ISSN 0006 8241 = Bothalia

Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 36,1 May 2006

TECHNICAL PUBLICATIONS OF THE SOUTH AFRICAN NATIONAL

BIODIVERSITY INSTITUTE PRETORIA

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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 Sw~vey of South Africa and Annals of Kirstenbosch Botanic Gardens.

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA

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

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

63.

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

Connell, Stella Gower, Rosemary Holcroft, Kathleen Lansdell, Cythna Lefty (over 700 plates),

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

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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.).

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 36,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

national

biodiversity

institute

S A N B I

2 Cussonia Avenue, Brummeria, Pretoria

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ISSN 0006 8241

May 2006

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

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

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

Bridson, Dr D. 21 Ferrymoor, Ham, Richmond, Surrey TWIO 7SD, England, UK.

Carter Holmes, Mrs S. Royal Botanic Gardens, Kew, UK.

Cope, Dr T. Royal Botanic Gardens, Kew, UK.

Craib, C. P.O. Box 67142, Bryanston, 2021 Sandton, RSA.

Crisp, Prof M.D. Australian National University, Canberra, Australia.

Daemane, M.E. Golden Gate National Park, South African National Parks, 8306 Hadison Park, RSA.

Duncan, G. South African National Biodiversity Institute, Cape Town, RSA.

Edwards, Prof T.J. University of Natal, Pietermaritzburg, RSA.

Geerinck, Dr D. Jardin Botanique National, Meise, Belgium.

Hilton-Taylor, C. lUCN Red List Programme Officer, Cambridge, UK.

Jordaan, Ms M. South African National Biodiversity Institute, Pretoria, RSA.

Klopper, Ms R. South African National Biodiversity Institute, Pretoria, RSA.

Lavranos, J. Apartado Postal 243, 8100 Louie, Portugal.

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

Lotter, M. Mpumalanga Parks Board, Lydenburg, RSA.

Meerow, Dr A.W. National Gemiplasm Repository, Miami, USA.

Nelson, Dr E.C. Tippitiwitchet Cottage, Hall Rd, Outwelj, Wisbech, UK.

Paterson-Jones, Dr D. South African National Biodiversity Institute, Cape Town, RSA.

Pfab, Ms M. Gauteng Nature Conservation, Johannesburg, RSA.

Phillipson, P.B. Museum National d’Histoire Naturelle, Paris, France.

Retief, Ms E. South African National Biodiversity Institute, Pretoria, RSA.

Simon, B. Queensland Herbarium, Brisbane Botanic Gardens, Toowong, Australia.

Smith, Prof G.F. South African National Biodiversity Institute, Pretoria, RSA.

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

Steyn, M. 10 Coniston Close, Bendor Place, 0699 Polokwane, RSA.

Turner, Ms Q. National Herbarium, National Museum, Monuments & Art Gallery, Gaborone, Botswana.

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

Welman, Ms W.A. South African National Biodiversity Institute, Pretoria, RSA.

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

Williamson, Dr G. 26 Starke Rd, Bergvliet, Cape Town, RSA.

CONTENTS

Bothalia 36,1

1 . A taxonomic revision of the genus Merciera (Campanulaceae). C.N. CUPIDO 1

2. Hypoxis (Hypoxidaceae) in Africa: list of species and infraspecific names. Y. SINGH 13

3. Sesotho names for exotic and indigenous edible plants in southern Africa. A. MOTEETEE and B-E.

VAN WYK 25

4. Two new species of Erica (Ericaceae) from the Langeberg, Western Cape, South Africa. R.C. TURNER

and E.G.H. OLIVER 33

5. Two new species of Nemesia (Scrophulariaceae) from southern Africa. K.E. STEINER 39

6. Two new species of Commiphora (Burseraceae) from southern Africa. W. SWANEPOEL 45

7. Notes on the systematics and nomenclature of Tritonia (Iridaceae: Crocoideae). P. GOLDBLATT and

J. C. MANNING 57

8. Notes on African plants:

Amaryllidaceae. A natural hybrid in the genus Clivia. Z.H. SWANEVELDER, J.T. TRUTER and

A.E. VAN WYK 77

Amaryllidaceae. A new variety of Clivia robusta. Z.H. SWANEVELDER, A. FORBES-HARDINGE,

J.T. TRUTER and A.E. VAN WYK 66

Apocynaceae. New records of Adenium boehmianum in the FSA region. S.P. BESTER 63

Asphodelaceae. Aloe kaokoensis, a new species from the Kaokoveld, northwestern Namibia. E.J.

VAN JAARSVELD, W. SWANEPOEL and A.E. VAN WYK 75

Asphodelaceae. Aloe vanrooyenii: a distinctive new maculate aloe from KwaZulu-Natal, South

Africa. G.F. SMITH and N.R CROUCH 73

Capparaceae. Maerua kaokoensis, a new species from Namibia. W. SWANEPOEL 81

Hyacinthaceae. Drimia montana (Urgineoideae), a new species from Eastern Cape, South Africa.

A.P. DOLD and E. BRINK 64

Hyacinthaceae. Ornithogalum kirstenii (Albuca group), a new species from Western Cape, South

Africa, and new combinations in the group. J.C. MANNING and P. GOLDBLATT 86

Poaceae. A long-awaited name change in Polypogon. L. FISH 70

Poaceae. A new species of Sporobolus (Sporobolinae) in South Africa. L. FISH 71

Poaceae. Concept of Stipagrostis uniplumis var. uniplumis redefined to include specimens with

hairy glumes. L. FISH 69

Rubiaceae. Correct author citations for names of three southern African species of Canthium. PM.

TILNEY and A.E. VAN WYK 68

Data deficient flags for use in the Red List of South African plants. J.E. VICTOR 85

9. Reappraisal and identification of Olinia rochetiana (Oliniaceae) in South Africa. R.J. SEBOLA and

K. BALKWILL 91

10. Floristic composition of gold and uranium tailings dams, and adjacent polluted areas, on South Africa’s

deep-level mines. I.M. WEIERSBYE, E.T.F. WITKOWSKI and M. REICHARDT 101

11. Obituary: Helen Joyce Vanderplank (1919-2005). E. BRINK 129

New combinations, hybrid, names, species, status and variety in Bothalia 36,1 (2006)

Aloe kaokoensis Van Jaarsv., Swanepoel & A.E.van Wyk, sp nov., 75

Aloe vanrooyenii Gideon F.Sni. & N.R.Crouch, sp. nov., 73

Clivia X nimbicola Z.H.Swanevelder, J.T.Truler & A.E.van Wyk, nothosp. nov., 78

Clivia robusta B.G. Murray, Y.Ran, P.J.De Lange, K.R.W.Hannnett, J.T.Tniter & Z.H.Swanevelder var. citrina

Z.H.Swanevelder, A.Forbes-Hardinge, J.T.Truter & A.E.van Wyk, var. nov., 67

Commiphora gariepensis Swanepoel, sp. nov., 52

Commiphora steynii Swanepoel, sp. nov., 45

Drimia montana A.P.Dold & E. Brink, sp. nov., 64

Erica euryphylla R.C. Turner, sp. nov., 35

Erica turneri E.G.H.Oliv., sp. nov., 33

Maerua kaokoensis Swanepoel, sp. nov., 81

Nemesia hemiptera K.E. Steiner, sp. nov., 42

Nemesia williamsonii K.E. Steiner, sp. nov., 39

Ornithogalum J.C. Manning & Goldblatt, nom. nov., 87

Ornithogalum kirstenii J.C. Manning & Goldblatt, sp. nov., 86

Ornithogalum neopatersonia /CMcwn/rtg cfe Goldblatt, nom. nov., 88

Ornithogalum soXent J.C. Manning & Goldblatt, nom. nov., 88

Ornithogalum \o\ui2Lrt J.C. Manning & Goldblatt, nom. nov., 88

Polypogon griquensis (Stapf) Gibbs Russ. & L.Fish, comb, nov., 71

Sporobolus oxyphyllus L.Fish, sp. nov., 71

Tritonia gladiolaris (Lam.) Goldblatt & J.C. Manning, comb, nov., 57

Tritonia securigera (Sol. in Aiton) Ker Gawl. subsp. watermeyeri (L.Bolus) J.C. Manning & Goldblatt, comb, et

stat. nov., 60

IV

Bothalia36,l: 1-11 (2006)

A taxonomic revision of the genus Merciera (Campanulaceae)

C.N. CUPIDO*

Keywords: Campanulaceae, Merciera A.DC., South Africa, taxonomy, Western Cape

ABSTRACT

A taxonomic account of Western Cape endemic genus Merciera A.DC. is presented. Six species, supported by recent phe-

netic studies, are recognized. M. brevifolia A.DC., M eckloniana H.Buek, M. leptoloba A.DC. and M. tenidfolia (L.f ) A.DC

are retained as species. M aziirea Schltr. is returned to species status and M. tetraloba C.N.Cupido was recently described.

Each species is described and illustrated. A key to the species, and distribution maps are provided.

INTRODUCTION

Merciera A.DC. is a poorly known genus of small

shrubs confined to Western Cape, South Africa. The

genus is classified in the Campanulaceae subfamily

Campanuloideae. This subfamily, often treated as a sepa-

rate family, comprises between 35 to 55 genera and 600

species (Cosner et al. 2004). In the southern hemisphere

only South Africa shows great diversity with 10 genera.

Eight genera are endemic to South Africa and of these

five are endemic to the Cape Floristic Region (CFR)

(Goldblatt 1978). Taxonomically, genera are separated

on account of capsule structure, particularly the mode

of dehiscence. Merciera is unique amongst the South

African genera in that its capsule is indehiscent. In addi-

tion to the unique capsule, the genus is characterized by

salverform, tetramerous or pentamerous, blue-violet or

white flowers and four basal ovules.

Species limits of Merciera were re-assessed in Cupido

(2003), employing phenetic methods. The results of the

phenetic studies support the recognition of six species.

A taxonomic account of Merciera based on the phenetic

studies is presented here.

MATERIAL AND METHODS

Sampling methods, preparation and examination of

study material used are set out in Cupido (2003). In addi-

tion to specimens from SAM, BOL, and NBG cited in

Cupido (2003), material from PRE, K and MO (acronyms

as in Holmgren et al. 1990) were also examined for this

revision.

Species concept

Taxonomists are frequently criticized for not being

explicit about the criteria used in species delimitation. No

universal species concept exists (Davis & Goldman 1993)

and it therefore depends on the individual taxonomist to

define species level taxa (Cupido 2003). In this study, the

criterion of overall similarity that is based on observed

patterns of character variation is employed. This criterion

has been formulated into the phenetic species concept.

The phenetic species concept considers discrete clusters

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

E-mail: Cupidocn{g!sanbi.org

MS. received: 2005-06-27.

in character hyperspace as species. Phenetic clusters are

recognized by the possession of a particular minimum

number of characters in common. Six phenetic clusters

were revealed by phenetic analysis (Cupido 2003). Each

of these clusters is considered a species.

TAXONOMIC HISTORY

De Candolle (1830) established the genus Merciera to

accommodate three species of Campanulaceae from the

Cape characterized by having four basal ovules in uniloc-

ular or incompletely two-locular ovaries, and indehiscent

capsules. One of these species, M. teniiifolia (L.f) A.DC.

had previously been placed in either Trachelium L. or

Roella L., the other species were newly described. The

genus was named in honour of botanist Phillip Mercier

who wrote a monograph on the family Polemoniaceae.

Initially De Candolle described the genus as "incertae

sedis' because of the unusual structure of the ovary, but

later classified it in the monogeneric tribe Merciereae

(De Candolle 1839). Buek (1837) added two more spe-

cies, but one of these, M. heterornorpha, was erroneously

placed in the genus, belonging in the Rubiaceae instead

(Sonder 1865). Species described by Buek (1837) appear

to have been overlooked by De Candolle (1839). Sonder

(1865) reduced the genus to two species, M. brevifolia

A.DC. and M. teniiifolia, which were divided into two

and four varieties respectively. More than three decades

later, Schlechter (1898) described a species, M. azurea

Schltr. from Sir Lowry’s Pass. In the last comprehensive

treatment of Merciera, Adamson (1954) recognized five

species, one of which, M. teniiifolia, was divided into

two varieties. In a remarkable coincidence, Adamson,

like Buek before him, described a species, M. vaginata

Adamson, which also belongs in the Rubiaceae (Adamson

1955). Since Adamson’s treatment, Cupido (2002) added

one more species from localities west of the Hottentots

Holland Mountains in the Western Cape. In the present

paper, six species are recognized in total, comprising De

Candolle’s (1830) original three, and one each described

by Buek (1837), Schlechter (1898) and Cupido (2002).

None of the species are divided into varieties.

Nomenclatural notes

Phillips (1951) cited Merciera leptoloba A.DC. as the

type species of Merciera, in effect, an act of lectotypifi-

cation. However, according to Stafleu & Cowan (1983),

Pfeiffer (1874) frequently indicated type species for

2

Bothalia 36,1 (2006)

generic names, which constitute in numerous instances,

the first selection of a lectotype. For the genus Merciera,

Pfeiffer (1874) cited only Trachelium tenuifolium, the

basionym of M. tenuifoUa (L.f) A. DC. Single species

are mentioned only when they serve as a type of new

genera or sections, as was done with Merciera. Article

9.17 of the Code (Greuter et al. 2000) is applied here. It

states that the author who first designated a lectotype or

a neotype must be followed. The species designated by

Pfeiffer should therefore be regarded as the lectotype of

Merciera.

The author citations of Merciera species described by

Buck in the Emimeratio (Ecklon & Zeyher 1837) were

given as H.Buek ex Eckl. & Zeyh. by Welman & Cupido

(2003), because the specific epithets are interpreted as

manuscript names. Nordenstam (2003) came to a dif-

ferent conclusion after studying Buck’s taxonomic con-

tributions to South African Campanulaceae, particularly

the typification of species described by Buck. He found

that the published descriptions attributed to the Buck MS

and the wording of the handwritten descriptions found

on the specimens in Stockholm (S), agree completely

with that of Buck. Buck is therefore ‘the author of the

Campanulaceae in the Emimeratio (Ecklon & Zeyher

1837) and names published therein must be attributed to

H.Buek alone (or H.Buek in Eckl. & Zeyh., if a biblio-

graphical citation is involved; cf Art. 46 Note 1 of the

ICBN, Greuter et al. 2000)’ (Nordenstam 2003). This

conclusion is followed here.

MORPHOLOGICAL CHARACTERS

Habit

All species are resprouting, dwarf shrubs with branched

stems. Stems are decumbent to suberect, occasionally

with groups of branches at the end of a year’s growth.

The resprouting habit of Merciera is an adaptation to

survive fires, and the genus is eonfmed to the fire-prone

fynbos vegetation of the Cape Region. Vegetatively,

Merciera species resemble species of Roella, particularly

of the series Ciliatae.

Leaves

The leaves are alternate, linear and sessile, often

appearing dead due to the brown colour. The margins are

entire, and usually ciliate. The abaxial surface is hairy in

all species, except in Merciera tetraloba C.N. Cupido.

Leaves are variable in size and insertion along the

stem. In Merciera leptoloba the leaves become smaller

towards the top of the stem. In M. temiifolia and M.

azurea the leaves are subequal and crowded, but in M.

eckloniana H.Buek, they are widely spaced.

Clusters of smaller green leaves are always present in

the axils of leaves in all species except Merciera azurea,

in which they are seldom present. These leaf clusters are

in fact highly reduced short shoots developed in the axils

of long shoot leaves.

Inflorescence

In an account on the inflorescence morphology of the

Campanulaceae, Philipson (1953) described the inflores-

cenee of Merciera as interealary. The flowers are solitary

in the axils of foliage leaves and after producing a zone

of flowers, the axis continues to grow vegetatively. This

arrangement is particularly marked in M. leptoloba.

Careful examination of the inflorescence reveals that

there are in fact three flowers per axil. Only the termi-

nal one develops however, and the two lateral flowers

remain rudimentary on highly reduced lateral branches

with bract-like leaves. The tenninal flower lacks a bract-

like leaf The reduction of the lateral branches gives the

flowers an axillary appearance. This basic structure is

repeated in the entire flowering zone, forming a spike-

like synflorescence towards the end of the main branches.

The order of flowering is acropetal.

Flowers

Phenotypic plasticity is common in the Campanulaceae

(Eddie & Ingrouille 1999) and Merciera is no exception

to the rule. Numbers of floral parts are sometimes vari-

able in the same species. Additional floral parts tend to

develop, and flowers that are normally pentamerous,

may become hexamerous. Both flower types are usually

present on the same plant.

Calyx

The calyx is 4- or more commonly 5-lobed. Hairs are

often present on the hyaline tips, but only on the margins

in Merciera tetraloba. The lobes are fused at the base to

fonn a short tube.

Corolla

The corolla is actinomorphic with a conspicuous tube

and 4 or 5 spreading lobes. The corolla colour is white or

blue to violet, and occasionally white-flowered species

have purple tips to the corolla lobes. Tube length and

flower colour is correlated, dividing the genus into two

groups. White-flowered species have tube lengths less

than 7 mm, whereas blue to violet-flowered species have

tube lengths exceeding 7 mm.

Androeciwn

Stamens are 4 or 5, free, inserted at the base of the

corolla tube and included in the corolla. The filaments

are flattened, wider and pilose about the middle becom-

ing narrower towards the apex. Anthers are linear and

basifixed.

Gynoeciwn

The inferior ovary is surrounded by a hispid hypan-

thium. In accordance with the trichome tenninology

of Payne (1978), four trichome types are found on

the hypanthium: filifonn, clavate, uncinate or circinate

(Figure lA-D). The locule number has been described

as 1- or incompletely 2-locular (De Candolle 1830).

However, careful dissecting of the ovary reveals a

complete but delicate septum, dividing the ovary into 2

locules. Each locule contains 2 basal ovules. The style

is filifonn, exserted, glabrous and inserted in a convex

disc. The style is shortly divided, with the number of

stigmatic lobes corresponding to the number of locules

in the ovary.

Bothalia36,l (2006)

3

ABC D

1 mm

FIGURE 1. — Merciera. A-D, trichome types: A, clavate; B, filiform;

C, uncinate; D, circinate. E, capsule. Scale bar: 1 mm. Artist:

W.A. Hitchcock.

Fruits and seeds

Merciera has indehiscent, hispid capsules, crowned

with a persistent calyx (Figure IE). Only 1 ovule devel-

ops into a seed, which occupies the entire capsule cavity.

Seeds are elliptic to ovate, pale brown with a darker

hilum.

TAXONOMIC TREATMENT

Merciera A. DC., Monographie des Campanulees.'

369 (1830); H.Buek: 372 (1837); Sond.; 530 (1865);

Adamson: 157 (1954). Type species: M. temiifolia (L.f.)

A. DC. [Trachelium teniiifoliiim L.f.: 143 (1782)]. (lecto-

type, designated by Pfeiffer 1 874).

Subshrubs; branches hispidulous to hispid. Stems

decumbent or suberect, branched. Leaves alternate, ses-

sile, linear, subulate, entire, scattered or crowded, ascend-

ing or spreading, glabrous or hairy abaxially, margins ±

ciliate, axillary clusters of smaller, glabrous leaves often

present. Inflorescence 3-flowered, with 1 terminal, and

2 rudimentary flowers lateral, on highly reduced lateral

branches with bract-like leaves, aggregated into spike-

like synflorescences towards ends of main branches.

Flowers sessile, axillary, actinomorphic; bract-like leaves

2, succulent, subtending each rudimentary flower, absent

in terminal flower; hypanthium obconical, hispid with

clavate, filiform, uncinate or circinate trichomes; calyx

4- or 5-lobed, often fused at base to form short tube, gla-

brous or hairy on hyaline tips and margins; corolla nar-

rowly tubular or funnel-shaped, white, occasionally with

purple tips, or violet-blue, or very rarely pale blue, lobes

4 or 5, ovate or linear-lanceolate, occasionally unequal,

glabrous, or hairy on back. Stamens 4 or 5, free, inserted

at base of corolla tube; filaments flattened, wider and

pilose ± in middle, narrower towards apex; anthers linear,

basifixed. Ovafy inferior, 2-locular, containing 4 erect

basal ovules; style filiform, bifid, exserted, glabrous,

swollen at base; stigmas glabrescent. bluish purple. Fruit

a hispid capsule, crowned with persistent calyx, 1 -seed-

ed, indehiscent. Seed elliptic to ovate.

The six species are endemic to the southwestern parts

of the CFR and grow in open, sandy, clayey or rocky soil,

often in disturbed habitats. Fire is important in the growth

and survival of the genus. After fire, the plants resprout

from the base and a period of four to six years of vigorous

vegetative growth and flowering follows. After six years,

the plants become moribund and start disappearing when

the veld remains unbumt for very long periods.

1. Merciera tenuifolia (L.f.) A. DC., Monographie

des Campanulees: 370 (1830); Sond.: 596 (1865);

Adamson: 159 (1954). Trachelium tenuifolium L.f:

143 (1782); Thunb.: 38 (1800). Roella tenuifolia (L.f)

Thunb.: 174 (1823). Type: South Africa, without precise

locality, Thunberg 4774 (UPS-THUNB, lecto., here des-

ignated, NBG, microfiche!).

M temiifolia (L.f.) A. DC. var. candotleana Sond.: 596 (1865).

Type: South Africa, Western Cape, Houwhoek Mountains, Ecklon &

Zeyher 2417 (SAM\).

M. temiifolia (L.f) A.DC. var. thunbergiana Sond.: 596 (1865).

Type: South Africa, without precise locality, Thunberg 4 773 (UPS-

THUNB, holo.).

4

Bothalia36,l (2006)

Key to species

la Flowers pentamerous, blue, violet or purple, rarely white; corolla tube more than 7 mm long:

2a Plants slender (stem equal to or less than 1 mm thick); leaves scattered; corolla lobes glabrous adaxially;

distributed from Groenlandberg (Grabouw, 3419AA) northwards to Tulbagh (3319AC) 3. M eckloniana

2b Plants stout (stem more than 1 mm thick); leaves crowded; corolla lobes hairy adaxially; distributed south

of Groenlandberg (Grabouw, 3419AA):

3a Stems suberect; leaves ascending, abaxial surface hairy, axillary clusters of smaller leaves always present;

corol-la tube 11-26 mm long, five times as long as lobes 1. M temiifolia

3b Stems decumbent; leaves spreading, abaxial surface glabrescent, axillary clusters of smaller leaves occa-

sionally present on lower parts of stem; corolla tube 7-14 mm long, less than three times as long as

lobes 2. M aziirea

lb Flowers tetramerous or pentamerous, white, occasionally with purple tips; corolla tube less than 7 mm long:

4a Flowers tetramerous; margins of calyx lobes ciliate; hypanthium trichomes uncinate to circinate; plants

growing in clayey soil; distributed west of Hottentots Holland Mountains 6. M tetraloba

4b Flowers pentamerous; margins of calyx lobes glabrous; hypanthium trichomes clavate or filiform; plants

growing in sandy or stony soil; distributed southeast of Hottentots Holland Mountains:

5a Plants decumbent, stout; lower leaves more than 8 mm long, crowded; corolla lobes, linear-lanceolate;

2-6 mm long, almost as long as tube; hypanthium trichomes clavate 4. M. leptoloba

5b Plants suberect, slender; lower leaves less than 8 mm long, scattered; corolla lobes ovate, 2-3 mm long,

up to half as long as tube; hypanthium trichomes filiform 5. M. brevifoUa

Stems suberect, sparsely or profusely branched, occa-

sionally with group of branches at end of each season’s

growth. Leaves crowded, ascending, hairy on abaxial

surface, axillary cluster of smaller leaves occasionally

present; bract-like leaves 4-10 mm long. Flowers violet-

blue, rarely white; hypanthium 1-3 mm long, hispid with

clavate trichomes; calyx lobes 5, 0.8-1 mm long, hairs

on hyaline tips; corolla tube narrow, 10.0-25.5 mm long;

lobes 5, ovate, 2. 5-4.0 mm long, hairy on back. Stamens

5; filaments 7-21 mm long. Style 13.0-30.5 mm long.

Flowering time: December to January. Figure 2.

Distribution and habitat: the distribution of Merciera

tenuifolia (Figure 3) is limited to Bot River, Houwhoek

and Kogelberg where it is found on stony soil at altitudes

ranging between 110 and 600 m.

The locality of the specimen MacOwan 3103 (SAM)

collected at Tulbagh Nuwekloof is suspect. It has the

same locality, collecting date and number as a specimen

of M. azurea. No recent collections of M. tenuifolia have

been made in the Tulbagh area.

Conservation status: Vulnerable D2 (World Conser-

vation Union [lUCN] 2001 ).

2. Merciera azurea Schltr. in Botanische Jahrbiicher

24: 447 (1898). M tenuifolia (L.f.) A. DC. var. azurea

(Schltr.) Adamson: 160 (1954). Type: South Africa,

Western Cape, Sir Lowry’s Pass, Schlechter 7263 (B,

holo.; MO, SAM, iso.!).

Stems decumbent, stout, occasionally with group

of branches at end of each season’s growth. Leaves

crowded, spreading, glabrous or hairy on abaxial surface,

axillary cluster of smaller leaves occasionally present;

bract-like leaves 4. 0-9. 5 mm long. Flowers violet-blue,

rarely white; hypanthium 1.5-2. 7 mm long, hispid with

clavate or filiform trichomes; calyx lobes 5, 1.0-1. 9

mm long, hairy on hyaline tips; corolla tube wide, 7-14

mm long; lobes 5, ovate, 3. 0-5. 5 mm long, glabrous

or hairy on back. Stamens 5; filaments 7-11 mm long.

Style 12.0-19.5 mm long. Flowering time: November to

February. Figure 4.

Distribution and habitat: Merciera azurea ranges from

Sir Lowry’s Pass to Bredasdorp (Figure 5) and occurs on

sandy or stony soil at altitudes ranging between 100 and

650 m.

The locality of the specimen MacOwan 3103b (SAM)

collected at Tulbagh Nuwekloof is suspect. It has the

same locality, collecting date and number as a specimen

of Merciera tenuifolia. No recent collections of M. azurea

have been made in the Tulbagh area.

During 1896 MacOwan made several collecting trips

to Tulbagh, Caledon, Houwhoek and the Hottentots

Holland Mountains (Gunn & Codd 1981) and it could

have happened that specimens from the different locali-

ties were unknowingly mixed up resulting in incorrect

locality information.

Consefwation status: Vulnerable D2 (World Conser-

vation Union [lUCN] 2001).

3. Merciera eckloniana H.Buek in Eckl. & Zeyh.,

Enumeratio plantarum Africae Australis extratropicae:

387 (1837); Adamson: 160 (1954). M. tenuifolia (L.f.)

A. DC. var. eckloniana (H.Buek) Sond.: 596 (1865).

Type: South Africa, inter rupes (altit. IV) montium supra

‘Waterfall’ in valle ‘Tulbagh’ (Worcester), December,

Ecklon & Zeyher 2420 (S, holo.; SAM, iso.!).

Stems semi-erect, slender, occasionally with group of

branches at end of each season’s growth. Leaves scat-

tered, spreading, glabrous, or hairy on abaxial surface,

axillary cluster of smaller leaves occasionally present;

bract-like leaves 2-6 mm long. Flowers violet-blue,

rarely white; hypanthium 1 .0-2.8 mm long; hispid with

filifonn trichomes; calyx lobes 5, less than 1 mm long,

hairs on hyaline tips; corolla tube narrow, 7.5-16.0 mm

long, lobes 5, ovate, 1.5-3. 5 mm long, glabrous on back.

Stamens 5; filaments 5.5-10.0 mm long. Style 8.5-17.5

mm long. Flowering time: October to February. Figure

6.

Distribution and habitat: this species is distributed

from the Groenlandberg northwards to Tulbagh (Figure

5). It is found on sandy or stony soil at altitudes ranging

from 450 to 1 500 m.

Conservation status: Vulnerable D1 (World Conser-

vation Union [lUCN] 2001).

FIGURE 2. — Merciera tenuifolia, Cupido 289. A, portion of plant; B, flowering branch; C, flower with prophylls. Scale bars: A, 10 mm; B, C, 3

mm. Artist: W.A. Hitchcock.

4. Merciera leptoloba A. DC., Monographic des

Campanulees: 371 (1830); H.Buek in Eckl. & Zeyh.:

387 (1837); Adamson: 162 (1954). M brevifolia A.DC.

van leptoloba (A.DC.) Sond.: 596 (1865). Type: South

Africa, Cape of Good Hope (Caput Bonae Spei), Hooker

s.n. (K, holo.!).

Stems decumbent with groups of branches at end of

each season’s growth. Leaves scattered to crowded, lower

leaves more than 8 mm long, glabrous to hairy on abaxial

surface, with axillary cluster of smaller leaves; bract-like

leaves 2-8 mm long. Flowers white; hypanthium 1-3

mm long; trichomes clavate; calyx lobes 5, 0.7-1. 8 mm

long, hairy on hyaline tips, rarely on back; corolla tube

3. 0-5. 5 mm long, occasionally shorter than lobes; lobes

5, linear, 2-6 mm long, glabrous on back. Stamens 5;

filaments 3-5 mm long. Style 4-12 mm long. Flowering

time: November to March. Figure 7.

Distribution and habitat: Merciera leptoloba is a com-

mon species of the Cape southeast coast, from Kogelberg

to Bredasdorp (Figure 8). This species is found on sandy

or stony flats and hills at altitudes ranging between sea

level and 400 m.

FIGURE 3. — Known distribution of Merciera tenuifolia.

Bothalia36,l (2006)

l^/.A

FIGURE 4. — Merciera azurea, Cupido 68. A, portion of plant; B, flowering branch; C, flower with prophylls. Scale bars: A, 10 mm; B, C, 2 mm.

Artist: W.A. Hitchcock.

FIGURE 5. — Known distribution of Merciera azurea, •; M. eckloni-

ana, ■.

Conservation status: Vulnerable D1 D2 (World

Conservation Union [lUCN] 2001).

5. Merciera brevifoiia A. DC., Monographie des

Campanulees: 371 (1830); H.Buek in Eckl. & Zeyh.:

387 (1837); Adamson: 161 (1954). Type: South Africa,

without precise locality, Masson s.n. (BM, holo. -NBG,

photo.!).

Stems semi-erect, slender with groups of branches at

end of each season’s growth. Leaves scattered to crowded,

less than 8 mm long, glabrous to hairy on abaxial surface,

with axillary cluster of smaller leaves; bract-like leaves

2-A mm long. Flowers white; hypanthium 0.8-1. 6 mm

long; trichomes filiform; calyx lobes 5, 0. 5-1.0 mm long,

hairy on hyaline tips, rarely on back; corolla tube 3-6 mm

long; lobes 5, ovate, 1. 5-3.0 mm long, glabrous on back.

Stamens 5; filaments 2-4 mm long. Style 4. 0-8. 5 mm

long. Flowering time'. November to February. Figure 9.

Distribution: Merciera brevifoiia is a montane spe-

cies occurring on the Babylons Tower, and on the Bot

Bothalia 36,1 (2006)

7

FIGURE 6. — Merciera eckloniana, Cupido 71. A, portion of plant; B, flowering branch; C, flower with prophylls. Scale bars: A, 10 mm; B, C, 3

mm. Artist: W.A. Hitchcock.

River, Houwhoek, Shaw’s Mountains and the Caledon

Swartberg (Figure 10).

On the Houwhoek Mountains where this species

occurs in sympatry with Merciera leptoloba, possible

hybrids between the two species are formed.

Conservation status: Vulnerable D2 (World Conser-

vation Union [lUCN] 2001).

6. Merciera tetraloba C.N. Cupido in Bothalia 31:

74 (2002). Type: Western Cape, Strand, Harmony Flats,

Tortoise Nature Reserve, offDisa Road, 17 January 2000,

C.N. Cupido 77 (NBG, ho lo.!; BM!, K!, MO!, NY!,

PRE!, iso.).

Stems decumbent or suberect, slender, occasionally

with groups of branches at end of each season’s growth.

Leaves scattered, ascending, the older spreading, gla-

brous on abaxial surface, margins ciliate; axillary cluster

of smaller leaves present; bract-like leaves, 1-4 mm long.

Flowers tetramerous, white, occasionally with purple

tips, or very rarely pale blue; hypanthium 1-2 mm long,

hispid with uncinate or circinate trichomes; calyx lobes

4, 0.6-1. 2 mm long, often hairy on hyaline tips and mar-

gins; corolla tube 4-6 mm long; lobes 4, ovate, 2-3 mm

long, glabrous or hairy on back. Stamens 4; filaments

3. 0^.5 mm long. Style 6-10 mm long. Flowering time:

November to January. Figure 11.

Distribution and habitat: this species is found in Faure,

Gordon’s Bay, Sir Lowry’s Pass, Somerset West, Strand,

Dal Josaphat, Du Toitskloof, Stellenbosch, Hermon and

Malmesbury on flats and lower mountain slopes at alti-

tudes between 30 and 350 m (Figure 10). It grows in open

clayey soil, often in disturbed habitats.

When originally described, this species was thought to

be restricted to Faure, Gordon’s Bay, Sir Lowry’s Pass,

Somerset West, Strand, Dal Josaphat, Du Toitskloof, and

Stellenbosch. Subsequently, the author came across a

specimen collected by Elsie Esterhuysen {Esterhuysen

34802, BOL) at Michiel Heyns Kraal near Malmesbury

in 1977 and misidentified as Lightfootia. Attempts to

verify this collection in the field have so far been unsuc-

cessful. John Manning recorded another new locality for

this species {Manning 2941, NBG) at Hermon on the

Farm Bosplaas in 2005.

Conservation status: Endangered B1 a (i, ii) b (iii) 2 (i,

ii) b (iii) D (World Conservation Union [lUCN] 2001).

Large areas of the habitat of this species in the

Helderberg and Stellenbosch areas have been destroyed

because lower mountain slopes and lowland areas are

sought after for urban development. On the Harmony

8

Bothalia 36,1 (2006)

FIGURE 7. — Merciera leptoloba, Cupido 66. A, portion of plant; B, flowering branch; C, flower with prophylls. Scale bars: A, 10 mm; B, C, 2

mm. Artist: W.A. Hitchcock.

Flats in Strand, the few existing populations are under

serious threat of extinction.

Excluded species

Merciera heteromorpha H.Buek = Carpacoce hetero-

morpha (H.Buek) Bolus

When Buek (1837) described the species, he noted that

it most likely constituted a distinct genus. Sonder (1865)

considered it a member of Rubiaceae, but did not treat it

taxonomically. A few decades after Sonder, Bolus (1896)

transferred the species to the genus Carpacoce in the

Rubiaceae, where it is currently classified.

Merciera vaginata Adamson = Carpacoce heteromor-

pha (H.Buek) Bolus {Merciera heteromorpha H.Buek)

Adamson erroneously thought that Stokoe s.n. (SAM)

from the Somerset Sneeukop represented a new spe-

cies of Merciera and consequently described it as such.

After it had been brought to his attention that the plant

described as M. vaginata appeared to be the same as

FIGURE 8. — Known distribution of Merciera leptoloba.

Bothalia 36,1 (2006)

FIGURE 9. — Merciera brevifolia, Cupido 235. A, portion of plant; B, flowering branch; C, flower with prophylls. Scale bars: A, 10 mm; B, 2

mm; C, 1 mm. Artist: W.A. Hitchcock.

FIGURE 10. — Known distribution of Merciera brevifolia, •; M.

tetraloba, ■.

certain specimens in the herbarium of the South African

Museum, he re-examined the specimen. He conceded that

the plant was the same as Zeyher 2421, the type of M

heteromorpha (Adamson 1955) and is therefore correctly

referred to Carpacoce heteromorpha.

Merciera muraltioides Schltr.

Specimens collected by R. Schlechter (Schlechter

7372, BOL, MO, SAM) from Houwhoek appear under

the manuscript name Merciera muraltioides Schltr. sp.

nov. This name was never published and is therefore not

valid. The inclusion of these specimens in M brevifolia

A.DC. by Adamson (1954) is supported by the phenetic

studies of Cupido (2003) and accordingly upheld in this

paper.

SPECIMENS EXAMINED

Adamson 4095 (4) PRE; 4098 (1) BOL; 4773, 4774, 4898 (4) BOL;

4780 (4) K; 478J (4) SAM; 4895 (6) BOL; 4904 (2) SAM.

Barker 286 (3) NBG; 7776 (4) NBG; 8802 (5) NBG; 8865 (6) NBG.

Barker (Belle) s.n. (2) MO. Barnard 40469 (2) SAM. Bayer SA01108

10

Bothalia36,l (2006)

FIGURE 1 1. — Merciera tetmloba, Cupido 77. A, portion of plant, life size; B, flowering branch; C, flower with prophylls. Scale bars: B, 4 mm;

C, 2 mm. Artist: Inge Oliver.

(2) MO. Bayliss 4089 (6) NBG, MO. Bolus 6i4 (2) SAM; 4679, 5105

(5) BOL; 6948 (I) NBG; 7402 (5) BOL; BOL98256 (6) BOL. Bond

1684 (3) NBG. Boucher 167, 1781 (2) NBG; 932, 1769 (4) NBG; 3447

(6) NBG, PRE; 5279 (\) NBG. Brenan 14048 (4) NBG, MO, K. Burger

2859 (4) NBG. Burman 1079 (2) BOL; 1255 (4) BOL.

Compton 6116, 10223, 14225, 18951, 19016, 23238 (4) NBG; 10372

(6) NBG; 10603 (5) NBG; 14234, 16835 (2) NBG; 16832 (1) NBG;

21897 (3) NBG. Cupido 66, 69. 72 (4) NBG; 67, 71 (3) NBG; 68. 70.

73 (2) NBG; 75, 77. 117(6) NBG.

De Vos 475 (4) NBG; 1161 (2) NBG. Drege SAMI 7297 (6) SAM.

Ecklon & Zeyher s.n. (5) SAM; 2417 (1) SAM; 3154 (5) NBG.

E.sterhuysen 4229, 10007 (2) BOL; 4954 (4) NBG; 11424, 14361 (3)

BOL; 19594, 33722 (4) BOL.

Forsyth 394 (4) NBG.

Galpin 11316 (4) PRE, K. Goldblatt 5381 (4) PRE, K; 7623, 11262 (3)

MO. Gill s.n. (1) BOL. Gillet s.n. (5) NBG; 670 (3) NBG. Guthrie s.n.

(4) NBG; 2275 (1) NBG; 2792 (6) NBG.

Hafstriim & Acocks 2006 (5) PRE. Haynes 1543 ( 1 ) NBG.

.lordaan s.n. (6) NBG; 832, 18402 (2) NBG.

Kensit 13469 (4) BOL, MO. Kruger 90 (2) NBG; 91 (4) NBG.

Leighton 906 (2) BOL; 2465, 2587 (4) BOL. Le Maitre 177 (\) NBG.

Levyns 4046 (4) BOL; 5372 (2) MO, BOL; 11265 (4) BOL, PRE. Lewis

3194 (4) SAM; 3532 (5) SAM. Liede 16447 (4) MO.

MacOwan 3103 (1) SAM; 3103b (2) SAM. Manning 2941 (6) NBG.

Markotter 8639 (6) NBG. Martin s.n. (2) NBG. McDonald 603, 1735

(3) NBG.

Nilsson 120 (4) PRE.

Oliver & Oliver 11866 (6) NBG. Orchard 341 (4) NBG, MO; 358, 524

(2) NBG, MO.

Pappe s.n. (4) K. Parker 3455 (6) K, MO; 3550 (6) NBG, MO, K; s.n.

(6) MO. Pillans 6749 (3) BOL. Potts 5054 (4) SAM.

Rourke 998 (1) NBG. Rycroft 3149 (4) NBG.

Salter 5136 (3) BOL, K. Schlechter 7211 (6) PRE, MO, K; 7263 (2)

MO, SAM; 7370 (1) MO; 7372 ( 5) BOL, MO, PRE; SAM, 9228 (3)

MO. Steiner 2445 (2) NBG. Stokoe 8653, 9113, 69735, 358443 (1)

BOL; 58444 (4) SAM; 58445, 64365, 67099 (3) SAM; 64366, 65581

(2) SAM.

Taylor 3016, 3877 (3) NBG; 3793 (4) NBG; 4401, 10251 (2) NBG;

Bothalia36,l (2006)

9557 (4) K, NBG, PRE, Thompson 2303 (3) NBG; 3872 (4) NBG.

Tyson 899 (6) SAM.

Van der Merwe 1787 (2) NBG. Van der Schijff 7437 (2) MO; 7457 (4)

MO. Van Jaarsveld & Bean 6411 (3) NBG. Viviens 81 (2) NBG; 775

(3) NBG.

Wallers 51, 1048 (4) NBG. Williams 65 (4) MO, K; 2937 (2) NBG;

2949 (4) NBG, PRE; 3389 (4) NBG.

Zeyher 3152 (1) NBG; 3154 (5) BOL.

ACKNOWLEDGEMENTS

This Study formed part of an M.Sc. (Systematics &

Biodiversity Science) thesis obtained at the University of

Cape Town. I wish to express my thanks and appreciation

to many people and institutions for their assistance and

support: Prof H.P. Linder for supervising the study; dr

J.C. Manning for valuable comments on the final draft

of this manuscript; the curators of BOL, K, NBG and

PRE for permission to examine their collections; P.B.

Phillipson for providing information for specimens at

MO; my colleagues at the Kirstenbosch Research Centre

for their support and encouragement; the South African

National Biodiversity Institute for financial support; and

the Western Cape Nature Conservation Board for grant-

ing permission to collect plants.

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■tesj

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Bothalia 36.1: 13-23 (2006)

Hypoxis (Hypoxidaceae) in Africa: list of species and infraspecific

names

Y. SINGH*t

Keywords: Africa, distribution, Hypoxis L., list, synonyms

ABSTRACT

A list of species and synonyms is presented for the African Hypoxis L. An abbreviated reference to the original publication

of a species is included in the list. Distribution of taxa is indicated by the country in which they occur. A list of synonyms

with accepted or suggested names and the reference to change in status of a taxon is provided. Subsequent publications on the

treatment of a taxon are given to denote congruous and differing resolution of taxa by authorities.

INTRODUCTION

Numerous species names appear in the genus Hypoxis

L., causing much confusion in nomenclature. Some

redundant epithets are due to species in the related genera,

Ciirculigo Gaertn., Spiloxene Salisb., Sanieila Hilliard &

B.L.Burtt and Rhodohypoxis Nel being initially recog-

nized and described as species of Hypoxis. Others arose

through new descriptions for species with names already

in use. This is understandable for a group that displays

great polymorphism during its growing season. Leaves in

most species elongate after flowering, giving the plant a

different appearance. Descriptions were often based on a

single morph of the plant and these gave rise to a number

of names for a single species. A further problem in the

genus is that the reduction of species to synonymy was

not always formalized in earlier studies on Hypoxis in

Africa. For example, Nel (1914) in his major treatment of

the Hypoxidaceae in Africa, indicated some synonyms by

citing the type specimens of the synonym under what he

considered to be valid species. He recorded other syno-

nyms in the index at the end of his treatment. In addition,

a few species reduced to synonymy appear as valid spe-

cies in later publications without reference to its earlier

reduced status or explanation for change in status.

To clarify the nomenclature of Hypoxis, a list of

species in Africa is presented. The intention is to pro-

vide a consolidated list of all names applied to African

Hypoxis since the description of the first African spe-

cies, H. villosa L.f in 1782. The list is proposed as a

start towards a possible World checklist for the small

family Hypoxidaceae, similar to that produced for the

Araceae (Frodin & Govaerts 2002). Data for the southern

African (South Africa, Swaziland, Lesotho, Botswana

and Namibia) species are based on a taxonomic revision

by the author (Singh in prep.) and on the contribution on

Hypoxidaceae by Snijman & Singh (2003) in Plants of

southern Africa: an annotated checklist. For the species

in tropical Africa, data were extracted from published

literature. Having examined most of the type material

and early literature, a few discrepancies have come to

* South African National Biodiversity Institute, KwaZulu-Natal

Herbarium, Botanic Gardens Road, 4001 Durban,

t Student affiliation: Department of Botany, University of Pretoria,

0002 Pretoria.

MS. received: 2005-02-28.

light: for example, the confusion with the concept of the

distinct species H. obtusa Burch, ex Ker Gawl. and H.

villosa L.f where the names were used interchangeably

for specimens belonging to the taxa. These discrepancies

will be elaborated upon in subsequent publications.

The IPNI electronic database (www.ipni.org./index.

html) was used as the starting point to compile a list

of all species names applied to Hypoxis on the African

continent. Accepted names were extracted into a list and

all synonyms and uncertain names were placed in Table

1 . All names proposed in manuscripts, dissertations and

herbarium sheets were added to the list of names in the

table, and where possible, the status of taxa was verified.

References on Hypoxis in Africa including those given in

the IPNI database for each taxon were sought. All spe-

cies reduced to synonyms were cross-referenced in the

literature and noted with a reference in the table. Based

on knowledge gained through the study of African type

material and descriptions, the reduction of species by

authors was either accepted or rejected.

Structure of the list

The list includes 69 species and 21 infraspecific taxa

of African Hypoxis, with synonymy, reference and dis-

tribution. Species are listed alphabetically with accepted

names in bold. Synonyms are listed per species in italics

and are also in alphabetical order. The reference to the

original description of a species is given in abbreviated

format stating the author, page number and year of pub-

lication (see references for details). Author names are

according to Brummitt & Powell (1992). Distribution

of taxa are indicated by the countries in which they

occur and the countries are abbreviated here alphabeti-

cally and in the list as follows: A, Angola; B, Burundi;

Bot, Botswana; Cam, Cameroon; CAR, Central African

Republic; Ch, Chad; DRC, Democratic Republic of

Congo; E, Eritrea; Eth, Ethiopia; G, Gabon; Gu, Guinea;

K, Kenya; L, Lesotho; Lib, Liberia; Mad, Madagascar;

Mai, Malawi; Mau, Mauritius; Moz, Mozambique; N,

Namibia; Nig, Nigeria; Re, Reunion; Rwa, Rwanda;

SA, South Africa; SL, Sierra Leone; Som, Somalia; Sud,

Sudan; Swa, Swaziland; T, Tanzania; Lf, Uganda; Zam,

Zambia; Zim, Zimbabwe. Where the locality of a taxon

is uncertain, a question mark is used after the suggested

country e.g. Mai?.

14

Bothalia 36,1 (2006)

Structure of the accompanying table

Table 1 is a listing of synonyms with accepted or sug-

gested names based mainly on literature and the reference

to where the status of a species was changed. Column 1

of the table is a list of synonyms and uncertain species in

alphabetical order. Column 2 gives the accepted names

in bold and unaccepted names in italics. For taxa where

the status could not be confirmed, suggested names are

marked with an asterisk. Column 3 gives the publica-

tion in which the status of a taxon was first changed.

Subsequent publications on the treatment of a taxon are

given in chronological order and this is included to denote

congruous and differing resolution of taxa by authorities.

The references are restricted mainly to revisions, regional

treatments and enumeration lists for Hypoxis in Africa.

References

References are restricted to taxonomic works on

African Hypoxis. These are abbreviated in the list and

table, and are arranged chronologically in the reference

column of the table. A list of references is appended.

Hypoxis L.

Subgenus Hypoxis Baker: 99 (1878b)

Section Hypoxis Geerinck: 75 (1969)

aculeata Nel: 327 (1914); Mai

acuminata Baker: 3 (1889); Les, SA, Swa

angolensis Baker: 266 (1878a); Ang, DRC, T, Zam

angustifolia Lam.: 182 (1789)

biflora Baker: 181 (1876), non De Wild.

djalonensis Hutch, in Hutch. & Dalziel: 394 (1931)

var. angustifolia; Mau, Re, SA

too Nel: 324 (1914)

var. buchananii Baker: 111 (1878b); SA, Swa

obliqua Jacq. var. woodii (Baker) Nel: 309 (1914)

woodii Baker: 3 ( 1 889)

var. luzuloides (Robyns & Toumay) Wiland: 148 (2002); B, Cam,

CAR, Ch. DRC, Eth, G, GU, K, Mad, Moz, Nig, Rwa,

SA?, SL, Sud, T, U, Zam, Zim

luzuloides Robyns & Toumay: 254 (1955)

var. madagascariensis Wiland: 148 (2002); Mad

araneosa Nel: 310 (1914); K

arenosa Nel: 325 (1914); T

argentea Harv. ex Baker: 110 (1878b)

var. argentea; Les, SA, Swa

var. sericea Baker: 110 (1878b); Les, N, SA, T

dinteri Nel: 302 (1914)

parviflora Dinter ex Soldi: 115 (1960), nom. nud.

sericea Baker: 111 (1 878b)

sericea Baker var. dregei Baker: 112 (1878b), pro parte quoad

specim. Drege 8525 (K)

sericea Baker vax. flaccida Baker: 112(1 878b)

bampsiana Wiland: 207 (1997c); DRC, Mai, T, Zam

multiceps sensu Zimudzi: 14 (1996), non Buchinger ex Baker

sp. A pro parte quoad Bullock 2045 Nordal et al,: 29 (1985)

camerooniana Baker 7: 577 ( 1 898); Cam

lanceolata’HeV. 335 (1914)

ledermannii Nel : 314 (1914)

petrosa Nel: 325 (1914)

recui-va Hook.f: 223 (1864)

thorbeckei Nel: 328 (1914)

villosa L.f. var. recurva (Hook.f.) Baker: 114(1 878b)

villosa auct. non L.f \ar. Jbliis recurvis ' Hook.f: 223 (1864)

campanulata Nel: 314 (1914); T

canaliculata Baker: 265 (1878a); A, DRC, Mai, Zam, Zim

colchicifolia Baker: 3 (1889); SA

dislachya Nel: 322 (1914)

gilgiana Nel: 322 (1914)

latifolia Hook.: t. 4817 (1854)

oligotricha Baker: 3 ( 1 889)

costata Baker: 119 (1878b); Les, SA, Swa

cryptophyila Nel: 316 (1914); Rwa, T

cuanzensis Welw. ex Baker: 265 (1878a); A, K, T, Zam, Zim

macrocarpa E.M.Holt & Staubo in Nordal et. al.: 25 (1985)

demissa Nel: 328 (1914); T

exaltata Nel: 331 (1914); SA

filiformis Baker: 109 (1878b); A, B, DRC, Les, Mai, Moz, SA, Swa,

T, U, Zam, Zim

caespitosa Baker: 858 (1901)

dregei (Baker) Nel: 306 (1914)

dregei Baker var. biflora (De Wild.) Nel: 306 (1914)

miinznerinAsV. 307 (1914)

sericea Baker var. dregei Baker: 112 (1 878b), pro parte quoad

specim. Cooper 1811 (LCD)

fischeri Pax: 143 (1893); Cam. Zam

flanaganii Baker: 179 (1896); SA

floccosa Baker: 357 (1894); SA

ecklonii Baker: 859 (1901) as eckloni

galpinii Baker: 188 (1896); SA, Swa, Zim

coitoto sensu NorL: 164(1937)

puugwensis’Hori.: 165 (1937)

strictalAtV. 321 (1914)

gerarrdii Baker: 1 1 0 ( 1 878b); Les, S A, Swa

junodii Baker: 859 ( 1901 ) as junodi

goetzei Hanus: 276 (1901); DRC, K, Mai, Moz, T, Zam, Zim

eia//e«to De Wild.: 537 (1913)

rubiginosalAeV. 320 (1914)

turbinata Nel: 329 (1914)

graminea Willd. ex Schult.: 768 (1829); Mad

gregoriana Rendle: 407 (1895); K

hemerocallidea Fisch., C.A.Mey. & Ave-Lalf: 64, 65 (1842); Bot,

Les, Moz, SA, Swa, Zim

elata Hook.f: t. 5690 (1868), non Schult.f

obconica Nel: 318 (1914)

pam/flNel: 333 (1914)

phoenica Nel, ined, based on specim. Wood 184 (K), nom. nud.

rigidida Baker var. hemerocallidea (Fisch., C.A.Mey. & Ave-Lall.)

Heideman: 892 (1983)

rooperii T.Moore: 65 ( 1 852) as rooperi

rooperii T.Moore var. forbesii Baker: 1 1 8 ( 1 878b)

hockii De Wild.: 537 (1913)

pedicellata Nel ex De Wild.: 315 (1914)

var. hockii Wiland: 321 (2001); DRC

var. colliculata Wiland: 321 (2001); DRC

var. katangensis (Nel) Wiland: 322 (2001); DRC, Zam

katangensis Nel ex De Wild. : 312(1914)

infausta Nel: 319 (1914); T

interjecta Nel: 321 (1914); SA

pretoriensis Goossens, ined. description attached to specim. Goos-

sens 91 (K), nom. nud.

kilimanjarica Baker: 378 (1898)

subsp. kilimanjarica; B, DRC, K, Rwa, T

alpina R.E.Fr.: 78 (1948)

iucisa Nel: 301 ( 1914)

subsp, prostrata E.M.Holt & Staubo in Nordal et al.: 26 ( 1985); K, T

kraussiana Buchinger: 311 (1845); DRC?, SA, Swa

laikipiensis Rendle: 407 (1895); T?

lejolyana Wiland: 418 (1997b); DRC, Zam

longifolia Baker ex Hook.f: t. 6035 (1873); Les, Moz, SA

filifolia Nob., ined. name applied to specim. Mimd & Maire s.n. (B)

by Nel, nom. nud.

longifolia Baker var. thunbergii Baker: 1 16 (1878b)

villosa L.f var. 5 Thunb. ex Baker: 1 16 (1878b)

zuluensis S.E.Wood, ined. name applied to specim, Gerstner 4936

(PRE)

zululandensis S.E.Wood: 66 ( 1976), MS.*

ludwigii Baker: 181(1 876); Les, SA

lusalensis Wiland: 421 (1997b); DRC

malaissei Wiland: 418 (1997b); DRC

malosana Baker: 284 (1897); B, Bot, DRC, Mai, Moz, SA?, T, U,

Zam, Zim

hiflora De Wild. : 537 (1913), nom. illeg.

matengensis G.M. Schulze: 376 (1939); T

membranacea F3aker: 106 (1878b); A, Swa

monanthos Baker: 266 (1878a); A, B, DRC, Mai

Bothalia 36,1 (2006)

15

muhilensis Wiland: 412 (1997b); DRC

subsp. kansimbensis Wiland: 414 (1997b); Z

subsp. muhilensis Wiland; 412 (1997b); Z

multiceps Buchinger: 31 1 (1845); Les, SA, Swa

neliana Schinz: 136 (1926); Les, SA

nyasica Baker: 284 (1897); Mai, Moz, T, Zam, Zim

canaliculata sensu Brenan: 86 (1954) quoad Brass 1 7598

engleriana'Nel: 315 (1914)

engleriana Nel var. sco/»7Nel: 315 (1914)

ingrafa'NeV. 311 (1914)

probata^eV. 317 (1914)

rerracto Nel: 312 (1914)

v///osa sensu Zimudzi: 17 (1996)

obliqua Jacq.: 54 (1796): t. 371 (1786-1793); Les, SA

villosa var, obliqua (Jacq.) Baker: 1 14 (1878b)

obtusa Burch, ex Ker Gawl.: t. 159 (1816); A, Bot, K, Les, Mai, Moz,

Nam, SA, Swa, T, Zam, Zim

iridifolia Baker; 117 (1878b)

nitida l.Verd.: t. 1058 (1949)

obtusa Burch. MS. ined. et auct. Plur. — vix Ker Gawl.

obtusa Burch, ex Ker Gawl.

var. chrysotricha Nel; 334 (1914)

var. nitida (l.Verd.) Heideman: 892 (1983)

var. obtusa'. Heideman: 892 (1983)

villosa L.f. var. obtusa (Burch, ex Kew Gawl.) T.Durand & Schinz:

236 (1895)

villosa sensu Eyles: 328 (1916), quoad specim. Gibbs 192

villosa sensu Zimudzi: 17 (1996)

parvifolia Baker: 183 (1896); Mai, SA, Swa, Zim

parvula Baker: 113 (1878b)

var, parvula; Les, SA, Swa

brevifolia Bstker: 183 (1896)

limicola B.L.Burtt: 188 (1988)

membranacea auct. non Baker

var. albiflora B.L.Burtt: 190 ( 1988); SA

polystachya Welw. ex Baker: 266 ( 1 878a)

completa Nel, ined, name applied to specim. Allen 30 (B), nom.nud.

multiflora Nel: 317 (1914)

obtusa auct. non Burch, ex Ker Gawl. (Nordal & Zimudzi 200 1:14)

orbiculatalAQV. 313 (1914)

var. polystachya; A

var. andongensis Baker: 266 (1878a); A

protrusa Nel: 336 (1914); T

rigidula Baker: 116 (1878b)

acuminata sensu Norl: 163 (1937), quoad specim. 4833

longifolia Dinter ex Solch: 2 (1960), nom. nud.

oblongaUtV. 332 (1914)

obtusa auct. non Burch, ex Ker Gawl. (Nordal & Zimudzi 2001 : 12)

volkmanniae Dinter: 257 (1931)

var. pilosissima Baker: 117 (1878b); SA, Swa, Les

arnottii Baker: 552 (1877)

var. rigidula Baker: 116(1 878b); Les, Moz, Nam, SA, Swa, T?, Zim

cordatalAtV. 331 (1914)

elliptical4e.\'. 332 (1914)

longifolia Baker: 176 (1904) based on specim. Jiinod 1445 (Z),

nom. illeg.

rigidula Baker var. hemerocallidea (Fisch., C.A.Mey. & Ave-Lall.)

Heideman: 892 (1983)

robustaNel: 313 (1914); DRC

sagittata Nel: 323 (1914); SA

schimperi Baker: 110 (1878b); Eth

villosa eaxci. non L.f. (Cufodontis 1971: 1578)

sobolifera Jacq.: 53 (1796): t. 372 (1786-1793)

var. pannosa (Baker) Nel: 309 (1914); SA

pannosa Baker: 130 (1874)

villosa L.f. var. pannosa (Baker) Baker: 1 1 4 ( 1 878b)

var. sobolifera (Jacq.) Nel: 309 (1914); SA

canescens Fisch. in Fisch. & C.A.Mey.: 50 (1845)

decumbens P & y Thunb. ex Baker: 114 (1878b)

krebsii Fisch, in Fisch. & C.A.Mey.: 72 (1846)

sclrweinfurthiana Nel: 329 (1914)

sobolifera Jacq, var. accedens Nel: 310 (1914)

villosa L.f. var. canescens (Fisch.) Baker (1878b)

villosa L.f. var, schweinfwthii Harms: 72 (1895)

villosa L.f var. sobolifera (Jacq.) Baker: 114 (1878b)

sp. A Nordal & Zimudzi; 15 (2001); Mai, Moz, Swa?, Zim

stellipilis Ker Gawl,: t. 663 (1822); SA

lanata Eckl. ex Baker: 118 (1878b)

subspicata Pax; 143 (1893); A, DRC, Mai, Zam

sp. A of Nordal et al., (pro parte quoad Davies 742) Nordal et al.:

29 (1985)

suffruticosa Nel: 335 (1914); Cam

urceolata auct. non Nel (Nordal & Iversen 1987: 37)

symoensiana Wiland: 421 (2001); DRC

tetramera Hilliard & B.L.Burtt: 299 (1983); Les, SA

uniflorata Markotter: 15 (1930); SA

upembensis Wiland: 414 (1997); DRC

urceolata Nel: 336 (1914); DRC, Rwa, Zim

apiculatalAel'. 327 (1914)

bequaertii De Wild.: 49 (1921b)

crispa Nel: 334 (1914)

villosa L.f: 326 (1781) sensu lato; K?, Les, Mai, Moz, SA, Swa, Moz,

Zam, Zim

abyssinica Hochst.: 32 (1844)

boranensis Cufod.: 328 (1939)

bowriana Baker, ined. name applied to specim. Bowie s.n. (BM),

nom. nud,

decumbens Lam.: 172 (1789)

jacquinii Baker: 112 (1878b)

microsperma Lallem. in Fisch. & C.A.Mey.: 50 (1845)

petitiana A. Rich.: 315 (1851)

scabraLoAd.'. t. 970 (1824)

simensis Hochst.: 32 (1844)

textilis Nel: 326 (1914)

tomentosa Lam.: 112 (1789)

tysonii Schonland ex Bruce-Miller: 36 (1995), MS.*, nom. nud.

var.yimin'ata Nel: 310 (1914)

var. scabra (Lodd.) Baker: 1 14 (1878b)

volkensii Harms ex Engl.: 733, 734 (1906), nom. nud.

zernyi Schulze: 375 (1939); T

zeyheri Baker: 112 (1878b); SA

setosaBdktx'. 113 (1878b)

* MS., manuscript.

TABLE 1 . — Synonyms in Hypoxis and reference to new combinations by various authors

16

Bothalia 36,1 (2006)

TABLE 1. — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

Bothalia36, 1 (2006)

17

TABLE 1. — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

Bothalia 36,1 (2006)

TABLE 1 . — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

Bothalia 36,1 (2006)

19

TABLE 1 . — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

C.A.Mey. & Ave-Lall.) Heideman based on

Junod 1445 (Z)

20

Bothalia36,l (2006)

TABLE 1. — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

Bothalia36,l (2006)

21

TABLE 1 . — Synonyms in Hypoxis and reference to new combinations by various authors (cont.)

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Bothalia 36,1; 25-32 (2006)

Sesotho names for exotic and indigenous edible plants in southern

Africa

A. MOTEETEE*t and B-E. VAN WYK*

Keywords: edible plants, exotic plants, indigenous plants, Sesotho, southern Africa

ABSTRACT

A comprehensive checklist of Sesotho names of both indigenous and exotic food plants is presented, based on a literature

survey and the first author’s experience. The list includes the scientific names, English common names and parts of the plant

that are used. Where possible, the origin or meaning of the Sesotho names is given. Exotic edible plants for which the English

common names are in general use have been excluded. The list includes 164 indigenous and 39 names of exotic species.

INTRODUCTION

The aim of this paper is to compile, for the first time,

a comprehensive checklist of Sesotho names of both

indigenous and exotic edible plants. With diminishing

vocabulary of many languages (Sesotho included), it has

become necessary to preserve and update these names so

that they are not lost. We believe such a list would be of

value to translators and compilers of dictionaries.

The extensive use of wild plants as food in southern

Africa has been well documented in a number of publi-

cations including the books of Fox & Norwood Young

(1982), Peters et al. (1992), and Van Wyk & Gericke

(2000). Lists of common names of plants in vernacular

languages have also been written. For example, a com-

prehensive list of Setswana common names of plants

by Cole (1995) is available. A list of some Zulu names

has also been published (Ngwenya et al. 2003). Phillips

(1917) and Jacot Guillarmod (1971) published Sesotho

names of indigenous plants. Smith (1966) also included

some Sesotho names. However, the lists are not com-

prehensive and do not include Sesotho names for exotic

food plants.

Many of the scientific names used in these publica-

tions are now outdated and it has become necessary to

update them. Furthermore, the list of Phillips (1917) is

only applicable to the Leribe Plateau region in Lesotho.

While Jacot Guillarmod’s (1971) work is much more

comprehensive, it did not include the meanings of the

Sesotho names. This checklist has been compiled based

on plants used by Sesotho-speaking people in southern

Africa, who live mainly in Lesotho and in the Free State,

South Africa.

MATERIALS AND METHODS

The checklist was compiled using the available litera-

ture and the first author’s own experiences while growing

up in a rural area (Ha Thuube, Qacha’s Nek District) in

Lesotho. Authorities for scientific names are given in

* Department of Botany and Plant Biotechnology, University of

Johannesburg, P.O. Box 524. 2006 Auckland Park, Johannesburg,

t Corresponding author e-mail address: amoti@rau.ac.za

MS. received: 2005-04-26.

Appendix 1 and are not repeated elsewhere in the text.

The nomenclature follows that of Germishuizen & Meyer

(2003).

The list is arranged alphabetically according to family

and genus starting with the monocots. Where possible,

English common names and the origin or meaning of

the Sesotho names are presented. The edible parts of the

plants have also been included in the list. Many exotic

plants including fruits (e.g. banana, kiwi, mango, among

others), nuts (e.g. almonds, cashews, macadamias, among

others), spices and herbs (e.g. coriander, oregano, parsley,

and others) do not have Sesotho names and are therefore

excluded.

DISCUSSION

Sesotho names of 164 indigenous and 39 exotic plant

species of edible plants are given in Appendix 1 .

Some Sesotho names of plants are expressed in short

sentences to indicate either the habit of the plant (e.g.

its appearance, striking features or size), habitat, or its

use (mostly for medicinal purposes). Many exotic plants

also have Sesotho names, although these names have

been taken directly from either English or Afrikaans.

For example, the Sesotho name tamati for tomato

{Lycopersicon escidentim) has been taken directly from

the Afrikaans word tamatie. To highlight the typical deri-

vation of names, some examples of plant names are given

below (for full meanings see Appendix 1).

Names based on characteristics of the plant

Diospyros austro-africana: Ntlo-ea-lekhoaba — the house

of a crow, with reference to its flower shape.

Empodium plication: Leihlo-la-khomo — eye of the cow,

so named because its flower resembles an eye of a

cow.

Sisymbrium capense: Tlhako-ea-khomo — hoof of the

cow, the shape of the leaves resembles a hoof of a

cow.

Watsonia densiflora (and other members of Iridaceae):

Khahla — the flower which pleases, this is beeause

the flowers of these plants look pretty.

Tragopogon porrifolius: Moetse-oa-pere — mane of a

horse, in reference to the thin, elongated leaves.

26

Bothalia 36,1 (2006)

Names based on the size of the plant

Gladiolus dalenii: Khahla-e-kholo — the big plant/flower

which pleases.

Hesperantha baurii: Khahla-e-nyenyane — the small

plant/flower which pleases.

Hypoxis argentea: Leihlo-la-khomo-le-leholo — the big

eye of the cow.

Mentha aquatica: Koena-e-nyenyane — the small croco-

dile.

Names based on the habitat of the plant

Eiilophia Mans: Mametsana-a-manyenyane — mother of

little water, because the plants are found in wet

places.

Gladiolus cruentus: Khahla-ea-loti — the mountain plant

which pleases, so called because the plants usually

grow in mountainous areas.

Oxalis corniculata: Bolila-ba-thaba — the sour plant of

the mountain; these plants are more frequent at

higher altitudes.

Names based on the taste of the plant

Epilobium hirsutum: Letsoai-la-balisana — salt of the

shepherds, so called because the leaves taste salty.

Nasturtium officinale: Liababa — they are bitter, due to

the bitter leaves.

Oxalis species: Bolila — the sour plant, because the

leaves are extremely sour.

The replacement of indigenous food plants with

seemingly more attractive exotic ones is just one aspect

of a broader pattern of aculturization that affects many

aspects of Sesotho culture. The impoverishment of

Sesotho plant nomenclature is very evident in rural areas

where only the elderly still know and use the traditional

names of plants.

ACKOWLEDGEMENTS

The second author acknowledges financial support

from the National Research Foundation (NRF).

REFERENCES

BRUMMITT, R.K. & POWELL, C.E. 1992. Authors of plant names.

Royal Botanic Gardens, Kew.

COLE, D.T. 1995. Setswana animals and plants. Macmillan Publishing,

Botswana.

FOX, F.W. & NORWOOD YOUNG, M.E. 1982. Food from the veld:

edible wild plants of southern Afiica. Delta Books, Craighall,

Johannesburg.

GERMISHUIZEN, G. & MEYER, N.L. (eds). 2003. Plants of southern

Africa: an annotated checklist. Strelitzia 14. National Botanical

Institute, Pretoria.

JACOT GUILLARMOD, A. 1966. A contribution towards the econom-

ic botany of Basutoland. Botaniska Notiser 119: 209-2 1 1 .

JACOT GUILLARMOD, A. 1971. Flora of Lesotho. Cramer, Lehre.

MASEFIELD, G.B., WALLIS, M., HARRISON, S.G. & NICHOLSON,

B.E. 1969. The Oxford book of food plants . Oxford University

Press, Great Britain.

NGWENYA, M.A., KOOPMAN, A. & WILLIAMS, R. 2003. Zulu

botanical knowledge: an introduction. National Botanical

Institute, Durban.

PETERS, C.R., O’BRIEN, E.M. & DRUMMOND, R.B. 1992. Edible

wild plants of sub-Saharan Africa. Royal Botanic Gardens,

Kew.

PHILLIPS, E.P. 1917. A contribution to the flora of the Leribe Plateau

and environs. Annals of the South African Museum 16: 1-379.

SMITH, C.A. 1966. Common names of South African plants. Memoirs

of the Botanical Survey of South Africa No. 35. Botanical

Research Institute, Pretoria.

VAN WYK, B-E. & GERICKE, N. 2000. People’s plants. Briza

Publications, Pretoria.

APPENDIX 1, — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references

* exotic taxa; 1 , Phillips ( 191 7); 2, Jacot Guillarmod (1971 ); 3, Fox & Norwood Young (1982); 4, Peters et al. (1992); 5, Masefield et al. (1969);

6, Van Wyk & Gcricke (2000); 7, Jacot Guillarmod ( 1966). Author abbreviations according to Brummitt & Powell 1992.

Bothalia36,l (2006)

27

APPENDIX 1 . — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

* exotic taxa; 1 , Phillips ( 1 9 1 7); 2, Jacot Guillarmod ( 1 97 1 ); 3, Fox & Norwood Young ( 1 982); 4, Peters et al. ( 1 992); 5, Masefield et al. ( 1 969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

28

Bothalia36,l (2006)

APPENDIX 1. — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

* exotic taxa; 1, Phillips (191 7); 2, Jacot Guillarmod ( 1971); 3, Fox & Norwood Young (1982); 4, Peters e! al. (1992); 5, Masefield et al. (1969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

Bothalia36,l (2006)

29

APPENDIX 1 . — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

* exotic taxa; 1 , Phillips ( 1 9 1 7); 2, Jacot Guillarmod ( 1 97 1 ); 3, Fox & Norwood Young ( 1 982); 4, Peters et al. ( 1 992); 5, Masefield et al. ( 1 969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

30

Bothalia36,l (2006)

APPENDIX 1 . — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

* exotic taxa; I, Phillips (191 7); 2, Jacot Guillarmod (1971 ); 3, Fox & Norwood Young (1982); 4, Peters et al. (1992); 5, Masefield et al. (1969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

Bothalia 36,1 (2006)

APPENDIX 1 . — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

31

* exotic taxa; 1, Phillips (1917); 2, Jacot Guillarmod (1971); 3, Fox & Norwood Young (1982); 4, Peters et al. (1992); 5, Masefield et al. (1969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

32

Bothalia 36,1 (2006)

APPENDIX 1. — List of edible plants; their common names (Sesotho and English); the parts that are used for food; and references (cont.)

* exotic taxa; 1 , Phillips (1917); 2, Jacot Guillarmod (1971 ); 3, Fox & Norwood Young (1982); 4, Peters e! al. (1992); 5, Masefield et al. (1969);

6, Van Wyk & Gericke (2000); 7, Jacot Guillarmod (1966). Author abbreviations according to Brummitt & Powell 1992.

Bothalia 36,1; 33-37 (2006)

Two new species of Erica (Ericaceae) from the Langeberg, Western

Cape, South Africa

R.C. TURNER* and E.G.H. OLIVER*

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

ABSTRACT

Two new species of the genus Erica L. from the north-facing slopes of the Langeberg are described — E. turneri, known

only from the type locality on Zuurbraak Mountain and E. euryphylla, occurring on the same mountain slope, as well as on

the middle north-facing slopes of Hermitage Peak near Misty Point in the Marloth Nature Reserve above Swellendam.

INTRODUCTION

The two species described in this paper are placed in

section §Ceramia in which there are many species asso-

ciated with damp, shaded or wet habitats. Most are soft,

low shrublets, either erect and compact or difftise and

sprawling, with long, delicate main branches and often

with open-backed leaves (Oliver & Oliver 2002). Both

new species possess recaulescent bracts and bracteoles

and have broad, flat leaves with distinctly thirmed mid-

ribs towards the apices, allying them morphologically

with E. oxycoccifoUa and E. cordata respectively, rather

than with the E. planifolia group, in which the bract is

not recaulescent and is leaf-like and the leaves have api-

cally thickened midribs.

Erica turneri E.G.H. Oliv., sp. nov., foliis 3-natis,

ramis foliis bractea bracteolisque pilis glandulosis sim-

plicibusque, bractea recaulescenti, corolla ± 3-4 x 2.5-

3.5 mm pilis brevibus simplicibus, ovario pilis sparsis.

Figura 1.

TYPE. — Western Cape, 3320 (Montagu): Langeberg

Range, Zuurbraak Moimtain west of Tradouw Pass, north-

facing slopes above Farm Sandrift, 774 m, (-DC), 30

August 2003, Turner 792 (NBG, holo.; BOL, K, iso.^

Plants up to 450 mm tall, laxly erect to sprawling,

entwined, single-stemmed reseeders. Branches: several

lax, spreading main and numerous entwined secondary

branches; stems, younger and older with sparse, short,

simple and gland-tipped hairs, no infrafoliar ridges,

intemodes + 5-10(-17) mm long. Leaves 3-nate, ovoid to

obovoid, ± 3^ X 1.0-2. 5 mm, flat, open-backed, abaxi-

ally with short, simple and gland-tipped hairs, midrib

slightly thickened in basal and median portions, adaxi-

ally with short, simple and occasional short, gland-tipped

hairs, green, margins slightly thickened abaxially, ciliate,

with short simple and gland-tipped hairs; petiole ± 0.5-

0.75 mm long, yellowish green. Inflorescence: flowers

1 to 3-nate in 1(2) whorls at ends of main branches and

secondary branchlets, the latter long or highly reduced;

pedicel ± 8 mm long, green turning reddish, with short,

simple and gland-tipped hairs; bract partially recaules-

cent, basal to median, leaf-like, narrowly obovate, ±

*Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

MS. received: 2005-07-29.

0.7 mm long, open-backed, abaxial and adaxial surfaces

and margins with short, dense, simple and gland-tipped

hairs, pale green; bracteoles 2, basal to median, longer

than bract, linear, ±0.8 mm long, open-backed, leaf-like,

abaxial and adaxial surfaces and margins with short,

dense, simple and gland-tipped hairs, pale green. Calyx

4-lobed; sepals adpressed, ovate, open-backed, ± 1. 5-2.0

mm long, adaxially glabrous, abaxially with short, sim-

ple and gland-tipped hairs, margins slightly thickened,

ciliate, with short simple and gland-tipped hairs, green.

Corolla 4-lobed, broadly cup-shaped to slightly ovoid,

3-4 X 2. 5-3. 5 mm, with short simple hairs (± 0. 1-0.4

mm long), translucent white tinged pink in upper half

and lobes, becoming deeper pink upon exposure to sun-

light and when older, lobes erect, acute, margins smooth.

Stamens 8, free, included; filaments linear, ± 1.5 mm

long, with a slight apical bend, glabrous, white; anthers

dorsally fixed at base, bipartite, thecae erect, subfalcately

rectangular-elliptic in lateral view, ± 0. 8-1.0 mm long,

sparsely aculeate, golden brown with reddish tinge on

dorsal ridge; appendages pendulous, dentate, narrowly

obcuneate, ± 0.7 mm long, sparsely aculeate, dorsally

fixed at bases of thecae, white, often tinged red, pores

± 0.25-0.3 mm long; pollen in tetrads. Ovary 4-locular,

subturbinate, slightly flattened, ± 0.85 mm long, with

sparse lanate hairs, green turning red; ovules 7 or 8 per

locule, placenta apical, nectaries, basal, green; style

filiform, ± 2 mm long, glabrous, exserted, occasionally

with sparse lanate hairs on upper 0.4 mm, white to pale

green, tinged red apically; stigma subcapitate, reddish.

Fruit a dehiscent capsule, broadly cylindric-ellipsoid,

± 0.85 mm long, sparsely lanate, pale cream-coloured,

valves thin and brittle, spending to ± 45°, septa ± 60% on

valve, 40% on columella. Seeds ellipsoid, ± 0.3 mm long;

testa yellowish brown, smooth, shiny; cells irregularly

elongate, 50-80 x 25^0 pm, anticlinal walls unevenly

jigsawed, periclinal walls with numerous small pits.

Flowering time: May to August. Figure 1.

Diagnostic features and discussion: Erica turneri is

remarkably similar in superficial appearance to E. oxy-

coccifolia Salisb., a species endemic to the Cape Peninsula,

as well as to the following species described in this paper,

E. euryphylla. Character similarities include 3-nate, broad,

open-backed leaves, 3-5 x 1-3 mm, small, finely hairy,

cup- to open cup-shaped, white to pink flowers, 3^ mm

long, as well as a soft, lax, spreading, intertwined habit.

Upon closer inspection however, E. turneri possesses

FIGURE 1 . — Erica titmeri. Turner 792. A, flowering branch, natural size; B, stem; C, leaf, abaxial view; D, flower; E, bract; F, bracteole; G, sepal,

abaxial view; H, stamen, front and side views; I, gynoecium; J, ovary with one valve removed; K, testa cells. Scale bars: B-D, G-J, 2 mm;

E, F, 1 mm; K, 50 pm. Artist: R.C. Turner.

stems with short, simple and gland-tipped hairs, abaxial

leaf surfaces with short, simple and gland-tipped hairs,

pedicels with short, predominantly gland-tipped, occa-

sionally simple hairs, a turbinate, sparsely hairy, 4-locular

ovary with 7 ovules per locule, and sparsely aculeate,

prognathous anthers with ± 0.7 mm long, pendulous,

dentate appendages. E. oxycoccifolia differs in having a

glabrous stem, glabrous abaxial leaf surfaces, glabrous

pedicels, smooth, muticous anthers and a globose, gla-

brous, 4-locular ovary with 10 ovules per locule, whereas

E. eitryphylla has stems with long, lanate and occasionally

gland-tipped hairs, abaxial leaf surfaces with long woolly

hairs, pedicels with long, lanate and occasionally gland-

tipped hairs, smooth, muticous anthers, and a globose,

4-locular ovary with 1 6 ovules per locule.

Pollination syndrome: the pollination syndrome of

Erica turneri is unresolved. The presence of anther

appendages and well-developed nectaries suggest ento-

mophily, although no potential pollinators, flying or crawl-

ing, have been observed during visits to the ten known

stands. It seems improbable, however, that a flying insect

would be able to penetrate the tangled, glandular leaves

and stems of the species, to reach flowers that are often

pressed against rock faces, or entirely contained by the

plants’ aforementioned habit. Preliminary studies into the

pollination syndromes of E. limosa L.Bolus and E. salteri

L. Bolus, suggest that large ants may play an important role

in the pollination of moisture-loving Erica species with a

low, diffuse, entwined habit, as well as with small (1-5

mm long), cup-shaped flowers (Turner pers. obs.).

Distribution and habitat: Erica turneri appears to be

confined to the catchment area of the Klein-Sandrivier on

the middle north-facing slopes of Zuurbraak Mountain in

the Langeberg Range, ± 1 1 km southwest of Barrydale

(Figure 2) (Turner pers. obs.). The species has been seen

at altitudes ranging from 580-950 m, on a substratum of

quartzitic Table Mountain Sandstone (Turner pers. obs.).

Plants occur in seasonally damp or wet, mostly shady

crevices and recesses at the bases of rocks and rock

FIGURE 2. — Known distribution of Erica turneri.

Bothalia 36,1 (2006)

35

ledges. This type of microhabitat often provides only

a small amount of derived quartzitic sand and darker,

peatty, organically derived accumulate in which plants

may grow. It is unusual, although not unique, for an

Erica species with a delicate growth form and markedly

open-backed leaves such as E. lurneri, to inhabit hotter,

drier, north-facing mountain slopes, as well as to enjoy

a generally north-facing aspect. However, the species

grows in seasonally wet or damp, mostly shady crevices

and recesses at the bases of rocks and rock ledges, result-

ing in the majority of specimens receiving potentially

direct sunlight only in autumn and midwinter, the period

over which the species flowers. Morphologically allied

taxa such as E. oxycoccifolia, E. physophylla Benth. and

E. utriculosa L.Bolus, which are by contrast confined to

damp, shady, montane, south-facing habitats on gener-

ally wetter, south-facing slopes, would receive their most

direct sunlight as well as optimum moisture availability

in spring to early midsummer, the seasons in which their

flowering occurs. Although the south and upper north-

facing slopes of Zuurbraak Mountain receive much

precipitation and resulting seepage, the middle and lower

north-facing slopes are comparatively dry.

Stands of E. tumeri growing in the non-perennial,

eastern tributaries of the Klein-Sandrivier do not receive

direct runoff or seepage from the wetter, upper slopes,

and plants in these more exposed habitats display red-

dish green leaves and produce flowers with a pink tinge

at an earlier stage than well-shaded specimens. Stands

growing in the Klein-Sandrivier Kloof enjoy a cooler,

moister microclimate and the only south-facing speci-

mens of the entire population are found in this kloof, in

a steep, damp side gully. Here plants form low, matted

‘hedgerows’ at the base of rock faces, individual plants

attaining dimensions of up to 450 x 400 mm. Given the

populations’ general aspect and habitat however (Turner

pers. obs.), it is apparent that the species is capable of

surviving some relatively dry periods. It is therefore

surprising, given its postulated relationship with other

species of §Ceramia, that in this situation, the species

has almost totally open-backed leaves. In its specialized

microhabitat, E. turneri grows in association with other

delicate, lax, shade-loving species such as Troglophyton

capillaceum (Asteraceae), Centella macrodis (Apiaceae)

and Gleichenia polypodioides(G\eicheniaceae), as well

as other moisture-loving species such as Lobelia neglecta

(Lobeliaceae), Drosera capensis (Droseraceae), Todea

barbara (Osmundaceae), E. cajfra L., E. cubica L.,

E. hispidula L., E. tenuis Salisb. (Ericaceae), Berzelia

lanuginosa and Raspalia virgata (Bruniaceae) and sev-

eral low Restionaceae species. The following species

described in this paper, E. euryphylla, although growing

on the same mountain slope, does not occur in direct

association with E. turneri. Furthermore, the north- and

south- facing slopes of Zuurbraak Mountain together sup-

port at least 34 Erica species (Turner pers. obs.), nine of

these endemic to the Langeberg.

Etymology, this species is named after its discoverer,

Ross Turner, who is also the co-author of this paper. He

has devoted considerable time and energy to the tracking

down, recording and studying of many Erica species,

especially the rare ones.

Paratype material

WESTERN CAPE. — 3320 (Montagu); Langeberg Mountains,

Zuurbraak Mountain west of Tradouw Pass; steep gully in Klein-

Sandrivier Kloof, above Farm Sandrift, 616 m, (-DC), 26 August 2004,

Turner 107 (NBG).

Erica euryphylla R.C. Turner, sp. nov., folds 3-

natis costa in mediano basique parum crassiore, ramis

foliis bractea bracteolisque pilis brevibus sparsis glan-

dulosis et simplicibus, bractea recaulescenti, corolla ±

3-4 X 2. 5-3. 5 mm pilis brevibus simplicibus et interdum

pilis glandulosis, antheris muticis interdum calcaribus

minutis, ovario pilis sparsis. Figura 3.

TYPE. — Western Cape, 3320 (Montagu): Swellendam

Hiking Trail, in shelter of overhanging rock along path

between Boskloof and Goedgeloof huts, Langeberg Moun-

tains, 3500-4000 ft [1 060-1 220 m], (-CD), 5 December

1983, Esterhuysen 36152 (BOL, holo.; NBG, iso.).

Plants up to 300 mm tall, soft, laxly erect, single-

stemmed reseeders. Branches: several, erect, lax main

with short, lax, secondary branches; stems, younger and

older with sparse, short, simple and gland-tipped hairs,

no infrafoliar ridges, intemodes ± 5-10(-14) mm long.

Leaves 3-nate, ovate to obovate, 3-5 x 1-3 mm, flat,

open-backed, green, abaxially with long, dense, woolly

hairs, midrib slightly thickened especially in basal and

median portions, adaxially with long, simple and gland-

tipped hairs, margins slightly thickened abaxially with

long, simple and gland-tipped hairs; petiole ± 0. 8-1.0 mm

long, yellowish green. Inflorescence: flowers (l-)3-nate in

1 to 5 whorls, umbel-like at ends of main and secondary

branchlets; pedicel ± 3. 5-5.0 mm long, rosy pink, with

long, lanate, simple and gland-tipped hairs; bract par-

tially recaulescent, basal to median, when basal leaf-like,

± 2-3 X 1-2 mm, open-backed, green, abaxially with long,

woolly, simple hairs and adaxially with long, simple and

gland-tipped hairs, when median narrowly obovate, ±0.7

X 0.5 mm, abaxially glabrous, adaxially with long, simple

and gland-tipped hairs and margins with long, simple

and gland-tipped hairs; bracteoles 2, ± median, narrowly

obovate, ± 0.8 mm long, slightly longer than bract when

bract in median position, partially open-backed, abaxially

glabrous, margins and adaxial surface with long, simple

and gland-tipped hairs. Calyx 4-lobed; sepals partially

fused at bases, adpressed, narrowly spathulate, ± 1. 2-2.0

mm long, open-backed, pink, green apically, abaxially

with short, woolly and long, simple and gland-tipped

hairs, adaxially glabrous, margins slightly thickened with

long, pilose hairs with red apical glands. Corolla 4-lobed,

open cup-shaped to slightly um-shaped, 3-4 x 2. 5-3. 5

mm, viscid, with sparse, simple and occasionally gland-

tipped hairs, ± 0. 1-0.4 mm long, pale to rosy pink, mar-

gins smooth, lobes erect, acute, entire. Stamens 8, free,

included; filaments linear, ± 1.8 mm long, erect, with

a slight apical bend, glabrous, white; anthers basifixed,

bipartite, erect, muticous, occasionally with minute, acule-

ate, spreading appendage near apex of filament, glabrous,

golden brown, thecae ± 0. 8-1.0 mm long, pores apical,

ovoid, ± 0.25-0.32 mm long; pollen in tetrads. Ovary 4-

locular, ovoid, ± 1 mm long, viscid, green turning red, with

short, dense, simple lanate and occasionally gland-tipped

hairs; nectaries basal, yellowish green turning red; ovules

± 16 per locule, placenta apical; style simple, + 3 mm

36

Bothalia 36,1 (2006)

FIGURE 3. — Erica euryphylla. Turner 799. A, flowering branch, natural size; B, stem; C, leaf, abaxial view; D, flower; E, bract; F, bracteole; G,

sepal, abaxial view; H. stamen, front, side and back views; I, anther, side and back views showing minute appendages; J, gynoecium; K,

ovary with one valve removed; L, testa cells. Scale bars: B-D, H, I, 2 mm; E-G, J, K, 1 mm; L, 50 pm. Artist: R.C. Turner.

long, exserted, glabrous, pink; stigma capitellate, reddish

purple. Fruit a dehiscent capsule, ± 1 mm long, ovoid,

pale brown to cream-coloured with short, lanate hairs,

valves spreading to ± 45°, thin, brittle, septa ± totally on

valve. Seeds ellipsoid, ± 0.4 mm long; testa smooth, pale

creamish brown, cells subequal to slightly elongate, 50-70

X 20^0 pm, anticlinal walls slightly thickened, unevenly

undulate, inner periclinal walls with numerous small pits.

Flowering time: May to September. Figure 3.

Diagnostic features and discussion: within §Ceramia,

Erica euryphylla shares several characters in common

with species of the E. cordata complex, in particular E.

cordata Andrews, E. macrophylla Klotzsch ex Benth. and

E. ocellata Guthrie & Bolus. These species also possess

3-nate, broad, open-backed leaves with densely woolly

abaxial surfaces, adaxial surfaces with long, simple and

gland-tipped hairs and a thickened midrib in the basal and

median portions but in all instances not protruding beyond

the lamina at the apex of the leaf, stems with long, simple

and gland-tippe*d hairs, sepals with red, stalked glands

on the margins, glabrous adaxial surfaces, abaxial sur-

faces with long, simple and gland-tipped hairs, as well as

woolly hairs in the sulcus, globose, 4-locular ovaries with

lanate hairs, and manifest, muticous anthers. Variation in

the anther morphology of E. euryphylla has been noted

however, with the occasional flower displaying two

minute, aculeate, spreading appendages on the margins

of the filament just below the attachment to the thecae.

These vestiges of appendages are only clearly visible at a

magnification of 25x or more. Such variation pertaining to

the presence or absence of anther appendages within a spe-

cies is not unique, examples being E. anguliger (N.E.Br.)

E.G.H.Oliv. in which appendages may be present or absent

within a single flower (Oliver 2000) and E. argentea

Klotzsch ex Benth. (Turner in prep.). Significant char-

acter differences separating E. euryphylla from the E.

cordata complex include a sparsely hairy, cup- to shortly

um-shaped corolla; variation in the type and placement of

the bract on the pedicel (either partially recaulescent, basal

to median and reduced, or partially recaulescent, basal and

leaf-like) and leaves with only very slightly thickened and

rolled-under margins as well as less dense, woolly hairs

on the abaxial surface. The habit is generally far more

lax than those of the compared species, with the excep-

tion of E. ocellata, which may have a sprawling habit

when mature (Turner pers. obs.). Character similarities

and differences between E. euryphylla and E. turneri are

discussed under the latter species in this paper.

Pollination syndrome: the pollination syndrome of E.

euryphylla is unknown. Although the species has mostly

muticous anthers, only occasionally displaying minute

anther appendages, it does possess well-developed nec-

taries and a capitellate stigma, suggesting some form of

entomophily. A lack of obvious wind-borne pollen dis-

charge when the plants are disturbed, as occurs in wind-

pollinated species such as E. hispidula L. and E. muscosa

(Sol.) E.G.H.Oliv., and the colour of the species’ flowers,

suggest that it is not wind pollinated.

Bothalia 36,1 (2006)

37

Distribution and habitat. E. euryphylla appears to be

endemic in the middle and upper north-facing slopes of

the Langeberg Range, on Zuurbraak Mountain and in

the Marloth Nature Reserve (Figure 4). It was recorded

at the latter locality by Elsie Esterhuysen in 1983 on the

Swellendam Trail between Boskloof and Goedgeloof huts

and by Dave McDonald in 1989 from the western end of

the Langkuilen Valley near Misty Point. Both Esterhuysen’s

and McDonald’s specimens were found growing on similar

north-facing slopes, in near-identical habitats to specimens

from Zuurbraak Mountain — ‘in shelter of overhanging rock’

and ‘in deep shade of rocks’ respectively, approximately 15

km west of Zuurbraak Mountain. McDonald cited a latitude

and longitude with his collection and a projection of this

point would appear to place Esterhuysens’ collection no

more than 2 km distant.

E. euryphylla has been recorded at altitudes between

1 060-1 340 m, always on a substrate of quartzitic Table

Mountain Sandstone (Turner pers. obs.). Plants grow in

pockets of quartzitic sand and darker, peatty, organically

derived accumulate, in seasonally damp or wet, mostly

shady crevices and recesses at the bases of rocks and rock

ledges. McDonald’s specimen indeed cites a substrate of

‘light grey soil with humus’. Associated species are similar

to those of E. tumeri above but with E. ardens Andrews,

E. triceps Link and several low Restionaceae species.

The first stand of the Zuurbraak subpopulation of E.

ewyphylla was found by ecologist and walking partner

Nick Helme, only minutes after E. turner i was discov-

ered by the author of this species.

Etymology: E. euryphylla is named for its broad, open-

backed leaves from the Greek words, eurys, broad/wide,

phyllon, leaf — a character displayed by only a few Erica

species.

Paratype material

WESTERN CAPE. — 3320 (Montagu): Marloth N.R., Langeberg.

western end of Langkuilen Valley on approach to Misty Point, in deep

shade of rocks. 1 300 m. (-CD), 22-11-1989, McDonald 1862 (NBG);

Zuurbraak Mountain, Langeberg, shady rock crevices on upper north-

facing slopes below summit. 1 287 m, (-DC), 30-08-2003, Turner 794

(NBG); ibid., 1 333 m, 31-08-2003, Turner 799 (NBG); ibid., 1312 m,

27-07-2004, Turner 1075 (NBG).

ACKNOWLEDGEMENTS

We wish to thank the Western Cape Nature Conser-

vation Board for permission to collect plants in the prov-

ince and the Buys Family of Sandrivier Farm, Barrydale,

for access to their mountain areas. Thanks from the first

author are due to Nick Helme for the many and con-

tinuing shared journeys made to botanically signifieant

localities throughout the southwestern Cape region, as

well as for his willingness to share his extensive field

knowledge of the Cape Flora.

REFERENCES

OLIVER. E.G.H. 2000. Systematics of Ericeae (Ericaceae-Ericoideae):

species with indehiscent and partially dehiscent fruits.

Contributions from the Bolus Herbarium 19: 1^83,

OLIVER. E.G.H. & OLIVER. I.M. 2002. Six new species of Erica

(Ericaceae) from Western Cape, South Africa. Bothalia 32:

167-180.

TURNER. R.C. in prep. Studies in Erica argentea Klotzsch ex Benth.

Bothalia.

LIST OF ERICA SPP. RECORDED FROM ZUURBRAAK

MOUNTAIN

albescens Klotzsch ex Benth.*

anguliger (N.E.Br.) E.G.H.Oliv.

ardens Andrews *

articularis L.

bracteolaris Lam.

brevifolia Sol. ex Salisb.

caffra L.

cerinthoides L.

chartacea Guthrie & Bolus*

conferta Andrews

corifolia L.

cubica L.

citrviflora L.

daphniflora Salisb.

denticulata L.

dianthifolia Salisb.

euryphylla R.C. Turner*

gracilis J.C.Wendl.

grata Guthrie & Bolus

hispidula L.

imbricata L.

intermedia Klotzsch ex Benth. subsp. intermedia*

longimontana E.G.H.Oliv.*

melanthera L.

midtumbellifera P.J.Bergius

muscosa (Sol.) E.G.H.Oliv.

ocellata Guthrie & Bolus *

plukenetii L. subsp. plukenetii

podophylla Benth. *

polifolia Salisb. ex Benth*

regerminans L.

rosacea subsp. rosacea (L. Guthrie) E.G.H.Oliv.

tenuicaulis Klotzsch ex Benth.

triceps Link

tumeri E.G.H.Oliv.*

versicolor Andrews

FIGURE 4. — Known distribution of Erica euryphylla.

Langeberg endemic species.

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Bothalia 36,1: 39^4(2006)

Two new species of Nemesia (Scrophulariaceae) from southern

Africa

K.E. STEINER*

Keywords: Kamiesberg, Namibia, Nemesia Vent., new species, Richtersveld, Scrophulariaceae, seed dimorphism, southern Africa

ABSTRACT

Two new annual species of Nemesia Vent, are described from southern .frfrica. N. williamsonii is characterized by bright

orange flowers with an inflated yellow palate. It differs from the related N. maxii Hiem by having a spur that projects back-

wards, not downwards, and bracts that are lanceolate with a truncate base, not triangular or cordate. It is unusual for the genus

in having dimorphic seeds. N. williamsonii occurs almost exclusively in the Richtersveld. but has been collected in a few

localities across the Orange River in southern Namibia. N. hemiptera is a delicate, wiry-stemmed annual with small white

flowers. The flowers are characterized by a tiny nipple-like spur and a seed that is winged on only one side. N. hemiptera is

endemic to the Kamiesberg from near Caries to Kamieskroon.

INTRODUCTION

Nemesia Vent, is a genus of ± 60 species of annual

and perennial herbs endemic to southern Africa (Steiner

1994). It has been over a hundred years since the last

revision of the genus (Hiem 1904) and since that time

many new species have come to light (Steiner 1989,

1994). A partial revision for species occurring chiefly

in KwaZulu-Natal has been published (Hilliard & Burtt

1986), but there are many new species that need to be

described from the Cape Floral Region and southern

Namibia, where ±15% of the species occur. The purpose

of this paper is formally to describe two distinctive spe-

cies, one restricted to Northern Cape and one occurring

in the Northern Cape and Namibia.

Nemesia williamsonii K.E. Steiner, sp. nov., N.

maxi Hiem proxima, sed differ! lobomm inferiomm

aurantiaco nec mbro, palato majoribus calcare corollae

inflexo nec deflexo aut recto et bracteis lanceolatis nec

cordatis.

TYPE. — Northern Cape, 2816 (Oranjemund):

Richtersveld National Park, road to Kouams Camp, 2.3

km SE of turnoff to Pokkiespram, ± 180 m, (-BB), 28

Sept. 2002, Steiner 3954 (NBG, holo.; CAS, K, iso.).

Annual herb up to 280 mm tall; stems angular in cross

section with 4 or 5 sides, up to 2.5 mm wide, comers

ridged, glandular pubescent. Leaves simple, opposite,

mostly sessile, lance-ovate to ovate, 10^3 x 5-20 mm,

sparsely glandular pubemlous to nearly glabrous, base

rounded to cuneate, apex acute, margins entire to toothed;

petioles up to 8 mm long, glandular pilose. Flowers

axillary or in lax, terminal racemes, racemes up to 150

mm long; bracts alternate, sessile, lanceolate, lowermost

leaf-like, uppermost reduced to ± 3.6 x 1 mm, base tmn-

cate, apex acute, margins entire; pedicels ± 5-14 mm

long, glandular pubescent. Calyx 5-lobed, central upper

lobe ± 3.5^. 1 X 0.95-1.00 mm, lateral upper lobes ±

3. 3-3. 6 X 0.8-1. 1 mm, lower two lobes ± 2. 8-3. 3 x

1.0-1. 3 mm, all lobes lanceolate, acute, sparsely glandu-

* Department of Botany, California Academy of Sciences, 875

Howard St, San Francisco, CA 94103, USA.

MS. received: 2005-02-21.

lar pilose. Corolla bilabiate, 10.1-16.1 x 9.6-14.6 mm,

upper lip four-lobed, two inner (upper) lobes oblong to

obovate, 2.7-5. 1 x 2. 8-3. 8 mm, base strongly oblique,

apex rounded to emarginate, two outer (lateral) lobes

oblong, 3. 1-5.1 X 3.2^. 1 mm, base strongly oblique,

apex rounded to emarginate; upper lip orange (rarely yel-

low, see Thompson & Le Roux 364) except for a bright

yellow patch (2.8 x 2.8 mm) just above corolla open-

ing, pale orange reverse, lower lip oblong to obcordate,

4. 9-6. 7 X 4. 5-5. 6 mm, orange (occasionally yellow, see

Thompson & Le Roux 364), pale orange reverse, basal

portion inflated into a convex projecting palate; palate

± 2. 9-3. 2 X 3.5^. 1 mm, bright yellow, longitudinally

grooved, glabrous; hypochile (floor of corolla tube) ±

3. 1-3.3 mm long, sides invaginated to form a narrow

channel, base drawn out into a spur, (2.4-)3.1-A.2(-5.3)

mm long, ± straight or curving downward in distal third,

orange-white. Stamens four, whitish, lying in a shallow

depression in upper surface of corolla tube; filaments

of anticous pair (twisted into posticous position) ± 2.5

mm long, ± straight, except at base and apex, glabrous

or with a few glandular trichomes; posticous filaments

± 0.85 mm long, ± straight except at base, sparsely glan-

dular pubescent; anthers 0.5-0.65 mm long, each pair

strongly coherent. Ovary oblong-ovate in outline, 1 .2-1 .3

X 1.0- 1.1 mm, laterally compressed; style ±0.75 mm,

slightly curved, compressed contrary to the ovary, apex

wider than base, lying between anther pairs, stigma a

crescent-shaped apical band. Capsules oblong in outline,

± 4.5-13.1 X 3. 6-6. 9 mm, laterally compressed contrary

to the septum, apex emarginate to bilobed, lobes rounded

to acute. Seeds dimorphic, plants with either winged or

wingless seeds; winged seeds widely ovate, ± 1.8-2. 3

X 1.8-2. 4 mm, brown, verruculate, wing membranous

with numerous parallel, brownish veins, wingless seeds

oblong, ± 1.5-1. 8 X 0. 8-1.0 mm, brown, verrucate.

Flowering time: June to September. Figure 1.

Diagnostic features: Nemesia williamsonii is easily

recognized by its bright orange and yellow flowers. It can

be distinguished from the similar and related N. maxii by

its colour (white vs violet), the difference in orientation

of the spur, and the shape of the bracts. In N. william-

sonii, the spur projects straight backwards or back and

then downwards distally, whereas in N. maxii, the spur

40

Bothalia36,l (2006)

FIGURE 1, — Nemesia wUliamsonii. Steiner 3954 (NBG). A, habit. B-E, flower: B, C, front and rear views; D, E, side views, intact and partially

cut away. F, posticous and anticous stamens; G, pistil; H, calyx; 1, capsule with calyx; J, K, seed: J, winged type; K, wingless type. Scale

bars: A, 10 mm; B-E, 5 mm; F, G, J, K, 1 mm; H, I, 2 mm. Artist: John Manning.

projects straight downwards or down and forwards dis- tme flower colour. N. wilUamsonii exhibits an interesting

tally. The bracts of N. wilUamsonii are lanceolate with a seed dimorphism at the plant level, even within a single

truncate base, whereas those of N. maxii are triangular or population. A plant produces either winged or wingless

cordate. In some individuals of N. wilUamsonii, flowers seeds (Figure IJ, K). The wingless seeds, however, are

turn blue upon pressing, giving a false impression of the not simply winged seeds that lack the wing. Instead,

Bothalia36,l (2006)

41

there are other morphological features that distinguish

the two seed types. The winged seed minus its wing is

smaller than a wingless seed. The body of the wingless

seed is wider and has much larger and broader verrucae

on the seed coat (Figure IK). A similar dimorphism has

been observed in populations of N. anisocarpa E.Mey.

ex Benth. (W. Metelerkamp unpubl.; K.E. Steiner pers.

obs.). The proportion of winged to wingless seeds in

populations of these species remains unknown.

Etymology’: this plant is named in honour of Dr

Graham Williamson who first brought this plant to my

attention and who has made a major contribution to

knowledge of the natural history of the Richtersveld

(Williamson 2000).

Distribution and habitat: Nemesia williamsonii is

a short-lived annual that comes up sporadically after

winter or early spring showers. These showers are often

localized, making it difficult to predict where the plants

will come up. It is known from the northernmost areas of

Northern Cape and adjacent areas in southern Namibia

(Figure 2). It has been collected most commonly from the

FIGURE 2. — Known distribution of Nemesia williamsonii, •; and N.

hemiptera, A , in southern Africa. Border between South Africa

and Namibia is indicated by dotted line.

Richtersveld National Park and from the adjacent Fish

River Canyon Park in southern Namibia. Together, these

areas now form the Ai-Ais/Richtersveld Transfrontier

Conservation Area. In the Richtersveld, the author has

observed N. williamsonii at the base of small, rocky hills

or koppies in or adjacent to dry streambeds or drainage

lines associated with sandy quartzitic soils. Quartzite

pebbles and stones were obvious at the type locality.

A few plants were also found associated with a lone

Pachypodium namaquanum on the slopes of a koppie. G.

Williamson (pers. comm.) has observed N. williamsonii

associated with Pachypodium namaquanum on south-

facing slopes near Oena. He also found (NBG5161) that

plants are common in black tillite and on brown-yellow

dolomite on the west side of the Dreigratberg.

Like many desert annuals, germination and flower-

ing are largely dependent on localized rainfall patterns.

Similarly, the ultimate size of individuals depends, to

a large extent, on available moisture and nutrients. The

largest individual of N. williamsonii at the type locality

occurred along the roadside in loose sand. It had obvi-

ously received additional rainfall in the form of runoff

from the road.

Breeding systems: Nemesia williamsonii, like many

Nemesia species, is self-incompatible. This is based on

the absence of capsule formation in cultivated plants.

Fruit set in the type locality was good, but no pollina-

tors were observed. Related species with similar look-

ing flowers are pollinated by anthophorid bees (Steiner

unpubl.). The spurs of N. williamsonii do not secrete

nectar, but pollen may act as an attractive food source for

the pollinators.

This plant was brought to my attention by Dr Graham

Williamson who first encountered it on one of his many

excursions into the Richtersveld (Williamson 2000).

He showed it originally to Prof E.A. Schelpe at the

Bolus Herbarium at the University of Cape Town, who

recognized it as an undescribed species. However, Prof

Schelpe was unable to describe this species before his

death and over the years. Dr Williamson made addi-

tional collections in the hope that someday it would be

described. Although it was clear from herbarium material

that this species was new, the author described it only

after seeing living material in the field. The annotation

'Nemesia marlothii Grant’ appears on some specimens

of N. williamsonii at NBG and in the literature (Range

1935), but this is a nomen nudum.

Other specimens examined

NAMIBIA. — 2717 (Chamaites): hills E of Dabimub River, ± 400 m,

(-CC), 4 Sept. 2000, P. Brnyns 8865 (NBG); Ai-Ais Reserve, NW of

camp, (-CD), 30 June 1986, Van Jaarsveld 8786 (NBG); Ai-Ais area.

400 m, (-DC), 26 July 1989, Oliver 9153 (NBG); Karibis, Ai-Ais, 800

m, (-DC), Aug. 1909, Marloth 4785 (NBG). 2816 (Oranjemund): West

Dreigratberg, N of old Sendelingsdrif police post, (-BB), 5 Aug. 1993,

Williamson 5161 (NBG); Olienhoutplaas, Karagaskloof, 400 m. (-BB),

3 Sept. 1977, Thompson & Le Roux 364 (NBG). 2817 (Vioolsdrif):

± 15 km E of Visriviermond on Jan Haak road, (-AA), 7 Aug. 1986,

Williamson 3551 (NBG); near Aussenkehr, 80 km W of Vioolsdrif,

(-AD), 9 Aug. 2000, Goldblatt & Manning 11365 (NBG).

NORTHERN CAPE. — 2816 (Oranjemund): top of Kodaspiek,

(-BB), 20 Sept. 1981, Van Jaarsveld & Kritzinger 6240 (NBG). 2817

(Vioolsdrif): Likkewaankloof, Richtersveld National Park, (-AA), 23

Aug. 1993, Zietsman 2350 (PRE); Abiekwarivier Mountain. (-AC),

20 Aug 1987, Jurgens 22386 (PRE); Rosyntjieberg, close to base.

42

Bothalia 36,1 (2006)

above Goennakouriep River, (-AC), July 1989, Williamson 4273

(NBG); Zebra Kloof, just NE of Rosyntjieberg, (-AC), 9 Oct. 1991,

Gennishiiizen 5582 (PRE); E slope of mountain overlooking Koerogab

Vlakte. 400 m. (-AC), Sept. 1994, Williamson 5516 (NBG); Tatasberg

River mouth, (-AC), 15 Aug. 1982, Williamson 3074\ northeastern

Richtersveld National Park. De Toon, 500 m, (-AD), Sept. 1994,

Williamson 5519 (NBG); Richtersveld, 10 km W of Springbokvlakte

on road to Grasdrift. ± 400 m, (-AD), Sept. 1990, Williamson 4361

(NBG); Richtersveld, Springbokvlakte, (-AD), July 1989, Williamson

4272 (NBG); Richtersveld, Tatasberg, Giant’s Playground. (-AD), 24

Aug. 1986, Williamson 3594 (NBG); Richtersveld, Tatasberg (-AD),

July 1989, Williamson 4271 (NBG); Richtersveld, in granite gravel in

stream bed below Tatasberg, 400 m, (-AD). Sept. 1990, Williamson

4360 (NBG).

Locality uncertain: Noaisobis, northern Richtersveld, 3 Sept. 1957,

Herre s.n. (NBG); Ganaquib, JJelskloof 21 Sept. 1933. Herre 19005

(NBG).

Nemesia hemiptera K.E. Steiner, sp. nov., N. maxi

Hiem proxima, sed differt florum albo nec rubro, cal-

carato brevissisimo, seminibus testis in alis expansa

unilateribus.

TYPE. — Northern Cape, 3018 (Kamiesberg):

Kainiesberg, Roodeberg’s Kloof, 21.3 km from N7 via

Farm Doringkraal, 640 m, (-CA), 27 Sept. 2002, Steiner

3946 (NBG, holo.; CAS, K, iso.).

Annual herb up to 310 mm tall; stems rectangular in

cross section, comers ridged, sides up to 0.9 mm wide,

minutely glandular pilose, lateral stems up to 250 mm

long. Leaves simple, opposite, sessile to shortly petiolate;

petioles up to 6 mm long, glandular pilose; lamina ovate to

lanceolate, 8-21 x 2-8 mm, subglabrous to sparsely glan-

dular pubemlous, base rounded to cuneate; apex rounded

to acute; margins entire to shallowly dentate. Flowers

axillary or in lax terminal racemes, racemes up to 225

mm long; bracts alternate, sessile, lanceolate, lowermost

leaf-like, uppermost linear, reduced to ± 2 x 0.3 mm, base

tmncate, apex acute; margin entire, glandular pubescent;

pedicels ± 5-14 mm long, glandular pubescent. Calyx 5-

lobed, spreading, central upper lobe ± 2. 6-2. 8 x 0.6 mm,

lateral upper lobes ± 2.6 x 0.5-0. 7 mm, lower two lobes

± 2.2 X 0.6-0. 8 mm, all lobes narrowly lanceolate, acute,

glandular pilose. Corolla bilabiate, 6.2-9. 6 x 6.7-9. 1

mm, upper lip four-lobed, two upper lobes oblong to

ovate, 3. 2-3. 7 x 1.2-2. 5 mm, base ± perpendicular to

margins, apex rounded to emarginate, two lateral lobes

oblong to elliptic, 3. 3^. 4 x 3. 1-3.3 mm, base strongly

oblique, apex rounded to emarginate; lobes white except

for a brown rectangular nectar guide, ± 0.5 x 0.7 mm,

below sinus of upper two lobes; lower lip with one lobe,

widely obovate and emarginate or obcordate, 3. 6-5. 7

X 5. 0-5. 8 mm, white, tube below lip strongly inflated

into a convex palate; palate ± 2.1 x 2.8 mm, white, with

dense patch of capitate, non-glandular trichomes at base

of palate near corolla opening, hypochile ±2.4 mm long

with a raised central ridge, densely pubescent with cla-

vate trichomes; trichomes on distal 7a of ridge brown,

remainder white, base of hypochile drawn out into a short

nib-like spur, 0.5-0. 8 mm long, entrance to spur flanked

by three brown spots visible on outside of corolla around

base of spur; inside of corolla tube white with lilac tinge.

Stamens 4, whitish, lying in a shallow depression in upper

surface of corolla tube; filaments of anticous pair (twisted

into posticous position) ± 1 .6 mm long, ± straight except

at base, glabrous; posticous filaments ± 0.4 mm long,

± straight except at base, glandular pubescent; anthers

0.4-0. 5 mm long, each pair strongly coherent. Ovary

widely ovate in outline, 0.6-0. 7 x 0.6-0. 7 mm, laterally

compressed; style ± 0.3-0. 4 mm, compressed contrary

to ovary, apex wider than base, lying between anther

pairs, slightly deflexed, stigma a crescent-shaped apical

band. Capsules ovate to oblong in outline, ± 3. 4-4. 6 x

3. 5^. 5 mm, laterally compressed contrary to septum,

apex emarginate to bilobed, lobes acute. Seeds ovate, ±

1. 3-2.0 X 0. 7-1.0 mm, verruculate, winged on one side

only, wing membranous, pale brown with numerous par-

allel veins. Flowering time: (July-) August to September

(-December). Figure 3.

Diagnostic features: Nemesia hemiptera is easily rec-

ognized by its small, white flowers with a tiny spur and

its seed that is winged on only one side. It can be distin-

guished from the related N. maxii by its colour (white vs

violet), the size of the corolla < 10 mm long, the smaller

spur (< 1 mm vs > 3 mm).

Etymology: the name refers to its unique, partially

winged, seeds.

Distribution and habitat: Nemesia hemiptera is a short-

lived, wiry-stemmed annual that comes up after winter or

early spring showers. It is endemic to Namaqualand and

restricted to the Kamiesberg and adjacent rocky hills

(Figure 2). It ranges in elevation from about 560 m to

1 250 m. It occurs in sandy, loam soils in Namaqualand

Broken Veld, under and around shrubs at the foot of large

granite outcrops. Annual rainfall in this area is probably

between 200 and 300 mm, but runoff from the surround-

ing granite outcrops increases the effective rainfall signi-

ficantly. The late flowering time for this annual is surpri-

sing, since the surrounding plants are mostly drying off

when these plants flower. Like many annuals from arid

areas, the ultimate size of individuals depends to a large

extent on available moisture and nutrients. Flowering is

largely dependent on localized rainfall patterns.

Breeding systems: based on cultivated specimens

transplanted from the field, Nemesia hemiptera, despite

its small flowers, is self-incompatible. In the wild, it is

probably pollinated by small pollen-collecting bees (e.g.

halictids or allodapines). The spurs do not secrete nectar,

but pollen may serve to lure pollinators to the flowers.

Nemesia hemiptera was first collected by Rudolf

Schlechter near Brakdam, ± 16 km north of Garies, on an

expedition to Namaqualand that he made with his brother

Max in 1 897 (Gunn & Codd 1981). Since that time, it has

been collected at least a dozen times, but it has never been

described formally.

Other specimens examined

NORTHERN CAPE. — 3017 (Hondeklipbaai): Bowesdoqj, (-BB),

Aug. 1929, L. Bolus 19054 (BOL); summit of Sneeukop, (-BB), 11

Dec, 1909, Pearson & Pillans 5818 (K); slopes of Sneeukop, NE of

Kamieskroon, ± 1 250 m, (-BB), 12 Sept. 1993, Goldhiatt & Manning

9724 (NBG); 4.8 km E of Kamieskroon. 1 090 m, (-BB), 24 Sept.

1948, Acocks 22616 (PRE); 6.3 km E of Kamieskroon, 1 070 m,

(-BB), 27 Sept. 2002, Steiner 3953 (CAS, NBG); between Garies and

Kamieskroon, (-BD), Aug. 1929, Pillans 6251 (BOL); Brakdam, north

of Garies, ± 560 m, (-BD), 7 Sept. 1897, Schlechter 11103 (PRE);

Brakdam, (-BD), 24 Aug. 1941, Barker 1939 (NBG); Brakdam, (-BD),

4 Sept. 1945, Barker 3653 (NBG); Garies, (-DB), 24 July 1941,

Esterhuysen 5428 (BOL). 3018 (Kamiesberg): Studer’s Pass. 22 km E

of Garies, 625 m, (-AC), 27 Aug. 1967, Thompson 421 (NBG, PRE),

Bothalia 36,1 (2006)

43

FIGURE 3. — Nemesia hemiptera, Steiner 3946. A, habit; B, C, flowers, front and rear views; D, calyx; E, F, flowers, side view, intact and partially

cut away; G, capsule with calyx;. H, seed; I. posticous and anticous stamens; J, pistil. Scale bars: A, 10 mm; B-F, H-J, 1 mm; G, 2 mm.

Artist: John Manning.

ACKNOWLEDGEMENTS

I thank Northern Cape Nature Conservation and the

Richtersveld National Park for permission to work in

their respective areas, Graham Williamson for collect-

ing N. williamsonii on numerous occasions over the

years, and John Manning for his excellent illustrations.

I also thank the Compton Herbarium for the use of their

facilities and NBG and PRE for the loan of Nemesia

specimens.

REFERENCES

GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern

Afi-ica. Balkema, Cape Town.

HIERN, W.R 1904. Scrophulariaceae. In W.T. Thiselton-Dyer, Flora

capensis 4,2: 121^20. Reeve, London.

HILLIARD, O.M. & BURTT, B.L. 1986. Notes on some plants of

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

Botanic Garden Edinburgh 43: 345-405.

RANGE, R 1 935. Die Flora des Namalandes. VIII. Feddes Repertorium

38: 263.

44

STEINER, K..E. 1989. A new perennial Nemesia (Scrophulariaceae)

from the western Cape. South African Journal of Botany 55:

405^08.

STEINER, K.E. 1994. A new Nemesia (Scrophulariaceae) from the

Bothalia 36,1 (2006)

interior of the southern Cape, South Africa. South African

Journal of Botany 60: 211-213,

WILLIAMSON, G. 2000. Richtersveld, the enchanted wilderness: an

account of the Richtersveld. Umdaus Press, Hatfield, Pretoria.

Bothalia36,l: 45-56 (2006)

Two new species of Commiphora (Burseraceae) from southern

Africa

W. SWANEPOEL*

Keywords: Burseraceae, Commiphora Jacq., endemism, Gariep Centre, Kaokovetd, morphology, Namibia, new species, South Africa, taxonomy

ABSTRACT

Commiphora steynii Swanepoel and C. gariepensis Swanepoel, here described as new species, are known only from

the Kaokoveld and Gariep Centres of Endemism respectively. Illustrations of the plants and distribution maps are provided.

Diagnostic characters of C. steynii include the pale ashy grey, non-peeling bark and the lack of wart-like projections around

the large lenticels. Diagnostic characters of C. gariepensis include the stamen number which varies between four and eight,

and the milky-watery latex which does not squirt when branches are damaged. When without leaves or fruit, C. gariepensis

can easily be confused with several other species. Comprehensive tables with diagnostic morphological features to distinguish

between the new species and closely related taxa are presented.

INTRODUCTION

Thirty-four species of Commiphora Jacq. are pres-

ently known from the Flora of southern Africa Region

of which twenty-six occur in Namibia (Craven 1999;

Germishuizen & Meyer 2003; Swanepoel 2005). Ten of

these species are more or less restricted to the Kaokoveld

Centre of Endemism, northwestern Namibia and four

to the Gariep Centre of Endemism along the Namibia-

South Africa border (Van Wyk & Smith 2001; Curtis &

Mannheimer 2005).

In this contribution, two new species of Commiphora

are described, C. steynii Swanepoel from the Kaokoveld

Centre of Endemism and C. gariepensis Swanepoel

from the Gariep Centre of Endemism. Collections of C.

steynii were formerly regarded to be conspecific with C.

merkeri Engl. (Van der Walt 1986) and more recently

with C. viminea Burtt Davy (Coates Palgrave 2002). C.

gariepensis is a newly discovered, rare species, with a

limited geographical distribution.

MATERIALS AND METHODS

Diagnostic morphological features to differentiate

between C. steynii and C. viminea (Van der Walt 1973)

and between C. gariepensis, C. oblanceolata Schinz

and C. dinteri Engl. (Van der Walt 1974), are presented.

Apart from examining the herbarium collections of

Commiphora in the WIND and PRE herbaria, live mate-

rial from numerous populations were studied in the field.

Unless indicated otherwise, morphological characters

were all determined from mature leaves and flowers and

from ripe fruit.

Comprehensive comparative tables of salient diag-

nostic morphological characters to differentiate between

C. steynii and C. viminea and between C. gariepensis,

C. oblanceolata and C. dinteri were compiled (Tables

1 & 2). Selected morphological differences between C.

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

University of Pretoria, 0002 Pretoria. Postal address: P.O. Box 21168,

Windhoek, Namibia. E-mail: monteiro@iway.na

MS. received: 2005-02-25.

Steynii, C. viminea, C. merkeri, C. habessinica (O.Berg)

Engl, and C. spathulata Mattick are presented in Table

3. For C. steynii and C. viminea, the diagnostic features

were determined from herbarium specimens and plants

in the field. In addition, some information regarding C.

viminea was sourced from the literature. Regarding C.

merkeri, C. habessinica and C. spathulata, all informa-

tion presented was sourced from Gillett (1991).

Commiphora steynii Swanepoel, sp. nov., C.

vimineae Burtt Davy cortice cum lenticellis magnis,

ramis ramulisque glabris levibus nitidis, foliis simplici-

bus vel trifoliolatis similis, sed trunco simplice vel e terra

multicaule, cortice sine prominentiis verruciformibus

circum lenticellos, non vel exigue deglubenti, foliolis lat-

eralibus usque ad terminalis longitudine, pseudo-arillo

camoso, e basi cum brachiis 4 insignitis differt.

TYPE. — Namibia, 1713 (Swartbooisdrif): Otjirova,

south of Steilrandberg, 1 000 m, (-CD), 08-01-2004,

Steyn & Swanepoel 1 (WIND, holo.!; PRE, iso.!).

Illustrations: Steyn: 45, 46 & 87 (2003).

Dioecious small tree up to 3.5 m high, with or with-

out spines; single or multi-stemmed from ground level;

trunk and stems cylindrical, up to 200 mm in diam. Bark

on trunk and older stems pale ashy grey, yellowish grey,

greyish brown or khaki, smooth, peeling insignificant,

in some specimens peeling in places in small, tough,

flake-like pieces or in short, narrow, transverse strips,

not papery, occasionally with few dark patches in places,

lenticels transversely elongated, often almost completely

encircling trunk and stems. Branches and branchlets gla-

brous, smooth, with small lenticels, shiny brown to dark

brown, rarely maroon-brown or blackish grey, often with

transversely alternating rings of dark and pale bark on

older branches, often spine-tipped, spines slender; spines

or spine-tipped lateral branchlets rarely in clusters of up

to 5 or branched into 2 or 3 spines or spine-tipped lateral

branchlets; new growth red or green, often with few glan-

dular hairs, otherwise glabrous; dwarf lateral branchlets

often scarred. Exudate milky, glutinous, not aromatic,

drying to form a soft to hard yellowish cream or caramel-

brown resin, often in beads, not transparent.

46

Bothalia36,l (2006)

TABLE 1 . — Salient morphological differences between Commiphora steynii and C. viminea

Bothalia 36,1 (2006)

47

TABLE 1. — Salient morphological differences between Commiphora steynii and C. viminea (cont.)

Leaves simple or trifoliolate; clustered on branches

and dwarf lateral branchlets, spirally on shoots; glaucous

or pale green with a dullish lustre, bright green when

young, occasionally retaining bright green colour, differ-

ent shades of green occasionally together on same tree

but on different branches; simple leaves and terminal

leaflets with few glandular and long, shaggy, flexuous

hairs adaxially at base, few short, glandular hairs on both

sides of lamina, especially adaxially on midrib, very

short glandular hairs clustered in comers of serrations

and occasionally on lamina margin, otherwise glabrous;

lateral leaflets with few glandular hairs adaxially on mid-

rib, with or without long flexuous hairs ad- and abaxially

on midrib, short glandular hairs clustered in comers of

serrations, otherwise glabrous; minute lateral leaflets

with few glandular and long flexuous hairs abaxially,

otherwise glabrous; all long flexuous hairs achromatous

at first, becoming brown with age; lamina of simple

leaves and terminal leaflets usually narrowly obovate

to broadly obovate, elliptic to broadly elliptic or rarely

broadly oblanceolate or suborbicular, (5-)12-35(-58)

X (3-)6-22(-36) mm. apex acute, obtuse, tmncate or

retuse, minute tip usually acute, base cuneate, acuminate

or shortly attenuate onto the petiole; margin usually cre-

nate-serrate with (l-)5-9(-15) teeth on each side, basal

third to half usually entire, occasionally crenate-serrate

from base, margin occasionally almost entire; lamina of

lateral leaflets narrowly elliptic to elliptic or narrowly

obovate, often asymmetric especially over basal part, up

to two thirds length of terminal leaflets, (2-)4-20(-30) x

(l-)2-10(-15) mm. apex acute, base cuneate or acumi-

nate, margin crenate-serrate with (l)2-4(5) teeth on each

side, basal third to half entire, margin occasionally almost

entire; where clustered, lamina of lateral leaflets very

small, lanceolate, oblanceolate or narrowly elliptic, often

asymmetric, (0.5-)5.0-12.0(-15.0) x (0.2-)3.0-6.0(-8.0)

mm, apex acute, base cuneate, margin entire; midrib con-

spicuous abaxially, broadest at base, gradually narrow-

ing towards apex, prominently raised adaxially, less so

abaxially, on lateral leaflets often curved, especially over

basal part, in minute lateral leaflets often obscure; leaves

subsessile or petiolate; petiole with few glandular hairs

and long flexuous hairs adaxially, otherwise glabrous,

from less than 1 mm up to 5 mm long, crescent-shaped

in t/s with 7-15 vascular bundles, sectional dimensions

(0.9-)l.l-1.5(-1.8) X (0.6-)0.8-l.l(-1.3) mm; petiolules

up to 1 mm long or leaflets sessile.

Inflorescence', flowers borne in clusters, solitary or

male flowers rarely in much-reduced, simple or dichasial

cymes with peduncle up to 1 mm long; axillary. Flowers

sessile, subsessile or pedicellate; pedicel often with short

glandular hairs; unisexual, hypogynous, precocious or

appearing with or after leaves. Bracteoles and bracts nar-

rowly triangular, apex acute, with glandular and long hairs,

bracteoles up to 1.5 mm long, bracts up to 2.3 mm long.

Calyx greenish yellow, yellow or greenish red, often with

scattered, short, glandular hairs, otherwise glabrous, lobes

triangular, apex acute. Petals greenish yellow, yellow,

greenish red or red, glabrous, recurved towards top but

minute tip inflexed. Disc cylindrical with 4 distinct lobes,

not adnate to calyx or corolla. Male flowers 4.9-12.5 mm

long; pedicel 0. 3-5.0 mm long; calyx 3. 1^.5 mm long;

calyx lobes 0. 5-2.0 mm long; petals cultrate, ensiform,

linear or oblong, 3. 0-8. 5 x 0. 6-1.0 mm; disc with indenta-

tion between lobes deep, disc lobes very fleshy, not bifid

at apex, deeply grooved on inside over basal part; stamens

8, 4 long ones with filaments 1.4-4. 7 mm long, inserted

on outside of disc lobe just below apex; 4 short ones with

filaments 0.6-2. 8 mm long, inserted on outside of disc

48

Bothalia 36,1 (2006)

FIGURE 1. — C. steynii. A, multi-

stemmed tree, 2.5 m tall; B,

tree with single stem, 3 m

tall.

between lobes; anthers on long stamens 0.8-1. 6 mm long;

anthers on short stamens 0.7-1. 2 mm long, usually 0. 1-0.2

mm shorter than anthers on long stamens or anthers rarely

equal in size; filaments slender, thread-like, not flattened

or broadened over lower part; gynoecium rudimentary.

Female flowers 3. 5^. 7 mm long; pedicel 0. 3-0.8 mm

long; calyx 1.6-3. 3 mm long, calyx lobes 0. 5-1.0 mm

long; petals linear or oblong, 2. 8^.4 x 0.6-0. 9 mm; disc

lobes not very fleshy, bifid at apex, grooved on outside

over apical part; staminodes 8, 4 long and 4 short; ovary

superior; style relatively long, sutures deeply grooved;

stigma obscurely 4-lobed, either protruding up to 0.5 mm

above petals, level with petals or 1.4 mm below top of

flower; pistil 1.7-2. 8 mm long.

Fruit a drupe, obovoid, ellipsoid or oblong-ellipsoid,

apiculate, slightly flattened, asymmetrical, 10-14 x 8-9 x

7-8 mm, apex occasionally bent over towards sterile lo-

cule; pericarp 2-valved; exocarp glabrous, not glutinous,

reddish green or red in ripe fruit; mesocarp fleshy; puta-

men flattened, asymmetrical, ovoid, obovoid, ellipsoid or

oblong-ellipsoid, with one fertile and one sterile locule,

rugose; fertile locule often very rugose, 6. 3-8.2 x 3. 9-5. 9

X 3. 5-4. 2 mm, fertile locule convex in sutural view and

convex or triangular in apical view; sterile locule dorsally

ridged, variable in sutural view, either rectilinear and

tapering to apex, rectilinear with a hump near apex and

occasionally with an additional smaller hump near base,

or rectilinear with an indentation in centre, triangular in

apical view; suture rectilinear or convex towards fertile

locule; angle between locules at apex (44-)65-75(-83)°;

pseudo-aril greenish yellow or yellow, fleshy, with 4

long narrow arms from base of putamen, commissural

arms reaching the 2 large apical pits, facial arms reaching

apex, arm on fertile locule usually broader than arm on

sterile locule. Flowering time: August to March. Figures

1^.

Diagnostic characters and affinities: Commiphora stey-

nii probably is most closely related to C. viminea (until

recently misidentified as C. merkeri in southern Africa),

the species with which it has hitherto been confused (Table

Bothalia 36,1 (2006)

49

TABLE 2. — Salient morphological differences between Commiphora gariepensis, C. oblanceolata and C. dinteri

50

Bothalia 36,1 (2006)

TABLE 2. — Salient morphological differences between Commiphora gariepensis, C. oblanceolata and C. dinteri. cont.

1). It also shows morphological resemblance to a number

of Central and East African species, in particular to C.

merkeri, C. habessinica and C spathulata (Gillett 1991).

C. steynii differs from these taxa (Table 3) mainly in habit,

bark, exudate, internal features of the flowers, in the fruit

and in geographical distribution. C. steynii differs conspicu-

ously from C. viminea in that its bark does not peel, or peels

FIGURE 2.— C steynii. Bark.

FIGURE 3. — C. steynii, leaves: A, B, simple; C, trifoliolate. Scale

bars: 20 mm.

Bothalia36,l (2006)

51

TABLE 3. — Selected morphological differences between Commiphora steynii, C. viminea, C. merkeri, C. habessinica and C. spathulata

arms and reaching apex

insignificantly only and is without the characteristic wart-

like projections around the large lenticels. At a distance, the

bark appears pale ashy grey to white. Young branches are

brown or blackish grey in colour. In C. viminea the bark

peels around stems in significant, yellow, papery strips and

young branches are purple. C. steynii is a small, single or

multi-stemmed tree up to 3.5 m tall, with or without slender

spines, whereas C. viminea is a larger tree up to 5 m tall,

single-stemmed, always spinescent with robust spines. In C.

steynii the exudate is non-aromatic, whereas in C. viminea

it is aromatic. The lamina margin on simple and terminal

leaflets of C. steynii are with up to 15 teeth on each side and

the lateral leaflets are up to two thirds the length of terminal

leaflets. The petiole in C. steynii is usually thicker in t/s,

0.9-1. 8 X 0.6-1. 3 mm, and it has 7-15 vascular bundles.

In C. viminea, the lamina margin on simple and terminal

leaflets has only up to 6 teeth on each side and the lateral

leaflets are only up to half the length of terminal leaflets, the

petiole is usually smaller, 1.0-0. 8 mm in transverse section

and with 10 or 11 vascular bundles. The flowers and fruit

provide additional distinguishing features. In C. steynii the

pedicel and calyx are often with short glandular hairs and

the pseudo-aril is yellow and fleshy, with four long, nar-

row arms. In C. viminea the pedicel and calyx are always

glabrous, the putamen has the fertile locule narrowly ridged

transversely one third from the base and the pseudo-aril is

membranous, yellow, orange or red and covers the stone

completely (Table 1 ). A study of herbarium material in PRE

shows that collections from the Zambezi River Valley are

tme C. viminea and not C. steynii. These plants also have

the typical peeling bark with black patches characteristic of

C. viminea.

Eponymy: the specific epithet honours Marthinus H.

Steyn, bom in 1935, tree enthusiast, amateur botanist,

author and publisher of A field guide, southern Africa,

Commiphora (2003), and various other field guides on

southern African trees. Marthinus was the first person

to propose that C. steynii should be regarded as a dis-

tinct taxon (Steyn 2003). I would like to propose the

names ring-bark corkwood and ringbaskanniedood as the

English and Afrikaans vernacular names, respectively.

Distribution: C. steynii is only known from the

Kaokoveld Centre of Endemism (Van Wyk & Smith

2001 ), in northwestern Namibia (Figure 5). It most prob-

ably also occurs in southern Angola, as it was collected

within only 1 km from the Namibian/ Angolan border

near Ruacana in the Kunene River Valley. C. steynii va-

ries from locally common to uncommon or rare within

its range. It is absent from many areas with seemingly

suitable habitat.

Habitat and ecology: C. steynii occurs in the

Kaokoveld, including the pro-Namib Desert, the escarp-

ment and to the east on the inland plateau. It occurs

70-260 km from the coast at altitudes of 800-1 200 m,

where the annual rainfall is 75-300 mm. It grows mainly

in Colophospermum-Commiphora woodland, where it

prefers rocky areas and mixed soil and gravel substrates

on hill slopes and plains. In the extreme south of its range

at Palm and Gomakukous, it occurs on Etendeka basalt of

the Damaraland Igneous Province. In the Sesfontein area,

it grows on sedimentary dolomite and metasedimentary

schist of the Damara Supergroup and in the Omuhiva

area on calcrete. South of the Steilrandberg and in the

Rooidrom area it occurs on quartzite of the Damara

Supergroup. In the northeast of its range at Ruacana, it is

found on limestone of the Karoo Supergroup (Miller &

Bothalia 36,1 (2006)

FIGURE 4. — C. steynii. A-E, male

flower: B, calyx and corolla

partly removed. C-E, disc: C,

from outside; D, from inside;

E, from above to depict posi-

tion of stamens (black) and

rudimentary ovary (circle).

F-H, female flower: G, calyx

and corolla partly removed;

H, disc as seen from above

to depict position of stamens

(black). I-L, putamen with

pseudo-aril: I, lateral view,

fertile locule right, sterile loc-

ule left; J, fertile locule; K,

sterile locule; L, apical view.

A, B, Steyn & Swanepoel 1;

F, G, Swanepoel 109; I-L,

Swanepoel 145. Scale bars:

A, B, 5 mm; F, G, 2 mm;

I-L, 4 mm. Artist: Charmaine

Baardman.

Schalk 1980; Mendelsohn et al. 2002).

Other specimens examined

NAMIBIA.— 1712 (Posto Velho): 5 km W of Rooidrom, (-CD),

Swanepoel 135 (WfND); 2 km S of Rooidrom. (-DC), Swanepoel

109 (WIND). 1713 (Swartbooisdrif): Etanga, (-CC), Swanepoel 99,

100 (WIND); 19°50.3' S, 13°16.2' E, (-CD), Swanepoel 101 (WIND).

1714 (Ruacana Falls): 8 km W of Ruacana Falls in Kunene River

Valley, (-AC), Swanepoel 133. 1812 (Sanitatas): between Sanitatas

and Otjikongo, (-BA), Merxmiiller & Giess 1457 (PRE, WIND).

1813 (Opuwo): Omuhiva, (-AD), Swanepoel 88 (WIND); along road

from Opuwo to Marienfluss via Etanda, (-BB), Hilbert 177 (WfND);

Outuwo, (-CB), Swanepoel 136 (WIND); 13 km SW of Tomakas, (-

CC), Swanepoel 134 (WIND); 2 km S of Okovikuti, (-CD), Swanepoel

141 (WIND); Orurupiza, (-CD), Swanepoel 144 (WIND); 3 km S

of Robbiespas, (-DA), Swanepoel 142 (WIND); Otjikondavirongo,

(-DC), Owen-Smilh 298 (WIND), Swanepoel 145 (WIND). 1913

(Sesfontein): 3 km N of Otjondundu Fountain, (-BB), Swanepoel 143

(WIND); near Gomakukous on Palmwag to Sesfontein road, (-DB),

Swanepoel 127 (WIND); 32 km N of Palmwag on Sesfontein road,

(-DB), Swanepoel 128, 129, 130 (WIND); near Gomakukous, N

of Palmwag on Palmwag-Sesfontein road, (-DB), Swanepoel 131,

138, 158 (WIND); near Gomakukous, (-DB), Swanepoel 137, 139,

140 (WIND). 1914 (Kamanjab): 49 km SE of Sesfontein on road to

Otjovasandu, (-AC), Van der Walt 239 (PRE, WIND); Farm Palm OU

708, (-CC), Giess 7727 (PRE, WIND).

Commiphora gariepensis Swanepoel, sp. nov., C.

oblanceolatae Schinz habitu et foliis plerumque trifoliatis

cum paucis simplicibus similis; exsudato lacteo-aquoso,

non emicanti ubi rami laesi vel secti, foliolis terminali-

bus oblanceolatis ve; cultratis, lateralibus oblanceolatis,

anguste ellipticis vel cultratis, lamina pro ratione latiori,

staminibus 4-8 in quoque flore differt.

TYPE. — Namibia, 2819 (Ariamsvlei): escarpment of

Blydeverwacht Plateau, 3 1 km SSW of Ariamsvlei, 950 m,

(-BD), 09-12-2003, Swanepoel 148 (WIND, holo.!; PRE,

iso.!).

Dioecious shrub or small tree, 0. 6-3.0 x 0.7-2. 2 m;

trunk short, branching repeatedly above ground level

into thick stems with succulent appearance; younger

branches slender. Bark greenish brown, greenish grey or

pale grey with small dark spots and longitudinal, narrow,

dark markings in places, transverse folds at base of stems

and at bends of older branches, usually with few parallel

longitudinal ridges on stems and older branches in places,

otherwise smooth, not peeling. Branches glabrous with

few small lenticels, glutinous when young, not spine-

Bothalia36,l (2006)

53

FIGURE 5. — Known distribution of C. steynii.

tipped. Exudate watery-milky (initially watery, followed

by a milky secretion), not squirting upon branches or

branchlets being damaged or cut, glutinous, aromatic,

forming a soft, transparent, pale cream-yellowish resin.

Leaves trifoliolate with in addition a few scattered,

simple and occasionally few, intermediate leaves, clus-

tered on dwarf lateral branchlets, spirally on shoots,

glabrous, green; lamina of terminal leaflets oblanceolate

or rarely cultrate, (5-)10-18(-36) x (2-)4— 8{-13) mm,

apex acute to obtuse, base cuneate or slightly acuminate;

lamina of lateral leaflets narrowly elliptic, oblanceolate

or cultrate to broadly cultrate, often asymmetrically, up

to (50-)60-70(-100)% as long as terminal leaflet, (3-)6-

14(-25) X (l-)3-6(-10) mm, apex acute to obtuse, base

cuneate or slightly acuminate; lamina of simple leaves

lanceolate-oblong, oblong, narrowly elliptic, oblanceo-

late or deeply trilobed, (10-)12-18(-20) x (5-)6-8(-10)

mm or when trilobed (10-) 17-26 (-30) x (15-)19-23(-

26) mm, apex acute to obtuse, base cuneate or obtuse;

margin serrate, serrate-dentate or rarely crenate-serrate

or margin rarely subentire with teeth on terminal leaflets

3-7(-ll), on lateral leaflets 2-6(-9) and on simple leaves

5-9(-13) on each side, entire near base; midrib either

slightly raised, plane or sunken ad- and abaxially; petiole

usually slightly grooved adaxially especially over basal

part, 1-18 mm long on trifoliolate and 1-8 mm long on

simple leaves, variable in t/s: circular, reniform, elliptic,

triangular, crescent-shaped or pentagonal with 3-5 vas-

cular bundles, sectional dimensions (0.4— )0.6-0.7(-0.9)

X (0.4— )0.5-0.6(-0.7) mm, leaflets sessile or subsessile.

Inflorescence', flowers borne in much reduced or short

simple dichasial cymes, up to 3 mm long, glandular,

or flowers solitary, axillary. Flo'wers sessile, or when

solitary, subsessile or pedicellate, unisexual, perigynous,

appearing before leaves and often continuously while in

leaf Bracteoles ovate, up to 0.4 mm long, apex acute,

glandular. Calyx green, continuous with hypanthium,

glandular otherwise glabrous, lobes triangular to ovate,

apex acute. Petals green to yellowish green, occasionally

sparsely glandular, otherwise glabrous, narrowly elliptic

or narrowly obovate, recurved apically but the minute tip

inflexed, inserted on hypanthium. Disc cylindrical, with 4

fleshy lobes, adnate to hypanthium but distal part of lobes

free. Male flowers 2.6— A.l mm long with pedicel up to

0.3 mm long; calyx 2. 1-3.3 mm long; calyx lobes 0.8-1 .3

mm long; petals 2. 6^. 8 x 0.8-1. 6 mm; disc lobes with

apices distinctly to obscurely bifid; stamens 4-8, 4 long

ones with filaments 1.1-2. 9 mm long, inserted on top of

disc lobes, 1^ short ones with filaments 0.6-1. 8 mm

long, inserted on top of disc between lobes, short stamens

rarely completely absent; anthers 0.7-1. 3 mm long, equal

in length on short and long stamens; filaments rarely

flattened and broadened over lower part; gynoecium

rudimentary. Female flowers 2. 5^.0 mm long; pedicel

up to 0.7 mm long; calyx 2. 3-2. 9 mm long; calyx lobes

0. 8-1.0 mm long; petals 1.3-2. 5 x 0.7-1 .2 mm; disc

lobes distinctly bifid to entire; staminodes present; ovary

half inferior; style variable in length from relatively

short to relatively long, sutures deeply grooved; stigma

obscurely 4-lobed; pistil with stigma from 0.9 mm below

top of flower to level with top of flower, 1.7-2. 4 x

0.9-1. 2 mm.

Fruit a drupe, ovoid or ellipsoid, 8. 2-9. 3 x 6.1-1 .A x

5. 6-6.0 mm, flattened, asymmetrical, pericarp 2-valved;

FIGURE 6. — C. gariepemis. A, natu-

ral habitat, tree ± 2.5 m tall;

B, bark.

54

Bothalia36,l (2006)

FIGURE 7. — C. gariepensis, leaves:

A, trifoliolate; B. simple.

Scale bar: 20 mm.

exocarp glabrous, glutinous, green, greenish brown or

red when ripe; mesocarp not very fleshy; putamen flat-

tened, asymmetrical ellipsoid with one fertile and one

sterile locule, 6. 8-7.4 x 4. 8-5. 2 x 3. 9^. 3 mm, slightly

rugose; fertile locule convex in sutural and apical view;

sterile locule convex in sutural view, convex or triangular

in apical view, often broadly ridged dorsally; suture con-

vex towards fertile locule, especially towards apex; angle

between locules at apex 70-105°; pseudo-aril orange,

fleshy, cupular, covering 24-30% of fertile locule and

40-50% of the sterile locule, with 2 commissural arms

and short facial lobe on sterile locule, extent of commis-

sural arms relative to length of putamen (with pseudo-

aril removed) 80-90%, facial lobe convex or triangular,

0.6-0. 8 mm long, apical pits small. Flowering time\

September to February. Figures 6-9.

Diagnostic characters and affinities'. C. gariepensis

has the same habit as C. oblanceolata and C. gracilifron-

dosa Dinter ex J.J.A.van der Walt, but differs from them

mainly in the shape of the leaves, in the morphology of

the flowers and in the type of exudate. All three of these

taxa have trifoliolate leaves, but C. gariepensis and C.

FIGURE 8. — Trifoliolate leaves: left, C gariepensis', centre, C. oblan-

ceoiaia', right. C. gracilifrondosa. Scale bar: 20 mm.

oblanceolata have simple leaves in addition. Trifoliolate

leaves in C. gariepensis have the lamina on terminal

leaflets oblanceolate or rarely cultrate with up to 11 teeth

on each side of the margin, whereas the lamina in lateral

leaflets is narrowly elliptic, oblanceolate or cultrate to

broadly cultrate with up to nine teeth on each side of the

margin. In C. oblanceolata the lamina on both terminal

and lateral leaflets is always oblanceolate, with up to 45

and 38 teeth respectively on each side of the margin. In

C. gracilifrondosa, the linear or cultrate leaflets, very

variable in size and form, distiguish it from the other two

taxa. The leaves of some specimens of C. gariepensis

and C. gracilifrondosa resemble each other superficially

(Figures 7 & 8). However, the leaves of C. gariepensis

are broader than those of C. gracilifrondosa for corre-

sponding length. In C. gariepensis the flowers are borne

in much reduced or short cymes up to 3.0 mm in length,

in C. oblanceolata in thyrses or cymes up to 13 mm long

and in C. gracilifrondosa on cymes up to 50 mm long. The

flowers of C. gariepensis have 4-8 stamens per flower,

whereas those of C. oblanceolata and C. gracilifrondosa

have four stamens only. Unlike C. oblanceolata and C.

gracilifrondosa, the latex of C. gariepensis is milky-

watery (not watery) and does not squirt when branches

or branchlets are damaged or cut. When without leaves

or flowers, C. gariepensis is virtually indistinguishable

from C. gracilifrondosa with which it shares the same

habitat. The only notable difference then between the

two taxa, is the milky-watery exudate which does not

squirt in C. gariepensis, as opposed to being copious

and only watery in C. gracilifrondosa. Although they

share the same habitat throughout the known range of

C. gariepensis, C. gracilifrondosa is common and much

more widespread. Some specimens of C. gariepensis

could easily be mistaken for C. oblanceolata due to the

similarity of the leaves. However, the geographical dis-

tribution of the two taxa differs markedly, C. gariepensis

being restricted to the Gariep Centre of Endemism and

C. oblanceolata to the northern Central Namib and the

Kaokoveld Centre of Endemism (Van der Walt 1986).

Herbarium specimens of C. dinteri and C. gariepensis

with leaves only, could also be confused. However, the

leaves of C. dinteri are ± twice as broad as those of C.

Bothalia36,I (2006)

55

FIGURE 9. — C. gariepensis. A. male

inflorescence. B-E, male

flower: C, caljoc and corol-

la partly removed; D, disc,

depicting insertion of stamens;

E, disc as seen from above

to depict position of stamens

(black) and rudimentary ovary

(circle). F-H, female flower:

G, calyx and corolla partly

removed; H, disc as seen from

above to depict position of

stamens (black). I, fruit. J-M,

putamen with pseudo-aril: J,

sterile locule; K, lateral view,

fertile locule right, sterile loc-

ule left; L, fertile locule; M,

apical view. A-C, Swanepoel

148', F, G, I-M, Swanepoel

157. Scale bars: B, C, F, G,

2 mm; I-M, 5 mm. Artist:

Charmaine Baardman.

gariepensis for corresponding length. The distribution of

the two taxa does not overlap, with C. dinteri occurring

from south central Namibia northwestwards (Van der

Walt 1986). For a comprehensive comparative table of

diagnostic characters to differentiate between C. gari-

epensis, C. oblanceolata and C. dinteri see Table 2.

Etymology’-, the specific epithet refers to the Gariep

Centre of Endemism along the lower Orange River in

western southern Africa, the region to which C. gari-

epensis is endemic. Gariep is the Khoekfioe name for

the Orange River. As English and Afrikaans vernacular

names, 1 propose Gariep corkwood and Gariep-kannie-

dood, respectively.

Distribution-. C. gariepensis is known from three

localities, all within the Gariep Centre of Endemism.

It is rare in these areas, growing in loose colonies of a

few plants each, in association with C. gracilifrondosa

(Figure 10).

Habitat and ecology-, habitat preferences of C. gari-

epensis are quite specific. It occurs in the hot, arid

Orange River Valley, at altitudes of 300-1 000 m,

where the annual rainfall is 50-150 mm, at the base

of rocky outcrops and on rocky slopes of escarpments

and river valleys. It is restricted to biotite rich gneiss

of the Namaqua Metamorphic Complex/Province at the

Blydeverwacht Plateau and at the Bak River, and to the

gneisses of the Haib Group along the Goodhouse Poort

(Miller & Schalk 1980; Van der Walt 2000; Mendelsohn

et al. 2002).

FIGURE 10. — Known distribution of C gariepensis.

56

Other specimens examined

NAMIBIA.^2818 (Warmbad): near river, road D208, (-CC),

Mannheimer CM 2318 (WIND); up valley off road D208, (-CD),

Mannheimer CM 2312 (WIND). 2819 (Ariamsvlei): 32 km SSW of

Ariamsvlei at base of rocky outcrop, escarpment of Blydeverwacht

Plateau, (-BC), Swanepoel 146 (WIND); 31 1™ SSW of Ariamsvlei on

hill slope, escarpment of Blydeverwacht Plateau, (-BD), Swanepoel 98,

147, 157 (WIND).

NORTHERN CAPE. — 2820 (Kakamas); slopes of Bak River

Valley, 21 km SW of Naroegas Station, (-AC), Swanepoel 149 (PRE);

slopes of Bak River Valley 22 km SSW of Naroegas Station, (-AC),

Swanepoel 150 (PRE).

ACKNOWLEDGEMENTS

I would like to thank Prof. A.E. van Wyk, University

of Pretoria, for advice and support, Dr H.F. Glen, SANBI,

for translating the diagnoses into Latin, Ms H. Steyn,

SANBI, for preparing the distribution maps and Ms C.

Baardman for the line drawings. The curator and staff of

the National Herbarium of Namibia are thanked for their

assistance during visits to the herbarium. The National

Herbarium of Namibia and the South African National

Biodiversity Institute are also thanked for the use of

information from their databases: SPMNDB, Flora DB

and PRECIS. The curator, National Herbarium, Pretoria,

is thanked for access to their collections; the assistance of

Dr C.L. Bredenkamp and Mrs M. Jordaan during visits to

the herbarium is acknowledged with thanks. Ms V. Noble

from the National History Museum, London, is thanked

for images of Angolan material. I am especially grateful

Bothalia 36,1 (2006)

to my wife Hannelie for assistance and support during

field trips.

REFERENCES

COATES PALGRAVE, M. 2002. Keith Coates Palgrave Trees of south-

ern Africa, edn 3. Struik, Cape Town.

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.

GERMISHUIZEN, G. & MEYER, N.L. (eds). 2003. Plants of southern

Africa: an annotated checklist. Strelitzia 14. National Botanical

Institute, Pretoria.

GILLETT, J.B. 1991. Burseraceae. Flora of tropical East Africa.

Balkema, Rotterdam/Brookfield.

MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T.

2002. Atlas of Namibia. Philip, Cape Town.

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 Affica/Namibia.

STEYN, M. 2003. A field guide, southern Africa Commiphora/’«

Veldgids, suider-Afrika Commiphora. Published by the author,

Polokwane, Limpopo.

SWANEPOEL, W. 2005. Commiphora kaokoensis (Burseraceae), a new

species from Namibia, with notes on C. dinteri and C. namaen-

sis. Bothalia 35: 47-53.

VAN DER WALT, J.J.A. 1973. The South African species of

Commiphora. Bothalia 11: 53-102.

VAN DER WALT, J.J.A. 1974. A preliminary report on the genus

Commiphora in South West Africa. Madoqua 1: 5-23.

VAN DER WALT, J.J.A. 1986. Burseraceae. Flora of southern Africa

18,3: 5-34.

VAN DER WALT, P.T. 2000. Augrabies weelde. Info Naturae,

Totiusdal.

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

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Hatfield, Pretoria.

Bothalia36,l: 57-61 (2006)

Notes on the systematics and nomenclature of Tritonia (Iridaceae:

Crocoideae)

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

Keywords: Iridaceae, southern Africa, systematics, Tritonia Ker Gawl., Tritonia crispa (L.f.) Ker GawL, Tritonia flava (Aiton) Ker GawL,

Tritonia securigera subsp. watermeyeri (L. Bolus) J.C. Manning & Goldblatt, Tritonia undulata (Burm.f.) Baker

ABSTRACT

Study of some early types of species now known to belong to the genus Tritonia Ker Gawl., a member of Iridaceae, sub-

family Crocoideae, comprising some 28 species of southern and south tropical Africa, has shown the need for some nomen-

clamral adjustments. Ixia undulata Burm.f (1768) is an earlier name for T. crispa (L.f) Ker Gawl. based on Gladiolus crispus

L.f (1782) and the combination T. undulata (Burm.f) Baker must be used for the species, which is native to the western half

of Western Cape, South Africa. The variety T. crispa van parviflora is also reduced to synonymy. The type specimen of Ixia

gladiolaris Lam. (1789), currently considered a synonym of Tritonia securigera (Aiton) Ker Gawl., has flowers that lack the

characteristic tooth-like ridges on the lower tepals of the latter, and corresponds closely to the eastern southern African T.

lineata (Salisb.) Ker Gawl., based on Gladiolus lineatus Salisb. (1796). The new combination T. gladiolaris (Lam.) Goldblatt

& J.C. Manning is made and T. lineata is reduced to synonymy. Montbretia lacerata and Tritonia lacerata. erroneously

regarded as synonyms of T. crispa, are combinations based on Gladiolus laceratus Burm.f, a species that remains unidenti-

fied because no type is known and the description is too vague to permit its identification. Lastly, field studies have shown

that the crisped-leaved T. watermeyeri is connected by a series of morphological intermediates to typical T. securigera, which

has straight leaves and identical flowers. The new combination T. securigera subsp. watermeyeri (L. Bolus) J.C. Maiming &

Goldblatt is proposed for this taxon.

INTRODUCTION

Our continuing studies of the systematics and biology

of the African Iridaceae led us to examine type material

of several species thought to belong to the genus Tritonia

Ker Gawl. but excluded by De Vos (1982, 1983) from

her revision of the genus as insufficiently known because

of the difficulty in relating type material to any known

species. Among the names in question are Gladiolus fla-

vus [Sol. in] Aiton (1789), G. laceratus Burm.f. (1768),

G. undulatus Burm.f (1768), and Ixia gladiolaris Lam.

(1789). The type of Ixia gladiolaris is what is currently

called Tritonia lineata (Salisb.) Ker Gawl., which thus

becomes T gladiolaris. Gladiolus flavus was based on

a specimen of T flabellifolia (D.Delaroche) Ker Gawl.,

to which it is now assigned as a later synonym. The type

of G. undulatus matches T crispa (L.f.) Ker Gawl. The

latter, based on G. crispus L.f (1782), closely resembles

the shorter-tubed T crispa var. parviflora Baker, and the

species must therefore be known by the earlier name T.

undulata, a combination made by Baker in 1877. We

have, however, failed to locate authentic material that

can be associated with G. laceratus and this species

must continue to be excluded from Tritonia. Taxonomic

adjustments required as a result of our investigation are

made below. Lastly, new collections of T. securigera

(Aiton) Ker Gawl. have led us to re-examine the distinc-

tion between this species and the western Little Karoo

plants referred to as T. watermeyeri. The latter species,

defined by its crisped and undulate leaves, is now known

to be connected by a series of intermediates to the wide-

spread T. securigera, which has plane leaves but flowers

identical to those of T. watermeyeri. We conclude that it

* B.A. Krukoff Curator of Afncan Botany, Missouri Botanical Garden,

RO. Box 299, St. Louis, Missouri 63166, USA.

** Compton Herbarium, South Afncan National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

MS. received: 2005-10-21.

is most appropriate to treat the latter as a subspecies of

T. securigera.

1 . Tritonia flabellifolia (D.Delaroche) G.J.Lewis in

Journal of South African Botany 7: 30 (1941). Ixia flabel-

lifolia D.Delaroche: 24 (1766). Type: South Africa, with-

out precise locality or collector (L-Herb. Van Rooyen,

neo.!, designated by Goldblatt & Barnard 1970: 310).

Gladiolus flavus [Sol. in] Aiton: 65 (1789). Tritonia flava (Aiton)

Ker Gawl.: 228 (1804). Type: South Africa, without precise locality,

cultivated in Great Britain, Paterson s.n. (BM, holo.!), syn. nov.

The type of Gladiolus flavus is readily identified as a

specimen in the British Museum (Natural History) col-

lected in early bloom (Figure 1). The long-tubed flowers

are quite evident, and the characteristic dry, brown, long-

attenuate and acuminate bracts immediately identify the

plant as the Western Cape species, Tritonia flabellifolia.

This plant has white to cream-coloured flowers and it

is puzzling that Solander, who drew up the description,

should have called it G. flavus (actually describing the

flower as intensely yellow). Possibly the flower buds

were sufficiently cream-coloured for him to have thought

they deserved the epithet he chose. Despite the apparent

inconsistency in flower colour we are confident that G.

flavus is a synonym of T. flabellifolia.

2. Tritonia gladiolaris (Lam.) Goldblatt &

J.C. Manning, comb. nov.

Ixia gladiolaris Lam., Encyclopedic methodique 3: 341 (1789).

Type: South Africa, Cape of Good Hope, cultivated in Paris, flower-

ing March and April, original collector unknown (P-Herb. Lamarck,

holo.!).

Gladiolus lineatus Salisb.: 40 (1796). Tritonia lineata (Salisb.) Ker

Gawl.: 228 (1804); M.P.de Vos: 369 (1983); M.P.de Vos: 111 (1999).

Montbretia lineata (Salisb.) Baker: 169 (1877). Tritonixia lineata

(Salisb.) KJatt: 357 (1882). Type: South Africa, without precise locality,

grown at the Royal Botanic Gardens, Kew in 1781 (BM, lecto., desig-

nated by M.P.de Vos 1983: 371), syn. nov.

58

Bothalia 36,1 (2006)

D1 23456789 10

cm copyright reserved

THE NATURAL HISTORY MUSEUM

DEPARTMENT OF BOTANY

BM000838886

LONDON

FIGURE 1. — Holotype of Gladiolus flavus [Sol. in] Aiton (BM),

Ixia gladiolaris, represented by a single sheet in the

Herbarier Lamarck in the Laboratoire de Phanerogamic,

Paris, was included in the synonymy of T. securigera by

De Vos ( 1 983, 1 999) in her accounts of Tritonia. The nar-

rowly sword-shaped leaves have prominent submarginal

veins, and the four or six flowers on the two preserved

spikes have large, finn, dry bracts that are rust-coloured

in the upper half. The flowers still show the rather

prominent darker venation on the tepals characteristic of

the genus, particularly of T. Uneata, as well as the low,

thickened ridges (raised, yellow, truncate scales accord-

ing to Lamarck) on the lower tepals. Lamarck described

the corolla as orange or yellow, lightly flushed with red.

The association by De Vos (1983, 1999) of Ixia gladio-

laris with T. securigera is refuted by two features: the

presence of prominent submarginal veins on the leaves.

and the low ridges on the lower tepals. The leaves of T.

securigera have no obvious submarginal veins and the

lower tepals of the flower bear large, tooth-like ridges,

3^ mm high, that resemble elaborate axeheads.

We regard the type and associated description of Ixia

gladiolaris as representing one of two largely eastern

southern African species, either Tritonia disticha (Klatt)

Baker or T. lineata. Both these species have leaves with

prominent submarginal veins, unusual in Tritonia, and

flowers with low median ridges on the lower tepals. The

two species are distinguished largely by flowering time,

flower size and perianth colour (De Vos 1999): T. disticha

blooms in summer, mainly January to March, and has

red to pink (or rarely yellow) flowers 20-30 mm long,

whereas T. lineata flowers earlier, mainly September to

Bothalia 36,1 (2006)

59

November, and has yellow to apricot (pale orange) flow-

ers (25-)30-40 mm long. Ixia gladiolahs most closely

matches the latter morphologically and is evidently also

early flowering. Plants grown in Paris bloomed in spring,

March and April in the northern hemisphere, whereas we

would expect T. disticha to flower in June or July, even

under glass. I. gladiolahs is an earlier name for T. lineata,

which thus falls into synonymy under the new combina-

tion T. gladiolahs.

3. Tritonia undulata (Burtn.f) Baker in Journal

of the Linnean Society 16: 163 (1877). Ixia undulata

Burm.f.: 1 (1768). Thtonixia undulata (Bunn.f.) Klatt:

357 (1882). Type: South Africa [Western Cape], without

precise locality, collector unknown (G-BU, holo.!).

Gladiolus crispus L.f.: 94 (1782). Tritonia crispa (L.f.) Ker Gawl.

18: t. 678 (1803); M.P.de Vos: 144 (1982); M.P.de Vos: 99 (1999). Type:

South Africa [Western Cape], without precise locality, A. Sparrmann

s.n. (LINN 59.18, holo.-microfiche!), syn. nov.

Tritonia crispa vdx. parviflora Baker: 192 (1892); M.P.de Vos: 149

(1982); M.P.de Vos: 100 (1999). Type: South Africa [Western Cape],

Winterhoek. Tulbagh, C.L. Pappe s.n. (K. holo. not seen), syn. nov.

The identity and generic disposition of Ixia undulata,

described by N.L. Butman in 1768, has proved enduring-

ly controversial. It was referred to Tritonia, as T. undulata

by Baker ( 1 877), and remained so treated in Flora capen-

sis (Baker 1896). Klatt (1882), however, referred/, undu-

lata to his new genus Thtonixia, a segregate of Tritonia

erected for the shorter-tubed members of the genus (but

including the type of Tritonia, T. crocata). Brown (1929)

actually made a new combination, T. undulata, because

although Baker had already made the same combination,

he had included plants of Ixia crispa L.f. (now I. erubes-

cens Goldblatt) under the same.

Later, Lewis (1962) in her monograph of Ixia excluded

Ixia undulata from that genus but did not explicitly say

to what genus she thought the plant belonged. Then, De

Vos (1983, 1999) excluded the species from Tritonia in

her accounts of the genus, commenting that although it

superfically resembled a Tritonia, the short, stout anthers

recalled Ixia subgenus Dichone, although the funnel-

shaped tube did not. She therefore suggested that the

type might represent a hybrid between I. crispa and I.

vanzijliae M.P.de Vos.

We have examined the type specimen of Ixia undulata,

a single plant without a corm, in the Burman Herbarium

at Geneva (Figure 2), and are amazed that its identity

could have caused such confusion. It is an exact match

for plants from the Tulbagh District of Western Cape,

referred by both Baker (1892, 1896) and De Vos (1983,

1999) to Tritonia crispa var. parviflora. The leaf margins

are tightly crisped and the entire blade is loosely undulate

and twisted, while the salver-shaped flower has a narrow,

almost cylindric (not funnel-shaped as De Vos stated)

perianth tube, ± 1 8 mm long with [apparently] spreading

tepals, stamens with filaments exserted ± 5 mm from the

tube, and anthers ± 3 mm long.

We have no hesitation in regarding Ixia undulata as

eonspecific with Tritonia crispa (basionym Gladiolus

crispus L.f. (1782)). I. undulata is the earlier name, and T.

crispa now falls into the synonymy of T. undulata.

We also question the merit of recognizing infraspe-

cific taxa in this species. Perianth tube length is variable,

ranging from 18-85 mm across its range. Flowers with

the longest tubes are recorded from the Olifants River

Mountains (85 mm in Goldblatt & Manning 10345,

NBG), south to Joostenberg (60 mm in Lewis 5901, NBG).

Plants with shorter-tubed flowers, 25-40 mm long, occur

inland and in the south, as at Sir Lowry’s Pass (35^0 mm

in Barker 3378, NBG), and Somerset West (25-30 mm in

Lewis 5675, NBG). Some plants from the Tulbagh Valley

fall at the lower extreme of the range with a tube 22-25

mm (Hansford 4 from Welgelegen House, NBG). In con-

trast, plants from the slopes of Roodezandberg, not far

from Tulbagh, have a tube ±50 mm long (Compton et al.

1882/36, NBG). The pattern is not entirely clear but there

is a trend for shorter tubes in populations in the south

and interior of its range and any division into subspecific

taxa based on tube length would be arbitrary. Tube length

is closely associated with the local pollinators, long-

proboscid flies (Manning & Goldblatt 1997). Where the

long-proboscid species Moegistorhynchus longirostris

occurs within the range of T. undulata, the perianth tube

is longest but inland and on the Cape Peninsula where

M. longirostris does not occur, shorter-proboscid flies,

presumably either Philoliche giilosa or P. rostrata, are

the inferred pollinators. Predictably, tube length tracks

the proboscis length of the pollinator, a situation that has

been documented in Disa draconis and Geissorhiza con-

fusa (Johnson & Steiner 1997). We therefore include var.

parviflora in T. undulata.

4. Tritonia lacerata (Burm.f.) Klatt, Erganzungen

und Berichtigungen zu Baker’s Systema Iridacearum.

Abhandlungen der Naturforschenden Gesellschaft zu

TYPUS

FIGURE 2. — Holotype of Ixia undulata Burm.f. (G-BU).

60

Bothalia 36,1 (2006)

Halle 15: 358 (1882). Gladiolus laceratus Burm.f.:

2 (1768). Montbretia lacerata (Burm.f.) Baker: 168

(1877). Type: unknown, not in G-BU (Brown 1929) or L

(J.Veldkamp pers. comm.).

No type for this species, described by Burman fil. in

1768, has ever been identified, and the description is too

brief to permit its association with any known species.

Baker (1877), who referred the species to Montbretia

(now included in Tritonia), believed it to be an earlier

name for T. crispa, now T. undidata. Both Baker and

Klatt, who made the combination in Tritonia, are silent

on their reasons for identifying Burman ’s species as a

Tritonia. In her most recent account of Tritonia, De Vos

(1999) placed Montbretia lacerata Baker (sic) and T

lacerata (Baker) Klatt (sic) as synonyms of T crispa,

and designated the type of T crispa as the lectotype of

M. lacerata. She thus ignored the fact that Baker’s name

is a combination based on Burman ’s Gladiolus laceratus

(and thus has no type of its own but is nomenclaturally

identical to G. laceratus). De Vos did not specifically

deal with the identity of the latter. In our search for type

material we have confirmed that no type is located at G

(Herb. Burman) or L (the Leiden Herbarium), the only

places where it is likely to be preserved.

5. Tritonia securigera (Sol. in Aiton) Ker Gawl.

in Annals of Botany 1: 228 (1804); M.P.de Vos: 384

(1983); M.P.de Vos: 1 15 (1999). Gladiolus securiger Sol.

in Aiton: 65 (1789). Type: South Africa, without precise

locality or date, cultivated at Chelsea Gardens in 1778, F.

Masson s.n. (BM, lecto., designated by De Vos 1983).

Ixia thimbergii Roem. & Schult.; 391 (1817), replacement name for

/, squalida Thunb. in Hoffmann, Phytogeographische Blatter: 4 (1803),

nom. illeg. non Aiton (1789). Type: South Africa, Cape of Good Hope,

without precise locality, Thunberg s.n. (UPS-//er6. Thimberg 996,

microfiche!).

There is no question about the identity of the type of

this predominantly Little Karoo species, to our knowl-

edge first collected by Masson and Thunberg in the

early 1770s. Masson’s collection was later grown at the

Royal Botanic Gardens, Kew, where plants in flower

were described as Gladiolus securiger a decade later

by Solander for Alton’s Hortus kewensis (Aiton 1789).

Plants of the same collection were illustrated in Curtis s

Botanical Magazine in 1797 under that name (Ker

Gawler 1797) but the species was later transferred to

Tritonia (Ker Gawler 1804).

Distinctive features of the species, in the narrow sense,

are a basal fan of firm, plane leaves (4-)6-12 mm wide

with acute tips, a typically unbranched stem (robust

plants may have one or more lateral branches), and an

erect spike of (3)4—8 orange, rarely yellow, and evidently

unscented flowers. The dorsal tepal is largest, 16-18 x

13-15 mm, and the lower tepals each bear a pronounced

bright yellow median tooth, ± 4 x 2.5 mm, shaped like

an axehead (securiger is Latin for axe-bearer). These

teeth, called calli in the early literature and calluses by

De Vos (1982, 1983, 1999) in her accounts of Tritonia,

are not unique to the species but are frequent in Tritonia.

They are particularly well developed in most species of

section Montbretia, but in the remaining sections usually

fonn low, thickened ridges, and are not developed at all

in some species, such as T. tugwelliae L. Bolus. The teeth

are especially well developed in T. securigera and its

western Little Karoo relative, T. watermeyeri L.Bolus. In

fact there is little difference between the flowers of the

two taxa, apart from the slightly smaller tepal teeth (±

3.0 X 1.3 mm) and faint woody scent in T. watermeyeri.

The major difference between the two are in their leaves,

and the crisped and undulate leaf blades of T. watermey-

eri contrast strikingly with the plane leaves of typical T.

securigera. Scent in flowers of Iridaceae is notoriously

variable, even within a species (Goldblatt & Manning

1998) and little reliance can be placed on it alone as a

specific character.

The distinction between Tritonia securigera and T.

watermeyeri becomes even less clear when plants with

flowers matching the two species from sandy habitats

in the Anysberg and at the foot of the Little Swartberg

in the western Little Karoo are taken into considera-

tion. These plants are often dwarfed, up to 0.6 m high,

and have either fairly soft-textured, undulate leaves

(Goldblatt & Porter 12060, NBG, from the foothills

of the Little Swartberg), or narrow, twisted leaves with

straight to slightly crisped margins (Martin 60, NBG,

from Anysberg Nature Reserve) while the spikes may

have 1-3 flowers in the most dwarfed individuals, or

up to 4 flowers in taller plants. Both T. securigera and

typical T. watermeyeri are usually taller plants, at least

150 mm high, and grow on shale slopes. The plants with

undulate leaves seem perfectly intermediate between

typical examples of T. securigera and T. watermeyeri

and maintenance of the latter as a separate species seems

arbitrary. Existence of plants with leaves of intermediate

form, either undulate or loosely twisted, make it more

appropriate to treat T. watermeyeri as a subspecies of T.

securigera. Typical T. securigera ranges widely through

the southern Cape, from the Touwsberg in the western

Little Karoo to Bedford in the Eastern Cape, whereas

subsp. watermeyeri is restricted to the extreme west,

between Montagu and Anysberg (Figure 3).

subsp. watermeyeri (L.Bolus) J.C. Manning &

Goldblatt, comb, et stat. nov.

Tritonia watermeyeri L.Bolus in Annals of the Bolus Herbarium 4:

44 (1926); M.P.de Vos: 395 (1983). Type: South Africa [Western Cape],

allegedly Vanrhynsdorp, without date, Watermeyer s.n. {BOLJ8050,

holo.l).

Specimens examined

WESTERN CAPE.^3320 (Montagu): Allemorgens, 1 km N of

T-junction near Syfer-se-Kop on road to Bloutoring, red sandy loam,

(-CB), 7 Sept. 1979, Malan 112 (NBG); 8 km from Montagu on road to

Ouberg Pass, (-CB), 27 Sept. 1981, Periy 1682 (NBG); 26 Sept. 1995,

Goldblatt & Manning 10329 (MO, NBG). 3321 (Ladismith): sandy

slopes north of Anysberg, (-BC), 18 Aug. 2002, Goldblatt & Porter

12066 (MO, NBG, PRE); Anysberg Nature Reserve, NW boundary,

(-BC), 3 Sept. 1989, Martin 60 (NBG); Laingsburg-Ladismith road, E

of Anysberg turnoff, (-BD), 17 Aug. 2002, Goldblatt <& Porter 12060

(MO, NBG). Without precise locality: Hoeveld naby Touwsrivier, Apr.

1930, Huysteen s.n. (NBG179592).

ACKNOWLEDGEMENTS

Support for this study from the National Geographic

Society (grants 6704-00, 7103-01, 7316-02, and 7799-05)

is gratefully acknowledged. We thank the Curators of the

following herbaria for loan of type specimens: BM, G,

and L; Pete Phillipson, Missouri Botanical Garden, who

Bothalia36,l (2006)

61

FIGURE 3. — Distribution of Tritonia seciirigera subsp. secuhgera. •;

and subsp. watermeyeri, O.

helped us identify Ixia gladiolaris in the Herbier Lamarck

at P; and J. Veldkamp, Leiden Herbarium, for searching

for a possible type of Gladiolus laceratus. We are also

grateful to Mary Stiffler who cheerfully helped solve

bibliographic questions and provided copies of numerous

articles not readily available to us.

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7-171. Reeve, London.

BOLUS, L. 1926. Novitates atficanae. Annals of the Bolus Herbarium

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DE VOS, M.R 1999. Tritonia. Flora of southern Africa 7,2, fasc. 1:

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HOFFMANN, G.F. 1803. Phytographische Blatter. Schroder, Gottingen.

JOHNSON, S.D. & STEINER, K.E. 1997, Long-tongued fly pollination

and evolution of floral spur length in the Disa draconis complex

(Orchidaceae). Evolution 51: 45-53.

KER GAWLER, J. 1797. Gladiolus securiger. Curtis’s Botanical

Magazine 11: t. 383.

KER GAWLER, J. 1803. Tritonia crispa. Curtis's Botanical Magazine

18: t. 678.

KER GAWLER, J. 1804. Ensatorum ordo. In J. Kdnig & J. Sims,

Annals of Botany 1; 219-247. Taylor, London.

KLATT, F.W. 1882. Erganzungen und Berichtigungen zu Baker’s

Systema Iridacearum. Abhandlungen der Naturforschenden

Gesellschaft zu Halle 15: 335-404.

LAMARCK, J.B.A.P.M. 1789. Encyclopedie methodique Botanique

3. Paris.

LEWIS, G.J. 1941. Iridaceae. New genera and species miscellaneous

notes. Journal of South African Botany 7: 19-59.

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

South African Botany 28: 45-195.

LINNAEUS, C. (fil.) 1782 (1781). Supplementarum plantarum.

Braunschweig.

MANNING, J.C. & P. GOLDBLATT. 1997. The Moegistorhynchus

longirostris (Diptera: Nemestrinidae) pollination guild: long-

tubed flowers and a specialized long-proboscid fly pollination

system in southern Africa. Plant Systematics and Evolution 206:

51-69.

ROEMER, J.J. & SCHULTES, J.A. 1817. Systema vegetabilium. Cotta,

Stuttgart.

SALISBURY, R.A. 1796. Prodromus stirpium in horto ad Chapel

Allerton vigentium. London.

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Bothalia 36,1: 63-89 (2006)

Notes on African plants

VARIOUS AUTHORS

APOCYNACEAE

NEW RECORDS OF ADENIUM BOEHMANUMYN THE ESA REGION

Currently four species of Adenium Roem. & Schult. are

recognized for the Flora of southern Africa (FSA) region,

namely A. boehmiamim Schinz, A. multiflonim Klotzsch,

A. oleifoliiim Stapf and A. swazicum Stapf (Codd 1963;

Victor et al. 2003). In the region, A. boehmiamim has a

northwestern distribution and the other species a central to

eastern distribution (Figure 1). A. boehmiamim was previ-

ously only known from the northern parts of Namibia and

extending just over the border into Angola. In May 2005,

Mr J.H. Vahrmeijer gave a specimen to PRE [Vahrmeijer

HV00702), collected at the ‘Sonop Koppies’ near the

northwestern border of the Central Kalahari Game Reserve

in Botswana on 15 April 2005. At the time, it was thought

to be the first collection of this taxon from Botswana.

On further investigation, Ms Q. Turner from the

Botswana National Herbarium (GAB) informed me

of another collection of Adenium boehmiamim from

Botswana (M Kabelo 2), collected on 17 February 2005,

at the base of the Pimple hill in the vicinity of the Tsau

Hills. This locality is near to that of the Vahrmeijer speci-

men.

Setshogo (2005) listed this taxon for Botswana, and

indicated that the voucher specimen was housed in

the National Herbarium in Zimbabwe (SRGH). When

requesting the precise locality for this specimen (no

collector or number was cited) from SRGH, the curator

informed me that the specimen could not be traced. Apart

from the listing, Adenium boehmiamim has not been

reported from Botswana in the literature.

All known Botswana records come from the north-

western borders of the Central Kalahari Game Reserve,

in central Botswana. These records .extend the known

distribution range of the species by at least 650 km to

the east (Figure 1). The distribution map was compiled

from information obtained from the National Herbarium

of Namibia (WIND) database: SPMNDB and Flora DB;

records from the Namibian Tree Atlas project; and the

National Herbarium Pretoria Computerised Information

System (PRECIS) database. Information on one of the

new records was obtained from GAB.

Vahrmeijer indicated on the collecting label that

plants were found locally, frequently growing on rocky

hillsides. Kabelo (Q. Turner pers. comm.) indicated that

the species was rare and grew only at the base of hills.

Succulents are generally sought after and sometimes

threatened by plant collectors and because of this and the

limited distribution in Botswana, the species was listed

as rare and a possible candidate for Red Data listing by

Setshogo (2005). Adenium boehmiamim is an exception-

al, attractive plant when in full flower, and with its glossy

foliage, it has great horticultural potential (Figure 2).

The disjunct distribution may indicate that Botswana

as a whole is undercollected, or at least that the flora of

that country is inadequately covered in PRECIS. Only

9.8% of the quarter-degree grids in Botswana have

more than 50 specimens at PRE, according to PRECIS

records in 2005, and almost a third of all Botswana

quarter-degree grids do not have any specimens in PRE.

12 14 16 18 20 22 24 26 28 30 32

FIGURE I. — Distribution in the FSA region

of Adenium boehmiamim: from WIND

and PRE, •; from Namibia Tree Atlas

Project, O; new records, □. A. oleifoliiim,

from GAB and PRE, ★; A. multiflonim,

from PRE, ■; and A. swazicum, from

PRE, ▲.

64

Bothalia 35,2 (2005)

FIGURE 2. — Flowering Ade-

nhim boehmiamtm photo-

graphed in the nursery

of the Pretoria National

Botanical Garden, x 0.5.

Plant grown from a cutting

collected in 1976 by Leistner

et al. PNBG294.

Further investigation may prove whether the occurrence

of Adenium boehmiamim in Botswana indicates a dis-

junct distribution or not. It is probably an indication that

Botswana is still undercollected, or not well represented

at PRE.

ACKNOWLEDGEMENTS

Thanks go to: Mr Vahraieijer for the gift of the speci-

men; Ms Turner (GAB), Mr Muzila (Curator UCBG)

and Ms Nobanda (curator SRGH) for checking speci-

mens in their respective herbaria; Dr Maggs-Kolling and

Ms Shubert, National Herbarium, National Botanical

Research Institute in Namibia for the use of infonnation

from databases; Ms B. Curtis for supplying the Namibian

Tree Atlas distributions for Adenium boehmiamim; the

South African National Biodiversity Institute for the use

of data from PRECIS; and Ms H. Steyn, SANBI, for

compiling the distribution map.

REFERENCES

CODD, L.E. 1963. Adenium. Flora of southern Africa 26: 278-282.

SETSFIOGO, M.P. 2005. Preliminary checklist of the plants of

Botswana. Southern African Botanical Diversity Network

Report No. 37. SABONET, Pretoria and Gaborone.

VICTOR, J.E., NICHOLAS, A., BRUYNS, P.V., VENTER. H.T.J, &

GLEN, H.F. 2003. Adenium. In G. Germishuizen & N.L. Meyer.

Plants of southern Africa: an annotated checklist. Strelitzia 14:

133. National Botanical Institute, Pretoria.

S.P. BESTER*

* National Herbarium, South African National Biodiversity Institute,

Private Bag XlOl, 0001 Pretoria. E-mail: besteri@sanbi.org

MS. received: 2005-07-19.

HYACINTHACEAE

DRJMIA MONTANA (URGINEOIDEAE), A NEW SPECIES FROM EASTERN CAPE, SOUTH AFRICA

Drimia montana A.P.Dold & E. Brink, sp. nov.,

D. marginatae (Baker) Jessop affmis sed foliis numero

3-6, usque ad 80 mm longis, bracteis pedicellorum usque

ad 1.4 mm longis cum calcari complanato pluteiformi

usque ad 0.8 mm longo, filamentis erecto-patentibus

fusiformibus flexum sigmoideum lenem externum usque

ad 3.6 mm longum formantibus, ovario conico truncato

trigono usque ad 2 mm longo, lineis sex longitudinalibus

purpureo-guttatis angulos flavidos delineantibus, atque

seminibus quinque in quoque loculo, differt.

TYPE. — Eastern Cape, 3226 (Fort Beaufort): The

Hoek Farm, Groot Winterberg, 2 150 m, (-AD), 01-01-

2004, Bold 4633 (GRA, holo.).

Deciduous geophyte, colonial, ± (30-)40-60(-80) mm

tall, densely aggregated in matted, cushion-forming

colonies of more than 100 individuals. Bulb globose

to ovoid, 10-15 X 10-15 mm, hypogeal, firm, tinged

purplish within; roots many, spreading, white, fleshy,

bulbiliferous; inner tunic reddish purple, forming a short,

loose neck; outer tunic loosely flaking, papery, greyish

brown. Leaves (2-)3-6, linear, 15-50 x 1.0-1. 5 mm (2)3-,

erect, entire, somewhat flattened, ± concave adaxially,

apices acute, dry and withered but persistent at flowering,

glabrous, glossy green. Inflorescence solitary, subcapi-

tate-racemose; peduncle slender, erect, (45-)50-60(-80)

X 0.7-0. 8 mm at base, with minutely white-puberulous

hairs in vertical rows, dark, glossy, purple-red, swollen

around and below pedicel fomiing a saddle-shaped, whit-

ish cushion of tissue subtending pedicel and bract; bracts

thinly flattened, up to 1 .4 x 1 .0 mm wide at base, loosely

clasping, somewhat cupped, with a flattened, horizontal

fold up to 0.8 X 0.6 mm broad at base, smooth, fawn-

coloured, drying quickly but persistent; pedicels spread-

ing-erect, (3.0-)5.0-6.0(-6.5) X 0.5-0. 7 mm, dark, glossy.

Bothalia36,l (2006)

65

FIGURE 3. — Drimia montana, Dold 4633: A, plant habit; B, inflorescence; C, pedicel bract; D, perianth; E, gynoecium with a single stamen; F,

anther; G, capsule; H, dehiscent capsule. Scale bars; A, B, D, G, H, 5 mm; C, E, 1 mm; F, 0.5 mm. Illustrations; L. de Wet.

purple-red, becoming erect as fruit develops; base thick-

ened. Flowers (6-)8-12(-15), opening one or two per day,

aborted flowers abscising 0.4 mm below insertion of tepals

leaving a skirt of tissue as pedicel dries. Perianth reflexed,

tepals elliptic, convex, 6.0-6.4 x 2. 2-2.4 mm, fused for

0.8 mm at base, abaxial surface maroon with broad white

recurved margins, adaxial surface white, apices truncate,

penicillate. Stamens spreading-erect; filaments fusiform,

weakly sigmoid, 3.2-3. 6 x 0.8 mm, flattened, 0.6 mm

thick, white; anthers broadly ovoid, bilobed, 1 x 0.7 mm

diam., dorsifixed, versatile, yellow-green, dehiscing longi-

tudinally; pollen ellipsoid, 60 x 20 pm, monosulcate, yel-

low. Ovary a truncate obtusely angled trigonous cone, up

to 2.0 X 1.6 mm at base, up to 1 mm diam. at apex, white,

with six longitudinal lines of purple speckling delineating

yellow tinged comers; style triangular in cross section,

1.4— 2.4 X 0.4 mm, white; stigma swollen, 3-angled with

stalked stigmatic papillae in three rows, white. Capsule

obovate, 8-10 x 6-8 mm, pale yellow-brown with the

persistent circumscissile perianth capping apex and leav-

ing a basal annulus, lobes spreading and twisting outwards

at dehiscence, leathery with thickened margins, cream-

coloured. Seeds ± 5 per locule, flat, oval in outline, 5-8

X 3. 8-5.0 mm, up to 1.2 mm thick; testa glossy, black,

shallowly reticulate; cotyledon fusiform, 4—7 x 1.0- 1.2

mm, yellow. Flowering time: ± 14.30 to ± 19:00 each day

in December to January. Figures 3, 4.

Distribution and biology, based on an ongoing study

of herbarium specimens, wild populations and cultivated

material of Drimia Jacq., D. montana is known to us from

only two mountain peaks in the Eastern Cape (Figure

5) growing in Karroid Merxmuellera Mountain Veld

(Acocks 1988), latterly known as South-eastern Mountain

Grassland (Lubke et al. 1996), at ± 2 000 m. The annual

rainfall is between 450-600 mm, falling predominantly in

the summer months, with severe frost and only occasional

snow in winter (Lubke et al. 1996). The habitat of these

two localities corresponds closely with each other [typi-

cal habitat is illustrated by Acocks (1988) in Figure 95:

112]. Plants form large mats, up to 0.5 m in diameter, of

densely aggregated bulbs on flat, exposed sandstone rock

slabs where, together with several dwarf succulent species,

they are partly concealed by lichen and moss. Associated

species include Cheilanthes hirta, Crassula corallina.

Euphorbia aggregata, E. epicyparissias, Felicia filifolia,

Melolobium candicans, and Pelargonium aridum. Plants

are dormant for two to three weeks in late January. It is

likely that D. montana occurs elsewhere on the Stormberg

and Winterberg where the inaccessible habitat is under no

immediate threat.

Diagnosis and relationships: following Goldblatt &

Manning (2000), D. montana falls within the Urginea

group of the genus Drimia within the Urgineoideae,

Hyacinthaceae. The new species appears to be related

to the D. depressa alliance comprising D. depressa,

D. marginata and D. sphaerocephala, all with ± capitate

inflorescences. The last two named species have cor-

responding recurved tepals (Jessop 1977), a character

previously used to separate Drimia from Urginea (Baker

1897; Mauve 1976), but they are separated by less dis-

tinct characters including the length of the pedicel bracts.

The new species is, however, clearly differentiated by

its dwarf habit, the small pedicel bract with flattened

horizontal fold, the fusiform, weakly sigmoid filaments

and the truncate, trigonous ovary with distinct purple

and yellow markings. In addition, the presence of bul-

bils forming at the base of the bulb and along the roots,

appears to be an unusual character within the genus.

66

Bothalia 36,1 (2006)

FIGURE 4. — Dfimia montana. Dold 4633: A, seed; B, seed testa; C,

pollen. Scale bars: A, 1 mm; B, 100 pm; C, 10 pm.

Additional material examined

EASTERN CAPE. — 3126 (Queenstown): summit of Andriesberg,

Carnarvon Estate, Black Eagle Nature Reserve. 1 885 m, (-DA), 15-

12-2002, Dold & Cocks 4700 (GRA).

FIGURE 5. — Known distribution of Drimia montana.

ACKNOWLEDGEMENTS

Thanks to Rhodes University Joint Research Council

for support; Dr Hugh Glen for providing the Latin diag-

nosis; the Morgan family, Redcliffe, Tarkastad and the

Halse family, Carnarvon Estates, Sterkstroom for their

hospitality; Leigh-Anne de Wet for line drawings.

REFERENCES

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

the Botanical Survey of South Afiica No. 57.

BAKER. J.G. 1897. Liliaceae. In W.T. Thiselton-Dyer. Flora capensis

6: 253-528.

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.

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: 265-319.

LUBKE, R.A., BREDENKAMP, G. & VAN ROOYEN, N. 1996.

South-eastern Mountain Grassland. In A.B. Low & A.G. Rebelo,

Vegetation of South Africa, Lesotho and Swaziland: 47, 48.

Department of Environmental Affairs & Tourism. Pretoria.

MAUVE (OBERMEYER), A.A. 1976. Liliaceae. In R.A. Dyer, The

genera of southern A frican plants, vol. 2: 915-944. Botanical

Research Institute. Pretoria.

A.P. DOLD* and E. BRINK**

*Selmar Schonland Herbarium, Rhodes University Botany Department,

P.O. Box 101, 6140 Grahamstown, South Africa.

**Selmar Schonland Herbarium, Albany Museum, P.O. Box 101,

6140 Grahamstown, South Africa.

MS. received: 2005-03-14.

AMARYLLIDACEAE

A NEW VARIETY OF CLIVIA ROBUSTA

Recently, the genus CUvia (Lindl.) Regel saw the intro-

duction of two new species, C. mirahilis Rourke and C.

rohusta Murray et al., as well as a new variety, C. gardenii

var. citrina Swanevelder et al. (Rourke 2002; Mumay et

al. 2004; Swanevelder et ai 2005). All six currently rec-

ognized species, including four varieties, are indigenous to

South Africa, with two species extending into Swaziland

(Watson 1899; Phillips 1931; Duncan 1985, 1992, 1999;

Rourke 2002; Snijman & Archer 2003; Murray et ai 2004;

Swanevelder 2003; Swanevelder et al. 2005).

Bothalia36,l (2006)

67

FIGURE 6. — Known distribution of C. robusta var. robusta, •; and C.

robiista var. citrina as well as the typical variety.

Clivia robusta, the tallest member in the genus, is

confined to the Pondoland Centre of Endemism, South

Africa (Figure 6). Plants occur as isolated populations

distributed mainly from Port St Johns (Eastern Cape),

through to Port Edward (KwaZulu-Natal), with a few

outliers as far north as Oribi Gorge (Van Wyk 1994; Van

Wyk & Smith 2001; Swanevelder 2003; Murray et al.

2004). Restricted to isolated patches of forest, C. robusta

is usually closely associated with swamps or seepage

areas, though individuals have been found in well-

drained, humus-rich soils, in shallow soil on rocky ledges

of cliffs and even on rocks (Swanevelder 2003, Murray

et al. 2004). The forest type with which it is associated

is Swamp Forest, or in most instances a floristically

enriched form of Afromontane Forest sometimes referred

to as Coast Scarp Forest (MacDevette et al. 1989).

In the description of Clivia robusta, Murray et al.

(2004) noted that yellow-flowered individuals are rarely

encountered in this species, the flowers of which are

usually orange or red. Such sporadic occurrences of

yellow-flowered forms are preferably named at forma

level (Stuessy 1990). However, Watson (1899) described

the sporadic yellow-flowered form of C. miniata as a

variety, namely C. miniata var. citrina, a name that has

been widely adopted. A second yellow-flowered infra-

specific taxon in the genus, C. gardenii var. citrina, was

also described at variety level (Swanevelder et al. 2005).

Following these precedents, we here formally describe

the yellow-flowered form of Clivia robusta Murray et al.

as a new variety.

Clivia robusta B.G. Murray, Y.Ran, P.J.De Lange,

K.R.W.Hammett, J.T.Truter & Z.H. Swanevelder var. cit-

rina Z.H. Swanevelder, A.Forbes-Hardinge, J.T.Truter

& A.E.van Wyk, var. nov., floribus pallide luteis vel

citrinis, apicibus laete vel atro-viridibus, non aurantiacis

vel rubris apicibus viridibus ut in varietate typico distin-

guitur.

TYPE. — KwaZulu-Natal, 3030 (Port Shepstone):

Maringo Flats, (-CC), 7 June 2003, Forbes-Hardinge

FHOl (PRU, holo.).

Flowers with perianth pale yellow or lemon yellow

with light to dark green apices, not dark orange-red, pale

orange or pink-orange with green apices as in the typical

variety.

Clivia robusta has flowers that are usually various

shades of red and orange (Murray et al. 2004). We

regard plants with flowers in all shades of these colours

(at anthesis) as belonging to Clivia robusta var. robusta,

with only the rare yellow-flowering form comprising var.

citrina.

The holotype of C. robusta var. citrina was collected

in an area known as Maringo Flats, located ± 20 km

inland from Port Edward on the KwaZulu-Natal south

coast (Figure 6). A single yellow-flowered specimen was

collected at the time, but more than one yellow indi-

vidual was observed. The frequency of the yellow form

in this particular stand is intermediate to the large stands

of Clivia gardenii var. citrina in nature (Ngome Forest,

Ngotshe District, KwaZulu-Natal) and the single yellow-

flowering specimen on which Watson based C. miniata

var. citrina (Watson 1 899; Swanevelder 2005). Hitherto,

C. robusta var. citrina has only been recorded in this one

population.

The habitat at the type locality is typical for Clivia

robusta, in this case, a swamp-like area with forest cover-

ing ± 2 ha. Most of the C. robusta population grows as

dense stands in very heavy mud on a stream bank. The

type specimen was growing in a silt deposit on the side

of the stream that runs through the forest on its way to

the Umtamvuna River. Associated forest species include

Strelitzia nicolai, Protorhus longifolia, Erythrina cajfra,

Macaranga capensis, Voacanga thouarsii, Syzygium cor-

datum, Phoenix reclinata, Zantedeschia aethiopica and

Cyathea dregei.

Even though Clivia robusta is present in a number of

conservation areas throughout its range, the distribution

of individual populations is very localized due to the

species’ specialized habitat requirements (Swanevelder

2003). In this particular case, even the inhospitable

marshy habitat does not prevent the removal of plants

by traditional healers and illegal plant collectors; it does,

however, restrain the complete removal of whole popula-

tions. All known plants of C. robusta var. citrina occur

on private land and enjoy the protection of the current

landowner.

ACKNOWLEDGEMENTS

Our thanks to Dr Hugh Glen for the Latin translation

of the diagnosis and Ms Hester Steyn for the distribution

map.

REFERENCES

DUNCAN, G. 1985. Notes on the genus Clivia Lindley with particular

reference to C, miniata Regel var. citrina Watson. Veld & Flora

71: 84, 85.

DUNCAN, G. 1992. Notes on the genus Clivia Lindley with particular

reference to C. miniata Regel var. citrina Watson. Herbertia

48: 26-29.

DUNCAN, G. 1999. Grow clivias. Kirstenbosch Gardening Series.

National Botanical Institute, Cape Town.

68

Bothalia 36,1 (2006)

SWANEVELDER, Z.H., VAN WYK, A.E. & TRUTER, J.T. 2005. A

new variety in the genus Clivia. Bothalia 35: 67, 68.

VAN WYK, A.E. 1994. Maputaland-Pondoland region. In S.D. Davis,

VH. Heywood & A.C. Hamilton, Centres of plant diversity. A

guide and strategy for their conservation 1: 227-235. lUCN

Publications Unit, Cambridge.

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

in southern Africa. A review with emphasis on succulents.

Umdaus Press, Hatfield, Pretoria.

WATSON, W. 1899. Clivia miniata var. citrina. The Garden 56; 388,

t. 1246.

MACDEVETTE, D.R., MACDEVETTE, D.K., GORDON, I.G. &

BARTHOLOMEW, R.L.C. 1989. Floristics of the Natal indig-

enous forests. In C.J. Geldenhuys, Biogeography of the mixed

evergreen forests of southern Africa: 124—145. Ecosystem

Programmes Occasional Report No. 45. Foundation for Research

Development, Pretoria.

MURRAY, B.G., RAN, Y, DE LANGE, P.J., HAMMETT, K.R.W.,

TRUTER, J.T. & SWANEVELDER, Z.H. 2004. A new species

of Clivia (Amaryllidaceae) endemic to the Pondoland Centre

of Endemism, South Afiica. Botanical Journal of the Linnean

Society 146: 369-374.

PHILLIPS, E.P. 1931. Clivia miniata war. flava. The Flowering Plants

of South Africa 11: t. 411.

ROURKE, J.P. 2002. Clivia mirabilis (Amaryllidaceae: Haemantheae)

a new species from Northern Cape, South Afiica. Bothalia 32:

1-7.

SNIJMAN, D.A. & ARCHER, R.H. 2003. Clivia. In G. Germishuizen

& N.L. Meyer, Plants of southern Africa: an annotated checklist.

Strelitzia 14: 958, 959. National Botanical Institute, Pretoria.

STUESSY, T.F. 1990. Plant taxonomy. The systematic evaluation of

comparative data. Columbia University Press, New York.

SWANEVELDER, Z.H. 2003. Diversity and population structure of

Clivia miniata Lindl. (Amaryllidaceae): evidence from molecu-

lar genetics and ecology. M.Sc. thesis. University of Pretoria,

Pretoria.

Z.H. SWANEVELDER*, A. FORBES-HARDINGE**, J.T.

TRUTER*** and A.E. VAN WYK^

* Department of Botany/Forestry and Agricultural Biotechnology Institute

(FABI), University of Pretoria, 0002 Pretoria.

** P.O. Box 14964, Trafalgar 4275.

*** P.O. Box 5085, 1502 Benoni South, Benoni. t H.G.W.J. Schweickerdt

Herbarium, Department of Botany, University of Pretoria, 0002 Pretoria.

MS. received: 2005-07-27.

RUBIACEAE

CORRECT AUTHOR CITATIONS FOR NAMES OF THREE SOUTHERN AFRICAN SPECIES OF CANTHIUM

There is confusion in the literature concerning the

correct author citations for the names of the southern

African taxa Canthium ciliatum, C. pauciflorum (now C.

kuntzeanum Bridson) and C. spinosum. All three taxa are

based on type material from South Africa. C. ciliatum is

confined to the Flora of southern Africa {FSA) region, C.

kuntzeanum extends from here to the Eastern Highlands

of Zimbabwe and adjacent parts of Mozambique and C.

spinosum just enters southern Mozambique (although not

treated in Flora zambeziaca). C. ciliatum and C. spino-

sum have received only brief mention in Bridson’s (1992)

revision of Canthium in tropical Africa. Here we clarify

the author citations for thg relevant correct names as well

as their synonyms, based in part on the work of Tilney

(1986) and Bridson (1992, 1998).

One cause of author citation variation is provisions

of the International Code of Botanical Nomenclature

[ICBN] itself. For example, the use of ‘ex’ in author cita-

tions is not mandatory (ICBN, Art. 46.4; Greuter et al.

2000). An author is free to use or not to use it. It is never-

theless essential in all instances to cite the author(s) after

the ‘ex’. Citations with ‘ex’ are cumbersome, and for the

sake of brevity the ‘ex’ and the author names preceding

it are often dropped, resulting in a shorter but still correct

citation. For the sake of completeness we have retained

‘ex’ in all the author citations supplied below. This is the

option we prefer when author citations are supplied in

scholarly publications.

Canthium ciliatum (Klotzsch ex Eckl. & Zeyh.)

Kuntze in Revisio generum plantarum 3, 3: 545 (1898).

Psilostoma ciliata Klotzsch ex Eckl. & Zeyh. : 362 ( 1 837). Plectronia

ciliata (Klotzsch ex Eckl. & Zeyh.) D.Dietr.: 856 (1839), non Sonder:

18 (1865).

Apparently unaware of the earlier work of Dietrich

(1839), Sonder (1865) also made the combination

‘'Plectronia ciliata (Klotzsch ex Eckl. & Zeyh.) Sond.’,

resulting in many subsequent authors incorrectly attribut-

ing the combination in Plectronia to Sonder.

Canthium kuntzeanum Bridson in Kew Bulletin

47: 393 (1992).

Plectronia pauciflora Klotzsch ex Eckl. & Zeyh.: 363 (1837).

Canthium pauciflorum (Klotzsch ex Eckl. & Zeyh.) Kuntze: 545

(1898), non Blanco: 165 (1837), nec Baill.: 189 (1878), nec King &

Gamble: 58 (1904), nec Pierre ex Pitard: 291 (1924).

Over the years the specific epithet pauciflorum, mean-

ing few-flowered, has been applied in combinations with

Canthium by Blanco (1837), Baillon (1878), King &

Gamble (1904) and Pierre ex Pitard (1924). However,

in all these instances, a different plant from the southern

African taxon was being referred to. Because of the ear-

lier names of Blanco and Baillon, Kuntze’s combination

is a later homonym, hence a new name, C. kuntzeanum,

was proposed for the southern African taxon by Bridson

(1992).

Canthium spinosum (Klotzsch ex Eckl. & Zeyh.)

Kuntze in Revisio generum plantarum 3, 3: 545 (1898).

Plectronia spinosa Klotzsch ex Eckl. & Zeyh.: 362 (1837), non

K.Schum.: 459 (1897).

P. klotzschiana K.Sehum.: 460 (1897), nom. superfi.

Based on a specimen from Sierra Leone (‘Guinea’),

Schumacher & Thorming (1827) published the name

Phaliaria spinosa for the taxon currently known as

Vangueriella spinosa (Schumach. & Thonn.) Verde.

Bentham (1849) gave the name Canthium thonningii

Benth. to the same taxon. Schumann (1897: 459) pro-

posed that C. thonningii be transferred to Plectronia, with

the new combination P. spinosa (Schumach. & Thonn.)

K.Schum. Realizing that this name had already been used

Bothalia36,l (2006)

69

by Ecklon & Zeyher (1837) for a different taxon from

southern Africa, Schumann proposed a new (yet superflu-

ous) name for the latter, namely Plectronia klotzschiana

K.Schum.

REFERENCES

BAILLON, H.E. 1878. Canthium paticiflomm. Adansonia 12: 189.

BENTHAM, G. 1849. Canthium thonningii. In W.J. Hooker, Niger

Flora: 410. Hippolyte Bailliere, London.

BLANCO, F.M. 1837. Canthium pauciflorum. Flora de Filipinas, edn

1: 165. Manila.

BRIDSON, D.M. 1992. The genus Canthium (Rubiaceae-Vanguerieae)

in tropical Africa. Ke\^’ Bulletin 47; 353-401.

BRIDSON, D.M. 1998. Rubiaceae (tribe Vanguerieae). In G.V. Pope,

Flora zambesiaca 5,2; 211-377. Royal Botanic Gardens, Kew,

London.

DIETRICH, D.N.F. 1839. Synopsis plantarum, vol. 1 . Bernard Friedrich

Voight. Weimar.

ECKLON, C.F. & ZEYHER, K.L.P. 1837. Enumeratio plantarum afri-

cae australis extratropicae, part 3. Perthes & Besser, Hamburg.

GREUTER. W., MCNEILL, J., BARRIE, F.R., BURDET, H.M.,

DEMOULIN, V., FILGUEIRAS, T.S., NICOLSON, D.H.,

SILVA, P.C., SKOG, J.E., TREHANE, R, TURLAND, N.J. &

HAWKSWORTH, D.L. 2000. International Code of Botanical

Nomenclature (Saint Louis code). Regnum vegetabile 138.

Koeltz Scientific Books, Konigstein.

KING, G. & GAMBLE, J.S. 1904. Canthium pauciflorum. Journal of

the Asiatic Society of Bengal 73: 58.

KUNTZE, C.E.O. 1898. Revisio generum plantarum, part 3. Felix,

Leipzig.

PITARD, J. 1924. Rubiaceae. In M.H. Lecomte, Flore generale de

I’Indo-Chine, vol. 3, fasc. 3: 289-432. Masson, Paris.

SCHUMACHER, C.F. & THONNING, P. 1827. Phaliaria spinosa.

Beskrivelse af Guineeiske planter: 113. Popp, Copenhagen.

SCHUMANN, K. 1 897. Beitrage zur Flora von Afirika XIII. Rubiaceae

africanae. Botanische Jahrbiicher 23: 412-470.

SONDER, O.W. 1865. Order LXXIII Rubiaceae Juss. In W.H. Harvey

& O.W. Sonder, Flora capensis 3: 1-39. Reeve, London.

TILNEY, P.M. 1986. The taxonomic significance of anatomical and

morphological characters in the southern African species

of Canthium Lam. (Rubiaceae). Ph.D. thesis. University of

Pretoria.

P.M. TILNEY* and A.E. VAN WYK**

* Department of Botany, University of Johannesburg, P.O. Box 524,

2006 Auckland Park, Johannesburg.

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

sity of Pretoria, 0002 Pretoria.

MS. received: 2005-07-13.

POACEAE

CONCEPT OF STIPAGROSTIS UNIPLUMIS VAR. UNIPLUMS REDEFINED TO INCLUDE SPECIMENS WITH HAIRY GLUMES

At present, following Smook (1990), the varieties of

Stipagrostis imiphtmis (Licht.) De Winter and S. hir-

tighima (Trin. & Rupr.) De Winter in the Flora of south-

ern Africa (FSA) region are characterized as follows:

Stipagrostis uniplumis van:

imiplumis: perennial; inflorescence with spikelets

numerous; glumes glabrous, up to 10 mm long; callus of

the uniplumis-type (short hairs continuous from naked

tip up entire length of callus to long hairs at junction of

callus and lemma); widespread in FSA region.

neesii (Trin. & Rupr.) De Winter; perennial; inflor-

rescence with spikelets few; glumes glabrous, usually

longer than 10 mm; callus of the iiniphimis-type', occurs

in northwestern Northern Cape, western Free State,

North-West and adjoining area of Botswana and extreme

northern Limpopo.

intermedia (Schweick.) De Winter: annual; inflor-

escence with spikelets numerous; glumes hairy, usually

up to 8 mm long; callus of the w«/p/z/wfr-type; dry north-

western Namibia.

Stipagrostis hirtigluma van:

hirtigluma: annual; inflorescence narrow; glumes

hairy; callus of the hirtigluma-type (with a distinct gla-

brous break between short hairs on body of callus and

long hairs at junction of lemma and callus, best seen at

back of callus).

pearsonii (Henrard) De Winter: annual; inflorescence

open; glumes hairy; callus of the hirtighma-ty\)Q.

patula (Hack.) De Winter: perennial; inflorescence

open; glumes densely hairy; callus of the hirtigluma-type.

The main characters separating Stipagrostis imiplumis

from S. hirtigluma are the hairs on the callus (see above)

and the presence of a distinct pencil of hairs at the branch-

ing point of the awns in the former. In the latter species, the

column is sometimes hairy, but without a distinct pencil

of hairs.

Perennials with hairs on the glumes and a uniplumis-

type callus have been referred to as a hybrid, Stipagrostis

uniplumis x hirtigluma — see De Winter (1965) and note

under S. uniplumis var. intermedia in couplet 30 of the

key in Smook (1990).

Re-examination of all the specimens at PRE that key

out as Stipagrostis uniplumis x hirtigluma showed that a

large number of the specimens fitted the description of S.

uniplumis var. uniplumis except for the hairs present on

the glumes. It is therefore proposed, for the reasons given

below, that the description of var. uniplumis be expanded

to accommodate those specimens that fulfill all the other

criteria for S. uniplumis var. uniplumis but have hairs on

the glumes:

1, hairy glumes are not unknown in S. uniplumis, also

occurring in var. intermedia, an annual from the more

arid northwest of Namibia;

2, the density of hairs, even the presence of hairs on the

glumes, can vary on spikelets of the same inflorescence.

The hairs are often difficult to see (a black background

makes them more visible). Hairiness appears not to be a

stable character in S. uniplumis. De Winter (1965) points

out that the presence or absence of hairs on glumes has

proved to be diagnostically unreliable in other species

of the genus. Therefore, not too much weight should be

placed on this character;

3, there are a large number of specimens from across the

entire distribution area of var. uniplumis that differ from

70

Bothalia36,l (2006)

the typical fonn only in having hairs on the glumes (see

specimens examined below);

4, the callus is of the imiplumis-, not hirtigluma-type.

In the Flora zambesiaca region, Melderis (1971) also

placed most specimens from Botswana and Zimbabwe

differing only by hairy glumes in var. imiplumis, acknowl-

edging that some specimens in Namibia could possibly be

hybrids.

The new expanded description of S. uniplumis var.

uniplumis is as follows: perennial; inflorescence with

spikelets many; spikelets 8-10(-ll) mm long (measured

from base of spikelet to apex of longest glume, excluding

awns); tuft of hairs at branching point of awns and often

hairy partly down column, hairs longer than 1.5 mm;

glumes glabrous or long-hairy, hairs dense to sparsely

scattered but at least always present on margins; callus of

the uniplumis-\y^Q.

There are still a number of perennial specimens with a

pencil of hairs at the branching point of the awns, hairy

glumes and a callus of the uniplumis-\Y'pQ that are diffi-

cult to place in either var. neesii or the new extended var.

uniplumis and will key out as the hybrid S. imiplumis x

5. hirtigluma (see key below).

The emended key in Gibbs Russell et al. (1990: 319,

320) is given below, starting at couplet 28 (22) of the

original key:

28(22) Glumes with long hairs, sometimes sparse and along margins

only 29

Glumes puberulous, scabrid or glabrous 35

29(28) Inflorescence spiciform, subsecund; culms not visibly or

obviously striate, usually densely scrabid

S. gonatostachys

Inflorescence open or contracted but not spiciform; culms

obviously striate, smooth 30

30(29) Callus with short hairs along entire length (except for na-

ked tip) until long hairs at junction of lemma and callus

(uniplumis-type) 31

Callus with distinct glabrous break between short hairs

along length of callus and long hairs at junction of lemma

and callus (hirtigluma-type, best seen at back) 33

31(30) Plant annual S. uniplumis var. intermedia

Plant perennial 32

32(31) Glumes 8.0-10(-l 1) mm long ... ,5'. uniplumis var. uniplumis

Glumes longer than 12 mm S. uniplumis x S. hirtigluma

33(30) Inflorescence narrow, when fully exserted much longer than

wide; annual S. hirtigluma subsp. hirtigluma

Inflorescence open, spreading, when fully exserted not

much longer than wide; annual or perennial 3

34 (33) Annual with very few leaves at base

S. hirtigluma subsp. pearsonii

Perennial with dense tuft of basal leaves

S. hirtigluma subsp. patula

35(28) Branching point of awns and short distance down column

with hairs longer than 1 .5 mm 36

Branching point of awns and short distance down column

glabrous or with hairs shorter than 1.5 mm 37

36(35) Inflorescence with numerous spikelets; glumes 8-10 mm long;

central awn usually straight S. uniplumis var. uniplumis

Inflorescence with a few spikelets; glumes 10 mm or longer;

central awn bent at right angles . ... S. uniplumis var. neesii

37 (35) Inflorescence subsecund, branched only in the lower part, spike-

lets in upper part solitary continue key from couplet

35 in Gibbs Russell et al. (1990) advancing the numbers appro

priately.

Specimens of S. uniplumis var. uniplumis with hairy

glumes examined:

NAMIBIA. — 1714 (Ruacana Falls): Kaokoland, about 4 km S

of Ombamndu, (-CC), Smook 7830. 1812 (Sanitatas): 8 miles W

of Orowanja Fountain on road to Orupembe, (-BA), De Winter &

Leistner 5649. 1813 (Ohopoho): Kaokoland, S of Opuwa on road to

Sesfontein, (-DA), Smook 7841. 2014 (Welwitschia): Farm Driekrone

OU 516, (-BC), Giess 7915. 2114 (Uis): Messumberge, (-AC), Giess

9662. 2117 (Otjosondu): Farm Omupanda, (-DD), Gibbs Russell &

Smook 5371. 2118 (Steinhausen): Farm Merx, (-AB), Gibbs Russell &

Smook 5408. 2216 (Otjimbingwe): Farm Friedenau, (-DB), Muller &

Kolberg 2112. 2317 (Rehoboth): 24 km S of Rehoboth on main road

to Kalkrand, (-CA), Gibbs Russell & Smook 5480. 2417 (Mariental):

20 km W of Mariental on road to Maltahohe, (-DB), Gibbs Russell cfe

Smook 5 509 A.

BOTSWANA.— 1821 (Andara): Tsodilo Hills, (-DC), Guy 121\64.

1922 (Nokoneng): Mojeye area, (-CD), Smith 1302. 1923 (Maun):

Chiefs Island, (-CA), Biggs M462. 2021 (Koanaka Hills): on sides of

Groot Laagte fossil river valley, (-CD), Smith 3205.

MPUMALANGA. — 2431 (Acomhoek): Farm Rietvley, (-AB), Zam-

batis 573. 2331 (Phalaborwa): 15 km SE of Shingwidsi, (-AB), Ellis

537.

LIMPOPO. — 2229 (Waterpoort): Langjan Nature Reserve, (-CC),

Zwanziger 522.

NORTFIERN CAPE. — 2820 (Kakamas): Augrabies National Park,

(-CB), Werger 378. 2824 (Kimberley): Olie River, (-CC), Leistner 1292.

ACKNOWLEDGEMENTS

I wish to thank Ms E. du Plessis for helping with the

text, G. Germishuizen for editing and the referees for

their input.

REFERENCES

DE WINTER, B. 1965. The South African Stipeae and Aristideae.

Bothalia 8: 201^04.

GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK,

L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J,

1990. Grasses of southern Africa. Memoirs of the Botanical

Survey of South Africa No. 58: 275.

MELDERIS, A. 1971 Aristideae 'm Flora zambesiaca 10,1: 133-136.

SMOOK, L. 1990. Stipagrostis . In G.E. Gibbs Russell, L. Watson, M.

Koekemoer, L. Smook, N.P. Barker, H.M. Anderson & M.J.

Dallwitz, Grasses of southern Africa. Memoirs of the Botanical

Survey of South Africa No. 58.- 31 8-329.

L. FISH*

* National Herbarium, South African National Biodiversity Institute,

Private Bag XlOl, 0001 Pretoria.

MS. received: 2005-05-26.

POACEAE

A LONG-AWAITED NAME CHANGE IN POLYPOGON

In 1955 Chippindall noted that Agrostis griquensis Gibbs Russell (1990) placed this taxon correctly, but the

Stapf should be classified as a species of Polypogon Desf combination was never validly published.

Bothalia36,l (2006)

71

According to Clayton & Renvoize (1986), the genus

Polypogon is characterized by the spikelet falling entire

with the upper part of the pedicel still attached. In contrast,

the spikelet in Agrostis disarticulates above the glumes.

Polypogon griquensis (Stapf) Gibbs Russ. &

L.Fish, comb. nov.

Agrostis griquensis Stapf in Kew Bulletin, Additional ser. 1897:

290 (1897). Type: Northern Cape, 2823 (Griquastad): near Griquatown,

(-CC), December 1811, Burchell 1863 (K, holo.; PRE fragment of

holo.).

Other specimens examined

FREE STATE. — 2926 (Bloemfontein): Bloemfontein, near Tempe

Farm. (-AA), Geo Potts 2467, fragment of Geo Potts 2470; both

October 1917.

NORTHERN CAPE. — 2824 (Kimberley): in Barkly West area,

Holpan, (-CB or -BC; there are two places with the same name in this

area), October 1937, Acocks 2466.

Gibbs Russell (1990) also placed Polypogon minit-

tifloms Pilg. into synonymy under P. griquensis. Pilger

(1941) described P. minutiflorus based on the specimen

‘Rusch. Jun. in herb. Dinter 7944, Leutwein’. The local-

ity is Namibia and may refer to a locality near Windhoek

and date of collecting as October 1934. Although the type

seems to have disappeared and has not been traced, Gibbs

Russell (pers. comm.) felt that the description matched

that of P. griquensis and that the two entities were there-

fore conspecific.

At PRE the only specimen from Namibia was collected

in the Naukluft River by VoUc {Volk 848) in October 1939.

It was originally identified as ^Polypogon minutiflorus ex

descr.’ but according to Gibbs Russell it is a good match of

the type of Agrostis griquensis housed at K.

Is Polypogon griquensis extinct or an alien grass?

According to the information gathered from collections

at PRE, Polypogon griquensis grows under wet conditions.

flowers in October and has not been collected since 1939.

The question is: is this an endemic species that is extinct,

rare or threatened, or was it an alien that never really

became naturalized? Some 18 species of Polypogon are

found in the warm temperate regions of the world and the

tropical mountains. Three species occur in the FSA {Flora

of southern Africa) region of which two, P. monspeliensis

(L.) Desf and P. viridis (Gouan) Breistr., are naturalized,

whereas P. strictus from the Eastern and Western Cape is

widely regarded as endemic. The author has not seen any

species other than those occurring in southern Africa.

ACKNOWLEDGEMENTS

1 wish to thank Dr G.E. Gibbs Russell for having done

most of the work, Ms E. du Plessis for helping with the

text, G. Germishuizen for editing and the referees for

their input.

REFERENCES

CHIPPfNDALL, L.K.A. 1955. A guide to the identification of grasses

in South Africa. In D. Meridith, The grasses and pastures of

South Africa. Trustees of the Grasses and pastures of South

Africa Book Fund, Johannesburg.

CLAYTON, W.D. & RENVOIZE, S.A, 1986. Genera gramineum.

Grasses of the World. Kew Bulletin, Additional Series XIII.

Royal Botanic Gardens, Kew.

GIBBS RUSSELL, G.E. 1990. Polypogon griquensis (Stapf) Gibbs

Russell ined. In G.E. Gibbs Russell, L. Watson, M. Koekemoer,

L. Smook, N.P. Barker, H.M. Anderson & M.J. Dallwitz,

Grasses of southern Africa. Memoirs of the Botanical Survey of

South Africa tSo. 58: 275.

LAUNERT, E. 1970. Prodromus einer flora von Siidwestafrika 160.

Gramineae: 155.

PILGER. R.K.F. 1941. Notizblatt des Botanischen Gartens und

Museums zu Berlin-Dahlem 15: 452.

L. FISH*

* National Herbarium, South African National Biodiversity Institute,

Private Bag XIOI, 0001 Pretoria.

MS. received: 2005-05-26.

POACEAE

A NEW SPECIES OF SPOROBOLUS (SPOROBOLINAE) IN SOUTH AFRICA

Sporobolus sp. {Smook 3429) in Gibbs Russell et al.:

313 (1990) is hereby named.

Sporobolus oxyphyllus L.Fish, sp. nov., S. vir-

ginico (L.) Kunth affmis sed foliis convolutis, paniculo

aperto axe principali visibili, habituque differt; etiam S.

pungenti (Schreb.) Kunth affmis, a quo partibus omnibus

parvioribus breviorbusque differt.

TYPE. — ^North-West, 2625 (Delareyville): Barberspan,

(-DA), Ellis 3628 (PRE, holo.).

Mat-forming perennial with robust rhizomes. Culms

erect, up to 500 mm tall, branched at base; nodes pale

to dark purple or black, glabrous, commonly covered by

leaf sheaths. Leaf sheaths persistent, strongly ribbed in

upper half, veins becoming less numerous and obvious

nearer base, sometimes flushed with purple, glabrous.

except sheath mouth with hairs up to 1.5 mm long.

Ligule with a fringe of hairs 0.2 mm long. Leaf blades

stiff, pungent, convolute, 20-90(-115) x 1.5^. 5 mm,

apex boat-shaped, veins numerous, midrib usually not

distinct, glabrous, adaxial surface with prickles and papil-

lae; margins smooth or scrabrid. Inflorescence an open,

ovate panicle, 30-100 x 10-30 mm, rarely contracted,

extending above main leaves, but often closely associated

with uppermost leaf; branches smooth or with scattered

prickles, primary branches not in whorls, ascending to

spreading at about 45° from main axis, lower part with-

out spikelets; yellow to red, glabrous pulvinus present in

axils of branches. Spikelets 1.8-2. 4 mm long, pallid to

grey-green sometimes flushed purple, these colours often

mixed on same spikelet, palea usually obvious, being

darker than rest of spikelet. Glumes unequal, obtuse to

acute, sometimes apiculate, 1 -nerved, nerve thickening

towards apex, upper V2 scabrid; lower glume 1 .0-1 .6 mm

72

Bothalia 36,1 (2006)

FIGURE 7. — Habit of Sporobolus oxyphyllus, x 0.7. Artist: Sibonela

Chiliza.

long; upper glume nearly as long as or as long as spike-

let, but length of both glumes variable even within same

panicle. Lemma ovate-elliptic, acute at tip, 1 -nerved.

Palea as long as spikelet to slightly longer, 2-keeled,

with a deep median groove. Stamens 3; anthers 1.2-1 .4

mm long, pallid, yellow to greenish often flushed purple.

Caryopsis ellipsoid, strongly laterally compressed, up

to 1.1 mm long. Flowering time'. November to February

with one specimen collected June. Figure 7.

Distribution and habitat'. Sporobolus oxyphyllus occurs

in the drier parts of the interior of South Africa, namely

in the southern North-West, western Free State and the

Eastern and Northern Cape bordering the Free State

FIGURE 8. — Known distribution of Sporobolus oxyphyllus.

(Figure 8). It forms dense stands and is locally abundant

in areas of high ‘sodic ‘soils especially at the edges of

salt pans and in salt vleis. It has been recorded growing

in water and is often found growing in association with

Cynodon dactylon.

Relationships'. Sporobolus oxyphyllus is similar to S.

virginicus (L.) Kunth which is found in saline areas in

coastal localites throughout the world in the tropics and

subtropics with some inland localities with sodic condi-

tions. In the Flora of southern Africa {FSA) region, S.

virginicus occurs along the coast, whereas inland locali-

ties lie mostly east and northeast of the distribution area

of S. oxyphyllus in KwaZulu-Natal (2632CD; 2632AB),

Mpumalanga (2231 BA, -CC) and Limpopo (2228DA,

2230CA). It also differs in having a dense, spike-like

panicle and leaves which are cauline and not mostly

basal. S. pungens (Schreb.) Kunth [5. arenarius (Gouan)

Duval-Jouve] from the Mediterranean, appears to be

even more similar, with open panicles and a comparable

growth form, but in S. oxyphyllus, all parts especially the

plant itself and the spikelets are larger and longer.

Etymology', the specific epithet oxyphyllus refers to

the sharply pointed leaves.

Other specimens examined (all housed in PRE)

NORTH-WEST.— 2625 (Delareyville): Paardefontein, {-CB), Allan

174', Farm Boskop, (-CB), Smook 627i\ Barberspan, (-DA); Davidse

& Loxton 6026', Zimbatis 4, 10.

FREE STATE. — 2727 (Kroonstad): Middelspruit Noord, (-CA),

Scheepers 1654. 2825 (Boshof): Du Plessis Dam, (-AD), Peyer 1077',

Berlyn, (-BC), Muller 1299', Myburgs Pan, Lynfontein Farm, (-DB).

Edwards 4152. 2826 (Brandfort): Bulfontein, (-AC), Smook 272P,

Krugersdrift Dam Nature Reserve, (-CC), Muller 1906.

NORTHERN CAPE. — 3024 (De Aar): near Potfontein station,

(-AA), Acocks 13532', Farm Biesiespan, (-DA), Smook 3429.

EASTERN CAPE. — 3025 (Colesberg): Oviston, (-DA), Anderson

349.

ACKNOWLEDGEMENTS

Thanks go to Dr O.A. Leistner for the Latin description.

Dr T. Cope of the Herbarium, Royal Botanic Gardens,

Bothalia 36,1 (2006)

73

Kew (K) for his advice, Ms H. Steyn for the map, Mr S.

Chiliza for the illustration, G. Germishuizen for the editing

and all those that helped in finalizing this paper.

REFERENCE

GIBBS RUSSELL, G.E., WATSON, L„ KOEKEMOER, M., SMOOK,

L„ BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J.

1990. Grasses of southern Africa. Memoirs of the Botanical

Survey of South Africa No. 58.

L. FISH*

* National Herbarium, South African National Biodiversity Institute,

Private Bag XlOl, 0001 Pretoria.

MS. received: 2005-05-26.

ASPHODELACEAE

ALOE VANROOYENll. A DISTINCTIVE NEW MACULATE ALOE FROM KWAZULU-NATAL, SOUTH AFRICA

INTRODUCTION

Although material of Aloe vanrooyenii Gideon F.Sm.

& N.R.Crouch from Weenen was known to well-known

aloe expert, Dr Gilbert W. Reynolds, he noted (Reynolds

1950: 266) in a somewhat noncommittal way that

plants ‘appear to be outlying forms of A. barbertoniae

However, Reynolds acknowledged that both morphologi-

cal and phonological differences were evident: Weenen

plants flowered later in the year (October and November)

and bore more crowded marginal teeth on the leaf. Our

view is that A. barbertoniae Pole Evans is a distinctly

subtropical entity centred on Nelspruit and Barberton in

Mpumalanga and that material from the KwaZulu-Natal

Midlands is sufficiently different to warrant recognition at

species rank. Pending a full pan- African revision of Aloe

sect. Pictae Salm-Dyck, we regard the closest affinities

of the new taxon to lie with A. parvibracteata Schbnland.

Although Van Wyk & Smith (2004) followed the pro-

posed inclusion of A. barbertoniae in A. greatheadii

Schbnland var. davyana (Schbnland) Glen & D.S. Hardy

(Glen & Hardy 1987), we consider that A. barbertoniae

will likely be reinstated in the course of a re-evaluation of

A. sect. Pictae. On the other hand. A. panibracteata has a

geographical distribution range centred in the mountain-

ous areas to the east and south of Nelspruit, Mpumalanga.

Although extending into northeastern KwaZulu-Natal,

this allied species is a distinctly winter-flowering entity,

which has longer, less deltoid-shaped leaves which are

paler green or distinctly purple, and smaller fruit.

One of the main distinguishing characters of A. van-

rooyenii is the large size of its erect, matt green to purple-

brown, cylindrical-oblong fruit. In the fresh condition,

shortly before dehiscence and seed dispersal, individual

capsules are typically 25-28 x 14—18 mm. In the dry

condition, and on herbarium specimens, the fruit shows

slight shrinkage, yielding capsules of (20-)21(-22) x

(10-)12 mm. In fact, this is the only maculate aloe in

which the robust inflorescence peduncle and side branch-

es carrying developed fruit, cannot support the weight of

the large, mature capsules and invariably bend towards

the ground.

Aloe vanrooyenii, flowering between October and

November, is the only early summer-flowering maculate

aloe from KwaZulu-Natal (Van Wyk & Smith 2004).

To the north of its distribution range, predominantly in

the Gauteng and Limpopo Provinces, A. transvaalensis

Kuntze, which is often included in the synonymy of A.

zebrina Baker, flowers from November to April. These

species respectively occupy similar summer-reproductive

niches in what is essentially more open grassland and

savanna vegetation.

Aloe vanrooyenii Gideon F.Sm. & N.R.Crouch, sp.

nov., apparenter parvibracteatae Schbnland arctissime

affmis, a qua fructibus capsularibus constanter multo

majoribus, fasciis perlatis albis longitudinalibus secus

margines secus quos capsulae dehiscunt omatis, mensi-

busque aestivus florens differt.

TYPE. — KwaZulu-Natal, 2830 (Dundee): alongside

road in hilly country, 25 km from Muden towards

Weenen (-CC), (S 28.93140, E 30.25385), N.R. Crouch

& G.F. Smith 2 (NH, holo.).

Small, slow-growing, herbaceous, succulent, perennial

herb, very rarely branching from base, not forming clus-

ters, consisting of small to medium-sized, densely foliate,

open rosettes, 240-300 mm diam. Roots terete, 4—5 mm

diam. Stems usually absent, if rarely present then up to

80 mm long and 45-55 mm diam., erect. Leaves 15-20,

rosulate, attenuate, tapering to apex, 120-150 mm long,

60-80 mm broad at base, basally sheathing, distinctly

spreading, apex dry, reflexed, dry leaves persistent; upper

surface shallowly and broadly canaliculate, flat near

base, shiny pale green, spotted, spots pale milky green to

whitish, variously shaped and sized, often ± confluent in

transverse bands; lower surface convex, uniformly milky

green, rarely with longitudinal darker greenish striations,

sometimes with small teeth arranged in a central row near

leaf apex; margins with a whitish, near-translucent edge,

armed with very pungent teeth; teeth brownish orange,

recurved like shark’s teeth, 3^ mm long, 3^ mm

apart, ± evenly spaced; leaf sap drying translucent, cut

end eventually turning purple. Inflorescence a sparsely

branched panicle, 500-800 mm tall, branched at or near

lower third with 1 or 2 branches, lowest branches re-

branched; 2 or 3 panicles produced consecutively, pedun-

cle and branches below racemes sometimes sterile brac-

teate, branches subtended at base by thin, scarious, pale

brown, many-nerved bracts up to 40 mm long, 1 5-22 mm

broad at base. Peduncle basally plano-convex, 10-20 mm

broad at base, matt greenish brown with a soft, whitish

bloom, sometimes sterile bracteate. Racemes cylindri-

cal to slightly conical, laxly flowered, flowering portion

250^70 X 70-90 mm; buds suberect, horizontal or

subpendulous, somewhat congested at apex, lowest open

flowers horizontal to subpendulous, racemes varying

74

Bothalia 36,1 (2006)

FIGURE 9. — Aloe vanrooyenii. A,

habit, X 0.8; B, dried capsules

showing prominent white

marginal strips along which

fruit dehisces, x 0.8; C, seeds

showing short wings, x 0.8; D,

raceme removed from an inflo-

rescence, X 0.8. Artist: Gillian

Condy.

in size according to age of plants, larger in old plants,

smaller in young plants. Bracts amplexicaul, small, thin,

scarious, dirty brownish white, margins creamy white,

8-10 mm long. Pedicels 8-10 mm long. Flowers 33-38

mm long, swollen at base, basal swelling 8-10 mm diam.,

cylindric-trigonous, usually monochrome; buds with

creamy white tips, otherwise uniformly orange or red;

mature flowers 8 mm diam. in middle, distinctly trigo-

nously indented above ovary, thence enlarging towards

throat and forming wide open mouth; outer segments

free for 8-15 mm from apex, free portion with 1 distinct,

central, somewhat more intense orange or red section,

with 1 mm broad creamy white border and subacute,

recurved apex; inner segments broader than outer ones,

with 1.0-1. 5 mm broad creamy white border and more

obtuse, spreading apex, dorsally adnate to outer ones for

their greater length. Stamens 6, hypogynous; filaments

slightly flattened, pale lemon-yellow, 30-33 mm long,

all 6 of equal length, withering concertina-like with

age; anthers small, purple-black, versatile, exserted for

up to 3 mm. Ovary 6-7 x 3 mm, pale greenish yellow;

style 25-30 mm long, minutely capitate; stigma small.

Fruit a large capsule, erect, matt green to purple-brown,

cylindric-oblong, 25-28 x 14-18 mm, apically truncate,

trilocular, dehiscing loculicidally, chartaceous to woody

when dry, margins of valves, along which split occurs,

broad, white, very conspicuous, for some time wrapped

in remains of dry perigone. Seed angled, 2.0 x 2.5 mm,

dark brown with pale brown to greyish white wing.

Chromosome number: unknown. Figure 9.

Distribution: the species is centred around Ladysmith,

and is known from the Dundee, Harrismith and Underberg

Districts (Figure 10). Within this region it is commonly

found as a thomveld savanna component.

Eponymy: the species is named after Mr Gert van

Rooyen of Greytown who prompted further investigation

into wild populations of the species.

Bothalia 36,1 (2006)

75

FIGURE 10. — Known distribution of Aloe vanrooyenii based on speci-

mens at NH and NU.

ACKNOWLEDGEMENTS

We thank the following staff members of the South

African National Biodiversity Institute, Pretoria, South

Africa: Dr Otto Leistner for providing the Latin diagno-

sis, Ms Emsie du Plessis for commenting on a draft of

this paper and Ms Gill Condy for preparing the accompa-

nying line drawing. Mr Brian Tarr of the KwaZulu-Natal

National Botanical Garden in Pietermaritzburg assisted

in the field and kindly facilitated communication with

Mr van Rooyen. We are also grateful to Messrs Charles

Craib, John J. Lavranos and an anonymous referee for

helpful comments.

REFERENCES

GLEN, H.F. & HARDY, D.S. 1987. Nomenclatural notes on three

southern African representatives of the genus Aloe. South

African Journal of Botany 53: 489-492.

REYNOLDS, G.W. 1950. The aloes of South Africa. The Trustees of

the Aloes of South Africa Book Fund, Johannesburg.

VAN WYK, B-E. & SMITH, G. 2004. Guide to the aloes of South

Africa, edn 2. Briza Publications, Pretoria.

G.F. SMITH* andN.R. CROUCH**

* South African National Biodiversity Institute, Private Bag XI 01,

0001 Pretoria/Schweickerdt Herbarium, Department of Botany, Univer-

sity of Pretoria, 0002 Pretoria.

** Ethnobotany Unit, South African National Biodiversity Institute,

P.O. Box 52099, Berea Road, 4007 Durban/School of Chemistry, Uni-

versity of KwaZulu-Natal, Private Bag XOl, 3209 Scottsville, South

Africa.

MS. received: 2005-09-09.

ASPHODELACEAE

ALOE KAOKOENSIS, A NEW SPECIES FROM THE KAOKOVELD, NORTHWESTERN NAMIBIA

Aloe kaokoensis Van Jaarsv., Swanepoel & A.E.van

Wyk, a new species allied to^. littoralis Baker, is described

from the Otjihipa Mountains, Kaokoveld, northwestern

Namibia. It is one of 28 currently recognized species

of Aloe L. in Namibia (Rothmann 2004). Nine of them

have been recorded in the Kaokoveld. This is the second

new species recently described from the Kaokoveld, an

indication of the poor state of botanical exploration in

this arid, mountainous region. The first. Aloe omavandae

Van Jaarsv. & A.E.van Wyk, was described from the cliff

faces of the eastern Baynes Mountains (Van Jaarsveld &

Van Wyk 2004; Van Jaarsveld et al. 2005).

Aloe kaokoensis Van Jaarsv. , Swanepoel &A.E. van

Wyk, sp. nov., A. littorali affinis, sed caulibus senectis

decumbentibus, folds dense rosulatis laete glaucis cum

dentibus nigris sursum spectantibus, perianthio 35 mm

longo differt.

TYPE. — Namibia, 1712 (Swartbooisdrif): Kaokoveld,

Otjihipa Mountains, ± 5 km ESE of Otjinhungwa grow-

ing on rocky outcrops, 1 000 m, (-BC), Van Jaarsveld &

Swanepoel 19504 (WIND, holo.).

Plants solitary, rosulate, up to 0.73 m tall and 1.3

m diam., shortly caulescent; stem up to 1 m long and

100 mm in diam., erect when young, becoming decum-

bent with age, covered with remains of old dry leaves;

bark grey. Roots grey, 3^ mm thick. Juvenile leaves

distichous at first, lorate-triangular and biconvex, both

surfaces distinctly spotted, some spots with short spines;

apex mucronate. Mature leaves 35^0, in a dense rosette.

arcuate-ascending, lower leaves spreading, tough, with

asymmetric keel near apex, triangular-lanceolate, 380-670

X 70-140 mm; upper surface flat, but becoming chan-

nelled in upper half, pale glaucous, slightly rough to

touch, sparsely white-spotted in basal third, occasionally

without spots, with lenticular spots irregularly arranged;

lower surface flat to slightly convex at first, becoming

convex, copiously white-spotted, with lenticular spots

arranged in obscure white bands; margin yellowish green,

cartilaginous, translucent, armed with deltoid-acuminate,

small, black (yellowish to reddish brown at first) teeth

3^ X 4-5 mm, 7-15 mm apart (8-9 mm apart near apex),

projecting towards apex; apex acute, ending in 1-3 teeth.

Leaf sap copious, yellowish, drying honey-coloured to

dark brown to black. Inflorescence: 1 or 2 per plant, a

much-branched panicle, 1.23-1.42 x 0.6-0.85(-1.12) m,

bearing many lateral branches in upper half; peduncle

biconvex, grey-green, up to 40 mm diam. at base and

unbranched for up to 220 mm, with powdery bloom,

flattened and marginiform (bearing an acute margin

opposite to each other) at base for 1 50-200 mm; racemes

cylindrical-acuminate, 330^70 x 700 mm; flowers laxly

arranged; pedicels 11(-14) mm long; bracts ascending,

navicular, linear-lanceolate, 10-12 x 3 mm, channelled,

acuminate, whitish, thin, scarious. Perianth orange-red

becoming slightly yellowish when open, cylindrical-trig-

onous, 35 X 6. 0-6. 5 mm, rounded at base; outer segments

free to halfway down; inner segments free but dorsally

adnate to outer over basal half; apices acute to subacute.

Stamens included but sometimes shortly exserted; anthers

oblong, 3 X 1 mm; filaments filiform-flattened, pale yel-

low, inner 3 narrower and slightly longer; pollen orange.

76

Bothalia 36,1 (2006)

Ovary green, cylindrical, 6-grooved, 8 x 2-3 mm; style

25 mm long, shortly exserted (sometimes up to 7 mm);

stigma capitate. Capsule cylindrical-trigonous, erect,

20-26 X 8-11 mm. Seed 3x2 mm, angular, winged,

grey-black. Figure 11.

Aloe kaokoensis is distinguished by its large, solitary

rosettes (up to 1.3 m in diameter) of pale, glaucous leaves

with black teeth, of which the apices project towards the

leaf apex, and large dense panicles (young erect plant

depicted in Figure 11) and stems that are invariably

decumbent in old plants. It is clearly related to Aloe litto-

ralis in its floral features. A. littoralis is found throughout

Namibia in suitable habitats and also occurs in Angola,

Zimbabwe, Mozambique and Botswana. A. littoralis

always has an erect stem up to 3 m tall and greyish green

leaves that are mottled or spotted when young, but usu-

ally without spots in adult plants. Furthermore, the leaves

have brown to reddish brown teeth. The perianth of A.

kaokoensis is 35 mm long, thus slightly longer than in^.

littoralis (23-34) (Reynolds 1974). A. kaokoensis con-

tains copious quantities of yellow leaf sap, which turns

honey-coloured to dark brown when exposed to air. Its

large, branched, paniculate inflorescence bears ascending

racemes of orange-red flowers, 35 mm long. A. kaokoen-

sis may also be confused with two other aloes with pale

glaucous leaves (Table 1). The first, A. namibensis Giess,

is a much smaller species occurring to the south and its

flowers are coral-pink. The second, A. dewinteri Giess, is

a cliff dweller from the Sesfontein region, bearing coral-

pink flowers that turn white when fully open.

Aloe kaokoensis grows at altitudes of 700 to almost

2 000 m on the northwestern extreme of the granitic

Otjihipa Mountains (eastern margin of the Marienfluss)

(Figure 12). The plants are sometimes locally abundant

where they grow in granitic soil and quartz fragments,

usually in full sun. A. kaokoensis is a constituent of arid

Colophospermum mopane (mopane) woodland, with

several species of Commiphora prominent, for example.

Commiphora glaiicescens, C. multijuga, C. tenuipetiolata,

C. virgata and C. wildii. Other associated species include

Adenium boehmianum. Aloe dinteri, Boschia tomentosa,

Ceraria longipedimculata, C. carrissoana. Euphorbia

guerichiana, E. monteiroi, E. virosa, Ledebouria sp.,

Lycium sp., Rhigozum virgatum, Sterculia africana, S.

STeAOH<^N '06

FIGURE II. — Aloe kaokoensis.

A, young adult plant (stem

not yet decumbent) with

inflorescence, x 0.08; B,

mature and juvenile leaf

apex, X 0.8; C, raceme,

showing flowers, x 0.6;

D, infructescence show-

ing capsules, x 0.6. Artist:

Lisa Strachan.

Bothalia36,l (2006)

77

TABLE 1 . — Main differences between Aloe kaokoensis, A. dewinteri, A. namibensis and A. littoralis

FIGURE 12. — Known geographical range of Aloe kaokoensis.

quinqueloba and a Polygala sp. The discovery of Aloe

kaokoensis brings to four the number of Aloe species

endemic or largely confined to the Kaokoveld Centre of

Endemism (Van Wyk & Smith 2001), the others being A.

corallina I. Verd., A. dewinteri and A. omavandae.

The flowering time of A. kaokoensis ranges from

March to May. Dusky sunbirds {Nectariina fusca) have

been observed visiting the plants, and appear to be

the main pollinators. Average annual rainfall in the

Kaokoveld varies from less than 50 mm along the coast

to ± 350 mm in the highlands (Mendelsohn et al. 2002).

Rainfall is erratic and occurs mainly in the form of thun-

dershowers in summer. On the Otjihipa Mountains where

the new species is found, the average annual rainfall is

estimated at ± 150-200 mm. The coastal mountains are

also subject to fog from the Atlantic Ocean (60 km from

the Otjihipa Mountains).

ACKNOWLEDGEMENTS

We thank Gerrit Germishuizen and Emsie du Plessis

for editing an early draft of the text, Hugh Glen for trans-

lating the diagnosis into Latin and Lisa Strachan for pre-

paring the illustration. The Ministry of Environment and

Tourism in Namibia is thanked for providing the neces-

sary plant collecting permits. Mr Koos Verwey of Syncro

Camp along the Kunene River (Marienfluss region) is

thanked for his help and assistance.

REFERENCES

MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T.

2002. Adas of Namibia. David Phillip, Cape Town.

REYNOLDS, G.W. 1974. The aloes of South Africa, edn 3. Balkema,

Cape Town.

ROTHMANN, S. 2004. Aloes. Aristocrats of Namibian flora. Creda,

Cape Town.

VAN JAARSVELD, E.J. & VAN WYK, A.E. 2004. Aloe omavandae,

a new species from the Kaokoveld, northwestern Namibia.

Haseltonia 10: 41-^3.

VAN JAARSVELD, E.J., VAN WYK, A.E. & CONDY, G. 2005. Aloe

omavandae (Asphodelaceae). Flowering Plants of Africa 59:

2-6, t. 2201.

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

in southern Africa. A review with emphasis on succulents.

Umdaus Press, Hatfield, Pretoria.

E.J. VAN J AARS VELD* t, W. SWANEPOEL * * and A.E. VAN WYK* ** ***

* South African National Biodiversity Institute, Private Bag X7, 7735

Claremont, Cape Town.

t Student affiliation: Department of Botany, University of Pretoria,

Pretoria.

** Department of Botany, University of Pretoria, 0002 Pretoria; P.O.

Box 21168, Windhoek, Namibia.

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

sity of Pretoria, 0002 Pretoria.

MS. received: 2005-05-04.

AMARYLLIDACEAE

A NATURAL HYBRID IN THE GENUS CLIVIA

The genus Clivia, with six described species, is The group is of considerable horticultural significance

endemic to South Africa and Swaziland (Duncan 1999; and its members, including many cultivars, are exten-

Rourke 2002; Swanevelder 2003; Murray et al. 2004). sively grown all over the world (Swanevelder 2003).

78

Bothalia36,l (2006)

Artificial hybridization between the different species

of Clivia results in attractive progeny, currently highly

sought after in eultivation (Koopowitz 2002; Swanevelder

2003). Numerous references to such hybrids between C.

miniata (Lindl.) Regel and C. nobilis Lindl.; C. miniata

and C. garden a Hook.; and C. miniata and C. caiilescens

R. A. Dyer have been recorded in the literature (Koopowitz

2002; Rourke 2003; Swanevelder 2003). Natural inter-

specific hybridization in the genus has, however, rarely

been documented. Rourke (2003) reported a natural

hybrid between C. miniata and C. caulescens from the

Bearded Man Mountain near Barberton, Mpumalanga,

and its subsequent cultivation at Kirstenbosch National

Botanical Garden. Swanevelder (2003) reviewed both the

natural and artificial hybrids known in the genus.

The recognition of hybrids in nature is sometimes

regarded as somewhat speculative and the existence of

such entities is usually based on circumstantial evidence.

Generally, the possession of intermediate morphological

features, proximity to the putative parents, hybrid fertility

with segregation recognizable in the F2 progeny, prefer-

ably supplemented by the artificial hybridization of the

putative parents, are applied as criteria in support of a

natural hybrid (Stewart & Manning 1982).

Hitherto, no natural hybrids (nothotaxa) have been

formally described in Clivia — despite the existence of

numerous records indicating that at least two species

grow sympatrically in different localized populations

(over the full distribution range of the genus). Here, for

the first time, we formally describe a natural hybrid in

Clivia. The identity of the hybrid is supported by the

four criteria stipulated above (Stewart & Manning 1982).

The new nothospecies is intended to eover all hybrids

between C. miniata (both var. miniata and var. citrina)

and C. caulescens.

Clivia X nimbicola Z.H. Swanevelder, J.T.Truter

& A.E.van Wyk, nothosp. nov., hybrida naturalis inter

Cliviam miniatam et C. caulescentem. Rhizoma aeria

inter illos specierum parentalium intermedia. Folia arcu-

ata, 250-350 mm longa, 55-70 mm lata, marginibus

integris, apice acuta. Flores numero 10-20(-30); pedi-

cellis suberecto, perianthio tubiformi, 30-60 mm longo,

suberecto, segmentis patulis.

TYPE. — Swaziland, 2531 (Barberton): Bearded Man

Mountain, (-CB), 30-05-2003, Pearton TPOl (PRU,

holo.).

Rhizomatous, solitary or clumping, stout, evergreen

perennial, 0.4-1. 2 m tall. Leaf sheath green to pale red.

Leaves long-lived, arching, strap-shaped, 250-350 x 55-

70 mm, apex acute; margins usually entire, rarely ± ser-

rated. Scape 200-600 x 1 0-30 mm, green. Inflorescence

umbel-like, 10-20(-30 (-flowered, usually loose/open and

± flat-topped; pedicels stiff, erect to drooping, 1 5^0 mm

long, green. Flowers semi-erect to drooping. Perianth

tubular to trumpet-shaped, 30-60 mm long, segments

spreading, pastel orange to pastel pink, occasionally with

green apices. Stamens 6, variable in degree of exsertion

at anthesis. Style 30-55 mm long; stigma trilobed, apex

penicillate. Ovary ovoid, green, trilocular. Fruit 10-30

berries, green maturing to red. Seeds \-A, subglobose, ±

10-15 mm in diam., pearly white. Figure 13.

Clivia X nimbicola is intermediate between Clivia cau-

lescens and Clivia miniata with regards to rhizome, leaf,

umbel and flower morphology (Table 2). Flower colour

ranges from pastel orange to pastel pink, with green tepal

apiees in some speeimens. Flowering is erratie, oceurring

all year round, mainly from July through to December,

with some speeimens flowering twice yearly — February

to May. This long flowering period eonnects the flower-

ing periods of the two putative parents in the Bearded

Man Mountain locality, namely October-November in

C. miniata and October-Deeember in C. caulescens. The

extended flowering period of C. x nimbicola is regarded

as further evidence in support of the taxon’s hybrid

origin. The formal deseription of C. x nimbicola is also

supported by the observation that the hybrid plants bear

berries in the wild, thereby inferring fertility and the pos-

sibility to maintain populations by means of subsequent

breeding among hybrid plants.

Field observations suggest some introgression between

Clivia X nimbicola and its putative parents. Where popula-

tions of C. X nimbicola occur close to or amongst C. cau-

lescens, back-crossing of the hybrid with C. caulescens

produces umbels with fewer flowers which are tubular,

yet more open than in typieal C. caulescens. Likewise,

where the hybrid occurs close to or amongst C. miniata,

the umbels are less floriferous and the flowers are more

funnel-shaped, yet not as open as in typical C. mini-

ata. It is suggested that from the inferred initial progeny

cross, subsequent generations have resulted from various

backcrossings, resulting in a hybrid-swarm. Artificial

hybridization between C. miniata and C. caulescens is

usually sueeessful with records dating back to 1945 (e.g.

1945/66, R. Marais PRE37 106). Morphologically the resul-

tant hybrids closely mateh the plants of C. x nimbicola

in the wild.

The holotype of Clivia x nimbicola was eolleeted on

the Bearded Man Mountain near Barberton (Figure 14),

on the border between South Africa and Swaziland. In this

area these natural hybrids are quite eommon (50 or more

individuals) in sympatric stands of C. miniata and C.

caulescens. Judging by plant sizes and the height of aerial

stems, original hybrids are as old as their putative parents.

Toppled plants with long aerial stems, from either parent-

age of hybrid, freely produce suckers when in contaet

with the soil. Seedlings, when present, were usually dis-

tributed around its putative parent. The natural distribu-

tion range of C. x nimbicola is confined to the Barberton

Centre of Endemism (Van Wyk & Smith 2001), the only

known region in which the distribution ranges of C. cau-

lescens and C. miniata overlap (Swanevelder 2003). At

least three separate, well-established populations of C.

X nimbicola were recorded, with stands extending into

both Swaziland and South Africa. C. caulescens prefers

steep cliff faces or steep rocky embankments, whereas

C. miniata generally prefers gentler scree embankments

or flatter forest habitats. The C. x nimbicola plants are

distributed between and amongst both parents, occupying

both specific habitats found in the Afromontane Forest.

The epithet nimbicola means dweller in the mist, and

Bothalia 36,1 (2006)

79

FIGURE 13. — Type specimen of Clivia x nimbicola: A, inflorescence; B, flower in 1/s. Scale bars: A, 30 mm; B, 15 mm. Artist: Magda Nel.

80

Bothalia 36,1 (2006)

TABLE 2. — Comparison of Clivia x nimbicola with its putative parents at Bearded Man Mountain, Mpumalanga and Swaziland

FIGURE 14. — Known geographical range of Clivia ^ nimbicola in

nature.

refers to the mist belt habitat in which this hybrid and its

putative parents are found.

ACKNOWLEDGEMENTS

Our thanks to Dr Hugh Glen for the Latin translation

of the diagnosis, Ms Hester Steyn for the distribution map

and Mr T.N. Pearton for the type specimen.

REFERENCES

DUNCAN, G. 1999. Grow clivias. Kirstenbosch Gardening Series.

National Botanical Institute, Cape Town.

KOOPOWITZ, H. 2002. Clivia. Timber Press, Singapore.

MURRAY, B.G., RAN, Y, DE LANGE, P.J., HAMMETT, K.R.W.,

TRUTER, J.T. & SWANEVELDER, Z.H. 2004. A new species

of Clivia (Amaryllidaceae) endemic to the Pondoland Centre

of Endemism, South Africa. Botanical Journal of the Linnean

Society 146: 369-374.

ROURKE, J.P. 2002. Clivia mirabilis (Amaryllidaceae: Haemantheae)

a new species from Northern Cape, South Africa. Bothalia 32:

1-7.

ROURKE, J.P. 2003. Natural interspecific hybrids in Clivia — C. mini-

ata X C. caulescens hybrids from Mpumalanga. In M. Dower,

C. Felbert, J. van der Linde & J. Winter, Clivia 5: 78-80. Clivia

Society South Africa, Cape Town.

STEWART, J. & MANNING, J.C. 1982. A new Disa hybrid in Natal.

The South A frican Orchid Journal 13: 35-41.

SWANEVELDER, Z.H. 2003. Diversity and population structure of

Clivia miniata Lindl. (Amaryllidaceae): evidence from molecu-

lar genetics and ecology. M.Sc. thesis. University of Pretoria.

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

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Pretoria.

Z.H. SWANEVELDER*, J.T. TRUTER** and A.E. VAN WYK***

* Corresponding author: Department of Botany & Forestry and Agri-

cultural Biotechnology Institute (FABI), University of Pretoria, 0002

Pretoria

**P.O. Box 5085, 1502 Benoni South, South Africa.

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

sity of Pretoria, 0002 Pretoria.

MS. received: 2005-05-23.

Bothalia 36,1 (2006)

81

CAPPARACEAE

MAERUA KAOKOENSIS, SPECIES FROM NAMIBIA

INTRODUCTION

Maerua kaokoensis Swanepoel, a new species con-

fined to the Kaokoveld Centre of Endemism (Van Wyk

& Smith 2001), is described. During fieldwork for

the Namibian Tree Atlas Project, the author encoun-

tered Maerua trees in the remote Okakora Mountains,

Kaokoveld. with the peculiar habit of being tall, erect

and extremely slender, with a few long, thin branchlets

near the apex, all of which are distinctly drooping or pen-

dulous. At a distance, the trees superficially resembled

Acacia robynsiana Merxm. & A.Schreib., a Kaokoveld

endemic with a remarkably similar habit. Subsequently,

another population of this Maerua was discovered in the

Otjihipa Mountains further to the west where flowers and

fruit were collected.

A study of the Maerua holdings in PRE and WIND

revealed several earlier collections of the new species,

all filed under M. schinzii Pax. When without flowers,

herbarium specimens of M. kaokoensis can easily be

mistaken for M. schinzii or M. angolensis DC. (Killick

1970). This resemblance is due to similarities in leaf and

fruit morphology. In the field, however, M. kaokoensis

is quite conspicuous due to its unusual weeping habit

(Figure 15), which is unlike that of any other member of

Maerua in southern Africa.

Maerua kaokoensis Swanepoel, sp. nov., M. schin-

zii Pax similis foliis nonnullis lamina elliptica, disco flori

coronato, impariter laciniato, ffuctu moniliformi; sed

caule pergracillimo, foliorum lamina non solum elliptica

sed etiam oblanceolata, lanceolata, lineari-elliptica, line-

ari-oblonga vel oblonga, non solum flavoviridi sed etiam

prasina vel atro-olivacea marronino-suffusa, coriacea

vel chartacea, petiolo semper gracillimo, saepe longiori,

margine disci semper cum fibrillis longis irregularibus;

androphora filamentisque staminum longioribus, gyno-

phora plerumque longiori, ovulis pluribus, tota semper

glabra praeter sepalorum faciem adaxialem er sutura,

Novembri usque ad Julio florenti differt.

TYPE. — ^Namibia, 1712 (Swartbooisdrif): Otjihipa

Mountains, 8 km ESE of Otjinhungwa, 1 850 m, (-BC),

17-01-2005, Swanepoel 172 (WIND, holo.!; PRE, iso.!).

Slender tree up to 10 m tall. Trunk single or rarely with

2 or 3 stems from ground level, occasionally branching

into 2 or 3 stems, erect and ± straight, extremely slen-

der with no lateral branches except for few drooping or

pendulous branches near apex, apex usually drooping;

stems 20^0(-60) mm in diam. Bark smooth, pale ashy

grey to reddish grey, with scattered, small indentations

in places. Branches glabrous with numerous, scattered,

small, whitish lenticels, young branches pale ashy grey,

reddish brown or yellowish brown, new growth yellow-

ish green or maroon. Leaves simple, petiolate, alternate,

spirally arranged, drooping or pendulous, glabrous, yel-

lowish green, green or dark olive-green with a maroon

tinge, emitting audible clatter when flapping against each

other in wind; lamina lanceolate, oblanceolate, narrowly

elliptic to elliptic, linear-elliptic, linear-oblong or oblong.

(16-)25-60(-95) X (5-)7-17(-30) mm, apex acute or

obtuse, rarely truncate or emarginate, mucronate, mucro

small, up to 0.8 mm long, base cuneate or cuneate to

rounded, rarely abruptly attenuate onto petiole, charta-

ceous to coriaceous; margin entire; midrib conspicuous

and prominently raised abaxially, yellowish green or

maroon; lateral veins 4-10, looping before margin, usu-

ally somewhat or often completely immersed abaxially;

petiole very slender, ( 10-)21^3(-50) x 0.3-1. 2 mm,

often slightly swollen over basal part, channelled in basal

part, yellowish green, reddish brown or maroon, glabrous.

Inflorescences short corymbose racemes, borne termi-

nally or on short lateral branches. Flowers pedicellate;

pedicel glabrous, 4-13 mm long. Receptacle cylindrical,

9-13 mm long, 2-3 mm wide at mouth, slender, ribbed,

glabrous; disc square in surface view, shortly coronate,

solid basal portion 0.4-2. 4 mm high, margin unequally

laciniate with long, irregular fimbrillae; fimbrillae usu-

ally branched as well as irregularly curved and recurved,

up to 2.6 mm long. Sepals 4, elliptic or spathulate, often

somewhat cucullate, 9.8-14.5 x 4. 0-5. 7 mm, apex acute

or obtuse, green, puberulous adaxially, glabrous abaxi-

ally, margin woolly. Petals absent. Androphore equal in

length to receptacle, or extending to 1 mm below or 2.5

mm above its upper rim, 11-13 mm long. Stamens 32-42,

pale yellowish green; filaments 18-35 mm long; anthers

oblong, ovate-elliptic or narrowly elliptic, basifixed,

1.8-2. 5 mm long. Gynophore 18-28 mm long, yellow-

ish green. Ovary cylindrical, 4.8-7. 1 x 0.9 mm, green;

ovules 48-52; stigma capitate. Print moniliform, up to

180 X 5-8 mm, green, faintly colliculate. Seeds globose,

4—5 mm diam., testa thinly textured, rather fragile, faintly

granulate, yellowish cream-coloured. Flowering time:

November to July. Figures 15, 16.

Diagnostic characters and affinities: Maerua kaokoen-

sis differs from M. schinzii and M. angolensis in habit

as well as in leaf and flower characters. Plants of M.

kaokoensis are markedly different from M. schinzii and

M. angolensis in being extremely slender, yet tall, with

only a few drooping or pendulous branches at the apex.

Usually the apical part of the main stem (leader shoot)

also droops. Trees of M. schinzii and M. angolensis have

a thick stem(s), which branches repeatedly to form a dis-

tinct, rounded crown.

The leaf lamina of Maerua kaokoensis is lanceolate,

narrowly elliptic to elliptic, linear-elliptic, linear-oblong,

oblong or oblanceolate, with 4—10 lateral veins on eaph

side of the midrib. In M. schinzii the lamina is elliptic to

broadly elliptic or ovate to narrowly ovate with only 4

or 5 lateral veins on each side. In the Flora of southern

Africa [FSA'\ region, M. angolensis has the lamina ellip-

tic, ovate, or obovate, also with only 4 or 5 lateral veins

on each side. Outside the FSA region, M. angolensis has

leaves, in addition to those mentioned, with lamina lan-

ceolate, ovate-lanceolate, linear-lanceolate or suborbicu-

lar, with 5 or 6 lateral veins on each side. Furthermore,

the lamina in M kaokoensis is coriaceous to chartaceous,

whereas in M. schinzii it is coriaceous only and in M

angolensis softly chartaceous only, although outside the

82

Bothalia 36,1 (2006)

FIGURE 15. — M kaokoensis in its natural habitat: A, ± 5 m tall; B, ± 7 m tall.

FSA region, M. angolensis can be coriaceous too. The

petiole of M kaokoensis is conspicuously slender, up to

50 mm long, whereas in M schinzii and M. angolensis

it is thicker, much more sturdy and up to 30 mm long.

The leaf lamina of Welwitsch 968b in BM (holotype of

M angolensis var. heterophylla Welw. ex Oliv.), a shrub,

1.0- 1.3 m high, from Luanda, Angola, superficially

resembles those in one collection of the new species,

namely Swanepoel 173. However, the lamina in the lat-

ter collection is linear-elliptic or linear-oblong with the

petiole long and very slender, whereas the lamina in the

Welwitsch specimen is linear-lanceolate and the petiole is

much shorter and not very slender.

The disc margin in Maenia kaokoensis differs from

M. schinzii by the apices being consistently fimbriate.

In M schinzii the fimbrillae are often absent and in M

angolensis {FSA region) they are always absent. In M

kaokoensis, the 11-13 mm long androphore is equal in

length to the receptacle, or extends to just above or below

its rim. In M. schinzii, the androphore is much shorter,

5-7 mm long and equal to or exserted above the recepta-

cle, whereas in M. angolensis {FSA region), it usually is

longer (12-17 mm) than in M kaokoensis and projected

beyond the receptacle. The gynophore is usually longer

in M. kaokoensis (18-28 mm) than in M. schinzii (15-20

mm) and shorter than in M. angolensis (35-37 mm).

All parts of Maerua kaokoensis are glabrous, except

for the sepals, which are puberulous adaxially and woolly

on the sutures. On the other hand, all parts of M schinzii,

are usually puberulous, whereas M. angolensis is gla-

brous or rarely puberulous. Flowering time is also diag-

nostic: November to July in M. kaokoensis, September

and October in M schinzii and July to December in M.

angolensis (Coates Palgrave 2002). As to habitat prefer-

ence, M kaokoensis occurs in mountains only, whereas

M schinzii and M angolensis grow on plains and in river

valleys as well.

Some of the more prominent morphological features

to differentiate Maerua kaokoensis, M. schinzii and M

angolensis are compared in Table 3. Diagnostic features

were determined through examination of herbarium

specimens and for M kaokoensis and M. schinzii, plants

were also examined in the field. For M angolensis, plants

in the Weenen District (KwaZulu-Natal) were examined.

Additional information for M schinzii and M. angolensis

(in the FSA region) is mainly from Killick (1970) and

Roessler (1966). Information on M angolensis from out-

side the FSA region was sourced from literature (Oliver

1868; Exell & Mendonga 1937; Hauman & Wilczek

1951; Wild 1960; Elffers et al. 1964).

Etymology, the specific epithet refers to the Kaokoveld

of northwestern Namibia. The distribution of M. kaokoen-

sis falls within the previous politically demarcated

Kaokoland, now called the Kunene Region.

Distribution: M. kaokoensis is presently known only

from the Okakora (part of the Baynes Mountains) and

Otjihipa Mountains, south of the Kunene River in north-

western Namibia (Figure 17). It is localized and uncom-

mon to rare in these areas. The species almost certainly

Bothalia 36,1 (2006)

83

FIGURE 16. — Maerua kaokoensis. A, mature leaves from different plants to show variation in size and shape; B, flower, Swanepoel 175\ C, recep-

tacle and disc, Swanepoel 175', D, fruit, Swanepoel 172. Scale bars: A, 20 mm; B, D, 10 mm; C, 5 mm. Artist: Julia Kreiss.

occurs in the adjacent mountainous parts of southwestern Habitat and ecology: M. kaokoensis grows on dolomite

Angola as well, especially the Serra Cafema range, and of the Otavi Group in the Okakora/Baynes Mountains and

may eventually prove to be more widespread on the high on paragneiss of the Epupa Metamorphic Complex in the

mountains of the Kaokoveld Centre of Endemism (Van Otjihipa Mountains (Miller & Schalk 1980; Mendelsohn

Wyk & Smith 2001), most of which remain botanically et al. 2002). It occurs on steep mountain slopes and less

poorly explored. often on plateaus and mountaintops. Its distribution

84

Bothalia 36,1 (2006)

TABLE 3. — Prominent differences between Maerua kaokoemis, M. schinzii and M. angolensis

ranges from 75-120 km from the Atlantic Ocean, from

the edge of the Great Escarpment eastwards, at altitudes

ranging from 700-1 850 m. Average annual rainfall var-

ies from 100-200 mm in these areas.

Specimens examined

NAMIBIA. — 1712 (Swartbooisdrif): Otjihipa Mtns, 5 km ESE of

Otjinhungwa, (-AD), Swanepoel 173, 174 (WIND); Otjimborombonga,

(-BB), Leistner, Oliver, Steenkamp & Vorster 142 (PRE); Baynes Mtns,

Okombambi, (-BB), Rusch 77 (WIND); Okakora Mtns, 2 km NW of

Okombambi, (-BB), Swanepoel 121 (WIND); Otjihipa Mtns, (-BC),

Craven 945 (WIND); Otjihipa Mtns 7 km SE of Otjinhungwa, (-BC),

Swanepoel 171 (WIND); Otjihipa Mtns, 8 km ESE of Otjinhungwa,

(-BC), Swanepoel 172 (PRE, WIND); Otjihipa Mtns 7.8 km ESE of

Otjinhungwa, (-BC), Swanepoel 175 (WIND); Orukatoa, Otjihipa

Mtns, (-BC), Viljoen 575 (WIND); NW of Otjitanda, (-DB), Meyer

1289 (WIND).

ACKNOWLEDGEMENTS

I would like to thank Prof A.E. van Wyk, University

of Pretoria, for advice and support. Dr H.F. Glen, SANBI,

for translating the diagnoses into Latin, Ms H. Steyn,

SANBI, for preparing the distribution map and Ms J.

Kreiss for the line drawings. The curator and staff of the

National Herbarium of Namibia are thanked for their

assistance during visits to the herbarium. The curator.

National Herbarium, Pretoria, is thanked for access to

their collections; the assistance of Dr C.L. Bredenkamp

and Mrs M. Jordaan during visits to the herbarium

is acknowledged with thanks. Ms V. Noble from the

National History Museum, London, is thanked for imag-

es of Angolan material. I am especially grateful to my

wife Hannelie and to Mr E. van Jaarsveld for assistance

and support during field trips.

REFERENCES

COATES PALGRAVE, M. 2002. Keith Coates Palgrave Trees of south-

ern Africa, edn 3, Struik, Cape Town.

Bothalia36,l (2006)

85

ELFFERS, J., GRAHAM, R.A., DEWOFF, G.P. & HUBBARD, C.E,

1964. Capparidaceae. Flora of tropical East Africa: 28: 1-88.

Crown Agents for Oversea Governments and Administrations,

London.

EXELL, A.W. & MENDONt^A, F.A. 1937. Capparidaceae. Conspectus

florae angolensis 1: 53-57. Junta de Investigafoes Cientificas

do Ultramar, Lisbon.

HAUMAN, L & WILCZEK, R. 1951. Spermatophytes: Capparidaceae.

Flore dll Congo Beige et dii Ruanda-Unmdi 2: 454— 521 . Institut

National pour I'etude Agronomique du Congo Beige, Brussels.

KJLLICK, D.J.B. 1970. Capparaceae: Maenia. Flora of southern

Africa 13: 159-171. Botanical Research Institute, Pretoria.

MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T.

2002. Atlas of Namibia. David Philip, Cape Town.

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 Affica/Namibia.

OLIVER, D. 1868. Order IX: Capparidaceae. Flora of tropical Africa

1: 73-101. Reeve, Ashford, Kent.

ROESSLER, H. 1966. Capparaceae. Prodromus einer flora von

Sudwestafrika 47: 1-16. Cramer, Lehre.

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

in southern Africa: a review with emphasis on succulents.

Umdaus Press, Hatfield, Pretoria.

WILD, H. 1960. Capparidaceae. Flora zambesiaca 1: 194-245.

Crown Agents for Oversea Governments and Administrations,

London.

W. SWANEPOEL*

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

sity of Pretoria, 0002 Pretoria. Postal address: PO. Box 21168,

Windhoek, Namibia. E-mail: monteiro@iway.com. na

MS. received: 2005-06-03.

DATA DEFICIENT FLAGS FOR USE fN THE RED LIST OF SOUTH AFRICAN PLANTS

The first Red Data List for southern African plants

was published in 1980 (Hall et al. 1980), and was fol-

lowed 16 years later by an update (Hilton-Taylor 1996).

These publications classified plants as Rare, Vulnerable,

Endangered, Indeterminate or Insufficiently Known.

Since then the Species Survival Commission (SSC) of

lUCN — World Conservation Union has introduced a

new system with improved methods of assessing extinc-

tion rates of taxa. This system makes use of prescribed

quantitative criteria to place taxa into different categories

according to their extinction risks (lUCN 1994, 2001).

Many taxa that were previously classified as Rare are

now in the category Least Concern (LC) since they are

not facing increased extinction risk. However, they may

still require conservation attention. For this reason, Victor

& Keith (2004) introduced the Orange List concept and

proposed a quantitative system of assessing, recording

and documenting taxa that should be considered for legal

protection and conservation. The Orange List includes

taxa that are rare but not declining, as well as taxa that are

declining but not fast enough to trigger a threatened list-

ing according to the lUCN Red List Criteria. Two other

categories that are considered under the Orange List are

Data Deficient (DD) and Near Threatened (NT).

According to lUCN (2001), a taxon qualifies for

the category Data Deficient when ‘there is inadequate

information to make a direct, or indirect, assessment

of its risk of extinction based on its distribution and/or

population status’. Although this usually applies to taxa

that are poorly known, this category might also contain

well-known taxa that lack sufficient data required for

using the lUCN Red List Criteria. Whereas DD is not

considered to be one of the categories of threat, listing

of taxa in this category acknowledges the possibility that

future research may show that threatened classification is

appropriate.

During the seven years of compilation of South

Africa’s Red List of threatened plants according to the

revised Red List assessment process, it has become

apparent that there is a need to distinguish between differ-

ent scenarios for listings within the DD category. A set of

flags is proposed to distinguish between the different rea-

sons for listing, with the aim of facilitating conservation

planning and highlighting research needs for the taxa.

Three main reasons for listing taxa in the DD category

are apparent. In the first scenario, taxa have been listed

as Data Deficient but are suspected to have taxonomic

problems (such as being indistinguishable from closely

related taxa) that make it difficult for them to be accu-

rately assessed. Only once these taxonomic problems are

sorted out, can a proper assessment be made of the taxon.

It is proposed that the taxa that are unable to be assessed

due to unclear taxonomic delimitation, or suspected to

be synonymous with other taxa, are listed as DD with a

flag of ‘Taxonomically uncertain’ (abbreviated as DDT).

Because these taxa are often thought to be synonymous

with more widespread taxa, they are usually unlikely

to warrant conservation attention. An example is Erica

obconica, which is probably conspeciflc with the wide-

spread Erica mucronata. However, this has not yet been

formalized so the species is classified as DDT for now.

The second flag deals with taxa that could very well

qualify for a category of threat but have insufficient

information required for the assessment process (such

as distribution or rate of decline). It is proposed that

such taxa are classified as DD with the flag ‘Distribution

and/or other information lacking’ (abbreviated as DDD).

Taxa classified as DDD are likely to be of high conserva-

tion importance and high research priority. An example

is Phylica apiculata, a shrub found on mountain slopes

of the Caledon District. Since much of the natural land in

the Caledon area is transformed, it is likely that this spe-

cies is threatened with extinction. It is therefore classified

as DDD until more information becomes available.

A third flag is proposed for taxa that are so poorly

known that it is impossible to determine whether or not

they could be classified as threatened. Whereas most

DD taxa are suspected to be threatened, some taxa have

so little information that it is not known whether they

are undercollected, rare, taxonomically problematic or

poorly known; but there is no cause to suspect that they

are threatened with extinction. These taxa are represented

by very few collection records in herbaria and have insuf-

ficient information about them in the literature. These

taxa are flagged to indicate that they are of high research

priority, but low conservation priority until such time

more information becomes available. These taxa are

flagged DDX. An example is Anderbergia fallax, which

86

Bothalia 36,1 (2006)

is known from a single collection made from Goedgeloof

Peak in the Langeberg near Swellendam. It is likely to

be undercollected as it is a fairly inconspicuous plant;

furthermore it is unlikely to be threatened on the high

mountain peaks where it grows.

The advantage of flagging subsections of the Data

Deficient category is that conservationists will no longer

have to divest efforts into the taxa in the DDT and DDX

categories. The DDT flag will serve to highlight those

taxa that need taxonomic attention; whereas DDD and

DDX flags would serve to highlight those taxa in need of

more field work and research attention.

REFERENCES

HALL, A.V., DE WINTER, M., DE WINTER, B. & OOSTERHOUT,

S.A.M. 1980. Threatened plants of southern Africa. South

African National Scientific Programmes Report No. 45. CSIR,

Pretoria.

HILTON-TAYLOR, C. 1996. Red Data List of southern African plants.

Strelitzia 4: 1-117. National Botanical Institute, Pretoria.

lUCN 1994. lUCN Red List Categories. Prepared by the lUCN Species

Survival Commission. lUCN, Gland, Switzerland.

lUCN 2001. lUCN Red List Categories and Criteria: Version 3.1.

lUCN Species Survival Commission. lUCN, Gland, Switzerland

and Cambridge, UK. http://www.iucn.org/themes/ssc/redlists/

redlistcatsenglish.pdf

VICTOR, J.E. & KEITH, M. 2004. The Orange List: a safety net for

biodiversity in South Africa. South African Journal of Science

100: 139-141.

J.E. VICTOR*

* Threatened Species Programme, South African National Biodiversity

Institute, Private Bag XlOl, 0001 Pretoria, e-mail: victor@sanbi.org

MS. received: 2005-11-02.

HYACINTHACEAE

ORNITHOGALUM KlRSTENll (ALBUCA GROUP), ANEW SPECIES FROM WESTERN CAPE, SOUTH AFRICA, AND NEW

COMBINATIONS IN THE GROUP

INTRODUCTION

The circumscription of the sub-Saharan genera of

Hyacinthaceae has recently undergone substantial revi-

sion as a result of molecular studies (Manning et al.

2004). One of the more radical changes has been the

inclusion within a widely circumscribed Ornithogalum

L. of all genera previously assigned to the subfamily

Omithogaloideae (Speta 1998), among them the sub-

Saharan African and Arabian species segregated as the

genus Albuca L. Ornithogalum in this broad sense is

defined by its flattened or angular seeds and distinctly

trifid or trifurcate stigma. While sinking these genera into

synonymy in an expanded Ornithogalum has its critics,

the alternative taxonomy consistent with the molecular

phylogeny not only requires the recognition of Albuca,

Dipcadi, Galtonia, Neopatersonia and Pseudogaltonia,

but also requires several (the final number is uncer-

tain) additional genera to accommodate the sub-Saharan

African species currently assigned to Ornithogalum,

with Ornithogalum itself restricted to the Mediterranean

and Near East (Speta 1998). Any other taxonomy would

render Ornithogalum paraphyletic.

The taxonomy of the Albuca alliance remains one of

the least understood in the Hyacinthaceae. One estimate

places the total number of species in the group at ± 60

(Speta 1998) but the most recent listing of the south-

ern African species alone includes 72 current names

(Manning & Goldblatt 2003). Many of these will undoubt-

edly prove to be synonyms. The last complete revision of

the southern African species of Albuca remains that of

Albuca s'ubgenus Falconera, comprising 19 species

from southern Africa, mainly the winter rainfall region of

Western Cape, is distinguished by relatively unspecial-

ized inner tepals lacking hinged or hood-shaped apices

(Miiller-Doblies 1995). The subgenus is further divided

into two sections based on the condition of the anthers

of the inner whorl of stamens. In section Falconera all

six stamens are fertile, whereas in section Trianthera

U.Miill.-Doblies, the stamens of the inner whorl bear

rudimentary anthers. Populations of an unusual autumn-

and early winter-flowering taxon of section Falconera

were recently discovered near Swellendam in Western

Cape by Kirsten Louw, a young Cape Town naturalist.

They represent an unknown species, described here as

Ornithogalum kirstenii, in memory of his tragic and

untimely death in 2005, just weeks after he brought the

species to our attention (Cohen et al. 2005).

Ornithogalum kirstenii J.C. Manning & Goldblatt,

sp. nov.

Plantae deciduae (100-)200-300 mm altae, bulbo

solitario vel fasciculis parvis conico non profunde infosso

vel partim supra terram 15-20(^0) mm diam., tunicis

extemis tenuiter coriaceis griseis, intemis arete imbri-

catis albis vel pallide viridibus ubi expositis, folds 2,

inflorentia leviter brevioribus vel subaequalibus lineari-

convolutis sed ad apicem teretibus 10-20(-30) x 1.5-3. 5

mm succulentibus, in quarta vel tertia parte basali caulem

amplectentibus; inflorescentia racemus laxus erectus vel

inclinatus parum flexuosus ad apicem in alabastro nutans,

(2)3-1 8-florus; bfacteis ovato-acuminatis, 5-7(-10) x

Baker (1897) but the revisions of subgenus Albuca and^®^2-3 mm viridibus initio demum pallide brunneismargini-

subgenus Falconera (Salisb.) Baker by Miiller-Doblies

(1994, 1995), albeit incomplete, represent a substantial

advance in our understanding of the genus and provide a

basis from which to assess and identify about half of the

southern African species currently known. One of their

most significant contributions is the proposed division of

the species in this alliance into four infrageneric group-

ings (subgenera).

bus latis pellucidis, pedicellis patentibus anthesis initio

10-15(-20) mm longis, suberectis ad erectis ubi fructi-

cantibus ad finem 30-50 mm longis; floribus nutantibus

flavis carinis viridibus leviter vanillariodoris, tepalis

biseriatis laminis extemis connatis ad ± 1 mm patentibus

oblongo-oblanceolatis 13-15 x 4. 0^.5 mm ad apicem

papillosis, intemis suberectis leviter divergentibus ubi

apertis oblanceolatis concavis 13-14 x 4. 5-5. 5 mm ad

Bothalia36,l (2006)

87

lo apicem pauciter tumidis succulentibusque in pagina inte-

riore fascia longitudinali papillarum, staminibus parum

dimorphis, extemis subteretibus decrescentibus infra

leviter expansis canaliculatisque ± 1 1 mm longis, intemis

lateraliter expansis; antheris erectis ± 3 mm longis, ovario

oblongo ± 4 mm longo cristis paraseptalibus obscuris,

stylo columnar! trigono transverse ruguloso ± 8 mm

longo, stigmate obtuso-trigono papilloso.

TYPE. — Western Cape, 3420 (Bredasdorp); 2 km

WSW of Malgas, Farm Malgaskraal, W bank of gulley

leading into Breede River opposite Perdekloof, (-BC),

17 April 2005, J. Manning 2942 (NBG, holo.; K, MO,

iso.).

Deciduous geophyte, (100-)200-300 mm high. Bulb

solitary or in small clusters, conical, shallowly buried

or partially epigeal, 15-20(^0) mm diam.; outer tunics

thinly leathery, pale grey; inner tunics tightly overlap-

ping, white but pale green when exposed. Leaves 2, erect,

slightly shorter than or subequal to inflorescence, fleshy

but firm-textured, linear-convolute for most of length

but terete in apical 5-15 mm, 100-200(-300) x 1.5-3. 5

mm, dull green, clasping stem in basal fourth or third.

Inflorescence an erect or inclined, lax raceme, weakly

flexuose, (10-)20-30 mm long, apex nodding in bud,

(2)3-1 8-flowered; bracts ovate-acuminate, 5-7(-10) x

2-3 mm, green when young but soon drying pale brown,

with broad transparent margins; pedicels spreading, nod-

ding at tip at anthesis, 10-15(-20) mm long, becoming

subsecund in fruit and suberect to erect, lengthening and

ultimately 30-50 mm long. Flowers pendulous, canary

yellow with green tepal keels, faintly vanilla-scented;

tepals biseriate with blades of outer series overlapping

inner, joined at base for ± 1 mm, outer tepals spread-

ing, oblong-oblanceolate, 13-15 x 4. 0^.5 mm, apices

papillate, inner tepals suberect, weakly diverging when

fully open, oblanceolate, concave, 13-14 x 4. 5-5. 5 mm,

apices slightly swollen and fleshy with longitudinal band

of papillae on inner face. Stamens adnate to perianth for ±

1 mm, weakly dimorphic, outer three erect around style,

inner three suberect and lying against iimer tepals; outer

filaments subterete and tapering, slightly widened and

channelled in basal third, ± 11 mm long, inner laterally

expanded and pinched in lower fifth; anthers erect, ± 3

mm long, cream-coloured. Ovary oblong, slightly nar-

rowed at waist, green, ± 4 mm long, paraseptal ridges

obscure; style columnar, trigonous, transversely rugulose,

yellow, ± 8 mm long, obtuse with trigonous, papillate

stigma. Capsule narrowly ovoid to flask-shaped, 3-

angled, 15-16 x 5. 5-6. 5 mm. angular-pyriform or

D-shaped with flattened sides, minutely papillose, dull

black, 2. 0-2. 5 x 1 mm. Flowering time: April to June.

Figure 18.

tion on the cliffs is dominated by succulents, especially

various Crassulaceae; Crassula rupestris, Cotyledon

orbiculata and an Adromischus species. O. kirstenii is

restricted to this particular habitat and is not found on the

adjacent, more gentle, stony or gravelly slopes that sup-

port renosterveld shrubland.

Diagnosis and relationships', among the species of

Ornithogalum previously segregated as the genus Albuca,

the pendulous flowers with weakly cucullate inner tepals

(not apically hinged nor hooded) and six fertile stamens,

place O. kirstenii in section Falconera of subgenus

Falconera as defined by Miiller-Doblies & Miiller-

Doblies (1995). Of the fourteen species recognized in

this section by Muller-Doblies & Muller-Doblies, just six

lack glandular hairs. Among these, O. clanwilliamaeglo-

ria (U.Mull.-Doblies) J.C. Manning & Goldblatt and O.

fragrans (Jacq.) J.C. Manning & Goldblatt from Western

Cape are distinguished by their smooth styles, O. bifolia-

tum (R.A.Dyer) J.C. Manning & Goldblatt from Eastern

Cape by its unique, 3-homed stigma, and O. angolense

J. C. Manning & Goldblatt (= Albuca monophylla Baker)

from Angola and Namibia by the single leaf and well-

developed, diverging paraseptal ridges on the ovary. This

leaves just two species, O. hesquaspoortense (U.Miill.-

Doblies) J.C. Manning & Goldblatt and O. robertsoni-

anum (U.Mull.-Doblies) J.C. Manning & Goldblatt, with

which O. kirstenii may be confused. Both occur in the

Breede River Valley in the general vicinity where O.

kirstenii has been collected, and both have few or just

two leaves, and weakly developed or obsolete paraseptal

ridges. These features are all characteristics of O. kirst-

enii but the absence of the types of O. hesquaspoortense

and O. robertsonianum from the herbaria in which they

are purported to have been deposited, makes direct com-

parison with them impossible. The type material of both

of these species was, however, apparently collected in

the spring, between August and October, rather than in

the late autumn or early winter, when O. kirstenii flow-

ers. In addition, O. hesquaspoortense is distinguished in

the protologue from other species by the basally scabrid

scape, and O. robertsonianum by its thickly fibrous, outer

bulb tunics. The scape in O. kirstenii, like the leaves, is

completely glabrous and the outer bulb tunics are thinly

leathery and not at all fibrous. On the available morpho-

logical and phonological evidence, therefore, it appears

that O. kirstenii warrants recognition as a species, at least

until more is known about O. hesquaspoortense and O.

robertsonianum.

Other specimens examined

WESTERN CAPE. — 3420 (Bredasdorp): 23.5 km along road to De

Hoop, shale cliff 300 m S of road, (-AD), 16 May 2005, C. Cohen s.n.

(NBG); 2 km WSW of Malgas, Farm Malgaskraal, W bank of gulley

leading into Breede River opposite Perdekloof, (-BC), 10 April 2005,

K. Louw s.n. (NBG),

Among the extensive list of new names and combina-

tions in Ornithogalum that accompanied the synonymy of

Albuca (Manning et al. 2004), are several that are inad-

vertently preoccupied. New names for the taxa concerned

are provided here.

Ornithogalum glutinosum J.C. Manning &

Goldblatt, nom. nov., pro O. hallii (U.Mull.-Doblies)

Distribution and ecology, known from several popula-

tions along the lower reaches of the Breede River Valley

and its tributaries near Malgas and De Hoop, south

of Swellendam (Figure 19), Ornithogalum kirstenii is

unique among related species in flowering in the late

autumn and early winter. Populations of the species occur

on both sides of the Breede River (K. Louw pers. comm.),

growing on shale cliffs along gulleys and rivers with the

bulbs partially exposed or shallowly buried in the decay-

ing rock on the exposed banks and cuttings. The vegeta-

Bothalia 36,1 (2006)

FIGURE 18. — Ornilhogabm kirst-

enii. Manning 2942 (NBG).

A, whole plant; B, t/s leaf;

C, flower; D, outer tepal; E,

apex of outer tepal; F, inner

tepal; G, apex of inner tepal;

H, outer stamen; 1, inner sta-

men; J, gynoecium and two

stamens; K, gynoecium with

t/s style; L, capsule; M, seeds.

Scale bars: A, L, 10 mm;

B-D, F, H-K, M, 5 mm;

E, G, 2.5 mm. Artist: John

Manning.

J.C. Manning & Goldblatt in J.C. Manning et al. in

Edinburgh Journal of Botany 60: 548 (2004), non O. hal-

///Oberm. (1978).

Ornithogalum melleri Baker in Journal of the

Linnean Society, Botany 13: 280 (1873).

Ornithogalum abyssinicum (Jacq.) J.C. Manning & Goldblatt in

J.C. Manning et al. in Edinburgh Journal of Botany 60: 546 (2004), syn.

nov., non O. abyssinicum Fresen. (1835).

Ornithogalum neopatersonia J.C. Manning &

Goldblatt, nom. nov., pro O. uitenhagense (Schonland)

J.C. Manning & Goldblatt in J.C. Manning et al. in

Edinburgh Journal of Botany 60: 553 (2004), non O.

uitenhagense Poelln. (1944).

Ornithogalum soleae J.C. Manning & Goldblatt,

nom. nov., pro O. diphyllum J.C. Manning & Goldblatt

in J.C. Manning et al. in Edinburgh Journal of Botany 60:

548 (2004), non O. diphyllum Baker (1895).

Ornithogalum \o\ui2a!:e J.C. Manning & Goldblatt,

nom. nov., pro O. circinatum J.C. Manning & Goldblatt

in J.C. Manning et al. in Edinburgh Journal of Botany 60:

547 (2004), non O. circinatum L.f (1781).

ACKNOWLEDGEMENTS

We are indebted to Kirsten Louw for drawing our

attention to this species and to Elizabeth Parker for her

assistance in collecting the type material, which was

gathered under a permit from Western Cape Nature

Conservation.

Bothalia 36,1 (2006)

89

FIGURE 19. — Distribution of Omithogalum kirstenii in Western

Cape.

REFERENCES

BAKER, J.G. 1895. Diagnoses africanae. Kew Bulletin 1895: 141-

153.

BAKER, J.G. 1897. Albuca. In W.T. Thistelton-Dyer, Flora capensis 6:

451^62. Reeve, Ashford.

COHEN, C., SPOTTISWOODE, C., MILLS, M. & MURISON, G.

2005. Kirsten Louw. Promerops 263: 4, 5.

FRESENIUS, G. 1835. Semina horto botanico Franco furtensi anno

1834 collecta. IDC Microfiche 7835.

LINNAEUS, C. fil. 1782 (1781). Supplementum plantanm. Impensis

Orphanotrophei, Braunschweig.

MANNING, J.C. & GOLDBLATT, P. 2003. Hyacinthaceae. In G.

Germishuizen & N.L. Meyer, Plants of southern Africa: an aruio-

tated checklist. Stretozia 14: 1054—1071.

MANNING, J.C., GOLDBLATT, P. & FAY, M. 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.

MULLER-DOBLIES, U. 1994. EnumeratioAlbucarum (Hyacinthaceae)

Austro-Africanarum adhuc cognitarum 1. Subgenus Albuca.

Feddes Repertorium 105: 365-368.

MULLER-D0BLIES,U. 1995. EnumeratioAlbucarum (Hyacinthaceae)

Austro-Africanarum adhuc cognitarum 2. Subgenus Falconera

(Salisb.) Baker emend. U.M-D. 1987. Feddes Repertorium 106:

353-370.

OBERMEYER, A. A. 1978. Omithogalum: a revision of the southern

African species. Bothalia 12: 323-376.

SPETA, F. 1998. Hyacinthaceae. In K. Kubitzki, The families and gen-

era of vascular plants. Springer, Berlin.

VON POELLNITZ, K. 1944. Neue afrikanische Pflanzen. Berichte der

deutschen botanischen Gesellschaft 61: 204-209.

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

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

** B.A. Kiirkoff Curator of African Botany, Missouri Botanical Garden,

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

MS. received: 2005-05-04.

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Bothalia36,l: 91-99 (2006)

Reappraisal and identiHcation of Olinia rochetiana (Oliniaceae) in

South Africa

R.J. SEBOLA*t and K. BALKWILL*

Keywords: habit, hypanthium, morphology, Olinia rochetiana A.Juss., phenetic, population. South Africa, taxonomy, variation

ABSTRACT

A numerical phenetic analysis of data obtained from populations of the Olinia rochetiana A.Juss. complex occurring in

South Africa (Mpumalanga and Limpopo Provinces) revealed the existence of two forms: 1 , a shrubby form (up to 2.5 m tall),

with thick terminal branches, coriaceous leaves with a tinge of red on margins (towards the apices), short inflorescence axes,

peduncles and deeply red pedicels and floral tubes/hypanthia; and 2, a slender tree form, measuring more than 4 m tall with

slender terminal branches, glossy and slightly thin, papery leaves, margin colour the same as the entire lamina, and the inflor-

escence axes, peduncles, pedicels and hypanthia pale green to creamy white. Differences in floral features between the two

forms correlate with differences observed in vegetative features. The two forms occupy distinct ecological niches and show

tolerances and preferences for different environmental conditions such as soil type, elevation and humidity. An identification

key for the two forms is presented.

INTRODUCTION

Olinia rochetiana A.Juss. sensu lato is a morphologi-

cally variable and widespread forest species occurring

on foothills and mountain ravines in Angola, Zambia,

Zimbabwe, South Africa, Mozambique, Malawi, Tanzania,

Kenya, Rwanda, Burundi, Democratic Republic of the

Congo, Uganda, Sudan and Ethiopia. The plants prefer

moist habitats at the foot of mountains, often along

ravines. In this kind of habitat plants often reach quite

considerable heights (± 4-6 m), whereas in exposed areas

the plants are associated with rocky outcrops and quartz-

itic soils, and mostly measure just less than 3 m tall. The

description of O. rochetiana by Jussieu (1846) was based

on Rochet d’Hericourt 18, collected during his voyage in

Abyssinia (Ethiopia). The current taxonomic concept of

O. rochetiana s.l. includes O. aequipetala Gilg, O. dis-

color Mildbr., O. huillensis Welw., O. ruandensis Gilg,

O. usambarensis Gilg and O. volkensii Engl., all of which

Verdcourt (1975, 1978) and Verdcourt & Fernandes

(1986) treated as conspecific in their regional treatments

of Oliniaceae. Commenting on the nature of the morpho-

logical variation, Verdcourt (1975, 1978) remarked that

South Afncan populations of Olinia from the northern

and eastern parts of the former Transvaal (now split into

four provinces: Gauteng, Limpopo, Mpumalanga and

some eastern parts of North-West) with short, broad pet-

als could be recognized as a variety of O. rochetiana.

Since then material collected from the escarpment in

Mpumalanga and Limpopo has been referred to O.

rochetiana, despite Burtt Davy’s (1926) recognition of

the plants collected from the mist belt forests in Graskop

and Pilgrim’s Rest, Mpumalanga, as O. usambarensis.

Gilg ( 1 895) described this latter species based on Holst

9115, collected from the elevated forests of Usambara in

Tanzania. Cufodontis (1960) and Fernandes & Fernandes

* C.E. Moss Herbarium, School of Animal, Plant and Environmental

Sciences, Private Bag 3, University of the Witwatersrand, 2050 WITS,

Johannesburg.

t Corresponding author’s current address: Senior Manager, Scientific

Research and Development, Department of Agriculture, P.O. Box

3748, 0001 Pretoria. E-mail: Ramagwai@biology.biol. wits. ac.za or

SMSRD@nda.agric.za

MS. received: 2005-03-31.

(1962) recognized O. usambarensis, but their concepts

did not include the South African populations, which

represent the most southern limits of the present distribu-

tion of the O. rochetiana complex in Africa. Thus, the

confusion between O. rochetiana and O. usambarensis

has persisted in South Africa. Information gleaned from

herbarium sheets of material collected from Usambara,

Tanzania, consistently indicates O. usambarensis to be

a medium to large tree with large and broadly elliptic

leaves, and sparsely pubescent hypanthia, as opposed

to the shrubby form with shorter obovate leaves, robust

and glabrous hypanthia that occurs in misty heights and

windswept outcrops in Graskop and within the Pilgrim’s

Rest area in Mpumalanga.

During October/November to March/ April it is possi-

ble to observe significant morphological variation in flo-

ral and fruit characteristics among populations of Olinia

rochetiana occurring in Mpumalanga and Limpopo. In

undertaking field studies on populations of O. rochetiana

in South Africa, as part of an Africa-wide revision of this

species complex, it became apparent that what is cur-

rently referred to O. rochetiana in South Africa shows

considerable discontinuities in morphological variation

between populations. It was therefore decided to study

and analyse the morphological variation within and

between populations of O. rochetiana occurring in South

Africa, and determine whether the variation is sufficient-

ly discrete to justify splitting the populations into species

or infraspecific taxa. Population level data are often pre-

ferred over other kinds of data, and have the potential to

provide useful information on the extent and distribution

of variation within and between sympatric taxa (Balfour

& Linder 1990; Wiltshire et al. 1991; Astholm & Nyman

1994; Chandler & Crisp 1998; Hong et al. 1998). Most

populations of O. rochetiana in South Africa are allopat-

ric, separated by mountain blocks of varying elevation/

altitudes, rainfall and different soil formations. However,

in the Pilgrim’s Rest area, some of the populations occur

in close proximity to each other, suggesting that the

observed morphological variation may have a genetic

basis and that the populations are maintained possibly by

some sexual reproductive system.

92

Bothalia 36,1 (2006)

In this paper, only results from studies on morphologi-

cal variation among populations of Olinia rochetiana in

South Africa are presented. The study focused on com-

paring morphological variation among populations of

the O. rochetiana complex occurring in Limpopo and

Mpumalanga in order to examine to what extent the pat-

terns of morphological variation between populations

can be related to environmental factors. A comprehensive

taxonomic treatment of the O. rochetiana species com-

plex over its entire range of distribution is currently being

prepared, and will be published shortly.

MATERIAL AND METHODS

Morphological variation within and between popula-

tions was assessed during their flowering (December/

January) and fruiting periods (March/ April) from 1997

to 2001. A total of fourteen populations of the Olinia

rochetiana complex occurring in South Africa were

sampled (Table 1). In addition, six populations belong-

ing to O. emarginata Burtt Davy were also sampled and

included in the analyses as a standard taxon to aid in an

objective evaluation and interpretation of the variability

within and between the populations. O. emarginata is a

clearly defined species in the genus, and is endemic to

but widely distributed in South Africa. A Geographic

Positioning System (GPS) and the 1 : 50 000 topographi-

cal maps (published by the Chief Directorate, Surveys

and Maps, Cape Town) were used to determine altitude

as well as the geographic co-ordinates. Some characters

were studied in the field (those pertaining to colour, size

and shape of floral features as well as growth form char-

acteristics) and others in the laboratory, using the Zeiss

dissecting microscope. Measurements on vegetative and

floral parts were made to the nearest 0.5 mm. A minimum

of five measurements were made for all the quantitative

characters per specimen and averaged. In total, 35 charac-

ters, 16 of which are quantitative continuous (obtained by

measurements), three qualitative discontinuous (obtained

by counting) and 16 qualitative diseontinuous (obtained

by scoring each specimen into states) were measured per

specimen (Table 2) and entered into a t x « (taxon x char-

acter) data matrix. The character states of the variation in

the measurements or observations made for the characters

were discrete and assumed to be under unique genetic

control (Stevens 1991; Gift & Stevens 1997). Voucher

specimens were prepared and accessioned into J.

The data matrices were analysed using the eluster-

ing and ordination algorithms of NTSYS-pc (Numerical

Taxonomy and Multivariate Analysis System) version 2.0

(Rohlf 1998). Firstly, the analyses were performed using

individuals within populations as operational taxonomic

units (OTUs). Secondly, the means of each character for

all individuals within each population were averaged

and then used in subsequent phenetic analyses with the

populations as OTUs. As a recommended procedure in

phenetic analyses (Sneath 1976; James & McCulloch

1990; Crisp & Weston 1993), the data matrices were

first standardized using the ‘STAND’ algorithm to render

characters dimensionless and to reduce all characters to a

scale of comparable range so that each character contrib-

utes towards the overall resemblance in proportion to its

variability among the set of OTUs.

The data matriees were subjected to an ordination

procedure by means of Principal Coordinate Analysis

(PCoA), which is preferable and more reliable over the

Principal Component Analysis (PCA) when analysing

mixed data of both qualitative and quantitative characters

(Austin 1985; Kent & Coker 1992) as was the case in

this study (Table 2). PCoA was performed on the correla-

tion matrix obtained from the standardized data matrix

by using the procedure SIMINT, DCENTER, EIGEN,

and MOD3D available in the NTSYS-pc package (Rohlf

1998).

TABLE 1 . — Descriptions of localities and populations of plants of Olinia emarginata and O. rochetiana complex in South Africa

'A DS, quarter degree square; No., number plants sampled (either flowering or fruiting) in a population; Tot. no., total number plants observed in a

population; Height (m), mean height of plants in a population and ±, standard deviation.

Bothalia 36,1 (2006)

93

TABLE 2. — Descriptions of characters used in phenetic analyses of O.

rochetiana complex and O. emarginata populations. * Denotes

characters defined as in Hickey (1973), Hill (1980) and Herman

etal. (1987)

1 . Mean lamina length (mm).

2. Mean lamina width (mm).

3. Mean petiole length (mm).

4. Mean inflorescence axis length (mm).

5. Mean hypanthium length (mm).

6. Mean petal length (mm).

7. Mean petal width (mm).

8. Mean pedicel length (mm).

9. Lamina obovate, broadly elliptic or narrowly elliptic.

10. Apex emarginate, or acuminate.

1 1 . Base cuneate to round, or decurrent.

12. Lamina midrib channeled above, or not channeled above.

13. Secondary veins loop once, or more than once before margins.

14. Secondary veins branch at 30-45° or at more than 45° from midrib.

15. Lamina paler above, or below.

16. Margins on apex reddish, or same color as lamina.

17. Midrib pinkish to red, or pale green.

18. Terminal branches pink to reddish, or pale green.

19. *Leaf Shape Index (200MAV, where M = mean perpendicular

distance from the midrib to the margin, and W = maximum width

of the leaf).

20. *Mean number of secondary veins, counted on both sides of midrib

on adaxial surface.

2 1 . *Mean secondary vein branching angle (°) on adaxial surface.

22. *Mean areole length (mm) measured on adaxial surface.

23. *Mean areole width (mm) measured on adaxial surface.

24. *Mean number of veinlets per areole counted on adaxial surface.

25. *Mean number of branches per veinlet expressed as branching

orders.

26. *Leaf apex angle (°) measured on adaxial surface.

27. *Leaf base angle (°) measured on adaxial surface.

28. Leaf lamina coriaceous /leathery, or soft to slightly papery.

29. Mean hypanthium width (mm) i.e. narrow (2-3 mm) or broad (> 3

mm)

30. Hypanthium deep red, or pale green to cream-white, turning pink

with age.

3 1 . Petal pink to deep red, or cream-white.

32. Petal oblong to spathulate, or obovate.

33. Petal apex distinctly mucronate, or without a mucro.

34. Mean width of inflorescence axis (mm).

35. Habit tree form or shrubby.

Cluster analysis was used to test whether groups simi-

lar to those obtained in the ordination analyses could be

recovered, and also to visualize the level of morphologi-

cal similarity/dissimilarity using appropriate coefficients

between and within populations. Only those characters

that were effective in discriminating between populations

(as judged by high eigenvalues i.e. > 0.6) in the first three

axes of ordination analyses were used in cluster analyses.

This approach was followed since cluster analysis is

known to ‘impose’ a hierarchical structure on any data

set (Thorpe 1983), and often shows clusters that may not

be recoverable in ordination analyses (Chandler & Crisp

1998). Clustering was performed using the Euclidean dis-

tances among means through the Unweighted Pair Group

Method of Arithmetic Averages (UPGMA).

RESULTS

The PCoA of the data in which individual specimens

were used as OTUs, separated specimens of Olinia roche-

tiana from those of O. emarginata along the first axis

(Figure 1). Specimens belonging to O. emarginata are

grouped neatly into one unit on the left side of the plot,

whereas specimens of O. rochetiana occupy the right side

of the plot. Characters most strongly correlated with the

first PCoA axis were, in decreasing order, leaf dimen-

sions (leaf width, leaf base angle, leaf apex angle, number

of veins looping before margins, angle of branching of

veins from the midrib, and petiole length), petal length

and shape. Two groups of specimens belonging to O.

rochetiana are clearly discernible along the second PCoA

axis. Specimens belonging to O. emarginata are slightly

intermediate between the two subgroups of O. rochetiana

along the second PCoA axis, but overlap with the slender

tree form populations of O. rochetiana. To the right side

of specimens of O. emarginata at the top comer is a group

comprising populations of the O. rochetiana complex

from Lost (Tity, The Pinnacle, God’s Window, Quartz

Hill and Mariepskop, all in Mpumalanga, whereas at the

bottom comer is a group of specimens representing the

O. rochetiana complex populations from the Blouberg,

Soutpansberg and the Wolkberg in Limpopo as well as

Blyde and Themeda Hill in Mpumalanga. Characters

most strongly correlated with the second PCoA axis were,

in decreasing order, the habit, leaf texture, size and colour

of the hypanthium, petals and terminal branches. The

close phenetic similarity of members of the two groups

of the O. rochetiana complex to each other and their dis-

similarity to the O. emarginata group can be visualized

easily along the first axis. This indicates lower levels of

variation within the OTUs of the same populations than

between the OTUs of the two groups of the O. roche-

tiana populations. The large gaps and sharp differences

between the clusters along the phenetic spaces, as well

as the high density or compactness within the clusters

(Figure 1) suggest a predominance of discontinuous-state

quantitative characters over overlapping continuous-state

characters in the data set, thus casting aspersion on the

realness of the gaps or discontinuities and how accurately

they reflect the morphological differences between the

plants studied (Ratliff & Pieper 1981; Stevens 1991; Gift

& Stevens 1997). However, the data analysed (Table 2)

is a balanced mixture of both quantitative continuous and

qualitative discontinuous characters.

The UPGMA cluster analysis using only the fourteen

characters with high eigenvalues (> 0.6) in the first two

PCoA axes, also separates three groups (Figure 2) cor-

responding to those obtained in the PCoA analysis. The

Olinia emarginata specimens form a distinct uniform

cluster separate from the O. rochetiana specimens at the

dissimilarity levels indicated by the two phenon lines (a

and b). The clustering of individual specimens belong-

ing to O. emarginata did not reflect their representative

populations (seven in total). Instead, the specimens were

mixed among each other, indicating a low level of vari-

ability among the sampled populations of O. emarginata.

The O. rochetiana specimens are split into two major

subclusters at the 1.33 dissimilarity level (phenon line a

in Figure 2) for which the specimens of O. emarginata

remain coherent in a single cluster.

The means of each character for all individuals within

each population were averaged and cluster analysis per-

formed with the populations used as OTUs. The results

(Figure 3) revealed similar groupings and associations

between populations as in Figures 1 and 2. The popula-

tions of Olinia emarginata form a distinct cluster a sepa-

rate from the populations of the O. rochetiana complex.

94

Bothalia36,l (2006)

FIGURE 1 . — Plot of first two principal co-ordinate analyses obtained from analysing morphological data from populations of Olinia rochetiana

complex and O. emarginata. First and second PCoA axes explain 47.3% and 10.4% of total variation respectively.

which are split into clusters b and c (Figure 3). The sepa-

ration of populations belonging to O. emarginata (cluster

a) from those populations (clusters b and c) belonging

to the O. rochetiana complex, occurs at a high level of

dissimilarity (i.e. 1.83), indicating that these clusters rep-

resent morphologically dissimilar entities.

Cluster b comprises populations of Olinia roche-

tiana characterized by a slender, tree-like habit; slen-

der terminal branches and branchlets; thin, glossy and

papery leaves; slender inflorescence axes, peduncles,

pedicels and hypanthia which are pale green to creamy

white and turning slightly pink when mature. These are

plants from the Soutpansberg (on the Farms Letjuma,

Llewelyn, Ontmoet and Sussens), Tate Vondo in Venda,

the Blouberg, the Wolkberg in Limpopo, and at Blyde

(Potholes) and Themeda Hill in the Pilgrim’s Rest

area in Mpumalanga (Figure 4). With the exception of

populations from Tate Vondo in Venda, which occur in

well-shaded, less exposed areas and reaching up to 5 m

high, all others occur in fairly exposed environments and

at most reach 4 m high. Cluster c comprises plants col-

lected exclusively from Mpumalanga at Lost City, The

Pinnacle, God’s Window, Quartz Hill and Mariepskop

(Figure 5). These plants are morphologically distinct

from those in cluster b in their shrubby, often multi-

stemmed fonn; coriaceous leaves with a tinge of red on

margins; and strikingly short intlorescence units relative

to inflorescence axes, and hypanthia and petals which are

deep red.

The distinctions among the populations are further

illustrated by the box plots using selected characters con-

tributing most to the discriminations among populations

in the PCoA. The box plots (Figure 6A-D) illustrate vari-

ations among populations for leaf width, petiole length,

hypanthium length and petal width. These characters,

including habit and flower colour, are used in the key.

DISCUSSION

The phenetic analysis, using both cluster and principal

co-ordinate analyses, of the specimens of O. rochetiana

in South Africa indicate that there are two major groups

of populations which are identifiable largely by their

habit, leaf texture and colour of inflorescence units,

hypanthium and sepals. This partly supports the observa-

tions that size and shape of floral features are taxonomi-

cally significant in Olinia (Sebola & Balkwill 1999).

Data on floral features indicate that the two major groups

of populations can be reliably distinguished from one

another on the basis of petiole length, petal length, petal

width, and the floral colour.

The conspicuous red flowers in the shrubby fonn pro-

duce less or no scent compared to the creamy white flow-

ers observed in populations with a slender, tree-like habit.

The hypanthia (floral tubes) in the shrubby form, were

consistently damaged at the base (i.e. at the attachment

to the ovary), and their petals often chewed by crawling

insects. Both young and old flowers among the shrubby

Bothalia 36,1 (2006)

95

9

I

5

FIGURE 2. — Phenogram, based on analysis of only morphological characters with high eigenvalues (>0.6) in PCoA analysis, using individual

specimens from populations of Olinia rochetiana complex and O. emarginata as OTUs; cophenetic correlation (r) = 0.979. Vertical lines

indicate phenon lines a and b.

96

Bothalia 36,1 (2006)

Lost City

Pinnacle i

Mariepskop

Quartz Hill '

God’s Window-

Potholes -

Letjuma”

Ontmoet-

Themeda Hill “

Blouberg "

Llewelyn"

Wolkberg -

Sussens-

Tate Vondo-

Uitkijk

Naaupoort

Witpoortjie

Melville Koppies

Hekpoort

Haenerstburg

0.05

0.50

1 I I I I I

0.94

Euclidean distance

1.39

1.83

FIGURE 3. — Phenogram, based on analysis of only morphological characters with high eigenvalues (> 0.6) from PCoA analysis, using populations

as OTUs; cophenetic correlation (r) = 0.993. Letters a, b and c indicate distinct clusters corresponding to O. emarginata, slender, tree-like

form populations of O. rochetiana and shrubby dwarf form populations of O. mchetiana respectively.

FIGURE 4. — Olinia rochetiana in South Africa. Known geographical

distribution of slender, tree-like form populations, •.

fonn populations showed signs of damage. The creamy

white to pink llowers among the slender tree populations

produce sweet scent (observed from morning to mid-

day), and bees were observed visiting. No damage was

observed on the young and mature flowers. Once pollina-

tion has been effected, the hypanthia start wilting, and the

signs of ageing appear first on the white petals, which turn

brown. In all the populations studied, the flowers appear

superficially simple, but closer examination revealed

the existence of some structural differences between the

two forms of populations. The deep red hypanthia in the

FIGURE 5. — Olinia rochetiana in South Africa. Known geographical

distribution of shrubby dwarf form populations, •.

shrubby form are robust, strong, hard and short relative

to the petal lengths, and have a wider diameter allowing

insects as large as 5 mm access into the tube, whereas

among the slender tree populations, the creamy white to

pink hypanthia are slightly narrower, weak, twice as long

as the petals and the tube hardly opens until anthesis,

soon after which petals start wilting. The effectiveness of

this barrier to insects would be enhanced by the occur-

rence of some recognition and rejection system to avoid

hybridization among populations given their overlap

in flowering periods. Currently, there are no records of

Bothalia36,l (2006)

97

LC PN GWQH MK LJ LWOM TV PH TH BK BB SS UK NP MV HB HP

FIGURE 6. — Box and whisker plots showing variation between populations for A, leaf width; B, petiole length; C, hypanthium length; and D,

petal width. LC, Lost City; PN, The Pinnacle; GW, God’s Window; QH, Quartz Flill; MK, Mariepskop; LJ, Letjuma; LW, Llewelyn; OM,

Ontmoet; TV, Tate Vondo; PH, Potholes; TH, Themeda Hill; BK. Wolkberg; BB, Blouberg; SS, Sussens; UK, Uitkijk; NP, Naaupoort; MV,

Melville Koppies; HB, Haenertsburg; HP, Hekpoort.

sympatric populations for the two forms in South Africa.

The nature of the breeding systems among populations

of the O. rochetiana complex, and other species within

Olinia needs to be determined to provide clarity on exist-

ing isolation mechanisms among taxa.

Both the PCoA and the box and whisker plots indicate

that characters that contribute to this differentiation vary

significantly among populations and are from diverse

plant structures (habit, vegetative and reproductive struc-

tures). Leaf texture consistently distinguishes the shrubby

plants with coriaceous and/or leathery leaves from the

slender, tree-like plants with membranous and/or papery

leaves. The shrubby form populations occur in dry condi-

tions, and are found consistently in exposed, windswept

sites on rocky outcrops, whereas the slender tree forms

occur in moist conditions and in shaded, less exposed

areas or near ravines and streams. Plant height is a

vegetative feature known to show variation in response

to micro-environmental differences, and was thus not

included in the phenetic analyses. Of the slender tree-like

group of populations, only those from Tate Vondo grow-

ing in semi-shaded areas were between 5 and 6 m high,

whereas other populations measured between 3 and 4.5

m high. A possible explanation for the plants reaching up

to 6 m high could be due to phototropism as a result of

limited direct sunlight in shaded areas. There was a cor-

relation between general plant height, habit and altitude.

The shrubby form populations characteristically meas-

ured well below 2.5 m high and all occurred at elevations

between ± 1 547 m and ± 1 980 m above sea level, where-

as the slender tree form populations measured above 3.5

m high and occurred at altitudes below 1 500 m above sea

level. Thus, a trend exists for decreasing plant height at

high altitude for the two forms of populations.

The morphological differentiation of the two forms of

Olinia rochetiana in South Africa seems to reflect adap-

tations to the micro-climates and ecological conditions

in which they occur. Some general habitat characteristics

and ecological features such as rainfall, temperature and

soil types seem to influence the distributions and occur-

rence of the two forms of O. rochetiana populations in

South Africa. Of the slender, tree-like populations, those

on the Blouberg and Wolkberg are geographically isolated

from those on the Soutpansberg, which have intermedi-

ate populations between them along the southern slopes.

Materials collected from Leolo Mountains and the rocky

outcrops around the Steelpoort area in Sekhukhuneland

have particularly glaucous branchlets and young leaves,

and are often confused with O. emarginata (M. Lotter

pers. comm.). In their descriptions of Oliniaceae, Schmidt

et al. (2002) made reference to a unique form of

O. rochetiana populations from Sekhukhuneland with

young twigs densely hairy. However, the phenetic analy-

ses in this study indicate a strong phenetic similarity

between populations from Sekhukhuneland with those

from the Blouberg, Wolkberg, Leolo Mountains and the

Soutpansberg than they are to those from The Pirmacle

(Graskop) and Lost City (Pilgrim’s Rest). The larger size

of leaves (5-8 mm long) for O. rochetiana referred to

by Schimdt et al. (2002) appears dubious, and extremely

98

Bothalia 36,1 (2006)

small compared to O. rochetiana plants in Mpumalanga

(26-48 mm long) and those from Letjuma (Soutpansberg)

and Tate Vondo (Thohoyandou, Venda), which range

from 34.5-65.0 mm long.

The causes of the morphological variation between the

two forms of populations remain unclear. An explanation

of the apparent morphological variation on the basis of

phenotypic plasticity and considering the varying micro-

climates and the environmental differences, appears

inadequate to account for the variation in reproductive

structures (size, shape and colour) which are generally

understood to be less phenotypically plastic than veg-

etative features (Davies 1983). Variation in reproductive

structures is often associated with recognition/rejection

mechanisms and successful pollination (Oliveira 1998;

Prance 1998). Any hypothesis regarding the causes for

the morphological variation among the Olinia rochetiana

populations in South Africa, would be enriched by an

investigation of the mating patterns and reproductive

traits to provide insights into gene flow, taking into con-

sideration the geographic distances and plant-pollinator

interactions and relationships.

Our phenetic analyses indicate that at least two taxa

can be recognized among the populations of Olinia

rochetiana in South Africa. Formal taxonomic status

for these groups of populations will be made in a com-

prehensive study of the morphological variation within

O. rochetiana s.l. over its known range of distribution

(Sebola & Balkwill in prep.). This study will indicate if

any of the two forms of populations overlap with other

taxa that might be delimited within the O. rochetiana

complex. However, the two forms of populations in

South Africa can be identified as follows:

Shrub, up to 2.5 m tall, semi-deciduous; intemodes of terminal

branches shorter than 45 mm, thick and compact, deep red;

leaves coriaceous; inflorescence axes and peduncles 6-10 mm

long, thick; pedicels, hypanthia and petals deep red

shrubby dwarf form

Slender tree, up to 4 m tall; intemodes of terminal branches longer

than 45 mm, slender and loose, pale green to grey; leaves thin

and papery; inflorescence axes and peduncles longer than 10

mm, slender and narrow; pedicels, hypanthia and petals pale-

green to creamy white slender tree-like form

ACKNOWLEDGEMENTS

We would like to thank Ian Geiger for allowing

access and permission to study and collect plant mate-

rial of Olinia on his Farm Letjuma, in the Soutpansberg

(Limpopo). Mervyn Letter (Mpumalanga Parks Board)

and Pieter Winter (former Curator, Herbarium of the

University of the North) are also thanked for their com-

pany and assistance in locating populations of Olinia

in Mpumalanga and Limpopo respectively. S.A. Tebele

(graduate student of the Department of Botany, University

of the Witwatersrand) is thanked for assistance in the

field and scoring of the specimens. The directors of

Nature Conservation in Mpumalanga and Limpopo are

thanked for providing permits to collect plant mate-

rial. The University of the Witwatersrand, the National

Research Foundation and the Mellon Foundation are

thanked for financial assistance. The editors and two

anonymous reviewers are gratefully acknowledged for

thoughtful comments and pointing to the literature on

continuous- and discontinuous-state characters.

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Bothalia 36,1:101-127 (2006)

Floristic composition of gold and uranium tailings dams, and adja-

cent polluted areas, on South Africa’s deep-level mines

I.M. WElERSBYE*t, E.T.F. WITKOWSKPand M. REICHARDT*

Keywords: acid mine drainage (AMD), mine rehabilitation, natural plant establishment, pyrite tailings, slimes dams, tailings storage facilities

(TSFs), vegetation survey

ABSTRACT

Gold and uranium tailings (‘slimes’) dams and the adjacent polluted soils in the deep-level mining regions of South Africa

(Carletonville, Klerksdorp and Welkom) were surveyed for the frequency of occurrence of naturally colonizing, actively

introduced and persisting plant species. Fifty-six tailings dams with a combined area of 5864 ha, and a similar area of tail-

ings-polluted soils, were surveyed between July 1996 and March 1997. During the survey, 376 plant species and subspecies

were recorded from the dams and adjacent polluted soils, with an additional 86 records obtained between 1998 and 2003 (i.e.

a total of 462 taxa: species and infraspecific species). Overall, the most commonly represented families were the Poaceae ( 1 07

species and subspecies), Asteraceae (81), Fabaceae (55) and Anacardiaceae (16). with other families represented by just one

to 14 species. Only 60 species were common to all three regions, and of these 24 had been introduced during rehabilitation

attempts. Most of the species found on tailings were persisters or natural colonizers (53-88%, depending on substrate), with

the vast majority being indigenous and perennial taxa (76% and 85% respectively), with semi-woody to woody growth forms

(66% being resprouters, forbs, shrubs and trees). Less than 4% of the naturally-colonizing taxa found during the survey had

also been introduced by vegetation practitioners. The majority of introduced plants were alien herbaceous taxa. The number

and frequency of annuals was only high on recently vegetated sites, whereas annuals were rarely present on old-vegetated and

never-vegetated dams. This list includes a wide range of indigenous plant species that may be suitable for phytoremediation

of tailings dams and polluted soils due to their apparent tolerance of acid mine drainage and salinity.

INTRODUCTION

Tailings storage facilities (TSF) containing waste

rock or milled rock slurry from the gold and uranium-

mining industry, cover vast areas in South Africa. Gold

TSF and 'footprints’ (the area of contaminated soil and

residual slimes left behind after re-mining of the original

TSF) cover about 400 km^ in the Witwatersrand Basin

goldfields alone, comprising about 6 billion tonnes of

gold and uranium tailings (Chevrel et al. 2003), and

contain an estimated 430 000 tons of uranium (Council

for Geosciences 1998; Winde 2004a, b, c) and approxi-

mately 30 million tonnes of sulphur (Witkowski &

Weiersbye 1998). The volume of waste generated by

mining in South Africa increases at the rate of 3 1 5 mil-

lion tonnes per annum, mostly in the fonn of tailings, of

which 105 million tonnes per annum is generated by the

gold mining industry on the Witwatersrand Basin alone,

at the rate of 200 000 tonnes of waste per ton of gold

(Department of Tourism, Economic and Environmental

Affairs, 2002; Chamber of Mines of South Africa, 2004).

Environmental degradation from gold TSF spreads far

beyond the waste deposit sites in the form of air pollution

(Van As et al. 1992; Mizelle et al. 1995), soil pollution

(Coetzee 1995; Rosner & van Schalkwyk 2000; Rosner

et al. 2001; Witkowski & Weiersbye 1998a) and pollu-

tion of streams, rivers, dams and sediments (Funke 1990;

Pulles 1992; Hodgson et al. 2001; Naiker et al. 2003;

Tutu et al. 2003; Coetzee et al. 2004; Winde et al. 2004a,

b, c). Since the TSF for slurry (referred to as slimes

dams) are elevated above the natural ground contours

and have steep slope angles, they are particularly sus-

* Restoration & Conservation Biology Research Group, School of Animal,

Plant & Environmental Sciences, University of the Witwatersrand,

P.O. WITS, 2050 Johannesburg.

t corresponding author: e-mail: isabel@biology.biol. wits. ac.za

MS. received: 2002-07-26.

ceptible to erosion (Mizelle et al. 1995; 1996). Whereas

erosion from agricultural fields may be as high as 10 to

15 tons ha ’ year', losses from the slopes of gold slimes

dams may exceed 500 tons ha ' year’ (Blight 1991).

Erosion and acid mine drainage from gold slimes dams

have severe impacts on nutrient cycling in polluted soils

(Witkowski & Weiersbye 1998a), on the regeneration of

vegetation (Witkowski & Weiersbye 1998b; Weiersbye &

Witkowski 2003) and on the biogeochemical cycling of

potentially toxic elements (Weiersbye et al. 1999', Winde

et al. 2004a, b; Weiersbye & Cukrowska 2005).

Prior to 1991, South Africa had little legislation spe-

cifically directed towards environmental protection from

mining impacts, although recommendations and statutes

existed for the structure and abandonment of tailings

dams (James 1964; James & Mrost 1965; Chamber

of Mines of South Africa 1968, 1979; Blight 1969).

Mines did not have a legal obligation to prevent dust

pollution until the promulgation of the Atmospheric

Pollution Prevention Act 45 (1965), amended in 1973.

The Chamber of Mines Guidelines (1979) recommend

that wind and water erosion of dams be controlled by the

most practical means possible using the BATNEEC (Best

Available Technology Not Entailing Excessive Cost)

concept. Erosion control included covering the surface of

tailings dams with waste rock, or vegetating the tailings;

the latter (‘grassing’) is still considered the most effec-

tive means of reducing dust by the industry. This accept-

ability is based on the speed with which the grass cover

establishes, rather than on its long-term persistence or

effective erosion control. The earliest recorded attempts

at rehabilitation (dust control) of gold tailings dams on

the Witwatersrand occurred in 1 894, with the planting of

Ammophila sp. (seed from Kew Gardens, UK), and have

been followed by a series of vegetation trials between

1932 and the present day (Thatcher 1979; Weiersbye &

102

Bothalia 36,1 (2006)

Witkowski 1998). Slimes dams are inhospitable envi-

ronments for plant growth, and various combinations

of leaching, liming, fertilization and irrigation are used

to facilitate the growth of a small suite of herbaceous,

mostly pasture species. As predicted by Halliday (1978),

the grass cover achieved is temporary and the methods

have proven economically and ecologically unsustain-

able (Thatcher 1979; Weiersbye & Witkowski 1998;

Witkowski & Weiersbye 1998a, b). The cost of grassing

the steep (30°-45°) slopes of tailings dams ranged from

R70 000 to R 160 000 ha ' in the period 1994 to 2002,

whereas the cost of grassing the flat tops of dams and

dam ‘footprints’ (polluted areas left after the dam itself

has been removed for re-processing to recover residual

gold) was between RIO 000 and R60 000 ha ' during the

same period (Weiersbye & Witkowski 2002a; Weiersbye

et al. 2002). Although grassing and irrigation of tailings

dam slopes significantly abates wind-borne erosion in the

short term (Blight 1991), long-term erosion control and

containment of water pollution from gold TSF by grass-

ing has been unsuccessful (Blight 1991, 1998; Weiersbye

& Witkowski 1998; Rosner et al. 2001).

South Africa now has some of the most stringent

environmental legislation in the world, with the right

to a healthy environment elevated to a basic human

right in the Constitution of South Africa (Act No. 108

of 1996). A number of Acts stress the responsibility of

industry to prevent environmental damage, and pro-

vide for the prosecution of polluters. These include the

Environment Conservation Act (ECA) No. 73 of 1989

and ECA Amendment Act 50 of 2003, the Conservation

of Agricultural Resources Act No. 43 of 1983 and

amendments of 2001, the National Environmental

Management Act 107 of 1998 and amendments, the

National Water Act 36 of 1998, the National Nuclear

Regulator Act 47 of 1999, the National Environmental

Management: Biodiversity Act No. 10 of 2004, the

National Environmental Management: Air Quality Act

No. 39 of 2004, the Minerals and Petroleum Resources

Development Act 28 of 2002, the National Environmental

Management: Protected Areas Act No. 57 of2003, and the

National Environmental Management Amendment Act

No. 46 of 2003, which facilitated the ‘Green Scorpions’

unit to investigate environmental offences. In addition,

the new regulatory framework for water usage renders

industry liable for the cost of water used, and polluted,

as a result of operations and rehabilitation under the new

Waste Discharge Charge System of the Department of

Water Affairs and Forestry. This environmental legisla-

tion means that novel, sustainable and cost-effective

methods of containing pollution from tailings dams, have

to be established.

The aim of this study was to undertake a broad-scale

survey of the plant species composition on gold and

uranium slimes dams, and slimes-polluted soils, in the

deep-level mining regions of South Africa. The under-

lying rationale was to assess the feasibility of a more

sustainable ecological engineering and phytoremediation

approach to slimes dam rehabilitation, through identify-

ing a greater suite of suitable specie! The dams surveyed

ranged in age (from 9 to 58 years since commissioning),

in planted vegetation status (present or absent) and in

the time elapsed since planting of vegetation (from 3 to

±50 years). The survey objectives were: (i) to provide

a systematic list of indigenous and alien plant species

found on slimes dams, and slimes-polluted soils; (ii) to

distinguish natural colonizers and persisters (i.e. individ-

uals present prior to slimes deposition but still surviving

despite the new conditions) from species intentionally

introduced during vegetating attempts; (iii) to broadly

classify species according to functional groups; (iv) to

assess the number of plant species and their frequency on

tailings dams differing in vegetation history at each min-

ing locality; and (v) to assess the number of plant species

and their frequency on each of the different substrates

that together constitute a tailings dam. These substrates

were identified in a parallel study and are character-

ized mainly by differences in slope angle, elevation (i.e.

time since slurry deposition), texture, soil organic mat-

ter, water content, conductivity, pH and redox potential

(Witkowski & Weiersbye 1998a).

THE STUDY AREA

Vegetation, soils and climate

The study was carried out at Anglo American Ltd

(subsequently AngloGold, and FreeGold) mines in the

Gauteng, North-West and Free State Provinces. Tailings

dams situated around Carletonville (Gauteng), Klerksdorp

(North-West) and Welkom (Free State) were surveyed for

plant species composition. The survey covered ± 12 000

ha, situated within an overall area of 150 x 100 km over

the Upper Witwatersrand Basin, on the West Wits, Vaal

and Welkom Reefs (Figure 1). All tailings dams at the

West Wits and Elandsrand gold mines (Gauteng), at the

Vaal River and Afrikaander gold mines (North-West),

and at the Free State Gold, Freddies, Western Holdings,

President Brand, President Steyn, Free State Saaiplaas

and Free Gold mines (Free State) were included in

the survey. Most of the surveyed tailings dams occur

within the Grassland Biome (Acocks 1988; Rutherford

& Westfall 1994; O’Connor & Bredenkamp 1997), with

one dam in the east occurring in the transition zone

between grassland and savanna (Afrikaander Leases in

the North-West). The Vaal River and Afrikaander dams

are situated on doleritic and sandy soils within the A2

vegetation subdivision (O’Connor & Bredenkamp 1997)

of the Grassland Biome at an altitude of 1 300 to 1 350

m. The Free State dams occur on clayey to sandy soils

within the A2 and B3 subdivisions at 1 300 to 1 400 m.

The Carletonville dams are constructed on rocky quartz-

ite, shale and dolomitic soils in the C6±7 subdivisions

at 1 600 to 1 650 m. Tailings dams in the Carletonville

region are situated within various combinations of bank-

enveld, xeric grassland (klipveld) and Acacia karroo

savanna (Acocks 1988). The main veld type in the Vaal

River and Afrikaander mine areas is a combination of

dry transitional Cymhopogon-Themeda veld, with some

development of a mixed grassy false Karoo veld, and

dry Cymbopogon-Themeda veld. However, most of the

surveyed dams here were surrounded by xeric grassland

(klipveld) and Acacia karroo savanna (Bredenkamp &

Brown 1995a, b). In the Welkom mine lease area, most

dams are surrounded by mesic to seasonally inundated

hydromorphic grassland on clays, endorheic saline pans

supporting halophytic grasses, sedges and A triplex spe-

Bothalia36,l (2006)

103

FIGURE 1 . — Sketch map showing the location of the study sites (dark grey shading), in relation to the Witwatersrand Basin (light grey shading) (after

Anhauser 1987).

cies, and perennial swamps dominated by Phragmites

australis and Tamarix spp. Two tailings dams near

Carletonville, three near Klerksdorp, and most of the

Welkom dams are situated on pans, vleis or streams.

The climate of the region surveyed is highly seasonal

and falls within the Austral summer rainfall belt (Schulze

1997). Mean annual precipitation is 662, 630 and 604

mm for Carletonville, Klerksdorp and Welkom respec-

tively, with high inter-annual variability (25-30%). The

regions all experience seasonal extremes of temperature.

Mean daily minima (July) and maxima (January) were

0-2 °C and 25-27.5 °C respectively for Carletonville

and Klerksdorp, and < 0 °C and 27.5-30 °C respec-

tively for Welkom during the study period. Evaporation

is 2-2. 5x higher than rainfall, frosts occur frequently in

winter (mean frost days is 150-175) (Schulze 1997), and

frequent veld fires occur in winter. Regional land use

includes cattle and game farming (rangelands), maize

and sunflower cropping. Wastelands (derelict, degraded

lands with little plant cover) and swampy lands inundated

by seepage from slimes dams are common within these

landscapes.

Tailings dam construction and composition

The dams were constructed using the paddock system,

which involves construction of peripheral slimes dykes

during the day (‘day walls’), and filling of the central

dam Cnight pan’) with slimes slurry during the night

(Mcphail & Wagner 1987). Excess water is drained away

during construction and the construction rate of the dam

is limited by the drying rate of the day walls. Moisture

content is high on the tops and upper slopes of current

dams due to the deposition of fresh slurry, and decreases

with distance down the slope. Moisture content increases

sharply again at the base of the lower slope and in the toe-

paddock due to seepage from the dam. The slopes of the

dams surveyed ranged between 29° and 35°; these steep

slopes result in high erosive losses (Blight 1991).

The tailings are derived from gold and uranium-bear-

ing conglomerates associated with the sediments of the

Witwatersrand Basin up to 4 000 m below surface. Pyrite

is the dominant sulphide in the conglomerates, up to 3%

of the ore mass, with an additional 2% of other sulphides

namely, pyrrhotite, galena, cobaltite, arsenopyrite and

chalcopyrite (Anhaesser 1987). The ‘all sliming’ process

was introduced in 1921. Slimes particles are cohesion-

less, predominantly silicaceous and of the size range

associated with clays and silts (Clausen 1973); the depo-

sition of such fine particles in an aqueous slurry results

in dense compaction and poor aeration in the rooting

zone. However, the reactive clay content is negligible,

with slimes consisting primarily of unreactive quartz and

pyrophyllite. The virtual absence of organic matter con-

104

Bothalia 36,1 (2006)

tributes further to the negligible cation exchange capac-

ity. The chemical composition of the slimes surveyed

varied according to the parent substrate, the metallurgical

recovery (‘sliming’) process used, the composition of

the mined ore and the age of the deposits (Bosch 1987;

Witkowski & Weiersbye 1998a). The main geochemi-

cal divisions of slimes deposits in this study occurred

between high (up to 5%) and low (< 1%) pyrite dams.

Although slimes produced using older processing tech-

niques have high sulphur contents, slimes derived from

the more recent ‘Acid Plant’ process are lower in sulphur

(Bosch 1987). Freshly deposited slimes are grey in col-

our, saline, moist and alkaline (up to pH 10.0) due to the

addition of liming agents during processing. On exposure

to air and water, the oxidation of pyrite results in the

production of sulphuric acid and ferric hydroxide, with

the tailings substrate consequently becoming acidic and

yellow. As the substrate acidifies, ferric iron also con-

tributes to oxidation, and sulphur-utilizing Thiobacillus

bacteria that occur in the tailings facilitate further ferrous

oxidation (James & Mrost 1965; Bradshaw & Chadwick

1980). On the slopes of current dams there is conse-

quently a steep pH and acidity gradient between the top

(comprising recent, alkaline deposits) and the base com-

prising older deposits of increasing acidity (Witkowski &

Weiersbye 1998a).

METHODS

Site classification

The tailings dams varied in age (from 9 to 58 years

since commissioning), in planted vegetation status (veg-

etated or never vegetated) and in the time elapsed since

planting of vegetation (from 3 to ± 50 years ago).

The tailings dams were grouped according to region

(Carletonville, Klerksdorp, Welkom), and slopes were

classed according to their vegetation history. Vegetation

history classes were: (i) recently-vegetated (RV) slopes:

amelioration (liming, fertilizing, seeding and irrigation)

had ceased >1 to < 4 years previous to 1996; (ii) old-

vegetated (OV) slopes: amelioration had ceased > 4 < 50

years previously; and (iii) never-vegetated (NV) slopes:

slopes on record as never having been intentionally

ameliorated or vegetated. Vegetation records (dates and

duration of planting, method and lists of species used)

were obtained from vegetation contracts stored with

the individual mines and from vegetation contractors.

Most dams have had some form of vegetating attempted

during the last 50 years. Distinguishing very OV slopes

(> 20 years ago) from NV slopes was made difficult

both by poor record-keeping prior to the 1980s, and the

rapid reversal of vegetated slopes to eroded, seemingly

never-vegetated conditions. This necessitated a forensic

approach to determining whether or not a dam had previ-

ously been vegetated. Previously vegetated areas were

identified from old photographs (including aerial survey),

and the remains of old plantings, irrigation pipes and

chemical signatures in the slimes as a result of liming and

intensive fertilization (Witkowski & Weiersbye 1998a).

All the dams except NV had received similar liming and

fertilization regimes. In most cases vegetation had been

established using intensive irrigation, and in seven cases,

vegetation had been established using dry-land methods.

In many cases dam slopes had been subjected to repeated

grassing attempts over the years, as each attempt had

failed.

Each slope was further subdivided into substrate

classes based upon marked differences in physical and

chemical properties (Figure 2). From the top of the dam

downwards, substrates comprised the flat tops, upper-to-

mid slopes and berms, mid-to-lower slopes and berms,

retaining walls (rock and soil mixed or overlaid with

slimes) and toepaddocks (a strip of veld from 20 to 60 m

wide surrounding the base of the dam and bordered by

an earthen wall). The toepaddocks are heavily inundated

by slimes, strongly acidic and often damp (Witkowski

& Weiersbye 1998a). The tops, bernis and slopes were

further categorized according to whether they comprised

younger or recently ameliorated and marginally acidic to

alkaline (pH > 6.0) slimes deposits, or older and more

acidic (pH < 5.9) slimes deposits.

Sampling methods

Lists of species planted on each tailings dam, and

planting methods, were obtained from vegetation con-

tracts and assessments archived with Anglo American

Mines and from individual contractors, from unpublished

theses and reports (including those lodged with indi-

vidual mines and the Chamber of Mines of South Africa;

Thatcher 1979 (and references therein); Wiegenhagen

1996) and from publications (James & Mrost 1965;

Wild & Wiltshire 1971; Cresswell 1973; Grove 1974;

Clausen 1976; Bradshaw & Chadwick 1980). Two repli-

cate surveys were carried out on the same dams within a

nine-month period, in winter (June to September, 1996;

Witkowski & Weiersbye 1996) and subsequently in

summer (December 1996 to March 1997; Weiersbye &

Witkowski 1997). Fifty-six tailings dams comprising 738

different slopes with the same number of toepaddocks

were assessed. Intensive searches were carried out on

dams, retaining walls and toepaddocks. These comprised

(i) large-scale assessments of vegetation cover (data not

TOP

FIGURE 2. — Aijpect of a gold slimes

dam divided into the substrate

classes.

Bothalia36,l (2006)

105

shown, Weiersbye & Witkowski 1998), vegetation struc-

ture (proportions of trees and shrubs, forbs and herbs, and

grasses) and species presence on all 738 slopes, retaining

walls and toepaddocks; and (ii) 254 one hundred metre

wide belt transect surveys on a sub-suite of dams (those

which had > 0.5% aerial vegetation cover). Data from

these 1 00 m belt transects were then subdivided for each

substrate on the slimes dam (upper, middle and lower

slopes, tops, berms and toepaddocks) as these invariably

differed in species composition. All species present on

transects were identified and the number of individuals

present for most species recorded. Additional records of

species presence only on slimes dams, retaining walls

and toepaddocks were obtained from the same mines

during 1998 to 2003.

Representative specimens of each taxon were col-

lected and pressed. Plants were identified using the keys

of Dyer (1975, 1976), Gibbs Russell et al. (1985, 1987,

1991), Venter & Joubert (1985), Coates-Palgrave (1996)

and Relief & Herman (1997), and by comparison with

specimens at the C.E. Moss and National Herbariums (PRE)

of South Africa. Taxa were named according to Arnold

& De Wet (1993). A few taxa (reputedly introduced

from Namibia and the Northern Cape) defied identi-

fication beyond genus. Voucher specimens have been

lodged with the C.E. Moss Herbarium, University of the

Witwatersrand.

Data classification

Species, subspecies and varieties found in the 100 m

transect survey {n = 327) were categorized according to

their % frequency of occurrence: (i) overall, (ii) in each

of the three regions, (iii) on dams of different vegetation

history classes, and (iv) on each substrate class. The rela-

tive contribution of each family (in terms of component

species) was also calculated for each slimes dam sub-

strate class.

Using the 1998 PRECIS database and vegetation

distribution records (Arnold & De Wet 1993; Relief

& Herman 1997), all species, subspecies and varieties

found during the 1996-1997 survey {n = 376) were cat-

egorized for each dam substrate and overall according to:

(i) whether they were indigenous to South Africa or alien

(including naturalized species), and (ii) whether they

were indigenous to each of the three regions surveyed.

Species were also grouped on the basis of: (iii) annual or

perennial habit; (iv) broad growth habit (shrubs and trees,

forbs and perennial herbs, annual herbs, annual and per-

ennial grasses); and (v) whether they were persisters or

naturally colonizing species, or intentionally planted on

dams, or combinations. Persisters were generally consid-

ered to be plants that pre-dated slimes dam construction.

These were categorized as old woody plants present on

toepaddocks and/or growing through retaining walls, but

usually not present on other substrates. Naturally coloniz-

ing species were those present on other substrates (berms

and/or slopes and/or tops), either solely or in addition to

being present on toepaddocks and/or retaining walls. All

non-woody species present on retaining walls were cat-

egorized as natural colonizers. Species found subsequent

to the survey in 1998 to 2003 {n = 86) were not included

in the frequency analysis or categorization.

RESULTS

Broad species-compositional patterns

A total of 376 species, subspecies and varieties were

recorded during the intensive winter (1996) and summer

(1997) surveys, of which the frequency of 327 species

and subspecies was recorded using the detailed 100

m transects (Appendix 1). The other (49) species and

subspecies are listed in Appendix 2. However, all 376

taxa were included in the general analyses (Tables 1-5).

Thirty-six taxa could only be identified to genus level.

The number of species on dams in the three provinces

was in the order of Klerksdorp (216 species in 1 488 ha),

Carletonville (168 in 765 ha), Welkom (120 in 3 611 ha)

(Appendix 1). The highest number of species was found

on the 100 old-vegetated and 139 never-vegetated slopes

(260 and 231 species respectively). The 15 recently-vege-

tated slopes contained 86 species, of which 21 had been

intentionally introduced during grassing, with another 1 8

weedy annuals and short-lived perennials.

Most species occurred on toepaddocks and retaining

walls (287 and 264 species respectively, of which 246

and 231 respectively were natural colonizers and per-

sisters), followed by the acidic slopes (149 species of

which 106 were natural colonizers) and acidic tops and

berms (137 species of which 103 were natural coloniz-

ers). Species composition differed markedly between

substrates (Appendix 1). Only 32 species (14 of which

had been introduced) were common to more than five

substrates, and 161 (16 of which had been introduced)

occurred on just one or two substrates. When considering

species common to tops and berms and slopes, only six

species were common to both acidic (pH < 6.0) and mar-

ginally acidic to alkaline substrates (pH > 6.0), whereas

for tops and berms only, 12 species were common to both

acidic and marginally acidic to alkaline substrates. For

slopes, 28 species were common to both acidic and mar-

ginally acidic to alkaline substrates. Sixty species were

common to flat and sloped acidic substrates, and just

eight were common to fiat and sloped marginally acidic

to alkaline substrates. Of the 376 taxa recorded during

the survey, only 60 were common to all three mining

regions, and of these, 24 were introduced during vegetat-

ing attempts (Appendix 1). Only 10% of taxa overall

(including < 4% of those which were natural colonizers

and persisters) are known to have been introduced during

vegetating attempts (Table 1; Appendix 1).

In addition to the 376 taxa, 86 taxa (species, subspe-

cies and varieties) were recorded from 1998 to 2003

(Appendix 3), but were not included in the general

analyses (Tables 1-4) or frequency survey (Appendix J ).

Of the 86 taxa listed in Appendix 3, 40 were introduced

species (three were indigenous trees), and 46 natural

colonizers and persisters (39 were indigenous). Thus a

grand total of 462 taxa (species, subspecies and varieties)

have been identified on gold mine tailings and tailings-

polluted soils in the three regions.

Plant families and higher taxa

No gymnosperms were found on slimes during the sur-

vey (two species were found post-survey). Dicotyledons

dominated the vegetation on all substrates. The high-

106

Bothalia 36,1 (2006)

TABLE 1 . — Percentages of persisting, naturally colonizing and intentionally introduced plants on each substrate class for surveyed slimes dams.

Categories mutually exclusive and values in parentheses are no. taxa found (species, subspecies and varieties, n = 376). Note that category

‘introduced and colonizing’ is for those introduced species that can also colonize unaided

* Twenty-one taxa included here as persisters occurred only on retaining walls and may actually be natural colonizers.

est proportion of monocot taxa was observed on the

two moistest and on flat substrates (regardless of pH):

35% on tops and berms, and 31% on toepaddocks. The

majority of monocots on these two substrates were

grasses (up to 28%), with Asparagaceae, Hyacinthaceae,

Hypoxidaceae and Juncaceae each contributing up to 1 .8%

of the total taxa on the former substrate, and Alliaceae,

Amaryllidaceae, Asphodelaceae, Commelinaceae, Cyper-

aceae and Typhaceae only present on toepaddocks, with

each of the latter families containing just 0. 3-1.0% of the

total species detected (Appendix 1 & 2).

The dominant families represented overall were

the Poaceae (containing 23.4% of all species found),

Asteraceae (17.2%), Fabaceae (10.5%) andAnacardiaceae

(3.8%), with other plant families containing between

0.3% and 3.5% of the total (Table 2; Appendices 1 &

2). A similar pattern was observed on toepaddocks and

retaining walls. On acidic (pH < 6.0) substrates (tops,

berms and slopes), most species belonged (in descend-

ing order of frequency) to the Asteraceae, Poaceae,

Fabaceae and Chenopodiaceae, whereas on marginally

acidic to alkaline (pH > 6.0) substrates (tops, berms and

slopes) most species belonged to the Poaceae, Fabaceae

and Asteraceae, followed by equal representation of

the Anacardiaceae, Chenopodiaceae and probably the

Tamaricaceae. Identification of Tamarix spp. was dif-

ficult due to the presence of putative hybrids between T.

usneoides, T ramosissima, T chinensis, T. gallica and

T. aphylla. Genera represented by > 4 species on slimes

dams were Acacia (12 spp.), Aristida (7), Asparagus (4),

Eragrostis (8), Felicia (4), Helichrysum (6), Hermannia

(4), Paspalum (5), Rhus (12), Senecio (6), Solarium (6)

and Sporobolus (4) (Appendix 1). An additional two

Acacia spp., five Eragrostis spp., five Helichrysum spp.,

three Rhus spp., three Senecio spp. and two Sporobolus

spp. were recorded subsequent to the survey (Appendices

2 and 3).

Indigenous versus alien species

The survey yielded a total of 90 alien species (includ-

ing naturalized species) and 286 species indigenous to the

southern African region. Fifty-five alien and 152 indig-

enous species occurred on tailings dams (slopes, berms

and tops), whereas 59 alien and 143 indigenous species

occurred on retaining walls and toepaddocks (Appendix

1). Eight alien and 14 indigenous species (of which

four had been introduced to dams from other regions)

occurred only on dams, and not on polluted soils. Overall,

the majority of species growing on slimes (76.1%), and

the vast majority of natural colonizers and persisters were

indigenous to southern Africa (Table 3), with most (91%)

normally found in the local province (Table 4). With the

exception of marginally acidic to alkaline slopes (where

numbers of indigenous species only slightly exceeded

those of alien species), the same pattern prevailed on all

slime substrates.

Species characteristic of particular substrates

The number of naturally colonizing and persisting spe-

cies was higher on acidic (pH < 6.0) substrates, in order

of abundance: retaining walls and toepaddocks > slopes

and tops and berms > marginally acidic to alkaline (pH

> 6.0) substrates (Table 1). Although species number was

slightly higher on slopes than on fiat substrates (tops &

berms), overall vegetation cover was always much higher

on flat surfaces (Weiersbye & Witkowski 1998). On

marginally acidic to alkaline substrates, a relatively high

proportion of introduced taxa were also natural colonizers

(28-30%, Table 1 ). However, once the substrate became

more acidic (pH < 6.0), this proportion decreased to

10-23%. Few species (8-11%) on retaining walls and

toepaddocks had been introduced. These two substrates

had the highest levels of vegetation cover and number of

species, dominated by indigenous, naturally colonizing

and persisting perennial taxa (Appendix 1).

Ecological traits of species

The vegetation of tailings was dominated by peren-

nial species (Table 5), with most also characterized by a

deciduous habit. The majority of natural colonizers and

persisters were perennial plants, whereas the majority of

introduced species were annual and short-lived perennials

(Table 5, Appendices 1, 2 and 3). Very few indigenous,

perennial species had been intentionally introduced to

TABLE 2. — Percentages of alien (including naturalized) and indig-

enous plants on tailings dams. Values in parentheses are no. taxa

(species, subspecies and varieties) found

Bothalia36,l (2006)

107

TABLE 3. — Percentages of taxa occurring on particular substrates within plant families (n = 65) during 1996 to 1997 survey. Values in parenthe-

ses are actual no. taxa (species, subspecies and varieties) found

*Percentages could not be calculated for Tamaricaceae as some of the 5 species were not always distinguishable in the field.

108

Bothalia 36,1 (2006)

TABLE 4. — Percentages of occurrence of plant species found on tail-

ings dams (based on PRECIS database). Values in parentheses

are no. taxa (species, subspecies and varieties) found

slimes. Overall, 84.6% of species were perennials versus

15.4% for annuals and short-lived perennials. The vast

majority of species found on slimes overall had semi-

woody to woody growth forms: perennial forbs and

herbs (47.6%), followed by perennial grasses (18.6%),

and shrubs and trees ( 1 8.4%), with annual herbs ( 1 1 .2%)

and annual grasses (4.3%) forming minor components

(Table 5). A similar pattern was seen on each substrate.

The relatively high contribution of shrubs and trees to

the persisting vegetation of marginally acidic to alkaline

tops and berms (± 30%) was due to the low height (depth

of slimes) of three current (recently commissioned) dams

included in the survey. These dams contained large live

trees that were rooted in the underlying soil and had sur-

vived tailings dam construction and inundation by slimes

for a number of years. Islands of fertility had fonned

on the slimes under the canopies of these trees, and

contained a number of herbaceous species that were not

found on this substrate under any other conditions. Trees

and shrubs were often abundant on the lowest reaches of

dams (base of lower slope and retaining wall). The large

size and/or morphology of many of these plants sug-

gests that they pre-date dam construction, and had grown

through the slimes. In contrast, trees and shrubs on the

slopes, berms and tops appear to be rooting only within

the slimes.

The naturally colonizing and/or persisting taxa com-

prised mostly woody and semi-woody growth forms,

whereas the majority of introduced taxa were herbaceous

forms (Appendix 1 ). The majority of naturally colonizing

grasses were C4 species, while virtually all the introduced

grass taxa were pasture species comprising a mixture of

C3 and C4 taxa (Appendix 1; Gibbs Russell et al. 1991).

The introduced species mirror the commercial availabil-

ity of grass seed in South Africa. Most of these species

are intended for intensively managed pasture cultivation

on agricultural lands, and not for the rehabilitation of low

nutrient, saline and acidic tailings dams. In a previous sur-

vey of slimes dam tops in Johannesburg, Thatcher (1979)

recorded a total of 142 species, 94 of which were also

recorded in this survey. Thatcher (1979) also found that

the majority of species were natural colonizers and the

Asteraceae, Fabaceae and Poaceae were well represented.

Woody species diversity was higher on more acidic sites,

whereas grasses dominated on less acidic sites.

Few of the 376 species, subspecies and varieties

found in the 1996-1997 intensive survey are known to

have been intentionally introduced to slimes dams during

vegetating attempts over the last 50 years. Some species

were introduced to slimes dams in the surveyed region by

the Chamber of Mines Vegetation Unit in the 1950s (B.

Cook, B. Dawson, J. Easton, pers. comm.) However, it is

not known whether existing conspecifics are remnants of

the introduced populations, or individuals that have natu-

rally colonized tailings. Some species exhibiting unusual

regional distributions may be remnants of these attempts

by the Chamber of Mines Vegetation Unit (e.g. Bassia

salsoloides colonizing old slimes dams in the Welkom

region), whereas other species were collected in remote

regions and introduced to tailings dams by mine person-

nel (e.g. Riischia spp. in the Welkom region).

DISCUSSION

This survey found a surprisingly high number of plant

species growing on tailings and tailings-polluted soils.

Earlier (pre-1980) attempts at vegetating slimes utilized

a number of woody, alien species such as Australian

acacias (wattles), eucalypts and tamarix in addition

to herbaceous legumes and pasture grasses (Thatcher

1979), whereas more recent attempts utilized herbaceous

(pasture) species and cultivars. On the basis of old pho-

tographs, many of these planted trees still survive on the

slopes and tops of tailings dams. Although the contribu-

tion of natural colonizers and persisters to cover could be

substantial on the flatter surfaces of dams (tops, benns,

toepaddocks), the contribution to cover on slopes was

extremely low and individual plants were transient due to

the high rates of erosion (Weiersbye & Witkowski 1998).

The lower number of species found on marginally acidic

to alkaline (pH > 6.0) tops, benns and slopes could be due

to some influence of pH, but also to the much younger

TABLE 5. — Categories of plants found on gold slimes dams. Values are percentages, and values in parentheses are no. taxa (species, subspecies

and varieties) found

Bothalia36,l (2006)

109

age of these substrates in comparison to acidic (pH < 6.0;

i.e. older and more oxidized) substrates, the low number

of species introduced to recently ameliorated slopes by

contractors, and the fact that many dams were current

dams, with slurry still being deposited on the top. As a

consequence, the tops of current dams only supported

reeds and sedges (Phragmites sp. and Cyperaceae), if

any vegetation. Marginally acidic to alkaline substrates

are also usually further and higher (in altitude) removed

from seed sources (i.e. surrounding veld) than the older,

acidic substrates on the lower slopes of dams. In addition,

vegetating efforts were seldom undertaken on slimes of

more recent genesis as the Chamber of Mines vegetation

guidelines (1979) recommended a dormant period for

slimes dams prior to leaching and grassing.

According to species distributional databases, the pat-

tern of species number found on slimes dams is similar to

that for the provinces as a whole, with the North-West and

Gauteng having the highest number of species, and the

Free State the lowest (Arnold & De Wet 1993; Retief &

Herman 1997). At the local scale, many Free State dams

occur within a degraded agricultural and semi-industrial

setting, with the only natural seed source emanating from

wastelands, hydromorphic grasslands, perennial swamps

dominated by Phragmites australis, and alkaline pans.

This landscape context limits the diversity and availabil-

ity of natural colonizers. Most dams in the North-West

and Gauteng region were in close proximity to natural,

albeit degraded veld and private (mine) nature reserves,

and this environmental setting provides a wider diversity

of suitable species for natural colonization.

Of the species identified, < 5% had been actively intro-

duced during grassing. Despite the high species diversity

of natural colonizers and persisters, most individuals

were detected on toepaddocks, retaining walls and on the

flatter surfaces of the dams (berms and tops), with the

actual contribution to cover on slopes being extremely

low. In contrast, the number of species introduced during

vegetating efforts was extremely low, despite the high

cover achieved on recently grassed dam slopes. However,

most introduced species are herbaceous and weedy,

and both cover and number of species declines rapidly

once liming, fertilization and supplemental watering has

ceased. Watering occurs either in the form of irrigation

on dormant dams (slimes are no longer deposited) or

water from slurry deposition on current dams. Less than

five species remained on any particular tailings slope by

four years after amelioration had ceased (Weiersbye &

Witkowski 1998). Although we recorded an additional

14 introduced pasture species in 1998-2003 from the

same slimes dam slopes surveyed in 1996-1997, these

plants occurred on dams that were undergoing grassing.

These same species had not been detected in the previous

survey on post-amelioration grassed sites (despite having

been originally planted), which suggests that they lack

persistence.

Although introduced species on dams were predomi-

nantly pasture grasses, the naturally colonizing species

were predominantly perennials with woody and semi-

woody growth forms. In the case of tailings dams in the

Carletonville and Klerksdorp regions, the predominance

of indigenous, woody growth forms as natural colonizers

and persisters is expected as these growth forms are com-

mon in the localities surrounding the dams (Arnold &

De Wet 1993; Bredenkamp & Brown 1995a,b; Retief &

Herman 1997). However, tailings dams in the Free State

are surrounded by hydromorphic grasslands, degraded

wetlands and alkaline pans, with woody and semi-woody

plants largely restricted to low, dry rocky outcrops (Fuls

et al. 1992, 1993; Malan et al. 1998). The colonization

of tailings in this latter region by semi-woody/woody

species despite their restricted availability strongly sug-

gests that these are suitable growth forms for tailings dam

rehabilitation.

The dominance of Poaceae, Asteraceae, Fabaceae and

Anacardiaceae on acid slimes suggests an inherent toler-

ance to the prevailing conditions in species of these fami-

lies. These results are further reinforced by seed biology

studies (Witkowski & Weiersbye 1998b; Weiersbye &

Witkowski 2002b, 2003) and by plant growth and water-

use trials in acid slimes and AMD conditions in which

hard-seeded legumes and Rhus spp. perform especially

well (Weiersbye et al. 1998; Dharamraj et al. 1999; Dye

et al. 2005). More recently, the use of AFLP analysis

has demonstrated that there is genetic evidence for local

adaptation of some woody species to the slimes-polluted

soils around tailings dams (Angus 2005). Some species

examined during this surv^ey showed no signs of physio-

logical stress despite growth on slimes, e.g. Tamarix

spp.. Acacia spp., Lessertia spp. (= Sutherlandia spp.),

Rhus spp.. Asparagus spp. and perennial Eragrostis spp.

Seed production and seed viability levels in these taxa

approaches that of conspecifics growing in unpolluted

veld, and seed production and viability in Asparagus

spp., woody legumes and Rhus spp. on tailings is high,

with seedlings establishing around parent plants. In con-

trast, seed production and viability in most grasses and

Asteraceae growing on tailings is low and regeneration

on tailings would therefore be dependent on seed dis-

persal from beyond the dam (Witkowski & Weiersbye

1998b). A parallel survey found that the majority of

plants persisting on tailings were infected by arbuscular

mycorrhizal (AM) fungi (Straker et al. 2006a, b). In addi-

tion, plant growth experiments have demonstrated that

slimes-tolerant AM fungi, and, for most indigenous hard-

seeded legumes, compatible tolerant rhizobia, contribute

significantly to host plant survival and growth in acidic

slimes (Weiersbye et al. 1998; Straker et al. 2006c).

The dominant plant functional growth forms (i.e.

woody and semi-woody perennials, resprouters) of slimes

dams are typical of stressful environments (Grime 1979),

whereas many of the grass and forb species are character-

istic of nutrient-poor (especially nitrogen), low competi-

tion environments in the Grassland Biome (O’Connor &

Bredenkamp 1997). The distribution of species on par-

ticular tailings substrates appears to be associated with

known physiological tolerances to moisture and nutrient

availability regimes. For example, species of Eragrostis

and Sporobolus that were prevalent on the arid substrates

typical of old vegetated and never vegetated tailings can

tolerate dehydration of foliage to the point of air dryness

(Gaff 1971; Gaff & Ellis 1974). The grass Cenchrus

ciliaris is tolerant of high nitrogen and phosphorus

availability (O’Connor & Bredenkamp 1997) and is an

indicator of nutrient enrichment in natural ecosystems.

no

Bothalia 36,1 (2006)

being characteristic of environments such as iron-age

kraal sites (Blackmore et al. 1990). C. ciliaris was only

prevalent on recently vegetated sites, with few individu-

als surviving the transition to old vegetated sites and the

concomitant decline in nutrient availability. In contrast,

Stoebe vulgaris and some dominant indigenous grasses

on tailings are inhibited by high nitrogen availability

(Roux 1969), and these species were abundant only on

the low nutrient substrates typical of the oldest vegetated

and never vegetated sites. Populations of S. vulgaris, and

similar species present on never vegetated sites that were

subsequently grassed and fertilized have died out.

Tailings solution extracts have extremely high con-

ductivity and salinity (Witkowski & Weiersbye 1998a).

Halophytic plants such as Tamar ix spp. and Atriplex

spp. naturally colonize, and may form dense cover, on

the moist and marginally acidic to alkaline slopes and

tops of tailings dams. However, there is no analogous

natural environment to the combination of acidity, salin-

ity, high heavy metal availability and low macronutrient

(N, P, K, Ca and Mg) availability that is found on the

lower reaches of dams. Plants that naturally colonize

gold tailings, or persist on tailings-polluted substrates

are therefore exhibiting a remarkable combination of

adaptive or constitutive physiological tolerances and are

being subjected to massive selection pressures (Bradshaw

1952; 1970; Mehary 1994). For example, the majority

of species growing on slimes dams and polluted sub-

strates in the goldfields have depressed seed production

and viability levels in comparison to conspecifics on

unpolluted substrates (Weiersbye & Witkowski 2002b).

However, those species that occur frequently on slimes

and slimes-polluted soils are also the species that main-

tain regeneration potential on these same substrates

(Witkowski & Weiersbye 1998). Tree species such as

Acacia karroo, A. hereroensis, A. hebeclada and Rhus

lancea maintain relatively high levels of seed viability

and germination despite elevated inorganic contents, and

high frequencies of seedling abnormalities (Weiersbye

& Witkowski 2003). Rhus lancea is also capable of vig-

orous growth in acid mine drainage, maintaining high

evapotranspiration rates (Dye et al. 2005). Salinity and

acid-tolerant land-races of the grass Cynodon dactylon,

and local ecotypes of Hyparrhenia hirta are now used in

slimes dam rehabilitation (B. Dawson, EMPR Services,

pers. comm.). Apparently healthy C. dactylon growing

in gold tailings can contain 30 mg g ' dry mass of iron

in the root epidermis and cortex, and 3 mg g ' dry mass

of uranium within the root stele, with even higher Fe and

U concentrations associated with arbuscular mycorrhizal

structures (Weiersbye et al. 1999).

Some indigenous, naturally-colonizing or persisting

species that were frequently encountered on slimes dams,

have also been recorded from gold tailings in Zimbabwe

and Botswana (Wild 1974a, b), and from andalusite,

asbestos, gold, platinum and base metal tailings in the

Limpopo and Mpumulanga Provinces (I.M. Weiersbye,

K. Balkwill & E.T.F. Witkowski unpublished). Land-

races that persist and colonize gold tailings and acid

mine drainage-polluted soils can be expected to have

phytoremediation potential for the gold mining industry

in South Africa.

CONCLUSIONS

Of the 376 species found in the intensive survey, only

60 were common to all three regions, and of these 24 had

been introduced during rehabilitation attempts. Most of

the species found on tailings were persisters or natural

colonizers (53-88%, depending on substrate), with the

vast majority being indigenous and perennial taxa (76%

and 85% respectively) with semi-woody to woody growth

forms (66% being resprouters, forbs, shrubs and trees).

The present rehabilitation aims of mine management (i.e.

the requirement for rapid green cover) forces vegeta-

tion contractors to expend massive effort and expense in

modifying tailings dams to become temporarily suitable

substrates for high basal cover pastures. In contrast, those

species actually persisting on, and naturally colonizing

tailings and tailings-polluted soils, are non-pasture spe-

cies, of which < 5% have also been introduced during

vegetating efforts. Naturally colonizing and persisting

species are predominantly indigenous perennials com-

prised of resprouting, semi-woody and woody plants and

C4 tussock grasses, which, by virtue of the comparatively

longer life-span of individuals and apparent tolerance to

native slimes conditions are more likely to assist in the

establishment of self-sustaining cover and rehabilitation

of gold tailings. Finally, only multi-stemmed/shrubby or

ground-covering woody plants and other growth forms

with high basal cover are suitable for planting on the

slopes of the dams due to erosion foci that may develop

around large, single-stemmed-trees. However, plantings

of shrubs and trees are suitable for the berms and tops

of slimes dams, providing that planting densities are

optimized in order to survive on incoming rainfall alone

in an increasingly arid environment as the dam dries out.

Woody plantings on the berms and tops of dams could

minimize recharge of the phreatic surface within the dam,

and thus limit the potential for seepage, as well as lower-

ing the risk of fire that is ever-present in highveld winter

grasslands, provide large canopies in order to abate wind