted to drought condi-

tions. The species can be recognised by the flattened

discs of the carpocephala and by the horizontally pro-

truding pseudoperianths, their rather short segments

becoming free at the tips before the capsules dehisce.

SPECIMENS EXAMINED

South Africa

KWAZULU-NATAL, — 2929 (Underberg): Drakensberg Park,

Monks Cowl, near streambank on path to Crystal Falls, (-AB), 1 800

m, 21 April 1997, Duckett & Matcham 6049 (dupl. E, PRE), with A.

bachmannia and Targionia hypophylla.

Malawi

NYIKA NATIONAL PARK. — 1033: on path across river, leading

to Juniper Forest, on soil of vertical bank, (-DB), 15 April 1991, S.M.

FIGURE 15. — Distribution of Asterella abyssinica in South Africa.

Perold 2664 (PRE); at bridge over Chelinda River, on rock and stone

wall, (-DA), 16 April 1991, S.M. Perold 2672 (dupl. E, PRE).

ACKNOWLEDGEMENTS

I sincerely thank Prof. J.G. Duckett for generously

giving me a part of his and Matcham’s collection of

Asterella abyssinica from Monk’s Cowl to deposit at

PRE. Further thanks are due to Dr D.G. Long, for identi-

fying my two Malawi specimens of A. abyssinica and for

kindly refereeing this manuscript; also to the artist, Ms

G. Condy, the typist, Ms D. Maree and the photographer,

Mrs A. Romanowski, for their valued contributions.

REFERENCES

GOTTSCHE, C M., LINDENBERG, J B G. & NEES AB ESEN-

BECK, C.G. 1846. Synopsis hepaticarum. Meissner, Hamburg.

PEROLD, S.M. 1994. Studies in the Marchantiales (Hepaticae)

from southern Africa. 6. The genus Asterella (Aytoniaceae:

Reboulioideae) and its four local species. Botlialia 24:

133-147.

STEPHANI, F. 1899. Species hepaticarum. Bulletin de PHerbier

Boissierl: 122, 123.

TREVISAN DE SAINT-LEON, V 1877. Schema di una nuova classi-

ficazione delle Epatiche. Memo lie del Reale Istituto Lombardo

di Scienze e Lettere, Ser. 3, 4: 383 — 45 1 .

VANDEN BERGHEN, C. 1972. Hepatiques et Anthocerotees.

Resultats scientifiques de l’ exploration hydrobiologique du

Basin Lac Bangweolo & Luapula 8: 1-202.

WIGG1NTON, M. & GROLLE, R. Supplemented by A Gyarmati.

1996. Catalogue of the Hepaticae and Anthocerotae of Sub-

Saharan Africa. Bryophytorum Bibliotheca Band 50. Cramer,

Berlin, Stuttgart.

S.M. PEROLD*

* National Botanical Institute, Private Bag X 10 1 , 0001 Pretoria.

MS. received: 1997-06-01.

Bothalia 28,1 (1998)

57

FABACEAE

A NEW SPECIES OF PEARSONIA FROM MPUMALANGA, SOUTH AFRICA

When the artist Anita Fabian was illustrating Wild

flowers of northern South Africa (Fabian & Germishuizen

1997), she collected and illustrated a specimen of

Pearsonia Diimmer from the Farm Paardekraal, near

Dullstroom during 1991. The specimen was examined at

the National Herbanum and was found to represent an

undescribed species. For convenience, the caption of the

illustration was left as Pearsonia sp. in Fabian &

Germishuizen (1997). Subsequently more specimens of

this taxon were found housed in the collections of PRE.

Pearsonia hirsuta Gemiish., sp. nov. P. aristatae

(Schinz) Diimmer similis sed habitu minore, foliis ses-

silibus ad subsessilibus, linearibus, dense pubescentibus,

calycibus dense pubescentibus, area mediana alorum

vexillique pilis densis tecta, differt.

TYPE. — Mpumalanga, 2530 (Lydenburg): Dull-

stroom, Verlorenvallei Farm 95 JT, (-AC), 29-11-1980,

Drews 215 (PRE, holo.). Figure 16.

Small erect herb up to 150 mm tall, often resprouting

from a perennial woody rootstock after burning in early

spring; stems densely clothed with long, white to golden

brown (especially on younger parts), appressed to spread-

ing hairs. Leaves exstipulate, sessile to subsessile; leaflets

3, linear to narrowly lanceolate, narrowed to an acute tip,

densely white to golden brown, hairy on both surfaces,

margin inrolled; mid-leaflet (6— )7— 1 2(— 16) mm long, up to

1 mm wide; lateral leaflets 5— 8(— 1 2) mm long, up to 1 mm

wide (Figure 17A). Racemes lax, unbranched, few-flow-

ered, terminal or mostly axillary on lower stems; pedicels

up to 3 mm long; bracts ovate, 5-9 x up to 3 mm, persis-

tent; bracteoles linear to narrowly lanceolate, 5-9 x up to

1 mm. Flowers yellow. Calyx 15-20(-22) mm long,

densely covered with white appressed or spreading hairs;

lateral lobes paired, narrowly triangular, up to 4 mm long;

carinal lobe linear, up to 10 mm long; lateral sinus two to

four times as deep as upper sinuses (Figure 17B). Standard

elliptic to broadly ovate, 18-22 x 10-14 mm, lateral mar-

gins strongly reflexed, dorsal surface densely hairy in a

broad median band from middle of lamina to apex outside,

margin crinkled or crisped at apex (Figure 17C). Wing

petals narrowly obovate, 15-20 x 3-5 mm; sculpturing

present, upper middle comprising transcostal lamellae and

densely bearded with white to golden brown appressed

hairs. Keel petals small, narrow, 14-16 x 2 mm; shallow

basal pockets present on outside, up to 0.5 mm deep.

Stamens monadelphous; sheath 12-15 mm long, split

adaxially to base; anthers 1.5 mm long, alternately basi-

FIGURE 16. — Holotype of Pearsonia hirsuta Germish.

FIGURE 17. — Pearsonia hirsuta, Drews 215, holotype. A, flower-

ing plant, x 1; B, calyx, x 1.5; C, standard, x 1.5; D, wing petals, x 1.5;

E, keel petals, x 1.5; F, style, x 1.5; G, stamens, x 1.5. Drawing by G.

Condy.

58

Bothalia 28,1 (1998)

TABLE 2. — Differences between Pearsonia hirsuta and P. aristata

fixed and dorsifixed (Figure 17G). Ovary densely white-

pilose, many-ovuled; style straight, glabrous; stigma capi-

tate (Figure 17F). Fruit and seeds unknown.

Superficially Pearsonia hirsuta and P. sessilifolia

(Harv.) Dtimmer subsp .filifolia (Bolus) Polhill resemble

one another. Polhill (1974), in his revision of the genus

Pearsonia , describes the latter taxon as being a suffrutex

with generally long, elongate, many-flowered racemes

and the standard 8—1 3(— 1 5) x 3. 3-8.0 mm. P. hirsuta

however, comprises short, few-flowered racemes and the

standard is 18-22 x 10-14 mm. Differences between P.

hirsuta and P. aristata are listed in Table 2.

Pearsonia hirsuta and P aristata are allopatric. P. hir-

suta occurs in the Lydenburg District of Mpumalanga

(Figure 18), growing in low grassland between rocks in

humus-rich sandy soil, whereas P. aristata occurs

throughout the Northern Province, Mpumalanga, Swazi-

land, northern KwaZulu-Natal and into Zimbabwe.

Flowering takes place during November. The specific

epithet ‘ hirsuta ’ refers to the hairy nature of this species.

Specimens examined

MPUMALANGA. — 2530 (Lydenburg): lower foothills of

Steenkampsberg, ± 34.5 km from Lydenburg on road to Roossenekal,

(-AB), Burgoyne 2046 (PRE); Dullstroom, Verlorenvallei Farm 95 JT,

(-AC), Drews 215 (PRE); Long Tom Pass, near cannon, (-BA),

FIGURE 18. — Distribution of Pearsonia hirsuta Germish. in South

Africa.

Burgoyne 2011 (PRE); Dullstroom District, crown of Farm

Paardekraal, ‘Oribi Falls’, (-CA), Fabian 1396 (PRE).

ACKNOWLEDGEMENTS

The author would like to thank Adela Romanowski

for the photographs, Gill Condy for the drawing and Dr

Otto Leistner for translating the diagnosis into Latin.

REFERENCES

FABIAN, A. & GERMISHUIZEN, G. 1997. Wild flowers of northern

South Africa. Femwood Press, Vlaeberg, Cape Town.

POLHILL, R.M. 1974. A revision of Pearsonia Diimmer (Legum-

inosae-Papilionoideae). Kew Bulletin 29; 383-410.

G. GERMISHUIZEN*

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

MS. received: 1997-09-17.

FABACEAE

A NEW SPECIES OF RHYNCHOSIA FROM SOUTH AFRICA

The genus Rhynchosia is currently under revision.

Herbarium and field studies have led to the discovery of

an undescribed species which closely resembles R. carib-

aea.

Rhynchosia atropurpurea Germish., sp. nov. R.

carihaeae habutu similis sed foliis valde minoribus, stip-

ulis maioribus, vexillo et alis valde brevioribus quam

carina, et carina omnino atropurpurea. Figure 19.

TYPE.— North-West, 2527 (Ruslenburg): Pilanesberg

Game Reserve, near Saulspoort, (-AA), 5-1-1989,

Germishuizen 4977 (PRE, holo.).

A much-branched shrub, scrambling herb or perennial

twiner up to 1 m tall; stems densely clothed with thin

appressed soft hairs, especially on young parts, sometimes

interspersed with patent, bulbous-based glandular hairs

and raised golden glands, becoming glabrous and shiny

with age. Leaflets pinnately 3-foliolate; upper surface

sparsely pubescent, with a few scattered glands, glabres-

ccnt; lower surface with prominent nervation, densely

pubescent especially on margins and veins, with small

raised golden glands scattered in intervena] areas; terminal

leaflet 1 6— 25(— 33) x 16-22(-36) mm, broadly rhomboid

or deltoid, gradually narrowed to an obtuse or subacute

apex; lateral leaflets 1 5— 22(— 30) x 1 1 — 20(— 22) mm, with

lower margin distinctly gibbous; petiole 15-26 mm long;

petiolules 5-12 mm long, canaliculate, stipellate (Figure

20A). Stipules up to 3.5 mm long, subulate, appressed,

densely pubescent and glandular outside, glabrous inside.

Inflorescences axillary, lax, unbranched, up to 1 3-llowered

racemes; bracts up to 3 x 2 mm, ovate, reddish brown,

pubescent, caducous; ebracteolate. Flowers yellow with

purple venation, keel deep purple to maroon. Calyx finely

pubescent, glandular with raised golden glands and bul-

Bothalia 28,1 (1998)

59

FIGURE 19. — Holotype of Rhynchosia atropurpurea Germish.

bous-based hairs; tube 3.5— 4.5 mm long; lobes narrowly

triangular, carinal lobe up to 7 mm long, longer than other

lobes (Figure 20B). Standard 15—16 x 9-13 mm, broadly

obovate to suborbicular, narrowing into an auriculate claw,

sparsely sericeous with raised golden glands especially

near apex on outside (Figure 20D). Wing petals up to 8

mm long, obovate to slightly cymbiform, slightly pouched

(Figure 20C). Keel petals 17 mm long; pocket present on

outside towards base, up to 1 mm deep; auricles present

(Figure 20E). Stamens diadelphous; sheath split adaxially;

anthers up to 1 mm long, alternately basifixed and dorsi-

fixed (Figure 20F). Gynoecium up to 20 mm long; ovary

2-ovulate, densely white pilose; style bent in upper third,

pubescent to point of flexure (Figure 20G). Fruit 35-44

mm long, oblong-falcate, narrowed towards base, recurv-

ing downwards at tip, conspicuously veined, covered with

bulbous-based glandular hairs and raised golden glands.

Seeds unknown.

Although R. atropurpurea and R. caribaea are super-

ficially similar, R. atropurpurea differs from the latter in

having much smaller leaves, larger stipules, the standard

and wing petals much shorter than the keel petals and the

keel dark purple to maroon throughout (in contrast to the

yellow keel with maroon tip found in R. caribaea).

Rhynchosia atropurpurea occurs in the Northern

Province, North-West and Mpumalanga (Figure 21),

occupying a diverse range of habitats from ravine bush.

FIGURE 20. — Rhynchosia atropurpurea, G. Germishuizen 4977 .

A, flowering branch, x 1 . B-G, x 1.5: B, calyx; C, wing petals; D, stan-

dard; E, keel petal; F, stamens; G, style. Drawing by G. Condy.

bushveld, to dense grassland with scattered trees on steep

rocky granite hills. In contrast, R. caribaea occurs

throughout southern Africa, except Lesotho. Flowering

takes place from January to March, with fruiting concen-

trated in March and April.

The specific epithet ‘ atropurpurea ’ refers to the charac-

teristic dark purple keel of this species — in other species

the purple marking is restricted to the apex of the keel.

Specimens examined

NORTHERN PROVINCE. — 2327 (Ellisras): Mogol Nature Reserve,

(-DD), Fourie 2588 (PRE). 2427 (Thabazimbi): Thabazimbi, western

road, 5 km alongside road, (-CB), Nel 564 (PRU). 2430 (Pilgrims

FIGURE 21. — The known distribution of Rhynchosia atropurpurea

Germish.

60

Bothalia 28,1 (1998)

Rest): Abel Erasmus Pass, (-BC), Van Wyk, Dahlgren & Kok 5477

(PRE, PRU); beginning of Abel Erasmus Pass, (-BC), Strey 3281

(PRE); Hoedspruit Dist., Chester Farm, (-BD), Burgoyne 4090 (PRE);

NORTH-WEST. — 2527 (Rustenburg): Pilanesberg Game Reserve,

near Saulspoort, (-AA), Germishuizen 4977 (PRE).

MPUMALANGA. — 2430 (Pilgrims Rest): foot of Kaspers Nek, on

Ohrigstad road, (-DA), Hilliard 4722 (PRE); Mogaba California

228 KT, (-DA), Raal & Raal 1743 (PRE). 2529 (Witbank): 5 km from

Groblersdal on road to Marble Hall, (-AB), Germishuizen 3722 (PRE).

2531 (Komatipoort): Crocodile Valley Farm, 6 km from Nelspruit on

Friedenhau-Karino road, (-AC), Coetzer 823 (PRE, PRU); 15 km from

Barberton on road to Kaapmuiden, (-CA), Coetzer 122 (PRE);

Claremont Vale, 10 km north of Barberton, (-CA), De Sousa 229 (PRE).

ACKNOWLEDGEMENTS

The author would like to thank Adela Romanowski

for the photograph of the holotype, Gill Condy for the

drawing and Dr Hugh Glen for translating the diagnosis

into Latin.

G. GERMISHUIZEN*

* National Botanical Institute, Private Bag X 1 0 1 , 0001 Pretoria.

MS. received: 1997-08-08.

ASPHODELACEAE: ALOOIDEAE

ASTROLOBA CORRUGATA: DESCRIPTION OF A LONG-KNOWN SPECIES IN A SOUTHERN AFRICAN

ENDEMIC ALOOID GENUS

Whilst compiling an account of the small southern

African endemic genus Astroloba Uitewaal (Aspho-

delaceae: Alooideae) for Vol. 3 of the Synopsis Plantarum

Succulentarum Project, co-ordinated by the Stadtische

Sukkulenten Sammlung in Zurich, Switzerland, it came to

our attention that a well-known and frequently cultivated

species of the genus has not been validly named.

The species is most commonly referred to as

Astroloba aspera (Haw.) Uitewaal, but based on the orig-

inal description of Aloe aspera provided in the proto-

logue by Haworth (1804), this combination was estab-

lished for a species of the genus Haworthia Duval. This

is evident from the description of the leaf arrangement of

Aloe aspera as ‘foliis trifariis' (Haworth 1804: 6). The

leaves of species of Astroloba are arranged in five, not

three, distinct rows, as the stems are viewed from above.

This character has been adequately and unambiguously

recorded as ‘ foliis quinquefariis’ in the description of e.g.

Aloe pentagona (Aiton) Haw., which also appeared in

Haworth (1804: 7).

If Haworth (1804) had given an adequate description

of the flowers of the species, it could have given further

clues to the exact identity of the novelty that he described

as Aloe aspera. At the time, the flowers of species of

Haworthia were usually simply described as 'in labio

duo', referring to the two-lipped flowers which are rather

consistently encountered in Haworthia (see for example

Haworth 1812: 90). Exceptions to this rule are represen-

tatives of Haworthia. subg. Robustipeduncularis Uite-

waal ex Bayer where the perianth somewhat resembles

those of some species of Astroloba.

Eight years later, in 1812, the leaf arrangement of Aloe

aspera, then clearly treated as a species of Haworthia,

was again given as Hilarious by Haworth. Furthermore, in

his enumeration of the species he grouped his Haworthia

aspera with the caulescent species of Haworthia, for

example H. viscosa, and not with the other species nowa-

days included in Astroloba. The floral morphology was,

however, clearly given as bilabiate, supporting the view

that Haworthia aspera (Haw.) Haw. was in fact a species

of Haworthia as generally circumscribed today. This

name, H. aspera, was recently re-instated by Parr (1971),

following his proposal to include Astroloba in Haworthia.

We do not uphold that interpretation.

Earlier, in 1811, Willdenow had the same view as

Haworth (1812), although he included this species in the

newly created genus Apicra Willd., a group which, in his

concept, included all the species of Haworthia and

Astroloba. The genus name Apicra, which was therefore

an illegitimate renaming of Haworthia (Duval 1809),

was unjustifiably taken up by Haworth in 1819 more or

less solely for the species of Astroloba, as circumscribed

today, and again the leaf arrangement of Apicra aspera

(Haw.) Willd. was given as trifarious.

Salm-Reifferscheid-Dyck (1817), who preferred the

Linnean concept of Aloe L. for all species currently dis-

persed amongst the genera Aloe, Astroloba , Gasteria

Duval and Haworthia, accepted Haworth’s (1804) inter-

pretation of the trifariously leaved Aloe aspera. However,

in 1840 Salm-Reifferscheid-Dyck extensively adapted

the original description of Haworth by, amongst others,

referring to the leaves of his Aloe aspera as 'foliis spi-

raliter quinquefariis' . By so doing he essentially

described a new species, but made the critical mistake of

adopting Haworth’s (1804) name, Aloe aspera. As argued

above, this was undoubtedly a species of Astroloba and

not Haworthia, since the flowers were unambiguously

described as 1 limbo regulari'. The species as known

today therefore does not have a legitimate name.

The final combination of the epithet aspera in the

genus Astroloba was made by Uitewaal in 1947 (Smith

et al. 1994; Smith & Van Wyk 1996). As basionym for

this new combination he cited Haworth’s (1804) concept

of the species.

More recently two researchers working independent-

ly came to the same conclusion as us. Roberts Reinecke

(1965), in an unpublished M.Sc. thesis proposed the

name Astroloba rugosa for the species, whereas Groen

(1987) proposed the name A. muricata. Neither of these

names have been validly published and to prevent further

confusion, we decided to choose the name A. corrugata

N.L.Mey. & G.F.Sm. for this unnamed species.

A single variety, var. major, has been described in

Apicra aspera by Haworth (1819). This variety was later

Bothalia 28,1 (1998)

61

transferred to the genus Astroloba , as Astroloba aspera

var. major (Haw.) Uitewaal (1947). In our opinion this

entity does not warrant recognition at any rank, and

should be included in the synonymy of A. corrugata.

In summary, the following names have been misap-

plied to this species: Aloe aspera Haw. (1804) and sensu

Salm-Reifferscheid-Dyck (1817) not of Salm-Reiffer-

scheid-Dyck (1840); Apicra aspera (Haw.) Willd.

(1811); Astroloba aspera (Haw.) Uitewaal (1947); and

Haworthia aspera (Haw.) Haw. (1812) and sensu Parr

(1971). The names Astroloba rugosa Roberts Reinecke

ined. (1965) and Astroloba muricata L.E.Groen nom.

prov. ined. (1987) have been proposed for Astroloba cor-

rugata, but were never formally published.

Astroloba corrugata N.L.Mey. & G.F.Sm., sp.

nov. ab aliis speciebus tuberculato-foliatis Astrolobae

apicibus non-marginatus foliorum, distribtione aequiore

densioreque tuberculorum staturaque parviore foliorum

differt.

TYPE. — Western Cape, 3320 (Montagu): Warm-

waterberg, 7.5 km west of Warmwaterberg turnoff on

Montagu-Ladismith road, (-BD), 27-02-1994, E.J. van

Jaarsveld 13913 (PRE. holo.).

Herbaceous, succulent perennial; caulescent, with

leaves carried in ± 5 straight rows on vertical stem, up to

300-600 mm tall, 20-25 mm diam., solitary or prolifer-

ous from base and forming clusters. Leaves rigid, the

young erect, the old spreading to more or less horizontal,

14—25 mm long, 11-18 mm broad at base, up to 5 mm

thick, deltoid-ovate or lanceolate, sharply tapering, often

twisted to one side in a spiral arrangement, tip pungent,

light to dark green; upper surface flat to concave, tuber-

cles concolorous, shiny, fairly evenly distributed, but

tending to form raised longitudinal bands, up to 0.5 mm

diam.; single distinct or indistinct keel occurring central-

ly or obliquely in distal third of leaf on upper surface,

tubercled or with transverse ridges, keel not forming mar-

gin at apex; lower surface convex, similar to upper sur-

face; margins upcurved, acute or rounded, rough.

Inflorescence up to 430 mm long; peduncle simple, terete,

2-4 mm diam. at base, bracteate; sterile bracts membra-

nous, ovate, acuminate, 4-8 mm long, erect, centrally

keeled with reddish brown vein; raceme up to 270 mm

long, lax, with 9-30 spirally arranged flowers and buds,

2-5 opening simultaneously; floral bracts membranous, ±

5 mm long, deltoid, acute, keeled with reddish brown

vein, clasping pedicels, shorter than or as long as

pedicels; pedicels erect, up to 9 mm long, up to 1 mm

diam., green. Flowers subactinomorphic, funnel-shaped,

white or cream with pink or greenish tinge, midribs of

perianth segments green with beige or pink tinge; tube

regular, ± straight, slightly decurved apically, ± 4 mm

across, constricted to 3 mm above; segments closely

coherent, fused towards base; outer segments not adja-

cent, spoon-shaped, retuse at tips; tips flared, < 1 mm

long. Stamens 6 of two lengths, 5 and 6 mm long, insert-

ed within perianth tube. Ovary' 4x2 mm, green; style 4

mm long, slightly curved, subcapitate, yellowish green.

Fruit a trilocular capsule, cylindrical, apically retuse, ± 1 2

x 5 mm diam. Seed dark grey, angled, shortly winged.

FIGURE 22. — Astroloba corrugata. Artist: Marietjie Steyn.

Chromosome number: 2 n = 14 (Ferguson 1926; Ma-

jumdar 1968; Snoad 1951).

Diagnostic characters

Astroloba corrugata differs from the other tubercu-

late-leaved species, although glabrous members have

been observed, in the genus by the non-marginate leaf

62

Bothalia 28,1 (1998)

ta.

tips and a more even, denser distribution of tubercles, as

well as smaller leaf size. Furthermore, the affinities of A.

corrugata would seem to lie with those species of the

genus that lack a marked inflation of the perianth tube,

viz. A. bullulata (Jacq.) Uitewaal, A. congesta (Salm-

Dyck) Uitewaal and A. foliolosa (Haw.) Uitewaal. Only

two smooth-leaved species, A. herrei Uitewaal and A.

spiralis (L.) Uitewaal show this trait.

Etymology

The epithet corrugata refers to the wrinkled appear-

ance of the leaves, imparted by the thickly dispersed,

rough tubercles (Figure 22).

Distribution

Astroloba corrugata is widely distributed in the

Ladismith, Montagu, Riversdale, Swellendam and

Worcester Districts of the Western Cape Province of

South Africa (Figure 23).

Conservation status

Astroloba corrugata is not threatened in any way

(Hilton-Taylor & Smith 1994; Hilton-Taylor 1996).

ACKNOWLEDGEMENTS

The authors are indebted to Dr O.A. Leistner for the

Latin diagnosis, and Mrs E. du Plessis for comments on

a first draft of the paper.

REFERENCES

DUVAL, H.A. 1809. Plantae succulentae in horto Alenconio. Apud

Gabon, Paris.

FERGUSON, N. 1926. V. The Aloineae: a cytological study, with espe-

cial reference to the form and size of the chromosomes.

Philosophical Transactions of the Royal Society of London, B

215: 225-253.

GROEN, L.E. 1987. Astroloba Uitew. (111). Succulenta 66: 82-87.

HAWORTH, A H. 1804. A new arrangement of the genus Aloe , with a

chronological sketch of the progressive knowledge of that

genus, and of other succulent genera. Transactions of the

Linnean Society of London 7: 1-28.

HAWORTH, A H. 1812. Synopsis plantarum succulentarum. Richard

Taylor, London.

HAWORTH, A H. 1819. Supplementum plantarum succulentarum.

Harding, London.

HILTON-TAYLOR, C. 1996. Red Data List of southern African plants.

Strelitzia 4: 1-117. National Botanical Institute, Pretoria.

HILTON-TAYLOR, C. & SMITH, G.F. 1994. The conservation status

of Aloaceae in southern Africa. In B.J. Huntley, Botanical diver-

sity in southern Africa. Strelitzia 1: 287-303. National

Botanical Institute, Pretoria.

MAJUMDAR, S.K. 1968. Morphology, cytogenetics and evolution of

the tribe Aloineae. Ph D. dissertation, Graduate School,

University of Kentucky, Lexington.

PARR, C.A.E. 1971. Revision of the genus Astroloba. Part II. Bulletin

of the African Succulent Plant Society 6: 145-150.

ROBERTS REINECKE, P. 1965. A revision of the genus Astroloba.

M.Sc. thesis. University of Cape Town.

SALM-REIFFERSCHEID-DYCK, J.M.F.A.H.I. 1817. Catalogue

raisonne des especes et varietes dAloes. Diisseldorf.

SALM-REIFFERSCHEID-DYCK, J.M.F.A.H.I. 1840. Aloe aspera.

Fasc. 3, Fig. 1; [Sect. 2, Fig. 2], Monographiae generum Aloes

et Mesembryanthemi. Bonn.

SMITH, G.F., MEYER, N.L. & GLEN, H.F. 1994. Little-known gener-

ic names in the family Aloaceae. South African Journal of

Science 90: 489, 490.

SMITH, G.F. & VAN WYK, A.E. 1996. Asphodelaceae. Generic num-

bers in the subfamily Alooideae. Bothalia 26: 158.

SNOAD, B. 1951. Chromosome numbers of succulent plants. Heredity

5: 279-283.

UITEWAAL, A.J.A. 1947. Revisie van de nomenclatuur der genera

Haworthia en Apicra. Succulenta No. 5, Sept./Oct. 1947:

51-54.

WILLDENOW, C.L. 1811. Bemerkungen iiber die Gattung Aloe. Der

Gesellschaft naturforschender Freunde zu Berlin Magazin fur

die neuesten Entdeckungen in der gesammten Naturkunde 5

(Neue Schreibe): 163, 164, 265-283.

N. L. MEYER' & G. F. SMITH'

‘Research Directorate, National Botanical Institute, Private Bag X 1 0 1 ,

0001 Pretoria.

MS. received: 1997-09-09.

HYACINTHACEAE

A NEW SPECIES OF ORNITHOGALUM FROM THE RICHTERS VELD, SOUTH AFRICA

INTRODUCTION

Obermeyer (1978) recognised 54 species of Orni-

thogalum L. from southern Africa of which about 15

species, including the newly described species, occur

within the Richtersveld. In a recent publication U. & D.

Miiller-Doblies (1996) have extended the number of

recognised species to 123 of which about 30 are recorded

as occurring in the Richtersveld. These plants survive in

all evolutionary niches from the highest mountains to the

Bothalia 28,1 (1998)

63

FIGURE 24. — Ornithogalum decus-montium, G. & F. Williamson 5803 (NBG). A, plant; B, leaf flattened (including basal sheathing portion); C,

flower; D, inner and E, outer tepal; F, inner and G, outer stamen; H, ovary, style and stigma; I, capsule; J, seeds, side and front views. Scale

bars; A, B, 45 mm; C, 16 mm; D, E, 7 mm; F, G, 2.25 mm; H, 2.5 mm; I, 4 mm; J, 1.5 mm. Drawn by G Williamson.

low, more arid sandy plains. O. decus-montium appears to

be restricted to the northern Richtersveld mountains.

Ornithogalum decus-montium G.Will., sp. nov.

ad subgenus Aspasiam et turmam Aspasiae pertinens.

Species insignis apice ovarii nigrovirenti, a speciebus

nobis notis bene distincta. O. geniculatum Oberm. primo

adspectu simile, sed non-geniculato pedunculo satim

diagnoscenda.

Planta bulbosa 40-100 mm alta (bulbo incluso); foliis

2, coaetaneis, anguste linearibus, subacutis, 60-100 mm

longis (vagina inclusa); scapo 60-100 mm alto, racemo

ad 30 x 25 mm; seminibus commaformibus.

TYPE. — Northern Cape, 2816 (Oranjemund): Rich-

tersveld National Park, ± 20 km E of Sendelingsdrift,

(-BB), Sept. 1995, G. & F. Williamson 5803 (NBG, holo.).

Plant a dwarf geophyte 40-100(-120) mm high

including bulb (Figure 24A). Bulb ovoid, 7x7 mm, outer

tunic thin and brittle, pinkish brown with aerenchyma-

tous layer ± 1 mm thick. Leaves 2, emerging from basal

sheathing papyraceous cataphylls 3 mm long, synan-

thous, longest as long as scape, sheathing at base for

20-30 x 4 mm at widest, free portion light green, of soft

texture, drooping to suberect, linear narrowing towards

apex, subacute, 30-70 x 1. 5-3.0 mm, glaucous with a

smooth margin. Scape fine, wire-like, up to 100 mm;

raceme up to 30 x 25 mm, 2-8-flowered, flowers subco-

rymbosely clustered and mostly facing skywards

(Figures 24A, C; 25), bracts very narrowly linear, mem-

branous, acute, 4-14 mm long; pedicels ± 10 mm long,

suberect to erect, thin, wire-like, elongating up to 14 mm

when capsule fully developed. Tepals shiny satin-white,

outer narrowly elliptic, acute, 7x3 mm, inner elliptic,

acute, 7x4 mm (Figure 24D, E). Stamens: filaments ±

64

Bothalia 28,1 (1998)

FIGURE 25. — Ornithugalum decus-montium. Flower photographed

east of Sendelingsdrift.

4.5 mm long, narrowing towards anthers, terete; inner

with an oblong base; anthers rounded oblong, yellowish,

all facing stigma, 0.8 mm long (Figure 24F, G). Ovary

green, ovoid with apex very distinctly coloured luminous

dark greenish black, 2.5 x 2.0 mm; style terete, ± 1 mm

long; stigma with 3 decurrent, papillate lobes, ± 0.5 mm

long (Figure 24H). Capsule ellipsoid, green with longitu-

dinal white stripes and darkened apex, 4.0 x 2.8 mm wide

(Figure 241); seeds coal black, comma-shaped, densely

tuberculate, 1.5 x 0. 5-0.8 mm (Figure 24J).

rounding decomposing orange-brown igneous rocks. The

main geological formation is the Vioolsdrif Suite. Plants

only occur on the southwest aspect in full sun.

O. decus-montium falls into subgenus Aspasia group

Aspasiae by possessing narrowly boat-shaped bracts

with a smooth margin and as long as the pedicels, broad

perianth segments with no dark rib and the style shorter

than the ovary. It is synanthous with narrow leaves. The

new species shows some similarities with O. diphyllum

Baker but O. diphyllum has up to 3 leaves with inrolled

margins and an inflorescence which usually has more

Bowers (up to 10), the perianth segments are more round-

ed-ovate and the stamens are usually longer (5 mm).

Furthermore O. decus-montium has the unique feature of

a luminous greenish black ovary apex. O. diphyllum is

restricted to the Drakensberg range in the summer rain-

fall region of southern Africa, whereas the new species is

endemic to the arid mountains of the northern

Richtersveld just south of the Orange River. This area

receives about 25-50 mm winter precipitation per year.

Rainfall figures were obtained from the nearby Rosh

Pinah Mine in Namibia and from the recently established

weather station at Sendelingsdrift. The lower Orange

River valley and adjacent mountains are bathed by sea

fogs (mainly in winter) moving up the river in a north-

easterly direction. The fogs tend to dissipate in the region

of Lorelei where the river gorges wind towards the south-

east. The new species occurs in the area where the fogs

almost cease to have any marked effect.

Flowering time: in abundance in September.

Etymology, the specific epithet describes the scattered

decorative carpets of flowering plants which cover the

lower mountain slopes.

Distribution-. The species is known only from the type

locality and a second dense colony some 10 km to the

west of this (Figure 26).

Habitat: open, exposed, undisturbed, arid mountain

slopes at about 200 m altitude, in soils derived from sur-

O. decus-montium shares some characters with O.

multifolium Baker which also occurs in the Richtersveld,

viz. narrow leaves, long flower bracts, broad perianth

segments and the style shorter than the ovary. However

O. decus-montium occurs in a completely different habi-

tat niche, is synanthous not hysteranthous, always pro-

duces only two leaves ( O . multifolium has up to 6 leaves)

and the flowers are white as opposed to yellow.

The new taxon also bears a superficial resemblance to

O. geniculatum Oberm. but is immediately separated

from this species by the sharp-angled peduncle where it

FIGURE 26. — Known distribution of

Ornithogalum decus-motium.

Bothalia 28,1 (1998)

65

emerges from the uppermost leaf. Furthermore O. genic-

ulatum is found in the Richtersveld only under moist, fog

dependent conditions in shady places.

Plants associated with O. decus-montium include sev-

eral species which occur mainly in the arid eastern lower

Orange River floristic zone, viz. Tylecodon hallii ,

Schwantesia herrei, Conophytum loeschianum, Stoeberia

carpii and Astridia hallii. Arid associates which have dis-

tributions beyond the lower Orange River valley also

occur nearby: Pachypodium namaquanum, Euphorbia

virosa, Sarcocaulon flavescens, Hoodia gordonii, H.

alstonii, Crassula garibina , the asclepiad Pentarrhinum

abyssinicum subsp. angolense and the trees Maerua

schinzii and M. gilgii.

ACKNOWLEDGEMENTS

The Chief Directorate, Nature and Environmental

Conservation of the Northern Cape is acknowledged for

providing permits to collect and research the flora of

northern Namaqualand. The wardens of the Richtersveld

National Park and the National Parks Board are thanked

for allowing access and giving permission to work within

the Park confines. The curators and staff of the Compton

Herbarium, Kirstenbosch and Bolus Herbarium, Uni-

versity of Cape Town allowed access to research their

extensive collections of Ornithogalum species. Dr D.

Paterson-Jones has given careful advice and corrected the

manuscript. The contribution from the referees is grate-

fully acknowledged. Francoise Williamson, co-collector,

typed and corrected the manuscript.

REFERENCES

OBERMEYER, A. A. 1978. Ornithogalum: a revision of the southern

African species. Bothalia 12: 323-376.

MULLER-DOBLIES. U. & MULLER-DOBLIES. D. 1996. Revi-

sionula incompleta Ornithogalorum austro-africanorum

(Hyacinthaceae). Feddes Repertorium 107,5-6: 361-548.

G. WILLIAMSON*

* Bolus Herbarium, University of Cape Town, 7700 Rondebosch.

MS received: 1997-02-05.

OXALIDACEAE

TAXONOMIC DELIMITATION OF OXALIS ENGLERIANA

In the most recent revision of Oxalis L. in southern

Africa, Salter (1944) regards O. engleriana Schltr. and

O. henrici F.Bolus as two separate species in the section

Angustatae subsection Multifoliatae. He does, however,

mention that O. henrici was only known to him from the

type specimen [Bolus 13079 , Worcester Division; near

De Dooms. (BOL)] and comments on its close morpho-

logical affinity with O. engleriana. The type specimen of

O. henrici appears to be starved and the leaves are under-

developed. Based on field observations, Bayer (pers.

comm.) regards O. engleriana and O. henrici as conspe-

cific. No constant interspecific morphological evidence

could be found to justify their separate taxonomic status.

In a comprehensive palynological review of the

southern African members of Oxalis , Dreyer (1996)

found the pollen of these two species to be identical, but

quite different to that of the rest of the genus. Both have

reticulate grains with large, open luminae, prominent

intraluminary bacules and very distinct supratectal spin-

ules. Elsewhere in the genus, such unique deviations

from the basic reticulate pattern always occur in mono-

typic groups. Although palynological evidence alone is

mostly not enough to substantiate taxonomic delimita-

tions, we believe that in this instance it provides conclu-

sive support to place O. henrici in synonymy under the

older name O. engleriana.

REFERENCES

DREYER, L.L. 1996. A palynological review of Oxalis (Oxalidaceae)

in southern Africa. Ph D. thesis. University of Pretoria,

Pretoria, South Africa.

SALTER, T.S. 1944. The genus Oxalis in South Africa. Journal of

South African Botany Suppl. Vol. 1: 1-355.

L.L. DREYER* & A.E. VAN WYK**

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

address: Department of Botany, University of Cape Town, Private Bag,

7700 Rondebosch, Cape Town.

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

of Pretoria, 0002 Pretoria.

MS. received: 1997-07-31.

'

Bothalia 28,1: 67-82 (1998)

New combinations in Antimima (Ruschioideae, Aizoaceae) from

southern Africa

H.E.K. HARTMANN*

Keywords: Aizoaceae, Antimima , new combinations, Ruschia , Ruschioideae, southern Africa

ABSTRACT

As a consequence of the emended description of Antimima N.E.Br. by Dehn (1989) and based on extensive comparison

of all types of the genera Antimima and Ruschia Schwantes, and on studies of about 250 fresh collections of Antimima , 95

species are tranferred to that genus from Ruschia. The arguments for this step, and brief diagnoses of the 100 known species

of Antimima are given. One species is excluded.

INTRODUCTION

Brown (1930) described the genus Antimima with a

single species, A. dualis (N.E.Br.) N.E.Br. — this species

was first named Mesembryanthemum duale N.E.Br.

(Brown 1920) and was transferred to the genus

Argyroderma N.E.Br. (Brown 1922), based on similari-

ties of leaf morphology and the light greyish white

colouring of the leaves. When, after 18 years of cultiva-

tion of the plant, Brown saw a flower and realized that

the species was not a member of the genus Argyroderma

since it lacked the hypanthium characteristic of that

genus (Figure 1 A, B).

The species had meanwhile also been described as

Ruschia dualis L. Bolus (1929), but based on a different

type. This name has been used for about 60 years, imply-

ing that the genus Antimima was a synonym of Ruschia

Schwantes.

In the course of his studies in the Ruschiinae, Dehn

(1989) found that not only Antimima dualis but also ±

100 species formerly in the genus Ruschia differed so

markedly from typical forms of the latter genus, that he

re-established the genus Antimima. He distinguished five

subgenera and made four new combinations: A. albo-

rubra (L.Bolus) Dehn, A. longipes (L. Bolus) Dehn, A.

microphylla (Haw.) Dehn, and A. virgata (Haw.) Dehn.

Later, Antimima hantamensis (Engl.) H.E.K. Hartmann

& Stiiber (1993) and A. aurasensis H.E.K. Hartmann

(1996) were added.

In a comprehensive survey covering all species of the

Aizoaceae, material of types of all species in Ruschia

and fresh material of ± 250 collections of Antimima was

examined. As a result, 95 species are here transferred to

Antimima. With A. virgata transferred back to Ruschia ,

based on comparisons of fruits and the type, Antimima at

present comprises 100 species.

The subdivision of the genus into subgenera will have

to be dealt with on a much broader basis. M. Dehn (pers.

comm. 1992) noticed that the circumscriptions of the

* Institut fiir Allgemeine Botanik, Ohnhorststr. 1 8, D-22609 Hamburg,

Germany.

MS. received: 1997-09-03.

subgenera need reconsideration, in the light of data

which must be derived from extensive studies in flowers

and leaves, especially heterophylly and Alicean mor-

phology of the epidermis. Features of flowers and leaf

anatomy are therefore given sporadically in this treat-

ment, due to lack of data for complete comparisons.

CRITERIA FOR INCLUSION OF SPECIES IN ANTIMIMA

Fruit

As demonstrated by Dehn (1989: 199), typical fruits

of Antimima possess very large closing bodies filling and

blocking the distal exit of the locule completely, i.e. no

seeds can be expelled through this opening (Figure 1 C,

F-H). In contrast, fruits of Ruschia have small, hook-

shaped closing bodies which never form a complete

blockage or barrier for the seeds.

In addition, expanding keels in fruits of Antimima are

broad and laciniate, diverging distinctly in a ± radial

direction and reaching almost to the tip of the valve

(Figure 1C, H). In Ruschia, the expanding keels are typ-

ically shorter and are spread into an almost tangential

direction, visible in the open capsule on the valve. Due

to these structural differences, fruits of Antimima nor-

mally open completely (Figure IF), the valves often

even recurving below the horizontal plane, whereas cap-

sules of most species of Ruschia open their valves into

an erect position only.

Covering membranes possess additional closing

devices at their distal ends in both genera in principle. In

fruits of Ruschia these devices appear mostly in the

shape of closing rodlets, rarely as closing ledges or clos-

ing bulges. Closing rodlets can also occur in Antimima,

but in this genus closing ledges (Figure 1C, F) or the lack

of any additional closing devices are more common

(Figure 1G, H). The last-mentioned condition is absent

in Ruschia.

In seven species of Antimima , the covering mem-

branes possess radial wings (Figure 1G, H) or other pro-

trusions on top, clearly visible in younger fruits. Similar

features have been found in genera like Cheiridopsis

N.E.Br., but never in typical fruits of Ruschia.

68

Bothalia 28,1 (1998)

FIGURE 1. — A-D, Antimima dualis: A, l.s. through flower showing flat ovary surface and free stigmata, in contrast to

Argyroderma: B, flowering branch with persisting old leaves below; C, part of open capsule with big closing

bodies (CB), no valve wings, expanding keels (EK) diverging moderately and covering membranes (CM), with

distinct closing ledge (CL) below near opening; D, side view of fruiting branch of herbarium material. A, B, figs

of Brown 1930; C, Hartmann & Ihlenfeldt 4157 (HBG); D, Peers NBG 370/33 (BOL!), holotype of A. villetii.

E, A. pumila , Hartmann 30325 (HBG!), side view of fruiting heterophyllous branch: long connate sheath is

papery and smooth, short erect free parts papillate. F, A. dolomitica, Dinter 3782 (BOL, iso.!), radial l.s. through

capsule in middle of locule: straight covering membrane (CM) lying distally on big closing body (CB), which

is also touched by closing ledge (CL). G, A. fenestrata, Hartmann & Dehn 15626 (HBG!), radial l.s. through

capsule in middle of locule: covering membranes sloping down from a high centre ending in a distal recurving

pressed against big white closing body (CB); erect wing in nearly radial position stabilizing covering membrane.

H, /! eendornensis, Dinter 5207 (B, holo!), part of open capsule with basally distant moderately diverging

expanding keels (EK), small valve wings (VW) broadest in middle, very big closing bodies (CB) and covering

membranes (CM) pressed against them; covering membranes with radial wings on top lowering towards centre.

I, A. distans, Hartmann et al. 20682: heterophyllous long shoot with several leaf pairs with short connate sheaths

and free long parts (FLP) and connate subterminal leaf pair (CLP) forming long sheath with short free parts

enclosing terminal leaf pair. ES, expanding sheet; P, placenta; V, valve; VR, valve rim; W, wing on top of cover-

ing membrane.

Bothalia 28,1 (1998)

69

Fruits of the majority of species of Ruschia lack valve

wings altogether, with a few exceptions such as Ruschia

subgenus Spinosae. Capsules of Antimima are also most-

ly devoid of valve wings (Figure 1C), but in a number of

species narrow, awn-shaped or even broader valve wings

are found, always broadest in the middle and tapering

towards both ends (Figure 1H).

Finally, fruits of Ruschia develop a long, deep fruit base

whereas those of Antimima have rather shallow, mostly

bowl- or funnel-shaped capsule bases (Figure ID, E).

Locule numbers are mostly five in both genera, six

occurring in several species in Ruschia and in Antimima.

Leaves

Isophylly (i.e. all leaves of a plant are ± equal), is pre-

sent in all species of Ruschia. In contrast, the species of

Antimima develop isophylly (Figure IB, D) or hetero-

phylly on a plant, the latter either as a result of two leaf

pairs (Figure IE) of different leaf shapes developing in

one season, or as a result of differing leaf shapes devel-

oping in sequence along a side branch (Figure II). The

first condition resembles that in Cheiridopsis subg.

Cheiridopsis (Hartmann & Dehn 1987) or Mitrophyllum

(Poppendieck 1976). Leaves developing in sequence

along a side branch resemble those of Vanzijlia L. Bolus

(Hartmann 1983). In both cases, the long-sheathed leaf

pair forms a papery, persisting sleeve-like protective

cover enveloping the subsequent, assimilating leaf pair

during the dry' season. The dry. persistent sheath consists

mostly of the connate lower parts of the leaves of a pair

(Figure IE, I), but in a few cases the leaves are separated

to the base but stand closely together forming a protective

sleeve-like cover. In these cases, the broad lower parts

resemble sheaths that have been slit open, but they were

never connate, and the upper parts resemble the free parts

of sheathed leaves closely. In general, the epidermis of

the sheaths is smoother than that of the free tips, and often

the different leaf types differ in their epidermal structures.

Flowers

Flowers of both genera are rather small in the majori-

ty of species, rarely exceeding 20 mm in diameter

(Figure 1A, B). The petals are white to pink or purple

and often arranged in five, rarely six, distinctive groups.

Filamentous staminodes are present, surrounding the

central cone of stamens. In many cases, the tips elongate

and recurve, and are often dark purple to almost black in

colour. Characters of the flowers do not differ in princi-

ple between Ruschia and Antimima , neither do inflores-

cences, which can be solitary or in cymes of various

sizes, typical of species but not of either genus.

Growth forms

After the removal of a number of compact species

from Ruschia by Glen (1986: Ebracteola Schwantes,

Acrodon N.E.Br.) and the re-establishment of Mar-

lothistella Schwantes (Hammer 1995), Ruschia compris-

es almost exclusively, shrubby to creeping growth forms

with long, visible internodes and branching at the distal

ends of the stems.

In Antimima, growth forms are variable: compact to

caespitose, rarely creeping, or with a compact centre

with long shoots climbing into other shrubs. All shrubs

with a marked distal branching, proposed for inclusion in

Antimima (by Dehn on herbarium sheets), had to be

excluded from that genus because of their fruit morphol-

ogy, which resembles that of Ruschia.

CORRELATIONS BETWEEN CHARACTER STATES

The possession of the typical fruit permits a definite

inclusion of a given plant in Antimima, but if fruits on the

type sheet are absent or incomplete, the knowledge of the

correlation of characters can also help to establish the

identity of the material.

In all heterophyllous species with capsules, the

expanding keels are basally separated, broad and lacini-

ate, and diverging towards the tips of the valves, not in a

tangential position as found in Ruschia. Valve wings are

absent or present, mostly forming flanges tapering

towards the tips. The closing bodies are mostly large, but

in some species small white ones are present. In one

species no closing body is developed but the capsules

show so many similarities with those of other species

that the taxon is included here in Antimima. Furthermore,

it is deduced that heterophylly is correlated with the

Antimima type of capsule morphology, and as a conse-

quence, all heterophyllous species formerly belonging to

the genus Ruschia are transferred to Antimima.

It should be noted that a revision of Ruschia down to

species level is not yet available, the circumscription of

that genus might therefore require some adjustment at a

later stage. Nevertheless, based on the criteria given

above, 95 species are transferred here from Ruschia to

Antimima.

TAXONOMY

Antimima N.E.Br., in The Gardeners’ Chronicle ser.

in, 87: 211 (1930); Dehn: 189 (1989); Hartmann &

Stiiber: 1 (1993); Hartmann: 229 (1996). Type species:

A. dualis N.E.Br.

Plants compact, caespitose or with compact centre

and long shoots, isophyllous or heterophyllous. Leaves :

cymbiform with convex sides to nearly finger-shaped,

often keeled in upper part; or mostly oval and very often

connate for at least part of the length, leaf pairs differing

in this feature, mucronate; epidermis cells flat to papil-

late, wax cover smooth or papillae, with a smooth later-

al channel and wax flakes or platelets on rest of surface.

Flowers: 1 (—3), rarely in well-developed cymes; bracts

and bracteoles present; petals pink to purple, rarely

white, often arranged in 5 (rarely 6) groups. Filamentous

staminodes: mostly in a central cone surrounding sta-

mens closely. Capsules: 5, rarely 6 locules, stalked,

always with persisting bracteoles, top with high or low

valve rims, base mostly bowl-shaped, sometimes funnel-

shaped; covering membranes undulate, more rarely

straight, mostly elevated in centre, in several species

with radial or tangential wings or other protrusions on

them; closing body large, rarely small, absent in one

species; expanding keels broad, diverging and radial in

distal part, reaching to tip of valve; valves with or with-

out mostly narrow valve wings.

70

Bothalia 28,1 (1998)

DISTRIBUTION

From Liideritz and Warmbad, Namibia; southwards to

the Northern Cape, Free State, Western Cape, and

Eastern Cape, South Africa.

The following species are included in Antimina as cir-

cumscribed above, most of them transferred from the

genus Ruschia.

1. Antimima addita (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia addita L Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 284 (1931). Type: M. Schlechter 8427 (BOL, holo.!).

Plants compact with long shoots, isophyllous. Leaves :

terete, mucronate. Capsules: typical in all respects.

Typical of species: rich cymes.

2. Antimima alborubra (L.Bolus) Dehn in Mit-

teilungen aus dem Institut fiir Allgemeine Botanik

Hamburg 22: 212 (1989). Type: Herre 8861 (BOL, holo.!).

Ruschia alborubra L.Bolus: 494 (1934).

Plants with compact centre from which ± 5 long

shoots rise, isophyllous. Leaves: trigonous to cymbiform.

Capsules: typical in all respects. Typical of species: very

big basal leaves and rather small flowers, smaller than

calyx.

3. Antimima amoena (Schwantes) H.E.K. Hart-

mann, comb. nov.

Ruschia amoena Schwantes in Monatsschrift der Deutschen

Kakteengesellschaft 2: 66 (1930). Type: Anon, s.n., photograph of flow-

ering ‘type plant’ in collection Schwantes (HBG, lecto.!, here designat-

ed).

Plants compact to caespitose, isophyllous. Leaves:

trigonous, aristate. Capsules: typical in all respects.

Typical of species: distinct darker lines along keel and

margins.

4. Antimima androsacea (Marloth & Schwantes )

H.E.K. Hartmann, comb. nov.

Ruschia androsacea Marloth & Schwantes in Zeitschrift fiir

Sukkulentenkunde 3: 299 (1928). Mesembryanthemum androsaceum

Marloth: 19 (1927) nom. nud. Ruschia androsacea (Marloth)

Schwantes: 19 (1927) nom. nud. Mesembryanthemum androsaceum

(Schwantes) N.E.Br.: 32 (1930). Type: Marloth 9682 (HBG, holo.!).

Plants forming flat mats, isophyllous. Leaves : form-

ing white bodies in the resting state. Capsule: top almost

flat, base short, funnel-shaped, closing body round,

white, rather small for genus; covering membranes con-

vex with a low distal rim, a closing ledge below it, bases

of expanding keels just not touching each other at their

bases, moderately diverging and tapering into short

awns. Typical of species: leaves of a pair form small,

brightly white bodies in the dry state.

5. Antimima argentea (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia argentea L.Bolus in Journal of South African Botany 27: 52

(1960). Type: Hall 1883 (BOL, holo.!).

Plants compact, isophyllous. Leaves: triquetrous with

pointed tip, grey from papillae. Capsules: typical of

genus. Typical of species: awl-shaped silvery to grey

leaves.

6. Antimima aurasensis H.E.K. Hartmann in

Kakteen und andere Sukkulenten 47: 231 (1996). Type:

Hartmann et al. 20865 (HBG, holo.!).

Plants caespitose, elongate branches formed when not

eaten, isophyllous. Leaves: triquetrous. Capsules: typical

of genus. Similar to A. perforata in growth and leaves,

but capsules with 5 locules only.

7. Antimima biformis (N.E.Br. ) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum biforme N.E.Br. in Kew Bulletin 1929: 58

(1929). Ruschia biformis (N.E.Br.) Schwantes: 230 (1938). Type: Muir

s.n. (K, holo.).

Plants caespitose, heterophyllous. Leaves: sheath of

one leaf pair forming a body with two very short lobes,

other leaf pair almost free, conspicuously dotted.

Capsules: with expanding keels diverging and transgrad-

ing into expanding sheets, closing body knob-shaped.

Typical of species: dotted, connate leaves of one pair.

8. Antimima bina (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia bina L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 282 (1931). Type: Pillans BOL19610 (BOL, holo.!).

Spreading shrubs to 150 mm high, internodes reddish,

heterophyllous. Leaves: sheath of one leaf pair oval,

white, keel of each leaf running down sheath as a narrow

wing, only tips papillate; second leaf pair basally connate

for only ± 2 mm, densely papillate, leaves free on long

shoots, oval sheaths always at base of short shoots,

where both types of leaf pairs are formed alternatively.

Capsules: with inconspicuous closing ledges, closing

body medium-sized. Typical of species: leaves.

9. Antimima bracteata (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia bracteata L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 93 (1929). Type: Erni BOL18921 (sheet I) (BOL,

lecto.!, here designated).

Plants are loose cushions, isophyllous. Leaves: trigo-

nous, smooth. Capsules: typical of genus; covering mem-

branes with remains of stiff wings in radial position.

Typical of species: radial wings on covering membranes.

Similar to A. compacta.

10. Antimima brevicarpa (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia brevicarpa L.Bolus in Notes on Mesembryanthemum and

allied genera, part 3: 279 (1954). Type: Esterhuysen 20444 (BOL, holo.!).

Bothalia 28,1 (1998)

71

Plants creeping, later forming a mat, heterophyllous.

Leaves', sheath of one leaf pair long, yellowish, rather

smooth, enclosing larger part of subsequent leaf pair

with a shorter connate part and longer free parts with

papillae, in particular along margins and keel. Capsules:

with closing bodies broader than thick, almost bean-

shaped, covering membranes ± straight with a low rim

and closing ledges. Typical of species: creeping habit.

Similar to A. prostrata , but larger-leaved.

11. Antimima brevicollis (N.E.Br.) H.E.K. Hart-

mann, comb. nov.

Meseinbryanthemum brevicolle N.E.Br. in Kew Bulletin 1929: 58

(1929). Ruschia brevicollis (N.E.Br.) Schwantes: 230 (1938). Type:

Muir 4064 (K, holo.).

Plants caespitose, heterophyllous. Leaves: sheath of

one leaf pair connate for half its length, other pair with a

short connate part. Capsule: not known.

12. Antimima buchubergensis (Dinter) H.E.K.

Hartmann , comb. nov.

Ruschia buchubergensis Dinter in Monatsschrift der Deutschen

Kakteengesellschaft 2: 219 (1930). Type: Dinter 6477A (HBG. lecto. !,

here designated: the three sheets of Dinter 6477 at HBG have been

marked A, B, and C: A is chosen as the lectotype because it shows the

typical caespitose growth and possesses fruits).

Plants compact to caespitose, isophyllous. Leaves: leaf

pairs on lateral branches as pea-shaped bodies. Capsules:

typical of genus, with very narrow valve wings. Typical

of species: pea-shaped leaf pairs on proleptic shoots.

13. Antimima compacta (L. Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia compacta L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 144 (1929). Type: L.Bolus BOL18977 (BOL,

holo.!).

Plants compact, isophyllous. Leaves: trigonous with

recurved tip. Capsules: typical of genus; covering mem-

branes with stiff wings in radial position and tiny valve

wings. Very similar to A. bracteata.

14. Antimima compressa (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia compressa L.Bolus in Journal of South African Botany 27:

54 (1960). Type: Hall NBG 1056/48 (BOL, holo.!).

Plants caespitose with some long shoots, isophyllous.

Leaves: trigonous, papillate. Capsules: typical of genus,

valve wings very narrow. Typical of species: protruding,

diagonal long shoots.

15. Antimima concinna ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia concinna L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 123 (1929). Mesembryanthemum comptum

N.E.Br.: 32 (1930). Type: Pillans NBG370/16 (BOL, lecto.!, here des-

ignated); Mathews 1802/21 (BOL, paralecto!).

Plants compact, heterophyllous. Leaves: persisting

yellow, papery sheath of one leaf pair around stem, free

parts triquetrous, other leaf pair connate for 2-3 mm

only, ending in recurved long awn; keels of leaves with

distinct, large, long papillae. Capsules: typical of genus.

16. Antimima condensa ( N.E.Br. ) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum condensum N.E.Br. in Kew Bulletin 1929: 58

(1929). Ruschia condensa (N.E.Br.) Schwantes: 230 (1938). Type:

Muir 4065 (K, holo.!).

Plants caespitose, heterophyllous. Leaves: sheath of

one leaf pair as a connate body with only short free tips

during the resting period, whitish yellow, smooth, with a

protruding roll along line where both leaves are fused in

dry state; second leaf pair hardly connate, with elevations

above subhypodermal idioblasts and a papillate epider-

mis. Capsules: typical of genus, valve wings narrow.

17. Antimima crassifolia (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia crassifolia L Bolus in Notes on Mesembryanthemum and

allied genera, part 3: 338 (1958). Type: Hall BOL25758 (BOL, holo.!).

Low shrubs with some erect flowering branches, iso-

phyllous. Leaves: trigonous, thick. Capsules: typical of

genus, 6-loculed.

18. Antimima dasyphylla (Schltr.) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum dasyphyllum Schltr. in Botanische Jahrbiicher

27: 127 (1899). Ruschia dasyphylla (Schltr.) Schwantes: 20 (1927).

Type: Schlechter 8773 (B, lecto.!, here designated: one of the two

sheets kept at B bears the note ‘holotype’ by Glen; it is here chosen as

the lectotype).

Ruschia concava L.Bolus: 124 (1929). Mesembryanthemum ciliola-

tum N.E.Br.: 32 (1930). Type: Levyns 1375/26 (BOL, holo.!).

Shrubs, heterophyllous. Leaves: sheath of one leaf

pair long, orange-brown, free parts only 2-3 mm long,

papillate; other leaf pair connate for 4 mm, free parts

spreading, longer papillae on keel and margins. Cap-

sules: typical of genus but expanding keels transgrading

into expanding sheets and with distal awns. Typical of

species: inconspicuous expanding keels.

19. Antimima defecta (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia defecta L.Bolus in Journal of South African Botany 31: 85

(1965). Type: Van Heerde BOL27654 (BOL, holo.!).

Ruschia deflecta L.Bolus ex Jacobsen: 49 1 (1970) orthographic variant.

Plants cushion-forming, heterophyllous. Leaves: yel-

low sheaths of one leaf pair on short shoots connate for ±

half their length, those on long shoots only connate for ±

Vs of their length, free parts of both types with distant, big

papillae. Capsules: typical of genus, but closing bodies

hood-shaped and broader than thick. A deviating single

fruit on the type sheet with broad valve wings and a

longer base belongs to Amphibolia, but not to the remain-

ing type material, which agrees with the description.

72

Bothalia 28,1 (1998)

20. Antimima dekenahi (N.E.Br.) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum dekenahi N.E.Br. in Kew Bulletin 1929: 58

(1929) Ruschia dekenahi (N.E.Br.) Schwantes: 230 (1938). Type:

Dekenah 23 (K, lecto.!, here designated); Muir 4079 (K, paralecto.!).

Plants compact to caespitose, isophyllous. Leaves'.

trigonous, mucronate. Capsules: typical of genus. Typi-

cal of species; fruits raised above plant and the thick root.

21 . Antimima distans (L.Bolus) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum distans L.Bolus in Annals of the Bolus

Herbarium 4: 89 (1927a). Ruschia distans (L.Bolus) L.Bolus: 236

(1950). Type: Anon. BOL18550 (BOL, holo. ! ).

Plants compact with erect, long shoots, heterophyl-

lous. Leaves: sheath of one leaf pair long, almost smooth,

with a bulge running down side where both leaves meet,

and long, ± triquetrous free parts with some large, long

papillae along keel only; other leaf pair only slightly con-

nate, free parts densely papillate. Capsules: typical of

genus. Typical of species: stout, dark brown or maroon,

long internodes (Figure II).

22. Antimima dolomitica (Dinter) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum dolomiticum Dinter in Feddes Repertorium 19:

150 (1923). Corpuscularia dolomitica (Dinter) Schwantes: 186 (1926).

Ruschia dolomitica Dinter& Schwantes: 69 (1929). Type: Dinter 3782,

Klinghardtberge (B, lecto.!, here designated); from Alicetal, Buntfeld-

schuhplateau (B, BOL, paralecto.!).

Plants with compact base with erect, stiff long shoots

bearing lateral proleptic bodies consisting of a hazelnut-

shaped leaf pair each; ± isophyllous. Capsules: typical of

genus, closing bodies only medium-sized, as seen from

above (Figure IF). Typical of species: hazelnut-shaped

leaf pairs.

23. Antimima dualis (N.E.Br.) N.E.Br., Garden-

ers’ Chronicle, ser. Ill, 87: 211 (1930). Type: Pearson &

Pillans 5483 (K, holo.!).

Mesembryanthemum duale N.E.Br.: 89 (1920). Argyroderma duale

(N.E.Br.) N.E.Br.: 105 (1922).

Ruschia dualis L.Bolus: 104 (1929). Type: Hutchinson s.n. (BOL,

lecto.!, here designated from three syntypes cited and placed on one

sheet).

Ruschia villetii L.Bolus: 50 1 ( 1 934) Type: Peers 1370/33 (BOL, holo. !).

Plants compact, highly branched, isophyllous.

Leaves: trigonous but keeled, sides convex to straight,

whitish grey. Capsules: typical of genus; covering mem-

branes almost straight. The species can easily be con-

fused with A. turneriana. Characteristic of A. dualis are

the strictly compact growth, retained also in cultivation,

and the strictly solitary flowers (Figure 1A-D).

24. Antimima eendornensis (Dinter) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum eendornense Dinter in Feddes Repertorium

Beiheft 53: 86 (1928). Type: Dinter 5207 ( B, holo.!).

Semi-compact dwarf shrublets, isophyllous. Leaves:

triquetrous with long papillae. Capsules: typical of genus

with narrow valve wings at the point where they touch

edge of valve and with oblique wings on top of covering

membranes, 6-loculed (Figure 1H).

Dinter (1928: 86, 87) did not give a formal descrip-

tion, only a short diagnosis in contrast to Mesem-

bryanthemum hospitale (now a member of Ruschia sub-

genus Spinosae). The type sheet at B had been wrongly

placed with A. quarzitica up to now.

25. Antimima elevata (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia elevata L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 305 (1931). Type: Luckhoff BOL19873 (BOL, holo.!).

Plants caespitose, isophyllous. Leaves: with broadly

trigonous, rough to warty free parts, greyish. Capsules:

typical of genus; covering membranes raised high in cen-

tre and closing bodies bean-shaped and rather small.

26. Antimima emarcescens (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia emarcescens L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 281 (1954). Type: Acocks 16995 (BOL, holo.!). Ruschia

emarcidens L.Bolus ex Jacobsen: 492 (1970) orthographic variant.

Plants with compact centre with long shoots, hetero-

phyllous; roots thickened and forming elongate, potato-

shaped tubers. Leaves: one leaf pair with dry, papery

sleeve-like covers longer than free parts, but not connate;

other leaf pair with sheaths shorter than free parts, smooth,

papery during dry season, free parts triquetrous, with low

papillae, turning dark when drying, both types 7-10 mm

long. Capsules: typical of genus with high valve rims.

27. Antimima erosa (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia erosa L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 279 (1954). Type: Acocks 17126 (BOL, holo.!).

Plants densely shrubby, isophyllous. Leaves: terete,

apically rounded. Capsules: typical of genus; covering

membranes straight; valve wings very narrow. Resembles

species of Rhinephyllum in growth form, both occur in

the same area, so that they can easily be confused.

28. Antimima evoluta (N.E.Br.) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum evolution N.E.Br. in Kew Bulletin 1913: 120

(1913). Ruschia evoluta (N.E.Br.) L.Bolus: 178 (1928a). Type:

Pearson 5946 (K, holo.!).

Plants compact, white in the resting state, heterophyl-

lous. Leaves: sheath of one leaf pair forming white body

in dry season, smooth, keel markedly smooth, margins

ciliate, completely embracing subsequent leaf pair with

longer free parts with short papillae on them. Capsules:

with closing body pointing into locule like a finger;

expanding keels very similar to the short ones of A.

dualis.

Bothalia 28,1 (1998)

73

29. Antimima exsurgens (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia exsurgens L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 280 (1931). Type: L.Bolus BOL19334 (BOL, holo.!).

Plants with compact centre and long shoots, each

internode with papery whitish apical rim, appearing

corky below and drying into rings, stems therefore

resembling cones put one into the other, heterophyllous.

Leaves : sheath of one leaf pair turning papery for most of

its length (± 6 mm) in the resting state, free apical part of

± 2 mm with very low papillae; second leaf pair en-

veloped during the resting period, ± 6-8 mm long, tri-

quetrous, epapillate; in both leaf types keel and margins

rounded. Capsules : typical of genus.

30. Antimima fenestrata ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia fenestrata L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 281 (1931). Type: Luckhoff BOL19612 (BOL, holo.!).

Compact shrubs forming tiny tree-like shapes, hetero-

phyllous. Leaves-, sheath of one leaf pair long, smooth,

white, embracing subsequent leaves at bases, these latter

free for ± ‘/2 their length and withering in a peculiar way:

hardened bundles along keel (and into the apical tooth)

and margins form a triangular window frame and persist

much longer than intermediate tissue so that finally tri-

angular openings are formed. Capsules : typical of genus;

highly undulate covering membranes with very broad,

horizontally arranged wings on top almost completely

hiding covering membranes when seen from above

(Figure 1G). Typical of species: windowed leaves drying

white, with frames remaining on plants.

31. Antimima fergusoniae (L.Bolus) H.E.K.Hart-

mann, comb. nov.

Ruschia fergusoniae L.Bolus in South African Gardening and Country

Life 18: 279 (1928b). Mesembryanthemum fergusoniae (L.Bolus)

N.E.Br.: 32 (1930). Type: Ferguson BOLI882 (sheet I) (BOL!).

Plants caespitose, heterophyllous. Leaves: both pairs

forming sheaths, one pair with pronounced bulges above

tannin idioblasts and low papillae; second pair with lower

bulges but longer papillae. Capsules : typical of genus.

After the first description in English in September

1928 (Bolus 1928b), Bolus published a Latin description

of the species as well in November 1928 (Bolus 1928d:

7), where she cited the type. Nevertheless, the descrip-

tion in September fulfills all requirements for the new

combination to be validly published, at that time.

32. Antimima gracillima (L.Bolus) H.E.K. Hart-

mann, comb. nov.

R. gracillima L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 203 (1930). Type: Frames BOL 19208 (BOL, holo.!).

Plants compact with decumbent long shoots, hetero-

phyllous. Leaves: sheaths of leaf pairs on short shoots

connate for ± V2 their length, those on long shoots for ±

'/4, free parts of both types with distant, big papillae, aris-

tate. Capsules: not known.

33. Antimima granitica (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum graniticum L.Bolus in Annals of the Bolus

Herbanum 4: 88 (1927a). Ruschia granitica (L.Bolus) L.Bolus: 221

(1950). Type: Pillans BOL17772 (BOL, holo.!).

Plants caespitose, isophyllous. Leaves: trigonous, apical-

ly roundish. Capsules: typical of genus; valve wings very

narrow.

34. Antimima hallii (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia hallii L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 218 (1950). Type: Hall BOL24059 (BOL, lecto.!, here

designated); Peers BOL 24060 (BOL, paralecto.!).

Plants with compact centre with long shoots, erect at

first, decumbent later, rooting at nodes some distance

from the primary centre and developing a secondary com-

pact centre, heterophyllous. Leaves: sheaths of both types

of leaves papery, white, disintegrating early; bases of sec-

ond leaf pair almost separated by a triangle of stem tissue,

with low papillae, old leaves closely set on short shoots.

Capsules: typical of genus. Typical of species: formation

of secondary compact centres with adventitious roots.

35. Antimima hamatilis (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia hamatilis L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 106 (1929). Mesembryanthemum hamatile

(L.Bolus) N.E.Br.: 32 (1930). Type: Pillans 6080 (BOL, holo.!).

Plants with decumbent branches, heterophyllous.

Leaves: sheath of one leaf pair connate for ± V2 its length,

free parts papillate, mucro to 1 mm long; second leaf pair

connate for '/6 of its length, free parts papillate, longer

papillae along keel and margins, these leaves forming

repeatedly on fresh long shoots. Capsules: unknown.

36. Antimima hantamensis ( Engle r) H.E.K. Hart-

mann & Stiiber, Contributions from the Bolus Herbarium

15: 68 (1993). Mesembryanthemum hantamense Engl.:

190 (1909) Type: Meyer s.n. (B, holo.!).

Ruschia disarticulata L.Bolus: 286 (1931). Eberlanzia disarticulata

(L.Bolus) L.Bolus: 387 (1958) Type: L.Bolus 19335 (BOL, holo.!).

Ruschia stellata L.Bolus: 140 (1927/1928). Mesembryanthemum

stellans (L.Bolus) N.E.Br.: 32 (1930). Eberlanzia stellata L.Bolus

nom. nud. Type: Compton 1414/27 (BOL, holo.!).

Flat shrubs smelling of fish when damaged, covered

in spines derived from big dichasial inflorescences, iso-

phyllous. Leaves: trigonous. Capsules: with high valve

rims extending over edge of capsule, otherwise with typ-

ical features. Plants always densely covered with spines,

making it easy to recognise them.

37. Antimima herrei (Schwantes) H.E.K. Hart-

mann, comb. nov.

74

Bothalia 28,1 (1998)

Ruschia herrei Schwantes in Zeitschrift fur Sukkulentenkunde 3: 301

(1928a). Mesembryanthemum herrei (Schwantes) N.E.Br.: 32 (1930).

Type: Karrooport Ceres, H. Herre in Bat. Gard. Stellenbosch 1876. As

this could not be traced, the following lectotype is designated: Anon. s.n.

(HBG!, photograph of ‘my type plant’ in collection Schwantes HBG).

Plants compact, heterophyllous. Leaves : sheath of one

leaf pair papery for most of its length; other leaf pair free

(flowering plant on photograph showing free leaves

only), as is the case during the growing and flowering

period. Capsules : typical of genus; covering membranes

with low protrusions in radial direction near centre of

fruit. Placed by Schwantes (1928a: 302) in a group called

‘ Ruschiella ’, not described formally, together with

ventricosa’ and 'R. schlechteri' , both also heterophyl-

lous.

38. Antimima intervallaris (L. Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia intervallaris L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 91 (1929). Type: Pillans 6056 (BOL, holo.!).

Shrubs with decumbent branches, heterophyllous.

Leaves : first leaf pair with long, loose sheaths enclosing

subsequent leaves; other leaf pair hardly connate, both

glabrous, green; epidermis papillate, epidermis cells with

lower papillae on smaller leaves. Capsules : typical of

genus; valve wings or very narrow bands appearing only

when a fresh fruit is opening. In the leaf sequence, the

species resembles that of Vanzijlia annulata. The citation

of the name Ruschia intervallaris L. Bolus under Mossia

intervallaris is an error (Brown 1930; 71, 151).

39. Antimima ivori (N.E.Br.) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum ivori N.E.Br. in Kew Bulletin 1929: 58 (1929).

Ruschia ivori (N.E.Br.) Schwantes: 230 (1938). Type: Dekenah 67 (K,

lecto.l, here designated); Muir 4285 (K, paralecto.!).

Plants with flat mats, heterophyllous. Leaves : sheath

of one leaf pair smooth, white in its lower half, often suf-

fused purplish, tips disintegrating into thread-like protu-

berances, sheath enclosing subsequent leaf pair of papil-

late leaves connate only for ± 'L of its length; surface of

epidermis of outer leaf pair of elongate cells with cutic-

ular folds covered by an almost continuous wax layer.

Capsules: typical of genus, but top with low rims, 6-

loculed. Typical of species: laciniate upper parts of high-

ly connate leaf pairs persisting on plant for many years.

In this feature, and in leaf surfaces, it resembles A. pyg-

maea most closely, differing in the truncate leaf tips and

in fruit morphology from the latter.

40. Antimima karroidea (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia karroidea L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 306 (1931). Type: Archer 364 (BOL, holo.!).

Plants compact with long decumbent shoots, numer-

ous short shoots on them, heterophyllous. Leaves: sheath

of one leaf pair whitish, keel of shorter free part continu-

ing as a row of long papillae on sheath and internode

below; alternating leaf pair with a sheath only 1.5 mm

long, free parts subulate. Capsules: typical of genus;

expanding keels merging into expanding sheets. Similar

in leaf shapes and sequence to A. fergusoniae.

41. Antimima klaverensis (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum klaverensis L.Bolus in Annals of the Bolus

Herbarium 4: 96 (1927a). Ruschia klaverensis (L.Bolus) Schwantes: 58

(1949). Type: Compton NBG291/22 (BOL, holo.!).

Plants caespitose, smelling of fish, isophyllous.

Leaves: trigonous to terete. Capsules: typical of genus.

Leaves, fruit and growth form agree well with those of A.

watermeyeri, which lacks a fishy smell.

42. Antimima koekenaapensis (L.Bolus) H.E.K.

Hartmann, comb. nov.

Ruschia koekenaapensis L.Bolus in Journal of South African Botany

28: 299 (1962). Type: Hall 2398 (BOL, holo.!).

Plants compact with long shoots, these in turn with

short shoots bearing flowers appearing in bundles or

dense groups in the following year, isophyllous. Leaves:

triquetrous, pointed. Capsules: typical of genus. Typical

of species: tufted short flowering shoots.

43. Antimima komkansica (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia komkansica L.Bolus in Journal of South African Botany 28:

298 (1962). Type: Hall 2550 (BOL, holo.!).

Shrubs up to 600 mm high, isophyllous. Leaves: trigo-

nous. Capsules: typical of genus; very narrow valve wings

and low radial wings on covering membranes. Plants very

tall for genus.

44. Antimima lawsonii (L.Bolus) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum lawsonii L.Bolus in Annals of the Bolus

Herbarium 4: 85 (1927a). Ruschia lawsonii (L.Bolus) L.Bolus: 219

(1950). Type: Lawson 18551 (BOL, holo.!).

Plants compact, isophyllous. Leaves: triangular to tri-

quetrous, light grey, hard. Capsules: most similar to

those of A. dualis, but closing body smaller and almost

hook-shaped. Grows far from most other species of

genus, settling in distinct, stony habitats on limestone

and seemingly unable to explore adjacent areas with a

sand cover where grass and trees predominate.

45. Antimima leipoldtii (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia leipoldtii L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 61 (1929). Mesembryanthemum leipoldtii (L.Bolus)

N.E.Br.: 32 (1930). Type: Leipoldt BOL18905 (BOL, holo.!).

Low shrubs, heterophyllous. Leaves: sheath of one

leaf pair long, smooth, long free parts with some big

papillae along keel only; other leaf pair short, with long

papillae. Capsules: typical of genus; valves with distinct

narrow wings broadest at middle.

Bothalia 28,1 (1998)

75

46. Antimima leucanthera (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum leucantherum L.Bolus in Annals of the Bolus

Herbarium 4: 7 (1925). Ruschia leucanthera (L.Bolus) L.Bolus: 239

(1927b). Type: Tugwell BOL17108 (BOL, lecto.!, here designated);

Russell NBG8/24 (BOL, paralecto.!).

Plants compact to caespitose, isophyllous, hypocotyl

and roots thickened. Leaves : trigonous, mucronate. Cap-

sules: typical of genus, interior falling off persistent old

outer bundles of capsule, leaving a prickly star on plant

where a fruit has been; in young fruits narrow valve wings

in angle where expanding keel meets edge of valve. In

habitat, plants adorned by whitish capsules for most of

year. Similar to A. dekenahi, but differing in wings on cover-

ing membranes and longer papillae along keel and mar-

gins of leaves.

47. Antimima limbata (N.E.Br.) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum limbatum N.E.Br. in Kew Bulletin 1929: 59

(1929). Ruschia limbata (N.E.Br.) Schwantes: 230 (1938). Type:

Marloth 13173 (K, holo!).

Plants creeping, isophyllous. Leaves', trigonous with

convex sides, keel and margin reddish brown. Capsules:

typical of genus. Leaves dry beautifully with reddish

margins. Creeping growth forms are rare in Antimima.

48. Antimima lodewykii ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia lodewykii L.Bolus in Notes on Mesembryanthemum and

allied genera, part 3: 218 (1950). Type: L.van Heerde (3) BOL 23640

(BOL, lecto.!, here designated: chosen from several sheets).

Plants compact, heterophyllous. Leaves: sheath of one

leaf pair white, smooth, free tips papillate; second leaf pair

also highly connate, but entirely papillate, only bulging

keel and margins apically smooth, enclosed during the rest-

ing state. Capsules: typical of genus; closing bodies shaped

like heads of snakes and small. Typical of species: very tight

arrangement of old leaves, like pieces of meat on a spit.

49. Antimima loganii (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia loganii L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 61 (1937). Type: Logan 31 (BOL, holo.!).

Plants caespitose, white during the dry season, hetero-

phyllous. Leaves: sheath of one leaf pair long, smooth,

with short, free parts with papillae; other leaf pair only a

little connate, free parts trigonous, papillate, apiculate,

both ± 9 mm long. Capsules: typical of genus; narrow

valve wings. Typical of species: raised, star-shaped fruits

above the white tiny shrub during dry season.

50. Antimima lokenbergensis (L.Bolus) H.E.K.

Hartmann, comb. nov.

Ruschia lokenbergensis L Bolus in Journal of South African Botany

30: 239 (1964). Type: Esterhuysen 30699 (sheet I) (BOL, lecto.!, here

designated: chosen from 2 sheets).

Shrubs to 450 mm high, isophyllous. Leaves: elon-

gate-ovate. Capsules: typical of genus. Almost free

leaves unusually rounded for genus.

51. Antimima longipes (L.Bolus) Dehn in Mit-

teilungen aus dem Institut fur Allgemeine Botanik

Hamburg 22: 211 (1989). Type: Pillans 5811 (BOL,

holo.!).

Ruschia longipes L.Bolus: 256 (1927b). Mesembryanthemum longi-

pes (L.Bolus) N.E.Br.: 32 (1930).

Plants are caespitose cushions to ± 80 mm high, iso-

phyllous. Leaves: slender, trigonous. Capsules: on

pedicels 60 mm long; covering membranes with large

horizontal wings forming a second layer over them, clos-

ing ledges low and inconspicuous; closing body typical.

Typical of species: fruit morphology with valve wings

and radial wings on covering membranes plus presence

of big white closing bodies.

52. Antimima luckhoffii (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia luckhoffii L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 283 (1931). Type: Luckhoff 19611 (BOL, holo.!).

Plants caespitose, heterophyllous. Leaves: one leaf

pair with brown sheath at base and with 2-3 mm long

free parts densely covered with low papillae, tip thus

forming a ± triangular top; other leaf pair with short

sheath 5-6 mm long, free parts with translucent margins

and keel. Capsules: not known.

53. Antimima maleolens (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia maleolens L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 22 (1928d). Mesembryanthemum maleolens

(L.Bolus) N.E.Br.: 32 (1930). Type: Pillans BOL17796 (BOL, holo.!).

Shrubs up to 250 mm high smelling of salted fish, iso-

phyllous. Leaves: trigonous. Capsules: typical of genus.

54. Antimima maxwellii (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia maxwellii L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 106 (1929). Mesembryanthemum maxwellii

(L.Bolus) N.E.Br.: 32 (1930). Type: M. Bolus BOL18939 (BOL, holo.!).

Plants with compact centre with long shoots and high-

ly branched, tufted short shoots, bearing flowers (and

fruits) in groups, heterophyllous. Leaves: smooth, papery

sheath of one leaf pair of ± 5 mm length enclosing base

of subsequent leaf pair with sheath of ± 4 mm and free

parts up to 15 mm long; epidermis cells of free parts

almost flat, only slightly protruding. Capsules: typical of

genus; covering membranes raised high near centre and

with indications of radial protrusions, but these appear as

if folded and not forming a wing proper. Typical of

species: dense groups of short shoots developing on long

shoots.

55. Antimima menniei (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia menniei L.Bolus in Journal of South African Botany 31: 86

(1965). Type: Mennie BOL27660 (BOL, holo.!).

76

Bothalia 28,1 (1998)

Plants caespitose with erect branches, heterophyllous.

Leaves : short, smooth sheath of one type of leaf pair per-

sisting as envelopes around stem, long, free parts with big,

distant papillae; bracteoles without a sheath, leaves almost

separated by tissue of pedicel. Capsules: typical of genus;

expanding keels transgrading into expanding sheets.

56. Antimima mesklipensis (L.Bolus) H.E.K.Hart-

mann, comb. nov.

Ruschia mesklipensis L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 281 (1931). Type: Mathews (Ryder) NBG2250/30

(BOL, holo.!).

Plants caespitose, heterophyllous. Leaves: one leaf

pair with greyish brown sheath, 2-3 mm long, free parts

distantly papillate; other leaf pair connate for < 1 mm,

free parts broadly trigonous, densely papillate. Capsules:

not known. Resembles A. persistens in persistence of old

(longer) leaves.

57. Antimima meyerae (Schwantes) H.E.K. Hart-

mann, comb. nov.

Ruschia meyerae Schwantes in Monatsschrift der Deutschen

Kakteengesellschaft 2: 64 (1930). Type: Anon. s.n. (HBG, lecto.!, here

designated: photograph of the ‘type plants’ in collection Schwantes

HBG).

Compact shrublets, heterophyllous with 3 different

types of leaves. Leaves: most commonly sheaths whitish

yellow with reddish brown free tips; epidermis at tips

with rounded, low papillae; leaves up to 8 mm long; at

bases of side branches first leaf pair with a white-yellow

sheath, but without any different free tips, ± 5 mm long;

second leaf pair hardly connate, free parts with promi-

nent, long papillae giving leaves a rough appearance, ± 5

mm long. Capsules: very hard, differing from typical

form by possession of small finger-shaped closing bod-

ies. Typical of species: formation of at least three differ-

ent leaf forms and development of four new branches

under terminal fruit, in contrast to common pattern of

two or single branch.

58. Antimima microphyiia (Haw.) Dehn in Mit-

teilungen aus dem Institut fur Allgemeine Botanik

Hamburg 22: 213 (1989). Type: drawing by Duncanson

[K, lecto.!, designated by Dehn: 213 (1989)].

Mesembryanthemum microphyllum Haw.: 73 (1803). Ruschia micro-

phyiia (Haw.) Schwantes: 20 (1927).

Low shrublets, heterophyllous. Leaves: set densely,

keels cartilaginous. Capsule: not known.

59. Antimima minima (Tischer) H.E.K. Hartmann ,

comb. nov.

Cheiridopsis minima Tischer in Succulenta 9: 145 (1927). Type:

Anon. s.n. (HEID).

Plants compact, isophyllous. Leaves: trigonous.

Capsules: typical of genus. Description of fruit permits the

conclusion that the species belongs to Antimima , where in

several species, centrally raised covering membranes

occur combined with deep-set, white, flat closing bodies.

60. Antimima minutifolia (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia minutifolia L.Bolus in Journal of South African Botany 30:

239 (1964). Type: Hall 2829 (BOL, holo.!).

Dark shrublets, outer long shoots decumbent, hetero-

phyllous. Leaves: long sheath of one leaf pair turning

white, basally maroon, free parts 1-2 mm long, papillate;

second leaf pair hardly connate, free parts 3^1 mm long,

papillate and ciliate along margins, latter leaves on long

shoots, sheaths always at bases of short side shoots, on

which both leaf types alternate regularly. Capsules: typi-

cal of genus. Typical of species: dark appearance as a

result of dark stems which are well visible because

leaves are very short.

Distinguished from A. distans from the same area by

much more slender stems and a broad cushion shape with

age. The first description appeared in June 1928, being

validly published at that time; the Latin description

appeared in August 1928 (Bolus 1928c), but was super-

fluous at that time.

61. Antimima modesta ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia modesta L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 105 (1937). Type: Holloway 63 (BOL, holo.!).

Ruschia modesta L.Bolus forma glabrescens L.Bolus: 54 (1960).

Type: Geyer SUG13973 (BOL, holo.!).

Plants compact with long, erect, stiff shoots, isophyl-

lous. Leaves: boat-shaped with convex sides. Capsules:

typical of genus. The name should be used for stout

plants with few to some flowers per inflorescence from a

restricted area north of the Gariep River in Namibia.

62. Antimima mucronata (Haw.) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum mucronatum Haw. in Miscellanea naturalia: 73

(1803). Ruschia mucronata (Haw.) Schwantes: 20 (1927). Type: draw-

ing 980/203 of Duncanson (K, lecto.!, here designated).

Ruschia mathewsii L.Bolus: 139 (1927/1928) Type: Mathews NBG

1893/24 ( BOL, holo.!).

Plants compact with decumbent long shoots, hetero-

phyllous. Leaves: sheath of one leaf pair smooth, later

white, thin, papery, keel markedly horny, ciliate to serru-

late; second leaf pair connate for 1-2 mm only, free parts

± 10 mm long, with distinct awn borne apically, with low

papillae, keel ciliate to serrulate, wax in dense layers of

vertical platelets on free parts. Capsules: typical of

genus; expanding keels merging gradually into expand-

ing sheets.

The drawing of Duncanson (K!) shows a plant in the

growing season with several long shoots (with prominent

internodes) with shortly connate, spreading leaves,

resembling those of long shoots of A. mathewsii very

closely.

63. Antimima mutica (L.Bolus) H.E.K. Hartmann,

comb. nov.

Bothalia 28,1 (1998)

77

Ruschia mutica L. Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 61 (1929). Mesembryanthemum muticum (L.Bolus)

N.E.Br.: 32 (1930). Type: Pillans BOL18898 (BOL, holo.!).

Compact shrublets, heterophyllous. Leaves : sheath of

one leaf pair white, ± 4 mm long, free tips ± 3 mm long,

covered by fine papillae, longer ones on margins; second

leaf pair with a 1-2 mm long sheath, free parts ± 6 mm

long, papillate, the latter on long shoots, short shoots

beginning with a sheathed leaf pair, later both types alter-

nating regularly. Capsules : typical of genus. Smaller-

leaved version of A. peersii.

64. Antimima nobilis (Schwantes) H.E.K. Hart-

mann, comb. nov.

Ruschia nobilis Schwantes in Monatsschrift der Deutschen Kakteen-

gesellschaft 2: 65 (1930). Type: M. Schlechter s.n. (HBG, collection

Schwantes!).

Plants compact with long shoots, isophyllous. Leaves :

trigonous with convex sides, thick. Capsules : typical of

genus; closing bodies sometimes smaller; covering

membranes sometimes a little reduced, 6-loculed.

65. Antimima nordenstamii (L. Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia nordenstamii L.Bolus in Journal of South African Botany 30:

241 (1964). Type: Nordenstam 799 (BOL, holo.!).

Plants compact with long, erect, or decumbent shoots

with dark purple to blackish internodes, several to many

short shoots developing on distal nodes, arrangement

resembling mistletoe on a branch, heterophyllous.

Leaves : one leaf pair with a wide, yellow sheath with

only short free parts, ± 6 mm long overall; other pair

united for ± ‘A of its length, trigonous, obtuse, up to 12

mm long; epidermis of free parts papillate. Capsules:

typical of genus. Typical of species: blackish internodes

and crowded short shoots bearing flowers.

66. Antimima oviformis (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia oviformis L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 261 (1931). Type: Herre SUG9311 (BOL, holo.!).

Low shrubs with long shoots only, heterophyllous.

Leaves: sheath of one leaf pair 5-8 mm long, free papil-

late tips shortly triquetrous, breaking off easily leaving

only sheaths on plants; other leaf pair with a short con-

nate base, triquetrous, papillate. Capsules: unknown.

Typical of species: bare, golden brown shoots.

67. Antimima papillata (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia papillata L Bolus in South African Gardening and Country

Life 17: 256 (1927b); L.Bolus: 129 (1927/1928). Mesembryanthemum

papillatum (L.Bolus) N.E Br.: 32 (1930). Type: Pillans 5713 (BOL,

holo.!).

Ruschia meyeri Schwantes: 300 (1928a). Mesembryanthemum mey-

eri (Schwantes) N.E.Br.: 32 (1930). Type: Anon, s.n., photograph of

type material (HBG, lecto.!, collection Schwantes, here designated).

Plants compact with bundles of erect long shoots, het-

erophyllous. Leaves: white, smooth sheath of one leaf

pair connate for at least '/2 its length; second pair connate

for ± 3 mm, free parts elongate-trigonous with rather

large, dense papillae, with a dense cover of wax platelets

transgrading into threads, underlying continuous wax

breaking eventually into big plates. Capsules: without

closing bodies, but with typical broad expanding keels

and narrow valve wings broadest at middle, both features

typical of genus.

68. Antimima paucifolia ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia paucifolia L.Bolus in Journal of South African Botany 30:

240 (1964). Type: Esterhuysen 30700 (BOL, holo.!).

Shrubs with diagonal long shoots, heterophyllous.

Leaves: sheath of one leaf pair white, basally maroon,

with short free parts ± 2 mm long, upper part breaking

away after resting period along pre-formed ring, leaving

a clear-cut edge surrounding stem; second leaf pair tri-

quetrous, connate part only ± 1 mm long, all free parts of

leaves with papillae of medium height, keel and the mar-

gins with distant, longer papillae. Capsules: typical of

genus. Similar to A. exsurgens.

69. Antimima pauper ( L.Bolus ) H.E.K. Hartmann,

comb. nov.

Ruschia pauper L.Bolus in Journal of South African Botany 27: 259

(1961). Type: Littlewood KG211/61 (BOL, holo.!).

Plants compact with long shoots on which short

shoots are developed in tufted bundles, isophyllous.

Leaves: rounded at tip. Capsules: typical of genus.

70. Antimima peersii (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia peersii L Bolus in Notes on Mesembrianthemum and allied

genera, part 1: 139 (1927/1928). Mesembryanthemum formulosum

N.E.Br.: 32 (1930). Type: Peers NBG3725/I5 (= BOL44507, not

3825/15 as in one description, BOL, lecto.!, here designated).

Plants compact with long shoots, heterophyllous.

Leaves: sheath of one leaf pair white, often with a cop-

per-coloured tinge at its upper end, ± 8 mm long, free

parts ± 10 mm long; subsequent leaf pair only a little

connate with ± 20 mm long free parts appressed to each

other during resting state, long shoots with these latter

leaves only, short shoots starting with sheathed leaf pair,

leaf forms alternating later; epidermis of free parts papil-

late. Capsules: typical of genus; narrow valve wings and

medium-sized closing bodies.

71. Antimima perforata (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia perforata L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 80 (1929). Mesembryanthemum perforatum

(L.Bolus) N.E.Br.: 32 (1930). Type: Kolle in Pillans 6097 (BOL,

lecto.!, here designated: chosen from several sheets).

Ruschia paripetala L.Bolus var. occultans L.Bolus: 125 (1929).

Type: Pillans 5827 (BOL, holo.!).

Plants with erect, stiff branches with proleptic short

shoots from a rather dense, caespitose centre. Capsules:

typical of genus with 6 locules.

78

Bothalia 28,1 (1998)

72. Antimima persistens (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia persistens L.Bolus in Journal of South African Botany 29: 16

(1963) nom. illeg., non L.Bolus: 334 ( 1932). Type: Van Breda 1750/62

(BOL, holo.!).

Plants compact with long shoots, heterophyllous.

Leaves : sheaths of one leaf pair ± 4 mm long and free

parts ± 2 mm long, these dominant on long shoots; other

leaf pair connate for ± 2 mm, free parts ± 4 mm long, all

free parts with elevations and papillae, on short shoots,

types of leaf pairs alternating regularly, longer leaves

persisting over several years. Capsules: typical of genus;

closing bodies medium-sized.

The name Ruschia persistens was used for two differ-

ent species by Bolus (1932, 1963), the latter being an

illegitimate name in Ruschia. However, the epithet is

available in the genus Antimima.

73. Antimima pilosula (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia pilosula L.Bolus in Notes on Mesembryanthemum and allied

genera, part 3: 259 (1954). Type: Hall NBG285/54 (BOL, holo.!).

Shrubs with many scars from old leaves, isophyllous.

Leaves: subfalcate to subclavate. Capsules: typical of

genus; valve wings in their basal halves broader than

expanding keels.

The broad, tapering valve wings, resembling those of

Eberlanzia schneideriana (A.Berger) H.E.K.Hartmann,

are unusual in the genus. The species is placed here

based on the broad, typical closing bodies and the undu-

late covering membranes with inconspicuous closing

ledges only.

74. Antimima piscodora (L.Bolus) H.E.K.Hart-

mann, comb. nov.

Ruschia piscodora L.Bolus in Notes on Mesembrianthemum and

allied genera, part 1: 141 (1927/1928). Mesembryanthemum piscodor-

um (L.Bolus) N.E.Br.: 32 (1930). Type: Tugwell BOL17109 (BOL,

holo.!).

Plants compact with long shoots. Capsules: incom-

pletely described: covering membranes touching closing

body; expanding keel denticulate, but these features are

typical of genus.

75. Antimima prolongata (L.Bolus) H.E.K.Hart-

mann, comb. nov.

Ruschia prolongata L.Bolus in Notes on Mesembryanthemum and

allied genera, part 3: 280 (1954). Type: Acocks 16937 (sheet I) (BOL,

lecto.!, here designated: chosen from 2 sheets).

Plants compact with very long, thin, yellow long

shoots in other bushes, heterophyllous. Leaves: one type

of leaf pair forming a papery sheath-like cover of two

parts ± 10 mm long, connate only for ± 1 mm, apical part

papillate, subulate; second leaf pair ± 8-10 mm long, tri-

quetrous, subulate, papillate, mucro recurved. Capsules:

typical of genus. Typical of species: pseudo-sheaths

formed by one leaf pair consisting of almost free leaves,

but turning papery for dry season like connate sheaths in

other species.

76. Antimima propinqua (N.E.Br.) H.E.K.Hart-

mann, comb. nov.

Mesembryanthemum propinquum N.E.Br. in Kew Bulletin 1929: 59

(1929). Ruschia propinqua (N.E.Br.) Schwantes: 230 (1938). Type:

Muir 4071 (K, holo ).

Plants compact, heterophyllous. Leaves: sheath of one

leaf pair connate; second leaf pair almost free, 5-6 mm

long, free parts keeled, margins and keel finely ciliate, all

glaucous-green. Capsules: not known.

77. Antimima prostrata (L.Bolus) H.E.K.Hart-

mann, comb. nov.

Ruschia prostrata L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 61 (1929). Mesembryanthemum prostratum

(L.Bolus) N.E.Br.: 32 (1930). Type: Leipoldt BOL18899 (BOL, holo. !).

Plants creeping with proleptic side branches as tiny

lateral bodies, later forming a mat, heterophyllous.

Leaves: sheath of one leaf pair long, yellowish, rather

smooth, enclosing larger part of subsequent leaf pair

with a shorter sheath and longer free parts with papillae,

in particular along margins and keel, often two or more

leaf pairs of second type following each other, ± 3 mm

long, body ± 2.5 mm diam. Capsules: typical of genus.

Small-leaved version of A. brevicarpa sharing similar

character states with that species, in particular the same

type of fruit and ecology.

78. Antimima pumila (Fedde & Schuster) H.E.K.

Hartmann, comb. nov.

Mesembryanthemum pumilum L.Bolus ex Fedde & Schuster in Just’s

Botanischer Jahresbericht 41: 92 (1918). Ruschia pumila (Fedde &

Schuster) L.Bolus: 60 ( 1929). Mesembryanthemum pumilum (L.Bolus)

N.E.Br.: 32 (1930) nom. illeg. Type: Pearson 3917 (BOL, holo.!).

Ruschia levynsiae (L.Bolus) Schwantes: 58 (1949). Mesembryanthe-

mum levynsiae L.Bolus: 256 (1927b). Type: Levyns NBG1 373/26

(BOL, holo.!).

Plants compact, heterophyllous. Leaves: smooth

sheath on one leaf pair with short, spreading, papillate,

rough tips; second leaf pair enclosed which unfolds dur-

ing rainy season showing two almost oval, spreading

leaves, connate slighty at bases and exhibiting upper sur-

faces, a rare condition in compact Ruschioideae; epider-

mis papillate. Capsules: typical of genus.

The difference in appearence between the resting

state, when the yellowish green leaf pair forming a round

body with the free tips protruding, and the growing phase

is striking — in season, the bright green leaf pair spreads

so completely that the persistent sheaths are hardly visi-

ble. This difference is the reason why Bolus described

the species twice within two years, having seen Ruschia

pumila in the resting state only, but Ruschia levynsiae

growing, developing the described heterophylly (Figure

IE). Bolus (1913: 150) had described ‘M. pumilum, circa

5 cm altum’, using the adjective as a descriptive word

only, not as a formal cpilheton to name a species, as she

stated herself in 1929, when describing Ruschia pumila.

Meanwhile, Fedde & Schuster (1918) had taken

Mesembryanthemum pumilum L.Bolus as a validly

described name, which it became only through their cita-

Bothalia 28.1 (1998)

79

tion of the basionym and the type, hence the authorship

‘L. Bolus ex Fedde & Schuster’.

79. Antimima pusilla (Schwantes) H.E.K. Hart-

mann, comb. nov.

Ruschia pusilla Schwantes in Zeitschrift fiir Sukkulentenkunde 3:

300 (1928a). A lesembryanthemum parvum N.E.Br.: 32 (1930). Type:

photograph of type (HBG, lecto.l, in collection Schwantes HBG, here

designated).

Plants compact, heterophyllous. Leaves: very similar

in so far as they both develop long, smooth sheaths, one

leaf pair with low distant papillae on short free parts (this

one persisting during dry season enclosing next leaf

pair); other with distinctly longer papillae thus looking

rougher, papillae elongated along margins. Capsules :

with expanding keels merging into expanding sheets and

both together covering lower half of valve (when open);

closing body small with a hollow underneath into which

placenta extends. The heterophylly may be overlooked

easily, but the fact that during the resting time one leaf

pair encloses the subsequent one indicates the different

roles the two leaf pairs play.

80. Antimima pygmaea (Haw. ) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum pygmaeum Haw. in Supplementum plantarum

succulentarum: 98 (1819). Ruschia pygmaea (Haw.) Schwantes: 92

(1928b). Type: drawing no. 996/204 by Duncanson (K, lecto.l, here

designated).

Plants caespitose, forming mats, heterophyllous.

Leaves: sheath of one leaf pair developing into a conical

body enveloping subsequent leaf pair, upper part disinte-

grating into a cracked cover through which the following

leaf pair grows when rain starts to fall, tips of both leaf

types pointed and triangular as seen from above; epider-

mis very smooth, covered by a thin rugose layer of wax.

Capsules: typical of genus; very narrow valve wings, 6-

loculed. Typical of species: conical white bodies formed

by leaf pairs in resting period becoming cracked with

time; laciniate old leaves occur also in A. ivori, but those

are truncate and appear like a trimmed edge.

81. Antimima quarzitica (Dinter) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum quarziticum Dinter in Feddes Repertorium 19:

151 (1923). Ruschia quarzitica (Dinter) Dinter & Schwantes: 69

(1929). Corpusculariu quarzitica (Dinter) Schwantes: 186 (1926).

Type: Dinter 3866 (B. holo.l).

Plants compact with erect to spreading long shoots,

bearing short shoots with several leaf pairs, isophyllous.

Leaves: triquetrous, apically recurved. Capsules: typical

of genus; closing bodies medium-sized.

82. Antimima roseola (N.E.Br.) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum roseolum N.E.Br. in Kew Bulletin 1929: 60

(1929). Ruschia roseola (N.E.Br.) Schwantes: 230 (1938). Type: Muir

4067 (not 4062, as in description) (K, holo.l).

Plants caespitose, heterophyllous. Leaves: sheath of one

leaf pair long, whitish with rather short free tips; epidermis

nearly smooth; margins and keel turning white with age,

keel continuing in a fold or narrow wing down sheath and

stem below; second leaf pair with longer free parts with

medium long papillae. Capsules: typical of genus; expand-

ing tissue thick and short at base of valve, rising to inner

side, but not forming a distinct expanding keel.

83. Antimima saturata ( L. Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia saturata L Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 122 (1929). Mesembryanthemum atrocinctum N.E.Br.:

32 (1930). Type: Pillans BOL18952 (BOL, holo.l).

Plants compact with long shoots, heterophyllous.

Leaves: white, papery sheath of one leaf pair 3-4 mm

long, free parts 4—5 mm long, with low papillae; other

leaf pair united for ± 2 mm only, free parts up to 10 mm

long, with long papillae on keel and margins, low papil-

lae of different sizes on sides, wax cover dense, consist-

ing of flakes often connected. Capsules: unknown.

The heterophylly had not been recognized in the orig-

inal description, nor the differences in papillae. In both

features, the species resembles A. mathewsii, which has

thicker and more robust stems but is similar in the irreg-

ular branching pattern.

84. Antimima saxicola (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia saxicola L Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 46 (1929). Mesembryanthemum saxicola (L.Bolus)

N.E.Br.: 32 (1930). Type: Smith 4090 (sheet II) (BOL, lecto.l, here des-

ignated from several sheets of that collection).

Plants compact with long shoots blackish brown with

age, isophyllous. Leaves: triquetrous, keel and margins

cartilaginous. Capsules: typical of genus; covering mem-

branes deeper in centre. Plants resemble those of Ruschia

orientalis L.Bolus and Ruschia putterillii (L.Bolus)

L.Bolus with tiny closing bodies, in growth form and

ecological preferences.

85. Antimima schlechteri (Schwantes) H.E.K. Hart-

mann, comb. nov.

Ruschia schlechteri Schwantes in Zeitschrift fiir Sukkulentenkunde

3: 301 (1928a). Mesembryanthemum schlechteri (Schwantes) N.E.Br.:

32 (1930). Type: Anon, s.n., photograph of ‘type plant’ (HBG, lecto.l,

here designated, collection Schwantes HBG).

Plants compact, with few longer shoots with dense

heads of leaves and fruits, heterophyllous. Leaves:

longer sheaths of leaf pairs persisting as a series of tubes

around old stem for several years, free parts of leaves

short; other type of leaf pair trigonous, epidermis almost

smooth, ± 9 mm long, connate for ± 5 mm. Capsules:

typical of genus; narrow valve wings and radial wings on

covering membranes.

86. Antimima simulans (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia simulans L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 260 (1931). Type: Frames BOL19375 (BOL, holo.l).

Low shrubs, heterophyllous. Leaves: sheaths of one

leaf pair 4-5 mm long, free parts 1-2 mm long, papillae

80

Bothalia 28,1 (1998)

low; other leaf pair with sheath ± 4 mm long, free parts

up to 9 mm long, on long shoots at each node in one sea-

son. Capsules : unknown.

87. Antimima sobrina (N.E.Br.) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum sobrinum N.E.Br. in Kew Bulletin 1929: 60

(1929). Ruschia sobrina (N.E.Br.) Schwantes: 230 (1938). Type: Muir

4014 (K, holo.).

Plants caespitose, heterophyllous. Leaves', sheaths of

one type of leaf pair connate for just over half their

length; other leaf pair almost free, free parts trigonous,

keel scabrous, 3-5 mm long. Capsules: not known.

88. Antimima solida (L.Bolus) H.E.K. Hartmann,

comb. nov.

Mesembryanthemum solidum L.Bolus in Annals of the Bolus

Herbarium 3: 136 (1922). Ruschia solida (L.Bolus) L.Bolus: 239

(1927b). Type: Pillans 2421 (sheet I) (BOL, lecto.!, here designated:

chosen from two sheets).

Ruschia solida (L.Bolus) L.Bolus var. stigmatosa L.Bolus: 94 (1929).

Type: Pillans 6048 1, cited as NBG147/28 , marked ‘ = 147/25’ on sheet

(BOL, lecto.!, here designated: chosen from two sheets).

Plants compact to forming big mats, isophyllous.

Leaves: finger-shaped to terete. Capsules: typical of

genus; only dorsal bundles of fruit present, interior hav-

ing fallen out as a tumble fruit, tiny valve wings some-

times present. Resembles Argyroderma fissum (Haw.)

L.Bolus in its decumbent growth and finger-shaped

leaves; both species grow sympatrically and can easily

be confused in their vegetative state. Since fruits remain

on the plants in both species, a distinction in habitat is

easier.

89. Antimima stayneri ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia stayneri L.Bolus in Journal of South African Botany 27: 260

( 1961 ). Type: Stayner KG450/61 (BOL, holo. !).

Shrubs with stout stems, heterophyllous. Leaves:

papery protective cover of two parts derived from one

type of leaf pair, these leaves elongate, free, to 10 mm

long, tips slender, pointed, dark when dry; second pair

almost free, finely papillate, ± 8 mm long. Capsules: typ-

ical of genus. In leaves and stems A. stayneri resembles

A. emarcescens and A. subtruncata, but the latter species

grow as climbers in other bushes.

90. Antimima stokoei (L.Bolus) H.E.K. Hartmann,

comb. nov.

Ruschia stokoei L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2 160 (1929) Type: Stokoe NBG28/25 (BOL, holo.!).

Plants compact, heterophyllous. Leaves: long sheath

in one type of leaf pair, free parts with some short, dis-

persed papillae; other leaf pair with short sheaths and

long, papillate free parts. Capsules: with very narrow

valve wings and small closing bodies. The type looks

very similar to that of A. verruculosa, differing mainly in

the diameter of the flower, a feature known to change

during anthesis.

91. Antimima subtruncata (LBolus) H.E.K. Hart-

mann, comb. nov.

Ruschia subtruncata L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 211 (1930). Type: L.Bolus BOL19231 (BOL,

holo.!).

Ruschia subtruncata L.Bolus var. minor L.Bolus: 286 (1954). Type:

Acocks 17185 (BOL, holo.!).

Plants compact with long shoots twining in other

bushes, heterophyllous; side branches with scars and

somewhat spiny remains of closely set leaf pairs. Leaves:

one leaf pair with papery protective cover of two parts ±

10 mm long, connate for only ± 2 mm, apical part papil-

late; second leaf pair enveloped during resting period, ±

6-10 mm long, free parts triquetrous, papillate, with

some bigger papillae along margins, mucro recurved.

Capsules: typical of genus; expanding sheets rising to

form keels. Typical of species: pseudo-sheaths formed by

one leaf pair consisting of almost free leaves, but turning

papery for the dry season like connate sheaths in other

species', these sheaths are about as long as broad in A.

subtruncata. Similar in general appearence to A.

emarcescens and A. stayneri, both with slender leaves.

92. Antimima triquetra (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia triquetra L.Bolus in South African Gardening and Country

Life 18: 178 (1928a); L.Bolus: 231 (1928d). Mesembryanthemum tri-

quetrum (L.Bolus) N.E.Br.: 33 (1930). Type: Banks NBG1935/15

(BOL, holo.!).

Plants compact with long shoots, erect at first, later

decumbent, heterophyllous. Leaves: one type of leaf pair

with a papery protective cover of two parts ± 8 mm long,

connate for only ± 2 mm, apical part subulate, with low

papillae; second leaf pair ± 6-8 mm long, triquetrous,

with long papillae along keel and margins, mucro erect.

Capsules: typical of genus. Typical of species: pseudo-

sheaths formed by one leaf pair consisting of almost free

leaves, but turning papery for the dry season like connate

sheaths in other species — in this feature, the species

resembles A. subtruncata.

93. Antimima tuberculosa (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia tubercolusa L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 1 10 (1929). Mesembryanthemum enorme N.E.Br.:

33 (1930). Type: Leipoldt BOL18940 (BOL, holo.!).

Plants caespitose, forming mats with age, isophyllous.

Leaves: trigonous. Capsules: typical of genus with very

narrow valve wings. The epithet tuberculosa refers to the

big closing body Bolus noticed when describing the

species.

94. Antimima turneriana (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia turneriana L.Bolus in Journal of South African Botany 29:

175 (1963). Type: Van Breda 1912/63 (BOL, holo.!).

Plants compact in habitat, developing some long

shoots in cultivation, isophyllous. Leaves: triquetrous

with convex sides. Capsules: typical of genus; radial pro-

trusions ± wing-shaped, disintegrating later, forming

rows of long teeth; valve wings at first as broad as

Bothalia 28.1 (1998)

expanding keels, disintegrating later quickly. Similar to

A. dualis , both compact in habitat, but the latter never

changing shape.

95. Antimima vanzylii (L.Bolus) H.E.K. Hart-

mann, comb. nov.

Ruschia vanzylii L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 209 (1930). Type: Van Zyl in Fuller 94 (BOL, holo.!).

Plants compact with long shoots when not eaten, iso-

phyllous. Leaves : triquetrous. Capsules : typical of genus.

In spite of their strong smell of soda, plants are obvious-

ly rather heavily browsed by animals.

96. Antimima varians ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia varians L.Bolus in Notes on Mesembrianthemum and allied

genera, part 2: 209 (1930). Type: L.Bolus BOL19212 (BOL, holo.!).

Plants compact with long shoots developing shorter

side shoots in bundles or groups. Capsules : typical of

genus.

97. Antimima ventricosa ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Mesembryanthemum ventricosum L.Bolus in Annals of the Bolus

Herbarium 3: 128 (1922). Cheiridopsis ventricosa (L.Bolus) N.E.Br.:

73 (1926). Ruschia ventricosa (L.Bolus) Schwantes: 106 (1927). Type:

Pillans NBG475/16 (BOL, holo.!).

Plants compact, heterophyllous. Leaves : one type of

leaf pair with a long sheath connate for almost half its

length; second with a longer free part; free parts trigo-

nous and papillate in both types, but more densely so in

younger, less connate leaves, 40-80(-120) mm long,

13-17 mm broad, 10-13 mm diam. Capsules: without

valve wings, expanding keels distant, 4-6-loculed, in-

completely known. The low number of locules excludes

the species from Cheiridopsis, where it might otherwise

belong.

98. Antimima verruculosa ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia verruculosa L.Bolus in Notes on Mesembrianthemum and

allied genera, part 2: 125 (1929). Mesembryanthemum verruculosum

(L.Bolus) N.E.Br.: 33 (1930). Type: Haymes NBG592/26 (BOL.

holo.!).

Low shrublets, heterophyllous. Leaves : sheath of one

leaf pair long, smooth with a short, papillate, free part;

other leaf pair with a short sheath and longer papillae.

Capsules : with a small closing body, partly destroyed.

The ‘type plant’ is very similar to A. stokoei, which has

larger flowers.

99. Antimima watermeyeri ( L.Bolus ) H.E.K. Hart-

mann, comb. nov.

Ruschia watermeyeri L.Bolus in Notes on Mesembrianthemum and

allied genera, part 1: 146 (1927/1928). Type: Watermeyer NBG554/23

(BOL, holo.!).

Ruschia stenopetala L Bolus: 53 (1960) Type: Littlewood KG276/60

(BOL, holo.!).

Ruschia obtusifolia L.Bolus: 298 (1962) Type: Hall 2467 (BOL,

holo.!).

Plants caespitose, isophyllous. Leaves: trigonous to

roundish. Capsules: typical of genus; extremely narrow

valve wings at areas where expanding keel meets edge of

valve.

The type material of all species included here is very

similar to that of A. klaverensis in growth pattern, leaf

shape, and fruit morphology, differing only in the

absence of the fishy smell reported for the latter species.

100. Antimima wittebergensis (L.Bolus) H.E.K.

Hartmann, comb. nov.

Mesembryanthemum wittebergense L.Bolus in Annals of the Bolus

Herbarium 4: 88 (1927a). Ruschia wittebergensis (L.Bolus) Schwantes:

106 (1927). Type: Compton NBG 1920724 (BOL, holo.!).

Plants caespitose, isophyllous. Leaves: cymbiform or

boat-shaped. Capsules: with small closing bodies, excep-

tionally broad and thick closing ledges and narrow valve

wings.

SPECIES EXCLUDED FROM ANTIMIMA

Ruschia virgata (Haw.) L.Bolus in South African

Gardening and Country Life 17: 239 (1927); L.Bolus: 72

(1927/1928).

Mesembryanthemum virgatum Haw.: 88 (1803); Antimima virgata

(Haw.) Dehn: 213 (1989). Type: Bortenschlag s.n. (K, lecto.!, desig-

nated by Dehn 1989: 213).

Plants with erect, thin branches from a denser centre,

often in other bushes. Leaves: triquetrous, hardly con-

nate, pair at its base leaving a triangle of stem visible

between them. Capsule: base elongate, funnel- to bell-

shaped, top almost semi-orbicular from high valve rims;

covering membranes forming a roof, ridge being the con-

tact line, closing ledges sharp and long; closing body

nearly hook-shaped, hollow; expanding keels distant at

bases. Material matching the collection of Bortenschlag,

which was seen by Haworth, possesses fruits of the

Ruschia type, with deep locules and small, hook-shaped

closing bodies. The species is therefore placed back in

Ruschia.

ACKNOWLEDGEMENTS

The basic work has been supported financially over

many years by the Deutsche Forschungsgemeinschaft;

additional travel grants were received from the

University of Hamburg, the Hansische Universitats-

stiftung, and the Cactus and Succulent Society of

America. To all donors I am most grateful. I should also

like to thank many friends, colleagues, students, and col-

laborators during field trips and in connection with the

extensive studies on the material. The directors of the

herbaria B, BOL, K are thanked for extensive loans of

type sheets, for making available prints of drawings, and

for much additional information. My thanks go also to

several referees who have helped to reduce mistakes in

citations and explanations.

82

Bothalia 28,1 (1998)

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Both alia 28,1: 83-90 (1998)

Meiotic chromosome behaviour in Cenchrus ciliaris (Poaceae: Pani-

coideae)

N.C. VISSER*, J.J. SPIES* + and H.J.T. VENTER*

Keywords: Cenchrus ciliaris , chromosomes, meiosis, Panicoideae, Poaceae, polyploidy

ABSTRACT

A basic chromosome number of x = 9 has been confirmed for Cenchrus ciliaris L. Polyploidy is common and levels vary

from tetraploid to hexaploid. Aneuploidy is reported for a single specimen, where two chromosomes of a single genome

were lost. Various meiotic irregularities were observed. The highest incidence of meiotic abnormalities was observed in the

pentaploid specimens. This was attributed to their uneven polyploid level. All specimens varied from segmental alloploid to

alloploid.

INTRODUCTION

The classification of the genus Cenchrus L. is com-

plex (De Lisle 1963) and it is sometimes difficult to

describe species on gross morphological characters alone

(Chapman 1990). Meiotic chromosome behaviour can

make a vast contribution to the classification process.

The aim of the present study was to use meiotic chro-

mosome behaviour in C. ciliaris in order to deduce the

polyploid origin of the specimens studied, as well as the

chromosomal variation within and among populations of

C. ciliaris in South Africa.

MATERIALS AND METHODS

Specimens were collected and fixed in the veld.

Voucher herbarium specimens are housed in the Geo

Potts Herbarium, Department of Botany and Genetics,

University of the Orange Free State, Bloemfontein

(BLFU) (Table 1).

Slides were prepared for meiotic analysis (Visser &

Spies 1994). A minimum of 20 cells of each of the fol-

lowing stadia were studied: diakinesis, metaphases I and

II, anaphases I and II and telophases I and II. The fol-

lowing were recorded: gametic chromosome numbers,

the number of univalents, laggards and micronuclei dur-

ing metaphase I, anaphase I and II, and telophase I and

II, respectively. The number of chiasmata per cell was

inferred from the chromosome configurations observed

during diakinesis or metaphase I.

The genomic constitution of some of the tetraploid

plants was calculated, based on the models proposed by

Kimber & Alonso (1981). Expected chromosome config-

urations for each of the proposed models (4:0, 3:1, 2:1:1

and 2:2) were calculated and compared with those

observed. The average sum of squares (SS) between the

expected and the observed values were calculated. The

* Department of Botany and Genetics, University of the Orange Free

State, RO. Box 339, 9300 Bloemfontein,

t To whom correspondence should be addressed.

MS. received: 1996-12-10.

relative affinity between the pairs of genomes was

expressed as an x-value. An x-value of 0.5 indicated a

close affinity between corresponding (homologous)

genomes, whereas non-homologous genomes were rep-

resented by an x-value of 1 . The model with the lowest

SS-value provided the best ‘fit’ for that particular speci-

men. The specific model was then considered to repre-

sent the genomic constitution of the specimen.

RESULTS

From all the meiotic stages studied, only metaphase

I, anaphase I and telophase I, proved to contribute to

meiotic analyses. Four ploidy levels, namely aneu-

ploidy (n = 2x - 1 = 17), tetraploidy (n = 2x = 18), pen-

taploidy (n = 5/2x = 45/2) and hexaploidy (n = 3x = 27)

were observed, based on a basic chromosome number of

x = 9 (Table 2).

A wide range of meiotic abnormalities were observed

for Cenchrus ciliaris. These abnormalities included the

presence of univalents during metaphase I, chromosome

and chromatid laggards during anaphases I and II respec-

tively, uneven segregation of chromosomes during

anaphase I, anaphase I bridges and micronuclei during

telophases I and II. Two additional meiotic abnormalities

were observed, namely precocious disjunction of chro-

mosomes into chromatids during anaphase I and the

absence of cytokinesis at the end of telophase I.

The presence of univalents, chromosome and/or chro-

matid laggards and micronuclei, were compared among

the tetraploid specimens. A histogram was plotted, based

on the percentage of cells containing the various num-

bers of univalents, chromosome and/or chromatid lag-

gards and micronuclei (Figure 1). The topography of the

curves corresponded. This fact indicated that, for C. cil-

iaris, the number of univalents present during metaphase

I, is the most likely reason why there is an increase in the

chromosome and/or chromatid laggards observed during

anaphase I. This in turn, could have contributed to the

formation of micronuclei during telophase I. The

absence of various peaks in the hyperboles indicates that

no distinct chromosomal groups exist in the tetraploid

specimens. Therefore, the average percentages for each

84

Bothalia 28,1 (1998)

TABLE 1. — List of Cenchrus ciliaris specimens studied, voucher specimen numbers and localities according to the degree reference system

(Edwards & Leistner 1971)

n Locality

Voucher

17 NORTH-WEST. — 2627 (Potchefstroom): in Potchefstroom, on route to Orkney, (-CA)

18 GAUTENG. — 2528 (Pretoria): near Pretoria, (-CC)

NORTH-WEST. — 2627 (Potchefstroom): in Potchefstroom, on route to Orkney, (-CA)

GAUTENG. — 2628 (Johannesburg): Grassmere Garage, Johannesburg, (-AB)

NORTH-WEST. — 2724 (Taung): 101 km from Kuruman to Vryburg, (-AB)

NORTH-WEST. — 2725 (Bloemhof): in Amalia, on route to Schweizer-Reneke, (-AA)

NORTH-WEST— 2725 (Bloemhof): 2 km from Britten to Christiana, (-CB)

FREE STATE. — 2726 (Odendaalsrus): 8 km from Wesselsbron to Bultfontein, (-CD)

FREE STATE — 2726 (Odendaalsrus): 46 km from Bothaville to Wesselsbron, (-DA)

FREE STATE. — 2727 (Kroonstad): 61 km from Kroonstad to Parys, (-AC)

FREE STATE. — 2727 (Kroonstad): 7 km from Kroonstad to Kroonvaal, (-CA)

KWAZULU-NATAL. — 2732 (Ubombo): Mhlosinga, on route to Sordwana, (-CC)

NORTHERN CAPE. — 2822 (Glen Lyon): 7 km from Smidtsdrift to Postmasburg, (-DA)

FREE STATE. — 2826 (Brandfort): 57 km from Wesselsbron to Bultfontein, (-AA)

FREE STATE. — 2826 (Brandfort): 30 km from Wesselsbron to Bultfontein, (-BB)

FREE STATE. — 2826 (Brandfort): 25 km from Bloemfontein to Brandfort, (-CD)

FREE STATE. — 2826 (Brandfort): 27 km from Bloemfontein to Brandfort, (-CD)

FREE STATE. — 2826 (Brandfort): 38 km from Bloemfontein to Brandfort, (-CD)

FREE STATE. — 2826 (Brandfort): 32 km from Bloemfontein to Abrahamskraal, (-CD)

FREE STATE. — 2925 (Jagersfontein): 44 km from Petrusburg to Kimberley, (-AB)

FREE STATE. — 2926 (Bloemfontein): near Bloemfontein, (-AA)

FREE STATE. — 2926 (Bloemfontein): 16 km from Bloemfontein to Winburg, (-AA)

EASTERN CAPE. — 3125 (Steynsburg): 30 km from Steynsburg to Hofmeyr, (-BC)

EASTERN CAPE. — 3125 (Steynsburg): near Hofmeyer, (-DC)

EASTERN CAPE. — 3125 (Steynsburg): 12 km from Hofmeyer to Cradock, (-DC)

WESTERN CAPE. — 3222 (Beaufort West): 5 km from Beaufort West, (-BC)

EASTERN CAPE — 3224 (Graaff-Reinet): 58 km from Jansenville to Graaff-Reinet, (-BC)

EASTERN CAPE. — 3224 (Graaff-Reinet): 131 km from Uitenhage to Graaff-Reinet, (-DC)

EASTERN CAPE. — 3224 (Graaff-Reinet): 145 km from Uitenhage to Graaff-Reinet, (-DC)

EASTERN CAPE. — 3224 (Graaff-Reinet): 122 km from Patensie to Willowmore, (-DD)

EASTERN CAPE. — 3225 (Somerset East): 57 km from Cradock to Cookhouse, (-DB)

EASTERN CAPE. — 3225 (Somerset East): Kokskraal, Cookhouse, (-DB)

WESTERN CAPE. — 3320 (Ladismith): 4 km from Calitzdorp to Oudtshoorn via Kuilsrivier, (-DC)

EASTERN CAPE. — 3324 (Steytlerville): 102 km from Uitenhage to Graaff-Reinet, (-BD)

EASTERN CAPE. — 3325 (Port Elizabeth): 40 km from Uitenhage to Graaff-Reinet, (-CD)

18+0-2B NORTH-WEST. — 2624 (Vryburg): near Vryburg, on route to Kuruman, (-DC)

NORTH-WEST. — 2627 (Potchefstroom): 10 km from Parys to Potchefstroom, (-CD)

FREE STATE. — 2925 (Jagersfontein): 60 km from Petrusburg to Kimberley, (-AA)

FREE STATE. — 2926 (Bloemfontein): 25 km from Bloemfontein to Winburg, (-AA)

EASTERN CAPE. — 3125 (Steynsburg): 10 km from Steynsburg to Hofmeyr, (-BC)

EASTERN CAPE. — 3125 (Steynsburg): 24 km from Steynsburg to Hofmeyr, (-BC)

WESTERN CAPE. — 3222 (Beaufort West): 5 km from Beaufort West, (-BC)

EASTERN CAPE. — 3225 (Somerset East): Kokskraal, Cookhouse, (-DB)

EASTERN CAPE. — 3324 (Steytlerville): 68 km from Uitenhage to Graaff-Reinet, (-DA)

EASTERN CAPE. — 3325 (Port Elizabeth): 30 km from Uitenhage to Graaff-Reinet, (-CD)

45/2 NORTH-WEST. — 2522 (Sanie): in the riverbed at Walersend, (-DB)

FREE STATE. — 2925 (Jagersfontein): Spitskop farmyard, Fauresmith, (-DA)

NORTHERN CAPE. — 3024 (Colesbcrg): 27 km from Verwoerddam to Venterstad, (-DA)

EASTERN CAPE — 3224 (Graaff-Reinet): 39 km from Jansenville to Graaff-Reinet, (-DA)

EASTERN CAPE. — 3224 (Graaff-Reinet): 15 km from Jansenville to Graaff-Reinet, (-DC)

EASTERN CAPE. — 3224 (Graaff-Reinet): 76 km from Patensie to Willowmore, (-DD)

27 NORTHERN CAPE. — 2824 (Kimberley) 1 km from Kimberley to Griekwastad, (-DA)

FREE STATE. — 2925 (Jagersfontein): 44 kin from Petrusburg to Kimberley, (-AB)

27+0- IB NORTHERN CAPE. — 2824 (Kimberley) I km from Kimberley to Griekwastad, (-DA)

Spies 5883

Spies 5645

Spies 5653, 5654

Spies 5646

Spies 5527

Spies 5538

Spies 5542, 5543

Spies 5659

Spies 5657

Spies 5650

Spies 5649

Venter 9286

Spies 5521

Spies 5662

Spies 5660

Spies 5576, 5577

Spies 5574, 5575

Spies 5849, 5850

Spies 5638

Spies 5508

Spies 5643, 5664

Spies 5847

Spies 5669

Spies 5587

Spies 5670

Spies 5487, 5489

Spies 5240

Spies 5236

Spies 5237

Spies 5215

Spies 5591

Spies 5594, 5676

Spies 5226

Spies 5232

Spies 5230

Spies 5531

Spies 5652

Spies 5512

Spies 5848

Spies 5584, 5585

Spies 5586

Spies 5488

Spies 5675

Spies 5231

Spies 5229

Spies 5497

Du Preez 2758

Spies 5581. 5583

Spies 5239

Spies 5238

Spies 5210

Spies 5513, 5514

Spies 5510

Spies 5517

Bothalia 28,1 (1998)

85

TABLE 2. — Meiotic chromosome behaviour of Cenchrus Lilians specimens showing voucher specimen no.; gametic chromosome no. (n); aver-

age frequency of univalents (I); frequency of chromosome laggards; percentage of cells studied containing anaphase I bridges; frequency

of micronuclei during telophase I. All ranges are included in brackets

Percentage of cells exhibiting chromosome abnormality

86

Bothalia 28,1 (1998)

TABLE 2. (cont.) — Meiotic chromosome behaviour of Cenchrus ciliaris specimens showing voucher specimen no.; gametic chromosome no. (n);

average frequency of univalents (I); frequency of chromosome laggards; percentage of cells studied containing anaphase I bridges; fre-

quency of micronuclei during telophase I. All ranges are included in brackets

cells in the particular meiotic stage have not been observed, or a complete meiotic analysis (involving at least 10 cells of the particular stage)

could not be concluded.

of the three chromosome abnormalities will be represen-

tative of all the tetraploid specimens. This process was

not repeated for the pentaploid and hexaploid specimens,

due to the limited number of specimens available.

The average numbers of univalents observed in the

specimens investigated were 1.5, 1.9, 4.0 and 2.8 respec-

tively for the n = 17, n = 18, n = 45/2, and n = 27 speci-

mens. The variation in the number of univalents is indi-

cated in Table 2. The highest number of univalents (0-18)

was observed for Spies 5230 (n = 18) (Figure 2B, C).

The univalents present were usually situated near the

equator (Figure 2A-D) and moved onto the plate where

they orientated themselves both syntelicly and amphi-

telicly. This led to the centromeres of the two chromatids

either undergoing reductional distribution or equational

distribution. The chromatids segregated to opposite

poles.

Since the B-chromosomes were mostly indistinguish-

able from the normal euchromosome complement, spec-

imens containing B-chromosomes were excluded during

the calculation of the univalent, laggard and micronuclei

averages.

Chromosome and chromatid laggards were frequently

observed during anaphase I (Figure 3A-E). The average

numbers of chromosome laggards observed, were 2.9,

1 .7, 9.2 and 3.3 for the n = 17, n = 18, n = 45/2, and n =

27 specimens, respectively (Table 2). The variation in

numbers is indicated in Table 2. The highest number of

laggards (1-32) was observed for Spies 5581 (n = 45/2).

The number of micronuclei observed during telophase

I varied (Figure 4A-D). The average numbers of

micronuclei observed were 0.9 (n = 17), 1.1 (n = 18), 4.1

(n = 45/2) and 3.8 (n = 27) (Table 2). The variation in

numbers is indicated in Table 2. The highest number of

micronuclei (11) was observed for Spies 5210 (n = 45/2)

(Figure 4B). The size of these micronuclei varied (Figure

4B, D).

Genome interpretation was performed on three

tetraploid specimens ( Spies 5215, 5240 and 5649). These

analyses revealed that the 2:2 model (Kimber & Alonso

1981) fitted the specimens to the greatest degree, with x-

values of 1 or approximately 1 (Table 3).

Anaphase bridges were observed in 26 specimens of

C. ciliaris (Table 2) (Figure 5A-C). The average number

of cells per specimen which contained bridges, varied

from nought to a maximum of 71.4% ( Spies 5652) (Table

2). The acentric fragment could most often be observed

(Figure 5 A, B), and varied in size among the various

specimens.

(3 Univalents

^Chromosome laggards

□ Micronuclei

y

X-1 0 2 4

X-2 0 1 2 3 4 5 6

Number of chromosomes undergoing meiotic abnormality

FIGURE I — Histogram, indicating

percentage of tetraploid cells

exhibiting univalents during

metaphase I, chromosome

laggards during anaphase I,

and micronuclei during telo-

phase I. X- 1 , number of chro-

mosomes involved in forma-

tion of univalents and chro-

mosome laggards; X-2, num-

ber of micronuclei observed

per cell.

Bothalia 28,1 (1998)

87

DISCUSSION

Basic chromosome numbers constitute the core of any

meiotic study, as they are essential for confirming the

presence of polyploidy. Cenchrus ciliaris has a basic

chromosome number of x = 9 and polyploidy is present.

Three polyploid levels have been observed, with the

most abundant being the tetraploids (82.9%). The penta-

FIGURE 2. — Variation in number of

univalents observed during

metaphase I in Cenchrus cil-

iaris. A, Spies 5240, n = 2x =

18, with two univalents; B,

Spies 5230, n = 2x = 18, with

two univalents; C, Spies

5230, n = 2x = 18, with eight

univalents; D, Spies 5514, n

= 3x = 27, with numerous

univalents. Scale bar: 10 pm.

and hexaploids were observed at much lower frequencies

(9.2% and 6.6% respectively).

Polyploidy is prominent in the plant kingdom

(Stebbins 1982). Polyploid levels for C. ciliaris , taking

the published aneuploid chromosome numbers into

account, include diploidy, triploidy, tetraploidy, penta-

ploidy, hexaploidy and nanoploidy. References for these

FIGURE 3. — Variation in number of

chromosome and chromatid

laggards observed during

anaphase I in Cenchrus cil-

iaris. A, Spies 5883, n = 17;

B, Spies 5508, n = 2x = 18;

C, Spies 5583, n = 5/2x =

45/2; D, Spies 5514, n = 3x =

27; E, Spies 5514, n = 3x =

27. Scale bar: 10 pm.

Bothalia 28,1 (1998)

FIGURE 4. — Variation in number

and size of micronuclei ob-

served during telophase I and

II in Cenchrus ciliaris. A,

Spies 5542 , n = 2x = 18, telo-

phase II, with 0-4 per cell; B,

Spies 5210 , n = 5/2x = 45/2,

telophase I, with nine micro-

nuclei; C, Spies 5583, n =

5/2x = 45/2, telophase I, with

minimum of four micro-

nuclei; D, Spies 5514, n = 3x

= 27, telophase I, with 10 mi-

cronuclei. Scale bar: 10 pm.

chromosome numbers are presented in Visser et al. (in

prep.). The wide range of polyploid levels and very low

frequency of diploids indicate that this species forms a

mature polyploid complex.

The meiotic chromosome behaviour of the polyploid

specimens varied from being normal to highly irregular,

depending on the number of genomes present and their

homology. Polyploids with an even number of genomes

are influenced less dramatically during meiosis than

those with an uneven number of genomes. Uneven poly-

ploid levels have more meiotic abnormalities, due to the

presence of an uneven number of sets of chromosomes

which complicates chromosome pairing. Genome

homology also plays an important role in the normality

of meiosis. It can affect chromosome pairing to such an

extent that from univalents to multivalents are formed.

In order to attest the proposed basic chromosome

number of x = 9 for C. ciliaris , a comparison was made

between the observed and the expected meiotic chromo-

some behaviour of each of the polyploid levels studied.

The presence of univalents during metaphase I was reg-

ularly observed. A variation in the number of univalents

present occurred within and among the polyploid levels

(Figure 2A-D). The highest average numbers of univa-

lents were observed in the pentaploid specimens (4.0),

whereas for the hexaploid specimens, they were 2.8

(Table 2). Approximately similar average numbers of

univalents were observed for the aneuploid and

tetraploid specimens (1.5 versus 1.9).

The increased averages of univalents present during

metaphase I for the pentaploids and the hexaploids were

expected, as a higher incidence of meiotic irregularities

is closely associated with uneven and higher polyploid

levels (Stebbins 1950). The variation in the number of

univalents observed within each of these polyploid lev-

els, indicates chromosomal differences among the speci-

mens.

The number of univalents present during metaphase I

corresponds with the numbers of chromosome laggards

and micronuclei observed (Figure 1). The data suggest

that the univalents observed during metaphase I were

mostly lagging during anaphase I (Figure 3A-E), and led

TABLE 3. — Genomic constitution of tetraploid Cenchrus ciliaris specimens, according to models of Kimber & Alonso (1981): voucher nos,

chiasma frequencies, relative x-values and the appropriate average sum of squares for each of the various models in brackets (M)

M

Bothalia 28,1 (1998)

89

FIGURE 5. — Some anaphase I bridges observed in Cenchrus ciliaris. A,

2x = 18. Scale bar: 10 pm.

to the formation of micronuclei during telophase I. The

average number of laggards was the highest for the pen-

taploid specimens (9.2 laggards per cell), followed by the

hexaploid and aneuploid specimens (3.3 and 2.9 respec-

tively) (Table 2). The number of chromosome and chro-

matid laggards varied among the specimens of the vari-

ous polyploid levels, emphasizing genetic differences

among specimens within each of the polyploid levels.

The average number of micronuclei also varied

among the specimens of each polyploid level (Table 2).

The micronuclei were not incorporated into the daughter

nuclei at the time of cell division (Figure 4A-D). The

highest average number was observed in the pentaploid

specimens (4.1), followed by the hexaploid (3.8),

tetraploid (1.1) and aneuploid (0.9) specimens respec-

tively. The highest number and greatest variation of

micronuclei were observed in the pentaploid specimens

(Figure 4B, C) (Table 2).

The highest average numbers of univalents, chromo-

some laggards and micronuclei have been observed in

the pentaploid specimens (Table 2). This polyploid level

was followed by the hexaploids, whereas for the

tetraploids and the single aneuploid specimen, the level

was approximately similar. The high incidence of meiot-

ic abnormalities in the pentaploid specimens was expect-

ed, due to their uneven polyploid levels. An average

number of 9.2 chromosomes lagged during anaphase I

and was representative of an entire genome. A basic

chromosome number of x = 9 for C. ciliaris , due to the

presence of a fifth genome lagging during anaphase I, is

hereby confirmed.

Genome variation in the tetraploid specimens was

confirmed during this study. Meiosis was normal in some

specimens ( Spies 5522), whereas for others, it was high-

ly irregular ( Spies 5230) (Table 2). The average numbers

of univalents present and chromosome laggards

observed, were almost similar (1.9 and 1.7, respectively).

These averages suggest two univalents observed during

metaphase I, lagging during anaphase I, and finally form-

ing a single micronucleus (1.1 per cell) (Table 2).

Genome interpretation of three tetraploid specimens

(Spies 5215, 5240 & 5649) revealed that the 2:2 model

(Kimber & Alonso 1981) agreed best with these speci-

mens, with x-values of 1 ( Spies 5215 & 5240) and 0.84

(Spies 5649) (Table 3). An x-value of 1 inferred two dis-

Spies 5652, n = 2x = 18; B, Spies 5594, n = 2x = 18; C, Spies 5574, n =

tinctly different sets of genomes, therefore. Spies 5215

and 5240 were classified as allotetraploids (for example

AABB). Although the 2:2 model fitted Spies 5649 the

best, a lower x-value of 0.84 was calculated (Table 3).

This specimen was classified as a segmental alloploid,

based on the lower x-value and the occasional presence

of quadrivalents (for example AAA’A').

The specimens’ genomic constitutions were attested

by their meiotic chromosome behaviour. The specimens

differed in respect of their meiotic behaviour.

Quadrivalents were observed in Spies 5215 and 5649,

whereas for Spies 5240, univalents and bivalents were

observed. The number of quadrivalents observed in the

two specimens varied. One to three quadrivalents were

observed for Spies 5649, whereas for Spies 5215, an

infrequent quadrivalent was observed.

Spies 5240 was confirmed as an alloploid (for exam-

ple AABB), based on the presence of bivalents only,

observed during metaphase I. The presence of an occa-

sional quadrivalent in cells of Spies 5215 and one to

three quadrivalents per cell in Spies 5649, justify seg-

mental alloploidy (for example AAA'A') for both these

specimens. However, the presence of genes, controlling

homoeologous chromosome pairing in Cenchrus, should

be studied before these genomic constitutions can be

accepted.

The two alloploid specimens were collected in the

Eastern Cape, whereas Spies 5649 was collected in the

Free State (Table 1). Taking their different geographical

localities into consideration, it is suggested that the

Eastern Cape specimens represent a genetically different

group (or hybrid swarm) from those in the Free State.

Therefore, chromosomal variation in these three

tetraploid specimens confirm the presence of genetic

variability in C. ciliaris in South Africa.

The highest incidences of meiotic abnormalities were

recorded in the pentaploid specimens (Table 2). These

abnormalities were mostly the result of an uneven poly-

ploid level. The average number of univalents observed,

was approximately half of the average number of chromo-

some laggards observed (4.0 versus 9.2) (Table 2). There

was an increase in the average number of laggards

observed (Table 2). This could have been due to amphitel-

ic orientation and equational distribution of the univalents

observed during metaphase I, as various chromatid lag-

90

Bothalia 28,1 (1998)

gards were observed during anaphase I (Figure 3C). These

anaphase I laggards were mostly included in more than

one micronucleus per cell (Figure 4B, C), as an average

frequency of 4.1 micronuclei per cell has been observed.

The average number of laggards observed for this poly-

ploid level, was representative of a complete genome lag-

ging during anaphase F It is suggested that the genomic

constitution of the pentaploid specimens includes one

unrelated genome. This suggestion is based on the average

number of univalents present, the absence of trivalents and

the high average number of laggards observed during

anaphase I. A genomic constitution of, for example

AAA'A'A", is proposed for this polyploid level. Segmental

alloploidy is justified by the high and the low occurrence

of bivalents and quadrivalents respectively, observed dur-

ing prophase and metaphase I. Chromosomal variation is

evident in this polyploid level, for meiotic chromosome

behaviour varied among the specimens studied (Table 2).

The average number of univalents present, the chro-

mosome and chromatid laggards and the micronuclei

observed for the hexaploid specimens (2.8, 3.3 and 3.8

respectively), were relatively low when compared to that

of the pentaploid specimens (Table 2). A genomic con-

stitution of, for example AAA'A’A"A", is proposed for

the hexaploids. Segmental alloploidy, based on the abun-

dant bivalent and occasional quadrivalent chromosome

configurations found (Figure 2D), is proposed. The pres-

ence of quadrivalents indicates a degree of homology

between the A and A' genomes.

The genomic constitutions of the various polyploid

levels (n = 17 — segmental alloploidy; n = 18 — segmen-

tal polyploidy to alloploidy; n = 45/2 — alloploidy and n

= 27 — segmental alloploidy) indicate the presence of

hybridisation in this species.

Hybridisation among plant individuals is usually

characterised by various changes in chromosome struc-

ture (Darlington 1937; Dobzhansky 1941). For C. cil-

iaris, these changes include the presence of paracentric

inversions. These inversions were mostly observed in the

tetraploid specimens (Figure 5A-C). The higher occur-

rence of these inversions could be due to the high num-

ber of tetraploid specimens which were cytogenetically

studied. The highest average number of cells containing

anaphase I bridges was observed in Spies 5652 (71.4%)

(Figure 5 A) (Table 2). Different paracentric inversions

were found among the specimens, as the chromosome

fragments differed in size.

Occasional or recurrent hybridisation and the com-

plete local breakdown of reproductive isolation between

sympatric species result in the production of hybrid

swarms. These swarms include the whole range of genet-

ic variability of the parental species. This scenario could

be representative of C. ciliaris, for a wide range of chro-

mosomal, morphological and genetic variation is evident

in this species.

CONCLUSIONS

With the aid of meiotic analyses, a basic chromosome

number of x = 9 has been confirmed for Cenchrus cil-

iaris. Polyploidy is common and varies from tetraploid

to hexaploid. Aneuploidy was observed in a single spec-

imen. It is suggested that this specimen is the result of

loss aneuploidy from two chromosomes of a single

genome.

Various meiotic irregularities were observed for this

species. The highest incidences of meiotic abnormalities

were observed for the pentaploid specimens. This was

attributed to their uneven polyploid and chromosome

number.

The chromosome abnormalities observed during

meiosis were an indication of genomic relationships.

These relationships varied among the specimens and the

polyploid levels. Segmental alloploidy was suggested

for the aneuploid specimen, whereas for the tetraploid

specimens, it varied from segmental alloploidy to allo-

ploidy. A genomic constitution of alloploidy and seg-

mental alloploidy is suggested for the pentaploid and

hexaploid levels respectively. The nature of the genom-

ic relationships indicated the presence of hybridisation.

Hybridisation in C. ciliaris was confirmed by the chro-

mosomal variation observed among specimens in each

of the polyploid levels.

ACKNOWLEDGEMENTS

The University of the Orange Free State and the

Foundation for Research and Development are thanked

for financial assistance during this study.

REFERENCES

CHAPMAN, G.P. 1990. Reproductive versatility in grasses.

Cambridge University Press, Cambridge.

DARLINGTON, C.D. 1937. Recent advances in cytology. Churchill,

London.

DE LISLE, D.G. 1963. Taxonomy and distribution of the genus

Cenclirus. Iowa State Journal of Science 37: 259-351.

DOBZHANSKY, T. 1941. Genetics and the origin of species.

Columbia University Press, New York.

EDWARDS, D. & LEISTNER, O.A 1971. A degree reference system

for citing biological records in southern Africa. Mitteilungen

der Botanischen Staatssammlung, Miinchen 10: 501-509.

KIMBER, G. & ALONSO, L.C. 1981. The analysis of meiosis in

hybrids. III. Tetraploid hybrids. Canadian Journal of Genetics

and Cytology 23: 235-254.

STEBBINS, G.L 1950. Variation and evolution in plants. Columbia

University Press, New York.

STEBBINS, G.L. 1982. Major trends of evolution in the Poaceae and

their possible significance. In J R Estes. R.J. Tyrl & J.N.

Brunken, Grasses and grasslands: systematics and ecology.

University of Oklahoma Press, Norman.

VISSER, N.C. & SPIES, ,I J 1994. Cytogenetic studies in the genus

Tribolium (Poaceae: Danthonieae). Ill Section Tribolium.

South African Journal of Botany 60: 31-39.

VISSER, N.C. SPIES, J.J. & VENTER. H J.T. in prep. Aneuploidy in

Cenchrus ciliaris (Poaceae, Panicoideae, Paniceae): truth or fic-

tion'? South African Journal of Botany.

Bothalia 28,1: 91-112 (1998)

Invasive alien woody plants of the southern and southwestern Cape

region, South Africa

L. HENDERSON*

Keywords: alien invasive plants, Forest Biome, Fynbos Biome, roadside survey, Savanna Biome, southern and southwestern Cape, Succulent

Karoo Biome

ABSTRACT

The frequency and abundance of invasive alien plants were recorded along roadsides and at watercourse crossings in

82.9% (145/175) of the quarter degree squares in the study area (31-35°S, 17-25°E and covering ± 90 000 km2). The sur-

vey yielded 102 species of which the most prominent (in order of prominence) in roadside and veld (natural and modified)

habitats were: Acacia cyclops, A. saligna and A. mearnsii. The most prominent species (in order of prominence) in stream-

bank habitats were: A. mearnsii, A. saligna and Populus x canescens.

The greatest intensity of invasion was recorded in forest and fynbos vegetation types in the relatively narrow belt

stretching from the coastline to the tops of the coastal mountain ranges. In the coastal lowlands Acacia cyclops and A.

saligna form the most extensive and continuous stands of alien vegetation recorded anywhere in South Africa. In the and

interior invasion was largely confined to watercourses.

CONTENTS

Introduction 91

Survey history and objectives 91

The study area 91

Method 94

Sampling method 94

Abundance ratings 94

Sampling level achieved 94

Data treatment— formulae used 94

Frequency 94

Prominence value 94

Mean species abundance rating in roadside and

veld habitats 95

Mean abundance of invaders per km in roadside

and veld habitats 95

Results 95

The streambank habitat 95

The whole study area 95

Analysis according to veld type 95

Analysis according to species 95

Frequency 95

Prominence 95

Roadside and veld habitats 97

The whole study area 97

Analysis according to veld type 97

Analysis according to species 99

Frequency 99

Prominence 99

Patterns of invasion 99

Discussion 99

Prominent and potentially important species .... 99

Relation of invasion to historical and environmen-

tal factors 109

Some ideas for the future 110

Acknowledgements 110

* Plant Protection Research Institute, Agricultural Research Council;

stationed at National Botanical Institute, Private Bag X101, Pretoria

0001.

MS received: 1997-07-10.

References 110

Appendix Ill

INTRODUCTION

Survey history and objectives

This study of the southern and southwestern Cape is

the sixth of eight regional surveys which together are

designed to reflect invasion by woody alien plants in

South Africa as a whole. Surveys have been completed

for the former Transvaal (Henderson & Musil 1984),

Natal and northeatem Orange Free State (Henderson

1989), Orange Free State (Henderson 1991a), northern

Cape (Henderson 1991b) and eastern Cape (Henderson

1992). The survey of this area was undertaken from east

to west and during the months of March and October

1990, January and December 1991, February and

November 1992, and May 1993.

The objectives of the survey were: to produce a check-

list of the major invasive alien woody plants of stream-

bank, roadside and veld habitats in the study area; to

determine the pattern of alien woody invasion as a whole

and for individual species; to attempt to relate distribution

to environmental factors and to determine which are the

most prominent and potentially important invaders.

The study area

The study area lies between latitudes 31° and 35°S

and longitudes 17° and 25°E (Figure 1 ), and occupies an

area of approximately 90 000 km2. It is bounded in the

south by the Indian Ocean and in the southwest by the

Atlantic Ocean. The topography is dominated by the

mountains of the Cape Fold Belt. These mountains occur

for the most part in subparallel ranges with an average

height of 1 000 to 1 500 m and with individual peaks

reaching over 2 000 m. In the south these ranges strike

from east to west whereas in the west the strike is more

92

Bothalia 28,1 (1998)

FIGURE 1. — The study area showing survey routes and intensive sites.

nearly north-north-west (Taylor 1978). The ranges are

separated by wide intermontane valleys. There are many

river systems in the study area which contains about 17%

of the major mountain catchments of South Africa,

Lesotho and Swaziland (Van der Zel 1981).

Rainfall ranges from 200 mm in the arid interior to

3 600 mm per annum on some of the higher mountains.

To the west of 20°E a Mediterranean-type climate pre-

vails with cool wet winters and warm dry summers. In

the east, rainfall is more consistent through the year.

FIGURE 2. — The biomes in the study area after Rutherford & Westfall (1986).

Bothalia 28,1 (1998)

93

TABLE 1. — Veld type categories in study area and equivalent Acocks Veld Type groupings and Veld Type numbers

* according to Henderson; f according to Rutherford & Westfall 1986.

Changes in the rainfall can be very abrupt, and are usu-

ally associated with the topography. During winter, snow

falls regularly on the higher mountains, but the lowlands

enjoy an equable climate and frost is rare except in some

of the deep valleys of the interior (Taylor 1978; Linder

1991; Jury 1993).

For the purposes of this survey and in keeping with

previous surveys, the vegetation of the study area has

been subdivided according to the biomes of southern

Africa defined by Rutherford & Westfall (1986) and

Acocks’s Veld types of South Africa (1988). There are

four biomes in the study area, namely the Succulent

Karoo, Savanna, Forest and Fynbos Biomes (Figure 2).

Eleven Acocks Veld Types occur in the study area and

have been grouped into nine broad veld type categories

for the purposes of this survey (Table 1; Figure 2).

Succulent karoo occupies the region commonly

known as the Little Karoo. The vegetation has a predom-

inance of succulents, and dwarf trees and shrubs are

numerous. It occupies rocky, hilly country, at elevations

ranging from 300-600 m. receiving 150-300 mm of rain

per annum (Acocks 1988).

Savanna is represented by a vegetation type common-

ly known as succulent mountain scrub which occurs on

steep mountain slopes. It is typically a dense scrub dom-

inated by Portulacaria afra. The altitude ranges from

400-1 060 m, and rainfall ranges from 250-300 mm per

annum (Acocks 1988; Low & Rebelo 1996).

The only forest area of biome dimension in South

Africa occurs in the southern Cape at Knysna

(Rutherford & Westfall 1986). Before the southern Cape

forests were heavily exploited and reduced to their pre-

sent remnant areas, they covered most of the southern

Cape coastal region between Mossel Bay and

Humansdorp (Von Breitenbach 1972). Today the Knysna

Forest and many smaller forest patches occur in a nar-

row, irregular belt along the southern coastal shelf and

foothills of the Outeniqua and Tsitsikamma Mountains

(Von Breitenbach 1972). Forest growth is favoured by

mild temperatures and a high, well-distributed rainfall

(Von Breitenbach 1972). The mean annual rainfall is

800-1 000 mm (Schulze & McGee 1978), but ranges

from 500-1 200 mm (Geldenhuys 1993). Altitude ranges

from sea level to 1 220 m at the tops of the mountains

only 10-37 km inland (Geldenhuys 1993). Small forest

outliers occur in fire-free habitats westwards towards the

Cape Peninsula (Taylor 1978).

Strandveld is a low-growing, semisucculent and

shrubby vegetation type confined to the sandy coastal

plains of the west coast. It is transitional between coastal

fynbos and succulent karoo. It receives 50-300 mm of

rain per annum, mainly in winter (Acocks 1988).

Mountain renosterveld is a karroid-like veld type

dominated by renosterbos, Elytropappus rhinocerotis

(Acocks 1988). It is usually situated between succulent

karoo and mountain fynbos and receives 250-400 mm of

rain per annum (Low & Rebelo 1996).

Coastal renosterveld occurs in a western and southern

belt and is situated between the fynbos of the mountains

and of the coastal plain. The terrain is undulating and the

soils are clayey. Rainfall ranges from 300-600 mm per

annum. Most of the original vegetation has been

ploughed up for the growing of wheat. Scrub relics that

remain are dense, thorny and semisucculent. Elsewhere

the scrub has been replaced mainly by renosterbos

(Acocks 1988).

Coastal fynbos occurs on sand and limestone in the

west and south coastal belts. The vegetation is a more or

less open scrub lacking the dense thorniness and semi-

succulence of the previous veld type. The altitude ranges

from 0-300 m and rainfall from 300-500 mm per annum

(Acocks 1988).

Mountain fynbos is the most widespread vegetation

type in the Fynbos Biome, occurring mainly along the

Cape Fold Belt from north of Nieuwoudtville to Cape

Town and Cape Agulhas and to Humansdorp in the east.

Altitude ranges from 0-2 200 m. and rainfall from 200 to

more than 2 000 mm per annum, occurring mainly in the

winter months (Low & Rebelo 1996). Summers are hot

and dry and conducive to veld fires. The vegetation is an

open to closed shrubland.

94

Bothalia 28,1 (1998)

Mountain fynbos & forest occurs outside the Forest

Biome defined by Rutherford and Westfall (1986) but

within the original limits of the Knysna Forest, Acocks

Veld Type 4, mapped by Acocks ( 1 988). This region con-

tains wet mountain fynbos (National Committee for

Remote Sensing, CSIR 1983), numerous small patches

of forest, including the Tsitsikamma Forest, and exten-

sive timber plantations of mainly pine, but also gums and

Acacia melanoxylon (Van der Zel 1988). The mean annu-

al rainfall is 800-1 000 mm (Schulze & McGee 1978)

and uniformly distributed throughout the year.

METHOD

Sampling method

The method used in this survey was the same as in the

previous survey of the eastern Cape (Henderson 1992).

The presence and abundance of all alien trees, large

shrubs and conspicuous climbers which appeared to be

spreading spontaneously (naturalised) were recorded for

each veld type category, habitat type (roadsides and

adjoining veld, and streambanks) and quarter degree/fif-

teen minute square traversed by road. Thirty three quar-

ter degree squares were selected for more intensive sur-

veying (Figure 1). They may be used at a later date for a

quick resurvey of the study area to assess any changes

that may have taken place.

Recordings of roadside and veld invaders were made

from a moving vehicle along road transects of between

five and 10 km in length. The average transect length

was 6.4 km for the general survey area and 5.0 km for the

intensive sites. Recordings of streambank invaders were

made at virtually all watercourse crossings on the survey

route. Details of the roads traversed are lodged in the

Plant Protection Research Institute (PPRI), Pretoria. As

on the previous occasions the survey was undertaken in

a minibus, with one driver and one recorder (the author).

The average speed was 60 km/h but ranged from about

20 km/h in densely vegetated areas to 100 km/h in

sparsely vegetated areas.

All the raw data for this survey as well as the previ-

ously completed regional surveys have been computer-

ized using the data management system DataEase. This

database is housed in Pretoria at the PPRI.

Abundance ratings

The abundance ratings for roadside and veld habitats

and streambank habitats are given in Table 2.

Sampling level achieved

The sampling level achieved was 82.9% (145 out of

the total 175 quarter degree squares) at an average of

44.5 km travelled per square. An average of 29.3 km of

road transects were sampled per quarter degree square

for abundance estimates of roadside and veld invaders.

The veld type coverage in terms of quarter degree

squares and road transects sampled, kilometres travelled

and watercourse recordings made, is given in Table 3.

Statistics for streambank, roadside and veld habitats are

given in Tables 4 and 5.

Data treatment — formulae used

Frequency

The percentage frequency of occurrence of a species

x in a given category (veld type, biome or study area) y

was calculated as follows:

no. of watercourse recordings/road transects

in category y having species x

frequency = : — r ; ; x 100

total no. ot watercourse recordings/road

transects in category y

Prominence value

The prominence value is a combined measure of a

species’ frequency and abundance relative to that of all

TABLE 2. — Abundance ratings

* approximate numbers of individuals or groups per 10 km transect.

Bothalia 28.1 (1998)

95

other species, within a given vegetation category (veld

type, biome or study area).

In streambank habitats the prominence value for a

species x in category y was calculated as follows:

total weighted abundance of species x

sum of the weighted abundances of

all species in category y

prominence value = +

frequency of species x in category y

sum frequency of all species in

category y

The abundance ratings were weighted according to

the minimum percentage cover in each scale rating (see

Table 2). Thus ratings 7. 6, 5 and 4 had weighted values

of 75, 50, 25 and 5 respectively. Ratings 1, 2 and 3 each

had weighted values of 1.

In roadside and veld habitats the prominence value for

a species x in category y was calculated as follows:

RESULTS

The survey yielded 102 naturalised alien species.

These species are listed in the Appendix together with a

further 34 species which were obtained from various lit-

erature and other sources. The distributions of 20 of the

most prominent species are given in Figures 6 and 7 and

a further 10 potentially important species are given in

Figure 8.

The streambank habitat

The whole study area

One thousand and thirty-six watercourse crossings

were sampled in which 75 species were recorded, with

up to eight species in one sample. Invaders were present

at 73.5% of all crossings and 30.9% of all crossings were

heavily invaded (Table 4).

Analysis according to veld type

total abundance* of species x in

category y

sum of the abundances* of all

species in category y

prominence value = +

frequency of species x in category y

sum frequencies of all species in

category y

The highest prominence values in a given category

which add up to approximately 160 points out of a total

of 200 are printed in bold in Tables 6, 7, 8 and 9. The cut-

off point of 160 points is arbitrary but represents 80% of

the summed prominence values.

Mean species abundance rating in roadside and veld

habitats (Tables 8 & 9)

The mean species abundance rating** of a species x

in a given category (veld type, biome or study area) y

was calculated as follows:

x 100

X 100

mean no. of

individuals

or groups

per 10 km

total no. of individuals or groups of species x

in category y

total distance along which species x was

rated in category y

Mean abundance of invaders per km in roadside and

veld habitats (Table 5)

The mean abundance of invaders per km in a given

category (veld type, biome or study area) y/quarter

degree square z was calculated as follows:

Overall the Fynbos Biome was the most heavily

invaded in terms of percentage crossings invaded and

percentage crossings heavily invaded. Invasion was most

intense in mountain and coastal fynbos where the highest

percentages of heavily invaded crossings were recorded.

Very few watercourses were sampled in the Forest

Biome. Most recordings were of small streams and it was

difficult to see beyond 10 m.

Analysis according to species

Frequency

Acacia mearnsii was the most frequently recorded

invader in the study area (29.4%). Only this species, A.

saligna (20.7%) and Populus x canescens (13.6%) were

recorded at 10% or more crossings in the whole study

area (Table 7).

In the Fynbos Biome the most frequently recorded

species were Acacia mearnsii (40.4%), A. saligna

(30.7%), Populus x canescens (17.1%) and A. cyclops

(11.2%). In the Forest Biome A. melanoxylon (70.6%),

A. mearnsii (35.3%) and Eucalyptus diversicolor

(23.5%) were the most frequent invaders. In the Savanna

Biome Nicotiana glauca (14.1%), Arundo donax

(11.1 %), Populus x canescens (11.1%) and Ricinus com-

munis (11.1%) were the most frequently recorded

species. In the Succulent Karoo Biome Acacia saligna

(15.6%) and Nicotiana glauca (15.6%) were the most

frequent invaders.

total abundance* of all species in category y/quarter

degree square z

total kilometres rated for abundance estimates in

category y/quarter degree square z

* each abundance rating was expressed in numbers of individuals or

groups recorded per transect (Table 2). To be both conservative and

consistent the minimum number was used in each instance, e.g. an

abundance rating of 5 over 10 km = 50 and an abundance rating of

5 over 5 km = 25.

** mean no. of individuals or groups per 10 km converted to rating

(Table 2).

Prominence

The most prominent invader in the whole study area

was Acacia mearnsii with a prominence value of 61.6

out of a combined total for all species of 200 (Table 7).

The next most prominent invaders were A. saligna (36.9)

and Populus x canescens (19.5).

In the Fynbos Biome Acacia mearnsii was the most

prominent invader in four of the six veld type categories,

namely mountain fynbos, mountain fynbos & forest,

96

Bothalia 28,1 (1998)

TABLE 3. — Sampling coverage of each biome, veld type category and study area

* this represents the distance along which abundance recordings were made. Total distance along which observations were made is approximately

one and a half times that given; * according to Henderson; * according to Rutherford & Westfall 1986.

TABLE 4. — Statistics for streambanks in each biome, veld type category and study area

* one or more species scored an abundance rating of 5 or more; ** invaders present; According to Henderson; * according to Rutherford &

Westfall 1986.

TABLE 5. — Statistics for roadside and veld habitats in each biome, veld type category and study area

* one or more species scored an abundance rating of 5 or more; ** see data treatment-formulae used; * according to Henderson; * according to

Rutherford & Westfall 1986.

Bothalia 28,1 (1998)

97

TABLE 6. — Alien species occurring in streambank habitats of Succulent Karoo, Savanna and Forest Biomes and in strandveld and mountain

renosterveld of Fynbos Biome

F, % frequency of occurrence; I, % crossings heavily invaded; P, prominence value; * species occurring in the given category but not included

in a formal recording at a watercourse crossing. Bold numbers: the highest prominence values in a given category which add up to ± 80% of the

summed values (see text).

coastal renosterveld and mountain renosterveld. A.

saligna was the most prominent invader in the remaining

two categories, namely strandveld and coastal fynbos. A.

cyclops was a close second in coastal fynbos.

Acacia melanoxylon was the most prominent invader

in the Forest Biome, followed by Acacia mearnsii.

Populus x canescens was the most prominent invader in

the Savanna Biome and A. saligna was the most promi-

nent invader in the Succulent Karoo Biome.

Roadside and veld habitats

The whole study area

One hundred and forty-five quarter degree squares

and 660 road transects were sampled in which 96 species

were recorded. Up to 45 species were recorded per quar-

ter degree square. Naturalised species were recorded in

93.8% of all transects sampled and 45.8%’ of all transects

were heavily invaded (Table 5).

Analysis according to veld type

The most intense invasion was recorded in mountain

fynbos & forest in the Fynbos Biome, closely followed

by the Forest Biome where the highest percentages of

transects (96.1% and 88.9% respectively) were heavily

invaded. The mean abundance of invaders per km

reached a maximum in coastal fynbos (29.8). The least

invasion was recorded in the Savanna Biome, where no

transects were heavily invaded.

It must be noted here that although invasion was intense

in the Forest Biome, most invasion was recorded in the dis-

98

Bothalia 28,1 (1998)

TABLE 7. — Alien species occurring in streambank habitats in coastal renosterveld, coastal fynbos, mountain fynbos, mountain fynbos & forest

of Fynbos Biome and study area

Bothalia 28,1 (1998)

99

TABLE 7 (cont.). Alien species occurring in streambank habitats in coastal renosterveld, coastal fynbos, mountain fynbos, mountain fynbos &

forest of Fynbos Biome and study area

F, % frequency of occurrence; I, % crossings heavily invaded; P, prominence value; * species occurring in the given category but not included

in a formal recording at a watercourse crossing. Bold numbers: the highest prominence values in a given category which add up to ± 80% of the

summed values (see text).

turbed areas between the remaining indigenous forest

patches. Only small sections of forest are accessible by

road. The following species were recorded along roadsides,

margins or other gaps in indigenous forest: Acacia meam-

sii, A. melanoxylon, Cinnamomum camphora, Cortaderia

selloana, Eucalyptus diversicolor, Pinus pinaster, P. radia-

ta, and Rubus fruticosus. They were never abundant but

occurred as single plants or small groups.

Most of the higher parts of the mountain fynbos were

inaccessible by road and therefore undersampled in this

survey. As a consequence the mountain species such as

Hakea spp., particularly H. sericea, and Pinus spp. were

no doubt under-recorded in this survey. The accessible

parts of mountain fynbos were the valleys and this is

where most recordings were done. The best data on the

extent of woody plant invasions in the higher altitude

areas can be obtained from studies of the Cape Peninsula

mountains (Moll & Trinder-Smith 1992; Richardson etal.

1996).

Analysis according to species

Frequency

The most frequently recorded species in the whole

study area were Opuntia ficus-indica (71.5%), Acacia

saligna (67.0%), A. cyclops (63.6%), Nicotiana glauca

(63.6%), A. meamsii (52.9%), Pinus pinaster (38.8%)

and A. melanoxylon (33.5%).

The most frequently recorded species in the Fynbos

Biome wer e Acacia cyclops, A. saligna and A. mearnsii.

In the Forest Biome A. melanoxylon, A. mearnsii, Pinus

pinaster and P. radiata were the most frequent species.

In the Savanna Biome Nicotiana glauca, Opuntia ficus-

indica and Agave americana were the most frequent

species. In the Succulent Karoo Biome Nicotiana glau-

ca, Opuntia ficus-indica and A. saligna were the most

frequent species.

Prominence

Acacia cyclops scored the highest prominence value

of 45.2 in the study area. The next most prominent

species were Acacia saligna (24.8) and A. mearnsii

(24.0) (Table 9).

In the Fynbos Biome, the three aforementioned

species were the most prominent invaders, followed by

Pinus pinaster. In the Forest Biome Acacia melanoxylon

was the most prominent invader followed by A. mearn-

sii, Pinus pinaster and P. radiata. In the Savanna Biome

Nicotiana glauca and Opuntia ficus-indica were the

most prominent species. In the Succulent Karoo Biome

A. saligna was the most prominent species.

Patterns of invasion

Alien plant invasion was recorded in streambank,

roadside and veld habitats throughout the southern and

southwestern Cape (Figures 3, 4 & 5). Most invasion,

however, was encountered within the relatively narrow

belt stretching from the coastline to the tops of the

coastal mountain ranges.

A comparison of Figures 3 and 4 shows that similar

patterns of invasion were recorded in streambank, road-

side and veld habitats except that in the dry inland areas

of the Little Karoo, centred around Ladismith and Oudts-

hoorn, there was more severe invasion of the streambank

habitat than of roadside and veld habitats.

DISCUSSION

Prominent and potentially important species

Acacia species were overall the most prominent

invaders in the study area with one or more species being

the most prominent in every vegetation category with the

100

Bothalia 28,1 (1998)

TABLE 8 — Alien species occurring in roadside and veld habitats of Succulent Karoo, Savanna and Forest Biomes and in strandveld and moun-

tain renosterveld of Fynbos Biome

F, % frequency of occurrence; A, mean abundance rating; P, prominence value; * species occurring in the given category but not included in a

formal recording in a road transect. Bold numbers: the highest prominence values in a given category which add up to ± 80% of the summed values

(see text).

Bothalia 28.1 (1998)

101

TABLE 9. — Alien species occurring in roadside and veld habitats in coastal renosterveld, coastal fynbos, mountain fynbos, mountain fynbos &

forest of Fynbos Biome and study area

102

Bothalia 28,1 (1998)

TABLE 9 (cont.). — Alien species occurring in roadside and veld habitats in coastal renosterveld, coastal fynbos, mountain fynbos, mountain

fynbos & forest of Fynbos Biome and study area

F, % frequency of occurrence; A, mean abundance rating; P, prominence value; * species occurring in the given category but not included in a

formal recording in a road transect. Bold numbers: the highest prominence values in a given category which add up to ± 80% of the summed values

(see text).

FIGURE 3. — Invasion in stream-

bank habitats in terms of the

intensity of invasion of

watercourse crossings and

species diversity per quarter

degree square.

Bothalia 28,1 (1998)

103

FIGURE 4. — Invasion in roadside

and veld habitats in terms of

the intensity of invasion of

road transects and species

diversity per quarter degree

square.

exception of savanna. Acacia meamsii, A. saligna and A.

cyclops were the top three most prominent invaders with

their combined prominence values amounting to 201 out

of a total of 400 points for all species. In streambank

habitats the Acacia species were dominant with five

species (A. meamsii, A. saligna, A. cyclops, A. longifolia,

A. melanoxylon) amongst the top six most prominent

species. In roadside and veld habitats four of the top six

most prominent species were Acacia species (A. cyclops,

A. saiigna, A. meamsii and A. melanoxylon).

Acacia meamsii (Figure 6D) was a prominent invad-

er in all vegetation categories with the exception of

savanna, strandveld and coastal fynbos. It was recorded

predominantly along watercourses and was the most

prominent riverine invader in the Fynbos Biome and

the whole study area. Unlike in Mpumalanga (of the

former Transvaal) and KwaZulu-Natal, where it has

been cultivated on a grand scale for tannin, it has been

cultivated only on a small scale in the Cape, yet has

spread widely. This is a reflection of its aggressiveness

as an invader.

Acacia saligna (Figure 6G) was a prominent invader

in all vegetation categories with the exception of savanna

and forest. It was the next most prominent riverine invad-

er in the study area after A. meamsii and unlike A. meam-

sii was prominent in strandveld and coastal fynbos. It was

the second most prominent invader of roadsides and veld

in the Fynbos Biome, after A. cyclops. Together with A.

cyclops it formed extensive stands on the Cape Flats,

where it was used for the stabilization of driftsands and

was grown for tannin from 1850 until 1910 (Stirton

1978). It was most abundant in a broad band stretching

from Bredasdorp northwestwards to Clanwilliam. It

scored a highest abundance rating of 8 in the vicinity of

Robberg (Plettenberg Bay) and Bredasdorp. A gall-form-

ing rust fungus Uromycladium tepperianum, introduced

for the biocontrol of A. saligna (Morris 1991), was only

noted as abundant north of Cape Town between Atlantis

and Kalkbaskraal in May 1993. Four years later, in 1997,

it is abundant throughout the range of A. saligna.

Acacia cyclops (Figure 6A) was the most prominent

invader in roadside and veld habitats in the Fynbos

FIGURE 5. — Invasion in roadside

and veld habitats in terms of

the mean abundance of in-

vaders per kilometre in each

quarter degree square.

104

Bothalia 28,1 (1998)

FIGURE 6. — Distribution of the most prominent species: A, Acacia cyclop s\ B, A. dealbata ; C, A. longifolia ; D, A. mearnsii ; E, A. melanoxylon ;

F, A. pycnanthcr, G, A. saligna ; H, Arundo donax\ I, Eucalyptus diversicolor, J, £. lehmannii. Highest abundance rating of 4 or less, O;

highest abundance rating of 5 or more, •

Bothalia 28,1 (1998)

105

FIGURE 7. — Distribution of the most prominent species: A, Hakea sericecr, B, Leptospermum laevigatum ; C, Nicotiana glauca', D, Opuntia ficus-

indica\ E, Pinus pinaster-, F, P radiata', G, Populus x canescens ; H, Ricinus communis ; I, Rubus fruticosus', J, Sesbania punicea. Highest

abundance rating of 4 or less, O; highest abundance rating of 5 or more, •.

106

Bothalia 28,1 (1998)

FIGURE 8. — Distribution of less prominent and potentially important species: A, Acacia elata\ B, Eucalyptus cf. camaldulensis ; C, Hakea dru-

pacea, D, H. gibbosw, E, Myoporum tenuifolium\ F, Nerium oleander', G, Paraserianthes lophantha', H, Pinus halepensis', I, Quercus robur,

J, Spartium junceum. Highest abundance rating of 4 or less, O; highest abundance rating of 5 or more, •.

Bothalia 28,1 (1998)

107

Biome and in the whole study area. It was prominent

along watercourses only in coastal fynbos. It formed

very extensive stands on the Cape Flats, where it was

established as a sand-binder from 1876-1886 (Stirton

1978). In the vicinity of Mitchell’s Plain, Strandfontein

and Muizenberg it formed almost pure stands for dis-

tances up to 10 km or more, thereby scoring the maxi-

mum abundance rating. It also scored the maximum rat-

ing in the vicinity of Cape Agulhas, Struisbaai and Quoin

Point. It was exceedingly abundant (scoring an abun-

dance rating of 7 or more) in coastal areas stretching all

the way from Saldanha Bay in the west to Humansdorp

in the east, a distance of approximately 1 000 km.

Acacia melanoxylon (Figure 6E) was concentrated in

the forested area defined by the limits of Acocks’s Veld

Type 4, Knysna Forest. It was exceedingly abundant and

the most prominent invader in the Forest Biome. The

only other areas where it was recorded as abundant were

in the vicinity of Wolseley and in the Riviersonderend

Valley near Greyton. The abundance of this species

around Knysna can be attributed to its use as a commer-

cial timber tree and its consequent spread. It was used

extensively from 1856 as a forest replacement species in

the Knysna Forest, where it was planted in gaps created

by the felling of indigenous trees (Stirton 1978). The

indigenous forest pioneer tree, Virgilia oroboides was a

widespread weed of roadsides and probably provides

strong competition for A. melanoxylon and A. meamsii.

Acacia longifolia (Figure 6C) was concentrated in a

broad belt from Bredasdorp westwards to Cape Town

and northwards as far as Ceres. Two other foci occurred

eastwards around Knysna and Klipdrif, east of Storms

River. It was absent from succulent karoo, savanna,

strandveld and mountain renosterveld. In the remaining

wetter vegetation categories, particularly coastal and

mountain fynbos, it was more prominent in streambank

habitats than roadside and veld habitats. Overall for the

Fynbos Biome, it rated fifth and sixth most prominent in

streambanks and roadsides/veld respectively. Galled

plants, caused by the introduced wasp Trichilogaster

acaciaelongifoliae for biocontrol (Dennill & Donnelly

1991), were very much in evidence throughout its distri-

bution.

The biocontrol programme against A. longifolia has

reduced seed production and the overall biomass of pop-

ulations; in some instances it has caused mortality of

adult trees (Dennill & Donnelly 1991). Now there also

appears to be a reduction in the extent and abundance of

this invader. Comparison of the distribution of A. longi-

folia in Richardson et al. (1992), which was based on

reports from the mid 1980’s, and with the present survey,

shows a marked reduction in the extent and abundance of

A. longifolia in the southern Cape.

Acacia dealbata (Figure 6B) was virtually restricted

to watercourses in mountain fynbos. Although recorded

at very few localities it was almost always abundant,

forming dense stands. These localities were: Bergkloof

River, near Herbertsdale; Uniondale Poort, Holdrif

River; Riviersonderend; Dwars River tributary, near

Stellenbosch. Acacia pycnantha (Figure 6F) was restrict-

ed to the Fynbos Biome and was most prominent in road-

sides and veld habitats in mountain fynbos. It was

exceedingly abundant i.e. scoring a 7 in the vicinity of

Wolseley.

Populus x canescens (Figure 7G) was a widespread

invader of watercourses and was recorded in all vegeta-

tion categories with the exception of the Forest Biome. It

was the third most prominent riverine invader in the

whole study area and in the Fynbos Biome.

Pinus pinaster (Figure 7E) was a prominent invader

of mountain slopes in both the Forest and Fynbos

Biomes. It has been used extensively for afforestation

and its distribution is a reflection of planting patterns

(Stirton 1978). Pinus radiata (Figure 7F) has been plant-

ed on a smaller scale and was less prominent than P.

pinaster. These two species were the third and fourth

most prominent invaders of roadside and veld habitats in

the Forest Biome. They are the most prevalent species on

the Cape Peninsula mountains (Moll & Trinder-Smith

1992).

Rubus fruticosus (Figure 71) occurred mainly in the

wetter vegetation types. It was most prominent in moun-

tain fynbos & forest where it formed thickets along road-

sides and on the margins of plantations or forest. In pre-

vious publications (Henderson 1989; Henderson 1992)

reference was made to R. affmis. This is one of about

2 000 species belonging to the Rubus fruticosus complex

(Tutin et al. 1968) and was regarded by Spies & Du

Plessis (1985) to occur in South Africa. In this publica-

tion reference is made only to R. fruticosus L. aggregate

which is in keeping with the Pretoria National

Herbarium.

Hakea sericea (Figure 7 A) was only rated tenth most

prominent invader in mountain fynbos. This is an under-

estimate and resulted from the undersampling of moun-

tain slopes. From a distance Hakea spp. are also more

difficult to distinguish from the fynbos than for example

the Pinus spp., which invade the same habitats.

Richardson et al. (1992) gives a very different picture of

the distribution and abundance of H. sericea , which they

said occurred in 30% of the quarter degree squares in the

Fynbos Biome and formed dense stands in 19% of

squares. These records are based on Macdonald et al.

(1985) but updated from various sources (D.M.

Richardson pers. comm.).

Eucalyptus spp. were prominent invaders of the

Forest Biome and mountain fynbos & forest. E. diversi-

color (Figure 61) has been cultivated commercially

around Knysna and was the most prominent species in

this region. It was often difficult to distinguish the

Eucalyptus spp. and then they were recorded collective-

ly. In the Knysna area it was not clear how abundant E.

cladocalyx was. It was, however, regarded as one of the

three problem species in the area, together with E. diver-

sicolor and E. microcorys (forester at Woodville State

Forest pers. comm.).

Eucalyptus lehmannii (Figure 6J) was the third most

prominent invader in roadsides and veld habitats in

coastal fynbos. It was most abundant in the coastal belt

stretching from Bredasdorp westwards to the Cape

108

Bothalia 28,1 (1998)

Peninsula. E. cf. camaldulensis (Figure 8B) was mainly

an invader of watercourses and was most prominent in

the Succulent Karoo Biome along the Bree River. It was

also abundant along some watercourses in the Fynbos

Biome, such as the Olifants River near Clanwilliam,

Riviersonderend and Berg Rivers. E. exserta, which

could be mistaken for E. camaldulensis was abundant on

the Berg River near Langebaan and locally prominent on

the Bree River near Robertson.

Leptospermum laevigatum (Figure 7B) was the sixth

most prominent invader in roadside and veld habitats in

coastal fynbos. It has been used mainly as a hedge or

windbreak in coastal towns from where it has spread into

the adjacent fynbos. It was most abundant on sandy soils

from Bredasdorp westwards to Fish Hoek on the Cape

Peninsula. It was exceedingly abundant around Herma-

nus and Kleinmond.

Sesbania punicea (Figure 7J) was recorded in all veg-

etation categories except savanna, strandveld and moun-

tain renosterveld. It was most abundant along water-

courses and in particular the Bree River in the Succulent

Karoo Biome near Worcester. This species could have

been underestimated in this survey as it is easily over-

looked when not in flower.

Opuntia ficus-indica (Figure 7D) was by far the most

widespread invader in the study area, being recorded in

71.5% of all road transects. It rated as the fifth most

prominent invader in roadside and veld habitats in the

study area but it was rarely abundant. It was only in the

dry coastal bush around Mossel Bay that it scored an

abundance rating of 5; elsewhere it occurred only as scat-

tered individuals or small clumps.

Nicotiana glauca (Figure 7C) was the second most

widespread invader, being recorded in 63.6% of all road

transects. It was only recorded in disturbed sites such as

along roads, railway lines, river banks, dry river beds,

and in quarries and rubble heaps. It was a prominent

invader of watercourses in the Succulent Karoo and

Savanna Biomes but was never abundant. The only

places where it was recorded as very abundant was on

the roadside and floodplain adjacent to the Verlorevlei,

north of Piketberg and near Elandsbaai.

Arundo donax (Figure 6H) was a widespread invader

and was absent only from strandveld and forest. It was a

prominent invader of watercourses in coastal renoster-

veld, succulent karoo and savanna. Flowering plants

were only seen in the coastal belt near Humansdorp,

Knysna and Wellington.

Ricinus communis (Figure 7H) was a widespread

invader of roadside, veld and streambank habitats and

was prominent in coastal renosterveld, savanna and suc-

culent karoo. It was recorded in all vegetation categories

but was only abundant in the Kogmanskloof near

Montagu and in the Kafferkuilsrivier valley near Stilbaai.

Agave americana was prominent in succulent karoo

and savanna but was never abundant. Its distribution is a

reflection of where it has been planted. It appeared to

have spread from seed in two localities within the succu-

lent karoo. These were north of Barrydale between the

Anysberg and Warmwaterberg and in the Touws River

valley south of Ladismith.

Atriplex nummularia was most frequently recorded in

roadside and veld habitats in the succulent karoo, but it

was never abundant. Judging from observations in the

Great Karoo in the central Cape (Henderson in prep.) it

is likely to become a prominent invader of seasonal and

episodic watercourses in the succulent karoo of the

Western Cape.

Salix babylonica was recorded along watercourses in

all vegetation categories with the exception of strand-

veld. It was seldom abundant and its distribution was

largely a reflection of where it has been planted. It was

most prominent in mountain renosterveld where it was

ranked third after Acacia mearnsii and Populus x cane-

scens.

Species which scored abundance ratings of 5 or more

but were not rated as prominent were: Hakea gibbosa

and H. drupacea (= H. suaveolens), Paraserianthes

lophantha , Pin us cf. canariensis, Prosopis spp. and

Lavatera arborea. Hakea gibbosa (Figure 8D) was only

recorded as abundant near Stanford on the Akkedisberg

Pass. H. drupacea (Figure 8C) was only recorded as

abundant on the Cape Peninsula between Simonstown

and Smitswinkelbaai. If one compares the distributions

of H. gibbosa and H. drupacea with those in Stilton

(1978) they appear to have changed little in the past

twenty years.

Paraserianthes lophantha (Figure 8G) was most fre-

quently recorded in mountain fynbos & forest and moun-

tain fynbos. It has a preference for moist, low-lying sites.

Pinus cf. canariensis was only recorded as abundant on

dry mountain slopes near Paarl on the road to

Franschhoek. It was locally common around Ceres and

Tulbagh. Prosopis spp. were only locally abundant on

the plains to the east of Piketberg. They have been plant-

ed in this area and were just starting to spread as mostly

small plants were seen along roadsides and along water-

courses. Lavatera arborea was recorded along roadsides

in many coastal towns; it was common to abundant near

Saldanha Bay on the west coast.

Species that were only locally common (i.e. scoring

abundance ratings of 4) were: Tamarix spp. along water-

courses in succulent karoo and savanna near Oudtshoorn

and in coastal renosterveld on the Gouritz River south of

Mossel Bay; Quercus robur (Figure 81) and Quercus spp.

along watercourses near Stellenbosch and Swellendam;

Eucalyptus cladocalyx and E. gomphocephala in several

localities in mountain fynbos and coastal renosterveld;

Opuntia vulgaris in coastal renosterveld near Mossel

Bay; Passiflora caerulea along watercourses in coastal

renosterveld near Riversdale; Pinus pinea in coastal

renosterveld near Stellenbosch; and P halepensis (Figure

8H) in mountain fynbos near Joubertina in the Lang

Kloof. According to D.M. Richardson, pers. comm., P.

halepensis has been underestimated in this study and it

forms dense stands in the Jonkershoek Valley and on the

Cape Peninsula near Miller’s Point.

Bothalia 28,1 (1998)

109

Several other species were locally common but were

not included in formal recordings. These were:

Hedychium sp., Ipomoea cf. purpurea , Solatium mauri-

tianum and Spartium junceutn on the Constantia Nek

Road near Hout Bay on the Cape Peninsula. Spartium

junceum (Figure 8J) was recorded in several localities,

but mainly close to habitation and plantings. This species

should be closely watched as it is showing signs of

becoming invasive. Phytolacca dioica was locally com-

mon around Knysna.

Metrosideros excelsa is a potentially important invad-

er of fynbos on moist, peaty soils. It was locally common

at Betty’s Bay where it has spread from plantings and

threatens about three km of fynbos on the seafront, but

has been controlled at least since the early 1980’s (R.

Attwell pers. comm.). It has also spread from plantings

around Hermanus (M.J. Wells pers. comm.). Small

groups of naturalised plants were recorded on roadsides

in the Cape Peninsula. These were on the Ou

Kaapseweg, on the Steenberg; and on Boyes Drive

between Muizenberg and Kalkbaai. In both cases they

were in close proximity to suburban gardens and culti-

vated plants.

Pittosporum undulatum is a potentially important

invader of fire-free, wooded or forested areas. It is a

common hedge and ornamental garden plant in the

Western Cape. In this survey it was seldom recorded as

naturalised, but this could partly be due to its inconspic-

uousness in the habitats it invades. It was recorded in the

river valley between Stellenbosch and Jonkershoek, and

also on the Cape Peninsula on Boyes Drive at Kalk Bay.

According to D.M. Richardson, pers. comm., it forms

dense stands in the Jonkershoek State Forest and also at

Newlands and Tokai State Forests on the Cape Peninsula.

This species has become an important invader of forests

in many parts of the world including Jamaica, New

Zealand and even in Australia, where it is indigenous, but

has invaded vegetation outside of its natural range

(Gleadow & Ashton 1981). Recent reports of diseased

plants caused by a pathogen in the Western Cape may

help to curb the invasiveness of P. undulatum in South

Africa (M.J. Morris pers. comm.).

Myoporum tenuifolium (Figure 8E), a tree with some-

what succulent leaves and adapted to windy, salt-laden

air, has been planted as a windbreak and for shade in all

coastal towns in the study area. Occasional seedling

spread was recorded throughout its range but was most

noticeable in strandveld from the Cape Peninsula north-

wards to the west coast. It appeared to be better adapted

to the more arid and semisucculence of the strandveld

rather than coastal fynbos.

Nerium oleander (Figure 8F) has invaded watercours-

es in dry mountain valleys in several parts of the study

area. It was most evident along the Gamka, Huis and

Gouritz Rivers in the Ladismith and Calitzdorp Districts.

It was also recorded near Robertson along the Bree

River, at Citrusdal through the Piekenierskloof (Grey’s

Pass), and at Wuppertal on the Tra-Tra River. It is known

to occur in the Cederberg (Stirton 1978) but this region

was not surveyed due to bad weather conditions at the

time.

Acacia elata (Figure 8A) was most frequently record-

ed in the mountain fynbos & forest and forest vegetation

categories. Virtually all recordings were of seedling

spread from plantings in urban areas or close to habita-

tion. On a few occasions, however, seedlings were noted

on roadsides far from any plantings e.g. on the

Franschhoek and Du Toits Kloof Passes.

Relation of invasion to historical and environmental

factors

Alien plant invasion in the southern and southwestern

Cape has been greatly influenced by the deliberate intro-

duction and large scale planting of alien plant species

which were adapted to the prevailing environmental con-

ditions. For the first 150 years after the colonisation of

the Cape in 1652, plants of mainly European origin were

introduced. Only a few of these plants have become

invasive such as Pinus pinaster a native of the

Mediterranean and adapted to the climate and soils of the

southern Cape. It was only after 1830, when tree and

shrub species were intentionally imported from areas of

similar climate, especially southern and western

Australia, and were extensively propagated, that most of

the important invaders of natural vegetation became

established (Richardson et al. 1992).

By 1865 all of the most prominent Acacia, Hakea and

Pinus species mentioned in this survey, had been intro-

duced to the Cape Town region (McGibbon 1858;

Shaugnessy 1986). The government forestry authority

played a major role in attempting to establish many of

these plants both on the Cape Flats and on Table

Mountain and adjoining mountains (Shaugnessy 1986).

They also encouraged private growers to establish alien

plantations. The reasons for the plantings were various:

timber, fuel, shelter, driftsand stabilisation, tannin pro-

duction and simply beautification of a landscape that was

perceived at the time to be ‘bleak and naked’

(Shaugnessy 1986). Forestry officials even justified the

afforestation of Table Mountain in terms of improved

water supply and fire control (Shaugnessy 1986) which

is exactly opposite to today’s thinking.

In the southern Cape forests, fast-growing alien trees,

particularly Acacia, Eucalyptus, Pinus and Quercus

species, were planted in forest gaps and in plantations

along the margins of forests. Acacia melanoxylon, the

most prominent invader in the forested areas of the

southern Cape today, occurred as large trees around

George and Knysna by 1876 (Geldenhuys et al. 1986).

The pattern of invasion of several of the most wide-

spread species today, such as Pinus pinaster, Acacia

cyclops and A. saligna is a reflection of where they were

planted. The disturbance of the soil and destruction of

indigenous vegetation when establishing plantations of

these alien species would have created conditions

favourable for their further spread and also for the spread

of other alien species. Later the abandonment of many of

the plantations left stands of aliens which could act as a

seed source for future generations of these species

(Shaugnessy 1986).

110

Bothalia 28,1 (1998)

Successful invaders in the Fynbos Biome have to be

either tolerant of, or adapted to, nutrient-poor sandy soils

and periodic high-intensity fires, particularly in moun-

tain fynbos. Fire may encourage invasion by activating

mass seed release from serotinous cones (as in Hakea

sericea and Pinus pinaster ), and by stimulating germina-

tion of soil-stored seeds as in Acacia saligna and A.

longifolia (Richardson et al. 1992). The aforementioned

Acacia species show a further adaptation which enables

their seeds to avoid destruction by fire on the soil sur-

face. Their seeds, with fleshy attachments that are eaten

by ants, are buried in caches below the soil surface (Dean

et al. 1986). A. saligna also has the ability to coppice

after fire.

The Fynbos Biome is subjected to strong winds

throughout the year and this has facilitated the rapid

spread of species with winged seeds, mainly those of the

Hakea and Pinus species. Whereas seed dispersal in

mountain fynbos is mainly by wind and water; in the

lowlands birds, mammals (including humans and their

implements), ants and water are important. For example,

Acacia Cyclops seed is dispersed by mammals, such as

the striped field mouse and the chacma baboon (Stirton

1978), and many indigenous birds as well as the intro-

duced European starling (Glyphis et al. 1981). The large

seeds of Pinus pinea are dispersed by the grey squirrel

(Richardson et al. 1994).

Watercourses have played an important role in the

long-range dispersal of alien plants throughout the study

area. Species that have depended mainly on water disper-

sal include the hard-seeded legumes which are not bird-

or ant-dispersed and are otherwise relatively immobile

such as Acacia dealbata, A. mearnsii, Paraserianthes

lophantha and Sesbania punicea. Watercourses have also

enabled some species, for example, A. saligna and A.

mearnsii, to penetrate the dry interior of the Succulent

Karoo and Savanna Biomes. Nerium oleander is confined

to riverbeds and is dependant on moisture for the germi-

nation and survival of its seedlings.

SOME IDEAS FOR THE FUTURE

Since this survey was completed in 1993, a national

programme for the removal of alien plant invaders in the

water catchments of South Africa has been initiated. The

project is government-aided and is called the Working for

Water Programme of the Department of Water Affairs

and Forestry. The 1997/98 budget stands at R I 1 5 million

and the prospects for further funding are promising

(Preston 1997). It is envisaged that a large proportion of

the funds will be derived from water levies which are

likely to become a long-term source of funding (H.G.

Zimmermann pers. comm.). The proposed ‘interception

of water levy’ is aimed primarily at the forestry industry

which is one of the major water users in the country

(Yeld 1997). Clearing operations were started in October

1995 and by the end of March 1997, 71 289 hectares had

been cleared (Willems 1997). However the ultimate suc-

cess of the project will depend on very strict follow-up

operations over an extended period of time. If these con-

ditions are not met then the problem could be exacerbat-

ed rather than improved.

Many of the problem species are also useful plants

and several are major commercial forestry crops. There

is a growing call for instituting the ‘polluter pays’ princi-

ple, whereby the parties that benefit through the propa-

gation of these species must contribute to the costs of

controlling their spread from sites of propagation (Ivey

& Heydenrych 1995; Richardson et al. 1997; Yeld 1997).

Prevention is better than cure! Every attempt should

be made to prevent other, new species from becoming

invasive. The possibility of developing sterile cultivars

of commercially important species needs investigating.

The most important step should be the screening of alien

plant species for potential invasiveness before they are

introduced and widely planted. To this end, an expert

system has been developed to assist authorities in screen-

ing plants for their invasive potential (Tucker &

Richardson 1995).

Biological control using mainly introduced insects

and pathogens could play an increasingly important role

in the long-term and sustainable control of invasive

plant species in the study area. Some programmes have

already been so successful, especially those against

Acacia saligna and A. longifolia, that they could ulti-

mately lead to the destruction of all the dense infesta-

tions of these species and severely restrict any further

spread.

This paper has focused on woody plants which are the

most prominent invaders of fynbos. However herbaceous

alien plants are also important in some systems (Vlok

1988). There is an urgent need to assess the extent of

invasions by alien herbs, especially in the remaining veg-

etation on the lowlands, and to determine what impacts

they are having.

ACKNOWLEDGEMENTS

I thank Mrs H. Joffe and Mr N. Klapwijk of the

National Botanical Institute, and Mrs H. Klein of the

Plant Protection Research Institute for their assistance

and companionship in the field. I thank Dr D.M.

Richardson for many valuable comments which have

been included in this paper.

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