ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 37,2

Oct. 2007

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE PRETORIA

Obtainable from the South African National Biodiversity Institute (SANBI), Private Bag XlOl,

Pretoria 0001, Republic of South Africa. A catalogue of all available publications will be issued on

request.

BOTHALIA

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

the Union of South Africa. This house journal of the South African National Biodiversity Institute,

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

ecology, anatomy and cytology. Two parts of the journal and an index to contents, authors and sub-

jects are published annually.

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

STRELITZIA

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

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

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

Published: Vols 14—19 (earlier volumes published as supplementary volumes to the Journal of South

African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

mainly by the artists at the South Afncan National Biodiverity Institute. Many botanical artists have

contributed to the series, such as Fay Anderson, Peter Bally, Auriol Batten, Gillian Condy, Betty

Connell, Stella Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Letty (over 700 plates^ Claire

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

interest or of economic value for illustration.

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

An index to Vols 1^9 is available.

FLORA OF SOUTHERN AFRICA (FSA)

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

and Botswana, the FSA contains descriptions of families, genera, species, infraspecific taxa, keys to

genera and species, synonymy, literature and limited specimen citations, as well as taxonomic and

ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

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

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

available:

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

Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), 1983, by J.M. &

H.M. Anderson.

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

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

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

J.M. Anderson (ed.).

SANBI BIODIVERISITY SERIES

A series of occasional reports on projects, technologies, workshops, symposia and other activities

initated by or executed in partnership with SANBI.

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 37,2

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

national

biodiversity

institute

S A N B I

2 CUssonia Avenue, Brummeria, Pretoria

Private Bag XlOl, Pretoria 0001

ISSN 0006 8241

Oct. 2007

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Applequist, Ms W.L. Missouri Botanical Garden, St Louis, USA.

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

Archer, Dr R.H. South African National Biodiversity Institute, Pretoria, RSA.

Bruyns, Dr P. Department Mathematics, University of Cape Town, RSA.

Burgoyne, Ms P. South African National Biodiversity Institute, Pretoria, RSA.

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

Esler, Prof K.J. University of Stellenbosch, Stellenbosch, RSA.

Geldenhuys, Prof C.J. Forestwood cc, P.O. Box 228, La Montagne, Pretoria, RSA.

Glen, Dr H.F. South African National Biodiversity Institute, Durban, RSA.

Henderson, Ms L. Agricultural Research Council, Pretoria, RSA.

Ihlenfeldt, Prof. Dr H-D. Waabs/Langholz, Gennany.

Knevel, Dr I.C. University of Groningen, Haren, The Netherlands.

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

McNeill, Dr J. Royal Botanic Garden, Edinburgh, Scotland, UK.

Miller, Dr A.J., Saint Louis University, St. Louis, USA.

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

Randall, Dr R.P. Department of Agriculture & Food, Bentley Delivery Centre, Western Australia.

Rejmanek, Dr M. University of California, Davis, Califomia,USA.

Rutherford, Dr M.C. South African National Biodiversity Institute, Cape Town, RSA.

Saez, Dr L. Universitat Autonoma de Barcelona, Bellaterra, Barcelona, Spain.

Smithies, Mrs S.J. South African National Biodiversity Institute, Pretoria, RSA.

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

Turland, Dr N.J. Missouri Botanical Garden, St Louis, USA.

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

CONTENTS

Bothalia 37,2

1. A revision of Ornithogalum subgenus Aspasia section Aspasia, the chincherinchees (Hyacinthaceae).

J.C. MANNING, M. MARTINEZ-AZORIN and M.B. CRESPO 133

2. Name changes in the Old World Rhus and recognition of Searsia (Anacardiaceae). R.O. MOFFETT .... 165

3. New species and notes on Hesperantha (Iridaceae) in southern Africa. P. GOLDLATT and J.C. MAN-

NING 177

4. New species of Drimia (Hyacinthaceae: Urgineoideae) allied to Drimia marginata from Western and

Northern Cape, South Africa. J.C. MANNING and P. GOLDBLATT 183

5. Notes on African plants:

Boraginaceae. Lobostemon lasiophyllus: discovery of a link specimen in Stockholm, the correct

author citation and synonymy. M.H. BUYS, B. NORDENSTAM and R. VOGT 196

Crassulaceae. Bryophyllum proliferum naturalized in KwaZulu-Natal, South Africa. N.R. CROUCH

and G.F. SMITH 206

Crassulaceae. Crassula streyi recorded from the Eastern Cape, South Africa. N.R. CROUCH and

T.J. EDWARDS 208

Didiereaceae/Portulacaceae. Ceraria kaokoensis, a new species from Namibia, with notes on

gynodioecy in the genus. W. SWANEPOEL 202

Iridaceae. Aristea nigrescens (subgenus Pseudaristea), a new species from Western Cape, South Africa

with a novel strategy for pollinator attraction. J.C. MANNING and P. GOLDBLATT 189

Iridaceae. Nivenia argentea misunderstood, and the new species Nivenia inaeqiialis (Nivenioideae).

J.C. MANNING and P. GOLDBLATT 192

Pedaliaceae. Dewinteria, a new semisucculent, cliff-dwelling genus endemic to the Kaokoveld,

Namibia. E.J. VAN JAARSVELD and A.E. VAN WYK 198

6. Stem diameter and bark surface area of the fluted trunk of Balanites maughamii (Balanitaceae). VL.

WILLIAMS, K. BALKWILL and E.T.F. WITKOWSKI 211

7. Invasive, naturalized and casual alien plants in southern Africa: a summary based on the Southern African

Plant Invaders Atlas (SAPIA). L. HENDERSON 215

8. Seasonal variation in soil seed bank size and species composition of selected habitat types in Maputaland,

South Africa. M.J.S. KELLERMAN and M.W. VAN ROOYEN 249

9. Obituary: Ian Frederick Garland (1925-2007). E. POOLEY 259

10. South African National Biodiversity Institute: administration and research staff 31 March 2007, publi-

cations 1 April 2006-31 March 2007. Compiler: B.A. MOMBERG 261

11. Guide for authors to Bothalia 283

New combinations, species, statuses and varieties in Bothalia 37,2 (2007)

Aristea nigrescens J.C. Manning & Goldblatt, sp. nov., 189

Ceraria kaokoensis Swanepoel, sp. nov., 202

Dewinteria VanJaarsv. & A.E.van Wyk, gen. nov., 198

Dewinteria petrophila (De Winter) VanJaarsv. & A.E.van Wyk, comb, nov., 198

Drimia ligulata J.C. Manning & Goldblatt, sp. nov., 186

Drimia pulchromarginata J.CMaww/wg & Goldblatt, sp. nov., 185

Drimia vermiformis J.C. & Goldblatt, sp. nov., 184

Hesperantha acuta subsp. tugwelliae (R.C. Foster) Goldblatt & J.C. Manning, comb, et slat nov., 182

Hesperantha helmei Goldblatt & J.C. Manning, sp. nov., 179

Hesperantha XWhKoXn J.C. Manning & Goldblatt, sp. nov., 180

Hesperantha longistyla J.C. & Goldblatt, sp. nov.. Ill

Nivenia inaequalis Goldblatt & J.C. Manning, sp. nov., 195

Ornithogalum corticatum Mart.-Azorln, sp. nov., 140

Searsia acocksii (Mojfett) Moffett, comb, nov., 166

Searsia acuminatissima (R.Fern. & A. Fern.) Moffett, comb, nov., 166

Searsia albida (Schousb.) Moffett, comb, nov., 167

Searsia albomarginata (Sond.) Moffett, comb, nov., 167

Searsia anchietae (Figalho & Hiern ex Hiern) Moffett, comb, nov., forma anchietae, 167

Searsia anchietae (Figalho & Hiern ex Hiern) Moffett forma mendon^ae (Meikle) Moffett, comb, nov., 167

Searsia anchietae (Figalho & Hiern ex Hiern) Moffett forma suffruticosa (Meikle) Moffett, comb, nov., 167

Searsia angolensis (Engl.) Moffett, comb, nov., forma angolensis, 167

Searsia angolensis (Engl.) Moffett fonna glabrescens (R.Fern.) Mojfett, comb, nov., 167

Searsia arenaria (Engl.) Moffett, comb, nov., 167

Searsia aucheri (Boiss.) Moffett, comb, nov., 167

Searsia batophylla (Codd) Moffett, comb, nov., 167

Searsia blanda (Meikle) Moffett, comb, nov., forma blanda, 167

Searsia blanda (Meikle) Moffett forma exelliana (Meikle) Moffett, comb, nov., 167

Searsia bolusii (Sond. ex Engl.) Moffett, comb, nov., 167

Searsia brenanii (Kokwaro) Moffett, comb, nov., 167

Searsia burchellii (Sond. ex Engl.) Moffett, comb, nov., 167

Searsia carnosula (Schonland) Moffett, comb, nov., 167

Searsia chirindensis (Baker f) Moffett, comb, nov., 167

Searsia crenata (Thiinb.) Moffett, comb, nov., 167

Searsia crenulata (A.Rich.) Moffett, comb, nov., 167

Searsia discolor (E.Mey. ex Sond.) Moffett, comb, nov., 168

Searsia dissecta (Thiinb.) Moffett, comb, nov., 168

Searsia divaricata (Eckl. & Zeyh.) Moffett, comb, nov., 168

Searsia dracomontana (Moffett) Moffett, comb, nov., 168

Searsia dregeana (Sond.) Moffett, comb, nov., 168

Searsia dumetorum (Exell) Moffett, comb, nov., 168

Searsia engleri (Britten) Moffett, comb, nov., 168

Searsia erosa (Thiinb.) Moffett, comb, nov., 168

Searsia fanshawei (R.Fern. & A. Fern.) Moffett, comb, nov., 168

Searsia fastigata (Eckl. & Zeyh.) Moffett, comb, nov., 168

Searsia flexicaulis (Baker) Moffett, comb, nov., 168

Searsia gallagheri (Ghaz.) Moffett, comb, nov., 168

Searsia gerrardii (Harv. ex Engl.) Moffett, comb, nov., 168

Searsia glauca (Thunb.) Moffett, comb, nov., 168

Searsia glaucescens (A.Rich.) Moffett, comb, nov., 168

Searsia glutinosa (Hochst. ex A.Rich.) Moffett . abyssinica (Oliv.) Moffett, comb, nov., 168

Searsia glutinosa (Hochst. ex A.Rich.) Moffett, comb, nov., subsp. glutinosa, 168

Searsia glutinosa (Hochst. ex A.Rich.) Moffett subsp. neoglutinosa (M.G.Gilbert) Moffett, comb, nov., 168

Searsia gracilipes (Exell) Moffett, comb, nov., 168

IV

Searsia gracillima (Engl.) Mojfett var. glaberrima (Schdniand) Moffett, comb, nov., 168

Searsia gracillima (Engl.) Moffett, comb, nov., var. gracillima, 168

Searsia grandidens (Harv. ex Engl.) Moffett, comb, nov., 169

Searsia grossireticulata (Van der Veken) Mojfett, comb, nov., 169

Searsia harveyi (Moffett) Moffett, comb, nov., 169

Searsia horrida (Eckl. & Zeyh.) Moffett, comb, nov., 169

Searsia humpatensis (Meikle) Moffett, comb, nov., forma humpatensis, 169

Searsia humpatensis (Meikle) Moffett forma subglabra (R.Fern.) Moffett, comb, nov., 169

Searsia incisa (L.f.) FA. Barkley var. effusa (Presl) Moffett, comb, nov., 169

Searsia keetii (Schdniand) Moffett, comb, nov., 169

Searsia kirkii (Oliv.) Moffett, comb, nov., 169

Searsia krebsiana (Presl ex Engl.) Moffett, comb, nov., 169

Searsia kwangoensis (Van der Veken) Moffett, comb, nov., 169

Searsia kwazuluana (Moffett) Mojfett, comb, nov., 169

Searsia laevigata (L.) FA. Barkley var. laevigata forma cangoana (Moffett) Moffett, comb, nov., 169

Searsia laevigata (L.) FA. Barkley var. villosa (L.f.) Moffett, comb, nov., 169

Searsia leptodictya (Diels) T.S. Yi, A.J.Mill. & J. Wen forma pilosa (R.Fern. & A. Fern.) Moffett, comb, nov., 169

Searsia longipes (Engl.) Moffett, comb, nov., var. longipes, 170

Searsia longipes (Engl.) Moffett var. elgonensis (Kokwaro) Moffett, comb, nov., 170

Searsia longipes (Engl.) Moffett var. schinoides (R.Fern.) Moffett, comb, nov., 170

Searsia longispina (Eckl. & Zeyh.) Moffett, comb, nov., 170

Searsia iucens (Hutch.) Moffett, comb, nov., 170

Searsia lucida (L.) FA. Barkley forma elliptica (Sond.) Moffett, comb, nov., 170

Searsia lucida (L.) F. A. Barkley forma scoparia (Eckl. & Zeyh.) Moffett, comb, nov., 170

Searsia magalismontana (Sond.) Moffett, comb, nov., subsp. magalismontana, 170

Searsia magalismontana (Sond.) Moffett subsp. coddii (R.Fern. & A. Fern.) Moffett, comb, nov., 170

Searsia magalismontana (Sond.) Moffett snhsyi. trifoliolata (Baker f.) Moffett, comb, nov., 170

Searsia maricoana (Moffett) Moffett, comb, nov., 170

Searsia marlothii (Engl.) Moffett, comb, nov., 170

Searsia montana (Diels) Moffett, comb, nov., 170

Searsia monticola (Meikle) Moffett, comb, nov., 170

Searsia mysorensis (G.Don) Moffett, comb, nov., 170

Searsia nebulosa (Schdniand) Moffett, comb, nov., forma nebulosa, 170

Searsia nebulosa (Schdniand) Moffett forma pubescens (Moffett) Moffett, comb, nov., 170

Searsia nitida (Engl.) Moffett, comb, nov., 171

Searsia obtusata (Engl.) Moffett, comb, nov., 171

Searsia ochracea (Meikle) Moffett, comb, nov., var. ochracea, 171

Searsia ochracea (Meikle) Moffett var. saxicola (R.Fern. & A. Fern.) Moffett, comb, nov., 171

Searsia pallens (Eckl. & Zeyh.) Moffett, comb, nov., 171

Searsia paniculata (Wall, ex G.Don) Moffett, comb, nov., 171

Searsia pendulina (Jacq.) Moffett, comb, nov., 171

Searsia pentheri (Zahlbr.) Moffett, comb, nov., 171

Searsia pondoensis (Schdniand) Moffett, comb, nov., 171

Searsia populifolia (E.Mey. ex Sond.) Moffett, comb, nov., 171

Searsia problematodes (Merxm. & Rdssl.) Moffett, comb, nov., 171

Searsia pterota (Presl) Moffett, comb, nov., 171

Searsia puccionii (Chiov.) Moffett, comb, nov., 171

Searsia pygmaea (Moffett) Moffett, comb, nov., 171

Searsia pyroides (Burch.) Moffett, comb, npv., var. pyroides, 171

Searsia pyroides (Burch.) Moffett var. dinteri (Engl.) Moffett, comb, nov., 171

Searsia pyroides (Burch.) Moffett var. gracilis (Engl.) Moffett, comb, nov., 171

Searsia pyroides (Burch.) Mojfett var. integrifolia (Engl.) Moffett, comb, nov., 171

Searsia refracta (Eckl. & Zeyh.) Moffett, comb, nov., 172

Searsia rehmanniana (Engl.) Moffett, comb, nov., var. rehmanniana, 172

Searsia rehmanniana (Engl.) Moffett var. glabrata (Sond.) Moffett, comb, nov., 172

Searsia retinorrhoea (Steud. ex Oliv.) Moffett, comb, nov., 172

Searsia rigida (Mill.) FA. Barkley var. dentata (Engl.) Moffett, comb, nov., 172

V

Searsia rigida (Mill) FA. Barkley var. margaretae (Burtt Davy ex Moffett) Moffett, comb, nov., 172

Searsia rimosa (EckJ. & Zeyh.) Moffett, comb, nov., 172

Searsia rogersii (Schonland) Moffett, comb, nov., 172

Searsia rudatisii (Engl.) Moffett, comb, nov., 172

Searsia ruspolii (Engl.) Moffett, comb, nov., 172

Searsia scytophylla (Eckl. & Zeyh.) Moffett, comb, nov., var. scytophylla, 172

Searsia scytophylla (Eckl. & Zeyh.) Moffett \&x. dentata (Moffett) Moffett, comb, nov., 172

Searsia sekhukhuniensis (Moffett) Moffett, comb, nov., 172

Searsia somalensis (Engl.) Moffett, comb, nov., 172

Searsia squalida (Meikle) Moffett, comb, nov., 172

Searsia stenophylla (Eckl. & Zeyh.) Moffett, comb, nov., 172

Searsia tenuinervis (Engl.) Moffett, comb, nov., 172

Searsia tenuipes (R.Fern. & A.Fern.) Moffett, comb, nov., 173

Searsia thyrsiflora (Balf.f) Moffett, comb, nov., 173

Searsia transvaalensis (Engl.) Moffett, comb, nov., 173

Searsia tridactyla (Burch.) Moffett, comb, nov., 173

Searsia tripartita (Ucria) Moffett, comb, nov., 173

Searsia tumulicola (S. Moore) Moffett, comb, nov., var. tumulicola, 173

Searsia tumulicola (S. Moore) Moffett var. meeuseana (R.Fern. & A.Fern.) Moffett, comb, nov., forma meeuseana, 173

Searsia tumulicola (S. Moore) Moffett var. meeuseana (R.Fern. & A.Fern.) Moffett forma pumila (Moffett) Moffett,

comb, nov., 173

Searsia volkii (Siisseng.) Moffett, comb, nov., 173

Searsia wellmanii (Engl.) Moffett, comb, nov., 173

Searsia wildii (R.Fern. & A.Fern.) Moffett, comb, nov., 173

Searsia wilmsii (Diels) Moffett, comb, nov., 173

Searsia zeyheri (Bond.) Moffett, comb, nov., 173

VI

Bothalia 37,2: 133-164(2007)

A revision of Ornithogalum subgenus Aspasia section Aspasia, the

chincherinchees (Hyacinthaceae)

J.C. MANNING*, M. MARTINEZ- AZORIN** and M.B. CRESPO**

Keywords: chincherinchees, Hyacinthaceae, Ornithogalum L., South Africa, subgenus Aspasia (Salisb. ) Oberm., taxonomy

ABSTRACT

The species of Ornithogalum L. subgenus Aspasia section Aspasia are revised. Section Aspasia is defined by a rosette of

lanceolate to oblong leaves; large, boat-shaped, ± petaloid bracts; moderately-sized white, yellow or orange flowers, sometimes

with dark central markings; thin-textured, ellipsoid capsules that are enclosed by and concealed within the persistent, papery

perianth; and angular, colliculate to echinulate seeds. Twelve species are recognized in the section, separable into three series

based on seed morphology. O. conicum is redefined to exclude specimens from the Eastern Cape, which are recognized as O.

synanthifoliiim, and O. conicum subsp. strictiim is raised to species stams as O. strictwn. The circumscription of O. dubium

is expanded to include O.fimbrimarginatum and O. subcoriaceum, previously distinguished on account of their longer styles.

Collections from the Roggeveld Escarpment and Klein Roggeveld that were previously included in O. fimbrimarginatum are

recognized as the new species O. corticatum Mart.-Azorin, on the basis of their unusual, thick, cartilaginous outer tunics and

puberulous adaxial leaf surface. O. ceresianum is removed from the synonomy of O. thyrsoides and recognized as a distinct

species on account of its extensive glossy black tepal markings, winged inner filaments, and glossy black ovary. The poorly

known O. puberulum is more fully described based on several recent collections, and O. leeupoortense is neotypified in

the absence of any original type material. O. nipestre and O. multifolium are regarded as colour forms of the same species,

for which O. nipestre is the older name. Similarly, O. roussouwii is a depauperate, pale form of O. maculatum and is thus

included in the synonomy of that species. The circumscription of O. pruinosum remains unchanged. The species O. baurii, O.

diphyllum and O. sephtonii from the Drakensberg Mountains of Eastern Cape and KwaZulu-Natal are excluded from section

Aspasia on the basis of their turbinate capsules that are exposed by the reflexed tepals. Each species is fully described, with

accompanying discussion of variation and relationships, a distribution map. and an illustration.

CONTENTS

Abstract 133

Introduction 133

Materials and methods 134

Taxonomy 134

Ornithogalum L 134

Subgenus Aspasia (Salisb.) Oberm. section Aspasia 135

Key to species 135

Series 1 Aspasia 135

1 . Ornithogalum conicum Jacq 135

2. O. synanthifoliiim E.M. height 138

3. O. corticatiim Mart..- Azonn 140

Series 2 Thyrsoides 143

4. O. L. Bolus 143

5. O. thyrsoides Jacq 143

6. O. ceresianum Y.M..hQigh\. 146

Series 3 Maculata 147

7. O. dubium Houtt 147

8. O. pruinosum F.M. height 152

9. O. maculatum Jacq 152

10. (9. nipestre L.f. 155

11.0. leeupoortense U.Mull.-Doblies & D.MiilL-

Doblies 156

12. O. puberulum Oberm $ 158

Excluded species 160

Acknowledgements 161

References 161

Other specimens examined 162

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

E-mail: manning@sanbi.org.

** CIBO (Instituto de la Biodiversidad), Universidad de Alicante, P.O.

Box 99, E-03080 Alicante, Spain.

E-mail: mmartinez@uz.es; crespo@ua.es.

MS. received: 2006-11-28.

INTRODUCTION

Hyacinthaceae, a predominantly Old World family,

is distributed throughout Africa and the Mediterranean,

extending through central to eastern Asia, with a single,

small genus in the Andes Mountains in South America

(Speta 1998). Around 400 of an estimated total of 700-

900 species are endemic to southern Africa, making it

one of the most important geophyte families in south-

ern Africa, exceeded in numbers only by the Iridaceae.

Hyacinthaceae are especially numerous in the south-

western winter rainfall region, which is one of the main

centres of diversity for the family.

Among the more common and conspicuous members

here are the large-flowered species of Ornithoglaum L.

subgenus Aspasia (Salisb.) Oberm., known colloquially

as chincherinchees, an onomatopoeic sobriquet derived

from the sound produced when the stems are rubbed

together (Smith 1966). Several species in this group,

especially O. thyrsoides, are a characteristic part of the

southwestern Cape spring, occurring in enormous popu-

lations in lowland areas around Cape Town and further

north into Namaqualand. Their extreme toxicity (Van

Wyk et al. 2002) enables them to colonize overgrazed

lands with impunity, and their ease of cultivation and

long vase-life have made them important horticultur-

ally. Several selections of O. dubium and O. thyrsoides

are available commercially as cut-flowers and further

breeding programmes are ongoing. The taxonomy of this

group, however, is far from fully understood and several

of the species are notoriously difficult to identify.

The southern African species of Ornithogalum have

been revised three times in the last sixty years, beginning

with the work of Leighton (1944, 1945). This review was

the first complete regional treatment of the genus since

Baker’s (1897) account for the Flora capensis, with the

134

Bothalia 37,2 (2007)

apparently significant advantage gained from examina-

tion of living material collected in the wild. Leighton,

like Baker before her, was impressed by the great vari-

ability among populations in the fonn of the inner fila-

ments, which may be slender and awl-like, lanceolate,

with or without small basal expansions, or conspicuously

winged. Flower colour is another source of variation,

ranging from pure white through various milky-white or

buff shades to yellow, orange or reddish orange, with or

without small to large dark central markings. Defining

her taxa very narrowly, Leighton (1944, 1945) described

numerous species to accommodate the different combi-

nations of these characteristics, ultimately recognizing

21 species and many forms in the group (Leighton 1945).

With further study, however, it became clear that many

of these species represented nothing more than local

populations. Obermeyer (1978), who was the first to pro-

pose a more formal recognition of the chincherinchees,

as the Aspasiae group of subgenus Aspasia, adopted a

much broader view of the species in her monograph on

the genus, reducing their number to 10, with two subspe-

cies recognized in O. conicwn. Ornithogahtm diphyUum

Baker from the KwaZulu-Natal Drakensberg, unaccount-

ably left out of Leighton’s treatment, was included as the

eleventh member of the group. This treatment was largely

followed by Muller-Doblies & Muller-Doblies (1996),

who formalized the group as section Aspasia. The sec-

tion was expanded slightly to include two additional

species, O. constrictum F.M.Leight. and O. inclusiim

F.M.Leight., which had been placed by Obemieyer in

the Hispidaspasiae group, and was subdivided into two

subsections and six series. The species in Obenneyer’s

Aspasiae group were dispersed among four series in two

subsections: Aspasia, Leenpoortensia U.Mull.-Doblies

& D.Miill.-Doblies and Maculata U.Mull.-Doblies &

D.Miill.-Doblies of subsection and series Riispes-

tria U.Mull.-Doblies & D.Miill.-Doblies of subsection

Teretaspasia U.Mull.-Doblies & D.Miill.-Doblies. A

total of 1 3 species was recognized, two of which, O. lee-

upoortense and O. roiissoiiwi , were newly described and

another two resuscitated from synonomy.

Despite the relatively high level of taxonomic study

to which it has been exposed, the taxonomy of the group

remains unclear, and the identification of specimens is

often problematical. Although some of the species are

well circumscribed and may be identified without much

difficulty, the boundaries of others, especially those

around O. duhium, remain poorly defined. Increased col-

lecting over the past decades has greatly improved our

appreciation of the natural variation among wild popula-

tions, indicating the need for another review of the spe-

cies in this group. A comprehensive, illustrated account

of the species is presented here for the first time.

It is evident that many of the characters that have tra-

ditionally been used to separate species in the group are

much more variable than has been realized. This is par-

ticularly true of flower colour, the degree of basal expan-

sion of the inner filaments, the length of the style rela-

tive to the ovary, and the surface sculpturing of the seeds.

Most species are reliably diagnosed by a combination of

characters, supplemented with distribution and eeological

data. A previously underappreciated character, the size

of the seeds, appears to be a useful indicator of relation-

ships among the species. Three seed size classes can be

distinguished; large (2. 0-3. 5 mm long), in O. conicum,

O. corticatiirn and O. synanthifolium; medium (1-2 mm

long), in O. ceresiamim, O. strictum and O. thyrsoides;

and small (0. 5-1.0 mm long), in the remaining speeies.

The species are arranged in three series aeeording to

these size classes.

MATERIALS AND METHODS

This study is based on an examination of dried her-

barium specimens as well as living plants studied during

extensive field work undertaken thoughout the southwest-

ern Cape. The herbarium specimens studied inelude the

complete collections in BOL, K, NBG and SAM, the types

of all names, and selected specimens from PRE. Specimens

examined are listed at the end. Seeds were examined with

both light and scanning electron microscopy.

TAXONOMY

Ornithogalum L., Species plantarum: 306 (1753).

Type: O. iimbellatiini L. (vide Steam 1983).

Deciduous or rarely evergreen perennials. Bulb subter-

ranean or epigeal, subglobose or rarely poorly developed

and rootstock rhizomatous, tunics sometimes scale-like

or loosely overlapping, usually white but rarely pinkish;

outer bulb tunics membranous, papery, or leathery. Leaves

1 -several, green or dry at flowering, erect or spreading,

linear to oblong or filiform, sometimes very sueculent,

usually glabrous but sometimes pubescent or glandular-

pubeseent, margins smooth, eiliate, fringed, or hyaline,

sheaths sometimes persistent and forming a papery or

weakly to strongly fibrous, sometimes horizontally barred

sheath around base of stem. Inflorescence a several- to

many-flowered raceme, sometimes subcorymbose with

shortened axis, or secund, usually solitary but sometimes

more than one; peduncle rarely papillate; bracts membra-

nous, leafy or petaloid, small or large, not spurred; bracte-

oles usually lacking but sometimes thread-like and borne

on alternate sides of pedicels at base; pedicels short or

long. Flowers white, yellow, orange, or yellowish green,

without darker keels, scented or unscented, sometimes

closing at night or more rarely nocturnal, suberect or pat-

ent, rotate or campanulate; tepals ovate to narrowly lan-

ceolate, persistent, ± shortly united at base or rarely united

into a short tube, spreading to erect, sometimes with tips

reflexed. Stamens suberect or slightly spreading; fila-

ments free or rarely united below, fused to base of tepals

or inserted at top of tube, filiform to lanceolate, all similar

or inner usually broader, both whorls'or more usually only

inner variously expanded or toothed below. Ovary ovoid

to globose or turbinate, sometimes shortly stipitate; ovules

few to many per locule; style vestigial to long, subcylindri-

cal, erect or sometimes slightly deflexed; stigma small and

3-lobed or -angled. Capsule fusiform, ovoid to subglobose,

± 3-angled or -lobed, membranous or leathery, enclosed in

dry perianth or exposed, dehiscing loculicidally. Seeds few

to many per chamber, flattened or angled, black, the testa

tightly adhering, laevigate, rugulate, papillate or echinate.

Base chromosome number x = 9.

About 250 species in Africa, Madagascar, Mediterra-

nean, Saudi Arabia and India, mainly in the winter rainfall

areas of southern Africa, including ± 40 species in Western

Cape, South Africa.

Bothalia 37,2 (2007)

135

Subgenus Aspasia (Salisb.) Oberm. in Bothalia 12;

333 (1978). Type: O. conicum Jacq.

Section Aspasia

Leaves radical, lanceolate to oblong, usually glabrous

but sometimes partially or entirely pubescent, mar-

gins usually fringed or ciliate. Inflorescence sometimes

subcorymbose; bracts large, boat-shaped, foliaceous

or petaloid, margins entire or ciliate but not denticu-

late. Flowers medium-sized to large, white, yellow or

orange, without darker keels but sometimes with dark

centre, closing at night, shallowly bowl-shaped; tepals

free, ovate, persistent and papery in fruit. Stamens free;

filaments filiform to lanceolate and all similar or inner

broader, both whorls or more usually only inner vari-

ously expanded or winged basally. Ovary ovoid; style

well-developed or vestigial. Capsule fusiform or ellip-

soid, thin-walled, enclosed and concealed within dry

perianth. Seeds many per chamber, pyriform, comma-

shaped or cuneate, 1-3 mm long, testa papillate or echi-

nulate. Base chromosome number x = 6.

Species 12, mainly endemic to the winter rain-

fall region of the southwestern Cape, with two species

extending eastwards to the Eastern Cape.

Key to species

la Style up to 2.5 mm long; leaves glaucous, margins smooth; flowers yellow to orange, rarely whitish or pinkish, sometimes with dark

markings at tips of tepals; bracts brownish apically:

2a Leaves 2-5, narrowly lanceolate to oblong; outer tepals usually with pale or dark marks at tips; flowers larger, tepals 1 1-25 x

5-14 mm 9.0. macidatum

2b Leaves 3-10. subterete or linear-canaliculate; tepals never maculate; flowers smaller, tepals 6-12 x 3. 5-5.0 mm 10. O. nipestre

lb Style usually more than 2.5 mm long but if shorter then leaf margins ciliate:

3a Leaves 2 or 3, soft-textured, glabrous or pubescent but margins always with soft cilia 1-2 mm long; lower leaf base clasping and

inflated; plants from southern Namibia and Richtersveld 12.0. pubenilum

3b Leaves usually more than 3, ± leathery or subsucculent, margins glabrous to densely ciliate with short, stiff cilia less than 0.7 mm

long; plants from South Africa:

4a Robust plants 350-950 mm high, ± evergreen with leaves 200-400 mm long and glabrous or minutely ciliate on margins; lowermost

pedicels 15-30 mm long in flower, lengthening up to 35-75 mm in fruit; capsules 12-15 mm long and seeds 2-3 mm long; plants

from Eastern Cape, east of Grahamstown 2. O. synanthifoliwn

4b Not as above:

5a Filaments ± monomorphic, filiform to awl-shaped, rarely inner with small basal expansion in lower 1.5 mm; bulbs large, 25-35

mm diam.. outer tunics pale and papery; plants restricted to southwestern Cape coastal districts; flowering from late November

to January 1.0. conicum

5b Filaments ± dimorphic, awl-shaped to lanceolate, inner wider than outer and usually with ± prominent basal expansions:

6a Outer bulb tunics soft-textured or papery, pale grey or whitish; leaves suberect and clasping at base, lanceolate, bright green;

seeds 1-2 mm long:

7a Raceme narrowly cylindrical, 70-150 mm long in flower; lowermost pedicels 5-15 mm long at flowering, scarcely elongating

in fhiit and then 15-25 mm long; flowers rarely with dark centre; inner filaments expanded only in lower third, expansion

rhomboidal or inconspicuously toothed; style often deflexed 4. O. strictum

7b Raceme conical to subcorymbose, 30-80 mm in flower; lowermost pedicels 14-37 mm long at flowering, elongating up to

20-70 mm long in fruit; flowers with darker centre, although this sometimes small; inner filaments expanded in lower half

and clasping ovary, expansions with conspicuous apical wings that often cover top of ovary; style erect:

8a Inner filaments expanded and winged in lower half, outer filaments linear to subulate; flowers usually with small greenish or

brownish centre; ovary dull brownish to black 5. O. thyrsoides

8b Both inner and outer filaments expanded and winged in lower half, inner more prominently so; flowers with large, dark centre

covering lower half of tepals; ovary glossy dark green to black and conspicuously 3-lobed 6. O. ceresianum

6b Outer bulb tunics firm-textured and leathery or cartilaginous, grey or blackish; leaves elliptical or shortly lanceolate; seeds usu-

ally 0. 5-1.0 mm long:

9a Leaf margins glabrous, minutely ciliate or obscurely papillate; plants from drier parts of Northern and Western Cape north of

Klawer;

10a Leaves bright green, distichous, falcate, attenuate, as long as or longer than inflorescence; inflorescence usually laterally

displaced and thus apparently axillary 11.0. leeupoortense

10b Leaves glaucous, rosulate, oblong-lanceolate, leathery, usually less than half as long as inflorescence, margins

sometimes crisped 8. O. pruinosum

9b Leaf margins densely ciliate; plants from Northern Cape south of Calvinia, Western and Eastern Cape:

1 la Leaves shortly pubescent adaxially near apex, withered at flowering; bulbs relatively large, globose, outer tunics cartilagi-

nous or thick and leathery, accumulating in thick layers; flowers white, without dark centre; inner filaments slightly keeled;

plants from Roggeveld Escarpment in Northern Cape 3. O. corticatum

lib Leaves glabrous except along margins, often still green at flowering; tunics not accumulating in thick layers; flow-

ers orange, yellow, cream-coloured or white, often with dark centre; bracts sometimes ciliate in upper half; plants

from Western and Eastern Cape 7. O. dubitim

Series 1 Aspasia

Flowers without dark centre; seeds 2. 0-3. 5 mm long.

1. Ornithogalum conicum Jacq., Collectanea 3: 232

(1791). O. lacteum var. conicum (Jacq.) Baker: 284 (1873).

Type: South Africa, Cape, without precise locality, in Jacq.,

leones plantarum rariorum 2, t. 428 (1789b)(icono.!).

O. lacteum Jacq.: 76 (1797). Type: South Africa, Cape, with-

out precise locality, in Jacq., leones plantarum rariorum 2, t. 434

(1789b)(icono.!).

O. aestivum L. Bolus: 55 (1934). Type: South Africa, [Western

Cape], between Malmesbury and Mamre, L. Bolus s.n. BOL20974

(BOL, holo.!).

Plants 300-600(-900) mm high. Bulb subglobose,

25-45 mm diam., flesh sometimes pale pink, outer

tunics whitish, papery. Leaves spreading or suber-

ect, 5-10, one quarter to one third as long as flow-

ering stem, partially or completely dry at flowering,

oblong-lanceolate, 45-100(-120) x 6-25(-30) mm,

glabrous but densely ciliate on margins. Raceme sub-

136

Bothalia 37,2 (2007)

corymbose to conical-cylindrical, compact, 30-80 mm long

at flowering, elongating to 60-100(-150) mm long in fruit,

(ll-)15-30(-35)-flowered; lowermost pedicels 10-17 mm

long, elongating to 15-28 mm in fruit; bracts petaloid, whit-

ish and papery, ovate, acute or acuminate, usually exceed-

ing pedicels, lowennost 14—20 mm long. Flowers white

with small greenish yellow centre, unscented or faintly

honey-scented; outer tepals ovate- lanceolate, inner obovate,

14-17(-20) X 6-9(-12) mm. Stamens half as long as tepals;

filaments filiform or subulate, white, 6-8 mm long, rarely

inner widened at base. Ovary ovoid, 4-5 mm long, green-

ish yellow but brighter yellow at apex; style white, 2-3 mm

long. Capsule fusiform to oblong-ovoid, 3-lobed, apicu-

late, 10-13 mm long. Seeds angular and irregularly folded,

papillate to echinulate, 2-3 mm long. Flowering time: late

November or December to mid-January. Figures 1 A, 2.

J

K

L

FIGURE 1. — Filament morphology in Ornithogaliim section Aspasia. Filaments are shown in pairs from same flower, outer on left and inner on

right. A, Ornilhogaltim conicunr, B, O. symmihifoliiinv, C, O. corticalunr, D, O. siriclum; E, O. thyrsoides', F, O. ceresianiiirr, G, O. duhiwrv,

H, O. pruinosnm', I, O. maadalion; J, O. rupesire, K, O. leeiipoorten.se, L. O. puhenilum. Not to scale.

Bothalia 37,2 (2007)

137

V

\

*

^raxti.dgalxxn oonicuA Jacq«

dat. it. i» Mauvq^ 1975

NATIONAL BOTANIC GARDENS OF

SOUTH AFRICA,

Distribution and ecology: restricted to coastal areas

in the southwestern Cape, from Graafwater southwards

along the west coast to the Cape Peninsula and Gordon’s

Bay (Figure 3), occurring on coastal limestone pavement,

stony hills and granitic outcrops, typically in coarse-

grained sandy or gravelly soils but also shale.

Discussion: Omithogalum conicum is a summer-flow-

ering species from coastal areas in the extreme southwest-

ern Cape. It is characterized by its very large bulb, some-

times with pale pink flesh, whitish to light grey, papery

tunics, and a rosette of short, densely ciliate leaves less

FIGURE 2. — Omithogalum conicum

Jacq.

than one third as long as the inflorescence and partially or

completely dry at flowering. Both whorls of filaments are

typically filiform or awl-shaped, or the inner may rarely

be slightly widened at the base but never into the api-

cally lobed, oblong expansions that occur in other species

in the group. The species is variable in stature and size

with the largest plants, up to 900 mm high, recorded from

Paleisheuwel. Plants from the Cape Peninsula are shorter,

never more than 500 mm high.

Both Omithogalum thyrsoides and O. dubium occur

with O. conicum on the Cape Peninsula. O. thyrsoides.

138

Bothalia 37,2 (2007)

FIGURE 3. — Known distribution of Ornilhogalum conicum, •; O. corticatiim, O; O. synanthifoliiim. ▲.

which often forms large colonies along road verges

and in waste ground, differs in its much smaller bulb,

seldom more than 20 mm in diameter, longer, suber-

ect, ± synanthous leaves, and conspicuously winged

inner filaments. It also flowers earlier in the season,

in September and October, and is in fruit by the time

that O. conicum begins to bloom in early November.

White-flowered forms of O. dubiurn that occur on Table

Mountain are distinguished by their smaller bulb with

dark tunics, mostly subcorymbose inflorescence, and

basally expanded or winged inner filaments. In addi-

tion, the ovary in O. dtibiwn is typically blackish, unlike

the yellow ovary in O. conicum, and the seeds are much

smaller (± 1 mm vs 2-3 mm long). On the Peninsula,

O. dubiurn is restricted to sheltered plaees on sandstone

cliffs, whereas O. conicum is found at lower altitudes

on exposed granite or shale slopes. The flowers in the

populations of O. conicum on Lions Head on the Cape

Peninsula are open throughout the day from early morn-

ing, and are slightly scented, unlike those of O. dubiurn,

which only open around mid-morning and lack fra-

grance.

Although the species has been doeumented as toxic,

the actual identity of the specimens tested is uncertain in

view of the much broader eircumscription of the species

that was eurrent in the past. Given its relatively restricted

geographic range, however, it seems more likely that

these results were based on tests of O. strictum rather

than true O. conicum.

History: Ornithogalum conicum was illustrated and

described by Jacquin from cultivated plants that were

almost certainly originally collected on the Cape Penin-

sula, where the species is still found on the slopes of

Signal Hill and the foot of Lions Head. At the same

time, Jacquin illustrated and named O. lacteum but this

appears to be nothing more than a luxuriant form with

the bases of the inner filaments slightly expanded. This

form was reduced to synonomy by Baker ( 1 897), whose

concept of O. conicum, later followed by Obermeyer

(1978), included not only plants from the Eastern Cape

with narrow filaments, which are segregated here as O.

synanthifolium, but also eollections from the Olifants

River Valley that are treated here as a distinct species O.

strictum. Both of these taxa have relatively long leaves

that are still green at flowering, leading Louisa Bolus to

describe O. aestivum for plants from the West Coast that

produced a rosette of short leaves which were dry and

withered by the time that the plants flowered in summer.

It is now elear that these populations represent true O.

conicum, which is more narrowly defined here than in

the past.

2. Ornithogalum synanthifolium F.M.Leight. in

Journal of South African Botany 10: 176 (1945). Type:

South Africa, [Eastern Cape], King William’s Town

Distriet, Perie [Pirie] Mtns, Galpin 2528 (PRE, holo.!).

Plants (200-)350-950 mm high. Bulb ovoid, some-

times not well developed, (20-)25-35 mm diam., outer

tunies not always persisting but then greyish to blackish,

leathery. Leaves suberect, 5-11, synanthous, oblong-lan-

ceolate, ± half as long as inflorescenee, (150-)200^00

X (8-)10-25(-30) mm, dark green, soft-textured, gla-

brous or ciliolate on margins. Raceme subcorymbose or

cylindrical, compact, (50-)70-150 mm long but elon-

gating to 100-150 mm long in fruit, (10-)15-40(-50)-

flowered, sometimes two per bulb; lowermost pedi-

cels 15-30 mm long, elongating to 35-72 mm in fruit;

bracts pale and papery, ovate, acuminate, lowermost

(1 2-) 15^0 mm long, longer or shorter than lower pedi-

cels in flower and shorter in fruit. Flowers white; outer

tepals ovate-lanceolate, inner obovate-lanceolate, 13-

18(-20) X 6-8(-10) mm. Stamens half as long as tepals;

filaments 5-7 mm long, white, outer linear to nanowly

subulate, inner either subulate or expanded and oblong

in lower half, rarely winged. Ovaiy ovoid, 3-5 mm long,

greenish; style erect, 2-3 mm long. Capsule oblong-

ovoid to obovoid, 3-lobed, obtuse-apiculate, 12-15 mm

long. Seeds angular, irregularly folded, 2. 5-3. 5 mm

long, colliculate to papillate, especially along margins.

Chromosomes: 2n = 12 (De Wet 1957; Pienaar 1963).

Flowering time: September to January. Figures IB, 4.

Bothalia 37,2 (2007)

139

:ro,it:.0($alua oo&ieun Jacq*

let. A. i. Xauve, 1973

Distribution and ecology, endemic to the Eastern Cape,

where it is scattered along the higher ground fringing the

coast, from Umtata in the north to Grahamstown in the

south (Figure 3), occurring in moist or marshy grassland

along streams and in damp valleys, sometimes in dense

colonies. The species is typically evergreen and flowers

throughout the wet summer season.

Discussion: Ornithogalum synanthifolium is typically

a large, often evergreen species 350-950 mm high with

relatively long leaves, usually at least half as long as the

inflorescence. The leaves are soft-textured and bright

FIGURE 4. — Ornithogalum synan-

thifolium F.M.Leight.

green, with the margins glabrous or at most ciliolate. The

plants produce a subcorymbose to cylindrical raceme of

pure white flowers with elongate lower pedicels, 15-30

mm long in flower but lengthening to 35-72 mm in fruit.

It is not uncommon for the plants to produce two inflor-

escences in a single season (e.g. Pienaar 8) or for the

inflorescence of the previous season to persist alongside

that of the current year due to the ± continuous vegeta-

tive growth in the species.

The bulb in O. synanthifolium may not be well devel-

oped as a result of both its evergreen habit and its pref-

140

Bothalia 37,2 (2007)

erence for moist habitats, and the outer bulb tunics may

not persist for the same reasons. When present, the outer

tunics are greyish or blackish and somewhat leathery in

texture. The inner filaments, like those of most species

in the section, are rather variable in shape, ranging from

lanceolate to basally expanded or winged, sometimes

{Taylor 4212) to the degree that they resemble those of

O. thyrsoides. The large capsules, 12-15 mm long and

large seeds, 2-3 mm long, are shared with O. conicum.

Other members in the section have smaller capsules, and

seeds less than 2 mm long.

There is no doubt that the Eastern Cape plants are

quite distinct from O. conicum, which occurs in sea-

sonally dry situations on the Cape Peninsula and adja-

cent parts of the southwestern Cape. The climate here is

Mediterranean and summer-dry, unlike the summer rain-

fall climate in which O. synanthifolium occurs. True O.

conicum invariably produces a well-developed, subglo-

bose bulb and a rosette of short, spreading, densely cili-

ate leaves that are dry and withered at flowering, which

takes place in early summer after the rainy season.

In the wild, Ornithogalum synanthifolium may be

confused with O. dubiurn, the only other species of sec-

tion Aspasia to occur in the Eastern Cape, but the lat-

ter is deciduous, grows in more rocky situations, and

produces a rosette of spreading leaves that are densely

ciliate on the margins and usually shorter than half the

length of the inflorescence. The raceme in O. dubiurn is

also typically shorter and more densely corymbose than

in O. synanthifolium, and the seeds are much smaller, ±

1 mm long.

Histoty. Ornithogalum synanthifolium has been much

misunderstood in the past. The first collection of the spe-

cies appears to have been made by the Rev. F. Baur near

Baziya in Eastern Cape. It was identified as O. lacteum

Jacq. (now regarded as a synonym of O. conicum Jacq.)

by Baker (1897) on account of its awl-shaped inner fila-

ments. Another early collection made by Ernest Galpin,

a local businessman with an intense interest in botany,

probably around the turn of the centui'y, fonned the basis

of O. synanthifolium, which was described by Leighton

(1944) for several collections of plants from the Eastern

Cape that have the inner filaments expanded at the base

but which otherwise closely match those collected by

Baur. The latter she retained in O. lacteum, albeit as a

distinct form restricted to the Eastern Cape.

Ornithogalum synanthifolium was subsequently inclu-

ded in O. conicum by Obenneyer (1978), despite the

fact that the latter species was otherwise restricted to the

extreme southwestern Cape, creating an extraordinary

and inexplicable disjunction of some 800 km between

the two areas of occurrence. This decision was based

largely on the large stature of the plants and their lanceo-

late or minutely expanded inner filaments. Specimens

from the Eastern Cape with more prominently expanded

inner filaments were referred to O. fimhrimarginatum,

here included within O. dubiurn. The inner filaments of

O. .synanthifolium, like so many of the species in this

group, are now known to be variable in shape, either

lanceolate without basal lobes or with an apically lobed,

oblong expansion in the lower portion.

3. Ornithogalum corticatum Mart.-Azorin, sp. nov.

Planta habitu cum O. conico congruens, sed valde dif-

fert tunicis bulbi incrassatis corticem fuscum formanti-

bus facile discedentibus, folds brevibus ad adaxialem

apicem pilosis marginibus dense ciliatis per anthesin

omnino emarcidis, et staminibus interais filamentis

linearibus basi triangularibus carinatis valde dilatatis.

Nomen O. corticatum a characteribus peculiaribus tuni-

corum bulbi corticem formatium proveniens.

TYPE. — Northern Cape, 3220 (Sutherland): Klein

Roggeveld, Skaapberg, (-DC), 12 November 2005, M.

Martinez- Azorin & J. Manning 96 (NBG, holo.; K, MO,

iso.).

Plants (250-)300^50 mm high. Bulb depressed-glo-

bose, 20-30(-35) mm diam., outer tunics thick, hard

and leathery or cartilaginous, pale greyish, brownish

or black, accumulating in loose layers. Leaves spread-

ing, 5-7, dry at flowering, oblong-lanceolate, up to one

quarter as long as inflorescence, 30-60 x (5-) 10-1 5

mm, densely puberulous adaxially in distal half and

densely ciliate on margins. Raceme conical to cylindri-

cal, compact, 30-90 mm long at flowering, elongating

to (35-)70-160 mm in fruit, (6-) 16-3 5 -flowered; lower-

most pedicels 10-12 mm long, elongating up to 15 mm

in fruit; bracts white, petaloid, pale and papery, ovate,

acute or acuminate, usually exceeding pedicels, lower-

most 13-20 mm long. Flowers white; outer tepals ovate-

laneeolate, inner tepals obovate-lanceolate, (13-)14-

18(-19) X (3-)5-7 mm. Stamens ± half as long as tepals;

filaments (6-)7.0-8.5 mm long, white, outer subulate,

inner triangular and keeled, slightly incurved over ovary.

Ovary oblong, obtuse or truncate, (3-)4.0-5.5 mm long,

greenish below but yellow in upper part; style 3^ mm

long, white. Capsule oblong-ovoid, 8-11 mm long, 3-

lobed, apiculate. Seeds angular-pyrifonn and irregularly

folded, 2. 0-2. 5 mm long, papillate-echinulate but echi-

nate on margins. Chromosomes'. 2n = 10 (Pienaar 1963

[as O. lacteum forma nov., Sutherland]). Flowering time:

November. Figures 1C, 5, 6.

Distribution and ecology: endemie to the Roggeveld

and Klein Roggeveld near Sutherland in Northern Cape

(Figure 3), where it favours heavy clay soils derived

from dolerite, growing in open renosterveld shnibland.

On the Roggeveld Escarpment, the species may occur

in dense colonies numbering thousands of plants that

cover large swathes of country. This suggests that it may

be toxic to stock. On the more rocky slopes of the Klein

Roggeveld, plants tend to be more scattered, sometimes

on sandstone pavement along drainage lines.

The old bulb tunics of Ornithogalum cor'ticaturn

are exceptionally firm, accumulating around the bulb

in thick, rigid layers. Cartilaginous or thickly mat-

ted tunics are a feature of other geophytes from the

Roggeveld Escaipment, including Lachenalia cornptonii

(Flyacinthaceae) and Devia xer'ornorpha (Iridaceae). The

region experiences extreme climatic conditions through

the year, with bitterly cold winters, during which tem-

peratures fall below freezing, contrasting with hot sum-

mers when temperatures reach the high thirties, and the

thickened covering of the underground parts may act as

insulation.

Bothalia 37,2 (2007)

141

FIGURE 5. — Ornhhogalum cortica-

tum Mart.-Azorin, M. Martinez-

Azorin & J. Manning 96 (NBG),

A, whole plant. B, foliage; C,

androecium and gynoecium; D,

outer stamen; E, inner stamen;

F, gynoecium; G, seed. Scale

bar: A, B. 10 mm; C-F, 5 mm;

G, 1 mm. Artist: John Manning.

Ornithogalum corticatim flowers in early sum-

mer, at which time the leaves are completely withered.

Shrinkage of the bulb at this time results in the flesh

pulling away from the stiff outer tunics, which form

loose, easily detachable layers.

Discussion: Ornithogalum corticatum is readily dis-

tinguished by its depressed-globose bulb surrounded by

firm, cartilaginous tunics, and rosette of short, spreading

leaves, less than one quarter the length of the inflores-

cence, that are shortly and densely hairy on the upper

surface in the distal half, and thickly fringed on the mar-

gins. These are completely withered at flowering but

the pubescence is still visible. The pure white flowers

are borne in conical or narrowly cylindrical racemes on

short pedicels up to 12 mm long. The relatively large,

strongly echinate seeds are also distinctive. Additional

support for its recognition as a distinct species comes

142

Bothalia 37,2 (2007)

FIGURE 6^ — Oniithogahim cortica-

tum Mart.-Azorin.

from the cytological studies of Pienaar (1963) and Roos

& Pienaar (1966), which identified these plants as a dis-

tinctive fonn of O. lacteum (a synonym of O. conicum)

with the unusual chromosome number of 2n = 10, other

members of the section typically having 2n = 12.

In its narrow raceme Ornithogalum corticatum resem-

bles O. strictum, with which it has been recorded grow-

ing on the Klein Roggeveld, but the tunics in that species

are pale and membranous, the leaves are green at flow-

ering, much longer and glabrous, and the inner filaments

are oblong in the lower half The short, densely fringed

leaves of O. corticatum recall those of O. conicum and O.

duhium but in both these species the leaf surface is gla-

brous (rarely apically pubescent in O. dubium) and the

tunics, although leathery, are not as finn-textured as in O.

corticatum. These two species also differ from O. cortica-

tum in details of the flowers. The filaments in O. conicum

are linear and erect, not curved over the ovary, and O.

duhium is characterized by a subcorymbose inflorescence

with longer, lower pedicels, and inner filaments that are

typically expanded and oblong or winged in the lower

part, not triangular. The small seeds of O. dubium, 0.5-1

mm long vs 2.5 mm also serve to distinguish the species.

Bothalia 37,2 (2007)

143

History: although first gathered almost seventy years

ago by Pauline Bond in 1939, the early collections of

Ornithogaliim corticatum lack the characteristic bulb

and were thus referred to O. conicutn on account of their

narrow filaments. Later collections by the horticulturist

Harry Hall were assigned to O. fimbrimarginatum, now

included in O. dubiiim. It is now obvious that they repre-

sent a distinct species.

Series 2 Thyrsoides

Outer bulb tunics membranous, pale; flowers with or

without dark centre; leaves suberect, stem-clasping below;

seeds 1.5-2. 5 mm long.

4. Ornithogalum strictum L.Bolus in Journal of

Botany, London 71: 72 (1933a). O. conicutn Jacq. van

strictum (L.Bolus) F.M.Leight.: 104 (1944). O. conicum

Jacq. subsp. strictum (L.Bolus) Oberm.: 334 (1978). Type;

South Afiica, [Northern Cape], Vanrhyn’s Pass, September

1931, P. Ross Frames BOL20072 (BOL, holo.!).

Plants (300-)500-800(-1000) mm high. Bulb subglo-

bose, (8-)ll-25(-30) mm diam., outer tunics whitish,

membranous. Leaves suberect to erect, 3-8, synanthous,

reaching halfway or more up flowering stem, often to base

of raceme, oblong-lanceolate, attenuate, clasping stem in

lower part, (80-)130-300(-350) x (6-)8-25(-35) mm, gla-

brous but slightly to densely ciliate along margins. Raceme

narrowly conical, compact, (50-)70-150 mm long at flow-

ering, elongating to (70-)100-200(-300) mm in fruit,

(9-)15^0(-65)-flowered; lowermost pedicels 5-14 mm

long, elongating to 9-20 mm in fruit; bracts petaloid, pale

and papery, ovate, acute or acuminate, usually exceeding

pedicels, lowermost 15-30(-45) mm long. Perianth pure

white, rarely with small brownish stain in centre; outer

tepals ovate-lanceolate, inner obovate, (15-)16-22(-25)

X 6-10 mm. Stamens half as long as tepals; filaments fili-

form to subulate, (5-) 7-10 mm long, white, inner usually

with square basal expansion in lower V^-V^, sometimes

bilobed or slightly winged above. Ovary oblong, 3-6 mm

long, truncate, 3-lobed, usually yellow or greenish yellow,

rarely dark greenish; style white, often deflexed, 3-6 mm

long. Capsule fusiform to oblong-ovoid, apiculate, 10-12

mm long. Seeds angular-pyriform and irregularly folded,

1. 5-2.0 mm long, colliculate to rugulose but echinulate

on margins. Chromosomes: 2n = 12 (Pienaar 1963 [as

O. conicum van strictum]). Flowering time: September-

November. Figures ID, 7.

Distribution and ecology: common along the Bokke-

veld Mountains around Nieuwoudtville and through the

Olifants River Valley, with outlying populations at the

foot of the Piketberg and in the Ceres Valley in the south,

and scattered collections from the Roggeveld and Klein

Roggeveld in the east, from 100-800 m above sea level

(Figure 8). The species occurs mainly on fine-grained clay

or loam, typically on moist slopes or in drainage lines or

vleis, where it can occur in large concentrations. On the

Klein Roggeveld the species grows in washes along sea-

sonal streams.

Ornithogalum strictum is the common white chinche-

rinchee in the Olifants River Valley, sometimes forming

large swathes in seasonal vleis on the valley bottom.

Discussion: Ornithogalum strictum is recognized by

its thin-textured, pale grey or whitish bulb tunics, long,

erect leaves that often reach to the base of the raceme

and are green at flowering, and by the narrowly conical

raceme with short lower pedicels, typically less than 15

mm long. The flowers are usually pure white, although

populations near Citrusdal may include individuals with

a brownish centre. The ovary is mostly yellow or yellow-

ish green and the style is typically deflexed in a highly

characteristic manner unique to the species. The inner

filaments are usually widened in the lower third into an

oblong expansion, sometimes bilobed above but not dis-

tinctly winged. As in most species of section Aspasia, O.

strictum shows some variation in the degree of expan-

sion of the inner filaments. Plants from the Bokkeveld

Escarpment have the irmer filaments invariably expanded

in the basal third but populations from further south, in

the Olifants River Valley and around the Piketberg, may

have almost linear or subulate inner filaments.

Ornithogalum strictum resembles O. thyrsoides in

its pale outer tunics, slender leaves clasping the base of

the stem, and expanded inner filaments, and both seem

to occupy similar, seasonally moist habitats. O. thyrsoi-

des can usually be distinguished by its subcorymbose

to broadly cylindrieal inflorescence with longer lower

pedicels, especially in fruit, and inner filaments that are

broadly expanded in the lower half (not third) and always

conspicuously winged, with the wings curved over the

ovary, which is typically dark greenish or blackish.

The distinction between the two is not always clear,

however, especially in the Olifants River Valley, where

occasional plants with the narrowly cylindrical raceme of

Ornithogalum strictum have winged irmer filaments like

those of O. thyrsoides. A similar situation arises further

north in Namaqualand. An alternative treatment would

be to regard O. strictum as the northern subspecies of O.

thyrsoides but intermediate types are rare and we prefer

to maintain the two taxa as separate species, especially in

view of the unusual declinate style that appears to char-

acterize O. strictum.

History: described by Louisa Bolus in 1933 from

plants collected on Vanrhyn’s Pass on the Bokkeveld

Escarpment, Ornithogalum strictum was included in O.

conicum by subsequent authors (Leighton 1944; Ober-

meyer 1978), although it continued to be distinguished

at an inffaspecific level by its slender, narrowly conical

raceme. The decision to include it in O. conicum was

made on account of the relatively weakly expanded inner

filaments, which are quite unlike the broadly winged fila-

ments that are diagnostic of O. thyrsoides. It is now clear

that O. conicum, with its large bulb, short, densely ciliate

leaves, and larger seeds, is quite a different species, and

on vegetative and seed characters it appears that O. stric-

tum is actually more closely allied to O. thyrsoides.

5. Ornithogalum thyrsoides Jacq., Hortus botani-

cus vindobonensis 3: 17 (1776). Type: South Africa,

Cape, without precise locality, in Jacq., Hortus botanicus

vindobonensis 3: t. 28 (icono.!).

O. coarctatum Jacq. 2: t. 435 (1795). Type: South Africa, Cape,

without precise locality, in Jacq., leones plantarum rariorum 3: t. 435.

(icono.!).

144

Bothalia 37,2 (2007)

FIGURE 7. — Oniithogalum strictum

L. Bolus.

O. bicolor Flaw.: 177 (1803). Type: South Africa, [Western Cape],

Cape of Good Hope, ex Whitley’s Nursery (type not traced).

O. hermannii F.M.Leight.: 71 (1933b). Type: South Africa, [Western

Cape], Clanwilliam Dist., 9 miles [14.4 km] N of Citrusdal, 1 October

1932, T.M. Salter 2806 (BOL, holo.!; K, iso.!).

Plants (1 10-)150-600(-700) mm high. Bulb subglo-

bose, comparatively small, 7-20(-23) mm diam., outer

tunics usually whitish, sometimes greyish or brownish,

membranous. Leaves erect or suberect, 3-7(-9), synan-

thous or withered at anthesis, narrowly oblong-lanceo-

late, attenuate, 100-300(-380) x 5-20(-25) mm, clasp-

ing stem below, glabrous but minutely to densely ciliate

on margin. Raceme conical-cylindrical to subcorymbose,

(40-)45-80(-95) mm long at flowering, elongating to

80-150(-200) mm in fruit, (6-)8^0-flowered, some-

times two per bulb; lowemiost pedicels (8-) 14-24 mm

long, elongating to (8-)15-50(-70) mm in fruit; bracts

pale, ovate, acute or acuminate, shorter or longer than

pedicels, lowennost 15^0(-50) mm long. Flowers

milky white or pure white, usually with small greenish

or brownish centre; outer tepals ovate-lanceolate, inner

obovate, 12-25(-27) x (6-)8-12(-15) mm. Stamens half

as long as tepals, sometimes shorter; filaments (4-)5-8(-

1 1 ) mm long, white, outer subulate and slightly broader

at base, inner greatly expanded in lower two thirds

Bothalia 37,2 (2007)

145

FIGURE 8. — Known distribution of Ornithogalwn strictum.

with paired, wing-like outgrowths curved inwards over

ovary. Ovary ovoid, obtuse, 3-6 mm long, grey, green or

brown; style erect, 2. 5^.0 mm long, white or pale green-

ish. Capsule fusiform to oblong-ovoid, (9-) 10-1 5(16)

mm long, 3-lobed, apiculate. Seeds ovoid-angular and

irregularly folded, 1. 5-2.0 mm long, rugulose or collicu-

late and then echinulate on margins. Chromosomes: 2n =

12 (Neves 1953; De Wet 1957; Pienaar 1963). Flowering

time: late September to early November. Figures IE, 9.

Distribution and ecology’: widespread through the

southwestern Cape, from Bredasdorp in the south to

Vanrhynsdorp on the West Coast, with outlying popula-

tions recorded from further north on the higher ground

in central Namaqualand, between Caries and Steinkopf

(Figure 10). The species is typically found at lower alti-

tudes and has not been recorded from the Bokkeveld,

Cedarberg or Cold Bokkeveld Mountain ranges. We have

not encountered it in the Warm Bokkeveld either, and the

only known collection from Ceres {Walters 904) prob-

ably refers to the general region and not the town itself.

The species is common on shale or loamy soils, espe-

cially in seasonally moist sites and ditches along road-

sides. It appears to flourish under slight disturbance or

overgrazing, and in such conditions may occur in dense

colonies numbering hundreds or thousands of plants.

Ornithogalum thyrsoides is the most commonly

encountered species in the southwestern Cape, especially

along the west coast between Piketberg and Gordon’s

Bay, and is the original florists’ chincheriiichee.

It is known to be highly toxic to stock (Van Wyk et al.

2002), which may contribute to its rather weedy propen-

sity. The toxic compound in the plants has been identi-

fied as prasinode G, a steroid glycoside (Van Wyk et al.

2002).

Discussion: Ornithogalum thyrsoides is characterized

by its relatively small bulb with pale, membranous outer

tunics, and slender, erect leaves that clasp the base of the

stem. The outer filaments are slender and awl-shaped but

the inner filaments are conspicuously expanded in the

lower two-thirds, the expansions with prominent apical

wings that are curved over the dark greyish to brownish

ovary. O. thyrsoides is usually readily distinguished from

other species, including O. conicum, by the combination

of thin-textured, pale tunics and winged filaments but

may be difficult to differentiate from O. strictum in the

Olifants River Valley where the two species co-occur. O.

strictum typically has a longer, narrower raceme, inner

filaments that are only expanded in the lower third and

not winged, and an ovary that is usually yellow or pale

greenish with a deflexed style.

Further south, in the Swartland and Boland, Ornitho-

galum thyrsoides occurs with O. dubium but that species

is distinguished by its black outer tunics, shorter, more

densely cilate leaves and often shorter style. In addition,

the filaments in O. dubium are often thicker-textured,

and the outer may also be somewhat expanded rather

than awl-shaped. Around Tulbagh, where the two spe-

cies co-occur, O. thyrsoides favours moister, loamy soils

while O. dubium prefers drier, stonier situations. Careful

examination of mixed populations has failed to reveal

a single hybrid plant. In the Warm Bokkeveld around

Ceres, O. thyrsoides is replaced by O. ceresianum.

Although vegetatively similar and occupying similar

habitats, O. ceresianum is recognized by the very large,

dark centre to the flower, covering the lower half of the

tepals, and by having all six filaments winged, not just

the inner three.

The species is rather variable in the shape of the inflor-

escence, which may range from narrowly cylindrical to

subcorymbose, and in the size of the flowers. Many of

these forms were accorded species status by Leighton

(1944) but we prefer to treat them as part of the nor-

mal variation of the species. Among the more distinc-

tive local forms are those from Langebaan and Saldanha

{Boucher 6476, 6477', Craven 175), which are anoma-

lous in their dark, brownish tunics. A particularly attrac-

tive form, with dense, subcorymbose racemes of flowers

with distinctive, brownish centres occurs around Darling

and Saldanha. In the Olifants River Valley some individ-

uals have unusually long, narrowly cylindrical racemes

(e.g. Boucher 2617), thus resembling O. strictum. The

dark ovaries or bases to the tepals, strongly winged inner

filaments, and erect styles, however, are more consistent

with O. thyrsoides. Further north, populations between

Klawer and Kamieskroon are robust, with arcuate, rather

woody pedicels in fruit. Finally, Barker 3332 from Aries

Kraal near Elgin includes plants with unusually short

styles and scarcely winged inner filaments. Although

resembling O. dubium in these features, the long, pointed

leaves and small bulbs of these plants are more consis-

tent with O. thyrsoides. It is possible that occasional

hybridization may be the origin of some of these anoma-

lous plants.

History: Ornithogalum thyrsoides was among the ear-

liest species in the genus to be described from southern

Africa and the first in section Aspasia. The winged inner

filaments led Baker (1897) to conflate the species with

O. dubium, and even Obermeyer (1978) was misled into

treating O. gilgianum from Bainskloof (here included in

O. dubium) as a synonym despite its dark tunics. Since

the revision of Leighton (1944), however, the species

has mostly been correctly understood.

146

Bothalia 37,2 (2007)

>ir

FIGURE 9. — Ornilhogalum thyrsoi-

des Jacq.

6. Ornithogalum ceresianum F.M.Leight. in

Journal of Botany, London 71: 72 (1933a). Type: South

Africa, [Western Cape], near Ceres, November 1931,

Cook suh Nat. Bot.Gard. 1756/25 (BOL, holo.!; K, iso.!).

Plants 200-350 mm high. Bulb subglobose, (1 1-)13-

20 mm diam., outer tunics whitish, sometimes brown-

ish grey, somewhat leathery or membranous. Leaves

3-7, suberect, more than half as long as inflorescence,

sometimes reaching to base of raceme, partially or

mostly dry at flowering, oblong-lanceolate, 40-90( 1 20)

X 7-14(-18) mm, attenuate, clasping stem in lower part,

glabrous. Raceme subcorymbose to conical, compact,

30-80 mm long at flowering, elongating to 60-100 mm

long in fruit, 7-15(-20)-flowered, sometimes two per

bulb; lowermost pedicels 27-37 mm long, elongating

to 30-40 mm in fruit; bracts pale brownish and papery,

ovate, acute or acuminate, lowermost 20-26 mm long,

shorter than lowermost pedicels. Flowers creamy white

with large, dark olive green central stain covering basal

third of tepals, outer tepals ovate-lanceolate, inner

obovate lanceolate, 16-21 x 7-H mm. Stamens half as

Bothalia 37,2 (2007)

147

FIGURE 10. — Known distribution of Omithogabim thyrsoides, •; O.

ceresianum, O.

long as tepals; filaments 7-9 mm long, white, expanded

and winged in lower two thirds, outer with shorter nar-

row wings, inner with longer wings. Ovary obovoid,

4-6 mm long, truncate, 3-lobed, glossy dark green to

black in upper half, greyish below; style erect, 3. 0-3. 5

mm long, black. Capsule oblong-ellipsoid, 3-lobed,

10-12 mm long. Seeds angular-tetrahedral, 1. 5-2.0 mm

long, rugulose. Chromosomes: 2n = 12 (Pienaar 1963).

Flowering time: October, rarely November. Figures IF,

11.

Distribution and ecology: a local endemic of the

Warm Bokkeveld around the town of Ceres (Figure 10),

where it occurs in loamy soils in seasonally moist vleis

and along seasonal streams, usually in dense colonies.

Discussion: although vegetatively similar to Ornitho-

galum thyrsoides in its small bulb with pale, membra-

nous tunics and slender, suberect leaves clasping the

stem at the base, O. ceresianum is distinguished by the

large, dark olive central eye that covers the basal third

of the tepals, the glossy black ovary, and by having all

six filaments expanded and winged in the lower two

thirds. The inner filaments are more broadly winged

than the outer and in some individuals may be petaloid

with a dark base. In O. thyrsoides the outer filaments are

at most slightly expanded, the dark centre of the flower

is much smaller, covering no more than the base of the

tepals, and the ovary is ovoid and matt. In addition the

racemes are typically longer, with the 16wer pedicels

shorter in fruit.

This striking species deserves to be in cultivation,

and there is little doubt that it can be as easily grown as

Ornithogalum thyrsoides.

History: first collected in the early years of the twen-

tieth century, Ornithogalum ceresianum was described

some twenty years later by Frances Leighton (1933a)

but was subsequently treated as merely an unusual

form of O. thyrsoides (Obermeyer 1978). Having seen

the taxon in the wild, however, we are convinced that

it is sufficiently distinct from O. thyrsoides in several

respects to warrant recognition as a distinct species.

Series 3 Maculata

Outer bulb tunics often dark; flowers white, yellow or

orange, with or without dark centre; seeds 0. 5-1.0 mm

long.

7. Ornithogalum dubium Houtt. in Natuurlijke

historic 2,12: 309, t. 82, fig.3 (1780). Type: South

Africa, Cape of Good Hope, without precise locality, in

Houttuyn herbarium (G, holo.-photo.!).

O. flavissimum Jacq.: t. 436 (1789b). Type: South Africa, without

precise locality, in Jacq., leones plantarum rariorum 2: t. 436 (icono.!).

O. fiavescens Jacq.: 20, t. 437 (1789b). Type: South Africa, without

precise locality, in Jacq., leones plantarum rariorum 2: t. 437 (icono.!).

O. miniatnm Jacq.: t. 438 (1789b). Type: South Africa, without pre-

cise locality, in Jacq., leones plantarum rariorum 2: t. 438 (icono.!).

O. aureum Curtis: t. 190 (1792). Type: South Africa, without pre-

cise locality, in Curtis’s Botanical Magazine: t. 190 (icono.!).

O. vandermerwei Barnes: 14 (1931). O. miniatum var. vandermer-

wei (Barnes) F.M. freight.: 89 (1944). Type: South Africa, [Western

Cape], 8 miles [12.8 km] from Bonnievale on road to Swellendam, 8

October 1929, N.J.S. van der Merwe BOL1814/29 (BOfr, holo.!).

O. vandermerwei Barnes var. album Barnes: 14 (1931). Type: South

Africa, [Western Cape], between Worcester and Robertson, October

1930, Hurling & McNeil BOLI9296 (BOfr, holo.!).

O. fergusoniae fr. Bolus: 57 (1932). Type: South Africa, [Western

Cape], near Still Bay, November-December 1931, E. Ferguson

NBG45/31 (BOfr!, lecto., designated in Obermeyer 1978; K, isolecto.!).

O. brownleei F.M. freight.: 62 (1933b). Type: South Africa, Eastern

Cape, King William’s Town Div., Middledrift, without date, Brownlee

BOL20366 (BOL, holo.!).

O. leipoldtii fr. Bolus: 71 (1933a). Type: South Africa, [Western

Cape], Clanwilliam Div., between Klawer and Clanwilliam, Olifants

River Valley, April 1929, C.L. Leipoldt BOL19941 (BOfr, holo.!; K,

iso.!).

O. subcoriaceum fr. Bolus: 55 (1934), syn. nov. Type: South Africa,

[Northern Cape], near Nieuwoudtville, L. Bolus BOL20090 (BOfr,

holo.!).

O. alticolum F.M. freight.: 93 (1944). Type: South Africa, [Western

Cape], Paarl Div., top of Dutoitskloof, without date, Pillans 8384 (BOfr,

holo.!).

O. pillansii F.M.freight.: 96 (1944). Type: South Africa, [Western

Cape], Piketberg Div., near Het Kruis, 29 September 1943, Leighton

134 (BOfr, holo.!; K!. PRE, iso.).

O. fimbrimarginatum F.M.freight.: 105 (1944), syn. nov. Type:

South Africa, [Western Cape], Montague, Whitehill Ridge, 25 October

1944, F.M. Leighton 273 (BOfr, holo.!; K. iso.!).

O. citrinum Schltr. ex Poelln.: 22 (1944). Type: South Africa,

[Western Cape], Genadendal, without date, Schlechter 9797 (B, holo.;

GRA, K!,PRE, iso.).

O. gilgianum Schltr. ex Poelln.: 23 (1944), syn. nov. Type: South

Africa, [Western Cape] Bainskloof, 3000' [915 m], 12 November 1896,

without date, Schlechter 9131 (B, holo.; BOfr!, K!, PRE, iso.).

O. perpulchrum Schltr. ex Poelln.: 24 (1944), syn nov. Type: South

Africa, [Western Cape], Rietfonteinpoort (10 miles [16 km] SSE of

Elim), 10 December 1896, Schlechter 9681 (B, holo.; BM, BTU, GRA,

K!, NSW, PRE, S, Z, iso.).

Plants (50-)100-450(-650) mm high. Bulb subglo-

bose, (7-)10-20(-25) mm diam., outer tunics leathery

to cartilaginous, dark brown or blackish. Leaves spread-

ing, rarely suberect, (2)3-7(-9), green or dry at an the-

148

Bothalia 37,2 (2007)

thyraol4«

:auve

NATIONAL BOTANJC GARDENS Oi

SOOTH AFRICA.

FIGURE 1 1, — Ornilhogaliimceresia-

mim F.M.Leight.

sis, oblong-lanceolate to ovate, (20-)25-160(-200) x

(3-)5-l 8(-20) mm, obtuse to acute, glabrous but thinly

to densely ciliate on margins, rarely ciliae extending

onto adaxial surface near apex and blade thus puberu-

lous apically. Raceme corymbose to subeorymbose,

rarely shortly cylindrical, (15-)20-60(-100) mm long

at flowering, elongating to 30-100(-150) mm in fruit,

(2)3-20(-30)-flowered; lowermost pedicels (8-) 10-35

mm long, elongating to (12-)15^5(-55) mm in fruit;

bracts greenish or whitish but dry and brown apieally,

ovate, acute or acuminate, usually shorter than pedicels.

sometimes ciliate along margins in upper third or near

apex, lowermost (6-)10-30(-35) mm long. Flowers

orange, yellow, cream-coloured or white with a brown-

ish or greenish centre, or plain orange or yellow; outer

tepals ovate-lanceolate, inner obovate, (7-)10-20(-23)

X (3-)5-ll(-13) mm. Stamens one third to half as long

as tepals; filaments (3.5-)4.0-6.0(7.0) mm long, usually

with brown or blackish stain in middle, or upper part

orange, yellow or brownish, sometimes entirely white,

often thick-textured, usually outer subulate and inner

broader and winged but sometimes all subulate to trian-

Bothalia 37,2 (2007)

149

gular or variously expanded and winged. Ovary ovoid,

(2.5-)3.0-6.0 mm long, blue-greyish, greenish or yellow,

usually narrowing into style, sometimes truncate; style

short and thick or longer and slender, (1.5-)2.0-3.5(-

4.0) mm long, white, yellow or blackish, erect. Capsule

fusiform to oblong-ovoid, (5-)8-15 mm long, 3-lobed,

apiculate. Seeds angular-pyriform, 0.5-1 .0 mm long,

echinulate to long-echinate. Chromosomes'. 2n = 10, 12

(De Wet 1957), 12, 12 + 5B, (18, 24) (Pienaar 1963).

Flowering time'. September-November at lower altitudes

but December-January at higher altitudes. Figures IG, 12.

FIGURE 12. — Ornithogalum dubiwn

Houtt,

Distribution and ecology, widespread through the

southwestern and southern Cape, from the Bokkeveld

escarpment southwards to the Cape Peninsula and thence

eastwards along the coast and through the Little Karoo

as far as Kentani and Stutterheim in the Eastern Cape

(Figure 13). The species typically grows on stony flats

or rocky slopes in pockets of shallow soil or humus in

rock crevices, sometimes along rocky stream banks, in

a wide variety of soils, from fine-grained clay to limestone

or sandstone and in a range of vegetation types, including

renosterveld and fynbos.

150

Bothalia 37,2 (2007)

28°-

30°-

32°-

FIGURE 13. — Known distribution of Ornithogalum dubitim: white-flowered populations, O; yellow-flowered populations, •; yellow- and orange-

flowered populations, 3.

Discussion: as circumscribed here, Ornithogalum

dubiiun is defined by the firm-textured, dark brown to

black bulb tunics, spreading, rather blunt leaves, seldom

up to half as long as the infloresence and typically densely

ciliate on the margin, and corymbose to subcorymbose

raceme of white, yellow or orange flowers, usually with a

dark centre. The filaments are at least partially concolor-

ous with the tepals, often with a dark zone in the middle or

upper part, and the dark-coloured ovary typically narrows

into the style, which may be shorter than or subequal to the

ovary. The seeds are minute, 0. 5-1.0 mm long, and echinu-

late. Similar small seeds are characteristic of all members

of series Maculata.

Yellow- or orange-flowered plants with dark outer bulb

tunics and ciliate leaves are readily assigned to Ornitho-

galum dubium but white-flowered individuals may pose

some problems. The dark tunics, relatively short, spreading

leaves and corymbose inflorescence serve to distinguish

such plants from O. thyrsoides and O. strictum, which also

have larger seeds, 1.5-2 mm long. O. pruinosum, from

further north in Namaqualand, has similar dark tunics and

small seeds but the leaves are distinctly glacous and usually

glabrous or minutely puberulous on the margins. O. cor-

ticatum from the Roggeveld Escarpment is distinguished

by its thick, almost corky bulb tunics, leaves that are dis-

tinctly pubescent on the upper surface near the ends, plain

white tepals and filaments, and yellow ovary. On the Cape

Peninsula, O. conicum is recognized by its large bulb and

linear-filiform filaments and larger seeds. In the Eastern

Cape, white-flowered O. dubium is distinguished from O.

synanthifolium by its deciduous habit, shorter leaves with

ciliate margins, subcorymbose inflorescence and much

smaller seeds (0. 5-1.0 mm vs 2-3 mm).

Ornithogalum dubium is easily the most variable spe-

cies in the section, displaying almost the entire range of

flower colours and filament forms found among the spe-

cies of sect. Aspasia. Populations are often constant in

flower colour, comprising plants that are either white-flow-

ered, or yellow- to orange-flowered, sometimes with less

than one kilometre separating populations of different

colours. However, varicoloured populations are known

from throughout the range of the species. We have seen

a good example of such a population at Hottentotskloof,

east of Ceres, comprising individuals displaying the entire

range of different flower colours, from orange and yellow

through cream-coloured to white with brownish centres.

Plants that match the type of Ornithogalum dubium

and from which the original collections are most likely

to have been made, occur around the town of Tulbagh at

the head of the Breede River, on the original track into

the interior. These populations have bright yellow or

orange flowers and orange filaments, the inner expanded

at the base and apically pointed but not winged, and

short styles. Populations from the Swartland west of

Tulbagh in contrast, are white-flowered with brownish

orange centres. The filaments in these plants are darkly

marked in the middle, and the inner are slightly winged.

These populations often co-occur with O. thyrsoides,

which is common on the coastal side of the mountains,

but hybrids between them have not been seen.

Further down the Breede River Valley, between

Worcester and Robertson, populations may be either white-

or orange-flowered, and the inflorescences, although usu-

ally corymbose, may be longer and more conical. In some

of these plants all six filaments are winged, the outer fila-

ments with acute, divergent wings and the inner with lon-

ger, convergent wings. Both filament whorls are often

stained dark in the middle or in the upper parts. A simi-

lar degree of variation is evident in the Little Karroo and

southem Cape, where populations may be white- or yel-

low-, or sometimes orange-flowered, and the filaments

vary from almost linear to winged.

Yellow-flowered populations are known from south

of the Riviersonderend Mountains, between Caledon and

Bredasdorp, and formed the basis of Ornithogalum citri-

num, described by Von Poellnitz from plants collected

by Rudolph Schlechter at Genadendal in the southem

Bothalia 37,2 (2007)

151

foothills of the Riviersonderend Mountains. Plants from

higher altitudes throughout the mountains of the south-

western and southern Cape are invariably white-flow-

ered, typically with longer styles, and were previously

segregated as O. fimbrimarginatum and O. subcoria-

ceum.

Although white-flowered, longer-styled plants typi-

cally occur on sandstone soils throughout the moun-

tains of the southwestern Cape, and short-styled plants

on inter-montane shale soils east of the Bokkeveld-

Hottentots Holland Mountain axis, this ecological-

geographical differentiation is not absolute. Yellow-

and orange-flowered plants occur, for instance, along

the Tsitsikamma Mountains and in the Longkloof on

sandstone substrates. Plants from the Witteberg near

Laingsburg, including the type of Ornithogalum fimbri-

marginatum, are long-styled but otherwise match typical

O. dubiiim perfectly in their dark centre, blackish green

ovary, and filaments with the outer portion flushed olive-

green. Apart from the length of the style there is nothing

to separate these plants from white-flowered O. dubium.

It is now apparent that there are all transitions between

short-styled and longer-styled plants, and that style

length alone is an insufficient basis for distinguishing

between species in the O. dubium complex. In the light

of the extensive variation in flower colour and filament

morphology that is now known to occur within even O.

dubium sensu stricto, we therefore prefer to recognize

a single, variable species in the complex pending more

thorough investigation into the population genetics in

the group.

The status of Ornithogalum subcoriaceum in particu-

lar deserves further study. This species was recognized

for small plants, rarely up to 200 mm tall, from the inte-

rior mountains of the West Coast, typically bearing rather

small flowers with tepals 8-14 mm long and bracts that

are ciliate towards the apex. In addition, the seeds of

collections that we have been able to examine are papil-

late rather than echinulate as is typical in O. dubium/0.

fimbrimarginatum. These distinctions, however, are not

absolute and larger plants from the Cedarberg that are

referable to O. dubium/0. fimbrimarginatum on this

basis may also have ciliate bracts and papillate seeds.

We thus provisionally include the smaller plants within a

broadly circumscribed O. dubium.

Ornithogalum dubium, unlike O. thyrsoides, has been

shown to be non-toxic (Obermeyer 1978).

History’-. Ornithogalum dubium, described by Houttyn

in 1780, was the second member of segtion Aspasia

known to science after O. thyrsoides. Other yellow- and

orange-flowered plants formed the basis of Jacquin’s

(1789b) O. fiavescens, O. fiavissimum and O. miniatum.

All of these were treated as varieties of a very broadly

defined O. thyrsoides by Baker (1897). The first modem

treatment of the species is that of Leighton (1944) but

the high degree of variation in flower colour and fila-

ment morphology evident in this widespread species led

her to distinguish almost a dozen different taxa, based on

small differences in the shape and size of the filaments

and length of the style. Many of these species were

reduced to synonomy under O. dubium by Obermeyer

(1978), who defined the species more broadly to include

all plants with dark bulb tunics, ciliate leaves, and yel-

low, orange or white flowers with a dark centre and short

style. Plants with dark tunics and white flowers with lon-

ger styles were excluded from this circumscription and

treated as separate species, either O. subcoriaceum or O.

fimbrimarginatum .

Ornithogalum subcoriaceum was described by

Louisa Bolus in 1934 from small plants collected on

the Bokkeveld Escarpment near Nieuwoudtville that

had ciliate bracts near the apex. Although maintaining

the species in her treatment of the genus, Obermeyer

(1978) recognized its similarity to O. dubium, and sug-

gested that the dwarfing might be the result of the colder

climate at high altitudes. Plants that match the type of

O. subcoriaceum occur throughout the mountains of the

southwestern Cape inland of the West Coast, from near

Nieuwoudtville to Ceres. Ornithogalum fimbrimargi-

natum, described by Leighton in 1944 from plants col-

lected at Whitehill near Matjiesfontein, has never been

well understood, and Obermeyer (1978) even considered

the possibility that it was a hybrid between O. dubium

and O. conicum, inheriting the dark tunics and expanded

inner filaments from the former, and the white flowers

and long style from the latter.

Ornithogalum dubium was separated from O. fimbri-

marginatum and O. subcoriaceum by Leighton (1944) and

Obermeyer (1978) on the length of the style, which was

taken to be less than 2 mm long in O. dubium and more

than 2 mm long in O. fimbrimarginatum and O. subcoria-

ceum. Long-styled plants with dark tunics and white flow-

ers are scattered throughout the mountains of the south-

western and southern Cape, typically growing in moist,

shallow soils in sandstone outcrops. Populations with

the outer filaments variously broadened were referred

by Leighton (1944) severally to O. leipoldtii, based on

plants from Clanwilliam, O. piUansii from Piketberg,

and O. alticolum from Du Toitskloof, whereas those with

the outer filaments linear or lanceolate were determined

respectively as O. fimbrimarginatum or O. subcoria-

ceum. The distinctly winged inner filaments of the plants

referred to O. leipoldtii and O. piUansii have a dark zone

in the middle, thus closely resembling white-flowered

forms of O. dubium, and both were therefore reduced

to synonomy under O. dubium by Obermeyer (1978),

despite the fact that the length of their styles exceeds the

upper limit accepted by her for that species.

There has been some confusion about the identity of

Ornithogalum perpulchrum, described by Von Poellnitz

(1944) from leafless plants collected by Rudolph

Schlechter on 10 December 1896 near Elim in the Over-

berg. Obermeyer (1978) associated these plants with a

collection of O. dubium made in the vicinity on the same

day by Harry Bolus {Bolus 8696), who was collect-

ing with Schlechter at the time, but Miiller-Doblies &

Muller-Doblies (1996) treated O . perpulchrum as a syno-

nym of O. rupestre (as O. multifolium) on the basis that

duplicates of both Schlechter 9681 and Bolus 8696 that

they examined had the leaves of O. multifolium. This is,

however, most definitely not the case with the Bolus col-

lections in both the Bolus and Kew herbaria, which have

the short, oblong, ciliate leaves and dark tunics of O.

dubium. Similarly, we have no hesitation in identifying

152

Bothalia 37,2 (2007)

the Kew isotype of Schlechter’s collection as O. diibhim,

despite the rather small stature of the plants. There is

no doubt that Schlechter’s Rietfonteinkloof is correctly

located near Elim (Jessop 1964), and Ornithogaliim

nipestre (= O. multifolium) is not known from south of

the Langeberg, whereas O. dubium is common here. All

evidence thus confirms that Ornithogalum perpulchrum

is correctly placed in synonomy under O. dubium.

8. Ornithogalum pruinosum F.M.Leight. in Journal

of South Atfican Botany 10: 104 (1944). Type: South

Africa, [Northern Cape], between Garies and Kamieskroon,

September 1934, L.Bolus BOL22781 (BOL, holo.!).

O. glaiicophylliim Schltr. ex Poelln.: 23 (1944). Type: South Africa,

[Northern Cape], Arakup [Arkoep, N of Kamieskroon], 14 September

1897. R. Schlechter 11249 (Breslau, holo.; BOL!, GRA, Kl, PRE, iso.).

Plants (40-)100^00(-650) mm high. Bulb some-

times epigeal or nearly so, subglobose, (8-)10-25(-27)

mm diam., outer tunics dark brown to black, sometimes

reddish apically, firm-textured or leathery. Leaves erect,

(2)3-6(7), oblong-lanceolate to ovate-lanceolate, (20-)

30-200(-300) X (5-)7-25(-35) mm, sometimes crisped,

less than half as long as inflorescence, leathery, glaucous,

glabrous or margins ciliolate. Raceme subcorymbose

to cylindrical, (15-)20-50(-70) mm long at flowering,

elongating to (30-)50-100(-150) mm in fruit, (4— )8^0

(-60)-flowered; lowermost pedicels (5-)6-17(-20) mm

long, elongating to (9-)12-25(-30) in fruit; bracts whitish

or pale greenish, ovate, acuminate, shorter or longer than

pedicels, lowermost (7-) 10-30(^5) mm long. Flowers

glossy white, sometimes with a small greenish or brownish

centre; outer tepals ovate-lanceolate, inner tepals obovate,

(8)9-1 5(-20) X (3)4-9(10) mm. Stamens half as long

as tepals; filaments (3.5-)4.0-7.0(8.0) mm long, white,

outer linear, sometimes slightly widened at base, irmer

either lanceolate or expanded and obovate (sometimes

apically lobed) in lower half. Ovary ovoid, (2.5-)3.0-5.0

(-5.5) mm long, green, yellow or brownish; style erect,

(1.5-)2. 0-3.0 mm long, yellow. Capsule ellipsoid to

oblong-ovoid, (5)6-10 mm long, 3-lobed, apiculate. Seeds

angular-pyriform, 0. 5-1.0 mm long, papillate or echinu-

late. Chromosomes'. 2n = 12 (Pienaar 1963). Flowering

time'. (July) late August to late September or mid October,

rarely into early November. Figures IH, 14.

Distribution and ecology, widespread through Nama-

qualand, especially along the western edge of the escarp-

ment between 500-1 200 m, from the Richtersveld in the

north to Klawer in the south, and extending eastwards

to Calvinia on the Bokkeveld Plateau and Aggenys

in Bushmanland, with two records from as far east as

Kumman and Augrabies in Gordonia (Figure 15). In

the south of its range, in the Knersvlakte, the species

occurs along the western foothills of the escarpment

below the Bokkeveld plateau, and only extends onto the

plateau itself, around Calvinia, from the more arid val-

leys of southern Bushmanland. Plants typically grow on

dry, open stony flats and slopes, and on rocky hillsides,

mostly in clay and granite.

Discussion'. Ornithogalum pruinosum is recognized

by its dark, blackish outer bulb tunics, characteristic

glaucous, leathery, often undulate leaves that are seldom

up to half as long as the inflorescence, and a subcorym-

bose inflorescence of pure white flowers (rarely with a

small dark centre). The yellow ovary and style in many

plants is also diagnostic. The small seeds ± 1 mm long,

and dark, leathery tunics place the species in the group

that includes O. dubium, O. puberulum, O. maculatum

and O. nipestre.

Although centred south of the range of Ornithogalum

pruinosum, in the southwestern Cape south of Klawer,

outlying populations of O. thyrsoides have been recorded

from the higher-lying parts of central Namaqualand, in

the Kamiesberg and near Steinkopf The two cannot be

confused, however, as O. thyrsoides is readily distin-

guished from O. pruinosum by the slender, bright green

leaves, small bulb with pale tunics, and inner filaments

that are conspicuously winged in the lower half and par-

tially obscuring the ovary. In the south, the distribution

of O. pruinosum also approaches that of O. strictum and

O. dubium but the species do not overlap and confusion

between them is unlikely. O. strictum, which is common

in seasonally damp sites along the Bokkeveld Mountains,

has similar flowers but slender, bright green leaves and

cylindrical racemes with short pedicels, and O. dubium,

which has been recorded from similar habitats on the

Gifberg, has bright green leaves that are densely ciliate

on the margins.

Ornithogalum pruinosum is very variable in stature,

ranging in size from small, few-flowered plants scarcely

50 mm high to robust individuals almost 700 mm tall.

Plants from Aggenys in the arid eastern Bushmanland

are especially dwarfed. This variation is to be expected

in a species from arid environments with variable pre-

cipitation. Although usually lanceolate, the inner fila-

ments are occasionally expanded at the base, the expan-

sions with small, pointed apical wings. Plants with these

winged filaments are scattered through the range of the

species, and are known from the Richtersveld, central

Namaqualand and Loeriesfontein. The range in form of

the inner filaments in the species was evident to Leighton

(1944), who noted that they varied in the same popula-

tion or individual, even in the same flower.

History. Ornithogalum pruinosum appears to have

been first collected in the late 19th century, first near

Kamieskroon by R. Schlechter in the spring of 1 897 and

a year later at Okiep by W. Morris. Although Schlechter

intended describing the species under the name O. glau-

cophyllum, it was only formally recognized in 1944,

when it was independently described by both Leighton

and Von Poellnitz, the latter using Schlechter’s material

and his proposed epithet. Since then it has been consis-

tently recognized, and its characteristic glaucous, often

undulate leaves have spared it the further taxonomic

fragmentation on the basis of the shape of the inner fila-

ments that has been the fate of other species from the

southwestern Cape.

9. Ornithogalum maculatum Jacq., Collectanea

3: 368 (1791). Type: South Africa, without precise local-

ity or date, Jacq., Collectanea 3: t. 18, f. 3 (icono.!).

O. maculatum Thunb.: 62 (1794). illegitimate homonym. O. thim-

bergianum Baker: 269 (1873), new name for O. maculatum Thunb.,

not of Jacq. (1791 ). Type: South Africa, [Western Cape], Saldanha Bay,

Thimherg 8289 (UPS, holo.-microfiche!).

Bothalia 37,2 (2007)

153

FIGURE 14. — Ornithogalum pru-

inosmn F.M.Leight.

O. speciosum Baker: 72 (1891), illegitimate homonym. O. insigne

F.M.Leight.: 113 (1933b), new name for O. speciosum Baker, not of

Salisb. (1796) or Rafin. (1810). O. magnificum Poelln.: 214 (1945),

new name for O. speciosum Baker. O. macidatum Jacq. var. speciosum

(Baker) F.M.Leight.: 110 (1944). Type: South Africa, [Northern Cape],

Namaqualand, without precise locality or date, W. Scully 175 (K, holo.!,

SAM, iso.!).

O. thunbergianum var. concolor Baker: 496 (1897). Type: South

Africa, without precise locality or date, Forster s.n. (K, holo.!).

O. splendens L. Bolus: 14 (1931). O. maculatum Jacq. var. splen-

dens (L. Bolus) F.M.Leight.: 110 (1944). Type: South Africa, [Northern

Cape], near Nieuwoudtville, November 1930, Buhr 2654/30 (BOL,

holo.!).

O. rossouwii U.Miill.-Doblies & D.Mull.-Doblies: 394 (1996), syn.

nov. Type: South Africa, [Northern Cape], 2 km N of Buffelsrivier and

32 km N of Laingsburg on Sutherland road. ± 800 m, 4 August 1986

(in leaf), Miiller-Doblies 86037b (PRE, holo., not located; B, BTU, K,

NBG, iso., not located).

Plants (40-)60^00(-550) mm high. Bulb subglobose,

(7-)8-20(-25) mm diam., outer tunics whitish or grey-

brownish, thin-textured. Leaves suberect, (1)2-5, lin-

ear-lanceolate to oblong-lanceolate, (15-)20-110(-170)

X 3-15(-20) mm, obtuse or acute, usually clasping at base,

glaucous and entirely glabrous. Raceme corymbose to sub-

corymbose or shortly cylindrical, (5-) 10-30(^0) mm long

154

Bothalia 37,2 (2007)

FIGURE 15. — Known distribution of Oniithogalum pruinosum.

at flowering, elongating to 25-55(-80) mm long in fruit,

l-6(-9)-flowered; lowermost pedicels 3-15 mm long,

elongating to 5-27 mm in fruit; bracts whitish but brown-

ish in outer part, ovate, acute or acuminate, longer than

pedicels, lowermost (6-)10-20(-30) mm long. Perianth

reddish, orange or yellow (rarely white), outer tepals

usually with black spot or transverse bar near apex,

sometimes unmarked; outer tepals obovate-lanceolate,

inner obovate, tepals (10)ll-25(-27) x 5-14(-18) mm.

Stamens ± half as long as tepals; filaments (4-)5-8(-9)

mm long, usually orange or yellow, subulate or inner

lanceolate. Ovary ovoid, 4-8 mm long, yellow; style

short and thick, 1.5-2. 5 mm long. Capsule fusiform to

oblong-ovoid, 8-10 mm long, 3-lobed, apiculate. Seeds

angular-pyriform or comma-shaped, 1 mm long, rugu-

lose. Chromosomes'. 2n = 12 (24) (Pienaar 1963), 14 (De

Wet 1957). Flowering time', (late August) September to

October. Figures II, 16.

Distribution and ecology', localized to the higher-

lying parts of central Namaqualand between Springbok

and Caries, but widely scattered through the drier parts

of the southwestern Cape, from the Bokkeveld Plateau

southwards through the Cedarberg and Cold Bokkeveld

to Langebaan and Malmesbury on the west coast, thence

eastwards along the fringe of the Cape Floral Region to

Prince Alfred (Figure 17). Plants are typically restricted

to rock outcrops, typically in shallow humus on granite

or sandstone pavements, but also outcroppings of shale.

Discussion'. Ornithogalum maculatum is one of the

most striking species in section Aspasia, readily recog-

nized by its glaucous leaves lacking any marginal vesti-

ture, orange or yellow flowers, usually with some dark

marking at the tips of the outer tepals, and subulate or

lanceolate filaments. Forms with pure yellow or orange

flowers may be confused with O. rupestre but that spe-

cies has narrow, subterete leaves. The minute seeds and

orange flowers might lead to confusion with forms of

O. duhium, but O. maculatum lacks the dark tunics and

ciliate leaves that are characteristic of that species.

Ornithogalum maculatum is very variable in stature,

flower size and in the development of markings on the

flowers. Large forms, with pure orange flowers lack-

ing markings and with broad filaments, originally seg-

regated as O. splendens, occur around Calvinia and in

Namaqualand between Kamieskroon and Flondeklipbaai.

Plants of similar size but with well-marked outer tepals

bearing triangular or diamond-shaped markings were

distinguished as O. speciosum and have been recorded

from Namaqualand, between Springbok and Garies,

from the Olifants River Valley around Clanwilliam,

and from the western Little Karoo, around Karoo Poort,

Touws River and Montagu.

The presence and degree of marking may vary within

a population, which can thus include both marked

and unmarked individuals (e.g. Van der Merwe 246).

Although typically blackish, the markings on the outer

tepals range in colour from green to black, sometimes

within a single population (e.g. Horrocks 12; Hall s.n.

NBG68697). Occasional plants or populations from

Karoopoort { Viviers 1568), Laingsburg (Bayliss 736) and

near Steytlerville may lack yellow or orange pigment,

producing whitish flowers with darker markings. Such

plants formed the basis of O. rossouwii.

Anecdotal evidence suggests that the species is not

toxic to animals since goats have been recorded as con-

suming it with impunity (Obermeyer 1978).

History', first described in 1791 by Nicolaas von

Jacquin from plants grown in Vienna, Ornithogalum

maculatum was redescribed a few years later by

Thunberg from wild material gathered near Saldanha.

Struck by the distinctive marks on the outer tepals, he

chose the same epithet for his species. The species is

now known to vary not only in flower colour but also

in the degree of maculation. Some of the more distinc-

tive of these forms were segregated as distinct spe-

cies or varieties in the past. Well-marked forms from

Namaqualand were segregated as O. speciosum by Baker

(1891), and an attractive form with large, unmarked yel-

low or orange flowers from near Nieuwoudtville was

named O. splendens by Bolus (1931). Both of these

forms were later reduced to varietal status by Leighton

(1944) but we are in accordance with Obermeyer (1978)

in preferring to regard them as local forms within a

much greater spectrum of variation and thus not deserv-

ing of taxonomic rank. Most recently, Miiller-Doblies &

Mtiller-Doblies ( 1 996) described O. rossouwii from five

greenhouse-grown plants collected between Laingsburg

and Janseville, and distinguished from O. macula-

tum by their small size and whitish or pinkish flowers.

Although the type material of this species has not been

deposited in any herbaria, the accompanying illustra-

tions make its identity clear. The differences between

this material and typical O. maculatum are trivial when

compared to the variation within the species. Several

collections of depauperate plants with the yellow, spot-

ted flowers of typical O. maculatum (e.g. Pretorius 92)

are indistinguishable from O. rossouwii apart from the

colour of the flowers, and these white or cream-coloured

plants are most appropriately treated as colour forms of

O. maculatum. A similar range in flower colour is com-

Bothalia 37,2 (2007)

155

HERBARIUM STELLENBOSCH

Northern Bokicevcld: flat* at west

b^ae of Geelberf

S.^indy open fl't.n: eh-illow s'lni on

rock surfaces.

fwo ^oups of plants;

flowftrs onne* with bliek p^itch on

reverse of outer segments.

K.F.Thoapaon -p^] 1089/10

Ornithog«lujr. naoul^tufi Jac<^.

/

det. A. i. Meuv«» 197?

FIGURE 16. — Ornithogalum wacii-

latiim Jacq.

mon within O. diibiim and it is now evident that it also

occurs in O. nipestre.

10. Ornithogalum rupestre L.f., Supplementum

plantarum: 199 (1782). Type: South Africa, [Western

Cape], Malmesbury Division, Witteklip, Thunberg 8302

(UPS, holo.-microfiche!).

O. multifolium Baker; 271 (1873), syn nov. Type: South Africa,

[Northern Cape], Namaqualand. Modderfontein, Whitehead s.n. (TCD,

holo.!).

O. virgineum Soland. ex Baker: 271 (1873). Type: South Africa,

without precise locality [probably Witteklip], Masson s.n. (BM, holo.!).

O. aurantiacum Baker: 748 (1878), syn. nov. Type: South Africa,

[Western Cape], Malmesbury District, Groenekloof [Mamre], 1878, H.

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

O. ranunculoides L. Bolus: 71 (1933b), syn nov. Type: South

Africa, [Northern Cape], Steinkopf, H. Metre STE3978 (BOL!, lecto.,

here designated).

O. witteklipense F.M. height.: 175 (1945). Type: South Africa,

[Western Cape], Witteklip near Vredenburg, Leighton 655 (BOL, holo.! ).

O. saxatile Schltr. ms.

Plants (30-)50-150(-200) mm high. Bulb subglobose,

(7-) 10-1 5 mm diam., outer tunics whitish or brownish.

Leaves (2)3-10, subterete or canaliculate, 20-110 x 1-3

mm, glabrous. Raceme corymbose to subcorymbose,

5-30 mm long at flowering, elongating to 10-50 mm in

fruit, (l)2-10(-15)-flowered; lowermost pedicels 2-18

mm long, elongating to 6-25 mm in fruit; bracts ovate,

whitish with upper part brownish, acute or acuminate.

156

Bothalia 37,2 (2007)

FIGURE 17. — Known distribution of Ornithogalmn macidatum: white-

flowered populations, O; orange-flowered populations, •.

usually shorter than pedicels but longer in small plants,

lowermost (4-)6-15 mm long. Flowers usually yellow

or orange, rarely white or pinkish; outer tepals ovate-

lanceolate, inner obovate, 6-12 x 3. 5-5. 5 mm. Stamens

± half as long as tepals; filaments 3-5 mm long, whitish

or yellowish, outer linear to subulate, inner lanceolate.

Ovary ovoid, 3-4 mm long, yellowish; style short, erect,

1-2 mm long. Capsule fusiform to oblong-ovoid, 10

mm long, 3-lobed, apiculate. Seeds angular-pyriform or

comma-shaped, 0.75 mm long, rugulose. Chromosomes:

2n = 12 (Pienaar 1963), 20 (Johnson & Brandham 1996).

Flowering time: late August to early October. Figures IJ,

18.

Distribution and ecology: widely distributed through

Namaqualand and the more arid southwestern Cape,

from the Richtersveld in the north through the higher-

lying parts of Namaqualand to the Bokkeveld Plateau,

southwards through the northern Cedarberg to Darling

in the southwest, and through the Cold Bokkeveld to

Barrydale and Laingsburg in the southeast, with isolated

records from the Roggeveld Escarpment (Figure 19).

Plants grow in shallow humus and soil on rock sheets,

often on granite or sandstone, where they may occur in

large numbers fringing depressions on the rocks that

accumulate water in the rainy season.

Discussion: Ornithogalum rupestre is one of the

most recognizable species in the section, identified by

the few to many, subterete, glabrous leaves. Other spe-

cies in section Aspasia have plane, oblong to lanceo-

late leaves. It is typically less than 100 mm high, with

pure yellow or orange flowers with linear to lanceolate

filaments and a very short style. O. rupestre shares gla-

brous leaves, mostly yellow or orange flowers with a

rudimentary style, and minute, rugulose seeds with O.

maculatum and it is likely that the two are immediately

related. The flowers of O. rupestre are typically bright

yellow or orange but populations from Langebaan and

the Cold Bokkeveld have pale yellow flowers, and plants

with cream-coloured flowers have been collected around

Pakhuis Pass. The flowers of populations from the

Vredenburg Peninsula are uniformly white or pinkish.

History: Ornithogalum rupestre was described by

Linneaus f (1782) from plants collected on Witteklip

near Vredenburg by Carl Peter Thunberg. Although not

mentioned in the original description, the plants from

Witteklip are uniformly white-flowered, sometimes with

a pinkish flush. Later collections of the more common

and widespread yellow-flowered forms were described

as O. multifolium, based on short, floriferous plants

from Namaqualand (Baker 1873), and O. aurantiacum,

from few-flowered plants collected near Mamre (Baker

1 897). Yet another name, O. ranunculoides, was applied

to taller forms from Namaqualand by Louisa Bolus

(1933b). All of these forms were subsequently included

within a single yellow-flowered taxon, O. multifo-

lium, by both Obermeyer (1978) and Muller-Doblies &

Muller-Doblies (1996). White-flowered O. rupestre from

Vredenburg has until now been retained as distinct from

yellow-flowered O. multifolium. The application of the

name O. rupestre was confused by Leighton (1944), who

used it for yellow-flowered plants (Obermeyer 1978),

but there is no doubt that the type is white-flowered.

White-flowered Ornithogalum rupestre was separated

from yellow-flowered O. multifolium on nothing more

than flower colour, a minute difference in style length,

and the smaller stature of the former. It is now clear

that flower colour varies within the species in the group

and is an insufficient basis for separating species. Even

within yellow-flowered plants, flower colour may range

from pale lemon-yellow through bright canary yellow

to orange, whereas paler forms may be white, cream-

coloured or pinkish. The purported differences in stat-

ure and style length are likewise trivial and we have no

hesitation in including the various colour forms within

a single species. It is unfortunate that O. rupestre is the

earlier name for this taxon, which is better known under

the name O. multifolium.

11. Ornithogalum leeupoortense U.Miill.-

Doblies & D.Miill.-Doblies in Feddes Repertorium

107; 396 (1996). Type: South Africa, Northern Cape,

Bushmanland, Naip Mountains, (-AD), June 1988, E.J.

van Jaarsveld 9478 (NBG!, neotype, here designated).

Plants 60-150 mm high. Bulb subglobose, 10-15

mm diam., outer tunics dark brown to black, somewhat

leathery. Leaves falcate, distichous, (2-)5 or 6, lanceo-

late-attenuate, canaliculate, ± as long as inflorescence

or longer, (25-)40-15 x (4-)5-10 mm, leathery, bright

green, margins translucent and papillate, sometimes

thickened. Raceme often laterally displaced by develop-

ment of second inflorescence, corymbose, 20-50 mm

long at flowering, ( l-)4-15-flowered; lowermost pedi-

cels ( 1 5-)20-30 mm long; bracts pale greenish, lanceo-

late-attenuate, shorter or longer than pedicels, lowennost

( 1 5-)20-25 mm long. Flowers glossy white; outer tepals

elliptic-ovate, inner tepals elliptic-obovate, 13-20 x 5-8

mm. Stamens less than half as long as tepals; filaments

5. 0-6. 5 mm long, white, outer linear or subulate, inner

linear-lanceolate. Ovaiy ovoid, 4-6 mm long, green;

style erect, 2-3 mm long. Capsule ellipsoid to oblong-

ovoid, 6-7 mm long, 3-lobed, apiculate. Seeds angular-

pyriform, 0. 5-1.0 mm long, echinulate. Flowering time:

August to September. Figures IK, 20.

Bothalia 37,2 (2007)

157

Orsitbo^aluM nultifoliuA Bak»

dot, 4. 4. Mauv«, 1975

FIGURE 18. — Ornithogahun rupes-

tre L.f.

Distribution and ecology, currently known from the

hills northeast of Springbok, with most of the collections

from the inselberg Naip se Berg, and from further north

on the higher mountains of the Richtersveld, between

800-1 000 m (Figure 21). Plants grow wedged in crev-

ices and fissures in quartzite rock and scree on sheltered,

south-facing slopes, where they may be locally abundant.

Discussion: still poorly collected and understood,

Ornithogalum leeupoortense is distinguished by the fan-

like arrangement of its leaves, which are falcate, pointed

and somewhat channelled. The leaf margins are obscurely

papillate and in the populations around Springbok they

are also distinctly thickened. Plants typically produce a

second inflorescence in a growing season and the older

inflorescence is characteristically displaced, appearing to

arise in the axil of one of the lower leaves. The relatively

short inflorescence is characterized by large, often foli-

ose, green bracts, the lowermost 15-25 mm long, with

the tips drawn out and attenuate.

Collections from the Richtersveld, although lack-

ing the thickened leaf margins of plants from around

Springbok, match the southern populations in all other

158

Bothalia 37,2 (2007)

essentials, including the falcate, glossy green leaves

with attenuate tips, and lateral displacement of the pri-

mary inflorescence through development of a later one,

and we have no hesitation in regarding them as conspe-

cific.

The minute, echinulate seeds of Ornithogalum leeu-

poortense indicate an alliance with the members of the

O. diibiiim group but it is unlikely to be confused with

any other species in series Maculata on account of the

distinctive glossy green, distichous leaves. Most other

species of the group from Namaqualand, particularly O.

pruinosum and O. maculatiim, have rosulate, glaucous

leaves. Pressed specimens of O. pruinosum may be dis-

tinguished by their erect, proportionally broader leaves

with ciliolate, often crisped margins. The leaves are also

typically shorter, less than half as long as the inflores-

cence, and thick-textured.

History, the species was described by Muller-Doblies

& Muller-Doblies (1996) from a collection of non-

flowering plants made on 11 August 1988 on the Farm

Leeupoort northeast of Springbok and subsequently

grown on in the glasshouse. None of the cited material

has been deposited in any herbarium but three collec-

tions of plants, all from Naip se Berg on the edge of

Bushmanland northeast of Springbok, match the pro-

tologue in the diagnostic essentials, viz. the distichous

leaves with thickened, obscurely papillate margins, and

are accordingly referred to this species. Since no type

material has been lodged in the cited herbaria, we select

one of these specimens to act as a neotype. The earliest

collection of the species that we have traced is that made

by Margaret Thompson and Annelise le Roux in August

1977 in the Richtersveld and until now referred either to

O. subcoriaecum (now a synonym of O. dubium) or O.

pruinosum.

12. Ornithogalum puberulum Oberm. in Bothalia

12: 337 (1978). Type: Namibia, Kahanstal, 5 miles [8

km] N of Loreley [Lorelei], 30 August 1963, H. Mueller

& W. Giess 3365 (PRE, holo.!).

O. merxmueUeri Roessler: 389 (1979). Type: Namibia, Farm

Namuskluft, 29 September 1977, Merxmuller & Giess 32350 (M,

holo.; PRE!, WIND, iso.).

O. puberulum subsp, chhs-bayeri U.Miill.-Doblies & D.Miill.-

Doblies: 390 (1996), syn. nov. Type: Namibia, mountain 2 km west

of Rosh Pinah, ± 450 m, 26 September 1989, Muller-Doblies 89120b

(WIND, holo., not deposited).

Plants 50-80(-90) mm tall. Bulb subglobose, 8-15

mm diam., outer tunics leathery or somewhat cartilagi-

nous, whitish or dark grey, sometimes forming a short

neck of narrow fragments. Leaves suberect or spreading,

2(3), synanthous, oblong to ovate-lanceolate, rarely lan-

ceolate, uppermost usually only half as large as lower,

40-80 X 1 0-25 mm, soft-textured, glabrous or pubescent

on both surfaces, margins with slender cilia 0.75-2.0 mm

long, base shortly tubular, somewhat inflated and clasp-

ing scape. Raceme corymbose or racemose, 15-100(-

130) mm long at flowering, 4-10(-25)-flowered; lower-

most pedicels 10-20 mm long in flower, elongating to 30

mm; bracts whitish, ovate, acute or acuminate, glabrous

or lowermost ciliate, shorter or longer than pedicels,

lowermost 8-16 mm long. Flowers white; outer tepals

FIGURE 19. — Known distribution of Ornithogalum rupestre: white- or

pale yellow-flowered populations, O; deep yellow- or orange-

flowered populations, •.

ovate-lanceolate, inner eliptical-ovate, 6-10 x 3-5 mm.

Stamens ± half as long as tepals; filaments lanceolate, 4-

5 mm long, white, sometimes marked with yellow, inner

with quadrate or apically auriculate expansion in lower

2 mm. Ovary ovoid, ± 4 mm long, sometimes greenish

black apically; style 1. 5-3.0 mm long. Capsule ovoid,

6-8 mm long. Seeds angular, ± 1 mm long, colliculate

to echinulate. Flowering time: August and September.

Figures IL, 21.

Distribution and ecology, largely restricted to a small

area of high ground in southern Namibia, between 600-

950 m, in the Klinghardt Mountains and nearby Huib

Hoch Plateau (Figure 22), with a southward extension

into the Richtersveld in Northern Cape, South Africa

around Lekkersing. This is an extremely arid region that

is summer-dry, although coastal fogs supply some pre-

cipitation in the autumn.

The leaves of Ornithogalum puberulum are remark-

ably soft-textured for a species from such an arid area

and plants are restricted to rock outcrops and crevices,

on south-facing exposures or at the foot of boulders,

where they are sheltered from the afternoon sun in win-

ter and benefit from the cooler, m,oist conditions during

the growing season. The species has been recorded as

locally common where it occurs but leafing and flower-

ing is dependent on adequate rain.

Discussion: this highly distinctive species is immedi-

ately recognizable by the two (rarely three) soft-textured,

ovate to elliptical (rarely lanceolate) leaves that are either

glabrous or pubescent but always with soft cilia 0. 7-2.0

mm long on the margins. The leaves of Ornithogalum

puberulum are typically rather tubular and inflated at

the base where they clasp the stem, and the upper leaf is

mostly only half the size of the lower one or two.

The species shows considerable variation in the

pubescence of the leaves, sometimes within populations

Bothalia 37,2 (2007)

159

5 11 3

COMriO> Hlr.KHA,Rn.;\l |VBC,I

NATlOAl KfrAMCM IvyriTlTK 0Al'»;T(JVk-V

yizv

FIGURE 20. — Ornithogalum leeii-

poortense U.Mull.-Doblies &

D.Mull.-Doblies.

but more usually between them. The type collection

comprises four plants, all of which have leaves that are

entirely pubescent on both surfaces, apart from the upper

leaf on one of the plants, which is pubescent only api-

cally. The leaf and bract margins in all four plants are

densely ciliate. The type of Ornithogalum merxmueUeri

has similarly pubescent leaves but the bracts are entirely

glabrous. All other collections that we have examined

have leaves that are glabrous with sparsely pubescent

margins. Although both the type and Bruyns 2668 have

ciliate margins to the bracts, other populations from

near Rosh Pinah {Bruyns 8359, 8866) have entirely gla-

brous bracts.

Described from near Rosh Pinah in southern Nami-

bia, Ornithogalum piiberulum subsp. chris-bayeri Miill.-

Doblies & Miill.-Doblies was distinguished from the

typical subspecies by the shorter cilia on the leaf mar-

gins, hairy bracts, yellow-spotted filaments and shorter

style (1.6 mm vs 2. 5-3.0 mm). More recent collections

of O. puberulum, with somewhat variably ciliate leaf

margins, glabrous or ciliate bracts and short styles, that

160

Bothalia 37,2 (2007)

FIGURE 21. — Known distribution of Ornithogalwn leeupoortense, O;

O. pubenilum, •.

have subsequently been made near Rosh Pinah (Briiyns

2778, 8359, 8866) indicate that the species is more vari-

able than was previously supposed and that the recogni-

tion of this subspecies is unwarranted.

Typical Ornithogalum puberuhim from southern

Namibia is characterized by a rather corymbose inflores-

cence with up to 15 flowers. Populations of vegetatively

similar plants from near Lekkersing in the Richtersveld

{Oliver, Tolken & Venter 776\ Thompson & Le Roux 85),

however, have elongate, cylindrical inflorescences with

up to 25 flowers, giving them a very different appear-

ance. These plants were associated with O. puberulum

without comment by Miiller-Doblies & Muller-Doblies

(1996). Although a similar range in inflorescence fonn

is shown by other species in the group, including O. pru-

inosum, the status of these populations requires further

investigation.

The characteristic leaves of Ornithogalum puberulum,

clasping at the base and with long-ciliate margins, have

led to confusion with O. hispidum in the herbarium.

The latter species, a member of section Hispidaspasia,

is readily distinguished by its small bracts, and narrow

raceme of smaller flowers with filiform filaments.

History. Ornithogalum puberulum was described by

Obermeyer (1978) from plants collected fifteen years

earlier in southern Namibia. Ornithogalum merxmuel-

leri, described the following year by Roessler (1979)

from fruiting material collected in the same area of

southern Namibia, does not appear to differ in any sig-

nificant way and was synonymized under O. puberulum

by Muller-Doblies & Muller-Doblies (1996).

At the same time, Muller-Doblies & Muller-Doblies

(1996) described O. puberulum subsp. chris-bayeri from

plants that they had collected near Rosh Pinah in south-

ern Namibia, distinguishing it by the shorter cilia on the

leaf margins, hairy bracts, and yellow-spotted filaments

and short style. These differences are, however, not sig-

nificant in the light of more recent collections. We owe

much of our knowledge of the species to the recent col-

lections made by the Cape Town succulent specialist,

P.V. Bruyns.

Excluded species

1. Ornithogalum baurii Baker in Flora capensis 6:

504 (1897). Type: South Africa, Eastern Cape [Transkei],

Baziya Mountain, 4000' [1 220 m], November without

year, Baur 552 (K, holo.!; SAM, iso.!).

O. hygrophihim Hilliard & B.L.Burtt: 195 (1988). Type: Eastern

Cape [Transkei], NW of Umtata, hill above Mhlahlane Forest Station,

10 December 1985, Hilliard & Burtt 19768 (E, holo.; Kl, NU, PRE,

iso.).

2. Ornithogalum diphyllum Baker in Kew Bulletin

1895: 153 (1895). Type: South Africa, [KwaZulu-Natal],

Ntabamhlope Mountain, Evans 374 (K, holo.!; PRE-

photo.!).

3. Ornithogalum sephtonii Hilliard & B.L.Burtt in

Notes from the Royal Botanic Garden Edinburgh 41 : 308

(1983). Type: South Africa, Eastern Cape, Barkly East

Dist., Ben Mcdhui, ± 8100' [± 2 466 m], 3 December

1981, Hilliard & Burtt 14665 (E, holo.; NU, iso.).

These three taxa comprise a group of poorly under-

stood species known from isolated localities along the

mountains of the Eastern Cape and KwaZulu-Natal.

They are all small or dwarf plants with two or three

leaves and subcorymbose racemes of pure white flowers

subtended by foliar bracts. The seeds of Ornithogalum

diphyllum are club-shaped and echinulate (Obermeyer

1978) and those of O. sephtonii tetragonal and reticulate

(Hilliard & Burtt 1983). In the light of this, Obermeyer

(1978) and Muller-Doblies & Muller-Doblies (1996) had

little hesitation in associating this group of species with

section Aspasia from the Western and Eastern Cape,

despite the obvious anomaly in their distribution. They

were interpreted by Obermeyer (1978) and others as iso-

lated montane relics of the Cape section Aspasia.

At the same time Obermeyer (1978) included O. bau-

rii and O. synanthifolium, both of which had been col-

lected by Baur at Baziya in Eastern Cape, in a broadly

circumscribed O. conicum. There is little doubt, how-

ever, that O. baurii and O. synanthifolium are distinct

species (see Discussion under the latter), an opinion that

was first aired by Hilliard & Burtt (1988, 1989). What

is not as evident, however, is where the affinities of O.

baurii, O. diphyllum and O. sephtonii lie. While there is

certainly a superficial similarity in vegetative and floral

morphology between these three species and those of

section Aspasia, particularly in their foliar bracts and

moderate-sized, white flowers, and while the seeds of

O. diphyllum at least have some resemblance to those of

other species in the section, the capsules of this group

are not at all like those that characterize section Aspasia.

All three species have distinctly turbinate, 3-angled

capsules that are exposed by the reflexing of the tepals.

Such capsules are anomalous in section Aspasia, which

is characterized by ellipsoid or ovoid, obscurely angled

capsules that are concealed by the persistent, papery

perianth. On this basis alone the three species of the O.

baurii group should be excluded from section Aspasia.

Bothalia 37,2 (2007)

161

FIGURE 22. — Ornithogalum pubem-

Iwn Oberm.

Support for this interpretation is provided by DNA

sequence data (Manning et al. in prep.), which places

O. sephtonii among the residue of species previously

grouped within subgenus Urophyllon, where such turbi-

nate capsules are common, and not among the species of

subgenus Aspasia. We accordingly exclude these three

species from section Aspasia. The differences between

them are relatively minor and further study is likely to

reduce the number of taxa.

ACKNOWLEDGEMENTS

Our thanks to Elizabeth Parker and Dave Gwynne-

Evans for assistance in the field and to Michelle Smith

for scanning the specimens used in the illustrations. We

are also grateful to the curators of the various herbaria

for access to their collections. Clare Archer is thanked

for her assistance with specific queries. Additional mate-

rial was collected under permits from the Northern and

the Western Cape Nature Conservation authorities.

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Jacq. Journal of South African Botany 32: 325-333.

Bothalia 37,2 (2007)

SALISBURY, R.A. 1796. Prodromus stirpium in horto ad Chapel

AUerton vigentium. London.

SMITH, C.A. 1966, Common names of South African plants. Memoirs

of the Botanical Survey of South Africa No. 35. Botanical

Research Institute, Pretoria.

SPETA, F. 1998. Hyacinthaceae. In K. Kubitzki, The families and gen-

era of vascular plants. Springer, Berlin.

STEARN, W.T. 1983. The Linnaean species of Ornithogahm (Lilia-

ceae). Annals of the Musei Goulandris 6: 139-170.

THUNBERG, C.P. 1794. Prodromus plantarum capensium: 62. Edman,

Uppsala.

VAN WYK, B-E., VAN HEERDEN, F. & VAN OUDSHOORN, B.

2002. Poisonous plants of South Africa. Briza Publications,

Pretoria.

OTHER SPECIMENS EXAMINED

Acocks 14767 (8) K. Andreae 177, 764 NBG, Axelson 105, 359 (5)

NBG.

Barker 9738 (1) NBG; 1916 (2) NBG; 295, 4766, 9478, 9796, 10330

(4) NBG; 1919 (5) BOL, NBG; 1917,1923, 1935, 2570, 2606, 2646,

3332, 4198, 4858, 10534, 10801 (5) NBG; BOL22845 (6) BOL; 87

(7) BOL, K; BOL27865 (7) BOL; 292, 1576, 1922, 2406, 2581, 4259,

4266, 4898, 5347, 6035, 6814, 6859, 6879, 6994, 7292, 7579, 7605,

9106, 9135, 9585 (7) NBG; 1928, 5705, 7323, 7324, 7415, 9486, 9503

(8) NBG; 294, 1921, 1925, 2656, 2662, 3069, 4758, 6574, 7446, 9194,

10752 (9) NBG; 1079, 1932, 1933, 6761, 9024, 10310, 10763 (10)

NBG. Batten 2.100.82 (2) NBG; 91 (7) NBG. Baur 508 (2) K, SAM,

Bayer 3629 (7) NBG. Bayliss 6164 (7) NBG; 736 (9) NBG; 275, 276

(10) NBG. Bohnen 4733, 7052, 7536, 7537, 7966, 8732 ( 7) NBG.

Bolus 4350 (5) BOL; NBG73561 (5) BOL, NBG; 8696, BOL410/33

(7) BOL; 1256/32 (7) NBG; 20349 (7) BOL, K; 6597 (8) BOL; 4346

(9) K; BOL12892, BOL13065 (10) BOL. Bond 183 (3) BOL, NBG;

BOL22801 (3) BOL; 522 (5) NBG; 733 (7) NBG; 548 (9) NBG. Botha

(3)109 (7) NBG. Boucher 2617 (5) K, NBG; 4117, 6361, 6476, 6477

(5) NBG; 2699, 3024 (7) NBG; 3136 (8) NBG; 2838, 6562 (9) NBG;

3338, 4717, 56^7 (10) NBG. Browne NBGl 23/37 (7) NBG. Brownlee

BOL 22175, BOL22714 (7) BOL. Bruvns 1848 (7) NBG; 9218 (10)

NBG; 2771, 2778, 8359, 8866 (12) NBG. Biihr STE119639 (9) NBG.

Burchell 6198 (7) K. Burgers 1279, 1390 (7) NBG. Buys 481 (7) NBG.

Cassidy 127 (1) BOL, NBG. Compton 12546, 16589 (1) NBG;

17769 (2) NBG; 3171/34, 9514, 11781, 12335, 15259, 15260, 15261,

15263, 16374, 18815 (5) NBG; 12082 (6) BOL, NBG; 3587, 9623,

9738, 9967, 11680, 12151, 12457, 12772, 139/45, 15238, 16388,

16491, 16742, 16885, 17861, 18718, 18982, 21151, 21681, 21732,

22232, 23179, NBG24/34, NBG261/44 (7) NBG; 5251, BOL55470,

NBGl 39/45 (7) BOL; 12058 (7) BOL, NBG; 22072 (8) NBG; 3565,

9571, 9692, 11700, 14846, 16138, 20839, 20876, 24200, 24302 (9)

NBG; BOL1528 (10) BOL; 3585, 14848 (10) BOL, NBG; 9288, 9611,

11773, 11891, 16301 (10) NBG, Compton et al. 2011/36 (4) NBG.

Cook BOLl 766/25 (7) BOL. Cowling 1906, 1907 A (1) NBG. Craven

179(5) NBG. Cross 71, 72 (5) NBG. Cruz 98 (9) NBG.

Davison 32 (7) SAM. De Villiers 949/62 (1) NBG. De Vos 154 (5)

NBG, Desmet 361 (8) NBG; 1941 (10) NBG; 3063 (11) NBG. Dobay

81 (7) NBG. Drege 1512, 2662(10) K. Du Plessis 58, 59 (7) NBG.

Ebersohn 370 (5) NBG, Ecklon 569 (1) K. Edwards 93/29 (7) NBG.

Emdon 174, 253 (7) NBG. Esterhuysen 11497 ( 1) BOL; 1433 (5) BOL;

71402 (5) NBG; 7777, 4390, 7278, 9317, 9392, 11207, 11280, 18327,

BOL7/42 (7) BOL; 2941, NBG73514, NBG73568 ( 7) NBG; 3781,

14957 (7) BOL, NBG; 4384, 22100 (10) BOL.

Fellingham 474 (5) NBG; 1089/5 (1) NBG; 176 (9) NBG. Flanagan

788 (1) BOL; 2238 (7) BOL, K. Forrester 102, 931 (7) NBG.

Fourcade 512 4175 (7) BOL, K, NBG; 5159A, 5866 (1) BOL.

Galpin 312 (2) K; 311 (7) K. Germishuizen 4063, 4156 (7) NBG.

Gillett 1914, 4289 (1) BOL; 3384 (1) NBG; 380 (9) NBG. Glass

380 (2) NBG; 379, 381 (7) NBG. Goatcher BOL13726 (6) BOL,

K. Goldblatt, Manning & Savolainen 11526 (8) NBG. Goulimis

BOL22 775 (1) BOL; BOL22809 (1) BOL. Grobler 491 (7) NBG.

Guthrie 2390, 2768 (7) NBG,

Hall 4542 (1) NBG; NBG88092 (3) NBG; 3110 (4) K, NBG; 272,

1504, 4511, NBG73519 (1) NBG; 325, 833, 4929 (8) NBG; 958,

NBG68697 (9) NBG; 475/52, 4930, 5195, 5217, NBG73664 (10)

NBG; 5062 (10) K, NBG. Hanekom 2616 (5) K, NBG; 3108 (5)

NBG. narrower 3001 (7) NBG. Heathie BOL3710 (6) BOL. Heese

Bothalia 37,2 (2007)

163

STEI0173 (7) NBG. Heginbotham 164 (7) NBG; 43 (9) NBG. Herre

BOL3966 (4) BOL; STEl 235-4. STE17465 (8) NBG; STEI7458,

STE19638 (10) NBG). Hiemstra 470 (8) NBG. Hihon-Taylor 1990,

2036 ( 7) NBG; 1991 (9) NBG. Horrocks 12, 29 (9) NBG. Hugo 2061

(7) K, NBG; 2915 (8) NBG. Hurling NBG73531 (7) NBG. Hutchinson

1116(1) BOL, K; 767(10)BOL, K.

IBSA NBG207216 (3) NBG. Isaac 7234 (10) BOL.

James 134044, 233844. BOL 3277/32 (7) BOL. Jones STE30397 (7)

NBG. Jordan STE25508 (5) NBG. Joubert (e) (7) NBG; (c.) (9) NBG.

Kellerman 11 (9) NBG. Kemper IPC740 (7) NBG. Kensit BOL22843

(7) BOL. Ketfoot 5907 (9) NBG. Kolbe BOL14312 (8) BOL. Kruger

44 (5) NBG. Kurzweil 1271 (5) NBG.

Lamb BOL1644/30 (4) BOL; BOL1619/30 (7) BOL. Laughton

NBG22685 (7) NBG. La\4s BOL22152 (5) BOL; BOL22831 (7) BOL.

Le Roilx 2837 (5) NBG; 2505 (8) NBG; 2781A (8) BOL. Le Roux &

Ramsey 326 (10) NBG. Lechmere-Oertel 355 (9) NBG. Leighton

175 (\) BOL; 1495/33 (4) NBG; BOL1492/33 (4) BOL; 135, 137 (5)

K, NBG; 138, 163, 164, 170, 189, 195, 196 (5) NBG; 188 (5) BOL,

NBG; BOL22700, BOL22701 (5) BOL; 134, 136, 1503/33 (7) NBG;

694, 717. 2200 (7) BOL; 275/45 ( 8) NBG; 3155 (8) BOL; 161, 655,

3159 (10) BOL; 1051 (10) BOL, NBG. Leipoldt 4374 (4) BOL; 4028

(8) BOL, NBG; BOL1048/36 (8) BOL; NBG73654 ( 8) NBG; 782 (9)

NBG; 3357, 4090, 4379 (10) BOL). Lexyns BOL1292 (4) BOL; 3470,

4795, BOL5763a, BOL7233 (7) BOL; 1776, BOL4631, BOL7232 (9)

BOL; 11652, BOL7235 (10) BOL. Lewis 2185 (1) SAM; BOL22799

(4) BOL; 5070 (5) NBG; 5701 (7) NBG; BOL22088, BOL22853,

NBG2696/32 (7) BOL; 279 (8) NBG; 1425, SAM55743, SAM62315,

SAM62316 (8) SAM; 5899 (9) NBG; BOL22856, BOL22858 (10)

BOL). Lombard s.n. (8) NBG. Louw 2689 (5) NBG. Low 8630 (9)

NBG.

MacKinnon s.n. (7) NBG. MacOwan 2652 (1) K, SAM; 20 (2) BOL;

940 (5) BOL, K; 505, 1819 (7) BOL, K. Maguire 2004 (8) NBG; 238

(9) NBG. Malherbe STE30398 (7) NBG. Manning 2655 (4) NBG;

2654 (5) NBG; 2650 (7) NBG; 1036 (9) NBG. Manning & Martinez-

Azorin 3 (4) NBG; 13. 63. 64. 83 ( 5) NBG; /, 12 (6) NBG; 4, 5, 6.

8, 10, 60, 61, 62, 82 (7) NBG. Manning & Snijman 2722 (7) NBG.

Marais 1 (\) NBG; BOL71519 (2) BOL. Marloth 11486 (5) NBG;

9572, 10781 (7) NBG; 12350, 12855 (8) NBG; 11497 (9) NBG;

11496 (\Q) NBG. Marsh 1043 (5) NBG; 462 (8) NBG; 479 ( 9) NBG;

BOL1667/30 (9) BOL. Martin 836 (8) NBG; 69 (9) NBG. Martinez-

Azorin 95 (4) NBG. Martley BOL22776, BOL22777 ( 1 ) BOL. Mathews

BOL22704 (7) BOL; BOL27883 ( 9) BOL. Mauve & Hugo 19. 51 (5)

NBG. Mauve. Reid & Wikner 91 (7) NBG. McDonald 1641 (7) NBG.

Meyer STE9067 (9) NBG. Middlemost NBG73521 (7) NBG. Mitchell

7/54 (10) NBG. Moffett 37 5. 451 (1) NBG. Montgomery 232 (5) NBG;

75, (7) NBG. Morley 230 (5) NBG; 325 (7) NBG. Morris BOL5805 (8)

BOL, K. Mucina 080905/14 (11) NBG. Muir 1164, 4989 (7) NBG.

NBG Expedition 175/65 (4) NBG. Nieuwoudt s.n. (9) BOL, K. Norden-

stam 2^53 (10) NBG.

O'Callagham 1303 (5) NBG. Oliver 3691. 5133 (5) NBG; 5179 (7)

K. NBG; 4801, 5028. 5160. 5188. 5298. 5475. 5501. 9759 (7) NBG;

3868. 4024. 9567. 9711 (9) NBG; 10191 (12) NBG. Oliver. Tolken

& Venter NBG169104 (8) NBG; 776 (12) NBG. Olivier 834, 874 (7)

NBG. Orchard 323 (7) K, NBG.

Palmer 244 (7) NBG. Pappe SAM23321 (1) SAM. Paterson-Jones

663. 704 (5) NBG; 683. 719 (7) NBG. Peers BOL27889 (1) BOL;

BOL27929 (5) BOL. Pegler 219 (2) BOL. K, SAM. Pells BOL22706

(7) BOL. Perry 3366 (3) NBG. Perry & Snijman 2423 (4) NBG.

Phillips 1080 (7) BOL; 2084 (7) SAM; >624 (.7) NBG; 7543 (9) NBG.

Pienaar 8 (2) NBG. Pillans 8370 (1) BOL; 8959, 9851 (5) BOL;

10890 (5) NBG; 8115 (7) BOL; 9566 (7) BOL, K, NBG; 4935, 5448

(8) BOL. Plowes NBG120374 (9) NBG. Pocock 467, 647 (7) NBG;

162 (9) NBG. Pond 117 (9) NBG. Pretorius 626 (4) NBG; 547 (7)

NBG; 92 (9) NBG; 374 (10) NBG.

Rode & Boucher 0180 (5) NBG. Rogers 27540 (7) BOL, K; 28224 (6)

BOL, K, SAM; BOL12941 (7) BOL; STE12770 (1) NBG. Rosch 273

(3) NBG; 664 (4) NBG. Ross-Prames 1241/26 (5) NBG; BOL1259/26

(8) BOL. Rouoc STE18987 (5) NBG. Rourke 617. 650 (7) NBG.

Rycroft 2372, 2760. 3020 (7) NBG. Ryder 2272/30 (4) NBG; 3169/34

(7) NBG.

Salter BOL869/32 (1) BOL; 2755, 2805 (4) BOL, K; 590/32, 1213/33

(5) NBG; 4978 (5) BOL; BOL7816 (5) BOL, NBG; 2718 (7) BOL;

2779 (7) BOL, K; 2746, (10) BOL, K; 6226 (10) BOL. Schlechter

9131 (7) BOL; STE9011. STE10883 (8) NBG; 1421 (9) K; STE10882

(9) NBG; 11368, 11487 (10) BOL, K. Schonken 307 (7) NBG. Scott

BOL2212/31 (7) BOL. Smith 6447 (4) NBG; 737, 2658 (5) NBG; 2659,

2869, 2870 ( 7) NBG; 6446 (8) NBG. Smuts s.n. (7) NBG. Snijman

376, 1370 (7) NBG; 7725 (9) NBG; 269, 7922 (10) NBG. Solomon 106

(5) NBG. Starket BOL17831 (9) BOL. Stayner NBG7354 7 (7) NBG.

Steiner 2914. 3024 (7) NBG; 2917 (7) K, NBG. Stephens & Glover

8632 (9) K. NBG. Steyn 488 (9) NBG. Stobie 24. 3S (10) NBG. Stokoe

7605 (7) BOL; SAM63139. SAM64702 (7) SAM; SAM60648 (8) SAM.

Stoty 4424 (4) K. Strauss 56 (8) NBG.

Taylor 3594. 4214 (2) NBG; 1202 (5) BOL; 3149/35. 4010 (5) NBG;

4325. 4348. 5171. 11137. 11887 (7) NBG; 3957, 5859. 5909. 10616.

11088 (9) NBG. Theron STE10181 (5) NBG. Thode STE6082 (5)

NBG; 5460. A709, All 35, A2096, STE9494 (7) NBG. Thomas 27864

(7) BOL. Thompson 807 (9) NBG; 2894 (10) NBG. Thompson & Le

Roux 224 (8) NBG; 105 (11) NBG; 85 (12) NBG. Thomson 2143,

3200, 3317. (7) NBG; 1288 (8) NBG; 303 (NBG) (9), 65, 2663 (10)

NBG. Thorne SAM48847 (8) SAM. Thorns NBG73539 ( 7) NBG.

Van Berkel 445 (8) NBG; 259, 446 (9) NBG; 452 (10) NBG. Van der

Merwe 278 (7) NBG; 246 (9) NBG. Van der Riet s.n (5) NBG. Van

der Westhuizen s.n. (10) NBG. Van Jaarsveld 8444 (8) NBG; 8189

(11) NBG. Van Niekerk s.n. (10) BOL. Van Rooyen, Steyn & de Villiers

497 (9) NBG. Van Wyk 344, 2059 (7) NBG. Van Zyl 2776/27 (5) NBG;

3101. 3150. 3191 (7) NBG; 3223, 3302. 3549 (9) NBG. Visser 44 (7)

NBG. Viviers 3. 1176 (7) NBG; 1568 (9) BOL. Vtviers & Vlok 45, 64

(7) NBG.

Wagener 208 (9) NBG. Walgate 328 (7) NBG; BOL22850 (7) BOL.

Walters 2308 (5) NBG; 498, 1219, 1229. 1438, 1781, 2730 (7) NBG;

45 (10) NBG). Wasserfall 779 (1) NBG; 622, 792 (7). Wells 2804

(2) K. Werdermann & Oberdieck 1068 (2) K. Willems 81 (7) NBG.

Williams 2913. 3343 (7) NBG. Williamson 3164 (8) BOL; 3795, 5451,

5453 (10) NBG. Williamson & Williamson 5825 (8) NBG. Wilman

NBG405/37 (7) NBG. Winkler 152 (5) NBG. Wolfardt STE10788 (10)

NBG. Woodvine 31 (7) NBG. Wurts 410. 417, 1204. 1584, 2266 (7)

NBG.

Zeyher 5047 ( 1 ) BOL, K, NBG, SAM; 1065 (7) NBG. Zinn SAM62230

(7) SAM.

INDEX

Aspasia, section, 135

Aspasia, series 1, 135

Aspasia, subgenus, 135

Omithogalum L., 134

Section Aspasia, 135

Series 1 Aspasia, 135

Series 2 Thyrsoides, 143

Series 3 Maculata, 147

Subgenus Aspasia (iSa/wi.) Oberm., 135

aestivum L. Bolus, 135

alticolum F.M.Leight., 147

aurantiacum Baker, 155

aureum Curtis, 147

hdiux'n Baker, 160

6/co/or Haw., 144

6row«/ee/ F.M.Leight., 147

CQxes,\wmm F.M.Leight., 146

citrinum Schltr. ex Poelln., 147

coarctatum Jacq., 143

conicum Jac(?., 135

subsp. strictum (L. Bolus) Oberm., 143

var. strictum (L. Bolus) F.M.Leight., 143

coxX\cat\xxx\ Mart.-Azorin. 140

diphyllum Baker, 1 60

dubium 77omH., 147

fergusoniae L. Bolus. 147

fimbrimarginatum F.M.Leight., 147

flavescens ]&cc\., 147

flavissimum Jacq., 147

gilgianum Schltr. ex Poelln., 147

glaucophyllum Schltr. ex Poelln., 152

/;ermo«ff/7 F.M.Leight., 144

hygrophilum Hilliard & B.L.Burtt, 160

/w5/gnc F.M.Leight., 153

164

Bothalia 'il ,1 (2007)

Omithogalum Z,. (cont.)

lacteum Jacq., 135

var. coniciim (Jacq.) Baker, 135

leipoldtii L.Bolus, 147

leeupoortense U.Mull.-Doblies & D-Miill.-Doblies, 156

Maculata, series 3, 147

maculatum 152

var. speciosum (Baker) F.M.Leight., 153

var. splendens (L.Bolus) F.M.Leight., 153

macidatiim Thunh., 152

magnificum Poelln., 1 53

we/'.rw;/e//en Roessler, 158

miniatum Jacq., 147

var. vandermerwei (Barnes) F.M.Leight., 147

midtifolium Baker, 155

perpidchntm Schltr. ex Poelln., 147

F.M.Leight., 147

prmnosum F.M.Leight., 152

puberulum Oberm., 158

subsp. chris-bayeri U.Mull.-Doblies & D.Miill.-Doblies, 158

ramincidoides L.Bolus, 155

roMOt/wn U.Mull.-Doblies & D.Miill.-Doblies, 153

rupestre L./, 155

saxatile Schltr., 155

Section Aspasia, 135

sephtonii Hilliard & B.L.Burtt, 160

Series 1 Aspasia, 135

Series 2 Thyrsoides, 143

Series 3 Maculata, 147

speciosum Baker, 153

splendens L.Bolus, 153

strictum L.Bolus, 143

subcoriaceum L.Bolus, 147

Subgenus Aspasia (Salisb.) Oberm., 135

synanthifolium FlMLe/g/??., 138

thimbergianum Baker, 152

var. concolor Baker, 1 53

Thyrsoides, series 2, 143

thyrsoides Jac^., 143

vandermerwei Buvaos, 147

var. album Barnes, 147

virgineum Soland ex Baker, 155

witteklipenseL.yi.Lol^l., 155

Bothalia 37,2: 165-175 (2007)

Name changes in the Old World Rhus and recognition of Searsia (Ana-

cardiaceae)

R.O. MOFFETT*

Keywords: Anacardiaceae, nomenclature, Rhus L., Searsia F.A, Barkley, taxonomy

ABSTRACT

The background to and status of the genus Searsia F.A. Barkley (Anacardiaceae) is discussed and reasons given as to why

it is the correct name for those Old World species in the Rhus complex formely regarded as subgenus Thezera (DC.) K.Koch

(section Gerontogeae Engl.). An annotated list of all the accepted 111 species and 28 further infraspecific taxa in Searsia is

presented, and where necessary, new combinations are made and types are designated.

INTRODUCTION

WEen the author revised the southern African spe-

cies of Rhus L., Anacardiaceae, for the Flora of southern

Africa (Moffett 1993), he stated that he was retaining the

species in Rhus pending the results of ongoing research

on the generic status of this heterogeneous genus.

Although he was aware that Barkley had previously pub-

lished the name Searsia for the Old World Rhus species,

he preferred to follow Brizicky (1963) in treating this

group as subgenus Thezera (DC.) K.Koch.

Recent phylogenetic analyses using DNA and gene

spacers of the Rhus complex have shown that Searsia is

clearly monophyletic and widely separated from Rhus s.

str (Miller et al. 2001; Yi et al. 2004) and there is there-

fore no further reason not to uphold Searsia F.A. Barkley

as the correct name for the ‘Old World’ Rhus species.

BRIEF HISTORY OF THE NOMENCLATURE OF THE ‘OLD

WORLD’ RHUS SPECIES

The Old World Rhus species are found in the Mediter-

ranean, Africa and Asia. Linnaeus included three of these

from the Cape in his list of 12 species in Species planta-

rum (Linnaeus 1753). Of the other nine, four have subse-

quently been retained as Rhus L., three as Toxicodendron

Mill., one as Cotinus Mill, and one as Allophyhts L.,

(Sapindaceae) [Index Nominum Genericorum (planta-

rum) (ING) 2006], epitomizing the heterogeneous nature

of Rhus sensu lato. The type species of Rhus and there-

fore that which governs the application of Rhus sensu

strict o is Rhus coriaria L., the spmach of southern

Europe, Mediterranean region and the near East. With its

pinnately compound leaves, drupes with prominent red

glandular hairs and resinous mesocarp, it is outwardly

markedly different to the species of subgenus Thezera,

which are characterized by temate, rarely simple or pal-

mate leaves and drupes pale, glabrous or tomentose with

a resinous mesocarp adhering to the bony endocarp.

* Department of Plant Sciences, Qwaqwa Campus, University of the

Free State, Phuthaditjhaba, South Africa.

E-mail: moffettro(§;qwa.uovs.ac.za

MS. received: 2006-12-21.

The heterogeneity of Rhus was recognized early,

and Bemhardi (1838) remarked at the end of a paper on

Laurophyllus Thunb., that the temate species of Rhus

from the Cape were different to the tme Rhus species and

seemed to form a distinct genus which one could name

Terminthia Bemh. Despite this name being used by Wu &

Ming (1979) in Flora Yunnan, it is illegitimate, as accord-

ing to Art. 34.1 (b) of the St Louis Code (Greuter et al.

2000), it is a provisional name. McNeill & Greuter (pers.

comm. October 2006) are of the opinion that Bemhardi ’s

use of the words ‘welche man Terminthia nennen kormte’

is a clear indication of provisional status and nonaccep-

tance of the name.

The first person to lump all the Old World Rhus spe-

cies together was Engler (1881), who placed them in his

new section Gerontogeae (the old rhusses), and who soon

after provided the first detailed account of the genus in

A. & C. De Candolle’s Monographiae phanerogamarum

(Engler 1883). His encyclopaedic treatment of Rhus with

its four sections, Trichocarpae, Venenatae, Gerontogeae

and Melanocarpae, formed the foundation by which Rhus

was measured for the next 60 years until Barkley (1942,

1943, 1950, 1965) criticized the traditional concept of the

genus. Engler ’s use of section Gerontogeae was actually

illegitimate as it was predated by section Thezera of De

Candolle (1825).

The name Searsia first appeared in a footnote to a key

to the genera of the Anacardiaceae where Barkley (1942)

stated 'Searsia n.gen. = (Rhus) Section Gerontogeae.

Named after Paul B. Sears’. Sears (1891-1990), who was

to become a renowned stratigraphic palaeontologist, eco-

logist and head of the Yale School of Botany, was one

of a number of botanists thanked by Barkley for encour-

agement and assistance during the early part of his Ph.D.

studies on American species (Barkley 1937).

The above publication, however, was illegitimate as

there was no description of the genus and this was cor-

rected the following year when Searsia appeared as genus

six in Flora of Texas, with a full description and includ-

ing the cultivated Searsia lancea (L.f) F.A. Barkley, a

South African species (Barkley 1943). In that publication,

Barkley also made the new combination of S. tomentosa

(L.) F.A. Barkley and designated this species as the type

species.

166

Bothalia 37,2 (2007)

Eight flirther combinations in Searsia were made by

Barkley in 1950, and eleven more in a 1965 Iraq pub-

lication. In this latter paper, titled ‘A criticism of the

traditional concept of the genus Rhus’, Barkley rec-

ognized ten genera in the Rhus complex and in place

of Searsia tomentosa, designated Searsia pentaphylla

(Jacq.) F.A. Barkley as the type of the genus. In separa-

ting the genera of the Rhus complex, he chose not to fol-

low Brizicky, who two years previously, in a paper on

the generic limits of Rhus, preferred to retain the name

Rhus at genus level and recognized six subgenera, one

of which was subgen. Thezera (DC.) K.Koch and which

included Engler’s Gerontogeae (Brizicky 1963).

Despite Barkley’s criticism, the name Rhus has been

maintained in all the various African regional floras pub-

lished subsequently, as well as in the few new species,

the latest being Rhus pygmaea Moffett (Moffett 1999)

and Rhus gallagheri Ghaz., (Ghazanfiir 2002).

CONFIRMATION OF THE STATUS OF SEARSIA

Recent research in the USA on the generic status of

Rhus has provided conclusive evidence that the ‘Old

World’ species of Rhus are sufficiently different to

warrant generic status of their own, thus vindicating

Barkley. Miller et al. (2001) examined the sequences

of the internal transcribed spacer region (ITS) of the

nuclear ribosomal DNA of six genera in the Rhus sensu

lato complex in order to determine the monophyly of

Rhus sensu stricto and to provide insight into the phy-

logenetic and biogeographical history of the genus. Two

species from the Old World, viz. Searsia ciliata and S.

quartiniana, both ex hort., were included. As outgroups

in their analysis, Pistacia vera and Schimis molle were

used. The results showed that Rhus sensu stricto is

monophyletic and that the other genera, Actinocheita

F.A. Barkley, Cotinus, Malosma (Nutt.) Raff, Searsia

and Toxicodendron were distinct from Rhus sensu stricto,

but the relationships between these other genera were

not well resolved. Interestingly Schinus L., Searsia and

Toxicodendron formed a clade (bootstrap value of 91%).

The authors suggested that the use of additional charac-

ters such as chloroplast genes, should help to resolve the

intergeneric relationships.

This suggestion was followed by Yi et al. (2004), who

carried out a phylogenetic analysis of the Rhus complex

using ITS of nuclear ribosomal DNA and chloroplast

{ndh¥ and trnL-¥). Among the species included, but not

used in the Miller et al. (2001) study, were the southern

African Searsia lancea, S. leptodictya, S. pyroides and

S. undulata, all ex hort. The phylogenetic analysis of the

ITS, the chloroplast and the combined ITS and chloro-

plast datasets confinned the monophyly of Rhus sensu

Barkley (1937) and that Searsia from Africa was mono-

phyletic and distinct from the Rhus clade. The ITS data

also showed that Searsia lancea, S. leptodictya and S.

undulata fonned a clade, which was sister to the clade

composed of S. ciliata, S. pyroides and S. quartiniana.

A chronogram indicating diversion times compiled by

these authors based on the maximum likelihood tree of

the combined ITS and cpDNA data showed that Searsia

undulata diverged 55 Ma, Rhus 49 Ma and R. coriaria

24 Ma.

SEARSIA F.A.BARKLEY: SPECIES AND TYPES

Barkley’s original designation of Searsia tomentosa

as the type species of the genus (Barkley 1943), and sub-

sequent replacement by S. pentaphylla (Barkley 1965)

requires some explanation, and I am indebted to Prof

John McNeill of Ontario and Edinburgh for clarifying the

issue.

Article 7.4 of the Code states ‘A new name formed

from a previously published legitimate name (stat. nov.,

comb, nov.) is, in all circumstances, typified by the type

of the basionym, even though it may have been applied

erroneously to a taxon now considered not to include the

type’. As Barkley, 1943, cited Rhus sect. Gerontogeae

Engl. (1881) as a synonym of his new genus, his des-

ignation of S. tomentosa as the type was superfluous,

as the type of Searsia is the type of sect. Gerontogeae.

Section Gerontogeae is, however, illegitimate as Engler

acknowledged in his publication that it was based on

the earlier sect. Thezera DC. (1825). As Engler did not

indicate a type for sect. Gerontogeae, according to Art.

7.5, Searsia is automatically typified by the type of

sect. Thezera. Neither De Candolle nor Koch (1853),

who treated Thezera as a subgenus, designated types

and it was left to Brizicky (1963) to choose a lectotype,

viz. Rhus pentaphylla (Jacq.) Desf, which explains

why Barkley accepted this in 1965. Despite this, ING

(2006), however, still cites R. tomentosa L. as the type

of Searsia.

The following list is based on research undertaken on

the taxonomy of all the Old World Rhus species when

revising Rhus in southern Africa between 1976 and 1992

(Moffett 1993), and on the relevant literature since then.

The few differences between this list and that of the

accepted names for sub-Saharan Africa (Klopper et al.

2006) are intentional. Based on the notes made when the

author visited many European herbaria in 1982 and cor-

respondence with respected taxonomists, types have been

designated for those taxa as yet untypified.

Searsia F.A. Barkley in Flora of Texas 3: 104 (1943).

Rhus subgen. Thezera (DC.) K.Koch., Hortus den-

drologicus: 197 (1853); Rhus sect. Thezera DC. (1825);

Rhus sect. Gerontogeae Engl. 379: (1881) nom. illeg.

Type: Searsia pentaphylla (Jacq.) F.A. Barkley: 57

(1965). Basionym Rhus pentaphylla (Jacq.) Desf lecto.,

designated by Brizicky: 63 (1963); Rhamnus pen-

taphylla Jacq.: 27 (1767); Rhamnus Siculus pentaphyl-

los Boccone: 43, t. 2 1 , ( 1 674).

Searsia acocksii (Moffett) Moffett, comb. nov.

Rhus acocksii Moffett in South African Journal of Botany 54:

172 (1988). Type: Eastern Cape, Lusikisiki District, southern edge of

Msikaba Gorge, Acocks 1 3250 (PRE, holo.!).

South Africa.

Searsia acuminatissima (R.Fern. & A. Fern.) Moffett,

comb. nov.

Rhus acwiiiiia/i.ssiimi R.Fern. & A. Fern, in Boletim Sociedade Brote-

riana, ser. 2, 38: 183 (1965b). Type: Malawi, Mulanje Mtn, Likabula,

Clements 111 (FHO, holo,).

Malawi, Mozambique,

Bothalia 37,2 (2007)

167

Searsia albida (Schousb.) Moffett, comb. nov.

Rhus albida Schousb.. lagttagelser Vextriget i Marokko: 142 ( 1 800).

Type: Marocco [Morocco], Schousboe s.n. (C!, lecto., here designated).

Algeria, Libya, Morocco.

Searsia albomarginata (Sond.) Moffett, comb. nov.

Rhus albomarginata Sond. in Flora capensis 1:519 (1860). Type:

Eastern Cape, Slaaykraal, Burke s.n. [K!, lecto., designated by Moffett:

88 (1993)].

South Africa.

Searsia anchietae (Figalho & Hiern ex Hiern) Mof-

fett, comb, nov., forma anchietae

Rhus anchietae Fipalho & Hiem ex Hiern, Catalogue of

Welwitsch’s African plants 1,1: 184 (1896). Type: Angola, Huilla,

Monhino, Welwitsch 4424 (LISU!, S, lecto., here designated).

Angola, Malawi, Tanzania, Uganda, Democratic Republic of the Congo,

Zambia.

Searsia anchietae (Figalho & Hiern ex Hiern) Mof-

fett forma mendon^ae (Meikle) Moffett, comb. nov.

Rhus anchietae Fifalho & Hiem ex Hiem forma mendongae

(Meikle) R.Fem. in Garcia de Orta (Lisboa) 14,3: 360 (1966). R. men-

dongae Meikle: 284 (1952). Type: Angola, Benguela, Rio Quito, prox.

de Quipeio, Exell & Mendonga 1877 (COI, holo.!).

Angola.

Searsia anchietae (Figalho & Hiern ex Hiern) Mof-

fett forma suffruticosa (Meikle) Moffett, comb. nov.

Rhus anchietae Figalho & Hiem ex Hiem forma suffruticosa

(Meikle) R.Fem. in Garcia de Orta 14,3: 362 (1966). R. suffruticosa

Meikle: 287 (1952). Type: Zambia, Mwinilunga District, near Dobeka

Bridge, Milne-Redhead 3609 (K, holo.!).

Angola, Democratic Republic of the Congo (DRC), Zambia.

Searsia angolensis (Engl.) Moffett, comb, nov., forma

angolensis

Rhus angolensis Engl, in Monographiae phanerogamamm 4: 448

(1883). Type: Angola. Huilla, Catumba, Lopolo, Welwitsch 4429 [G-

DC (Mon. Phan.), holo.!].

Angola.

Searsia angolensis (Engl.) Moffett forma glabre-

scens (R.Fem.) Moffett, comb. nov.

Rhus angolensis Engl, forma glabrescens R.Fem. in Garcia de Orta

(Lisboa) 14,3: 366 (1966). Type: Angola, Cuanza Sul, Cela, Barbosa &

R. Correia 8873 (LISC, holo.!).

Angola.

Searsia angustifolia (L.) F.A.Barkley in Monographs

of the Biological Society of Iraq 3: 54 (1965).

Rhus angustifolia L.: 267 (1753). Type: Aethiopia (Africa). Herb.

LINN 378.21 [LINN!, lecto., designated by Moffett: 100(1993)].

South Africa.

Searsia arenaria (Engl.) Moffett, comb. nov.

Rhus arenaria Engl, in Botanische Jahrbiicher 32: 132 (1903).

Type: Angola, Huilla, alongside Nene River, Dekindt 685 [P!, o, neo.,

designated by R.Fem.: 363 (1966)]. ,

Angola.

Searsia aucheri (Boiss.) Moffett, comb. nov.

Rhus aucheri Boiss., Diagnoses plantamm orientalium novamm

2: 5 (1843). Type: Mascate [Muscat & Oman], Aucher-Eloy 4324 [G

(Herb. Boiss.!), lecto., here designated)].

Arabia (Sultanate of Oman).

Searsia batophylla (Codd) Moffett, comb. nov.

Rhus batophylla Codd in Bothalia 6: 539 (1956). Type: Eastern

Transvaal [Mpumalanga], Steelpoort, Mooihoek Chrome Mine, Codd

& Dyer 7699 (PRE, holo.!).

South Africa.

Searsia blanda (Meikle) Moffett, comb, nov., forma

blanda

Rhus blanda Meikle in Boletim Sociedade Broteriana, ser. 2, 26:

286 (1952). Type: Angola, Benguela, Nova Lisboa and Texeira da

Silva, Exell & Mendonqa 1822 (COL holo.!).

Angola.

Searsia blanda (Meikle) Moffett forma exelliana

(Meikle) Moffett, comb. nov.

Rhus blanda Meikle forma exelliana (Meikle) R.Fem. in Garcia

de Orta 14,3: 371 (1966). R. exelliana Meikle: 102 (1954). Type:

Angola, Bie, Rio Cubango, Vila da Ponte. Gossweiler 3648 (BM,

holo.!).

Angola.

Searsia boiusii (Sond. ex Engl.) Moffett, comb. nov.

Rhus boiusii Sond. ex Engl, in Monographiae phanerogamamm 4:

436 (1883). Type: Eastern Cape, Graaff-Reinet, Cave Mountain, Bolus

737 (S, holo.!).

South Africa.

Searsia brenanii (Kokwaro) Moffett, comb. nov.

Rhus brenanii Kokwaro in Kew Bulletin 34: 754 (1980). Type:

Tanzania, Rungwe District, Siwago, Pangundutani, Brenan & Green-

way 8202 (EA, holo.).

Tanzania.

Searsia burchellii (Sond. ex Engl.) Moffett, comb. nov.

Rhus burchellii Sond. ex Engl, in Monographiae phanerogamamm

4: 412 (1883). Type: Northern Cape, confluence of Vaal and Orange

Rivers. Burchell 1722 (K, holo.!).

Lesotho, Namibia, South Africa.

Searsia carnosula (Schonland) Moffett, comb. nov.

Rhus carnosula Schonland in Bothalia 3: 41 (1930). Type: Eastern

Cape, Gekau, Drege 5569f3 [P!, lecto., designated by Moffett: 33

(1993)].

South Africa.

Searsia chirindensis (Baker f.) Moffett, comb. nov.

Rhus chirindensis Baker f. in Botanical Journal of the Linnean So-

ciety 40: 49 (1911). Type: Zimbabwe, near Chirinda, Swynnerton 168

(BM, holo.!).

Mozambique, South Africa, Swaziland, Zimbabwe.

Searsia ciliata (Licht. ex Schult.) A.J.Mill. in A.J.

Mill et al. in International Journal of Plant Sciences 162:

1403 (2001).

Rhus ciliata Licht. ex Schult. in L.: 661 (1820). Type: Northern

Cape. Grootte Rivier Poort, Liechtenstein in Herb. Willd. 6016 [B

(WILLD.), holo.!].

Botswana, Namibia, South Africa.

Searsia crenata (Thitnb.) Moffett, comb. nov.

Rhus crenata Thunb. in Phytographische Blatter: 28 (1803). Type:

Cape of Good Hope, Thunberg in Herb. Thunberg 7321 [UPS!, Rhus

crenatum a , lecto., designated by Moffett: 75 (1993)].

South Africa.

Searsia crenulata (A. Rich.) Moffett, comb. nov.

Rhus crenulata A. Rich., Tentamen Florae Abyssinica 4: 142 ( 1 847).

Type: Ethiopia, Tcheleukote, Petit ex Herb. Rich. (P!, lecto., R. crenu-

lata nob., here designated).

Ethiopia.

Searsia cuneifolia (L.f) F.A.Barkley in Monographs

of the Biological Society of Iraq 3: 54 (1965).

Rhus cuneifolia L.f: 183 (1781). Type: Cape of Good Hope,

Thunberg in Herb. Thunberg 7323 [UPS!, lecto., designated by Moffett:

92 (1993)].

South Africa.

168

Bothalia 37,2 (2007)

Searsia dentata (Thimb.) FA. Barkley in Monographs

of the Biological Society of Iraq 3: 54 (1965).

Rhus dentata Thunb. : 52 (1794). Type; Cape of Good Hope, Thun-

berg in Herb. Thunberg 7325 [UPS!, lecto., designated by R.Fem.: 124

(1967)].

Lesotho, Mozambique, South Africa, Swaziland, Zimbabwe.

Searsia discolor (E.Mey. ex Sond.) Moffett, comb.

nov.

Rhus discolor E.Mey. ex Sond. in Flora capensis 1: 507 (1860).

Type: Eastern Cape, Katberg, Drege 3449 [Si, lecto., designated by

Moffett: 52 (1993)].

Lesotho, South Africa. Swaziland.

Searsia dissecta (Thunb.) Moffett, comb. nov.

Rhus dissecta Thunb. in Phytographische Blatter 29 (1803). Type:

Cape of Good Hope. Masson in Herb. Thunberg 7330 (UPS, holo.l).

South Africa.

Searsia divaricata (Eckl. & Zeyh.) Moffett, comb. nov.

Rhus divaricata Eckl. & Zeyh., Enumeratio plantarum Africae aus-

tralis extratropicae 1: 146 (1836). Eastern Cape, Tambukiland, Klip-

plaatrivier, Ecklon & Zeyher 1106 [SI, lecto., designated by Moffett:

53 (1993)].

Lesotho, South Africa.

Searsia dracomontana (Moffett) Moffett, comb. nov.

Rhus dracomontana Moffett in Flora of southern Africa 19,3: 41

(1993). Type: Natal [KwaZulu-Natal], Van Reenen, Schlechter 6754

(BOL, holo.l).

South Africa.

Searsia dregeana (Sond.) Moffett, comb. nov.

Rhus dregeana Sond. in Flora capensis 1: 516 (1860). Type: East-

ern Cape, Stormberg, Mooiplaats, Drege s.n. [SI, lecto., designated by

Moffett: 65 (1993)].

Lesotho, South Africa.

Searsia dumetorum (Exell) Moffett, comb. nov.

Rhus dumetorum Exell in Journal of Botany 66, Supplementum

Polypetalum: 93 (1928). Type; Angola, Bie, Cassuango, Cuiriri, Goss-

weiler 3692 (BM, holo.l).

Angola.

Searsia engleri (Britten) Moffett, comb. nov.

Rhus engleri Britten in Journal of Botany 38; 316 (1900). Type:

Transvaal [Limpopo], Klippan, Rehmann 5325 [Zl, lecto., designated

by Moffett: 61 (1993)].

South Africa.

Searsia erosa (Thunb.) Moffett, comb. nov.

Rhus erosa Thunb., Flora capensis 2: 212 (1818). Type: Cape of

Good Hope, Sparrman ? in Herb. Thunberg 7333 [UPS!, lecto., desig-

nated by Moffett: 69 (1993)].

Lesotho, South Africa.

Searsia fanshawei (R.Fern. & A. Fern.) Moffett,

comb. nov.

Rhus fanshawei R.Fem. & A. Fern, in Boletim Sociedade Brote-

riana, sen 2, 38: 185 (1965b). Type: Zambia, Nkloemfumu, Fanshawe

4767 (K. holo.l).

Zambia.

Searsia fastigata (Eckl. & Zeyh.) Moffett, comb. nov.

Rhus fastigata Eckl. & Zeyh., Enumeratio plantannn Africanae

australis extratropicae 2: 146 (1836). Type: Eastern Cape, Albany and

Uitenhagc, Ecklon & Zeyher 1107 [SI, lecto., designated by Moffett:

51 (1993)].

South Africa.

Searsia flexicaulis (Baker) Moffett, comb. nov.

Rhus flexicaidis Baker in Kew Bulletin 108: 316 (1895). Type; south-

ern Arabia, Hadhramaut, Hirsch 153 (K, holo.l).

Arabia, Egypt, Sudan, Yemen.

Searsia gallagheri (Ghaz.) Moffett, comb. nov.

Rhus gallagheri Ghaz. in Kew Bulletin 57: 492 (2002). Type; Sulta-

nate of Oman, Dhofar, hills above Sharbitat, Hughes & Gallagher

7895/2 (K, holo.).

Arabia (Sultanate of Oman).

Searsia gerrardii (Harv. ex Engl.) Moffett, comb. nov.

Rhus gerrardii (Harv. ex Engl.) Diels in Botanische Jahrbiicher 24:

588 (1898). R. viminalis Vahl van gerrardii Harv. ex Engl.: 442 (1883).

Type: Natal [KwaZulu-Natal], Gerrard & McKen 1396 [Kl, lecto., des-

ignated by R.Fem.: 131 (1967)].

South Africa, Swaziland.

Searsia glauca (Thunb.) Moffett, comb. nov.

Rhus glauca Thunb. in Phytographische Blatter: 27 (1803). Type:

Cape of Good Hope. Thunberg in Herb. Thunberg 7339 [UPS!, lecto.,

designated by Moffett: 82 (1993)].

South Africa.

Searsia glaucescens (A. Rich.) Moffett, comb. nov.

Rhus glaucescens A. Rich., Tentamen Florae Abyssinica 1: 143

(1847). Type: Ethiopia, Crescit in provincie Choa, A. Petit (P, holo.).

Ethiopia, West, Central and East Africa.

Searsia glutinosa (Hochst. exA.Rich.) Moffett subsp.

abyssinica (Oliv.) Moffett, comb. nov.

Rhus abyssinica Oliv. in Flora of tropical Africa 1: 438 (1868).

Type: Ethiopia, Tigray Region, Gennis, near Adowa, Schimper 259 (K,

holo.l).

Eritrea, Ethiopia, Sudan.

Searsia glutinosa (Hochst. exA.Rich.) Moffett, comb,

nov., subsp. glutinosa

Rhus glutinosa Hochst. ex A.Rich., Tentamen Florae Abyssinica

1: 144 (1847). Type: Ethiopia, Gunder Region, near Tschenausa,

Schimper 11: 851 [PI, lecto., designated by Gilbert: 572 (1986b)].

Ethiopia.

Searsia glutinosa (Hochst. ex A.Rich.) Moffett subsp.

neoglutinosa (M.G. Gilbert) Moffett, comb. nov.

Rhus neoglutinosa M.G. Gilbert in Nordic Journal of Botany 6:139

(1986a). Type; Ethiopia, Shewa Region, between Addis Abeba and

Ambo, near Menegesha Village, Jackson 712 (K, holo.l).

Ethiopia.

Searsia gracilipes (Exell) Moffett, comb. nov.

Rhus gracilipes Exell in Journal of Botany 66, Supplementum

Polypetalum: 93 (1928). Type: Angola, Benguela, Caconda, prox. de

Bissapa, Gossweiler 4269 (BM, holo.l).

Angola.

Searsia gracillima (Engl.) Moffett var. glaberrima

(Schonland) Moffett, comb. nov.

Rhus gracillima Engl. var. glaberrima Schonland in Bothalia 3: 86

(1930). Type: Eastern Transvaal [Mpumalanga], Witbank District, hills

near Wilge River, Schlechter 3746 (Z, holo.l).

South Africa.

Searsia gracillima (Engl.) Moffett, comb, nov., var.

gracillima

Rhus gracillima Engl, in Monographiae phanerogamamm 4: 445

(1883). Type: Transvaal [Limpopo], Boshveldt, between Menaar’s

Farm and Elands River, Rehmann 4882 [Zl, lecto., designated by

Moffett: 113 (1993)].

South Africa.

Bothalia 37,2 (2007)

169

Searsia grandidens (Harv. ex Engl.) Mojfett, comb.

nov.

Rhus grandidens Harv. ex Engl, in Monographiae phanerogamarum.

4: 440 (1883). Type: Natal [KwaZulu-Natal], Gerrard & McKen 1399

(K. holo.l).

South Africa, Swaziland.

Searsia grossireticulata (Van der Veken) Moffett,

comb. nov.

Rhus grossireticulata Van der Veken in Bulletin Jardin Botanique

Etat Bruxelle 29: 241 (1959). Type: Democratic Republic of the

Congo, Haut-Katanga, Kisenge-Kapolo, R Duvigneaud 2336 (BRLU,

holo.l).

Democratic Republic of the Congo.

Searsia gueinzii (Sond.) F.A.Barkley in Lilloa 23:

253 (1950).

Rhus gueinzii Sond.: 515 (1860). Type: Natal [KwaZulu-Natal],

Port Natal [Durban], Gueinzius s.n. [TCDl, lecto., designated by

Moffett: 76(1993)].

Mozambique, South Africa, Swaziland. Zimbabwe.

Searsia harveyi (Moffett) Moffett, comb. nov.

Rhus harveyi Moffett in Flora of southern Africa 19,3: 95 (1993).

Type: Natal [KwaZulu-Natal], Zululand, Gerrard & McKen 1406

(TCD, holo.l).

South Africa, Swaziland.

Searsia horrida (Eckl. & Zeyh.) Mojfett, comb. nov.

Rhus horrida Eckl. & Zeyh., Enumeratio plantarum Africanae aus-

tralis extratropicae 2: 146 (1836). Type: Northern Cape. Namaqualand.

Kamiesberg. Ecklon & Zeyher 1135 [SI. lecto., designated by Moffett:

115 (1993)].

South Africa.

Searsia humpatensis (Meikle) Moffett, comb, nov.,

forma humpatensis

Rhus humpatensis Meikle in Boletim Sociedade Broteriana, ser. 2,

26: 284 (1952). Type: Angola, Huila. Humpata, Carriso & Mendonga

609 (COl holo.l).

Angola.

Searsia humpatensis (Meikle) Moffett forma sub-

glabra (R.Fern.) Moffett, comb. nov.

Rhus humpatensis Meikle forma subglabra R.Fern. in Garcia da

Orta (Lisboa) 14,3: 367 (1966). T}q5e: Angola. Sa da Bandeira, Fenda

da Tunda-Vala, R. Santos & Henriques 1143 (LISC. holo.l).

Angola.

Searsia incisa (L.f.) F.A.Barkley var. effusa (Presl)

Moffett, comb. nov.

Rhus incisa L.f. var. effusa (Presl) R.Fern. in Boletim Sociedade

Broteriana. ser. 2, 42: 128 (1967). Type: Eastern Cape. Ecklon &

Zeyher 1111 (PR. holo.-photol).

South Africa.

Searsia incisa ( L.f.) F.A.Barkley var. incisa in Mono-

graphs of the Biological Society of Iraq 3: 54 (1965 ).

Rhus incisa L.f: 183 (1781). Type: Western Cape, near Paardeberg,

Thunberg in Herb. Thunberg 7341 [UPSl, lecto., designated by

R.Fern.: 128 (1967)].

South Africa.

Searsia keetii (Schonland) Moffett, comb. nov.

Rhus keetii Schonland in Bothalia 3: 87 (1930). Type: Eastern

Transvaal [Mpumalanga], Lydenburg District, Klip River, Steelpoort

Park. Keet 1435 (GRA, holo.l).

South Africa.

Searsia kirkii (Oliv.) Moffett, comb. nov.

Rhus kirkii Oliv. in Flora of tropical Africa 1: 439 (1868). Type:

Zimbabwe, near Victoria Falls, Kirk s.n. (K, holo.l).

Angola, Democratic Republic of the Congo, Namibia, Zambia,

Zimbabwe.

Searsia krebsiana (Presl ex Engl.) Moffett, comb. nov.

Rhus krebsiana Presl ex Engl, in Monographiae phanerogamarum

4: 409 (1883). Type: Cape of Good Hope, Krebs s.n. [G-DC (Mon.

Phan.), holo.l].

South Africa.

Searsia kwangoensis (Van der Veken) Moffett, comb.

nov.

Rhits kwangoensis (Van der Veken) Kokwaro in Kew Bulletin

34: 753 (1980). R. kirkii Oliv. var. kwangoensis Van der Veken: 242

(1959). Type: Democratic Republic of the Congo, Lukuni, Callens

3015 {BR. holo.l).

Democratic Republic of the Congo, Uganda.

Searsia kwazuluana (Moffett) Moffett, comb. nov.

Rhus kn’azuluana Moffett in Flora of southern Africa 19.3: 71

( 1993). Type: Natal [KwaZulu-Natal], St Lucia, Mbomvini, MacDevette

353 (PRE, holo.l).

South Africa.

Searsia laevigata (L.) F.A.Barkley var. laevigata

forma cangoana (Moffett) Moffett, comb. nov.

Rhus laevigata L. forma cangoana Moffett in Flora of southern

Africa 19,3: 38 (1993). Type: Western Cape, Oudtshoom District,

Boomplaas, Moffett 118 (PRE, holo.l).

South Africa.

Searsia laevigata (L.) F.A.Barkley var. laevigata

forma laevigata in Monographs of the Biological Society

of Iraq 3: 54(1965).

Rhus laevigata L.: 1672 (1763). Type: Cape of Good Hope, Herb.

LINN 378-23 [LINN!, lecto., designated by R.Fern.: 129 (1967)].

South Africa.

Searsia laevigata (L.) F.A.Barkley var. villosa (L.f)

Moffett, comb. nov.

Rhus laevigata L. var. villosa (L.f) R.Fern. in Boletim Sociedade

Broteriana, ser. 2, 42: 130 (1967). Type: Cape of Good Hope, Herb.

LINN 378.26 [LINN!, lecto., designated by Schonland: 17 (1930)].

South Africa.

Searsia lancea (L.f.) F.A.Barkley in Flora of Texas

3: 104(1943).

Rhus lancea L.f: 184 (1781). Type: Cape of Good Hope, Thunberg

in Herb. Thunberg 7348j3 [UPS!, lecto., designated by Moffett: 24

(1984)].

Botswana. Lesotho. Namibia, South Africa, Zimbabwe.

Searsia leptodictya (Diels) T.S.Yi, A.J.Mill. & J.Wen

forma leptodictya in Molecular Phylogenetics & Evolution

33: 861 (2004).

Rhus leptodictya Diels: 86 (1907). Type: Transvaal [Gauteng],

Pretoria, Reck 13 [GRAl, neo., designated by R.Fern. & A. Fern.: 699

(1965a)].

Angola, Mozambique, Namibia. South Africa, Swaziland, Zimbabwe.

Searsia leptodictya (Diels) T.S. Yi, A.J.Mill. & J. Wen

forma pilosa (R.Fern. & A.Fem.) Moffett, comb. nov.

Rhus leptodictya Diels forma pilosa R.Fern. & A.Fem.: 187

(1965b). Type: Zimbabwe, Ndanga, zlm/toge 104/55 (SRGH, holo.l).

Malawi. Zimbabwe.

170

Bothalia 37,2 (2007)

Searsia longipes (Engl.) Mojfett var. elgonensis

(Kokwaro) Moffett, comb. nov.

Rhus longipes Engl. var. elgonensis Kokwaro in Kew Bulletin 34:

754 (1980). Type: Kenya, Mt Elgon, T.H.E. Jackson 3l2a (EA, holo.).

Kenya, Uganda.

Searsia longipes (Engl.) Moffett, comb, nov., var. lon-

gipes

Rhus longipes Engl, in Monographiae phanerogamarum 4: 431

(1883). Type: Angola, Cuanze Norte, Carengwe, Queta, Welwitsch

4413 [G-DC (Mon. Phan.), holo.!].

From Sierra Leone in the west to Ethiopia in the east and as far south

as Angola and Zimbabwe.

Searsia longipes (Engl.) Moffett var. schinoides (R.

Fem.) Moffett, comb. nov.

Rhus longipes Engl. var. schinoides R.Fem. in Memoire Junta

Investigado da Ultramar, ser. 2, 38: 39 (1962). Type: Zambia, 13 km

NW of Abercom, Hutchinson & Gillett 4010 (K, holo.!).

Kenya, Tanzania, Zambia.

Searsia longispina (Eckl. & Zeyh.) Moffett, comb. nov.

Rhus longispina Eckl. & Zeyh., Enumeratio plantarum Afficae

australis extratropicae 2: 148 (1836). Type: Eastern Cape, Ecklon &

Zeyher 1116 p.p. [SAM!, lecto., designated by Moffett: 83 (1993)].

South Africa.

Searsia lucens (Hutch.) Moffett, comb. nov.

Rhus lucens Hutch., A botanist in southern Africa: 480 (1946).

Type: Zimbabwe, near Victoria Falls, Hutchinson & Gillett 34 73 (K,

holo.!).

Botswana, Namibia, Zimbabwe.

Searsia lucida (L.) FA. Barkley forma elliptica

(Sond.) Moffett, comb. nov.

Rhus lucida L. forma elliptica (Sond.) Moffett in Flora of southern

Africa 19,3: 81 (1993). R. elliptica Sond.: 517 (1860). Type: Western

Cape, mouth of Onrustrivier, Zeyher 2248 [K!, lecto., designated by

Moffett: 81 (1993)].

South Africa.

Searsia lucida (L.) F.A.Barkley fornia lucida in Mo-

nographs of the Biological Society of Iraq 3:54 (1965).

Rhus lucida L.: 267 (1753). Type: Cape of Good Hope, Herb. Cliff.

Ill, 6d(32) [BM!, lecto., designated by Moffett: 79 (1993)].

South Africa, Zimbabwe.

Searsia lucida (L.) F.A.Barkley forma scoparia

(Eckl. & Zeyh.) Moffett, comb. nov.

Rhus lucida L. forma scoparia (Eckl. & Zeyh.) Moffett in Flora

of southern Africa 19,3: 79 (1993). R. scoparia Eckl. & Zeyh.: 149

(1836). Type: Eastern Cape, Ecklon & Zeyher 1122 [SAM!, lecto., des-

ignated by Moffett: 79 (1993)].

South Africa.

Searsia magalismontana (Sond.) Moffett subsp.

coddii (R.Fern. & A. Fern.) Moffett, comb. nov.

Rhus magalismontana Sond. subsp. coddii (K.¥em. & A. Fern.) Mof-

fett in Flora of southern Africa 19,3: 57 (1993). R. coddii R.Fem. &

A. Fem.: 251 (1965c). Type: Northern Transvaal [Limpopo], Venda,

Sambandou, Codd 6902 (PRE, holo.!).

South Africa.

Searsia magalismontana (Sond.) Moffett, comb,

nov., subsp. magalismontana

Rhus magalismontana Sond. in Flora capensis 1 : 510 (1860). Type:

Transvaal [Gauteng], Magaliesberg, Crocodile River, Zeyher 341 (S,

holo.!).

Botswana, South Africa.

Searsia magalismontana (Sond.) Moffett subsp. tri-

foliolata (Baker f.) Moffett, comb. nov.

Rhus magalismontana Sond. subsp. trifoliolata (Baker f.) Mof-

fett in Flora of southern Africa 19,3: 59 (1993). R. trifoliolata

Baker f.: 429 (1899). Type: Zimbabwe, Bulawayo, Rand 66 (BM,

holo.!).

South Africa, Zimbabwe.

Searsia maricoana (Moffett) Moffett, comb. nov.

Rhus maricoana Moffett in Flora of southern Africa 19,3: 110

(1993). Type: Transvaal [North-West], Zeemst, Marico chrome mine,

Moffett 3566 (PRE, holo.!).

South Africa.

Searsia marlothii (Engl.) Moffett, comb. nov.

Rhus marlothii Engl, in Botanische Jahrbiicher 10: 37 (1888). Type:

South West Africa [Namibia], Otjimbingue, Marloth 1394 [K!, lecto.,

designated by Moffett: 61 (1993)].

Namibia.

Searsia montana (Diels) Moffett, comb. nov.

Rhus montana Diels in Botanische Jahrbiicher 40: 86 (1907). Type:

Eastern Cape, Transkei, near Engcobo, Bolus 8837 [K!, lecto., desig-

nated by R.Fem.: 131 (1967)].

Lesotho, South Africa.

Searsia monticola (Meikle) Moffett, comb. nov.

Rhus monticola Meikle in Memoirs of the New York Botanical

Garden 8: 242 (1953b). Type: Malawi, Mulanje Mtn, Luchenya Plateau,

Brass 16656 (K, holo.!).

Malawi.

Searsia mysorensis (G.Don) Moffett, comb. nov.

Rhus mysorensis G.Don, A general system of gardening and botany

2: 74 (1832). Type: India, Herb. Wallich 997 [K (Wallich)!, sheet with

4 elements: a, Rhus indicum nob. from the Governor’s Garden, Jan. 17,

1804; b, Rhus mysorensis ex herbareo Heyneano in Horto Botanico

Calcutta; b here designated as neo.]* **.

India.

Searsia natalensis (Bernh. ex Kraiiss) F.A.Barkley

in Lilloa23: 253 (1950).

Rhus natalensis Bemh. ex Krauss: 349 (1844)’*’*. Type: Durban,

forests around Natal Bay, Krauss 395 [TUB!, lecto., designated by

Moffett: 75 (1993)].

Mozambique, Reunion, South Africa.

Searsia nebulosa (Schonland) Moffett, comb, nov.,

fonna nebulosa

Rhus nebulosa Schonland in Bothalia 3: 33 (1930). Type: Natal

[KwaZulu-Natal], near Durban, Schlechter 2858 [GRA!, lecto., desig-

nated by Moffett: 39 (1993)].

Mozambique, Reunion, South Africa.

Searsia nebulosa (Schonland) Moffett fonna pube-

scens (Moffett) Moffett, comb. nov.

Rhus nebulosa Schonland forma pubescens Moffett in Flora of

southern Africa 19,3: 40 (1993). Type: Eastern Cape, Alexandria

Forest, Olifantshoek, Jo/;w5on 649 (PRE, holo.!).

South Africa.

* No specimen annotated by G.Don could be found, thus requiring the

designation of a neotype. Both a and b on the above sheet are almost

identical and any one could have been chosen.

** Bemhardi is sometimes cited as the author of the new names based

on Krauss’s specimens. In this particular case, however, Krauss himself

should be recognized as he used the words ‘in specimen meo’ when

citing Cissus natalensis Bemh. olim in sched.

Bothalia 37,2 (2007)

171

Searsia nitida (Engl.) Moffett, comb. nov.

Rhus nitida Engl, in Monographiae phanerogamamm 4: 434 (1883).

Type: Angola, Cuanza Norte, entre Mutoto e Candumba, Pungo An-

dongo, Welwitsch 4417 [G-DC (Mon. Phan.), holo.!].

Angola.

Searsia obtusata (Engl.) Mojfett, comb. nov.

Rhus obtusata (Engl.) Meikle in Boletim Sociedade Broteriana, ser,

2, 26: 287 (1952). R. villosa L.f. var. obtusata Engl.: 425 (1883). Type:

Angola, Mopamedes, Base da Serra da Chela, Welwitsch 4419 [G-DC

(Mon. Phan.), holo.!].

Angola.

Searsia ochracea (Meikle) Moffett, comb, nov., var.

ochracea

Rhus ochracea Meikle in Kew Bulletin 8: 107 (1953a). Type: Mala-

wi, Fort Hill, Whyte s.n. (K, holo.!).

Malawi, Tanzania, Zambia.

Searsia ochracea (Meikle) Mojfett var. saxicola

(R.Fern. & A.Fern.) Moffett, comb. nov.

Rhus ochracea Meikle var. saxicola R.Fern. & A. Fern, in Boletim

Sociedade Broteriana, ser. 2, 38: 189 (1965b). Type: Zambia, Mpika,

Muchinga Escarpment, Angus 865 (K, holo.! ).

Zambia.

Searsia pallens (Eckl. & Zeyh.) Mojfett, comb. nov.

Rhus pallens Eckl. & Zeyh., Enumeratio plantarum africanae austra-

lis extratropicae 2: 147 (1836). Type: Eastern Cape, Uitenhage, Ecklon

Zeyher 1114 [SAM!, lecto., designated by Moffett: 85 (1993)].

Lesotho, South Africa.

Searsia paniculata (Wall, ex G.Don) Moffett, comb.

nov.

Rhus paniculata Wall, ex G.Don, A general system of gardening

and botany 2: 73 (1832). Type: Burma [Myanmar], Yenanghuen, Herb.

Wallich 993 [K (Wallich)!, sheet with two elements: right hand ele-

ment, S, Sept. 1826, neo., here designated]*.

Bhutan, China, India, Myanmar.

Searsia parviflora (Roxb.) FA. Barkley in Lilloa 23:

253 (1950).

Rhus parviflora Roxb.: 100 (1832). Type: Nepal & Hort. Calcutta,

Roxburgh (BM!, sheet marked Rhus paniflonim Roxb., Patria Nepaul,

HB1815, lecto., here designated].

India, Nepal.

Searsia pendulina (Jacq.) Mojfett, comb. nov.

Rhus pendulina Jacq., Plantarum rariorum horti caesarei schoenb-

runnensis 4: 24, t. 449 (1804). Type: ex hort Schonbrunn. Jacquin s.n.

[M!, lecto., designated by Moffett: 91 (1993)].

Namibia, South Africa.

Searsia pentaphylla (Jacq.) FA. Barkley in Mono-

graphs of the Biological Society of Iraq 3:57 (1965).

Rhamnus pentaphylla Jacq.: 27 (1767), (Rhamnus Siculus pen-

taphyllos Boccone: 43, t. 21, 1674). Type: Sicily, Boccone s.n. ex Dr

Jacquin (BM!, sheet 92. lecto., here designated.).

Algeria, Israel, Morocco, Palestine, Sicily.

Searsia pentheri (Zahlbr.) Moffett, comb. nov.

Rhus pentheri Zahlbr. in Annalen des Naturhistorischen Museums

in Wien 15,1: 52 (1900). Type: Natal [KwaZulu-Natal], Colossa, Krook

sub. Penther 2290 (W, holo.!).

Mozambique, South Africa, Swaziland.

* Don cited ‘(Wall. mss. in herb. Lin. soc.)’. No Wallich specimen was

found in LINN, and as no other specimens annotated by him could be

traced, a neotype had to be chosen.

Searsia pondoensis (Schonland) Mojfett, comb. nov.

Rhus pondoensis Schonland in Bothalia 3: 95 (1930). Type: Natal

[KwaZulu-Natal], near Murchison, 7. Medley Wood 3002 (SAM, holo.!).

South Africa, Swaziland.

Searsia populifolia (E.Mey. ex Sond.) Moffett,

comb. nov.

Rhus populifolia E.Mey. ex Sond. in Flora capensis 1: 508 (1860).

Type: Northern Cape, mouth of the Gariep [Orange] River, Drege s.n.

[TCD!, lecto., designated by Moffett: 105 (1993)].

Namibia. South Africa.

Searsia problematodes (Merxm. & Rossi.) Moffett,

comb. nov.

Rhus problematodes Merxm. & Rbssl. in Mitteilungen der Bota-

nischen Staatssamlung Miinchen 11: 66 (1973). Type: South West

Africa [Namibia], near Aus, Plateau/Aar, Wiss 3001 (M, holo.!).

Namibia.

Searsia pterota (Presl) Mojfett, comb. nov.

Rhus pterota Presl in Botanische Bemerkungen: 44 (1884). Type:

Eastern Cape, Ecklon & Zeyher 1116 (PRC, holo.-photo.l).

South Africa.

Searsia puccionii (Chiov.) Mojfett, comb. nov.

Rhus puccionii Chiov. in Flora Somala 1: 132 (1929). Type:

Somalia, coast of Migiurtini, near mouth of Nogal River, Puccioni &

Stefanini 849 (FT-photo.!, lecto, here designated).

Somalia.

Searsia pygmaea (Moffett) Mojfett, comb. nov.

Rhus pygmaea Moffett in Botanical Journal of the Linnean Society

130: 39 (1999). Type: Mpumalanga, Barberton, near Agnes Mine,

Moffett 4905 (PRE, holo.! ).

South Africa.

Searsia pyroides (Burch.) Moffett var. dinteri (Engl.)

Moffett, comb. nov.

Rhus pyroides (Burch.) Moffett var. dinteri (Engl.) Moffett in Flora

of southern Africa 19,3: 43 (1993). R. dinteri Engl.: 211 (1921). Type:

South West Africa [Namibia], Schaaprivier, Dinter 1898 [SAM!, lecto.,

designated by Moffett: 43 (1993)].

Namibia.

Searsia pyroides (Burch.) Mojfett var. gracilis (Engl.)

Moffett, comb. nov.

Rhus villosa L.f var. gracilis Engl, in Monographiae phaneroga-

marum 4: 425 (1883). Type: Transvaal [Gauteng], Pretoria, Rehmann

4742 [Z!, lecto., designated by Moffett: 45 (1993)].

Lesotho, South Africa. Swaziland.

Searsia pyroides (Burch.) Moffett var. integrifolia

(Engl.) Moffett, comb. nov.

Rhus tridentata Engl. var. integrifolia Engl, in Monographiae pha-

nerogamarum 4: 426 (1883). Type: Natal [KwaZulu-Natal], Inanda,

Rehmann s.n. [Z!, lecto., designated by Moffett: 45 (1993)].

South Africa, Swaziland.

Searsia pyroides (Burch.) Moffett, comb, nov., var.

pyroides**

Rhus pyroides Burch, var. pyroides. Travels in the interior of

southern Africa 1: 344 (1822). Type: Northern Cape, Asbestos Mtns,

Burchell 1796 (K. holo.!).

Ethiopia, Kenya, Lesotho, South Africa, Swaziland, Tanzania, Uganda,

Zambia, Zimbabwe.

** In Central and East Africa this taxom is known as R. vulgaris Meikle.

172

Bothalia 37,2 (2007)

Searsia quartiniana (A.Rich.) A.JMill. inA.J. Mill,

et al. in International Journal of Plant Sciences 162: 1403

(2001).

Rhus quartiniana A.Rich.: 141 (1847). Type: Ethiopia, Tigray,

Shire, Quartin-Dillon & Petit s.n, (P, holo.!).

Angola, Democratic Republic of the Congo, Ethiopia, Kenya, Namibia,

South Africa, Tanzania, Zambia, Zimbabwe.

Searsia refracta (Eckl. & Zeyh.) Mojfett, comb. nov.

Rhus refracta Eckl. & Zeyh., Enumeratio plantarum africanae

australis extratropicae 2: 145 (1836). Type: Eastern Cape, Uitenhage,

Zwartkops River, Ecklon & Zeyher 1103 [S!, lecto., designated by

Moffett: 73 (1993)].

South Africa.

Searsia rehmanniana (Engl.) Moffett var. glabrata

(Sond.) Moffett, comb. nov.

Rhus pyroides Burch, var. glabrata Sond. in Flora capensis 1:511

(1860). Type: Natal [KwaZulu-Natal], Oomcomas [Mkomaas], Drege

5580 [S!, lecto., here designated, replacing the Drege 6800 in P lecto-

type chosen by Moffett: 49 (1993)]*.

South Africa, Swaziland, Zambia, Zimbabwe.

Searsia rehmanniana (Engl.) Moffett, comb, nov.,

var. rehmanniana

Rhus rehmanniana Engl, in Monographiae phanerogamarum 4: 422

(1883). Type: Transvaal [Limpopo], Houtbosch, Rehmann 5560 [Z!,

lecto., designated by Moffett: 49 (1993)].

Mozambique, South Africa, Swaziland.

Searsia retinorrhoea (Steud. ex Oliv.) Moffett, comb.

nov.

Rhus retinorrhoea Steud. ex Oliv. in Flora of tropical Africa 1: 438

(1868). Type: Ethiopia, Tigray Region, Dscheladscheranne, Schimper

lit: 1627 (K, holo.!).

Ethiopia, Somalia, Sudan, Yemen.

Searsia rigida (Mill) F.A.Barkley var. dentata (Engl.)

Moffett, comb. nov.

Rhus zeyheri Sond. var. dentata Engl, in Monographiae phanero-

gamarum 4: 433 (1883). Type: Eastern Transvaal [Mpumalanga],

Transvaal Drakensberg, Laingsnek, Rehmann 6942 (Z, holo.!).

South Africa.

Searsia rigida (Mill.) F.A.Barkley var. margaretae

{Bur tt Davy ex Moffett) Moffett, comb. nov.

Rhus rigida (Mill.) Moffett var. margaretae Burtt Davy ex Moffett

in Flora of southern Africa 19,3: 36 (1993). Type: Transvaal [Gauteng],

Elsburg, Schlechter 3539 (PRE, holo.!).

South Africa, Swaziland.

Searsia rigida (Mill.) F.A.Barkley var. rigida in

Monographs of the Biological Society of Iraq 3: 54 (1965).

Rhus rigida Mill.: 14 (1768). Type: ex Herb. Miller (BM, holo.!).

South Africa, Swaziland.

Searsia rimosa (Eckl. & Zeyh.) Moffett, comb. nov.

Rhus rimosa Eckl. & Zeyh., Enumeratio plantarum africanae aus-

tralis extratropicae 2; 150 0836). Type: Western Cape, Clanwilliam,

Meerenlogement, Ecklon & Zeyher 1124 [SAM sheet 2557!, lecto.,

designated by Moffett: 91 (1993)].

South Africa.

*Although both Drege specimens are the same taxon, a referee pointed

out that 5580 is preferable as it has been annotated as [3 glabrata by

Sonder.

Searsia rogersii (Schonland) Moffett, comb. nov.

Rhus rogersii Schonland in Bothalia 3: 42 (1930). Type: Eastern

Transvaal, [Mpumalanga], Barberton, Rogers 18270 [GRAl, lecto.,

designated by Moffett: 35 (1993)].

South Africa, Swaziland.

Searsia rosmarinifolia (Vahl) F.A.Barkley in Mono-

graphs of the Biological Society of Iraq 3: 53 (1965).

Rhus rosmarinifolia Vahl: 50 (1794). Type: Locality unknown, ex

Herb. Hoffman-Bang e coll. Vahl, Bulow s.n. [C!, lecto., designated by

Moffett: 99 (1993)].

South Africa.

Searsia rudatisii (Engl.) Moffett, comb. nov.

Rhus rudatisii Engl., Die Pflanzenwelt Afrikas 3,2: 217 (1921).

Type: Natal [KwaZulu-Natal], Alexandra County, Friedenau, Mgai

Flats, Rudatis 698 [K!, lecto., designated by Moffett: 109 (1993)].

South Africa.

Searsia ruspolii (Engl.) Moffett, comb. nov.

Rhus ruspolii Engl, in Annuario Regius Istituto Botanico di Roma

7: 18 (1897-1898). Type: Ethiopia, Biddume, Ruspoli & Riva 1317

(FT, holo.).

Democratic Republic of the Congo, Ethiopia, Kenya, Uganda.

Searsia scytophylla (Eckl. & Zeyh.) Moffett var.

dentata (Moffett) Moffett, comb. nov.

Rhus scytophylla Eckl. & Zeyh. var. dentata Moffett in Flora of south-

ern Africa 19,1: 93. Type: Western Cape, Ceres District, Agterwitzenberg,

Modderrivierskloof. Van Jaarsveld 1538 (NBG, holo.!).

South Africa.

Searsia scytophylla (Eckl. & Zeyh.) Moffett, comb,

nov., var. scytophylla

Rhus scytophylla Eckl. & Zeyh., Enumeratio plantarum Africae

australis extratropicae 2: 150 (1836). Type: Western Cape, between Sir

Lowry’s Pass and Palmiet River, Grietjiesgat, Ecklon & Zeyher 1130

[S, sheet 1 !, lecto., designated by Moffett: 93 (1993)].

South Africa.

Searsia sekhukhuniensis (Moffett) Moffett, comb. nov.

Rhus sekhukhuniensis Moffett in Flora of southern Africa 19,3: 77

(1993). Type: Eastern Transvaal [Mpumalanga], Lydenburg District,

Steelpoort Park Pass, Moffett 2000 (PRE, holo.!).

South Africa.

Searsia somalensis (Engl.) Moffett, comb. nov.

Rhus somalensis Engl, in Abhandling der Koniglike Preussiche

Akademie fur Wissenschaft 2: 289 (1891-1892). Type: Somalia, near

Mei'd, Semit Mtns, Hildebrand 1542 (K!, lecto., here designated).

Ethiopia, Somalia.

Searsia squalida (Meikle) Moffett, comb. nov.

Rhus squalida Meikle in Boletim Sociedade Broteriana, ser. 2, 26:

283 (1952). Type: Angola, Calumba, Welwitsch 4421 (BM, holo.!).

Angola, Zambia.

Searsia stenophylla (Eckl. & Zeyh.) Moffett, comb.

nov.

Rhus stenophylla Eckl. & Zeyh., Enumeratio plantarum Africae

australis extratropicae 2: 144 (1836). Type: Western Cape, Table

Mountain and Hottentotshollandberge, Ecklon & Zeyher 1094 [S!,

lecto., designated by Moffett: 99 (1993)].

South Africa.

Searsia tenuinervis (Engl.) Moffett, comb. nov.

Rhus tenuinervis Engl, in Monographieae phanerogamarum 4: 423

(1883). Type: Angola, between Benguela and R. Catumbela, Welwitsch

4418 [G-DC (Mon. Phan.), holo.!].

Angola, Botswana, Democratic Republic of the Congo, Ethiopia, Kenya,

Malawi, Moqambique, Namibia, South Africa, Sudan, Tanzania, Zambia,

Zimbabwe,

Bothalia 37,2 (2007)

173

Searsia tenuipes (R.Fern. & A.Fem.) Moffett, comb.

nov.

Rhus tenuipes R.Fem. & A.Fem. in Boletim Sociedade Broteriana,

ser. 2, 38; 191 (1965b). Type: Zimbabwe, Great Dyke, Mhlaba Hills,

Charter, Wild 5667 (SRGH, hole.!).

Mozambique, Zimbabwe.

Searsia thyrsiflora (Balf.f.) Moffett, comb. nov.

Rhus thyrsiflora Balf.f. in Proceedings of the Royal Society,

Edinburgh 11: 507 (1882). Type: Socotra, Balfour369 (BM!, lecto.,

here designated).

Socotra.

Searsia tomentosa (L.) F.A.Barklev in Flora of Texas

3:104(1943).

Rhus tomentosa L.: 266 (1753). Type: Cape of Good Hope, Herb.

LINN 378.20 [LINN, lecto.!, Wijnands: 42 (1983)].

South Africa, Zimbabwe.

Searsia transvaalensis (Engl.) Moffett, comb. nov.

Rhus transvaalensis Engl, in Monographiae phanerogamamm 4:

440 (1883). T)q)e: Northern Transvaal [Limpopo], Houtbosch, Rehmann

5559 [Zi, lecto., designated by Moffett: 29 (1993)].

South Africa, Swaziland.

Searsia tridactyla (Burch.) Moffett, comb. nov.

Rhus tridactyla Burch., Travels in the interior of southern Africa 1 :

340 (1822). Type: Northern Cape, Asbestos Mtns, Burchell 1667 (K,

holo.l).

South Africa.

Searsia tripartita (Ucria) Moffett, comb. nov.

Rhamnus tripartitus Ucria in Nuovo race, opusculum autore cata-

logus plantamm Siciliae 6: 249 (1793), Sicily. [Icono., here designated;

left hand figure of plate Rhamnus polytriphyllus, p. 2 1 of reprint by

Carta (1959) of Plantae ad Linneanum opus addendae — Beraadino da

Ucria].

Algeria, Egypt. Israel, Jordan, Libya, Morocco, Palestine, Niger, Sicily,

Sudan, Syria, Tunisia. Western Sahara.

Searsia tumulicola (S. Moore) Moffett var. meeuse-

ana (R.Fem. & A.Fem.) Moffett, comb, nov., forma meeu-

seana

Rhus synstylica R.Fem. & A.Fem. var. meeuseana R.Fem. &

A.Fem. forma meeuseana in Boletim Sociedade Broteriana, ser. 2,

39: 251 (1965c). Type: Northern Transvaal [Limpopo], Venda, Lake

Funduzi, Meeuse 9409 (PRE, holo.l).

South Africa.

Searsia tumulicola (S. Moore) Moffett var. meeuse-

ana (R.Fem. & A.Fem.) Moffett forma pumila (Moffett)

Moffett, comb. nov.

Rhus tumulicola (S. Moore) Moffett var. meeuseana (R.Fem. &

A.Fem.) Moffett forma pumila Moffett in Flora of southern Africa

19,3: 25 (1993). Type: Eastern Transvaal [Mpumalanga], Carolina,

Galpin 12493 (PRE, holo.l).

South Africa.

Searsia tumulicola (S. Moore) Moffett, comb, nov.,

var. tumulicola

Rhus tumulicola S. Moore in Journal of Botany 59: 227 (1921).

Type: Northern Transvaal [Limpopo], The Downs, Rogers 22033 (BM,

holo.l).

South Afnca, Zimbabwe.

Searsia undulata (Jacq.) T.S.Ti, A.JMill. & J.Wen

in Molecular Phylogenetics and Evolution 33: 861 (2004).

Rhus undulata Jacq.: 52 (1798). Type: ex Hort. Schdnbmnn,

Jacquin 379 [Wl, lecto., designated by R.Fem.: 133 (1967)].

South Africa.

Searsia volkii (Siisseng.) Moffett, comb. nov.

Rhus volkii Siisseng. in Mitteilungen der Botanischen Staatssamlung

Miinchen 1,8: 343 (1953). Type: South Africa [Namibia], Great

Namaland, slopes of Tsaris Mtns, Volk 752 (M, holo.l).

Namibia.

Searsia wellmanii (Engl.) Moffett, comb. nov.

Rhus wellmanii Engl., Die Pflanzenwelt Afrikas 3,2: 213 (1921).

Type; Angola, Benguella District, De Moco, Gossweiler 12357 (LISCI,

neo., designated by R.Fem.: 356 (1966)].

Angola.

Searsia wildii (R.Fern. & A.Fem.) Moffett, comb. nov.

Rhus wildii R.Fem. & A.Fem. in Boletim Sociedade Broteriana,

ser. 2, 38: 192 (1965b). Type: Zimbabwe, Sipolilo, Mpingi Pass, Wild

5776 (SRGH. holo.l).

Zimbabwe.

Searsia wilmsii (Diels) Moffett, comb. nov.

Rhus wilmsii Diels in Botanische Jahrbucher 24: 501 (1898). Type:

Eastern Transvaal [Mpumalanga], Lydenburg District, Wilms 249

[AMD!, lecto.. designated by Moffett: 111 (1993)].

South Africa.

Searsia zeyheri (Bond.) Moffett, comb. nov.

Rhus zeyheri Sond. in Flora capensis 1 : 514(1 860). Type: Transvaal

[Gauteng], among shmbs at Magalisberg, Zeyher 345 (S, holo.l).

South Africa.

ACKNOWLEDGEMENTS

I am grateful to the following persons who have assis-

ted me in completing this paper: Laura Settesoldi of the

Erbario Tropicale, University of Florence and Wang

Yuhua of Kunming Institute of Botany, China for help

in tracing obscure publications and for providing me

with images of important specimens; Raul Puente of the

Desert Botanic Garden, Phoenix, Arizona for sending

me images of the voucher specimens used in the recent

phylogenetic studies; Robert Archer and Hugh Glen of

SANBI, Ted Oliver, formerly SANBI, Ashley Nicholas

(when S.A. Liaison Officer at Kew), Peter Linder of the

Institute of Systematic Botany, University of Zurich and

John McNeill of the Royal Botanic Garden, Edinburgh,

who all gave nomenclatural advice and three referees

whose recommendations improved the paper. I am also

grateful to the Department of Plant Sciences (Main &

Qwaqwa Campus) of the University of the Free State,

who provided logistical support.

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Bothalia 37,2: 177-182 (2007)

New species and notes on Hesperantha (Iridaceae) in southern Africa

P. GOLDLATT * and J.C. MANNING**

Keywords: Hesperantha Ker Gawl., Iridaceae, southern Africa, taxonomy

ABSTRACT

Field studies of the sub-Saharan African and largely southern African Hesperantha conducted since 2003 have resulted in

the discovery of three new species in this genus, bringing the total to 82. Hesperantha longistyla J.C. Manning & Goldblatt,

known from one collection from the mountains of SW Namibia, is a dwarf plant with moderately long-tubed, purple flowers

and unusually long style branches, possibly allied to the Kamiesberg species, H. latifolia. A second species. H. helmei

Goldblatt & J.C. Manning, also known from a single collection from the interior mountains of Eastern Cape near Graaff-

Reinet, has terete leaves and small flowers with tepals ± 8 x 2.5 mm, about as long as the perianth tube. It is apparently

most closely allied to the Roggeveld species, H. ciliolata. A third novelty, H. lithicola J.C. Manning & Goldblatt, restricted

to the Swartruggens range in the eastern Cold Bokkeveld. has bell-shaped corms with toothed margins, leaves 1-2 mm

wide, and white flowers with a tube 10-12 mm long, and appears most closely allied to the widespread H. falcata. A new

collection of H. karooica from northeast of the Hantamsberg represents a small but significant range extension for this local

endemic previously known from just two collections near Calvinia, south of these mountains. The flower size, especially

dimensions of the tepals, confirms its status as a separate species allied to H. vaginata. Lastly, new collections of the relatively

uncommon, yellow-flowered variant of H. acuta show that this plant, confined to the eastern portion of the range of the

species, differs consistently from the white-flowered form in several floral features, and it is raised to subspecies rank as H.

acuta subsp. tugwelUae.

INTRODUCTION

Hesperantha Ker Gawl. (Iridaceae: Crocoideae) is dis-

tributed across sub-Saharan Africa but is most diverse and

species-rich in southern Africa. The genus has two cen-

tres of diversity here, the winter rainfall west (Goldblatt

1984), and the coast and adjacent mountains of the east-

ern, summer rainfall half of the subcontinent (Hilliard

& Burtt 1986). In the most recent revision of the genus,

79 species were recognized (Goldblatt 2003). Although

Hesperantha is now well understood taxonomically,

three collections made since the publication of this

revision represent novelties, all from areas of southern

Africa that are poorly collected. We describe them here.

In addition, a new collection and important range exten-

sion of the poorly known H. karooica Goldblatt confirm

that flower size, especially dimensions of the tepals, con-

sistently differs from that in the closely allied H. vagi-

nata (Sweet) Goldblatt, removing doubts about its status

as merely a depauperate form of the latter. In H. acuta

new collections have provided convincing evidence that

the eastern, yellow-flowered populations of the species

consistently have larger flowers that differ in several

additional features from the western, white-flowered

populations. The yellow-flowered populations clearly

constitute a separate race of the species, which we recog-

nize as H. acuta subsp. tiigH’elliae. The total number of

species in Hesperantha is thus increased to 82: 43 of

these occur in the southern African winter rainfall zone;

39 in the southern African summer rainfall zone south of

the Limpopo; and four in tropical Africa.

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

RO. Box 299. St. Louis, Missouri 63166, USA.

E-mail: peter. goldblatt(§mobot.org.

** Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

E-mail: manning(g!sanbi.org.

MS. received: 2007-01-09.

TAXONOMY

Hesperantha \or\gi%ty\a J.C. Manning & Goldblatt,

sp. nov.

Plantae ± 30 mm altae, conno campanulato ± 10 mm

diam. margine spinis radiatis amiato, caule ad 8 mm

longo, foliis 3 falcatis 30-50 x 2. 5-3.0 mm, spica 1- vel

2-flora, bracteis viridibus 7-10 mm longis, floribus mani-

feste purpureis ad faucem tubi flavis, tubo perianthii

20-25 mm longo, tepalis subaequalibus anguste ovatis

patentibus, staminibus adscendentibus, antheris ± 5 mm

longis, ramis styli ± 10 mm longis laxe patentibus.

TYPE. — Namibia, 2716 (Witputz): south of Sebra-

fontein, 1 400 m, (-DD), 28 August 2003, P. Bruvns

9484A (NBG, holo.).

Plants ± 30 mm high; coma bell-shaped with flat base,

± 10 mm diam. at base, margin with radiating spines.

Stem reaching up to 8 mm above ground, unbranched.

Leaves 3, falcate, 30-50 x 2. 5-3.0 mm; cataphylls mem-

branous. Spike 1- or 2-flowered; bracts green, 7-10 mm

long, outer about as long as inner or slightly longer,

inner forked at apex. Flowers evidently purple, yellow

in mouth of tube; perianth tube 20-25 mm long, slender,

widening near apex; tepals subequal, narrowly ovate, ±

10x6 mm, outer slightly wider than inner, spreading

horizontally. Stamens symmetrically disposed, slightly

spreading; filaments ± 4 mm long, exserted ± 3 mm from

tube; anthers ± 5 mm long. Ovary ovoid, ±1.5 mm long;

style dividing at mouth of perianth tube, branches ± 10

mm long, spreading laxly. Capsules and seeds unknown.

Flowering time: mid-August to early September. Figure

lA.

Distribution and ecology, found in rock crevices

at higher elevations in the mountains of southwestern

178

Bothalia 37,2 (2007)

Namibia near Rosh Pinah, in the winter rainfall part of

that country (Figure 2). Plants were growing among

dolomite rocks and in crevices on the summit of a high

ridge, where they would have benefited from any addi-

tional moisture that condensed from sea fog. It is likely

that the species occurs elsewhere in suitable places in

this rugged, largely inaccessible, and botanically poorly

explored region. Known only from the type collection,

made by Cape Town succulent specialist Peter Bmyns in

2003, Hesperaniha longistyla is a surprising discovery,

representing the only species of the genus known from

Namibia.

FIGURE 1. — A, Hesperantha longi-

styla, Bruyns 9484A; B, H. hel-

met, Helme 3144. Scale bars:

10 mm. Artist: J.C. Manning.

Diagnosis and relationships', distinctive in its short

stature, falcate leaves, relatively long perianth tube, 20-

25 mm long, and disproportionately long style branches,

Hesperantha longistyla has the bell-shaped corm with a

flat base that characterizes a handful of other species in

the genus from the southern African winter rainfall zone

to the south, including white-flowered H. falcata (L.f )

Ker Gawl. as well as the two purple-flowered species,

H. latifolia (Klatt) M.P.de Vos and H. paucifiora (Baker)

G.J.Lewis. The last two species occur in Namaqualand,

the former restricted to the Kamiesberg and the latter

reaching its northern limit there, well to the south of

Bothalia 37,2 (2007)

179

28°-

30°-

32°-

34°-

FIGURE 2. — Known distribution of Hesperantha longistyla. A; H. helmei. A; H. Uthicola, ■; H. acuta subsp. acuta, O; and H. acuta subsp.

tugH'elliae, •.

Namibia. Nevertheless, the proximity of their ranges,

and their similar flower colour suggest that H. longistyla

is allied to these two species. H. longistyla is most simi-

lar to the long-tubed H. latifolia but differs in its smaller

flowers, with tepals ± 10 mm long, and in the marked

difference in the relative proportions of stamens and

style. The filaments in H. longistyla are relatively short,

3-4 mm long and the style branches project ± 3 mm

beyond the tips of the anthers, thus appearing unusually

long. The flowers of H. latifolia, in contrast, are gener-

ally much larger, with tepals 15-23 mm long, the fila-

ments (4-)7-10 mm long, and the style branches do not

extend much beyond the tips of the anthers.

Hesperantha helmei Goldblatt & J.C. Manning,

sp. nov.

Plantae 180-250 mm altae, cormo globoso asymmet-

rico ± 9 mm diam., caule eramoso in quarta superiore

folium squamiformem gerente, foliis 3, inferioribus

dua teretibus costatis, superiore vaginanti, spica 1- vel

2-flora, bracteis viridibus, ± 9 mm longis, floribus pal-

lide malvinis, tubo perianthii 6-8 mm longo, tepalis sub-

equalibus anguste ovatis ± 8 x 2.5 mm patentibus, fila-

mentis ex tubo ± 2 mm exsertis, antheris ± 4 mm longis,

ramis styli ±3.5 mm longis.

TYPE. — South Africa, Eastern Cape, 3224 (Graaff-

Reinet): ± 50 km E of Graaff-Reinet, upper northern

slopes of Nardouwsberg, 1 km N of main peak. Annex

Stijlfontein 250, 2300' [700 m], (-BB), 19 November

2004, N.A. Helme 3144 (NBG, holo.).

Plants 180-250 mm high; corm globose, ± 9 mm

diam., asymmetric, with concentric tunics. Stem slen-

der, unbranched, bearing short, scale-like leaf in upper

fourth. Leaves 3, lower 2 terete, conspicuously grooved,

blades 8-15 x 1.2 mm, uppermost leaf sheathing lower

half to two thirds of stem; cataphylls membranous. Spike

1- or 2-flowered; bracts green, ± 9 mm long, inner about

as long as outer and forked at apex. Flowers pale mauve.

darker at mouth of tube, outer tepals with broad purple

streak on reverse; perianth tube 6-8 mm long, slender,

widening near apex; tepals subequal, narrowly ovate, ±

8.0 X 2.5 mm, outer slightly larger than inner, spread-

ing slightly above horizontal. Stamens symmetrically

disposed, slightly spreading; filaments exserted ± 2 mm

from tube; anthers ± 4 mm long. Ovary’ ovoid, ± 2 mm

long; style dividing at mouth of perianth tube, branches

± 3.5 mm long, ascending, ultimately reaching to

about middle of anthers. Capsules and seeds unknown.

Flowering time: November and probably December.

Figure IB.

Distribution and ecology’: apparently restricted to

the southern edge of the interior escarpment, where it is

known from a single collection made at high elevations

in rocky grassland in the Sneeuberge, east of Graaff-

Reinet (Figure 2). The plants were found in open areas in

Merxmuellera grassland among dolerite boulders. These

grasslands bum regularly every few years during the dry

season and are regularly covered with snow in winter

(N.A. Helme pers. comm.) The tips of the long, basal

leaves in all the specimens have been burned or frosted

off, indicating that the species begins leafing early in the

growing season when the danger of fire or frost is not

yet over.

Diagnosis and relationships: Hesperantha helmei is

named for its discoverer, the Cape Town ecologist and

energetic and wide-ranging plant collector, N.A. Helme.

The species is distinctive in its relatively small, pale

mauve flower with a perianth tube 6-8 mm long, narrow

tepals ± 8.0 x 2.5 mm, and terete, finely ribbed leaves.

Another unusual attribute of the species is the small,

sheathing, scale-like leaf in the upper part of the stem.

This distinctive feature is characteristic of a small group

of species allied to H. pilosa (L.f ) Ker Gawk, all native

to the winter rainfall zone of southern Africa. Among

these species, H. pilosa is largely a species of the Cape

floristic region, whereas H. ciliolata Goldblatt, H. tereti-

folia Goldblatt and the hairy-leaved H. pseiidopilosa

180

Bothalia 37,2 (2007)

Goldblatt are centred on the Roggeveld Escarpment of

the Western Karoo (Goldblatt 1987). Among the spe-

cies in this group, H. helmei is probably most closely

related to H. ciliolata and H. teretifoUa, which also have

terete, prominently ridged leaves, the grooves lined with

fine ciliate hairs that are absent in H. helmei. The flow-

ers of H. ciliolata and H. teretifoUa are slightly larger

than those of H. helmei, with a perianth tube ± 8 mm

long and tepals ± 10 mm long. Hesperantha teretifoUa

in particular is broadly similar to H. helmei in general

appearance, but has minutely ciliolate leaf ribs, a small,

scale-like leaf, 2-A mm long, and larger anthers, ± 8 mm

long. H. helmei, in contrast, has glabrous leaves, a larger

scale-like leaf, 8-10 mm long, and smaller anthers, ± 4

mm long.

While the presence of an endemic species of Hespe-

rantha in the Sneeuberge is surprising, it is not without

precedent in the Iridaceae. A handful of species of the

family occur there, including taxa from both the winter

rainfall zone to the west and the summer rainfall zone

to the east. Those from the winter rainfall zone include

Moraea ciliata, M. crispa and M imguiculata. The

Iridaceae from the summer rainfall zone include one

Dierama species, Romulea macowanii, three species of

Syringodea, including the near endemic S. pulchella,

and a distinctive short, broad-leaved form of Babiana

bainesii. This mix of summer and winter rainfall species

reflects the transitional position of these mountains, that

have a grassland-dominated flora characteristic of the

summer rainfall region at higher elevations but which

receive predictable amounts of winter rainfall that permit

the persistence of a handful of species from the winter

rainfall zone to the west.

Hesperantha lithicola J.C. Manning & Goldblatt,

sp. nov.

Plantae (40-)80-200(-290) mm altae, cormo campanu-

lato, 7-10 mm diam., ad basem dentato, caule eramoso,

foliis 4, inferioribus 2 suberectis vel falcatis (30-)50-

100(-160) X 1-2 mm, spica 1^-flora, floribus albis,

tepalis exterioribus extra rubris, tubo perianthii 10-12

mm longis, tepals subaequalibus, 10-15 x 4. 0-6. 5 mm,

filamentis 3-5 mm longis, antheris ± 6 mm longis, ramis

styli 4. 0-5. 5 mm longis.

TYPE. — South Africa, Western Cape, 3219 (Wupper-

tal): Swartruggens, Farm Knolfontein, beween Katbak-

kies Pass and Skitterykloof, in loam and leaf litter on

sandstone pavement, (-DC), 12 September 2006, P.

Goldblatt & J.C. Manning 1 28 17 A (NBG, holo.; MO,

PRE, iso.).

Plants (40-)80-200(-290) mm high; corm bell-shaped

with flat base, 7-10 mm diam., breaking into segments

below, margins fringed and often with radiating spines.

Stem unbranched or rarely with one branch from near

base. Leaves 4, lower 2 basal, suberect or falcate, often

undulate or loosely coiled, midrib prominently raised

and margins squared and minutely scabrid along both

edges, blades (30-)50-l00(-160) x 1-2 mm, upper-

most leaf entirely sheathing; cataphylls membranous.

Spike 1 ^-flowered; bracts green, flushed reddish and

becoming dry apically, 11-20 mm long, inner about as

long as outer and forked at apex. Flowers white, outer

tepals red outside, opening in evening and producing

rosy citrus fragrance; perianth tube 10-12 mm long,

not or slightly exserted beyond bracts, slender, widen-

ing near apex; tepals subequal, elliptic, 10-15 x 4.0-6. 5

mm, outer slightly larger than inner, spreading horizon-

tally when fully open. Stamens symmetrically disposed,

slightly spreading; filaments 3-5 mm, exserted 2-3 mm

from tube; anthers horizontal, pale yellow, ± 6 mm long.

Ovaty ovoid, ± 3 mm long; style dividing at mouth of

perianth tube, branches 4. 0-5. 5 mm long. Capsules

ellipsoid, 9-10 x 4 mm. Seeds subglobose with flattened

chalaza and persistent funicle, golden brown, 1.0-1. 5

mm diam., rugose, testa cells domed. Flowering time:

September. Figure 3.

Distribution and ecology, known from sandstone rock

pavement in the Swartruggens Mountains in the eastern

Cold Bokkeveld, where the plants grow in rock crevices

or shallow humus, often in the shelter of small shrubs

(Figure 2).

Diagnosis and relationships'. Hesperantha lithicola

is distinctive in its symmetrical, bell-shaped corm and

narrow, often twisted or coiled leaves. It is probably

most closely allied to H. falcata, which it resembles in

general appearance, but from which it differs in its con-

sistently narrower leaves, 1-2 mm wide (vs. 4-8 mm

in H. falcata), and longer perianth tube, 10-12 mm vs.

4-9 mm. The two species differ also in their habitats.

Hesperantha lithicola is restricted to shallow soils on

sandstone rock pavement, whereas H. falcata is a com-

mon and widespread species of seasonally moist, deeper

clay or sandy soils. This ecological difference is very

evident at the type locality, where H. falcata occurs on

sandy flats not far from rocky outcrops to which H. lithi-

cola is restricted.

Hesperantha lithicola is remarkably similar in gen-

eral appearance to H. acuta (Licht. ex Roem. & Schult.)

Ker Gawl. but this species has an asymmetrical corm

with one side flattened and extended downward. Without

conns, however, the two taxa are essentially indistin-

guishable, although H. acuta does not occur on the Cold

Bokkeveld and is typically found on clay soils, rarely

limestone or in pockets of loam on sandstone outcrops.

Other material examined

WESTERN CAPE. — 3219 (Wuppertal): Katbakkies 139 Farm,

exposed sandstone slope, (-DC), 20 September 1991, C. Reid 1389

(NBG, PRE); Swartruggens, 60 km NE of Ceres, Knolfontein, 1 205 m,

(-DC), 14 September 2005, 1. & C. Jardine 126 (NBG); Groenfontein,

Zeekoegat, west of Riet River, 900 m, (-DC), 15 September 2001, M.S.

Stohie 7 (NBG).

TAXONOMIC NOTES

Hesperantha karooica Goldblatt

Known from just a handful of plants collected at two

locations around Calvinia in Northern Cape, Hesperantha

karooica is recognized among the species of the genus

with large, asymmetric corms with overlapping tunics

by its low stature, 30-50 mm high, and spikes of 1 or

2 yellow flowers with a tube ± 5 mm long and tepals ±

20 mm long (Goldblatt 1984, 2003). In general aspect

Bothalia 37,2 (2007)

181

FIGURE 3. — Hesperantha lithicola. Goldblatt & Manning 1 2817 A. A,

whole plant; B, flower; G, capsule; D, seed. Scale bars: A-C, 10

mm; D, 1 mm. Artist: J.C. Manning.

the species is very like the more widespread Bokkeveld

Plateau species, H. vaginata (Sweet) Goldblatt, which

is normally taller, 120-180 mm high, and has spikes of

(1)2^ yellow flowers, usually marked with contrasting

brown markings in the centre and on the tips of the outer

tepals. Plain yellow-flowered plants are known from a

few sites but both these and plants with marked tepals

have a perianth tube 5-8 mm long and tepals 30-35 x

15-17 mm. Although the two species have been recog-

nized in recent revisions of the genus (Goldblatt 1984,

2003), there has remained some doubt as to whether they

are really distinct.

New collections of Hesperantha vaginata from near

Loeriesfontein and of yellow-flowered plants that accord

with H. karooica from well to the east of Calvinia, con-

tribute to our understanding of the two taxa. The latter

population, found some 50 km NNE of Calvinia on the

road to Klipwerf in the area known as the Agter Hantam

{Goldblatt & Porter 12747), consists of plants with

bright yellow flowers, the outer tepals shaded brown out-

side, and ranging in height from 50 to 100 mm, with one

plant found in the shade of a low shrub attaining a height

of 160 mm. The flowers have a perianth tube 6-7 mm

long, thus longer than until now reported for H. karoo-

ica and matching H. vaginata, but the tepals are ± 20

X 9-10 nun long and the anthers ± 10 mm long, thus ±

as recorded for the two other collections of H. karooica

(Goldblatt 1984).

The collection of Hesperantha vaginata from near

Loeriesfontein {Goldblatt & Porter 12775), from the

Farm Rietfontein, and which comprises a modest range

extension (Goldblatt 1984), consists of plants 50-120

mm high, the smallest of these unusual for the species.

All individuals have flowers with the brown markings

on the inside of the tepals that are typical of the species

and anthers that are ± 12 mm long and tepals 30 x ± 12

mm. Even the smallest plants have flowers typical of the

species.

We conclude that plant height and perianth tube

length are not factors that consistently distinguish the

two species, but that anther length and tepal size always

differ. The tepal size especially is substantially smaller in

Hesperantha karooica. Both this species and H. vaginata

typically grow on heavy red clay derived from dolerite,

although populations of H. vaginata from the west of its

range also occur on tillite-derived clay or a mix of clay

and sand derived from Table Mountain Sandstone.

Range extensions

Hesperantha karooica

NORTHERN CAPE.— 3119 (Calvmia): Agter Hantam, NNE of

Calvinia on road between Moordenaarspoort and Klipwerf, stony dol-

erite clay flats, (-BB), 1 September 2006, Goldblatt & Porter 12747

(MO, NBG, PRE).

Hesperantha vaginata

NORTHERN CAPE. — 3119 (Calvinia): Farm Rietfontein, ± 48 km

SE of Loeriesfontein, low dolerite clay hill, (-BC), 5 September 2006,

Goldblatt & Porter 12775 (MO, NBG, PRE).

Hesperantha acuta (Licht. ex Roem. & Schult.) Ker Gawl.

New and well-preserved collections of the yellow-

flowered variant of Hesperantha acuta (Goldblatt 1984),

until now believed to be rare, have provided us with

additional information about the species. Typical white-

flowered H. acuta is common in the western Karoo,

and blooms proliflcally in years of adequate rainfall.

The flowers, including the anthers, are white, although

the outer tepals are brown to red on the outside. Careful

measurements of the stamens and styles show that the

182

Bothalia 37,2 (2007)

TABLE 1 , — Comparison of yellow- and white-flowered variants of Hes-

perantha acuta. Only well-pressed, fully open flowers were mea-

sured

filaments are exserted 1.5-2. 5 mm from the tube and that

the style branches reach from ± 2 mm below the anther

tips to 1 mm above them. The tube itself is 8.5-12.0 mm

long and the tepals 9.5-16.0 rmn long (Table 1). Plants

with flowers falling in the upper range of these dimen-

sions are most common along the Cederberg but also

occur over the entire range of the white-flowered variant

of the species (Figure 2). Smaller-flowered plants occur

mainly in the northwest of the range, along the foothills

of the Bokkeveld and Matsikamma-Gifberg Mountains.

Yellow-flowered plants, referred by R.C. Foster (1948) to

H. tiigwelliae, occur in the east of the range of H. acuta,

in the northern foothills of the Swartberg Mountains,

along the Kammanassie Mountains, and in the Long

Kloof to the south in the vicinity of Joubertina (Figure

2). New collections made since 2002 (listed below) show

that plants have a perianth tube 12.5-15.0 mm long and

tepals 12.5-16.0 mm long, thus closely matching the

white-flowered plants, but the filaments are exserted ± 4

mm and the anthers are 6-8 mm long, whereas the style

branches reach only to about the middle of the anthers

(Table 1 ).

These consistent differences in the dimensions of

anthers and style provide proof, in addition to flower

colour, that the eastern populations constitute a sepa-

rate, larger-flowered race, and we believe it is useful to

recognize the yellow-flowered populations as a separate

taxon. Because of the small differences between the two,

and complementary geographic ranges, we recommend

subspecies rank for the taxon, and thus make the combi-

nation H. acuta subsp. tiigwelliae.

Hesperantha acuta subsp. tugwelliae (R.C. Foster)

Goldblatt & J.C. Manning, comb, et stat nov. Hesperan-

tha tugwelliae R.C. Foster, Contributions from the Gray

Herbarium 166: 26 (1948). Type; South Africa, [Western

Cape], Prince Albert, Tugwell s.n. (Nat.Bot.Gard. 1 1 5 1/29

in K, holo.!, BOL, MO, iso.!).

Selected specimens

WESTERN CAPE. — 3321 (Ladismith): 30 miles beyond Seven

Weeks Poort along Ladismith-Laingburg road, (-AC), 9 September

1971, Thomas s.n (NBG93821); Swartberg, between Matjesvlei

and Gamkaspoort, (-BC), Oct. 1986, Vlok s.n. (MO); Swartberg

Mtns, road to Gamkaskloof, (-BD), 9 September 2001, Goldblatt c6

Porter 11859 (MO, NBG). 3322 (Oudtshoom): Prince Albert, Farm

Kleinsleutelfontein, (-AB), 1 August 1995, Marincowitz sub Snijman

1 492 (NBG); Kareedouw Pass between Klaarstroom and Prince Albert,

(-AD), 10 September 2002, Goldblatt & Porter 12191 (MO, NBG);

Groot Doom River, (-CD), August 1931, Thorne s.n. (SAMS 17 10).

3323 (Willowmore); eastern slopes of Kammanassie Mtns, Farm

Wildepaardefontein, (-CA), 18 August 1986, Viviers & Vlok s.n. (MO);

Kammanassie Mtns, lower steep slopes above Buffelsdrif, (-DB), 8

August 1983, Matthews 1128 (NBG).

EASTERN CAPE. — 3323 (Willowmore): 8 km west of Joubertina,

(-DD), 24 August 1981, Snijman 450 (NBG); hill north of Joubertina,

(-DD), August 1923, Fourcade 2690 (BOL, K, NBG).

ACKNOWLEDGEMENTS

Support for this work by grants 7316-02 and 7799-

05 from the National Geographic Society is gratefully

acknowledged. We thank Peter Bruyns, Nicholas Helme,

and Ivor and Cora Jardine for drawing our attention to

the novelties described here. We also acknowledge our

debt to Ingrid Nanni and Tendon Porter for their assis-

tance and companionship in the field and to Roy Gereau

for checking our Latin descriptions. Collecting pennits

were provided by the Nature Conservation authorities of

Western Cape Province, South Africa.

REFERENCES

FOSTER, R.C. 1948. Studies in the Iridaceae V. Some new or note-

worthy species of Hesperantha. Contributions from the Gray

Herbarium 166: 3-27.

GOLDBLATT, P. 1984. A revision of Hesperantha (Iridaceae) in the

winter rainfall area of southern Africa. Journal of South African

Botany 50: 15-141.

GOLDBLATT, P. 1987. New species and notes on southern African

Hesperantha (Iridaceae). South African Journal of Botany 53:

459^63.

GOLDBLATT, P. 2003. A synoptic review of the African genus

Hesperantha (Iridaceae: Crocoideae). Annals of the Missouri

Botanical Garden 90: 390—443.

HILLIARD, O.M. & BURTT, B.L. 1986. Hesperantha (Iridaceae)

in Natal and nearby. Notes from the Royal Botanic Garden

Edinburgh 43: 407^38.

Bothalia 37.2: 183-187 (2007)

New species of Drimia (Hyacinthaceae: Urgineoideae) allied to Drimia

marginata from Western and Northern Cape, South Africa

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

Kej'words: Drimia Jacq., Hyacinthaceae, South Africa, taxonomy, Urgineoideae

ABSTRACT

Plants until now identified as Drimia marginata (Thunb.) Jessop on account of their leathery, oblong to elliptical leaves

with thickened, cartilaginous margins and capitate inflorescences of campanulate flowers, are shown to comprise three sets

of populations separable on leaf morphology, ecology and distribution. Typical D. marginata produces 1(2) oblong, apiculate

leaves with retrorsely-scabridulous margins and occurs in fine-grained clay soils on the Hantam and Roggeveld Plateaus.

Plants from Namaqualand and the Richtersveld. described here as Drimia puichromarginata J.C. Manning & Goldblatt.

occur in sandy or gravelly soils and produce 1-A, elliptical to suborbicular, apiculate leaves with an ornate, duplex margin:

the dorsal surface bears a submarginal band of dense, velvety trichomes fringing the thickened, colliculate margin. A third

series of populations from seasonally moist sandstones at higher altitude on the interior mountains of the West Coast produces

2 or 3(4) narrowly oblong, obtuse leaves with a simple, papillate or colliculate margin and are recognized as D. ligulata

J.C.Manning & Goldblatt. A fourth taxon with a similar capitate inflorescence of campanulate flowers produces a solitary,

subterete or subclavate leaf, elliptical in section. Recorded from scattered localities in the Northern and Western Cape', it is

here described as D. vermiformis J.C.Manning & Goldblatt.

The genus Drimia Jacq. comprises ±100 species of

largely deciduous geophytes distinguished from other

members of subfamily Urgineoideae, one of four sub-

families of the Hyacinthaceae, by their apomorphic,

short-lived flowers with the tepals ± united at the base

(Manning et al. 2004). Each flower lasts just several

hours or up to a day, and the perianth is caducous and

circumscissile, abscising at the base and withering as a

cap on the developing capsule. The genus includes sev-

eral smaller groups of obviously related species with

spurred floral bracts, some of which were previously

treated as separate genera, including Rhadamanthiis

Salisb., Schizobasis Baker and Tenicroa Raf (Jessop

1977). Among those taxa traditionally retained in the

poorly defined genus Urginea itself, is a small assem-

blage of species from the winter rainfall region of the

Western and Northern Cape Provinces characterized

by their capitate inflorescence of campanulate flow-

ers with spreading tepals. Florally rather uniform, the

species in the group are distinguished by their foliage:

D. barkerae J.C.Manning & Goldblatt by a rosette of

oblanceolate, ± ciliate leaves with simple margins; D.

marginata (Thunb.) Jessop by two or three, oblong to

elliptical leaves with heavily thickened margins; and

D. minor (A.V.Duthie) Jessop by fililform or subterete

leaves (Jessop 1977; Manning el. al. 2002; Manning &

Goldblatt 2003). Another taxon in this group, with a soli-

tary, subcylindrical or clayate leaf, has been referred to

D. Virens Schltr. (Goldblatt & Manning 2000; Manning

et al. 2002).

It is now clear that three distinct species are included

within the concept of Drimia marginata as currently cir-

cumscribed (Jessop 1977; Goldblatt & Manning 2000).

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

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

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

MS. received: 2007-05-07.

These three entities are separable by differences in the

number, shape, apex, and especially the marginal orna-

mentation of the leaves, as well as ecology and distribu-

tion. Typical D. marginata is characterized by oblong,

apiculate leaves with thickened, minutely retrorsely sca-

bridulous margins, and occurs in fine-grained clay soils

on the Hantam and Roggeveld Plateaus. A second series

of populations from seasonal sandstone rock flushes at

moderate altitudes in the mountains of the southwest-

ern Cape, described here as D. ligulata J.C.Manning &

Goldblatt, produces narrowly oblong, truncate leaves

with a papillate margin. A third group of populations,

described here as D. puichromarginata J.C.Manning &

Goldblatt, occurs further north, in Namaqualand and the

Richtersveld, in sandy or gravelly soils, and is charac-

terized by its broader, elliptical to suborbicular leaves

with a unique duplex margin comprising an outer rim of

papillae, edged on the inner side by a dense, narrow, vel-

vety band of minute trichomes.

Furthemiore, examination of the syntypes of Drimia

virens. Schlechter 10127 and Leipoldt s.n., reveals that

the species falls within the current circumscription of D.

minor (Duthie) Jessop, a species characterized by several,

erect or spreading, linear or filiform leaves (Manning

et al. 2002). In his review of the genus Drimia, Jessop

(1977) treated Urginea virens as a synonym ofD. modesta

(Baker) Jessop, a species from the summer rainfall region

with an elongate inflorescence but it is clearly misplaced

here. Drimia virens is a combination based on Urginea

virens Schltr. (1897) and as such is an earlier name for

this species and we formally synonymize D. minor

below.

The plants with a single, fleshy, subclavate leaf that

were previously referred to D. virens by us (Goldblatt

& Manning 2000; Manning et al. 2002) comprise a dis-

tinct species, which we describe here as D. vermiformis

J.C.Manning & Goldblatt.

184

Bothalia 37,2 (2007)

Drimia vermiformis J.C. Manning & Goldblatt,

sp. nov.

D. virens sensu Goldblatt & Manning (2000), non D. virens

(Schltr.) J.C. Manning & Goldblatt.

Species Drimiae minori similis sed folio uno, 50-80

X 2—^ mm, suberecto vel expanso, camoso subclavato,

floribus campanulatis, pallide bmnneis tepalis carinis

atrobninneis, tepalis biseriatis per ± 1 mm connatis.

TYPE. — Western Cape: 3218 (Clanwilliam), Clan-

william Dam, picnic site along N7 near wall, (-BB), 3

August 1987 (fl. in cult. 11 October 1988), P.L. Perry 3587

(NBG, holo.).

Deciduous, bulbous herb. Bulb solitary, subglobose,

15-20 mm diam.; outer tunics pale brown, thinly leath-

ery; inner tunics tightly overlapping, white or flushed

pink. Leaf dry and withered at flowering, 1(2), spreading

or suberect, leathery, dark green; blade falcate or subcla-

vate, subterete or ellipsoid in section, 50-80 x 2-4 mm,

glabrous or minutely hispidulous. Inflorescence nodding

in bud; scape erect or flexuose at base, (20-)50-120 mm

long, glabrous; raceme corymbose-capitate, 2-5 mm

long, densely 5-20-flowered; bracts elliptical, ± 2 mm

long, lower with spur 1-2 mm long; pedicels spread-

ing, 5-10 mm long at anthesis. Flowers campanulate,

1 or 2 open at a time, pale brownish with darker keels,

opening in the late afternoon and fading in the evening,

apparently unscented; tepals biseriate with blades of

outer series overlapping inner, penicillate at apex, fused

for ± 1 mm, erect below forming a cup ±1.5 mm deep,

spreading above; blades of outer tepals ovate, ± 5.0 x

2.0 mm, inner oblong, ± 4.5 x 1.8 mm, weakly canalicu-

late at top of cup. Stamens adnate to perianth for ± 0.5

mm; filaments erect, subterete and tapering, ±2.5 mm

long; anthers erect, dorsifixed, dehiscing by longitudi-

nal slits, ± 1 mm long, yellow with yellow pollen. Ovary

ellipsoid, ± 2 mm long, truncate, greenish yellow; style

columnar, ± 1.5 mm long, white, apically truncate with

trigonous, papillate stigma. Capsules subglobose, 5-6

X ± 5 mm, erect on suberect, slightly curved pedicels.

Seeds compressed, elliptical or rectangular, 2-3 mm

diam., glossy black, irregularly folded, testa finely retic-

ulate. Flowering time: October and November. Figure 1.

Distribution and ecology, widely distributed through the

arid parts of the southwestern Cape, from Bushmanland

in Northern Cape southwards onto the Roggeveld and

Nuweberg Escarpments and into Western Cape, extend-

ing westwards through the arid During River Basin into

the lower Olifants River Valley around Clanwilliam, and

into the Little Karoo as far east as Oudtshoom (Figure

2). The species is certainly more common than records

indicate. Plants occur on exposed, mostly shale flats and

lower slopes in fine-grained clay or loam. Populations in

the Little Karoo around Calitzdorp have been recorded

from quartz patches.

Additional specimens examined

NORTHERN CAPE.— 2919 (Pofadder): Pofadder, (-AB), 14

October 1954, E. Esterhiiy.?en 23640a (BOL). 3120 (Williston):

Roggeveld Escarpment, Middelpos, hill behind school, (-CC), 12

September 2001 (in bud), D. Snijman & Van cier Westhuysen 1853

(NBG).

FIGURE 1 . — Drimia vermiformis. Manning 3092. A, whole plant; B,

flower. C, D, stamen: C, front view; D, side view. E, gynoecium;

F, capsule; G, seed; H, c/s leaf Scale bar: A, F, H, 10 mm; B, G,

2 mm; C-E, 1 mm. Artist: John Manning.

WESTERN CAPE. — 3118 (Vanrhynsdoip): Trawal, steep slopes

west ofN7, (-DC), 31 August 2003 (in bud), P.V. Bruyns 9523 (NBG);

Trawal, right hand slope below N7 leading to Olifants River, (-DC), 15

April 2007, J. Manning 3092 (NBG). 3218 (Clanwilliam): Zwart Vley,

near upper waterfall east of Uitspankraal on Doring River, (-AB), 16

September 1992 (in cult.), D. Snijman 1223 (NBG). 3222 (Beaufort

West); Stolshoek, [Karoo National Park, west of Beaufort West],

(-AD), 13 September 1989, P. Bruyns 3977 (BOL). 3320 (Montagu);

near Montagu, (-CC), September 1933, M.R. Levyns 4600 (BOL); 13.5

km east of Bonnievale, (-CC), without date, P.L. Peny s.n. (NBG).

3321 (Ladismith): Farm Droogkraal, 30 km NW of Oudtshoom, (-BD),

1 October 1980, A. Bean s.n. (NBG); road to Rooiberg Pass, SW of

Radleigh, (-DA), E.G.H. Oliver 3665 (NBG).

Drimia virens (Schltr.) J.C. Manning & Goldblatt in

Strelitzia 9: 712 (2000), Urginea virens Schltr.: 433 (1897).

Type: [Western Cape], Cold Bokkeveld, Tweefontein, 24

January 1897, Schlechter 10127 (BOL, lecto.!, here desig-

nated; E!, GRA!, BM, K!, L, P, PRE!, S!, Z, isolecto.).

D. minor (Duthie) Jessop: 306 (1977). Urginea minor Duthie: 11

(1928), syn nov. Type: [Western Cape], Stellenbosch Flats, March

1924, Duthie s.n. STE1546 (NBG, holo.!).

Bothalia 37,2 (2007)

185

FIGURE 2. — Known distribution of Drimia vermiformis, ■; D. ligit-

lata, •; D. marginala, O; D. pidchromarginata. A.

THE DRIMIA M4RG1NATA COMPLEX

Drimia marginata (Thimb.) Jessop in Journal of

South African Botany 43: 295 (1977). Anthericum mar-

ginatum Thunb.; 63 (1794). Idothea marginata (Thunb.)

Kunth.: 346 (1843). Urginea marginata (Thunb.) Baker;

218 (1873). Type: [Northern Cape], Hantam, without

exact date, [November 1774], Thunberg s.n. {UPS-8393,

holo.-microfiche!).

Deciduous, bulbous herb. Bulb solitary, subglobose,

15-30 mm diam.; outer tunics pale brown, thinly leath-

ery; inner tunics tightly overlapping, white. Leaf dry and

withered at flowering, 1(2), spreading or prostrate, leath-

ery, dark green, base amplexicaul; blade oblong to oblan-

ceolate, apiculate, 30-50(-60) x 9-15 mm, glabrous or

minutely hispidulous, margin adaxially thickened, carti-

laginous and densely and minutely retrorsely scabridu-

lous, ± 0.5 mm wide. Inflorescence erect or flexuose at

base; scape (100-) 150-200 mm long, glabrous; raceme

capitate, 2-10 mm long, densely 10-20-flowered; bracts

elliptical, 2-3 mm long, lower spurred with spur 1-2

mm long; pedicels spreading, 5-10 mm long at anthesis.

Flowers campanulate, 1 or 2 open at a time, pale brown-

ish with darker keels; tepals biseriate with blades of

outer series overlapping inner, penicillate at apex, fused

for ± 1 mm, erect below and forming a cup ± 1.5 mm

deep, spreading above, blades of outer tepals ovate, ±

5.0 X 2.0 mm, inner oblong, ± 4.5 x 1.8 mm. Stamens

adnate to perianth for ± ,0.5 mm; filaments erect, subte-

rete and tapering, ± 2.5 mm long; anthers erect, dorsi-

fixed, introrse, dehiscing by longitudinal slits, ± 1 mm

long, yellow with yellow pollen. Ovary ovoid, ± 2 mm

long, truncate, greenish yellow; style columnar, ± 1.5

mm long, white, apically truncate with trigonous, papil-

late stigma. Capsules and seeds unknown. Flowering

time: October and November.

Distribution and ecology, apparently restricted to the

Hantam Plateau in Northern Cape, between Loeriesfontein

and Calvinia (Figure 2), on open clay flats in renosterveld

vegetation.

Discussion', the first species in the complex to be

described, D. marginata is still relatively poorly col-

lected. It is distinguished by its oblong to oblanceolate,

distinctly apiculate leaves with a thickened margin that

is densely ornamented with minute, stiff, retrorse tri-

chomes 0.2-0. 2 mm long (Figure 3C, D). The blade may

be glabrous or minutely hispidulous and typically just a

single leaf is produced (rarely two).

The species appears to be restricted to fine-grained

clay soils derived from shale or dolerite. No infonna-

tion is recorded on the time of anthesis, nor on floral fra-

grance.

Additional specimens examined

NORTHERN CAPE. — 3019 (Loeriesfontein): 15 km from Loeries-

fontein on road to Kliprand, (-CD), 13 September 2000 (in leaf), P.

Goldblatt. J. Manning & V. Savolainen II 525 (NBG). 3119 (Calvinia):

Farm uitvlug NW of Calvinia, (-BC), 6 July 2000 (fl. in cult, 5 October

2000), J. Manning 2270A (NBG); Hantam Mtn, Akkerdam, (-BD), 22

July 1961 (fl. in cult. 14 November 1961), W.F. Barker 9343 (NBG).

3120 (Williston): northern foot of Klein Tafelberg, (-CA), 22 October

1991, E L Brtiyns 4293 (BOL).

Drimia pulchromarginata J.C. Manning & Gold-

blatt, sp. nov.

Species Drimiae marginatae similis sed decidua vel

sempervirens, tunicis bulbi interdum laxe dispositis,

foliis 2-4, latioribus ellipticis ad suborbicularibus usitate

1 5-25 mm latis, marginibus incrassatis parte dorsali per-

omatis supra colliculosis infra dense scabridulis, parte

ventrali simplicibus colliculosis.

TYPE. — ^Northern Cape: 3018 (Kamiesberg), Farm

Draaiklip, (-AA), 31 October 1983 (fl. in cult.), C.H.

Stirton 9226 (NBG, holo.!).

Evergreen or deciduous, bulbous herb. Bulb solitary,

subglobose, 20-30 mm diam.; outer tunics pale brown,

thinly leathery; inner tunics sometimes loosely arranged,

white. Leaves green or drying at flowering, (1)2^, pros-

trate or erect, leathery, dark green, sometimes purple

beneath, base flat; blade elliptical to broadly elliptical

or suborbicular, acute, 25-60 x (1 3-) 15-25 mm, gla-

brous or minutely hispidulous adaxially, margin thick-

ened, cartilaginous, 0. 5-1.0 mm thick, colliculate, with

adaxial submarginal band ± 0.5 mm wide of dense,

suberect or weakly retrorse trichomes ± 0.1 mm long.

Inflorescence erect or flexuose at base; scape (60-)150-

300 mm long, glabrous; raceme capitate, 2-5(-20) mm

long, densely 10-30-flowered. Flowers as in D. mar-

ginata, usually opening in late afternoon and fragrant.

Capsules and seeds unknown. Flowering time'. October

and November.

Distribution and ecology, endemic to the west-

ern parts of Northern Cape, from the Richtersveld and

higher-lying parts of northern and central Namaqualand

as far south as Garies, rarely near the coast (Figure 2).

Plants typically occur on sandy or gravelly flats among

granite outcrops, sometimes on quartz patches. Anthesis

has been variously recorded as in the afternoon (Hall

4172) or morning (Harrower 1521) and the floral fra-

grance described as either unpleasant (Stirton 9226) or

freesia-like (Harrower 1521).

186

Bothalia 37,2 (2007)

V

FIGURE 3. — Leaf shape and leaf

margin in Drimia marginata

complex. A, B, Drimia pid-

chromarginata, Stirton 9226:

C, D, D. marginata, Goldblatt,

Manning & Savolainen 11525;

E, F, D. ligulata, Esterhuysen

14487. Scale bar: A, C, E, 10

mm; B, D, F, 0.5 mm. Artist:

John Manning.

Discussion-, the northernmost member of the Drimia

marginata complex, D. pulchromarginata is distin-

guished by its elliptical to suborbicular, apiculate leaves,

mostly 15-25 mm wide, with a highly ornamented,

duplex margin comprising a narrow, colliculate rim

edged internally with a broader band, ± 0.5 mm wide,

of closely packed, suberect or weakly retrorse trichomes

± 0.1 mm long (Figure 3 A, B). This uniquely complex

margin is developed only on the adaxial surface, and the

margin on the ventral surface is simple and colliculate,

as found in D. ligulata. The leaves of D. pulchromargin-

ata are also characteristically broader and more elliptical

than in the other species in the complex. Typically 2-A

leaves are produced, and the plants remain evergreen if

conditions permit.

The development of dense trichomes along the leaf

margins in Drimia marginata and D. pulchromarginata

is evidently a derived character and suggests that these

two taxa are allopatric sister species.

Additional specimens examined

NORTHERN CAPE.— 2817 (Vioolsdrif): Richtersveld, Chubiesies,

(-AB), 15 October 2006 (fl. in cult.), A. Narrower 1521 (NBG). 2917

(Springbok): Springbok, (-DB), January 1977, B. Jeppe s.n. PRE57508

(PRE). 2918 (Gamoep): Vaalkoei, 1 000 m, (-CD), 10 July 1991 [fl.

4 November 1991], P.V. Bruyn.'s 4713 (PRE). 3017 (Hondeklipbaai):

Riethuis, (-AB), 11 July 1989 [as 1898] [fl. 21 October 1991], P.

Bruyn.s 3879 (BOL); 6 km NNE Riethuis on road to Springbok, (-AB),

without date, / Manning 1040 (NBG); Kamieskroon, (-BB), 9 Novem-

ber 1950, W.F. Barker .i.n. NBG 1181/50 (NBG); Kharkams, (-BD),

18 October 1971, H. Hall 4172 (NBG); Darter’s Grave, 22 miles N

of Garies, (-BD), August 1932, J. Mathews s.n. NBG1891/31 (BOL);

4 May 1963 (fl. in cult. 18 November 1963), L. Booysen 13 (NBG).

Without precise locality or date: [Northern Cape], Namaqualand, 1924,

Giffen s.n. NBG1051/24 (BOL).

Drimia ligulata J.C. Manning & Goldblatt, sp. nov.

Species Drimiae marginatae similis sed foliis 2-3(^),

angustioribus oblongis usitate 6-10, raro ad 15 mm latis,

expansis vel prostratis obtusis, marginibus simplicibus

papillosis, papillis truncatis vel acutis, racemo corym-

boso-capitato, florescentia Decembri Januarique.

TYPE. — Western Cape: 3218 (Clanwilliam), Piket-

berg Mtns, Zebra Kop, (-DB), 23 May 1948 (fl. Dec-Jan

1948 and 1949), E. Esterhuysen 14487 (BOL, holo.).

Deciduous, bulbous herb. Bulb solitary, subglobose,

15-30 mm diam.; outer tunics pale brown, thinly leath-

ery; inner tunics tightly overlapping, white. Leaves dry

and withered or emergent at flowering, 2 or 3(4), pros-

trate or spreading, leathery, dark green, base amplexi-

caul; blade oblong, obtuse, (20-)30-90 x (4-)6-10(-15)

mm, glabrous, margin thickened, cartilaginous, 0.5 mm

thick, papillate or colliculate. Inflorescence erect or flex-

uose at base; scape (60-)80-200 mm long, glabrous;

raceme corymbose-capitate, 2-20 mm long, densely

5-20-flowered. Flowers as in D. marginata, usually fra-

grant. Capsules spreading or suberect on pedicels 5-18

mm long, ovoid to subglobose, 6-8 x 5-7 mm. Seeds

elliptical to reniform, peripherally winged, 3^(-6) x

1.8-2. 5 mm long, glossy black, irregularly folded, testa

finely reticulate. Flowering time-. December and January,

rarely as early as October at lower altitudes.

Distribution and ecology-, recorded from most of

the western mountain chains of the Cape Fold Belt of

Western Cape, from the northern Cedarberg southwards

through the Cold Bokkeveld Mountains and the Skurwe-

Bothalia 37,2 (2007)

187

berg, and also on the higher parts of the Piketberg to the

west (Figure 2). Plants grow in seasonally moist rock

flushes or shallow rock basins on sandstone.

Discussion: the most commonly collected of the three

species of the Drimia marginata complex, D. ligidata is

distinguished by its generally narrower, oblong leaves,

usually 6-10(rarely up to 15) mm wide, with an obtuse

apex and a simple, papillate margin (Figure 3E, F).

Typically two or three leaves are produced, rarely up

to four. The leaf margin is thickened on both dorsal and

ventral surfaces and the papillae may be blunt or acute.

Other members of the complex have broader, oblong or

elliptical leaves that are distinctly apiculate and have

margins that are partially or entirely ciliolate.

Drimia ligulata is geographically and ecologically

distinct from other members of the complex, being con-

fined to seasonally moist sandstone substrates at mod-

erately high altitudes, between 500-1 500 m, in the

western mountains of the Cape Fold Belt. At these alti-

tudes the species flowers later than D. marginata and D.

piilchromarginata, typically in December and January,

rather than October and November.

Additional specimens examined

WESTERN CAPE. — 32 1 8 (Clanwilliam): Piketberg, (-DC ), Decem-

ber 1950. E. Esterhuysen 14487 (BOL); Piketberg. S entrance to

Kapteinskloof, (-DC), 22 October 1935, N. Pillans 8092 (BOL).

3219 (Wuppertal): Cedarberg, between Pakhuis and Heuning Vlei,

(-AA), 28 December 1941, E. Esterhuysen 7426 (BOL); Cedarberg,

Tafelberg. (-AC), 29 December 1947, E. Esterhuysen 14337 (BOL);

Cedarberg. Wolfberg, (-AC), 3 October 1952 (fl. December 1952), E.

Esterhuysen 20587 (BOL); Elands Kloof, (-CA), 29 September 1944

(leafing), fV.F. Barker 3072 (NBG). 9 September 1946 (leafing), W.F.

Barker 3822 (NBG), 24 September 1956 [fl. 18 November 1955], T.R

Stokoe SAM68464 (SAM); W slopes of Cold Bokkeveld Mtns near

Keerom, (-CC), 4 December 1950, E. Esterhuysen 17922 (BOL). 3319

(Worcester): Kliphuis Vlakte on the Skurweberge, (-AA), 10 September

1989 (leaflng), D. Snijman 1236 (NBG); Schurweberg, between

Bokkeveld Tafelberg and Bokkeveld Sneeuwberg, ± 5000' [1 500 m],

(-AA). 11 October 1952 (fl. December 1962), E. Esterhuysen 20662

(BOL); Hansiesberg, (-AB), 16 December 1944, R.H. Compton 16689

(NBG); top of Gydo Pass, (-AB), 7 December 1940, E. Esterhuysen

3952 (BOL); Mosterthoek Twins, 400' [1 370 m], (-AD), 8 January

1944, E. Wasserfall 810 (NBG), E. Esterhuysen 9892 (BOL).

REFERENCES

BAKER, J.G. 1873. Revision of the genera and species of Scilleae and

Chlorogaleae. Botanical Journal of the Linnean Society. Botany

13:209-292.

DUTHIE, A.V. 1928. Contribution to our knowledge of the Stellenbosch

flora. The species of Urginea of the Stellenbosch Flats. Annals of

the Stellenbosch University 6, A, 2: 1-14.

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

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

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

JESSOP, J.P. 1977. Studies in the bulbous Liliaceae in South Africa: 7.

The taxonomy of Drimia and certain allied genera. Journal of

South African Botany 43: 265-3 1 9.

KUNTH, C.S. 1843. Enumeratio plantanim 4. Tubingen. Stuttgart.

MANNING, J.C. & GOLDBLATT, P. 2003. A new species and new

combinations in Drimia (Urgineoideae). Bothalia 33: 109-1 1 1 .

MANNING, J.C., GOLDBLATT, P. & FAY, M.F. 2004. A revised

generic synopsis of Hyacinthaceae in sub-Saharan Africa, based

on molecular evidence, including new combinations and the

new tribe Pseudoprospereae. Edinburgh Journal of Botany 60:

533-568.

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

encyclopedia of Cape bulbs. Timber Press, Oregon.

SCHLECHTER, R. 1 897. Decades plantanim novamm austro-afficanum.

Journal of Botany. London 35: 428^33.

THUNBERG, C.P. 1794-1800. Prodromus plantanim capensium. Edman.

Uppsala.

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Notes on African plants

VARIOUS AUTHORS

IRIDACEAE

ARISTEA NIGRESCENS (SUBGENUS PSEUDARISTEA), A NEW SPECIES FROM WESTERN CAPE, SOUTH AFRICA WITH A NOVEL

STRATEGY FOR POLLINATOR ATTRACTION

INTRODUCTION

Aristea Aiton comprises ± 50 species of evergreen,

rhizomatous perennials distributed through sub-Saharan

Africa as far north as Ethiopia and Senegal, with seven

species in Madagascar (Manning et al. 2002). The

genus is especially diverse in the Cape Floral Region,

where some 33 species are recorded. Recent advances

in our understanding of morphology and relationships

among the species have led to the recognition of three

subgenera based on differences in fruit, seed and pollen

(Goldblatt & Le Thomas 1997; Goldblatt et al. 2004). In

contrast, the flowers of most species are very similar in

form and colour, displaying little of the remarkable vari-

ability that is associated with other genera of African

Iridaceae. With few exceptions, flowers of Aristea are

rotate, uniformly deep blue, and fugacious, lasting a sin-

gle morning. Significant exceptions are many species of

subgenus Pseudaristea Pax, which is characterized by a

relatively high degree of floral diversification. Subgenus

Pseudaristea is endemic to the southwestern Cape, and

is well defined by several vegetative and floral charac-

ters. The inflorescences are sessile, with large rhipidial

spathes concealing the flower buds, and the ovaries are

elongate and subcylindrical, mamring into elongate, 3-

lobed capsules that can measure more than 80 mm long.

These are thick-walled and slow to dry, remaining closed

for up to a year in some species, only releasing the seeds

gradually over an extended period. The distinctive seeds

are triangular-columnar in shape with obliquely trun-

cate ends, and are shortly fringed or papillate along the

angles. Florally, the subgenus is characterized by promi-

nent, deeply fringed stigma lobes, and dizonosulculate

pollen that is unique for the entire Iridaceae.

Unlike other species of Aristea, which are pollinated

by pollen-collecting bees, most species of subgenus

Pseudaristea have developed highly specialized pollina-

tion systems, relying primarily on hopbines, or monkey

beetles (Scarabaeidae: Hopliini) (Goldblatt & Manning

1997). This dnusual pollination system, which is well

developed in lowland habitats in the southwestern Cape,

is often associated with the evolution of highly localized

endemic plant species. Beetle-pollinated members of

subgenus Pseudaristea are no exception, and the ranges

of most species extend little more than 50 km and may

be substantially smaller than that. The local nature of the

species and their small population size puts them at high

risk of extinction. In addition, the pyrophilic nature of

many of the species, which flower only in the spring fol-

lowing a bum, means that they are rarely seen. This has

led to several species being overlooked for decades or

even centuries despite their proximity to well-populated

areas (Goldblatt & Manning 1997). Currently, nine spe-

cies are recognized in subgenus Pseudaristea (Manning

et al 2002; Goldblatt et al. 2005). Three of these have

been described during the past decade (Goldblatt &

Manning 1997; Goldblatt et al. 2005), all of them from

well-botanized areas near large towns. The new species

Aristea nigrescens, described here, is another example,

and increases the number of species in the subgenus to

ten. The species is extremely vulnerable to extinction

through transformation of the habitat to agriculture.

Aristea nigrescens J.C. Manning & Goldblatt, sp.

nov.

Plantae (100-)200-300(-500) mm altae vivaceis caes-

pitosis, folds linearibus 3-5 mm latis, caule angulato sim-

plex vel raro uniramoso, inflorescentia terminal! 4— 6-flora,

spathis late lanceolatis infra viridibus supra siccis brun-

neisque (10-)12-15 mm longis, floribus albis vel cae-

ruleis, tepalis extemis partim nigris nitidisque, tepalis

patentibus oblique obovato-spathulatis subaequalibus

vel interioribus leviter latioribus (20-)25-35 x 13-14

mm, filamentis 5-7 mm longis, antheris 6-7 mm longis,

ovario cylindrico ± 10 mm longo, stylo 12-15 mm longo

ecentrico, capsulis pedicellis ad 12 mm longis, cylindri-

cis 40-50 mm longis.

TYPE. — Western Cape, 3319 (Worcester); Wolseley,

Farm Romansrivier, southwestern slopes of hill above

dam behind farmhouse, alluvium, (-AC), 2 September

2006, J. Manning 3054 (NBG, holo.; K, MO, iso.).

Plants (100-)200-300(-500) mm high, evergreen,

tussock-forming. Stem erect, angled to slightly winged,

bearing two or three short, reduced leaves, simple or

rarely with one axillary rhipidium. Leaves in a basal fan,

linear, one third to ± half as long as stem, loosely twisted,

3-5 mm wide. Inflorescence: rhipidia, terminal or lateral,

if present sessile, in axil of uppermost stem leaf, 4-6-

flowered; rhipidial spathes paired, broadly lanceolate,

(10-)12-15 mm long, acute, green at base but dry and

papery brown in distal two thirds with paler margins,

sometimes irregularly tom; floral bracts 5-7 mm long,

entirely dry and papery. Flowers shortly pedicellate,

pedicels ± 1.5 mm long, actinomorphic with style eccen-

tric, upright, white or pale to mid-blue but paler in cen-

tre, outer tepals partially glossy blackish on reverse, last-

ing one day, opening mid-moming ± 10:00 and fading in

afternoon ± 17:00; tepals spreading, obliquely obovate-

spathulate, cucullate, connate at base for ± 1 mm, sub-

equal or inner slightly broader, (20-)25-35 x 13-14 mm.

190

Bothalia 37,2 (2007)

FIGURE \.—Arislea nigrescens, Manning 3054. A, flowering stems and base of plant; B, fruiting stem. Scale bar: A, B, 10 mm. Artist: John

Manning.

Bothalia 37,2 (2007)

191

Stamen filaments suberect, straight, 5-7 mm long, white

to pale blue; anthers basifixed, 6-7 mm long before

anthesis, yellow; pollen yellow, grains dizonasulculate,

exine reticulate. Ovary cylindric, ± 10 mm long, elon-

gating rapidly after fertilization; style 12-15 mm long,

white or flushed blue in distal half, dividing into three

short, broad, fringed lobes 1.5-2. 5 mm long. Capsules

on pedicels up to 12 mm long, cylindric, 40-50 mm

long, 3-lobed in transverse section. Seeds numerous per

locule. Flowering time: late August to mid-September.

Figure 1.

Distribution and ecology: a highly local endemic

known from two sites along the foot of the Waaihoek

Mountains overlooking the Breede River near Wolseley

in Western Cape (Figure 2). Plants flower only in the

season following a summer bum. At the type locality,

Romansrivier Farm, the species is still abundant, occur-

ring in large numbers in a narrow ecological zone on

ferricrete, or sandstone alluvium overlying Malmesbury

shale. Most plants at this locality have pale blue flow-

ers but flower colour through the population varies

from white to mid-blue. The vegetation is transitional

fynbos-renosterveld. A second population located on a

neighbouring farm has been reduced to just a handful

of plants through cultivation of the slopes for vineyards.

Both plants seen at this locality had white flowers but the

small sample makes it impossible to ascertain if this was

typical of the entire population. It is likely that the spe-

cies once occurred in a band all along the foothills of the

Waaihoek Mountains but much of the natural habitat has

now disappeared under cultivation or sylviculture and

the species must therefore be considered to be highly

endangered.

The species is adapted to pollination by hopbine bee-

tles (Scarabidae: Hopliini), and we observed and cap-

tured individuals of two species, Anisonyx ditus and A.

ursus as frequent visitors to the flowers, which open in

the mid-moming at around 10:00 and fade in the early

afternoon around 15:00. The beetles crawl over the flow-

ers, become dusted with pollen in the process and sub-

sequently transfer it to the stigma. In the post-fire envi-

ronment, Aristea nigrescens flowers with several other

beetle-pollinated species, including Drosera cistiflora

(Droseraceae), Moraea versicolor, M. villosa and Ixia

viridiflora (all Iridaceae). These species show the char-

acteristics of flowers adapted to this pollination system,

notably a brightly coloured, salver-shaped perianth with

dark central markings that act as pollinator attractants.

Aristea nigrescens is anomalous in lacking dark mark-

ings in the centre of the flower, either on the tepals or on

the stamens, ^and only the underside of the outer tepals

is flushed blackish. The dark underside of the tepals is

highly visible in the buds surrounding the open flower

and in older, withered flowers, however, and the form of

the buds especially is highly reminiscent of the beetles

themselves. The buds in A. nigrescens are well exposed

above the spathes for several days prior to anthesis and

thus quite visible among any open flowers. In other spe-

cies in the subgenus the buds are concealed among the

spathes and floral bracts until the morning of anthesis.

In A. nigrescens, therefore, it appears that the function

of pollinator attraction has been transferred from the

mature flower to the buds and withered flowers, which

FIGURE 2. — Known distribution of Aristea nigrescens.

thus act as accessory pollinator attractants. The use of

accessory structures for pollinator attraction has been

identified in several southern African bird-pollinated

taxa: floral buds and immature flowers function as

accessory floral attractants in bird-pollinated species of

Melianthus (Melianthaceae) (Linder at al. 2006), and in

Iridaceae the inflorescence spathes in Klattia or the floral

bracts in certain species of Gladiolus such as G. abbre-

viatus (Goldblatt et al. 1999) play a similar role. This is

the first record, however, of secondary pollinator attrac-

tion among insect-pollinated members of the Iridaeeae.

A distinct indole-dominated fragrance was detected at

anthesis in a single flower, fading within an hour or so,

but investigation of several other stems failed to detect

any scent and this remains an anomalous observation.

Diagnosis and relationships'. Aristea nigrescens is

characterized by unbranched stems bearing a solitary,

terminal flower cluster, or very rarely with a single sec-

ond cluster in the axis of the upper stem leaf; broadly

lanceolate rhipidial spathes, (10-)12-15 mm long,

that are papery brown in the distal two thirds; and by

its large, white to blue flowers without dark markings.

The underside of the outer tepals is flushed blackish and

the buds thus appear dark. In its unbranched stems and

broad, rather blunt rhipidial spathes, the species resem-

bles A. lugens (L.f.) Steud. and it is likely that the two

are allied. Aristea lugens is distinguished by its strongly

dimorphic tepals, with the outer tepals ± half as long

as the inner and dark brown or black over most of the

surface and strongly incurved, exposing both upper and

lower surfaces. Thus A. lugens, like A. nigrescens, uti-

lizes the underside of the tepals for pollinator attraction

but, unlike the latter, only in the mature flower and not

in bud. Other species in subgenus Pseudaristea typi-

cally have branched stems bearing several lateral flower

clusters and smaller, acute or acuminate rhipidial spathes

that are ± entirely leathery and green.

Other material examined

WESTERN CAPE. — 3319 (Worcester): Wolseley, Waaihoek Road,

Silver Leaf Mountain Vineyards, (-CB), 12 September 2006, J. Manning

& P. Goldblatt 3055 (MO, NBG).

192

Bothalia 37,2 (2007)

ACKNOWLEDGEMENTS

Thanks to Rupert Koopman for alerting us to the spe-

cies. Material was gathered with a permit from Western

Cape Nature Conservation on a trip sponsored by

Elizabeth Parker. Mr Lategaan was kind enough to allow

us to visit his farm.

REFERENCES

GOLDBLATT, R, DOLD, A.P. & MANNING, J.C, 2005. Three cryp-

tic new species of Aristea (Iridaceae) from southern Africa.

Bothalia 35: 1-6.

GOLDBLATT, P. & LE THOMAS, A. 1997. Palynology, phylogenetic

reconstruction and the classification of the Afro-Madagascan

genus Aristea (Iridaceae). Annals of the Missouri Botanical

Garden 84: 263-284.

GOLDBLATT, P, LE THOMAS, A. & SUAREZ-CERVERA, M. 2004.

Phytogeny of the Afro-Madagascan Aristea (Iridaceae) revisited

in the light of new data on pollen morphology. BotanicalJonrnal

of the Linnean Society 144: 41-68.

GOLDBLATT, P. & MANNING, J.C. 1997. New species of Aristea

(Iridaceae) from South Africa and notes on the taxonomy and

pollination biology of Section Pseiidaristea. Novon 1: 137-144.

GOLDBLATT, P, MANNING, J.C. & BERNHARDT, P 1999.

Evidence of bird pollination in Iridaceae of southern Africa.

Adansonia,s€'c.3,2\:25-Af>.

LINDER, H.P, DLAMINI, T., HENING, J. & VERBOOM, G.A.

2006. The evolutionary history of Melianthus (Melianthaceae).

American Journal of Botany 97 : 1 052-1 064.

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

encyclopedia of Cape bulbs. Timber Press, Oregon.

J.C. MANNING* and P GOLDBLATT**

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

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

Garden, PO. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2007-01-09.

IRIDACEAE

NIVENIA ARGENTEA MISUNDERSTOOD, AND THE NEW SPECIES NIVENIA INAEQUALIS (NIVENIOIDEAE)

INTRODUCTION

The genera Nivenia Vent. (10 spp.), Klattia Baker

(3 spp.) and Witsenia Thunb. (I sp.) are the only truly

shrubby members of the family Iridaceae, and comprise

the monophyletic subfamily Nivenioideae (Goldblatt

et al. in press). All are endemic to the southwestern

Cape, South Africa, and were recently monographed

by Goldblatt (1993). Since then a single additional spe-

cies, Nivenia parviflora Goldblatt, has been described

(Goldblatt 1 997). Species of Nivenia are largely restricted

to montane habitats at middle elevations and most are

narrow local endemics, occurring on a single mountain

range or at most on adjacent ranges. A striking excep-

tion is N. argentea Goldblatt, which was until now

considered to occur on both the coastal Langeberg and

Riviersonderend Mtns as well as inland on the Rooiberg

range in the Little Karoo (Goldblatt 1993). Our current

knowledge of this species, however, was based almost

entirely on the populations from the Rooiberg, which are

readily accessible by road.

In May 2005 an unidentified Nivenia was photo-

graphed on the Riviersonderend Mountains by plant

enthusiast David Gwynne-Evans. The photographs

showed a species with the congested, pseudoracemose

inflorescence typical of N. argentea but the dark (rather

than pale) blue flowers appeared to lack the short, sub-

orbicular style anns that were regarded as characteristic

of that species, as well as the conspicuous silvery bracts

that gave the species its name (Goldblatt 1993). In addi-

tion, the inflorescence appeared to be well exserted from

the leaf cluster, unlike the Rooiberg populations in which

it is typically only shortly extended beyond the leaves.

The form of the inflorescence in the Riviersonderend

plants is, however, a close match for the Langeberg popu-

lations of N. argentea, as well as the type of the species,

which was collected on the Riviersonderend Mtns.

Further examination of the Langeberg material at our

disposal raised the strong possibility that it was not in

fact conspecific with the Rooiberg collections after all,

although most specimens available lacked well-preserved

flowers. Efforts were therefore made to obtain good

flowering material of the Langeberg or Riversonderend

plants and in December 2006 members of the Friends of

the Outeniquas, under the guidance of Di Turner, suc-

ceeded in locating ample flowering plants. The following

year Martin Grantham secured material from both the

Langeberg and Riviersonderend Mountains. These col-

lections confirm that the Langeberg and Riviersonderend

plants are a close match for the type of Nivenia argentea

and are distinct from the Rooiberg populations.

This confirms that Nivenia argentea is typical of the

other members of the genus in having a limited distri-

bution and that the Rooiberg populations comprise an

undescribed species that we name N. inaequalis, for the

markedly unequal stamens that characterize the species.

We also provide a full description of true N. argentea,

flowers and fruits of which have not been available for

detailed examination until now.

Nivenia now comprises 1 1 species, all of rocky sand-

stone habitats in the southern African winter rainfall

zone. Of these, seven species are distylous and four are

homostylous.

Nivenia argentea Goldblatt, The woody Iridaceae:

54 (1993), as a substitute name for Nivenia capitata (Klatt)

Weim.: 366 (1940), nom illeg. non N. capitata R.Br.

(Proteaceae). Witsenia capitata Klatt: 546 (1866). Type:

South Africa, [Western Cape], banks of the Riviersonderend,

Bothalia 37,2 (2007)

193

Appelskraal, Eksteen, and surrounding mountains,

500^000' [150-1 200 m], Ecklon & Zeyher Ir id. 62 (B,

holo.; MO!, SI, iso,).

Rounded shrubs up to 400 mm high, branching from

base, forming dense cushions. Stems ascending, com-

pressed, leafy portions 2-A mm diam. Leaves crowded

apically, linear-lanceolate, (20-)40-80 x 1. 5-3.0 mm.

Inflorescence a congested, branched pseudoraceme, up

to 15 -flowered, lateral branches short, 3 -flowered, each

node bearing a short-stalked, 1 -flowered rhipidium;

peduncle axis compressed, extending (10-)20-40 mm

from leaf cluster, ultimate branches ± 2 mm long; inflor-

escence bracts lanceolate-attenuate, keeled, outer bracts

15-25 mm long, dry and brown with brown membranous

margins, inner bracts about two thirds as long as outer,

with narrow transparent membranous margin; spathes

tightly enclosing flower, lanceolate-attenuate, 13-18 mm

long, dry and papery, brown below and in midline, mar-

gins and apices transparent-membranous; floral bracts

tubular, 18-25 mm long, dry and membranous, usually

silvery transparent in upper half or third, brown in lower

half to two thirds, sometimes ± entirely shining brown,

becoming tom apically. Flowers salver-shaped, hetero-

distylous, deep blue, darker in centre and white in throat

and tube; perianth tube 14-20(-25) mm long, widening

gradually towards mouth; tepals oblong-oblanceolate,

13- 18 X 4-6 mm, spreading. Stamen filaments inserted

in mouth of tube, either 4-5 mm long (thrum flower) or

1.0-1. 5 mm long (pin flower); anthers 1.0-1. 5 mm long

before anthesis, yellow; pollen yellow. Ovary’ ± 1.5 mm

long; style either included 0.5^.0 mm below mouth of

tube (thrum flower) or exserted ± 7 mm and reaching ±

5 mm beyond anthers, dividing into filiform branches ±

1.5 mm long. Capsules obovoid, 6-7 x 3-4 mm, typi-

cally with just a single fertile locule containing a solitary

seed. Seeds one per locule, ±4.5 x 1.8 mm, scutiform,

papillate-rugulose. Flowering time: mainly October

to December, occasionally out of season in March and

April. Figure 3.

Distribution and ecology: a montane species occur-

ring at middle to upper elevations, 500-1 000 m, along

the Riviersonderend and adjacent Langeberg Mountains

(Figure 4), Nivenia argentea has been recorded from

the eastern Riviersonderend Mtns above the town

of Riviersonderend and from several sites along the

southern length of the Langeberg, from Tradouwberg

and Grootvadersbos north of Heidelberg in the west to

Garcia’s Pass and Paardeberg near Riversdale in the east.

Plants grow on exposed, rocky ridges, forming densely

leafy, rounded shrubs up to 400 mm high. Populations

are usually small.

Diagnosis^and relationships: Nivenia argentea is dis-

tinguished by its congested, pseudoracemose inflores-

cence with lanceolate-attenuate subtending bracts that

are entirely brown and dry. The ultimate branches bear

single-flowered rhipidia, and each flower is surrounded

by a tubular bract 1 8-25 mm long that is dry and mem-

branous, usually silvery transparent in the upper half and

brown in the lower half but sometimes entirely shining

brown. The species is heterodistylous, as are most spe-

cies of Nivenia, and the flowers have a perianth tube

14- 25 mm long with the anthers exserted on filaments

either 4—5 mm long (thrum flowers) or 1.0-1. 5 mm long

(pin flowers). The style branches are slender and typical

of most other species in the genus, and differ from those

of N. inaequalis, which are short and rounded.

In the structure of the compound inflorescence

Nivenia argentea closely resembles N. inaequalis from

the Rooiberg but in this species the inflorescence bracts

and spathes have broad, silvery margins and the floral

bracts are entirely silver-transparent. The rhipidia in

N. inaequalis, unlike those of N. argentea, are 2-flow-

ered and the flowers themselves are distinctive in their

longer tubes, 30^0 mm long vs 14-20(-25) mm long

in N. argentea', in their unequal stamens in which one

filament is notably shorter than the other two; and in

the unique, short, rounded style branches. The tubu-

lar floral bract of N. argentea, in which the lower mar-

gins are fused to form a closed cylinder (Figure 3F),

have not been recorded in the genus before. The bract

margins in Nivenia species are usually free to the base,

although overlapping and enclosing the perianth tube.

Closer examination of the bracts of the remaining spe-

cies reveals that N. fruticosa (L.f ) Baker, also from the

Langeberg Mountains, is the only other species with

tubular floral bracts, possibly indicating a close relation-

ship between it and N. argentea.

History: Nivenia argentea has been misunderstood

almost since its first description by F.W. Klatt in

1866, and its early history has been well documented

by Goldblatt (1993). First described under the name

Witsenia capitata, the species was based on imma-

ture specimens collected in bud by C.F. Ecklon & C.L.

Zeyher in the Riviersonderend Mountains. Although

subsequently included in a second species, N. fruti-

cosa, by both J.G. Baker (1877, 1896) and N.E. Brown

(1933), the species was later resuscitated by Weimarck

(1940), unfortunately under the illegitimate combina-

tion Nivenia capitata (Klatt) Weim., a homonym for

N. capitata R.Br. [now Paranomus capitatus (R.Br.)

O. Kuntze, Proteaceae]. By this time, further specimens

from Garcia’s Pass in the Langeberg had been collected

by E.E. Galpin in 1897, enabling Weimarck to describe

the flowers more fully. Weimarck’s illustration (1940:

fig. 2A) clearly shows the extremely short filaments

(of the pin morph), and his interpretation of the inflo-

rescence as comprising several solitary flowers led him

to place the species in his section Singulares, erected

for those taxa with single-flowered rhipidia. The spe-

cies remained poorly collected, however, and it is thus

not surprising that when a Nivenia species with a simi-

lar congested, pseudoracemose inflorescence and silvery

floral bracts was collected in 1957 from the Rooiberg in

the Little Karoo, inland of the Langeberg, it was referred

to the same species (Goldblatt 1993). The Rooiberg

locality is easily reached by vehicle and plants from

there thus served as the basis for the description and

fine illustrations that appeared under the new name N.

argentea Goldblatt, coined to replace the illegitimate N.

capitata (Goldblatt 1993). In the absence of any further

collections of the species from west of the Langeberg,

Goldblatt followed Weimarck in concluding that the

locality of the type collection on the Riviersonderend

Mountains was most probably incorrect. More recent

collections of the species from the Langeberg, largely

lacking flowers, did nothing to change this interpreta-

tion but the photographic evidence of plants from the

194

Bothalia 37,2 (2007)

FIGURE 3. — Nivenia argentea, Custodians of Rare and Endangered WUdflowers s.n. A. two flowering branches; B, outer inflorescence bract; C,

inner inflorescence bract; D, single rhipidium; E, spathes; F, floral bract; G, floral bract flattened; H, pin flower flattened; I, thrum flower

flattened; J, style branches. Scale bars, A-I, 10 inm; J, 1 mm. Artist: John Manning.

Bothalia 37,2 (2007)

195

FIGURE 4. — Known distribution of Nivenia argentea, •; N. inaequa-

lis, O.

Riviersonderend Mountains suggests that the type local-

ity of the species is indeed this mountain range.

The name Nivenia argentea is thus correctly

applied only to the populations of the species from the

Langeberg and Riviersonderend Mountains, and the

Rooiberg populations represent an unnamed species that

we describe here as N. inaequalis.

Other material seen

WESTERN CAPE. — 3320 (Montagu): Swellendam. Tradouwberg,

(-DD), without date, J. Bowie s.n. (K); Langeberg East MCA, Com-

partment 8, S slopes of Eloringberg. 3800' [1 160 m], (-DD), December

1984, T.J. van der Merwe 31 (NBG, WIGHT). 3321 (Ladismith):

Garcia’s Pass, (-CC), October 1926, C. Thorne SAM38856 (NBG,

SAM); Garcia State forest Reserve, approaching Aasvoelkrans, 1 151

m. (-CC), 18 December 1988, D.J. McDonald 1797 (NBG); Garcia

Forest Reserve, Rooiwaterspruit overlooking Oudebosch. 600 m. (-CC),

21 November 1991, D.J. McDonald 2107 (NBG); near Rooiwaterspruit

huts, (-CC), 9 January 2007, M. Grantham s.n. (NBG); mountains at

Garcia’s Pass, (-CC), 2 October 1897, Galpin 4664 (PRE), October

1931, M.A. Pocock s.n. (BOL); southern slope next to path from

Rooiwaterspruit huts heading east, about 1 km from huts, 517 m, (-CC),

13 December 2006, Custodians of Rare and Endangered Wildfiowers

s.n. (MO, NBG); Sandkraal, Rooielsberg, (-CD), 5 December 1982,

Viviers 292 (PRE); northern Langeberg, northeast of Langkloof, 2 km

west of Waterval Peak and 12 km east of Garcia’s Pass, 1 100 m, (-CD),

3 July 2000 (fr.), N.A. Helme 1730 (NBG); eastern Langeberg, summit

ridge 1,5 km west of Doodkisberg, 900 m, (-DC), 29 April 2006, N.A.

Helme 3922 (NBG); summit ridge of Perdeberg [Paardeberg], northwest

of Bergfontein Farm. 1 120 m, (-DC), 27 April 2001, N.A. Helme 1972

(NBG). 3420 (Bredasdorp): eastern Riviersonderend Mms, 34°05’41.7"S

19 °55’47.5"E, (-BB), 17 January 2007, M. Grantham s.n. (NBG).

Without precise jpcality: Cape of Good Hope, J. Niven s.n. (BM), W.

Roxburgh s.n. (BM); ‘very high in the Swellendam Mountain’, Mackrill

s.n. (BM).

Nivenia inaequalis Goldblatt & J.C. Manning, sp.

nov.

Nivenia argentea sensu Goldblatt (1993), in part.

Plantae 400-800 mm altae, sempervirentes, caulibus

pluribus e caudice lignoso compressis ellipticis, foliis

distichis anguste lanceolatis (40-)55-90 x 2-3 mm,

inflorescentia composita ex rhipidiis binatis bifloris in

pseudoracemum congestum dispositis constanter, spathis

13-15 mm longis, bracteis siccis papyraceis albotrans-

lucentibus, floribus caeruleis sessilis heterodistylis, tubo

perianthii 30^0 mm longo, tepalis patentibus lanceo-

lato-ellipticis ± 18 x 5-6 mm, filamentis 7-8 mm longis

(plantis brevistylis) vel 3 mm longis (plantis longisty-

lis), antheris 1.0-1. 5 mm longis flavis, ovario ± 1.5 mm

longo, stylo in tubo incluso (plantis brevistylis) vel 6-8

mm exserto (plantis longistylis), lobulis styli 0.5 mm

longis.

TYPE. — Western Cape, 3321(Ladismith): Rooiberg

Mtns, next to forestry track near Bailey’s Peak, 3300^600’

[1 000-1 400 m], (-CB), 29 December 1986, J.H.J. Vlok

1794 (NBG, holo., K, MO, PRE, iso.).

Rounded evergreen shrubs, 400-800 mm high, branch-

ing from base. Stems ascending, with short spur-shoots in

upper part, leafy portions ± 3 mm diam. Leaves crowded

apically, narrowly lanceolate, (40-)55-90 x 2-3 mm.

Inflorescence a congested, branched pseudoraceme, up

to 20-flowered. lateral branches short, each node bearing

a short-stalked, 2-flowered binate rhipidium; peduncle

axis compressed, exserted (10-) 15-30 mm from leaves,

ultimate branches 3-4 mm long; inflorescence bracts lan-

ceolate-attenuate, keeled, outer bracts ± 17 mm long, dry

and white-translucent with brown midline, inner bracts

white-translucent membranous and somewhat shorter;

spathes tightly enclosing flower, lanceolate-attenuate,

13-15 mm long, dry and membranous, brown below

and in midline, margins white-transparent; floral bracts

enveloping tube, open to base, 22-25 mm long, dry and

white-translucent throughout, becoming tom apically.

Flowers salver-shaped, heterodistylous, blue, darker at

tepal bases and white in throat and tube; perianth tube

30^0 mm long, widening gradually towards mouth;

tepals elliptic-lanceolate, ± 18 x 5-6 mm, spreading.

Stamen filaments inserted in mouth of tube, unequal with

one shorter than others, either two 7-8 mm long and

one ± 5 mm long (thmm flower) or two 3 mm long and

one 1.5-2. 5 mm long (pin flower); anthers 1.0-1. 5 mm

long before anthesis, yellow; pollen yellow. Ovary ± 1.5

mm long; style either included 2-3 mm below mouth of

tube (thmm flower) or exserted 6-8 mm and reaching

2-A mm beyond anthers, dividing into rounded lobes 0.5

mm long. Fruit a subglobose capsule, ± 6 x 4.0^. 5 mm.

Seeds one per locule, shield-shaped, mgose, ± 4.5 x 3.5

mm. Flowering time: mainly November and December,

occasionally out of season in Febmary and March.

Distribution and ecology'. Nivenia inaequalis is restric-

ted to the upper reaches of the Rooiberg (Figure 4), an

isolated massif southeast of Ladismith, where it occurs

above 1 000 m on rocky ridges and summits in arid fyn-

bos, usually in exposed sites in rock outcrops.

Diagnosis and relationships: Nivenia inaequalis is

instantly recognized in the genus by its unequal stamens,

with one filament shorter than the other two, and by

the short, rounded, rather than linear or ± filiform style

branches. The species is heterodistylous and in the pin

morph the shorter stamen is included in the elongate

perianth tube, which is 30^0 mm long. The floral bracts

are very conspicuous, being silver-translucent through-

out their length.

Nivenia inaequalis is most likely to be confused with

N. argentea from the Langeberg and Riviersonderend

196

Bothalia 37,2 (2007)

Mountains. Both species have a similar general aspect,

forming rounded, leafy, cushion-like shrubs, and similar

dry, lanceolate-attenuate bracts and a distinctive, com-

pact, pseudoracemose inflorescence. This inflorescence

type, however, is probably ancestral, being matched

among the woody Iridaceae in Witsenia and, moreover,

most like the inflorescence of Aristea, which is sister to

the Nivenioideae-Crocoideae clade (Reeves et al. 2001).

The 2-flowered, binate rhipidia of N. inaequalis repre-

sents the ancestral condition, whereas the single-flow-

ered rhipidia of N. argentea are clearly derived. Nivenia

argentea also differs from N. inaequalis in its slightly

shorter perianth tube, 14-25 mm long, and specialized

tubular bracts that are partially or completely brown.

The equal stamens and linear style branches of N. argen-

tea are unspecialized in the genus.

History, although first collected by T.M. Wurts in

1957, Nivenia inaequalis was invariably confused with

N. argentea Goldblatt, and the fine illustrations published

under the name N. argentea in The woody Iridaceae

(Goldblatt 1993) actually represent this species.

Other material seen

WESTERN CAPE. — 3321 (Ladismith): crest of Rooiberg west

of Rooiberg Pass, 3600' [1 100 m], (-CB), 13 November 1957, T.M.

Wurts 1618 (NBG); Rooiberg crest road, 1 159 m. (-CB), 10 May

1983, A.H. Marshall 30 (NBG); Ararat Ridge, 4100' [1 250 m], (-CB),

18 December 1977, H.C. Taylor 9773 (NBG); south-facing slopes

approaching Mount Ararat, 900 m, (-CB), 15 December 1989, D.J.

McDonald 1903 (NBG); upper north slopes of Rooiberg, forestry

track, (-CB), 12 Eebruary 1997, P. Goldblatt 10614 (MO); ridge

west of Teeboskop above Assegaaiboskloof, 3000' [915 m], (-DA), 9

November 1974, E.G.H. Oliver 5351 (NBG, PRE); Teeboskop area,

3000' [915 m], (-DA), 2 November 1987, P. Goldblatt & J.C. Manning

8558 (MO, NBG).

ACKNOWLEDGEMENTS

We thank David Gwynne-Evans for alerting us to the

existence of plants later found to be Nivenia argentea in

the Riviersonderend Mountains, Jan Vlok for assisting in

locating living plants from the Langeberg, plant enthusi-

asts Di Turner and the Friends of the Outeniquas for col-

lecting plants and forwarding them to us for study, and

Martin Grantham for visiting the populations to secure

capsules and seeds.

REFERENCES

BAKER, J.G. 1877 [as 1878]. Systema iridearum. Journal of the

Linnean Society, Botany 16: 61-180.

BAKER, J.G. 1896. Irideae. In W.T. Thiselton-Dyer, Flora capensis 6:

6-171. Reeve, London.

BROWN, N.E. 1933. Nivenia Vent, and Nivenia R.Br. Transactions of

the Royal Society of South Africa 2 1 : 250-270.

GOLDBLATT, P. 1993. The woody Iridaceae: systematics, biology and

evolution o/Nivenia, Klattia Witsenia. Timber Press, Portland,

Oregon.

GOLDBLATT, P. 1997. A new species of Nivenia (Iridaceae). Bothalia

27: 101-103.

GOLDBLATT, R, RODRIGUEZ, A., POWELL, M.P, DAVIES, T.J.,

MANNING, J.C., VAN DER BANK, M. & SAVOLAfNEN, V.

in press. Iridaceae ‘Out of Australasia’? Phylogeny, biogeog-

raphy, and divergence time based on plastid DNA sequences.

Systematic Botany.

KLATT, F.W. 1866. Revisio Iridearum (Conclusio). Linnaea 34: 537-

689.

REEVES, G., CHASE, M.W., RUDALL, P.J., FAY, M.F., COX, A.V,

LEJEUNE, B. & SOUZA-CHIES, T. 2001. Molecular system-

atics of Iridaceae: evidence from four plastid DNA regions.

American Journal of Botany 88: 2074-2087.

WEIMARCK, H. 1940. A revision of the genus Nivenia Vent. Svensk

Botanisk Tidskrift 34: 356-372.

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

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont. Cape Town.

E-mail: manning(§sanbi.org.

** B. A. Kmkoff Curator of African Botany, Missouri Botanical Garden,

P.0, Box 299, St. Louis, Missouri 63166, USA.

E-mail: peter.goldblatt(§mobot.org.

MS. received: 2007-01-09.

BORAGINACEAE

LOBOSTEMON LASIOPHYLLUS: DISCOVERY OF A LINK SPECIMEN IN STOCKHOLM, THE CORRECT AUTHOR CITATION AND

SYNONYMY

Relief (2003), following Levyns (1934), cited Lobos-

temon lasiophyllus DC. as a synonym of L. fruticosus

(L.) H.Buek. Link (1821) described Echium lasiophyllum

as new and De Candolle (1846) accepted the species and

transferred it to Lohostemon, whence the correct author

citation must be L. lasiophyllus (Link) DC. The view that

L. lasiophyllus is a synonym of L. fruticosus can be ques-

tioned when the characters mentioned in the protologue

such as calyx lobes as long as corolla, corolla white and

style glabrous are contemplated. These do not charac-

terize L. fruticosus, but rather L. trichotomus (Thunb.)

DC. However, the leaf width given (6 lines wide, i.e. ±

12 mm) excludes L. trichotomus again, which typically

possesses leaves (1 .0-)l .5-3.5(-5.0) mm wide; L. fruti-

cosus possesses leaves (4-)5-10(-12) mm wide. In addi-

tion, reference to ‘Folia ... basi attenuata’ reminds us of

the oblanceolate or obovate leaves found in L. fruticosus

and not the narrowly oblong to lanceolate leaves of L.

trichotomus.

According to Stafleu & Cowan ('1981), Link’s types

were all housed in Berlin (B), but no specimens per-

taining to Echium lasiophyllum exist there — these were

destroyed in World War II. Link material also exists in a

number of other places, but all attempts to find relevant

specimens have failed for the herbaria listed in Stafleu &

Cowan (1981).

Link presumably considered Echium longifolium

Defile (1813) a barrier to his publishing ‘£. longifolium

(ex hort.)’ and supplied a new name, E. lasiophyllum to

avoid homonymy. Wright’s (1904) and the International

Plant Name Index (2004) reference of the authorship of

E. longifolium as ‘hort. ex DC.’ is considered incorrect

because De Candolle (1846: 10) merely cites ‘£’. longi-

Bothalia 37,2 (2007)

197

folium (ex hort.)’ as a synonym of L. lasiophylliis . No

material labelled 'E. longifoUum hort.’ has been found

either.

However, in the Swedish Museum of Natural History

(S) there is a specimen associated with Link and which

can shed light on the identity of E. lasiophyllum. It is a

single flowering specimen with an old, small handwrit-

ten label in two different hands. In fainter ink there is:

‘Echium lasio... Link Enumerat’. Only ‘lasio’ in the epi-

thet is clearly written, the rest is a wavy line indicating a

longer name {lasiophyllum). Another hand has added in

darker ink: ‘ded Otto Berol 1823’ (Figure 5).

We believe the first writer is J.H.F. Link himself His

way of writing his own name is characteristic, espe-

cially with the enlarged ‘K’ in ‘Link’. The second writer

is J.G.C. Lehmann. Apparently C.F. Otto provided Leh-

mann in Hamburg with this specimen from Berlin. The

reference to 1823 is probably the date of acquisition of

the specimen by Lehmann, and not the date of collection

(‘dedit’ = given by, furnished by). Christoph Friedrich

Otto (1783-1856) was long active at the Berlin Botanic

Garden and Museum (B), appointed first as gardener and

inspector by Willdenow in 1 805 and he became a close

collaborator with Link up to the year 1843. According to

Hiepko (1987), Link’s herbarium, purchased by B after

his death in 1851, contained many type specimens from

the botanical garden. Although there is no evidence that

the specimen in S was made from the cultivated plant on

which Link based the new name, it gives an indication

of the taxonomic identity because of the label in Link’s

hand. A perusal of collections in S suggests that this is a

unique case of a specimen associated with Link having

found its way there via Hamburg.

Echium lasiophyllum Link in Enumeratio planta-

rum horti regii botanici berolinensis altera 1: 170 (1821).

Lobostemon lasiophyllus (Link) DC.: 10 (1846); C.H.

Wright: 42 (1904). Neotype (here designated): ex hort.,

Ottos.n. (8-06-47281).

This specimen agrees with L. fruticosus as currently

understood on account of its oblanceolate to obovate

leaves possessing dimorphic hairs as well as infundibu-

lar flowers with hairs on the abaxial corolla surface. The

discrepancy in the protologue mentioned above cannot

be accounted for.

Details of the accepted name are:

Lobostemon fruticosus (L.) H.Buek in Linnaea

11: 134 (1837). Echium fruticosum L.: 139 (1753).

Lectotype (Buys & Van der Walt in Taxon 45: 515.

1996): [icon] "Echium Africanum frutescens foliis pilo-

sis’ in Commelin, Horti Med. Amstelod. 2: 107, t. 54

(1701).

ACKNOWLEDGEMENTS

We thank Johannes Lundberg (S) for providing digi-

tal images of the specimen in question and the curators

of BR, C, FI, H, LIV, P, PH and W for assisting us in

searching for original material. We are grateful to John

McNeill for useful comments on the manuscript.

FIGURE 5. — Neotype in S of Echium lasiophyllum Link, annotated by

J.H.F. Link and J.G.C. Lehmann.

REFERENCES

BUEK. H.W. 1837. Echia capensia. Linnaea 11: 129-149.

BUYS, M.H. & VAN DER WALT, J.J.A. 1996. Lectotypicfication of

the names of Linnaean Echium species now in Lobostemon

(Boraginaceae). Taxon 45: 515-517.

COMMELfN, J. 1701. Horti medici amstelodamensis 2: 107, t. 54.

Amsterdam.

DE CANDOLLE, A.R 1846. Prodromus systematis natiiralis regni

vegetabilis. 10. Masson, Paris.

DELILE, A.R. IS\3. Flore d'Egypte: 192,1. 16, f 3. Panckouke, Paris.

HIEPKO, P. 1987. The collections of the Botanical Museum Berlin-

Dahlem (B) and their history. Englera 7: 219-252.

INTERNATIONAL PLANT NAMES ENDEX (IPNI). 2004. http://

www.ipni.org [accessed 1 June 2006].

LEVYNS, M.R. 1 934. A revision of Lobostemon. Journal of the Linnean

Society, Botany 49: 393^51.

LINK, H.F. 1821. Enumeratio plantarum horti regii botanici berolinen-

sis altera 1: 170. Reimer, Berolini.

LINNAEUS, C. 1753. Species plantarum. Stockholm.

RELIEF, E. 2003. Boraginaceae. In G. Germishuizen & N.L. Meyer,

Plants of southern Africa: an annotated checklist. Strelitzia 14:

313-320.

STAFLEU, F.A. & COWAN, R.S. 1981. Taxonomic literature: Lh-0,

vol. 3. Bohn, Scheltema & Holkema, Utrecht & The Hague.

WRIGHT, C.H. 1904. Boragineae. In W.T. Thiselton-Dyer, Flora cap-

ensis 4,2: 2^5. Reeve, London.

M.H. BUYS*, B. NORDENSTAM** and R. VOGT***

* Compton Herbarium, South African National Biodiversity Institute,

Kirstenbosch, Private Bag X7, 7735 Claremont, and Department

of Botany & Zoology, University of Stellenbosch, Private Bag XI,

7602 Matieland, South Africa, email: buys@sanbi.org (author for

correspondence).

** Department of Phanerogamic Botany, Swedish Museum of Natural

History, P.O. Box 50007, SE-104 05 Stockholm, Sweden.

*** Botanischer Garten und Botanisches Museum Berlin-Dahlem,

Freie Universitat Berlin, Konigin-Luise-Strasse 6-8, D-14195 Berlin,

Germany.

MS. received: 2006-06-23.

198

Bothalia 37,2 (2007)

PEDALIACEAE

DEWINTERIA, A NEW SEMISUCCULENT, CLIFF-DWELLING GENUS ENDEMIC TO THE KAOKOVELD, NAMIBIA

While studying cremnophilous plants in the northern

Namib Desert (Kaokoveld), Namibia, specimens of the

hitherto poorly known Rogeria petrophila De Winter, a

small, herbaceous and semisucculent chasmo-cremno-

phyte, were collected in July 2002 and January 2005.

This enabled us to record, for the first time, the repro-

ductive behaviour of the species in its natural cliff-face

habitat, and to collect ample sterile and reproductive

material for a comparative morphological study. A re-

evaluation of the taxonomic position of the species was

considered necessary in the light of its unique dimorphic

fruiting capsules, texture and morphology of the seeds

and specialized filiform basal branches by which it dif-

fers markedly from the other three species of Rogeria.

We concluded that R. petrophila warrants separate

generic status and it is here formally described as a new

monotypic genus.

Dewinteria Van Jaarsv. & A.E.van Wyk, genus

novum, Rogeriae J.Gay ex Defile affine, sed differt char-

acteribus sequentibus: herba biennis vel perennis, sae-

pius caulibus procumbentibus 100-200 mm longis. Folia

cordata, margine grosse dentato. Capsula 2.0-2. 5 mm

longa, lateraliter compressa chartacea. Semina 2. 0-2. 2

mm longa, anguste oblonga ad clavata, leviter laterali-

ter compressa, pagina minuta reticulata. Ad basim rami

filiformes 0.25 mm diametri, ad 200 mm longi, ferentes

folia integera ovata 2-A x 0.6-1. 7 mm, et flores axillares

cleistogamos, 2 mm longos et capsulas complanatas ova-

tas ad ovato-cordatas 5-8 x 4. 0-5. 5 mm; semina lineari-

obovoidea 2. 5-3.0 mm longa.

Type species: D. petrophila (De Winter) Van Jaarsv.

&. A.E.van Wyk.

Rogeria J.Gay ex Defile subgen. Microrogeria Ihlenf.:

73 (1967).

Description partly based on De Winter (1961).

Soft, somewhat trailing, branched, biennial or peren-

nial herb, up to 200 mm long; most parts covered with

mucilage glands; base of stem slightly swollen, semisuc-

culent, somewhat ovate, up to 5 mm diam., often com-

pressed due to narrow crevices. Roots fibrous. Main

branches 3-4 mm diam. at base. Specialized branchlets

annual, usually dying back after fruiting, filiform, 0.25

mm diam., basally produced, trailing, negatively photo-

tropic. Leaves on main branches opposite (intemodes 8-

14 mm long), broadly cordate to kidney-shaped, up to 40

X 55 mm, grey-green; margin coarsely dentate; petiole

20-60 mm long; mostly with a paired or single extraflo-

ral nectary in the axil. Leaves on specialized branchlets

(arise as accessory shoots below the extrafloral nectary

and flower) small, 2-4 x 0.6-1. 7 mm, entire, ovate; peti-

ole 3-4 mm long, sometimes becoming slightly longer

but then leaves becoming broader and coarsely toothed.

Flowers on main branch in axils of leaves, mostly

single, rarely in pairs, conspicuous, trumpet-shaped,

up to 30-70 mm long; pedicel 1.2^.0 mm long. Calyx

slightly zygomorphic, persistent, 5-partite; lobes oblong-

triangular, up to 3 mm long. Corolla slightly swollen at

base, somewhat 2-lipped, sparsely covered with muci-

lage glands; cream-coloured (pale yellow in bud stage)

and maroon-purple in throat and tube; lobes 5, broadly

ovate, emarginate, lower pair slightly larger than upper

3. Stamens 4, arising from base of corolla tube, with

short staminode between the pairs; filaments filiform,

slightly flattened, up to 12 mm long, pilose; anthers basi-

fixed. Ovary elongate-conical, 2-chambered, placenta-

tion axile, with 3-5-seriate ovules; style up to 23 mm

long; stigma capitate, up to 1.5 mm diam. Cleistogamous

flowers on specialized branchlets, 2 mm long, fight yel-

lowish green; pedicels up to 1.5 mm long. Capsules on

main branches lanceolate in side view, 18-25 mm long,

tapering into a curved apex, laterally flattened, dehisc-

ing loculicidally; valves 2, chartaceous. Specialized cap-

sules flattened, ovate to ovate-cordate, 5-8 x 4. 0-5. 5

mm, brown; both carpels dehiscing loculicidally, false

septa nearly completely reduced to small seams at base

of capsule. Seeds of main branch capsules linear-oblong

to club-shaped, slightly flattened, 2. 0-2. 2 mm long,

minutely foveate, brownish. Specialized seeds oblong-

obovoid, slightly flatened, 2. 5-3.0 mm long, minutely

fringed. Figures 6-8.

The new genus is named in honour of Dr Bernard

de Winter, retired Director of the former Botanical

Research Institute in Pretoria (now part of the South

African National Biodiversity Institute), who collected

and described Rogerha petrophila and who first men-

tioned the possibility of fruit dimorphism in this species

(De Winter 1961).

The genus includes a single species:

Dewinteria petrophila (De Winter) Van Jaai'sv. &

A.F.van Wyk, comb. nov.

Rogeria petrophila De Winter in Kirkia 1: 106-108 (1961). Type:

South West Africa [Namibia] 1712 (Orokatuwo): Kaokoveld, 30 miles

S of Kunene River on road to Orupembe, (-DA), 10-05-1957, De

Winter & Leistner 5790 (PRE, holo.; K, M, SRGH, iso.).

Description as for genus.

DISCUSSION >

Distribution and ecology. Dewinteria petrvphila is

a semisucculent, biennial or perennial chasmo-crem-

nophyte, at present only known from crevices and As-

sures on granite cliffs of the Otjihipa Mountains in the

western Kaokoveld of northern Namibia (Figure 9). It

grows on all aspects but is more abundant on south-fac-

ing ones and at altitudes of about 600-1 700 m on the

northwestern peaks of these mountains (eastern margin

of the Marienfluss). The upper slopes receive fog and

are cooler, with Cape floristic elements such as species

of Friocephalus, Othonna and Pelargonium. The aver-

age annual rainfall in the Kaokoveld varies from less

than 50 mm along the coast to 350 mm in the highlands

(Mendelsohn et al. 2002). This species is a constitu-

Bothalia 37.2 (2007)

199

FIGURE 6. — Dewinteria petrophila:

A, plant growing in crevice,

X 0.7; B, plant in flower, with

stem bearing nonnal flowers

and fhiiting capsule as well as

thread-like branchlets. x 0.7;

C, normal fhiiting capsule, x

0.7; D, specialized seed, x 5;

E, normal seed, x 5. Artist: Lisa

Strachan.

ent of arid Colophospermum mopane woodland, with

several species of Commiphora prominent. It shares its

habitat with other cremnophytes such as Aeollanthns

haumannii, Kalanchoe lanceolata, Plectranthus dinteri

and Tetradenia kaokoensis. The range of D. petrophila

almost certainly extends to the adjacent high moun-

tains of southwestern Angola, especially the botani-

cally poorly known Serra Cafema range just north of the

Otjihipa Range. Dewinteria is one of about seven genera

endemic to the Kaokoveld Centre of Endemism, a bio-

geographical region in arid northwestern Namibia and

adjacent southwestern Angola (Van Wyk & Smith 2001).

Cliffs provide a stable, safe environment for plants

in the absence of larger herbivores and there is often

a defence relaxation tendency (disarmament) in obli-

gate cliff dwellers (Van Jaarsveld & Van Wyk 2003).

However, the vertical habitat (extreme water runoff)

and a lack of space to grow, demand a shift in habit and

reproductive output. This often results in specialist fea-

tures such as succulence, compact growth fonn, dwarf

clustering, cylindrical shape, pendent habit, profuse

flowering and an increase in vivipary (vegetative repro-

duction) (Snogerup 1971; Van Jaarsveld & Van Wyk

2003). There is also a shift towards wind-dispersed seed

(anemochory).

In his original description of Rogeria petrophila,

De Winter (1961) mentions thread-like branches and

smaller capsules of which the function was uncertain.

His surmise is correct. Dewinteria petrophila does have

a remarkable and unique dimorphic facultative repro-

duction strategy — an adaptation that helps it to survive

under the extreme desert conditions of its cliff habitat.

In addition to its conventional aerial branches bearing

insect-pollinated flowers and wind-dispersed seeds, it

produces specialized shoots that are negatively photo-

tropic, entering hairline cracks and other crevices. These

bear small cleistogamous flowers with ± heart-shaped

capsules with fewer but larger, differently shaped seeds

200

Bothalia 37,2 (2007)

FIGURE 7. — Dewinteria petroph-

ila in flower in a fissure on

granite cliffs of the Otjihipa

Mountains in northwestern

Namibia.

(Figures 6, 8). This unique adaptation ensures that seeds

are buried deep into crevices near the mother plant.

Dewinteria petrophila displays a unique amphicar-

pous condition, with both atelechorous and anemocho-

rus dispersal methods. This is a remarkable adaptation

and the first of its kind recorded for an obligate cliff-

dwelling species. Its larger seeds are a self-preserving

strategy, ensuring its long-term survival in its present

habitat. This atelechorous seed dispersal ensures self-

cloning, and the normal smaller anemochorous seeds

(from insect-pollinated flowers) ensures interbreeding

and dispersal to new sites. The larger seed carries larger

reserves that the seedling needs after germination. The

thread-like branches in the crevices ensure an almost

100% survival rate if the mother plant should die owing

to drought or natural causes. The smaller capsules pro-

duce about five or fewer seeds per capsule, whereas the

normal and larger aerial seed capsules produce more

than 50 seeds per capsule. Winds on the cliffs are often

strong and updrafts ensure the seeds’ effective dispersal

to other crevices. D. petrophila flowers in the rainy sea-

son, dispersing its seeds in autumn. Seeds are covered

with mucilage, which is often associated with plants

from desert or semi-desert regions, helping to anchor the

seeds to the substrate (Van der Fiji 1982).

Systematic affinities', the genera of the Pedaliaceae (a

small family of ± 13 genera and 70 species, mostly Afri-

can) are delimited largely by their fruit type (Ihlenfeldt

1964, 1967; Kadereit 2003). They occur mainly in semi-

arid and arid habitats of the Old World tropics (Smithies

& Herman 2003). Life fonns in the family are remarkably

diverse, varying from erect annuals or biennials (Rogeria),

trailing perennials (Dicerocaiyinn), thickset chamaephytes

(Pteroclisciis), to dwarf trees {Sesarnothamniis). Members

have mainly opposite leaves and attractive, gamopetalous,

tubular flowers, the pedicels bearing characteristic extra-

floral nectar glands at the base. The faiits are very diverse,

varying from winged to heavily annoured capsules, that

are dehiscent or indehiscent. Mucilage glands are com-

mon in the family. Perhaps the best known member is

FIGURE 8. — Dewinteria petrophila. Van Jaarsveld & Swanepoel

1 94 1 3: A, nomial seed; B, specialized seed (SEM).

Bothalia 37,2 (2007)

201

Harpagophytum, a tuberous-rooted trailing plant of

the Kalahari, used world-wide as a medicine for arthri-

tis (Van Wyk et al. 1997). Plants of the genus Rogeria

(three species and exclusive to Africa) are robust, erect

annuals or weak perennials from semidesert to des-

ert regions of southern Africa (Smithies & Hennan

2003). Rogeria lorigiflora (Royen) J.Gay ex DC. occurs

in southern Namibia and up the Orange River Valley

as far as Upington in South Africa, and two species in

Namibia, R. adenophyUa J.Gay ex Delile (also extends

to the southern edge of the Sahara from Djibouti to

Senegal and the Cape Verde Islands), and R. bigibbosa

Engl. (Merxmuller & Schreiber 1968) on the escarpment

in central Namibia. They usually grow in disturbed sites

and dry watercourses. The main morphological differ-

ences between the new genus and Rogeria are summa-

rized in Table 1 .

Specimens examined

NAMIBIA. — 1712 (Orokatuwo); east of Ezorotuuo, 17°23'32.0"

S 12°35'0.8" E, (-BC), Van Jaarsveld, Voigt & Cilliers 17520

(Wind); east of Otjihungwa, west-facing slopes of Otjihipa, (-BC),

Van Jaarsveld & Swanepoel 19413, 19527 (Wind); northwestern

Kaokoveld, (-DA), Becker & Ihlenfeldt 107560 (Wind).

ACKNOWLEDGEMENTS

Norval Geldenhuys is thanked for the Latin transla-

tion of the diagnosis. Out thanks are due to Koos Verwey

of Syncro Camp for his hospitality and help during vis-

its to the area, to Wessel Swanepoel for assistance in the

field and to Dr Gillian Maggs-Kolling of the Botanical

TABLE 1 . — Main differences between Dewinteria and Rogeria

FIGURE 9. — Known distribution of Dewinteria petrophila.

Research Institute in Windhoek for her support of our

botanical work in Namibia. Gerrit Gennishuizen and

Emsie du Plessis are thanked for editing the text. Ms M.

Waldron of the Electron Microscope Unit, University of

Cape Town, is thanked for help with the SEM microgra-

phy.

REFERENCES

DE WINTER, B. 1961. Rogeria petrophila. In J.G. Anderson, L.E.

Codd, R.A. Dyer, M.D. Henderson, D.J.B. Killick & B. de

Winter. New and interesting taxa from southern Africa. Kirkia

1: 106-108.

IHLENFELDT, H.-D. 1964. Uber die systematische Gliederung der

Pedaliaceae nach makroskopish erkermbaren Merkmalen.

Berichte der Deutschen Botanischen Gesellschaft 77: 27-31.

IHLENFELDT, H.-D. 1967. Uber die Abgrenzung und die natiirli-

che Gliederung der Pedaliaceae R.Br. Mitteilimgen ans dem

Staatsinstitut fiir Allgemeine Botanick Hamburg 12: 43-128.

KADEREIT, J.W. (ed.). 2003. Families and genera of vascular plants.

vol. 7: 307.

MENDELSOHN, J., JARVIS, A., ROBERTS. C. & ROBERTSON, T.

2002. Atlas of Namibia. Phillip, Cape Town.

MERXMULLER, H. & SCHREIBER, A. 1968. 131. Pedaliaceae.

Prodromus einer flora von Siidwestafrika. Verlag von Cramer,

Lehre.

SMITHIES, S.J. & HERMAN, P.P.J. 2003. Pedaliaceae. In G.

Germishuizen, & N.L. Meyer, Plants of southern Africa: an

annotated checklist. Strelitzia 14: 772-774. National Botanical

Institute, Pretoria.

SNOGERUP, S. 1971. Evolutionary and plant geographical aspects

on chasmophytic communities. In P.H. Davis, P. Harper & I.C.

Hedge, Plant life in southwest Asia: 157-170. The Botanical

Society of Edinburgh.

VAN DER PUL, L. 1982. Principles of dispersal in higher plants.

Springer- Verlag, Berlin.

VAN JAARSVELD, E.J. & VAN WYK. A.E. 2003. Adaptations with-

out barriers: succulent plants shaped by cliffs. Aloe 40: 98-103.

VAN WYK, A.E. & SMITH, G.F: 200 1 . Regions offloristic endemism in

southern Africa: a review with emphasis on succulents. Umdaus

Press, Hatfield, Pretoria.

VAN WYK. B.-E., VAN OUDTSHOORN, B. & GERICKE, N. 1997.

Medicinal plants of South Africa. Briza Publications, Pretoria.

E.J. VAN JAARSVELD* and A.E. VAN WYK**

* South African National Biodiversity Institute, Private Bag X7, 7735

Claremont, Cape Town. Student affiliation: Department of Botany,

University of Pretoria, Pretoria.

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

sity of Pretoria, 0002 Pretoria.

MS. received: 2006-06-06.

202

Bothalia 37,2 (2007)

DIDIEREACEAE/PORTULACACEAE

CERARIA KAOKOENSIS, A NEW SPECIES FROM NAMIBIA, WITH NOTES ON GYNODIOECY IN THE GENUS

INTRODUCTION

Traditionally, Didiereaceae s.str: has been treated as a

family endemic to Madagascar. Based on morphological

similarities, a close affinity between Didiereaceae and

the African portulacaceous genera Ceraria El. Pearson

& Stephens and Portiilacaria Jacq. has been suggested

(Rauh & Scholch 1965). Molecular evidence indicates

that Ceraria and Portiilacaria, as well as the African

genus Calypti'otheca Gilg, are more closely related to

Didiereaceae than to other Portulacaceae (Hershkovitz &

Zimmer 1997; Applequist & Wallace 2000, 2001). The

circumscription of Didiereaceae was therefore enlarged to

accommodate these three genera (Applequist & Wallace

2003). Didiereaceae s.I. is divided into three subfami-

lies, of which Portulacarioideae comprises Ceraria and

Portiilacaria.

Ceraria is endemic to the arid western parts of south-

ern Africa. The infrageneric classification of the group is

not satisfactory and what appear to be still undescribed

taxa are encountered in the wild. Herbarium specimens

of the group are generally incomplete, fragmentary and

not very useful for comparative morphological studies.

Depending on the authority, from four (Craven 1999;

Germishuizen & Meyer 2003) to six (Rowley 2002)

species are recognized at present. Two of the described

species, C. carrissoana Exell & Mendonqa and C. lon-

gipedunculata Merxm. & Podlech, are restricted to the

Kaokoveld Centre of Endemism in northwestern Namibia

and southwestern Angola (Van Wyk & Smith 2001),

whereas the ranges of the other species are ± centred on

the Gariep Centre of Endemism in southwestern Namibia

and northwestern South Africa (Van Wyk & Smith 2001;

Curtis & Mannheimer 2005).

In this contribution, a new species of Ceraria from

the Kaokoveld, Namibia, is described. Gynodioecy — a

sexual state in which plants of the same species bear

either functionally female or bisexual flowers — is also

recorded in the genus for the first time. During a botani-

cal expedition to the Otjihipa Mountains in northwest-

ern Namibia, the author noticed a Ceraria that superfi-

cially resembles C. friiticiilosa H. Pearson & Stephens

from southern Namibia and the Northern Cape Province

of South Africa. This taxon was subsequently found in

several other localities in the northern Kaokoveld. It has

apparently not been collected before, as no herbarium

specimens of it could be found in either NBG, PRE,

SAM or WIND.

Live material from the various known populations

of the new species was studied in the field and from

plants in cultivation. Moiphological characters were all

determined from mature leaves, fresh flowering mate-

rial and from ripe fruit. Diagnostic features for all the

above-mentioned species and C. friiticiilosa were deter-

mined through examination of herbarium specimens

and live plants in the Kaokoveld and southern Namibia

(Rosh Pinah area), respectively. Additional infonna-

tion was found in Pearson & Stephens (1912), Exell &

Mendon9a (1938-1939), Merxmiiller & Podlech (1961)

and Podlech ( 1 967).

Ceraria kaokoensis Swanepoel, sp. nov., C. fru-

ticulosam H. Pearson & Stephens tangit ob arborem fruti-

cosam et corticem brunneolam heterochromam, foliorum

aliquorum laminam oblanceolatam, flores singulares vel

fasciculatos, roseos. Differt cortice fissurata longitudina-

liter ramis ramulisque, alba incremento recente et viridi

flavo-virenteve novo incremento, prunia carente; novo

incremento obtecto pilis brevibus, conicis, papilliformi-

bus; foliis petiolatis vel subsessilibus; lamina saepe line-

ari-oblanceolata vel lineari, angustata, 3^ plo longiore

quam latiore, tenuiore simili longitudine, bevibus, coni-

cis, papilliformibus pilis, viridi, prunia carente; florente

profuse, ovario et ovulo parvulis.

TYPE. — Namibia, 1712 (Posto Velho): Otjihipa Moun-

tains, 4 km ESE of Otjinhungwa, 760 m, (-AD), 17-01-

2005. Swanepoel 224 (WIND, holo.!; PRE, iso.!).

Gynodioecious, semisucculent, densely branched,

shrub-like tree, 0. 3-2.0 x 0. 3-2.0 m. Trunk branch-

ing repeatedly just above ground level into many stems,

rarely up to 0.2 m high, up to 0.5 m wide when wedged

between rocks. Bark smooth, longitudinally fissured,

often peeling in small flake-like pieces, dark brown, red-

dish brown or greyish brown, new growth green to yel-

lowish green, bark on more recent growth creamy white,

with short, brittle, conical hairs, becoming glabrous with

age. Branches and branch! ets with small, raised, cush-

ion-like nodes, opposite and decussate at intervals of

2-13 mm; new branchlets often growing from cushions,

glabrous; younger growth with short, conical, papilla-like

hairs, some hairs bifid at apex; ultimate branchlets 1-2

mm diam.; when dried, irregularly winged or grooved

in places when viewed under magnification, younger

growth usually ± square in t/s. Leaves deciduous, single

or clustered, opposite and decussate on new growth,

green or yellow-green; lamina fleshy, flat or falcate

towards abaxial side, oblanceolate, linear-oblanceolate

or linear, (0.8-)3.0-12.0(-14.5) x (0.4-)1.2-3.0(-3.8)

X (0.3-)0. 6-1.1 (-1.3) mm, ratio of length versus width

3^:1, in t/s narrowly elliptic, crescent-shaped, reniform

or oblong, adaxial side flat, convex or concave, abaxial

side convex or flat, apex obtuse, acute or emargin-

ate, often minutely apiculate, base cuneate or cuneate

and abruptly rounded onto petiole, with short, conical,

papilla-like hairs; margin entire; midrib or veining not

visible; petiolate or subsessile, petiole up to 1 mm long,

in t/s ± circular, 0. 1-0.5 x 0. 1-0.3 mm long, glabrous or

with very short, conical, papilla-like hairs. Inflorescence:

flowers borne on cushion-like nodes in clusters of 2-14

or solitary, usually flowers profusely. Flowers bisexual

or female, pedicellate, glabrous, appearing before or with

leaves; bracts ± ovate, flat to cucullate, up to 0.6 mm

long, glabrous; pedicel very slender, 0. 1-0.2 mm diam.,

green or reddish green, inserted on a short, peduncle-like

structure, up to 0.3 mm long, involucrated by bracts;

receptacle conical, green or reddish green, fleshy. Calyx

of 2 sepals, short, broad, membranous, contiguous or

distant, bract-like, persistent, hemispherical or triangular,

white with pinkish tinge or pink, apical part often dic-

ing soon to conspicuous reddish brown colour. Corolla:

Bothalia 37,2 (2007)

203

petals 5, persistent, oblanceolate, linear-oblanceolate or

elliptic, often cucullate, especially towards apex, pink-

ish white, pink or pinkish red. Ovary ftask-shaped, in t/s

slightly flattened, triquetrous or rarely elliptic, pinkish

white, pink or cherry red; style none; stigmas 3, yellow-

ish white or white; ovule oblong-ellipsoid, up to 0.3 x 0. 1

mm long. Bisexual flowers 1 .8-2.8 mm long; pedicel 1 .9-

3.6 mm long; receptacle 0. 3-0.4 x 0. 5-0.6 mm; sepals ±

0.4 X 0.6 mm; petals 1. 9-2.4 x 0.4-0. 8 mm; stamens 5;

anthers conspicuously pinkish red, 0.3-0. 5 mm long; fila-

ments 1.3-2. 3 mm long, subterete, pinkish white or pink;

pollen orange-yellow or cherry-red; pistil 0.9-1. 5 mm

long; ovary 0.9-1. 3 x 0. 3-0.4 x 0.2 mm; stigmas ± linear,

pustulate above, short, up to 0.1 mm long, rarely patulous

and triangular. Female flowers 1. 1-2.0 mm long; pedicel

1. 2-2.0 mm long; receptacle 0.3 x 0.6 mm; sepals ± 0.3 x

0.6 mm; petals ± 1.4 x 0.5 mm; stamens 5, rudimentary;

anthers not developed; pistil 1.2-1. 5 mm long; ovary

1.0-1. 3 X 0. 3-0.4 X 0.3 mm; stigmas linear or triangular,

pustulate above, spreading, relatively long when linear,

up to 0.4 mm long. Fruit asymmetrically elliptic or hemi-

spherical, 3. 3^. 4 X 1.4-1. 8 X 0. 9-1.1 mm, apiculate,

almost flat, very narrowly triquetrous in t/s, soft, pink,

soon drying to brown, pod-like, one-seeded, indehiscent;

pedicel extremely thin, brittle; probably wind-dispersed.

Flowering time: November to March. Pollination', vari-

ous species of flies, blowflies and bees, including mopane

bees, were observed visiting the flowers. Figures 10-12.

Diagnostic characters and affinities'. Ceraria kao-

koensis is very attractive when in flower and as such

would be a valuable addition to succulent gardens.

During the flowering season it is easily recognized, even

from a distance, due to the profuse pink flowers. C. kao-

koensis is probably most closely related to C. fruticu-

losa, from which it differs mainly in bark, leaf and inflor-

escence characters, as well as geographical distribution.

C. kaokoensis can also be confused with the sympatric

C. longipedunculata, a species with which it shares a

much-branched habit and narrow, superficially similar

leaves. However, the leaf lamina is ± cylindrical in C.

longipedunculata, but distinctly flattened in C. kaokoen-

sis. Vegetatively C. longipedunculata is most similar to

C. namaquensis (Sond.) H. Pearson & Stephens, a spe-

cies from southern Namibia and adjacent parts of South

FIGURE 10. — Ceraria kaokoensis in

its natural habitat, 1 .2 m tall.

Africa (Namaqualand). Some of the more prominent

morphological characters to differentiate between C. kao-

koensis and C. fruticulosa are summarized in Table 2.

Etymology: the specific epithet refers to the Kaokoveld

of northwestern Namibia. The distribution of Ceraria

kaokoensis falls within the previous politically demar-

cated Kaokoland, now part of the Kunene Region.

Distribution and habitat: Ceraria kaokoensis is pres-

ently known from a few isolated localities, all within

FIGURE 1 1 . — Bark of Ceraria kaokoensis.

204

Bothalia 37,2 (2007)

the Kaokoveld Centre of Endemism in northwestern

Namibia (Figure 13). More specific localities include

the Otjihipa and Hartmann Mountains to the east and

west of the Marienfluss respectively; the rocky area to

the south of the Engo River Valley and the mountain-

ous area to the west and north-northwest of Sesfontein.

It is localized and common to rare in these areas. The

species almost certainly occurs in the adjacent moun-

tainous parts of southwestern Angola as well, especially

the Serra Cafema range, and may prove to be more

widespread on the mountains of the Kaokoveld Centre

of Endemism, most of which remains botanically poorly

explored (Van Wyk & Smith 2001). It is found 40-110

km from the coast at altitudes of 700-1 100 m, where

the mean annual rainfall is 50-150 mm (Mendelsohn

et al. 2002). It appears to be habitat specific as it was

only found in rocky places on mountain slopes, plateaus

and on rocky outcrops. It is limited to the granites and

gneisses of the Epupa Metamorphic Complex and to

mica schist of the Damara Supergroup (Miller & Schalk

1980; Mendelsohn et al. 2002). At all the presently

known locations, C. kaokoensis was found to be sympat-

ric with C. longipeduncidata and in some instances also

with C. carrissoana.

Other specimens examined

NAMIBIA. — 1712 (Posto VeUio): Hartmann Valley Mtns, highest

peak (-AC), Swanepoel 225 (WIND), 6 km NW of Ombivango, (-

AD), Swanepoel 226 (WIND), Hartmann Valley, peak of 1 039 m high

mountain, (^A), Swanepoel 265 (WIND), 4 km SW of Ombivango (-

CB), Swanepoel 227 (WIND); 10 km south of Engo River Valley, near

Skeleton Coast Park boundary, (-CC), Swanepoel 233 (WIND). 1813

(Opuwo): 10 km SE of Ozombari, (-DC), Swanepoel 228 (WIND).

1913 (Sesfontein): Ganamub River, 10 km N of junction with Hoanib

River, (-AB), Swanepoel 260 (WIND), 10 km NNW of Sesfontein,

(-BA), Swanepoel 229, 231, 232 (WIND).

Key to species

la Leaves relatively large; lamina broad, flattened, elliptic, obovate, broadly oblanceolate or suborbicular, 10-37 x 10-31 mm. Sparsely

branched shrubs or small trees. Otjihipa and Baynes Mountains, Kaokoveld, Namibia; also in southwestern Angola C. carrissoana

lb Leaves relatively small; lamina narrow to broad, up to 15 x 9 mm, ± circular in t/s or flattened. Much-branched shrubs or small trees.

Kaokoveld (northwestern Namibia) or Gariep Centres of Endemism (southern Namibia en adjacent parts of Northern Cape, South Africa):

2a Leaf lamina cylindrical (± circular in t/s), narrowly cuneate, conspicuously succulent. Inflorescences racemose or subpaniculate; flow-

ers red, pink, pinkish white or white. Shrubs or small trees up to 5 m tall. Bark on stems dark- to pale-coloured: blackish brown,

dark brown, reddish brown, cream-coloured or greyish white, smooth but peeling horizontally in tough, papery strips or pieces.

Widespread in northern half of Kaokoveld Centre of Endemism, Namibia and in southwestern Angola C. longipediinculata

2b Leaf lamina flattened (narrowly elliptic in t/s), slightly succulent. Flowers solitary or in clusters, pink. Shrubs or shrub-like trees up to

2 m tall. Bark on stems dark-coloured. Kaokoveld and Gariep Centres of Endemism:

3a Bark on branches/branchlets longitudinally fissured, younger growth with short, conical, papilla-like hairs. Leaves with lamina nar-

row, ratio of length versus width 3—4:1, linear, linear-oblanceolate or oblanceolate, with short, conical, papilla-like hairs. Flowers

produced in profusion, usually borne in dense clusters on cushion-like nodes along stems. Shrub-like trees up to 2 m tall. Locally

in northern half of Kaokoveld Centre of Endemism, Namibia; probably also in southwestern Angola C. kaokoensis

3b Bark on branches/branchlets smooth, glabrous. Leaves with lamina narrow to broad, ratio of length versus width 1-3:1, oblanceo-

late, obovate, oblong-obovate or obcordate, glabrous. Flowers few, sparsely scattered along stems. Shrubs up to 1 .5 m tall. Mainly

Gariep Centre of Endemism, southern Namibia and Northern Cape (South Africa) C. fruticulosa

TAXONOMIC SIGNIFICANCE OF GYNODIOECY IN CERARIA

The genus Ceraria was described by Pearson &

Stephens (1912) and distinguished from Portulacaria

in being dioecious, whereas the latter is hermaphroditic

(flowers bisexual). Subsequently this sexual distinction

has been widely employed in identification keys to dif-

ferentiate between the two genera (Dyer 1975; Carolin

1993; Jordaan 2000; Eggli 2002). Dyer (1975), Carolin

(1993), Rowley (2002) and others noted that flowers in

Ceraria may be rarely bisexual. However, functionally

unisexual flowers in Ceraria bear rudiments of the recip-

rocal organs, hence flowers may easily be mistaken for

being either structurally or functionally bisexual.

As in other members of the genus, flowers of Ceraria

kaokoensis appear structurally bisexual, although the sta-

mens are smaller with indehiscent anthers in female flow-

TABLE 2. — Prominent differences between Ceraria kaokoensis and C. fhtticulosa

Bothalia 37,2 (2007)

205

FIGURE 12. — Ceraria kaokoensis. A, branch with leaves; B, flowering

branch; C, female flower; D, bisexual flower; E, fhiit. C & E,

Swanepoel 226\ D, Swanepoel 227. Scale bars: A, 10 mm; B, 3

mm; C, D, 0.5 mm; E, 1 mm. Artist: Julia Kreiss.

FIGURE 13. — Known distribution of Ceraria kaokoensis.

ers and the pistil is smaller in male flowers. However, in

C. kaokoensis both types of flowers were seen to develop

fruit with seed and the species is best described as gyno-

dioecious. Plants of the new species are either function-

ally female or hermapliroditic and this is supported by

observations on plants grown in the author’s garden in

Windhoek. However, in angiosperms, all fomis of dio-

ecy are rarely absolute (Policansky 1982). The possibil-

ity that at times some structurally bisexual flowers in

C kaokoensis may still be functionally male cannot be

excluded.

Following the confirmation of gynodioecy in at least

some plants of Ceraria kaokoensis, a critical re-assess-

ment of the sexual state in other species traditionally

referred to Ceraria is required as more members may

be predominantly gynodioecious, and not dioecious (or

rarely hermaphroditic) as have hitherto been reported.

More field work, preferably involving long-tenn moni-

toring of specific plants, is required to fully elucidate

patterns of sexual expression in the group. Moreover,

observations on the sexual state of Portiilacaria armiana

Van Jaarsv. is required to confirm its current generic

placement; it may well be better classified as a Ceraria

(E.J. van Jaarsveld pers. comm.).

The presence of gynodioecy in at least one mem-

ber of Ceraria is of considerable phylogenetic signifi-

cance. Current knowledge would indicate that all other

Portulacaceae are hermaphroditic, with the exception of

Talinella Baill., a Madagascan genus (12 species) of ±

woody shrubs with lax and slender branches. Most spe-

cies of Talinella are dioecious, with either the stamens

or the gynoecium vestigial. Indications are that some of

the species may at least be morphologically gynodioe-

cious (Applequist 2005). Although Hershkovitz (1993)

associated Talinella with Portulacaria and Ceraria, sub-

sequent molecular studies have clearly shown that it is

most closely related to Talinum Adans. (Hershkovitz &

Zimmer 1997; Applequist & Wallace 2001). Gynodioecy

has, however, also been reported in the monotypic

Decaryia Choux, one of the more basal lineages of the

206

Bothalia 37,2 (2007)

Otherwise dioecious Didiereaceae s.str. (Applequist &

Wallace 2000; Schatz 2001). The presence of gynodio-

ecy in Decatyia might be a plesiomorphy, an interpreta-

tion supported by its presence in the even more distantly

related genus Ceraria. This argument presupposes rever-

sion to hennaphrodite flowers in Calyptrotheca. It also

provides support for the suggested placement of Ceraria

in an expanded Didiereaceae.

ACKNOWLEDGEMENTS

I would like to thank Prof A.E. van Wyk, University

of Pretoria, for advice and support, Prof T.V. Jacobs,

UNISA, for translating the diagnosis into Latin, Ms

Hester Steyn, SANBI, for preparing the distribution

map and Ms Julia Kreiss for the line drawings. The

curator and staff of the National Herbarium of Namibia

are thanked for their assistance during visits to the her-

barium. The National Herbarium of Namibia and the

South African National Biodiversity Institute are also

thanked for the use of infonnation from their databases;

SPMNDB, Flora DB and PRECIS. The curator. National

Herbarium, Pretoria, is thanked for access to their collec-

tions; the assistance of Ms Marie Jordaan during visits

to the herbarium is acknowledged with thanks. Mr Koos

Verwey of Otjinhungwa is thanked for logistical support

during visits to the Otjihipa Mountains. The University

of Pretoria is thanked for financial support. I am espe-

cially grateful to my wife Hannelie for assistance and

companionship during field trips.

REFERENCES

APPLEQUIST, W.L. 2005. A revision of the Malagasy endemic

Talinella (Portulacaceae). Adans'onia, ser. 3, 27; 27-80.

APPLEQUIST, W.L. & WALLACE, R.S. 2000. Phylogeny of the

Madagascan endemic family Didiereaceae. Plant Systematics

and Evolution 221: 157-166.

APPLEQUIST, W.L. & WALLACE, R.S. 2001. Phylogeny of the por-

tulacaceous cohort based on ndh¥ sequence data. Systematic

Botany 26: 406^19.

APPLEQUIST, W.L. & WALLACE, R.S. 2003. Expanded circum-

scription of Didiereaceae and its division into three subfamilies.

Adansonia, ser. 3, 25: 13-16.

CAROLIN, R.C. 1993. Portulacaceae. In K. Kubitzki, J.G. Rohwer &

V. Bittrich, The families and genera of vascular plants — dicoty-

ledons 2: 544-555. Springer- Verlag, Berlin.

CRAVEN, P. (ed.). 1999. A checklist of Namibian plant species.

Southern African Botanical Diversity Network Report No. 7.

SABONET, Windhoek.

CURTIS, B.A. & MANNHEIMER, C.A. 2005. Tree atlas of Namibia.

National Botanical Research Institute, Windhoek.

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

vol. 1 : dicotyledons. Department of Agricultural Technical Servi-

ces, Pretoria.

EGGLI, U. (ed.). 2002. Illustrated handbook of succulent plants: dicoty-

ledons. Springer- Verlag, Berlin.

EXELL, A.W. & MEND0N(;A, F.A. 1938-1939 [published 1939].

Contribui'foes para o conhecimento da flora de Africa. Boletim

da Sociedade Broteriana 13: 309, 310.

GERMISHUIZEN, G. & MEYER. N.L. (eds). 2003. Plants of southern

Africa: an annotated checklist. Strelitzia 14. National Botanical

Institute, Pretoria.

HERSHKOVITZ, M.A. 1993. Revised circumscriptions and subgeneric

taxonomies of Calandrinia and Montiopsis (Portulacaceae) with

notes on phylogeny of the portulacaceous alliance. Annals of the

Missouri Botanical Garden 80: 333-365.

HERSHKOVITZ, M.A. & ZIMMER, E.A. 1997. On the evolutionary

origins of the cacti. Taxon 46: 217-232.

JORDAAN, M. 2000. Portulacaceae. In O.A. Leistner, Seed plants of

southern Africa: families and genera. Strelitzia 10; 453^56.

MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T.

2002. Atlas of Namibia. Philip, Cape Town.

MERXMULLER, H. & PODLECH. D. 1 961 . Miscellanea. Portulacaceae,

Ceraria longipedunculata Merxm. & Podlech spec. nov. Mittei-

lungen der Botanischen Staatssammlung Miinchen 4: 73, 74.

MILLER, R. McG. & SCHALK, K.E.L. 1980. Geological map of

South West Africa/Namibia. Geological Survey of the Republic

of South Africa and South West Africa/Namibia.

PEARSON, H.H.W. & STEPHENS, L. 1912. 3. List of the plants coll-

ected in the Percy Sladen Memorial Expedition, 1908-1909,

1910-1911: Portulacaceae. Annals of the South African Museum

9: 30-35.

PODLECH, D. 1967. Portulacaceae. Prodromus einer flora von

Siidwestafrika 29: 2-7. Cramer, Lehre.

POLICANSKY, D. 1982. Sex change in plants and animals. Annual

Review of Ecology and Systematics 13: 471 -M95 .

RAUH, W. & SCHOLCH, H.F. 1965. Weitere Untersuchungen an

Didiereaceen. 2. Teil. Inflorescenz-, blutenmorphologische

und embryologische Untersuchungen mit Ausblick auf die

systematische Stellung der Didieeraceen. Sitzungsberichte der

Heidelberger Akademie der Wissenschaften 1965, 3: 221^34.

ROWLEY, G.D. 2002. Ceraria. In U. Eggli, Illustrated handbook of suc-

culent plants: dicotyledons'. 379-380. Springer- Verlag, Berlin.

SCHATZ, G.E. 2001. Generic tree flora of Madagascar. Royal Botanic

Gardens, Kew & Missouri Botanical Garden.

VAN WYK, A.E. & SMITH, G.F. 2001 . Regions offloristic endemism in

southern Africa. A review with emphasis on succulents. Umdaus

Press, Hatfield, Pretoria.

W. SWANEPOEL*

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

sity of Pretoria, 0002 Pretoria, South Africa. Postal address: P.O. Box

21168, Windhoek, Namibia.

MS. received: 2006-08-11.

CRASSULACEAE

BRYOPHYLLUM PROLIFERUM ^NATURALIZED IN KWAZULU-NATAL, SOUTH AFRICA

The most efficient method of multiplication by suc-

culents is via adventitious plantlet proliferation. A num-

ber of alien succulents that use this method of propaga-

tion has become naturalized or weedy in South Africa.

Well-known examples include Agave sisalana Perrine

(Smith & Mossmer 1996) and Opimtia ficiis-imUca L.

(Obenneyer 1976). Abnormal (adventitious) develop-

ment is reflected in the panicle of A. sisalana becoming

bulbiferous after flowering, and shooting and rooting of

the detached fruits occurring in O. ficus-indica. Amongst

the few Crassulaceae that have been recorded naturalized

in southern tropical Africa, the genus Biyophylhiin Salisb.

features prominently (Fernandes 1983). However, in the

Flora of southern Africa region, Biyophylhiin delagoense

(Eckl. & Zeyh.) Schinz is the only member of this family

to have been recorded as such (Toelken 1985; Dreyer &

Bothalia 37,2 (2007)

207

FIGURE 14. — A. leaves; B. plant showing adventitious buds.

Makwarela 2000) although Wells et al. (1986) listed B.

proliferum Bowie ex Hook, as a potential problem plant.

Recent field work in KwaZulu-Natal and subsequent

consideration of herbarium collections have revealed

that the hardy stem and leaf succulent Bryophyllum pro-

liferum has escaped from cultivation. Although these

populations have not been shown to be self replacing

for a period of ten years, we categorize this hardy inva-

sive as naturalized, for the Botha’s Hill population is

evidently expanding through recruitment. The popula-

tion status at the original 1985 collection site on Ismont

Farm has not been reassessed. Significantly, in defining

‘naturalized’ alien invasive taxa, Pysek et al. (2004)

admitted that how long a species must persist to be con-

sidered naturalized is inevitably arbitrary.

This tall species, a native of Madagascar, is grown

predominantly for its foliage (Figure 14) rather than

its rather unattractive greenish yellow flowers; detract-

ing further from its horticultural appeal are the inflor-

escences (terminal panicles) that are usually disfigured

FIGURE 15. — Known distribution oi Bryophyllum proliferum based on

specimens at NH, NU and PRE.

by the numerous adventitious buds that spring from the

pedicel bases of aborted flowers. Plants grow as clump-

forming subshrubs, with decussate leaf pairs borne

along the upper third of the four-angled, somewhat

woody stems. The impari-pinnatisect leaves are a ver-

dant pale green and marginally crenate, each with 3-7

opposite pairs of sessile leaflets that are subdecurrent

and asymmetric at the base. Small plantlets may addi-

tionally be produced in the crenatures of the entire leaf,

and along the upper grooved midrib. Wells et al. (1986)

described further proliferation via root suckers. The

leaves are borne more or less horizontally, effectively

shading out low-growing indigenous vegetation.

It is likely that the species has been naturalized

(Figure 15) for some time but has gone unnoticed. This

phenologically plastic species presents variable leaf

morphologies and coloration; for example, leaf margins

turn red under high light intensities, whereas this trait is

absent from specimens established in shade.

Specimens examined

KWAZULU-NATAL. — 2930 (Pietermaritzburg): 1.5 km east of

The Valley Trust, Portion 585 of the Farm Assagay Kraal, Botha’s

Hill, colony alongside stream in granite gully. Growing in semi-shade

along with Adiantum capillus-veneris and Pteris vittata, 450 m, S

29°, 735 169; E 30°, 748894, (-DA), 05-05-2007, N. Crouch 1123 (NH).

3030 (Port Shepstone): edge of grassy track in Eucalyptus plantations

between Mount Langford and Loni River (mid-Illovo area), on Ismont

Farm, (-BA), 15-07-1985,5. Culcross s.n. (NH).

ACKNOWLEDGEMENTS

Ms L. Henderson and Mr G. Nichols are thanked for

helpful discussions on the identity of B. proliferum.

REFERENCES

DREYER, L.L. & MAKWARELA, A.M. 2000. Crassulaceae. In O.A.

Leistner, Seed plants of southern Africa: families and genera.

Strelitzia 10; 235, 236. National Botanical Institute, Pretoria.

208

Bothalia 37,2 (2007)

FERNANDES, R. 1983. Crassulaceae. In E. Launert, Flora zambe-

siaca 7, 1 : 3-7 1 .

OBERMEYER. A. A. 1976. Cactaceae. In J.H. Ross, Flora of southern

Africa Tl\ 144-156. Botanical Research Institute. Pretoria.

PYSEK, P, RICHARDSON, D.M., REJMANEK, M., WEBSTER.

G.L., WILLIAMSON, M. & KIRSCHNER, J. 2004. Alien

plants in checklists and floras: towards better communication

between taxonomists and ecologists. Taxon 53: 131-143.

SMITH, G.F. & MOSSMER, M. 1 996. FSA contributions 4: Agavaceae.

Bothalia 26: 31-35.

TOELKEN, [as Tdlken] H.R. 1985. Crassulaceae. Flora of southern

Africa 14: 1-244.

WELLS, M.J., BALSINHAS, A.A., JOFFE, H., ENGELBRECHT,

V.M., HARDING, G, & STIRTON, C.H, 1986. A catalogue of

problem plants in southern Africa. Memoirs of the Botanical

Survey of South Africa No. 53. Botanical Research Institute,

Pretoria.

N.R. CROUCH* and G.F. SMITH**

* Ethnobotany Unit, South African National Biodiversity Institute,

P.O. Box 52099, 4007 Berea Road/School of Chemistry, University of

KwaZulu-Natal, 4041 Durban.

E-mail: Crouch@sanbi.org.

** Biosystematics Research and Biodiversity Collections, South

African National Biodiversity Institute, Private Bag XIOl, 0001

Pretoria/Acocks Chair, Schweickerdt Herbarium, Department of

Botany, University of Pretoria, 0002 Pretoria.

E-mail: SmithG@sanbi.org.

CRASSULACEAE

CRASSULA STREYI RECORDED FROM THE EASTERN CAPE, SOUTH AFRICA

The most recent checklist of the Crassulaceae in

South Africa (Burgoyne 2006) delimits, as have earlier

accounts (Hunt 1979; Dreyer 1993; Burgoyne 2003),

Crassula streyi Toelken as a KwaZulu-Natal endemic.

However, recent field work in the Transkei region of the

Eastern Cape Province and subsequent consideration of

herbarium materials has revealed that this species has

been known from the Eastern Cape for over 20 years.

It was first collected there by Mr Tony Abbott in 1985,

in forest adjacent to the Msikaba River Mouth, near

where the Sao Bento was wrecked in 1554. It has more

recently been collected to the north of the Mkambati

Game Reserve, alongside the Mtentu River (Figure 16).

It was considered a highly restricted endemic, occurring

at a few localities within the Mtamvuna Nature Reserve

(Toelken 1973, 1985); its range now extends southwest-

wards by approximately 25 km. This species evidently

occurs in suitable rocky forest habitat in the intervening

Mzamba Gorge system, and will likely be found along

the lower Mnyameni Gorge. The Eastern Cape collec-

tions confirm C. streyi sight records, and its status as

endemic to the Msikaba sandstone-defined Pondoland

Centre (Scott-Shaw 1999; Van Wyk & Smith 2001;

FIGURE 16. — Known distribution of Crassula streyi based on speci-

mens at GRA, KEI, NI I, NU and PRE.

Abbott 2006). Furthermore, the Izotsha Falls (within

5 km of Oribi Gorge) voucher extends its range north-

eastwards by 40 km. It is restricted to altitudes ranging

from 2-280 m and is therefore always found close to the

coast.

Previously uncited KwaZulu-Natal vouchers extend

substantially the documented phenology of C. streyi

(Toelken 1985): plants flower from spring to late summer

as well as in mid-winter.

Specimens examined

KWAZULU-NATAL.— 3030 (Port Shepstone): Izotsha Falls view

site, ex hort., (-CB), 25-03-1974, Hilliard 5493 (NU); Umtamvuna

Nature Reserve, forest walk, rock face in forest, (-CC), 24-12-1983, A.

Abbott 1586 (NH). 3130 (Port Edward): Umtamvuna Nature Reserve.

Clearwater River Trail, (-AA), 04-02-1983, A. Abbott 766 (NH);

Umtamvuna Nature Reserve, (-AA), 08-03-2001, P.M. Gavhi, P.J.H.

Hurter & E. Van Wyk 61 (PRE).

EASTERN CAPE. — 3129 (Port St Johns): Mkambati, Sao Bento

crossing, rock in forest, (-BD), 24-03-1985, A. Abbott 2553 (KEI;

NH). 3 1 30 (Port Edward): Ikubu River Gorge, tributary of the Mzamba

River, riverine forest in shade on forest floor, (-AA), 22-06-1986, A.E.

van Wyk 7305 (KEI); Transkei, Mzamba District, Sikuba River, in for-

est, (-AA), 22-06-1986, A. Abbott 3176 (KEI); rocky ledge alongside

northern bank of Mtentu River, 1 km inland from mouth, opposite

Mkambati Game Reserve, ex hort., (-AA), 10-06-2007, N. Crouch,

T. Edwards & D. Bellstedt 1129 (NH); Mkambati Reserve, Kebengeni

River (tributary of Mtentu River), riverine scrub, growing on a rock on

sandy soil on streambank, in sun on a level area, undisturbed, Msikaba

formation sandstone, (-AA), 15-09-1998, E. Cloete 5236 (GRA); com-

mon in cliffs of estuary of Mtentu River, (,-AA), 06-2006, Edwards,

Crouch & Bellstedt 3275 (NU).

ACKNOWLEDGEMENTS

The trip during which the most recent material was

obtained, was financed by the University of KwaZulu-

Natal, with D. Bellstedt kindly assisting with transport

in the field. Mr A. Dold of GRA and Dr K. Immelman of

KEI are thanked for making their speeimen data available.

REFERENCES

ABBOTT, A. 2006. The story of the Pondoland Centre. PlantLife 33

& 34: 1-62.

Bothalia 37.2 (2007)

209

BURGOYNE, RM. 2003. Crassulaceae, In G. Germishuizen & N.L.

Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 378-413.

BURGOYNE, P.M. 2006. Crassulaceae. In G. Germishuizen, N.L.

Meyer, Y. Steenkamp & M. Keith, A checklist of South African

plants. Southern African Botanical Diversity Network Report

No. 41: 336-369. SABONET, Pretoria.

DREYER, L. 1993. Crassulaceae. In T.H. Arnold & B.C. De Wet,

Plants of southern Africa: names and distribution. Memoirs of

the Botanical Siin’ey of South Africa No. 62: 328-341 . National

Botanical Institute, Pretoria.

HUNT, D.R. 1979. Crassula streyi. Curtis's Botanical Magazine 182,

part 3: 88-90, t. 770.

SCOTT-SHAW, C.R. 1999. Rare and threatened plants of KwaZulu-

Natal and neighbouring regions. KwaZulu-Natal Nature

Conservation Service, Pietermaritzburg.

TOELKEN, [as Tblken] H.R. 1973. Crassula streyi. The Flowering

Plants of Africa 42: t.l672.

TOELKEN, [as Tblken] H.R. 1985. Crassulaceae. Flora of southern

Africa 14: 1-244.

VAN WYK, A.E. & SMITH, G.F. 2001 . Regions offloristic endemism in

southern Africa. A review with emphasis on succulents. Umdaus

Press, Pretoria.

N.R. CROUCH* and T.J. EDWARDS**

* Ethnobotany Unit, South African National Biodiversity Institute, P.O.

Box 52099, 4007 Berea Road/School of Chemistry, University of

KwaZulu-Natal, 4041 Durban.

** School of Biological and Conservation Sciences, University of Kwa-

Zulu-Natal, Private Bag XOl, 3209 Scottsville.

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Bothalia 37,2: 211-214 (2007)

Stem diameter and bark surface area of the fluted trunk of Balanites

maughamii (Balanitaceae)

V.L. WILLIAMS*t, K. BALKWILL* and E.T.F. WITKOWSKI*

Keywords: Balanites maughamii Sprague, commercial bark harvesting, diameter at breast height (dbh), traditional medicine

ABSTRACT

Balanites maughamii Sprague (Balanitaceae) is a woodland tree used and harvested for bark products in the traditional

medicine trade of South Africa. The tree has a distinctively fluted and buttressed stem, especially in mature individuals. This

short communication quantifies the relationship between two diameter measurements D1 and D2 that respectively exclude

and include the bark surface contained in the convolutions of the flutes at five height intervals up the stem to 2 m. Regressions

show D 1 to be an accurate predictor of D2 (r = 0.97-0.99) over a range of tree sizes, hence obviating the necessity to measure

both D1 and D2. The circumference and bark surface area on the stem was determined to estimate the quantity of bark that

can potentially be harvested. At least 69% of the stem circumference and bark surface area was estimated to be contained

within the convolutions of the flutes.

INTRODUCTION

Balanites maughamii Sprague (Balanitaceae) is a

medium to large, slow-growing deciduous tree ranging

from 8-20 m tall (Pooley 1993). The stem is straight

and the trunks of older trees are distinctively fluted and

buttressed (Pooley 1993; Van Wyk & Van Wyk 1997).

The grey bark has medicinal value and is harvested

and sold to consumers in traditional medicine markets

in KwaZulu-Natal (KZN), Gauteng and Mpumalanga

(Botha et al. 2001; Grace 2002; Williams 2003) (Figure

1). Based on the total amount of bark harvested (m^),

B. maughamii was ranked third out of 36 tree spe-

cies harvested for bark in the woodlands of southern

Maputaland, KZN (Twine 2004). A detailed population

study there revealed that 55% of all individuals [diam-

eter at breast height (dbh) > 10 cm] had harvest wounds,

and the mean amount of bark harvested per individual

was 1.09 m- (Twine 2004).

In KZN, the species is classed as declining and consid-

ered to be heavily exploited for bark products (Cunning-

ham 1988; Netshiluvhi 1999; Grace 2002). Its legal sta-

tus in KZN is described as ‘controlled’ by Von Ahlefeldt

et al. (2003), i.e. written permission is required from

the land owner/holder for this species to be harvested

or collected from the wild. The turnover from 23 trad-

ers in the Isipingo and Victoria Street informal herbal

medicine markets in Durban was estimated to be 187

fifty kg bags per annum (± 1995) (Netshiluvhi 1999).

On the Witwatersrand, 56% of the muti shops sold the

bark (Williams et al. 2001), and a volume equivalent to

= seven 50 kg bags were present between 17 of the 100

traders surveyed in the Faraday Street traditional medi-

cine market in Johannesburg in January 2001 (Williams

2003). On the western boundary of the Kruger National

Park, 29% of the vendors sold Balanites maughamii bark

and considered it a readily available resource (Botha et

al. 2001). The mean price per 50 kg bag of B. maughamii

bark bought by muti shops in Johannesburg in 1 995 was

* School of Animal, Plant and Environmental Sciences, University of

the Witwatersrand, PO Wits, 2050.

t Corresponding author e-mail address: vivwill(^planetac.co.za

MS. received: 2006-07-27.

R66.70 ± R33.50 [± standard deviation (SD)] (n = 15),

and in 2001 a bag cost ~ RIOO.

As part of an extensive investigation into the rela-

tionship between tree size and bark thickness of six tree

species, including Balanites maughamii'. 1, to determine

the size of trees targeted by commercial bark harvest-

FIGURE 1 . — Balanites maughamii individual repeatedly harvested for bark

on communal land in the Ingwavuma region of KwaZulu-Natal in

1998. Parts of stem, including buttresses have been removed.

212

Bothalia 37,2 (2007)

ers from the thickness of the bark sold in the miiti mar-

kets; 2, the mean wet and oven-dry bark thickness per

tree size class; and 3, the mean harvestable bark mass

per stem (Williams et al. 2005, in press a, b), various

aspects of the tree stem profile were measured. These

aspects included: 1, approximate height of the tree and

branch-free bole length; and 2, diameter of the stem at

five height intervals. Bark thickness was also measured.

Data collected for B. maughamii are a subset of the origi-

nal study. This short communication describes specific

aspects of the B. maughamii tree stem profile related to

the fluted trunk, including: 1, the relationship between

two measurements around the stem that respectively

include and exclude the bark surface area contained in

the convolutions of the flutes; 2, the number of flutes

observed at 1.3 m above ground (where dbh is nonnally

measured); and 3, the percentage of the stem enclosed

within the flutes.

METHODS

Between March and May 1998, 39 Balanites maugh-

amii stems were measured at six sites in three South

African provinces (Table 1). At each sample site, a

population of trees was located and individuals were

selected from five stem diameter classes based on diam-

eter at breast height (dbh) ranging between 10 cm and

60 cm. A minimum of five and a maximum of ten trees

were measured per diameter class (not per site). None of

the individuals had suffered any prior harvesting dam-

age, and the bark on the bole was intact. Balanites indi-

viduals larger than 60 cm dbh were found in communal

lands; however, these trees were not sampled as bark

harvesters had previously removed whole sections of the

bark, fluted stems and buttresses. The method used for

assessing vertical height was a direct estimate using a 2

m height pole, with 0.5 m intervals. The number of pole

lengths was counted by eye to estimate tree height and

branch-free bole length. After the 22 Balanites stems

were measured, the number of flutes at dbh were counted

and their depth was categorized (subjectively, shallow or

deep).

It is standard practise in forestry to measure tree stem

girth with a forestry diameter tape. The tape is calibrated

in n centimetres so that a circumference measurement

is converted directly to a diameter measurement (Philip

1983), and the measurement is thus recorded as a diam-

eter dimension rather than a circumference. Two diam-

eter readings were taken at five height intervals (0.5

m, 1.0 m, 1.3 m, 1.5 m and 2.0 m, abbreviated as

TABLE 1. — Sample sites and no. individuals sampled per site

FIGURE 2. — Schematic representation of a cross section through

Balanites maughamii tree stem showing measurements D1 and

D2. Dl; measurement around stem that excludes bark surface

in flutes; D2: measurement in concave convolutions of flutes,

thereby measuring entire bark surface. Measurements were made

using a forestry diameter tape, calibrated in ti centimetres, which

converts a circumference measurement of a stem directly into a

‘diameter’ measurement. Measurements of Dl at 1.3 m above

ground (dbh) were used to construct stem diameter classes, s,

subjective classification of shallow flutes; d, deep flutes.

D|p, D|j, and D,p respectively) from the Balanites

maughamii stem: 1, a circumference measurement

around the stem that excludes the area inside the flutes

(diameter 1, Dl); and 2, a circumference measurement

into the convolutions of the flutes, measuring along the

entire bark surface (diameter 2, D2) (Figure 2). Hence

Dl is the typical stem diameter measurement taken by

foresters, usually at breast height (1.3 m, dbh), whereas

D2 is a hypothetical diameter, where the flutes are

pushed out to fonn a circle. Initially, only Dl was mea-

sured, but after six samples, D2 was also measured.

RESULTS AND DISCUSSION

The Balanites maughamii individuals measured,

ranged in height from 4 to 1 2 m, with a mean of 8 ± 2 m

(SD). Branch-free bole length was 2.9 ± 1.4 m (SD). The

dbh of the largest tree sampled was 01,^ = 59.2 cm and

D2| 3 = 260.0 cm (circumference equals 186 cm and 817

cm respectively), from a site in a private protected area

in KZN.

There was a very strong positive relationship between

Dl and D2 at all height intervals 'up the stem (Figure

3A-E), especially at D^^ (r* = 0.988, p < 0.0001, Figure

3A). No branching occurred on the stem below 0.9 m,

hence results for D^^ were not affected by the response

of the tree to branching. The quadratic regressions

were only slightly better fits than the linear regressions

(results not shown). For example, r" = 0.988 for the qua-

dratic equation at D^^, whereas r^ = 0.979 for the linear

equation at the same height.

These results show that by measuring Dl at a par-

ticular stem height, D2 can be accurately estimated,

hence obviating the necessity to measure both Dl and

D2. When compared with the observed D2, the D2 pre-

dicted by the quadratic regression equations was slightly

Bothalia 37,2 (2007)

213

D1o5(cm) D1,o(cm)

D1v3(cm)

D1i 5(cm)

FIGURE 3. — Relationship between stem diameter 1 (Dl)and stem diameter 2 (D2) measured at A, 0.5 m; B, 1.0 m; C, 1.3 m; D, 1.5 m; and E, 2.0

m above ground [n = 28 (all graphs)]. To obtain circumference, multiply D1 or D2 by tc.

overestimates [mean percentage error = 0.35 ± 5.99%

(SD), n = 154]. By contrast, the linear regression equa-

tions tended to underestimate the predicted D2 [-1.22 ±

9.04%(SD), n= 154].

By converting the observed D2 measurements back to

circumferences, the area of bark (m^) on the stem could

be estimated. The mean amount of bark up to 2 m on the

stem ranged from 3.3 ± 0.6 m^ (SD) (n = 10) on trees in

the 10-19 cm diameter class (Dl), to 16.1 ± 0.7 m^ (SD)

(n = 4) on trees in the 50-59 cm diameter class (Table

2).

As the dbh of Balanites maiighamii individuals in-

creased, the number and depth of flutes at Dj ^ was

observed to increase (Table 3), thus increasing the pro-

portion of the bark surface area within the convolu-

tions of the flutes. Trees in the 10-20 cm and 20-30 cm

stem diameter classes (Dl) generally had two shallow

flutes and one deep one. As tree size increased, the shal-

low flutes became deeper until there were 2 or 3 and 4

or 5 deep flutes in the 30^0 cm and 40-50 cm classes

respectively. Trees larger than 50 cm had more than six

deep flutes with sometimes as many as 10 per stem as

the trees approached 60 cm dbh.

214

Bothalia 37,2 (2007)

TABLE 2. — Estimated mean bark area (m^) up to 2 m height per stem

size class

dbh, diameter at breast height; SD, standard deviation.

Most of the trunk circumference is contained within

the concave sections of the flutes (Figure 4). At 0.5 m

above ground, 73.0 ± 4.0% (SD) of the stem was within

the flutes. The percentage decreased gradually with

increasing height up the stem until it was 70.3 ± 4.3%

(SD) at 2 m (Figure 4). Furthenuore, in trees with larger

dbh, a greater percentage of stem was enclosed within

the flutes. Similarly, more than two thirds of the bark

surface area is within the flutes [mean = 72 ± 3% (SD), n

= 31]. The proportion of the bark inside the flutes varied

according to tree size, with up to 79% of the bark area

found in the flutes of trees in the 50-59 cm stem diam-

eter class (Dl), and decreasing to 69% in flutes of trees

in the 10-19 cm stem diameter class.

CONCLUSION

Despite the buttresses in the Balanites maughamii

stems, it appears that the diameter measurement Dl is an

acceptable predictor of D2. Hence, the bark surface area

can be estimated as well as the amount of bark that can

potentially be removed from the stems. Because most

of the bark area is contained within the convolutions of

the flutes, the tree trunks are difficult to ring-bark. Even

when harvesters remove whole sections of the flutes/but-

tresses, including the timber, they usually leave behind

some of the bark at the base of the flute. This may poten-

tially enable wound recovery following harvesting and

probably makes the species more resilient to harvesting.

ACKNOWLEDGEMENTS

Thanks are due to Megan Whelan for her field

assistance; the Eddie Young Memorial Bursary of the

Endangered Wildlife Trust and the National Research

Foundation for funding.

TABLE 3. — Observed number and depth of flutes per measured tree at

D| j(dbh). Individual trees are enclosed in parentheses in column

three

Size-class No. trees measured Observed number and depth of flutes

(dbh, cm) (n = 15, out of 39) per tree at (dbh)

(Dl)

depth).

FIGURE 4. — Mean percentage of stem circumference contained within

concave sections of flutes at five height intervals up stem to 2 m,

including diameter at breast height (dbh, 1.3 m). Means calcu-

lated for trees ranging from 11.7 cm to 52.7 cm dbh. SD, stan-

dard deviation.

REFERENCES

BOTHA, J., WITKOWSKI, E.T.F. & SHACKLETON, C.M. 2001. An

inventoiy of medicinal plants traded on the western boundary of

the Kruger National Park, South Africa. Koedoe 44: 7-46.

CUNNINGHAM, A.B. 1988. An investigation of the herbal medicine

trade In Natal/KwaZulii. Investigational Report No. 29. Institute

of Natural Resources, Pietermaritzburg, South Africa.

GRACE, O.M. 2002. Bark in traditional healthcare in KwaZulu-Natal.

South Africa: usage, authentication and sustainability. M.Sc.

thesis, University of Natal, Pietermaritzburg.

NETSHILUVHI, T.R. 1999. Demand, propagation and seedling estab-

lishment of selected medicinal trees. South Afi-ican Journal of

Botany 65: 331-338.

PHILIP, M.S. 1983. Measuring trees and forests. Aberdeen University

Press, UK.

POOLEY.E.S. 1993. The complete field guide to trees of Natal, Zululand

& Transkei. Natal Flora Publications Trust, Durban.

TWfNE. W. 2004. Medicinal bark harvesting and yields in woodlands:

a case study from southern Maputaland. In M.J. Lawes, H.A.C.

Eeley, C.M. Shackleton & B.G.S. Geach, Indigenous forests and

woodlands in South Africa: policy, people and practice: 533-

537. University of Natal Press, Pietermaritzburg.

VAN WYK, A.E. & VAN WYK, P. 1997. Field guide to trees of south-

ern Africa. Struik. Cape Town.

VON AHLEFELDT, D., CROUCH, N.R., NICHOLS, G., SYMMONDS,

R., MCKEAN, S„ SIBIYA, H. & CELE, M.P. 2003. Medicinal

plants traded on South Africa’s eastern seaboard. Porcupine

Press, Durban.

WILLIAMS, V.L. 2003. Hawkers of health: an investigation of the

Faraday Street traditional medicine market in Johannesburg,

Gauteng. Plant Ecology and Conservation Series No 15. (Report

to Gauteng Directorate of Nature Conservation, DACEL).

University of the Witwatersrand, Johannesburg.

WILLIAMS, V.L., BALKWILL, K. & WITKOWSKI, E.T.F. 2001.

A lexicon of plants traded in the Witwatersrand umuthi shops.

South Africa. Bothalia 31 : 71-98.

WILLIAMS, V.L., BALKWILL, K. & WITKOWSKI, E.T.F. in press

a. Bark mass estimates for six tree species used medicinally in

South Africa. Advances in Economic Botany.

WILLIAMS, V.L., WITKOWSKI, E.T.F. & BALKWILL, K. 2005.

Height, branch-free bole length and bark thickness for six tree

species used medicinally in South Africa. Koedoe 48: 57-65.

WILLIAMS, V.L., WITKOWSKI, E.T.F. & BALKWILL, K. in press

b. Relationship between bark thickness and diameter at breast

height for six tree species used medicinally in South Africa.

South African Journal of Botany.

Bothalia 37,2: 215-248 (2007)

Invasive, naturalized and casual alien plants in southern Africa: a sum-

mary based on the Southern African Plant Invaders Atlas (SAPIA)

L. HENDERSON*

Keywords: biomes, casual alien plants, invasive plants, Lesotho, naturalized plants, roadside surveys, SAPIA mapping project. South Africa,

Swaziland

ABSTRACT

The primary objective of this publication is to provide an overview of the species identity, invasion status, geographical

extent, and abundance of alien plants in South Africa, Swaziland and Lesotho, based on field records from 1979 to the

end of 2000. The dataset is all the species records for the study area in the Southern African Plant Invaders Atlas (SAPIA)

database during this time period. A total of 548 naturalized and casual alien plant species were catalogued and invasion was

recorded almost throughout the study area. Most invasion, in terms of both species numbers and total species abundance, was

recorded along the southern, southwestern and eastern coastal belts and in the adjacent interior. This area includes the whole

of the Fynbos and Forest Biomes, and the moister eastern parts of the Grassland and Savanna Biomes. This study reinforces

previous studies that the Fynbos Biome is the most extensively invaded vegetation type in South Africa but it also shows

that parts of Savanna and Grassland are as heavily invaded as parts of the Fynbos. The Fabaceae is prominent in all biomes

and Acacia with 1 7 listed species, accounts for a very large proportion of all invasion. Acacia mearnsii was by far the most

prominent invasive species in the study area, followed by A. saligna, Lantana camara, A. cyclops, Opimtia ficus-indica,

Solanum mauritiamim, Popultis albaMcanescens, Melia azedarach, A. dealbata and species of Prosopis.

INTRODUCTION

History of roadside surveys in South Africa

Roadside surveys of invasive plants in South Africa

were pioneered by Henderson and Musil (nee Duggan)

starting in 1979 in the central Transvaal, now Gauteng

(Wells, Duggan & Henderson 1980), with the remainder

of the Transvaal surveyed in 1982 and 1983 (Henderson

& Musil 1984). Surveys of the rest of South Africa were

conducted by Henderson from 1986, starting with Natal

(Henderson 1989), followed by the Orange Free State

(Henderson 1991a), northern Cape (Henderson 1991b),

eastern Cape (Henderson 1992), western and central

Cape (completed in 1993 but unpublished), and southern

and southwestern Cape (Henderson 1998a).

All terminology used in this paper relating to invasive

plants such as ‘alien’, ‘invasive’, ‘naturalized’, ‘casual

alien’, ‘weed’ and ‘environmental weed’ conforms, as

far as possible, to the definitions provided by Richardson

et al. (2000) and Pysek et al. (2004). The method used

in these surveys was designed initially to make use of

otherwise unproductive travelling time whilst engaged

in other research projects. The method was refined as

the surveys progressed until a standardized method was

developed (see Henderson 1992, 1998a). The presence

and abundance of all alien trees, large shrubs and con-

spicuous climbers which appeared to be naturalized or

occurring outside of cultivation were recorded for each

veld type category, habitat type (roadsides and adjoining

veld, and streambanks) and quarter-degree/fifteen minute

square traversed by road.

Recordings of species on roadsides and in the adja-

cent veld were made from a moving vehicle along road

* Agricultural Research Council: Plant Protection Research Instimte.

c/o SANBI. Private Bag XI 01, 0001 Pretoria.

E-mail: henderson(g!sanbi.org

MS. received: 2006-10-06.

transects of between five and 10 km long. Recordings

of streambank species were made at virtually all water-

course crossings on the survey route.

The Southern African Plant Invaders Atlas mapping

project (SAPIA)

The Southern African Plant Invaders Atlas (SAPIA) is

a mapping project, launched in January 1994, to collate

information on the distribution, abundance and habitat

types of invasive and naturalized alien plants in southern

Africa (Henderson 1998b). The first phase of SAPIA,

involving volunteer participants, was scheduled for a

five-year period, ending in December 1998. The atlas

region covered South Africa, Lesotho and Swaziland.

Infonnation was recorded on two standardized atlas

sheets, with slightly different species lists, covering the

western and eastern halves of the atlas region. One hun-

dred plant taxa were listed on each sheet, with a com-

bined total of 161 species. A pocket field guide was com-

piled to help with the identification of all listed species

(Henderson 1995).

SAPIA database

A computerized SAPIA database was created by

incorporating all Henderson survey data (± 23 000

records) and SAPIA phase one project data (± 20 000

records). The SAPIA project continued on an ad hoc

basis and by the end of 2000 a total of ± 48 000 records

had been accumulated. Thereafter, the SAPIA initiative

dwindled due to lack of funding. Only 10 000 records

were added in the five year period from 2001 to the

end of 2005. The SAPIA project was revived in 2006

with funding from the Department of Water Affairs and

Forestry’s Working for Water Programme. The SAPIA

database has been computerized using Microsoft Access

and is housed at the Plant Protection Research Institute

in Pretoria.

216

Bothalia 37,2 (2007)

Objectives of this study

• To provide an overview of the species identity, inva-

sion status, geographical extent, and abundance of

alien plants in South Africa, Swaziland and Lesotho,

based on field records from 1979 to the end of 2000.

• To highlight the most prominent invaders in the

region as a whole, in each of the biomes, and in

riparian and wetland habitats.

• To compare invasion and provide species profiles for

each of the biomes.

METHODS

Sampling method

The dataset for this study is all the species records for

South Africa, Swaziland and Lesotho in the SAPIA data-

base collected from 1979 until the end of 2000 (± 48 000

records). During this period a concerted effort was made

to gather as much data from as wide an area as possi-

ble. The information gathered is the best available data

concerning the extent of invasion and species composi-

tion, at least of the larger trees, shrubs and conspicuous

climbers, in the study area over this time period.

The SAPIA dataset was subdivided on a quarter-

degree square (QDS) basis into six datasets representing

the biomes of southern Africa. According to Rutherford

(1997) there are seven biomes in southern Africa: Savanna,

Fynbos, Forest, Grassland, Nama-Karoo, Succulent Karoo

and Desert. The Forest Biome in southern Africa is min-

iscule, only occurring in the Knysna area. However, if

all the forest patches elsewhere are included, its area

increases several-fold (Rutherford 1997). In this study

Forest refers to the Forest Biome and also forest habitats

within the Savanna, Fynbos and Grassland Biomes. The

Desert Biome occurs almost exclusively in Namibia,

except for a very small patch along the Orange River

bordering on South Africa that has been excluded from

this study.

Data treatment

Abundance

Species abundance ratings in the SAPIA database are

qualitative estimates. Table 1 shows the abundance rat-

ings used in the SAPIA database and the equivalent rat-

ing used in Henderson surveys. For the purposes of this

study, species abundance ratings were converted to a

numerical value as done in previous surveys (Henderson

1998a) and each abundance rating was expressed in

numbers of individuals or groups per 10 km transect/

recording (Table 1).

Prominence

A similar formula was used in this study to calcu-

late prominence as in previous studies by Henderson

(1998a). The prominence value of a species x in category

y (biome or study area) was calculated as follows:

total abundance of species x in categoty y

X 100

sum of the abundances of all species in category y

prominence

value

total species records of species x in category y

X 100

sum of the records of all species in category y

The highest prominence values in a given category

which add up to ± 160 points out of a total of 200 are

printed in bold in Appendices 1-3. The cut-off point is

arbitrary but represents the upper 80% of the summed

prominence values.

RESULTS

A total of 548 naturalized and casual alien plant spe-

cies were catalogued in the SAPIA database for South

Africa, Swaziland and Lesotho from 1979 to the end of

2000 (Appendix 4). At least 119, mainly herbaceous,

taxa are considered to have been under-recorded and

TABLE 1 . — Abundance ratings used in Henderson surveys, SAPIA and this study

t, very abundant extensive stands; abundant; many clumps or stands: frequent, many sightings of single plants or small groups: occasional, a few

sightings of one or a few plants: rare, one sighting of one or a few plants.

#, weighted abundance, numbers of individuals or groups per 1 0 km transect/recording.

Bothalia 37,2 (2007)

217

the results presented are not a true reflection of their sta-

tus (see asterisked species in Appendix 4). A further 45

species were recorded in the study area after 2000 and

are asterisked in the species checklist (Appendix 5). A

total of 601 species are listed in the full checklist given

in Appendix 5 — this is estimated to be about half the

total number of naturalized and casual alien plant spe-

cies in southern Africa. The most comprehensive listing

of naturalized species in southern Africa, compiled by

Wells et al. (1986), contains approximately 965 species,

predominantly herbaceous. The SAPIA database, with a

bias towards trees and shrubs, has an additional 231 spe-

cies not listed by Wells et al. (1986).

Geographical extent of invasion

Alien plant invasion was recorded almost through-

out the study area. Figure lA shows invasion in terms

of species numbers per QDS and Figure IB shows

the severity of invasion per QDS based on the total

weighted abundance of all species per QDS. Most inva-

sion, in terms of both species numbers and total species

abundance, was recorded along the southern, southwest-

ern and eastern coastal belts and in the adjacent interior.

This corresponds with the regions of highest rainfall

(Schulze 1997), urban development, and cultivation of

agricultural and silvicultural crops. It also includes the

whole of the Fynbos and Forest, and the moister east-

ern parts of the Grassland and Savanna Biomes (Figure

1C). Distribution maps of 234 species, which include all

declared species under the Conservation of Agricultural

Resources Act, Act 43 of 1983, and amended in 2001,

are given in the field guide Alien weeds and invasive

plants (Henderson 2001).

Prominent invasive species

There were 97 prominent invasive species in the

study area and each of the biomes (Appendices 1-3).

All these species were invading natural and semi-natu-

ral habitats.

Study area

Fifty species account for most invasion (the upper

80% of the summed prominence values) in the study

area (Appendix 1). Acacia mearnsii (black wattle) was

the most prominent species by far, with a value of 18.37

(out of a maximum of 200) which is more than double

the value of the second-ranked species, A. saligna (Port

Jackson). The remaining top ten most prominent invad-

ers in the study area were in order, Lantana camara

(lantana), A. cyclops (rooikrans), Opuntia ficus-indica

(sweet prickly-pear), Solanum mauritianum (bugweed),

Populus alba/xcanescens (white/grey poplars — values

of these two taxa were combined where they were diffi-

cult to distinguish at a distance during roadside surveys),

Melia azedarach (seringa), A. dealbata (silver wattle)

and Prosopis spp. (P glandulosa var. torreyana, P. velu-

tina and their hybrids)(mesquite trees). Together these

species cover almost the entire study area (Figures 2, 3).

Savanna Biome

Forty-eight species were the most prominent invaders

in the Savanna Biome (Appendix 2). Lantana camara

FIGURE 1. — A, species numbers per quarter-degree square in study

area; B, severity of invasion per quarter-degree square. Light

invasion: < 1 individual or group per km. Moderate invasion: up

to 5 individuals or groups per km; some species forming stands.

Heavy invasion: up to 50 individuals or groups per km; many

species forming stands; some completely dominating landscape.

C, heavy invasion in relation to biomes in study area.

was the most prominent species with a prominence value

of 20.6, followed by Chromolaena odorata (triffid weed)

with a value of 14.2 and Melia azedarach with a value

of 12. The remaining top ten invaders were, in order,

Solanum mauritianum. Acacia mearnsii, Opuntia ficus-

indica, Ricinus communis (castor-oil plant), Psidium

218

Bothalia 37,2 (2007)

FIGURE 2. — Distribution and severity of invasion in study area: A, Acacia cyctops\ B, Acacia dealbata', C, Acacia nieanisii\ D, Acacia saligna; E,

Lantana cainara; F, Melia azedarach. Light invasion. A; moderate invasion. A; heavy invasion, ■.

guajava (guava), Eichhornia crassipes (water hyacinth)

and Jacaranda mimosifolia (jacaranda).

Fynhos Biome

Twenty species were the most prominent invaders in

the Fynbos Biome (Appendix 2). Acacia mearnsii was

the most prominent species with a prominence value of

31.5, followed by A. saligna and A. cyclops with val-

ues of 30.4 and 27.2, respectively. The remaining top

ten most prominent invaders in order, were, Pinus pin-

aster (cluster pine). Acacia melanoxylon (Australian

blackwood), A. longifolia (long-leaved wattle), Popuhis

xcanescens (grey poplar), Paraserianthes lophantha

(stinkbean), Riibiis friiticosiis (European blackbeiTy) and

Opuntia ficus-indica. Plakea sericea (silky hakea) and

Pinus radiata (radiata pine), both invaders of mountain

fynbos, were most likely under-recorded because of the

inacessibility and under-sampling of this habitat.

Forest habitats

Forty species were the most prominent invaders in

forest habitats (Appendix 2). Chromolaena odorata was

Bothalia 37,2 (2007)

219

FIGURE 3. — Distribution and severity of invasion in study area: A, Opiintia ficus-indica; B, Popiihis albaMcanescens', C, Prosopis spp.; D,

Solamim mauntianum. Light invasion, A; moderate invasion. A; heavy invasion, ■.

the most prominent species with a prominence value

of 23.9, followed by Solamim mauritiamim and Acacia

mearnsii with values of 19 and 16.7, respectively. The

remaining top ten prominent invaders were, in order.

Acacia melanoxylon, Lantana camara, Cestrum laevi-

gatiim (inkberry), Caesalpinia decapetala (Mauritius/

Mysore thorn), Melia azedarach, Finns pinaster and

Psidium gnajava. Pereskia acnleata (pereskia) ranked

eleventh and could have been vastly underestimated

because of the difficulty of observing this forest canopy,

climbing species.

Grassland Biome

Thirty-two species were the most prominent invaders

in the Grassland Biome (Appendix 3). Acacia mearnsii

was the most'prominent species with a prominence value

of 21.3, followed by A. dealbata and Salix babylonica

(weeping willow) with values of 20.9 and 17.3, respec-

tively. The remaining top ten most prominent invaders

were, in order, Popuhis alba/xcanescens (white/grey

poplars), Solamim mauritiamim, Riibiis spp. (mainly R.

ciineifoliiis){hramh\es), Pyracantha angiistifolia and P.

creniilata (yellow and Himalayan firethoms), Eucalyptus

spp. (eucalypts), Melia azedarach and Opiintia ficiis-

indica. Campulocliniiim macrocephalum (pompom

weed) which did not feature as a prominent invader in

this study showed an explosive rate of increase after

2000 and currently would be rated as one of the most

prominent invaders in the Grassland Biome (Henderson

et al. 2003).

Nama-Karoo Biome

Fourteen species were the most prominent invaders in

the Nama-Karoo Biome (Appendix 3). Prosopis spp. (P.

glandulosa var. torreyana, P veliitina and their hybrids)

were the most prominent species with a prominence

value of 60.6, followed by Atriplex inflata (sponge-fruit

saltbush) and Opuntia ficiis-indica with values of 21 and

14 respectively. The remaining top ten prominent invad-

ers were, in order, Salsola kali/tragus (Russian tumble-

weed), Azolla filiculoides (red water fern), Nicotiana

glaiica (wild tobacco), Atriplex nummularia (old man

saltbush), Schinus mode (pepper tree). Agave americana

(American agave) and Solamim elaeagnifolium (silver-

leaf bitter-apple).

Succulent Karoo Biome

Twelve species were the most prominent invaders in

the Succulent Karoo Biome (Appendix 3). Nicotiana

glauca was the most prominent invader with a promi-

nence value of 26.8, followed by Acacia cyclops and

Prosopis spp. (P. glandulosa var. torreyana, P. veliitina

and their hybrids) with values of 26.3 and 25.9, respec-

220

Bothalia 37,2 (2007)

tively. The remaining top ten most prominent invaders

were, in order, Acacia mearnsii, A. saligna, Atriplex

inflata, Arundo donax (giant reed), Atriplex monmularia,

Opuntia ficus-indica and Popuhis xcanescens.

Riparian and wetland habitats

Fifty-five species had more than 50 records in ripar-

ian and wetland habitats (Appendix 4). Salix babylonica

was the most frequently recorded riparian and wetland

species with 1 323 records, followed by Populus alba/

xcanescens with 1 176 records Acacia mearnsii with

953 records. The remaining top ten riparian and wetland

invaders were, in order, Melia azedarach, Riciniis com-

munis, Arundo donax. Acacia dealbata, Sesbania puni-

cea (red sesbania), Prosopis spp. and Nicotiana glauca.

Biome comparison

The Savanna Biome, which occupies the largest

number of QDS (645) in the study area, had the great-

est number of species (358) and the most invasion in

tenns of total abundance of all species (Table 2). The

Fynbos Biome, however, which occupies the least QDS

(139), was the most heavily invaded in terms of average

abundance of all species per QDS, average abundance of

individual species per QDS and % QDS heavily invaded.

The Grassland Biome ranks third after Fynbos for total

abundance of all species, followed by Forest, Nama-

Karoo and the Succulent Karoo Biome was the least

invaded.

Biome profiles

Appendix 6 provides species characteristics of the

prominent invasive species. Table 3 analyses the promi-

TABLE 2. — Biome comparison in terms of extent, numbers and abun-

dance of species and severity of invasion

QDS, quarter-degree squares in Fynbos, Savanna, Grassland, Nama-

Karoo and Succulent Karoo according to Rutherford (1997); QDS in

forest habitats according to SAPIA database.

*, total weighted abundanee of all species (see text).

Prominent invasive species: species with highest prominence values

adding up to ± upper 80% of summed values (see text).

#, % QDS lightly invaded: less than I individual or group per km; #,

% QDS moderately invaded: up to 5 individuals or groups per km;

some species forming stands; #, % QDS heavily invaded: up to 50

individuals or groups per km; many species fonning stands; some

completely dominating landscape.

FB, Fynbos Biome; Fh, Forest habitats; SB, Savanna Biome; GB,

Grassland Biome; NKB, Nama-Karoo Biome; SKB, Succulent

Karoo Biome,

nent invasive species in each of the biomes and the study

area in tenns of region of origin, taxonomy, growth

fonn, perennation, type of reproduction, dispersal mecha-

nism and cultivated use.

Savanna Biome speeies are predominantly of tropical

origin; members of the Fabaceae, Solanaceae, Asteraceae

and Rosaceae; woody trees and shrubs, followed by

herbs and climbers; perennial evergreen and evergreen/

deciduous; seed-producers; water and bird dispersed;

ornamentals and agricultural crops.

Fynbos Biome species are predominantly of temperate

origin (particularly southern temperate); members of the

Fabaeeae, Myrtaceae, Pinaceae and Salicaceae; woody

trees and shrubs; perennial evergreen; seed-producers;

water, bird and wind dispersed; silvicultural crops, orna-

mentals and cover/binders.

Forest habitat species are predominantly of tropical

origin; members of the Fabaceae, Asteraceae, Myrtaceae,

Solanaceae, Pinaceae and Zingiberaceae; woody trees

and shrubs, followed by herbs and climbers; perennial

evergreen; seed-producers; bird and water dispersed;

ornamentals, barriers and silvicultural crops.

Grassland Biome species are predominantly of north-

ern temperate origin and the tropics; members of the

Rosaceae, Fabaceae and Salieaeeae; woody trees and

shrubs, followed by herbs; perennial evergreen/decidu-

ous and deciduous; seed-producers, but a greater per-

centage of species coppice and sucker than in other veg-

etation categories; water and bird dispersed; barriers,

ornamentals and agricultural crops.

Nama-Karoo Biome species are predominantly of

northern temperate origin and the tropics; members of

the Chenopodiaceae, Salicaceae, Cactaceae, Fabaceae,

Solanaceae and Tamaricaceae; woody trees and shrubs,

followed by herbs and succulent trees and shrubs; peren-

nial evergreen/deciduous and deciduous; seed-producers,

but a greater percentage of species reproduce by vegeta-

tive division than in other vegetation categories; water

and wind dispersed; agricultural crops and ornamentals.

Succulent Karoo Biome species are predominantly of

temperate origin; members of the Fabaceae, Chenopo-

diaceae and Tamaricaceae; woody trees and shrubs;

perennial evergreen and evergreen/deciduous; seed-pro-

ducers and reproduce vegetatively by coppicing; water

and wind dispersed; agricultural crops, ornamentals and

cover/binders.

DISCUSSION

Biome comparison: extent of invasion

No previous studies have enabled a direct comparison

of the extent of invasion in the different biomes using the

same parameters. This study reinforces previous studies

that the Fynbos Biome is the most extensively invaded

vegetation type in South Africa (Richardson et al. 1997)

but it also shows that parts of Savanna and Grassland are

Bothalia 37,2 (2007)

221

TABLE 3. — Analysis of region of origin, taxonomy, growth forms, perennation, reproduction, dispersal mechanisms and cultivated uses of promi-

nent invasive species in each of the biomes, forest habitats and study area

222

Bothalia 37,2 (2007)

as heavily invaded as parts of the Fynbos. These find-

ings have important implications for the management

of alien plant invasions in South Africa. Without inter-

vention we can expect invasion to increase in all parts

of South Africa and particularly in the Grassland and

Savanna Biomes where large areas are yet to be invaded

and many species are only starting to invade.

Biome comparison: prominent invaders

Each biome has a different suite of prominent invad-

ers. In part, this can be explained by their pre-adap-

tion to the prevailing environmental conditions, but

also to their history of planting. Most of these species

were deliberately introduced and cultivated on a grand

scale as silvicultural and agricultural crops e.g. Acacia

mearnsii, A. melanoxylon, Finns pinaster and species of

Prosopis, as barriers e.g. Acacia dealbata, Hakea seri-

cea and Pyracantha angiistifolia, as cover/binders e.g.

Acacia cyclops, A. saligna and Popidus ^canescens, and

ornamentals e.g. Melia azedarach and Lantana carnara.

Some species which have become prominent invad-

ers were not cultivated widely or on a grand scale e.g.

Solanum mauritianwn, Chromolaena odorata and

Nicotiana glaiica. Although the latter species have on

occasion been cultivated as ornamentals they have man-

aged to disperse very efficiently without human assis-

tance— C. odorata by wind, S. mauritianum by birds

and N. glaiica by wind, soil and water.

Some species, although widely planted, have become

prominent invaders in only one biome, indicating that

environmental factors have limited their distribution.

Examples are members of the family Rosaceae, such

as Pyracantha angiistifolia, P crenulata, Cotoneaster

franchetii and C. pannosiis that are virtually restricted

to high-altitude grasslands where it appears that freez-

ing winter temperatures are needed to trigger seed ger-

mination (Henderson 1989). Jacaranda mimosifolia is

another species that has been planted throughout South

Africa yet is only invasive in the moister parts of the

Savanna and Forest Biomes. In its native northeastern

Argentina, J. mimosifolia occurs mainly on river banks

under wanner-temperate, subhumid conditions (Poynton

1973) — environmental conditions which are similar to

those in its naturalized range in southern Africa. A pre-

vious study by Henderson (2006b) showed that the cur-

rent distributions of invasive plants in southern Africa

are a reflection of the climatic zones of their origin.

There are considerable differences in the species pro-

files of the biomes but shared features are the promi-

nence of the family Fabaceae, woody trees and shrubs,

reproduction by seed and water dispersal. Within the

Fabaceae the Acacia species are the most numerous

with 1 7 listed species and account for a very large pro-

portion of all plant invasion in South Africa. They are

important invaders of all the major vegetation types

except for those in the arid interior, where other legu-

minous invaders take over, namely species of Prosopis.

The most widespread and abundant acacias are Acacia

mearnsii, A. cyclops and A. saligna. Acacia mearnsii has

invaded the widest range of vegetation types in South

Africa and is the most widespread riverine invader,

occurring almost continuously from Louis Trichardt

in the Limpopo Province down the eastern seaboard to

Cape Town, a distance of ± 2 500 km. Acacia cyclops

stretches along almost the entire Cape coastline from

Port Nolloth in the northwest to beyond East London in

the east, a distance exceeding 2 000 km. Acacia saligna

stretches along the Cape coastline from Saldanha Bay

in the west to the Kei River in the east, a distance of ±

1 500 km.

Sixty-eight per cent of prominent invaders are peren-

nial trees or shrubs. There are only two grasses listed as

prominent invaders and only 14 species as nonperennial

(annual, biennial or variable). Grasses and herbaceous

species are under-represented in the SAPIA database

largely as a consequence of biassed recording of the

larger, more conspicuous species. In southern Africa

the Poaceae is one of the largest plant families with

847 indigenous species and 115 (12%) naturalized spe-

cies (Gibbs Russell et al. 1990). However, only 30 grass

species are listed in this publication. There is definitely

a lack of expertise in identifying grasses in South Africa

and this is one of the reasons for the under-represen-

tation of alien grasses in weed surveys. There is simi-

larly an under-representation of the alien herbaceous

Asteraceae. The South African National Biodiversity

Institute’s online species checklist at http://posa.sanbi.

org/searchspp.php lists 125 alien herbaceous species in

South Africa, yet only 44 alien herbaceous species have

been listed in this publication.

Comparison with other studies

Versfeld et al. (1998) provide the only other assess-

ment of the extent and importance of invasive plants on

a national level. This study combined expert knowledge

of local landowners and managers with existing data-

bases such as those of provincial conservation authori-

ties and national departments. The SAPIA database

was used as a means of data verification particularly for

areas where expert knowledge was lacking. Overall the

assessment by Versfeld et al. (1998) relating to impor-

tance rankings and the distribution of dense infesta-

tions concurs with this study. Eight of the top ten invad-

ing species or groups of species, ranked by condensed

invaded area, also appear within the top ten ranking in

this study — these are: Acacia cyclops, Prosopis spp., A.

mearnsii, A. saligna, Solaniim mauritianum, Opuntia

spp., Melia azedarach and Lantana carnara. Versfeld

et al. (1998) include Pinus spp. and Hakea spp. within

the top ten ranking, whereas this study includes Popidus

albaMcanescens and Acacia dealbata. The lower rank-

ing of Pinus spp. and Hakea spp. in this study can be

explained by the under-sampling of mountain habitats,

which are largely inaccessible by road, in which these

species are invasive.

Abundance data presented in this study suggests that

Versfeld et al. (1998) may have underestimated the

area of invasion of Salix babylonica and Popidus alba/

xcanescens. In the present study these species were not

only the most frequently recorded invaders in riparian

and wetland habitats but their total weighted abundance

Bothalia 37,2 (2007)

223

was in both instances more than Melia azedarch and

Eucalyptus spp. (Appendix 1) which were rated above

Salix spp. and Populus spp. by Versfeld et al. (1998).

Other riparian species which may also have been under-

estimated include Arundo donax (giant reed). Morns

alba (common mulberry) and Ricinus communis.

Looking to the future

The Working for Water Programme (WfW) and biologi-

cal control

Alien plant invasion is a dynamic process and there

will undoubtedly be changes in species composi-

tion and prominence of invaders in the future. Many

of the large tree species — mainly Acacia, Eucalyptus,

Pinus, Populus, Prosopis species and Melia azedarach

have been targeted by a national clearing programme,

Working for Water (WfW), which started in October

1995 (Marais et al. 2004). To date there has not been

an assessment of the affects of the WfW programme on

the status of invasive alien infestations. The programme

has been proposed for 20 years but Marais et al. (2004)

indicate that even with the existing generous levels of

funding, it is unlikely that the problem will be contained

within the next half century.

Biological control of invasive plants using introduced

insects and pathogens is the only sustainable, effec-

tive and inexpensive solution to the most intractable of

the invasive alien plant problems (Marais et al. 2004).

When they are successful, the damage inflicted by bio-

logical control agents causes a decline in population

densities, distribution and, or, rates of spread of inva-

sive plants, and reduces the costs of other management

practices (Zimmermann et al. 2004). There have been

some outstanding successes with biocontrol in South

Africa, dating back to the early and mid- 1900s with

Opuntia monacantha (drooping prickly pear) and O.

ficus-indica, and in more recent years with several of

the Acacia spp. (Zimmermann et al. 2004). Population

monitoring of A. saligna in the Western Cape has shown

marked decreases in population densities caused by the

gall-forming rust fungus, Uromycladium tepperianum

(Morris 1997; Wood & Morris 2007).

New invaders

Since 2000 a further 45 species have been added to

the SAPIA database for the study area (Appendix 5).

Another eight species, two of which are indigenous to

South Africa, are naturalized in neighbouring Zimbabwe

and Malawi (Appendix 5). All but three of the additional

species have been listed as weeds in A global compen-

dium of weeds (Randall 2002) and 28 species are envi-

ronmental weeds elsewhere in the world and therefore

have the potential to become invasive in South Africa.

Fourteen of the new species are ‘noxious weeds’ or

restricted in California, Florida, Hawaii, New Zealand

and Australia — places with similar climates and with

which South Africa has many invasive species in com-

mon. We should be especially wary of these species

which include some of the most damaging and costly

invaders such as Hydrilla verticillata (hydrilla), a sub-

merged aquatic plant that has invaded much of the USA

since the 1960s, and Chondrilla juncea (skeleton weed),

a terrestrial herb that has become a major agricultural

weed in the USA, Canada and Australia.

CONCLUSION

The main objective of this paper was to provide a

historical overview of the extent and species composi-

tion of alien plant invasion in southern Africa from 1979

until the end of 2000. This snapshot of invasion will pro-

vide a yardstick by which we can measure our progress

or failure in the management of invasive alien plants in

southern Africa.

This publication will also contribute to the global

knowledge of invasive alien plants. One of the most

useful predictors of invasiveness is whether a species

is invasive elsewhere in the world (Richardson et al.

2004a). The lists of prominent invaders and other natu-

ralized species provided here will serve as a warning

to neighbouring countries and to those as far afield as

Australia, New Zealand and the USA of potentially inva-

sive species in their regions.

The results presented here are but a summary of the

more than 50 000 records of invasive alien plants in

the SAPIA database. Much more can be gleaned from

the SAPIA data. SAPIA has provided the raw data for

analyses that have been used to prioritize invasive alien

species for management (Robertson et al. 2003; Nel et

al. 2004), to map the potential spread of invasive plants

(Rouget et al. 2004), to look at broad-scale distribution

patterns of invasive species (Richardson et al. 2004b),

to correlate patterns of alien plant species richness

with the environment and indigenous species richness

(Richardson et al. 2005), to correlate patterns of invasion

with interactions between environment, species traits and

human uses (Thuiller et al. 2006) and to look at potential

range and residence time (Wilson et al. 2007). SAPIA

has also played a crucial role in providing infomiation

on invasive plants for the revision of the Conservation

of Agricultural Resources Act, Act 43 of 1983, and the

drafting of the National Environmental Management:

Biodiversity Act, Act 10 of 2004.

Alien plant invasion is a dynamic process and there-

fore it is essential that the SAPIA database be kept up-to-

date with current information. From October 2006 a sec-

ond phase of the SAPIA mapping project was launched

and all the SAPIA data will be available online at the

Weeds and Invasive Plants (WIP) website, www.agis.

agric.za/wip (Henderson 2006a).

ACKNOWLEDGEMENTS

I thank all the people who have contributed in many

different ways to the compilation of the SAPIA data-

base and have made it one of the most comprehensive

databases on invasive alien plants in southern Africa.

Les Powrie and Mike Rutherford of the South African

National Biodiversity Institute are thanked for the QDS

224

Bothalia 37,2 (2007)

coverage of the biomes of South Africa which was used

to subdivide the SAPIA dataset into the six biome data-

sets for this publication. Special tribute is made to Mike

Wells of the Botanical Research Institute for his inspi-

ration and mentorship during the development of road-

side survey techniques from 1979 to the mid-1980s,

and to Helmuth Zimmermann of the Plant Protection

Research Institute for his motivation and support of the

SAPIA mapping project. SAPIA is an initiative of the

Agricultural Research Council (ARC): Plant Protection

Research Institute which has provided the infrastruc-

ture, basal funding and support since the mid 1980s to

the present. External funding of SAPIA has been grate-

fully received from the Departments of Agriculture,

Environmental Affairs and Tourism, and Water Affairs

and Forestry (Working for Water Programme).

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APPENDIX 1 . — Prominent invaders in study area

Combined taxa e.g. Ageraliim conyzoides/hoiistoniamim indicate uncertainty of identification.

QSp, quarter-degree squares present; QSa, quarter-degree squares abundant; Tr, total records; A, total weighted abundance (see text); Pv,

prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values — see text); R, ranking of

top ten taxa (taxa that are difficult to distinguish are grouped together).

APPENDIX 2. — Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identification. QSp, quarter-degree squares present; QSa, quarter-

degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values

which add up to ± upper 80% of summed values — see text).

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