Bothalia

A JOURNAL OF BOTANICAL RESEARCH

Vol. 33.1

May 2003

TECHNICAL PUBLICATIONS OF THE NATIONAL BOTANICAL INSTITUTE,

PRETORIA

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

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

BOTHALIA

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

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

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

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

published annually.

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

STRELITZIA

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

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

MEMOIRS OE THE BOTANICAL SURVEY OE SOUTH AERICA

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

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

ANNALS OF KIRSTENBOSCH BOTANIC GARDENS

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

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

South African Botany). Discontinued after Vol. 19.

FLOWERING PLANTS OF AFRICA (FPA)

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

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

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

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

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

or of economic value for illustration.

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

An index to Vols 1^9 is available.

FLORA OF SOUTHERN AFRICA (ESA)

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

and Botswana. The FSA contains descriptions of families, genera, species, infraspecific taxa, keys

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

and ecological notes.

Contributions to the FSA also appear in Bothalia.

PALAEOFLORA OF SOUTHERN AFRICA

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

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

available:

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

Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), 1989, by J.M.

& H.M. Anderson.

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

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

RSA.

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

by J.M. Anderson (ed.)

BOTHALIA

A JOURNAL OF BOTANICAL RESEARCH

Volume 33,1

Scientific Editor: G. Germishuizen

Technical Editor: B.A. Momberg

NATIONAL

.Botanical

INSTITUTE

NASIONALE BOTANif^S

INISTITUUT ~

Private Bac- x in-' po-y.^n , ,

2003 -G8- 1 3 ' C

2 Cussonia Avenue. Brummeria, Pretoria

Private Bag XIOl, Pretoria 0001

ISSN 0006 8241

May 2003

Centenary Year

IN CELEBRATION

OF

THE CENTENARY OF THE NATIONAL HERBARIUM (PRE)* IN 2003 AND

ITS IMMEASURABLE CONTRIBUTIONS TO BOTANY IN SOUTH AFRICA AND FURTHER AFIELD

Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

J.P. Rourke

M.J.A, Werger

Royal Botanic Gardens, Kew, UK

National Botanical Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

Compton Herbarium, NBI, Cape Town, RSA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Acknowledgements to referees

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

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

Bohs, Dr L. University of Utah, Salt Lake City, USA.

Brown, Dr E. Royal Botanic Gardens, Sydney, Australia.

Brummitt, Dr R.K. Royal Botanic Gardens, Kew, UK.

Burgoyne, Ms P. National Botanical Institute, Pretoria, RSA.

Craven, Ms P. P.O.Box 399, Omaruru, Namibia.

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

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

Hammer, S. Sphaeroid Institute, 2077 Monte Vista Drive, Vista, USA.

Hilliard, Dr O.M. Royal Botanic Garden, Edinburgh, Scotland, UK.

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

Kativu, Dr S. University of Zimbabwe, Harare, Zimbabwe.

Leistner, Dr O.A. National Botanical Institute, Pretoria, RSA.

Manning, Dr J.C. National Botanical Institute, Cape Town, RSA.

Nicholas, Dr A. University of Durban-Westville, Durban, RSA.

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

Oliver, Dr E.G.H. National Botanical Institute, Cape Town, RSA.

Paterson-Jones, Dr D. National Botanical Institute, Cape Town, RSA.

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

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

Smith, Dr G.E. National Botanical Institute, Pretoria, RSA.

Stedje, Prof. B. University of Oslo, Norway.

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

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

Victor, Ms J.E. National Botanical Institute, Pretoria, RSA.

Werger, Prof. M.J.A. University of Utrecht, The Netherlands.

.See Bothalia 28: 271-297 (1998).

CONTENTS

Volume 33,1

1. The genus Solanum (Solanaceae) in southern Africa: subgenus Leptostemomim, the introduced sections

Acanthophora and Ton’a. W.G. WELMAN 1

2. Studies in the genus Machairophyllwn (Mesembryanthemaceae), with notes on some related genera.

H. KURZWEIL and P. CHESSELET 19

3. A new serotinous species of Cliffortia L. (Rosaceae) from Northern Cape, South Africa and section

emended. A. C. EELLINGHAM 41

4. A revision of Ledebouria (Hyacinthaceae) in South Africa. 3. The reinstatement of L. ensifolia, L. gol-

pinii and L. sandersonii. T.J. EDWARDS and S. VENTER 49

5. An annotated checklist of the pteridophyte flora of Swaziland. J.P. ROUX 53

6. Taxonomy of the genus (Thymelaeaceae). C.L. BREDENKAMP and A. E. VAN WYK .... 59

7. Studies in the liverwort family Aneuraceae (Metzgeriales) from southern Africa. 5. Riccanlia amazonica.

S.M. PEROLD 99

8. Notes on African plants:

Apocynaceae. A new subspecies of Brachystelma from Eastern Cape, South Africa(Asclepia-

doideae-Ceropegieae). A.P DOLD and PV BRUYNS 105

Asteraceae. A new species of Philyropliylliim (Gnaphalieae) from Namibia. PPJ. HERMAN .... 118

Hyacinthaceae. A new species and new combinations in Drimia (Urgineoideae). J.C. MANNING

and P. GOLDBLATT 109

Hyacinthaceae. A new species of Ornithogaliim from Eastern Cape, South Africa. A.P. DOLD

and S. A. HAMMER 112

Hyacinthaceae and Crassulaceae. Two new cremnophilous taxa from semi-arid regions in South

Africa. E.J. VAN JAARSVELD and A.E. VAN WYK 115

Lamiaceae. Tetradenia kaokoensis, a new species from Kaokoland, Namibia. E.J. VAN JAARS-

VELD and A.E. VAN WYK 107

Lamiaceae. The correct name for Salvia thermara. E.J. VAN JAARSVELD 112

9. A comparison of Mopaneveld vegetation in South Africa. Namibia and Zimbabwe. F. SIEBERT, G.J. BRE-

DENKAMP and S.J. SIEBERT 121

10. Miscellaneous notes:

Hyacinthaceae. Chromosome studies on African plants. 19. New chromosome counts for three

taxa. P.D. LEBATHA, J.J. SPIES and M.H. BUYS 135

11. Book reviews 139

New name, species, subspecies and combinations in Bothalia 33,1 (2003)

Albuca therniarum Van Jaarsv., sp. nov., 116

Brachystelma franksiae N.E.Br. subsp. grandiflorum A.PDold & Bniyns, subsp. nov., 105

Cliffortia dichotoma Fellingham, sp. nov., 41

Crassula foveata Van Jaarsv., sp. nov., 116

Drimia barkerae Oberni. ex J.C. Manning <& Goldblatt, sp. nov., 109

Drimia ciliata (Lf.) J.C. Manning & Goldblatt, comb, nov.. Ill

Drimia hesperantha J.C. Manning & Goldblatt, nom. nov.. Ill

Drimia kniphofloides {Baker) J.C. Manning & Goldblatt, comb, nov., Ill

Drimia nana (Snijinan) J.C. Manning & Goldblatt, comb, nov.. Ill

Drimia saniensis (Hilliard & B.L.Burtt) J.C. Manning & Goldblatt, comb, nov.. Ill

Ledebouria ensifolia (Eckl.) S. Venter & T.J. Edwards, comb, nov., 49

Ledebouria galpinii (Baker) S.Venter & T.J. Edwards, comb, nov., 50

Ledebouria sandersonii (Baker) S.Venter & T.J. Edwards, comb, nov., 50

Ornithogalum perdurans A.P.DoW & S. A. Hammer, sp. nov., 112

Passerina esterhuyseniae Bredenkamp & A.E.van Wyk, sp. nov., 75

Philyrophyllum brandbergense P.P.J. Herman, sp. nov., 118

Tetradenia kaokoensis Van Jaarsv. & A.E.van Wyk, sp. nov., 107

iv

Bothalia 33.1: 1-18 (2003)

The genus Solanum (Solanaceae) in southern Africa: subgenus

Leptostemonum, the introduced sections Acanthophora and Torva

W.G. WELMAN*

Keywords: Acanthophora Dunal, ethnobotany, Leptostemonum (Dunal) Bitter, morphology, Solanaceae, Solanum L., southern Africa, taxonomy,

Ton'a Nees

ABSTRACT

In the genus Solanum L. (Solanaceae), subgenus Leptostemonum (Dunal I Bitter, section Acanthophora Dunal has four

representatives in the Flora of southern Africa region (South Africa, Lesotho, Swaziland, Namibia, Botswana), namely S.

aculeatissimum Jacq., S. Allioni. S. mammosum L. (cultivated only) and S. viarum Dunal. Section Tona Nees

has two representatives in southern Africa, namely S. chrysotrichum Schltdl. (S. hispidum auctt. non Pers.) and S. tonmni

Sw.; both are naturalized weeds. Solanum capsicoides, S. viarum and S. ton'um have not been listed before for southern

Africa. All are introduced species native to the New World. Descriptions, discussions, illustrations and distribution maps of

the naturalized species are presented, as well as keys to the species of both sections.

CONTENTS

Abstract 1

Introduction 1

Descriptions and discussions 1

Section Acanthophora Dunal 1

Solanum aculeatissimum Jacq 3

S. capsicoides Allioni 6

S. mammosum L 8

S. viarum Dunal 10

Section Torx’a Nees 11

Solanum chrysotrichum Schltdl 12

S. tonnim Sw 13

Specimens examined 15

Acknowledgements 16

References 16

INTRODUCTION

In the genus Solanum L., the prickly subgenus Lepto-

stemonum (Dunal) Bitter is represented by eight sections

in southern Africa. Two sections contain only indigenous

taxa, one section contains mainly indigenous taxa, and

the remaining sections have only introduced species.

Section Acanthophora Dunal is represented by four

species in southern Africa. All are originally from the

Americas, namely the rather common and widespread

weed S. aculeatissimum, the relatively rare weed S. cap-

sicoides, the cultivated ornamental S. mammosum, and S.

viarum which is becoming a fairly common weed in

KwaZulu-Natal and neighbouring areas. S. capsicoides

and S. viarum have not been listed before for southern

Africa (Welman 1993). Section Totra Nees is represent-

ed by two species in southern Africa; both are originally

from the Americas. S. chrysotrichum is so far naturalized

only in Eastern Cape, while 5. tonnun is confined to a

fairly small area north of Durban, KwaZulu-Natal.

* National Herbarium. National Botanical Institute, Private Bag XlOl,

0001 Pretoria.

E-mail: wgw(s>nbipre.nbi.ac.za

MS. received: 2001-06-28.

Detailed descriptions and discussions are given for both

sections and all six species, together with keys, illustra-

tions and distribution maps. S. mammosum, being culti-

vated only, has no distribution map.

DESCRIPTIONS AND DISCUSSIONS

Section Acanthophora Dunal, Histoire naturelle,

medicale et economique des Solanum: 131, 218 (1813).

Lectotype species: S. mammosum L. (D’Arcy 1972).

Section Aculeata G.Don subsection Acanthophora

(Dunal) G.Don: 434 (1838).

Psilocarpa, grad, ambig., Dunal: 216 ( 1852).

Section Simplicipilum Bitter: 147 (1923). Lectotype

species: S. aculeatissimum Jacq. (Seithe 1962).

Description (based on Nee 1991)

Small shrubs, 0. 3-3.0 m tall, sprawling or more often

erect and few-branched, annuals or short-lived perenni-

als, heavily armed with straight acicular (needle-shaped)

prickles, in some species stems with additional robust

recurved prickles from enlarged compressed bases.

Stems usually glandular-puberulent, often also eglandu-

lar-pilose with few-celled, simple hairs. Leaves petiolate,

alternate or usually geminate, with one about twice the

size of the other; blades usually ovate to orbicular and

with 2-4 pairs of lobes, or repand, coarsely and irregu-

larly toothed, membranous, spiny along main veins

above and below, usually glandular-puberulent above

and usually also eglandular-pilose with simple few-

celled hairs, these sometimes with vestigial rays at base,

below with simple hairs and sessile few-rayed stellate

hairs (these lacking in S. capsicoides). Inflorescences lat-

eral, simple or few-branched, sessile or shortly peduncu-

late, remote from leaves, axis often spiny; flowers few to

many, fewer than half fertile, often only lowest fertile in

large-fruited species. Calyx seldom much accrescent;

lobes triangular to linear. Corolla white, yellowish, green-

2

Bothalia33,l (2003)

ish or blue-puiple, glandular and pilose on outside, stel-

late, without interstitial pleating. Ovary glabrous or

minutely stipitate-glandular, puberulent in few species

with simple hairs that drop off early. Fruit usually glo-

bose, small to large, strongly dark green and pale green

marbled when young, yellowish, orange or reddish at

maturity, glabrous at maturity, exocarp ± tough, meso-

carp juicy, in some species becoming dry at full maturi-

ty, placentae simply columnar, divided or variously

intruded, the fruit becoming effectively or completely

unilocular at maturity. Seeds numerous, subreniform to

orbicular, seed coat closely investing endosperm or

extending around periphery of seed into a flattened wing.

Distribution and ecology

According to Whalen (1984), section Acanthophora

occurs throughout the American tropics except in

Amazonian Brazil; it is most diverse in the northern

Andes and in southeastern Brazil. About 20 species grow

in open savanna and disturbed sites. Several species are

established in the Old World tropics.

Nee (1979) noted that this section, in common with

almost all groups of Solanum, has great weedy tenden-

cies; the species are found in areas of high light intensi-

ties and also where the natural vegetation has been dis-

turbed. Under natural circumstances, landslides, eroded

stream banks and forest openings provide suitable habi-

tats. Man’s agricultural and industrial practices are still

expanding the ranges of numerous species of Solanum.

The succulent berries of many species are popular with

birds and ensure seed dispersal.

Taxonomy

Dunal (1813) appropriately named this group of

prickly species section Acanthophora (thorn-bearing). In

1923, Bitter referred to the characteristic hairs of this

section in the name Simplicipilum. However, Dunal’s

name for this section must take precedence.

In 1972, D’Arcy elevated subsection Lasiocarpum

Dunal of section Acanthophora to section Lasiocarpum

(Dunal) D’Arcy, with S. lasiocarpum Dunal as type

species. Section Acanthophora can be distinguished

from section Lasiocarpum by the lack of stellate hairs on

the ovaries, the more general coverage with multi-celled

simple hairs and the less substantial habit (Jaeger 1985).

Section Lasiocarpum is indigenous to South America

and no representatives have so far been recorded from

southern Africa, though a few species are naturalized

and/or cultivated in East Africa; see Jaeger (1985).

Whalen (1984) placed S. aculeatissimum, S. capsicoi-

des, S. mammosum and S. viarum in his 5. mammosum

Group.

Microscope studies

Section Acanthophora has distinctive simple hairs

that are long, multicelled, uni,seriate, shiny and translu-

cent. These simple hairs are identical with the central ray

of the stellate hairs on the same plant; the latter often

only present on juvenile plants or leaves. During evolu-

tion the basal side rays of the stellate hairs have appar-

ently disappeared. Single-celled simple hairs, often

gland-tipped, may also occur on the upper surface of the

leaf as well as elsewhere on the plant (Nee 1979).

Diagnostic characters

According to Whalen (1984), the S. mammosum Group

of species is distinguished by an essentially herbaceous

habit, difoliate-geminate sympodia, vestiture of predom-

inantly simple hairs, variously lobed or dentate, prickly

leaves, lateral, often simple inflorescences, stelliform

corollas and dryish, glabrous berries.

Nee (1979) stated that section Acanthophora, as a

member of the prickly subgenus Leptostemonum (Dunal)

Bitter, deserves its name; all the species are heavily

armed with sharp, slender prickles and some species

have stouter, broad-based prickles as well. He defined

the section by the combination of the upper leaf surface

bearing only simple hairs, glabrous fruit and the non- to

only slightly accrescent calyx.

This section can be divided into two subsections

based on the seed morphology (Nee 1979). One subsec-

tion, with seeds that do not differ much from those of

almost all other Solanum species, includes S. aculeatissi-

mum, S. mammosum and S. viarum. The other subsection

is characterized by seeds surrounded by a broad, flat-

tened margin or wing, a character unique in Solanum.

This subsection is probably monophyletic and includes

S. capsicoides. The evolutionary significance of the

winged seed is not clear; it could aid in wind or water

dispersal.

Sex forms and branching

Whalen & Costich (1986) described section Acan-

thophora as weakly to strongly andromonoecious; having

both perfect and male flowers, but no female flowers.

Species of this section have some short-styled flowers,

and are self-compatible. The fruits are small to large; it

was found that andromonoecy in section Acanthophora

is most strongly developed in large-fruited species.

Child (1979) and Child & Lester (1991) studied the

branching patterns in the Solanaceae. All sections of the

subgenus Leptostemonum have 2- or 3-foliate sympodial

units often with the subtending leaf paired with the small-

er first leaf of the daughter shoot (anisophyllous geminate

sympodia). Section Acanthophora has mostly simple

few-flowered inflorescences with only the basalmost

flower and/or the few proximal flowers of the subsidiary

inflorescence hermaphrodite; the distal flowers are usual-

ly smaller, brachystylous or short-styled (andromonoe-

cious; Symon 1979) and are often inserted with a spatial

gap from the hermaphrodite flowers. Species with prick-

ly calyces in the hermaphrodite flowers usually have

unarmed, functionally male, brachystylous flowers.

Note

Nee (1991) noted that this section contains a number

of promising candidates for screening for alkaloids.

Bothalia 33.1 (2003)

3

Key to species of section Acanthophora in southern

Africa (from Jaeger 1985)

la Seeds 4-6 mm diam., prominently winged; branches and

leaves almost glabrous to sparsely hairy; ripe fruit Ver-

million (sometimes cultivated) S. capsicoides

lb Seeds 2-5 mm diam., not winged; branches and lower leaf

surface hairy; ripe fruit yellow:

2a Stems with acicular prickles only; branches with scattered

simple hairs S. acideatissimiim

2b Stems prickly with both compressed and recurved and

long, straight acicular prickles; branches hirsute with

dense, simple hairs:

3a Hairs on stem of variable lengths, 2^ mm long; calyx

not prickly; corolla violet; fruit globose, usually

with an apical nipple and several basal protuber-

ances; seeds 5 mm diam. (cultivated only)

S. mammosum

3b Hairs on stern of uniform lengths, up to 5 mm long; calyx

prickly; corolla white; fruit globose; seeds ± 2 mm

diam S. viarum

1 . Solanum aculeatissimum Jacq., Collectanea aus-

triaca ad botanicum 1: 100 (1787a); Jacq.: t. 41 (1787b);

Bitter: 148 (1923); Heine; 334 ( 1963); Pearse: 173(1978);

Gbile: 115 (1979); Jaeger: 478 (1985); Bukenya & Hall: 84

(1988); Nee: 265 (1991); Bukenya & Carasco: 51 (1995);

Relief & P.P.J. Herman: 622 (1997); Shaw: 234, 239

(2000). Type: cultivated plant at Vienna, ‘Patria in zona tor-

rida est’, Jacquin s.n. (W).

S. reflexion Schrank: t. 81 (1822). Type: cultivated plant in Hortus

Monacensis (Munich), von Schrank s.n. (M).

S. khasianum C.B. Clarke: 234 (1883). Type: India, Khasia Mts.

Nunklow. J. D. Hooker & TThomson 14 (CAL).

S. cavaleriei Leveille & Vaniot: 207 (1908). Type: from China.

J.Cavalerie 2722 (E).

Annual, sparsely branched herb or undershrub up to

1 .5 m high, pubescence highly variable, dense to sparse

(plant rarely nearly glabrous). Stef7is tinged purple and

green, rarely densely pubescent, with simple, uniseriate,

multicellular hairs of unequal lengths, up to 4 mm long,

heavily armed with many slender, acicular, straight,

spreading or slightly reflexed prickles up to 12 mm long

with base 0.5 mm broad. Leaves ovate, up to 160 x 180

mm, base truncate or slightly cordate, lobation variable,

usually doubly or dentately lobed to 'A width of leaf,

with 3 or 4 pairs of major acute, triangular lobes, middle

lobe up to 55 mm long, dark green when fresh, usually

drying dull dark green, with simple pilose hairs on both

surfaces of leaves, rarely with stellate hairs on underside,

prickles on midrib and petiole slender, ± 15 mm long,

base ± 2 mm broad, prickles on primary lateral veins

slender, ± 7 mm long, purple near base, upper part yel-

low, the base 0.8 mm broad; petiole 4-80 mm long.

Inflorescences simple, 3-5(6)-flowered; peduncles up to

0.5 mm long; pedicels ± 10 mm long. Calyx: lobes often

unequal, elongating immediately after anthesis and con-

cealing immature fruits with subcaudate lobe tips,

pubescent, becoming prickly in fruit. Corolla 10-15 x

25-30 mm, stellate, white, greenish or yellow-green,

sometimes with a purple tint, rarely with purplish hairs.

Stamens: filaments green to yellow; anthers yellow. Style

white; pistil pale green. Fruit nodding, globose, 15-30

mm diam., smooth, pale green with medium green

stripes or white with green reticulation when young,

dirty yellow or brownish when ripe. Seeds 2-4 mm

diam., brownish, moderately flattened, seed coat closely

investing endosperm, with margin sometimes thickened

but not wing-like. Figure 1 .

Distribution

Whalen (1984) gave the distribution of this species as

southea.stem Brazil (natural) and Central Africa (natural-

ized). Jaeger (1985) stated: The wide distribution of this

species in Africa, and its scarcity or even absence from the

New World, together suggest that S. aculeatissimum is

native to Africa. This gives section Acanthophora an inex-

plicably disjunct distribution. However, S. aculeatissimum

grows as a weed of secondary vegetation, indicating that it

is more likely to be an introduction to Africa; and when it

is considered that trade routes between the New World and

Africa have been established for over 400 years (Roe

1979), an early introduction of this species would allow

time for its subsequent dispersal. Its hideously spiny habit

must deter grazing and contribute to its success.’

Jaeger & Hepper ( 1986) confirmed that S. aculeatissi-

mum seems to have been in Africa for a long time, hav-

ing achieved a very wide distribution in the Afromontane

regions in western, eastern and southern Africa. Nee

(1991) reported that S. aculeatissimum occurs naturally

mainly from Rio Grande do Sul, Parana and Santa

Catarina states in Brazil. He also reported that this

species has long been grown in European botanical gar-

dens where it is very uniform and may represent a single

introduction from South African populations, which they

strongly resemble.

Because of confusion with S. capsicoides and other

heavily armed species of Solanum, distribution records

of S. aculeatissimum in the literature are often not reli-

able. In southern Africa, this species is fairly common in

Limpopo [Northern Province], Mpumalanga, KwaZulu-

Natal, Eastern Cape and rare in the Free State, Lesotho

and Western Cape, with one isolated record in the North-

West almost on the border with Gauteng (Figure 2). It is

also sometimes cultivated as a curiosity in these areas,

e.g. in botanical gardens; Forbes NH34644 (NH) was

grown at the Botanical Station, Durban in 1944.

Ecology

Nee (1991) recorded that S. aculeatissimum is a weedy

shrub that grows in disturbed or pastured forest, in grazing

fields and along roadsides, from 400-1 200 m altitude. Tn

sub-Saharan Africa it is found at forest edges and in savan-

nas, mostly in the eastern uplands of the great lakes region,

but scattered through the rain forest and into South Africa,

from 1 000-2 400 m, rarely descending almost to sea

level.’ Gbile ( 1979) described this as a highland species in

Nigeria that grows mostly on rough ground and amongst

rocks. According to Bukenya & Carasco (1995), S. aculea-

tissimum generally grows in forest clearings in Uganda;

according to Bukenya & Hall (1988), it is a rare species in

secondary forest in western Ghana.

In southern Africa, S. aculeatissimum grows in forest

margins and clearings, along forest paths, and among

undergrowth in closed natural forest, but also in Finns and

4

Bothalia 33,1 (2003)

FIGURE 1 . — A, Solcmum aculeatissi-

mum, Acocks 10787, fruit

Nienaber 9 (PRE); B, S. tor-

vum, Henderson 737 (PRE).

Artist: G. Condy.

Eucalyptus plantations. It prefers damp, shady places

under shrubs and trees. It is also found along roadsides and

in other disturbed areas, as a weed in gardens, ploughed

fields as well as grassland. Wells et al. (1986) listed S.

aculeatissimurn as a ruderal, silvicultural and pastoral

weed of the temperate and subtropical summer rainfall

areas in southern Africa. It is found on gentle and steep

mountain and hill slopes, in gorges and valleys, but also on

flats; it can grow on moist humus-rich and red-brown

loamy soil and also on well-drained sandy or stony soil. It

is common in mountainous areas and is recorded from

about sea level to ± 2 200 m altitude, with a rainfall of up

to 2 000 mm annually. Acocks (1988) listed S. aculeatissi-

miiin as one of the generally occurring smaller plants of

the forest floor and margin in his Northeastern Mountain

Sourveld which is one of the Inland Tropical Forest Types.

He also listed it as one of the smaller plants of general

occurrence in the Highland .Sourveld which is one of the

Temperate and Transitional Forest and Scrub Types.

Nee (1991 ) stated that in Brazil, its natural distribu-

tion area, flowering is confined from October to

February, similarly in East Africa, but throughout the

year in the Democratic Republic of the Congo. In south-

FIGURE 2. — Distribution of Solanum acideatisshmim in the FSA

region.

Bothalia 33.1 (2003)

5

em Africa it flowers from about November to March and

fruits from about November to April.

Chromosome number and chemistry

Both Fedorov (1969) and Bukenya-Ziraba ( 1996) list-

ed 2n = 24.

Dalziel (1937) reported that the stem contains sola-

nine, an alkaloidal glucoside with physiological action

like saponin, but much less poisonous. Watt & Breyer-

Brandwijk (1962) also reported that S. aculeatissimum

contains solanine. The immature fruit is said to contain

more solanine than the mature. Reports that the plant is

toxic to cattle, horses and even humans are suspect as

they may not be referring to the true 5. aculeatissimum.

Hepper (1978) stated that the steroidal alkaloid solaso-

dine always occurs in the fruits of S. aculeatissimum, but

little is found in the leaves. Roddick (1986) gave figures

for the amounts of solasodine in the leaves and fruit, con-

firming Flepper. Hutchings et al. (1996) reported that

solasodine, solamargine and solasonine have been isolat-

ed from leaves, stems, fruit pericarps and seeds. The

highest concentration of glycoalkaloids was found in

seeds, followed by fruit pericarps, leaves and stems.

Solasonine accounted for about 80% of the total glyco-

alkaloids, whereas solamargine accounted for about

13%, and was not found in stems.

Medicinal and other uses

Mehra (1979) reported that in Malaysia the leaf juice

of S. aculeatissimum is used in a ritual preparation to be

taken at childbirth. In Puerto Rico the sliced fruit of S.

aculeatissimum (or plants known as) is fatal when con-

sumed by cockroaches (Mabberley 1997). In Liberia, the

fluid prepared by boiling the fruits of S. aculeatissimum

is applied in an enema for constipation (Dalziel 1937).

Gbile & Adesina (1988) reported the same for Nigeria. In

West Africa the fruit and the root are used as remedies for

cough and dysmynorrhoea.

Hutchings et al. ( 1996) reported that in Transkei, South

Africa, the ripe fruit is applied to the forehead for

headaches and is used for healing the navels of new-born

babies, and smoke from burning plants is used for

toothache (also in Nepal); root decoctions are taken for

backache and impotence. An infusion of the root is a Zulu

snake-bite remedy (Watt & Breyer-Brandwijk 1962). In

Eastern Cape, South Africa, it was noted on the voucher,

Bradley 49 (GRA), that the poisonous fruit is used for skin

infections and the root for flatulence and as a purgative.

Microscope studies

Gbile & Sowunmi (1979) described the pollen of S.

aculeatissimum as subprolate and triangular, polar axis ±

28.0 pm, equatorial diameter ± 22.5 pm, with the sexine

pattern faintly distinct. Nee (1986) investigated the pla-

centation pattern of S. aculeatissimum. The placenta is

H-shaped in cross section, allowing the seeds to fill the

total interior of the large fruit. At maturity the septum is

a very thin film, resulting in a more or less unilocular

berry. The ‘ribs’ running down the sides of the inner peri-

carp at the carpellary midvein strongly suggest traces of

the false septum of S. sisymbriifolium Lam., but does not

indicate a close relationship. Gbile (1986) observed

straight upper epidermal walls in the leaves of S. aculea-

tissimum, which is an uncommon occurrence in a meso-

morphic montane species. Groth (1989) studied the seeds

of S. aculeatissimum-, they are ellipsoid or subdiscoid

with a finely punctate surface. The embryo is circinate

and the cotyledons lanceolate. Ogundipe & Daramola

( 1997) also investigated the leaf epidermis: the anticlinal

cell walls have a wavy pattern, the adaxial cells (± 38.0

X 15.5 pm) have an iiTegular and the abaxial cells (± 33

X 15 pm) an isodiametric cell shape; the abaxial stomata

(± 28.5 X 17.0 pm) are anomocytic.

Taxonomy and diagnostic characters

S. aculeatissimum (seeds not winged) has often been

and still is confused with S. capsicoides (= S. ciliatum

Lam.) (seeds winged) e.g. in Wright (1904); Wright (1906);

Bailey & Bailey (1977); Mabberley ( 1997). D’ Arcy (1972)

chose S. aculeatissimum as the lectotype species of Ar-

matae (subgenus?) and of Macrophyllae (section?) and of

Pannflorae (subsection?), all of Wright (1906).

In southern Africa, S. aculeatissimum can be distin-

guished from related species by its yellow ripe fruit,

seeds without wings and by its stems with scattered sim-

ple hairs and acicular prickles.

Potential for crop improvement

Several investigators have looked at hybridization

with S. melongena L. (aubergine, brinjal, eggplant). Rao

(1979) reported successful crosses between S. melongena

and S. aculeatissimum. Pearce & Lester (1979) showed

that S. acideatissimum has very little serological rela-

tionship with the taxa of the eggplant complex. Hybrids

produced seed which did not germinate. Daunay, Lester

& Laterrot (1991) recorded that S. aculeatissimum is

resistant to Verticillium, but susceptible to Meloidogyne

spp., which are both eggplant pests.

Notes

The specific epithet refers to the large number of

sharp prickles. Numerous common names have been

recorded e.g. Bailey & Bailey (1977): cockroach berry,

love apple, soda apple nightshade; Wells et al. (1986):

apple-of-Sodom and devil’s apple for southern Africa;

Jeffery RUH4919 (GRA): Peru apple.

The photograph and description published by Rhind

(1975) in a popular journal for South African farmers,

belong to S. aculeatissimum and not to the indigenous

S. panduriforme E.Mey., as claimed in the discussion

accompanying the photograph.

2. Solarium capsicoides Allioni, Auctuarium ad

synopsim methodicam stirpium horti regii Taurinensis:

64 (1773); Dunal: 156 (1852); Symon: 101, t. 29 (1981);

Symon: 118, t. 26A, B (1982); Whalen: 252, t. 28 (1984);

Carvalho: 77 (1985); Jaeger: 477 (1985); Troupin: 378, t.

6

Bothalia 33,1 (2003)

121.1 (1985); Bukenya & Hall: 84 (1988); Lemke: 365

(1991); Nee: 261 (1991 ). Type: cultivated at Turin, origin

unknown, Allioni s.n. (TO, holo.).

S. ciliatum Lam.: 21 ( 1794). Type: sheet with label 'S. ciliatum Lam.

illustr. Die. No. 55’ (P-LA).

S. aculeatissimum auett. non Jacq., C.H. Wright: 97 (1904).

S. macowanii Fourc.: 101 ( 1934). Type: Eastern Cape, Humansdorp

Div., Fourcade 1224 (SAM, holo,; K).

Annual or short-lived perennial, herbaceous shrub, erect

to sprawling, occasionally woody at base, sometimes

spreading by rbizomes, up to 1 .5 m high. Stems green or

purplish; stems, petioles, veins of upper and lower leaf sur-

faces, pedicels and calyces bearing scattered to many

prickles, very sharp, straight, pale or straw-coloured, ± nar-

rowly (up to 2 mm) based, 2-14 mm long; sparsely pilose

with 1 .5-5.0 mm long, simple, shiny, uniseriate, few-celled

hairs (stellate hairs absent) and minute, simple, glandular

hairs. Leaves membranous or papery, concolorous green,

broadly ovate in outline, up to 150 x 135 mm, 5-7-lobed,

slightly cordate at base, sinuses reaching about halfway or

less to midrib, lobes and leaf apex acute or obtuse, major

FIGURE 4. — Distribution in the FSA region, of Solanuw capsicoides,

O; and S. viantm, •; (both species 3).

Bothalia 33,1 (2003)

7

lobes often entire or slightly repand but scarcely pinnately

lobed, margins ciliate; petiole up to 80 mm long. Flowers

1-7 together on short peduncle, 2-4 mm long, or pedicel-

late on stem in intemodal position; first flowers staminate;

pedicels 10-25 mm long at anthesis. Calyx often prickly;

tube 2-5 mm long; lobes 2-5 mm long, broadly lanceolate

or triangular-ovate, acute, slightly accrescent. Corolla

10-15 X 20-30 mm, deeply stellate with lobes ± 10 mm

long, glabrous inside and outside, white or rarely pale

mauve. Stamens subequal; filaments 1-2 mm long,

glabrous; anthers 3-7 mm long, tapered upwards, erect in

cone, pale yellow; pores minute; pollen 24.0-30.5 pm.

Ovary with some glandular hairs; style 5-8 mm long, erect,

pale, styles of perfect flowers exserted, those of staminate

flowers equalling filaments; stigma green. Fruit slightly

depressed-globular, 20-40 mm diam., bright, matte orange-

scarlet when mature, flesh thick, white, spongy, sweetish

tasting (tasteless?), dryish when ripe, containing ± 300

seeds. Seeds orbicular-ovoid, 4—6 mm diam., flat, minutely

pitted, reticulate, area over embryo pale yellowish or light

brown, bordered by a distinct, pale wing ± 1 .5 mm wide.

Cotyledons broadly ovate-lanceolate, ± 13 x 7 mm, first

true leaves almost orbicular, petioles without spines, later

leaves increasingly lobed. Figure 3.

Distribution

The native range is ‘coastal Brazil south of the Amazon’

(Nee 1979). S. capsicoides is adventive on the Caribbean

Islands, in Central and South America, also in the southern

United States. It is a common weed in southern China and

elsewhere in Asia; also in New South Wales and Queens-

land in Austraha where it is called devil’s apple.

Solanum capsicoides is introduced in tropical and

South Africa (probably as an ornamental) where it is so

far found naturalized only around Durban in KwaZulu-

Natal and in Eastern Cape (Figure 4). It was first collect-

ed in the Durban District in 1895, J.M. Wood 5718 (K,

NH). It was cultivated at the Natal Herbarium in Durban

in 1933, McClean NH26927 (NH), and in a garden in

Cape Town in 1955 (seed from KwaZulu-Natal), Primos

SAM684203 (SAM).

Ecology

Solanum capsicoides is native to coastal Brazil near

Rio de Janeiro and is apparently adapted to the restinga

vegetation of the coastal sand dunes (Nee 1979). It

prefers high rainfall, coastal lowlands of tropical and

subtropical regions. It becomes a naturalized weed in

disturbed sites, clearings, forest margins and other open

habitats. It seems to have a wide ecological range and

Morton (1976) claimed that S. capsicoides (S. ciliatiim)

is hardy as far north as Baltimore, Maryland in the USA.

Man is probably responsible for its initial wide dissemi-

nation; it has been in European botanical gardens for

more than two centuries.

For some unknown reason it has spread fairly slowly

in Africa. In KwaZulu-Natal it was collected at ± 15 m

altitude in shade at the edge of a forest in stony sand on

a disturbed site. It seems to flower and fruit throughout

the year.

Chromosome number, toxic properties and potential for

crop improvement

Fedorov; 695 ( 1969) reported 2n = 24 (as S. ciliatum).

The berries were reported to be poisonous to calves in

Australia (Symon 1982) and Nee (1991) noted that the

fruit is considered to be poisonous in northeastern Brazil.

The fruit of S. capsicoides is used as a cockroach poison

in parts of tropical America and this plant is sometimes

called cockroach berry in the United States (D’Arcy

1974). Its toxicity to humans is unknown. According to

Daunay, Lester & Laterrot ( 1991 ), 5. capsicoides is resis-

tant to Pseudomonas and is a non-host plant for potato

cyst-nematodes. They also reported that S. capsicoides is

graft-compatible with S. melongena, but is not used as a

rootstock as it contains toxic alkaloids.

Taxonomy and diagnostic characters

This species has often been confused with the quite

distinct S. aculeatissimum which has neither winged

seeds nor vermillion fruit. It was also previously known

under the synonym, S. ciliatum (Hepper 1979). Other

synonyms are listed by Lebrun & Stork (1997) and

Whalen (1984). Nee (1979) believed that this species,

which has been known and grown for a long time and has

been described several times under various names, might

have an older name which could emerge from some

obscure publication. He also noted that the slightly

accrescent, stoutly prickly calyx of S. capsicoides might

indicate a relationship with S. sisymbriifolium Lam. of

section Cryptocarpinn Dunal.

Fourcade collected the type of his S. macowanii at

700 ft (± 235 m) altitude in the forest near Storms

River, Humansdorp Div., Eastern Cape, in April 1921,

and quoted another specimen. MacOwan 1493, from

the Somerset East Div., also in Eastern Cape. He stated

that S. macowanii was close to S. acutissimum Jacq.

(sic) ‘from which it differs by the branches and the

lower surface of the leaves being glabrous, by the entire

leaf lobes, and by the flowers being geminate.’ Jaeger,

Nee and Lester all identified the isotype of S. macow-

anii in K as S. capsicoides: PRE has a colour photo-

graph of this specimen. The specimen of MacOwan

1493 in GRA has a much more spiny stem than the type

and one unripe fruit with immature seeds. Apparently S.

capsicoides has not been collected again in Eastern

Cape since 1921 and it could have died out in that area.

Ross (1972) quoted Ward 4888 as S. aculeatissimum

Jacq. in his checklist of the Flora of Noted, but this

specimen has orange-red ripe fruit and winged seeds

and is clearly S. capsicoides.

The large orange-red fruit with winged seeds and the

absence of stellate hairs make S. capsicoides unique in

the southern African context.

Notes

S. capsicoides is occasionally cultivated for its orna-

mental vermillion dry fruits which keep their colour for

a long time. [The large-fruited form common in cultiva-

Bothalia 33,1 (2003)

tion is var. macrocarpiim Hort. (Morton 1976).] This

species could have a ‘shaker’ mechanism for seed dis-

persal (Nee 1991). The ripe dry fruit splits irregularly

into 2^ lobes and releases the dry winged seeds, which

adhere neither to each other nor to the pericarp. The inner

fruit wall of the unilocular ovary is ± 4 mm thick, slight-

ly juicy and spongy; it is nearly tasteless and dries to a

virtually invisible thin film soon after the fruit begins to

split. The conspicuous fruit colour could indicate a very

recent derivation from ancestors with bird-dispersed

seeds. Nee ( 1986) noted that the placentas are split into

two ‘horns' and at maturity are attached only to the base

of the fruit. The fruit and seed dispersal method of S.

capsicoides probably is an adaptation to its natural sandy

habitat; the placentation ensures space for a great num-

ber of large seeds. The evolutionary significance of the

winged seed is not clear (Nee 1979). It could perhaps aid

in local dispersal by wind or water.

The species name refers to the resemblance of the

fruit to that of the solanaceous genus Capsicum.

3. Solanum manunosum L., Species plantamm, edn

1: 187 (1753); Dunal; 250 (1852); D’Arcy: 712 (1973);

D’Arcy: 851 (1974); Morton: 201 (1976); Bailey & Bailey:

1055 (1977); Symon: 103 (1981); Jaeger: 479 (1985);

Bukenya & Hall: 83 (1988); Bukenya & Carasco: 51 (1995);

Shaw; 234, 239 (2000). Lectotype: t. 226, fig. 1 in Plukenet,

Phytographia, pars tertia (1692), ‘Habitat in Virginia,

Barbados.’ Typotype: Herb. Sloane vol. 98, fol. 59 (BM); see

Jaeger: 479 ( 1985) and Knapp & Jarvis: 342 (1990).

Annual or short-lived perennial, herbaceous shrub up

to 2 m high. Stems and twigs white or yellow viscid-vil-

lous with 2^ mm long simple hairs, armed with scattered

yellow or green, large acicular prickles, 10-25 mm long,

which with age become flattened, recurved, woody and

brownish. Stems hollow. Leaves suborbicular to broadly

ovate, 60-150 x 120-200 mm, blade pinnatifid with 5-7

main broadly triangular lobes (each lobe ± coarsely, den-

tately lobed), apices acute, bases truncate to cordate,

sinuses rounded; both sides white- or yellow-villous,

upper surface with mainly 2^ mm long, simple glandu-

lar hairs, lower surface with sessile stellate hairs mixed

with long, simple, glandular hairs; armed on principal

veins on both sides with flattened, acicular prickles, 1-2

mm broad, up to 17 mm long; petioles 30-150 mm long,

covered with simple hairs and some straight prickles up to

13 mm long. Inflorescences lateral, extra-axillary, 1^-

flowered cymes/racemes bearing both staminate and her-

maphrodite flowers with only one flower fruiting per

inflorescence, ± sessile or peduncles up to 10 mm long;

pedicels 5-15 mm long, becoming stout and somewhat

longer in fruit, viscid-villous to lanate, sometimes armed

with few small prickles. Calyx 8-10 mm long, deeply

lobed, prickles absent; tube 2 mm long; lobes subulate-

acuminate, pubescent outside. Corolla showy, pale blue

to violet, up to 20 mm long, 40 mm wide, deeply stellate,

lobes linear-lanceolate; hirsute outside, glabrous within.

Stamens equal; filaments 1 mm long; anthers 8-12 mm

long, linear-oblong, tapering with small, apical pores.

Ovary and style glabrous. Fruit globose to broadly ovoid,

30-100 mm long, often with a 20 mm long nipple-like

contraction at apex and 1-5 rounded protrusions at base,

orange to yellow, spongy, pulp white. Seeds compressed-

lenticular, 5-7 mm wide, finely rugose, shiny, purple-red

to dark brown, lacking marginal wing. Figure 5.

Distribution and ecology

Solanum mammosum is native to the Caribbean (Nee

1979), but has spread by long distance dispersal, direct

migration or with the casual or deliberate assistance of

humans to the tropical lowlands of South and Meso-

America. It is now often naturalized in disturbed, lowland

habitats in the tropical and subtropical regions of both hemi-

spheres, though not yet in Africa (Jaeger 1985) or Australia

(Symon 1981). Nee (1991) stated that S. mammosum now

occurs mostly from sea level to 100 m altitude in tropical

America, but can occur as high as 1 800 m. It flowers and

fruits throughout the year with no special season. It can be

expected to become naturalized or to escape locally from

cultivation in areas of high rainfall, whether this is seasonal

or year-round. It has been known as a curiosity in botanical

gardens for several centuries and is occasionally cultivated

as an ornamental in tropical Africa; also in southern Africa

in frost-free or frost-protected localities.

Chromosome number and chemistry

Heiser (1971) gave references of reports for both n =

12 and n = 11; the latter was the first report of n = 11 in

the genus Solanum. Dan & Dan (1984) recorded the

occurrence of nuatigenin and iso-nuatigenin (two rare

isomeric non-nitrogenous sapogenins) as the main root

constituent of S. mammosum and some other species of

Solanum. Roddick (1986) listed 1% of solasodine (dry

weight) in the fruit of S. mammosum, while Mabberley

( 1997) referred to the molluscicidal glycoalkaloids of the

fruit. Nee (1991) stated that the immature fruits contain

the highest concentration of glycoalkaloids and that

other plant parts contain varying amounts or none.

Medicinal and other uses, potential for crop improvement

Solanum mammosum (macaw bush, nipple fmit, pig’s

ears) is widely cultivated in tropical and warm temperate

regions for its unique ornamental but also poisonous fruit.

In the Americas, these fruits are widely used for killing rats

and cockroaches, also for catching fish, extracting maggots

and curing common colds (Nee 1979). Heiser (1971) quot-

ed the use of the leaves as a narcotic in Bolivia. Daunay,

Lester & Laterrot (1991) listed that S. mammosum is resis-

tant to Fusarium wilt and several other pests and diseases

of S. melongena cultivars. They also regarded it as frost

resistant, but susceptible to Meloidogyne incognita and

Verticillium, both pests of S. melongena.

Microscope studies

The glandular villous pubescence in this species is

composed of reduced stellate hairs in which the rays are

fused with the leaf lamina and the mid-point remains as

an apparently simple hair (D’Arcy 1974). The structure

of this hair type is best observed in seedlings. Similar

hairs occur in S. capsicoides (= S. ciliatum), but they

are not so dense and are less commonly glandular.

Whalen & Costich (1986) listed a report on female

Bothalia 33,1 (2003)

9

HGURE 5. — A, Solanum nuimmoswn.

Code! 6569 (PRE); B, S. via-

niin, T. Dickers & S. Neser

s.n., grown from seed collect-

ed in Brazil (Cedara Weeds

Laboratory). Artist: G. Condy.

Sterility of short-styled flowers in S. maminosiim, and

noted that embryo sac development in these flowers

was abnormal.

Taxonomy and diagnostic characters

Whalen (1984) and Lebrun & Stork (1997) listed a

number of synonyms for this species. Wright (1904)

expressed doubt whether the S. marnmosiim Thunb. in his

revision of the South African species of Solanum, is the

same as S. mammosum L. The description does not men-

tion the unique fruit. A specimen (Thunherg s.n.) collect-

ed in the Swellendam Div. (Western Cape) in the late

1 8th century is quoted; probably before S. mammosum L.

was introduced into South Africa. However, no specimen

of it was found in Thunberg’s herbarium in 1883.

Solanum mammosum has a pubescent calyx without

prickles, a showy pale blue to violet corolla, yellow.

ripe mammiform fruit and seeds at least 5 mm wide,

lacking marginal wings. In southern Africa, S. mam-

mosum has so far only been recorded in a cultivated

state, and not yet as naturalized or even as a garden

escape.

Notes on the fruit

Miller (1969) made a thorough study of the morphol-

ogy and development of the unusual mammiform fruit of

S. mammosum. Wild specimens in the Western Hemis-

phere show considerable variation in fruit shape, from

entirely globose to mammiform with an apical nipple and

five protuberances at the base. The more unusual forms

were probably selected by man from a teratological

abnormality and used as an ornamental and for its

extremely poisonous fruit. There has apparently also

been a selection for plants less prickly than the wild form

(Nee 1991 ).

10

Bothalia 33,1 (2003)

Nee (1986) studied placentation patterns in the

Solanaceae. The basic fruit in Solamtm is 2-locular with

a slightly enlarged placental area in the centre of the sep-

tum, from which the seeds radiate into the usually pulp-

filled locules between the septum and the pericarp. In the

mature fruit of S. mammosum, the septum can hardly be

seen, and is only a thin film, easily destroyed; the pla-

centas are connected only at the base. Nee (1979) sug-

gested that the spongy mesocarp may adapt the fruit to

dispersal by water flotation.

4. Solanum viarum Diinal in DC., Prodromus sys-

tematis naturalis regni vegetabilis 13,1: 240 (1852);

Carvalho: 83 (1985); Jaeger: 482 (1985); Nee: 264

(1991); Bryson & Byrd: 382 (1994); Wunderlin et al.:

606, t. 1, 2 (1993). Type: Sao Paulo, Brazil, Lund 799 (G-

DC, holo.,-IDC microf. 2080: I.l).

S. khasianum C.B, Clarke var. chatterjeeanum Sen Gupta: 413 (1961).

Type: Nilgiri District, Madras State, India, K. Submmanyam 10413 (CAL,

holo., iso.).

Straggling, short-lived, perennial herbs or subshrubs,

1-2 m high. Stems much branched at and above base;

stems, petioles, pedicels and leaves densely glandular-

puberulent with ± uniform, multicelled, straight, simple

hairs less than 0.5 mm long, some hairs eglandular; stem

prickles few, most with thickened and flattened bases

and conspicuously recurved, up to 6 mm long. Leaves

solitary or in pairs, membranous, blades broadly ovate to

ovate-triangular, base subcordate, 80-150 x 50-150 mm,

sinuate-lobed up to '/a of way to midrib, lobes 4-6, sub-

obtuse or subacute, lower surface also with 4 or 5-rayed

stellate hairs, rays nearly always in one plane, one ray

longer than rest, up to 0.5 mm long; armed on midrib and

minor veins with straight, acicular, laterally compressed,

broad-based, whitish to yellowish prickles up to 20 mm

long; petioles 20-90 mm long, armed with similar prick-

les. Inflorescences axillary, 4- or 5-flowered; peduncles

stout, 15-20 mm long. Calyx pubescent on outside, 4-7

mm long, 5-lobed to about middle; lobes triangular-

ovate, enlarging and concealing immature fruit, then

prickly on outside. Corolla white or greenish, 15-27 mm

diam., deeply stellately 5-lobed; lobes broadly lanceo-

late, 8-12 mm long, recurved, apex acute to mucronate,

outer surface pubescent with glandular and eglandular

hairs. Stamens white to cream; filaments ± 1 mm long;

anthers linear-lanceolate, attenuate, 8-10 mm long, pores

minute. Ovary globose, densely and minutely puberu-

lent; stigma capitate. Fruit globose, 15-30 mm diam.

when mature, minutely puberulent when young, later

smooth and glabrous, immature fruit pale and dark green

with white mottling, yellow at maturity with leathery

skin surrounding thin-layered, pale green, mucilaginous,

scented pulp. Seeds numerous, 190-385, moderately

compressed, suborbicular, ± 2 mm long, lacking marginal

wing, surface reddish brown, very minutely rugulose,

seed coat closely investing the endosperm. Figure 5.

Distribution

Originally from southern Brazil, Paraguay, Uruguay

and northern Argentina, but S. viarum has spread to other

areas of South America, also Central America, the West

Indies and the southern United States (tropical soda

apple). It was introduced into India and Nepal and also

into Africa; it is recorded in the Democratic Republic of

the Congo and possibly also in Cameroon (Jaeger 1985).

Probably accidentally and sporadically introduced into

Africa and can be expected to spread to many other sub-

tropical areas (Nee 1991).

In southern Africa, specimens have been collected in

Swaziland, KwaZulu-Natal and Eastern Cape (Figure 4).

The earliest record seen for this study, was collected in

1962: Marr & Harding 54 (PRE) from the Natal Agri-

cultural Institute, Pietermaritzburg, KwaZulu-Natal.

Ecology

Wunderlin et al. (1993) reported that in Florida S.

viarum is usually found in soils belonging to the order of

Spodosols, that means nearly level, somewhat poorly

drained soils with a spodic horizon 1-2 m below ground

level. In South Africa it has been collected on well-

drained, reddish brown, sandy loam soil, with a western,

northwestern or eastern slope, from 30-800 m in alti-

tude; elsewhere it is reported from low altitudes, mostly

below 100 m. S. viarum prefers full sunlight, but will

also grow in the shade.

It is often a common weed of natural areas, improved

pastures, croplands, roadsides and fence rows, waste

places, open disturbed sites, secondary growth and forest

edges. In southern Africa it was also collected in open

woodland and pine plantations. In more tropical areas,

flowering and fruiting occurs throughout much of the

year (Nee 1991), but in southern Africa flowering and

fruiting material has been collected from November to

April, with one fruiting specimen in August.

Weedy characters, dispersal and control

Solanum viarum can produce up to 50 mature fruits

per plant, each fruit containing numerous seeds. Accord-

ing to Bryson & Byrd (1994), it has been on the Florida

noxious weed list since 1994. It was present in that state

since about 1980 and spread very rapidly and aggres-

sively, being able to form near monocultures in certain

situations, although its opportunistic adaptations may

restrict it to disturbed areas.

Its foliage is generally not grazed by cattle, probably

because it is heavily armed with prickles and glandular

hairs, although they will sometimes eat the bitter mature

fruit. It can be transported from pasture to pasture in con-

taminated hay and farm machinery. The primary means of

dispersal seems to be livestock and wildlife, including

birds. Scarification of seeds by digestive systems of ani-

mals seems to promote seed germination. Movement of

livestock that have recently fed on S. viarum fruit will has-

ten its spread. Bryson & Byrd (1994) found that in mature

plants the root systems were up to 300 mm deep and lat-

eral roots up to I m long. They also found that experi-

mentally detached green fruit more than 20 mm in diame-

ter will ripen in sunlight. Seed gemiination from these

fruits was more than 70%. Fruit from less than 10 mm in

Bothalia 33.1 (2003)

diameter up to mature yellow or dry fruit are extremely

buoyant and can then also be dispersed by water. Control

of this plant therefore requires elimination of immature

and mature fruit as well as the whole plant including the

root system.

Chromosome number, chemistry, medicinal use and poten-

tial for crop improvement

Fedorov (1969) gives 2n = 24 as the chromosome num-

ber for S. khasianum.

The section Acanthophora contains several species with

a high alkaloid content (Nee 1991 ). Of these S. viariim is the

species most studied, as it contains a high percentage of sola-

sodine, concentrated in the berries. Dan & Dan (1984) listed

S. vianim as a high solasodine yielding plant (the alkaloid of

pharmaceutical interest). Roddick ( 1986) listed 0.7-3.0% of

solasodine (dry weight) in the fruit. It is a common weed in

India since at least 1932 where it is also grown as a com-

mercial source of this glycoalkaloid which is a key interme-

diate in the synthesis of steroid drugs (Maiti et al. 1979).

Saini ( 1966) found that the ‘fleshy cover’ and washed seeds

of S. khasianum var. chatterjeeanum do not contain any

alkaloid; most is found in the mucilaginous layer around the

seeds. When fmits are dried, the mucilaginous layer dries to

a white powder which is easily lost or is capable of contam-

inating the seed or pericarp assay. This provides an explana-

tion for the discrepancies in the literature of the glycoalka-

loid content from different parts of the fmit. Daunay, Lester

& Laterrot (1991) reported that S. vianim is resistant to the

following diseases of S. melongena: Leiicinodes orbonalis,

Epilachna vigintioctopunctata, Phomopsis blight, ‘little leaf'

and partially resistant to Meloidogyne incognita. It is also a

non-host plant for potato cyst-nematodes.

Taxonomy and diagnostic characters

Morton (1976) regarded S. vianim as a synonym of S.

reflexiim Schrank. However, 5. refle.xiim is a synonym of 5.

aciileatissimiirn, a different but closely related species, see

above (Nee 1991). Lester (1978) pointed out that the descrip-

tion given by Morton for S. refle.xiim, is that of S. vianim.

Lester (1978) also pointed out that careful examination

showed that S. khasianum var. chatterjeeanum (which is

commonly, but incorrectly called simply S. khasianum ) is 5.

vianim. More synonyms are hsted by Lebrun & Stork ( 1 997 ).

A good character for distinguishing S. vianim from all

other species of section Acanthophora is the presence of

glandular hairs on the ovary in young flowers.

In the past, specimens in South African herbaria have

often been misidentified as S. aculeatissimiim, but the

combination of long straight prickles on the leaves and

petioles and short curved prickles on the stems distin-

guishes S. vianim from all other members of Solanum in

southern Africa. The density of the prickles, however, is

an unreliable taxonomic character, particularly in domes-

ticated species of Solanum subgenus Leptostemonum.

Notes

The species name probably refers to the habitat as

described on the type specimen: ‘Ad vias et circa domos

vulgare in provincia Brasiliana S. Pauli’ (Dunal 1852).

The photograph and description published by Rhind

(1969) in a popular journal for South African farmers,

belong to S. vianim and not to the indigenous S. pan-

diiriforme E.Mey. ex Dunal, as claimed in the discussion

accompanying the photograph.

Section Torva Nees in Transactions of the Linnean

Society of London 17: 37-82 ( 1 837). Type species: S. tor-

viim Sw. (D’Arcy 1972).

Description', based on Whalen (1984); Jaeger (1985);

Hepper & Jaeger ( 1986).

Large shrubs or small trees, 2-10 m high; early growth

prickly on both stems and leaves; later growth usually

unarmed or only sparsely prickly or bristly; sympodial

units difoliate, commonly with geminate leaves; cauline

pubescence of stalked or sessile stellae, varying in densi-

ty, often with reduced midpoints; cauline prickles, when

present, usually straight, subulate or broad-based, lateral-

ly compressed. Leaves petiolate, large and repand on

early growth; later leaves medium to large, ovate to

broadly ovate, 30-140 x 70-300 mm, entire, repand or

lobed, usually unarmed, stellate-pubescent on both sur-

faces, with stalked stellae below, pubescence of upper

surface very variable in structure and diagnostic of spe-

cies. Inflorescences corymbose or cymose, lateral or ter-

minal and becoming lateral by continued shoot growth,

usually unarmed, remote from leaves, asymmetrically

several- to many-branched, rarely simple, inflorescence

branches monochasial, bearing female-sterile flowers

with reduced gynoecia distally; peduncles short or well

developed; pedicels unarmed, slender and flexuous in

flower. Calyx variable, from nearly truncate to attenuate-

lobed. Corolla white or lavender, stellate-pentagonal or

less commonly deeply stelliform with narrow, nearly sep-

arate lobes. Stamens with anthers equal, slender and

tapering. Fruit: berries small to medium, 5-15 mm diam.,

glabrous, 4-locular; pericarp tough and leathery, green,

dull yellow or brown when ripe; flesh saponaceous and

sticky. Chromosome number: n = 12, rarely 24, 36.

Distribution

Whalen (1984) included some 50 species in his S.

ton’iim group, occurring from Mexico to southeastern

Brazil, with many endemics, mostly in South America.

He believed that there may also be several species

indigenous to southeastern Asia and the East Indies. Two

species are widely introduced in the Old World tropics.

Solanum ton’iim Sw., originally from Mexico, Central

America and the West Indies, is now a common intro-

duced weed throughout the tropics of both hemispheres,

but it has not been listed before for southern Africa

(Welman 1993). S. chrysotrichiim Schltdl. (=5. hispidum

auctt. non Pers.) originally from Mexico to Panama, is

now widely introduced in the Palaeotropics. In the past,

this species was often misidentified as S. hispidum Pers.,

but true 5. hispidum Pers. is restricted to the Andes of

southern Colombia, Ecuador and Peru.

Ecology

According to Whalen (1984), the habitat preferences

of section Tor\’a are variable, ranging from sea level to

12

Bothalia 33,1 (2003)

over 3 000 m altitude, often in forest edges, grazed fields,

and other places where direct sunlight is available. Most

of the species show weedy tendencies. These large

shrubs or trees are characteristic of secondary vegetation

and grazed lands throughout the montane Neotropics.

Species grow in understoreys and edges of montane and

seasonal forest, in dry tropical woodland, in savanna, and

even in subalpine elfinwoods.

Taxonomy

Bitter (1921) placed the Afro-Asian plants (series

Giganteiformia Bitter) under the Neotropical section

Torvaria Bitter (section Torva Nees). Whalen (1984)

and Jaeger (1985) claimed that section Torva is cladis-

tically not closely related to the Afro-Asian species,

the former separating the Giganteum group, the latter

keeping them together for convenience. Jaeger (1985)

emphasized the particular differences between the

neotropical Torva (lobate leaves, bifoliate geminate sym-

podial units, white flowers and firmer, larger, green to

yellow fruits) and the African endemic species of the

series Giganteiformia Bitter, raised to sectional level

as Giganteiformia (Bitter) Child in 1998. Species of

that section have plurifoliate sympodia, inflorescences

with shortened, umbelloid branches, and succulent,

red, translucent berries. Nee (1991) suggested that sec-

tion Acanthophora may have been derived from the

large section Torva.

Morphology and reproductive biology

Child (1979) noted that section Tor\’a has many-flow-

ered corymbose cymes where most flowers are hermaph-

rodite and fully fertile. Nee ( 1979) recorded that it is com-

mon in section Torva for the flowering branches to lack

prickles, whereas the lower stem or young shoots are quite

prickly. Whalen & Costich (1986) stated that section Tonm

is weakly andromonoecious, short-styled flowers are pre-

sent, fruit size is small and the species are self-compatible.

Child & Lester (1991) gave the following summary of sec-

tion Torva: trifoliate sympodial units with anisophyllous

geminate leaf clusters, inflorescences subsessile to pedun-

culate, generally lateral, leaf remote to subopposed, pleioto

monochasial, pleiochasia with most flowers hermaphro-

dite, fully fertile, but fewer-flowered inflorescences with

often only proximal flower(s) hermaphrodite, distal flowers

brachystylous, functionally male, some taxa androdi-

oecious; mostly shrubs to trees, some climbers.

Potential for crop improvement

Daunay, Lester & Laterrot (1991) noted that sterile

hybrids have been obtained by artificial crosses between

S. melongena and species of section Torva.

Key to naturalized species in southern Africa

(after Jaeger 1985)

la. Inclumcnium ot young parts ol'f-white; inllorescence axes

with simple glandular hairs; calyx lobes in flower 4 mm

long, sinuses glabrous; corolla less than 30 mm diam. . . .

S. torviim

lb. Indumentum of young parts rust-coloured; inflorescence

axes and calyx sinuses not as above; calyx lobes in

flower 7 mm long; corolla 30-50 mm diam. . . S. chrysotrichum

5. Solanum chrysotrichum Schltdl. in Linnaea 19:

304 (1847); Shaw: 234, 238 (2000). Type: Mexico, Las

Trojes, Nov., C. Scliiede 81 (HAL).

S. hispidum auctt. non Pers. (1805); D’Arcy; 703 ( 1973); Gentry &

Standley; 121 (1974); Bailey & Bailey: 1055 (1977); Gbile: 116

(1979); Symon: 113, t. 35, 150, 161 (1981); Symon: 121 (1982);

Jaeger: 346 (1985).

5. warscewiczii Hort. ex Lambertye: 429 (1865). Type: cultivated,

L. Lambertye s.n. (W).

S. pynaertii De Wild.; 437, t. 119 (1907). Type: Boma, Congo, L.

Pynaert 302 (BR?).

Evergreen shrub or small tree up to 7 m high (in South

Africa up to 3 m high), trunk up to 200 mm diam. Prickles

2-6 mm long, stout, usually sparse on stems, petioles and

veins on upper and lower leaf surfaces, flattened towards

broad, often pubescent base, straight or slightly recurved,

rarely absent. Pubescence on all parts of stellate hairs (ses-

sile or long, multi-seriate-stalked, porrect-stellate with

medium or long central cell). Stems grooved and sparsely

stellate when old, stem and young parts with ferruginous

tomentum. Leaves solitary, usually broadly ovate to ellip-

tical, ± 200-300 X ± 250^00 mm, rarely entire, margins

mostly sinuate to lobed, with 7-13 lobes, sinuses cut one

quarter of way to midrib, lobe apices obtuse, acute or

acuminate, sinuses rounded, leaf base equal or oblique,

truncate or rounded to subcordate; upper surface scabrid

and covered with appressed hairs, lower surface densely

covered with stellate hairs and nerves sparsely prickly;

petiole up to 180 mm long, terete, with (0)3-10 prickles.

Inflorescences extra-axillary, branched, dense scorpioid

cymes with 30-50 flowers; peduncles 10-25 mm long,

variable; pedicels 10-15 mm long, enlarged in fruit up to

5 mm diam. below calyx with marked corky thickening.

Calyx densely stellate-pubescent; tube 3^ mm long; lobes

linear or lanceolate, 4—6 mm long, acuminate, in fruit

broader and thicker, at first appressed, later somewhat

reflexed. Corolla white, 30-50 mm diam., stellate; lobes

10-15 mm long, sparsely stellate-pubescent along mid-

vein inside, densely stellate-pubescent outside, apices

acute. Stamens with filaments 1-3 mm long; anthers 7-9

mm long, yellow, slender, erect, not coherent, tapered,

dehiscing by small terminal pores and also longitudinally

in basal portion of anther. Ovary sometimes tuberculate,

ovary and style base sparsely glandular-pubescent; style

9.0-13.5 mm long, glabrous; stigma bluntly 2-lobed, pale

green. Fruit globose, 10-15 mm diam., at first green, later

yellowish to drab orange-yellow, brown when dry,

glabrous, glossy, hard, not particularly fleshy, mucilagi-

nous. Seeds 2-3 mm diam., flat, light brown, shiny, with-

out obvious surface pattern, 100-250 per fruit. Cotyledons

4x10 mm, oblong; petioles 5 mm long, first true leaves

ovate, scarcely lobed, not prickly until about fourth leaf.

Figure 3.

Distribution

Solanum chrysotrichum is native to tropical Central

America from southern Mexico to Costa Rica and Panama.

Bothalia 33.1 (2003)

13

FIGURE 6. — Distribution in the FSA region, of Solaiium chrysotrichum,

•; and S. tonnim. U.

It had been introduced widely in the Old World tropics as

an ornamental plant, and has now been naturalized in

many areas, such as in disturbed sites in the Brisbane

area of Queensland (Australia) where it is known as giant

devil’s fig (Symon 1982). It has been collected in West

Africa (e.g. Nigeria), Zimbabwe and Malawi and also in

southern Africa.

In southern Africa, S. chrysotrichum seems to be natu-

ralized only in Eastern Cape from about the Albany to the

Libode Districts (Figure 6). The oldest record seen in this

study dates from 1909, collected near Grahamstown (Molly

A7788, GRA). It is sometimes cultivated in gardens in that

province or persists as a garden relic, e.g. Jacot Giiillarmod

4567 (GRA); Bums, Olyott & Potts 11 (GRA).

Ecology

In its native area. S. chrysotrichum is found in moist

or wet thicket, sometimes in oak or oak-pine forests or in

fields at 1 200-2 500 m in altitude (Gentry & Standley

1974). In South Africa it grows at an altitude of ± 250 m,

in full sun and scattered along roadsides and in other dis-

turbed areas. It is also occasional on riverbanks and near

water courses, in hillside scrub and valley vegetation

where it can be found with Zantedeschia, Senecio ilici-

folius and aliens like Acacia longifolius, A. mekmoxylon

and Melia azedarach. Wells et al. (1986) described S.

chrysotrichum as a summer rainfall ruderal and flora

weed, which does not seem to spread aggressively. In

South Africa, it appears to flower almost throughout the

year, while fruiting is from October to June.

Morphology

Roe (1971) described the stem hairs of S. chryso-

trichum from Mexico as multiseriate-stalked, porrect-

stellate with rather short central rays. Gbile & Sowunmi

(1979) described the pollen of S. chrysotrichum as oblate-

spheroidal with a triangular amb and indistinct sexine

pattern. The polar axis is ± 21.6 pm and the equatorial

diameter is ± 22.0 pm.

Chromosome number and chemistry

Fedorov ( 1969) reported the chromosome number n =

12 for S. chrysotrichum (as S. hispidum).

Dan & Dan ( 1984) reported that the fruits and leaves

of S. chrysotrichum contain non-nitrogenous spirostane

sapogenins and are devoid of any alkaloid. The major

leaf constituents are the spirostane sapogenins neo-

chlorogenin and paniculogenin. The chemical consti-

tuents of the roots of S. cluysotrichum are not known.

Uses and potential for crop improvement

No reports of any medicinal use of S. chrysotrichum

were found in the literature. It is a garden subject in the

warmer parts of the world. Khan (1979) reported that S.

chrysotrichum had been crossed with the economically

important 5. melongena L. and produced fertile hybrids.

Daunay, Fester & Faterrot (1991 ) reported that S. chry-

sotrichum is resistant to Pseudomonas, Meloidogyne and

Verticillium, and partially resistant to Thielaviopsis, all

of which attack S. melongena. It is also a non-host plant

for potato-cyst nematodes.

Taxonomy

Most literature references to S. hispidum refer to S.

chrysotrichum. True S. hispidum Pers. is from the Andes;

the upper surface of the leaves is hairless or almost so

(Shaw 2000). D’Arcy (1973) reported that S. chryso-

trichum can hybridize with S. torvum in nature, at least in

Panama.

Diagnostic characters and derivation of species name

Solanum chrysotrichum differs from all other species

of Solanum in southern Africa by its coarse, reddish brown

pubescence, particularly on the calyx, stems, young parts

and main veins on the underside of the leaves. The species

name refers to the indumentum of golden (reddish brown)

hairs.

6. Solanum torvum Sw., Nova genera et species

plantarum; 47 (1788); Dunal: 260 (1852); C.H. Wright:

231 (1906); Heine: 333 (1963); D'Arcy: 708 (1973);

Gentry & Standley: 139, t. 19 ( 1974); Gbile: 118(1979);

Symon: 1 15, t. 36 ( 1981 ); Symon: 122 ( 1982); Symon:

152 (1985); Jaeger: 344 (1985); Troupin: 382 (1985);

Bukenya & Hall: 86 ( 1988); D'Arcy & Rakotozafy: 129

(1994). Type; ‘India Occidentalis’ (Jamaica), O. Swartz

s.n. (S, holo.).

S. mannii C.H. Wright: 129 (1894). Types: Gabon. Soyaux 329 (P,

syn.j; Fernando Po, Mann 55 (?K, syn.).

S. mannii C.H. Wright var. compactum C.H. Wright: 129 ( 1894). S.

torvum Sw. var. compactum (C.H. Wright) C.H. Wright: 231 (1906).

Types: Angola, Welwitsch 6105 (COI, syn.); Welwitsch 6052, 6080

(COI, syn.); Monteiro s.n. (COI, syn.).

Shrub up to 3 m high; branches terete, tomentose

with yellowish brown, rarely reddish brown, stipitate-

stellate and/or sessile-stellate hairs; usually armed with

14

Bothalia 33,1 (2003)

few straight or recurved compressed prickles which are

usually pubescent basally, up to 7 x 8 mm. Leaves ± dis-

colorous, solitary or sometimes in pairs, similar in

shape, different in size, broadly ovate to oblong or ellip-

tic, up to 180 X 250 mm, apex acuminate or acute, base

mostly unequal, obtuse to subcordate, margins subentire

to deeply lobed, with 2-4 pairs of lobes (uncommon in

our area), upper surface scabrid with dispersed stipitate-

stellate or sessile-stellate hairs, lower surface densely,

softly tomentose with mostly stipitate-stellate hairs,

veins rarely armed; petioles 10-100 mm long, softly

stellate-hairy. Inflorescences lateral and internodal,

corymbose cymes, several- to ± 50-flowered, lateral,

distal flowers with short styles, functionally male; inflores-

cence axes with simple glandular hairs; peduncles 10-30

mm long, usually 1 -branched less than 100 mm from

base, tomentose, hairs sessile-stellate and shortly stipi-

tate-stellate; pedicels ± 5 mm long, purple, drying dark,

hispid-viscid with mostly glandular simple and sessile-

stellate hairs, elongating up to 22 mm and thickening in

fruit, then sometimes recurved. Calyx purplish, 3-6 mm

long, hispid-viscid outside with glandular sessile-stel-

late hairs, glabrous within, 5-lobed; lobes 1.0-2. 5 mm

long initially, linear or subulate, calyx splitting with age,

then lobes 2-5 mm long, lanceolate to ovate-lanceolate,

apex mucronate, sinuses glabrous. Corolla white, 15-30

mm diam., sessile-stellate pubescent outside, glabrous

within, 5-lobed; lobes lanceolate, each with prominent

dark lengthwise line inside, 5-12 mm long. Stamens

with filaments 1.0-1. 5 mm long, glabrous; anthers lin-

ear-lanceolate, 6-10 mm long, often oblique basally,

tapering above and dehiscing by minute terminal pores.

Ovary globose, with short glandular, simple hairs near

apex; style 8-1 1 mm long, curved, glabrous or sparsely

glandular-hairy at base. Fruit globose, 10-15 mm diam.,

glabrous, dull, mucilaginous, produced in clusters of

few to 10, yellow-green to dirty brown when ripe, dry-

ing black. Seeds flattened-discoid, 1.5-2. 5 mm across,

buff-coloured, indistinctly reticulate, ± 350 per fruit.

Figure 1 .

Distribution and ecology

Solanum torvum is native to the West Indies and

Meso-America, but is now a common weed throughout

the humid tropics. It grows in forest areas of west, east

and central tropical Africa, also in Madagascar; in south-

ern Africa it is found in the districts of Mtunzini, Lower

Tugela, Inanda and Durban in KwaZulu-Natal (Figure

6). The oldest record seen in this study dates from 1932:

Gcdpin 11868 (PRE) was collected on the Lower Tugela

River bank, Stanger. This species was cultivated in a gar-

den in Stellenbosch in 1937 [Herre NBGl 62978 (NBG)].

In the early 1950s the Durban Botanical Station cultivat-

ed S. torvum from seed received from Buitenzorg, Java

in 1949 \ Wager NH64099, 64100 (NH)].

In its native area, S. torvum is a very common weedy

shrub of the lowlands and grows in dry or wet thickets,

often in secondary growth, just above sea level to 1 500

m. Elsewhere it can be an aggressive weedy species in

humid areas, generally at low altitudes where it becomes

common in disturbed sites such as old gardens, pastures,

roadsides and other waste places, also near human habi-

tation. It thrives well on good soils and grows quickly in

abandoned clearings and becomes common in exploited

forest reserves. It is well established along the tropical

coastline of eastern Queensland (Australia) where it is

regarded as a noxious weed (devil’s fig) and suspected of

poisoning stock (Symon 1982).

Solanum torvum is a successful colonizer because of

its weedy and opportunistic characters: it fruits plentiful-

ly and produces large numbers of seeds per fruit. The

seeds germinate quickly in sunlit conditions, showing

that dormancy is controlled by shade (Hall & Swaine

1980). Husaini & Iwo (1990) noted that in Nigeria, S.

torvum grows in damp places, in soil with a pH of 6.1

such as ferrasols, ferruginous tropical soils and lithosols.

In southern Africa S. torvum grows from about sea

level to 200 m, often on red clayey sand, along river and

stream banks, in swamp forest, open shrubland, coastal

thorny bush and secondary scrub, often in clearings and

other disturbed places. It does not seem to be an aggres-

sive weed in this area. In KwaZulu-Natal flowering and

fruiting material have been collected from April to Oc-

tober.

Morphology

Ahmad (1964) noted that the stomata on the lower

leaf surface are 27 x 22 pm in size. The epidermal cell

walls on the lower leaf surface are sinuous, whereas

those on the upper surface are slightly sinuous. Ogundipe

& Daramola (1997) found that the stomata are narrowly

elliptical in surface view outline; stomata are present on

both leaf surfaces. They also found that the abaxial and

adaxial cells are both irregular in shape. The abaxial anti-

clinal cell walls are wavy while the adaxial anticlinal cell

walls are wavy-sinuous. The outer periclinal walls are

flat. The abaxial and adaxial cells are both ± 38 x 19 pm

in size.

Ahmad (1964) noted that the hairs on both the upper

and lower leaf surfaces are of two types: firstly unicel-

lular and dagger-shaped and secondly stellate with an

elongated many-celled, multi-seriate stalk bearing 4-9

rays. Bukenya & Hall (1988) found that the young

leaves have dense, stalked, stellate hairs on both sur-

faces; mature leaves have more or less sessile, stellate

hairs on the upper surface, 4-6-armed with one arm

much longer than the rest, whereas the lower surface has

stalked, stellate hairs of 8 or 9 more or less equal arms.

Gbile & Sowunmi (1979) described the pollen of S.

torvum as subprolate with a circular amb and distinct

sexine pattern. The polar axis is ± 25.9 pm and the equa-

torial diameter is ± 21.2 pm.

Chromosome number and chemistry

Eedorov ( 1 969) and Daunay, Lester & Laterrot (1991)

reported a chromosome number of n = 12. Randell &

Symon (1976) reported n = 24; this was confirmed by

Husaini & Iwo (1990).

The roots and leaves of this species yield steroid alka-

loids of the 3-aminospirostane group (Schreiber 1979).

Bothalia 33.1 (2003)

15

Dan & Dan ( 1984) reported that the major leaf constituents

of S. ton’iirn are the spirostane sapogenins neochlorogenin

and paniculogenin. The leaves, fruits and roots contain

non-alkaloidal as well as alkaloidal sapogenins. The roots

contain jurubidine which is a 3-amino-spirosolane alka-

loid. Jaeger (1985) noted that the steroidal alkaloids sola-

sodine and jurubidine as well as the sapogenins chloro-

genin and neochlorogenin/paniculogenin had been record-

ed for S. ton'um. Mabberley (1997) listed this species as

one of the commercial sources of solasodine.

Culinary uses

Jain & Borthakur ( 1986) noted that in India the ripe

fruits of S. tonnim are eaten as a vegetable. Bukenya &

Hall ( 1988) reported that the fruits are collected in some

parts of Ghana and used in soup and stew. D’Arcy &

Rakotozafy ( 1994) reported that this species is cultivated

in countries like India and Myanmar for its fruits which

are used as seasoning. Mabberley ( 1997) noted that in Sri

Lanka the fruits are used in curry after the toxic seeds

have been removed.

Medicinal and traditional uses

Jain & Borthakur (1986) recorded that in India S.

torx’um alone or in combination with other medicinal

plants, is used for chest pain (fruits), snake bite and

insect stings (leaves), stomach pain (fruits and roots),

antidote to poisoning (roots), malaria (bark), night blind-

ness (fruits), tooth infections (fruits). Lu (1986) recorded

that in China the roots of this species, containing juru-

bidine, jurustune and other compounds, are used to ‘dis-

perse extravasated blood,’ relieve pain and alleviate

coughs. Gbile & Adesina ( 1988) noted that the ripe fruits

are used for liver and spleen complaints; the fruits are

expectorant and sedative. Roddick (1991) listed S.

tor\’um as an antidote for various poisons, toothache,

snake and insect bites and as an ingredient in malarial

preparations in India and as a cough medicine and gen-

eral pain killer in China.

Potential for crop improvement

Solanum tor\mm is graft-compatible with S. melonge-

na and is resistant to Pseudomonas, Meloidogyne,

Verticillium, Thielaviopsis, Phytophthora parasitica and

the Phytophthora and Fusarium solani complex. It is

resistant to Epilachna vigintioctopunctata and to little

leaf disease, but highly susceptible to Colletotrichum

gloeosporioides. It is a non-host plant for potato cyst-

nematodes (Daunay et al. 1991). In Cuba 5. laciniatum

Alton, which is cultivated as an important source of the

alkaloid solasodine, has been grafted onto S. torxmm

(Esquivel & Hammer 1991).

Taxonomy

Gooding (1965) rejected the name S. tor\nim Sw. as an

illegitimate and superfluous substitute by Swartz for S.

indicum L. and replaced it with the later name S.ficifoli-

um Ortega. However, Heine (1976) and Hepper (1978)

pointed out that Swartz realized that Linnaeus had

included both Old and New World elements under his

S. indicum references. Swartz then grouped the West

Indian element under S. torvum, which is a legitimate

name, with his own material as type.

Solanum torxmm has become naturalized widely and

as a result, numerous synonyms have been recorded

(Bitter 1921; Gentry & Standley 1974; Symon 1981;

Lebrun & Stork 1997). Only synonyms based on African

types are listed in the present article. The fact that the

leaves of this species present a great range of leaf size

and lobing, depending on age and vigour, also caused

several superfluous varieties to be described. In Central

America it seems to hybridize with several other similar

species (D’Arcy 1974); this makes the taxonomy of the

section complicated.

Wright (1904) listed 5. ferrugineum for the Durban

area in KwaZulu-Natal, but described the fruit as black;

Bitter (1921) believed that he was not dealing with the

true S. ferrugineum. The specimens quoted were not seen

in this study. Wright (1906) listed S. ferrugineum as a

synonym of S. torvum, but Bitter (1921) disagreed with

that, as he regarded these two species as related but dif-

ferent. D'Arcy (1973) also regarded S. ferrugineum as a

synonym of S. torvum, but Nee (2001) viewed them as

two distinct species.

Notes on stx'lar heteromorphism

Hossain (1973) studied stylar heteromorphism in S.

torxmm, the term used when two forms of flower are

noticeable in each inflorescence, a character seen in sev-

eral species of Solanum. In one form, the style is long

and distinctly exserted; in the other, it is short and includ-

ed within the conically connivent anthers, so that it is not

visible from outside in an open flower. Hossain noted

that in S. torxmm the later-formed distal flowers have

slightly reduced ovaries and much shortened styles, and

so act as male flowers, forming a small proportion of the

inflorescence. Only the lower flowers in any inflores-

cence are structurally hermaphrodite which explains the

characteristic incomplete fruit set in this species.

Diagnostic characters and derivation of species name

In southern Africa S. torxmm can be recognized by its

rough, hairy leaves and its many-flowered inflorescences

with fairly large white flowers and numerous large yel-

lowish berries, drying black. Sterile specimens can be

confused with S. anguivi Lam., but S. torxmm has dense,

fairly short-rayed stellate hairs on both sides of the leaf,

whereas S. anguivi has fewer stellate hairs each with a

single long ray or midpoint. Symon (1985) stated that

since the specific epithet means savage, gloomy, grim

and fierce, it could refer to the drab aspect of the plant or

the stout prickles on the stems.

SPECtMENS EXAMINED

(southern Africa only)

Specimens held at PRE, unless otherwise indicated.

The numbers in brackets indicate the identity of the

specimens; (1) S. aculeatissimunr, (2) S. capsicoides; (3)

S. mammosunr, (4) S. viaruim, (5) S. chrysotrichum', (6) S.

torxmm.

16

Bothalia 33,1 (2003)

Acocks 10787 (1) NH, PRE. Acocks & Hafstrom 1375 (1). Arnold

NBG109203 (4) NBG; NBG109367 (5) NBG.

Barker 8253 (1) NBG. Bond A15 (5) GRA. Bos 1047 (1). Bottomley

s.n. (3). Bradley 49 (\) GRA. Braun 1843 (4). Bredenkamp 1430 (1).

Brink 276 (5) GRA, PRE; 793 (5) GRA. Burns, Olyott & Potts 11 (5)

GRA.

Carbiener PRE41371 (2). Codd 6569 0). Connell PRE41379 ( 1 ). Cooper

1147 (\). Crawford 441 (1).

Devenish 1001. 1789 (\). De Villiers NBG 1 0502 0) NBG. De Winter

8251 A ( 1 ). Dieterlen 887 (\) NBG, NH, PRE, SAM.

Eckhardt 465 (\) PRU.

Flanagan 1165 (1) GRA, PRE. Forbes NH34644 (1) NH. Forbes &

Obermeyer 64 (2) NH, PRE. Forester 2(1). Fourcade 992 ( 1 ) GRA.

Fraser & Burns 21 (5) GRA. Froneman 8 (4) Lowveld Botanical

Garden.

Galpin 11868 (6) NH, PRE; 14539 ( 1 ). Garrett 69 (4) NH. Geldenhuys

668 (1). Groenewald 146 (4) NH. Gerstner 5660 (1). Giffen s.n. (1)

GRA. PRE.

Harding 39N, PRE819 (1). Henderson 737 ( 6); 940 (5). Herre

NBG162978 (6) NBG. Hewitt A7789 ( 5) GRA. Hilliard 8229 (1).

Hilliard & Bunt 15098 (1). Hoole N6 (5) GRA; s.n. (1) GRA, PRE.

Hulley s.n. (5); A7790 (5) GRA; 22 (4) NH, PRE; 40. 46. 135, 232 (4)

Cedara Weeds Laboratory; 44 (1) GRA, PRE; 159 (1) Cedara Weeds

Laboratory; 231 (1) GRA. Hulley & Dickers 284 (4) Cedara Weeds

Laboratory, GRA; 335 (4) Cedara Weeds Laboratory. Huntley 464 ( 1 ).

Hutchinson, Forbes & Verdoorn 74 ( 1 ).

Jacot Guillarmod 4567, 6836 (5) GRA; 7904 (5) GRA, PRE; Jeffery

RUH4919(\)GKN.

MacDonald 25 (5). MacOwan 1424 (1) SAM; 1493 (2) GRA. Mally

A7788 (5) GRA. Marr & Harding 54 (4) NH, PRE. Martin

RUH8014(c) (5) GRA. Matthews 27 (1) PRU. McCallum PRE41368

(1). McClean NH26927 (2) NH. McClean & Bayer 55 (1). McDonald

& Smit 6(1) PRU. Moffett, Daemane & Taoana 5313 ( 1 ) PRE, QWA.

Mogg 1019 (1). Mohle 71 (1) Lowveld Botanical Garden, PRE. Moll

1880 (4) NH, PRE; 2191 (6) NH, PRE; 3452 (1) NH, PRE. Muller &

Scheepers 223 ( 1 ).

Ngwenya 1186 (6) NH, PRE, Nicholas 2033 (1) NH. Noel RUH 10623

(5) GRA.

Obenneyer 933 (1). Dickers s.n. (4) Cedara Weeds Laboratory. Dliver

7(6) NH, PRE.

Pctppe SAM 1 8603 (\) SAM, Pegler 142 (\) GRA; 142A, 142B, 142C

( 1 ). Pentz & Acocks 10344 (6) NH, PRE. Phillipson 1198 ( 1 ) GRA,

PRE. Pienaar 34 (3) NH, PRE. Pont s.n. (5). Pott TRV13649,

TRV14902, TRV16808 ( 1 ). Priinos SAM684203 (2) SAM.

Rogers 18078 ( 1 ). Rudatis NBG162972 ( 1 ) NBG.

Saainum 633 (1). Scheepers 538 (1) PRE, PRU. Schlechter 6528 (1)

GRA. Sim 19998 (1). Stahnans 542 (1). Stirton 6273, 6291 (1); 6306

(5); 7452 (A) NH; 8649 (A) NH, PRE.

Theron 993 ( 1 ). Thode 4218, 5134, 5135 ( 1 ) NBG. Trauseld 597 ( 1 ).

Tyson 2929(1) SAM.

Van der Schijff 4895 (1) PRE, PRU; 5636 (1). Van Vuiiren 95 (1).

A.E.yan Wvk 13111 (1 ) PRU, Pvan Wyk BSA2511 (6) PRU. Van Wyk &

Theron 4586 ( I ) PRE, PRU,

Wager NH64099, NH64100 (6) NH. Ward 2884 ( 1 ) NH; 4888 (2) NH,

PRE; 4897 (A); 11176 (6) NH, PRE. Watt & Breyer-Brandwijk 1205 ( 1 ).

Wells 4021 (A) NH, PRE, West 1148 ( 1 ) NH, PRE. Wood 5718 (2) NH.

Wylie NH22472 (1) NH.

ACKNOWLEDGEMENTS

Dr R.N. Le.ster, University of Biimingham, United King-

dom, is thanked for the identification of some S. capsi-

coides, S. tonntm and S. viariim specimens and also for

the colour photograph of the isotype of S. macowanii at

Kew. Ms G. Condy is thanked for the line drawings for

this article. The Curators of GRA, NBG, NH, PRE, PRU

and SAM are thanked for loans of their specimens; also

the curator of the herbarium at the Lowveld Botanic

Garden, Nelspruit (National Botanical Institute) and Mr

T. Dickers of the Cedara Weeds Laboratory, KwaZulu-

Natal. The referees are thanked for valuable advice.

REFERENCES

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

the Botanical Survey of South A frica No. 57: 30, 94.

AHMAD, K.J. 1964. Epidermal studies in Solanum. Lloydia 27:

243-250.

ALLIONI. C. 1773. Auctuarium ad synopsim methodicam stirpium

horti regii Taurinensis. Turin.

BAILEY, L.H. & BAILEY, E.Z. 1977. Hortus third, a concise dictio-

nary of plants cultivated in the United States and Canada.

Macmillan, New York.

BITTER, G. 1919. Die papuasischen Arten von Solanum. Botanische

Jahrhiicher 55: 59-113.

BITTER, G. 1921. Solana africana III. Botanische Jahrbiicher 57:

248-286.

BITTER, G. 1923. Solana africana IV. Repertorium specierum nova-

rum regni vegetabilis. Beiheft 16: 1-320.

BRYSON, C.T. & BYRD, J.D. 1994. Solanum viarum (Solanaceae),

new to Mississippi. Sida 16: 382-385.

BUKENYA, Z.R. & CARASCO. J.F. 1995. Solanum (Solanaceae) in

Uganda. Bothalia 25: 43-59.

BUKENYA, Z.R. & HALL, J.B. 1988, Solanum (Solanaceae) in Ghana.

Bothalia 18: 79-88.

BUKENYA-ZIRABA, Z.R. 1996. Uses, chromosome number and dis-

tribution of Solanum species in Uganda. In L.J.G. van der

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

The biodiversity of African plants: 33-37. Kluwer Academic

Publishers, Dordrecht.

CARVALHO, L. D" AVILA F. DE 1985. Flora fanerogamica da reserva

do parque estadual das fontes do Ipiranga (Sao Paulo, Brasil).

136. Solanaceae. Hoehnea 12: 67-85.

CHILD, A. 1979. A review of branching patterns in the Solanaceae. In

J.G, Hawkes, R.N. Lester & A.D. Skelding, The biology and

taxonomy of the Solanaceae: 345-356, Linnean Society of

London.

CHILD, A. 1998. Studies in Solanum and related genera (6). New infra-

generic taxa for the genus Solanum L. (Solanaceae). Feddes

Repertorium 109: 407^27.

CHILD, A. & LESTER, R.N. 199 1 . Life form and branching within the

Solanaceae. In J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada,

Solanaceae 111: taxonomy, chemistry, evolution: 151-159.

Royal Botanic Gardens Kew and Linnean Society of London,

CLARKE, C.B. 1883. Solanaceae. In J.D. Hooker, The Flora of British

India A: 228-246.

DALZIEL, J.E. 1937. The useful plants of West tropical Africa. Crown

Agents for Oversea Government and Administrations, London.

DAN, S. & DAN, S.S. 1984. Chemotaxonomy of the genus Solanum.

Journal of Economic and Taxonomic Botany 5; 875-879.

D’ARCY, W.G, 1972. Solanaceae studies II. Typification of subdivi-

sions of Solanum. Annals of the Missouri Botanical Garden 59:

262-278.

D’ARCY, W.G. 1973. Flora of Panama, part IX. Family 170, Solana-

ceae. Annals of the Missouri Botanical Garden 60; 573-780.

D’ARCY, W.G. 1974. Solanum and its close relatives in Florida.

Annals of the Missouri Botanical Garden 61: 819-867.

D’ARCY, W.G. & RAKOTOZAFY, A, 1994. Flore de Madagascar et

de Comores, Famille 176, Solanacees: 129, 130.

DAUNAY, M.-C., LESTER, R.N. & LATERROT, H. 1991 . The use of

wild species for the genetic improvement of brinjal eggplant

(Solanum melongena) and tomato (Lycopersicon esculentum).

In J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada, Solanaceae

111: taxonomy, chemistry, evolution: 389^12. Royal Botanic

Gardens Kew and Linnean Society of London.

DON, G. 1838. Solanum section Aculeata subsection Acanthophora.

General System of Gardening and Botany A. London.

DUNAL, M.F 1813. Histoire naturelle, me'dicale et economkjue des

Solanum et des genres c/ui out ete confondus avec eux. Paris,

Strasbourg, Montpellier.

DUNAL, M.F 1852. Solanaceae. In A.P. de Candolle, Prodromus sys-

tematis naturalis regni vegetabilis 13: 1-690. Masson, Paris.

ESQUIVEL, M. & HAMMER, K. 1991. The cultivated species of the

family Solanaceae in Cuba. In J.G. Hawkes, R.N. Lester, M.

Nee & N. Estrada, Solanaceae 111: taxonomy, chemistry, evolu-

tion: 357-364. Royal Botanic Gardens Kew and Linnean Society

of London.

FEDOROV, A. A. 1969. Chromosome numbers of flowering plants.

Solanum: 693-703. Academy of Sciences, USSR, Moscow.

FOURCADE, H.G. 1934. Contributions to the llora of the Knysna and

neighbouring divisions. Transactions of the Royal Society of

South Africa 2\ : 15-102.

Bothalia 33,1 (2003)

17

GBILE, Z.O. 1979. Solanum in Nigeria. In J.G. Hawkes, R.N. Lester &

A.D. Skelding, The biology and taxonomy of the Solanaceae:

113-120. Linnean Society of London.

GBILE, Z.O. 1986. Epidermal studies in the Solanum nigrum complex

in Nigeria. In W.G. D’ Arcy, Solanaceae'. biology and systemat-

ics: 159-168. Columbia University Press, New York.

GBILE, Z.O. & ADESINA, S.K. 1988. Nigerian Solanum species of

economic importance. Annals of the Missouri Botanical Garden

75: 862-865.

GBILE, Z.O. & SOWUNML M.A. 1979, The pollen morphology of

Nigerian Solanum species. In J.G. Hawkes, R.N. Lester & A.D.

Skelding, The biology and taxonomy of the Solanaceae:

335-342. Linnean Society of London.

GENTRY, J.L. & STANDLEY, P.C. 1974. Flora of Guatemala, Solana-

ceae. Fieldiana, botany: 24.10: 1-151.

GOODING, E.G.B. 1965. Flora of Barbados 9\. Solanaceae: 373-382.

GROTH, D. 1989. Caracteriza5ao morfologica das sementes e plantu-

las de seis especies invasoras do genero Solanum L. Acta botanica

brasilica 3: 25^8.

HALL, J.B. & SWAINE, M.D. 1980. Seed stocks in Ghanaian forest

soils. Biotropica 12: 256-263.

HEINE, H.H. 1963. Solanaceae. In F.N. Hepper, Flora of West tropical

Africa, edn 2, 2: 325-335. Crown Agents, London.

HEINE, H.H. 1976. Solanacees. Flore de la Nouvelle-Caledonie et

Dependances 1: 119-205.

HEISER. C.B, 1971. Notes on some species of Solanum (sect. Lepto-

stemonum) in Latin America. Baileya 18: 59-65.

HEPPER, F.N. 1978. Typification and name changes of some Old World

Solanum species. Botanical Journal of the Linnean Society 76:

287-292.

HEPPER, F.N. 1979. On typifying Linnaean names of Solanaceae. In

J.G. Hawkes, R.N. Lester & A.D. Skelding. The biology and

taxonomy of the Solanaceae: 131-133. Linnean Society of

London.

HEPPER, F.N. & JAEGER, P.-M.L. 1986. A review of the genus Sola-

num in Africa. In W.G. D’Arcy, Solanaceae: biology and sys-

tematics: 41-55. Columbia University Press, New York.

HOSSAIN, M. 1973. Observations on stylar heteromorphism in

Solanum torvum Swartz (Solanaceae). Botanical Journal of the

Linnean Society 66: 291-301.

HUSAINI, S.W.H. & IWO, G.A. 1990. Cytomorphological studies in

some weedy species of the family Solanaceae from Jos plateau

in Nigeria. Feddes Repertorium 101 (1, 2): 41^7.

HUTCHINGS, A.. SCOTT, A.H., LEWIS, G. & CUNNINGHAM. A.B.

1996, Zulu medicinal plants, an inventory. University of Natal

Press, Pietermaritzburg.

JACQUIN, N.J. VON 1787a. Collectanea austriaca ad botanicum,

chemiam et historiam naturalem spectantia cum figuris 1.

Kraus, Vienna.

JACQUIN, N.J. VON 1787b. leones plantarum rariorum 1,5. Wappler,

Vienna.

JACQUIN, N.J. VON 1798. Plantarum rariorum horti caesarei

schoenbrunnensis descriptiones et icones 3. Vienna.

JAEGER. P.-M.L. 1985. Systematic studies in the genus Solanum in

Africa: 1-540. Ph.D. thesis. University of Birmingham. Un-

published.

JAEGER, P.-M.L. & HEPPER, F.N. 1986. A review of the genus

Solanum in Africa. In W.G. D’Arcy, Solanaceae: biology and

systematics: 41-55. Columbia University Press, New York.

JAIN, S.K. & BORTHAKUR, S.K. 1986. Solanaceae in Indian tradi-

tion, folklore and medicine. In W.G. D’Arcy, Solanaceae: biology

and systematics: 577-583. Columbia University Press, New

York.

KHAN, R. 1979. Solanum melongena and its ancestral forms. In J.G.

Hawkes, R.N. Lester & A.D, Skelding. The biology and taxon-

omy of the Solanaceae: 629-636. Linnean Society of London.

KNAPP, S. & JARVIS, C.E. 1990. The typification of the names of

New World Solanum species described by Linnaeus. Botanical

Journal of the Linnean Society 104: 325-367.

LAMARCK. J.B.A.P.M. DE 1794. Tableau encyclopedique et metho-

dique des trois regnes de la nature. Botanique, suppl. 2. Paris.

LAMBERTYE, L. DE 1865. Solanum warsce'wiczii. Revue horticole: 429.

LEBRUN. J.P. & STORK, A.L. 1997. Enumeration des plantes a fleurs

dAfrique tropicale IV. Solanaceae: 386-400. Conservatoire et

Jardin Botaniques, Geneve.

LEMKE, D.E. 1991. The genus Solanum (Solanaceae) in Texas. Phy-

m/ogia 71: 362-378.

LESTER. R.N. 1978. The identity of S. viarum Dunal (S. khasianum

C.B. Clarke var. chatterjeeanum Sen Gupta). Solanaceae News-

letter 5: 5.6.

LEVEILLE, A.A.H. & VANIOT, E. 1908. Solanum cavaleriei. Bulletin

Societe Botanique de France 55: 207.

LINNAEUS, C. 1753. Species plantarum, edn 1. Salvius, Stockholm.

LU, A. 1986. Solanaceae in China. In W.G. D’Arcy, Solanaceae: biology

and systematics: 79-85. Columbia University Press, New York.

MABBERLEY. D.J. 1997. The plant-book, a portable dictionary of the

vascular plants, edn 2. University Press, Cambridge.

MAITL PC.. MOOKHERJEA, S„ MATHEW, R., DAN, S.S. 1979.

Studies on Indian Solanum I. Alkaloid content and detection of

solasodine. Economic Botany 33: 15-11 .

MEHRA, K.L. 1979. Ethnobotany of Old World Solanaceae. In J.G.

Hawkes, R.N. Lester & A.D. Skelding, The biology and taxon-

omy of the Solanaceae: 161-170. Linnean Society of London.

MILLER, R.H. 1969. A morphological study of Solanum mammosum

and its mammiform fruit. Botanical Gazette 130: 250-251 .

MORTON, C.V. 1976. A revision of the Argentine species o/ Solanum:

1-260, Academia Nacional de Ciencias, Cordoba, Argentina.

NEE. M. 1979. Patterns in biogeography in Solanum. section Acantho-

phora. In J.G. Hawkes, R.N. Lester & A.D. Skelding. The biology

and tcLxonomy of the Solanaceae: 569-580. Linnean Society of

London.

NEE, M. 1986. Placentation patterns in the Solanaceae. In W.G. D'Arcy.

Solanaceae: biology and systematics: 169-175. Columbia Uni-

versity Press, New York.

NEE, M. 1991 . Synopsis of Solanum section Acanthophora: a group of in-

terest for glycoaUcaloids. In J.G. Hawkes, R.N. Lester, M. Nee & N.

Estrada, Solanaceae HI: taxonomy, chemistry, evolution: 257-266.

Royal Botanic Gardens Kew and Linnean Society of London.

NEE, M. 2001. Index to Solanum epithets. Unpublished electronic list.

NEES VON ESENBECK, C.G.D. 1837. Monograph of the East Indian

Solaneae. Transactions of the Linnean Society of London 17:

37-82.

OGUNDIPE, O.T. & DARAMOLA, S.O. 1997. Epidermal studies in

some species of Solanum L. (Solanaceae). Boletim da Socie-

dade Broteriana, ser. 2, 68: 5-33.

PEARCE, K. & LESTER. R.N. 1979. Chemotaxonomy of the cultivat-

ed eggplant — a new look at the taxonomic relationships of

Solanum melongena L. In J.G. Hawkes, R.N. Lester & A.D.

Skelding, The biology and ta.xonomy of the Solanaceae:

615-628. Linnean Society of London.

PEARSE, R.O. 1978. Mountain spleiuJour The wild flowers of the Drakens-

berg. Timmins. Cape Town.

PERSOON, C.H, 1805. Synopsis plantarum, seu enchiridium botani-

cum I: 228. Paris, Tubingen.

RANDELL. B.R. & S'YMON, D.E. 1976. Chromosome numbers in Austra-

lian Solanum species. Australian Jounuil of Botany 24: 369-379.

RAO, N.N. 1979. The barriers to hybridization between Solanum mel-

ongena and some other species of Solanum. In J.G. Hawkes,

R.N. Lester & A.D. Skelding, The biology and ta.xonomy of the

Solanaceae: 605-614. Linnean Society of London.

RETIEF, E. & HERMAN. P.P.J, 1997. Plants of the northern provinces

of South Africa: keys and diagnostic characters. Strelitzia 6:

622-624.

RHIND, J. 1969. Weeds in South Africa. Farmer’s Weekly, July 16: 67.

RHIND, J. 1975. Weeds in South Africa. Fanner's Weekly, October 15:

133.

RODDICK, J.G. 1986. Steroidal alkaloids of the Solanaceae. In W.G.

D'Arcy. Solanaceae: biology and systematics: 201-222,

Columbia University Press, New York.

RODDICK, J.G. 1991. The importance of the Solanaceae in medicine

and drug therapy. In J.G. Hawkes, R.N. Lester, M. Nee & N.

Estrada, Solanaceae III: taxonomy, chemistry, evolution: 1-25.

Royal Botanic Gardens Kew and Linnean Society of London.

ROE, K.E. 1971 . Terminology of hairs in the genus Solanum. Ta.xon 20:

501-508.

ROE, K.E. 1979. Dispersal and speciation in Solanum, section Brevan-

therum. In J.G. Hawkes, R.N. Lester & A.D. Skelding, The

biology and ta.xonorny of the Solanaceae: 563-568. Linnean

Society of London.

ROSS, J.H. 1972. The flora of Natal. Memoirs of the Botanical Survey

of South Africa No. 39: 308.

SAINT A.D. 1966. Alkaloidal content of Solanum khasianum Clarke.

Current Science 35: 600.

SCHLECH'l ENDAL, D EL. VON 1847. Solanum chrvsotrichum. Linnaea

19: 304.

SCHREIBER, K. 1979. Steroid alkaloids of Solanum. In J.G, Hawkes,

R.N. Lester & A.D. Skelding, The biology and taxonomy of the

Solanaceae: 193-202. Linnean Society of London.

18

Bothalia 33,1 (2003)

SEITHE, A. 1962. Die Haararten der Gattung Solanum L. und ihre tax-

onomische Verwertung. Botanische Jahrhiicher 81: 261- 336.

SEN GUPTA, G. 1961. A taxonomic note on Solanum khasianum

Clarke and description of a new variety under it. Bulletin of the

Botanical Survey of India 3: 413.

SHAW, J.M.H. 2000. Solanum. In J. Cullen et al.. The European garden

flora, vol. VI, Dicotyledons (part IV). Cambridge University

Press, Cambridge.

SWARTZ, O.P. 1788. Nova genera et species plantarum. Stockholm.

SYMON, D.E. 1979. Sex forms in Solanum (Solanaceae) and the role

of pollen collecting insects. In J.G. Hawkes, R.N. Lester & A.D.

Skelding, The biology and taxonomy of the Solanaceae: 385-

398. Linnean Society of London.

SYMON. D.E. 1981. A revision of the genus Solanum in Australia.

Journal of the Adelaide Botanic Garden 4: 1-367.

SYMON, D.E. 1982. Solanum. In R.W. Purdie, D.E. Symon & L. Haegi,

Flora of Australia 29, Solanaceae: 69-175.

SYMON, D.E. 1985. Solanaceae of New Guinea. Journal of the Adelaide

Botanic Garden 8: 1-171.

TROUPIN, G. 1985. Solanum (Solanaceae). Flore du Rwanda 3: 372-382.

VON SCHRANK, F. VON P. 1822. Solanum reflexum. Plantae rari-

ores horti academici monacensis 2: t. 81.

WATT, J.M. & BREYER-BRANDWIJK, M.G. 1962. Medicinal and

poisonous plants of southern and eastern Africa, edn 2. Living-

stone, Edinburgh & London.

WELLS, M.J., BALSINHAS, A.A., JOFFE, H., ENGELBRECHT, V.M.,

HARDING, G.G. & STIRTON, C.H. 1986. A catalogue of

problem plants in southern Africa. Memoirs of the Botanical

Survey of South Africa No. 53: 483^91.

WELMAN, W.G. 1993. Solanaceae. In T.H. Arnold & B.C. de Wet,

Plants of southern Africa: names and distribution. Memoirs of

the Botanical Survey of South Africa No. 62: 617-621.

WHALEN, M.D. 1984. Conspectus of species groups in Solanum sub-

genus Gentes Herbarum 12: 179-282.

WHALEN, M.D. & COSTICH, D.E. 1986. Andromonoecy in Sola-

num. In W.G. D’Arcy, Solanaceae: biology and systematics:

284-302. Columbia University Press, New York.

WfLDEMAN, E.A.J. DE 1907. Solanum pynaertii. Mission Emile Laurent

2. F. Vandenbuggenhoudt, Brussels.

WRIGHT, C.H. 1894. Solanaceae. Kew Bulletin: 127-129.

WRIGHT, C.H. 1904. Solanaceae. In W.T. Thiselton-Dyer, Flora capen-

.sis 4,2: 87-121.

WRIGHT, C.H. 1906. Solanaceae. In W.T. Thiselton-Dyer, Flora of tropi-

cal Africa 4,2: 207-261.

WUNDERLIN, R.P., HANSEN, B.F, DELANEY, K.R., NEE, M. &

MULLAHEY, J.J. 1993. Solanum viarum and S. tampicense

(Solanaceae): two weedy species new to Florida and the United

States. Sida 15: 605-611.

Bothalia 33.1: 19-39 (2003)

Studies in the genus Machairophyllum (Mesembryanthemaceae), with

notes on some related genera

H. KURZWEIL* and P. CHESSELET*

Keywords: Bijlia N.E.Br., Carruanthus (Schwantes) Schwantes. Cerochlamys N.E.Br., fynbos, Machairophyllum Schwantes, Mesembryanthe-

maceae, morphology. South Africa, taxonomy, ultrastructure. Western and Eastern Cape

ABSTRACT

Machairophyllum Schwantes is a small genus of the family Mesembryanthemaceae, confined to the southern part of

central South Africa from the Little Karoo at Barrydale in the west to the Zuurberg near Kirkwood in the east. Machairo-

phyllum is unusual in the Mesembryanthemaceae-Ruschioideae in that fynbos and renosterveld are the preferred habitats.

The morphology, anatomy as well as the pollen, seed and leaf epidermis ultrastructure of the genus were examined. The

taxonomy of Machairophyllum is revised on the basis of data presented. Of the ten names originally described, only four

are upheld. M. albidum (L.) Schwantes, M. stayneri L. Bolus, M. bijliae (N.E.Br.) L. Bolus and M. brevifolium L. Bolus. All

four species are very similar in their vegetative habit, and differ mainly in inflorescence and flower characters. Several

presumably related genera, Bijlia N.E.Br.. Carruanthus (Schwantes) Schwantes and Cerochlamys N.E.Br., are briefly dis-

cussed.

INTRODUCTION

Machairophyllum Schwantes is one of the few genera

of the succulent plant family Mesembryanthemaceae

which is almost exclusively found in fynbos or renoster-

veld. The species mostly prefer rocky habitats (Eigure

lA, B), although occasionally they grow among grass

and shrubs (Eigure 1C). In its distribution Machairo-

phyllum is, with two possible exceptions, entirely con-

fined to Western and Eastern Cape, South Africa (Eigure

2). Two localities have been reported in Northern Cape,

but are doubtful and need to be verified (see chapter on

phytogeography).

Plants of the genus Machairophyllum are unmistak-

able, with their clump-forming habit and the smooth

trigonous leaves. The leaves have a sabre-like appear-

ance, which is the origin of their botanical name

CMachaira\ Greek, means sabre, and "phyllon means

leaf). Plants are seldom found in flower as flowering is

stimulated by fire. Furthermore, flowers are mostly ves-

pertine or nocturnal. The striking, fairly large flowers

(Figure ID, E) are yellow and frequently orange-red on

the outside and on the petal tips.

Ten species have been described of which only four

are recognized in the present study, Machairophyllum

albidum (L.) Schwantes, M. bijliae (N.E.Br.) L. Bolus, M.

brevifolium L. Bolus and M. stayneri L. Bolus. M. albi-

dum is known from several sites in the Little Karoo and

is easily recognized by the branched inflorescences and

the normally long basal sheaths of the primary bracteoles

(‘bracts’). M. stayneri also has branched inflorescences,

and is known only from the type collection made on the

Zuurberg. M. bijliae is the most common and widespread

species. Vegetatively it looks identical to M. albidum, but

once flowers appear, the difference is obvious since the

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

7735 Claremont, Cape Town.

MS. received: 2002-04—15.

inflorescences are always unbranched and have mostly

short, basal, bracteole sheaths. M. brevifolium, as its

name suggests, has short and rather wide leaves. It is

known only from the red conglomerate hills between

Oudtshoom and De Rust.

Machairophyllum albidum and M. bijliae are fairly

abundant. In some cases the populations that were visit-

ed by the first author comprised hundreds of individuals.

It has been suggested to rate M. vanbredai L. Bolus as

‘Critically Endangered’ (Kurzweil & Linder-Smith 2001 ),

but we now consider this taxon synonymous with the

widespread M. bijliae. M. stayneri is only known from

the type collection, and the fact that the species was

never recollected may indicate that it is indeed very rare

or extinct. Populations of the two dwarf species from the

crumbling silcrete hills around De Rust (M. brevifolium

and the synonymous M. latifolium) appear to be declin-

ing in numbers due to the erosion of their habitat (Smith

etal. 1998; 220).

Machairophyllum is not very popular among succu-

lent growers because of the often large plant size and the

sporadic flowering. Plants are therefore rather uncom-

mon in cultivation although well-grown specimens may

be remarkably showy. M. albidum was introduced into

Europe in 1715, but is rather scarce now (Hammer 1991 ).

The plants can endure fairly low temperatures, and due

to their fynbos habitat they can tolerate considerably

more watering than many other mesembs (Kurzweil

pers. obs.). Propagation can take place through seeds or

cuttings (Jacobsen 1977). M. brevifolium is not uncom-

mon in collections in California and England, probably

because seeds were sent overseas by Mrs Van der Bijl, a

well known South African succulent collector (Hammer

1991: 77).

The objective of this study is to examine the character

distribution in Machairophyllum with the aim to resolve

the taxonomy of the genus. For the purpose of phyloge-

netic interpretation, the presumed relatives in Hartmann’s

20

Bothalia 33,1 (2003)

FIGURE 1. — A-C. habitat. A, Macluiirophylluin alhiclum, Kiirzweil IH90. B, C, M. hijiicie: B, Kurzweil I896\ C, Kurzweil 1913. D, E, llowering

plants in previously burnt veld, photographed at around 17h00, Kurzweil 1902.

(1991 ) Bergerantim.s group, i.e. the minor genera Bijiia

N.E.Br., Carruanthu.s (Schwantes) Schwantes and Cero-

chUimys N.E.Br., which all .share smooth and dotless

leaves with Macluiirophylluiu. were briefly surveyed. Out

of these, Bijiia and Carruanthus were found to form a

very close-knit group with Macluiirophylluw, which is

here referred to as the "Macluiirophyllum complex’.

Taxonomic position and history

On account of having an inferior ovary with parietal

placentation and lophomorphic (crest-shaped) nectaries,

Macluiiropliylliwi is placed in Mesembryanthemaceae-

Ruschioideae (Herre 1971). The Ruschioideae is the

larger of the two subfamilies of Mesembryanthemaceae

with 107 genera mainly in .southern Africa. Ruschioideae

was classified into 12 tentative groups according to the

then state of knowledge (Hartmann 1991, 1993, 1998a).

Machaiwphylhim was listed in the Bergeranthiis group

which comprises perennial plants that mostly have free

leaves with homocellular xeromorphic epidermis, exsert-

ed inllorescences and flowers with separate lophomor-

phic nectaries. Distinguishing characters of the capsules

are the complete, firm and straight covering membranes

and the reduced valve wings. Closing bodies can be pre-

sent or absent. The genera included in this group are

Bergeranilius Schwantes, Bijiicp Carruanthus, Cerochla-

mys, Hereroa (Schwantes) Dinter & Schwantes, Rhomho-

phyllum (Schwantes) Schwantes and Machairophyllum.

Many members of the group also have semi-terete or

trigonous linear to lanceolate leaves. The genera includ-

ed in the Bergerauthus group share a similar distribution

in the southern and eastern parts of central South Africa.

Hereroa, however, has a wider distribution ranging into

Bothalia 33,1 (2003)

21

HQOHT ABOVE SEA LEVS.

^ Over 1500 m

Fn 900 - 1500 m

m 300 - 900 m

I 1 Under 300 m

FIGURE 2. — Distribution of the

genus Machairophyllunr. M,

albidiim, O; M. bijiiae. •; M.

brevifolium. A; M. stayneri, O.

Doubtful records in Northern

Cape: O, fide Herre (1971:

198; near Vanrhynsdorp, no

comment on source of infor-

mation or identity of species

was made); ©. Viviers s.n.

(NBG, hort.; Richtersveld;

probably M. bijiiae).

the Free State, Northern Cape, Gauteng and Namibia.

Most genera of the Bergeranthiis group share a similar

pollination syndrome — with Cerochlamys as the only

exception, all have yellow flowers which are open in the

afternoon, evening or at night.

The genus Machairophyilum was described by Gustav

Schwantes in May 1927, based on M. albidiim and M.

cookii (here also included in the concept of M. albidiim)

which he had previously incorporated into the genus

Carniantluis. The species were said to differ from Car-

nianthiis mainly by having many stigmas. Schwantes

selected the former as the type species. Four years later

M. stenopetalum was described by Louisa Bolus { 1931 ),

followed by M. acuminatum and M. baxteri in 1935 (all

three are here included in M. bijiiae). M. brevifolium and

M. latifolium (here regarded as synonymous) were

described by L. Bolus in 1938. In the same year L. Bolus

also transferred Perissolobus bijlii to Machairophyilum.

The last two additions to the genus Machairophyilum

were M. stayneri (Bolus 1960) and M. vanbredai (Bolus

1964) (here included in M. bijiiae).

In 1960 L. Bolus published a brief review of Ma-

chairophyilum together with a key to the species; M.

bijiiae was not included as it was insufficiently known.

Later this key was used by Jacobsen (1977) in Lexicon of

succulent plants. In her review. Bolus (1960: 156) point-

ed out that most species were known only from their type

collections, and that ‘it is quite possible that, when more

intensive collecting and study have been done, M. bijiiae

(in spite of the much smaller size of the type, perhaps due

to adverse conditions), M. stenopetalum, M. acuminatum

and M. baxteri may prove to be forms of one variable

species’. This statement is very much in agreement with

the results of the present study. It is interesting that Bolus

doubts the validity of her own species (except for the

species that M. bijlii is based on, all were described by

herself). However, this did not prevent her from describ-

ing another species a few years later, namely M. van-

bredai in 1964, which is also merely a form of this ‘vari-

able species’ (M. bijiiae).

The species that Machairophyilum bijiiae is based on

was first described in a monotypic genus Perissolobus

(Brown 1931 ). According to Brown, the main reasons for

the recognition of Perissolobus were its eight sepals;

affinities to Machairophyilum were not mentioned in his

paper. Bolus (1938) pointed out the striking similarity of

the two genera and transferred Perissolobus bijlii to

Machairophyilum. She regarded the eight sepals of the

species as being ‘but a further variation of the already

variable calyx of Machairophyilum' (Bolus 1938: 135).

Bolus (1950: 229) pointed out that the reason for there

not being any reference to Machairophyilum in Brown’s

paper was probably the fact that at the time Machairo-

phyllum was erroneously thought to occur in Namaqua-

land in Northern Cape, whereas Perissolobus bijlii was

described from Eastern Cape.

A synopsis of Machairophyilum was presented by

Hartmann (2001). Both species with five or six sepals

and ± double the number of locules were considered con-

specific, namely M. albidiim and M. cookii. Also M. bijii-

ae and M. vanbredai on the one hand as well as M. bre-

vifolium and M. latifolium on the other hand were united

as they were considered to differ only in minor charac-

ters. Thus the number of species was reduced to seven. It

was also pointed out that a study of the genus may show

that only four or five species are distinguishable, namely

M. albidiim, M. bijiiae, M. brevifolium, M. stayneri and,

possibly, a new species, allegedly from the Richtersveld.

MATERIAL AND METHODS

Dry and liquid-preserved material of Machairo-

phyilum, as well as live seeds, were collected in the field

in South Africa. For comparative purposes, material of

Bijlia, Carriianthus and Cerochlamys was also collected.

Herbarium material on loan from the Bolus Herbarium of

the University of Cape Town (BOL) and the Herbarium,

Royal Botanic Gardens Kew, London (K) was examined.

Additional material was obtained from the Compton

Herbarium of the National Botanical Institute, Cape Town

(NBG) and from living plants cultivated in the National

Botanical Garden, Kirstenbosch.

22

Bothalia 33,1 (2003)

Herbarium, live and liquid-preserved material was

used for gross-morphological investigations. The scan-

ning electron microscope (SEM) technique had to be

employed for the study of fine details. With this tech-

nique the examination of seeds, pollen and epidermal

surfaces was done without previous treatment of sam-

ples. SEM investigations of floral structures as well as

anatomical studies required the previous fixation of the

material in FAA (ethanol 70% : glacial acetic acid : for-

maldehyde = 18 : 1 : 1). Material was treated with FAA

for a few days and later stored in ethanol 70%. For SEM

studies it was dissected in ethanol. Subsequently, sam-

ples were chemically dehydrated in formaldehyde-

dimethylacetal and critical-point-dried directly from this

reagent without the use of an intermedium, using CO2 as

the earner gas (technique after Gerstberger & Leins 1978).

Samples were sputter-coated with AuPd and viewed and

photographed with a CAMBRIDGE STEREOSCAN

S200 scanning electron microscope at voltages ranging

from 10 to 30 kV. Anatomical studies were undertaken

using a standard dehydration and wax embedding tech-

nique; samples were stained with alcian blue and

safranin.

MORPHOLOGICAL CHARACTERS (SEE TABLE 1)

Growth forms

Machairophylhim species are vegetatively fairly uni-

form. The plants are compact, glabrous succulents with

short stem internodes and trigonous leaves (Figure

lA-C). Old plants may form mats of up to 1.2 m in

diameter. Even in large clumps the side branches nor-

mally do not form adventitious roots and are thus not

anchored to the ground. Superficially, plants of

Machairophylhim may be mistaken for Carpobrotus,

which also occurs in fynbos, although these have some-

what longer and greener leaves and have well-developed

stem internodes; however, the flowers and the fruit struc-

ture of Carpobrotus are very different.

Leaf

Morphology

The leaves are opposite with members of a pair ±

equal and basally connate to form a sheath 10-15 mm

long; in early ontogenetic stages leaves of a pair are often

weakly unequal (anisophylly). The lamina is linear to

lanceolate, entire, trigonous in the central portion but

basally semi-terete because the dorsal (abaxial) keel dis-

appears; the apex is acute to acuminate. Leaves are pale

green often with a whitish tinge, or frequently flushed

with red in sun-exposed specimens. Old leaves die off

and turn black, but persist on the plant for a long time.

Near the base, just above the union of a pair, an obscure

swelling is visible on the upper side, which stretches as a

curved structure over the entire width of the leaf. It

marks the point up to which the leaf is tightly adpressed

to the following leaf pair. However, this swelling is not

developed as a prominent pustule, which is found in

many other mesembs (Hartmann 1991).

Plants growing wild have leaves 10- ± 145 x 8-26 mm.

The variation in length and width is continuous and does

not allow a clear-cut separation of taxa (Figure 3).

Whereas M. brevifoliitm always has short leaves, a few

specimens referable to M. bijliae also have very short

leaves, namely Van der Bijl 93 (the type), Desmet 2158,

Pocock NBG627/26 and Kurzweil 1919. Particularly long

leaves are found in M. albidum. It appears that leaf length

and width are mainly the result of environmental factors

and are not genetically fixed. The leaf size was previously

considered a major key character (Bolus 1960).

Leaf surface

The leaf surface belongs to the ‘xeromorphic type’

defined by equal or nearly equal epidermis cells, a

strongly thickened outer cell wall and a prominent wax

cover (Ihlenfeldt & Hartmann 1982) (Figure 4); epider-

mis cells are irregularly polygonal-isodiametric to elon-

TABLE 1. — Important features of the Machairophyllwn species recognized

Bothalia 33,1 (2003)

23

25

23 *

19 *

£ °

£ 17 0

^'5 °

♦ ^

0 20 40 60 80 100 120 140

leaf length (mm)

FIGURE 3. — Diagram showing relationship of leaf length and width in

Machairophyllum. M. albidiim, □; M. stayneri, O; M. bijliae,

♦; M. brevifolium, A. Plotted measurements represent average

values found in examined specimens.

gate, ± 61-146 x 30-85 jam; anticlinal walls are straight;

stomata are superficial to slightly sunken (Figure 4F), on

both sides of the leaf, paracytic (seusii Dupont 1976),

39.32 X 22.00 pm.

Leaves lack teeth, dots or hairs, are covered by a con-

tinuous wax layer, which is either smooth or breaks up

into platelets (Figure 4A-F). Local wax projections

mainly take the shape of elongate rodlets (Figure

4B-D), but round granules or flakes are also found

occasionally (Figure 4B). In Kurzweil 1914 structures

resembling burst bubbles were observed next to a stoma

(Figure 4F), but nothing can be said about their nature at

this stage.

Anatomy

Leaves are mostly well differentiated into peripheral

assimilation tissue and central water storage tissue

(Figure 5 A), which are parenchymatic and have intercel-

lular spaces. Tannin idioblasts, a common feature in

xeromorphic Mesembryanthemaceae (Hartmann 1991),

as well as raphide idioblasts occur throughout the meso-

phyll, but are absent in the epidermis and hypodermis.

Epidermis is homocellular, cells with flat outer peri-

clinal walls without papillae; cuticle 1 to 2 pm thick;

outer wall is strongly thickened at 5-25 pm and contains

crystal sand made up of minute grains of calcium oxalate

as in other xeromorphic Mesembryanthemaceae (Ihlen-

feldt & Hartmann 1982; asterisk in Figure 6A); inner

periclinal walls and the anticlinal walls are 1-3 pm thick;

hypodermis cells are elongate or more rarely isodiamet-

ric, walls 1-3 pm thick.

Assimilation tissue is compact, 10-15-seriate (Figure

6B); vascular bundles as well as fibre bundles are absent;

a ring of small vascular bundles is present at the bound-

ary of assimilation tissue and water storage tissue,

referred to as secondary bundles (Dannemann 1883;

arrowhead in Figure 6C). The central water storage tissue

is 10-30-seriate, contains laxly arranged cells; large

water storage cells are scattered throughout. Primary

bundles: 1-3, are large with several smaller collateral

vascular bundles (Figure 6D); the main bundle is nor-

mally broad, straight or slightly curved. Most of the pri-

FIGURE 4. — SEM micrographs of

leaf surfaces in Machairo-

phylliim: A. E, M. cilhiduin:

B, M. cf. hijliae', C, D, M.

bijiiae', F, M. sp. A. Kurzweil

1900: B, Bayer 7027: C,

Kurzweil 1901: D, Kurzweil

1896: E. Kurzweil 1902: F,

Kurzweil 1914. Scale bars:

20 pm.

24

Bothalia 33,1 (2003)

FIGURE 5. — Semi-diagrammatic representation of the anatomy of leaf

and stem in Machairophyllunr. A, M. alhidiim. Kurzweil 1890,

leaf, B-D. M. hijiiac. Kurzweil /9/9: B, primary stem; C. D,

enlarged sections of B. A, assimilation tissue; C, cortex; COLL,

collenchyma; E, epidermis; H. hypodcrmis; R phloem; PB. pri-

mary bundle; PI, pith; SB, secondary bundle; VC, vascular

cylinder; W, central water storage tissue; X, xylem. Seale bars:

A, 5 mm; B, 20 mm; C, D. 0.1 mm. Drawn by P. Chesselet,

mary bundles have collenchymatic caps, and bundles

plus their caps are surrounded by parenchymatic sheaths.

Stem

The anatomy of the stem conforms essentially to that of

other Mesembryanthemaceae with similar compact growth

(Hartmann 1976, 1977; Hartmann & Golling 1993).

Epidermis and cortex are only visible as separate

structures in the primary stem (Figure 5B, D). The epi-

dermis is similar to that of the leaf, and the hypodermis is

insignificant. The massive parenchymatic cortex proba-

bly acts as a storage organ. In older stems the cortex is

successively replaced by a layered cork which originates

in its inner part. In several specimens a layer of small and

compactly arranged collenchymatic cells is found just

outside the vascular cylinder (Figure 5B, C), reminiscent

of a similar collenchymatic ring in Odontophonis

(Hartmann 1976) and Fenestraha (Hartmann 1982). The

vascular tissue of the primary stem consists of a ring of

collateral bundles, and takes the shape of a rounded rec-

tangle with its long axis parallel to the axis of the first

leaf. Feaf traces emerge in rapid succession from the

shorter sides of the rectangle. Each pair emerges at an

angle of 90° to the previous one, and the shape of the vas-

cular cylinder therefore changes according to this pattern.

Secondary growth starts in the second internode. It is

anomalous as in other Mesembryanthemaceae (Metcalfe

& Chalk 1950) in that numerous secondary bundles are

developed by new cambium formed on the outside of the

primary vascular cylinder (epifascicular cambium).

These eventually form several concentric rings around

the primary vascular cylinder (Figure 7A, B). An unusu-

al cell pattern was observed in the pith of several speci-

mens of MachaiwphyUiim bijiiae (Figure 7C), looking

like pith cells with minute rounded ‘cells’ between them.

It is possible that the latter are in fact intercellular spaces.

Root

Roots are mostly thin and slender, up to 2 mm in

diameter. Anatomically the roots resemble stems. They

also show an anomalous secondary growth (Metcalfe &

Chalk 1950). Roots are usually diarch (2-stranded) or

triarch (3-stranded). Prominent collenchyma caps are

found on the phloem of the bundles. In some specimens,

strongly thickened sclereid cells are found below the

xylem and form pronounced sclerenchyma caps.

Inflorescence and pedicel

In principle the inflorescence of Machairophylliim is a

dichasium (Figure 8A) which is typical of Mesem-

bryanthemaceae (Hartmann 1993). The number of flowers

ranges from three to ± seven. Frequently one of the side

branches is aborted, which is probably the result of envi-

ronmental factors, or may be caused by disease, or insect

attack. In a large number of specimens the flowering shoot

is consistently unbranched and has a single flower (Figure

8B), and lateral buds or vestiges thereof were not found.

This is obviously the result of a reduction which is also

found in some other mesembs and may be part of their life

Bothalia 33,1 (2003)

25

FIGURE 6. — Leaf anatomy in

Machairophylhim in t/s. A,

M. a I bid urn, Kurzweil 1902,

epidermis. Asterisk indicates

crystal sand in outer epider-

mis wall. B, D. M. bijliae,

NBG806/83: B, assimilation

tissue; D, central vascular

bundle, note the dark col-

lenchyma caps. C, M. sp.,

NBG44/83: secondary vascu-

lar bundles, indicated by ar-

rowheads. Scale bars: A, D,

50 pm; B, C, 100 pm.

Strategy, aimed at a synchronization of the flowering in

order to improve the probabihty of successful pollination

(Ihlenfeldt 1989: 80). It was found that the degree of

branching is remarkably constant in Machairophylhim. In

the present study the inflorescence type is considered a

character of major taxonomic importance, with the

branched inflorescence characterizing M. albidum and M.

stayneri, and the unbranched inflorescence characterizing

M bijliae and M. brevifoliiim. This corresponds exactly

with the concepts of Bolus (1960: 157) and J. Vlok (pers.

FIGURE 7. — Secondary stem in

Machairophylhim bijliae in

t/s, Kurzweil 1919. A, B, vas-

cular bundles; C, pith. PB,

primary bundles; SB, sec-

ondary bundles. Scale bars:

50 pm.

26

Bothalia 33.1 (2003)

FIGURE 9. — Length of bracteole sheath in Machairophylliim albidum,

□ ; and M. bijiiae, ♦. Average measurements were used for cal-

culation.

fruit stage, 3-6 mm thick and are smooth on the outside

or have pronounced ribs. Anatomically they are rather

uniform, consisting of an epidermis with a crystal sand

layer, a weakly-defined hypodermis, a massive cortex

followed by the vascular cylinder and a parenchymatic

pith (Figure lOA, B). The vascular cylinder is a closed

ring with inner xylem elements, multi-layered cambium

and an outer phloem. All specimens examined have a

closed collenchymatic ring on the outside of the phloem.

Fibre bundles in the cortex occur occasionally.

FIGURE 8. — Semi-diagrammatic representation of inflorescences in

Machairophyllum. A. M. albidum. Kurzweil 1900\ dichasium;

B. M. bijiiae. Kurzweil 1919; solitary flower. Scale bars: 10

mm. Drawn by R Chesselet.

comm.). In the vast majority of cases the distinction of

specimens with branched versus unbranched inflorescences

is clear-cut but, rarely, poorly grown specimens of M.

albidum were found to be single-flowered (however, they

normally reveal their affinity to M. albidum in floral char-

acters).

In all cases the flowers are subtended by an opposite

pair of short bracteoles which resemble foliage leaves

[These bracteoles have also been referred to as ‘bracts’,

and it needs to be noted that their homology is disputed.

However, the term ‘bracteoles’ was used in the most recent

treatment of the genus (Hartmann 2001), and is therefore

adopted here]. Bracteoles are 7-130 mm long, with their

free upper portion ranging from 6-60 mm; bases are con-

nate and form a sheath. The length of this sheath is strong-

ly intercorrelated with the degree of inflorescence branch-

ing: M. bijiiae and M. brevifolium, the two species with

unbranched inflorescences, have predominantly basal

bracteole sheaths of up to 25 mm long (total range 5-42

mm) whereas M. albidum, with branched inflorescences,

has predominantly primary bracteole sheaths which are

longer than 21 mm (ranging from 15 to almost 100 mm)

(Figure 9). The single specimen of M. stayneri (branched

inflorescence) has primary bracteole sheaths of 15 mm.

The term ‘primary’ is here used for the terminal flower

of the dichasium, i.e. here applying to the basal-most

bracteoles. M. albidum and M. stayneri have ‘secondary’

bracteoles below the flowers of the side branches.

Flowers are generally stalked in Machairophyllum.

The pedicel is up to 85 mm long in solitary flowers and

in terminal flowers of branched inflorescences, is usual-

ly shorter on side branches, sometimes elongate in the

Flower

Flowers are actinomorphic and bisexual, up to 65 mm

in diameter, with few sepals, numerous petals and sta-

mens, and up to 12 stigmas; filamentous staminodes are

absent. In M. albidum and M. bijiiae the number of sepa-

ls and locules is subject to slight variation (see also Bolus

1950: 229), and different numbers may even be found in

different flowers on the same plant. In contrast, in M.

brevifolium the number is constantly six. Pollination

studies were not earned out, but from a functional-mor-

phological point of view the flowers can be referred to

the ‘Phalaenophilous’ type (Hartmann 1991: 91). A

moth-pollination syndrome is suggested by the yellow

flowers with a pale centre which are open in the after-

noon, evening or at night.

Sepals five to eight, unequal, (Figure 11 A) succulent

and weakly keeled, with white, membranous margins,

apically acute, acuminate or rarely somewhat obtuse, fre-

quently with subapical mucro on the dorsal side; total

length + 3-22 mm. Sepals of M. albidum tend to be

wider than those of the other species, but a comparative

analysis of the sizes does not allow a grouping within the

genus (Figure 1 IB).

Petals develop as staminodes from the androecial pri-

mordium as in other Mesembryanthemaceae-Ruschioi-

deae (Hartmann 1993); are arranged in three to seven

series; are ligulate or linear-lanceolate with acute or

more often obtuse apex; up to 35 x ± 0.3-2 mm; are yel-

low or golden yellow and often have reddish, copper-

coloured or orange-coloured outsides and tips and paler

bases; petal sizes as well as the ratio of the lengths of

petals and sepals are largely continuous (Figure 1 1C, D).

Stamens remain erect during anthesis and form a cone;

their bases are epapillate or only the inner ones are obscure-

BothaIia33,l (2003)

27

FIGURE 10. — Anatomy of flower

stalk in MachairophxUum in

t/s. A, M. albidum, Kurzweil

1902. epidermis; B, M. bijliae,

Kurzweil 1925. close-up of

vascular cylinder. Scale bars:

50 pm.

ly papillate; are 5-19 mm long and generally longer in the

larger-flowered M. albidum than in M. bijliae.

Nectaries are meronectaries of the lophomorphic type

(Rappa 1912; Hartmann 1991), green. They appear as

five to twelve ± prominently raised portions of a ring at

the inner base of the stamens opposed to the locules

(Figure 12A-C). The number of nectaries is normally

identical with the number of carpels/locules. Numerous

modified stomata can be seen and are frequently partly

blocked (Figure 12F). Crystals were occasionally also

observed on these nectaries, both in transections and

SEM micrographs (Figure 12E). Their chemical nature is

not known, but they are probably not simply made up of

solidified nectar, as it is difficult to imagine that crystal-

lized nectar would not have dissolved during the chemi-

cal preparation for SEM or wax embedding. In a few

specimens there are also structures that resemble minute

craters surrounded by a pronounced circular ridge (Figure

12D). In the past, the nectaries of Machairophyllitm have

been misinterpreted, as they have been termed distant,

approximate or contiguous (Herre 1971) which is proba-

bly mainly the result of their different extent of formation.

The syncarpous ovary is epigynous and is made up of

five to twelve carpels which terminate in slender, free stig-

mas that nearly reach the height of the stamens. Each ovary

chamber contains many ovules on parietal placentae. M.

albidum has predominantly ten to twelve stigmas, whereas

most specimens of M. bijliae and the type of M. stayneri

have six to eight stigmas (erroneously 7 or 8 were cited in

the description of the species). The two specimens of M.

brevifolium examined here exhibit six stigmas (Figure 13).

Unlike many other Mesembryanthemaceae, the flow-

ers of MachairophyUum are afternoon-active, vespertine

or nocturnal and remain closed during the day.

Fruit

Obconic, 7-15 mm in diameter when closed, loculici-

dal, hygrochastic capsules as in most other Mesem-

bryanthemaceae-Ruschioideae (Hartmann 1991, 1993);

their base is funnel-shaped and the top strongly raised in

the centre; sutures are prominent and the high valve rims

are normally reflexed (Figure 14A). Fully expanded cap-

sules usually have the valves at right angles to the floral

axis. The capsules do not clearly fall into any of the mor-

phological fruit types distinguished by Hartmann (1988:

349) and the genus was therefore listed as a ‘genus of

uncertain position’. Chesselet et al. (2000) referred to the

capsule as ‘similar to that of Cylindwphyllum and therefore

perhaps close to the Leipoldtia type’, but with spongy tis-

sue in the covering membranes and small closing bodies.

Within the genus MachairophyUum the internal struc-

ture of the capsule is not uniform (Figure 14B-D).

Capsules normally have 5-12 locules with the numbers

6-10 being the most common. A larger locule number of

14 or 15 was reported by Salm Dyck (1849), but this

could not be confirmed. As pointed out before, the num-

ber of locules is often not stable in MachairophyUum.

Valves are broadly to narrowly triangular, depending on

the locule number, 3. 0-6. 6 mm long, basal width 3. 5-6.0

mm, shortly and narrowly winged with the exception of

M. brevifoliimp wings taper apically with their ends

drawn out into slender tips which are connate with the

awns of the expanding keels; expanding keels ± parallel

at the base and close to each other and diverge to some

degree in their distal part, serrate or lacerate on their

inner margin, though sometimes minutely so, tips are

slender awns, l-2(-3) mm long which nearly reach the

tip of the valves. These awns are generally cohering and

frequently cross each other. In M. brevifolium the awns

are shorter or hardly developed. Covering membranes

petal length (mm) ^ petal width (mm) 03 sepal width (mm) ^ number of specimens

28

Bothalia33,l (2003)

♦

♦ ♦ ♦

♦ ♦ ♦

□

♦ 0 4

□ alhicliim

o siayneri

♦ hijliae

A hrevifolium

sepal length (mm)

□ albiJum

o siayneri

♦ hijllae

A hrevifoluini

[K'tal length (mm)

30

25

20

15

10

D

□ alhichim

o siayneri

♦ hijliae

A hrevifolium

8 10 12 14 16 18 20

FIGURE 11. — Diagrams showing

sepal and petal characters in

Machairophylliwi. Plotted

measurements in B-D repre-

sent average values found in

examined flowers. A, num-

ber of sepals; B, relationship

of sepal length and width; C,

relationship of petal length

and width; D, relationship of

sepal length and petal length.

D

sc|):d length (mm)

Bothalia 33.1 (2003)

29

FIGURE 12. — SEM micrographs of nectaries in Machairophyllum. A-E. M. alhidiinr. A. receptacle with sepals, petals, stamens and stigmas

removed, showing ring of nectar glands; B. C. individual nectar gland; D. crater-like structures on surface of nectar gland; E. crystals. F. M.

bijiiae. modified stomata on nectary. A-C. Kiirzweil 1900', D. E. Kurzweil 1902'. F. Kurzweil 1919. Scale bars: A. B. 1 mm; C-E. 20 pm.

are concave in M. albiclum (Figure 14B), whereas in M.

hijliae and M. hrevifoliiim (Figure 14C. D) they are roof-

like in shape, ± translucent and rather stiff in texture. In

the latter species, they do not cover the locules entirely

and are somewhat short. In fact, M. brevifoliwn is quite

distinct from other Machairophyllum species in its fruit

morphology: the expanding keels are thick, without awns

or wings, closing bodies are reduced to tiny knobs and

the fruit is consistently six-locular. In the other species,

closing bodies are small and frequently deep-set. Their

shape varies from rotund to kidney-shaped. In one speci-

men they were found to be slightly two-lobed (Kurzweil

1919). Although fruit morphology has been considered

number of stigmas

FIGURE 13. — Diagram showing number of stigmas in Machairo-

phyllum.

30

Bothalia 33,1 (2003)

FIGURE 14, — Capsules in Machairopliyllimr. A, closed; B-D, open. A, B, M. ulbidimi, Kiirzweil 1900', C, M. bijiiae, Kiirzweil 1919', D, M. bre-

vifoliitm. Kiirzweil 1904. Drawn by R Chesselet.

as the most important complex of characters for the

delimitation of genera in the Mesembryanthemaceae,

this does not seem to be the case in the current interpre-

tation of the genus Machairophyllum.

Seeds

The micromorphology of 29 specimens was examined

in the present study. Seeds are pear-shaped with a pro-

truding micropyle which conforms with the seeds of

other Mesembryanthemaceae (Figure 1 3 A) (Hartmann &

Gbiling 1993), 0.87(0.73-1 .00) x 0.64(0.44-0.77) mm,

dark brown to light yellowish brown. In the central seed

portion the testa ceils are predominantly elongate with a

strongly convex periclinai wall and have iiregulariy

undulating anticlinal walls (Figure 15B-E); neighbour-

ing cells almost touch each other. Very rarely, the anticli-

nal depressions between the cobblestone-like convex

testa cells are rather wide. The outer periclinai wall is

normally smooth to slightly verrucose. In Desmet 2158

most of the periclinai surface is smooth but the marginal

areas are rough (Figure 15D). The periclinai walls of the

testa exhibit epicuticular formations of various density

and shape. Epicuticular formations mostly consist of

minute, loosely or densely arranged rodlets (Eigure

15E-G), but flake-like epicuticular formations also occur

Bothalia 33,1 (2003)

31

FIGURE 15. — SEM micrographs of

seed moiphology in Machairo-

phyllum. The close-up micro-

graphs in B-G were taken in

the central seed portion. A,

seed; B-G, testa cells. A, B,

M. cf. hijliae, Bayer 7027:

C-G, M. hijliae. C. Kurzweil

1898: D, Desmet 2158: E, F,

Kurzweil 1916: G, Kurzweil

1925. Scale bars: A, 100 pm;

B-F, 10 pm, G, 5 pm.

(e.g. Desmet 2158; Kurzweil 1901) (Figure 15D). The

occurrence of spinulose rodlets was reported in Me-

sembryanthemoideae and a few genera of Ruschioideae

(Bittrich 1986; Ehler & Barthlott 1978). A crystal of

unknown nature and function was found on the testa just

above an anticlinal wall (Figure 15F).

Pollen

Pollen was examined in 33 air-dried specimens and

proved to be uniform (Figure 16). Pollen grains are tri-

colpate which is the common condition in the family

(Dupont 1977), equiaxial to strongly longiaxial, average

equatorial diameter 14.80 pm; average of the polar dis-

tance 15.36 pm. The comparatively small size observed

(compare with e.g. Dupont 1977 and Chesselet et al.

1998) is probably the result of hydration state. The sur-

face of the pollen grains is microspinulose as in the

majority of Mesembryanthemaceae (Hartmann 1991),

and the size and density of spinules is subject to slight

variation in the genus. Irregular perforations through the

tectum are visible in all specimens (Figure 16C). In M.

brevifolium (Lategan 2367/36) these are rather prominent

and the pollen is thus approaching a reticulate condition.

Phytogeography and ecology

All confirmed collections of Mochairophyllum come

from the southern parts of central South Africa (Western and

Eastern Cape) the only area that the genus is known from

with certainty (Eigure 2). The distribution range is in the

Little Karoo from Barrydale eastwards and in the adjacent

mountain ranges of the Swaitberg in the north and the

Langeberg in the south, and stretches further to Oudtshoom,

Uniondale, the Baviaanskloof and Kouga Mountains and to

Salt Pans Nek (near Jansenville); it is also known from the

Zuurberg (near Kirkwood north of Port Elizabeth) which is

the easternmost occurrence of the genus. Unfortunately many

of the older herbarium collections indicate only the district

where they were found and their exact locality is therefore

unknown. Populations aie rather localized within the distri-

bution range, but in some areas this may be the result of the

lack of suitable habitats or of low collecting activity.

Specimens referred to MachairophyUiim were also collected

in Port Elizabeth, near Mossel Bay, in De Hoop Nature

Reserve near Bredasdorp and near Laingsburg, but these

were later correctly identified as belonging to different gen-

era. In addition, there is also a record from the Richtersveld

of Northern Cape but this isolated occurrence needs confir-

mation (Figure 2). This spectacular collection was made near

Lekkersing about 10 years ago (Viviers s.n., NBG, hort.) —

some 600 km from the well-known distribution area of the

genus and also in an area where the prefemed habitat of

Machairophyllum is scarce. No plants were later found

despite an extensive search in the area (E. van Jaarsveld pers.

comm.). The unbranched inflorescences, the short bracteole

sheaths, as well as the number of sepals and locules suggest

that the plant belongs to M. hijliae. Another locality in

Northern Cape, situated in the Vanrhynsdorp District, was

shown on the distribution map of Machairophyllum present-

ed in Herre (1971 : 198) but no comment on the origin of the

information was made (Figure 2). However, the occurrence

of Machairophyllum in Northern Cape would not be too sur-

prising, as up to the 1930s ‘Little Namaqualand’ was thought

to be the distribution area of M. alhidum (see below).

32

Bothalia 33,1 (2003)

Machairophyllum albidiim occurs in the western part

of the distribution area of the genus, ranging from

Barrydale to around Oudtshoorn. An outlying locality is

known near Klaarstroom in the eastern Swartberg. The

widespread M. bijliae is known from the George and

Oudtshoorn Divisions eastwards to the Zuurberg. The

distribution areas of these two species overlap in the

Oudtshoorn District. M. brevifoUiim has so far only been

recorded in the red conglomerate hills between

Oudtshoorn and De Rust, and M. stayneri is only known

from the Zuurberg.

In the past there was considerable confusion around

the distribution of Machairophyllum. Linnaeus (1762),

when describing Mesembryanthemiim albidum (now

Machairophyllum albidum) gave its locality as ‘Habitat

in Aethiopia’ which merely implied that the plants came

from Africa. Sonder (1894), in his account in Flora

capeusis, quoted Namaqualand as the distribution area of

the sole Machairophyllum species known at the time, M.

albidum (as Mesembryanthemiim albidum). Sonder cited

two collections, one made by A. Wyley and the other by

Drege. The former collection was examined by Mrs L.

Bolus who provisionally referred it to the genus

Cheiridopsis (Bolus 1960: 156 [in the footnote]). The

latter could not be traced in the present study, and there-

fore it cannot be ruled out entirely that it is indeed a

species of Machairophyllum. Several subsequent authors

quoted the distribution ‘Namaqualand’ for M. albidum or

the entire genus Machairophyllum (e.g. Berger 1908;

Jacobsen 1935). However, both Berger and Jacobsen

probably merely adopted the distribution given by

Sonder. All subsequent collections of Machairophyllum

were only made in the southern part of central South

Africa, and therefore the occurrence of Machairophyllum

in Northern Cape was increasingly doubted (Schwantes

1957; 93).

Machairophyllum albidum and M. bijliae are most

commonly found on rocky outcrops (Figure lA, B), on

bare rock on N-exposed slopes and on rocky ridges in

mesic to dry fynbos or in renosterveld, commonly also

FIGURE 16. — SEM micrographs of

pollen morphology and sur-

face ornamentation in Ma-

chairophyllum bijliae. A, B,

pollen grains; C, surface

ornamentation. A, Foiircade

NBG424/24- B. Holland

NBGI697/36-, C, Ba.xter

NBG3J66/14. Scale bars: A,

B. 10 pm; C, 1 pm.

where fynbos and renosterveld meet. On one herbarium

collection a fynbos-valley bushveld mix is also given.

The occurrence of Machairophyllum in comparatively

moist fynbos is rather atypical in the family, as most

Mesembryanthemaceae are found in arid or semi-arid

areas in southern Africa, although this unusual habitat is

also that of a few other genera (e.g. Erepsia, Carpobro-

tus). The few accurate collector’s notes available indicate

that the species grow in rocky soil and bedrock derived

from TMS. The plants rarely occupy grassy habitats

amongst rocks over deeper soils (mainly in Eastern

Cape) and are then nearly hidden in tall vegetation

(Figure 1C). No accurate ecological information is given

on the type sheet of M. stayneri, but the area where the

species was collected generally falls in the Savanna

Biome (Rutherford & Westfall 1994) although fynbos

and renosterveld occur at higher altitudes in this area. M.

brevifoUiim is only known from the red conglomerate

hills near Oudtshoorn where it grows in fields of pebbles

interspersed with small shrubs.

Altitudes for the genus range from 330 m in the Little

Karoo to just below 1 600 m in the Swartberg. M.

albidum is usually found at low altitudes ranging from

330 to 600 m. M. bijliae is very much a high-altitude

species, with almost all collections made between 800 m

and 1 560 m (top of Swartberg Pass). The only exception

is a collection of this species made at 570 m near Jouber-

tina (Fourcade 2384\ BOL). The type locality of M. bre-

vifoliuni is between 550 and 600 m.

Flowering in Machairophyllum albidum and M. bijli-

ae is often stimulated by burning of the habitat, which

has been observed during the present study and has also

been reported before (J. Vlok pers. comm.; notes on a

herbarium sheet of M. albidum, Blackburn s.n.,

BOL63638). The fact that veld fires have a profound

effect on the flowering of plants in the Cape Floristic

Region is well documented in other families (e.g. Ama-

ryllidaceae, Iridaceae, Orchidaceae), but is not known in

Mesembryanthemaceae. However, other effects of veld

fires on mesembs have been reported (e.g. the rapid

Bothalia 33.1 (2003)

33

spreading of Erepsia species in response to burning:

Liede 1989, 1990; Hartmann 2001 ). Plants of Maclmiro-

phyUiim are normally not seriously damaged by veld

fires. Most plants grow in rocky sites where the fire

intensity is low. Dense clump formation ensures that

only the outer leaves are singed during the burning and

most adult individuals therefore survive the fire.

Furthermore, the seeds of the previous season are well

protected by the comparatively hard and robust capsules,

and plenty of seed is therefore available in the post-fire

environment. Flowering in M. brevifoliiim is apparently

not stimulated by burning.

Phytogeny

Machaiwphylhim in its current delimitation is fairly

uniform in vegetative, floral and several capsule features

which seems to suggest that it is indeed a natural group.

At present, the genus can be diagnosed according to the

following characters: clump-foiTning habit without visi-

ble stems; smooth and dotless and often waxen leaves of

a green or whitish green colour; large yellow flowers

which are open in the afternoon, evening or at night;

absence of filamentous staminodes; long filiform stig-

mas; capsules with raised sutures and reflexed valve

rims; covering membranes present; closing bodies small;

expanding keels serrate or lacerate and apically ± awned.

However, in a strict sense the morphological and ana-

tomical data obtained here are considered insufficient as

a basis for a sound phylogenetic analysis, as a definite

synapomorphy for the genus cannot be identified at this

stage. We therefore suggest that further data be obtained

for the entire Bergeranthus group, including molecular

ones, to resolve the phylogeny of Machairophyllitm.

While M. alhidum and M. bijliae are fairly similar in

details of the internal capsule structure (valve wings pre-

sent; covering membranes complete; high and variable

number of locules), M. brevifoliiim deviates somewhat

(valve wings absent; covering membranes incomplete;

low and stable number of locules) thus leaving doubt

regarding its systematic position. Nevertheless, until new

data become available, MachairopliyUiim is retained in

its current delimitation for practical reasons despite the

distinctness of M. brevifoliiim and we consider it prema-

ture to require that genera in the mesembs be defined on

the basis of synapomorphies.

TAXONOMY

Machairophyllum Schwaiites in Mollers Deutsche

Gartner-Zeitung 42: 187 (1927); Bolus: 155-158 (1960)

(synopsis). Type species: Machairophyllum alhidum (L.)

Schwantes.

Perissolobus N.E.Br: 278 (1930); N.E.Br.: 294 (1931).

Type species: Perissolobus bijlii N.E.Br.

Nearly stemless and tuft-like perennial herbs, entire-

ly glabrous, basally branched with age, sometimes form-

ing mats of up to 1.2 m diam. Leaves succulent and

trigonous, decussate, simple and entire, sessile, estipu-

late, crowded, linear-lanceolate (‘sabre-shaped’, ‘dag-

ger-shaped’) or rarely subrhomboid, 10-145 x 8-26 mm

and up to V4 as deep, base of a leaf pair connate to form

a sheath 5-18 mm long, leaf apex acute or acuminate;

surface smooth and dotless, waxen in appearance,

whitish green to green, often with a reddish tinge; old

leaves persisting on plant for several years and turning

black; stomata superficial to slightly sunken, outer epi-

dermal wall with well-developed crystal sand layer, epi-

cuticular wax mostly in the form of a solid and smooth

layer, rarely breaking up into platelets, local wax pro-

jections mainly in form of rodlets, rarely granules or

flakes. Inflorescences unbranched with a solitary flower

or dichasially branched with up to seven flowers, rarely

monochasial due to loss of one side branch, bracteolate;

bracteoles leaf-like, 7-130 x ± 5-10 mm, bases connate

for (5-)10-70(-100) mm. Flowers actinomorphic and

bisexual, large, showy, up to 65 mm diam., open in

afternoon, evening or at night, on 12-100 mm long

pedicels; pedicels sometimes elongating in fruit. Sepals

5-8, triangular-lanceolate, green with purplish tinge,

with whitish membranous margins, (3-)8-18(-22) x

(3-)4-16 mm, unequal, acute, acuminate or rarely

obtuse, outside often apically keeled and mucronate.

Petals (petaloid staminodes) numerous, densely 3-7-

seriate, linear or lorate, (8-) 15-35 x 0. 3-2.0 mm. obtuse

or acute, yellow or golden and frequently orange-red or

copper-coloured on outside and near tip, often paler near

base. Filamentous staminodes absent. Stamens erect,

forming a cone; filaments epapillate or only inner ones

inconspicuously papillate at base. (5-)7-15(-19) mm

long, whitish; anthers dehiscing with longitudinal slits,

anther and pollen yellow. Ovary usually obconic, syn-

carpous, 5-12(-15?)-carpellate, placentation parietal,

with many ovules per chamber. Nectar glands 5-12,

apparent as ± raised portions on a ring. Stigmas

(5)6-12(-15?), slender or filiform, usually slightly

shorter than stamens, (4-)8-15(-16) mm long. Fruit a

hygrochastic loculicidal capsule, 5-12(-15?) locular,

base funnel-shaped with slightly concave sides, sutures

on top strongly compressed, valve rims high and usual-

ly reflexed, valves mostly shortly and narrowly winged,

wings basally broader and tapering into fine awns dis-

tally, which are connate with awns of expanding keels,

expanding keels at first parallel and close to each other,

but diverging above, serrate or lacerate and mostly ter-

minating in slender awns which are directed towards

median axis of valve (often crossing each other), nearly

as long as valves, loculi with complete or almost com-

plete translucent to whitish covering membranes, clos-

ing bodies small and frequently deeply set, no addition-

al closing devices present. Seeds ± pyriform, ±

(0.7-)0. 8-1.0 mm long. Flowering occurs from spring

to early summer.

Key to species

la Inflorescence branched; bracteole sheatb mostly longer than

25 mm:

2a Sepals 5 or 6; stigmas 8-12; widespread 1 . M. albidum

2b Sepals and stigmas 6-8; only Zuurberg 2. M. stayneri

lb Inflorescence unbranched; bracteole sheatb mostly shorter

than 25 mm:

3a Leaves (20-)32-70(-85) x 8-15(-18) mm; sepals 5-8;

stigmas 5-10; widespread 3. M. bijliae

3b Leaves 10-30 x 10-26 mm; sepals and stigmas 6; current-

ly only known in red conglomerate hills to east of

Oudtshoorn 4. M. brevifoliiim

34

Bothalia 33,1 (2003)

1. Machairophyllum albidum {L.) Schwantes in

Mollers Deutsche Gartner- Zeitung 42: 187 (1927). Me-

seiiibiyanthemiim albidum L: 699 (1762). Bergeranthus

albidiis (L.) Schwantes: 180 (1926). Carriiandius albi-

diis (L.) Schwantes: 181 (1926) nom. inval. Iconotype:

Dilleniiis, Hortus Elthamensis: 243, t. 189, fig. 232 (1732).

Mesembryanthemum cookii L. Bolus: 2 (1925). Bergeranthus cookii

(L. Bolus) Schwantes: 180 (1926). Carruanthus cookii (L. Bolus)

Schwantes: 181 (1926) nom. inval. Machairophyllum cookii (L. Bolus)

Schwantes: 187 (1927). Type: Western Cape, 3320DC, Barrydale and

15 miles west, October, Compton & Cook NBG 1995/23 (BOL, holo.!).

Compact, tufted and nearly stemless, succulent herb,

entirely glabrous, basally branched with age. Leaves

trigonous, linear-lanceolate, entire, sessile, 40-145 x

9-20 mm and up to V4 as deep, apically acute to acumi-

nate, connate sheath 7-17 mm long, surface smooth and

waxen in appearance, whitish green or green. Inflo-

rescence branched, up to 170 mm tall, with 2-7 flowers;

bracteoles leaf-like, (25-)50-130 x 5-8 mm, with a con-

nate sheath (17-)42-80(-100) mm long. Flowers large

and showy, 45-65 mm diam., vespertine, nocturnal or

open from 15h00 onwards and often closing again in the

evening; pedicels up to 100 mm long. Sepals 5 or 6, tri-

angular-lanceolate, 6-20 X 5-16 mm, unequal, green

with a purplish tinge, with whitish membranous margins,

apically ± acute. Petals ± 400, linear or lorate, 18-35 x

(0.3-)0.6-1.0(-1.5) mm, inside yellow, frequently orange-

red or copper-coloured on outside and near tip. Stamens

numerous, erect and forming cone, 10-15(-19) mm

long; filaments white; anthers yellow. Ovary obconic,

(8-)10-12(-15?)-carpellate, placentation parietal, with

many ovules per chamber. Nectar glands as ± raised por-

tions on whitish ring, (8-)10-l2, green. Stigmas

(8-)10-12(-15?), slender, 8-15(-16) mm long. Fruit a

loculicidal capsule 12-15 mm diam., base funnel-

shaped, sutures on top compressed, valve rims high and

diverging; with (8-)10-12(-15?) locules, valves 4. 0-5. 6

X 3. 2-3. 5 mm in size, narrowly winged; covering mem-

branes complete; closing bodies small; expanding keels

lacerate, parallel and adjacent at base, but soon diverg-

ing, apically awned. Flowering time: mainly November

and December.

Distribution, habitat and biology

The species is mainly known in the Little Karoo from

Barrydale to the George and Oudtshoorn Divisions

which is the western part of the distribution area of the

genus. On Cloetespas near Herbertsdale, the species

extends over the coastal mountain range. M. albidum has

also been collected near Klaarstroom in the eastern part

of the Swartberg {Burger BOL38796, BOL) which is an

outlying occurrence of the species (Ligure 2). M.

albidum grows in fynbos and renosterveld, mostly asso-

ciated with rocky habitats, from 330-600 m.

Similar species

Vegetatively Machairophyllum bijliae is largely iden-

tical to M. albidum which also occurs in the same type of

habitat although M. bijliae frequently has shorter leaves.

In the flowering stage the difference is immediately

obvious, as M. bijliae consistently has solitary flowers.

whereas inflorescences of M. albidum are branched. On

average M. bijliae has also slightly smaller flowers with

shorter petals. M. bijliae differs further by mostly having

shorter bracteole sheaths, a higher number of sepals, and

ovaries with fewer locules. While the centre of distribu-

tion of M. bijliae is further east, the distribution areas of

the two species overlap in the Oudtshoorn area. M. bijliae

is always found above 800 m, whereas M. albidum is

generally a low-altitude species.

Vouchers: Compton & Cook NBG1995/23 (BOL); Hall BOL32083

(BOL); Kurzweil 1890. 1900 (NBG); Muir 3937 (K).

2. Machairophyllum stayneri L.Bolus in Journal

of South African Botany 26: 155 (1960). Type: Eastern

Cape, 3325BC, summit of Zuurberg Pass, October 1958,

Stayner KG 196/59 (BOL, holo.!).

Densely compact, nearly stemless, entirely glabrous,

basally branched with age, sometimes forming mats up

to 1.2 m diam. Leaves trigonous, entire, linear-lanceo-

late, 70-80(-100?) X 13-16 mm and up to V4 as deep,

apex acute or acuminate, basal sheath 10-15 mm long;

surface smooth and waxen. Inflorescence branched, up to

150 mm tall, lateral buds develop ‘tardily’ (fide Bolus

1960); bracteoles up to 52 mm long, basally connate for

± 15 mm (fide Bolus 1960). Flowers nocturnal, 50-62

mm diam.; pedicels up to 33 mm. Sepals 6-8, long-

acuminate, 14—20 X 5-8 mm. Petals in 5 or 6 series, 17-27

X 0. 5-1.0 mm, yellow to golden and orange to pink on

outside, basally paler. Stamens 13-17 mm. Nectar glands

6-8. Stigmas 6-8, thick-filiform, 12-14 mm long. Fruit a

loculicidal capsule with 6-8 locules. Flowering time:

October.

Distribution, habitat and biology

The species is known only from the type collection

which was made on the Zuurberg in the Eastern Cape

(Ligure 2). No ecological information was given on the

type sheet.

Note

The only collection of this species, consisting of a

complete plant and several loose fragments, was made

over forty years ago. Despite an extensive search by the

first author, no plants could be located. The number of

sepals and locules associated with short bracteole sheaths

and branched inflorescences suggest that the species is

distinct. The branched inflorescence seems to point to a

close relationship with Machairophyllum albidum.

Alternatively, the short bracteole sheaths as well as floral

characters may indicate affinities to M. bijliae.

If the species is recollected, future studies based on

more material may well suggest that Machairophyllum

stayneri is merely a form of one of the other species. It

must be noted that all characters except the branched

inllorescence are well within the range of variation of M.

bijliae which also shares a similar distribution. Thus it

cannot be ruled out entirely that the specimen in question

is an abnormal plant of M. bijliae.

Vouctier: .Stayner KG 196/59 (BOL).

Bothalia 33,1 (2003)

35

3. Machairophyllum bijiiae (N.E.Br.) L.Bolus,

Notes on Mesembryanlhemum and allied genera 3: 135

( 1938), as bijUi. Perissolobiis bijiiae N.E.Br: 294 (1931),

as bijlii. Type: Eastern Cape, Somerset East Div., no

date. Van der BijI 93 (K, holo.-NBG, photo!).

Machairophyllum stenopetalum L.Bolus: 303 ( 1931 1. Type: Eastern

Cape, near Willowmore, August 1931, Brauns SUG8030 (BOL, holo.!).

M. acuminatum L.Bolus: 485 (1935). Type: Eastern Cape, near

Humansdorp, October 1934, Fourcade NBG424/24 (BOL, holo.!).

M. baxteri L.Bolus: 486 (1935). Type: Western Cape, near George.

October 1934, Baxter NBG3 1 66/1 4 (BOL, holo.!).

M. vanbredai L.Bolus: 238 (1964). Type: Eastern Cape. 3325BC,

Witwater Farm, Somerset East Div., August 1962. Van Breda 1749/62

(BOL. holo.!).

Plants compact, nearly stemless, entirely glabrous,

basally branched with age. Leaves trigonous, linear-lance-

olate, entire, (20-)32-70(-88) x 8-15(-20) mm and up to

V4 as deep, connate for (5-)8-13(-18) mm, apex acute or

acuminate; surface smooth, waxen in appearance, whitish

green to green, often with reddish tinge. Inflorescence

unbranched; bracteoles leaf-like, ( 18-)25-40(-60) x

5-8(-10) mm, connate for 7-18(-42) mm. Flowers soli-

tary, 30-55 mm diam.; pedicels ( 13.5-)30.0-70.0(-100.0)

mm and up to 38 mm diam., elongating up to 130 mm in

fruit; flowers mainly nocturnal (opening late afternoon or

evening and staying open all night). Sepals (5)6-8,

unequal, leaf-like, triangular-lanceolate, (3-)8-22 x

3-8(-ll) mm, apex acute, acuminate or rarely obtuse,

green with purplish tinge, with whitish membranous mar-

gins. Petals numerous, linear to lorate. (8-)15-25(-28) x

0.3-1. 0(-2.0) mm, yellow, outsides often orange-yellow

to copper-coloured, base pale yellow. Stamens erect, form-

ing cone, (5-)7-14(-15) mm long; filaments without sig-

nificant papillae, white; anthers yellow. Ovary obconic,

(5)6-8(-10)-carpellate. with many ovules per chamber.

Nectar glands as (5)6-8(-10) ± raised portions on ring,

green. Stigmas (5)6-8(-10), filiform, (4— )7-14 mm long.

Fruit a loculicidal capsule, up to 13 mm diam., up to 24

mm when expanded, base funnel-shaped, sutures on top

compressed, valve rims high and diverging, capsule with

(5)6-8(-10) locules, valves 4.0-5. 3 x 3. 8-5. 8 mm with

narrow wings with terminal awns; expanding keels serrate

to lacerate, parallel and adjacent at base, but soon diverg-

ing, with terminal awns; covering membranes complete,

translucent to whitish; closing bodies small. Seeds brown,

± 1 mm long. Flowering time: mainly between August and

November.

Distribution, habitat and biology

This is the most widespread Machairophyllum species.

It is known from many collections which were made in a

broad band from the Oudtshoom Division, the Swartberg

Mountains, the Willowmore District, the Baviaanskloof

and Kouga Mountains and Jansenville to the Zuurberg

near Kirkwood in Eastern Cape (Figure 2). M. bijiiae

grows in fynbos and renosterveld, mostly associated with

rocky habitats. Most collections were made at altitudes

of between 800 and 1 300 m, but one collection has been

made near Joubertina at 570 m (Fourcade 2384; BOL).

The highest record comes from near the top of Swartberg

Pass at 1 560 m.

Similar species

Machairophyllum bijiiae vegetatively resembles M.

albidum. Differences are given under this species.

Note

The original spelling "bijlii'' is a mistake in gender

because the species was named after Mrs D. van der Bijl.

Vouchers: Baxter 3166/14 (BOL); Desmet 2158 (NBG); Tavlor

NBG1556/33 (BOL); Van Breda 1761/61 (BOL).

4. Machairophyllum brevifolium L.Bolus, Notes

on Mesembryanthemum and allied genera 3: 126 ( 1938).

Type: Western Cape, 3322CB, near Oudtshoom, August-

September 1937, Lategan NBG2369/36 (BOL, holo.!).

Machairophyllum latifolium L.Bolus: 126 (1938). Type: Western

Cape, near Oudtshoom. February 1938 (hort?). Lategan NBG2367/36

(BOL. holo.!).

Plants nearly stemless, compact, sometimes forming

mats with age, entirely glabrous. Leaves trigonous,

lanceolate or subrhomboid, entire, 10-30 x 10-26 mm,

12-15 mm deep, apically acute, connate for ± 8-10 mm;

surface smooth, waxen in appearance, whitish green to

green. Inflorescences unbranched; bracteoles 7-23 mm

long, connate for ±5-10 mm. Flowers solitary on a pedi-

cel 12-15 mm long, open in the afternoon or vespertine,

30^5 mm diam. Sepals 6, unequal, broadly triangular-

lanceolate, 8-14 X 4-8 mm. Petals numerous, lorate, 3 or

4-seriate, 13-20 x 1-2 mm, yellow. Stamens 9-10 mm

long; filaments without conspicuous papillae at base;

anther and pollen yellow. Ovary 6-carpellate, with many

ovules per chamber. Nectar glands 6. Stigmas 6, filiform,

10-12 mm long. Fruit a loculicidal capsule with 6

locules, base funnel-shaped, sutures on top compressed,

valve rims diverging; valves ±3x4 mm. without wings;

expanding keels lacerate, basally parallel and adjacent to

each other but soon diverging, acute, but without awns;

loculi largely covered by translucent covering mem-

branes; closing bodies very small. Seeds brown, ± 1 mm

long. Flowering time: August and September.

Distribution, habitat and biology

The species is known from only three collections made

at or near the type localities of Machairophyllum brevi-

folium and the conspecific M. latifolium in the red con-

glomerate hills between Oudtshoom and De Rust (Figure

2). In the population which was visited by the first author

(at the Farm ‘Skuinspad’) ± 50 plants grew among small

pebbles in rather open shrub vegetation next to the edge

of a cliff. The type locality of M. brevifolium is at ±

500-600 m altitude. A few collections of this species

were also made elsewhere, but their identity is doubtful.

Note

Bolus (1938: 126) pointed out the striking resem-

blance of this species to the genus Faiicaria, referring to

it as a possible ‘connecting link’. This does not necessar-

ily imply a close affinity in a phylogenetical sense al-

though a relationship of the two genera would not be

36

Bothalia 33,1 (2003)

very unlikely (J. Vlok pers. comm.). Machaiwphyllum

and Faiicaria were not considered as being related in the

preliminary grouping presented by Hartmann (1991).

However, in the new classification of Chesselet et ah

(2002) they are grouped together in the Delospermeae.

Vouchers: Lategan NBG2367/36. NBG2369/36 (BOL).

Unknown species

Three names, ^ Machairophyllum minor L.B’ (‘L.B.’

probably stands for L. Bolus; Nel SUG8313; BOL), ‘M.

minus' (Tiigwell & Levenson NBG2634/14\ BOL) and M.

imiondalensis (Hammer 1991), could not be traced and

have never been validly described.

NOTES ON SOME PRESUMABLY RELATED GENERA (TABLE 2)

Bijlia and Carnianthus

Bijlia N.E.Br is a genus of two species known from

around the Great Karoo town of Prince Albert and from

kloofs on the northern slope of the Swartberg to the west

of it. A detailed explanation of the complicated nomen-

clature of the then sole species B. dilatata H.E.K. Hart-

mann was presented by Hartmann (1992). A second spe-

cies, B. tiigwelliae (L.Bolus) S. A. Hammer was added later

(Hammer 1995). The genus Carriiandms (Schwantes)

Schwantes, from the Karoo near Willowmore, comprises

two species, C. ringens (L.) Boom and C. peersii L.Bolus.

The latter species was placed in the monotypic genus

Tischleria Schwantes on account of having a slightly dif-

ferent capsule structure, but is now also included in

Carruanthus (Herre 1971; Dyer 1975; Smith et al. 1998;

Chesselet et al. 2000; Hartmann 2001).

Plants of Bijlia and Carruanthus are compact and

glabrous herbs like Machairophyllum, and have trigo-

nous leaves. In their morphology, the leaves of Carru-

anthus are similar to those of Machairophyllum but dif-

fer by having prominent teeth on the margins. Anatomi-

cally the leaves of Bijlia and Carruanthus are very simi-

lar to those of Machairophyllum. Their surface has a

solid wax cover which breaks up to form platelets

(Eigure 17A, B). Local wax projections in the form of

rodlets, granules or flakes (common in Machairo-

phyllum) appear to be scarce in Bijlia and Carruanthus.

Elowers of Bijlia and Carruanthus are solitary or are

arranged in few-flowered cymes, the short peduncles

usually hidden among the leaves. Pedicels of Bijlia are

up to 10 mm long and those of Carruanthus up to 100

mm long. Both genera have five subequal to unequal

sepals, linear-lanceolate petals and numerous stamens.

Eilamentous staminodes are absent. In Bijlia tiigwelliae

the nectaries are visible as raised portions of a ring

(Figure 17D-F) which is also the situation in Machairo-

phyllum. Capsules of Bijlia and Carruanthus are five-

locular, and have strongly raised sutures. In Carruanthus

these have reflexed valve rims which is shared with

Machairophyllum. In contrast, the rims are erect and

straight in Bijlia. While the capsule opens up, in Bijlia

the valves separate only slightly from each other, where-

TABLE 2. — Various features in the genera Machairophyllum, Carruanthus, Bijlia and Cerochlamys

Bothalia33,l (2003)

37

as they are fully or largely spreading in Carmanthus.

Valve wings are absent in Bijlia but are developed as

short and narrow flaps in Carnicmthiis. The covering

membranes are complete in Bijlia and Carnianthiis t in-

gens but are only developed as a small limb in C. peer-

sii. Closing bodies are large, white bulges in Bijlia but

are small or absent in Carmanthus. The general shape of

the seeds of Bijlia and Carmanthus does not differ

markedly from that of Machairophyllum. Seeds are

ovoid to pear-shaped and ± 0.6-0. 9 mm long. In both

Bijlia species the testa cells are extended into papillae

(Figure 17G). and have epicuticular formations in the

form of granules. Seeds of Carmanthus are very similar

to those of Machairophyllum in that they have elongate

and strongly convex testa cells with coarsely undulate

anticlinal walls (Figure 17H). Epicuticular formations in

Carmanthus are also very similar and mainly take the

shape of rodlets.

Concluding remarks: a similar leaf shape and anato-

my, elongate-pedicelled flowers, capsules with raised

sutures with reflexed valve rims and short and narrow

valve wings, as well as a strikingly similar seed ultra-

structure mark clear affinities of the genera Carmanthus

FIGURE 17. — SEM micrographs of

various features in Bijlia.

Carmanthus and Cerochla-

niys. A-C, leaf surface: A,

Bijlia tugwelliae, Kurzweil

1927: B, Carmanthus peer-

sii: C, Cerochkunys pachy-

phylla. D-F. nectaries of

Bijlia tugwelliae, NBG702/86.

G-I, seed surface: G, Bijlia

dilatata. NBG526/6H: H, Car-

ruanthus peersii: I. Cerochla-

mvs pachyphylla. B. H, Kurz-

weil 1926: C. I, Kurzweil

1905. Scale bars: A-C, F-I,

20pm;D, E, 500 pm.

and Machairophyllum and suggest that the two are sister

groups. The small size or absence of closing bodies in

both Carmanthus species and the lack of a complete

covering membrane in one (C. peersii) is probably the

result of reduction.

Cerochlamys

Cerochlarnys N.E.Br comprises three species in the

Swellendam, Oudtshoorn and Laingsburg Districts in

Western Cape. A fourth species was recognized by

Hartmann ( 1998b) but was referred to the genus Acrodon

N.E.Br (Burgoyne 1998). The following brief comments

refer to the actual examination of C. pachyphylla

(L. Bolus) L. Bolus, while information on the other species

was largely obtained from the literature (mainly Hart-

mann 1998b).

All three species are compact and nearly stemless

succulents with smooth, clavate or trigonous leaves. The

leaf surface of Cerochlamys pachyphylla and C. gemina

(L. Bolus) H.E.K. Hartmann has a solid wax cover which

breaks up to form platelets (Figure 17C) which also

occurs in C. trigona N.E.Br (Hartmann 1998b). In C. pachy-

38

Bothalia 33,1 (2003)

phylla {Kurzweil 1905) the main leaf vascular bundles

was found to be strongly curved or concentric with col-

lenchymatic sheaths.

The short-pedicelled flowers of Cerochlamys are soli-

tary or arranged in few-flowered cymes. Flowers are

pink, purple or white, and are open during the day.

Cerochlamys has mostly five subequal to unequal sepals

(except six in C. gemina), linear-lanceolate petals and

numerous erect stamens in a cone. All species have white

filamentous staminodes. Nectaries were reported as sep-

arate (Herre 1971) and as ‘... very broad, touching each

other, sometimes apparently in a ring’ (Hartmann 1998b:

53). The stigmas of the three species are comparatively

short. Capsules are five-locular in C. pachyphylla and C.

trigona, but six-locular in C. gemina. In C. pachyphylla

the sutures on top of the capsule are strongly raised, and

have straight valve rims. Valves are erect in the opened

position and lack wings. The roof-shaped covering

membranes are complete. Expanding keels are diverging

and terminate in long awns. Closing bodies are absent in

Cerochlamys although the placenta ends are often knob-

shaped (Hartmann 1998b). Seeds are ovoid to pear-

shaped and ± 0.6-0. 9 mm long. The isodiametric to elon-

gate testa cells are extended into papillae (Figure 171).

Concluding remarks: floral and fruit characters of

Cerochlamys, particularly, the filamentous staminodes,

the short stigmas, its flowering phenology with diurnal

pink, purple or white flowers are very unusual in the

Bergerantlnis group suggesting that the genus may not

be correctly placed here. While resolving the correct

taxonomic position of Cerochlamys is obviously beyond

the scope of the present paper, the genus is here not con-

sidered as part of the Machairophylliim complex.

SPECIMENS EXAMINED

Herbarium abbreviations in accordance with the latest

version of Index Herbariorum, BOL* = Pickle collection

at BOL; NBG = National Botanic Garden Kirstenbosch,

SUG = Stellenbosch University Garden.

Acocks 15999 (3) BOL.

Baxter NBG3166/14 (3) BOL. Bean 1091 (3) BOL. Blackburn 2,

BOL63638 (1 ) BOL. Brauns SUG8030 (3) BOL. Burger BOL38796 ( I )

BOL.

Cilliers SUG10742 ( 3) BOL. Clarke NBG930/20 (3) BOL.

Compton & Cook NBG1995/23 (1 ) BOL.

Desmet 2076, 2147, 2158 (3) NBG.

Esterhuysen 16231, 24894 (3) BOL.

Fourcade 2384, NBG424/24 (3) BOL.

Hall BOL25194. BOL32083. NBG477/53, NBG522/52 (1) BOL;

NBG223/68 (1) NBG; Herre SUG12178 (3) BOL. Holland

NBG 1697/36 (3) BOL.

Kurzmdl 1890, 1900, /902 (1) BOL*, NBG; 1896, 1898, 1899, 1901,

1913, 1916, 1918, 1925 (3) BOL*; 1919 (3) BOL*, NBG; 1904 (4) BOL*.

Lategun NBG2367/36, NBG2369/36 (4) BOL.

Murkoelier SUG 10661 (3) BOL. Muir 3937 ( I ) K.

Nel SUG8313 (3) BOL.

Peers BOL32084 (3) BOL. Pillans 915, NBG496/I6 (1) BOL.

Pocock NBG627/26 (3) BOL, S. 246 = NBG661/28 (3) BOL.

Stayner KG196/59 (2) BOL; NBG177/88 (3) hort. ‘student’

SUG1004\ (3) BOL.

Taylor BOL32085, BOL63637, NBG1556/33 (3) BOL; NBG858/30

(I) BOL. Tugwell & Levenson NBG2634/14 (3) BOL.

Van Breda 1196, 1749/62, 1761/61 (3) BOL. Van Jaarsveld 07082

= NBG 769/83 (3) NBG; 07264 = NBG 806/83 (3) hort. Van der Bijl

NBG1465/33 (1) BOL. Viviers s.n. (3) hort.

Wisura 2159 (3) NBG. Without collector: 467b, SUG82 74 (3)

BOL;St/G S9S4(l)BOL,

ACKNOWLEDGEMENTS

Most of all we would like to express our thanks to Mr

Jan Vlok (Oudtshoom) for sharing his invaluable experi-

ence and for providing input into the manuscript. The

collecting work was made possible by the Western Cape

Nature Conservation Board and the Department of

Economic Affairs, Environment and Tourism of the

Province of the Eastern Cape who issued the necessary

collecting permits. The first author would also like to

thank Dr R.J. Chinnock and Mr Deon Viljoen for their

assistance during the collecting work. Herbarium mater-

ial was obtained on loan from BOL and K, and thanks are

also due to the staff of the herbaria PRE, SBT and UPS

for sending photocopies/photographs of specimens.

Useful advice was also provided by Prof. Dr H.-D.

Ihlenfeldt and Dr H.E.K. Hartmann (both Germany). We

are also grateful to Ernst van Jaarsveld (Kirstenbosch,

Cape Town) for help in many ways. Bruce Bayer, Phillip

Desmet, Colin Patterson-Jones and Mike Viviers are

thanked for providing additional specimens. We are also

indebted to the staff of the Electron Microscope Unit at

the University of Cape Town for providing SEM time.

The study was carried out at the National Botanical

Institute, and we thank the curators of the Compton

Herbarium and the National Botanical Garden, Kirsten-

bosch for the use of their collections.

RELERENCES

BERGER, A. 1908. Mesembryanthemen und Portulacaceen. Ulmer,

Stuttgart.

BITTRICH, V. 1986. Untersuchungen zu Merkmalsbestand, Gliederung

und Abgrenzung der Unterfamilie Mesembryanthemoideae

(Mesembryanthemaceae Lenzl). Mitteilungen aiis dem Institut

fur Allgemeine Botanik Hamburg 21: 5-1 16.

BOLUS, H.M.L. 1925. Novitates Africanae. Annals of the Bolus

Herbarium 4: 1-30.

BOLUS, H.M.L. 1931. Notes on Mesembryantlienium and allied gen-

era, part 2: 303. University of Cape Town.

BOLUS, H.M.L. 1935. Notes on Mesembryanthemum and allied gen-

era, part 2: 485^87. University of Cape Town.

BOLUS, H.M.L. 1938. Notes on Mesembryanthemum and allied gen-

era, part 3: 126. 127, 135. University of Cape Town,

BOLUS, H.M.L. 1950. Notes on Mesembryanthemum and allied gen-

era, part 3: 228, 229. University of Cape Town.

BOLUS, H.M.L. 1960. Notes on Mesembryanthemum and allied gen-

era. Journal of South African Botany 26: 155-164.

BOLUS, H.M.L. 1964. Notes on Mesembryanthemum and allied gen-

era. Journal of South African Botany 30: 231-243.

BROWN, N.E. 1930. Mesembryanthemum and some new genera sepa-

rated from it. Gardeners’ Chronicle 88, ser. 3: 278.

BROWN, N.E. 1931. Mesembryanthemum and some new genera sepa-

rated from it. Gardeners’ Chronicle 89, ser. 3: 294, 295.

BURGOYNE, R 1998. Rinding a place in the sun: where does Ruschia

purpureostyla belong? Aloe 35: 60, 61,

CHESSELET, P., HARTMANN, H.E.K., HAHN, N., BURGOYNE, P.

& SMITH, G.L. 1998. Taxonomic notes on the genus Khadia

(Mesembryanthemaceae/Aizoaceae). Bothalia 28: 25-33.

Bothalia 33,1 (2003)

39

CHESSELET, R, SMITH, G.F., BURGOYNE, P.M., KLAK. C„

HAMMER, S.A., HARTMANN, H.E.K., KURZWEIL, H„

VAN JAARSVELD, E.J., VAN WYK, B.-E. & LEISTNER,

O.A. 2000. Mesembryanthemaceae. In O.A. Leistner, Seed

plants of southern Africa: families and genera. Strelitzia 10:

360-410. National Botanical Institute. Pretoria.

CHESSELET, P, SMITH. G.F. & VAN WYK, A.E. 2001. A new trib-

al classification of the Mesembryanthemaceae Eenzl based on

characters of the floral nectary. Aloe 38: 25-28.

CHESSELET. P. SMITH. G.F. & VAN WYK, A.E. 2002. A new trib-

al classification of the Mesembryanthemaceae: evidence from

floral nectaries. Taxon 51: 295-308.

DANNEMANN, J.F. 1883. Beitrdge zur Kenntniss der Anatomie imd Ent-

wickhing der Mesembryanthema. Ph.D. thesis. University of Halle.

DILLENIUS. J.J. 1732. Hortus Eltiumiensis: 243. London.

DUPONT, S. 1976. Nouvelles observations surdes stomates de Mesem-

bryanthemacees. Actes dii 97‘ congres National des Societes.

Savantes Nantes, section des sciences, tome IV: 371-390.

DUPONT. S. 1977. Notes on the pollen of the Mesembryanthemaceae.

Cactus and Succulent Journal of Great Britain 39: 57-63.

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

vol. 1 (Dicotyledones). Department of Agriculture, Pretoria,

EHLER. N. & BARTHLOTT, W. 1978. Die epicutikulare Skulptur der

Testa-ZeUwiinde einiger Mesembryanthemaceae. Botanische Jahr-

biicher 99: 329-340.

GERSTBERGER, P. & LEINS, P. 1978. Rasterelektronenmikroskopische

Untersuchungen an Bliitenknospen von Physalis philadelphica

(Solanaceae). Anwendung einer neuen Praparationsmethode.

Berichte der Deutschen Botanischen Gesellschaft 91: 381-387.

HAMMER. S.A. 1991. Friendly daggers, or notes on Machairo-

phyllum. Mesemb Study Group Bulletin 4: 77, 78.

HAMMER. S. 1995. Mastering the art of growing mesembs. Cactus

and Succulent Journal (US) 61: 195-247.

HARTMANN. H.E.K. 1976. Monographic der Gattung Odontophorus

N.E.Br, (Mesembryanthemaceae Eenzl) (Monographic der sub-

tribus Leipoldtiinae Schw. I). Botanische Jahrbucher 91: 161-225,

HARTMANN, H.E.K. 1977. Monographic der Gattung Argyroderma

N.E.Br. (Mesembryanthemaceae Eenzl). Mitteilungen aus clem

Instinct fiir Allgemeine Botanik Hamburg 15: 121-235.

HARTMANN. H.E.K. 1982. Monographien der Subtribus Leipoldtiinae.

III. Monographic der Gattung Fenestraria (Mesembryan-

themaceae). Botanische Jahrbiicher 103: 145-183.

HARTMANN, H.E.K. 1988. Fruit types in Mesembryanthema.

Beitrdge zur Biologic der Pflanzen 63: 313-349.

HARTMANN, H.E.K. 1991. Mesembryanthema. Contributions from the

Bolus Herbarium 13: 75-157.

HARTMANN, H.E.K. 1992. A new name for Bijlia cana (Mesembry-

anthema). Cactus and Succulent Journal ( US) 64: 173-180.

HARTMANN. H.E.K. 1993. Aizoaceae. In K. Kubitzki, J.G. Rohwer

& V. Bittrich, The families and genera of vascular plants, vol.

2: 37-69. Springer Verlag, Berlin.

HARTMANN, H.E.K. 1998a. A letter to the MSG. Mesemb Study

Group Bulletin 2: 32-34.

HARTMANN, H.E.K. 1998b. New combinations in Ruschioideae,

based on studies in Ruschia (Aizoaceae). Bradleya 16: 44—91.

HARTMANN, H.E.K. (ed.) 2001. Illustrated handbook of succulent

plants: Aizoaceae (in 2 parts). Springer Verlag, Berlin, Heidel-

berg, New York.

HARTMANN, H.E.K. & GOLLING, H. 1993. A monograph of the

genus Glottiphyllum (Me.sembryanthema, Aizoaceae). Bradleya

11: 1-49.

HERRE, H. 1971. The genera of the Mesembryanthemaceae. Tafel-

berg. Cape Town and Johannesburg.

IHLENFELDT. H.-D. 1989. Life strategies of succulent desert plants.

Excelsa 14: 75-83.

IHLENFELDT. H.-D. & HARTMANN, H.E.K. 1982. Leaf surfaces in

Mesembryanthemaceae. In D.F. Cutler, F.L. Alvin & C.E. Price,

The plant cuticle: 397M^23. Academic Press, London.

JACOBSEN, H. 1935. Succulent plants — description, cultivation and

uses of succulent plants, other than cacti. Williams & Norgate,

London.

JACOBSEN. H. 1977. Lexicon of succulent plants, edn. 2 [revised and

enlarged edition]. Blandford Press, Poole.

KURZWEIL, H. & LINDER-SMITH, C. 2001. Machairophyllum van-

bredai. Flowering Plants of Africa 57: 64—69. t. 2172.

LIEDE, S. 1989. Untersuchungen zum Merkmalsbestand und zur

Taxonomic der 'Erepsiinae' (Mesembryanthemaceae). Beitrdge

zur Biologie der Pflanzen 64: 391—479.

LIEDE, S. 1990. Studies in the genus Erepsia — on the ecology of a

fynbos succulent. Mitteilungen aus dem Imstitut fiir Allgemeine

Botanik Hamburg 23b: 759-766.

LINNAEUS, C. 1762. Species plantarum, edn 2. Stockholm.

METCALFE. C.R. & CHALK, L. 1950. Anatomy of the Dicotyledons.

Clarendon Press, Oxford.

RAPPA. F. 1912. Per una classificanzione naturale dei Mesembrian-

themi. Bolletin Reale Orto Botanico e Giardino Coloniale di

Palermo 11: 21-36.

RUTHERFORD. M.C. & WESTFALL, R.H. 1994. Biomes of southern

Africa: an objective categorization. Memoirs of the Botanical

Sun'ey of South Africa No. 63: 1-94,

SALM-DYCK, J.M.F.A.H.I. 1849. Mesembryanthemum albiduni.

Monographia generum Aloes et Mesembryanthemi. Bonn,

SCHWANTES. G, 1926. Bergeranthus. Zeitschrift fiir Sukkulentenkunde

II: 180. 181.

SCHWANTES, G. 1927. Einige neue Mesembriaceen. Mdllers Deutsche

Gdrtner-Zeitung 42: 187.

SCHWANTES, G. 1957. Flowering stones and mid-day flowers. Benn,

London.

SMITH. G.F., CHESSELET, P, VAN JAARSVELD, E.J., HART-

MANN, H.E.K., HAMMER, S., VAN WYK, B.-E., BUR-

GOYNE, P, KLAK, C. & KURZWEIL, H. 1998. Mesembs of

the world. Briza Publications, Pretoria.

SONDER, W. 1894. Mesembryaceae. In W.H. Harvey & O.W. Sonder,

Flora capensis 2: 386-479. Reeve, London.

Bothalia 33,1: 41-47 (2003)

A new serotinous species of Cliff ortia L. (Rosaceae) from Northern

Cape, South Africa and section Arboreae emended

A.C. FELLINGHAM *

Keywords: Clijfortia L., new species, serotiny, taxonomy

ABSTRACT

A new species, Clijfortia dichotoma Fellingham, from the Oorlogskloof Escarpment, near Nieuwoudtville, Northern

Cape, is described and compared to its closest allies in the genus, Clijfortia arborea Marloth, which is widespread along

the escarpment of the Great Karoo from Calvinia in the north to Beaufort West in the southeast, and C. conifera E.G.H.Oliv.

& Fellingham from the Anysberg near Laingsburg, further east in the Karoo. The description of the section Arboreae is

emended here to include the new species, and the description of C. aborea is corrected.

INTRODUCTION

The more than one hundred species of the genus

Clijfortia L. represent an amazing variety of growth

forms. These range from delicate, herbaceous ground

covers to shrubs of various shapes and sizes, as well as

trees. Until recently only two trees had been recognized

and placed in the section Arboreae Weim. emend.

(Oliver & Fellingham 1994).

A third tree-like species was brought to my attention

by Prof. D.J. von Willett, who took it to be the well-

known C arborea. It is indeed more closely related to C.

arborea than to the third species in this section viz. C.

conifera, but also distinctly different from both.

Not only does the growth form set these tree-like

species apart from the rest of the species in the genus, but

they are also distinct in having highly condensed female

inflorescences. Furthermore, these inflorescences are

borne terminally on main stems or main branches and

always near the top of the trees, whereas the male inflores-

cences, which are much smaller and less condensed, are

borne on the same plants but lower down and on lateral

branchlets. This arrangement of male and female flowers

is rare in wind-pollinated plants (Proctor et al. 1996).

The similarity in these three species also includes

their habitats. In each instance it is the edge of an escarp-

ment, in fynbos in the cases of C. conifera and C.

dichotoma and in karroid shrubland for C. arborea

(Marloth 1905; Oliver & Fellingham 1994).

According to the owners of the Farm Papkuilsfontein

where the new species occurs, the known population had

not been subjected to fire for more than 20 years. This

could account for the fact that there are no seedlings or

young trees present and many trees are partly dead.

In this paper the terminology used by Weberling

(1981, 1983) for describing the structure of the inflores-

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

7735 Claremont. Cape Town. Present address: 23 Moreson Avenue,

Valmary Park, 7550 Durbanville, South Africa.

MS. received: 2002-05-08.

cence, is followed. The ultimate flower-bearing element

is a highly condensed raceme (botryum), the co-flores-

cence. These racemes are aggregated into highly con-

densed double racemes (dibotrya) which are clustered

around the main axis of a compact synflorescence, form-

ing the cone-like female inflorescence. The positions in

which these female cones are borne, differ from each

other in the three species.

Cliffortia dichotoma Fellingham, sp. nov., C. ar-

borea proxime affinis, sed positione inflorescentiarum

feminarum strobiliformium in apicibus ramorum princi-

palium dichotomorum, non sequentialiter in ramis prin-

cipalibus, et foliis unifoliolatis parvis glabrescentibus,

non trifoliolatis multo longioribus et incano-pubescen-

tibus, differt.

TYPE. — 3119 (Lokenburg): Nieuwoudtville Dist.,

Oorlogskloof Nature Reserve, Farm Cipher Kop 804,

720 m, (-CA), 16-10-1996, Pretorins 396 (NBG, holo.;

BOL, K, MO, PRE).

Trees up to 5 m tall with main trunks butressed, up to

500 mm diam.; bark reddish brown, flaking; branching

pattern dichotomous, resulting in a spreading canopy.

Branches incano-floccose, glabrescent with age and

appearing segmented with leaf sheath residues from

obsolete primary leaves. Primary leaves unifoliolate,

stipulate, caducous except at swollen branch tips which

become female cone-like inflorescences; stipules lanceo-

late, up to 0.5 mm long, pungent, greyish brown, glabres-

cent; leaflets lanceolate, (10-)12(-15) mm long, pun-

gent, glabrescent, margins revolute. Secondary leaves

unifoliolate, exstipulate; vagina membranous, sheathing,

successive vaginas imbricate; leaflets ligulate, ( 1 .5-)

2.0-3.0(-3.3) X 0.9 mm, up to 0.5 mm thick; margins

revolute; upper surface glabrous greyish green, longitu-

dinally grooved, undersurface incano-floccose except for

revolute margins and distally raised main vein; apex

mucronate, mucro minute, translucent. Male inflores-

cence: a condensed raceme of 2 or 3, (rarely only 1)

flowers on a villous, highly condensed brachyblast in

axil of a secondary leaf subapically on a brachyblast on

lower part of a main branch bearing a female inflores-

cence apically. Male flowers: very shortly pedicellate.

42

Bothalia 33,1 (2003)

FIGURE 1. — Clifforlia dichoumui. A, initiation of female cone as apical swelling on main stem; B, young female cone with numerous strap-shaped

styles/stigmas, apical continuation of mtiin axis as vegetative main stem and two robust lateral branches basally; C, primary cone leaf from near

apex of cone; D, apex of cone with four primary cone leaves subtending involucrally arranged groups of secondary cone leaves; E, primary

cone leaf with much-extended sheath, from equator of cone, therefore, apparently upside down; F, two female Bowers with (3)4 calyx lobes and

stnip-shaped styles/stigmas, borne on minute pedicels; G, pedicel with two braeteoles from female Bower; H, one bracteole from female Bower;

I, pedicel; J, smaller bracteole from female Bower; K, male Bower with pedicel, braeteoles and stamens; L, vegetative branchlet with male Bow-

ers; M, vegetative leaf with amplexicaul sheath, adaxial view; N, vegetative leaf, abaxial view; O, secondary cone leaf bearing capitulum-like

botryum in axil; P, capitulum-like botryum, hairs removed to show pedicels and bracts; Q, mature female intlorescence cone with 'shroud' of

vegetative lateral branchlets and proliferating apical main axis; R, S, fruits, variously faceted and narrowly winged or ridged. A, C-E, M-S,

drawn from Felliiif’hani I6H4, B, F-L, from Felliiif’lwm 1689. Scale bars: I mm.

Bothalia 33,1 (2003)

43

creamy white; bract narrowly triangular, up to 0.5 mm

long, long ciliate, whitish; sepals 4, broadly elliptical to

broadly obovate, 1. 5-2.0 x 1.2-1. 8 mm, apex subacute,

thickened, dorsally villous; stamens 4; filaments fili-

form, up to 2 mm long, glabrous; anthers up to 0.75 x 0.6

mm. Female inflorescence', many condensed double

racemes (homothetic dibotrya) each comprising a few,

sessile, capitulum-like co-florescences (botrya), aggre-

gated spirally on condensed main axis of an oblong,

cone-like polytelic synflorescence apically on a main

branch, one of a pair originating from below a cone of

previous season; flowering cones (30-)35^0 x (15-)

20-25 mm with apical proliferation of main axis

(30-)40-50(-60) mm long, secondary cone leaves green-

ish white incanous, interspersed with long maroon styles;

fruiting cones (40-)45— 55 x (18-)22-25 mm with api-

cal proliferation of main axis plus proliferation shoots of

lateral dibotrya, these thin vegetative branches up to 80

mm long and shrouding cone, caducous; leaves of pri-

mary cone axis ( primary cone leaves) unifoliolate, vagi-

na grossly expanded, woody, flushed with red, inter-

spersed with longitudinal cartilaginous veins and with

pungent woody stipules up to 2 mm long, ciliate, vaginas

sheathing main cone axis and segregating individual

dibotrya, leaflets narrowly elliptical, up to 2.5 x 1 mm

woody, pungent to minutely bifid with remnants of short

lanate indumentum basally; leaves of secondary cone

axis (secondary cone leaves) trifoliolate, arranged

involucre-like interspersing florescences (botrya), vagina

3.0-3.5 X 4—5 mm, dorsi-ventrally flattened, woody, longi-

tudinally furrowed, exstipulate, leaflets 3.0-3. 5 x 2. 0-2. 5

mm, greyish green, incano-floccose, conical-mucronate,

mucro 1.5-1.75 mm long, pungent, becoming woody,

leaflets of each leaf arranged in triangle, reducing in size

towards centre of involucre; co-florescence (botryum)

7-12-flowered, highly condensed with axis a flattened

platform of irregular shape bearing minute pedicels sub-

tended by reduced and modified bracts interspersed with

villous hairs. Female flowers: hidden except for styles

and tips of calyx lobes; bract subulate, up to 0.5 mm

long, long-ciliate, translucent; pedicel 0. 1-0.2 mm long;

sepals (3)4, linear-acute, 2. 0-2. 5 x 0. 3-0.5 mm, reflexed,

glabrous; receptacle cylindrical, 1.5-1. 7 x 0.75 mm.

faintly longitudinally furrowed, glabrous; style 1, linear,

plano-convex, 5. 5-6.0 x 0.2-0.25 mm, acute, spirally

twisted, exerted from cone, deep maroon, upper half

edged with short irregular teeth. Fruit: irregularly longi-

tudinally angled. 2. 3-2. 5 x 1.0-1. 2 mm, apically con-

cave around style remnant, dark brownish grey and

glabrous with two narrow whitish longitudinal wings;

retained within cone for several years. Figures 1, 2.

This new species differs from its closest ally, C.

arborea, mainly in the ultimate position of the female

inflorescence cones, which are borne terminally on the

dichotomous main branches, whereas those in C. arborea

occur sequentially on main stems. In both species the

female inflorescence cones originate terminally on main

branches, which subsequently proliferate beyond the

cones. In C. arborea, the apical proliferation shoots are

permanent structures, promoting longitudinal growth by

continuation of the main stems, which bear sequential

cones. In C. dicliotoma, how'ever, the proliferation shoots

soon become obsolete, leaving the cones in apical posi-

tions on dichotomous branches; longitudinal growth being

achieved by means of the development of two lateral

branches, from just below each cone, into dichotomous

main stems. It is in their ultimate condition and position on

the stem, that there is a marked difference between the

cones in C arborea and C. dicliotoma, and where the sim-

ilarity between the inflorescences in C. conifera and C.

dicliotoma is more pronounced, being terminal in both

species. The markedly dichotomous branching pattern that

arises as a result of the final condition and position of the

cones, is another character that clearly sets this new

species apart from its two allies. Furthermore, the (sec-

ondary) vegetative leaves of C. dicliotoma, differ from

those in C. arborea in being unifoliolate, small, and

glabrescent, whereas those of C. arborea are trifoliolate,

much longer and incano-floccose.

As in the case of C. arborea, herbarium specimens of

the new species are not always representative of the actu-

al condition in the wild. A far larger specimen than what

is practicable will have to be collected to reflect the true

positioning of the inflorescences and the vegetative

branching pattern. In most cases herbarium specimens

contain a single branch with a single terminal inflores-

cence only. This would in fact, be less than half of the

specimen needed to give a true history over two seasons

of the phenology and branching pattern in this species. In

contrast to the series of two cones on a single main stem,

which would represent the development over two sea-

sons in C arborea, the normal pattern in C. dicliotoma

would be three inflorescences, with the older one (of the

previous season) in the angle between two dichotomous

branches, each of which bears a terminal inflorescence of

the present season (Figure 2).

During his physiological research on C. dicliotoma.

Prof, von Willert raised seedlings which he reported as

being ‘tri-foliolate’. Photocopies made of these seedlings

seem to substantiate this finding. Close examination of

the actual material, however, led to the discovery of two

types of vegetative leaves, primary and secondary. Prof,

von Willert’s observations were of very young, trifolio-

late and exstipulate, or unifoliolate and stipulate, prima-

ry leaves (see below), before the development of vegeta-

tive short shoots with unifoliolate and exstipulate sec-

ondary leaves and the subsequent loss of the caducous

primary leaves.

Primary vegetative leaves occur on long shoots and

secondary vegetative leaves on (vegetative) short shoots.

Secondary thickening of the long shoots, causes the early

loss of the vegetative primary leaves, as their sheaths do

not enlarge to accommodate the thickening. They are thus

best observed on the apical swellings on main branches,

where they are permanent structures, destined to become

primary cone leaves on the primary short shoot (main

axis) of the female inflorescence cone, changing in form

and dimension as the cone develops. Both the primary

vegetative and primary cone leaves, can be interpreted as

being trifoliolate and exstipulate, or unifoliolate with

leaf-like stipules (Figure 1 A, C, E). Anatomical examina-

tion will have to be done to settle this question.

Secondary vegetative leaves, borne on vegetative short

shoots, are exstipulate, and of quite a different shape,

size and texture to the primary leaves. On older stems

44

Bothalia 33,1 (2003)

C ^ D

only secondary leaves are visible, because of the

caducous nature of the primary vegetative leaves. The

secondary vegetative leaves are spirally arranged on

short shoots originating immediately above the residual

vaginas of the obsolete primary leaves and their vaginas

are imbricate (Figure IL-N).

Oliver & Fellingham (1994) noted ‘the development

of lateral branches from the cones’ in C. arhorea, but

erroneously interpreted these as ‘sterile, lateral, sec-

ondary branchlets within the cone’, that is to say, origi-

nating as vegetative branchlets and directly on the main

cone axis, independent of the secondary flower-bearing

axes of the cone. During the present study, however, it

became clear that these so-called ‘sterile, lateral, sec-

ondary branchlets’ of the female cone in C. arhorea are

FIGURE 2. — Longitudinal diagrams

of architecture and structure

of synflorescences. A, Clif-

fortia dichotomcv. female

synflorescence cone of previ-

ous growth season, terminal-

ly on a main branch, with

(younger) synflorescences on

two lateral branches. B, C.

arborea: three female synflo-

rescence cones on main

branch and two younger on

lateral branches originating

from older cones. C, part of

syntlorescence with three

dibotrya as in young synflo-

rescences in both C. dicho-

toma and C. arborea. D. E.

single co-florescence or

botryum (solid square) with

its individual flowers (open

circles); (similar in both spe-

cies). MA. main axis; SN,

syntlorescence; DB, dibot-

lyum; VB. vegetative branch;

CoF, co-florescence (botryum);

pci, primary cone leaf; scl,

secondary cone leaf; br, bract

of single flower; broken lines

in synflorescences A & B re-

present dibotrya; dotted zig-

zag lines in C-E represent

expanded highly condensed

axes.

in fact, the proliferation shoots of the condensed sec-

ondary flower-bearing axes (dibotrya) of the cone. This

means that they originate on the main axes as the basal

parts of the lateral (fertile) cone axes and not as separate

(sterile) vegetative shoots. It is only at the periphery of

the cone that the short shoots change into vegetative long

shoots — the proliferation shoots. As recorded in 1994,

these lateral branchlets on the cones are permanent struc-

tures and can develop into main branches bearing female

inflorescence cones. These in turn are capable of contin-

uing longitudinal growth in the same way as the main

axis of the cone from which they originated. The

schematic interpretation for the mature (lateral) cone in

C. dichotoina, before abortion of the lateral proliferation

shoots, also pertains to the structure in C. arborea

(Figure 2A-C).

Bothalia 33.1 (2003)

45

In C. dichotoma, anthesis is marked by the appearance

of concentric circles of long, maroon styles, and by the

proliferation of the main axis of the cone into an apical

vegetative stem, a few centimetres long. After anthesis

this proliferation shoot lengthens somewhat. At this stage

the cone also becomes shrouded in an outcrop of sec-

ondary branchlets up to about 80 mm long, originating as

the proliferation shoots of the secondary inflorescence

branches (dibotrya). Up to this point in the development

of the inflorescence, it is similar to that in C arborea. In

C. dichotoma, however, this shroud of side branches is

eventually shed, as is the main proliferation shoot, leav-

ing the cone bare and terminal on a main branch. At this

latter stage, it more closely resembles the condition in C.

conifera, bearing its cone terminally (apically) on a

branch and is devoid of any lateral proliferation shoots.

In both of the last-mentioned species the growth points

of the main as well as the lateral cone axes have been

lost — in the case of C. dichotoma through abortion and

in C. conifera through loss of viability of the apical buds

of both the main axis and lateral axes of the cone. In rare

instances in C. dichotoma the main axis proliferation

shoot is retained beyond the time of fruiting (Fellingham

1706) (Figures IB, S; 2A).

With the loss of the growth point of the main branch

in C. dichotoma, two lateral branches from just below the

inflorescence cone and close to each other develop into

main branches which curve around the cone. These

branches supply two growth points to compensate for the

one lost and thus longitudinal growth is continued while

giving rise to the dichotomous vegetative branching pat-

tern that distinguishes this species. The female cones of

the next season develop apically on these two branches

(Figure 2A).

Rare occasions where three cones were produced in a

cluster, occurred during a very dry season following an

exceptionally wet one. These clusters of three cones were

in fact, the terminal cone, plus two cones, terminally on

extremely short lateral branches, from just below the

central cone. The development of dichotomous branches

and their apical cones, which normally stretches over

two seasons, was contracted into the same season, with

the resultant foreshortening of the branches, and the

simultaneous maturing of all three cones. This condition

can clearly be seen in the collections of Fellingham

1705, 1707 and 1711, all collected in October 1997.

Another unusual feature noticed during the same dry sea-

son following a very wet one, is the development of mul-

tiple lateral branches from below an aborted cone

{Fellingham 1708),

The onset of anthesis, as indicated by the swelling of

the apices of the dichotomous branches with their cover-

ing of enlarging imbricately arranged primary (vegeta-

tive) leaves, coincides with the maturing of the fruits in

the (central) cone of the previous season. Initially these

primary leaves do not differ from the primary stem

leaves, found on all main stems, which become obsolete

with secondary growth of the main stems, but it soon

becomes evident that these primary leaves remain imbri-

cate and the apical swelling enlarges. In contrast to the

sheaths of the primary vegetative leaves, the sheaths of

the primary cone leaves expand as the cone develops and

at the same time, remain in close proximity to each other

basally, as the main axis of the cone remains contracted

— the axis of a fertile short shoot. Furthermore, each pri-

mary leaf surrounds a developing synflorescence and

thus becomes a true primary cone leaf. The curvature of

the developing cone causes the imbricately attached

sheaths to fan out, so that those below the equator of the

cone seem to grow up-side down (Figure lA, C-E).

In contrast to the condition in C. dichotoma with the

loss of the growth points of both primary and secondary

proliferation shoots, these growth points are both

retained in C. arborea. Here the primary proliferation

shoot continues the vegetative growth of the main

branch, to give rise to a series of female cones, one per

season. Any two cones are thus separated by a length of

vegetative main branch, proliferating from the main axis

of the older cone. Similarly, proliferation of the lateral

cone axes (main axes of the dibotrya), give rise to per-

manent vegetative lateral branchlets, which are capable

of developing into stout lateral branches bearing female

inflorescence cones, which in turn can continue longitu-

dinal growth of the plant, through proliferation of both

axes of the cones. In this way a series of cones can devel-

op out of what was initially the lateral cone branchlets

(Figure 2B).

Oliver & Fellingham (1994) considered the possibility

that cone-like inflorescences might still be discovered in

other species. Known species with recorded compound

inflorescences, viz. C. heterophylla Weim., C. odorata L.f.

and C. hirsuta Eckl. & Zeyh. were noted, though none of

these was found to be cone-bearing. Of these three

species, the first two have subsequently been studied in

detail (Fellingham 1999; Fellingham & Linder in press).

The initial apical arrangement of imbricate modified

leaves, obscuring the reduced female flowers in C. hetero-

phylla, relates to the primary leaves covering the swollen

tip of a main branch, marking the onset of cone develop-

ment in C. dichotoma. In this early stage of development

of the inflorescence in C, heterophylla, the condensed

character of the apex of the main stem is evident in the

imbricate anangement of the modified leaves on it. As the

development of the inflorescence progresses through the

various stages from the early female, through the bisexu-

al to the male stage, the secondary condensed inflores-

cence axes become evident, as do the proliferation shoots

on these secondary axes, again mimicking C, dichotoma.

There is, however, also an unmistakable difference

between the inflorescence structures in these two species,

mainly in the demonstration of a peculiar plasticity mani-

festing itself in the total disappearance by integration and

not loss of the inflorescence in C. heterophylla, whereas

the cone-like inflorescence in C. dichotoma is retained for

a number of years, but eventually discarded (Fellingham

1999; Fellingham & Linder in press).

The relatively large flowers in C. odorata obscure the

complex nature of the inflorescence, to such an extent

that Weimarck (1934) described the flowers as being

‘fascicled in leaf axils’ and in another context, in racemes

that ‘approach the form of heads’(Weimarck 1948).

These ‘heads’ were found to consist of a number of head-

like subclusters of flowers, each with a condensed, short-

46

Bothalia 33,1 (2003)

Stalked axis, and implanted on a highly condensed disc-

like axis, in the axil of a vegetative leaf. The disc-like

structure in this species, though totally flattened, and far

more pronouncedly modified, is analogous to the highly

condensed main axis of the cone in the three cone-bear-

ing species, even to the extent of retaining a viable

growth point in the centre of the disc. The stalked sub-

clusters are analogous to the secondary axes of the cone-

like inflorescences. The loss of the viability of the

growth point of these secondary axes, renders it closer to

C. conifera than to the other two species in the section.

The arrangement on the secondary axes of the flowers in

smaller, closely packed groups with modified or even

absent bracts, is analogous to that of the ultimate flower-

bearing platforms in the cones (Fellingham 1999; Fel-

lingham & Linder in press).

As a result of the discovery of the error in interpreta-

tion of the lateral branchlets on the female cone in C.

arborea as discussed above, it is necessary to emend the

description of the species thus: the phrase 'with second

order axes continuing growth laterally’ should read 'with

second order cone axes continuing growth laterally'.

Section Arboreae H.Weim. emend. Fellingham

Shrubs and trees up to 5 m tall. Leaves uni- or trifolio-

late; stipules 0-2, subulate to foliaceous; leaflets revolute

or flat, entire to dentately 2-4-lobed. Male inflorescence:

a condensed raceme of 2-5 flowers on a lateral, highly

condensed, short shoot in axil of vegetative leaf. Male

flowers: sepals 3 or 4; stamens 4-10. Female inflores-

cence: a conoid synflorescence subterminally on main

branches with continued apical growth (C. arborea), api-

cally on main branches after loss of initial apical prolif-

eration fC. dichotoma) or apically on determinate lateral

short shoots fC. conifera), the synflorescences composed

of double racemes (dibotrya) each consisting of con-

densed co-florescences (botrya). Female flowers: sepals

3 or 4; receptacle triangular, narrowed to base, naiTowly

3-winged or 3-ridged (C. arborea, C. conifera) or irreg-

ularly longitudinally furrowed or angled, naiTowly 2-

winged (C. dichotoma): style/stigma single, linear, edges

irregular and shortly dentate, creamy white (C. arborea,

C. conifera) or maroon (C. dichotoma): achene single.

Species included: Cliffortia arborea Marloth (type

species), C. conifera E.G.H.Oliv. & Fellingham, C. dicho-

toma Fellingham.

PHYTOGEOGRAPHY

Cliffortia dichotoma is known only from a small area

on the Oorlogskloof Escarpment, around the waterfall on

the Farm Papkuilsfontein, about 25 km south of Nieu-

woudtville. Northern Cape (Figure 3). It grows at 700 m

which is much lower than either C. arborea at

I 500-1 800 m or C. conifera at I 300 m. It grows on

west-facing, rocky ledges near the edge of the escarp-

ment and about 500 m onto the plateau. This locality is

jiKst south of the Oorlogkloof Nature Reserve. The annu-

al rainfall in this region is typically very low, occurring

in late winter to early spring, if at all.

FIGURE 3. — Known distribution of Cliffortia dichotoma, •; C. ar-

borea O; C. conifera, H.

PHENOLOGY

Like C. arborea and C. conifera, C. dichotoma is

monoecious with unisexual male and female flowers

borne on separate branches. The female flowers are

borne in terminal or subterminal cones near or on the

ends of the main branches. The male flowers are borne in

small clusters in the axils of vegetative leaves on vegeta-

tive short shoots on branches. As in the related two

species, in C. dichotoma these branches are situated

below the female cones and mostly much lower down on

the plant {Fellingham 1685, 1689, 1712). It is also clear-

ly wind pollinated like the two known species in the sec-

tion and presumed for all species in the genus (Koutnik

1987). While sharing all the adaptive characters

(Koutnik 1987; Proctor et al. 1996) with other wind-pol-

linated plants, these coniferous species of Cliffortia have

a rare reversal of location on the plant, of the male and

female elements, with the male below the female. This

rare condition is shared with only one gymnosperm viz.

Finns sylvestris (Proctor et al. 1996). The question

remains as to whether this condition could be an adapta-

tion to pollination by updrafts prevailing at the edges of

the escarpments that are the preferred habitats of these

species. Apart from the obvious parallelism between

these three coniferous species and the evergreen seroti-

nous gymnosperm, they also share the phenomena of

anemophily and monoecy.

Except for the tips of the sepals and the long, strap-

shaped styles/stigmas, the female flowers are totally hid-

den within the involucres of secondary cone leaves. The

styles/stigmas in C. dichotoma are more showy than in

either C. arborea or C. conifera by virtue of their maroon

colour in contrast to the creamy white organs in the lat-

ter species. The fruits mature over a period of about a

year, during which time the cone undergoes a number of

changes. Apart from becoming more woody, it also

develops an apical proliferation shoot and lateral prolif-

eration shoots on the lateral cone axes. Both these types

of proliferation shoots are, however, shed quite soon.

This leaves the cone temporarily in an apical position on

the branch. It is, however, soon over-topped by the two

Bothalia 33,1 (2003)

47

or more secondary branches originating from just below

the cone and destined to continue longitudinal growth of

the plant. The dichotomous branching pattern that results

is clearly marked by the retention of the cones for sever-

al seasons.

The flowering period in C. clichotoma is earlier and

shorter than in C. arborea or C. conifera, viz. between

late October and late November (Fellingham 1702,

1709- 1712). Male flowers were encountered in late

August 1995 (Fellingham 1684, 1685 & 1689) which is

rather surprising as female flowers were only found more

than a month later.

Paraty'pe material

C. clichotoma

NORTHERN CAPE. — 3119 (Lokenburg): Papkuilsfontein. rocky

ledges of Oorlogskloof Escarpment. SW aspect, 700 m, (-CA), 27-08- 1 99.S.

Fellingham 1684 (BOL, K, NBG. PRE); 28-11-96, Fellinghain 1702

(BOL male & female, NBG male & female, K male, PRE); Kranskloof,

rocky ledges, SW aspect, 700 m, (-CA), 24-10-1997, Fellingham 1705

(BOL, NBG, PRE); Fellingham 1706, (sheet 1 & 2 BOL); Fellinghain

1707 (BOL); near waterfall, 700 m, (-CA), 24-10-1997, Fellingham 1708.

1710- 1712 (BOL); 1709 (BOL, NBG. PRE); Oorlogskloof Nature

Reserve, Farm Cipher Kop 804, plateau at bottom of koppie on right of

Cipher Kop homestead, 720 m, (-CA), 16-10-1996, Pretorius 396

(BOL, K, MO, NBG, PRE); 'Papkuilsfontein’ on road to waterfall, on

rocky, sandy, flat plateau about 1 km from Oorlogskloof, and 2 km from

waterfall. (-CA), 20-08-1993. Van W\-k 626 (NBG); Niewoudtville area.

Papkuilsfontein, escarpment, from waterfall in De Hoop se Rivier,

northwards only to border with Kranskloof Farm, Von Willert ,r./r

(NBG).

Additional material examined

C. arborea

NORTHERN CAPE. — 3119 (Calvinia); Akkerendam, dolerite

screes and krantzes of Hantamsberg. 1 372 m, (-BD), 14-11-1955,

Acocks 18621 (BOL male & female, K male & female, NBG male);

summit of Hantam Peak. 1 660 m, (-BC). Wisura 3556 (NBG sterile);

Hantamsberg, Voetpadskloof, S-facing slope, renosterveld, 1 465 m,

(-BD), 3-09-1986. Oliver 8878 (NBG sterile); upper E slope of

Hantam Mtn. Vanrhynshoek Farm, kloof near stream, (-BD). 7-10-

1986. Thomas <& Van Jaarsveld 8961 (NBG sterile). 3220 (Sutherland):

Roggeveld. Farm Uitkyk, Sneeuwkrans, below krantz facing W. I 370

m, (-AD), 10-1920. Marloth 9730 (PRE stem & bark only, NBG male

& female); Hottentotsbank near Sneeukrans, W-facing .scree slopes,

1 433 m. (-AD), 22-9-1981, Ronrke 1728 (K .sterile, NBG sterile);

Roggeveld, southwestern krantzes on the Komsberg. I 550 m. (-DB),

04-1905. Marloth 3907 (BOL female, K male. NBG sterile);

Roggeveld Mtns. Komsberg. 1 525 m, (-DB). 10-1920, Marloth 9770

(PRE female. NBG female); just W of road at top of Komsberg Pass,

sandstone escarpment, (-DB). 22-09-1977, Moffett 1463 (NBG sterile);

plateau at top of Komsberg Pass, E of road, 1 600 m, (-DB), 02-1986,

Moffett & Steensina 4067 (NBG male); Komsberg Pass, lower slopes

ENE of Skurwekop. 1 500 m. (-DB), 4-06-1992, Oliver 10054 (BOL,

K. MO. NBG, PRE. S. all female). Without precise locality: mountains

near Sutherland. 12-1905, Du Toit sub BOL10057 (BM. BOL, K, all

sterile); Sutherland, shady side of very inaccessible kloofs, 13-10-1969,

De Villiers s.n. (NBG immature female). 3221 (Merweville): Sterk-

boomkloof near Vinkfontein, edge of kloof, 1 500 m, (-CA), 21-02-1986,

Moffett & Steensina 4060 (NBG sterile).

WESTERN CAPE. — 3222 (Beaufort West): Nuweveldberge, Karoo

National Park, mountain view area near EM tower, SW slope, 1 830 m,

(-BA), 17-11-1992, Fellingham 1624 (MO, NBG, PRE, all female);

Karoo National Park, mountain view, top of mountain, 1 830 m, (-BA),

3-01-1985, Shearing 893 (PRE female); Nuweveld Mtns, S slopes

above Beaufort West, 1 525 m, (-BC), 07-1940, Esterhuysen 2759a

(BOL sterile); Nuweveldberge, Karoo National Park, mountain view

area near the Look Out. 1 830 m. (-BC). 17-11-1992. Fellingham 1625

(BM, NBG. PRE, all male & female).

C. conifera

WESTERN CAPE. — 3320 (Montagu): Ladismith, Anysberg, E

end of Prinskloof, 1 300 m, (-BC). 23-09-1990, Oliver 9730 (NBG,

PRE. S, all female); ibid. 10-08-1991, Fellingham 1531 (NBG female);

ibid. 4-06-1992, Oliver 10055 (BOL, MO, NBG. PRE, all male and

female); Anysberg, E end, gully leading to Prin.sberg, (-DA), 6-10-1982,

M. van Wyk 1072 (NBG female).

ACKNOWLEDGEMENTS

I appreciate the help received from Dr E.G.H. Oliver

in preparing the Latin diagnosis and for editing. My

thanks go to Mrs I.M. Oliver for useful tips on preparing

the illustrations and for drawing the schematic diagram.

I want to thank Mr W.A. Pretorius and Mr J. Afrika, con-

servation officers of the Oorlogskloof area, for guiding

me to the outlying units of the population. Without the

friendly reception and accommodation received from the

owners of the Farm Papkuilsfontein, Mr and Mrs W.N.D.

van Wyk, this project could not have been undertaken.

REFERENCES

FELLINGHAM, A.C. 1999. Phenology and branching in eight select-

ed species of the genus Cliffortia L. M.Sc. thesis. University of

Cape Town. Cape Town. Unpublished.

FELLINGHAM, A.C. & LINDER, H P In press. Inflorescences of

Cliffortia L. (Rosaceae). Bothalia.

KOUTNIK. D. 1987. Wind pollination in the Cape flora. In A.G. Rebelo,

A preliminary synthesis of pollination biology in the Cape flora.

South African National Scientific Programmes Report No. 141:

126-133.

MARLOTH, R.H. 1905. Eine neue interessante Cliffortia vom Rogge-

veld. Botanische Jahrbiicher 29: 318, 319.

OLIVER, E.G.H. & FELLINGHAM, A.C. 1994. Anew serotinous spe-

cies of Cliffortia (Rosaceae) from the southwestern Cape with

notes on Cliffortia arborea. Bothalia 24: 153-162.

PROCTOR, M., YEO, P. & LACK. A. 1996. The natural histoiy of polli-

nation. Harper Collins, London.

WEBERLING, F. 1981. Morphologie der Bluten und der Blutenstande.

Eugen Ulmer, Stuttgart.

WEBERLING. F. 1983. Fundamental features of modern inflorescence

morphology. Bothalia 14: 917-922.

WEIMARCK, A.H. 1934. Monograph of the genus Cliffortia. Gleer-

upska, Lund.

WEIMARCK, A.H. 1948. The genus Cliffortia, a taxonomical survey,

Botaniska Notiser 1948: 167-203.

Bothalia 33,1:49-51 (2003)

A revision of Ledebouria (Hyacinthaceae) in South Africa. 3. The

reinstatement of L. ensifolia^ L. galpinii and L, sandersonii

TJ. EDWARDS* and S. VENTER*

Keywords: Hyacinthaceae, Ledebouria Roth, South Africa, taxonomy

ABSTRACT

Three taxa of Ledebouria Roth are raised from synonymy to species status, with notes, diagnostic characters and dis-

tributions. L. galpinii {Baker} S. Venter & T.J. Edwards is a narrow endemic in the Wolkberg Centre and was formerly placed

within L. cooperi (Hook.) Jessop. The other species, L. sandersonii (Baker) S. Venter & T.J. Edwards and L. ensifolia (Eckl.)

S. Venter & T.J. Edwards, are closely allied to each other, sharing a number of synapomorphies.

INTRODUCTION

In his revision of Scilla L. in South Africa, it was neces-

sary for Jessop (1970) to place a large contingent of

species into Ledebouria Roth. This subdivision of Scilla

saw the sinking of a number of valid species and result-

ed in some confusion in the taxonomy of Ledebouria. In

his paper Jessop (1970) alerted readers that the ‘lump-

ing’ involved may have been too drastic. His inclusion

of a wide range of morphologically divergent forms

within L. cooperi (Hook.) Jessop was necessary; however,

S. galpinii Baker and S. sandersonii Baker need to be re-

surrected.

In our opinion L. undulata (Jacq.) Jessop also em-

braces more than a single species and is subdivided into

L. ensifolia (Eckl.) S. Venter & T.J. Edwards and L. undu-

lata sensu stricto. This subdivision is necessitated by the

differences in gynoecium and bract morphology.

1. Ledebouria ensifolia (Eckl.) S.Venter & T.J. Ed-

wards, comb. nov.

Drimia ensifolia Eckl. in South African Quarterly Journal 1 : 364

(1830). Scilla ensifolia (Eckl.) Britten: 201 (1908). Type: Uitenhage.

Zwartkops River. Zeyher 10 (K!. lecto. selected here; -PRE. photo.!).

D. ludwigii Miq.: 39 (1839). Idothea ludwigii (Miq.) Kunth: 681

( 1843). Scilla ludwigii (Miq.) Baker: 9 ( 1870). Type: Caput bonae Spei,

on the sand hills near the Zwartkops River. Ecklon & Zeyher 1064 (U.

holo.; GRA!; PREI).

D. apertiflora Baker: t. 19 (1868) synon. nov. Ledebouria aperti-

flora (Baker) Jessop: 254 (1970) synon. nov. Type: Saunders Refugium

Botanicum 1 : t. 19(1 868 ).

Scilla prasina Baker: 10 (1870). Type: Kaffirland. Gill s.n. (Kl, holo.;

-PRE. photo.!).

S. pusilla Baker: 183 (1876). Type: Transkei. Bazeia. Baur293 (K!,

holo.; -BOL. drawing!; -PRE, photo.!).

S. ecklonii Baker: 7 (1892). Type: Tambukiland, mountains between

Silo and Windvogelberg. Ecklon & Zeyher 12 (B!, holo.).

* School of Botany and Zoology, University of Natal, Private Bag XOl,

3209 Scottsville, Pietermaritzburg.

MS. received: 2002-03-15.

Plants solitary. Bulb hypogeal, cylindrical, 40-60 x

20-30 mm; dead bulb scales hard, dark brown to purple,

apices attenuate; live bulb scales membranous. Leaves

5-10, fully developed at anthesis, spreading, narrowly

ovate to ensiform, 80-150 x 15^0 mm, fleshy, dull

glaucous, usually immaculate; base canaliculate; apex

acute; margin smooth. Inflorescences 1-3, dense, 8-320

mm long, 30-100-flowered; scape flaccid, terete, 50-200

mm long; bracts ±1x1 mm, deltoid, fleshy, green, with-

out bracteoles; pedicels patent, 3-4 mm long, green.

Perianth stellate; tepals green to pink with a dull green

keel, shai-ply reflexed, equal, oblong, 3. 0-3. 5 x 1.5 mm;

apex acute. Stamens erect, ± 3 mm long; filaments pink,

epitepalous; anthers ± 1 mm long. Ovary ellipsoid, 6-

lobed, ± 1.0 x 2.5 mm, carpels oblong with papillate

basal lobes; style triangular in section, ± 1.5 mm long,

glabrous, puiple; stipe up to 0.5 x 0.5 mm. Capsule 3-

lobed, symmetrical, globose; base truncate. Seed ellip-

soid, 3-5 mm long, surface wrinkled, brown.

L. ensifolia differs from other species of Ledebouria

in having fusiform roots. The species is widespread,

occurring on sandy soils derived from quartzites, gneiss

or rhyolite in woodlands or shrublands (Eigure 1). L.

cooperi and L. revoluta (L.f.) Jessop are commonly

misidentified as L. ensifolia but both have bracteoles and

50

Bothalia 33,1 (2003)

smooth carpel lobes; the latter also has elongate styles (±

6 mm). In addition, the torn leaves of L. revoluta reveal

‘threads’ (from unravelling xylem), no such ‘threads’

occur in the leaves of L. efisifolia.

Jessop ( 1970) placed L. ensifolia in synonymy with L.

imdulata which, in our opinion, is not a close ally. The

latter species has ovoid bulbs, lacks fusiform roots, lacks

papillate basal lobes on the ovary and has bracteoles. In

addition the undulate leaf margins, from which the

species was named, are consistently absent from L. ensi-

folia. Instead L. ensifolia appears to be closely allied to

L. sandersonii which also lacks bracteoles, lacks ‘threads’

in the torn leaves, has solitary bulbs and prominent papil-

late lobes on the carpel bases.

Vouchers: Acocks 16235 (PRE); Barker 6900 (NBG); Harrison 231

(NH); Moss & Rogers 540 (J); Rogers 28366 (GRA); Schlechter 18

(BOD.

2. Ledebouria galpinii (Baker) S. Venter & TJ. Ed-

wards, comb. nov.

Scilla galpinii Balcer in Flora capensis. 6: 487 (1896). Type: Eastern

Transvaal [Mpumalanga], summit of Duiwel's Kantoor, Galpin 672

(PRE!, lecto., designated here; BOLI, GRAI, NH!, SAM!, Z).

Plants usually gregarious. Bulb ovoid, hypogeal,

20-30 X 20-25 mm, neck 1-2 x 3-5 mm, usually with

bulblets on basal stem; dead bulb scales brown, apices

truncate; live bulb scales fleshy, aiTanged loosely; cata-

phylls 1-3, exserted above ground. Leaves fully devel-

oped at anthesis, 3 or 4, humifuse, oblong to ovate-

spathulate, 50-80 x 20-25 mm, fleshy, without threads

when torn, glossy purple to green-puiple, adaxial surface

lacunose, venation obscure; margins smooth; base

canaliculate; apex mucronate. Inflorescences 1-3, sub-

globose, 10-20 X 10-20 mm, flaccid, 20-30-flowered, as

long as or longer than leaves, extending in fruit; scape

usually twisted basally, purple, 20-50 mm long; bracts

fleshy, linear-lanceolate, up to 1 x 0.5 mm, pink to pur-

ple; bracteoles present; pedicels patent, 3^ mm long,

purple. Perianth stellate; tepals subequal, oblong, 4-5 x

2 mm, pink to purple, keel green; apex obtuse, cucullate.

Stamens spreading, 3^ mm long; filaments maroon;

anthers up to 0.5 mm long, violet. Ovary globose, 3-

lobed, ± 1.5 X 1 mm, lobes obtusely ovate; style up to 3

mm long, triangular in section, purple; stipe ± 0.5 x 0.5

mm. Capsule globose, base truncate. Seed elliptic,

1 .5-2.0 mm long, surfaces strongly wrinkled, red-brown.

L. galpinii occurs on Black Reef Quartzites of the

Wolkberg Group (SACS 1980). The species may be

added to the list of 130 endemics within the Wolkberg

Centre of Endemism (Van Wyk & Smith 2001) ( Figure

1 ). Plants occur in mistbelt sourveld often in association

with Streptocarpus galpinii Hook.

Jessop’s (1970) reduction of L. galpinii into syn-

onymy with L. cooperi is unwarranted. L. cooperi is a

plant of seepages and vleis, it is .soboliferous and quick-

ly forms dense populations. By contrast, L. galpinii is a

plant of lithosols and, while occasionally producing

basal bulbs, is never soboliferous. Bulb morphology dif-

fers significantly, L. galpinii has ovate bulbs with trun-

cate apices compared to the acuminate apices of L.

cooperi. L. galpinii frequently produces 3 cataphylls but

in L. cooperi cataphylls are always solitary. The leaves of

L. galpinii are fleshy, flat, broadly ovate and humifuse,

whereas those of L. cooperi are mesic, canaliculate,

lanceolate and erect. The most obvious distinguishing

character of L. galpinii is the corrugated adaxial leaf sur-

face, embossed with regular pits. Plants of L. cooperi are

always more floriferous, producing between 20 and 60

flowers per raceme, the inflorescences of L. galpinii have

less than 30 flowers. Lastly the pedicels of L. galpinii are

usually 3^ mm long, whereas those of L. cooperi are

6-12 mm. The abundance of distinguishing characteris-

tics, and the occunence of L. galpinii in an area re-

nowned for plant endemism, leaves us in no doubt of its

specific status.

Voucliei-s: Codd 9789 (PRE); Galpin 672 (BOL. GRA. NBG. NH,

PRE); Van der Merwe 2047 (PRE); Venter 13389 (UNIN).

3. Ledebouria sandersonii (Baker) S.Venter & T.J. Ed-

wards, comb. nov.

Scilla sandersonii Baker in Saunders Refugium Botanicum 3 (Ap-

pend.): 5 (1870). Type: Transvaal, without precise locality. Sanderson

s.n. (K!, holo.; -PRE. photo.!).

S. baiirii Baker: 484 (1896). Type: Tembuland. Bazeia Mountain,

Baar 550 (K!. holo.; -PRE, photo.!; SAM!).

S. rysonii Baker: 484 (1896). Type: Griqualand East, Tyson s.n. (K!,

holo.; -BOL!, drawing; -GRA!, drawing).

S. oostachys Baker; 487 (1896). Type: Upper Umkomaas, Wood

4627 (K!, holo,; -BOL!, drawing; -NH!, -PRE, photo.!).

S. dipinlla Baker: 489 (1896). Type: Barberton. Saddleback Range,

Galpin 1182 (K!. holo.; BOL!, GRA!, NH!. PRE!, SAM!).

S. hella Markbtter: 13 (1930), Type: Oliviers Hoek Pass, Thode

STE3372 (STE!, holo.; -PRE, photo.!).

Plants solitary. Bulb hypogeal, ovoid to subglobose,

10-30 X 10-15 mm; dead bulb scales membranous,

brown, apices attenuate; live bulb scales membranous.

Leaves 2-6, fully developed at anthesis, spreading or

appressed, ovate to broadly lanceolate, 15-75 x 8-30

mm, fleshy, dull glaucous; adaxial surface sometimes

blotched with purple; abaxial surface suffused with pur-

ple; base canaliculate; apex acute; margin smooth.

Inflorescences 1 or 2, lax, 12-22-flowered, 20-25 mm

long; scape erect to flaccid, terete, 20^0 mm long;

bracts deltoid, ± 1.0 x 0.5 mm, fleshy, pink to purple,

without bracteoles; pedicels patent, 6-8 mm long, pink.

Perianth stellate; tepals pink, sometimes with a dull

green keel, recurved, equal, oblong, ±3x1 mm, apex

acute. Stamens erect, ± 3 mm long; filaments purple

above, white below, epitepalous; anthers ± 1 mm long.

Ovary ovoid, 6-lobed, ±1x2 mm; carpels oblong with

papillate basal lobes; style ± 3 mm long, triangular in

section, glabrous, purple; stipe up to 0.5 x 0.5 mm.

Capsule 3-lobed, symmetrical, globose; base truncate.

Seed ellipsoid, ± 2.5 mm long, surface wrinkled, brown.

The inclusion of Scilla sandersonii within L. cooperi

obscures its true alliances. By his recognition of the

.sandersonii series it is clear that Jessop (1970) was

aware of divergence within his broad concept of L.

cooperi. Examination of the gynoecium of L. .sander-

BothaIia 33.1 (2003)

51

FIGURE 2. — Distribution of Ledeboitria sandersonii.

sonii reveals a synapomorphy shared with L. ensifolia. In

these species the carpel bases are strongly lobed and are

adorned with papillae. Although closely allied to L. ensi-

folia. L. sandersonii is easily distinguished by its small-

er bulbs (10-30 X 10-15 mm versus 40-60 x 20-30 mm)

and the absence of fusiform roots. In addition, the leaves

of L. sandersonii are fewer (1-6 vs. 5-10), smaller and

broader (15-75 x 8-30 mm vs. 80-150 x 15^0 mm). L.

ensifolia is more floriferous producing 30-100 flowers

per raceme, whereas inflorescences of L. sandersonii sel-

dom produce more than 20 flowers. Flowering time:

October to November.

L. sandersonii is commonly associated with montane

grassland in the eastern parts of southern Africa where it

grows in shallow soils, overlying rock outcrops (Figure

2). These soils act as seepages in summer and frequently

freeze in winter. Both maculate and immaculate forms of

L. sandersonii occur, but mixed populations have never

been encountered.

Vouchers; Brown 323 (BOL); Compton 27803 (NBG, PRE); Kerfoot

8168 (PRE); Strey 9362 (NH. PRE); Van der Mern-e 2602 (NU).

ACKNOWLEDGEMENTS

Financial assistance from the Natal University Research

Fund and the FRD is gratefully acknowledged. The cited

herbaria are thanked for the loan of material examined

during this study. Ms C. Potgieter and the anonymous

referees are thanked for suggestions made on the manu-

script.

REFERENCES

BAKER. J.G. 1868. Saunders Refugium Botanicum 1: t. 18. 19.

BAKER. J.G. 1870. Monograph of Scilla: sections Ledeboitria and

Drimiopsis. Saunders Refugium Botanicum 3 (Append.): 1-18.

BAKER, J.G. 1876. New bulbous plants from the Cape. Journal of

Botany, London 5: 183.

BAKER. J.G. 1892. Liliaceae novae Africae australis herbarii regii

Berolensis. Botanische Jahrbiicher 15 Beiblatt 35: 7.

BAKER. J.G. 1896. Scilla in Flora capensis 6; 478— f 94.

BRITTEN. J. 1908. Overlooked Cape plants. Journal of Botany 46: 201.

ECKLON. C.F. 1830. Plants found in the District of Uitenhage. South

African Quarterly Journal 1 : 364.

JESSOP. J.P. 1970. Studies in the bulbous Liliaceae; 1. Scilla. Sclilzo-

carplius and Ledeboitria. Journal of South African Botany 36:

233-266.

KUNTH. C.S. 1843. Enwneratio plantarum 4: 338-681 . Cotta. Stuttgart

& Tubingen.

MARKOTTER. E.I. 1930. In Annale van die Universiteit van Stellen-

bosch. Reeks A, 8, afl. 1:13.

MIQUEL, F.A.W. 1839. Drimia in Melanges Botaniques. Bulletin des

Sciences Physique et Naturelles en Neerlande: 39.

SOUTH AFRICAN COMMITTEE FOR STRATIGRAPHY (SACS)

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

Geological Survey of South Africa.

VAN WYK. A.E. & SMITH. G.F. 2001. Regions offloristic endemism

in southern Africa. Umdaus Press, Pretoria.

* .* ^ '” . . • ■■- :>.v^-

r;ra ■•.■' J'Tl. ■' '

""^^ir* » ‘ft

yfc,f Jv*-f • ’. p •'III-.’ -tmi f)< ‘I* ^..'X^MJ'if A* i *:•• «U^%R.:«'vW,T0W4LLvi»‘*i^

. i/j'fcWiwt .* * ' I ■■ .l 'Ifet 4 ^ "

’>J'' -t'< ‘i’

: vU‘;’»>y f . , 'fe-tittl.,. ,..- . g,|

.,.,M .. ;;"V' ;. . fF- ^

^ ’ *’ -■r^.’r-^f’ ' ■ ' ■•• '.s' '''-lb'

•_s;\'fi. ‘ i'*-;; V:. , ('it. ‘ V|. ,. . jifS'^'. -I

Mjj -' I fif j;

rvCfi*^ ^ ■^^’'\.‘V‘j^^

w>; ^i^'ifr- •■ ■*%.... .^

_ ■ ■ '**V ;1£ iw-ff ' ^»•iflfe'

<^»v R5 i:i,^«i^-'.ftSij|^;*i!;i|V " ■■'•4* -I' ■(w»','nU;»

•j,^>?iAf r * '••'■ >' ' "* ,4->'iV-'

"'■1; ,: rti*.- ' '*W X «.in^'jpi^; 1 >*- ••.f. vV^„. 4'- '

li' ''™

k« TffW, i' •K>il‘rVT«'Vf'

■f'-' ■>

—M if j, ,'ift.f s^ i

'V •'

" ■'"■ 'L^ '* ■ f*.i »• Y'*‘'-'*f-V**''’ ^■4''<" ■' ' WVf

‘t-4*- .,.„,i=^ „,;■ IF •'! 'i'l'i,; , ■ ' '■^*- ■'( *f ivV -^4, ,,n

)\,,;,"i6i>.‘ , '‘. (U>;ni. ■ V ,'• ; ..‘i •.?/., :vJ#> . i.. ,

.;• • ..I'*,**, i * -' ll. ■•%■..■ .im ;4«4y ^’iinMl '3^' >.Jef ",

* ^'ii' * ■• 4/.- . '■ ■'

' .. Y.-' ■

I «, ":' *'

i- -. ' • • ■j.'.’ifii*'

©

Bothalia 33,1: 53-57 (2003)

An annotated checklist of the pteridophyte flora of Swaziland

J.P. ROUX*

Keywords: checklist, flora, Pteridophyta, southern Africa, Swaziland

ABSTRACT

An herbarium and literature-based review of Swaziland pteridophytes showed the poor species record and distribution

data available for these plants. Field work carried out over a period of 17 days resulted in a further 15 species recorded and

this brings the number of pteridophyte taxa known for the country to 112.

INTRODUCTION

An herbarium and literature-based review of

Swaziland pteridophytes showed the poor distribution

data available for these plants in the country. This con-

clusion was reached having worked through the check-

lists of Compton (1966) and Kemp (1981, 1983), and

the collections at the Bolus Herbarium (BOL), Natal

Herbarium (NH), National Herbarium of Swaziland

(SDNH), and the Compton Herbarium (NBG). A list of

collections held at the National Herbarium, Pretoria

(PRE) was also consulted. The checklist of Compton

(1966) is primarily based on his own collections assem-

bled during his stay in Swaziland, whereas those of

Kemp (1981, 1983) are based on herbarium records and

field work.

To improve the distribution data of the known species,

and record new taxa for the country, 17 days of intensive

collecting was done, from 6th to 22nd May 2002, trying

to cover as much territory as possible. Well-collected

areas were largely avoided due to the time constraint.

The drought, which was experienced during the time,

limited collecting in especially the lowveld and eastern

regions of the country. In spite of these constraints, 15

pteridophyte species are here recorded for the first time

for the Flora of Swaziland. This brings the number of

taxa known for the country to 112. This number will

undoubtedly increase significantly as the flora of the

country becomes better known. A more detailed account

of the pteridophyte flora of Swaziland is currently in

preparation.

RESULTS AND DISCUSSION

The taxonomic arrangement followed is that of Roux

(2001). The numbers following the name refers to the

respective checklists available for the country: 1,

Compton (1966); 2, Kemp (1981); and 3, Kemp (1983).

Names that do not comply with the names in current use,

are in italics and placed in square brackets. A single

voucher, where possible the first recorded for the coun-

try, is cited to substantiate its occurrence in Swaziland.

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

7735 Claremont, Cape Town.

MS. received: 2002-06-19.

PSILOTACEAE

Psilotum nudum (L.) P.Beauv., 1, 3

-2632 (Bela Vista): Siteki, Blue Jay Ranch, rocks near stream in

shade, ± 152 m, (-AA), 15 Dec. 1959, Compton 29588 (NBG.

SDNH).

EQUISETACEAE

Equisetum ramosissimum Desf. subsp. ramosissimum, 1, 3

-2631 (Mbabane): Bulunga Poort, sandy streambed, ± 304 m, (-DA),

18 Dec. 1963, Compton 31842 (NBG, NH, PRE).

LYCOPODIACEAE

Huperzia

gnidioides {L.f.) Trevis. [Lycopodium gnidioides L.f.]. 1, 3

-2631 (Mbabane): falls 16 km from Manzini (Bremersdorp), (-AD),

3 July 1953, Schutte 48 (BOL).

verticillata (L.f.) Trevis. [Lycopodium verticillatum L.f.], 1, 3

-2631 (Mbabane): falls 16 km from Manzini (Bremer.sdorp), (-AD),

3 Apr. 1953, Schutte 49 (BOL).

Lycopodium

clavatum L., 1,3

-2531 (Komatipoort): Havelock Concession, (-CC). 15Sept. 1890, Stdt-

marshe 1022 (BOL, PRE).

Lycopodiella

sarcocaulon {A. Braun & Welw. ex Kuhn) Pic.Serm. [Lycopodium car-

olinianum L.; L. carolinianum L. var. grandifolium Spring], 2

-2631 (Mbabane): Stroma, wet black peat below spring, ± 1 216 m,

(-AC), 4 July 1961, Compton 30675 (SDNH).

cernua (L.) Pic.Serm. [Lycopodium cernuum L.], 1, 3

-2531 (Komatipoort): Komassan Ridge, (-CC), 22 July 1890, Sah-

marshe 982 (PRE).

SELAGINELLACEAE

Selaginella

dregei (C.Presl) Hieron., 1, 3

-2631 (Mbabane): Forbes Reef, (-AA), 2 Jan. 1905, Burn Davy 2794

(PRE).

imbricata (Forssk.) Spring ex Decne., 2, 3

-2632 (Bela Vista): Mbuluzi Gorge. Blue Jay Ranch, S of gorge, 3

mis NNW of Mhlumeni border post, (-AA), 12 Nov. 1977,

Culvenvell 1174 iPRE).

kraussiana (Kunze) A.Braun, 1, 3

-2531 (Komatipoort): 19.3 km from Bulembu (Havelock), moist kloof,

(-CC), 30 June 1953, Schutte 1 (BOL).

mittenii Baker. 1 . 3

-2631 (Mbabane): Mbabane, Ukutula, moist shady rocks, (-AC), 15

Eebr. 1955, Compton 24932 (PRE).

OPHIOGLOSSACEAE

Ophioglossum

lusoafricanum Welw. ex Prantl

-2631 (Mbabane): Malolotja Nature Reserve, wardens house, near

stream above road to house, (-AA), 5 March 1989, Braun 691 (PRE).

reticulatum L., 1,3

-2631 (Mbabane): Mbabane, Secreterial garden, ± I 216 m, (-AC), 5

Jan. 1956, Compton 25274 (SDNH).

MARATTIACEAE

Marattia fraxinea Sm. [M. fraxinea Sm. var. salicifolia (Schrad.) C.Chr.],

1,3

-2631 (Mbabane): Mbabane, Ukutula, (-AC), 7 May 1955, Compton

25120 (PRE).

54

Bothalia 33,1 (2003)

OSMUNDACEAE

Osmunda regalis L., 1,3

-2631 (Mbabane): Mbabane, by stream, (-AC), 18 Jan. 1951,

Compton 22441 (NBG),

Todea barbara (L) T.Moore, 2, 3

-2531 (Komatipoort): Horo Forest, (-CB), Wager 125 (PRE).

GLEICHENIACEAE

Gleichenia

polypodioides (L.) Sm„ 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, ± 1 520 m,

(-CC), 9 May 1958, Compton 27835 (NBG, PRE, SDNH).

umbraculifera (Kimze) T.Moore. 1, 3

-2631 (Mbabane): Mbabane, bushy places, (-AC), July 1951, Comp-

ton 22745 (NBG).

Dicranopteris linearis (Burm.f) Underw., 1, 3

-2631 (Mbabane): Mbabane, hilltops, (-AC), 1917, Lovemore .s.n.

(BOL).

HYMENOPHYLLACEAE

Hymenophyllum tunbridgense {L.) Sm., 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, occasion-

al on wet dripping rock faces in deep shade in forest, ± 1 520 m,

(-CC), 13 July 1956, Schelpe 6185 (BOL).

Crepidomanes

borbonicum (Bosch) J.P.Roux [Trichomanes borbonicum Bosch], 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, locally

common on wet rock faces in deep shade in forest, ± 1 368 m,

(-CC), 14 July 1956, Schelpe 6186 (BM, BOL, K).

inopinatum (Pic. Serin.) J.P.Roux

-2631 (Mbabane): Mbabane, hill NE of Mbabane, ± 1 368 m, (-AC),

21 Nov. 1957, Compton 27250 (NBG, PRE, SDNH). Material be-

longing to this species has often been identified as Trichomanes pyx-

idiferum L. var. melanotrichiim (now Crepidomanes melanotrichum

(Schltdl.) J.P.Roux).

melanotrichum (Schltdl.) J.P.Roux [Trichomanes pyxidiferum L. var.

melanotrichum (Schltdl.) Schelpe], 1, 3

-2531 (Komatipoort): Bulembu, King’s Forest, infrequent in deep shade

on rocks in forest, 1 259 m, (-CC), 7 May 2002, J.P Roux 3398 (NBG).

Cephalomanes rigidum (Sw.) K.lwats. [Trichomanes rigidum Sw.], 3

-2631 (Komatipoort): Mdzimba Mts, Manzimnyana Falls, infrequent

in wet mossy areas among Osmunda along main stream, shaded,

(-AD), 26'’25.140’S, 3n6.178’E, 10 May 2002, J.P. Roux 3514

(NBG). It is not clear if Wager 190 (PRE), made in Lomati Gorge,

originates from Swaziland or from South Africa.

SCHIZAEACEAE

Schizaea pectinata (L.) Sw.. 1, 3

-2631 (Mbabane): Mbabane, Ukutula, (-AC), 27 Febr. 1955, Compton

24971 (NBG).

ANEMIACEAE

Anemia dregeana Kunze. 2

-2531 (Komatipoort): Horo Forest, (-CB), Wager 126 (NBG).

Mohria vestita Baker [as M. caffrorum (L.) Desv.], 1, 3

-2531 (Komatipoort): Horo Forest, (-CB), Wager 117 (PRE).

PARKERIACEAE

Ceratopteris thalictroides (L) Brongn.. 2, 3

-2632 (Bela Vista): Lubombo Mts, Mbuluzi Poort, Mlawula Farm,

Nkumbane stream, (-AA), 22 May 1976, Ciilverwell 95 (PRE).

PTERIDACEAE

Pityrogramma calomelanos (L.) Link var. aureoflava (Hook.) Weath. ex

F.M. Bailey

-2631 (Mbabane): Malolotja Nature Reserve, Ngwenya Mine, park-

ing area between pits, (-AA), 24 May 1990, Braun 962 (PRE). This

species, a native of tropical America, has been introduced to various

parts of the world. In South Africa it is known from Mpumalanga,

KwaZulu-Natal and Eastern Cape. The first known collection from

Swaziland is that of K.P. Braun made on 24 May 1990 at the

Ngwenya Mine, now included in (he Malolotja Nature Reserve. The

species is now widespread in the western part of the country where

it mostly inhabits disturbed sites.

Actinioptcris radiata (J.Kdnig ex Sw.) Link. 2. 3

-2631 (Mbabane): Lubombo Mts, 17 km N of Siteki on Siteki/Mhlu-

meni road between Groenpan and Cyrildene Farms, (-BD), 8 May

1976, Culverwell 72 (PRE),

C’heilanthes

bergiana .Schltdl.. 2, 3

-2531 (Komatipoort): Horo Forest, (-CB), Wager 128 (PRE).

cckloniana (Kunz.e) Melt.. 3

-2631 (Mbabane): Hlatikhulu, (-CD), 13 Nov. 1956, Compton 26231

(PRE).

multifida (Sw.) Sw. [including C. multifida (Sw.) Sw. subsp. lacerata

N.C. Anthony & Schelpe], 3

-2631 (Mbabane): Mbabane, (-AC), 15 Jan. 1951, Compton 22384

(NBG).

quadripinnata (Fomk.) Kuhn [Pellaea quadripinnata (Forssk.) Prantl],

3

-2531 (Komatipoort): 19.3 km from Bulembu (Havelock), moist kloof,

(-CD), 30 June 1953, Schiitte 11 (BOL).

viridis (Forssk.) Sw.

var. viridis [Pellaea viridis (Forssk.) Prantl var. viridis]. 1, 3

-2531 (Komatipoort): 19.3 km from Bulembu (Havelock), moist

kloof, (-CC), 30 June 1953, Schutte 66 (BOL).

var. glauca (Sim.) Schelpe & N.C.Anthony [Pellaea viridis (Forssk.)

Prantl var. glauca (Sim) Sim], 1, 3

-2631 (Mbabane): Mbabane, rock crevices, (-AC), 5 Febr. 1952,

Compton 23295 (NBG, NH).

var. macrophylla (Kunze) Schelpe & N.C.Anthony [Pellaea viridis

(Forssk.) Prantl var. macrophylla (Kunze) Sim], 1, 3

-2632 (Bela Vista): Jilobi Forest, forest floor, 500 m, (-CA), 1 1 Jan.

1979, Kemp 1480 (SDNH),

Pellaea

calomelanos (Sw.) Link var. calomelanos, 1, 3

-2631 (Mbabane): Mbabane area, (-AC), 1 July 1953, Schutte 27 (BOL).

dura (Willd.) Hook., 1, 3

-2631 (Mbabane): Mbabane, Gobholo, ± 1 160 m, (-AC), 18 Apr. 1963,

DIamini s.n. (SDNH).

pectiniformis Baker

-2731 (Louwsburg): Mahamba, Mahamba Gorge, N-facing quartzitic

cliffs above Mkhondvo River, in narrow exposed rock crevices,

(-AA), 27°03.984’S, 31°03.359’E, 18 May 2002, J.P. Roux 3711

(NBG). In South Africa the species has until now been known from

Limpopo [Northern Province], Mpumalanga, Gauteng and North-

West and further south from south-central KwaZulu-Natal. This col-

lection narrows the gap between these northern and southern distri-

butions.

Doryopteris concolor (Langsd. & Fisch.) Kuhn [Doryopteris concolor

(Langsd. & Fisch.) Kuhn var. kirkii (Hook.) R.E.Fr.; Cheilanthes

concolor (Langsd, & Fisch.) R.M.Tryon & A.F.Tryon], 1, 3

-2631 (Mbabane): Mbabane, Mantenga Falls, rocks, (-AC), 29 Jan.

1964, Compton 31921 (PRE).

Adiantum

capillus- veneris L., 1,3

-2631 (Mbabane): Hlatikhulu, Mamiti River, shady streambank, + 608

m, (-CD), Compton 27956 (NBG, PRE).

poiretii J.E.Wikstr. [Adiantum thalictroides Willd. & Schltdl.], 1, 3

-2631 (Mbabane): Mbabane. Darkton, streambanks, ± 1 368 m, (-AC),

Compton 27869 (NBG, PRE, SDNH),

raddianum C.Presl

-2531 (Komatipoort): Bulembu, ± 1.5 km from Bulembu on Pigg’s Peak

road, common on moist, shaded roadcutting, (-CC), 25°56.661’S,

3 1 °08.672’E, 7 May 2002, J.P. Roux 3426 (NBG). This tropical Ameri-

can species has become naturalized in many parts of southern Africa.

Pteris

catoptera Kunze

var. catoptera, 1 , 3

-2631 (Mbabane): falls 16.09 km from Manzini (Bremersdorp),

(-AD), 3 July 1953, Schiitte 42 (BOL).

var. horridula Schelpe

-2631 (Mbabane): Ntondozi Summit, rare in deep shade in forest

along seasonal drainage line, (-CA), 26°37.897’S, 31°10.017’E,

1 274 m, 15 May 2002, J.P. Roux 3575 (NBG). This variety has

until now been known from only two collections in southern

Africa, the one from Mpumalanga and the other from the

Drakensberg in KwaZulu-Natal (Burrows & Crouch 1995), The

Swaziland collection narrows the gap between these disjunct sites,

cretica L.. 1,3

-263 1 (Mbabane): Mbabane, Stroma, forest, ± 1 216 m, (-AC), Comp-

ton 25828 (NBG, PRE, SDNH).

dentata Forssk.

-2531 (Komatipoort): ± 3.2 km from Bulembu on road to Pigg’s

Peak, on moist, shaded roadcutting, (-CC), 25°56.722’S, 31°084’E,

7 May 2002, J.P. Roux 3419 (NBG). Widespread in southern Africa,

occurring in Mpumalanga and neighbouring Swaziland, Until now

the species has gone unnoticed by collectors in Swaziland,

friesii Hieron., 3

-2631 (Mbahinc): Siteki, moist kkxif, (-BD), 8 July 1953, Schutte 58 (BOL).

viltata L., 1, 3

-2531 (Komatipoort): Bulembu (Havelock) mine dump, (-CC), 30

June 1953, Schiitte 21 (BOL).

Bothalia 33,1 (2003)

55

VITTARIACEAE

Vittaria isoetifolia Bory

-2531 (Komatipoort); Bulembu, King’s Forest, epiphytic on Cyathea

dregei in forest, deeply shaded, (-CC), 25°56.317’S, 31°08.028’E, 1

316 m, 7 May 2002, J.P. Roux 3397 (NBG). Occurs along moister

southern and eastern parts of South Africa with no collections

recorded between northern KwaZulu-Natal and southeastern

Mpumalanga. The Swaziland collection slightly narrows the gap

between these disjunct distributions.

CYATHEACEAE

Cyathea

capensis (Lf.) Sm„ 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, frequent

in wet forest, ± 1 520 m, (-CC), 13 July 1956, Schelpe 6181 (BOL).

dregei Kimze, 1, 3

-2631 (Mbabane): mts above Mbabane, (-AC), 19 May 1948, Rodin

4526 {BOL, PRE).

DENNSTAEDTIACEAE

Microlepia speluncae (L) TMoore

-2531 (Komatipoort): Songea Range, Hhoro Forest, in partial shade

along main stream, (-CB), 25°44.215’S, 31°24.218’E, 8 May 2002,

J.P. Roux 3442 (NBG). Has a somewhat unusual and scattered dis-

tribution in southern Africa with isolated collections known from

Limpopo [Northern Province], North-West and KwaZulu-Natal.

This new record for Swaziland occurs in a somewhat isolated ravine

and may be the reason why it has escaped recording.

Pteridium aquilinum (L.) Kuhn subsp. aquilinum, 1, 3

-2531 (Komatipoort): between Bulembu (Havelock) Mine and Pigg’s

Peak, (-CC), 30 June 1953, Schutte 19 (BOL).

Hypolepis sparsisora (Schrad.) Kuhn, 1, 3

-2531 (Komatipoort): 19.3 km from Bulembu (Havelock), moist

kloof, (-CC), 30 June 1953, Schutte 8 (BOL).

Blotiella glabra (Bory) R.M.Tryon, 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, ± 1 520

m, (-CC), 9 May 1958, Compton 27833 (NBG. PRE, SDNH).

POLYPODIACEAE

Lepisorus schraderi (Men.) Ching [Pleopeltis schraderi (Mett.) Tardieu],

1, 3

-2631 (Mbabane): Hlatikhulu, forest margin, ± 1 216 m, (-CD), Comp-

ton 26364 (NBG, SDNH).

Polypodium polypodioides (L.) Watt, subsp. ecklonii (Kimze) Schelpe,

1. 3

-2631 (Mbabane): The Caves. Mbuluzi 'Valley, shady rocks, ± 1 368

m, (-AC), 17 March 1956, Compton 25798 (NH, PRE).

Microgramma mauritiana (Willd.) Tardieu [as M. lycopodioides (L.)

Copek], 1, 3

-2632 (Bela Vista): Siteki, Jilobi Forest, along rocks and twigs, ± 608

m. (-CA), 14 Sept. 1961, Compton 30718 (NBG, PRE, SDNH).

Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. [as Pleopeltis lanceolata

(L.) Kaulf.], 1, 3

-2631 (Mbabane): Mbabane, (-AC), 1905. Burn Davy 2796 (PRE).

Loxogranune abyssinica (Baker) M.G. Price [as Loxogramme lanceola-

ta (Sw.) C.Presl], 1, 3

-2631 (Mbabane): hill NE of Mbabane, shady rocks in forest, ± 1 368

m, (-AC), 21 Nov. 1957, Compton 27251 (SDNH).

NEPHROLEPIDACEAE

Nephrolepis exaltata (L.) Schott

-2631 (Mbabane): Mbabane area, (-AC), 1 July 1953, Schutte 30

(BOL). This species is native to tropical America, but has natural-

ized in many parts of South Africa. Although it has been recorded as

early as 1953 from the Mbabane area, it has never been taken up in

any of the checklists. It is, however, cited by Schelpe & Anthony

( 1986). The species has now also become naturalized at Bulembu.

THELYPTERIDACEAE

Ampelopteris prolifera (Retz.) Copel.

-2631 (Mbabane): old bridge across Komati River, S bank, moist

area E of road, (-AA), 30 Jan. 1992, Braun 1300 (PRE).

Cyclosorus intemiptus (Willd.) H.lto [Thelypteris interrupta (Willd.)

K.Iwats.; T. totta (Thunb.) Schelpe], 1, 3

-2631 (Mbabane): 19.3 km from Manzini, (-AD), 1 July 1953, Schutte

35 (BOL).

Amauropelta bergiana (Schltdl.) Holttum var. bergiana [Thelypteris

bergiana (Schltdl.) Ching], 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, frequent

in forest undergrowth in moist conditions, ± 1 520 m, (-CC), Schelpe

6166 (BOL).

Thelypteris confluens (Thunb.) C.V.Morton [T. palustris Schott var.

squamigera (Schltdl.) Weath.], 1, 3

-2631 (Mbabane): Mbabane, Pholinjane River, vlei, ± 1 216 m, (-AC),

18 Apr. 1956, Compton 25897 (NBG, PRE, SDNH).

Stegnogramma pozoi (Lag.) K.Iwats.

-2531 (Komatipoort): Bulembu, King’s Forest, frequent in deep

shade along moist streambanks in forest, (-CC), 25°56.317’S,

31°08.028’E, 1 316 m, 7 May 2002, J.P. Roux 3406 (NBG). This

collection falls within the general north-south distribution of the

moisture -dependent species in South Africa. This collection narrows

the gap between the KwaZulu-Natal and Mpumalanga distributions.

Pneumatopteris unita (Kimze) Holttum [Thelypteris madagascariensis

(Fee) Schelpe], 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, locally

common in deep shade in wet forest undergrowth, ± 1 520 m, (-CC),

13 July 1956, Schelpe 6180 (BOL).

Christella

dentata (Forssk.) Brownsey & Jenny [Thelypteris dentata (Forssk.)

E.P.St.John], 1, 3

-2631 (Mbabane): Mbabane, forest, (-AC), July 1951, Compton 22748

(NBG).

hispidula (Decne.) Holttum

-2531 (Komatipoort): Songea Range, Hhoro Forest, along main stream

in forest, moist conditions, (-CB), 25°44.456’S, 31°23.667’E, 8 May

2002, J.P. Roux 3438 (NBG). This pantropical species has only

recently been recorded for the first time from KwaZulu-Natal, pre-

viously known from eastern Zimbabwe and further north (Burrows

& Burrows 2001). The Swaziland collections narrow these disjunct

distributions.

gueinziana (Mett.) Holttum [Thelypteris gueinziana (Mett.) Schelpe],

3

-2631 (Mbabane): 19.3 km from Manzini, (-AD), 1 July 1953, Schutte

34 (BOL).

DRYOPTERIDACEAE

Nothoperanema squamiseta (Hook.) Ching [Dryopteris squamiseta

(Hook.) Kuntze], 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, wet rock

faces in deep shade, ± 1 520 m, (-CC), 13 July 1956, Schelpe 6178

(BOL).

Dryopteris

athamantica (Kimze) Kuntze, 1, 3

-2631 (Mbabane): Mbabane, Ukutula, (-AC), 10 Nov. 1954, Comp-

ton 24689 (NBG).

lewalleana Pic. Serin.

-2631 (Mbabane): Mbabane, Ukutula, edge of forest, ± 1 216 m,

(-AC), 16 May 1956, Compton 25922 (NBG, PRE, SDNH). D.

lewalleana forms part of the D. inaequalis species complex with the

result that material belonging to it have always been misidentified.

pentheri (Krasser) C.Chr.

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, ± 1 520

m, (-CC), Compton 27829 (NBG, SDNH). Like D. lewalleana, D.

pentheri has generally also been included into D. inaequalis. It is a

tetraploid species more frequently inhabiting exposed environments.

Polystichum

luctuosum (Kimze) TMoore, 1, 3

-2631 (Mbabane): Stroma, (-AC), 23 March 1956, Compton 25822

(NBG, PRE, SDNH).

macleae (Baker) Diels, [as 'macleaii'], 1, 3

-2631 (Mbabane): Mbabane, Ngwenya Mtn, (-AA), 27 March 1962,

Compton 31405 (NBG, SDNH).

pungens (Kaulf.) C.Presl [as P. lucidum], 1, 3

-2631 (Mbabane): Miller’s Falls, ± 1 368 m, (-AC), 16 May 1956,

Compton 25922 (NBG, PRE, SDNH).

transkeiense W.Jacobsen

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, ± 1 520

m, (-CC), 9 May 1958, Compton 27831 (NBG, PRE). Material be-

longing to this species has always been erroneously identified as

Polystichum pungens.

Didymochlaena truncatula (Sm'.) J.Sm.

-2531 (Komatipoort): Bulembu, King’s Forest, frequent in deep

shade along stream in forest, (-CC), 25°56.317’S, 31°08.028’E,

1 3 16 m, 7 May 2002, J.P. Roux 3402 (NBG); Farm Wyldsdale, for-

est on S-face of Lufafa, common in streambed in deeply shaded for-

est, (-CD), 25°48.596’S, 31°16.341’E, 12 May 2002, J.P. Roux

3553 (NBG). This moisture-loving species is relatively common at

both sites, where it was collected, growing along perennial streams

in deep shade.

56

Bothalia 33,1 (2003)

TECTARIACEAE

Tectaria gemmifera {Fee) Alston

-2531 (Komatipoort): Songea Range, Hhoro Eorest, common in deep

shade on forest Boor, (-CB), 25°44.215'S, 31°24.218’E, 8 May

2002, Roux 3451 (NBG). Relatively localized in South Africa with

scattered disjunct populations occurring in Limpopo, Mpumalanga,

and KwaZulu-Natal. At both sites in Swaziland the species is com-

mon, often forming large monospecific stands on moist, not too

deeply shaded forest floors,

Megalastrum lanuginosum (Willd. ex Kaulf) Holttum

-2531 (Komatipoort): Bulembu, King’s Forest, frequent along stream

in deep shade in forest, (-CC), 25°56.785’S, 31°12.263’E, 1 259 m,

7 May 2002, J.P. Roux 3414 (NBG). It is remarkable that this majes-

tic fern, widespread in southern and East Africa, the Madagascan

region and the islands in the Gulf of Guinea, has gone unnoticed in

Swaziland. In southern Africa it falls within the north-south distrib-

ution of most mesophytic ferns,

WOODSIACEAE

Athyrium scandicinum (Willd.) C.Presl, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, rare on

wet rock-face in forest, ± 1 520 m, (-CC), 13 July 1956, Schelpe 6177

(BOL).

Deparia japonica (Thunb.) M.Kato

-2531 (Komatipoort): Farm Wyldsdale, forest in ravine on S face of

Lufafa, (-CD), 25°48.092’S, 31°16.852’E, 11 May 2002, J.P Roux

3549 (NBG). Deparia japonica, a native of Asia and Australia, has

become naturalized in South Africa, recorded from KwaZulu-Natal

and Mpumalanga. For Swaziland the species was first recorded from

the Pigg’s Peak area (Burrows 1990). The species now appears to be

more widespread in Swaziland with collections made near Mbabane

and at Hlatikhulu.

OLEANDRACEAE

Oleandra distenta Kunze. 1, 3

-2631 (Mbabane): Mbabane, The Caves, Black Mbuluzi Valley, rock

clefts, ± 1 368 m, (-AC), 17 March 1956, Compton 25799 (NBG, PRE,

SDNH).

Arthropteris monocarpa (Cordetn.) C.Clir.

-2531 (Komatipoort): Farm Wyldsdale, Ugutugulo River, on season-

ally moist S-facing cliffs and in screes above river, (-CD),

25°48.092’S, 31°16.852’E, 11 May 2002, J.P. Roux 3533 (NBG).

Known from KwaZulu-Natal and an isolated collection from

Mpumalanga, the Swaziland collection effectively narrows the gap

between these disjunct distributions.

LOMARIOPSIDACEAE

Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe, 1, 2

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, occasion-

al on mossy boulders on forest floor, ± 1 520 m, (-CC), 13 July

1956, Schelpe 6174 (BOL).

BLECHNACEAE

Blechnum

attenuatum (Sw.) Mett. [Blechnum giganteum (Kaulf.) Schltdk], 3

-2531 (Komatipoort): 19.03 km from Bulembu (Havelock), moist kloof,

(-CC), 30 June 1953, SchUtte 7 (BOL).

australe L, 1,3

-2631 (Mbabane): Mbabane, Miller’s Falls, forest floor, ± 1 368 m,

(-AC), 7 June 1956, Compton 25969 (NBG, SDNH).

capense Burm.f. [Blechnum sylvaticum Schelpe], 3

-2631 (Mbabane): 16.09 km from Manzini, (-AD), 3 July 1953,

Schiiile 52 (BOL).

intlexum (Kunze) Kuhn

-2631 (Mbabane), Ntondozi summit, infrequent in shade among

bushes and granite rocks, (-CA), 26°37.897’S, 31°10.017’E, 1 274

m, 15 May 2002, J.P. Roux 3580 (NBG). Restricted to the southern

and eastern mountains ranging from the western Cape to eastern

Zimbabwe. The Swaziland collection falls within this north-south

distribution,

punctulatum ,Sw.

var. punctulatum. 1, 3

-2631 (Mbabane): Mbabane. Ngwenya Mts, shady rocks, ± 1 672

m, (-AA), 18 Febr. 1957, Compton 26680 (NBG, SDNH).

var. alherstonei (Pappe & Raw.son) Sim

-2631 (Mbabane): Mbabane, Fonteyn, shade of boulders, ± 1 368

m, (-AC), 8 June 1956, Compton 25979 (NBG, NH, SDNH).

tabulare (Thunb.) Kuhn, 1, 3

-2531 (Komatipoort): 19.3 km from Bulembu (Havelock), moist kloof,

(-CC), 13 June 1953, Schiiile 5 (BOL).

ASPLENIACEAE

Asplenium aethiopicum (Burm.f. ) Bech.

subsp. aethiopicum, 1, 3

-2631 (Mbabane): near Forbes Reef, ± 1 520 m. (-AA), Compton

29638 (NBG, SDNH). Braithwaite (1986) showed that A.

aethiopicum subsp. aethiopicum forms part of an extensive poly-

ploid series, this subspecies being a sexual octoploid.

subsp, tripinnatum (Baker) A.F.Braithw.

-2631 (Mbabane): Mbabane, Duiker Bush, ± 1 520 m, (-AC),

Compton 25541 (NBG, PRE, SDNH). This subspecies is a sexual

tetraploid (Braithwaite 1986).

subsp. filare (Forssk.) A.F.Braithw.

-2631 (Mbabane): 16.09 km from Manzini (Bremersdorp). (-AD),

SchUtte 55 (BOL). This material belongs to octoploid apomict

pteudq/i/are (Braithwaite 1986)

anisophyllum Kunze, 1 , 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, occasion-

al on wet rock faces in shade of forest, ± 1 520 m, (-CC), 1 3 July 1956,

Schelpe 6175 (BOL).

boltonii Hook, ex Brause cfe Hieron.

-2531 (Komatipoort): Pigg’s Peak, Devil’s Bridge Forest, (-CD), 4

May 1960, Compton 30042 (NBG, PRE, SDNH). The collection

was initially erroneously identified as A. anisophyllum Kunze.

erectum Bory ex Willd.

-2631 (Mbabane): Mbabane, Malandela, ± 1 216m, (-AB), 10 Feb. 1965,

Compton 32260 (NBG, PRE). The collection was initially erroneous-

ly identified as A. lunulatum Sw.

inaequilaterale Willd., 3

-2531 (Komatipoort): Horo Forest, (-CB), Apr. 1932, Wager 127 (BWE).

lobatum Pappe & Rawson, 1 , 3

-2631 (Mbabane): Mbabane. Malandela, dense forest, ± 1 216 m (AB),

19 Febr. 1965, Compton 32271 (NBG, PRE, SDNH),

multiforme Krasser

-2731 (Louwsburg): ridge E of Grandis Mountains, near Galile school,

road between Hlatikhulu and Nhlangano, (-AA), 27°00.785’S,

31°14.568’E, 17 May 2002, J.P. Roux 3706 (NBG). This is an octo-

ploid belonging to the A. splendens Kunze polyploid complex

(Braithwaite 1972). The species is also ecologically distinct, as it

inhabits rock crevices in seasonally dry, more exposed environments,

rutifolium (P.J.Bergius) Kunze, 1, 3

-2631 (Mbabane): Forbes Reef, (-AA), 1 Jan. 1905, Burtt Davy 2731

(PRE).

sandersonii Hook.

-2531 (Komatipoort): Farm Wyldsdale, forest in ravine on the south-

ern face of Lufafa, (-CD), 25°48.596’S. 31°16.341’E, 12 May 2002,

J.P. Roux 3551 (NBG). Widespread in Central and East Africa

extending to Eastern Cape in the south. The Swaziland collection

slightly narrows the disjunct Eastern Cape/KwaZulu-Natal popula-

tions and the Mpumalanga/Limpopo populations,

splendens Kunze subsp. splendens. 1, 3

-2531 (Komatipoort): Bulembu (Havelock), King’s Forest, frequent

on forest floor, ± 1 520 m, (-CC), 13 July 1956, Schelpe 6155 (BOL).

theciferum (Humb. Bonpl. et Kunth) Mett., 1, 3

-2631 (Mbabane): Hlatikhulu, forest, ± 1 216 m, (-CD), 13 Nov. 1956,

Compton 26243 (NBG, SDNH).

unilaterale Lam.

-2531 (Komatipoort): Farm Wyldsdale. Ugutugulo River, on rocks in

seepage area, (-CD), 25'’48.092’S, 31°16.852’E, 11 May 2002, J.P.

Roux 3536 (NBG). Widespread in the palaeotropics, has recently

been recorded for the first time in southern Africa (Burrows &

Burrows 2002). By coincidence the same species was collected

independently on the same day in the same ravine.

MARSILEACEAE

Marsilea

ephippiocarpa A/.s-ro/), 1, 3

-2632 (Bela Vista): Mlawula Station, Mlawula Farm, Nkumbane stream,

3.2 km S of station, E of farm track, (-AA), 29 May 1977, Culver-

well 840 (PRE).

fenestrata Launert, 3

-2631 (Mbabane): Siteki Dist., Hlane, swamp, ± 304 m, (-BB), 18Jan.

1960, Compton 29735 (NBG, SDNH).

minuta L. var. minuta. 2, 3

-2632 (Bela Vista): Mbuluzi River, Mlawula Farm. 4 km NE of station,

(-AA), 29 May 1977, Culverwell 807 (PRE).

Species excluded

Anemia simii Tardieu, 1

Complon (1966) lists the species as ‘authentically recorded but not

represented in the herbarium’. Jacobsen ( 1983) also lists the species

Bothalia 33,1 (2003)

57

for Swaziland, but no herbarium collections from this country were

located.

Blechnum aitriculalum Cav., 1

A Compton collection lodged in the Compton Herbarium (NBG)

named as such belongs to Blechnum punctulatum Sw. var. punctula-

tum.

Cheilanthes involuta (Sw.) Schelpe & N.C. Anthony, 3

No collections belonging to this species could be traced to verify its

occurrence in Swaziland.

Cheilanthes involuta (Sw.) Schelpe & N.C. Anthony var. obscura

(N.C. Anthony) N.C. Anthony [Cheilanthes viridis (Forssk.) Sw.

var. obscura N.C. Anthony], 3

No collections belonging to this species could be traced to verify its

occurrence in Swaziland.

Marsilea aegyptiaca Willd., 2

The species was first listed by Kemp (1981). based on Culvenvell

840 (PRE). The collection has since been identified as belonging to

M. ephippiocarpa Alston.

ACKNOWLEDGEMENTS

I would like to express my gratitude to the Steering

Committee of the Southern African Botanical Diversity

Network (SABONET) and Prof. G.F. Smith for support-

ing this project. My thanks also go to Kate Braun who

shared and provided valuable information whilst in

Mbabane; the National Herbarium, Pretoria, for provid-

ing a printout of their Swaziland holdings; and the

Curators of the Bolus (BOL) and Natal (NH) Herbaria

for allowing me to work through their collections. The

assistance of the staff at the National Herbarium of

Swaziland is also appreciated, and the Swaziland

National Trust Commission is thanked for allowing

access to the national parks.

REFERENCES

BRAITHWAITE, A.F. 1972. The cytotaxonomy of the Asplenium splen-

dens complex in South Africa. Journal of South African Botany

38: 9-27.

BRAITHWAITE, A.F. 1986. TYis Asplenium aethiopicum complex in South

Africa. Botanical Journal of the Linnean Society 93: 343-378.

BURROWS, J.E. 1990. Southern African ferns and fern allies.

Frandsen Publishers, Sandton.

BURROWS, J.E. & BURROWS, S.M. 2001. New distribution records

for southern African Pteridophyta. Bothalia 31: 205-207.

BURROWS. J.E. & BURROWS, S.M. 2002. A new fern record for the

Flora of southern Africa region. Bothalia 32: 195, 196.

BURROWS. J.E. & CROUCH, N.R. 1995. New distribution records of

South African pteridophytes. Bothalia 25: 236-238.

COMPTON. R.H. 1966. An annotated check list of the flora of Swazi-

land. Journal of South African Botany, suppl. vol. 6.

JACOBSEN, W.B.G. 1983. The ferns and fern allies of southern Africa.

Butterworth, Durban.

KEMP, E.S. 1981. Additions and name changes for the flora of Swazi-

land. Swaziland National Trust Commission.

KEMP, E.S. 1983. A flora checklist for Swaziland. Occasional Papers 2,

Swaziland National Trust Commission.

ROUX, J.P. 2001. Con.spectus of southern A frican Pteridophyta. South-

ern African Botanical Diversity Network Report No. 13. Cape

Town.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. In O.A.

Leistner, Flora of .southern Africa. Department of Agriculture

and Water Supply, Pretoria.

Bothalia 33,1: 59-98 (2003)

Taxonomy of the genus Passerina (Thymelaeaceae)

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

Keywords: anatomy, cladistics, endemism, macromorphology, new species, palynology, Passerina, southern Africa, taxonomy, Thymelaeaceae

ABSTRACT

Passerina L, is mainly a southern African genus, comprising 20 species and four subspecies, A few species occur along

the Great Escarpment, two extend into Zimbabwe and Mozambique, but most are concentrated in the Cape Floristic Region.

Palynological, macromorphological and anatomical evidence was used in the delimitation of the genus and its infrageneric

taxa, A cladistic study supports Passerina as a monophyletic genus, A genus treatment, key to species and a full species

treatment are given. Each species treatment includes a taxonomic diagnosis, description and notes on taxonomy, etymolo-

gy. economic value and distribution. Illustrations of representative species are provided and distribution maps are included

for each species. P. esterhuyseniae Bredenk. & A.E. van Wyk is newly described. A list of excluded species names high-

lights the previous cosmopolitan taxonomic interpretation of Passerina, as many names are now in synonymy under other

genera of the Thymelaeaceae.

CONTENTS

Abstract 59

Introduction 59

Thymelaeaceae 59

Passerina 60

Materials and methods 61

Passerina E 61

Key to species 65

1 . P. paleacea Wikstr 66

2. P. trimcata (Meisn.) Bredenk. & A.E.van Wyk ... 68

[for a complete description see Bredenkamp

& Van Wyk in Bothalia 32: 66 (2002a)]

3. Passerimi qiiadrifaria Bredenk. & A.E.van Wyk ... 68

[for a complete description see Bredenkamp

& Van Wyk in South African Journal of

Botany 68: 304 (2002b)]

4. Passerina niontana Thoday 68

5. Passerina burchellii Thoday 70

6. Passerina ericoides L 71

7. Passerina rigida Wikstr 73

8. Passerina nivicola Bredenk. & A.E.van Wyk 74

[for a complete description see Bredenkamp &

Van Wyk in Bothalia 32: 77 (2002c)]

9. Passerimi esterlnryseniae Bredenk. & A.E.van Wyk ... 75

10. Passerina comosa (Meisn.) C.H. Wright 76

11. Passerina pendida Eckl. & Zeyh. ex Thoday .... 77

12. Passerina galpinii C.H. Wright 78

13. Passerina drakensbergensis Hilliard & B.L.Buitt ... 80

14. Passerina corymbosa Eckl. ex C.H. Wright 81

15. Passerina obtiisifolia Thoday 82

16. Passerina paludosa Thoday 84

17. Passerina montivaga Bredenk. & A.E.van Wyk ... 85

[for a complete description see Bredenkamp

& Van Wyk in Bothalia 32: 34 (2002d)]

* National Botanical Institute. Private Bag XlOl. 0001 Pretoria.

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

of Pretoria, 0002 Pretoria.

MS. received: 2002-1 1-08.

18. Passerina filifonnis L 85

[for a complete description see Bredenkamp &

Van Wyk in Bothalia 32: 29-34 (2002d)]

18a. subsp. filiformis 85

18b. subsp. glutinosa (Thoday) Bredenk. & A.E.van

Wyk 85

19. Passerina falcifolia (Meisn.) C.H. Wright 85

20. Passerina rubra C.H.Wright 87

Doubtful name 89

Excluded names 89

Specimens examined 92

Acknowledgements 94

References 95

Index to taxa (also doubtful and excluded names) 97

INTRODUCTION

Thymelaeaceae

Applying the sexual system of classification, Lin-

naeus (1754) placed the genus Passerina under Class

VIII, Octandria, 1. Monogynia. This system was followed

until De Jussieu (1789) instated the family Thymelae-

aceae, with the following genera: Dirca L., Lagetta Juss.,

Daphne L., Passerina L., Stellera L., Struthiola L. Dich-

naea L„ Dais L., Gnidia L., Nectandra Berg, and Quisqualis

L. Wikstrom (1818) accepted the Thymelaeaceae, but

based the infrafamilial classification on the number of

stamens, following Linnaeus. The most important con-

tributions towards the infrafamilial classification of the

Thymelaeaceae, based on morphological characters, were

made by Endlicher (1847), Meisner (1857), Bentham &

Hooker (1880) and Gilg (1894a). Using anatomical char-

acters, further contributions were made by Van Tieghem

(1893), Gilg (1894b) and Leandri (1930).

Domke (1934) proposed a widely adopted subfamilial

classification for the Thymelaeaceae and divided the family

into four subfamilies, namely Gonystyloideae, Aquilari-

oideae, Gilgiodaphnoideae and Thymelaeoideae. The genus

Passerina is classified under the Thymelaeoideae. Based

on palynological evidence Archangelsky (1971: fig. 10)

added the new subfamilies Octolepidoideae, Microsem-

60

Bothalia 33,1 (2003)

matoideae and Synadrodaphnoideae and raised the

Gonystyloideae to the family Gonystylaceae (also recog-

nized by Takhtajan 1997, amongst others). Bredenkamp

& Van Wyk (1996) published new evidence on the struc-

ture of the pollen wall in Passerina resulting in the ele-

vation of the subtribe Passerininae Endl. to the mono-

generic tribe Passerineae (Endl.) Bredenk. & A.E.van

Wyk. Evidence obtained from floral morphology, anato-

my, embryology and palynology indicates that the

Thymelaeaceae has a strong malvalean relationship, an

affinity also supported by molecular data (APG 1998;

Magallon et al. 1999). The possible phylogenetic rela-

tionships of the Thymelaeaceae are discussed by

Bredenkamp & Van Wyk (2001b).

The Thymelaeaceae is currently considered a family

of ± 58 genera and ± 720 species. (Mabberley 1989;

Brummitt 1992; Takhtajan 1997). It is subcosmopolitan

and the distribution of the genera is listed by Mabberley

( 1989), as follows:

Africa: temperate southern Africa — Dais L., Englero-

daplme Gilg, Gnidia L., Lachnaea L., Passerina L., Peddiea

Harv., Struthiola L., Synaptolepis Oliv.; tropical Africa —

Craterosiphon Engl. & Gilg, Dicranolepis Planch., Octo-

lepis Oliv., Synandrodaphne Gilg.

Asia: Aetoxylon Airy Shaw, Amyxa Tiegh., Drapetes

Lam., Eriosolena Blume, Pentathymelaea Lecomte, Rham-

noneuron Gilg, Restella Pobed., Wikstroemia Endl.

Australia: Arnhemia Airy Shaw, Drapetes Lam.,

Pimelea Banks & Sol., Oreodendron C.T.White.

Europe: Daphne L., Diarthron Turcz.

Japan: Daphnimorpha Nakai, Edgeworthia Meisn.

Madagascar: Stephanodaphne Baill.

Malesia: Aquilaria Lam., Enkleia Griff., Gonystylus

Teijsm. & Binn., Linostoma Wall, ex Endl., Phaleria Jack.

Mediterranean region: Thyme laea Mill.

New Ccdedonia: Deltaria Steenis, Lethedon Spreng.,

Solmsia Baill.

North and South America: Daphnopsis Mart. & Zucc.,

Dirca L., Eunifera Leandro ex C.A.Mey., Goodallia Benth.,

Digetta Juss., Lasiadenia Benth., Linodendron Griseb.,

D>phostoma Meisn., Ovidia Meisn., Schoenobiblus Mart.

Sri Lanka: Gyrinops Gaertn.

Perhaps the economically most important character in

the family is its tough fibrous bark. The bark of

Wikstroemia, Daphne, Edgeworthia, Thymelaea and

Daphnopsis is used for rope, and in the manufacturing of

bank notes and strong paper. Elexible shoots of Dirca are

used for baskets. Bark of Pimelea was used as a source

of twine by early settlers in Australia.

Many genera are also known for their medicinal

value. The wood of Wikstroemia is a source of incense

and that of W. ovata C.A.Mey. is a strong purge. In China

the bark of Daphne is used as an apparently safe and effi-

cient abortifacient; it contains the glycoside daphnin and

an acrid resin (mezerein) giving plants a bitter taste. The

decaying heartwood of Aquilaria malaccensis Lam. is

saturated with a resin which is the basis of incense and

when distilled it is used in perfume and medicine.

The genera Pimelea, Edgeworthia and Daphne are cul-

tivated for horticultural purposes. The scent of Daphne

flowers is camation-like and attractive to Lepidoptera;

some members are moth-pollinated. Gonystylus bancanus

(Miq.) Kurz. is a peat swamp-forest tree, with knee-roots.

Its lightweight commercial timber is used for dowelling

and is much exported from Indomalesia.

In southern Africa, the bark of various genera is used

for tying down thatch, for plaiting into whip thongs and

for twine. Dais cotinifolia L. is an ornamental tree with

attractive flowers, occurring mostly along the eastern

regions of the country.

Passerina

In his comprehensive work on the circumscription of

the Thymelaeaceae and infrafamilial taxa, Domke (1934)

gave a complete historical review of the intergeneric clas-

sification of Passerina. He included the southern African

genera Dais, Gnidia (= Lasiosiphon), Struthiola, Lachnaea

(= Cryptadenia) and Passerina in the tribe Gnidieae, sub-

tribe Gnidiinae of the subfamily Thymelaeoideae. Breden-

kamp & Van Wyk (1996) place Passerina in the mono-

generic tribe Passerineae on the basis of mainly pollen

characters. Currently Passerina is considered advanced at

the intergeneric level, as many of the advanced character

states present in other genera of the Thymelaeoideae are all

found together in this genus. The most prominent charac-

ters distinguishing Passerina are the exserted, extrorse

anthers and the unique anemophilous habit (Bredenkamp

& Van Wyk 1996, 2001b).

The infrageneric classification of Passerina is docu-

mented by Linnaeus (1753) in his Species plantarum, in

which he described P. filiformis, P. hirsuta, P. ciliata

and P. uniflora. P. filiformis is the only species that is

currently recognized in Passerina. Publications men-

tioned in the applicable protologue and in synonymy to

the various species that pre-date the nomenclatural start-

ing point for the Spermatophyta [International Code of

Botanical Nomenclature, Article 13.1 [Greuter et al. 2000)]

are Linnaeus’s Hortus Cliffortianus (1737), Van Royen

(1740), Plukenet (1700: 180), Breyne (1678) and Burman

(1739). The generic name Passerina appearing in Species

plantarum (Linnaeus 1753) is associated with the subse-

quent description given in Genera plantarum (Linnaeus

1754) (Greuter et al. 2000, Article 13.4).

Wikstrom (1818) recognized 41 species of Passerina

and the subspecies P. filiformis subsp. divaricata', of these

only four species are presently recognized in Passerina. In

the interim the subspecies was raised to species level and is

presently known as P. falcifolia. Thunberg (1825a) recog-

nized nine species of which only one is currently main-

tained. His concept of P. glomerata, P. ericoides and

Lachnaea conglomerata were completely incorrect and

caused confusion right up to the present study. Meisner

(1840; 1857: 563-565) redefined the genus by clarifying 92

‘species exclusae’ which were mostly synonymous with

Bothalia33.I (2003)

61

Other cosmopolitan genera in the Thymelaeaceae and he

retained only four species and six subspecies. The distribu-

tion of the remaining species clearly indicated that

Passerina was a smaller genus, largely confined to southern

Africa. At the beginning of the 20th centui7, Wright (1915)

revised the Thymelaeaceae for the Flora capensis and his

generic concept of Passerina was mostly based on that of

Meisner ( 1857). He recognized ten species, of which three

were new, as well as three subspecies. He recognized P. eri-

coides and Chymococca empetroides. We agree with

Thoday ( 1924a) that C. empetroides is a synonym of P. eri-

coides. Although Thoday (1924a) provided a much

improved classification of the group, the circumscription

and identification of several species remained problematic,

especially in the herbarium. Table 1 is a summai'y of taxa

recognized in the most comprehensive works on Passerina

from Linnaeus (1753) to the present study.

In his treatment of Passerina, Meisner (1840) divided

the genus into section I. Pentamerae and section II.

Tetramerae. P.polycepliala E.Mey., P. anthylloides L.f and

P. calocephala Meisn., with pentamerous flowers (section

I), were eventually all placed in the genus Gnidia (Meisner

1857; Gilg 1894a). Meisner (1857) did not divide Passerina

into infrageneric taxa, a pattern followed by all subsequent

treatments and no further mention was made of the relevant

sections. In the present genus treatment the sectional classi-

fication is not maintained.

MATERIALS AND METHODS

Material from the following herbaria was studied (acro-

nyms according to Holmgren et al. 1990); BM, BOL,

BREM, C, GRA, K. LINN, M, MEL, MO, NBG, P, PR,

PRC, PRE, PRU, S, SBT, TCD, UPS, W, WU. A data-

base of the specimens was compiled on the Microsoft

Access Relational Database Management System for

Windows, Version 2.0.

Live and preserved (dried and in liquid preservatives)

material of all the species and subspecies in Passerina

was studied. As far as possible, material was collected

from at least five different localities for every taxon.

Illustrations were made from herbarium material by

means of a drawing tube. Measurements were taken

using a dissection microscope and a calibrated eyepiece.

Because the laminas of most leaves and floral bracts are

cymbiform or rolled, the depth was measured, with

dimensions indicated as length x depth.

Light microscopy (LM) was used for general leaf anato-

my, epidermal studies and floral anatomy (Bredenkamp &

Van Wyk 1999, 2001a, 2001b). Leaf and floral material

was fixed and stored in a 0. 1 M phosphate-buffered solu-

tion at pH 7.4, containing 2.5% formaldehyde, 0.1% glu-

taraldehyde and 0.5% caffeine [modified Kamovsky fixa-

tive; Kamovsky (1965)]. The material was washed in

water, dehydrated and embedded in glycol methacrylate

(GMA) following the methods of Eeder & O’Brien (1968).

Embedded material was serially sectioned. Sections were

stained in toluidine blue ‘O’, subjected to the periodic acid-

Schiff’s (PAS) reaction and mounted in Entellan (Art.

7961, E. Merck, Darmstadt).

Scanning electron microscopy (SEM) was used to

study the epidermal surface features (including epicuti-

cular waxes) and to verify the structure of the cuticle

(Bredenkamp & Van Wyk 2000).

Transmission electron microscopy (TEM) was used

for the study of the structure of mucilaginous epidermal

cell walls in Passerina (Bredenkamp & Van Wyk 1999).

Terminology used in the descriptions of inflores-

cences and flowers is mentioned in Bredenkamp & Van

Wyk (2001b). General descriptive terminology follows

Steam (1973) and Radford etcd. (1974). Author citations

follow Brummitt & Powell (1992).

Passerina L. Species plantarum: 559 ( 1753); L.: 168

(1754); P.J.Bergius: 126 (1767); Mill.: ( 1768); Burm.f: 12

(1768); L.: 236 (1771); L.: 225 (1782); L.: 374 (1784);

Thunb.: 75 (1794); J.C.Wendl.; 18 (1798); Willd.: 429

(1799); Poir.: 39 (1804): Lam. & DC.: 359 (1805); Wikstr.:

319 (1818); Thunb.: 374 (1825a); Meisn.: 390 (1840);

Steud.: 273 (1841); C.A.Mey.; 45 (1843); Meisn.: 561

(1857); Harv.; 325 (1868); Gand.: 418 (1913); C.H. Wright:

9 (1915); Thoday: 146 (1924a); Marloth: 214 (1925);

Domke: 137 (1934); Palmer & Pitman: 1583 (1972);

Coates Palgrave: 648 (1977); Bond & Goldblatt: 432

(1984); Hilliard & B.L.Burtt: 182 (1987); Goldblatt &

Manning: 683 (2000). Type species: Passerina fdiformis L.

Sananmnda [Clus.: 89 (1601); L.: 146 (1737)] Adans.:

258 (1763); Lam. & DC: 359 (1805); Raf: 104 (1836).

Type species: as above.

Tliymelaea [Toum.: 594 (1719); L.: 146 (1737)] Adans.:

258 (1763); Juss.: 77 (1789); Lam. & DC.: 359 (1805).

Type species: Daphne laureola L.

Passerine Lam. & DC.: 359 (1805) orth. var.

Balendasia Raf: 105(1836). Type species: B. ericoides

(Burm.f.) Raf

Steiroctis Raf: 105 (1836). Type species: not desig-

nated (Parr et al. 1979).

Trimeiandra Raf: 105 (1836). Type species: T. spicata

Raf. nom. illeg.

Lonchostoma obtusiflorum Wikstr. nom illeg. = Pas-

serina pentandra Thunb. (Parr et al. 1979) s Loncho-

stoma Wikstr.: 350 (1818) nom. cons.

Chymococca Meisn.: 565 (1857); Harv.: 325 (1868);

Benth. & Hook.: 194 (1880); Thoday: 166 (1924a). Type

species: C. empetroides Meisn.

Passerina L. Sectio Pentamerae Meisn.: 390 (1840).

Type species: not designated.

Passerina L. Sectio Tetramerae Meisn.: 395 (1840).

Type species: not designated.

= the identity sign denoting nomenclatural synonymy for

names based on the same type species.

TABLE 1. — A summary of taxa in the most comprehensive works on Passehna from Linnaeus ( 1753) to the present study

62

Bothalia 33,1 (2003)

OU Ly

d,

(2 Q, Q, 3 3

cy cL

g b

: N

! I

i J "So

I Li < M Cy

S ^ 5

E

a. E

(U d-f

.>1

2 2

> >

J J

cii cC

S "Bo

o 'ir

8^

0-i ^

s

a: “

Li 'S

'■B

3 ai Q-

'M

o ^

^ —

^ lS

CO.

I-

c2

E

E 8

o .

’hfi

*2 '-v

S

d ^

<U ^

d S ^

(U (U

^-8

3 H

3 ^

■a 3

1 ^

u 21

■a S

■= j j=

a 7; c

g I f|

6 2? S

o 5

"bi) 8

R “L a;

^ ^ ?N

8

'S ^ a m

Co ^

cC ^

II II

TABLE I . — A summary of taxu in the most comprehensive works on Passerimi from Linnaeus ( 1 753) to the present study (cont. )

Bothalia 33.1 (2003)

63

*The complete list of excluded species described by Meisner (1857: 563) was not included in Table 1 as most names are dealt with under ‘Excluded names’.

64

Bothalia 33,1 (2003)

Shiubs or small trees. Stems greyish brown; bark tough

and stringy. Leaves decussate, imbricate on young branch-

lets, sessile, closely appressed to stem or spreading at an

angle of 5-20°(-60°), cymbifomi (boat-shaped), falcate or

cigar-shaped; plane shape lineai; oblong, lanceolate or nar-

rowly trullate; base sessile or cuneate; apex tmncate and

hump-backed, obtuse, rounded, acuminate or acute to almost

spine-tipped; margins sometimes ciliate; length x depth

( 1.5-)2.5-4.0(-8.0) X (0.8-)1.2-2.0(-3.0) mm (leaf shape

usually cymbifomi, depth of lamina is distance from adaxial

groove to main vein situated abaxially); lamina inversely eri-

coid, adaxial surface concave, tomentose, abaxial surface

convex, glabrous, seldom tomentose. Inflorescences com-

prising polytelic synflorescences [apex of main florescence

(main axis) not temiinating with a flower (indetenninate),

co-florescences (lateral branches) of the same structure |;

main florescences as well as co-florescences spicate; spikes^

reduced, resembling terminal subcapitulate inflorescences,

each chaiacterized by two terminal leaves with axillary

blind-ending mdimentary flowers, enveloping minute grow-

ing point, proliferating growth (inflorescence apex grow-

ing out and returning to vegetative growth) less common

(Figures 1 , 3, 4), or spikes mostly extended, number of spikes

reduced or multiflowered, main and co-florescences present,

proliferating growth common (Figures 10, 12, 13, 15, 16, 20,

23). Bracts enveloping flowers and fruits, largest after anthe-

sis of flowers, becoming more coriaceous and rounded at

fmit set, decussate, imbricate, cymbiforTn or helmet-shaped;

plane shape oblong, lanceolate, ovate and obovate to widely

ovate and obovate, rhombic and narrowly obtrullate to

obtrullate; base sessile, cuneate; main vein strongly devel-

oped, often keeled, extending to form a leaf-like point in

many species; apex obtuse, rounded or acute; texture mostly

coriaceous; lamina with adaxial surface (inside) concave,

abaxial surface (outside) convex, outside usually glabrous,

inside base or midrib tomentose or completely tomentose,

coriaceous or charlaceous, rrtgose or smooth, ± succulent or

thin, sometimes ± 3-5-ribbed, reticulately veined or ribbed

and reticulately veined on each side of main vein; wings

absent or bullate, cor iaceous, charlaceous or membranous;

margins often ciliate to setose; size variable, bracts without

leaf-like point, length x depth (2.5-)3.5^.5(-5.5) x (0.9-)

1 .0-1 .5(-2.4) mm or bracts with leaf-like point, length x

depth (4.0-)5.1-6.3(-7.3) x ( 1 .4— )1 .5-2.0(-2.6) mm.

Flowers actinomor'phic, hypogynoirs. Floral envelope

constituting hypanthium and sepals, (4.0-)5.3-7.3(-8.4)

mm long, membranous during pollination and yellowish

in P. rigida, P. paleacea, P. nivicola and P. esterhiiyse-

niae, slightly succulent and gr'eenish in P. ericoides,

mostly yellow-pink in all other species, dehydrated after

shedding of pollen, becoming papyraceous or coria-

ceous, yellow-pink tones turning red. Pedicel ver'y shor1

or absent. Receptacle very shorl. Hypanthium a membra-

nous to coriaceous cylindric lube (fused calyx and

androecium, differentiating into sepals and diplostemo-

nous androecium ar ising from hypanthium rim at separa-

tion of sepals, Figur'e I ); indumentum var iable in densi-

ty, trichornes nonglandular, uniseriate, often spiralled,

whitish, density of indumentum at ovary ranging from

glabrous to tomentose or slrigose; neck (narrowed tube

between apex of ovar-y and sepals) (().3-)().6-2.6(-3.0)

nmi long, density of indumentum ranging IVom glabrous

to tomentose on outside, inside often hairy, abscission

tissue not macroscopically discer nable, arliculation plane

absent, after fruiting fragmentation of neck base caused

by dehydration and torsification of tissue, sepals and an-

droecium being shed in most species. Sepals 4, petaloid,

imbricate in bud, flexed in flower, often setose with up to

5 long trichornes on outer surface and glabrous to tomen-

tose on inner surface; outer sepals cymbifomi or con-

cave; inner sepals oblong, elliptic or obovate. Corolla

absent. Disc absent. Androecium dimorphic diplostemo-

nous, arising from hypanthium at separation point of

sepals; filaments of antipetalous whori (0.4— )0.7-l.2(-l.5)

mm long, those of antisepalous whorl ( l.2-)l.4— 2.2(-2.4)

mm long; anthers (0.5-)0.7-0.9(-l.l) x (0.2-)0.3-0.4(-0.7)

mm, sub-basifixed, 2-thecous and 4-locular, exserted,

extrorse. Ovary superior, ( 1 .6-)2.0-2.5 (-2.7) x (0.5-)0.6-l .4

(-1.7) mm, bicarpellate during embryonic stage (Bun-

niger 1972), pseudomonomerous (Heinig 1951 ) at matu-

rity, placentation parietal, uniloculate, with I pendulous

ovule laterally attached near top of ovary; style separat-

ing laterally from top of ovary, maintaining lateral posi-

tion in hypanthium neck, reaching beyond hypanthium

rim; stigma ± globose, mop-like or penicillate (wind pol-

lination). Fruit enveloped by persistent, loosely arranged

hypanthium fragmented at neck base or, in some species,

fragmenting over widest circumference of fruit, the frag-

mented hypanthium, sepals and androecium being shed;

in P. ericoides and P. rigida a fleshy, l-seeded berry, 5.3

X 4 mm; in all other species an achene, pericarp mem-

brafious and dry, 2.5 x 1 .2 mm. Seed broadly fusiform

with outgrowths at both micropylar and funicular ends,

2.2(-2.9) X 1 .2(-l.6) mm; legmen black and shiny, often

with white spots; endosperm formation nuclear, but later

becoming cellular throughout.

Diagnostic characters: plants of Passerina are

shrubs or small trees, distinguished by the inversely eri-

coid leaves, that are inverse-dorsiventral in c/s. The in-

florescences are few- to multiflowered, simple or com-

pound spikes, often reduced, artificially resembling ter-

minal subcapitulate inflorescences. Each flower is

enveloped by a conspicuous bract, becoming more cori-

aceous and rounded at fruit set. The flowers are adapted

to wind pollination. During pollination the flower colour

is yellow-pink, the four petaloid sepals are flexed and

the anthers are exserted and extrorse (unique for Thy-

melaeaceae in southern Africa). All anthers open explo-

sively and the pollen is shed at once. The stigma is mop-

like. The fruit is enveloped by a persistent, loosely

arranged hypanthium. Because of the absence of an

articulation plane, the hypanthium fragments at neck

base or, in some species, over the widest circumference

of the fruit, the fragmented hypanthium, sepals and

androecium being shed. P. ericoides and P. rigida are

characterized by a fleshy, l-seeded berry and all other

species by an achene.

Etymology: Passerina refers to the Latin passer (= a

sparrow) as the seeds resemble a sparrow’s beak.

Common names: the vernacular name 'sparrow-wort’

was suggested by Miller ( 1 768) for all Passerina species

and Wendland (1 798) used the name fadenfdrmige

Vogelkopf. According to Smith ( 1 966) gonna is a collec-

tive name once used by the Khoekhoe for various mem-

bers of Thymelaeaceae, e.g. several species of Passerina

and Strnthiola.

Bothalia 33,1 (2003)

65

Uses: many Passeriiui species grow on sand dunes

and in sandy areas, with parts of the woody stem subter-

raneous, forming runners and developing an extended

root system. Most of these plants are pioneers and

resprouters, increasing their chances of survival in dis-

turbed areas. These plants are excellent sand binders and

are suitable for binding problematic sandy areas, espe-

cially after the clearing of invader species. Sim (1919)

recommended Passerina in his list of trees and shrubs for

coastal areas exposed to sea winds. Certain Passerina

species such as P. falcifolia are small trees and can be

used as ornamental garden plants. P. filiformis has been

cultivated in Britain and Europe since the time of

Linnaeus. P. obtusifolia is used in the wild flower indus-

try in the Robertson area. The bark is exceedingly tough

and is used for tying down thatch. According to Watt &

Breyer-Brandwijk (1962) it is also plaited into whip

thongs and used as twine. Members of the genus are not

browsed by stock as the plants are apparently unpalatable

(Story 1952). Ash from Passerina obtusifolia was tradi-

tionally used by the people of Genadendal in Western

Cape in the home industry of soap-making. Although

certain species have been recorded in cancer research,

these plants are not cun'ently known for their medicinal

value.

Flowering and fruiting: most Passerina species

flower profusely in spring, from September to October.

During this season the Cape Floristic Region is quite

windy and large amounts of pollen are produced, as

Passerina is wind-pollinated. Pollen is often wafted

away in clouds, causing a kind of hay-fever in sensitive

persons (Marloth 1925). Fruiting time is mostly from

December to January. The fleshy fruits of P. ericoides

and P. rigida are dispersed by birds or rodents inhabiting

the distribution ranges of these species along the South

African coast. Fruits of P. truncata subsp. truncata,

growing in the Karoo, passively fall to the ground, where

they are probably dispersed by ants or rodents. The fruits

of P. montana, occurring along the Great Escarpment.

are probably dispersed by birds as they are arranged at

the tips of branchlets, exposed, red, and beak-like, possi-

bly resembling the beaks of nestlings.

Distribution and ecology: in Passerina the highest

number of species per grid (nine) occurs in each of the

grids 3321 (Ladismith), 3322 (Oudtshoorn) and 3419

(Caledon). The highest diversity of species (six) occurs

in the False Bay area, from Seekoeivlei, including the

Cape Flats, to De Mond at the Palmiet River (341 8B).

After an extensive study of herbarium material in co-

operation with field work. Western Cape is regarded as

the centre of diversity for Passerina, from where certain

species extend west, north and east.

Thoday ( 1925) published an account of the geograph-

ical distribution and ecology of Passerina, based on 15

species. Of the 20 species currently studied, 10 are

endemic and 4 species are near-endemic to the Cape

Floristic Region. P. obtusifolia is widespread in the

Northern, Eastern and Western Cape, whereas P. corym-

bosa occurs in Western and Eastern Cape, with outliers

in KwaZulu-Natal. P. rigida is distributed from Western

Cape, along the coast to northern KwaZulu-Natal; all

these species are endemic to the southern African

provinces in which they occur. P. drakensbergensis is

endemic to the Bergville District in KwaZulu-Natal.

P. inontivaga is found from Mossel Bay and Oudtshoorn

to Eastern Cape and along the escarpment northwards to

Zimbabwe and P. montana is distributed from the east-

ern mountains and Great Escaipment of southern Africa

to Zimbabwe and Mozambique. P. montivaga and

P. montana are near-endemic to the Great Escarpment.

Consen’ation status: with the exception of Passerina

esterhuyseniae (from the northern Cederberg Mountains)

known from herbarium material only, all other species

and subspecies of Passerina were studied in the wild.

Assessments were done using the guidelines of the lUCN

Species Survival Commission (2000) and Victor (2002).

Key to species

la Inflorescences comprising terminal subcapitulate spikes; proliferating growth (inflorescence apex growing out

and returning to vegetative growth) uncommon (Figures 1, 3. 4):

2a Floral envelope yellow and membranous, up to 4 mm long, neck (portion of hypanthium between ovary and

sepals) very short, ± 0.3 mm long; bracts widely ovate, wings membranous and obscurely veined

\. P. puleacea

2b Floral envelope yellow-pink and papyraceous, 4. 9-6. 4 mm long, neck 0.7- 1.4 mm long; bracts variously

shaped, wings present or absent;

3a Leaves narrowly oblong to oblong; base sessile, dilated; apex truncate, or truncate to rounded, keeled, often

appearing humped on the back (abaxially) 2. P. truncata

3b Leaves linear-lanceolate; base diamond-shaped to rounded; apex rounded to acute;

4a Abaxial surface of young leaves tomentose; bracts ovate to widely ovate; lamina comose on inside, sparsely

hairy to tomentose on outside, ± 3-ribbed on each side of main vein, coriaceous and rugose; length x depth

(4.5- )4. 9 X 1.5(-L8) mm (leaf shape usually cymbiform, depth of lamina is distance from adaxial

groove to main vein situated abaxially) 3. P. qiuidrifaria

4b Abaxial surface of young leaves glabrous; bracts ovate to obovate; lamina villous on inside, glabrous on

outside, obscurely ribbed on each side of main vein, thinly coriaceous; length x depth (3.2-)4.0 x

0.9(-L6) mm 4. P. montana

lb Inflorescences comprising extended spikes, number of spikes often reduced, or many compound, multiflowered

spikes present; proliferating growth common (Figures 10. 12, 13, 15, 16, 18, 20, 21, 23);

5a Inflorescences reduced, often to solitary spikes; dwarf shrubs up to 300 mm high; bracts rhombic, dark green

when fresh, dark brown in dried specimens, coriaceous, membranous wings absent 5. P. hiirchellii

5b Inflorescences with multiflowered main and co-florescences; low shrubs, shrubs or small trees; bracts various-

ly coloured, textured and winged;

6a Fruit fleshy; floral envelope greenish or yellow, coriaceous or membranous; leaves greyish green:

7a Fruit a red berry; floral envelope greenish and coriaceous, hypanthium strigose; leaves oblong, apex obtuse

to subacute; bracts larger, oblong to lanceolate, leaf-like 6. P. ericoides

66

Bothalia 33.1 (2003)

7b Fruit a bright yellow berry; floral envelope yellow and membranous, glabrous at ovary, tomentose at neck;

leaves narrowly lanceolate to ovate, apex acute with main vein visible as a blunt keel; bracts widely

ovate, apex acute 1 . P. rigidu

6b Fruit dry (an achene); floral envelope yellow, yellow-pink or red, membranous or papyraceous; leaves variously

coloured;

8a Floral envelope membranous, yellow or yellow-pink;

9a Floral envelope yellow or yellow-pink; bracts chartaceous, widely obovate, wings membranous, often

bullate, tinged red at margins 8. P. nivicola

9b Floral envelope yellow; bracts thinly chartaceous, smooth and helmet-shaped with membranous rims , .

9. P. esterhuyseniae

8b Floral envelope papyraceous, yellow-pink or red:

10a Young leaves and bracts abaxially sparsely hairy, becoming tomentose towards apex, older leaves rugose

or warty with bases of fallen hairs 10. P. comosa

10b Young leaves and bracts abaxially glabrous;

1 la Bracts shorter than 4.5 mm:

12a Bracts rhombic in outline, softly coriaceous, with membranous wings, margins brownish ciliate . . .

1 1 . P. pendula

12b Bracts oblate in outline, chartaceous, with bullate membranous wings, margins glabrous

12 P. galpinii

11b Bracts longer than 4.5 mm: ^

13a Floral envelope ± 5.9 mm long; outer and inner sepals concave and lanceolate; bracts larger than

leaves, bracts and leaves lanceolate and glaucous 13. P. drakensbergensis

13b Floral envelope 6.0-8.4 mm long; outer and inner sepals variously shaped; bracts not as above, variously

shaped and coloured;

14a Midrib of bract shortly extended into an acute apex, lamina rhombic to obtrullate, distinctly angled,

4- or 5-ribbed; leaves with a distinct midrib, laterally compressed, greyish green, drying greyish

brown 14. P. corymbose/

14b Midrib of bract extending beyond lamina into a leaf-like point, lamina variously shaped and

ribbed; leaves with a distinct midrib, or midrib less obvious, abaxially convex or laterally com-

pressed, variously coloured:

15a Bracts with leaf-like point, obtuse at apex, lamina closely 2-ribbed at margin; hypanthium fragments

at circumference of ovary 15. P. obtusifolia

15b Bracts with leaf-like point variously shaped, but not obtuse, lamina (faintly ribbed in P. dnikens-

bergensis) extending into a membranous margin; hypanthium fragments at neck base;

16a Adaxial (inner) surface of bracts basally to centrally setose or tomentose over entire length of

midrib; wings glabrous;

17a Bracts with midrib and leaf-like point stout and strongly developed, apex acute

1 6. P. paludosa

17b Bracts with midrib forming a straight or filiform, leaf-like point, or midrib shortly extended;

18a Bracts with leaf-like point straight or slightly incurved; wings of bracts ovate, margins hairy

in distal half, or obtrullate, narrowing abruptly into midrib 17. P. moiitivaga

18b Bracts with leaf-like point shortly extended or extended into a filiform, slightly falcate

point; wings of bracts ovate-acuminate, gradually narrowing to a point or widely obovate,

narrowing abruptly into midrib 18. P . f/Ufon/zis

16b Adaxial (inner) surface of bracts completely villous:

19a Bracts with midrib extended, leaf-like point falcate; wings of bracts ± 4-ribbed; hypanthium

neck ± 3 mm long, tomentose, often arcuate; spikes lax, often arcuate, mottled grey-green,

with up to 16 fertile, enlarged bracts 19. P.falcifolia

19b Bracts with midrib shortly extended into a short point, apex acute; wings of bracts ± 5-ribbed;

hypanthium neck ± 2 mm long, glabrous to sparsely pubescent; spikes robust, rigid and

extended, glaucous, with up to 30 fertile, enlarged bracts 20. P. rubra

1 . Passerina paleacea Wikstr. in Kunglinga Svenska

Vetenskapsakademiens Handlingar 39: 323 (1818); Meisn.;

400 (1840); Meisn.; 562 (1857); C.H.Wright; 12 (1915);

Thoday: 164 (1924a); Thoday: 388 (1924b). Type: Caput

bonae Spei, Herb. Wikstromii, Speurmtm s.n. (S!, lecto.,

here designated; UPS!).

Lx/cimaea paleacea Herb. Banks, ined,, fide Wikstr.; 324 (1818);

Meisn.: 562 ( 1857); C.H.Wright: 12 (1915); Thoday; 164 ( 1924a). nom.

inval. in synonymy.

Passerii/a glo/nerata sensu Thunb.: 374 (1825a) pro parte quoad

specim. Herb. Thimherg 9596D, 9579.

L. congUm/erala L. sensu Thunb.; 374 (1825a) pro parte quoad

spccim. Herb. Thiu/berg 9596D.

P. ericoides sensu Thunb.: 374 (1825a) pro parte quoad specim.

Herb. Thu/iberg 9596D, 9579, non L.; P. ericoides sensu Meisn.: 401

(1840) pro parte, non L.; Meisn.; 562 (1857) pro parte quoad spccim.

Dreges./i. (G!, K!, P!, S!).

Shrubs or shrublets 0. 1-1.5 m high. Stems branching

from base up to growing points, branchlets from previous

growth persistent, arcuate, indurate; younger branchlets

ascending, densely white-tomentose, villous closer to

growing points; shredded bark of older branchlets grey-

ish brown, remains of tomentum forming lengthwise

strips; leaf scars conspicuous; older stems fissured

lengthwise exposing greyish white sclerenchyma fibres;

internodes shorter than leaves. Leaves imbricate on

young branchlets, closely appressed to stem, diverging at

an angle of ± 0-5°, cymbiform, often expanding, becom-

ing thickly chartaceous and bract-like towards inflores-

cences; lamina inversely ericoid, adaxial surface con-

cave, tomentose, abaxial surface laterally compressed

and glabrous, plane shape linear to linear lanceolate,

length X depth 1 .5-2.5(-4.0) x 0.6-0.8(-1.2) mm; base

sessile, dilated; apex acute, median vein forming a dis-

tinct keel incurved at apex; margins involute. Inflores-

cences subcapitulate, ± ellipsoid. Bracts decussate, im-

bricate, sessile, appressed, widely ovate in outline, length

X depth (2.5-)2.7 x 1.2(-1.9) mm; lamina adaxially (in-

side) concave and villous, abaxially (outside) convex and

glabrous, thickly chartaceous, smooth on each side of

main vein, concolorous, greyish green, senescing to yellow-

Bothalia 33.1 (2003)

67

FIGURE 1. — Passerina paleacea,

Bredenkamp 960. A, spike

reduced, resembling terminal

•subcapitulate inflorescence;

B. leaf; C. bract; D, Bower

clasped by bract in ventral

view; E. liypanthium frag-

menting at circumference of

ovary. F-H, achene; envelop-

ed by membranous pericarp;

G. lateral view; H. ventral

view. h. hypanthium; s, sepa-

ls. Scale bars; 2 mm. Artist;

A. Stabler.

ish brown; base cuneate; main vein extending into obtuse

apex; wings membranous, borders glabrous, obscurely

veined. Floral envelope membranous and yellow during

pollination, dehydrated after shedding of pollen, turning

red to brownish, ± 4.2 mm long. Hypanthium glabrous,

neck ± 0.3 mm. long, abscission tissue and articulation

plane absent. Sepals concave, elliptic or subrotund and

glabrous. Androeciiun with filaments of antipetalous

whorl ± 0.7 mm and antisepalous whorl ± 1.4 mm long;

anthers 0.6 x 0.5 mm. Ovary 2.4 x 1 .4 mm. Fruit an ach-

ene with pericarp membranous and dry, ± 2.3 x 1.2 mm,

enveloped by persistent, loosely ananged hypanthium

fragmenting over widest circumference of fruit, the frag-

mented hypanthium, sepals and androedum being shed.

Figure 1.

Nomenclatural notes: in the latter half of the eighteenth

century Lachnaea conglomerata L. (1753), Passerina

ericoides L. (1753) and P. glomerata Thunb. (1794) were

constantly confused by botanists, causing Wikstrom

(1818; 322) to place P. glomerata and L. conglomerata

in the synonymy of P. conglomerata Thunb. In the same

publication Wikstrom delimited and described P. paleacea.

However, P. paleacea is not mentioned in Thunberg’s

revision of 1825, in which he described P. glomerata

occurring in ‘Hautbay’, the currently known locality of

both P. paleacea and P. ericoides. This confusion is re-

flected on many herbarium specimens, e.g. the specimen

Herb. Thunherg 9579, bearing the inscriptions P. eri-

coides, P. glomerata (struck out) and the word ‘paleacea’

written in pencil. Although Thoday (1924b) cho.se the

specimen Herb Thunberg 9597 as the type of P. paleacea,

this specimen was not chosen as lectotype in the present

study, as the Sparrman specimen cited by Wikstrom

(1818; 324) was located at S. The specimen LINN 504.3,

positively identified as P. paleacea, bears the inscription

‘sp 161’, possibly referring to Spairman. However, there

will always be doubt whether it is a duplicate of the

Sparrman specimen cited by Wikstrom.

Diagnostic characters and relationships: Passerina

paleacea may easily be confused with P. rigida, as both

occur on sand dunes along the coast. The branches of P.

rigida are nodding and abundantly covered by pendulous

branchlets, spikes are extended and the fruits are fleshy,

yellow berries. Plants of P. paleacea are less robust,

reaching a maximum height of 1.5 m, and are character-

ized by an abundance of subcapitulate inflorescences and

dry fruit. The subcapitulate inflorescences at times led to

the confusion of P. paleacea with P. truncata (= P.

glomerata), but, these two species are morphologically

as well as geographically distinct. P. paleacea has a mari-

time habit and P. truncata is distributed from Vanrhyns-

dorp, along the Cederberg Mountains, to Malmesbury,

Ceres, Tulbagh and Matjiesfontein up to Seven Weeks

Poort. The earlier confusion between P. paleacea and

P. ericoides was probably due to their sympatric occur-

rence, but these two species are moiphologically quite

different.

Etymology: the Latin specific epithet paleacea (= chaffy)

probably refers to the chaff-like subcapitulate inllores-

cences.

Distribution and ecology: Passerina paleacea occurs

in both the Southwestern and the Agulhas Plain Centres

of the Cape Floristic Region (CFR) (Goldblatt & Man-

ning 2000) and is a typical fynbos element. It grows on

coastal dunes and in maritime habitats from Lange-

baan, round the Cape Peninsula to the Cape Flats, Kogel

Bay, Hermanus, Gansbaai, De Hoop, the Potberg coast,

Bredasdorp, Arniston, Vermaaklikheid and Puntjie up

to Stilbaai (Figure 2). The vegetation types dune fyn-

bos and dune thicket form a mosaic along many parts

68

Bothalia 33,1 (2003)

FIGURE 2. — Known distribution of

Passerina pateacea, •; P. tnm-

cata subsp. tnmcata, ■.

of the southern Cape coast (Lubke 1998a, 1998b;

Lubke & Van Wijk 1998). This same distribution pat-

tern is displayed by P. paleacea as it is found in the

dune scrub, amongst typical fynbos species, but not in

the dune thicket amongst larger shrubs or small trees

with mesophytic leaves such as species of Chry-

sanlhemoides, Mimusops, Morelia, Rhus and Sideroxy-'

Ion.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

2. Passerina truncata (Meisn. ) Bredenk. & A.E.van

Wyk in Bothalia 32: 66 (2002a). Type: Western Cape, near

Tulbagh Waterfall, April 1865, Zeyher 43 (K!, lecto.;

MEL!, MO!, NBG!, S!, W!).

The complete description of the species and sub-

species, the synonymy, relationships, etymology, distrib-

ution, habitat and key to subspecies is dealt with in

Bothalia 32: 66-71 (2002).

2a. subsp. truncata (Figures 2, 3)

Conseri’ation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

2b. subsp. monticola Bredenk. & A.E.van Wyk

Conserx’ation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

3. Passerina quadrifaria Bredenk. 8i A.E.van Wyk

in South African Journal of Botany 68: 304 (2002b).

Type: Eastern Cape, 3324 (Steytlerville): Uitenhage District,

Great Winterhoek Mountains, Cockscomb, (-BD), rocky

ridge, 30 Nov. 1958, Esterhuysen 28006 (PRE, holo.!;

BOL!, K!).

Passerina sp. nov. 3 Bredenk. & A.E.van Wyk: 70

(2000); Bredenk. & A.E.van Wyk: 56 (2001a); Bredenk.

& A.E.van Wyk: 217 (2001b).

The complete description of the species, relationships,

etymology, distribution and habitat appear in the South

African Journal of Botany 68: 304-307 (2002).

Conserx’ation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

4. Passerina montana Thoday in Bulletin of Mis-

cellaneous Information, Kew 4: 152 (1924a); Norl. &

Weim.: 630 (1958); Bond & Goldblatt: 432 (1984);

Hilliard & B.L.Burtt: 182 (1987). Type: Mooi River, Wood

FIGURE 3. — Passerina truncata

subsp. truncata. Bredcnkamp

985. A, spike reduced, re-

sembling terminal subcapilu-

late inllorescence; B, bract;

C, leaf; D, flower clasped by

bract, fragmenting at circum-

ference of ovary. E, F, ache-

ne: E, clasped by bract; F,

enveloped by membranous

pericarp. Scale bars; 2 mm.

Artist: A. Stadler.

Bothalia 33,1 (2003)

69

4036 [K!, lecto., designated by Thoday 10: 387 (1924b);

BOL!, GRA!, PRE!].

P. ericoides sensu Meisn.: 401 (1840) pro parte, non L.; Meisn.:

562 (1857) pro parte; C.H. Wright: 12 (1915), pro parte.

P. rigida Wikstr. var. tetragona Meisn.: 563 (1857) pro parte quoad

specim., circa Stormberg Drege s.n.

Small trees or shrubs, (0.2-)1.0-2.0(-2.5) m high. Stems

branching from base up to growing points, young stems

profusely branched, young branchlets ascending; branch-

lets terminally leafless and woolly, with conspicuous ter-

minal scars after dispersal of fruit; bark greyish brown,

younger branchlets densely white-tomentose, villous

closer to growing points; bark on older branchlets shred-

ding, greyish brown, remains of tomentum forming length-

wise strips; leaf scars conspicuous; older stems fissured

lengthwise exposing greyish white sclerenchyma fibres;

intemodes mostly shorter than leaves. Leaves imbricate

on young branchlets, closely appressed to stem, diverg-

ing at an angle of ± 0-5°, cymbiform, lamina inversely

ericoid, adaxial surface concave, tomentose, abaxial sur-

face laterally compressed and glabrous, plane shape lin-

ear to lanceolate, length x depth 1.5-2.5(-4.0) x 0. 6-0.8

mm; base sessile, dilated; apex acute, median vein

prominent in upper third of leaf, incurved at apex; mar-

gins involute. Inflorescences subcapitulate, ± ellipsoid.

Bracts decussate, imbricate, sessile, appressed, ovate to

obovate in outline, length x depth (3.2-)4.0 x 0.9(-1.6

mm); lamina adaxially (inside) concave and villous,

abaxially (outside) convex and glabrous, thinly coria-

ceous, obscurely ribbed, yellowish green, margins of

fruiting bracts turning red; base sessile; main vein

extending into subacute apex; wings membranous,

brownish. Floral envelope papyraceous and yellow-pink

during pollination, dehydrated after shedding of pollen,

turning red to brownish, ± 5.8 mm long. Hypanthinm

glabrous at ovary, neck tomentose, ±0.8 mm long, frag-

mentation at neck base. Sepals: outer sepals cymbiform,

adaxially scantilly tomentose, abaxially glabrous; inner

sepals obovate, adaxially tomentose, abaxially glabrous.

Androeciurn with filaments of antipetalous whorl

± 0.6 mm and those of antisepalous whorl ± 1.5 mm

long; anthers ± 0.8 x 0.4 mm. Ovary ± 2.1 x 0.6 mm.

Fruit an achene enveloped in beak-like, reddish, papyra-

ceous hypanthium, fragmented at neck base; pericarp

membranous and dry, ± 2.3 x 1.2 mm. Figure 4.

Nomenclatural notes: according to the concept of

Wright (1915), P. ericoides is not only distributed along

the southern coast of Western Cape (present interpreta-

tion), but also along the coast to Eastern Cape and further

inland up to Mpumalanga. However, most of the inland

specimens cited by him have been classified as P. mon-

tana by Thoday (1924a). The interpretation of P. corym-

hosa by Wright (1915) posed the same problem, as Wood

4036 (the lectotype of P. montana) was also placed in

this taxon.

Meisner (1857) described P. rigida var. tetragona cit-

ing two Drege specimens, one from Ezelsbank and the

other from Stormberg. The Ezelsbank specimen {Drege

2971, P, K) is P. trimcata, but the Stormberg specimen

could not be located. According to Gunn & Codd (1981),

Drege crossed the Stormberg (3126BC, Queenstown) on

17 December 1832. The first author suspected that the

Drege specimen would be P. montana, as it is common

in this area. This suspicion is supported by Sim 68 (from

the Pirie Mountains in the King William’s Town District),

a syntype of P. montana, bearing the inscription ‘P. rigida

WW^-tetragona' and the Drege specimen from Storm-

berg is consequently regarded as P. montana.

Diagnostic characters and relationships: Hilliard &

Burtt (1988) noted two rather distinct forms of P. mon-

tana in KwaZulu-Natal. The first form is characterized

by plants on rock platforms that are low, rounded bush-

es, 0.3-1 m high, with the tips of the branches erect,

whereas those of the second form inhabit valleys and are

riverside bushes of up to 2 m high, with open branches

and pendulous branchlets. The present study, taking the

whole distribution range of P. montana into considera-

tion, recognizes two forms. One, centred in the God’s

Window area of Mpumalanga, are rounded shrubs 0.5-2

m high, with many branchlets covered with smaller, de-

cussate, imbricate leaves, bluish green in colour. The

second form dominates in the Free State, Lesotho,

KwaZulu-Natal and Eastern Cape. These plants are more

robust, with open branches and larger, yellowish green

leaves and inflorescences, which are tinged pink.

However, the two forms are not geographically distinct

and intermediates are common. Both forms unequivocal-

ly show the specific characters and therefore we do not

propose to give them formal taxonomic recognition. Studies

FIGURE 4. — Passerina montana,

Bredenkump 893. A, spike

reduced, resembling terminal

subcapitulate inflorescence;

B. leaf; C, bract; D, flower

clasped by bract. E, F, ach-

ene: E, enveloped by mem-

branous pericarp; E, lateral

view. Scale bars: 2 mm.

Artist: A. Stadler.

70

Bothalia 33,1 (2003)

of the leaf epidermis, anatomy and floral morphology

(Bredenkamp & Van Wyk 2000, 2001a, 2001b) supplied

no further evidence on which the two forms could be

delineated.

Etymology, the specific epithet is derived from the

Latin montanus (= pertaining to or growing on moun-

tains). This is a very appropriate epithet as P. montana is

distributed along the Great Escarpment from Eastern

Cape to Zimbabwe.

Common names: Cooper 2302 (K), from Lesotho, re-

ported the vernacular name Likhabei and Staples 17

(PRE), from the Maluti Mountains in Lesotho, recorded

the name Lekaphu. Story (1952) mentions the name

pakaan. Von Breitenbach et al. t2001) used the names

berg-gonna and mountain gonna.

Uses: information on the specimen Watt & Breyer-

Brandwijk 1851, collected at Thabaneng, states that the

plants are used medicinally. However, Watt & Breyer-

Brandwijk ( 1962) supplied no further details.

Distribution and ecology: Passerina montana is a

near-endemic to the Great Escarpment of southern Africa,

with distant satellite populations in high mountain areas

of Angola, Namibia and Limpopo [Northern Province],

South Africa. It is distributed from Nyanga in Zimbabwe,

along the escarpment to Manica and Sofala in Mozam-

bique, Limpopo, Mpumalanga, Swaziland, KwaZulu-

Natal, Free State, Lesotho and the Eastern Cape (Figure

5). Outliers in Angola have been found on the escarp-

ment of the Huilla Plateau near Lubango and the Cheila

Mountains. Several specimens of this species have been

collected at Moltkeblick on the Auas Mountains in Nami-

bia. In Limpopo, P. montana is found in the Soutpansberg

area and on the Blouberg, as well as on the summit of

Krantzberg in the Waterberg Mountains. A single speci-

men (Goossens 375) was collected in the Pretoria

District, but the species is currently probably extinct in

this area, due to human impact.

FIGURE 5. — Known distribution of Passerina nunuana.

This species grows at altitudes of (900-)l 200-3 000 m.

At Nyanga, P. montana is associated with Erica niannii

and E. hexandra, bordering on Bracliystegia woodland and

montane forest. In Mozambique and South Africa it has

been found with Widdringtonia nodiflora and Erica

species, bordering on montane forest. It is common

amongst rocks on hills, mountain slopes, mountain tops,

cliff ledges and rocky ridges. It also frequents stream

courses and banks as well as riverbeds and banks, where

the growth form has been reported as a shrub amongst

rocks, a drooping bush over running water, a limply

spreading bush in sand or dense bushes. These plants also

grow in river valley forests and along plantations.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

5. Passerina burchellii Thoday in Bulletin of Mis-

cellaneous Information, Kew 4: 155 (1924a); Bond &

Goldblatt: 432 (1984); Goldblatt & Manning: 683 (2000).

Type; Cape, Caledon Div., mountain tops of Baviaans-

kloof near Genadendal, Biirchell 7761 [K, lecto.!, desig-

nated by Thoday: 387 (1924b); M!, P!].

P. tetragona Burch. MS. in Herb. Kew, non Steud. fide Thoday: 156

(1924a).

Low, erect, many-stemmed shrublets, branching mostly

on new growth, ± 0.3 m high, from a common rootstock.

Stems greyish brown, cork fissured, grey-brown, scabrous,

surrounding prominent leaf scars; indumentum at growing

point densely white-tomentose, flaking off with cork on

older branchlets, which become glabrous. Leaves imbri-

cate, overlapping ± 50%, diverging at an angle of 30°,

plane shape rhombic, length x depth 2.8(-3.5) x 1.5 mm,

adaxial surface concave, villous, abaxial surface convex,

glabrous; base sessile, cuneate; apex subacute, bearded;

margins brownish setose. Inflorescences with spikes

extended, number of spikes often reduced, spikes some-

times solitary, 6-12-flowered, arrangement subterminal,

axis white-tomentose, proliferating growth common.

Bracts appressed, rhombic, length x depth (3.2-)3.5 x 1.5

mm; lamina adaxially concave (inside), abaxially convex

(outside), villous inside, glabrous outside, coriaceous and

smooth, extending into a smooth wing, dark green when

fresh, dark brown in dried specimens; base cuneate; main

vein extending into acute, bearded apex; margins brown-

ish setose, involute. Eloral envelope ± 4.7 mm long,

papyraceous and yellow-pink during pollination, dehy-

drated after shedding of pollen, turning red to brown.

Hypanthium glabrous at ovary, neck tomentose, ± 0.8 mm

long. Sepals: outer sepals cymbiform, midrib adaxially

and apex abaxially setose; inner sepals obovate, adaxially

tomentose, apex abaxially setose. Androecium with fila-

ments of antipetalous whorl ± 0.7 mm and those of anti-

sepalous whorl ± 1 .5 mm long; anthers 0.5 x 0.3 mm, sub-

basifixed, 2-thecous and 4-locular. Ovary ± 1.6 x 0.6 mm.

Emit an achene, pericarp membranous and dry, ±2.5x1 .2

mm, enveloped by persistent, loosely arranged hypanthi-

um, breaking up at neck base due to dehydration and tor-

sification of tissue, resulting in sepals and androecium

being shed. Figure 6.

Nomenclatural notes: although Thoday (1924a) cited

P. rigida var. comosa Meisn. partly (ex MS. in Herb.

Kew) in synonymy under P. biirchellii, the specimens

cited by Meisner (1857) in the description of the var.

comosa all belong to P. pendida (Eckl. & Zeyh.) Thoday.

Diagnostic characters and relationships: bearded

sepals, leaves and bracts distinguish this species from P.

pendida.

Eponymy: this plant was named in honour of the

explorer and botanist W.J. Burchell, who collected in

Caledon and as far north as Tulbagh between 1810 and

1811. During this trip Burchell 7761, the lectotype of P.

biirchellii, was collected on the summit of the mountains

of Baviaanskloof near Genadendal.

Distribution and ecology: Passerina burchellii is en-

demic to the Southwestern and Langeberg Centres with-

in the CFR. It is common on mountain summits of the

Villiersdorp and Genadendal Districts (Figure 7), with

outliers on southeastern rocky slopes of Towerkop in the

Swartberg Mountains at Ladismith. This species occurs

at altitudes of 1 333-2 167 m, often covered in mist. It is

found in small groups on sandy loam, between boulders

and rocks on upper south- or southeast-facing slopes.

s: z mm.

Artist: A. Stadler.

Consen’ation status: Vulnerable ( VU D2) (Victor 2002)

because of small population size.

6. Passerina ericoides L., Systema naturae 12,2: 733

(1767); Bui-m.f.: 12 (1768); L.: 236 ( 1771 ); L.: 374 (1784);

Willd.: 430 (1799); Poir.: 41 (1804); Wikstr.: 325 (1818);

Meisn.: 401 (1840) pro parte minore; Steud.: 274 (1841);

C.A.Mey.: 49 (1843); Meisn.: 562 (1857) pro parte minore;

C.H.Wright: 12 (1915) pro parte; Thoday: 166 (1924a);

Bond & Goldblatt; 432 (1984); Hilliard & B.L.Burtt: 182

(1987); Goldblatt & Manning: 683 (2000). Type:

Passerina ericoides, Linnean Herbarium 504.5 [LINN,

lecto.!, designated by Thoday: 148 (1924a)].

Chvmococca empetroides Meisn.: 565 (1857); Harv.: 325 (1868);

Bolus &Wolley-Dod: 315 ( 1904); C.H.Wright: 15 ( 1915); Thoday: 166

( 1924a). P. fiUformis L. var. crassifolia Eckl. & Zeyh. fide Meisn.: 565

( 1857). Type: Eckl. & Ze\h. herb. no. 39 (G!, lecto., here designated;

BOL!, MO!. P!. W!).

P. glomerata sensu Meisn.: 562 (1857), non Thunb.: 75 (1794).

Lachnaea congloinerata L. sensu Meisn.: 562 ( 1857).

Low, rounded, many-stemmed shrublets, branching

profusely on new growth, 0.3-1. 2 m high, older branch-

FIGURE 7. — Known distribution of

Passerina burchellii. •; Passe-

rina ericoides, ■.

72

Bothalia33,l (2003)

lets indurate, young branchlets lax, often arcuate. Stems

light greyish brown, cork fissured lengthwise, grey-

brown, whitish scabrous, surrounding prominent leaf

scars; indumentum at growing point densely white

strigose, flaking off with cork on older branchlets, which

become glabrous. Leaves slightly succulent, imbricate,

overlapping ± 50%, diverging at an angle of 45°, plane

shape oblong, length x depth 2. 5-2. 8 x 0.6-0. 7 mm,

adaxial surface concave, villous, abaxial surface convex,

glabrous, greyish green, smooth; base sessile, dilated;

apex obtuse to subacute; margins glabrous, basally

sparsely setose. Inflorescences with spikes usually

extended, 6-12-flowered, arrangement subterminal, axis

white strigose, proliferating growth common. Bracts

spreading at an angle of 60° (flowering) or 90° (fruiting),

oblong to lanceolate, length x depth ± 3.6 x 1.5 mm;

lamina adaxially concave (inside), abaxially convex

(outside), villous inside, glabrous outside, smooth, wings

absent, greyish green, slightly succulent; base dilated;

apex obtuse to subacute; margins glabrous, basally

sparsely setose, involute. Floral envelope ± 5 mm long,

coriaceous and greenish during pollination, dehydrated

after shedding of pollen, turning red. Hypanthiiim ovate-

oblong at ovary, strigose, neck strigose, ± 0.6 mm long.

Sepals globose when young, concave, widely obovate,

outer and inner sepals adaxially puberulent, abaxially

glabrous. Androeciiim with filaments of antipetalous

whorl ± 0.5 mm and those of antisepalous whorl ± 1.3

mm long; anthers large, + 0.9 x 0.7 mm, subbasifixed, 2-

thecous and 4-locular. Ovary ±2.1 x 1.7 mm. Fruit a

fleshy red beiry, ± 5.3 x 4 mm, enveloped by persistent,

loosely arranged hypanthium, fragmenting over widest

circumference of fruit, the fragmented hypanthium,

sepals and androecium being shed. Seed ± 2.9 x 1.6 mm.

Noinenciatnral notes: in the Catalogue of the Lin-

naean Herbarium, Savage (1945) made the following

inscription ‘Tulb. list c. 1769. n.l. det. L. — Blaeria eri-

coides’. This refers to consignments of bulbs, seeds and

herbarium specimens that Rijk Tulbagh sent to Van

Royen, the Burmans at Amsterdam and Linnaeus at

Uppsala (Gunn & Codd 1981). Jackson (1917, 1918)

published a list of 203 of the specimens sent to Linnaeus

around 1769 and identified by him. The first inscription

on the list is the provisional name Blaeria ericoides,

which Savage (1945) believed to be the P. ericoides

specimen at LINN, but there is no numbering or any

other indication on the specimen to link it with Tulbagh’s

list (Jackson 1917-1918). As Linnaeus had already

described P. ericoides in 1767, the specimen at LINN is

probably not part of the Tulbagh collection. Thoday

(1924a) clearly regarded the specimen at LINN, named

by Linnaeus, as the type of P. ericoides. As no other origi-

nal elements exist, P. ericoides LINN 504.5 is regarded

as a lectotype designated by Thoday ( 1924a).

Thunberg ( 1825a) accepted Wikstrom’s concept of P.

glomerata, occurring at Hout Bay in the Cape, and cited

P. ericoides in synonymy, causing confusion about the

identity of the latter taxon. Meisner ( 1840) reinstated P.

ericoides, but the concept of this taxon became even

more doubtful in the light of the cited distribution. In

1 857 Meisner retained his concept of P. ericoides, occur-

ring at Uitenhage, Port Elizabeth, Witbergen and

Onderbokkeveld, and placed P. glomerata and Laclmaea

conglomerata in synonymy under P. ericoides. This revi-

sion by Meisner (1857) was largely followed by Wright

(1915). Because of his incorrect concept of the taxon,

Meisner ( 1 857) was confronted with material from Table

Bay and Standvallei with red berries, which he then

named Chymococca empetroides, based especially on

the fleshy fruit. Thoday (1924a) was justified in placing

this name in synonymy under P. ericoides, as the

descriptions of these taxa coincide and as the fleshy fruit

of C. empetroides is not unique, but is also found in

P. rigida. The concept of P. ericoides, occuning along

coastal dunes mainly in the Cape Peninsula and adjacent

coastal areas of the Western Cape, was clarified by

Thoday ( 1 924a) and is also accepted in the present study.

Diagnostic characters: Passerina ericoides is charac-

terized by greenish flowers, with a coriaceous, strigose

hypanthium and the fruits are fleshy red beiTies. The

leaves are greyish green and oblong, with an obtuse

apex. The bracts are leaf-like, larger and lanceolate.

Etymology: the specific epithet ericoides refers to the

ericoid appearance of this species indicated by the phrase

‘corollae tubus globosus, inflatus — unde et Ericam refert

flore’, which was used by Linnaeus (1767) in his origi-

nal description of the species.

Common names: Willdenow (1799) introduced the

vernacular name heideartiger Vogelkopf, and the com-

mon names ‘Christmas berry’ or dronkhessie were docu-

mented by Smith (1966).

Uses: Marloth (1925) remarked that P. ericoides was

laden with bright, scarlet fruits and that it was often

employed as a Christmas decoration. The juicy pulp has

a somewhat unpleasant taste, but appears to be harmless

(dronkhessie). As early as 1919, Sim recommended P.

ericoides as a useful shrub for planting in coastal areas

exposed to sea winds. This species occurs on coastal

dunes and on the banks of lagoons in the Cape Peninsula

and adjacent coastal areas of Western Cape. The plants

are excellent sand binders as they have an extensive root

system from which resprouting often takes place.

Because human impact and invasion of alien vegetation

along the coast of the Cape Peninsula are very high, reha-

bilitation and conservation of coastal dunes is of vital

importance. P. ericoides plants are ideally suited to com-

bat erosion of coastal dunes and can be used as a substi-

tute in coastal areas where alien vegetation is cleared. In

their research on the coastal erosion of the Milnerton

beaches, Biggs et al. (2001) made use of P. ericoides,

occurring on the mobile dunes of this area as a natural

monitor to indicate coastal erosion.

Distribution and ecology: Passerina ericoides ranges

from Melkbosstrand along the coast of the Cape Penin-

sula to De Mond in the Bredasdorp District (Eigure 7). It

is endemic to the Southwestern and Agulhas Plain

Centres within the CER. This species occurs on littoral

sand between rocks, or in dune valleys between the pri-

mary and secondary dunes.

According to a note on the specimen Taylor 4042, P.

ericoides forms part of the Coastal Eynbos (Acocks

1988), currently divided by Rebelo (1998) into the

Bothalia33,l (2003)

73

Laterite Fynbos of the Elim Flats, the Limestone Fynbos

on calcareous sands overlying the limestone and associ-

ated calcretes of the Bredasdoip Formation, and the Sand

Plain Fynbos from the Olifants River Mouth to Muizen-

berg on the West Coast lowlands. According to Rebelo

{ 1998) the southernmost centre of the Sand Plain Fynbos

is almost engulfed by the Cape Town Metropolitan Area

and the area between Milnerton and Malmesbury must

rank as one of the world’s hottest spots for the loss of

plant biodiversity.

Consen’ation status: Lower Risk (LR-nt) (Victor

2002), as a large portion of the coast along the Cape

Peninsula is affected by human impact and invasion by

alien vegetation associated with the Cape Town Metro-

politan Area.

7. Passerina rigida Wikstr. in Kunglinga Svenska

Vetenskapsakademiens Handlingar 39; 326 (1818);

Meisn.: 402 (1840); Steud.; 274 (1841); Drege; 208

(1843); C.A.Mey.; 49 (1843); Meisn.: 563 (1857);

C.H. Wright; 13 (1915); Thoday; 165 (1924a); Palmer &

Pitman; 1585 ( 1972); Coates Palgrave; 649 (1977); Bond

& Goldblatt; 433 (1984); A.E.van Wyk & P.van Wyk; 68

(1997). Type; Hab. ad Promont bonae spei [Cape

Peninsula], Prof. Sparrman s.n.. Herb. TInmh. 9578

(UPS, holo.!; M!, S!).

P. eriophora Gand.: 418 ( 1913); Thoday: 165 ( 1924a). Types: Natal

[KwaZulu-Natal], Stanger; Natal, in ora prope Durban [coast near

Durban], Wood 1712 (K!, lecto., designated by Thoday: 165 (1924a),

BM!,BOL!, M!,W!); Wood 6592 (S\).

P. ericoides sensu Meisn.: 562 ( 1857) pro parte quoad specim. Drege

s.n. (MO!), non L.

Many-Stemmed, much-branched, robust shrubs of

(0,6-)l,0-2,0(-3,04) m tall on coastal dunes; secondary

and tertiary branches ascending, conical in appearance,

formation of branchlets profuse, decussate, older branch-

lets self-pruning, lax or arcuate, 60-100 mm long, pro-

gressively shortening towards growing point, young

branchlets ascending, 5-60 mm long, growing point nod-

ding, fertile branchlets often pendulous and secund.

Stems greyish brown, bark stringy; cork grey-brown;

branchlets and growing points densely white-tomentose.

tomentum forming lengthwise patterns with cork on

older branchlets, which later become glabrous. Leaves

imbricate, overlapping ± 50%, appressed, plane shape

lanceolate to ovate, length x depth 1,6-2, 5 x 0, 1-1,1

mm, adaxial surface concave, villous, abaxial surface

convex, glabrous, greyish green, smooth, often covered

by salt crystals; base sessile, dilated; median vein in dis-

tal half visible as a keel, forming acute apex; margins vil-

lous, Inflorescences with spikes usually extended, 6-10-

flowered, arrangement subterminal, axis white-tomen-

tose, proliferating growth common. Bracts appressed,

ascending in fruit, widely ovate, length x depth (2,6-)3,4

X 1,4(-1,9) mm; lamina adaxially concave (inside),

abaxially convex (outside), villous inside, glabrous out-

side, smooth, with 2 or 3 shallow folds on each side of

main vein, wings absent, greyish green, coriaceous; base

dilated; apex with distinct, short, acute point; margins

tomentose, involute. Floral envelope ± 4 mm long, mem-

branous and yellow during pollination, dehydrated after

shedding of pollen, turning red, Hypanthium glabrous at

ovary, neck tomentose, ± 0,8 mm long. Sepals: outer

sepals cymbiform, midrib adaxially tomentose, abaxial

surface glabrous, inner sepals obovate, adaxially tomen-

tose, abaxially glabrous, Androeciiim with filaments of

antipetalous whorl ± 0,5 mm and those of antisepalous

whorl ± 1 ,5 mm long; anthers, ± 0,8 x 0.4 mm, sub-basi-

fixed, 2-thecous and 4-locular. Ovary 2.2 x 1.4 mm.

Fruit a fleshy yellow ben'y, ± 2.6 x 2.3 mm, enveloped

by persistent, loosely arranged hypanthium, fragmenting

over widest circumference of fruit, the fragmented

hypanthium, sepals and androecium being shed. Seed ±

1.4 X 1.1 mm. Figure 8.

Nomenclatural notes: in his description of P. rigida,

Wikstrom (1818) clearly indicated the specimen of

Sparrman, in the Thunberg Herbarium, as the type. This

specimen bears the inscriptions Passerina glomerata (3

and epithet rigida in pencil. Thoday (1924a) identified

the handwriting of the pencilled ‘rigida’ as Wikstrdm’s,

comparing it to signed letters in the library at Kew.

Wikstrom’s handwriting was also confirmed in the pre-

sent study, using examples published by Burdet (1979).

According to Stafleu & Cowan (1986), original speci-

mens of the Thunberg Herbarium (to which Sparrman

also contributed) were donated to UPS and the duplicates

FIGURE 8. — Passerina rigida. Bre-

denkump 1013. A, flowering

intlorescence; B, fruiting in-

florescence; C, leaf; D, bract;

E, Bower clasped by bract; F,

hypanthium fragmenting at

circumference of ovary. G,

H, fruit: G, clasped in tomen-

tum of bract; H. yellow,

Beshy beny. I, seed. Scale

bars: 4 mm. Artist: A. Stabler.

74

Bothalia 33,1 (2003)

were sent to S. Thus three other Sparnnan s.n. specimens

of Herb. Swartzii, Herb. Wikstromii and Herb. Gastromii,

housed at S, and a fourth one from Schreber’s herbarium,

housed at M, are duplicates. As Wikstrom clearly indi-

cated the specimen in the Thunberg Herbarium as the

type, we regard it as the holotype and the other four

SpaiTman specimens as isotypes.

Diagnostic characters and relationships: Passerina rigi-

da is easily distinguished as a robust, rigid shrub, usually

1-2 m high. The ascending branches are conical in shape

due to many branchlets that are pendulous when fertile.

The flowers are yellow and membranous and bright yel-

low berries are borne subterminally. The leaves are nar-

rowly lanceolate to ovate and the apex is acute, with the

main vein visible as a blunt keel. -The bracts are widely

ovate with the apex acute. This species is easily distin-

guished from P. paleacea which occurs on secondary

dunes and is distributed mainly along the southern coast

of Western Cape. Plants of the latter species are less

robust, reaching a maximun height of 1.5 m, character-

ized by an abundance of subcapitulate inflorescences and

the fruits are dry (achenes).

Etymology: the epithet rigida refers to the rigid, ascend-

ing branches, characteristic of the growth form of this

plant.

Common names: Smith (1966) recorded the vernacu-

lar names gonnabas and seekoppiesgonna, while both

Palmer & Pitman (1972) and Coates Palgrave (1977)

added the name ‘dune gonna’. Palmer & Pitman (1972)

also listed the names ishoba and imyenyevit. The names

‘dune-string’, duinetaaibos and gonnabos were used by

Lubke & Van Wijk (1998). Diiin-gonna, ‘dune gonna’,

inwele, imyenyevii are names given by Von Breitenbach

et cd. (2001).

Uses: Passerina rigida is a pioneer of the coastal

dunes along large portions of the South African coast.

Because these robust plants are excellent sand binders

and are completely adapted to maritime winds and salt

spray, they can be used in the rehabilitation of coastal

dunes in disturbed areas. P. rigida has an extensive root

system from which resprouting commonly takes place.

The yellow bemes are an important food source for ani-

mals inhabiting coastal areas, especially birds.

Distribution and ecology: Passerina rigida is distrib-

uted from Witsand River Mouth on the western coast of

the Cape Peninsula, along the coastline of South Africa

to Lake Sibayi on the northeastern coast of KwaZulu-

Natal (Figure 9). It is endemic to the coastlines of

KwaZulu-Natal, Eastern Cape and Western Cape. The

specimen Taylor 4143, recorded as far North as

Lambert’s Bay on the West Coast, is regarded as an out-

lier, as no other specimens have been recorded in the grid

3318. Thoday (1924a) mentioned Bowker s.n. from

Somerset, Cooper 2301 from Albany and Ecklon &

Zeyher s.n. (SAM 19801) as specimens from inland local-

ities. In recent years more cases of P. rigida growing

along sandy banks of rivers adjacent to the coast have

been noted.

This species occurs on littoral sand dunes and ham-

mock dunes just above the level of spring tide. It is also

found in marshy places and on sandy banks of river

mouths and lagoons. A stunted form is present on shal-

low marine sand over limestone and on rocky hills fac-

ing the sea. Lubke & Van Wijk (1998) regard P. rigida

on the southern and Eastern Cape coast as a pioneer

found in bush clumps or bush pockets on rear dunes.

According to them, there are often no pioneer communi-

ties on the vast dune sands and the first vegetation

encountered as one moves away from the shore is dune

thicket, in which P. rigida is one of the dominant shrubs.

Passerina species occurring on littoral dunes in Western

Cape are found mainly in Coastal Fynbos (Acocks

1988). From the southern Cape coast to Port Alfred, dune

fynbos and dune thicket form a mosaic as well as a suc-

cessional series between the two vegetation types (Lubke

& Van Wijk 1998).

Conser\>ation status: Least Concern (LC) (lUCN

Species Survival Commission 2000).

8. Passerina nivicola Bredenk. & A.E.van Wyk in

Bothalia 32: 77 (2002c). Type: Western Cape, 3319

(Worcester): Ceres Dist., Waboomberg, 1 760 m, (-DD),

FIGURE 9. — Known distribution of

Passerina rigida, •; P. nivi-

cola, ■ .

Bothalia 33.1 (2003)

75

FIGURE 10. — Passerina esterhuy-

seniae, Esterhuysen 26859.

A, inflorescence with apex

growing out, returning to

vegetative growth (prolifer-

ating growth); B, leaf; C,

bract; D. flower clasped by

bract showing fragmentation

of hypanthium at circumfer-

ence of fruit; E, fruit with

remnants of hypanthium. F,

G, achene: F, enveloped by

membranous pericarp; G, lat-

eral view. Scale bars: 2 mm.

Artist: A. Stadler.

12 November 1989, E.G.H. Oliver 9281 (PRE!, holo.;

NBG!, iso.).

Passerina sp. now I Bredenk. & A.E.van Wyk: 70 (2000); Bredenk.

& A.E.van Wyk: 56 (2001a); Bredenk. & A.E.van Wyk: 217 (2001b).

For a complete description see Bredenkamp & Van Wyk

in Bothalia 32: 76-79 (2002c). Distribution (Figure 9).

Consetyation status: Fower Risk [FR-lcj (Victor 2002).

This species is rare, but does not qualify for Red Fist sta-

tus under lUCN (2000) guidelines.

9. Passerina esterhuyseniae Bredenk. & A.E.van

Wyk, sp. nov., P. comosae C.H. Wright affinis. Bracteae

galeiformes, circumscriptione obovatae; lamina adaxi-

aliter concava, abaxialiter convexa, intra setosa, extra

glabra, tenuiter chartacea, laevis, concolorans, brunneo-

la, in marginem membranaceum vel in alas membran-

aceas expansa; basis cuneata; costa excurrens apicem

subacutum vel acutum faciens; margines ciliati dimidio

superiore. Elores membranacei, tempore pollinationis

flavidi, postea rubri ad brunnei.

TYPE. — Western Cape, 3218 (Clanwilliam): N Ceder-

berg Mountains, (-BB), Groenberg near Pakhuis, along

base of high rugged rock, rocky slopes, ±1 167 m, 27-

12-1956, Esterhuysen 26859 (BOF, holo.!).

Passerina sp. now 2 Bredenk. & A.E.van Wyk: 70 (2000); Bredenk.

& A.E.van Wyk: 56 (2001a); Bredenk. & A.E.van Wyk: 217 (2001b).

Shrubs or shrublets 0.3-0.5 m high. Stems greyish

brown, younger branchlets greyish tomentose; cork fine-

ly fissured, grey-brown, displaying whitish sclerenchy-

ma fibres at scars. Leaves imbricate on young branchlets,

closely appressed to stem, cymbiform, plane shape lin-

ear-lanceolate, length X depth ± 2.0 x 0.5 mm; lamina

inversely ericoid, adaxial surface concave, setose, abaxi-

al surface convex, glabrous; base sessile; apex rounded

into subacute point; margins sometimes ciliate. Inflores-

cences polytelic synflorescences; main florescences and

co-florescences spicate. Bracts enveloping flowers and

fruits, largest after anthesis of flowers, becoming more

coriaceous and rounded at fruit set, decussate, imbricate,

sessile, helmet-shaped, widely obovate in outline, length

X depth ± 3.1 X 2.4 mm; lamina adaxially concave

(inside), abaxially convex (outside), setose on inside,

glabrous on outside, thinly chartaceous, smooth, concol-

oroLis, brownish, extending into a membranous rim or

membranous wings; base cuneate; main vein extending

to form a subacute to acute apex; margins ciliate in dis-

tal half. Eloral envelope constituting hypanthium (fused

calyx and androecium) and sepals; membranous and yel-

lowish during pollination, dehydrated after shedding of

pollen, becoming papyraceous, turning red to brown, ±

4.6 mm long. Hypanthium a membranous cylindric tube,

indumentum at ovary and neck tomentose, neck ± 0.7

mm long, abscission tissue and articulation plane absent.

Sepals 4, petaloid, imbricate in bud, flexed in flower;

outer sepals concave oblong with apex adaxially tomen-

tose, abaxially setose; inner sepals concave, obovate

with apex adaxially glabrous, abaxially setose.

Androecium: filaments of antipetalous whorl + 0.4 mm

and those of antisepalous whorl ± 1 .2 mm long. Ovary ±

1.8 X 0.5 mm. Emit enveloped by persistent, loosely

ananged hypanthium fragmenting over widest circum-

ference of fruit, the fragmented hypanthium, sepals and

androecium being shed; an achene with pericarp mem-

branous and dry, ± 2.5 x 1.2 mm. Figure 10.

76

Bothalia33,l (2003)

Diagnostic characters and relationships: Passerina

esterhiiyseniae is easily distinguished from P. comosa by

its helmet-shaped bracts, which are widely obovate in

outline. The concolorous, brownish bracts are thinly

chartaceous and smooth in texture, the lamina extends

into a membranous rim or membranous wings and the

main vein elongates forming a subacute to acute apex.

The flowers are membranous and yellowish during polli-

nation and red to brown after shedding of the, pollen.

Eponymy: this species is dedicated to Elsie Ester-

huysen who diligently collected plants especially the

high-mountain flora of the Northern, Western and East-

ern Cape.

Distribution and ecology: Passerina esterhiiyseniae

has been collected on the northern Cederberg Mountains

at Groenberg near Pakhuis and at Konpoort (Eigure 1 1 ).

It is endemic to the Northwestern Centre within the CER.

The northern Cederberg area is covered by Mountain

Eynbos (Rebelo 1998). This species grows at the peaks

of mountain tops at altitudes of ± 1 167 m, or against

rocky slopes amongst high rugged rocks. Confined most-

ly to mountainous areas, this species is still undercollect-

ed. Pillans 7689 (BOL) collected on slopes near the road

SE of Redelinghuis has been classified under P. ester-

hiiyseniae, although these plants seem to be more robust

and grow at lower altitudes.

Conservation status: Lower Risk (LR-lc) (Victor

2002), as the population size of this species is probably

very small or restricted.

10. Passerina comosa (Meisn.) C.H. Wright in

Elora capensis 5,2: II (1915); Thoday; 158 (1924a);

Bond & Goldblatt: 432 (1984); Goldblatt & Manning:

683 (2000). Type: Little Namaqualand; Khamiesberg

Range, between Pedros Kloof and Leliefontein, 3000-

4()0() ft., Drege 2570 [K!, lecto., designated by Thoday:

388 (1924b); PRE!|.

R Jllifoniiis L. var. comosa Meisn.: 399 ( 1 840); Meisn.: 562 ( 1 857);

C.H.Wrighl: II ( 1915); Thoday: 158 (1924a).

R. fulcifonnis Drege: 68 (1843) nom. nud.

FIGURE 1 1 . — Known distribution of

Rasserina esterhuyseniae, •;

R. comosa. ■.

Small shrubs, (0.3-)0.6-0.9(-1.06) m high, branches

often virgate. Steins grey-brown; cork fissured length-

wise, grey-brown, suiTounding prominent leaf scars;

indumentum of young branches densely white or yellow-

ish tomentose, flaking off with cork on older branchlets,

which become glabrous. Leaves imbricate, overlapping ±

20%, diverging at an angle of 20°, plane shape linear to

narrowly lanceolate, length x depth (2.7-)3.0(^.0) x

0.6-0.9 mm, adaxial surface concave, villous, abaxial sur-

face convex, tomentose to villous, rugose or warty with

bases of fallen hairs; base sessile; apex obtuse to subacute;

margins glabrous, basally setose. Inflorescences with

spikes usually extended, 6-12-flowered, arrangement

subterminal, axis white-tomentose, proliferating growth

common. Bracts appressed, widely ovate, length x depth

(3.0-)5.5 X 1.5(-1.8) mm; lamina adaxially concave

(inside), abaxially convex (outside), villous inside,

tomentose to villous outside, coriaceous, rugose with

bases of fallen hairs, ± 3-ribbed on each side of main

vein, margins submembranous or wings extended and

abaxially setose to villous; base sessile; apex acute; mar-

gins glabrous, basally setose, involute. Floral envelope ±

7.3 mm long, yellow-pink during pollination, dehydrated

after shedding of pollen, turning red. Hypanthiiini

glabrous at ovary, neck tomentose, ± 0.9 mm long.

Sepals: outer sepals cymbiform, inner sepals elliptic,

outer and inner sepals adaxially glabrous, abaxially

tomentose. Androeciiun with filaments of antipetalous

whorl ± 0.7 mm and those of antisepalous whorl ± 1.5

mm long; anthers 0.7 x 0.4 mm, sub-basifixed, 2-thecous

and 4-locular. Ovary ± 2.7 x 1 mm. Fruit an achene with

pericarp membranous and dry, ±2.5x1 .2 mm, envelop-

ed by persistent, loosely arranged hypanthium, breaking

up at neck base due to dehydration and torsification of

tissue, resulting in the sepals and androecium being shed.

Figure 12.

Diagnostic characters and relationships: Passerina

comosa and P. qiiadrifaria both have abaxially hairy

bracts and are easily confused. However, these two

species are geographically segregated, with P. comosa

considered as a ‘north-western endemic’ of the Cape

Bora (Weimarck 1941 ), whereas P. qiiadrifaria is endemic

to the Karoo Mountain and Southeastern Centres. Mor-

Bothalia 33,1 (2003)

77

phologically P. comosa is less robust, interno(Jes are

longer, leaves a(Jhere closely to the stem and are gener-

ally more hairy and the bracts often have extended wings

that are abaxially setose to villous.

Etymology, the epithet comosa refers to the hairs on

the abaxial surface of the leaves, bracts and sepals, which

are characteristic of this species.

Distribution and ecology. Passerina comosa ranges

from mountain summits and slopes of the Kamiesberg to

Calvinia in the Northern Cape (Figure 11). In Western

Cape it is distributed in the area between 33° and 34°S lat-

itude and from 19° to 21°E longitude, with Primos 41

(PRE) as the most easterly outlier. This species is endem-

ic to the Northern Cape, as well as the Northwestern,

Southwestern and Karoo Mountain Centres within the

CFR. It occurs on the Roggeveld, Witteberg and the Klein

Swartberg Mountain Ranges. This species is found in sand

among rocks, on rocky ledges, on mountain summits, or

on SW-facing slopes at altitudes of 1 000-1 200 m.

Consenation status: Least Concern (EC) (lUCN Spe-

cies Survival Commission 2000).

1 1 . Passerina pendula Eckl. & Zeyh. e.x Tlioday in

Bulletin of Miscellaneous Information, Kew 4: 155

(1924a); Bond & Goldblatt: 433 (1984); Goldblatt &

Manning: 684 (2000). P. rigida Wikstr. var. comosa

Meisn.: 402 (1840); Drege: 584 (1847a); Meisn.: 563

(1857); C.H. Wright: 13 (1915) pro parte; Thoday; 155

(1924a). Type: Uitenhage Div., Zwartkops River, Eckloii

& Zexher 7381 [K!, lecto., designated by Thoday: 387

(1924b); BREM!, P!, PRE!, WU!].

P. pendula Eckl. & Zeyh. fide Drege: 210 (1847b) nom. nud.; Meisn.:

563 (1857) nom. nud.

Erect, many-stemmed shrubs, 1.0-1. 5 m high, young

branchlets often pendulous. Stems greyish brown; indu-

mentum of young stems densely white-tomentose, flak-

ing off on older branchlets, which become glabrous,

remaining indumentum forming lengthwise patterns with

FIGURE 12. — Passerina comosa.

Andraea 1288. A, inflores-

cence with apex growing out,

returning to vegetative growth

(proliferating growth); B,

leaf; C, bract; D, flower clasp-

ed by bract. Scale bars: 2 mm.

Artist: G. Condy.

grey-brown cork; brown leaf scars prominent. Leaves

greyish green, imbricate, overlapping ± 25%, appressed,

plane shape rhombic, length x depth 2. 5-3.4 x 1.0-1. 2

mm, adaxial surface concave, comose, abaxial surface

convex, softly coriaceous and smooth, glabrous; base

sessile, abruptly tapered; apex subacute; margins brown-

ish setose. Inflorescences with multiflowered main and

co-florescences; spikes usually extended, 6-16-flow-

ered, airangement subterminal, axis white-tomentose,

proliferating growth common. Bracts greyish green

when fresh, grey-brown in dried specimens, appressed,

rhombic, length x depth (3.0-)4.2 x 1.0(-1.5) mm; lami-

na adaxially concave (inside), abaxially convex (out-

side), comose inside, glabrous outside, softly coriaceous

and smooth, wings membranous; base cuneate; apex

obtusely angled; margins brownish setose. Eloral en-

velope ± 6.5 mm long, membranous and yellow-pink

during pollination, dehydrated after shedding of pollen,

turning red to brown. Hypanthium glabrous at ovary,

neck tomentose, ± 0.6 mm long. Sepals: outer sepals

cymbiform, adaxially scantily tomentose, abaxially

glabrous; inner sepals oblong, adaxially scantily tomen-

tose, abaxially glabrous. Androecium with filaments of

antipetalous whorl ± 1 .5 mm and those of antisepalous

whorl ± 2 mm long; anthers ± 0.8 x 0.4 mm, sub-basi-

fixed, 2-thecous and 4-locular. Ovaiy ± 2.0 x 0.7 mm.

Emit an achene with pericaip membranous and dry, ± 2.5

X 1.2 mm, enveloped by persistent, loosely arranged

hypanthium, breaking up at neck base due to dehydration

and torsification of tissue, resulting in the sepals and

androecium being shed. Eigure 13.

Nomenclatural notes: Passerina pendula, ascribed to

Ecklon & Zeyher, was first published as a nomen nudum

by Drege (1847b). This name was placed in synonymy

under P. rigida var. comosa by Meisner (1857). Wright

(1915) partly followed Meisner’s inteipretation of P. rigi-

da var. comosa, but in the citation of the specimens he

added all those that were later published as P. hurchellii

by Thoday ( 1924a). In his revision of Passerina, Thoday

( 1924a) reinstated the name P. pendula Eckl. & Zeyh. ex

Meisn., as the varietal name ‘comosa’ had already been

used at species level by Wright (1915). The present study

78

Bothalia 33,1 (2003)

FIGURE 13. — Passerina pendula,

Bredenkamp 908. A, inflores-

cence with apex growing out,

returning to vegetative growth

(proliferating growth); B, leaf;

C. bract; D, flower. Scale

bars: 2 mm. Artist: G. Condy.

regards Thoday’s publication of P. pendula as valid, as it

is accompanied by a Latin diagnosis, a description of the

species and it was published in 1924. As P. pendula Eckl.

& Zeyh. ex Meisn. was based on a nomen nudum, and

since Thoday ascribed the name to Ecklon & Zeyher, the

correct author citation for this species is P. pendula Eckl.

& Zeyh. ex Thoday. '

Diagnostic characters and relationships: Passerina

pendula is distinguished from P. burchellii by being

taller (up to 1.5 m), much-branched shrubs with pendu-

lous branchlets, with grey-green, softly coriaceous and

smooth leaves and yellow-pink membranous flowers that

are abaxially glabrous and adaxially scantily tomentose.

Etymology: the specific epithet pendula refers to the

pendulous branchlets of these shrubs as seen in their nat-

ural habitat.

Distribution and ecology: Passerina pendula is ende-

mic to the Southeastern Centre within the CFR. It is dis-

tributed on hills and slopes from the Kouga Mountains

in Western Cape to the Langkloof Mountains and the

Great Winterhoek Mountains in Eastern Cape (Figure

14). The species is also distributed along watercourses

as it occurs in the KwaZunga Catchment Basin and on

the banks of the Upper Swartkops River as well as the

Bushmans River at Port Elizabeth. P. pendula grows at

altitudes of ( 133-)383-600 m. On mountain slopes it is

often found in a belt above valley thicket and below

mountain fynbos. It grows in sand or shallow, gravelly,

sandy loam. The plants are frequent throughout the nat-

ural range of the species and a number of populations

are conserved in the Groendal Nature Reserve at Uiten-

hage.

Conservation status: Least Concern (LC) (lUCN

Species Survival Commission 2000).

12. Passerina galpinii C.H.Wright in Flora capen-

sis 5,2: 10 (1915) as P. galpini; Thoday; 161 (1924a);

Bond & Goldblatt: 433 (1984); Goldblatt & Manning:

683 (2000). Type: Cape [Western Cape], Riversdale

Dist., Melkhoutfontein, about 600 ft, 7 October 1897,

Galpin 4491 (K, holo.!; PRE!).

Bothalia 33.1 (2003)

79

c

FIGURE 15. — Passerina galpinii,

Bredenkamp 946. A, inflores-

cence with apex growing out.

returning to vegetative growth

(proliferating growth); B.

leaf; C. bract; D, flower

clasped by bract, hypanthium

fragmenting at circumfer-

ence of ovary. Scale bars: A,

3 mm; B-D, 2 mm. Artist: G.

Condy.

Stunted or erect, much-branched shrubs, (0.1-)0.3-1.0

(-1.2) m high. Stems light greyish brown; indumentum

of young stems greyish brown tomentose; cork grey-

brown, fissured lengthwise, with sclerenchyma fibres

protruding between fissures; leaf scars rounded. Leaves

greyish green, imbricate, overlapping 10%-25%, diverg-

ing at an angle of 30°, plane shape linear, incurved

towards apex, length x depth 2.9— 4.5 x 0.5-0. 8 mm,

adaxial surface concave, comose, abaxial surface con-

vex, chartaceous, glabrous; base sessile; apex subacute to

obtuse; margins glabrous, involute. Inflorescences with

spikes subterminal, usually congested, 8-10-flowered,

axis whitish tomentose, proliferating growth common.

Bracts appressed, oblate, length x depth (3.6-)4.5 x

1.1 (-2. 2) mm; lamina greyish green, adaxially concave

(inside), abaxially convex (outside), chartaceous,

glabrous outside, midrib extended into leaf-like point,

tomentose inside; wings straw-coloured, membranous,

broadly rounded, bullate; base cuneate; apex subacute;

margins glabrous, involute. Floral envelope ±5.3 mm

long, papyraceous and yellow-pink during pollination,

dehydrated after shedding of pollen, turning red to

brown. Hypanthium pubescent at ovary, neck pubescent,

± 1 mm long. Sepals: outer sepals concave elliptic, glabrous,

inner sepals obovate, abaxially glabrous, apex margin

adaxially tomentose. Androeciiim with filaments of anti-

petalous whorl ± 1 .4 mm and those of antisepalous whorl

± 2.1 mm long; anthers ± 0.6 x 0.5 mm, sub-basifixed, 2-

thecous and 4-locular. Ovary’ ± 2.0 x 1.1 mm. Fruit an

achene with pericarp membranous and dry, ± 2.5 x 1 .2

mm, enveloped by persistent, loosely arranged hypanthi-

um, breaking up at circumference of ovary, resulting in

the sepals and androecium being shed. Figure 15.

Nornenclatiiral notes: Passerina galpini C.H. Wright

(1915), published with a full description, but without a

Latin diagnosis, was accepted by Thoday ( 1924a). In the

present study the specific epithet is corrected to "gal-

pinii’ and the name accepted as validly published by

Wright, as the starting date for a Latin diagnosis as pre-

requisite for valid publication is 1 January 1935 (Greuter

et al. 2000).

Diagnostic characters and relationships: Passerina

galpinii is distinguished by its characteristic bracts,

which are oblate, with the midrib extended into a leaf-

like point; the lamina is cymbiform, greyish green, char-

taceous and glabrous, with the midrib adaxially tomen-

tose; the wings are straw-coloured, membranous, broad-

ly rounded and bullate. The distribution of this species is

also diagnostic as it is endemic to the Agulhas Plain

Centre within the CFR.

Eponymy: Passerina galpinii was named in honour of

the botanist E.E. Galpin. The holotype of this name,

Galpin 4491, was collected on 7 October 1897. At this

time, Galpin organized a collecting trip from Port Eliza-

beth via the Humansdoip, Knysna, George, Riversdale,

Swellendam and Caledon Districts to Cape Town and

increased his collecting numbers from 3531 to 4846. All

these specimens were probably identified at the Bolus

Herbarium in Cape Town, where he also spent a few

weeks (Gunn & Codd 1981).

Common name: Rebelo (1998) mentions the vernacu-

lar name Eliin gonna for this species.

Distribution and ecology: Passerina galpinii is ende-

mic to the Agulhas Plain Centre within the CER. It is dis-

tributed on stony flats, coastal limestone deposits and

limestone hills, from Elim to Bredasdorp, Arniston, Stil-

baai, Melkhoutfontein, Albertinia and Mossel Bay

(Figure 14); it grows at altitudes of 0-290 m. Plants reach

a height of ± 1 .2 m on stony flats, but become stunted on

southeast-facing slopes of limestone hills, overlooking

the sea. The plants are frequent in their natural environ-

ment. They are conserved in the De Hoop and Potberg

Nature Reserves and several private nature reserves. The

vegetation of the area is threatened by large stands of

Acacia cyciops (r-ooikmns), an alien invasive tree.

80

Bothalia 33,1 (2003)

Passerina galpinii is associated with Laterite Fynbos

(Rebelo 1998), occiiiTing on the Elim Flats of Western

Cape, which is characterized by gravelly, lateritic and

seasonally waterlogged soils. The present study also

indicates the presence of this species in Limestone Fyn-

bos, where it occurs on coastal limestone deposits.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

13. Passerina drakensbergensis Hilliard &

B.L.Burtt in Annals of the Kirstenbosch Botanic Gardens

15: 233 (1987); Hilliard & B.L.Burtt: 88 (1988). Type:

Natal [KwaZulu-Natal],' Harrismith, Bergville Dist.,

Cathedral Peak Forest Reserve, Ndedema River, 6000 ft,

5 November 1985, Hilliard & Burtt 18570 (E, holo.; K!,

M!, P!, PRE!, S!).

Erect, many-stemmed shrubs, ± 2 m high, much-

branched. Stems grey-brown; indumentum of young

stems closely white-tomentose, flaking off on older

branches, forming lengthwise patterns with patches of

yellowish brown cork; leaf scars prominent. Leaves

greyish green (glaucous), imbricate, overlapping ± 15%,

appressed, plane shape linear to lanceolate, longitudinaly

folded and somewhat keeled, length x depth (3.5-)

4.2-5.5(-6.5) X 0.8-2 mm, adaxial surface concave,

tomentose, abaxial surface slightly convex, rugose,

glabrous; base sessile, abruptly narrowed; apex subacute,

sometimes incurved; margins setose. Inflorescences with

multiflowered main and co-florescences; spikes usually

extended, 8-12-flowered, arrangement subterminal, axis

white-tomentose, proliferating growth common. Bracts

greyish green (glaucous), appressed, lanceolate, length x

depth 4.75-6.0(-7.0) x 1.5 mm; lamina folded length-

wise and keeled, adaxially concave (inside), abaxially

slightly convex (outside), villous inside, glabrous out-

side, coriaceous, obscurely ribbed and reticulately

veined; base cuneate; apex obtuse to acute; margins

sometimes narrowly membranous. Floral envelope ± 5.9

mm long; papyraceous and yellow-pink during pollina-

tion, dehydrated after shedding of pollen, turning red to

brown. Hypanthium scantily tomentose at ovary, neck

tomentose, ± 1.7 mm long. Sepals: outer and inner sepa-

ls concave lanceolate, outer sepals adaxially glabrous,

abaxially tomentose; inner sepals adaxially and abaxial-

ly tomentose. Androecium with filaments of antipetalous

whorl ± 1 .2 mm and those of antisepalous whorl ± 2 mm

long; anthers oblong, 0.9(-1.25) x 0.3 mm, sub-basifixed,

2-thecous and 4-locular. Ovary ± 2.4 x 0.9 mm. Fruit an

achene with pericarp membranous and dry, ± 2.5 x 1.2

mm, enveloped by persistent, loosely arranged hypanthi-

um, breaking up at neck base due to dehydration and tor-

sification of tissue, resulting in the sepals and androeci-

um being shed. Figure 16.

Diagnostic characters and relationships: Passerina

drakensbergensis is characterized by appressed leaves,

up to 6.5 mm long. The bracts are lanceolate, up to 7 mm

long, the apex is obtuse to acute without a leaf-like point

and membranous wings are absent. It may easily be con-

fused with P. montivaga and P. montana, both occuning

in the northern KwaZulu-Natal Drakensberg area.

P. montivaga has longer leaves (up to 8 mm) and bracts

with the midrib extended, forming a straight or filiform,

leaf-like point. The wings are ovate with margins hairy in

the distal half, or obtrullate, narrowing abruptly into the

midrib. P. montana can be separated by its terminal sub-

capitulate spikes and short leaves (up to 4 mm long),

which are linear to lanceolate, with a dilated base and

with a prominent median vein in the upper third of the

leaf, incurved at the acute apex. The bracts are ovate to

obovate in outline.

Passerina montivaga is a fynbos element which pos-

sibly originated in the southern Cape and dispersed east-

wards. Both this species and P. montana are distributed

from Eastern Cape via the Drakensberg Mountains

northwards to Zimbabwe. Although the distribution of P.

drakensbergensis, P. montivaga and P. montana overlap

in the northern part of the KwaZulu-Natal Drakensberg,

significant trends in the geographical and altitudinal

ranges of these species have been identified.

FIGURE 16. — Passerina drakens-

bergensis, Bredenkamp 1 02 1.

A, innoiescence with apex

growing out, retiiniing to

vegetative growth (prolifer-

ating growth); B, leaf; C,

bract; D, Bower clasped by

bract. Scale bars: 2 mm.

Artist: G. Condy.

Bothalia 33,1 (2003)

81

17° 18° 19° 20° 21° 22° 23° 24° 25° 26° 27° 28° 29° 30° 31° 32°

FIGURE 17. — Known distribution

of Passerina drakensbergen-

sis, O; P. coiyinbosa. ■.

Etymology, the specific epithet refers to the location

of this species in the northern KwaZulu-Natal Drakens-

berg.

Distribution and ecology: Passerina drakenshergen-

sis is endemic to the Bergville District in the northern

KwaZulu-Natal Drakensberg (Figure 17). It has been

collected from the Royal Natal National Park to Giant’s

Castle Game Reserve at altitudes between 1 500 and

1 980 m. It is frequent at Cathedral Peak above Ndedema

Gorge, where it is found in a marginal belt between for-

est and grassland. It also grows along streams and river-

banks and on mountain slopes.

Consen’ation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

14. Passerina corymbosa Eckl. ex C.H. Wright in

Flora capensis 5,2: 14 (1915); Thoday: 162 (1924a).

Type: Cape [Western Cape], Bredasdorp, Zwellendam,

Pupas Valley, Voormansbosch and Duivelsbosch,

Keurboomsrivier, October, Ecklon & Zeyher 40 ( 70.10)

(B, holo.t; W!, lecto., here designated; MO!, S!).

P. filifonnis L. var. vulgaris Meisn.: 399 (1840); Meisn.: 562 (1857);

Drege: 87 (1843). P. vulgaris (Meisn.) Thoday: 162 (1924a); Bond &

Goldblatt: 433 (1984); Goldblatt & Manning: 684 (2000). Type: locis

subhumidis secus rivulum. prope Paarlberg, alt. 1000-1500 ped. [sub-

humid locality along river, near Paarlberg. alt. 1000-1500 ftj. Nov.-

Dee. 1839. Drege s.n. [K, lecto., designated by Thoday: 388 (1924b);

S!, W!].

P. filifonnis sensu C.H, Wright: 11 (1915) pro parte, non L.

Much-branched shrubs. (0.3-)1.0-1.5(-2.5) m high,

younger branchlets leafy, older ones leafless, often arcu-

ate, indurate. Stems often arising from a woody, under-

ground rootstock, greyish brown; indumentum of young

stems closely whitish tomentose, forming lengthwise

patterns with cork on older branches, which gradually

become glabrous; cork fissured lengthwise, with scle-

renchyma fibres protruding between fissures; leaf scars

oblate. Leaves greyish green, drying greyish brown,

ascending, diverging at an angle of ± 40°, imbricate,

overlapping up to 50%, laterally compressed, plane shape

linear to narrowly lanceolate, length x depth (2.0-)3.2^.4

(-10.0) X 0.5-0. 8 mm, adaxial surface concave, tomen-

tose, abaxial surface glabrous; lamina longitudinaly fold-

ed, midrib distinct and somewhat keeled, with 2 or more

lateral veins visible as faint striae when fresh, base ses-

sile; apex tapered or acute; margins glabrous, involute.

Inflorescences with multiflowered main and co-flores-

cences; spikes usually extended, 10-16-tlowered, arrange-

ment subterminal, axis white-tomentose, proliferating

growth common. Bracts greyish green, ascending, rhom-

bic to obtrullate, distinctly angled, length x depth (4.4-)

5.1 X 1.4(-1.7)mm; lamina folded lengthwise and keeled,

adaxially concave (inside), abaxially slightly convex

(outside), shortly villous inside, glabrous outside, coria-

ceous; wings coriaceous, distinctly 4- or 5-ribbed; base

cuneate; midrib shortly extended into an acute apex;

margins glabrous, involute. Eloral envelope ± 6.2 mm

long, papyraceous and yellow-pink during pollination,

dehydrated after shedding of pollen, turning red to

brown. Hypanthium glabrous at ovary, neck scantily

tomentose, ± 1.6 mm long. Sepals: outer sepals cymbi-

form, inner sepals obovate, outer and inner sepals adaxi-

ally scantily tomentose, abaxially glabrous. Androeciiim

with filaments of antipetalous whorl ± 0.7 mm and those

of antisepalous whorl ± 1.4 mm long; anthers ovoid, ±

0.9 X 0.4 mm, sub-basifixed, 2-thecous and 4-locular.

Ovary ±2.1 x 1.1 mm. Emit an achene with pericarp

membranous and dry, ± 2.6 x 1 .5 mm, enveloped by per-

sistent, loosely aiTanged hypanthium, breaking up at

neck base due to dehydration and torsification of tissue,

resulting in the sepals and androecium being shed.

Diagnostic characters and relationships: Passerina

corymbosa is distinguished by its greyish green leaves

and grey-brown stems of which the older branchlets are

leafless, often arcuate and indurate. The leaves are lateral-

ly compressed with the distinct midrib somewhat keeled.

The rhombic to obtrullate (diamond-shaped) bracts are

always conspicuously angled and distinctly 4- or 5-

ribbed. The most diagnostic character in the leaf anato-

my of P. corymbosa is the presence of a hypodermal

sclerenchymatous sheath, illustrated by Bredenkamp &

Van Wyk (2001a). This species has always been con-

fused with P. filifonnis sensu lato, which has inconsis-

tently been distinguished by longer, filiform leaves. In

Western Cape P. filifonnis subsp. filifonnis is separated

82

Bothalia 33,1 (2003)

by widely obovate bracts, narrowing abruptly into a fili-

form point. Where the distribution of P. coiymhosa and

P. montivaga overlap in the southern Cape, the latter

species is distinguished by bracts with ovate wings and

margins that are hairy in the distal half. In Eastern Cape

P. montivaga is distinguished by obtrullate bracts nar-

rowing abruptly into a straight, leaf-like point.

Etymology, of all the species in the genus, Passerina

corymhosa is the most common, as it is adapted to a wide

range of habitats mostly in Western and Eastern Cape.

The specific Latin epithet corymhosa (= with a cluster of

flowers or of fruits) indicates the 10-16-flowered,

extended spikes usually arranged in multiflowered main

and co-florescences.

Common names: according to Van Wyk & Gericke

(2000), P. corymhosa (formerly known as P. vulgaris) is

also called hakkershos, a name that commemorates an

era when the official bakers in the Cape used this plant to

heat up their outdoor ovens.

Uses: from an agricultural point of view. Story (1952)

described P. corymhosa at Keiskammahoek as an un-

palatable bush, which remained undamaged from grazing,

among the few closely cropped specimens of Clijfortia

linearifolia and C. paucistaminea. However, the value of

P. corymhosa as a pioneer, and also in combatting ero-

sion, cannot be underestimated. This species is common-

ly found along roadsides and in other disturbed places. It

is one of the most successful species for the rehabilita-

tion of embankments along newly built roads in Western

and Eastern Cape. The plants are resprouters from

woody, underground rootstocks and are excellent sand

binders, often found on coastal sand dunes. Considering

the human impact and invasion of alien vegetation along

the Cape coast, P. corymhosa would be a natural pioneer,

combatting erosion in areas where alien vegetation is

cleared.

Distrihiition and ecology: except for a few outliers, P.

corymho.sa is endemic to Western and Eastern Cape, and

all the phytogeographic centres within the CFR.

Although this species is distributed from Clanwilliam to

Cape Town and eastwards to East London, it most com-

monly occurs in an area between the coast and the 33°S

latitude and from 18° to 29°E longitude (Figure 17).

Gerstner 105 (PRE), collected near Compasberg in the

Lady Grey District, represents the most northerly distri-

bution of P. corymhosa in Eastern Cape. The specimens

collected in KwaZulu-Natal are regarded as outliers, rep-

resenting remnants of a former wider distribution.

Hilliard 40H! (PRE), collected at the Ellesmere Farm in

Ngome (KwaZulu-Natal), is an anomalous specimen,

with a greyish appearance, infected by fungi and record-

ed from cliff faces. This specimen was classified as

P. corymhosa on the basis of the angular bracts and the

leaves that are laterally compressed. The other two speci-

mens, Herh. Poeppig s.n., probably collected before

1868, and Rudatis 1204 (PRE), collected in 1910, repre-

sent populations that have possibly succumbed to human

impact.

Passerina corymhosa is a species with a wide habitat

spectrum. It most commonly occurs as a pioneer along

roadsides over the whole range of its distribution. The

species is found in stony areas on mountain slopes, peaks

and mountain passes. Along the coastal region, it is often

found on the rear dunes. It also grows in river valleys and

on the banks of river mouths. This species is common in

the whole of the Fynbos Biome of the CFR. In Eastern

Cape it is found in all the above-mentioned habitats, but

also in grassland. Story (1952) reported that P. corym-

hosa is found in sourveld and mixed grassveld but that it

showed no sign of advancing into the sweetveld. In open

grassland this species is often clustered along stream-

banks or on rocky areas. P. corymhosa occurs at a range

of altitudes, from sea level up to 1 300 m.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

15. Passerina obtusifolia Thoday in Bulletin of

Miscellaneous Information, Kew 4.T57 (1924a); Bond &

Goldblatt: 433 (1984); Goldblatt & Manning: 683 (2000).

Type: Cape [Western Cape], Worcester Div., between

Osplaats and Tunnel Sidings 2000-3000 ft., Rogers 16703

[K!, lecto., designated by Thoday: 388 (1924b)].

Erect shrubs with several fairly stout stems, bare at

base, branching profusely higher up, (0.3-)0.8-1.8(-2.4)

m high. Stems ash-grey; indumentum of young stems

whitish tomentose, flaking off, becoming scabrid on older

branchlets and forming lengthwise patterns with cork,

which fissures lengthwise; sclerenchyma fibres protrud-

ing between fissures; leaf scars oblate. Leaves greyish

green, imbricate, spreading at an angle of 45°, plane

shape linear, straight or incurved, length x depth

4.0-8. ()(-l 2.0) X 0.9-1. 5 mm, adaxial surface concave,

comose, abaxial surface convex, coriaceous, glabrous;

base sessile; apex obtuse; margins glabrous, involute.

Inflorescences with multiflowered main and co-flores-

cences; spikes usually extended, 8-16-flowered, arrange-

ment terminal, axis whitish tomentose, proliferating

growth common. Bracts spreading, narrowly obtrullate,

length X depth (4.0-)5.8 x 1.5(-L7) mm; lamina greyish

green, adaxially concave (inside), abaxially convex (out-

side), coriaceous, glabrous outside, tomentose inside,

closely 2-ribbed at margins, midrib extending beyond

lamina into a leaf-like point, apex obtuse; wings absent;

base cuneate; margins glabrous, involute. Floral envelope

± 6.8 mm long, papyraceous and yellow-pink during pol-

lination, dehydrated after shedding of pollen, turning red

to brown. Hypanthium glabrous at ovary, neck tomentose,

± 1.8 mm long. Sepals: with outer sepals cymbiform,

adaxially tomentose, abaxially glabrous; inner sepals nar-

rowly oblong, adaxially tomentose, abaxially glabrous.

Androecium with filaments of antipetalous whorl ± 1.2 mm

and those of antisepalous whorl ± 2.2 mm long; anthers ±

1.1 X 0.5 mm, sub-basifixed, 2-thecous and 4-locular.

Ovary ± 2.5 x 1.1 mm. Fruit an achene with pericaip

membranous and dry, ± 2.3 x 1 .2 mm, enveloped by per-

sistent, loosely ananged hypanthium, breaking up at cir-

cumference of ovary, resulting in sepals and androecium

being shed. Figure 18.

Diagnostic characters and relationships: the growth

form of P. ohtusifolia may easily be confused with that

of P. corymhosa, but P. ohtusifolia is distinguished by

Bothalia 33,1 (2003)

83

FIGURE 18. — Passerina obtiisifolia. Bredenkamp 919. A, flowering

inflorescence; B. leaf: C. bract; D, flower clasped by bract,

fragmenting at circumference of ovary. E. F, achene: E, lateral

view, with basal fragment of membranous pericarp; F, ventral

view, with remnant of membranous pericarp. Scale bars: 2 mm.

Artist: G. Condy.

the spreading, linear, incurved leaves, with obtuse apices.

The bracts are narrowly obtrullate, with the midrib

extending beyond the lamina into a leaf-like point and

the lamina is closely 2-ribbed at the margins.

Intermediate forms between P. obtusifolia and P. trunca-

ta SLibsp. tnincata are found in the Karoo Desert

National Botanical Garden at Worcester. P. obtusifolia

usually occurs on the northern side of the southern Cape

mountain ranges in drier habitats and P.falcifolia is con-

fined to the summits of mountains and southwards

towards the southern Cape coast. Intermediates between

P. obtusifolia and P. falcifolia have been found on the

boundary between the two species, just north of the

Prince Albert Pass.

Etymology, the Latin specific epithet, obtusifolia, refers

to the obtuse apices of leaves and bracts, which are char-

acteristic of this species.

Common name: the vernacular name karoo gonna is

used by the local people at Genadendal.

Uses: according to the curator at the Museum in

Genadendal, P. obtusifolia was traditionally used by the

local people in the home industry of soap-making. The

plants were burnt and the alkaline ashes used to react

with the stearic acid in fat at boiling point, thus forming

soap. In the Robertson area these plants are used in the

wild flower industry. Bayliss 521 (PRE) is a voucher

specimen recorded in cancer research, but the results

must have been negative; these plants are not currently

known for their medicinal value.

Distribution and ecology: Passerina obtusifolia is

endemic to Northern, Western and Eastern Cape (Eigure

19). It is centred in a belt between 33° and 34°S latitude

and from 19° to 27°E longitude, comprising all the cen-

tres within the CFR — it is most common in the Karoo

Mountain, Langeberg and Southeast Centres. P. obtusi-

folia is distributed from Worcester in Western Cape, to

Grahamstown in Eastern Cape. The most northwesterly

distribution are the Vanrhyn’s Pass and the Hantamberg.

The Karoo National Park and the Nuweveld Mountains

in the Beaufort West District and the Lootsberg Pass at

Middelburg are the most northerly distribution in

Western and Eastern Cape. Hilliard & Burtt 14654 and

Acocks 20170 from the Lady Grey District are consid-

ered as outliers.

Passerina obtusifolia is well adapted to drier karoo

habitats and is often found on the north-facing aspect of

mountains in the southern Cape. It frequents the bound-

FIGURE 19. — Known distribution

of Passerina obtusifolia, •;

P. pahidosa. ■.

84

Bothalia33,l (2003)

FIGURE 20. — Passerina pahidosa.

Jangle 156. A, inflorescence

with apex growing out,

returning to vegetative growth

(proliferating growth); B,

leaf; C, bract; D, flower

clasped by bract. Scale bars:

2 mm. Artist: G. Condy.

ary of fynbos and karroid vegetation and is common in

the Little Karoo, growing at altitudes of (300-)670-l 400

(-1 700) m. Although this species occurs at high altitudes

on the summit of the Swartberg Pas's, it grows below the

snow line and does not occur on the highest peaks of

mountain ranges in its distribution range. It is common in

drier mountainous habitats, growing in shallow rocky

soil and between rocks on well-drained slopes. It also

grows amongst sandstone boulders of upper mountain

slopes and on stony ridges of mountain tops. On the

Hantamberg it has been recorded in renosterveld on the

flat, rocky, dolerite summit. On Jonaskop it grows in a

zone below the fynbos and is absent at the summit. This

species is also found amongst rocks in river valleys and

dry streambanks. The average height of these plants is

0.8-1. 8 m, but stunted forms have been recorded from

the arid Bergkwagga National Park, which is one of the

most northeasterly localities. P. ohtiisifolia is a very com-

mon species and amongst the dominant species within its

distribution range.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

16. Passerina paludosa Thoday in Bulletin of Mis-

cellaneous Information, Kew 4: 161 (1924a); Bond &

Goldblatt: 433 ( 1984); Goldblatt & Manning: 683 (2000).

Types: Cape [Western Cape], Simonstown, Cape Flats,

Riet Valley, in and near shallow vleis, November 1922,

Thoday 100 [K!, lecto., designated by Thoday: 388 (1924b);

C!, NBG!, PRElj.

Much-branched shrubs or small tree, up to 2 m high.

Stems fawn-coloured; indumentum of young stems

closely white-tomentose, forming lengthwise patterns

with cork on older branches, which gradually become

glabrous; leaf scars oblate, comose at upper rim. Leaves

erect, nearly straight, greyish green, imbricate, overlap-

ping ± 25%, appressed, plane shape narrowly lanceolate

to lanceolate, longitudinaly folded and somewhat keeled,

length X depth (3,0-)5.5-6.8(-10) x 0. 8-1.4 mm, adaxi-

al surface concave, tomentose, abaxial surface glabrous;

base sessile; apex acute, sometimes incurved, bearing a

persistent tuft of white, erect hairs; margins setose. In-

florescences with multiflowered main and co-florescen-

ces; spikes usually extended, 10-12-flowered, amange-

ment subterminal, axis white-tomentose, proliferating

growth common. Bracts greyish green, appressed, nar-

rowly obtrullate with leaf-like point, length x depth

(5.2-)7.0 X 1.8(-2.0) mm; lamina folded lengthwise and

keeled, adaxially concave (inside), abaxially slightly

convex (outside), basally setose inside, glabrous outside,

coriaceous, ± 2-ribbed; wings membranous with obscure

venation, glabrous; base cuneate; apex acute; margins

white setose. Floral envelope ± 7.2 mm long, papyra-

ceous and yellow-pink during pollination, dehydrated

after shedding of pollen, turning red to brown. Hypan-

thium shortly tomentose at ovary, neck strigose, ± 2.6

mm long. Sepals: outer sepals cymbiform, ad- and abax-

ially glabrous, apex setose; inner sepals oblong, ad- and

abaxially glabrous, apex setose with margins tomentose.

Androecium with filaments of antipetalous whorl

± 1.2 mm and those of antisepalous whorl ± 2.4 mm

long; anthers ovoid, ± 0.7 x 0.5 mm, sub-basifixed, 2-

thecous and 4-locular. Ovary ± 2.3 x 1 mm. Fruit an ach-

ene with pericaip membranous and dry, ± 2.4 x 1.2 mm,

enveloped by persistent, loosely arranged hypanthium,

breaking up at neck base due to dehydration and torsifi-

cation of tissue, resulting in sepals and androecium being

shed. Figure 20.

Diaf’uostic characters and relationships: Passerina

paludosa is a stout shrub up to 2 m high, occurring most-

ly in marshy ground on lowland flats. It is characterized

by erect, nearly straight, greyish green, imbricate, ap-

Bothalia33,l (2003)

85

pressed leaves, which are ± lanceolate. The bracts are

narrowly obtrullate, with the midrib and leaf-like point

stout and the apex acute. This species is distinguished

from P. filiformis subsp. filiformis which has filiform

leaves and widely obovate bracts, which narrow abrupt-

ly into a filiform point.

Etymology: the specific epithet paludosa refers to the

habitat of this species, namely marshy lowland flats;

from the Latin paliidosus (= marshy, swampy or boggy).

Distribution and ecology: Passerina paludosa is

endemic to the Southwestern Centre within the CFR

(Figure 19). Herbarium specimens dated from 1921 to

1995 show that this species used to be distributed from

sandy places along the Malmesbury Road (Acocks 2482),

along marshy areas of the Cape Flats and the Stellen-

bosch District to the Palmiet River at Elgin, the most

easterly locality. As P. paludosa was severely affected by

urbanization and invasion by alien vegetation in the Cape

Peninsula, it is currently confined to small marshy areas

east of Muizenberg.

According to Smuts ( 1996) the only three extant popu-

lations known, are at the Rondevlei Nature Reserve,

Zeekoevlei and along the Strandfontein Road. Label

information on Peterson 1263, collected in 1982, states

that the population at a housing estate site SE of Zee-

koevlei consisted of ± 400 plants, but Smuts (1966)

reported only 60 living plants. At the same time the popu-

lation at Rondevlei consisted of 35 plants and the one

along the Strandfontein Road of possibly a few hundred.

Currently both the Zeekoevlei and Strandfontein sites are

in danger of urban development and are being threatened

by invasive alien vegetation, primarily Port Jackson

{Acacia saligna) and rooikrans (A. cyclops). Conserva-

tion measures proposed by Smuts (1996) include an

environmental impact study at the Zeekoevlei site prior

to any development and a plea for urgent attention by

conservation authorities to ensure the conservation of the

Strandfontein population.

The Rondevlei Nature Reserve boasts more than 250

plant species of which many are rare and endangered.

Species associated with P. paludosa include Chondro-

petalum nudum, Juncus krausii and Leucadendron levi-

sanus. In recent years the management at the reserve

concentrated on restoring and managing its biodiversity.

Alien vegetation has been cleared, plant species that

occurred there historically have been re-introduced and

P. paludosa has been successfully propagated by cut-

tings to expand the population. As acidification is an

important effect of urbanization and as alien vegetation

impacts on the natural drainage system of an area, the

whole wetland east of Muizenberg can be conserved only

if it is included in the Rondevlei Nature Reserve (Smuts

1996).

Recently two new populations of plants, that appear to

be P. paludosa, were collected at Springfontein Earm

near Stanford [3419AD, Louw 7083 (NBG, PRE)], and

in seasonally wet clays at Heidehof, 5 km NW of Pearly

Beach [3419CB, Helme 2376 (NBG, PRE)]. These speci-

mens were not included in the distribution of P. paludosa

as further population studies need to be done. Taking

urbanization and invasion by alien vegetation into

account, the Red List status of P. paludosa was also not

changed.

Conserx’ation status: Vulnerable, [VU B 1 B2abcd] (Victor

2002).

17. Passerina montivaga Bredenk. & A.E.van Wyk

in Bothalia 32: 34 (2002d). Type: Natal [KwaZulu-

Natal], 2930 (Pietermaritzburg): hills above Pinetown,

2400 ft, (-DD), 3 December 1 89 1 , /M. Wood in PRE 49409

(PRE!, holo.; MO!).

R montivagus Bredenk. & A.E.van Wyk: 34 (2002d), sphalm.

P. filiformis L.; 559 (1753) pro parte, excluding type; Thunb.; 75

(1794): Wikstr.: 324 (1818); Thunb.: 374 (1825a); Meisn.: 562 (1857);

C.H.Wright: 10 (1915); Thoday: 159 (1924a); Bond & Goldblatt: 432

(1984); Hilliard & B.L.Burtt: 182 (1987); Goldblatt & Manning: 683

(2000).

Passerina sp. nov. 4 Bredenk. & A.E.van Wyk: 70 (2000); Bredenk.

& A.E.van Wyk: 56 (2001a); Bredenk. & A.E.van Wyk: 217 (2001b).

Eor a complete description see Bredenkamp & Van

Wyk in Bothalia 32: 34 (2002d).

Consen’ation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

18. Passerina filiformis L. in Species plantarum:

559 (1753); Thunb.: 75 (1794): J.C.Wendl.: 18 (1798);

Wikstr.: 324 (1818); Thunb.: 374 (1825a); Meisn.: 562

(1857); C.H.Wright: 10 (1915); Thoday: 159 (1924a);

Palmer & Pitman: 1587 (1972); Coates Palgrave: 648

(1977); Bond & Goldblatt; 432 (1984); Hilliard &

B.L.Burtt: 182 (1987); A.E.van Wyk & P.van Wyk : 68

(1997); Goldblatt & Manning; 683 (2000). Bredenk. &

A.E.van Wyk 32: 29 (2002d). Type: Passerina filiformis,

Linnean Herbarium 504.1 [EINN, lecto.!, designated by

Thoday; 388 (1924b)].

P. cupressina J.C.Wendl. nom. nud. Meisn.: 404 (1840); Meisn.:

563 (1857); Thoday: 159 (1924a). P. cupressoides Steud.: 273 (1841).

P. pectinata Lodd.: 18 (1816) nom. nud. Wikstr.: 347 (1818); Meisn.:

404 (1840); Meisn.: 562 (1857); Thoday: 159 (1924a).

Eor a complete description see Bredenkamp & Van

Wyk in Bothalia 32: 29-34, figs 1^ (2002d).

18a. subsp. filiformis

Eor a complete description see Bredenkamp & Van Wyk

in Bothalia 32: 31-33, figs 1-3 (2002d).

18b. subsp. glutinosa (Thoday) Bredenk. &. A.E.van

Wy’k

Eor a complete description see Bredenkamp & Van Wyk

in Bothalia 32: 33-34, fig. 4 (2002d).

Consen’ation status: Lower Risk, [LR-nt] (Victor 2002).

19. Passerina falcifolia (Meisn.) C.H.Wright in

Flora capensis 5,2; 10 (1915); Thoday: 158 (1924a);

Palmer & Pitman: 1585 (1972); Coates Palgrave: 648

86

Bothalia 33,1 (2003)

(1977); Bond & Goldblatt: 433 (1984); Goldblatt &

Manning: 683 (2000). P. filiformis L. var. falcifolia

Meisn.: 399 (1840); Drege: 118, 124 (1843); Meisn.': 562

(1857); C.H.Wright: 10 (1915); Thoday: 158 (1924a).

Type: Eastern Cape, Uitenhage, Vanstaadesberg [now

Van Staden’s Mountains], 1000-2000 ft, Drege s.n. (K!,

lecto., designated by Thoday: 388 ( 1924b)].

P.filifonnis L. var. divaricata Wikstr.: 325 ( l818);TlTOday: 158 (1924a).

Type: Caput bonae Spei [Cape Peninsula], Sparrman s.n.. Herb. Thimherg

9573 (UPS. holo.!). Herb. Swartzii (S, iso.!).

Much-branched, tall shrubs or small ornamental trees,

( 1.0-)1.2-2.0(-3.04) m high, branchlets often arcuate,

pendulous. Stems grey-brown; indumentum of young

stems closely whitish tomentose, forming lengthwise pat-

terns, with cork on older branches, which gradually

become glabrous, fissuring lengthwise, yellowish brown

lenticulae present; leaf scars oblate. Leaves greyish green,

drying greyish brown, falcate, linear to narrowly lanceo-

late, longitudinaly folded, triangular in section, length x

depth 3.9-5. 6 x 0.6 mm, adaxial surface concave, tomen-

tose, abaxial surface glabrous; base sessile; apex obtuse

or tapered; margins glabrous, involute; younger leaves

inclined, diverging at an angle of ± 60°, imbricate, over-

lapping up to 50%, older leaves horizontally spreading,

not imbricate. Inflorescences with multiflowered main

and co-florescences; spikes usually extended, lax, often

arcuate, up to 16-flowered, arrangement subterminal, axis

white-tomentose, proliferating growth common. Bracts

mottled grey-green, rose-tinted during flowering time,

ascending, imbricate, widely ovute, narrowing into a leaf-

like, falcate point, length x depth (5.3-)5.8 x 1.7(-2.0)

mm; older bracts folded lengthwise along midrib and

keeled, younger bracts adaxially concave (inside), abaxi-

ally slightly convex (outside), villous inside, glabrous

outside, coriaceous; wings chartaceous, distinctly ± 4-

ribbed and reticulately veined; base cuneate; apex obtuse

or tapered; margins ciliate in distal half. Floral envelope

± 8.4 mm long, papyraceous and yellow-pink during pol-

lination, dehydrated after shedding of pollen, turning red

to brown. Hypanthiinn scantily tomentose at ovary, neck

exserted, often arcuate, tomentose, ± 3 mm long. Sepals:

outer sepals concave, obovate, adaxially scantily tomen-

tose, abaxially glabrous; inner sepals obovate, adaxially

tomentose, abaxially glabrous. Androecium with fila-

ments of antipetalous whorl ± 1 .4 mm and those of anti-

sepalous whorl ± 2.1 mm long; anthers ovoid, ± 1 x 0.4

mm, sub-basifixed, 2-thecous and 4-locular. Ovary ± 2.5

X 0.7 mm. Fruit an achene with pericarp membranous and

dry, ± 2.5 x 1.4 mm, enveloped by persistent, loosely

arranged hypanthium, breaking up at neck base due to

dehydration and torsification of tissue, resulting in sepals

and androecium being shed. Figure 2 1 .

Nomenclatural notes: Wright (1915) overlooked the

combination P. filiformis L. var. divaricata Wikstr.

(1818), also indicated by Thoday (1924a), which is the

earliest name for the taxon. However, this name based on

Sparrman s.n. (Herb. Thunberg 9573) falls into syn-

onymy under P. falcifolia, as the name of a taxon does

not have priority outside the rank in which it was pub-

lished (Greuter et al. 2000).

Diagnostic characters and relationships: Passerina

falcifolia is distinguished by the mottled grey-green fal-

cate leaves, which are inclined or horizontally spreading.

The widely ovate bracts are villous inside and narrow

into a leaf-like, falcate point, with chartaceous wings that

are distinctly ± 4-ribbed and reticulately veined. The

most conspicuous floral character is the slender, often

arcuate, tomentose hypanthium neck, exserted from the

clasping bract. This species may be confused with P. fili-

formis suhsp. filiformis which has widely obovate bracts,

narrowing abruptly into a filiform point. The bracts are

basally to centrally setose on the inside and the wings are

glabrous. P. montivaga is another close species, but is

distinguished by bracts that are basally setose on the

inside, with glabrous wings.

Etymology: the specific epithet falcifolia is derived

from the Latin falcatus (= curved like a sickle), referring

to the falcate or sickle-shaped leaves of these plants.

FIGURE 21. — Pa.sserina falcifolia,

Bredenkamp 917. A. (lower-

ing intloreseence; B. leaf; C,

bract; D. flower clasped by

bract. E. F. anther: E. dorsal

view; F. ventral view. Scale

bars: 2 mm. Artist: G. Condy.

Bothalia33.l (2003)

87

17” 18” 19° 20° 21° 22° 23° 24° 25° 26° 27° 28° 29°

FIGURE 22. — Known distribution

of Passerimi falcifolia. •; P.

rubra, □.

Common names: Passerina falcifolia is also known as

the ‘Outeniqua gonna’ or the Oiiteniekwagonna, refer-

ring to the Outeniqua Mountains where it occurs (Coates

Palgrave 1977). Palmer & Pitman ( 1972) use the vernacu-

lar name forest gonna, as these plants are commonly seen

along roadsides on mountain passes of the southern Cape

forests. Von Breitenbach et al. (2001) use the names

Oiiteniekwa-gonna, gonnabas, gonnahos, kannabas and

Outeniqua gonna, gonna bush.

Uses: Passerina falcifolia is used for fuel or for mak-

ing cord (Palmer & Pitman 1972). According to label

information on Dahlstrand 1905 (PRE), the species is

cultivated by florists. Plants grow into small ornamental

trees and could be used more widely in horticulture.

According to Grobbelaar 63 (PRE), P. falcifolia is a host

to members of the insect genus Eremmis.

Distribution and ecology: Passerina falcifolia is asso-

ciated with forests and Mountain Eynbos (Rebelo 1998)

in the southern Cape and the southern parts of Eastern

Cape. It is a near-endemic to the CER and occurs in the

Karoo Mountain, Southwestern and Southeastern Centres,

as well as the Zuurberg, Blaauwkrantz and Alexandria

Eorests of Eastern Cape. It most commonly occurs in a

belt between the coast and the 33°S latitude and from 22°

to 26°E longitude (Eigure 22). The two specimens.

Brown 25975 and Rogers 28858, collected near Caledon

in October 1924, are regarded as outliers, possibly indi-

cating a wider previous distribution of the species into

areas with woody vegetation in Western Cape. P. falcifo-

lia occurs from Meiringspoort, in the Oudtshoorn area, to

Ruytersbosch in the Mossel Bay area, and along the

Outeniqua, Tsitsikamma and Great Winterhoek Mountains

to the Grahamstown area.

Passerina falcifolia is found on mountain plateaus

and southeast-facing slopes on Table Mountain Sand-

stone in shallow, sandy loam soil. Plants commonly occur

along forest margins, in open patches, or disturbed areas

along roadsides. This species is also found in coastal

regions and riverine fynbos. P . falcifolia grows at a range

of altitudes, from sea level up to 1 100 m.

Conservation status: Least Concern (LC) (lUCN Spe-

cies Survival Commission 2000).

20. Passerina rubra C.H. Wright in Elora capensis

5,2: 12 (1924a); Thoday; 156 (1924a); Bond & Gold-

blatt: 433 (1984); Goldblatt & Manning; 684 (2000).

Type; Cape [Western Cape], Ladismith, Muiskraal, near

Garcia Pass, 1200 ft, 3 October 1897, Galpin 4492

[K!, lecto., designated by Thoday; 388 (1924b); GRA!,

PRE!].

P. filifonnis L. var. squarrosa Meisn.: 562 (1857); Thoday: 156

( 1924a). Type: Eastern Cape, prope Cradockstad et Port-Elisabeth [near

CradockTown and Port Elizabeth], October, Zexher 3779 (S!, lecto., here

designated; BM!, MEL!, NBG!, W!).

Much-branched, erect shrubs, with rigid branchlets

and inflorescences or smaller, extensively branched, round-

ed shrublets under arid, calcareous habitat conditions,

(0.2-)0.3-0.75(-l . 1 ) m high. Stems: older ones grey-brown,

indurate, and sclerenchyma fibres exposed; young stems

reddish brown, indumentum whitish tomentose, forming

lengthwise patterns with cork on older branches, which

gradually become glabrous; cork Assuring lengthwise;

internodes longer than leaves during prolific lengthening

of branchlets or shorter under arid conditions. Leaves

greyish green, ascending, appressed, decussate and rigid,

or under arid conditions, imbricate (overlapping 5-30%),

appressed or ascending, diverging at an angle of up to

30°; lamina naiTowly lanceolate or oblong, longitudinaly

folded, triangular in section, length x depth 2.4-4.3 x 0.7

mm, adaxial surface concave, tomentose, abaxial surface

glabrous; base sessile; apex obtuse; margins glabrous,

involute. Inflorescences with conspicuous, multiflow-

ered main and co-florescences; spikes robust, rigid,

extended, narrowly ellipsoid, with rows of enlarged,

decussate, pointed bracts, 20-30-flowered, arrangement

subterminal, axis white-tomentose, proliferating growth

common. Bracts grey-green, rose-tinted during flower-

ing time, ascending, imbricate, widely ovate, midrib

shortly extended into a point, length x depth (4.3-)5.1 x

1.8(-2.0) mm; older bracts folded lengthwise along

midrib, younger bracts adaxially concave (inside), abax-

ially slightly convex (outside), villous inside, glabrous

outside, coriaceous; wings widely ovate, chartaceous, ±

5-ribbed, reticulately veined; base cuneate; apex acute;

margins ciliate in distal half. Floral envelope ±8.4 mm

long, papyraceous and yellow-pink during pollination,

dehydrated after shedding of pollen, turning red to brown.

Hypanthiiim glabrous at ovary, neck exserted, sparsely

pubescent, ± 2 mm long. Sepals: outer sepals cymbiform.

Bothalia33,l (2003)

FIGURE 23. — Passerina rubra, Bre-

denkamp 914. A, inflores-

cence with apex growing out,

returning to vegetative growth

(proliferating growth); B,

leaf; C, bract; D, tlower clasp-

ed by bract; E, fruit clasped

in tomentum of bract, envelop-

ed by hypanthium, fragment-

ed at neck base. F, G, achene:

F. enveloped by membranous

pericarp; G, side view. Scale

bars: 4 mm. Artist: A. Stadler.

ad- and abaxially glabrous, inner sepals obovate, adaxi-

ally scantilly tomentose, abaxially glabrous. Androecium

with filaments of antipetalous whorl ± 0.7 mm and those

of antisepalous whorl ± 1.7 mm long; anthers ovoid, +

0.9 X 0.3 mm, sub-basifixed, 2-thecous and 4-locular. Ovary

±2.1 X 1.1 mm. Fruit an achene with pericaip membra-

nous and dry, ± 2.1 x 1.2 mm, enveloped by persistent,

loosely arranged hypanthium, breaking up at neck base

due to dehydration and tofsification of tissue, resulting in

the sepals and androecium being shed. Figure 23.

Nomenclatural notes: as the starting date for a Latin

diagnosis is 1 January 1935 (Greater et al. 2000), P.

rubra is a valid name, although it was published with a

full description, but without a Latin diagnosis, by Wright

(1915). The combination P. filiformis L. var. squarrosa

(Meisner 1857), was overlooked by Wright (1915), but

mentioned in synonymy by Thoday (1924a). In the pre-

sent revision all the type material cited by both Wright

(1915) and Meisner ( 1857) was studied. Galpin 4492 (K)

was selected as the lectotype of P. rubra by Thoday

( 1924b) and Zeyher 3779 in S was selected as lectotype

for P. filiformis var. squarrosa as it is internationally

available in many herbaria.

Diagnostic characters and relationships: the distribu-

tion of P. rubra partly coincides with that of P. corym-

bosa, P. montivaga and P. falcifolia. P. rubra is a small-

er shrub (average height 0.3-0.75 m), often occurring in

calcareous soil. It is distinguished from the other three

species which are taller (average heights 1-2 m), and

especially from P. falcifolia, which is a tall shrub or a

small tree (up to 3.04 m), often associated with indige-

nous forests. P. rubra may also be separated by the inllores-

cences which have extended, robust spikes, with up to 30

fertile, enlarged bracts. The bracts are typified by the

midrib which is shortly extended into a point and by the

wings which are adaxially tomentose, widely ovate, char-

taceous, ± 5-ribbed and reticulately veined. Flowers are

distinguished by the exserted hypanthium neck, which is

± 2 mm long and glabrous to sparsely pubescent.

Etymology: the specific epithet rubra was derived